CONTENTS PART | (Issued 31 December 2000) ALLSOPP, P.G. Revision of the Australian genus Aramalamorpha Attow (Coleoptera: Scarabaeidae: * Dryaastisak) with anew species from the Wet Tropics of Queensland. ..............2.-0-.00 2-4-0, 1 BAEHR, M. Some genera and species of ground beetles new to Australia (Coleoptera: Carabidae). ......... 9 BALLANTYNE, L.A. & LAMBKIN, C. Lampyridae of Australia (Coleoptera: Lampyridae: Luciolinae: Luciolini).................. 15 BOUCHARD, P. Cuemus, 2 new genus of Tenebrionidae (Coleoptera) from the northern Queensland Wet Tropics... .........24 Bulaet sas Mae Talts «Ma Nal alle tl os Pans fle « oleePe dale Blan sal (= Mel och} ol eraBent 95 CERMAK, M. & HASENPUSCH, JW. Distribution, biology and conservation status of the peppermint stick insect, Megacrania batesii (Kirby) (Phasmatodea: Phasmatidae), in Queensland... ...... 0.00.0. e eee eee an en. KO CRANSTON, P.S. Three new species of Chironomidae (Diptera) from the Australian Wet Tropics. ..........4: 107 DAVIES, V.T. & LAMBKIN, C. Wabua, a new spider genus (Araneae: Amaurobioidea: Kababininae) from north Queenstand, Australia i420 24 pea¢ cs Sega Se be teat bate etaage peated iitestitess peters 12D GROVE. §,J, ‘Trunk window trapping: an effective technique for sampling tropical saproxylic beetles... .. 149 ITARVEY, M.S. A review of the Australian schizomid genus Notozomus (llubbardiidae). 2... 0 ee 161] HOWDEN, H.F, & STOREY, RL. New Stercomerini and Rhyparini from Avstralia, Borneo and Fiji (Coleopteray Scarabaeidae: AphOOunaey | os oes dt etd pk ete bee Bett Pieter 4g ee ee bt ta 175 KOHOUT, RJ. A review of the distribution of the Polyrhachis and Echinopla ants of the Qucensland Wet Tropics (Hymenoptera: Formicidae: Formicinae)... .,.....5. 062. 0e eee e ae ee, 183 O'KEEFE, S,T, & MONTEITH, GB. Clidicus abbotensis O'Keefe, a new spevics of Scydmaenidae (Coleoptera: Staphylinaides} from Australia with description of the larva. .,.........- PrepOat eas shames de ses 211 RAVEN, R.J. A new species of funncl-web spider (Hadronyche: Hexathelidac: Mygalomorphae) from MGrth Queensharid ook oi ec ee alse ensfecepeelecemebece ejere odnere baba jee sje a alceel lee alle eecabe 225 REID, C.A.M. A complex of cryptic species in the genus. Captedactyla Burmeister (Coleoptera: Scarabaeidae: Copang) 3b 2S Be ee alae Hilla geile oodles stellate gio dyithe ged ted 231 REID, C.A.M. & STOREY, RI, Reyision of the dung beetle genus Temboplectran Westwood (Coleoptera: Scarabaeidae: SATA BAB ENED foe alte pst tte daeeterm atone cM fo otal cata ut abe af alle ata} fal wale GMa [ew fo Ola] OMe afta EL yp ew . 253 SINCLAIR, D.P. Two hew genera of Tessaratomidae (Hemiptera: Heteroptera: Pentatomoidea).......5..5-55 299 SINCLAIR, DP. A generic revision of the Oncomerinae (Heteroptera: Pentatomoidea: Tessaratomidae)....... 307 SLANEY, D.P. New species of cave dwelling cockroaches in the genus Neotemnopteryx Princis (Blattaria: Blattellidae: Blattellinae).. 6... 6. ee eee cee eeu ete ee eee er eeee 331 STANISIC, J Taxonomy of the Australian rainforest snail, Helix bellendenkerensis Brazier, 1875 (Mollusca; Eupulmonata: Camaenidae)... 2... pe ee eee eect ety serene 337 STOREY, RJ. & MONTEITH, G.B. Five new species of Aptenocanthon Matthews (Coleoptera: Scarabaeidae: Scarabaeinae) from tropical Australia, with notes on distribution ......0.. 0.0.00. e cee eee tue a ees 349 CONTENTS (continued) LAMPRELL, K., STANISIC, J. & CLARKSON, P. Spondylids from the Mediterranean Sea and Atlantic Ocean (Mallyusca: Bivalvia: Spondylidae). . 611 LAMPRELL, K., STANISIC, J, & CLARKSON, P. Some spondylids from the Pacific Ocean (Mollusca: Bivalvia; Spondylidae)..,,.....2..444 623 LIMPUS,.C.J. A breeding population of the Yellow-bellicd sea-snake, Pelamiy platurus, in the Gulf of GarPenbarig i aie asd a ats 3d gia e's Hela i's ellad bet ates cee ee be At oe elected 629 LIMPUS, C.J., DE VILLIERS, D.L., DE VILLIERS, M.A., LIMPUS, D.J. & READ, M. A, The loggerhead turtle, Caretta careita in Queensland: feeding ecology in warm temperate WALES sta iiss c ecd oa pe baie eet ae ae tt oot He Pb obese bate ne Sta! eee elec: om 631 McGUIGAN, K.L. An addition to the rainbowfish (Melanotacniidac) fauna of North Queensland..,......,;... 647 JONINSON, I.W., RANDALL, J.E. & CHENOWETH, S.F. Diagramma melanacrum, new species of haemulid fish from Indonesia, Borneo and the Philippines with a generic review .. 2.0.2.2... wet iedes fatdactvtoipas sa dagel », 657 NORMAN, M.D. New octopus species from Queensland. 0.0.0.0... 0. 50sec cece cee erences 677 OTTO, L.C. tlalacaridae ftom the Great Barrier Recf lagoon and Coral Sea: Halacarellus and Halacarus (Acarina: Halacaroidea). 0.00... eet ent beeen ees 691 OTTO, LLC. Halacaridae of the Great Barrier Reef Lagoon and Coral Sea: the Copidognathus ornatus group (Acarina: Prostigmata: Halacaridae)............-.0-.6 beet do a-tty wo ¥ oh es 717 RICHARDS, 8.J, Anew species of torrent-dwelling frog (Anura: Hylidae: Litoria) from the mountains of Indonesian New Guinea (West Papua) ....... . - piles deed: 24 Pali ce tng tt de jebheye lao ROWE. C., STANISIC, J., DAVID, B. & LOURANDOS, H. The helicinid land snail Pleuropoma extineta (Odhner, 1917) as an environmental indicator im archaeology... 4 ps. ee pe ee cee ge pene eg ewe ss peer tepereeceeret i xbe oe 741 LOHMANN, U. & SACHS, S. Observations on the postcranial morphology, ontogeny and palacobiolagy of Sclerocephalus haeuseri (Amphibia: Actinodontidae) from the Lower Permian of Southwest-Germany , . . 771 TIMMS, B.V. A new species of Calamoecia (Copepoda: Calanoida) from arid Australia, with comments on the calanoid copepods of the Paroo, northwestern Murray-Darling Basin,...,.....- 783 NOTES HANNAH, D. & THURGATE, N.Y. Range extensions for two poorly known Queensland snakes... 0. ee eee 400 COVACEVICH, J.A,, BUFFETT, A-F., COUPER, PJ. & AMEY, AP. Herpetological ‘foreigners’ on Norfolk Island, an external territory of Australia ..;.,,...... 408 McGROUTHER, M,A, First record of the Larye-looth Cookie-Cutter Shark Jsistius plutodus trom Australian SALOES ss 2 oj ot ep yee oe Peat ye th ta ae alle tb te es fg tp ee oe rey! 442 PALMER, R.A. Southern range extension for the Delicate Mouse (Pseucomys delicatulus)....0..0.00 0.040. 460 McGROUTHER, M.A, First record of Elsman‘s Whipnose Anglerfish, Gigantactis elsmani{Lophiformes: Gigantactinidae), from Australian waters....3....0.e000085 prhtitpeta taste: 646 ARANGO, CP, Sea spiders (Pycnogonida) from the Great Barrier Reef, Australia, feed on fire corals and PRM TAS yg oe jae ee cjert- at a! |e “eof pre ct fo ahs] leo alincte alle b ajiete-e|iole i cris Settee He Hse este 656 PART 2 (Issued 30 June 2001) BAER, B. & JOCQUE, R. Revisions. of genera in the Asteron-complex (Araneae: Zodariidae): new genera Pentasteron, Phenasteron, Leptasteron and Subasteran ... 6. en eens 359 CANNON, L.R.G. & SEWELL, KB. A review of Temnosewellia (Platyhelminthes) ectosymbionts of Cherax (Crustacea: Parastacidae) in Australia... 6... ce ee eee eet te ete bette eteeeenes 385 COOK, A.G, & CAMILLERE, N. Additions to the rostroconch fauna of Australia and China .... 2... c ese e ete eee eta eee 401 DALL, W. Australian species of Aristeidae and Benthesicymidae (Penaeoidea: Decapoda). .,..,..,.... 409 DAVIES, V.E. & LAMBKIN, C. A revision of Provambridgea Forster & Wilton, (Araneae: Amaurobioidea: Stiphidiidae)..... 443 DETTMANN, MLE. & CLIFFORD, ELT. The fossil record of Elacocarpus L. fruits... 02... 0.6. eee ex ghetes USE e AST eT Le 461 EWART, A. Dusk chorusing behaviour in cicadas (Homoptera: Cicadidae) and a mole cricket, Brisbane, QueénSlard 44 5. te atatte ls bree tre er litet asec eet Cates Peto a eb rteeet 499 FAUBEL, A, & CAMERON, B. Turbellaria from salt marshes of Coomera River, southeastern Queensland, Australia ........ 511 FRITH, C.B. & FRITH, D.W. Bowerbird (Ptlonorhynchidac) biometrics, with observations on sexual dimorphism and intraspecific variation 2... 25. cs eee ae peer e ene cece bees ect ewe seas teataetes 521 GUILBERT, E. Western Pacific Tingidae (Heteroptera): new species and new records.................05 543 HALLETT, $.L., ERSEUS, C., O7 DONOGHUE, Pu. & LESTER, RG. Parasite fauna of Australian marine aligachactes 2.0.0... cece etre eee er eee eeeree 555 HEALY, J.M., LAMPRELL, K. & KEYS, LL. The ‘ water-trap’ spiny oyster, Spondylus varius Sowerby, |827.G.B. Rewer fs 1827 (Mollusca: Bivalyia: Spondylidae) from Australia,....... er. ed resraeremn,: pat E ety 577 KOFRON, C,P. & SMITH, R. Status of Estuarine Crocodiles in the populated coast of northeast Queensland. ............ 603 LAMPRELL, K., STANISIC, J. & CLARKSON, P. Spondylids from ihe Mediterranean Sea and Atlantic Ocean (Mollusca: Bivalvia: Spondylidae).. 611 LAMPRELL, K., STANISIC, J, & CLARKSON, P. Some spondylids ftom the Pacific Ocean (Mollusca: Bivalvia: Spondylidac)..........0....4 623 LIMPUS, C.J. ' A breeding population of the Yellow-bellied sea-snake, Pelamis platurus, in the Gulf of Catpentariat eM chet. i tettets Meet ete nee pate beep tate Pk ee ie OS eed 629 LIMPUS§, C.J., DE VILLIERS, D.L., DE VILLIERS, M.A., LIMPUS, D.J. & READ, M.A. The loggerhead turtle, Caretta caretta in Queensland: feeding ecology in warm temperate Waletey aes apr $4 dae lett et aentitiee aati ebowine ened acta nig ats 631 MeGUIGAN, K.L. An addition to the rainbowfish (Melanotacniidac) fauna of North Queenstand..,..,......, . 647 JOHNSON, J.W., RANDALL, J.F. & CHENOWETH, S.F. Diagramma melanacrum, new species of haemulid fish from Indonesia, Borneo and ihe Philippines with a genetic review - 6.6... eee ce ee net cnet eet e es 657 NORMAN, M.D. New octopus species from Queensland... ... tigen eyptea topes epee eee att 617 OTTO, IC. Halacaridae from the Great Barrier Reef lagoon and Coral Sea: Halacarellus and Halacarus (Acarina: Hafacaroidea), ..,. - etishepabteité Lteettseyadepnsc testis 691 OTTO, I.C. Halacaridae of the Great Barrier Rcef Lagoon and Coral Sea: the Copidognathus ornatus group (Acarina: Prostigmata: Halacaridae).... 00.0.2 0 cea eee eee sees 717 RICHARDS, S.J. A new species of torrent-dwelling frog (Anura: Hylidae: Litoria) from the mountains of Indonesian New Guinea (West Papua) .......... 0.00. c eee eee ee eee eens 733 ROWE, C., STANISIC, J., DAVID, B. & LOURANDOS, H. The helicinid land snail Pleuropoma extincta (Odhner, 1917) as an environmental indicator in archaeology .......... Sct Mapa dpb x ah Sob eyes duke, vlac else Ryle! satel eto dt adele a fdeatt And 741 LOHMANN, U. & SACHS, S. Observations on the postcranial morphology, ontogeny and palaeobiology of Sc/eracephalus haeuseri (Amphibia: Actinodontidae) from the Lower Permian of Southwest Germany ... 771 TIMMS, B.V. A new species of Calamoecia (Copepoda: Calanoida) from arid Australia, with comments on the calanoid copepods of the Paroo, northwestern Murray-Darling Basin........... 783 NOTES HANNAH, D. & THURGATE, N.Y. Range extensions for two poorly known Queensland snakes ........... 0.00 c cece ee eee 400 COVACEVICH, J.A., BUFFETT, A.F., COUPER, P.J, & AMEY, A.P. Herpetological ‘foreigners’ on Norfolk Island, an external territory of Australia ............ 408 McGROUTHER, M.A. First record of the Large-Tooth Cookie-Cutter Shark Jsistius plutodus from Australian TA RCE a at Sea ecap lo ora marion bag by briny ny 8 yea ocaaen Ag abba oe y tie gle fatarte Harp: 6 pabhcebicaes carp eh ag od agra bead any 442 PALMER, R.A. Southern range extension for the Delicate Mouse (Pseudomys delicatulus).. 0... 0.206 oe es 460 McGROUTHER, M.A, First record of Elsman’s Whipnose Anglerfish, Gigantactis elsmani (Lophiformes; Gigantactinidae), from Aus- tralian waters ,........-.- wh ants aPC Mabh iy b lets St bdatd pare i Tedeevs Shaker hs 646 ARANGO, C.P. Sea spiders (Pycnogonida) from the Great Barrier Reef, Australia, feed on fire corals and OAM vaste sie alle ahs oes bie a pe ks Sapetiotle oid ajtete bed jemubepeleld ely caine slie + slaps 656 REVISIONS OF GENERA IN THE 4STERON-COMPLEX (ARANEAE; ZODARIDAE): NEW GENERA PENTASTERON, PHENASTERON, LEPTASTERON AND SUBASTERON BARBARA BAEHR AND RUDY JOCQUE Baehr..B. & Jocqué, R. 2001 06 30; Revisions of genera in the 4sferen-complex (Araneae: Zodariidae); new genera Peniasferon, Phenasreron, Lepiasteron and Subusteren, Memoirs of the Queensland Muse 46(2); 359-385, Brisbane, ISSN 0079-8835 Four genera are erected to accommodate 13 new species in the Asferon-complex, a large yroup of Australian Zodariidae. Penresterar is characterised by a cymbial concavity. lis type species is simplex (6 2) and it is further represented by parasimpley (4), intermedi (22), securiter(d 2), ascitas (6 2). sardidum (a ), storosoides (2) and isohelae (a 2). Phenasieron is created [or the types species Jongiconductar (3) and machinasum (2). Leplasteran includes the type species plaryveanducior (2) and vexf/um (2), Both the later penera are characterised by a sclerolised basal swelling on the sublewulum anda huge DTA with refolded distql margin, They differ mainly by the structure of the carapace. cymbium and tegulum. Svbaxteran contains only the peculiar type species deaviesue (dF) diagnosed by shape of the carapace and of the J abdomen and a prolateral tegular apophysis on the nale-palp. T draneue, Zodariidie, Asteron-comiplex, new genera, dustraliu, &. Baehr. Queensland Musenm, PO Box 3300. Santh Brishune 4101, Australia (e-mail harbarubtagm.gid.gav.au); R. Jacque, Inveriebrate Section, Royal Africa Museum, B-3080 Terviven, Belgilin (e-mail jacquetvajricamuseum be); 13 February 2001. This ts the third contribution to the systematics of spiders belonging to the large As/eron- complex (Baehr & Jocqué, 1996). Jocque & Baehr (2001). and Baehr & Jocqué (2000) revised Asleron Joequé, 199]. 10 contain 8 species. and Pseudasteron, Cavasieron and Minasteron were erected with 1, 12 and 3 species, respectively. The present paper describes another 4 genera with §, 2, 2 and 1 Species. Whereas the former genera were fairly easily defined as they exhibit clear autapomorphies, it was much more difficult to unite the species that were placed in the basal groups of the cladogram for the As/eron-complex that was presented by Baehr & Joeque (1996). ‘The original idea was (0 create Pentasteren lor the 5 basal groups in the cladogram. It proved, however, that this was impossible due to the lack of synapomorphies for this grouping that would have created a paraphyletic taxon in the absence of Asferon s. str, Therefore, We have erected 3 new genera: Subasteron, is monotypic whereas Phenasteron and Leptasteron each contain 2 species united by characters of the male palp and carapace shape. MATERIAL AND METHODS Descriptions follow Jocque & Baehr (1992), Abbreviations: ALE, antertor lateral eyes: AME, anterior median eyes; AS, anterior spinnerets; E, embolus; EA, embolar apophysis; DTA, dorsal tegular apophysis; F, femur; FL. flange: LTA, lateral tegular apophysis; MS, median spinnerets; MT. metatarsus, MOQ, median eye quadrangle: P. patella; PE. prolateral extension of tegulum, PLE, posterior lateral eyes; PME, posterior median eyes; PS, posterior spinnerets: PTA, prolateral tegular apophysis; T, tibia; VTA, ventral legular apophysis. Abbreviations of institutions where material was borrowed: AM, Australian Museum, Sydney: KBIN, Koninklijk Belgisch Institul voor Natuurwetenschappen; QM, Queensland Museum, Brisbane; SAMA, South Australian Museum, Adelaide; VM, Victoria Museum, Melbourne; WAM, Western Australian Museum, Perth; ZSM, Zoologische Staatssammlung Muenchen. SYSTEMATICS Pentasteron gen, nov. TYPE SPECIES, Pentasteran simplex sp. nov, ETYMOLOGY, Greek pretix, penta = five; witli zenerje name, Asterar; referring to 5 basal groups of Astenon- votnplex (Baehr & Jocqué. 1996), Gender ts neuter: DIAGNOSIS. Member of dAsteroa-complex, with 2 palp having tibia with a deep retrolateral concavity combined with a pronounced concavity on the base of the cymbiuin, having tegulum with a broad base traversed by the seminal duct and ending in a typical median apophysis (VTA) with curved tip, usually having embolar apophysis (EA) of variable length. 360 MEMOIRS OF THE QUEENSLAND MUSEUM B “{ ‘ a f \ e « N _ Yr - EB AYN 5 iN \ 7 — em Ps, aaa =f I } \ 7 ——"\ — el AA Fe aes \3 ] ‘| y\ ( } \4 | \] i pp ] \y | / | \ NO, ar \ ) / \ | \ —L \ | f . } | \ fet / ‘a Y} vay, _ rh 4 ws | 4 “ EG fe 8 Roa | | aie © cae \ . /\ “Wy = ed FIG. 1. Pentasteron simplex. A, body dorsal; B-D, cephalothorax; B, ventral; C, frontal; D, lateral. Scale 1mm. DESCRIPTION. Small to medium-sized spiders (3.00-7.00) with slightly granulate tegument. Carapace widest at coxae II, narrowed to ().60-0.65 maximum width in both sexes. Profile flat with highest point between fovea and PME (Fig.1A,B). Colour: carapace dark brown, chelicerae and sternum medium brown; legs with strongly contrasting white to dark brown femora; coxae pale, trochanters dark, other legs yellow or brownish. Eyes (Fig. 1C, D) in 3 rows (2-4-2). Only ALE in first row, AME (in the middle) and PLE in second, third only PME. Eyes subequal or ALE smaller than rest. MOQ slightly longer than wide. Clypeus straight, about 3 times diameter of ALE; with few hairs. Chilum single, short wide, without setae. Chelicerae as usual for family with a few hairs in front and dense row on distal promargin; no teeth. Labium narrowed at base; sparsely haired. Gnathocoxae rather elongate; sparsely haired; with anteromesal scopula. Sternum flat; shield-shaped with straight anterior margin and tiny triangular extensions between coxae. No inter- nor precoxal sclerites. Legs: formula 4123. Spination: few spines on pairs I, II, more numerous on III, IV. Paired tarsal claws with numerous (+12) teeth on I and II, with + 7 on III and IV. Unpaired claw toothless, on very small onychium. Trichobothria in two rows on T, ina single row on Mt and t. Hinged hairs present but few, restricted to dorsal side of TI and II. Metatarsal preening brush on Mt I] and III poorly developed. Abdomen oval; mostly without sigilla but some species with two dorsal and a prolateral one on either side. Spinnerets: AS short, conical, with very short distal segment; MS and PS very short, absent in males. Colulus represented by group of setae. Tracheal spiracle hidden by well developed anterior lip; posterior lip sometimes sclerotised and protruding from under anterior lip. REVISION OF THE ASTERON-COMPLEX 361 Male palp (Fig. 2C,D): tibia with large lateral concavity delimited by solid dorsolateral apophysis and ventrolateral apophysis, most often swollen along its lower lateral margin. Cymbium with basal concavity, simple unmodified flange, several spines near distal tip. Tegulum with broad base carrying transverse section of seminal duct; distal part extended in typical median apophysis (VTA) with curved tip. Embolus emerging on prolateral part of tegulum. Several species with split relatively short, rigid embolus, dorsal prong of variable length. LTA usually short and thorn-like. DTA membranous, simple. Epigyne: external structure simple, with central depression, sometimes double: copulatory ducts starting near centre or slightly in front, running towards the side and backward to enter simple, thick-walled spermathecae. Female palp with finely dentate claw. KEY TO THE SPECIES OF PENTASTERON 14 -TWialesy ube s OA gle ass Goes Jd ae 2 Females : 024: tt baer pte ee bee 9 2. Embolusnotbifid ...............2005 3 Embolus bifid (Figs 3E,H,4C,F,I, prolateral view), . . 5 3, Tegulum with very large prolateral extension (PE), guiding embolus(Fig.5A) ............. P. securifer Tegulum without suchextension. 2.2, 2.2.0.0. 4 4. Tegulum with large half funnel-shaped VTA (Fig. SC,D) ¢ RP oe te Y pes Se + ERS P. isobelae VTA not half funnel-shaped, but straight (Fig. 3A) te fe Pe ace ad Hh fo fechas ae ol OU be eas P. parasimplex Cymbium with very large retro-basal cymbial concavity delimited by triangular flap(Fig.4B) .. . . P. oscitans Cymbium with small concavity (Figs 3D,G,4E,H). . . 6 6, EAatbasethickerthanembolus(Fig.3H)... 2... . ee a en ee P. intermedium EAat baseotthickerthantmbdlus (Figs 3E,4F 1) .. 7 7. EA bifid at distal end; dorsolateral tibial apophysis recurved(Fig.4D-F).. 20... 0.02. P. storosoides EA not bifid at distal end; dorsolateral tibial apophysis not recurved (Figs 3E, 41) & 8. EA longer than embolus, clearly visible in ventral view uw (Fig. 4G a & age Hp a ae ae P. sordidum EA shorter than embolus, not visible in ventral view (Fig. (9 14 a a ee P. simplex 9. Central part of epigyne clearly delimited, with inverted v-shaped ridge, or at least rebordered in front (Figs 6E,F, PRB sl es ae fee we ee a ee af by 11 Central part of epigyne poorly delimited, front never rebordttéd ..0. rey ate wee tn a Bde x 12 11. Epigyne with inverted u-shaped ridge only in front (Fig. PAA ii ele aoe ey ae oD ety yee P. scurifer Fpigyne with inverted v-shaped ridge for more than half ofepigyne(Fig.6E).........4.- P intermedium 12. Posterior margin of epigyne clearly indented (Fig. 6C) ee gs Pore gas ee eB ee aed es P. oscitans Posterior margin sometimes sinuous but not indented (Figs6A,7C) 13. Posterior margin of epigyne straight (lig. 7C) MS eee at de a tock 3" P. isobelae Posterior margin of epigyne sinuous (Fig. 6A) o ghee wh o Pale e ch lew oy SES P. simplex Pentasteron simplex sp. nov. (Figs 1A-D, 2A-D, 3C-E, 6A,B, 15) ETYMOLOGY. For the simple genitalia. MATERIAL. HOLOTYPE: ¢&, Lake Broadwater, via Dalby, SE Qld, pitfalls site 1, 13.i-25.i1.1986, QM & M. Bennie (QMS15746). PARATYPES: Queensland: 58 ¢, 62, together with holotype (QMS52610; 2d in KBIN; 1312 in ZSM); 1d, Christmas Creek, xi.1912, E. Mjéberg (RMS). NSW: 1d, Myall Lakes NP, 32°30°S 152°21°E, 14.xii.1996, L. Wilkie (MLCO/05) (AM KS 55653); 12, Myall Lakes NP, 32°37’S 152°12°E, 14.xi.1996, L. Wilkie (MLIO/03) (AM KS 55654); 13, as previous; 32°17’S 152°12°E, 15.xii.1996 (MLIO1/01) (AM KS55650); 1 2, Myall Lakes NP, 32°37’S 152°12°E, 14.xii.1996, L. Wilkie MLIO1/01) (AMKS55651); 1d, 15.xii.1996, further as previous (MLIO1/05) (AM KS55652); 3¢. Booti Booti NP, 32°16’S 152°31°E, 13.xi1.1996, L. Wilkie (BBIO1/09) Syprsanegt Id,as previous (BBIO1/06) (AMKS55644); 12, as previous, (BBIO1/01) (AMKS55645); 1d, as previous (BBIO1/05) (AM KS55648); 1 2, as previous, 13.xi.1996 (BBIO2/01) (AM KS55647); 23, as previous (BBIO1/09) (AM KS55659); 1 2, as previous (BBIO01/09) (AM KS55643); 12, as previous (BBIO2/09) (AMKS55649); 1d, as previous, 32°14°S 152°32°E, 14.xii.1996, L. Wilkie (BBCO2/07) (AM KS55646); 1 2, Munmorah State Rec., 33°13’S 151°34’E, 16.x1i.1996, L. Wilkie (MUNIO2 /04) (AM KS55655); 1 2, Wyrrabalong NP, 33°16’S 151°32°E, 16.xii.1996, L. Wilkie (WYRCO02/07) (AM KS55658); 1d, as previous, 16.xi.1996 (WYRCO02/10) (AM KS55657); 1d, as previous, 16.xi.1996 (WYRCOOI/09) (AM KS55656); 2d, Ramornie SF, Main Ck, track off Mt. Tindal, 29°43’S 152°38°E, 4.ii-9.iii,1993, M. Gray & G. Cassis (AMKS39135); 1¢, Ramornie SF, Mt Tindal, 29°42’S 152°35°E, , 4.ii-9.i11.1993, M. Gray & G Cassis (AMKS 39136); 5d 12, Ramornie SF, track off T- Ridge Rd, Mt Tindal, 29°33’S 152°38’E, 4.ii-9.i11.1993, M. Gray & G Cassis (AMKS391 34). DIAGNOSIS. ¢¢ are recognised by simple palpal organs: dorsolateral tibial apophysis with broad base, split embolus with thin and short EA; 2 epigyne simple with a longitudinal pale zone in middle; the copulatory openings are at margin of this zone just in front of centre. DESCRIPTION. Male (holotype). Total length 3.56; carapace 1.85 long, 1.22 wide; tibia+patella 11.59, Colour: Carapace medium brown with darker radiating striae; chelicerae and sternum reddish brown; coxae pale with dark pro- and retrolateral spots; trochanters dark; proximal half of femora white with dark proximal ring, distal half dark brown; patellae brownish yellow suffused with dark brown on sides and with dark distal ring; tibiae brownish yellow, II, IIT and IV darkened on ventral and lateral sides. Abdomen dark sepia to black; dorsum with narrow dark brown scutum in front and 7 white spots: 2 pairs in anterior half, 3 in a row in front of spinnerets; sides with large oblique white spot; venter uniform dark sepia with 2 small yellow spots in front of epigastric fold. Carapace finely granulated; sternum not granulated. Eyes: a: 0,10; b: 0.10; c: 0.11; d: 0.12; e: 0.02; f 0.02; g: 0.04; h: 0.08; AL-AL: 0.18. MOQ: AW = 0.84 PW; AW = 0.78 L. Clypeus 0,32 or 3,2 times ALE. Chilum single, 0.08 high, 0.38 wide. 186Km Aeoag4s6 MEMOIRS OF THE QUEENSLAND MUSEUM Spination: F P T Mt | plid2 - v2 v4 Il pl3d2 - v2 v5 dw3 If] pl2d3rl2 plidirll — pl2d2rl2vl-2-2 10disp dw5 IV pl2d4ril plidirll = pl2d2rl2vi-2-2. 10disp dw5 No hinged hairs. Male palp (Fig. 3C-E): tibia with large retrolateral concavity delimited by ventrolateral lamellate apophysis with swollen lateral margin and dorsal, short, blunt downpointing apophysis. Cymbium with shallow basal concavity and fairly long flat flange(FL). Embolus short, rigid, curved outward, split at base, EA thin, short, only visible from prolateral side. VTA short, sturdy; DTA membranous, attached to dorsal part of VTA, Female (paratype). Total length 4.31; carapace 1.98 long, 1.35 wide; tibiat+patella I: 1.58. 1SkU 4154 FIG. 2. Pentasteron simplex; A, cephalothorax dorsal; B, epigyne; C,D, left male palp; C, ventral; D, retrolateral. REVISION OF THE ASTERON-COMPLEX Colour: exactly as in male. Eyes: a: 0.10; b: 0.10; c: 0.12; d: 0.12; e: 0.04; f: 0.04; g: 0.04; h: 0.10; AL-AL: 0.20. MOQ: AW = 0.86 PW; AW = 0.71 L. Clypeus 0.34 or 3.4 times ALE. Chilum single, 0.07 high, 0.36 wide Spination: F P T Mt 1 plid2 - v2-2 v2-1-2 Il plid2 - v2-1-2 v4 dw3 Hl pl3d3rll plidirll = pl2d2rl2v2-2-2. 9disp dw5 IV pl2d3rl1 plidirll = pl2d3rl3v2-2-2 Odisp dw5 Hinged hairs: TI d1, TI dl. Epigyne(Fig. 6A,B): very simple: with hardly sclerotised plate with slightly concave posterior rim; copulatory openings in front, fairly closely set under semicircular darker shields; internal structure showing through translucent epigyne. Copulatory ducts very broad at entrance, describing more than one loop before entering small lateral spermathecae. Variation: colour pattern and size very stable: do carapace length 1.82-1.87 and width 1.20-1.22; 2 carapace length 1.92-1.98, width 1.33-1.40. DISTRIBUTION. Known only from type locality. Pentasteron parasimplex sp. nov. (Figs 3A,B, 15) ETYMOLOGY. Similar to P. simplex. MATERIAL. HOLOTYPE: 3, Wyperfield NP, Victoria, Dattuck track, Eucalyptus foecunda leaf litter, 2.vii.1982, M. Harvey & B. Roberts (WAM). DIAGNOSIS. Coloration very uniform; 3 d with swollen ventrolateral tibial apophysis and very short LTA. DESCRIPTION. Male (holotype). Total length 4.03; carapace 2.08 long, 1.40 wide; tibia+patella 11.85. Colour: Carapace, chelicerae and sternum pale brown, patternless; legs uniform yellowish brown; abdomen dark sepia: dorsum with narrow brownish frontal scutum and five pale spots, 2 pairs and single one in front of spinnerets; venter pale sepia. Carapace and sternum smooth. Eyes: a: 0.14; b: 0.10; c: 0.12; d: 0.12; e: 0.02; f 0.04; g: 0.06; h: 0.10; AL-AL: 0.26. MOQ: AW = 1.00 PW; AW = 0.75 L. 363 Spination: F P T Mt I plid2 v2-1-2 v2 i plid2 - v1-2-1 v4 HT pl3d3rl2—s plidirll ~— pl2d2rl2v2-2-2_7disp dw6 IV pl2d4rl| plidirl] pl3d3rl3v2-2-2. 9disp dw6 Hinged hairs: TI d1, TH d1. Abdomen with 2 dorsal sigilla and a lateral in front on either side. Male palp (Fig. 3A,B): tibia with deep retrolateral concavity; ventrolateral apophysis swollen, rounded at the back, blunt in front; dorsolateral one fairly short and thick, sharp, pointing outwards; cymbium with fairly long flat flange, with small proximal indentation; embolus short, rigid, curved outwards; without EA; VTA short, rigid, blunt; DTA membranous; LTA very short with thick base and sharp tip. Female unknown. DISTRIBUTION. Known only from type locality. Pentasteron intermedium sp. nov. (Figs 3F-H, 6E,F, 15) ETYMOLOGY. Refers to its intermediate taxonomic position. MATERIAL. HOLOTYPE: ¢, Augusta, Cave Break road, Western Australia, 34°20’S 115°09°E, Agonis & moss litter, 24.vii.980, S. & J. Peck (WAM 90/170-1). PARATYPES: 1 subadult 3, together with holotype; South Australia: 1¢, Blinman, 8-19.xii.1986, post office, on floor, 31°05’S, 138°40°E; M. Dykshoorn (SAMA); 1d, Kolay Hut, 32°33’S 135°36’E, 10.x1i.1989, on ground, D. Hirst (SAMA N1992120); New South Wales: I d , Federal Highway on NSW/ACT border; 35°12’S_ 149°12°E, 10.v.1992, J. Hunt (AM KS49459); Victoria: 29 2d, Barr Ck, Kervins Rd, Cohuna, 35°48°30"S_ 144°10°30"E, 1 May 1999, watering, J. Hooper, D. & J. Shield, J. Woodman (S30490); 1 2, Upper Lurg, 36°35’S, 146°11°E, col. J. Strudwick 14, Jan 1997 (JSt 529); 14d, same data as previous (JSt 550); 1 2, same data as previous, 4 Apr 2000 (JSt 717); 12 1d, Spring Gully, 36°37°48"S, 144°15°177E, J. Shield, 22-30 Dec 1993 (CVIC 777); 1°, same data as previous, 18 Jan 1994 (CVIC 790);22 26, Barr Creek, Cohuna, 35°48.5’S, 144°10.5’E col. J. Hooper, D & J Shield, J. Woodman 1 May 1999, watering (CVIC 785). DIAGNOSIS. Males have a unique combination of sclerites in the palp: large VTA, small spine-like LTA and large curved EA accompanying embolus along its dorsal side. Female has a simple epigyne with clearly delimited inverted v-shaped ridge in front, pale zone on posterior half. 364 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 3. Pentasteron spp. right male palps. A,B, P. parasimlex, A, ventral; B, retrolateral; C-E, P. simplex; C, ventral; D. retrolateral; E, prolateral; F-H, P. intermedium;, F, ventral; G, retrolateral; H, prolateral. Scale 0.5mm. DTA = dorsal tegular apophysis, E = embolus, EA = embolar apophysis, LTA = lateral tegular apophysis, ST = subtegulum, VTA = ventral tegular apophysis. REVISION OF THE ASTERON-COMPLEX DESCRIPTION. Male (holotype). Total length 3.42; carapace |.72 long, 1.18 wide; tibiatpatella 11.50. Colour: Carapace medium brown with faint radiating striae and V-shaped dark mark in front of fovea. Chelicerae and sternum medium brown, slightly suffused with black. Coxae pale, trochanters pale brownish yellow suffused with black on sides; femora dark brown with paler dorsal lines, in distal third, pale in proximal part; patellae, tibiae and metatarsi orange brown, tibiae with slightly darkened sides. Abdomen dark sepia to black; dorsum with 4 pale spots in frontal half and 3 pale chevrons, posterior 2 anatomising, in front of spinnerets; sides with oblique pale stripes, venter pale sepia with broad pale median stripe and a spot on either side in front of dark ring around spinnerets. Carapace very finely reticulated; sternum smooth. Eyes: a: 0.09; b: 0.10; c: 0.10; d: 0.10; e: 0.02; f: 0.02; g: 0.04; h: 0.08; AL-AL: 0.18. MOQ: AW= 0.83 PW; AW = 0.62 L. Clypeus: 0.3 or 3.0 times diameter of ALE. Spination: F P T Mt i plid2 - vi-l v2 IT plld2 - vi-l v2-1dw3 Il pild3 plidirll pl2d2rl2v2-2 8isp dw6 IV pl2d4 plidirll pl3d3rl3v2-2-2— Bisp dw6 One dorsal hinged hair on tibiae I and II. Male palp (Fig. 3F-H): tibia with large retrolateral concavity delimited by fairly flat ventrolateral lamellate apophysis with dorsal swelling and roughly triangular (as seen from above) flat dorsolateral apophysis with denticle at its dorsal base. Cymbium with fairly extensive flange. Embolus fairly short and rigid, curved outward, split at base, the dorsal prong (EA) thick and well developed. VTA well developed and strong; DTA membranous, attached to tegulum dorsad of VTA; LTA a short spine-shaped excrescence between sperm-duct and VTA. Female (paratype). Total length 4.83; carapace 2.08 long, 1.33 wide; tibiatpatella I: 1.58. Carapace medium brown with faint darker radiating striae; chelicerae and sternum medium brown; sternum pale brown; coxae pale, brownish towards base: trochanters brownish yellow suffused with black on sides; femora white with small basolateral ring in proximal half, dark brown in distal half; remainder of legs yellowish brown, tibiae suffused with dark on sides; abdomen dark: dorsum with 2 pairs of white spots in frontal halfand 3 pale chevrons ina row in front of spinnerets; sides with 2 or 3 pale stripes; venter pale sepia with pale median stripe and a spot on either side in front of dark ring around spinnerets. Lung covers yellow. Eyes: a: 0.10; b: 0.10; ¢: 0.14; d: 0.12; e: 0.04; f: 0.04; g: 0.04; h: 0.08; AL-AL: 0.18. MOQ: AW= 0.80 PW; AW = 0.66 L. Clypeus 0.36. Chilum single, 0.08 high, 0.48 wide. Spination: F P T Mt 1 plld2 - v2-2-2 v2 II plid2 - vl-2-2 v2-2 dw3 Il pl2d3rl2 plidirll = pl2d2rl2v2-2-2— &disp dw6 Hinged hairs: TI dl, TI dl. Preening brush on Mt II and II. Epigyne (Fig. 6E,F): clearly delimited inverted v-shaped ridge in front, pale zone on posterior half; copulatory ducts showing trough tegument; copulatory ducts large, strongly sclerotised, directed diagonal, ending in poorly delimited, adjacent caudal spermathecae. Variation: d carapace length: 1.72-1.84, carapace width: 1.18-1.30. Colour of carapace in South Australian specimens darker; leg colour stable; abdomen without spots or with 5 tiny spots; slight variations in shape of palpal sclerites, such as curvature of VTA and EA. DISTRIBUTION. Western and South Australia. Pentasteron securifer sp. nov. (Figs 5A,B, 7A,B, 15) ETYMOLOGY. Latin: securis, an axe, refers to shape of the huge, H-shaped VTA. MATERIAL. HOLOTYPE: <2, John Forrest NP, Western Australia, (31°52’S, 116°04’E) 1967, GH. Lowe (WAM 90/272). PARATYPES: 2 9, Jarrahdale Mine site, Western Australia, 32°13°S 116°04’E,12-18.iv.1999, KEG. Brennan (WAM 99/2378-2379). DIAGNOSIS. 6 ¢ have a very typical palp with very large, H-shaped VTA of which retrolateral distal part is axe-shaped; 2 2 have an epigyne with fairly deep, roughly diamond-shaped central depression, clearly delimited in front. DESCRIPTION. Male (holotype). Total length 4.79; carapace 2.40 long, 1.63 wide; tibiat+patella 12.45. Colour: Carapace medium brown with faint darker radiating striae and V-shaped pattern in 366 tront of fovea; chelicerae medium brown, paler alone median margins sternum pale brown, slightly darker along margin; coxae pale yellow, trochanters pale brown; femora pale in proximal half, brown in distal half; tibiae [ pale yellow, II, Ut and IV shghtly darker shghtly suffused with dark on sides; metatarsi and tarsi brownish yellow: abdomen sepia: dorsum with narrow brownish frontal scutum and six pale spots, | pair in front, one in the middle, and two in Iine in front of spinnerets; sides with large oblique pale area; venter pale, darker in front of epigastric furrow, Carapace and sterium smooth, Cyes:a: 0.72:b: 0.12; 0: 0.14: d: 0.14; 6: 0.02: 6: 0.04; 2: 0.06; h: 0.12; AL-AL: 0.28. MOQ: AW = 0,94 PW: AW = 0.83 L, Spmation: h P y Mi pid? v2 Welds iW plid2 : vie) vlele2e2 HW plidaek — pl2rh pl2d2ri2\22-2 Sdisp die Iv place) Hinged hairs: Thdl, Tid. Male palp (Fig. 5A,B): tibia with deep retro- lateral concavity, delimited by 2 large apophyses: ventrolateral apophysis flat, rebordered along margin, dorsolateral one roughly quadrangular with anterior comer drawn out into short sharp prong; cymbium with flat flange provided with backward directed prong, and with poorly defined distal haired ridge thus forming concavity; embolus long and slender, originating on posterior end of tegulum which has huge, roughly H-shaped VTA; its retrolateral distal prong is broad, axe-shaped and runs subparallel with membranous DTA; retrolateral part broadly connected with prolateral part which accom- panies the embolus on its ventral side; LTA tiny spine; without EA. Female (paratype). Total length 4.36; carapace 2.40 long, 1.62 wide; tibia+patella I: 2.06. Carapace dark brown with faint darker radiating striae; chelicerae and sternum medium brown; sternum pale brown; coxae pale, yellowish towards base; trochanters dark with pale yellow ventral patch; femora white with small basolateral spots in proximal half, dark brown in distal half} remainder of legs yellowish brown, tibiae suffused with dark on sides; abdomen dark; dorsum with 6 white spots, | pair in front, | in the middle, and 2 in a row in front of spinnerets: sides with large oblique spot; yenter fairly pale, paler centrally; lung covers yellow. plidirl) — plid2rjay2-2-2 iOigp dwe MEMOIRS OF THE QUEENSLAND MUSEUM Eyes: a: 0.10: b: 0.703.c: 0.01; di 0.12: e: 0.03: £ 0.03; g: 0.05; h; 0.08; AL-AL; 0.19, MOQ: AW = 0.86 PW; AW = 0.76 .L. Clypeus 0.32, Chilum single, 0.06 high, 0.22 wide. Spination I P U MI | plid2 - yl ‘2 iL plid2 - vl-2-2 v2 dwt th plad2ri2 so plidiril = pl2d2ri2v2-2-2 Bdisp dws Winged hairs: Tl dl, TH dl, Preening brush on Mt and IIL. Epigyne (Fig. 7A,B): with fairly deep, roughly diamond-shaped central depression, deeper in front, there delimited by posteriorly indented plate; copulatory ducts clearly showing trough tegument; copulatory ducts large, strongly sclerotised, ending in poorly dehmited, adjacent, caudal spermathecae. DISTRIBUTION, Known only from type locality. Pentasteron oscitans sp. noy. (Pigs 4A-C, 6C,D, 15) ETYMOLOGY. Latin aveitare. accommodating; relers lo the shape of the male palpal eymbiumn. MATERIAL, HOLOTYPE: ¢, Barrington Tops State Forest. 1.4 km S along Bungaree trail from Barrington Tops Forest Rd, NSW, 31°56°S 151°21°E, 4.ii-9,iv, 1993, LE80m. (NPWS survey), M. Gray & G Cassis (AM KS039485). PARATYPE: 19. together with holotype. DIAGNOSIS. ¢ have a very large cymbial fold, 29 differ in the shape of the posterior indentation of the epigyne. DESCRIPTION, Male (holotype). Length 3,56; carapace 1.85 long, 1.22 wide; tibiat+patella [ 1.59. Colour: Carapace chestnut brown; chelicerae medium brown; stermum medium brown with yellow posterior tip; coxae white with dark brown rim; trochanters dark with yellow ventral patch; femora white in proximal half, dark brown in distal half, patellae yellow with darker distal section; ibiae with white proximal part preceded by thin dark ring and dark distal part; metatarsi and tarsi yellowish brown. Abdomen shiny black; dorsum with two pairs of small white spots, sides with one oblique white spot; venter sepia, slightly paler in front of epigastric fold, Carapace finely granulated; sternum smooth. Eyes: a: 0,10; b: 0.103: 0.11; d: 0.12; e: 0.02; 0,02; 2: 0.04; h: 0.08; AL-AL: 0.18. MOQ: AW= REVISION OF THE ASTERON-COMPLEX 367 FIG. 4. Pentasteron spp, right male palps. A-C, P_ oscitans; A, ventral; B, retrolateral, C, embolus with embolar apohysis, prolateral. D-F, P, storasvides: D, ventral; E, retrolateral; F, embolus with embolar apohysis, prolateral. G-I, P. sardicdum; G, ventral; H, retrolateral; 1, embolus with embolar apohysis, prolateral. Scale 0.5mm, 0.84 PW; AW = 0.78 L. Clypeus 0.30. Chilum 0.03 high, 0.13 wide. Spination: F P T Mt I d2 - v2 v2 iv d2 - v2 v2 I pl3d3rl3—splidirll ~— pl2d2rl2v1-2-2 8disp dw5 IV pl3d3rl2 so plidirll = pl2d3rl2v1-2-2.—- 10disp dw5 Hinged hairs: TI and TIL: d1. Preening brush on Mt II and HI. Abdomen with large rounded lip in front of tracheal spiracle, Colulus with 3 setae. Male palp (Fig. 4A-C): tibia with large retro- lateral concavity delimited by elongate, roughly triangular, pointed dorsal apophysis and ventro- lateral lamellate apophysis. Cymbium with large proximal fold, forming concavity together with tibial concavity, dorsally delimited by large trian- gular flap; cymbial flange unmodified. Tegulum broad based, tapered toward rounded VTA; LTA short, truncated, broader at extremity than at base; DTA membranous, distally sharply curved outwards. Embolus short, thick, rigid, curved outward; with thin short EA, Female (paratype). Total length 4.31; carapace 1.98 long, 1.35 wide; tibiat+patella I: 1.58. Colour: As ¢ but sternum uniform medium brown, Palp: femora dark brown with pale ventral patch, other segments yellow. Eyes: a: 0.10; b: 0,10; c: 0.11; d: 0.12; e: 0.03; f: 0.03; g: 0.05; h: 0.08; AL-AL: 0.19. MOQ: AW= 0.86 PW; AW = 0.76 L. Clypeus 0.32. Chilum single, 0.06 high, 0.22 wide. Spination: F P T Mt I dl - v2-2-2 v2 if] dl - v2-1-2 v2 ut pl2d3ril —plidirll_ pl2d2rl2v2-2-2&disp dws IV pl2d3—pldirll_pl2d2rl2v2-2-2 1 0disp dw5 Hinged hairs: TI d1, TI d1. Preening brush on Mt II and IIT. Epigyne (Fig. 6C,D): simple, sclerotised plate with 2 depressions (entrance openings) centrally; posterior clearly indented. Copulatory ducts and spermathecae strongly sclerotised. DISTRIBUTION. Known only from type locality. Pentasteron sordidum sp. nov. (Figs 4G-I, 15) ETYMOLOGY. The name refers to the colour of the male which is rather ‘dirty’ (Latin: sordidus). MEMOIRS OF THE QUEENSLAND MUSEUM MATERIAL. HOLOTYPE: d, Lake Wytchugga, 6km w. Wilcannia,New South Wales, 21-22.xii.1998, M. Baehr (QM 846889). DIAGNOSIS. 6 ¢ have a palp with deep tibial concavity delimited by the large longitudinal swollen ventrolateral swelling and a ventrally ridged dorsolateral apophysis. DESCRIPTION. Male (holotype). Total length 4.88; carapace 2.38 long, 1.56 wide; tibiatpatella 11.74. Colour: Carapace chestnut brown; chelicerae and sternum medium brown; coxae pale; trochanters yellowish brown; femora I yellowish brown with darker patches at base and tip; femora II-[V white in proximal half, yellow overlaid with dark brown in distal half; other parts yellow. Abdomen grey mottled with white and black, yellowish in front of epigastric fold and on lip in front of tracheal spiracle. Carapace finely granulated; sternum smooth. Eyes: a: 0.15; b: 0.14; c: 0.14; d: 0.14; e: 0.03; f: 0.02; g: 0.04; h: 0.10; AL-AL: 0.26. MOQ: AW= 1.03 PW; AW = 0.87 L. Clypeus 0.42 or 3.0 times ALE. Chilum 0,08 high, 0.30 wide. Spination: F P T Mt | plid2 - y2-2-2 vl-l-1-l in d2 - v2-2-2 vl-1-1-ldw3 TH pl3d3rl2—pliidirlt— pl2d2rl2v2-2-2—8disp dws IV pl2d3rll_—pldirl1— pl3d3r13v2-2-2 lost Hinged hairs: TI and TII: d1, Preening brush on Mt I and III. Abdomen with large rounded lip in front of tracheal spiracle. Colulus a group ofc. 10 setae. Male palp (Fig. 4G-1): tibia with large retrolateral concavity delimited by thick longitudinal ventrolateral swelling and large dorsolateral, ventrally ridged apophysis; cymbium with basal fold linked up with tibial concavity. Tegulum broad at base, tapered toward strong VTA which has tip curved outward; LTA short, thorn-shaped; DTA membranous, widened toward broadly truncate extremity. Embolus fairly short, curved outward, thin but rigid; with EA longer than embolus proper, slightly widened at tip. Female unknown. DISTRIBUTION. Known only from type locality. REVISION OF THE ASTERON-COMPLEX 369 FIG. 5. Pentasteron spp. right male palps. A,B, ?. seeurifer: A, ventral; B, retrolateral. C,D, P. isabelae; C, ventral; D, retrolateral. Scale 0.5mm. DTA = dorsal legular apophysis, F = embolus, LTA = lateral tegular apophysis. VTA = ventral tegular apophysis. DIAGNOSIS. Males have a palp with a deep tibial concavity delimited by the large longitudinal! ventrolateral swelling and a dorsolateral apophysis with recurved tip, in combination with the bifid embolar apophysis which gives the impression that the embolar complex.is trifid. DESCRIPTION. Male (holotype), Total length 4.88; carapace 2.46 Jong, 1.74 wide; tibia*patella | 2.50. Colour: Carapace chestnut brown; chelicerae and sternum medium brown; coxae white with dark brown rim; trochanters dark with yellow yentral patch; femora white with dark patches at base in proxintal half, dark brown in distal half; remainder of legs yellowish brown, posterior tibiae with blackish lateral streaks. Abdomen shiny black; dorsum with two pairs of small white spots and 3 crescent-shaped spots in [ront of spinnerets; sides with one oblique white spot and pale mottling; venter sepia, with two yellow spots in Iront of epigastric fold. Carapace finely granulated; sternum smooth. Eyes: a: 0.15: b: 0.12; 0: 0.14; d: 0.145 e: 0.04; f 0.02; a: 0,07; h: 0.14; AL-AL: 0.28. MOQ: AW = 1.00 PW; AW = 0.86 L. Clypeus 0.48 or 4 times ALE. Chilum 0.10 high, 0.0.36 wide. Spination: Pentasteran storosoides sp. nov. ! h ! Mi (Figs 4D-F, 15) I pplid2 vie? v2-l-l i pltd3 - yl-2-2 y2-2dw3 ETYMOLOGY, Superticially like Starosa Jocqué in its Ht = plad3ri2pildiel? = pl2d2r12h2-2-2— 1odisp do deep tibial concavity and strong ventral tibial knob. V pi2ds —plldtil)—pl3d3pl3v2-2-2 —Lddisp dive MATERIAL. HOLOTYPE: 2, 30km SW of Wilcannia, Hinged hairs: Tl and TIL: dl, Preening brush on New South Wales, ca, 142°45°E, ca, 32°25°S, 22.Ni7.1998, black box fogging, LI. & M. Baehr (QM §46948). Mt I] and II. 370 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 6. Pentasteron spp. epigynes. A,B, P. simplex; A, ventral; B, dorsal (cleared). C,D, P. oscitans; C, ventral; D, dorsal. E,F, P. intermedium E, ventral; F, dorsal. Scale 0.5 mm. Abdomen: anterior lip of tracheal spiracle not contrasting from rest of venter; posterior lip larger, sticking out, brownish yellow. Colulus a small swelling with 8 setae. Male palp (Fig. 4D-F): tibia with large retrolateral concavity delimited by thick longitudinal ventrolateral swelling and large dorsolateral apophysis with strongly recurved tip; with short prolateral dorsal apophysis. Cymbium with shallow proximal fold and shallow basal concavity. Cymbial flange long, separated in front from cymbial rim by short bend. Tegulum broad at base, tapered toward fairly slender VTA; LTA short, thorn-shaped; DTA membranous, widened towards truncated, serrated extremity. Embolus fairly short, fairly slender; EA with bifid tip, giving the embolar complex a trifid impression. DISTRIBUTION. Known only from type locality. Pentasteron isobelae sp. nov. (Figs 5C,D, 7C,D, 15) ETYMOLOGY. In honour of Isobel Raven. MATERIAL. HOLOTYPE: ¢, Ramornie SF, Track off Mt Tindal Rd. NSW, 29°42’S 152°38’E, 4.ii-9.iv.1993, 220m, M. Gray & G Cassis (AM 039404). PARATYPES: New South Wales: | ¢ , together with holotype; 12°, NSW, 68AR, Maria River SF, NPWS survey, 1km along Northern Trail into rubbish dump, 31°08’S 152°28°E, 4.1i1-9iv.1993, 35m, M. Gray & G Cassis (AM KS 039403); REVISION OF THE ASTERON-COMPLEX 371 1d, Ramornie SF, Track off Mt Tindal Rd, 29°42’S 152°37°E, 4.i1-9.iv.1993, 220m, M. Gray & G Cassis (NPWS survey) (AM KS039402); 1d, Tindal Rd, 380m, further as previous (AM KS 039200); 1¢, 240 m east of junction of Kunderang East and Kunderang West Rds, 30°488 152°02’E, 4.ii-9.iv.1993, 900m, M. Gray & G Cassis (NPWS survey) (AM KS039120); 1d, Bundjalung National 20, near gravel quarry, 29°17°S 153°16’E, 4.ii-9.iv.1993, M. Gray & G Cassis (AM KS 039198); 1d, Chaelundi SF, 1.2km W along Stockyard Fire Trail from Chandlers Ck, 29°56S 152°31E, 450m, 4.ii-9.iv.1993, M. Gray & G. Cassis (AM KS 039199); SE Queensland: 2d, Expedition Ra NP, Amphitheatre camp, 25°12’S 148°59°E, 14-19.xii.1998, 560m, open forest, G. Montheith, G Cook & G Thompson (QM §52611). DIAGNOSIS. Male palp with a small cymbial concavity and ear-shaped embolar appendage; 2 2 with large central depressions and an almost straight posterior margin of epigyne. DESCRIPTION. Male (holotype; Pt in brackets). length 4.74 (4.46); carapace 2.30 (2.16) long, 1.62 (1.60) wide; tibia+patella ] 2.22 (2.11). Colour: Carapace chestnut brown; chelicerae medium brown; sternum medium brownish with darker lateral margins; coxae white; trochanters dark with yellow ventral spot; femora white in proximal half, dark brown in distal half; patellae yellow; tibiae yellow suffused with black on sides; metatarsi and tarsi yellow. Abdomen dark grey; dorsum with 2 pairs of small white spots followed by 2 white chevrons and spot with sinuous margins in front of spinnerets; sides with 1 white spot in front and 3 large oblique stripes; venter sepia with 2 small yellow spots in front of epigastric fold; lung covers yellow. Carapace finely granulated; sternum smooth. Eyes: a: 0.16; b: 0.12; c: 0.11; d: 0.12; e: 0.03; f 0.03; g: 0.08; h: 0.10; AL-AL: 0.32. MOQ: AW= 1.06 PW; AW= 1.00 L. Clypeus 0.38 or 3.2 times ALE. Chilum single 0.24 wide 0.08 high. Spination: F P T Mt I plid2 - v2-2-2 2-2-2 tl d2 v2-2-2 v2-2 dw3 HI pl2d3rll = plidirll = pl2d2rl3v2-2-2 10disp dw5 IV plid4 plidirll = pl3d3rl3v2-2-2.—-10disp dw5 Hinged hairs: TH and TIL: dl. Preening brush on Mt IT and III. Male palp (Fig. 5C,D): femur and patella pale, contrasting with remainder of palp; tibia with large retrolateral concavity delimited along post- erior end by solid, tapered and twisted, sharp- tipped apophysis pointing forward and thin, truncated and slightly indented ventrolateral apophysis. Cymbium with shallow basal con- cavity according with tibial concavity; flange unmodified. Tegulum broad, with large, tongue- shaped terminal VTA. Embolus short, flat, rigid, slightly twisted, almost straight, accompanied by large, flat transparent ear-shaped apophysis, provided with 2 semicircular ridges; DTA membranous, narrow, straight. Female (paratype). Total length 5.60; carapace 2.56 long, 1.70 wide; tibia+patella I: 2.53. Colour: almost as in male but generally paler. Eyes: a: 0.16; b: 0.15; c: 0.14; d: 0.15; e: 0.05; f: 0.03; g: 0.08; h: 0.15; AL-AL: 0.32. MOQ: AW = 1.00 PW; AW = 0.85 L. Clypeus: 0.44 or 3.0 times ALE. Chilum single 0.23 wide, 0.11 high. Legs: Spination: F P T Mt I plid2 - v2-2-2 v2-]-1-2 I d2 pl2y2-2-2 v2-1-2 dw3 in pl2d3rll ss plidirll = pl2d2rl2v2-2-2 1 Ndisp dw5 IV plid3rll— plldirll = pl3d3rl3v2-2-2. 1 0disp dwS Hinged hairs: TI dl, TIL dl. Preening brush on Mt I] and II. Epigyne (Fig. 7C,D): simple: suboval scler- otised with almost straight posterior rim and 2 large central depressions. Copulatory ducts semi- circular; spermathecae small, caudal, adjacent. Variation: colour pattern and size very stable: d carapace length 1.82-1.87, width 1.20-1.22; carapace 1.92-1.98 long, 1.33-1.40 wide. DISTRIBUTION. SE Queensland and NSW. Phenasteron gen. nov. TYPE SPECIES. Phenasteron longiconductor sp. nov. ETYMOLOGY. Greek phenomenon with Asteron; refers to ‘phenomenal’ male palps. Gender is neuter. DIAGNOSIS. 6 ¢ have an domed cephalothorax with highest point just in front of fovea (Fig. 9A), enormous T-shaped distal tegular apophysis (DTA) with refolded margin (Fig. 9B,D), course of the sperm-duct in the tegulum not transverse but oblique and the posterior sclerotised swelling of the subtegulum. 2 2 unknown. DESCRIPTION. Small spiders (2.90-3.5) with smooth or slightly granulate tegument. Carapace widest at level of coxae II (Fig. 8), narrowed to MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 7. Pentasteron spp. epigynes. A,B, P. securifer; A, ventral; B, dorsal (cleared). C,D, P. isobelae; C, ventral; D, dorsal. Scale 0.5mm. 0.60 maximum width in males. Profile domed with highest point just in front of fovea (Fig. 9A). Colour: carapace orange to medium brown, chelicerae and sternum yellowish brown. Eyes (Figs 8, 9A) in 3 rows (2-4-2). Only ALE in first row, second AME (in middle) and PLE, third only of PME. Eyes subequal or AME larger than remainder. MOQ slightly longer than wide. Clypeus slightly concave retreating, c. 5 times diameter of ALE; with few hairs. Chilum single, short and wide, without setae. Chelicerae as usual in the family with few hairs in front and dense row on distal promargin; no teeth. Labium narrowed at base; sparsely haired. Gnathocoxae rather elongate; sparsely haired; with anteromesal scopula. Sternum flat; triangular with slightly procurved anterior margin and slight triangular extensions between coxae. No inter- nor precoxal sclerites. Abdomen dark sepia with five pale spots. Legs: formula 4123. Spination: few spines on pairs I, II, more numerous on III, IV. Paired tarsal claws with numerous (12-14) teeth. Unpaired claw on small onychium. Trichobothria in 2 rows on T, single row on Mt and t. Hinged hairs few, restricted to dorsal side of TI and I. Metatarsal preening brush on Mt II and III poorly developed. Abdomen oval; with poorly developed trans- lucent anterior scutum; with faint dorsal sigilla and small lateral frontal sigillum on sides. Spin- nerets: AS, conical, with short distal segment; MS, PS small, in a row. Colulus represented only by some hairs. Tracheal spiracle ordinary, small. Male palp (Fig. 9B-E): tibia with large retro- lateral concavity delimited by solid dorsolateral apophysis and ventrolateral apophysis, with swollen lateral margin provided with macrosetae or row of hairs. Cymbium unmodified, flange simple, area above it sclerotised and slightly concave. Subtegulum with backward extended swelling. Base of tegulum narrowed toward origin of embolus; course of seminal duct oblique, not transverse; VTA large; DTA very large, T-shaped, distal margin of transverse bar refolded. Embolus emerging on lateral part of tegulum, long and slender. LTA a small, short thorn, or reduced. Females unknown. REVISION OF THE ASTERON-COMPLEX 373 FIG. 8. Phenasteron longiconductor body dorsal. Scale 1mm. KEY TO THE SPECIES OF PHENASTERON |. DTA and embolus almost as long as bulbus; VTA with long slender distal prong, at right angle with body of this apophysis; course of sperm-duct longitudinal (FIP-OBIG 35 3 babe RES P. longiconductor DTA and embolus much shorter than bulbus; VTA without slender distal part; course of sperm-duct oblique (Ri SDI BY patie he ae Plates ootes P. machinosum Phenasteron longiconductor sp. nov. (Figs 8, 9A-C, 16) ETYMOLOGY. Noun in apposition, refers to the very large DTA which appears to be the functional conductor. MATERIAL. Holotype, ¢, 12.3km SSW of Murrayville P.O. Victoria35°22’S 141°09°E; site 62, xi.1985, drift fence pitfall trap, A.L. Yen (VM). PARATYPES: Victoria: 1d, 6.5 km SW of junction of MV highway and Annuello Rd., 34°50’S 142°34’E, site 11, x.1985, drift fence pitfall trap, A.L. Yen (VM); 1 d, 16 km SE of Murrayville, 35°22’S 141°19’E, site 71, xi.1985, drift fence pitfall trap, A.L, Yen (VM); 1d, 15.5 km WSW of Hattah, 34°47’S 142°07’E, site 40, x.1985, drift fence pitfall trap, A.L. Yen (VM); WA: 1d, Nanga station, 26°35°31"S 113°53°22”E 16.x.1994-19.i.1995 (WAM 99/2379), N. McKenzie & J. Rolfe, wet pits WAM/CALM Carnarvon survey (NA5). DIAGNOSIS. Males have an enormous pick- shaped tegular apophysis at embolus base. DESCRIPTION. Male (holotype). Length 3.51; carapace 1.87 long, 1.31 wide; tibiatpatella I 1.33 long. Colour: Carapace medium brown with darker radiating striae and u-shaped darker pattern delimiting cephalic area; chelicerae and sternum medium reddish brown; coxae pale; trochanters pale with dark pro- and retrolateral spots; prox- imal half of femora white with dark proximal ring, distal half of femora I-IV medium brown suffused with black; femur I pale brown suffused with black; patellae uniform pale yellow; tibiae brownish yellow, darkened on ventral side. Abdomen dark sepia; dorsum with faint, narrow dark brown scutum in front and 5 white spots: 2 pairs in anterior half, and 1 spot in posterior half; 2 oblique white stripes on each side. Carapace and sternum finely granulated. Highest point of profile halfway between fovea and PME. Eyes: a: 0.12; b: 0.08; c: 0.08; d: 0.09; e: 0.03; f: 0.02; g: 0.10 h: 0.12; AL-AL: 0.28. MOQ: AW = 1.00 PW; AW = 0.87 L. Clypeus retreating, 0.34 or 4.2 times the diameter of an ALE. Chilum single, 0.08 high, 0.18 wide. Legs: Spination: F P T Mt [ dl II dl - dw2 ul d2 plidiril pl2d2rl2v2-2 4disp dw6 One hinged hair on tibiae | and I]. Male palp (Fig. 9B,C): tibia with large retro- lateral concavity delimited by 2 apophyses: ventrolateral one, lamellate with sharply bent distal tip and ventrolateral haired ridge, dorso- lateral one long, gradually tapered, sharp. Cym- bium dorsoventrally flattened, ventrally glabrous; basally shallowly concave; with retrolateral haired ridge and long flange distally curved and swollen. Subtegulum extended backwards, swelling strongly sclerotised, reaching special 374 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 9, Phenasteron spp, A-C, P. longiconductor; A, body lateral; B,C, right ¢ palp; B, ventral; C, retrolateral. D,E, P. machinosum, right 3 palp; D, ventral; E, retrolateral. Scales, B, 1mm; D, 0.5mm. REVISION OF THE 4STERON-COMPLEX concavity of tibia; tegulum broad at base, sperm- duet slightly bent backward. Embolus about as long as cymbium, fairly slender, curved outward. VTA. very large, pick-shaped; base long, broad distal part turned outwards over more than 90°, short, sturdy, DTA yery large: retrolateral margin rebordered, anterior prong broad and rounded. posterior prong long, tapered toward blunt apex. Female unknown, Size. colour pattern and spination very stable. DISTRIBUTION. Victoria and WA, Phenasteron machinosum sp. nov. (Figs 9D,E, 16) ETYMOLOGY, Latin: machinasus, provided with tools: reters to the toal-shaped palpal sclerites. MATERIAL, HOLOTYPE: 2, South Gap Station, Beda Hill, Sth Aust, 31°51°S )37°3S7"E: 4-6,xii, 1989, pitfall trap, D. Hirst (SAMA N199296), DIAGNOSIS. Males with large shovel-shaped tegular apophysis (VTA) al embolus base; sub- legulum with backwardly extended posterior swelling. DESCRIPTION. Mele (holotype). Total length 2.90; carapace 1.42 long, 1.12 wide; tibia+patella 11,06 long. Colour: Carapace orange brown with faint darker radiating striae and u-shaped darker pattern delimitmg slightly paler cephalic area; chelicerae and sternum pale yellowish brown, sternum darkened along margin: coxae pale: trochanters pale with dark pro- and reirolateral spots; proximal half of femora white with dark proximal ring, distal half of femora pale yellow suflused with black on sides; tibiae [and II pale yellow and suffused with black on venter and sides in proximal half, pale in distal half tibiae [1] and IV pale yellow, suffused with black on yenter and sides; metatarsi yellow: tarsi orange yellow, darkened towards tip. Abdomen dark sepia; dorsum with faint, narrow dark brown scutum m front and’3 white spots: 2 pairs in anterior half, and | spot in posterior half; 2 oblique white stripes on each side. Carapace and sternum finely granulated. Carapace lairly high, highest point of profile just in front of fovea, Eyes: a: 0.09; b: 0.08; c: 0.07; d: 0.06; &: 0.02; F 0.01; 2: 0.08 h: 0.10; AL-AL: 0,18, MOQ: AW = 0.18 PW; AW = 0,75 L. Clypeus slightly retreating, 0.28 or 3.5 limes ALE, Chilum single: 0.10 high, 028 wide. 375 Spination: ! r | felt | al | al iT dl vi ¥2 1! d2 plied plad2il20 1-2 Slings dye W I? mitrtivtt pl2d3r}2v 1-122 Tdisp atws One dorsal hinged hair on tibiae 1, Il. Male palp (Fig. 9D,E.). femur pale with dark retrolateral patch, patella pale, contrasting with rest of palp: tibia with large retrolateral concayity delimited by two apophyses; ventrolateral one fairly fat with group of strong setae; dorsolateral one ridge-shaped with sharp, proximal prong, slightly curved forward. Cymbium with thin bul well developed retrolateral Hange in proximal halt. Embolus about half as long as cymbium, fairly slender, curved outward. VTA large, in shape of curved shovel, base fairly broad but narrower than distal part ending im shghtly curved, distally rebordered Hat part. DTA large; retrolateral margin strongly rebordered, amterjor prong broad, rounded, posterior prong fairly short, sharp, curved imward. Femule unknown. Colour pattern and spination very stable, DISTRIBUTION, Known on!y from type lovality. Leptasteron gen. nov TYPE SPECIES. Leprisreron platcondyeter sp, nov, ETYMOLOGY. Greek leptos. hidden with dsveron refers to this genus remaining hidden in ihe Asteran-comples. Gender ts neuter, Bachr & Jocque (1996) mentioned these taxa as the brachycunductor-group. DIAGNOSIS. Males have flat cephalothorax, elongate flat cymbium, large distal tegular apophysis (DTA) with retolded margin, sperm-duct in the tegulum not transverse but longitudinal and posterior sclerotised swelling of the subtegulum, Females unknown. DESCRIPTION. Small to medium-sized (4.50) -6.60) with smooth or slightly granulate teg- ument. Carapace widest at level of coxae Tl. narrowed to 0.65 max. width in 3 2. Profile Nat with highest point just behind PME (Fig. 1 1A). Colour: variable; carapace colour varies. from dark brown (L. platveonductor) to bright vellow (L. verillyni), chelicerae and sternum and Jegs uniform legs (L, vexi//im) or with contrasting leg segments (/. plaiveonductar), Abdomen dark sepia with five pale spots. Sclerotised in front of 376 epigastric fold (LZ. platyconductor) or with pale booklung opercula (L, vexi/lum). Eyes (Figs 10A,B,11A) in 3 rows (2-4-2). ALE only in first row, second with AME (in middle) and PLE, the third of PME. Eyes subequal or AME larger than remainder. MOQ slightly longer than wide. Clypeus straight or slightly retreating, 2.5 to 4 times diameter of ALE; with few hairs. Chilum single (L, vexi//um) or double (L. platyconductor). Chelicerae as for family with a few hairs in front and a dense row on distal promargin; no teeth. Labium narrowed at base; sparsely haired. Gnathocoxae rather elongate; sparsely haired; with anteromesal scopula. Sternum flat; triangular with straight anterior margin and slight triangular extensions between coxae. No inter- nor precoxal sclerites. Legs: formula 4123. Spination: few spines on pairs I, 1, more numerous on III, IV. Paired tarsal claws with numerous (12-14) teeth. Unpaired claw on small onychium. Trichobothria in two rows on T, single row on Mt and T. Hinged hairs present but few, restricted to dorsal side of TI and II. Metatarsal preening brush on Mt II and III poorly developed. Abdomen oval; with poorly developed trans- lucent anterior scutum; with (L. platyconductor) or without (P. vexil/um) dorsal and lateral sigilla. Spinnerets: AS, conical, with very short distal segment; MS and PS small, in a row. Colulus represented only by some hairs. Tracheal spiracle hidden by well developed anterior lip. Male palp (Fig. 11 B-E): tibia with a large retro- lateral concavity delimited by a solid dorsolateral apophysis and ventrolateral apophysis, with swollen lateral margin provided with macrosetae or row of hairs. Cymbium elongate, flat , flange simple, area above it sclerotised and slightly concave. Subtegulum with backward-extended swelling. Base of narrowed tegulum toward origin of embolus; course of seminal duct longitudinal, not transverse; VTA large, either wide and with large recurved extremity or long and slender. Embolus emerging on posterior part of tegulum, very long and slender. LTA small: knob-shaped; DTA very large, membranous or sclerotised; distal margin refolded. Females unknown. KEY TO THE SPECIES OF LEPTASTERON |. Carapace uniform yellow; DTA membranous, T-shaped; VTA long and slender (Fig. 11D-E).. 2. . Lvexillum Carapace uniform dark brown; DTA sclerotised, sickle-shaped, VTA not long and slender but sharply bent (Fig VIB Cy. fies. s ¢-ne ¥ 3G + L. platyeonductor MEMOIRS OF THE QUEENSLAND MUSEUM Leptasteron platyconductor sp. nov. (Figs 10A,11A-C,16) ETYMOLOGY. Noun in apposition; refers to wide flat DTA which appears to be the functional conductor. MATERIAL. HOLOTYPE: ¢, Cape Range, WA, 22°05’S 114°00’E; 14. iii-6.v.1992, pitfall trap outside cave C56, R.D. Brooks (WAM BES:1103). PARATYPE: 1d, Station Creek, 127 km SSE Leinster, Western Australia, 28°45’S, 121°00’E, 8-9.xi1987, M. Baehr (QM 845244). DIAGNOSIS. Males with elongate palpal cymbium and very broad, large DTA (Fig. 11B); uniform dark colour. DESCRIPTION. Male (holotype, paratype in brackets). Total length: abdomen missing in holotype (6.60); carapace 3.11 (3.10) long, 2.04 (2.02) wide; tibiat+patella I 2.85 (2.80) long. Colour: Carapace dark brown with darker radiating striae and v-shaped darker pattern in front of fovea; chelicerae and sternum dark brown; coxae, trochanters and femora dark brown with some darker stripes; tibiae medium brown with darker ventral side; metatarsi and tarsi brownish yellow. Abdomen dark sepia with five small white spots two in front, two in middle, one in front of spinnerets. Sides sepia, mottled with pale; venter pale sepia; two pale spots in front of epigastric fold; lung covers yellow. Carapace and sternum smooth. Carapace fairly flat, highest point of profile just behind PME, provided with sparse but evenly dispersed cover of tiny setae. Eyes: a: 0.16; b: 0.14; c: 0.17; d: 0.14; e: 0.04: f: 0.02; g: 0.08 h: 0.12; AL-AL: 0.30. MOQ: AW = 0.86 PW; AW = 0.82 L. Clypeus slightly retreating, 0.70 or 5.0 times diameter of ALE. Chilum double: each part 0.12 high and 0.28 wide. Legs: Spination: F P T Mt I plid2 y2-1-2 v2-1-1-2 I d2 plivl-1-2 v2-1-ldw2 il pl3d2rl2 ss pildirll == pl2d2rl2v2-2-2—- &disp dw6 IV pl2d3rl2—piidtrll =~ pl3d2rl3v2-2-2— 8disp dw6 One hinged hair on tibiae [ and II. Preening brush on Mt I and III. Abdomen with 2 round dorsal sigilla and an elongate lateral one in front on either side. Tracheal spiracle with swollen anterior lip and sclerotised protruding posterior lip. Colulus a group of about 8 short setae. REVISION OF THE ASTERON-COMPLEX \\ ee) | | FIG. 10, Leptasteron. body dorsal. A, L. platvconductor. B, L. vexillum. Scale |mm Male palp (Fig. 11B,C): tibia with large retrolateral concavity delimited by two apophyses: ventrolateral one, Jamellate with blunt, rebordered, frontal extension provided with some large setae; dorsolateral one with proximal, tapered, sharply pointed prong, with triangular tooth at frontal base. Cymbium elongate, dorsoventrally flattened, long, flange fairly long, slightly concave at base; retrolateral ridge provided with hairs standing out. Embolus very long, whip-like, originating on posterior part of tegulum with base pointing backward; tegulum with long retrolateral ridge, ending in slender proximal knob; VTA large, basal part broad and concave, sharply curved outward at about half its length, ending in long, tapered prong; DTA large, very wide, broadly curved, rebordered along retrolateral edge. Female unknown. Variation: the two known males are very similar. DISTRIBUTION. Known only from type locality. Leptasteron vexillum sp. nov. (Figs 10B, 11D,E, 16) ETYMOLOGY. Noun in apposition; Latin; vexil/ium, flag; referring to the large DTA. MATERIAL. HOLOTYPE: d, Tindery Nature Res., southern entrance, NSW, 35°39°39"S 149°127°43"E, 14.ii1.1999, J. Tarnawski & S. Lassau, CBCRO003-032 (AM KS 55882). DIAGNOSIS. Males are unique in palp with enormous, terminal folded DTA, very long VTA and long whip-like embolus. DESCRIPTION. Male (holotype). Total length 4.86; carapace 2.24 long, |.64 wide, tibiatpatella 12.32. Colour: Carapace uniform yellow with small dark area on either side above condyle of yellow chelicerae; sternum pale yellow; femora yellow turning to orange distad; rest of legs orange. Abdomen dark grey, with orange tinge above pedicel; with two pairs of white spots and smaller triangular spot in front of spinnerets; sides and venter pale; area in front of epigastric area yellow. Teguments smooth. Highest point of carapace just behind PME. Carapace and legs provided with sparse but evenly dispersed cover of tiny setae. Eyes: a: 0.16; b: 0.14; c: 0.14; d: 0.14; e: 0.04; f: 0.04; g: 0.08; h: 0.08; AL-AL: 0.20 MOQ: AW = 1.00 PW; AW = 0.90 L. Clypeus slightly retreating, 0.34 high or 2.5 times diameter ALE. Chilum single: 0.14 high, 0.18 wide. Legs: Spination: F P pl Mt 1 dl y2-2-2 v2-2 dw3 i d - v 1-2-2 v2-2 dw3 ut pl2d3rl2_—plldirll pl2d2rl1wv2-2-2 8disp dw5 IV plild3rll — plldirll pl2d2rl2v2-2-2 Sdisp dw5 Hinged hairs: one dorsal on TI and Il. Epigastric area with triangular indentation. Large sclerotised area in front of tracheal spiracle with pronounced frontal lip. Colulus a row of setae. Male palp (Fig. 11D,E): tibia with large retrolateral concavity delimited dorsally by long, forward -directed slightly downcurved pointed apophysis, ventrally by slightly shorter, straight, pointed apophysis; prolaterally swollen with 2 macrosetae. Cymbium crescent-shaped, strongly tapered; tegulum with caudal, flattened extension bearing long, whip-shaped embolus which MEMOIRS OF THE QUEENSLAND MUSEUM originates on posterior part of tegulum. Long, slender, outward curved VTA originates on prolateral tegular ridge. DTA large, membranous, broad extremity with large fold accommodating extremities of both VTA and embolus. Female unknown. DISTRIBUTION. Known only trom type locality. Subasteron gen. nov. TYPE SPECIES. Subasteron daviesae sp. nov. (Fig. 14) Image from D. Knowles, mentioned in Lindsey (1998) as knobble spider. ETYMOLOGY. Subasteron, is referring to the slightly aberrant somatic morphology of the single species in this genus as compared to other members of the Asteron-complex. DIAGNOSIS. Recognised by the peculiar shape of the cephalothorax which reaches its highest point at the level of the PME and the accordingly high clypeus, up to ten times the diameter of the ALE. Further diagnostic characters are from male palp, first the presence of a prolateral tegular apophysis (PTA) which is unique in the Asteron-complex: tibia has a deep retrolateral concavity combined with more or less pronounced concavity on base of cymbium; cymbium has a prolateral basal extension fitting in aconcavity with membranous bottom of tibia. DESCRIPTION. Medium-sized spiders (7.00-9.00) with very finely granulate tegument. Carapace widest at coxae II (Fig. 12A), slightly narrowed to 0.8 maximum width in females, to ca. 0.68 maximum width in males. Profile raised toward front with highest point near PME (Fig. 12C); fovea deeper in males than females. Colour: carapace and sternum dark brown, chelicerae medium brown; legs with strongly contrasting, white to dark brown segments: coxae pale, trochanters dark, femora dark brown and white, tibiae brown with darker stripes; metatarsi pale, medium brown in distal part in females, uniform dark brown in males; tarsi brownish orange. Abdomen dark with contrasting pattern of white spots and patches. Males darker and with more contrasting pattern. Eyes (Fig. 12C,D) in 3 rows (2-4-2). ALE only in first row, second with AME (in the middle) and PLE, third with PME. Eyes subequal but ALE smaller than others. MOQ longer than wide. Clypeus slightly concave, high, 6 times ALE in females, 10 times ALE in males; with some setae. Chilum double; separation not complete in REVISION OF THE ASTERON-COMPLEX 379 I : <| \ FIG. 11. Leptasteron spp. A-C, L. platvconductor,; A, body lateral; B,C, right male palp; B, ventral; C, retrolateral, L. vexi/lum; D,E, right male palp; D, ventral; E, retrolateral. Scales, C, 1mm; E, 0.5mm. DTA = dorsal tegular apophysis, LTA = lateral tegular apophysis, VTA = ventral tegular apophysis. 380 MEMOIRS OF superior half; without setae. Chelicerae as usual in the family with few hairs in front and dense row on distal promargin; no teeth. Labium narrowed at base; sparsely haired. Gnathocoxae rather elongate; sparsely haired; with anteromesal scopula. Sternum flat; triangular with anterior margin slightly concave; with very small triangular extensions between coxae. No inter- nor precoxal sclerites, Legs: formula 4123. Spination: nine fartly long: up to 4 times of diameter Mt IV in d ¢, times diameter of Mt IV in females; few spines on pairs | and II, more numerous on III and IV. Paired tarsal claws with numerous (12-14) teeth on anterior leg pairs, with slightly fewer on those THE QUEENSLAND MUSEUM FIG. 12. Subasteron daviesae; A, body dorsal; B, abdomen dorsal variation; C-E, cephalothorax: C, lateral; D frontal; E, ventral. Scale Imm, of legs II] and [V. Unpaired claw toothless, on small onychium. Trichobothria in 2 rows on T, single row on Mt and t. Hinged hairs few, restricted to dorsal side of TI and II. Metatarsal preening brush on Mt II, IIL and IV, poorly developed. Abdomen oval, fairly elongate; in males with marked central dip; with 2 dorsal sigilla, poorly developed elongate frontal sigillum on either side and pair just behind epigastric gold, more strongly developed in d d. Spinnerets: AS, fairly long, slightly conical, with very short distal segment; MS and PS very short, absent in ¢ dé. Colulus represented by group of setae. Tracheal spiracle hidden by well developed anterior lip. REVISION OF THE ASTERON-COMPLEX Male palp (Fig, 13A-C); tibia with large retrolateral cancavity delimited by solid dorsolateral apophysis and ventrolateral apophysis, with swollen lateral margin and frontal tooth. Cymbium with well developed Nange and shallow concavity, several spines near distal tip. On prolateral side with basal extension fitting in concavity with membranous bottom of the tibia. Subtegulum stronyly developed; partly membranous. Tegulum with broad base carrying transverse section of seminal duct; behind 1 partly membranous, partly strongly sclerotised; VTA small but strongly selerotised: with strong prolateral apophysis more or less parallel with embolus, Embolus emerging on prolateral part of legulum, short, rigid, curved outward. DTA strongly developed, sclerotised. LTA, flat, thorm-shaped. Epigyne (Fig. 13D,E): strongly scleroused plate with central depression and roughly rectangular plate with rounded anterior margin. Internal structure obscure due to strong sclerotisation: entrance ducts starting near centre running toward the [ront then along sides backward enter simple. thick-walled spermathecae near centre. Female palp with fincly toothed claw. Subasteron daviesae sp. nov. (Figs 12A-b, ISA-E, 14, 16) ETYMOLOGY. th honour of Val Davies, one of the collectors and in recognition ol’ her impodant wark ot Australian spiders. MATERIAL. HOLOLYPE, ¢, SEO, 3669, Kroormbit Tops, Lower Dry Creek, 45kin SSW Calliope. Queensland, 9 19.xi1.1983 000m, open forest, V. Davies & J. Gallon (QM $3669). PARATY PES: Queensland: 42 12, together with holotype. 1d, SEQ, Braemar ‘SF, I7°13°S (50°S07E, 48.i.1980 R. Raven & OM (QM 83668); 1d 12, SEQ. Kroombii Tops, northern escarpment, 45 km SSW Calliope. 9-19.xi),1953, open forest. v. Davies. J. Ciallon (QM 84429), | a xi. 1983, B. Jahnke, further as previous (OM $4415): 14. SEQ. Clear Mt, Sumsonvule Lake, 27.xii.1984, Gi Anderson (OM $4275); 22, NO, Cairns, 1968, C. Coleman (AM KS15719}; 12, SEQ, Numinbah SF, 28°12S8 133°13E,xi,1979, under bark, T. Robinson (QM $3822); 14, SEQ, Fraser Island National Park HQ, }4.x,1978(QM 83767); 1d, Stony Ck. via Sanford, 27-2078 152°48"E. 2.11-6.1v. 1996, H. Janetzki & G Monteith (OM 837773); }d. SEQ. Clear ML, Samsonvale, 27.01.1984. G, Anderson (QMS 4275); (2 | 2, SEQ, Gurgeena Plateau, open forest, 25°27'S 151°22°E, 10.x.-19.5i1.1998, intereept trap 360m, 7511. G Monteith & G Cough (QM 847507); 14, SEQ, Gurgeena Plateau, evergreen forest, 25°27'S 151°23E. 10.x 19,11, 1998, intercept trap, 360m 7513, G Monteith & Ci Gough (QM 847508). aa DIAGNOSIS, Males have a unique combination of sclerites in palp: particularities of cymbium with a prolateral basal extension and of bulbus with poorly developed VTA, large DTA and mostly very well developed prolateral tegular apophysis (PTA). DESCRIPTION, Mule (holotype), Length 7.96; catapace 3,82 long, 2.60 wide: tibiatpatella | 4.38. Colour: carapace dark brown with very taint dark radiating striae and V-shaped dark mark in front of fovea, Chelicerae and stemum medium brown, slightly suffused with black. Coxae pule with dark, distal, prolateral triangles; trochanters medium brown with darker lateral spots: femora cach with different contrasting black and white pattern, obliquely divided between upper and lower parts; patellac medium brown, anterior une dorsally pale, second one with pale dorsal spot; tibiae medium brown, first one with pale dorsal side, second and fourth with pale proxipal, dorsal spot: metatarsi medium brown, paler towards proximal end: tarsi yellowish orange. Abdomen dark sepia to black; dorsum with 10 pale spots, 8 in 4 pairs, 2 in front of spinnerets; trontal peur reniform, second and third pair small and oval, tourth pair large, rounded; central spots in front o! spinnerets clongate; sides with large, oblique, drop-shaped white patch; venter with a pain of rounded white spots on pale sepia ackground, Carapace and sternum smooth, Byes: a: 0.10; b: 0.14; 0: 0.18; de 0.18:¢: 0.04; f 0.162: 0.20; b: 0.24; AL-AL: 0.26. MOQ: AW = 0.71 PW; AW © 0.63L Clypeus: 1,0 or 7.1 times diameter of ALL. Chilunr double each part 0.32 large, 0.14 high. Legs: Spination: F p { MI | pllcirll pil pla? dy F 1 pladarl9 pil pllw2-|-2 dud ( pldcddrl plarl plld2n2v2-2 Aitisp dad WwW pra pl2etl plddwrlsy2-2-2 Vdisp te 9 One dorsal hinged hair on tibiae [T and OL. several macroselae on ventral femora, Male palp (Pig, I3A-C): tibia with large retrolateral concavity delimited by swollen ventrolateral apophysis provided with pointed anterior part; dorsolateral apophysis with slightly ridged prong directed forward. Cymhium with well developed rebordered flange, Embolus fairly short and rigid, well delimited [rom 382 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 13, Subasteron daviesue. A-C, right male palp: A, ventral; B, retrolateral: C, prolateral; DE, epigyne; D, ventral; E, dorsal. Scales 0.5mm, PTA = prolateral tegular apophysis. tegulum; VTA hardly developed, nothing more than a shallow prominence near base of embolus; LTA a membranous thorn-like appendage, DTA broad, curyed, broadened towards extremity, concave in ventral view; PTA well developed, originating on dorsal part of tegulum separated from ventral part by shallow clefi; shape similar to that ofembolus, but no embolarapophysis, like in Penreisteron ssp. Female (Fig. 13D,E). Total length 9.00; carapace 3.60 long, 2.40 wide; tibia+patella I 3.30, Colour: very much as in male but less dark and vontrasl less strong. Palp pale yellow with distal part of femur and tarsus medium brown. Carapace and sternum smooth. Eyes: a: 0.14: b: 0.12; ¢: 0.14; d: 0.16;e: 0.062 f 0.123 g; 0.22; ht 0.22; AL-AL: 0.30. MOQ: AW = 0.64 PW; AW = 0.04. Clypeus: 1.11 or 9.2 times diameter of ALR. Chilum double each part 0.42 large, 0.16 high. Legs: Spination: F P T Me \ plbelorit plivi-2-2 vi-2-2dw3 u plad3rl2 pul pllyt v2-1-1-1dw it plddayl3 plarli pl2d2r2\y2-2-2 Bdisp dwé IV pl2d4rl | pla) pl2d2r13y2-2-2 disp dws One dorsal hinged hair on tibiae [ and II, several macrosetae on ventral side of femora, Epigyne (Fig. 13D,E): central part strongly sclerotised and almost black, provided with two narrow copulatory openings; posteriorly with paler part. Copulatory ducts run around epigyne margin, ending in small touching spermathecae. Variation: male size stable TL: 6,9-8.0; carapace length: 3.2-3.9, carapace width: 2.4-2.7. Colour pattern with slight variations: dorsal spots in front of spinnerets sometimes with transverse or longitudinal divisions or sometimes completely fused and forming one large white patch. Clypeus in male up to 10 times diameter of REVISION OF THE ASTERON-COMPLEX 383 FIG. 14. Subasteron daviesae, body dorsal, slide from Knowles. ALE; discrepancy is mainly due to delimitation of ALE since height of clypeus appears stable. DISTRIBUTION. Queensland. DISCUSSION Including species described herein, the Asteron-complex now contains 37 species in 7 genera. Asteron mas Jocqué, 1991 was not included in the revision of Asferon ss. as it belongs in another genus of the complex but keeps its binomen until the new speciose genus where it belongs is described. As stated in the introduction separation of the genera erected herein was problematic. Phylo- genetically basal taxa are often difficult to define due to the lack of synapomorphies. Jocqué (1991) described a number of Australian zodariid genera almost exclusively based on palpal morphology. Jocqué (1995a,b) erected a few more genera and foreshadowed more supraspecific taxa for the wealth of Australian zodariids. Definition of the genera will continue to be almost exclusively based on genitalia. Somatic characters are often stable within these taxa but they do not offer a reliable base, as they appear to be plesiomorphic or extremely homoplastic characters. Examples in the present paper are the shape of the carapace (clypeus height), and of the chilum (single, double), eye arrangement (proportions of MOQ), colour pattern, shape of tracheal spiracle. Definition of genera in the Australian zodariids and in that family in general, has therefore mainly been based on genitalia. Yet, the epigyne of these species is especially hard to study, mainly because of the thickness and strong sclerotisation of its internal structure. Also the epigyne is usually structurally simple offering few characters. In this group, ¢ palps therefore 384 0°s- Pentasteron ra @ intermedium @ isobelae @ oscitans A parasimplex | ¥ securiter ; * simplex | ye sordidum i$} storosoides au"s- | | | 120° 130°E 140°5 1S0°E MEMOIRS OF THE QUEENSLAND MUSEUM Ww Leptasteron vexillum T 7 10°S- Leptasteron platyconductor A l A tee 22 4d L @) Phenasteron machinosum ¢ 4 @ Phenasteron longiconductor ¢ a { A 1 Subasteron daviesae Ma i ~/ | ‘ | sy, 20°S- | a \y | | \ | , % L a Eee an , Les! \ A | ee fl | 30°S. = | a — e f | , - a4 i | f ar liad WS go. f A e v / H Sy aie ray 0 ENG 40°S- } aes | 120°E BOE 140°E 150°E FIG. 15. Records of Pentasteron species in Australia. remain the main characters to define species and genera. Great care has to be taken to use appropriate characters as it now becomes clear that increase in complexity (defined as addition of apophyses and modifications) is a general phenomenon and procedes in parallel in many, if not all spider taxa (Jocqué 1998). General characters such as ‘long and flexible embolus’, ‘bifid embolus’ should therefore be avoided. They tend to appear over and again in the course of the evolution of the palp. Appearances of new sclerites (e.g. the prolateral tegular apophysis in Subasteron, the basal cymbial concavity combined with the tibial concavity in Pent- asteron) are more likely to be reliable generic discriminators. The character on which Phen- asteron is based, the size and shape of the DTA, is less reliable because amplification of an apophysis is evidently less drastic than the addition of a new structure. ACKNOWLEDGEMENTS We thank M. Gray, M. Harvey, R. Raven and J. Waldock for the loan of material. We are indebted to M. Baehr, K. Brennan and J. Shield for giving additional material, A. Reygel for his skillful habitus drawings and D, Knowles for image of Subasteron daviesae. We are specially grateful to R. Raven for the preparation of the distribution maps. This work was supported by the Australian FIG. 16. Records for species of Leptasteron, Phenasteron and Subasteron in Australia. Biological Resources Study Participatory Program. LITERATURE CITED BAEHR, B. & JOCQUE R. 1996. A revision of Asteron starring male palpal morphology. Revue Suisse Zoologie. hors série: 15-28. 2000. Revisions of genera in the Asteron-complex (Araneae, Zodariidae).The new genera Cavasteron and Minasteron. Records of the Western Australian Museum 20: 1-30. JOCQUE, R. 1991. A generic revision of the spider family Zodariidae (Araneae). Bulletin of the American Museum of Natural History 201: 1-160. 1995a. Notes on Australian Zodariidae (Araneae), I. New Taxa and Key to the Genera. Records of the Australian Museum 47; 117-140. 1995b. Notes on Australian Zodariidae (Araneae), II. Redescriptions and new records. Records of the Australian Museum 47: 141-160. 1998. Female choice, secondary effect of ‘mate check’? A hypothesis. Belgian Journal of Zoology 128: 99-117. JOCQUE, R. & BAEHR, B. 2001. Revisions of genera in the Asteron-complex (Araneae, Zodariidae). A revision of the genus Asteron with description of the new genus Pseudasteron. Records of the Australian Museum 53(1): 21-36. LINDSEY, T. 1998. Green guide to spiders of Australia. (New Holland: Frenches Forest, Sydney). A REVIEW OF TEMNOSEWELLIA (PLATYHELMINTHES: TEMNOCEPHALIDA) ECTOSYMBIONTS OF CHERAX (CRUSTACEA: PARASTACIDAE) IN AUSTRALIA LESTER R.G. CANNON AND KIM B. SEWELL Cannon, L.R.G. & Sewell, K.B. 2001 06 30: A review of Temnosewellia (Platyhelminthes) ectosymbionts of Cherax (Crustacea: Parastacidae) in Australia. Memoirs of the Queensland Museum 46(2): 385-399. Brisbane. ISSN 0079-8835. New species are described and existing species reviewed of Jemnosewellia, worms living ectosymbiotically on parastacid crayfish, Cherax spp., in Australia. 0 Temnosewellia, Cherax, Australia, ectosymbionts, crayfish. Lester R.G. Cannon and Kim B. Sewell, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia; 3 June 2000, Temnocephalida are dalyellioid rhabdocoels found as ectosymbionts, especially on freshwater crustaceans; they are characterised by a syncytial epidermis divided into a series of plates and a tendency to lose locomotory ciliation (Cannon & Joffe, 2001). Cannon (1986) recognised three families, Scutariellidae from prawns in Europe and Asia, the monotypic Actinodactylellidae from burrowing crayfish from southern Australia and the Temnocephalidae — a large and diverse family with Gondwanan associations. Sewell & Cannon (1996) resolved the position of contro- versial Didymorchis, i.e. in the Temnocephalida and within the Didymorchidae Bresslau & Reisinger, 1933. Cannon & Joffe (2001) also recognised Diceratocephalidae to include Diceratocephala and Decadidymus, each with two anterior tentacles. By far the largest and most diverse family, Temnocephalidae, was first recorded in Australia in 1888 with Temnocephala fasciata Haswell, 1888 and 7. minor Haswell, 1888 from the crayfish Astacopsis serratus (Shaw, 1794) and A. bicarinatus Gray, 1845, respectively. Today these crayfish are known to be several species, respectively in the genera Ewastacus and Cherax. Haswell (1893) added temnocephalans from A. bicarinatus (i.e. Cherax), viz. Temnocephala dendyi Haswell, 1893 and Craspedella spenceri Haswell, 1893. Cannon & Sewell (1995) reviewed Craspedella adding new species and genera and recognising the subfamily Craspe- dellinae. With the exception of Dactylocephala from Madagascar which shows some differences (Cannon & Sewell, 2001), the remaining genera recorded within the Temnocephalidae, viz. Tennocephala, Temnohaswellia, Temnomonticellia, Notodactylus, Achenella and Craniocephala all display a similar facies and may be assigned confidently to the subfamily Temnocephalinae. The largest genus, Temnocephala, has species found on a wide variety of hosts. Recently, Damborenea & Cannon (2001) reviewed members of this genus from the Neotropics and concluded that the Australian representatives should be separated as Jemnosewellia. Here we review 7emnosewellia from Cherax spp. crayfish in Australia. Collection and processing of crayfish and worms and morphological terminology follow Cannon & Sewell (1995). All worms were highly mobile on crayfish, and unless otherwise stated, worms were recorded as collected on the surface of the crayfish exoskeleton, Several species of worms were commonly found in the branchial chamber of their crayfish hosts, but none were located there exclusively. Recognition that the cirrus is a most effective discriminator of species has led to taxonomic descriptions that are more succinct than in previous reports (Cannon & Sewell, 1995; Sewell & Cannon, 1998). In addition, many of the specimens we examine here were collected prior to our adoption of improved techniques requiring the use of live worms, i.e. the use of de Faure’s fluid to elucidate the structure of the cirrus, and the use of silver nitrate to examine the epidermal mosaic (Cannon & Sewell, 1995; Sewell & Cannon, 1998). TERMINOLOGY AND MEASUREMENTS Specimen data are listed in the order: QM registration number; specimen/slide preparation details (in parentheses); host scientific name; locality details; date collected; collector(s); histological fixation/staining procedures. Full registration details are provided for each holotype specimen and for each new locality. For all subsequent specimens listed in the Materials SRO sechion (including paratypes), the QM reg- istration number and specimen/shde preparation details are provided, followed by only thase data which are different from that of the preceding registration. Discrete blocks of registration data are separated by semicolons, Specimens recorded in the Materials section, other than type material, are grouped by craytish hous|, then Australian State. Unless otherwise stated, measurements provided for sofi structures are taken only from the taxonomic type series and those of the cirrus. from special cirrus preparations. Descriptions of the cirrus refer to the inverted State of the organ and exclude fine details of the introvert spines. All measurements were made in microns (jn) with the aid of a drawing tube, The sequence is: B, total body length to tip of tentacles * width al greatest dimension; LE, length from posterior to eyes; SD, sucker diameter; PD, sucker pedunele diameter; PH, pharynx length « width; LA, exeretory ampullae - length » width of left, then right, ampulla; E, diameter of lefl, then right, eye; AT, anterior testes - length >< width of lett, then right, lestis; PT, posterior testes - length « width of left, then right, testis; S, shaft length * width at proximal end; I, introvert length % width. The following other abbreviations are used; Ale,100% ethanol; Bouin, Bouin’s fixative; CP, cirrus preparation; deF, de Faure’s mounting medium; E, East; Form, 10% formalin buffered to pH 7.0 with phosphate: Form-Acetic, Avetic-Formalin-Aleohol (AFA): H, Holotype, H&E, haematoxylin & eosin stain: HF, hot 10% Formalin; HW, hot water; Hx, Mayer's or Harris’s haematoxylin stain; LS, longitudinal serial sections; NSW, New South Wales: NT, Northern Territory: P, Paratype: PP, pigment preparation; QLD, Queensland, QM, Queensland Museum; S, South; WA, Western Australia; WM, wholemount. SYSTEMATICS Order TEMNOCEPHALIDA Family TEMNOCEPIALIDAE Temnosewellia Damborenea & Cannon, 2001 TYPE SPECIES. Jemnosewelli niinor (Haswell, 1888) Damborenea & Cannon, 200), DIAGNOSIS. Temnocephalidae (Temnoceph- alinae) with 5 anterior tentacles, a posterior pedunculate adhesive disc, lacking conspicuous papillate ridges on tentacles or dorsal body, with dark (melanin?) pigment (if any) in body or eyes, MEMOIRS OF THE QUEENSLAND MUSEUM with 5 syncytial plates: 1, tentacular, 2. a characteristic single, saddle-like post tentacular: 3, body: 4, peduncular: and 5, adhesive dise. Furthermore, the excretory pores lie on the body plate, outside the single, post-tentacular plate. Other Australian species from Cherax: Tomnosewellii acirrur sp.nov. Temnosewellia chaerapsis (Hett, 1925) Damborenea & Cannon, 2001 Temnusewellia christincae 3p). Noy. Temnasewellia dendyi (Haswell, Cannon. 2001 Temnosewellia puncita sp. now. Temnosewellia phamusmella sy. nov. KEY TO SPECIES OF TEMNOSEWELLIA FROM AUSTRALLAN CHERAYX SPP, CRAYFISH: |. Lacking pigmentexceptforeyes. oo ey ee 2 Dark pigment tracery on dorsal and sometimes ventral, 1] ot fo a a a Lacking any male selecotic copulatory apparatus T, aeirra sp. nov. With sclerotic male copulatory apparatus bearing | or more large sharp spines at the base of the introvert TF dendvt Dense even pigment in blows emphasising, neural net, Without posterior marginal glands — 7 elisfinewe sp, nov, Dorsal pigment a tracery, with posterior marginal glands4 4+. Marginal glands pastels aca only. from @astern Australia 2. _ . - , 3 Murpicalalandapistesiae ftira WA, iat ate 9 5. Pigment an bie tae, extei wo ‘imaidolbas & cirrus curved H Tomlin Pigment very open. ‘cancertratedt suit excretory pores, fot extendiny, to Tentacles; cirrus shurt and coneal T. phantasmetla sp.nov. 6. Vipment decidedly clumped, mogi notiveable on body periphery and tentacles. several small glands ulong posterior bods margin. 7 panclidta sp, Noy, Piement a dense tracery, two prominent, adjacent glands — “ 5S alpusteriorextremity ofbody, . 2... DT chavrapsts 1893) Damborenen & mw ee Ww Temnosewellia acirra sp, nov, (Figs 1,2 Temnecephala sp. 2: Jolfe & Cannon, 199823, ETYMOLOGY, Latin, lacking a cirrus. MATERIAL. HOLOTYPE: QMGLI8689 (WM), ex Cherax destructor, dam, Eukey Qld (28°46.2’°S 151°59.2°E) 18/Apr/1990, $.D. Cook, Mornm/ttx. PARATYPES: QOMGLIS8690-18691 (WM); QMCLIS693 (LS[1]), Condaming R. Warwick Qld (28°11.4°S 151°57.5°E) 24/Oct'1 992, K.B. & S.G Sewell, Bouin H&E; QMGLIS702 (LS]2]), ex Cheray destructor, Bunyil Ck. Roma (26°30°S 148°487E) 2/Dec/1991, L.R.G Cannon & J.B, Jennings, Form/Hx; QMGL18710(LS[3]), TEMNOSEWELLIA, ECTOSYMBIONTS FROM CHERAX 387 FIG.1. Temnosewellia acirra, Longitudinal section through genital region of QMGLI8710. A-C,d.D, 2. A, gonopore (arrow head), distal muscular ‘penis’ (arrow) and peduncle (Ped). B,, junction of ejaculatory sac (arrowhead) with proximal copulatory bulb dorsal to the distal seminal vesicle junction (arrow), C. vas deferens (arrowhead) entering proximal seminal vesicle adjacent to. copulatory bulb (arrow). D, from vesicula resorbens (VR) to gonopore (arrowhead). A muscular duct connects the vesicula resorbens to the proximal vagina (P) where the vitelline duct (arrow) enters. The proximal vagina is more muscular than the distal vagina (D) opens to the common atrium and blind caecum or copulatory bursa (*), Scale = 100um. 12km SSE of Armidale (30°31°S 51°40°E) 26/Feb/1987, W. Higgins Form/H&E, OTHER MATERIAL. Ex Cherac destructor. QLD: QMGLI87I1 (WM), Marlong Ck, Mt Moflat Nat. Pk (25°02’S 147°54’k), 26/Feb/1986, N.C. Monteith, 70% Alce/Hx; QMGL18703-18705 (WM), Willows gemfield, in dam beside road near Emerald (23°45°S 147°25°E 20/Ocv1990, S.D. Cook, HW/Form/Hx; QMGL18706 (LS{S]). HW/Form/H&E; QMGL18700-18701 (WM), Bungil Ck, Roma (26°30’S 148°48"°E) 2/Dec/1991, L.R.G. Cannon & J.B. Jennings, Form/Hx; QMGL18695-18699 (WM), dam, Eukey (28°46.2°S 151°59.2’E) 18/Apr/ 1990, S.D, Cook, Porm/Hx; QMGL18692 (LS[2]), HW/PForn/ H&E, same locality. NSW: QMGL18707-18708 (WM), 12km SSE of Armidale (30°31°S 51°40°E) 26/Feb/1987, W. Higgins Form/tHx; QMCiL18709 (LS[3]), Form/H&e, SV VD “ik SSS PS FIG, 2. Temnosewellia acirra copulatory structures. A, 3d; B, 2. ES, MEMOIRS OF THE QUEENSLAND MUSEUM between the vesicula resorbens and the top of the muscular vagina; vagina has a large proximal chamber and a smaller distal chamber with ridged walls, this opens via a sphincter into a female antrum which in turn passes to the common genital atrium which is voluminous and opens also to acaecum (bursa copulatrix). Male. Testes with several large lobes when fully mature, anterior testis lies laterally at mid gut level, posterior testis at level of posterior of gut, seminal vesicle relatively thin walled, ejaculatory sac only slightly smaller, copulatory bulb, prostate without large reservoirs, totally lacking any sclerotic armature of male organ which consists of a muscular tube lined with a high epidermis; at the distal end is a sphincter and the duct opens to a large male antrum also lined with a high epidermis, a second sphincter lies at the mouth to the common genital atrium. HOST. Cherax destructor (Parastacidae). ES ejaculatory sac; g, gonopore; P, muscular ‘penis’; PS, prostate secretions; S, ) sphincter; SV, seminal vesicle; VD, vas deferens. DESCRIPTION. General Anatomy. A medium sized, rather thick-bodied worm without body pigment except for two eyespots set close together. A small cluster of posterolateral glands present. Selected measurements are: QMGL 18689 (H): B(1675x1041), LE(1302), SD(651), PD(385), PH(183x426), EA(89 x 59 & 89 x 59), ED(47 & 47); AT(266X219 & 296X237), PT(278X278 & 314 260); QMGL18690 (P): B(1858 x 1000), LE(1479), SD(657), PD(396); PH(195 x 473), EA(95 x 65 & 89 X 59), ED(41 & 41); AT(302 207 & 290201), PT(284x*284 & 308 x 266); QMGL18691 (P): B(1728 x 1087), LE(1361), SD(521), PD(308); PH(237 x 343), EA(IOI1 x77 & 11271), ED(41 & 41); AT(225 x 148 & 278 x 178), PT(278 X 219 & 272 X 237). Reproductive System. Female. No seminal receptacles, but a globular muscular duct lies LOCALITY. Known from tributaries of the Murray Darling system draining westward from Armidale in central NS W north to the Carnarvon region of central Queensland. REMARKS. This is the only species to lack a sclerotic stylus or cirrus. In some sections the muscular proximal tube is seen to push or evert into the male antrum and thus assume the role ofa penis. Another unusual feature is the large bursa copulatrix or caecum opening from the common atrium. This worm lives in the branchial chamber, or nearby sheltered body regions, of C. destructor and alongside Temnosewellia dendyi which it superficially resembles in size and lack of pigment. 7. dendyi has the following characters which serve to distinguish it from the new species: the pharynx is smaller, the posterior testis is set further back behind the gut and it has, apart from a prominent cirrus armed with a TEMNOSEWELLIA, ECTOSYMBIONTS FROM CHERAX 389 distinctive central stylet, a longer and more muscular copulatory bulb in which prostate vesicles are aligned along the duct, and a muscular ejaculatory sac, which is not reflexed, opening via a wide mouth at the back of the copulatory bulb. In the female, there are 3-4 seminal receptacles. Furthermore, the rhabdite glands are fewer and larger. Joffe & Cannon (1998: fig. 3E,J) identified and figured the epidermal mosaic of worms they identified as Temnocephala sp. 2 ex Cherax destructor from the Condamine R., Warwick. We can now formally identify these worms as Temnosewellia acirra. Temnosewellia chaeropsis (Hett, 1925) Damborenea & Cannon, 2001 (Figs 3A,H, 5A, 6A) Temnocephala chaeropsis Hett, 1925:569. MATERIAL, Ex Cherax tenuimanus. WA: QMGL18717- 18722 (WM), Inlet R., on South Western Highway to Walpole (34°55.2’S 116°34.2°E) 25/Jan/1992, L.R.G Cannon & K.B. Sewell, HW/Form/Hx; QMG18716 (LS[1]); QMG217457 (CP), HW/Form/deF; QMG217499- 217500 (CP); QMG217502 (CP); QMG217458 (PP); QMG217501 (PP); QMG217503 (PP). Ex Cherax cf. quinquecarinatus. WA: QMGL18712- 18715 (LS[1 each]), Inlet R., on South Western Highway to Walpole (34°55.2’S 116°34.2’E) 25/Jan/1992, L.R.G Cannon & K.B. Sewell, HW/Form/H&E. DESCRIPTION. General Anatomy. As described by Hett (1925), but with a fine tracery of pigment dorsally, becoming much less dense ventrally. Reproductive System. Female. The female reproductive system has a distinctly bipartite vagina: distally globose and strongly muscular with a powerful distal sphincter, proximally narrow, with distinct seminal receptacles as described by Hett (1925, text-fig. 7). Male. The cirrus is slightly curved (Fig. 5A), with a small, inconspicuous ejaculatory sac. Posterior gland reservoirs often contiguous (Fig. 3H). Selected measurements are: QMG217457: S(186 x 75); 1(50 X 31); QMG217499: S(166 x 62), 1(52x24); QMG217500: S(182 x 80), I(? x25); QMG217502: S(154 x 78), 1(52 x 30). HOSTS. Cherax teniumanus, C. cf. quin- quecarinatus (Parastacidae) LOCALITY. Southwestern WA. REMARKS. Originally known only from some preserved specimens conveyed to England. No types were designated, but the characteristic posterior gland reservoirs as two adjacent bulbs are quite distinctive. Hett (1925) said of T. chaeropsis that the entire animal showed no evidence of pigment (although they had been ‘ for some time in spirit’), but in sections dorsal pigment cells were apparent. Pigment is really quite extensive particularly dorsally. Furthermore, she said ‘the penis, which is straight, has no terminal dilation’, in fact the cirrus does curve slightly, though the introvert is hardly wider than the distal shaft. She claimed also ‘there is no distinct ejaculatory sac’: it is true that this sac is quite small and indistinct. Hett further remarked on the presence of seminal receptacles which she considered unique; in fact, Merton (1913) had described such structures from Temnocephala rouxi. Hett (1925) commented extensively on the paired posterior glandular organs which she believed the distal ends of rhabdite glands (cf Hett, 1925, text-fig. 8 with Fig. 3H). Until Cannon (1993) and Cannon & Sewell (1995) described similar structures from other temno- cephalans, 7. chaeropsis was considered unique in having such glands. This led Cannon to incorrectly identify worms which were taken from WA marron, Cherax tenuimanus, cultured in South Africa, as 7. chaeropsis. It has now been established that such glands are also found in 7, minor, but have been overlooked (Cannon & Watson, 1996). Reports of 7 chaeropsis from cultured marron in South Africa (Mitchell & Kok, 1988; Avenant-Oldewage,1993) are incorrect: the worms are 7: minor. Temnosewellia christineae sp. nov. (Figs 3B, 5B, 6B) ETYMOLOGY. In memory of Dr Christine Cannon. MATERIAL. HOLOTYPE: QMGL18901 (WM), ex Cherax depressus, Gap Ck, Gap Ck Reserve picnic ground, Brookfield, Qld (27°28.7’S 152°55.7’E) Nov/1984, L.R.G Cannon, AFA/Hx. PARATYPE: QMGL18902 (LS[7]); QMGL18903 (WM), 3/Sep/1984; Bouin/H&E. OTHER MATERIAL. Ex Cherax depressus. QLD: QMGL18904-18905 (WM), Gold Coast (28°00°S 153°25’E), Dec/1983, L.R.G. Cannon, Form/Hx; QMGL18906-18907 (LS[3,3]), Bouin/H&E; QMGL18908-18918 (WM), gully near Capalaba (27°32’S 153°12°E) 22/Sep/1988, L.R.GCannon, AFA/Mayer’s; QMGL18919-1892] (WM), Bouin/Hx; QMGL18922-18928 (LS[3,4,7,5,4,7,4]), Bouin/Hx; QMGL18929-18930 (WM), 19/Sep/1988, Alc/Hx; QMGL18931-18932 (WM), Winston Ck Rd, Sheldon (27°34’S 153°12’°E), 18/Mar/1990; |. Forrester & L.R.G. Cannon, Form/Hx; QMGL18933 (LS[5]), Bouin/H&E; 39() MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 3. Temnosewellia species showing pigment patterns (scales A-E= 500m) and posterior glands (scales F and H = 100um, scale G = 50um). A, 7 chaerapsis QMG217458; B, T. christineae QMG217467; C. T, minor QMGL18876: D, T punctatus QMG2174712E, T. phantasmella QMG217477; F, GT. punctatus QMGLI18861: H, T. chaerapsis QMG217458. QMGL, 18934-18937 (WM), Wallaby Ck, on Henderson Rd 0.2km from Mt Cotton Rd junction, Sheldon (27°34.2’8 153°12.8'E) 18/Mar/1990; L.R.G Cannon, Form/Hx; QMGL18938-18939 (L.S[9,7]), Bouin/H&E: QMGL.18940-18941 (WM), Gap Ck, Gap Ck Reserve picnic ground, Brookfield (27°28.7°S:152°55,7’E) Now/1984, L.R.G) Cannon, AFA/Hx,; QMGL18942 (WM), 2/Apr/1984, N. Hall; QMGL18943 (WM), Nov/1985. J. Jennings; QMGL.18944-18945 (WM), Ithaca Ck (27°29°S:152°S7°E) 22/Oct/1988, J. Short & L.R.G Cannon, AFA/Hx; QMGL.18946 (LS[5]), Bouin/H&E; QMGL18947-18948 (WM), gully on Mumford Rd, Narangba (27°12.1°S 152°57.3’E), 22/Apr/|990, J. Short & L.R.G. Cannon, AFA/Ix; QMGL.18949 (LS[9]), Bouin’ /H&E; QMG217459-217463 (CP), creek down stream from Kelly St crossing, Narangba (27°12’S 152°57°E) 19/Oct/1997, J.W, Short, HW/deF; QMG217464-217467 (PP); QMGLI8950-18951 (WM), Kroombit Ck, trib TAG4 (24°22.9°§ |50°59.8°E) 19/Sep/1990, L.R.G, Cannon & K.B. Sewell. HW/APA/Hx; QMGL18952-18953 (LS[4,6]), Kroombit Ck, trib TA47 (24°22.2°8:150°S8.5°E), 20/Sep/1990, L.R.G. Cannon & K.B, Sewell, Bouin/M&E; QMGL18954-18955 (WM), 20/Sep/1990, HW/AFA/Hx; TEMNOSEWELLIA, ECTOSYMBIONTS FROM CHERAX 391 QMGL18956-18960 (WM), Kroombit Ck, Beauty Spot 98 (24°23°S 150°59°E), 21/Sep/1990; QMGL18961- 18962 (LS[4,3]). Ex Cherax robustus. QLD: QMGL18877-18878 (WM), Sunshine Ck, Sunshine Beach (26°24.5’S 153°06.5°E) 16/Apr/1990, J.W. Short & L.R.G Cannon, AFA/Hx; OMGL18880 (WM); QMGL18882 (WM); QMGL18879 (LS[20]); QMGLI8881 (LS[2]); OMGL18883-18885 (LS[8,2,2]), Boutn/H&E. DESCRIPTION. General Anatomy. A medium sized worm with a distinctive pigment pattern dorsally appearing as a series of blocks emphasising the neural network. Selected meas- urements are: QMGL18901 (H): B(3650 x 1635), LE(2336), S$D(633), PD(314); PH(521 x 432), EA(207x 112 & 230X112), ED(65 & 65); AT(432 «284 & 432161), PT(336 263 & 373 X 260); QMGL18903 (P): B(1796 x 1460), LE(1460), SD(509), PD(314); PH(350 x 321), EA(201X 160 & 189142), ED(65 & 77); AT(260X 166 & 272x160), PT(284*172 & 337 X 172). Reproductive System. Female. The vesicula resorbens leads through a short tube and is joined by the ovary, four small seminal receptacles and the vitelline duct before opening to a muscular vagina with a prominent distal sphincter at the entrance to a commodious common atrium. Male. Swollen vasa deferentia enter separately a seminal vesicle so reduced as to resemble no more than a thickened ejaculatory canal. It and the ejaculatory sac, which is large, enter the base of the cirrus side by side together with the prostate gland ducts: there is no obvious copulatory bulb. Selected measurements are: QMG217459: S(468 x 86); 1(77x31); QMG217460: 8(337 x 59), 1(75 X30); QMG217461: S(465 x 80), (77 ¥ 32); QMG217462: S8(468 x 145), 1(75 X 28); QMG217463: S(479 x 109), I(74 x 30). HOSTS. Cherax depressus and C. robustus (Parastacidae). LOCALITY. Known from SEQ and Kroombit Tops CEQ. REMARKS. Potentially confused with T minor, but the pigment pattern is much more regular. This pattern is reminiscent of that illustrated by Haswell (1893) of a young 7 fasciata (Haswell, 1888) from crayfish now recognised as Euastacus spp. Furthermore, the cirrus of 7. christineae 1s at least twice as long with a more oval introvert when inverted than that of 7. minor. This cirrus is not dissimilar to that illustrated by Haswell (1893: pl. XII, fig. 14) as a variant of 7. fasciata, though we now know that a great diversity of worms occur on these crayfish (Cannon & Sewell, 1994). Small and large specimens of this worm swim using rapid dorso-ventral movements of the body and tentacles: it is the only temnocephalan known to swim. Temnosewellia dendyi (Haswell, 1893) Damborenea & Cannon, 2001 (Figs 5C, 6C) Temnocephala dendyi Haswell, 1893:135. MATERIAL. Ex Cherax albidus, VIC:QMGL18748 (WM), creek on Mathison Rd, 6km S of Winchelsea (38°15’S 143°59’E ), 30/Sep/1991, L.R.G Cannon & K.B. Sewell, HW/Form/Hx; QMGL18749 (LS[2]), HW/Form/H&E. Ex Cherax destructor, QLD: QMGL18723-18724 (WM), Bungil Ck, Roma (26°30’S 148°48’E) 3/Dee/1986, L.R.G. Cannon & J.B. Jennings, Form/Hx; QMGL18725-18727 (LS [5,6,8]), AFA/H&E; QMGL18728-18729 (WM), 2/Dec/1986; QMGL18730 (LS[2]), AFA/H&E; QMGL18731 (WM), Western R. 1.5km from Winton on Jundah Rd. (22°24.2’S 143°02.2°E) 22/Sep/1990, S.D. Cook, 70% Ale/Hx; QMGL18732 (LS[4]), Willows gemfield in dam beside road (23°45°S 147°25’°E) 20/Sep/1990, S.D, Cook, 70% Alce/H&E; QMGL18733 (WM), dam at Eukey (28°46.2°S 151°59.2°E) 18/Apr/1990, S.D. Cook, 70% Alc/Hx; QMGL18734- 18736 (WM), Accommodation Ck, near Bald Mountain (28°52.9°S 151°53.7°E) 10/Apr/1990, L.R.GCannon & K.B.Sewell, HW/AFA/Hx; QMGL18737 (LS[3]), HW/AFA/H&E; QMGL18738-18740 (LS[1,1,1)), Condamine R., Warwick (28°11.4°S 151°57.5’°E) 24/Oct/1992, K.B. & S.G. Sewell, Bouin/H&E; QMG?217489-217493 (CP), 4/Aug/1994, K.B. Sewell & B.I. Joffe HW/deF; QMG217494-217495 (CP), 25/Jun/1996, K.B. Sewell, R.D. Adlard & R.D. Sewell; QMGLI8900 (WM), creek by Marlborough Caravan Park, Marlborough (22°49.2’S 149°53.2°E), L.R.G. Cannon & K.B. Sewell, 20/Sep/1990, HW/AFA/Hx. NSW: QMGL18741-18743 (WM), Lake Madgwick, UNE campus, Armidale (30°31°S 151°40°E) 23/May/1991, Zoology Dept. UNE, Bouin/Hx; QMGL 18744 (LS[3]), Yarunga Ck tributary, 1.2km NW Fitzroy Falls Morton NP. (34°38.4°S 150°28.4’E) 19/Oct/1991, L.R.G. Cannon & K.B. Sewell, Bouin/H&E. SA: QMGL18750-18751 (WM), Kangaroo I. (35°50°S 137°15°E) 11/Aug/1995, S. Nichols, Form/Hx; QMGL18752 (WM), Coopers Ck (27°44’S 140°15°E) 26/Nov/1988, |. Beveridge, Form-Acetic/Hx; QMGL18753 (WM), Bool Lagoon (37°09°S 140°43’E) 15/Nov/1988, I. Beveridge, Form-Acetic/Hx; QMGL18754 (WM), Mt Jagged, 9/Feb/1988, 1. Beveridge, Form-Acetic/Hx; QMGL18755 (WM), Lake Merretti (34°01°S 140°46’E), 3/Dec/1988, I. Beveridge, Form-Acetic/Hx; QMGL18756 (WM), The Narrows via Clayton (29°17°S 138°23°E), 2/Dec/1988, I. Beveridge, Form-Acetic/Hx; QMGL18757-18758 (WM), Mt Benson ((37°02'S 139°49°E) 18/Oct/1988, L. Beveridge, Form-Acetic/Hx; QMGL18759 (WM), ane Alexandrina (35°25’S 139°10°E) 20/Jan/1989, |. Beveridge, Form-Acetic/Hx; QMGL18760 raha, Bordertown (36°18’S 140°46°E) 31/Oct/1988, |. Beveridge, Form-Acetic/Hx. VIC: QMGL18745-18746 (WM), creek on road to Ballan 1Skm § of Daylesford (37°21°S 144°09°E) 6/Oct/1991, L.R.G Cannon & K.B. Sewell, HW/Form/Hx; QMGL18747 (LS[2]), HW/Form/H&E. Ex Cherax robustus. QLD: QMG217496, G218301 (CP), trackside pond on McMahon Rd , Bribie I. (27°02.5’S 153°10.3°E) 31/Jan/1995, K.B. Sewell, L.R.G. Cannon, Z. Kalil & J. Short HW/deF. Ex Cherax depressus. QLD: QMG217487-88 (CP), Wallaby Ck, Sheldon, 27/Sep/1994, K.B. Sewell, B.I. Joffe, 1. Solovei & S, Solovei, 1! W/deF. DESCRIPTION. General Anatomy. Largely as described by Haswell (1893). Moderately large and without pigment except for the eyes. Reproductive System. Female. Vesiclar resorbens opening to duct into which open 4 seminal receptacles along with ovary and vitelline duct. Vagina reflexed proximally, muscular with distal sphincter. Common atrium large with a small posterior caecum (bursa copulatrix). Male, The vasa deferentia enter the seminal vesicle separately; the seminal vesicle has narrow distal region surrounded by massed prostate glands which enter the base of the copulatory bulb, their ducts continuing parallel to the ejaculatory duct. Selected measurements are: ex Cherax depressus, QMG217487: S(194 x75), 1(93 X 19); QMG217488: S(158 x95), 1(93 x 22); QMG217489: S(260 x 118), 11101 x 27); QMG217490: (182 x 62), 1(96 X25); QMG217491: $(274 x 139), (101 X27); QMG217492: $(271 x 104), 1(92 x21); QMG217493:; S(259 x 96), 1(95 x 22); QMG217494: (243 x 98), 1(90 x 18); QMG217495: S(158 x 52), (98 x 19); ex Cherax robustus QMG217496: 8(192 x 58), 1(99 x 19); QMG217497: S(183 x 75), 1(99 x 24). HOST. Cherax destructor, C. robustus (Parastacidae), LOCALITY. Known from the Murray-Darling system in Qld., NSW, Vic. and SA, but also from coastal Qld and Kangaroo Is. REMARKS. This and 7 acirra may co-inhabit the branchial chamber and nearby protected locations on the principal host, Cherax destructor. The worms are of similar size and since both lack pigment can be superticially confused (see remarks on T. acirra above). The details of the cirrus of specimens from different hosts and from widely separated regions are very albidus and C. MEMOIRS OF THE QUEENSLAND MUSEUM similar (Figs 5C, 6C show the cirri of worms from Cherax depressus and C. robustus are indistinguishable from those from C. destructor well illustrated by Haswell (1893; pl. 12, fig. 7). The pattern of the mosaic of 7) dendyi ex Cherax destructor trom the Condamine R. Warwick was described and figured by Joffe & Cannon (1998: figs 2J; 4A-C). Joffe & Cannon (1998) identified worms close to T. dendyi from Cherax dispar and C. depressus from Wallaby Creek, Sheldon and described the epidermal mosaic as identical to that of 7. dendyi: these worms were T- dendyi (Fig. 5Ci). Temnosewellia minor (Haswell, 1888) Damborenea & Cannon, 2001 (Figs 3C, 4, 5D, 6D) Temnocephala minor Waswell, 1888:284; Haswell, 1893:134. MATERIAL. Ex Cherax albidus. SOUTH AFRICA: OQMGL 18854-18855 (WM) between Dardanap and Lowden on Patterson Rd (in culture), 6/Sep/1991, HJ. Schoonbee, Form/Hx; QMGL 18856 (LS[2]) Form/H&E. Ex Cherax depressus. QLD: QMGL18963-18966 (WM), Gold Coast, (28°00’S 153°25°E), Dec/1983, L.R.G. Cannon, Form/Hx; QMGL18967-18968 (LS[2,2]), Form/H&E; QMGL18969 (WM), gully near Capalaba, (27°32’S 153°12’E) 22/Sep/1988, L.R.G. Cannon, AFA/Hx; QMGL18970-18972 (WM), Bouin/Hx; QMGL18973-18975 (LS[1,2,3]), Form/H&E; QMGL 18976-18977, 18985-18988 (WM), Wallaby Ck., on Henderson Rd 0.2km from Mt Cotton Rd junction, Sheldon (27°34.2’S 153°12.8’E) 18/Mar/1990, I. Forrester & L.R.G. Cannon, Form/Hx; QMGL18983 (WM), AFA/Hx; QMGL18984 (WM), Bouin/Hx; QMGL18978-18982 (LS[2,3,3,3,3]), Form/H&E; QMG217478-217479 (CP) 22/Sep/1994, K.B. Sewell, BLL. Jofie & LV. & S. Solovei, HW/deF; QMGL18989 (LS[3]), Winston Rd, Sheldon (27°34°S 153°12’E) 18/Mar/1990, I. Forrester & L.R.G Cannon, AFA/H&E; QMGL18990 (WM) Gap Ck, Gap Ck Reserve picnic ground, Brookfield, (27°28.7'S 152°55.7°E) Nov/1984, J. Jennings, Form/Hx; QMGL18991 (LS|2]), Form/H&E; QMGL18992 (WM), creek by Marlborough Caravan Park, Marlborough (22°49, 27S 149°53,2° E), L.R.G. Cannon & K.B. Sewell, 20/Sep/1990, Form/Hx. Ex Cherax destructor, QLD: QMGL18762-18763 (LS[2,2]), Western R., 1.5km from Winton on Jundah Rd (22°24.2°§ 143°02.2°E) 22/Sep/1990, S.D. Cook, 70% Alc/H&E; QMGL18764 (LS[1]), Thompson R. at Longreach waterhole. (23°24.7°S 144°13.8’E) 2/Oct/1990, L.R.G. Cannon & K.B. Sewell, HW/Form/H&E; QMGL18765-18767 (WM), in dam beside road, Willows gemfield (23°45’S 147°25’E) 20/Sep/1990, $.D. Cook, 70% Alc/Hx; QMGL18768- 18770, (LS[1,2,2]), 70% Alc/H&E; QMGLI8771 (WM), Marlong Ck, Mt Moffat NP (25°02°S 147°54’E), 26/Sep/1986, N.C. Monteith, 70% Ale/Hx; QMGL18772 (LS[4]), 70% Alc/H&E; QMGL18773 (WM), Dawson R. Taroom (25°39’S 149°48’E) 3/Dec/1986, L.R.G Cannon TEMNOSEWELLIA, ECTOSYMBIONTS FROM CHERAX & J.B. Jennings, Form/Hx; QMGL18774 (LS[3}), Form/H&E; QMGL18775-18777 (WM), Bungil Ck, Roma (26°30’S 148°48’E), 2/Dec/1991, L.R.G Cannon & J.B. Jennings, Form/Hx; QMGL18780-18781 (LS[2,2]), Form/H&E; QMGL18778-18779 (LS[2,3]), 3/Dec/1991, Bouin/H&E; QMGL18782 (WM), Wilson R. at Noccundra, W of Thargomindah (27°49°S_ 142°35’E) 17/Apr/1990, G.B. Monteith, 70% Alce/Hx; QMGL 18783-18784 (LS[1,1]), Condamine R., Warwick (28°11.4°S 151°57.5°E) 24/Oct/1992, K.B Sewell & S.G. Sewell, Bouin/H&E; QMGL18785-18786 (WM), Feb-Mar/1993, L.R.G.Cannon, silver nitrate; QMG217481-217482 (CP), 4/Aug/1994, Sewell & B.I. Joffe, HW/deF; QMG217483 (CP) 1/Sep/1994, K.B. Sewell; QMG217484 (CP) 25/Jun/1996, K.B. Sewell; R.D. Adlard & R.D. Sewell; QMG217485-217486 (PP); QMGL18787-18788 (WM), dam at Eukey (28°46.2S 151°59.2°E) 17/Apr/1990, S.D.Cook, 70% Ale/Hx; QMGL18789-18791 (LS[4.3,3]), 70% Alc/H&E. NSW: QMGL18804 (WM), Lake Madgwick, UNE campus, Armidale (30°31°S 151°40°E ), 26/Feb/1987, W. Higgins, Bouin/Hx; QMGL18805-18807 (WM) 23/May/1991, Zoology Dept. UNE; QMGL18808 (LS[4]), Bouin/H&E; QMGL18809-18811 (WM), Yarunga Ck trib., 1.2km NW Fitzroy Falls Morton NP. (34°38.4’S 150°28.4’E ), 19/Oct/1991, L.R.G.Cannon & K.B.Sewell, HW/Form/Hx; QMGL18812-18813 (LS[4,3]), Bouin/H&E; QMGL18814-18818 (WM), trib. of Murrumbidgee R. on Nanangro Rd, near Childowlah, outside Yass (34°51’S 148°55’E), 16/Oct/1991, L.R.G.Cannon & K.B.Sewell, HW/Form/Hx; QMGL18819 (LS[3]), HW/Form/H&E. VIC: QMGL18820 (WM), creek on Mathison Rd, 6km S of Winchelsea (38°15°S 143°59°E) 30/Sep/1991, L.R.G. Cannon & K.B. Sewell, HW/Form/Hx; QMGL18821 (LS[3]), HW/Form/H&E. SA: QMGL18822 (WM), Cowell (33°41’S_ 136°55°E) 8/Dec/1988, I. Beveridge, Form-Acetic/Hx; QMGL18823 (WM), Lake Merreti (34°01'S 140°46°E), 3/Dec/1988; QMGL18824 (WM), Avenue Ra. (37°05’S 140°18’E), 18/Oct/1988; QMGL18825 (WM), Onkaparinga R. (35°00’S 138°49’E) 22/Feb/1988; QMGL18826 (WM), Murray R., Murray Bridge (35°07’S_ 139°16’E) 10/Feb/1989; QMGL18827 (WM), Lake Alexandrina (35°25’S 139°10°E) 20/Jan/1989; QMGL18828 (WM), Inman R. (35°30°S 138°31°E) 19/Nov/1988; QMGL18829 (WM), Bordertown (26°18°S 140°46’E) 31/Oct/1988; QMGL18830-18831 (WM), Mt Benson (37°02’S 139°49’E), 18/Oct/1988; QMGL18832 (WM), Bool Lagoon (37°09’S 140°43’E), 15/Nov/1988. Ex Cherax dispar, QLD: QMG217480 (CP) Wallaby Ck, on Henderson Rd, 0.2km from Mt Cotton Rd junction, Sheldon (27°34.2’S 153°12.8°E) 22/Sep/1994, K.B. Sewell, B.I. Joffe & LV. & S. Solovei HW/deF; QMGL18792-18794 (WM), Woodgate Lagoons (25°07.4’°S 152°30.6°E) 6/Apr/1991, K.B.Sewell, Form/Hx; QMGL18795 (WM) 18/Sep/1990, L.R.G. Cannon & K.B.Sewell, Form/Hx; QMGL18796 (WM), Form/unstained. Ex Cherax quadricarinatus. QLD: QMGL18761 (WM), Deception Bay (in culture) (27°10’S 153°05°E) 393 FIG. 4. Temnoasewellia minor. Scale = 500um. 22/Oct/1986, B.Herbert, Form/Hx. Ex Cherax tenuimanus. QLD: QMGLI18797-18801 (WM), Ipswich (in culture) (27°37°S 152°47°E) 12/Oct/1982, L.R.G Cannon, Form/Hx; QMGL18802- 18803 (WM), Esk (in culture) (27°14’°S 152°25°E) 28/Sep/1982, L.R.G. Cannon, Form/Hx. SA: QMGL18833 (WM), Parilla (in culture) (35°18°S 140°40°E) 11/Oct/1988, I. Beveridge, Form-Acetic/Hx. WA: QMGL18834 (WM), Inlet R., SW hwy. to Walpole (34°55.2’S 116°34.2’E) 25/Jan/1992, L.R.G Cannon & K.B. Sewell, HW/Form/Hx; QMGL 18835-18836 (WM), 394 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5. Nomarski interference contrast photomicrographs of De Faure’s preparations of the whole cirri of Temnosewellia species. A, T. chaeropsis QMG217457. B, T. christineae QMG217462. C, T. dendyi - (i) QMG217488 ex Cherax depressus, (11) QMG217497 ex C. robustus. D, T. minor QMG217482. E, T phantasmella QMG217475. F, T. punctatus QMG217469. Scale = 100um. Dam near Byford (32°13’S 116°00’E), 22/Feb/1990, L. QMGL18841-18844 (LS[2,2,2,1]), Form/H&E; Evans, Form/Hx; QMGL18837-18838 (LS[3,2]), QMGL18845 (WM) Western Australia, Novw/1989, Form/H&E; QMGL18839-18840 (WM) WA Fisheries, L.Evans, Form/Hx; QMGL18846-18847 (LS[1,1]), Fish Health Dept., 6/Aug/1991, T. Thorne, Form/Hx; Form/H&E; QMGL18848-53 (WM) Margaret R. Marron TEMNOSEWELLIA, ECTOSYMBIONTS FROM CHERAY 39 Farm, 10km S Margaret R. (34°00.8°S JIS°09.9°E) 23/Jan/ 1992, L.R.G. Cannon & K.B. Sewell. Fonn/Hx. SOUTH AFRICA: QMGL18868-18872 (WM) fin culture), 1991, A. Avenant-Oldewave, Form/Carmine: OMGL [8873-18875 (WM) (in culture), 1982, DL. Rok, Form/Carmine. DESCRIPTION. General Anatomy, Medium sized temnocephalan with pigment dorsally very variable but usually with some open patches (Fig. 3C): itextends forward from the eyes mainly into the proximal or basal regions of the central 3 tentacles; ventral pement only a Lracery to about the level of the mouth. Although they can be overlooked, clusters of posterolateral glands oecur (Cannon & Watson, 1996). Reproductive System. Female, Vesicula resorbens Opens to a muscular chamber and then to the relative short muscular vagina with a powerful sphincter distally before opening to the commonatrium. No evident seminal receptacles. Male. Testes deeply notched on lateral borders, not generally overlapping. Cirrus gently curved with introvert only slightly inflated, Selected measurements are: OMG217478: $193 * 67), H25* 19); OMG217479° $(98 * (5B); (33. 19): QMG217480: S(72 * $2), (34% 22); QMG2I74B1: SUG05 S89), 134% 18); QMG217482: 5(294 * 61). W374 18); OMG217483; S(343~% 70). WAT 15); OMG217484: S(38 #21), 40 ® 19), HOSTS. Cherax desiruetor, C_ dispar, C depressus and, in culture, C albidus, C quddricarinutus and CC. tenuimans (Parastacidae). LOCALITY. Found in the whole of the Murray -Darling system Qld, NSW, Vic and SA, and also coastal Queensland from Marlborough south where Cheray depressuy and C. dispar occur. Found on cultured and Itee range marron. C. lenninnuias, ii WA and elsewhere where this crayfish 1s cultured, notably Japan (Oki, Tamura, Takai & Kawakatsu, 1995) and South Africa. This worm has been found also on European crayfish presumably contaminated in aquaria holding Australian crayfish (Nylander. 1997), but now 1s believed to have escaped Into streams in Bavaria, Germany (Xylander, pers. comm.). REMARKS. Haswell (1888, 1893) clearly characterised this species. No types exist, but there seeins little possibility now of confusion about the identity of this worm. which is so common and widespread, The most striking difference or diserepaney with the original accounts (Laswell, }888,1893) is the presence of - postero-lateral glands (Cannon & Watson, 1996), These glands do not stain consistently and can easily be overlooked. 7 minvr is now known to be the worm infecting cultured marron in South Africa. It appears to have a catholie choice of hosts and obyiously may become widespread. Reports of this species from Cherax punctatus by Cannon & Jennings (1987) should be referred to C. depressus, Details of the cirrus are essentially as presented by Haswell (1893) as can be seen from the de Faure’s preparation (Fig. 5D; 6D), We observed no more than minor differences in proportions in specimens from different localities. Furthermore, our specimens mostly resemble Llaswell's (1893, pl. 15, fig. |) regarding the distribution of the tesles ie. they are clearly separated and not overlapping, as his text states (though with equivecation). The patiern of the mosaic of 7 minor ex Cherax destructor fram the Candamine R., Warwick was described aiid figured by Joffe & Cannon (1998: figs 2C_F-G, 3C). Temnosewellia punctata sp, nov. (Figs 3D,F.G 5F, 6F) ETYMOLOGY. In reference to the punctate coalescerice of the dorsal pigment. MATERIAL. HOLOTYPE: QMGLIS&57 (WM), @s Chere, ct. qilinguecarinans, Carbutup R,, 2kin N of Carbunup at railway bridge, WA (33°40.8'°S 115° .8°E) 23/Jan/1992. L.R.G. Cannon & K.B. Sewell, HW/Form/Hx. PARATYPES: OMGL188S8-18859 (WM); OMGL1S860-T8861 (LS{1.3]), HW/FormV/HA&t. OTHER MATERIAL. Ex Cheray ef. guinguecarincatas, WA: OMGL 18862-18863 (WM). - Carbunup R.., 2km N of Carbunup at railway bridge (33°40.8°S [1S°1L.8°E) 23/Jan/1992, L.R.G. Cannon & K.B. Sewell, HW/Form/Hx; OMGLIS864 (LS[2]). HW/FornvHa&hs; QMG217468-217469 (CP) HW/70% Ale/deP: OMG217497 (CP): QMG217470-217471 (PP); QMG21 7498 (PP). Ex Cheray tenuimanus, WA: QMGL18865- 188G7 (WM). al catchment weir, Margaret R. (33°57,0°S. 115°05.2"E), 23/Jan/1992, L.R.G. Cannon & K.B. Sewell. HW/Porm/Hx DESCRIPTION. General Anatomy, Medium sized temnocephalan with distinctive pattern of pigment with punctiform clusters (Fig. 3D). Posterior gland reservoirs numerous and spread in a wide are along the posterior margin of the body. Selected measurements are; QMGL|8857 (H): B(I432% 858). LE(ISI4). SDi414), PRU 66); PH(225 4 260), EAU42 © 77 & LTR ® 77), ED(4L & 47); ATO89 © 130 & 237% 124), PT(QAT IIS & 2435 % 1601; OMGL] 8855 (P). Bi 1686 » 746), 396 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 6. Nomarski interference contrast photomicrographs of De Faure’s preparations of the introverts of cirri of Temnosewellia species. A, T. chaeropsis QMG217457. B, T. christineae QMG217462. C, T. dendyi - (i) QMG217488 ex Cherax depressus, (ii) QMG217497 ex C. robustus. D, T. minor QMG217482. E, T phantasmella QMG217475. F, T. punctata QMG217469. Scale = 50 um. LE(1065), SD(367), PD(142); PH(294 x 254), EA(89 x 65 & 89 X 65), ED(36 & 36); AT(124 x 112. & 154x101), PT(219x130 & 195x130); QMGL18859 (P): B(1941 x 805), LE(1219), SD(385), PD(148); PH(195 x 278), EA(I0I x 71& 95X77), ED(47 & 47); AT(189 X 118 & 183 x 118), PT(231 X 130 & 219 x 136). Reproductive System. Female. Vesicula resorbens with 4 small seminal receptacles at its base; ovary joins below these at the top of a long, narrow, proximal vagina which enters an extremely voluminous and highly muscular distal vagina. This in term opens via a strong distal sphincter to a small common atrium. Male, A moderately small ejaculatory sac joins the top of the cirrus bulb with the prostate glands and the ejaculatory duct. Cirrus gently curved with slightly inflated introvert (Fig. 6F). Selected measurements are: QMG217468: S(222 x 93), 1(72 x 31); QMG217469: S(223 x 100), 1(70 x 30); QMG217497: S(164 x 62), 1(75 x 30). HOSTS. Cherax cf. quinquecarinatus and C. tenuimanus (Parastacidae) LOCALITY. Southwestern WA. REMARKS. This species most closely resembles 7! chaeropsis with which it shares not only the posterior gland reservoirs, but also the large and muscular distal vagina which clearly must serve as an ootype or uterus. The most striking differences are the pigment pattern which in this species is distinctively punctate, not a tracery, and the much smaller, but more numerous and more broadly distributed posterior gland reservoirs. TEMNOSEWELLIA, ECTOSYMBIONTS FROM CHERAX 397 Temnosewellia phantasmella sp. nov. (Figs 3E, 5E, 6E) ETYMOLOGY. Latin, from phantasma =a spirit: literally a little spirit from the appearance of the pigment pattern. MATERIAL. HOLOTYPE: QMGL18893 (WM), ex Cherax rhynchotus, Lake Wicheura, Cape York, Qld (10°46’S_ 142°34’E) 27/Sep/1990, P.J.F. Davie & J.W.Short, 70% Alc/Hx. PARATYPES: QMGL18894- 18895 (WM); QMGL18891-18892 (LS[1,2]), Ale/H&E. OTHER MATERIAL. Ex Cherax rhynchotus. QLD: QMGLI8889 (WM), Lake Wicheura, Cape York, (10°46’S_ 142°34°E) 27/Sep/1990, P.J.F. Davie & J.W.Short, Alc/Hx; QMGL18896 (WM); G217472- 217475 (CP) Alc/deF; QMG217476-217477 (PP); QMGL 18890 (LS]2]), Lake Boronto, near Somerset, Cape York (10°45’S_ 142°35’E) 25/Sep/1974, GB. Monteith, Alc/H&E. DESCRIPTION. General Anatomy. Small slender worm with eyes set very close together and a distinctive pigment pattern concentrated anteriorly leaving lacunae about excretory ampullae. Pharynx robust and sucker set well posterior; prominent postero-lateral glands, Selected measurements are: QMGL18893 (H): B(1609 x 337), LE(680), SD(148), PD(71); PH(130 x 124), EA(S59 x 41 & 53 X 41), ED(24 & 24); AT(118 xX 107 & 136X107), PT(142*77 & 136 X 89); QMGL18894 (P): B(840 x 320), LE(562), $D(148), PD(77); PH(83 X77), EA(41 x36 & 41 x36), ED(18 x 18); ATU118 X59 & 118 x65), PT(83 X59 & 83X65); QMGL18895 (P): B(746 x 266), LE(485), SD(130), PD(53): PH(107 X 59), EA(41 x 36& 41 x30), ED(18 & 18); AT(95 X 53 & 77 X59), PT(65 X 53 & 71 X65). Reproductive System. Female. The vesicula resorbens opens to a short muscular vagina and then to a large common atrium with a low epithelium. Male. Seminal vesicle large and ejaculatory sac a small bulb enter the base of the cirrus side by side: no evident copulatory bulb. Selected measurements are: QMG217472: S(79 x45), (16 X 12); QMG217473: S(72 * 34), I(15 x 11); QMG217474: 8(64 x 29), I(16 x 11); G217475: 8(76 x 43), [17 x 12). HOST, Cherax rhynchotus (Parastacidae) LOCALITY. Known only from Cape York, Queensland. REMARKS. This small worm is characterised by a cirrus dramatically smaller than in any other Temnosewellia from Cherax. Its closest relative appears to be 7. butlerae known from the freshwater crab, Holthuisana transversa, taken from Augathella, western Queensland. However, the pigment pattern is dissimilar: in 7, phantasmella there is an anterior concentration spreading laterally and surrounding the excretory ampullae as two clear lacunae, but in 7) butlerae there is a small concentration before the eyes and then a straggling tracery in about 3 longitudinal bands, The eyes in 7: butlerae are set farther apart, the common atrium has very well developed epithelium and the seminal vesicle and ejaculatory sac are slender. Furthermore, the hosts and localities are very different. Temnosewellia sp. MATERIAL. Ex Cherax punctatus. QLD: QMGL18886 (WM), Dingo Ck near Traveston (26°19°S 152°47°E) Mar/1973, S.R. Monteith, Alc/Hx; QMGL18887 (LS[2]), Alc/H&E. DESCRIPTION. A small and apparently eyeless species. REMARKS. The single specimen QMGL18886 cannot be readily referred to another species. The host crayfish is a deep burrowing one and may therefore rarely emerge. An eyeless species of worm from a host that rarely emerges into daylight is thus not unexpected. Haswell (1893) reported the eyes to be ‘very small’ in 7: engaei (Haswell, 1893) from the burrowing land crayfish Engaeus fossor from Gippsland. Similarly, the temnocephalans reported from the burrowing isopod Phreatoicopsis terricola, T. caeca (Haswell, 1900) and 7. geonoma (Williams, 1980), are blind and without pigment. Teratological specimens of temnocephalans without eyes, however, are sometimes encountered, but usually can be readily placed. It is not possible to describe this species formally as there is far too little material and we are reluctant to infer too much about the absence of eyes in one specimen. DISCUSSION Cannon & Sewell (1995) showed that subtle differences in male copulatory structures can indicate species separation in temnocephalans (Craspedella spenceri and C. simulator); such differences are not readily apparent within Temnosewellia, e.g. T. minor. We recognise, however, that only a thorough meristic study from live preparations using de Faure’s fluid would reveal such differences. Furthermore, life history and ecology of Temnosewellia spp. suggests no more than regional variations will be 398 found: these worms are active external dwelling worms which colonise new hosts much more readily than do species of Craspedella which are found in the sheltered branchial cavity and consequently show higher host specificity. Clearly the two WA species of Temnosewellia are close as would be expected. Low host specificity is particularly manifest, however, with 7. minor which has the potential to spread around the world. The worms 7. acirra and T. dendyi which occur predominantly in the sheltered branchial habitat of Cherax destructor may be shown to exhibit niche separation or competition. This matter needs investigation, for the two species were confused from live study: only a microscopic examination of the male copulatory structures can distinguish them. Questions arise concerning the recent bio- geography of 7 minor from C. dispar and C. depressus. T. minor is widespread in the Murray Darling system and has been translocated around the country and the globe in aquaculture. Its occurrence on the two small Cherax from coastal Queensland seems to indicate a recent inter- mingling of streams across the watershed of the Great Dividing Range. Musyl & Keenan (1992) suggested this as an explanation for some freshwater fish in central Queensland being found on both sides of the divide, i.e. from both the western Warrego and coastal Dawson drainages. 7. minor could have spread south from here along the coast to include both C. dispar and C. depressus as hosts: molecular techniques may be able to indicate when this may have happened. On the coast, 7. christineae appears the ecological homologue to 7. minor. Is competition between these species evident? Other Australian species currently in 7emno- cephala s. l. and found, for example, on crabs, shrimps and crayfish have also been tentatively assigned to Temnosewellia by Damborenea & Cannon (2001), pending more thorough review. Further, the eyeless worm specimen from C. punctatus, a rarely encountered burrowing crayfish with a limited distribution, is likely to prove a new species (other eyeless species are known), but more specimens are needed to rule out any teratology. The exclusion of 7: rowxi from this account is tentative. Though previously known only from Aru Is. (Indonesia), Cannon (1991) reported it from cultured red-claw (Cherax quadri- carinatus) from the NT. It and Temnocephala MEMOIRS OF THE QUEENSLAND MUSEUM semperi Weber, 1889 from Asian freshwater crabs may belong to a separate, as yet undescribed, genus (B.I. Joffe, pers. comm.). ACKNOWLEDGEMENTS We would like to acknowledge the support of ABRS who funded the initial studies and to the Queensland Museum. Mrs Zeinab Khalil assisted ably and cheerfully with laboratory routines. LITERATURE CITED AVENANT-OLDEWAGE, A. 1993. Occurrence of Temnocephala chaerapsis on Cherax tenuimanus imported into South Africa, and notes on its infestation of an indigenous crab. South African Journal of Science 89: 427-428. CANNON, L.R.G. 1986, Turbellaria of the World - a guide to Families and Genera. (Queensland Museum: Brisbane). 1991. Temnocephalan symbiotes of the freshwater crayfish Cherax quadricarinatus from northern Australia. Hydrobiologia 227: 341-347. 1993. New temnocephalans (Platyhelminthes): ectosymbionts of freshwater crabs and shrimps. Memoirs of the Queensland Museum 33: 17-40. CANNON, L.R.G. & JENNINGS, J.B. 1987. Occurrence and nutritional relationships of four ectosymbiotes of the freshwater crayfishes Cherax dispar Riek and Cherax punctatus Clark (Crustacea: Decapoda) in Queensland. Australian Journal of Marine and Freshwater Research 38: 419-427. CANNON, L.R.G. & JOFFE, B.I. 2001. The Temnocephalida. Pp.83-91. /n Littlewood, D.T.J.& Bray, R.A. (eds) Interrelationships of the Platyhelminthes. (Taylor & Francis: London). CANNON, L.R.G. & SEWELL, K.B. 1994. Symbionts and biodiversity. Memoirs of the Queensland Museum 36: 33-40. 1995. Craspedellinae subfamily nov. (Platyhel- minthes:Temnocephalida) from the branchial chamber of Australian crayfish. Memoirs of the Queensland Museum 38: 397-418. 2001. Observations on Dactylocephala madagas- cariensis (Vayssiere, 1892) a temnocephalan with twelve tentacles from Madagascar. Zoosystema 23: 11-17. CANNON, L.R.G. & WATSON, N. 1996, On the posterior rhabdoid glands of Temnocephala minor. Australian Journal of Zoology 44: 69-73. DAMBORENEA, C. & CANNON, L.R.G. 2001. On neotropical Temnocephala (Platyhelminthes). Journal of Natural History (in press). HASWELL, W.A. 1888. On Temnocephala, an aberrant Monogenetic trematode. Quarterly Journal of Microscopical Science 28: 279-303. 1893. A Monograph on the Temnocephalae. Macleay Memorial Volume (Linnean Society of New South Wales): 94-152. TEMNOSEWELLIA, ECTOSYMBIONTS FROM CHERAX 399 HETT, M.L. 1925. On a new species of Jemnocephala (1. chaeropsis) (Trematoda) from West Australia. Proceedings of the Zoological Society of London: 569-575. JOFFE, B.I. & CANNON, L.R.G. 1998. The organisation and evolution of the mosaic of the epidermal syncytia in the Temnocephalida, (Plathelminthes: Neodermata). Zoologischer Anzeiger 237: 1-14. MERTON, H. 1913. Beitrage zur Anatomie und Histologie von Temnocephala. Abhandlungen Senckenbergische Naturforschende Gesellschaft 35: 1-58. MITCHELL, S.A. & KOK, D.J. 1988. Alien symbionts introduced with marron from Australia may pose a threat to aquaculture. South African Journal of Science 84: 877-878. MUSYL, M.K. & KEENAN, C.P. 1992. Population genetics and zoogeography of Australian freshwater Golden Perch, Macquaria ambigua (Richardson 1845) (Teleostei:Percichthyidae), and electrophoretic identification of a new species from the Lake Eyre Basin. Australian Journal of Marine and Freshwater Research 43: 1585-601. OKI, I., TAMURA, S., TAKAI, M. & KAWAKATSU, M. 1995. Chromosomes of Zemmocephala minor, an ectosymbiotic turbellarian on Australian crayfish found in Kagoshima Prefecture, with karyological notes on exotic turbellarians found in Japan. Hydrobiologia 305: 71-77. SEWELL, K.B. & CANNON, L.R.G. 1996. The taxonomic status of Didymorchis paranephropis Haswell, 1900. Memoirs of the Queensland Museum 42(2): 585-595. 1998. New temnocephalans from the branchial chamber of Australian Ewastacus and Cherax crayfish hosts. Proceedings of the Linnean Society of New South Wales 119: 21-36. XYLANDER, W.E.R. 1997. Epidermis and sensory receptors of Zemnocephala minor (Plathel- minthes, Rhabdocoela, Temnocephalida): an electron microscopic study. Zoomorphology 117: 147-154. 400 RANGE EXTENSIONS FOR TWO POORLY KNOWN QURENSLAND SNAKES (lemmas of He Oneensant Musenin dij) 400, 2007 — Purine diitnall/ (Worrell, 1955) and Sas dievert (Worrell 1956) are amongst the least known of Australia’s many poorly known clapids. The former, narrowly confined to mid and southeastern Queensland, is. known from paly 2 specimens tn the Australian Museum and isa ‘vulnerable’ species (Longmore, 1986: Ingram & Raven. 1991; MeDonald et al, 1997; Copmer etal, 1993; [ICN Council. 1994, Nature Conservation Ledislation Amendment Repulation (No, 2) 1997), Siderchever? occurs from southern Queensland to northern Vicloria ind, 1 museum collections areany guide, 1s merecommon, It has ne special conservation status (Longmore, 186; Ingran & Raven, 1990) but its tata status i$ uneertin (Cogger, 2000: Cogger et al. 993), urine recent surveys ny) Queersland’s Brigalow Gell and Desert Uplands Bioregions, one specinien of cach species was discovered find both extend the known distributions Furina dunmatii This snake, collected thom the outskires of Clermont (22°45°S, 147°35°E), in ihe Northern Brigalow Bell Bivregion, was preseded to the Emerald office of the Qutecnslind Parks und Wildlile Survive for identification (ate summer, 1999). The specimenowas identified as Fo dinmeall and lodged with ie Queensland Museum (QMI73000). I was melanistic, lacking pale dilfuse bloteles/burring on the upper lips. (Such barring eecurs on all specimens in the OM collection. P Couper. pers. comm), Vegetation communities adjacentia Clormont include Aweu/vprus caglaheh (cootibab ) and £ pepilned (poplar box) woodlands on alluvials and & crebre. (narrow-letyed ironburk) and dead rhadaxylon (rosewood) on shallow reky soils. Two specimens inthe \aistraliin Museum came trom the southern Brigalow Belt (27°15'S, 149°41°F) and the South-east Queensland Bioregion (2737'S, S37 by. The Queensland Muscun: has 7 specimens from the southern Brigalow Bell (Oakey 27°26'S, JUSS F; Miles 26°4°S, ISOTTER, Taroom 25°)8'S. 149°" E> and Maidenwell 267315. SIRE): two trom the South-east Queensland, Bioregion (Archookooru 26°44°S, 131°45"L and Gladstone 23°S0S. ISTP LAE); and one irom the Northern Brivalow Beli (Mt Archer, yia Rockhampton 23°28. (50°38), the Clermont specimen af & danmeall) extends the distribution approximately 300km to the north west et Mt Areher It is the first from the Central ightands at the northern Brigalow Bell Bioregion, an area assailed by tree elearing. Suu dwyert An adult Sue ahevert (nomenclature after Hutchinson, 1990) Was removed by the authors fron a pial trap on the 12/11/99, while surveying in poplar box open woodland on Monklunds Station via Alpha. Southern Desert Uplands Bioregion (23°2R48"S_ 146025 13°). Identification was vorilied by PJ. Couper of the Queensland Museum and the specimen Indeed in the (Queensland Muscum collection (OMITFIIZ1). The locality was a large (about 40,000ha) poplar box and silyer-leaved ironbark woodland with an open grassland understorey dominated by deride spp. (Wircurdsses). Bolleiwehlod ewartiand (desert blueprass). Chrvsepogon fallax (golden beardgrass), Heterapagon contortion (hack spear grass), and Triacia mitehellir (buck spinifex) on texture contrast spils, Swe cw (Worrell, 1956) hus un unstable taxonomic history. I) has heea placed in Denison. Unectis— Mamuplacephalis and Suite; und has been trédted os a subspecies ot Sufa snectabitiy (Kretl, 1869), ce Worrell, MEMOIRS OF THE QUEENSLAND MUSEUM 14956; Longmore, |YS6; Wilson & Krowles, UBS: Hutchinson, 1990; Ingram & Raven, 1991; Chmann, 1992; Cogper. 2000. The species Is poorly known and its distributian il-detined. Covacevich & Couper (in Ingram & Raven.199]) showed i) to he contined primarily (in Queensland) to the eastern Mulga lands, Southern Brigalow Bell, and South-east Queensland Bioregions. Two specimens have alsa been collected front the northern Brigalow Belt; Newlands Mine neur Muekuy (21°11 8S. 147°5a'R: QMI65065), and bowhsville (Mere S, 146 40 b- AMR] 19422), The Monklands specimen is the first record from Queens- jund’s Desert Uplunds Bioregion, The nearest So dyerd Joculity is al Mt Moflatt in the Carnarvon Ranges (20°35'S. 14013": QMJ59373). 225kor to Ue Southeast: the Newlands Mine reeord is 300kim to the northeast. Suni dave Js How koown to fige through the Malwa Lends, Soptheast Queenslund, Brigalow Bell and the Desert Uplands Bioregions. Acknowledgments We thank (he Natural Herifae ‘boost for finding support and thy Queensland Parks and Wildlile Service, Queensland Muscum. Queensland Herbarium, Grillith University, ‘Tropical Savanoas Cooperative Rescurch Centre and the Parks and Wildlife Commission of the Northern Lerritory for in-kind support for the survey component of this work Jeanette Coyacevigh, Patrick Couper and Andrew Amey of the Queensland Museum and Ross Sadlier of the Australian Museum kindly ussisicd with the preparation oF this ute, Thanks also to Juliana MeCosker and Richard Johnsen whe offered comments on an earlier drat Literature Cited COOGER, HAG. 2100, Reptiles end anpnibians of Ausualia Oth Lalition, (Reed New Hollabd: Ss dacyt COGGER, HL, CAMERON, PB SADLIRE RA & KOGLER P 1999) The action play tor Ausialian nepoles. (Australia Nature Conser ation Ageney Canberra) PUMANS, |) 1992. Lneyelupedia-of Agsuiiian annals ~ Reptiles (Angus & Roberson; Pymble, NSW); pp, 495 HUTCHINSON, MON 19900 The generic classification of Uye \uatralian terrestrial ehapid snukes Memoirs af the Ouvenstand Musewn 242); 397-405 TNORAM, G0 & RAVIEN, RO (eds) $99. Anatlas of Queensland's frogs, repules. binds and mranimals, (Queensland Museu, Brisbane) IUCN COUNCIL 1944 1UC'N Red List Categones. Prepated iy the TLIC Species Suevival Contmission as approves! by the 40th mectina of Hye ION Counesh Gland. Siwitverlund, 40 Now, 164 TLIC Copneily Gland), LUNGOMORE, R. 1986. Avbs of elapid- snakes ol Australia. Australia Plo und Fauna Serius 7) Australia Government Publishing Servpoes Cunber'ra), MLUGONALD. KR. COVACEVICH, TAL INGRAM, (1. & COUPER, IP )99T The status af frogs and repules.. Pp. 338-345. |i bngram, Gad, & Rayer, Ru feds), An atlas ot OQuversland's (roys. cepliles, birds und mammuls. (Qucensiand Museom: Brishunes WILSON, SK & KNOWLES, DG, (988, Australia's reptiles A pholopraphie reference to the terrestrial reptiles of Austrailia (Collins Sydney) WORRELL, E, 1956, A ney suake from Ouevnsland, Ausialian Zouldpist 12: 202-205. David Hannal and Nicole ¥, Thirgarw Oueenstund Marks avid Wildlife Seeview, PO Bax “th, Emerald 4724. Australia, 4 November 2000. ADDITIONS TO THE ROSTROCONCH FAUNA OF AUSTRALIA AND CHINA ALEX G. COOK AND NATALIE CAMILLERI Cook, A.G. & Camilleri, N. 2001 06 30: Additions to the rostroconch fauna of Australia and China. Memoirs of the Queensland Museum 46(2): 401-407. Brisbane. ISSN 0079-8835. Eight taxa of rostroconch are described from stratigraphically and geographically dispersed localities in Australia; Euchasma caseyi and Euchasma colliveri sp. nov from the Ordovician (Caradoc) of Tasmania; Kandosoconcha pembertoni gen. et sp. nov. from the Lochkovian- Pragian Yellowmans Creek Beds near Kandos, NSW; and Hippocardia sp. from the Lochkovian-Pragian Martins Well Limestone, NQ. Conocardium sowerbyi de Koninck, is decribed from the Emsian Brogans Creek Lst of the Capertee Valley, and a neotype for the species is designated. A single rostroconch is described from the Middle Devonian of Guangxi, China. Rostroconchs are described for the first time from the Carboniferous of Australia. 0 Rostroconchs, new taxa, Ordovician, Devonian, Carboniferous, Australia, China. Alex G. Cook & Natalie Camilleri Queensland Museum PO Box 3300, South Brisbane 4101, Australia (e-mail: alexe(@@qm.qld.gov.au); 20 November 2000. Ordovician rostroconchs are known from central Australia (Pojeta, et al., 1977) and Tasmania (Etheridge,1883; Pojeta, 1979). Euchasma caseyi Pojeta et al.,1977 and E. colliveri sp. nov. are described from the Gordon Group near Zeehan Tasmania, in shales from near the Zeehan Smelters Quarry which Banks & Burrett (1989) correlated with their OT17 or OT1I8 faunal assemblage (Caradoc). Known Australian Devonian rostroconchs come mainly from the Lower Devonian of SE Australia (de Konninck, 1876; Creswell, 1893); Etheridge, 1881; Chapman, 1908; Fletcher, 1943; Talent, 1956; Talent & Philip,1956; Tassell, 1982) with only C. gogoense Fletcher (1943), known from Upper Devonian of the Canning Basin WA Early Devonian material described herein comes from: 1)Yellowmans Creek Beds, near Kandos NSW; 2) Brogans Creek Lst in the upper Capertee Valley NSW, and 3) the Martins Well Limestone Member of the Shield Creek Fmn, Broken River Province, north Queensland. The Yellowmans Creek Beds are a thick sequence of shales with minor limestone and calcareous shales near the base (Pemberton et al 1994, Colquhoun 1996). Kandosoconcha pembertoni gen. et sp. nov. is derived from the lowermost 10m of the formation above the Kandos #1 limestone quarries. The locality has yielded Ozarkadina remschiedensis remschiedensis, Amydrotaxis sexidentata (Mawson in Cook, 1988) and is regarded as late Lochkovian to earliest Pragian. The Brogans Creek Limestone has been re- garded as lateral equivalent of the Carwell Creek Group (Pemberton et al, 1994). Colquhoun (1995) suggested an early Emsian age for the limestone based on conodonts. Shelly faunas currently under investigation show strong species-level similarity with the ‘Receptaculites’ Limestone of Taemas, and thus for the single horizon containing Conocardium sowerbyi de Koninck, a late Emsian age is probable. The Martins Well Limestone Member of the Shield Creek Formation is regarded as spanning the Lochkovian-Pragian boundary (Jell et al., 1993). Hippocardia sp. were collected in the uppermost part and are here regarded as Pragian. Conocardium gogoense Fletcher 1943 has been recovered from a silicified fauna in the Pillara Limestone (Frasnian) in the Hull Range, Canning Basin WA. (cf. Playford & Lowry, 1966). A paucity of biostratigraphically-useful fossils from the Pillara Limestone in the Hull Range make exact placement of the occurrence within the Frasnian impossible at present. Fletcher (1943) concluded there was only one vague reference to Australian Carboniferous rostroconchs. The specimens described herein are from the Utting Calcarenite, Utting Gap identified by Roberts (1971) as Viséan in age. Terminology follows Pojeta & Runnegar (1976) and Pojeta (in Boardmann et al., 1987). 402 SYSTEMATIC PALAEONTOLOGY Phylum MOLLUSCA Class ROSTROCONCHIA Pojeta, Runnegar, Morris & Newell, 1972 Order CONOCARDIOIDA Neumayr, 1891 Superfamily EOPTERIOIDEA Miller, 1889 Family EOPTERIIDAE Miller, 1889 Euchasma Billings, 1865 Euchasma caseyi Pojeta, Gilbert-Tomlinson & Shergold, 1976 (Fig. 1A-E) Euchasma caseyi Pojeta, Gilbert-Tomlinson & Shergold 1976: 27, pl. 20, figs 8-15, pl. 21, figs 1-9, pl. 23, figs 1-10, pl. 24, figs 1-5. MATERIAL. QMF37321-37323, two conjoined valves and a fragmentary right valve, from QML901, Zeehan Smelters Quarry, Zeehan Tasmania. DESCRIPTION. Shell medium sized, up to 10.8 mm long, 11.4mm high, 8.2mm wide; laterally expanded, posterior outline circular, oblique straight anterior face, very weakly concave. Rostrum prominent, extending at least two thirds along the dorsal margin. Anterior faces reclined 60° from horizontal, with modest ribs stronger towards the gape. Gape a characteristic ‘key hole’ shape, oval with invagination above and slit like with interlocking denticles below. Posterior shell dominated by evenly spaced ribs. REMARKS. The only slight difference between this and the Georgina Basin material Pojeta et al. (1976) is its coarser denticulation in the anterior gape. This species differs from E. colliveri by the architecture of the gape, weaker ribs and lack of the prominent carina. Euchasma colliveri sp. nov. (Fig 1F-H) ETYMOLOGY. For F.S.Colliver. MATERIAL. Holotype: QMF37324 from QML901, Zeehan Smelters Quarry, Zeehan Tasmania. DIAGNOSIS. Concave upper anterior face convex lower anterior face, strong carina and subtriangular outline. DESCRIPTION. Shell medium sized, oblique, sub-triangular in outline, moderately inflated, 11.6mm long, 9.8mm high, 8.4mm wide. Rostrum elongate, extending nearly to posterior margin. Dorsal margin slightly arcuate. Anterior face reclined at approximately 60° from the horizontal, bordered from posterior shell by MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. A-E, Euchasma caseyi Pojeta, Gilbert- Tomlinson & Shergold, 1976. A-C, QMF37322, right lateral, anteroventral and posterior views, x 2.4. D, QMF37323, left lateral view, * 2.4. E, QMF37324, internal view, 2.4. F-H. Euchasma colliveri sp. nov. Holotype QMF37321 x2.4, right lateral, posterior and dorsal views. prominent raised carina. Anterior face concave near the dorsum, lacking ornament dorsally, convex towards the venter where it is adorned by strong but thin ribs and finer comarginal growth ROSTROCONCH FAUNA OF AUSTRALIA AND CHINA 403 lines, Posterior shell bears 6 ribs of subequal strength and numerous fine lines continuing the rib-dominant reticulation on the shell anterioir to the main carina. Rostral clefis obscured. REMARKS. Reticulation and the carina suggest affinities with £. shwarkoi Pojeta et al., 1976, but that taxon bears finer more numerous ribs, more robust ribs on the anterior face, and a less pronounced carina, The specimen differs from the type E. blumenbachii (Billings) (Pojeta & Runnegar, 1976, pl. 27) by having coarser ornament and a prominent carina, Superfamily CONOCARDIOIDEA Miller, 1889 Family HIPPOCARDIIDAE Pojeta & Runnegar. 1976 Hippocardia Brown, 1845 TYPE SPECIES. Cardiwn hibernicum Sowerby 1815 trom the lower Carboniferous of Ireland. Hippocardia sp. (Fig. 2) MATERIAL. QMF33798-33801 from OQML541, Martins Well Limestone Member of the Shield Creek Formation, Broken River Province, north Queensland. DESCRIPTION, Medium to large, up to 25.2mm long, |}3.4mm high and approximately as wide as high. Dorsal margin straight with umbo mid-shell projecting slightly above hinge line. Prominent short rostrum situated just below dorsal margin. Snout long with wide anterior gape. A strong ridge and a smaller ridge posterior to it form a week hood separating the posterior and anterior parts of the shell, Posterior face at approximately 60< to vertical with a concavily to rostrum. Ornament reticulate, evenly so on anterior shell, ribs stronger on posterior. REMARKS. This poorly preserved material is generically assigned on the basis of the elongate snout and weak hood. ? Family HIPPOCARDUDAE Pojeta & Runnegar, 1976 Kandosoconcha gen. nov. TYPE SPECIES. Kanedosacancha pembertoni sp. nov. ETYMOLOGY. For the Kandos district, NSW. DIAGNOSIS. Minute, trapezoidal, with prom- inent mid-shell carinae and intercalated ridges. FIG. 2. Hippoeardia sp. A-C, QMF33798, * 2, lefi lateral, ventral and dorsal yiews. DE, QMF33800, * 2, right and lett lateral views. F, OQMF33801. * 2.4, right lateral view REMARKS, Gross anatomy, prominent carinae and reticulation suggests affinities with Mulceo- dens Pojeta & Runnegar or Bigalea Pojeta & Runnegar. \du/ceodens was established for taxa with a constriction in the ventral part of the snout. but the restriction 1s lacking in the Kandos species. Bigalea has 2 rostral hoods. and a slit-like aperture, The Kandos species is closest to and shares the reticulation and trapezoidal shape of B. visbevensis Pojeta & Runnegar from the Silurian of Gotland differing by intercalation of ridges, no development of hoods, rather having carina, 404 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. A-I, Kandosoconcha pembertoni gen. et sp. nov. A,B, Holotype QMF37325, x 20. C, Paratype QMF37326, X16, left valve internal view. D, E, Paratype QMF37328, X16, right valve. F,G, Paratype QMF37327, x 10, left valve. H,I, Paratype QMF37329, x 10, left valve. J-L, Conocardium sp. 1. QMF41130, x 3.2, left lateral, ventral and anterior views. Kandosoconcha pembertoni sp.nov. (Fig. 3A-D) ETYMOLOGY. For John Pemberton. MATERIAL. Holotype: QMF37325 (right valve), paratypes QMF37326-37330 from QML1026, top bench of Kandos #1 Quarry, W of Kandos NSW, Yellowmans Creek Beds, Early Devonian. DIAGNOSIS. As for genus. DESCRIPTION. Shell minute, up to 2.8mm long, 2.7mm high, trapezoidal,with 2 inter- mediate strength ribs between 2 major mid shell carinae. Dorsal margin nearly straight, umbones above dorsal margin. Anterior part of shell equantly triangular, with equally reticulate orna- ment, or on some specimens slightly dominant comarginal fine lines. Posterior portion smaller than anterior, with weak fine comarginal lines. Mid-shell with 3 or 4 strong ribs on the posterior ridge, the strongest forming a carina with next most robust ridge penultimate anterior also forming a carina. Minor ridges intercalated some originating from mid shell, not from umbones. Reticulation continuing on mid-shell. Gape nar- row with some denticulation. Rostrum unknown. Family CONOCARDUDAE Miller, 1889 Conocardium Bronn, 1835 TYPE SPECIES. Cardium elongatum Sowerby 1815. Conocardium sowerbyi de Koninck 1876 (Fig. 4) Conocardium sowerbyi de Koninck 1876:109; de Koninck 1898 (transl. ): XX. Conocardium sp. Tassell 1982: 2 pl. | fig.1. TYPE. This taxon was not illustrated by de Kon- inck, 1876 and destruction of the material by fire rendered the name dubious in Tassell’s (1982) opinion. Complicating this, the type locality was nebulously given by de Koninck as Yass district, with the host lithology as a black limestone. Thus the name is based on a non-existant type from an unclear locality. | here designate a neotype, ANU 36845 from the Receptaculites Limestone ROSTROCONCH FAUNA OF AUSTRALIA AND CHINA FIG. 4. Conocardium sowerbyi de Konnick, 1876. A-C, QMF37306, x 3.5, left lateral, posterior and dorsal views. D-F, QMF37307, X 3.5, right lateral, dorsal and anterior views. G,H, QMF37209, 2.7, left lateral and internal views. I, QMF37312, x 2.7, dorsal view of snout. J,K, QMF37310, x 2.7, right lateral and internal views. LLM, QMF37308, x 2.7, left lateral and internal views. described and figured as Conocardium sp. (Tassell, 1982). Given other taxa described by de Koninck, and Tassell (1982a), such as Mur- chisonia turris and Mitchellia striatula, are also abundant in the Receptaculites Limestone the most likely source of the conocardiid was this unit. Whilst the species was not illustrated by de Koninck and the name unused, there are reasonable grounds to conclude Tassell’s material is topotypic and conspecific. MATERIAL. QMF37306-QMF37312 from QML1027, Brogans Creek Limestone, Brogans Creek, Upper Capertee Valley, NSW. Devonian (Emsian). DESCRIPTION. Small to medium sized, up to 11mm long, 8mm wide and 7mm high. Long rostrum just below and at slight angle to hinge. Umbo subcentral, well above hinge line. Valves moderately inflated. Anterior gape slit-like wide- ning close to dorsum to form the oval snout, with elongate denticles. Short apertural shelves. Pegma at 45° to hinge. Broad flattened ribs on mid and posterior shell, widest and most pronounced on mid shell, ribs without intercalation. Anterior ribs finer. Fine growth lines present. Posterior-most shell and rostrum lacks ornament. REMARKS. The narrow slit, broad ribs, and aperture confined to the end of the shell demonstrate that this material is conspecific with that described by Tassell (1982) and de Koninck’s (1876). Conocardium gogoensis Fletcher, 1943. (Fig. 5A,B) MATERIAL. QMF36085, QMF42233 from QML1033, Pillara Limestone, Hull Range, Canning Basin, WA; Devonian (Frasnian). DESCRIPTION. Moderately inflated, up 7mm long, and 6.5mm high, umbones well above dorsal margin. Antero-dorsum straight, rostrum not preserved. Posterior face at 30° from horizontal, Anterior surface with wide numerous flattened ribs. Posterior at least partially ornamented by finer ribs. Fine detail lacking due to coarse silicification. REMARKS. Of the many hundreds of kilograms of limestone dissolved only these two specimens of rostroconch were recovered. Fletcher’s (1943) specimens are similarly coarse ribbed, and equally long as high. 406 MEMOIRS OF THE QUEENSLAND MUSEUM PIG. 5. A,B, Conocardium gogoense Fletcher, 1945. A, QMF36085 right valve, x 4; B, QMF42233 left valve, x 4. C-R, Conocardium sp. 2. C-G, QMF37317, * 3.2, right lateral, posterior, anterior, dorsal and ventral views; H-K, QMF37314, x 4 right lateral, dorsal, inclined posterior and ventral views; L, M. QMF37316, x 1.6 left lateral and internal views; N,O, QMF37317 x 3, posterior and right lateral inclined views;was not illustrated P-R, QMF37320, X 2.5, left lateral, dorsal and ventral views, ROSTROCONCILLAUNA GF AUSTRALIA AND CHINA Conocardium sp. | (Fig. 3J-L) MATERLAL, OMF41330 fram the Dongangling Formation, 7km NW of Ertang, Wuxian County, Guanxi Province China; Middle Devonian (Givetian). DESCRIPTION. Medium sized. tumid. quadrale in oulline, 7.3mm high, 7.1min long, 7.2mm maximum width, Rostrum well below dorsum, projecting from short sttep convex posterior face, Gape widening strongly dorsally, with long denticules. Shell reticulate with subequal ribs dominant on anterior face, comaryinal lines fine, very numerous. Posterior face with coarse ribs and much finer radial lines. Rostral cleits unknown. REMARKS, This stout taxon differs from the type species C. elonguituni (Sowerby) by the shortness of the shell. From Conocareium sewerhyi, fromthe Devonian of Taemas, it diflers in its more (umid shell form. [ can find no other records of Middle Devonim rostrovonchs from Guangxi. Conocardiom sp, 2 (Fig. SC-R) MATERIAL, QME3 7313-37320 Irom QM) L095, Uting Calcarenite, Utting Gap Bonaparte Gulf Basin, WA; Carbonilerous, Visean. DESCRIPTION. Shell, medium sized, up to 19mm long }3mm high and 9mm wide, moderately inflated, rosirum prominent, slightly below the hinge axis, gemly inclined upwards from the shell margin, Rostral cletts unknown. Anterior gape with weak denticles. Ornament of flattened wide ribs, with comarginal rugae vontined to posterior surface. LITERATURE CITED BANKS, MLR. & BURRETT, C.b, 1989 The Gordon Group (Early Ordovician to Early Silurian)- mainly platform carbonates, Geological Society of Australia Special Publication 15; 201-242, CHAPMAN, F. 1908, A monograph on the Silufan bivalved Moflusea of Victoria. Memoirs of the National Museum of Victoria 2: |-62. COLOQUHOUN, GLP, 1995. Carly Devonian cunodont faunas from the Capertee High, NE Lachlan Fold Belt, southeastern Australia. Courier Forschungsinstitut Senckenber | 82: 547-369. COOK. A.G, 1988. Aspects of the lower Kandos Limestone, New South Wales. Unpuhl BSe (Hons) thesis, University of Wollongong. NSW, CRESSWELL, A.W. 1893, Nates on the Lilydale Limestone, Proceedings of the Royal Society of Victoria N.S. 5S: 38-46. 407 ETHERIDGE. R. 1881. Sones on a collection of fossils trom the Palaeozoic rocks of New South Wales. Journal and Proceedings of the Royal Society of New South Wales |4: 247-258. 1883_ A deseription of the remains of trilobites from the Lower Silurian racks of the Mersey River District, Tasmania. Papers and Proceedings of the Royal Society of Tasmania 1882: 150-163, FLETCHER, HD. 1943. The genus Coneeardinn trom Australian Palaeozoic rocks. Records of the Australian Museuin 21: 231-243, JEL. S., SIMPSON, A., MAWSON. R. & TALUNT. LA. 1993, Biostratigraphic summary, Pp. 239-245. In Withnall, 1.W, & Lang, S.C. (eds) Geology of the Broken River Provinee. north Queensland, Queensland Geology 4, (Department of Minerals and Fnergy: Brisbane). KONINCK, LG. De, 1876, Recherches sur les fossiles paleozaiyues de la Nouvelle-Galles du Sud (Australie). Memoaires de la Societe Royale des Sciences de Liege. Detuxienie serie. 6; 1-135. PEMBERTON ,J.W., COLOQUHOUN. GP, WRIGHT, A... BOOTIL AN. CAMPBELL, 1.C,, COOK, AG. & MILLSTEED. B.D. 1994, Stratigraphy and depositional environments of the northern Capertee High. Proceedings of the Linnean Society of New South Wales E14: 105-134 PLAYFORD. TLE. & LOWRY, 1.0. 1966. Devonian reef complexes of the Canning Basin, Western Australia, Geological Survey of Western Austrilia Bulletin 118: 1-150. POJETA Jr, J) 1979 Geographic distribution af Cambrian and Ordovician rostraconch mollusks. In Gray, J, (ed) Historical biogeography, plate teclonies, and thechanging environment. (Oregon State University Press: Corvallis). POIJIBTA Jr J. GILBERI-TOMLINSON, J. & SHERGOLD, JH. 1977, Cambrian and Ordovician restroconch molluses trom northern Australia. Bureau ol Mineral Resources, Geology and Geophysics Bulletin 17): 1-54, pls (-27, POJETA, J. & RUNNEGAR, B. 1976, The pdeontolugy ofpostreonch mollusks und the curly history of the phylum Mollusca. Uniled States Geoloical Survey Prolessiunal Paper 968: |-S8, pls 1-54. ROBERTS, JR. L971. Devonian and Carboniferous brachiopods from the Bonaparte Gulf Basin. northwestern Australia. Bureau of Mineral Resources Geology and Geophysics 122:1-319- TALENT, J.4.1956. Devonian brachiopods and pelecypods of the Buchan Caves Limestone Victoria. Proceedings of the Royal Sociew of Victoria NS. 6821-56. TALENT, JA. & PHILIP, GM, 19564. Silura-Devonian Mollusea from Marble Creek, Thomson River, Victoria, Proceedings of the Royal Society of Victoria ns. 68> 57-72. TASSELL, C-B. 1982. Some Siluro-Devonian rostro- conch oelluses front southeastern Australia. Records of the Queen Vietaria Museum 79: If 1. 408 HERPETOLOGICAL ‘FOREIGNERS’ ON NORFOLK ISLAND, AN EXTERNAL TERRITORY OF AUSTRALIA. Memoirs of the Queerisland Museum 46(2): 408. 2001:- ‘Biotas of islands, especially oceanic islands, characteristically differ from continental biotas in four ways. They are relatively impoverished, unsaturated, and disharmonic, and they harbor a disproportionately high number of endemic species. This last trait~ high endemism — means that island species are crucially important to global biodiversity, while the first 3 traits are often seen as causing island species and communities to be particularly fragile. This is the island dilemma — great biodiversity, much of it not found elsewhere, but great danger’ (Simberlofi, 2000). The Norfolk group (29°02’S 167°57°E) is a yoleanic and sedimentary island cluster 1,367km east of Australia, 772km northeast of New Zealand and 675km south of New Caledonia (Schodde et al,, 1983). The extant and extinct biotas of these islands are highly endemic and, predictably, have affinities with those from Australia, New Zealand and New Caledonia/the tropical Pacific (Holloway, 1977; Cogger etal., 1983; Schodde et al., 1983), Reptiles are scant on the Norfolk group. Two species — a gecko, Christinus guentheri (Boulenger, 1885) and a skink, Oligosoma lichenigera (O° Shaughnessy, 1874) — occur there. Populations referred to these species oceur also on the Lord Howe | group, 900km to the east. Whether these taxa are conspecific with the Norfolk populations is uncertain. Norfolk and Lord lowe populations are known to be distinet to ‘some degree’ (Cogger et al.. 1993). Neither C. guentheri nor OQ. Jichenigera occurs on the main island (Norfolk) in the former group, although C. guentheri is reported from Holocene deposits there (Cogger et al,, 1983). In the Norfolk group, C_guentheri occurs on Phillip I, Nepean I., Moo’oo Rock and Bird Rock. O. lichenigera is known from only Phillip l. Both species are reputed to have ‘disappeared’ from Norfolk 1, due, largely, to the introduction (by Polynesian colonists some 900 ybp) of Ratius exulans. Reputed declines of populations of both species on Phillip 1. have been attributed to massive habitat destruction there, by introduced rabbits, pigs and goats (Cogger et al., 1993). Both C. guentheri and O. lighenigera are classed ‘endangered’ (Cogger et al., 1993), Should the Norfolk group populations prove to be distinct from those on the Lord Howe group, the already narrow occurrences of each will be halved and their vulnerability to extinction will increase concomitantly, Frogs and other amphibians are unknown in both fossil and modern records for all islands in the Norfolk group. Given the uniqeness of the Norfolk group’s biota and its high vulnerability to further population declines and extinction, we report a series of reptile and amphibian specimens recently introduced to the main island of the Norfolk group. Specimens of 3 coloniser species have been found either on the island or in cargo for the island on a vessel anchored there, since 1998: Bufo marinus (Linnaeus, 1754), Cane Toad QMJ74062 9 near the Colonial Hotel, Burnt Pine, Norfolk [.; Neil Tavener, mid 1999, This specimen was collected following its sighting by hotel staff and by a visitor from Cairns, NEQ. Literia caerulea (White, 1790) Green Tree Frog. QM J74063, near watertank by workshops, Norfolk [. airport. Charles Buffet, 18 Noy. 1999; QMJ74064 ‘Pot Pourri’ shop. Burnt Pine, Norfolk 1., Joan Kenny, early 1998; QM174065 in pallet of airfreight from Brisbane, SEQ, Norfolk L., ca 1998. Hemiductylus frenatus Duméril & Bibron, 1836. QMJ74060 Norfolk I, ona drum of telephone cable imported to the island from Australia, via Yamba, ca June 2000, Glen Williams and Tony Grant, 13 Sept., 2000. QMJ74061, amongst timber on board MV Southern Moana anchored al Norfolk L. to discharge cargo, Neil Tayener, early 2000. MV MEMOIRS OF THE QUEENSLAND MUSEUM Southern Moana serves Auckland, Norlolk 1, Raratonga, (Cook Islands) and, occasionally, docks in Aitutaki (Cook Islands) and Nuie Island. These are the first specimens of herptiles foreign to the Norfolk I. group to be recorded there. Cogger et al, (1983) conducted detailed herpetological surveys on all islands in the group and reported the presence ofonly C. guentheri and O. lichenigera. Since that time, no other herpetological specimens from the Norfolk group have been added to the collections of the Australian Museum, Sydney (R. Sadlier, pers. comm.). B. marinus, L. caerulea and H_frenatus are aggressive and highly successful colonisers elsewhere. B, marinus, intro- duced to a handful of sugar cane-growing districts in coastal Queensland between 1935 and 1938, now occurs from the Mann R., NT, to northeastern New South Wales (Covacevich & Archer, 1975; R. Alford, pers. comm., October, 2000). L. caerulea is nalive to, common, and widespread in northern and eastern Australia. It adapts exceptionally well to urban areas and to agricultural and grazing lands and is known to travel easily with produce and building material. /7. frenctus isa recent, very successful coloniser of Australia. It is presumed to have entered Australia with cargoes from ports in Asia and/or the Pacific Islands. Specimens were observed in Australia first in Darwin in 1964, but 1. frenatis is believed to have entered Australia before then, Horner, pers. comm; 2000; Wilson, 2000. 17. frenatus is now widely distributed in and near Queensland ports from Cooktown to the Gold Coast (Queensland Museum records) and has been reported recently from Adelaide, SA (M. Hutchinson, pers. comm., September, 2000). In Brisbane. SEQ, /7. frenatus has been extraordinarily successful, Earliest Brisbane specimens were collected on the wharves in 1983 (QMJ41978), This species is now probably the most common gecko in Brisbane, having moved from wharves, to inner city buildings and parks, to suburban homes/gardens with what appears to be astonishing success. Discovery of specimens of these 3 species on the Norfolk group has implications for its future. All are highly successful in new, especially disturbed, habitats; thrive in the subtropics: and, if they gain even a tiny ‘foothold’ on the island/s, will become conspicuous elements of the Norfolk fauna, already dominated by species introduced by Europeans; one species (H. frenatus) may have the potential to ‘out compete’ the gecko (C, guentheri) endemic to the group; and two species (B. marinus, L. caerulea) will fill the present vacant amphibian/riparian niche on Norfolk L. on a massive scale. Literature Cited COVACEVICH, J. & ARCHER, M. 1975, The distribution of the cane toad, Bufo marinus, in Australia and its effects on indigenous vertebrates. Memoirs of the Queensland Museum | 7; 305-310, COGGER, H.G., CAMERON, B.E.. SADLIER, R.A. & EGGLER, P. 1993, The Action Plan for Australian Reptiles. (Australian Nature Conservation Agency: Canberra). COGGER, H., SADLIER, R, & CAMERON, E. 1983. The terrestrial reptiles of Australia’s island territories. Special Publication II (Australian National Parks and Wildlife Service: Canberra), HOLLOWAY, J.D. 1977, The Lepidoptera of Norfolk Island: their bigreography und ecology. (Dr W. Junk B.V.: The Hague). SCHODDE, R., FULLAGAR, P. & HERMES, N. 1983. A review of Norfolk Island birds: past and present. Special Publication 8. (Australian National Parks and Wildlife Service: Canberra) SIMBERLOFF, D, 2000, Extinction ~ proneness of island species — causes and management implications, The Raffles Bulletin of Zoology 48(1):1-9. WILSON. S. 2000. Geckos. The eyes have it. Australian Geographic. 59; 72-89. JA, Covacevich', AF. Buffett, PJ Couper’ & AP. Amey’, ‘Queensland Museum, PO Box 3300, South Brishane 4111; “Health and Quarantine, Kingston, Norfolk Island 2899; 12 May 2001 AUSTRALIAN SPECIES OF ARISTELDAE AND BENTHESICYMIDAE (PENALOIDEA: DECAPODA) W. DALL Dall, W. 2001 06 30: Australian species of Aristeidae and Benthesicymidae (Penaevidea: Decapoda). Memoirs of the Queensland Ntusevm 46(2); 409-441, Brisbane. ISSN 0079-8835. Twelve species of Aristeidae Irom Australian seas, representing all genera in the Jamily, have been identified (*indicates new fees Arixtueomorpha foliacea, Arisiacopsis edwardsiuna*Aristeas mahahissae, A virilis, Austropenaens. nilidus, *Hemipenaeus carpeneri, *Hepamudus tener, * Paruhepomadus vaubeni, *Plesispendeus urmatus, *P coruscans, *Pseudaristeus kathleenae, *P sibogae. (Aristeix semidentatis has also been recorded from Aus{ralia, but its identity could not be contirmed in existing museum collections). In the Benthesicymidae ten species have heen identified: Benthesievmes investigetoris, Bo urinator, Gennadas hauveri, G capensis, G. gilchristi, G incertus, G kenipl. G prapinqueus, G. scutatus, G, tinavrei, plus anew Subspecies Benthesicyvmus urinate hawensis. Definitions of the 2 families and the genera represented, with keys. are included. Keys to the Indo-West Pacific species are viven, together with diagnoses of the Australian spec ies, Zoogeography of the 2 families is discussed briefly O /ado-Wesy Pacific. Aristeidae, Benthesicymidae, Australia, diagnoses, disiribution, zeogewgraphy: W. Dall. Queensland Museum, PO Box 3300, South Brixbane, Queensland 4101. Australia: J November 2000 Up to the late 19th century all penaeoid deca- pods were included in the Penacidae. [t was recognised, however, that there were major dif- ferences between some groups and Wood-Muson (1891) identified 3 distinet deep-water groups in the Penaeidae: Aristacina, Benthesicymina and Solenoverina. Later, the Aristaeina was raised to fainily status to include the Aristeinae. Benthesi- cyminae and Solenocerinae. These 3 sub- families have been raised to family level within the Penacoidea (Pérez Farfante & Kensley, 1997). Thus there are now 5 families within this super-family, distinguished as follows. KEY TO THE FAMILIES OF TILE PENAEOIDEA |. Postorbital*® spine presen . Solenoceridae Posturbilalspineabsent 2. 6. 2 ee 2. Third to fifih pleopods aniramous, . Steyoniidae Third to-fAilth pleopods biramous. 0... . 3 3. Dorsal rostral teeth plus postrostral teeth 0-2, rarely 3: rostrum truncate, deep und ranging fram short al to barely exceeding the cornea: adrostral carina absent; antero-ventral Eeraace usually emareinate . Benthesicymidae Dor si rostral teeth sus postrostral teeth more than 2; rostrum usually well exceeding the cornea: aurostral carina usually present; anlero-yentral region of the curapace noldeepls emarginaie. = 2 _.4 4. Upper antennular Vagellum much shorter than the lower and attached laterally to the third seement of the antennular pedal, Erie reduced 19 a selase knob. oy ey weet ot . Aristeidae Upper anfennular flagellum of similar length to (he lower and attached apically to the third segment of the pean PRAHA well developed and foliaceous ies ic) ee . Pengeidae * In same solenocerid wenera a Ve could be identilied as i postantennal sping, In the Aristeidac Parahepomadys Was a posGinlemnal spine, bai the antennulir Magella and long rostrum with only three teeth readily: distinguish f from the Solenoceridue. Twentyseven species of Solenoceridae have been identified from Australian seas (Dall, 1999), but although several species of Aristeidae are common in deeper water commercial prawn trawls, only 13 Australian species are described inthis paper . The Benthesteymidae are small and delicate and few were in the collections of Australian museums until the advent of extensive investigations using mid-water trawls, which collected 8 Gennadas species from Australian seas. While the Solenoceridae largely inhabit the continental slope, the Aristeidae are mostly found from the lower slope into deeper water. exceptionally down to 5,0Q0m. Some Benthesi- eymidae are mesopelagic or bathypelagic, others are benthic, often at depths below 1,000m. Key taxonomic papers on the Aristetdae and Benthesicymidae are by Crosnier (1978, 1985), Crosnier & Forest (1973), Kensley (1971), Kensley etal. (1987), Kikuchi & Nemoto (1991), Pérez, Farlante (1973, 1987), Pérez Farlante & Kensley (1997). Most of these publications and others describe specimens fram outside 410 Australian waters. This paper therefore attempts to cover the Australian species in sufficient detail to facilitate their identification by non- specialists. Definitions are given of the families and genera, with keys. Species keys include known Indo-West Pacific species, because it is likely that, in the future, some additional species will be found in Australian seas. The species diagnoses and figures are from specimens in the collections of the Australian (AM), Northern Territory (NT), Queensland (including Museum of Tropical Queensland) (QM), Tasmanian (TMH) and Victorian (MV) Museums and the CSIRO Marine Laboratories, Hobart. Generally synonomies are restricted to primary and key references, especially where the commoner species, such as Aristaeomorpha foliacea have a very extensive synonymy. General taxonomic features of the carapace and appendages are identified in Grey et al. (1983) or in Dall et al.(1990). The special taxonomic features of the Solenoceridae Dall (1999) are also applicable to the Aristeidae and Benthesicymidae, except for the nomenclature of the anterior branchiostegal region. In the Benthesicymidae the anterior end of the branchiostegal emargination is often mark- ed by an angular projection, the infra-antennal angle. Also, all members of both families have, near or on the margin of the carapace, a branchiostegal spine, which is continuous with a carina of varying length. It is similar to the hepatic carina of other families, particularly when it extends past the lower end of the cervical sulcus. In many other species it stops well short of this region and could equally be called a branchiostegal carina. For the sake of uniformity it will be described here as an hepatic carina. Length. Except when scale bars are included in figures, lengths are carapace length (CL) i.e. distance between the posterior rim of the orbit a os oe Pleurobranchs =| Arthrobranchs | Arthrobranchs | 2 Arthrobranchs MEMOIRS OF THE QUEENSLAND MUSEUM and the midline of the posterior rim of the carapace. SYSTEMATICS Superfamily PENAEOIDEA Family ARISTEIDAE Wood-Mason, 1891 Aristaeina Wood-Mason, 1891: 278. Aristaeinae Alcock, 1901: 27; Ramadan, 1938: 36; Kubo, 1949: 193. Aristeinae Bouvier, 1908: 6, 13; Balss, 1957: 1516; Crosnier, 1978: 14. Aristeidae Crosnier, 1978: 14; De Freitas, 1985; 3; Squires, 1990; 20; Pérez Parfante & Kensley, 1997:32. Aristaeidae Grey et al., 1983: 14; Dall et al., 1990: 58. DIAGNOSIS. Rostrum usually elongate, apparently sexually dimorphic in some genera, being shorter in adult d 3; with 3 or more dorsal teeth, without ventral teeth. Antennal and branchiostegal spines always present, postorbital and pterygostomian spines absent; postantennal and hepatic spines rarely present, cervical sulcus variable, sometimes reaching the dorsum, sometimes weak and restricted to the lateral surface of the carapace; postcervical sulcus sometimes present. Abdominal somites 4-6 always carinate, somite 3 sometimes carinate; telson apically acute with 3 or 4 pairs of movable lateral spines. Optic peduncle with a mesial tubercle, optic scale absent. Prosartema reduced to a setose boss; antennular flagella unequal, the dorsal flagellum very short and flattened and inserted proximally into the third segment; ventral flagellum long and sexually dimorphic in some genera; maxillulary palp unsegmented. Exopods on all maxillipeds, present or absent on pereopods; pereopods 4 & 5 usually more slender and longer than pereopod 3. Pleopods well developed, sometimes longer than the carapace. Petasma simple, open; second male pleopod with appendix masculina consisting of inner and outer projections (sometimes referred to as appendix Epipods | es ! Somites 3-8 Somite | | Somite2_—_|_—_— Somites 3-7 | Somites 1-7 _ Aristeomorpha | — +(3s)* a 8,1 - + + (also 6) _ et Aristaeopsis i ee | ee Sy See ee a + (also 6) oF | Aristeus _+ (sort, 3-7) | I | sd | | ee 1-6 only A ustropenaeus _ _ + | 7 at | Ss 1 - | ai + {also ron 6)- + 7 |Hemipenaeus | 7 PF J he 7 is & + | +(ron7) | Hepomadus | 4 ] | 2a + | ad +(ron7) Parahepomadus = + 1 + + + (son 7) Plesiopenaeus | - ra =e s | + + (also 6) +(son7) Pseudaristeus | + r | + + 1-6 only AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE interna and appendix masculina, respectively) and no distolateral projection. Thelycum open, seminal receptacle deep, thoracic somite 7 with a shield-like median plate. The arrangement of branchiae and epipods of the geneta of the Aristeidae are listed in Table 1, REMARKS, Pérez Farfante (1987) concluded that the relative length of the rostrum in some genera is size dependent as well as being sexually dimorphic. The prevailing view is that this is true sexual dimorphism and not due to an earlier breakage of this structure, which is often more slender in d 3, followed by regeneration ofanew lip. (Some adult 2 2 also show evidence of such a regeneration). The family as now detined, 1s very homo- geneous and consists of 9 genera, all of which have been recorded from Australian seas. Aristeomorpha stands a little apart from the other genera: with 6-11 fairly evenly spaced dorsal rostral tecth: basicerite with a prominent spine (like that in other penaeoid families); petasmal halves diverging distally and ventral costa attached along its length; and thoracic 8 sternite of the thelycum with a short rounded median prominence. The remainder are similar in uppearance and some are difficult to distinguish, sometimes relying on rather minor features of the vephalothorax or abdomen, For example, the rudimentary exopods (Fig. 1) which separate 2 groups of 3 genera each are often difficult to detect, Unfortunately, better criteria do not seem fo exist. Also, the trend in taxonomy of the Aristeidae has been to create small genera. Thus of the 9 genera 4 are monospecific, 2 contain 2 species and | includes 3. Arevision of the family is perhaps needed at this stage and certainly before any new genera are created. KEY TO GENERA OF THE ARISTEIDAE }. Tlepaticspine present, Hepatic spine absent . 2. Three dorsal rostral teeth. eluding “the nastnastral (epivastric}touth. . Sa ere . Flepamiactis More than three dursal rostral teeth, ‘eluding the postrostral taglh. 2... a Ce Jristaeontor ple 3. Postuntennal spine present — - tae et Paruhepamadius Posiuntennalspineabsent, -.- 5... -.--..---4 4. Exopods absent from all pereopods 2... . $ Lxopods (ustally reduced ar rudimentary. br ie I prsen onall pereapods- bis ew 3. A sub-distal nreral spine present on pereod ) #2 a seaphncerite in male nop modified, PR iin soobealh No meral spines on pereopads | & 2; scanner in male dislally clongate Iris fermrpeis 41) 6 Cervical suletis distinet and extending to the dorsum of the curapace: posiver ical salcus present... Preudursylens Cervien) sulcus reduced to a very shor ni in the hepaticresion:postcery ical sulcusabsent . Aristens Abdeminal somite J varinate: ipa on pereopud 4 ridimentary 2 ee, . Hemipenceris Abdominal somite 3 not carinutes évipol on purer 4 urge... . oo KO) 8 &. Podobranchia on pdieepdd, 3 rulizpentetty carimae the carapce Weak; ventral antennulur Hayellanyand duewy! af the third naxilliped madified in the male | stesomm | tm edehean | + ASMP O ICTS Podobranchia on pereopod 3 large; curinae on the cumupace well defined: ventral antennular flagellum and dacty! ol the third baseline nol mudified in the mule SG eee ee _ Plestapenueus Aristaecomorpha Wood-Mason, 1891 Penaeus Risso, 1827269 [part |; HL. Walne-Kewards. {8372 418 [part]. Afristeis Bate: (8812 189: TBRR: 317 [part]. fristasumerpha Wood-Nluson, ba¥l: 286: Anderson & Lindner, 1943: 301; Kuho, 1949; 200; Crosnier, 197%: 52) Hayashi, l98ab: 28Us de Freitas, P9832 135; Liu d& Zhome, L986: 33. Perex lartante & Kenstey. 1997: 33, Jriswomerpha Senna, 1903: 265; Bouvier, Ramadan, (948: 53, 1ohsu: 52; DIAGNOSIS. Integument finely pubescent, with a pair of large ventr ral photophores on the thoracic and abdominal somites with 3 pattern of smaller photophores on the ventral surface, the scapho- cerites, external uropods and most of the other appendages. Rostrum reaching well beyond the scaphocente in females and juveniles, usually shorter than the antennular peduncle in adult 3d @: with with 5-9 dorsal teeth in all; adrostral carina reaching between the first and second rostral teeth; postrostral carina low and reaching about half the carapace. Antennal and branchioslegal spines present, the latter on the margin of the carapace with a carina extending almost to the hepatic spine: hepatic spine large, eee OM FIG. 1. Rudimentary exopod on basis of pereopad 3 of Austropenceus nities (de Man, 1911). AM P53936. 34°49'S 1SV14VE. 1225 m. 2. 33 mm. B, basis: C, coxa; Ep, epipod: Ex. rudimentary exopod, 15, ischium; CM, margin ofcarapace. (Scalebar= linm). cervical sulcus reaching about halfway to the dorsum. A deep sulcus extends from a shallow emargination below the branchiostegal spine to its junction with the hepatic sulcus, which turns ventrad at the mid-carapace; hepatic carina short and blunt; branchiocardiac carina prominent, the sulcus shallow, the carina extending from the divergence of the hepatic sulcus almost to the posterior margin of the carapace. A low carina, sometimes barely visible, on the posterior two thirds of abdominal somite 3, 4-6 fully carinate, all 4 somites ending in a sharp spine. Telson with 4 pairs of small lateral movable spines. Eye large; stylocerite reaching about 0.75 the first segment of the antennular peduncle. Dorsal antennular flagellum flattened with terete tip, slightly longer than the third antennular segment, ventral flagellum long, not modified in the male. Lateral spine of the scaphocerite at about 3/4 the length of the lamella; scaphocerite not modified in the male. Pereopods without exopods or ischial and meral spines; pereopod 3 with a large podobranch. Petasma simple with diverging apices; thelycum with a deep hemicylindrical receptacle, formed by a shield-like anterior plate on thoracic somite 7, the rounded apex directed anteroventrally, the coxae of the pereopod 5 anda rounded median boss on somite 8. REMARKS. Armature of the rostrum sets Aristaeomorpha apart within the family. The 2 species, 4. foliacea (Risso, 1827) and A. woodmasoni Calman, 1925, are distinguished by the relative depths of the pterygostomian area, the length/depth ratios being 3.5-4.0 and <2.5, respectively. Other features are minor (relative lengths of the uropods and dactyls of the fourth and fifth pereopods). Barnard (1950) pointed out that Kemp & Sewell (1912) could find no differences in the petasmas and thelyca of the 2 species. A. woodmasoni has so far been recorded only from Indian waters and its specific status needs to be re-examined as A. foliacea has been recorded from the Maldive Islands, Sri Lanka, Indonesia and surrounding seas. While Aristeus and possibly other members of this family have photophores, at least on the appendages, A. foliacea appears to be the most luminescent (Crosnier, 1978: 57, fig 23e-f, for MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Aristaeomorpha foliacea (Risso, 1827) QMW10091, 26°20°S, 153° 53°E, 300m, 2, 42mm. full description). It is widely distributed (Crosnier, 1978 listed 99 references in his ‘Bib- liographie partielle’) and occurs in deeper-water prawn fisheries, including SE Australia. Aristaeomorpha foliacea (Risso, 1827) (Fig. 2) Penaeus foliacea Risso, 1827: 69, pl. 2, Milne-Edwards, 1837: 418; Miers, 1878: 307, Aristeus rostridentatus Bate, 1881: 189; 1888: 317, pl. ST. Aristaeomorpha Giglioliana Wood-Mason, 1892: pl. 2, fig 2. Aristaeomorpha foliacea (see Crosnier, 1978 for bibliography prior to 1976); Crosnier, 1978: 54, fig. 23; 1984: 21; 1985: 861; 1989: 42; 1994b: 369; Hayashi, 1983b: 280, fig. 53; Grey et al., 1983: 46, pl. 1; de Freitas, 1985: 16, fig. I-7; Liu & Zhong, 1986: 33, figs 12, 13; Kensley et al., 1987: 279; Hanamura, 1989; 51. MATERIAL. QM W10091, 26°20’S 154°E, 300m, 3, 43mm, 22, 42, 43mm;QMW11428, 23°46’S_ 153°E, 550m, 3, 40mm, 3°, 44, 44, 46mm; QMW14351, 26°30’°S 153°45°E, 390m, 3d, 43mm, °, 40 mm; QMW 15292, 28°S 153°54’E, 550m, 2 ¢, 28, 35mm, 2, 42mm. DIAGNOSIS. Apart from the differences in pterygostomian depths of the 2 species noted above, the genus diagnosis is also the specific definition. Colour. Uniformly deep pink to wine red (colour plate in Grey et al., 1983). DISTRIBUTION. E Australia and Tasmania, 18°-42°42’S, the western approaches to Bass Strait, Great Australian Bight, Arafura Sea, 8°34’S 131°E, through the Northwest Shelf to 28°S on the west coast; 250-700 m on mud to muddy sand. Thus it is likely to occur at these depths all round Australia. Known range: cosmopolitan, Indian Ocean from SW and E Africa, Madagascar, Réunion, Maldive Islands, Sri Lanka, Indonesia, Philippines, China Sea, Japan, Australia, New Zealand, New Caledonia, Wallis and Futima Islands, Fiji, W Atlantic Ocean, from Massachusetts to Venezuela, Mediterranean Sea, E Atlantic Ocean from Bay of Biscay to Rio de Oro. Depth range 170-810m, fig. 6: AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE although exceptionally caught up to 61m. Crosnier (1978) suggested that there may be a diurnal migration from deep to shallower water at night. Aristaeopsis Wood-Mason, 1891 Aristaeopsis Wood-Mason, 1891: 282; Bouvier, 1908: 61; Burkenroad, 1936: 94 [part]; Crosnier, 1978: 86; Pérez Farfante & Kensley, 1997: 36. Plesiopeneus Faxon, 1895: 199 [part]; Milne-Edwards & Bouvier, 1909: 200 [part]; Burkenroad, 1936; 94 [part]; Ramadan, 1938: 49; Barnard, 1950: 621 [part]; Crosnier, 1978: 85 [part]; Squires, 1990: 41. (non Plesiopenaeus Bate, 1881). Aristaeus (Plesiopenaeus) Alcock, 1901; 35 [part]. Aristeopsis de Man, 1911: 6. DIAGNOSIS. Integument glabrous, carapace firm. Rostrum with unarmed tip upcurved, as long as the carapace in 22 of 30mm CL, decreasing to around 0.55 of the carapace at 62mm CL, not sexually dimorphic although often damaged in d d ; with 3 dorsal rostral teeth; adrostral carina not quite reaching the first rostral tooth; postrostral carina low, but reaching about (0).8-0.9 the carapace. Antennal spine with a well- defined carina; hepatic spine absent; branchio- stegal spine on the margin of the carapace and continuous with a prominent carina running almost to the branchiocardiac carina; hepatic sulcus deep, occupying three-quarters the posterior part of the carina and descending to the submarginal carina. Gastro-orbital carina prominent, extending from just behind the orbit almost to the upper end of the prominent cervical carina; cervical sulcus deep in the lower part, but barely defined in the upper half, where it sometimes meets the dorsum; branchiocardiac carina and sulcus prominent and reaching the posterior margin of the carapace; two irregular carinae running from the cervical carina to the branchiocardiac sulcus. A dorsal carina beginning on the posterior quarter of abdominal somite 2 and continuing to somite 6, abdominal somites 3-6 each with a sharp carina and each ending in a prominent spine; telson with four pairs of small movable spines. Dorsal antennular flagellum short, flattened; ventral flagellum long, tapering and not modified in the male; scaphocerite in the adult male with an elongate distal projection, ovate in cross section, and sub-equal in length to the lamellar part. Pereopods without exopods, pereopod 3 with podobranchia; 413 pereopods | & 2 without distal movable ischial and meral spines. Petasma with dorsal lobule and median lobes of similar length, the ventral costa forming a projecting median hook. Thelycum with an acute anteriorly pointed prominence on thoracic somite 7, the posterior part forming a rounded hollow; eighth somite with a bluntly pointed median prominence.Uniformly scarlet to deep crimson REMARKS. 4ristaeopsis was originally created for Penaeus edwardsiansus by Wood-Mason & Alcock (18916), but Faxon (1895) synonymised it with Plesiopenaeus Bate, 1881 and this usage has persisted for most of the 20th century. Burk- enroad (1936) recognised 4 species of Plesio- penaeus (P. edwardsianus, P. armatus, P. cor- uscans and P. nitidus). Although accepting this classification, Crosnier (1978, table 11) listed a number of differences between P. edwardsianus and P. nitidus and the other 2 species. For P. edwardsianus the most important of these are lack of exopods on all pereopods; a dorsal carina and posterior spine on abdominal somite 3; the modified scaphocerite in ¢; absence of movable distal meral spines on pereopods 1 & 2. Pérez Farfante & Kensley (1997) considered these differences merited the resurrection of Arist- aeopsis (feminine gender) which thus contains only A. edwardsiana. Plesiopenaeus edwards- ianus has been extensively cited in the literature (Crosnier, 1978 listed 50 references) and because of long familiarity the general recognition of A. edwardsiana will no doubt take some time. Aristaeopsis edwardsiana (Johnson, 1867) (Fig. 3) Penaeus Edwardsianus Johnson, 1867: 897. Aristeus Edwardsianus Miers, 1878: 308, pl. 17, fig. 3. Aristeus coralinus Bate, 1888: 32, fig 10. Aristaeopsis Edwardsiana Wood-Mason & Alcock, 1891b: 283, figs 8-9; Wood-Mason, 1892: pl.1. figs 1,2: Alcock 1899. Aristaeopsis edwardsiana Alcock & Anderson 1894: 147: Pérez Farfante & Kensley 1997:37, fig. 7. FIG. 3. Aristaeopsis edwardsiana (Johnson, 1867) AMP26776, 33°40°S, 151° 56’E, 732m, 2, 45mm. 414 Plesiopenaeus edwardsianus Faxon 1895 (for full synonomy to 1974 see Crosnier, 1978: 88 ); Crosnier, 1978: 88, figs 3la-c, 32a-c, 33a; Kensley et al. 1987: 281; Hayashi 1983c: 368, fig. 59; Liu and Zhong 1986: 43, fig. 17; Crosnier 1985: 863; 1994b: 369; de Freitas 1985: 20, fig. 2-9; Grey et al. 1983; 38, pl. 2. MATERIAL. AM P26776, 33°40’S 152°E, 732m, 2d, 42, 53mm, 10°, 30-63mm, QMW 11307, 24°S 153°E, 550m, 22, 38, 77; QMW11461, 23°34’S 153°E, 650m, 3 48mm, 22 64, 82mm; QMW 15286, 28°S 154°E, 550m, 33, 37, 41, 42mm, &, 57 mm; QMW 15287, 28°S 154°E, 550m, 3d, 42, 44, 46mm, 22, 40, 50mm: W15291, 28°S 154°E, 550m, 6, 38mm. DIAGNOSIS. As for genus. REMARKS. A. edwardsiana occurs regularly in deep water prawn trawls off the Australian east coast, but at best is only of minor commercial importance. Crosnier (1978) reported that, off Madagascar, at the 700-800 depths preferred by the species, catches were barely 6-8 kg/h. DISTRIBUTION. East coast of Australia, 17°-34°S, Arafura Sea 8°S 132°E, off Rowley Shoals, through the Northwest Shelf to 29°S 113°E; 200-1800 m on muddy substrates. Cosmopolitan, Indo-West Pacific Ocean from South Africa to the Arabian Sea, Bay of Bengal, Andaman Sea, Indonesia, South China Sea, Japan, Australia, Wallis and Futuma Islands; Western Atlantic Ocean from the Grand Banks to French Guiana, Eastern Atlantic Ocean from Portugal to the western Sahara, the Azores, Madeira and Canary Islands, 200-1850m. Aristeus Duvernoy, 1840 Penaeus Risso, 1816: 96. Aristeus Duyernoy, 1840; 217; Bate, 1881: 171, 187; 1888: 228, 240, 309; Senna, 1903: 261; Bouvier, 1908: 69; de Man, 1911: 27; Ramadan, 1938: 36; Crosnier, 1978: 60; Hayashi 1983a: 188; Liu & Zhong, 1986: 37; de Freitas, 1985: 3; Pérez Farfante & Kensley, 1997: 39. Aristaeus Faxon, 1895: 197; Alcock, 1901: 27; Anderson & Lindner, 1943: 300; Kubo, 1949: 194. Aristaeus (Aristaeus) Alcock, 1901: 29. DIAGNOSIS. Integument glabrous or pubescent. Rostrum elongate, reaching well beyond the tip of the antennular peduncle in 2 2 and young 33, but usually shorter in adult ¢d; with 3 dorsal teeth. Carapace with an antennal and a MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4, A, Aristeus mabahissae Ramadan, 1938. AMP55934, 34°53’ S, 151° 15°E, 1116m, 2,37mm. B, A. virilis (Bate, 1881). NTCR006630, 8°39’S, 132°E, 540m, 2, 39mm, anterior carapace showing long hepatic carina. branchiostegal spine, the latter extending back as a carina, other spines absent; cervical sulcus marked only by a short lateral sulcus; branchiocardiac sulcus well-defined, the carina low. Only abdominal somites 4-6 with a distinct dorsal carina; telson with 4 pairs of movable lateral spines, apex acute. Cornea well developed, the peduncle with a prominent tubercle. Dorsal antennular flagellum short, flattened; ventral flagellum long, proximally sinuous in d. Pereopods without exopods; pereopods 1 & 2, and sometimes 3 with a movable subdistal meral spine; pereopods 4 & 5 slender; all pereopods with photophores. Of 8 species known worldwide, 6 occur in the Indo-West Pacific. A. antennatus is also an Atlantic species together with A. antillensis and A. varidens. So far A. mabahissae and A. virilis have been collected only from Australian waters; A. semidentatus is a possible third; A. antennatus is listed in the Australian Museum collection, but could not be found at the time of writing and is not described below. Among the Indo-West Pacific species A. virilis is readily identifiable because of its finely tomentose integument and pereopods, the spine on the merus of pereopod 3 and a long carina extending posteriorly from the branchiostegal spine. The remaining glabrous species are difficult to distinguish, almost impossible unless the material is in a good state of preservation. Unusually for penaeoids, the genitalia are of limited use for identification; Crosnier (1978) notes that the colouration is identical for A. anfennatus, A. mabahissae and A. AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE 415 TABLE 2. oe distinguishing features for identification of Aristeus alcocki, A. antennatus, A. mabahissae, A. pallicauda, A . semidentatus. * P l= first pereopod. oe Character _A 4. ale ocki _A.antennatus | A mabahissue A. pallicauda | A. semidentatus | iL Length of *P 1 chela/length of carpus _ 115-125 0) 13-148 | 1.07-1,25 14 10 ~1.06 | ih ength of P 5 carpus/length of merus _ 1.08 — 1.18 100-1.14 | 0.98— 1.06 124 | 1.28-—1.34 Number of photophores on the carpus of P 5 a al 65-117 alt 4B 36 : 14-21 | | Number of photophores on the propodus of P 5 | 2 al 73 —98 | 8-13 31 | 7-25 . 7 , | she third bdominal somite EY | OF Rome” | “amelispine | | Absent Absent semidentatus; the carapace is almost featureless, except for the carination behind the branchio- stegal spine. Ramadan (1938) recommends using the ratios between various segments of the pereopods, but often these are not sufficiently different for reliable identification. Crosnier (1978) found that the number of photophores on pereopod 5 could be used as specific characters, at least in a given area; more recently, he has found that they are reliable for most of the Indo- West Pacific (Crosnier, 1994b, & pers. comm.). The red colour of the photophores fades with preservation, but in recently- and well-preserved specimens they may be seen by oblique light as a series of small circular structures. Unfortunately, the photophores are difficult, if not impossible to see in old and poorly preserved material, even if the slender pereopods 5 are present, which is often not the case. Crosnier (1978) recommended that a number of characters be taken in conjunction to distinguish species and the more useful are shown in Table 2. A key is also given, but unless the prawns are fresh, is mostly of value in preliminary identification. KEY TO THE INDO-WEST PACIFIC SPECIES OF ARISTEUS Body and pereopods finely pubescent. A subdistal mobile spine near the inferior border of the merus of pereopod 3 (P3) A. virilis Body glabrous. No mobile spine on the merusofP3 . , 2 Ratio length of chela/length of carpus of P | approx. 1.0 ae Sere es ok. 4, semidentatus Ratio length of chela/length of carpus of P | greater than Bs re rE, Sram eee Rt te ee 3 A small posterior-dorsal spine on abdominal somite 3; number of photophores on carpus and propodus of P 5 greaterthan60.. 2.2... ee es 4, anlennatus Abdominal somite 3 usually without a spine; number of photophores on carpus and propodus of P 5 less than 60. 4 Ratio length of chela/length of carpus of P | greater than 1,3; photophores on carpus and merus of P 5 greater than os pha ie V8 dei aes] HE oe he 2 A, pallicauda Ratio length of chela/length of carpus of P | less than 1.3; photophores carpus and merus of P 5 less than 20 be ee ee af le dé ee sae A, mabahissae Ne It was not possible to include 4. alcocki Ramadan, 1938 in the key because the number of photophores on P 5 is not recorded in the literature. So far it appears to be limited to the Gulf of Aden, the Arabian Sea and the Bay of Bengal and thus may not occur in Australian seas. Similarly A. pallicauda Komai, 1993 is a rare species recorded only from northern Japan. Although A. semidentatus has been recorded from Australian seas (Kensley et al., 1987), | was unable to confirm this with relevant specimens in the Australian Museum. Some were undoubted A, mabahissae, others were probably this species and the remainder were unidentifiable as A. semidentatus, these 2 species being similar. The criteria listed by Kensley et al. (1987) are in fact identical or very close to those of 4. mabahissae (Crosnier, 1978) except forthe ratio of the lengths of the carpus and merus of pereopod 5. While it is likely that A. semidentatus is present in Australian seas, until bona fide specimens are collected, it is best to regard existing records as doubtful. Aristeus mabahissae Ramadan, 1938 (Fig. 4A) Aristeus mabahissae Ramadan, 1938: 43, figs 2b, 3b, 4a-c; Crosnier, 1978: 65, figs 25e-f, 26c-f; 1984: 22; 1994a: 352, 1994b: 369; Hayashi 1983a: 190, figs 49, 50; Komai, 1993: 22. MATERIAL. AMP39948, 33°40’S 152°E, 1115m, 9, 38 mm; AMP39977, 33°S 151°E, 1097m, 2, 42mm; AMP 55934, 34°53’S_ 151°E, 1116m, 49, 37-44mm; AMP 55938, 35°S 1S1°E, 1015m, 3, 20mm, 2 2, 39, 40mm; NT Cr 007070, 29°S 114°E, 880m, 3 2, 39, 43, 53mm; NT Cr 007086, 13°S 122°F, 900m, 24, 28, 33mm, 22, 34, 41mm; CSIRO, SS/1/91 #44, 27°07’S 112°22’E, 714m, 22, 42, 46mm. DIAGNOSIS. Carapace glabrous. Rostrum in 2 & and juvenile 3 variable in length, usually well exceeding the antennular peduncle, the unarmed part slender and upturned, sometimes strongly; often shorter in adult ¢ ¢; in both sexes with 3 dorsal teeth, the first smaller than the 416 second and third; adrostral carina ending at the first rostral tooth; postrostral carina reaching about half the carapace. A low gastro-orbital carina above a short, shallow orbito-antennal sulcus; antennal spine extending back as a short antennal carina. A prominent branchiostegal spine on the border of the carapace, extending posteriorly as a sharp carina, which reaches the base of the cervical sulcus and thereafter continues as a blunt prominence, almost meeting the low branchiocardiac carina; hepatic sulcus starting below the cervical sulcus and joining the deep branchiocardiac sulcus, which almost reaches the posterior margin of the carapace. Cervical sulcus restricted to the lateral area of the carapace. A subdistal meral spine on pereopods | & 2; pereopod 5 with 4-10 photophores on the carpus and 8-13 on the propodus. Colour. Variable, the body slightly whitish, with the branchial area, posterior part of the abdominal somites, the extremities of the rostrum and appendages generally rose to rose-orange. REMARKS. A. mabahissae appears to be fairly common in Australian seas. It is easily confused with A. semidentatus, but this species may be distinguished by the number of photophores on pereopod 5 (carpus 13-33 and propodus 20-31). DISTRIBUTION. Eastern Australia 28°-34°S, Western Australia 13°-29°S; 500-1100m. Known range: Madagascar, Maldive Islands, Indonesia, South China Sea, Japan, Australia, Wallis and Futuma Islands, 500-1100m. Aristeus virilis (Bate, 1881) (Fig. 4B) Hemipenaeus virilis Bate, 1881: 187; 1888; 303, pl. 44, fig. 4. Aristeus tomentosus Bate, 1881: 189; 1888: 307, pl. 49, figs 2,3, pl.S0. Aristaeus virilis Faxon, 1895: 198; Alcock, 1901; 30; Kubo, 1949; 194, figs 1, 6, 8, 11, 13, 14, 19, 23, 36, 44, 65, 69, 72, 78, 85, 86. Aristeus virilis Bouvier, 1908: 70; de Man, 1911: 6, 27; 1913: pl. 2, fig. 6; Ramadan, 1938: 39; Okada et al., 1966: 140, 141, 151, pl. 1, fig. 3; Crosnier, 1978: 61, figs 25a-b, 26a-b; 1984; 21; 1985: 861: 1989: 42; 1994a: 352; Hayashi 1983a: 190, figs 51, 52c.d: de Freitas: 3, fig, II-1, 1985; Liu & Zhong, 1986: 37, fig. 14: Kensley et al., 1987: 281. MATERIAL. AM P21683, 29°46’S 154°E, 500m, 32, 30, 38, 38mm, 42, 34-57mm; QMW11282, 23°S 154°E, 460m, 3,39mm, 2, 55mm; QMW11429, 23°34’S 154°E, 650m, 2, 48mm; QMW 14296, 23°40’S 154°F, 530m, 2, 38mm; QMW 18059, due E Brisbane 28°S, 700-900m, 32, 43, 57, 57mm; QMW20793, 18°S 118°E, 250-390m, 2, 53mm; NT Cr006630, 8°38’S 132°E, 22, 39, 40mm; CSIRO, SS/1/91 #43, 27°S 112°E, 2, 53mm. MEMOIRS OF THE QUEENSLAND MUSEUM DIAGNOSIS. Body, pereopods and other appendages with a fine pubescence. Adrostral carina ending just behind the first rostral tooth; postrostral carina low but reaching three-quarters the length of the carapace. A low gastro-orbital carina above a short, but distinct orbito-antennal sulcus. Branchiostegal spine on the border of the carapace and continuous with a prominent carina, which almost reaches the branchiocardiac carina; hepatic sulcus beside the posterior half of the branchiostegal carina. Branchiocardiac sulcus deep and almost reaching the posterior border of the carapace. A sub-distal, ventrolateral movable spine on the merus of pereopods 1-3. Carpus of pereopod 5 with 15-25 photophores, propodus with 16-22. Colour. Red-orange, deeper on the margin of the carapace and rostrum, the posterior edges of the abdominal somites, the uropods and telson; pereopods lighter in colour. REMARKS. This appears to be the commonest of the Australian species of Aristeuws and is readily identifiable by the key characters. DISTRIBUTION. Eastern Australia 10°-34°S, Arafura Sea 8°S 132°E, through Northwest Shelf to 29°S; 250-1050m. Known range: South Africa, Madagascar, Andaman Islands, Indonesia, South China, Philippines, Japan, Australia, New Caledonia, New Hebridies, Wallis and Futuma Islands, 250-1050m. Austropenaeus Pérez Farfante & Kensley, 1997 Austropenaeus Pérez. Farfante & Kensley, 1997: 4, figs 11, 12 DIAGNOSIS. Integument glabrous. Rostrum slender and upcurved, approx. |.3-1.4 the length of the carapace in adult ° 2, slightly shorter, but still exceeding the length of the carapace in adult 3 4; with 3 dorsal teeth; adrostral carina short, starting at the base of the third tooth and ending behind the first tooth; postrostral carina reaching about one third of the carapace. Carapace with an antennal and a prominent branchiostegal spine on. the margin of the carapace, with a short carina; hepatic sulcus wide and almost reaching the branchiocardiac sulcus; the latter barely defined, but almost reaching the posterior margin of the carapace; cervical sulcus short and indistinct; carapace otherwise almost featureless. Abdominal somite 3 without a distinct carina, but with a postero-dorsal spine; somites 4-6 each dorsally carinate and ending in a spine; telson acute with 4 pairs of lateral movable spines. AUSTRALIAN ARISTEIDAE AND BENTHESICY MIDAE FIG. 5. A, Austropenaeus nitidus (Barnard, 1947) AMP55939, 33°51’S, 152°15°E, 1200m, 2, 26mm. B, AMP55395, 34°53’S, 151°1S’E, 1116m, 3, 21mm, antennular flagella, lateral view. Cornea slightly flattened, peduncle with a prominent mesial tubercle. Dorsal antennular flagellum flattened and reaching as far as the spine of the scaphocerite in females, slightly longer in males; ventral flagellum slender, terete and in the male with a proximal enlargement (Fig. 5B); scaphocerite not modified in the adult male, but with the dactyl of the third maxilliped thicker basally, with a finger-like distal part and shorter than in the female. Rudimentary exopods on all pereopods (Fig. 1), pereopods 1-3 slender and each with a sub-distal movable spine on the merus; the fourth and fifth more slender than the first to third. Dorsolateral lobule of petasma produced as a rounded process, well exceeding the ventromedian lobule; distal ventrolateral lobule semicircular and rounded; thelycum with apically pointed shield-shaped sternal plate on thoracic sternite 7, sternite 8 with low median prominence and paired anterolateral ridges. REMARKS. This genus contains only A. nifidus, formerly included in Plesiopenaeus. The key characters enable Austropenaeus to be easily separated from Plesiopenaeus, notably by the almost featureless carapace. However, the rudimentary exopods on the pereopods, which separate it from other genera, are easily missed. Austropenaeus nitidus (Barnard, 1947) (Figs 1, 5A,B) Plesiopendeus nitidus Barnard, 1947: 383, 1950: 622; Crosnier, 1978: 89; Kikuchi & Nemoto, 1986: 52. MATERIAL, AMP39951, 33°40’S 152°E, 1115m, &, 30mm; AMP 39979, 33°S 153°E, 1095m, 23, 21, 24mm, 5 2, 22-24mm; AM K24-20-03, E Sydney, 969-1006m, 52, 16-26mm; AMP55392, 38°S 150°21E, 960m, °, 26mm; AMP 55395, 35°S 151°E, 11l6m, 24, 21mm, 24mm, °, 26mm; AMP55396, 34°50’°S 151°E, 1225m, 417 42, 25-33 mm; AMP55939, 33°51°S, 152°15’E, 1200m, 2, 26mm; SAM (unregistered, Great Australian Bight approx. 34°S, 128 to 132°E, 927-1249), 52, 30-37.5mm; CSIRO, SS/1/91 # 33, #47, #61, 29°35’°S 111-113°E, 1101-1277m, d 25 mm, 6%, 25-31mm. DIAGNOSIS. As genus. REMARKS. Kikuchi & Nemoto (1986) recorded 1 2A. nitidus (as Plesio- penaeus nitidus) from the Northwest Pacific. [t was collected around 22°N in a 0-5700m oblique tow, whereas all Southern Hemisphere A, nitidus have been collected in bottom trawls S of 26°S, nearly all between 1000-1300m. It is moderately common in the southern part of Australia, but to date it has not been found anywhere between 22°N and 26°S. Thus the identity of the Northwest Pacific species needs to be confirmed. DISTRIBUTION. Australia S of 26°S, WA, through the Great Australian Bight and app- roaches to Bass Strait (approximately 40° S), to 27°S on the NSW coast, usual depth range 1000-1300m, where it appears to be moderately common (one recorded depth of 457m, may be an error). Known range: with the possible exception noted above A. nitidus appears to be an inhabitant of latitudes higher than 26° S: south Atlantic Ocean; off Cape of Good Hope and Natal, South Africa; Amsterdam and St. Paul Islands, south Indian Ocean; southern Australia, 457-1300m. for Hemipenaeus Bate, 1881 Hemipenaeus Bate, 1881: 171, 186; 1888: 299; de Man, 1911: 23; Burkenroad, 1936: 90; Ramadan, 1938: 47; Anderson & Lindner, 1943; 300; Roberts & Pequegnat, 1970: 43; Crosmer, 1978: 74; Hayashi, 1983c: 366; Pérez. Farfante & Kensley, 1997: 43, Hemipeneus Faxon, 1985: 198, Aristeus (Hemipeneus) Alcock, 1901: 31 [part]. Hemypenaeus Kikuchi & Nemoto, 1986: 52. DIAGNOSIS. Integument glabrous. Rostrum usually short in both sexes, occasionally of moderate length in 2 2; with 3 dorsal teeth; a short adrostral carina present; postrostral carina pronounced, reaching at least to the midlength of the carapace. Antennal and branchiostegal spines present, the latter on the margin of the carapace and with a prominent carina; cervical and 418 postcervical sulci present or absent; branchiocardiac carina and sulcus well-defined. Abdominal somite 3 with a prominent downcurved spine; somites 4-6 each with a dorsal carina; telson with 4 pairs of small movable spines in the posterior half, apex acute. Cornea moderately small, dorsoventrally flattened; ventral antennular flagellum not modified in the male. All pereopods with rudimentary exopods (Fig. 1); no movable meral spines on the per- eopods. Petasma with distal margin of the dorsolateral lobule oblique and about the same length as the dorsomedian lobule; distal half of ventral costa free, the tip flattened with laterally directed projection. Thelycum with broad, anteriorly pointed plate on thoracic somite 7, somite 8 short and broad, roughly rectangular. REMARKS. The genus contains only H. carpenteri and H. spinidorsalis. Both are deep water inhabitants, usually below 2000 m, which probably explains their apparent rarity. H. carpenteri 1s easily distinguished by its cervical and postcervical sulci (although these may be quite faint dorsally), which are totally lacking in H. spinidorsalis. Only H. carpenteri has been collected in Australian waters, although both species have a similar range. Hemipenaeus carpenteri Wood-Mason, 1891 (Fig. 6) Hemipenaeus Carpenteri Wood-Mason, 1891: 189: Wood-Mason & Alcock, 1891 b: 286; de Man, 1911: 6, 23. Hemipeneus triton Faxon, 1893: 215; 1895: 202, pl, 50, 1896: 163. Aristeus (Hemipeneus) carpenteri Alcock & McArdle, 1901: pl. 49, fig. 4. Aristeus (Hemipeneus) Carpenteri Alcock, 1901; 32. Hemipenaeus carpenteri Burkenroad, 1936: 91; Ramadan, 1938: 48; Anderson & Lindner, 1945: 301; Roberts & Pequegnat, 1970: 43; Pequegnat & Roberts, 1971: 8, pl. 5D; Crosnier, 1978: 76, figs 27 c-d, 28 a-b, 29a, 1985; 862, 1994b: 369; Hayashi 1983c: 366, fig. 57. MATERIAL. QMW13261, 16°54’S 147°E, 1473-1590m, 2, 8mm; QMW13451, 17°S 148°E, 1500m, 2, 35mm; TMH G4044, 21°S 113°E, 1139-1128m, 2,20 mm; G4045, 23°S 113°E, 1460-1700, 23, 24, MEMOIRS OF THE QUEENSLAND MUSEUM and second rostral teeth; postrostral carina prominent, ending at 0.7-0.8 the length of the carapace. Gastro-orbital carina present, almost reaching the cervical sulcus, Antennal and branchiostegal spines present, the latter on the margin of the carapace and with a prominent carina reaching below the lower extremity of the cervical sulcus. A shallow hepatic sulcus, continuous with the deep branchiocardiac sulcus, which almost reaches the posterior margin of the carapace. Lower half of the cervical sulcus well-defined, with a very short carina at its lower end; upper part faint, but reaching the mid-dorsum; a faint postcervical sulcus present; submarginal carina sharp. Abdominal somite 3 with a large down-curved postero-dorsal spine; a dorsal carina on each of abdominal somites 4-6, that on the fourth weak. Ventral costa of the petasma free distally, the apex markedly flattened, with a laterally directed point. Thelycal plate on thoracic somite 7 with a long tapering anterior point. Colour, Carapace deep blue, abdomen lighter blue anteriorly, changing to pinkish towards the telson; appendages pink to red. REMARKS. The 20mm & (Fig.6) appeared to be close to sexual maturity, the thelycum being similar to that figured by Crosnier (1978, fig. 28a, b). The petasma of a mature 3 does not appear to have been figured, but although one 24mm ¢ had well-developed petasmal halves they were not joined medially and it was therefore immature. Consequently, it has not been figured. The other 3, although of identical size, was decidedly immature. DISTRIBUTION. Western Australia, 20-22°S 113°E, 1100-1700m; NE Australia, 16-17°S 147°E; 1473-1590m. Known range: throughout the Indian Ocean, Indonesia, Japan, Australia, Wallis and Futuma Islands; Gulf of Panama and Galapagos Islands; W Atlantic, 900-3900m. 24mm. DIAGNOSIS. Rostrum short in both sexes, not exceeding the tip of the cornea in adults, slightly longer in the 8mm juvenile, with 3 dorsal teeth; a short adrostral carina present, ending between the first FIG. 6. Hemipenaeus carpenteri Wood Mason, 1891 TMHG4044, 20°55’S, 112°51°E, 1139m, 2, 20mm. AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE Hepomadus Bate, 1881 Hepomadus Bate, 1881: 171, 189, 188: 319; Bouvier, 1908: 56; Milne Edwards & Bouvier, 1909: 194; Burkenroad, 1936: 86; Ramadan, 1938: 55; Pérez Farfante, 1973: 441; Crosnier, 1978: 47; Liu & Zhong, 1986: 28; Pérez Farfante & Kensley, 1997: 46. Aristeus (Hepomadus) Alcock, 1901: 42. DIAGNOSIS. Cuticle glabrous. Rostrum exceeding the antennular peduncle, with 3 dorsal teeth, including the epigastric. Postrostral carina variable; adrostral carina present. Antennal spine present; branchiostegal spine on the margin of the carapace at the end of a sharp carina; hepatic spine present, hepatic sulcus well-defined below the spine; branchiocardiac sulcus deep, the sulcus prominent and accompanying the carina almost to the margin of the carapace; cervical sulcus well defined and reaching to at least the dorsal region of the carapace. Abdominal somite 3 with or without a prominent posterodorsal spine; somites 4-6 dorsally carinate; telson with 4 pairs of lateral movable spines. Eye small, cornea flattened; ventral flagellum of antennule not sexually dimorphic; scaphocerite exceeding the antennular peduncle, its apex well exceeding the distolateral tooth. Pereopods with rudimentary exopods, sometimes lacking on the fifth; a distal movable spine on the merus of pereopods | & 2. Dorsomedian lobule of the petasma nearly as long as the dorsolateral lobule, which is rounded distally; ventral costa with distal half free, the tip tapering. Thelycum with large elongate median protuberance on thoracic somite 7, with pointed anterior tip reaching the anterior rim of sternite 6. REMARKS. This genus includes H. glacialis Bate, 1881, South Atlantic; H. inermis Bate, 1881, south- central Pacific Ocean; H. tener Smith, 1884, Atlantic, Indian and Pacific Oceans. All 3 species inhabit deep water, which may explain their apparent rarity. H. inermis appears to have been collected only once (type locality, depth 4665 m) and H. glacialis is almost as rare FIG. 7. Hepomadus tener Smith, 1884, TMH4046, 22°S, 113°8’E, 1460-1700m (aborted trawl), d, 31mm, 419 (depth ca. 3400 m). However, H. tener, with a depth range of 765-5400m, mean 2400m, has been collected on a number of occasions, although still considered rare. KEY TO THE SPECIES OF HEPOMADUS 1. A prominent posterodorsal spine on abdominal somite 3 = No posterodorsal spine on abdominal somite 3 ee ee re ee rs ee ee H. inermis 2. Carapace with a pronounced hump behind the cervical sulcus, which interrupts the postrostral carina AA OW, Ord FOE E ST . at ee SE H. glacialis Carapace only slightly arched behind the cervical sulcus, postrostral carina continuing uninterrupted almost to the posterior marginofthecarapace........ H. tener Hepomadus tener Smith, 1884 (Fig. 7) Hepomadus tener Smith, 1884: 409, pl. 9, fig. 7-8: 1887: Bouvier, 1908: 57, pl. 1, fig. 5, pl. 13, figsl-12:; Burkenroad, 1936: 86; Ramadan, 1938: 55; Roberts & Pequegnat, 1970: 43, fig. 3 ; Pequegnat & Roberts, 1971: 9: Pérez Farfante, 1973: 442, figs 1-8; Crosnier, 1985: 860, 1994b: 369; Liu & Zhong, 1986: 28, fig. 10. Hepomadus tener? Wood-Mason & Alcock, 1981: 189. Aristaeus (Hepomadus) tener? Alcock, 1901: 42. Hepomadus glacialis Milne Edwards & Bouvier, 1909: 194, figs 13-19, pl. 1 fig 3. MATERIAL. TMH G4046, 22°S 113°E, 1460-1700m, 44, 21-34mm, 22, 29, 37mm. DIAGNOSIS. Cuticle flexible and glabrous. Rostrum styliform, upturned, its length relative to that of the carapace increasing with size, being shorter in smaller specimens and longer in larger specimens; with 3 dorsal teeth including the epigastric; adrostral carina well defined, reaching at least the base of the epigastric tooth. Postrostral carina almost reaching the posterior margin of the carapace, the postrostral region of the carapace slightly convex. Antennal, hepatic and branchiostegal spines present, the latter on the margin of the carapace at the end of a prominent carina, which extends as far as the hepatic spine. An orbito-antennal sulcus present, continuous with a deep hepatic sulcus, which meets the branchiocardiac sulcus, before turning ventrad. Branchiocardiac sulcus deep, the carina prominent, both almost reaching the posterior margin of the carapace, before turning ventrad. Cervical sulcus well-defined and reaching the postrostral carina; postcervical sulcus faint. 420 Abdominal somite 3 with weak dorsal carina on its posterior half ending in a large, down-curving spine; somites 4-6 each with a sharp dorsal carina; telson with 4 small movable spines. Optic peduncle long, cornea small and slightly flattened. Scaphocerite exceeding the antennular peduncle, the tip longer than the distolateral spine, Pereopods | & 2 each with a prominent movable meral spine. Dorsomedian lobules of petasma diverging distally; dorsolateral lobule rounded distally, about as long as the dorso- median lobule; ventral costa prominent, the distal half free and ending in bluntly pointed tip, directed laterally. Thelycum with an elongate plate on thoracic sternite 7, the sides rounded and pointed apex reaching almost to sternite 5; sternite 8 subrectangular, more or less convex. REMARKS, Pérez Farfante (1973) found a good deal of variation in specimens from the Atlantic, but nevertheless H. tener is a distinctive species. DISTRIBUTION. NW Australia, 22°S 113°E; 1460-1700m. Known range: Zanzibar, Madagascar, Réunion, central Indian Ocean, Bay of Bengal, Australia, Wallis and Futuma Islands, W and E Atlantic Ocean, 765-5400m. Parahepomadus Crosnier, 1978 Parahepomadus Crosnier, 1978: 47; Liu & Zhong, 1986: 30; Pérez Farfante & Kensley, 1997; 48. TYPE SPECIES. Parahepomadus vaubuni Crosnier, 1978 (monotypic). DIAGNOSIS. Cuticle thin and flexible, finely pubescent. Rostrum often short in adults of both sexes, not reaching beyond the second segment of the antennular peduncle, but may be slender and well exceeding the peduncle. Adrostral carina reaching just past the first rostral tooth; postrostral carina reaching about half the carapace. Postorbital spine present, continuous with the gastro-orbital carina, which almost reaches the shallow cervical sulcus, the latter limited to the lateral region of the carapace. A very small antennal spine may be present, absent in the larger specimens, but with the low antennal carina ending in a minute MEMOIRS OF THE QUEENSLAND MUSEUM tubercle just behind the carapace margin; orbito-antennal sulcus deep, continuous with the hepatic and branchiocardiac sulci. A large branchiostegal spine on the margin of the carapace, the branchiostegal carina reaching the branchiocardiac carina; the latter almost reaching the posterior margin of the carapace. Eye dorsoventrally flattened, with a small median tubercle on the peduncle; lower antennular flagellum with a slight sigmoidal flexure in males. Scaphocerite very wide; exopods usually absent from pereopods; minute subdistal meral spines on pereopods | & 2. Petasma very wide with rounded median lobe; thelycum a large concave setose plate, with prominent anterior point, on thoracic somite 7. Uniformly orange. Parahepomadus vaubani Crosnier, 1978 (Fig. 8A, B) Parahepomadus vaubani Crosnier, 1978: 48, figs 20-22, 1989, 1994a; Liu & Zhong 1986: 30, fig. 11. MATERIAL. NT Cr007084, 13°S 122°E, 900-1000m, 3, 48mm, °, 67mm; NTCr007085, 13°06’ 122°18’E, 900-1000m, 3, 53, 54mm, 22, 57, 58mm. DIAGNOSIS. As for genus. REMARKS. Two specimens had rudimentary exopods on pereopods 1-5, which could confuse identification, but the postantennal spine sets this apart from other members of the family. The normal very long slender rostrum is apparently subject to breakage, since most adults have a shortened rostrum (Crosnier, 1978, fig. 20). DISTRIBUTION. Off the Northwest Shelf of Australia, 13°S 122°E; 900-1000m. Known FIG. 8. Parahepomadus vaubani Crosnier, 1978 NT CR007085, 13°6’S 122°18’E, 900-1000m. A, 2, 58 mm, rostrum. B, ¢, 54mm, short rostrum. AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE range: Madagascar, Indonesia, Australia, Philippines, 750-1525m. Plesiopenaeus Bate, 1881 Aristeus Bate, 1881: 187 [part]; 1888: 309 [part]. Aristaeus Wood-Mason 1891: 278 [part]. Plesiopenaeus Bate, 1881: 188; Bouvier, 1908: 63 [part]; Burkenroad, 1936: 94; Ramadan, 1938: 49 [part]; Barnard, 1950: 621; Crosnier, 1978: 85: de Freitas, 1985: 20; Pérez Farfante & Kensley, 1997:50. Plesiopeneus Faxon, 1895; 199, Aristeopsis Bouvier, 1908: 61; Milne Edwards & Bouvier, 1909, 197. DIAGNOSIS. Integument firm, glabrous. Rostrum longer than the carapace in juveniles and adult 2 2, shorter in ¢ ¢; with 3 dorsal teeth. Antennal and branchiostegal spines prominent, the latter on the margin of the carapace and continuous with a sharp carina; orbital, postantennal, pterygostomian and hepatic spines absent; gastro-orbital carina present; cervical sulcus variable; branchiocardiac carina well-defined. Posterior half of abdominal somite 3 and 4-6 dorsally carinate. Telson with 4 pairs of movable lateral spines, apex acute. Cornea slightly flattened; a strong tubercle at about midlength of the peduncle. Dorsal antennular flagellum flattened, ventral flagellum elongate, slender and not modified in the adult d. Scaphocerite with thickened margin ending in a short spine at about 0.8 the length of the lamella. Merus of pereopod | or 1 & 2 with a sub-distal movable spine; exopods on all thoracic appendages, those of the pereopods rudimentary. Ventral costa of petasma free distally, the apical part hooked with tip directed laterally; thelycum with sternum of thoracic somite raised and shield-like, with acute apex. REMARKS. This genus contains P. armatus and P. coruscans, P. edwardsianus and P. nitidus having been transferred to the monotypic genera Aristaeopsis and Austropenaeus, respectively (Pérez Farfante & Kensley, 1997). P. armatus is readily distinguished by a prominent dorsal spine FIG, 9. Plesiopenaeus armatus (Bate, 1881) AMP39939, 11°42’S, 145°37°E, 2053m, 2, 39mm. 421 on abdominal somite 3, whereas P. coruscans has none. Plesiopenaeus armatus (Bate, 1881) (Fig. 9) Aristeus armatus Bate, 1881: 188, 1888: 312, pls. 45, 46. Aristeus ?tridens Smith 1884: 104, pl. 9, 1886b: 189,192, 193, 1887: 689, pl. 19. Aristaeopsis armata Wood-Mason & Alcock, 1891b: 285. Plesiopenaeus armatus, Vaxon, 1895: 199, 1896: 163; Burkenroad, 1936: 95; Ramadan, 1938: 51, Anderson & Lindner, 1945: 301; Roberts & Pequegnat, 1970: 46; Pequegnat & Roberts, 1971: 8, pl. 5; Wasmer, 1972: 259; Crosnier & Forest, 1973: 294, fig. c-d; Crosnier, 1978:93, fig. 33b, 1985: 863, 1994b: 369; Hayashi, 1983c: 368, fig. 58a-d; Pérez Farfante & Kensley, 1997: fig. 19. Aristaeus (Aristaeopsis) armatus Alcock, 1901: 41. Aristeopsis armatus Bouvier, 1905: 983; de Man, 1911: 6, Balss, 1925: 222. Aristeopsis armatus var. tridens Bouvier, 1908: 62, pl. 11, fig. 6; Milne Edwards & Bouvier, 1909; 197, figs 4-7, 20-27; de Man, 1911: 6; Bouvier, 1922: 12. Plesiopenaeus armatus tridens Burkenroad, 1936. MATERIAL. AMP39932, 20°42’S 160°E, 2450m, 6, 57mm, 2, 56mm; AMP39939, 12°S 146°E, 2053m, 2, 35mm; QMW13666 14°27°S 146°34’E, 1930-1942m, 4°, 47-57mm;QMW136667, 13°40°S 147°43°E, 2884-2932m, 3, 63mm, 2, 54mm; QMW13668, 13°29°S 147°13’E, 2490-2542m, 3d, 60mm, 32, 55, 70 80mm; QMW 13669 13°40°S 146°57E, 1880-1921m, 34, 57, 61, 65mm, 2, 43mm. DIAGNOSIS. Carapace glabrous. Rostrum long, well exceeding the scaphocerite, with 3 dorsal teeth; adrostral carina ending in front of the first tooth; postrostral carina low and reaching about half the carapace. Antennal spine with a short carina; branchiostegal spine prominent, on the margin of the carapace and with a carina extending past the beginning of the branchiocardiac carina; the latter reaching the posterior margin of the carapace, both carina and sulcus well defined; orbito-antennal sulcus deep and continuous with the wide hepatic sulcus which becomes indistinct at the level of the branchiocardiac carina. A low gastro-orbital carina present; cervical carina and sulcus present in the lateral region of the carapace only, the latter very wide and shallow. Abdominal somite 3 without a distinct dorsal carina, somites 4 -6 each with a carina; somites 3-6 each ending ina large tooth, that of somite 6 smaller; pleura of somites 3-5 postero-ventrally mucronate. Eye small, flattened; stylocerite 422 exceeding the second antennular segment; scaphocerite not sexually dimorphic; exopods on pereopods rudimentary, without setae; a subdistal spine on the merus of pereopods | & 2. Thelycal plates on thoracic somite 5-8 with a prominent median ridge. Colour. Uniformly wine red. REMARKS. This is one of the largest aristeids, the 80 mm @ being in the upper size range. Arist- aeopsis edwardsiana reaches a comparable size. DISTRIBUTION. NE Australia 11-21°S, Lord Howe Rise 27-30°S 159°40°E; 1880-2930m. Cosmopolitan, W and E Atlantic, Indo-Pacific from Madagascar and Zanzibar, Bay of Bengal, Australia, Philippines, off Japan, off Hawaian and Tuamotu Islands, W coast USA, 752-5413m. Plesiopenaeus coruscans (Wood-Mason, 1891) Aristeus coruscans Wood-Mason in Wood-Mason & Alcock, 1891b; 280, fig, 6. Aristaeus coruscans Wood-Mason, 1892; pl. 2, fig. 3; Faxon, 1895: 198. Aristaeus (Plesiopenaeus) coruscans Alcock, 1901: 37. Plesiopenaeus coruscans Bouvier, 1908: 69; de Man, 1911: 6; Burkenroad, 1936: 95, fig. 61; Roberts & Pequegnat, 1971: 46; Crosnier, 1978:94, fig. 33e-f; Liu & Zhong 1986: 47, fig. 19. MATERIAL. NTCR006994, 13°S 122°F, 900-1000m, 2, 63mm. DIAGNOSIS. Rostrum exceeding the antennular peduncle in juveniles and adult ° °, shorter in adult dd; with 3 dorsal teeth; adrostral carina extending past first rostral tooth; postrostral carina nearly reaching the posterior margin of the carpace. Antennal spine with a carina almost reaching the orbito-antennal sulcus; gastro- orbital carina prominent and reaching the cervical sulcus; branchiostegal spine large and on the margin of the carapace, with prominent carina reaching the branchiocardiac carina. Hepatic sulcus continuous with the branchiocardiac sulcus, which, with the carina almost reaches the posterior margin of the carapace; cervical sulcus shallow, but almost reaching the mid-dorsum; a short carina on the lower part. A dorsal carina on the posterior half of abdominal somite 4 and on the full length of somite 5 & 6, Posteroventral pleura of abdominal somites 3-5 not mucronate. Eye dorsoventrally flattened, the peduncle long and with a prominent mesial tubercle. Pereopod 1 only with a sub-distal spine on the merus; rudimentary exopods on all pereopods. Thelycum with prominent anteriorly pointed flat MEMOIRS OF THE QUEENSLAND MUSEUM sternal plate on thoracic somite 7; somite 8 witha transverse low triangular projection. Colour. Clear orange. REMARKS. The sole specimen available was badly damaged and unsuitable for illustration (Crosnier, 1978). Its integument was exceptionally thin and flexible, more so than most other members of this family, and it could have been newly moulted. It is a large species and so far rare (Crosnier, 1978 noted that there were only 4 specimens known besides the 2 from Madagascar). Other features which distinguish it from P armatus besides the absence of a large dorsal abdominal spine, are the cervical sulcus which almost reaches the mid-dorsum, lack of a spine on the merus of pereopod 2, lack of any armature on the pleura of the abdomen and a median carina on the thelycum. DISTRIBUTION. Northwest Shelf of Australia, 900-1000m. Known range: Indian Ocean from Madagascar, the Arabian Sea, Bay of Bengal to NW Australia and the Atlantic Ocean, near the Bahamas, 900-2367m. Pseudaristeus Crosnier, 1978 Hemipenaeus Bate, 1881: 186 [part]; Ramadan, 1938; 47[part]. Aristaeus Wood-Mason, 1891: 278 |part]. Aristaeus (Hemipenaeus), Alcock, 1901: 31 [part]. Pseudaristeus Crosnier, 1978: 81; de Freitas, 1985: 11; Pérez Farfante, 1987: 312; Pérez Farfante & Kensley, 1997: 52. DIAGNOSIS. Integument pubescent. Rostrum elongate, when undamaged reaching well beyond the scaphocerite; with 3 dorsal teeth; postrostral carina defined. Carapace with antennal and marginal branchiostegal spines, the latter continuous with a strong carina; hepatic sulcus present; orbital, pterygostomian and hepatic spines absent; cervical sulcus reaching the mid-dorsum, a postcervical sulcus present; branchiocardiac carina and sulcus well defined. Abdominal somites 1-3 dorsally rounded, somites 4-6 dorsally carinate, somite 4 carinate in its posterior half only. Telson with slender acute apex, with 4 pairs of movable lateral spines. Eye well developed, dorsoventrally flattened; peduncle with a small tubercle on the mesial margin. Scaphocerite not sexually dimorphic. Pereopods without exopods, 1 & 2 each with a subdistal movable spine on the merus. Pereopod 3 without a podobranch. Petasma with distal margin of dorsolateral lobule oblique, reaching or slightly surpassing the median lobe; ventral costa distally free for ca. half its length, turning mesially. Thelycum open, with large lanceolate AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE plate on thoracic sternite 7; a broad plate on sternite 8, produced into anterolateral hoods. The genus includes P. crassipes (Wood-Mason, 1891), P. gracilis (Bate, 1888), P kathleenae Pérez Farfante, 1987, P. protensus Pérez Farfante, 1987, P. sibogae (De Man, 1911), and P. speciosus (Bate, 1881). The last is an Atlantic species and P. gracilis is known only from the Philippines. P. sibogae and P. kathleenae have both been collected from Australian seas, while P. crassipes has been found in Indonesia and could also be an Australian species. Pérez Farfante (1987) erected P. protensus on the differences in proportions of the thelycal plates, based on 2 ? 2 from Indian seas, but Crosnier (1994a) considered it a doubtful species. More specimens, particularly a mature d will be necessary to validate P protensus. KEY TO THE INDO-WEST PACIFIC SPECIES OF PSEUDARISTEUS 1, Pereopods covered with minute setae. Optic peduncle long, mesial margin at least 1.4 the width of the distal ONION 0 WS af PW Se ee ene P. sibogae Pereopods not covered with setae. Optic peduncle relatively short, mesial margin 1.3 or less the width of the distalextremity. 2... 2... 2 ee ee 2 2, Third segment of antennular peduncle expanded laterally, forming a prominent subtriangular projection in males and asmaller rounded structure in females . . . P. kathleenae Third segment of antennular peduncle not expanded laterally’ $"P te oR F end ge hae leg: 3 3. Males with ventral antennular flagellum sinuous proximally, with a narrow band of small dense setae, distal to the tip of the dorsal flagellum. Median thelycal plate of thoracic somite 7 expanded in a pair of posterolateral prominences ......... P. gracilis Males with ventral antennular flagellum straight proximally and without a band of small setae. Median thelycal plate of thoracic somite 7 without posterolateral prominences.. 2... 2.2.0.0 0005 P. crassipes Pseudaristeus kathleenae Pérez Farfante, 1987 (Fig. 10A,B) Aristaeus crassipes Wood-Mason, 1891 [part]; Alcock, 190 1a [part]. Hemipenaeus crassipes De Man, 1911: 24 [part]; 1913: pl. 2, fig. 4a-c; Kemp & Seymour Sewell, 1912: 17, pl 1, fig. 8 [part]; Balss, 1925 [part]. Pseudaristeus kathleenae Pérez Farfante, 1987: 314, figs 1-9; Crosnier, 1994a: 353; Pérez Farfante & Kensley, 1997: figs 21-23. FIG. 10. Pseudaristeus kathleenae Pérez Farfante, 1987. A, NT CR 006629, 9°17’S 131°8’E, 297m, 2°, 36.5mm, antennule. B, ¢, 23.5mm, antennule (after Pérez Farfante, 1987, fig. 2). MATERIAL. NT Cr006629, 9°17’S 131°E, 297m, 3, 29, 31, 36mm (all damaged and unsuitable for illustration). DIAGNOSIS. Carapace finely pubescent, cuticle thin. Rostrum usually exceeding the antennular peduncle in 2 2 andimmature 3 3, much shorter in mature dd; with 3 dorsal teeth; adrostral carina ending just behind the first rostral tooth; postrostral carina distinct up to the cervical sulcus, becoming indistinct thereafter and reaching about 0.75 the carapace. Antennal spine prominent with the carina reaching the shallow orbito-antennal sulcus; gastro-orbital carina well defined and reaching the cervical sulcus. Branchiostegal spine large, on the margin of the carapace and continuous with a prominent carina, which almost reaches the branchiocardiac carina. Hepatic sulcus continuous with the orbito-antennal sulcus and turning sharply downwards at its junction with the branchiocardiac sulcus. Branchiocardiac carina forming a wide arc, which runs downward into the posterior margin of the carapace. Cervical sulcus shallow, but crossing the mid-dorsum, the carina limited to a short prominent shallow are in its ventral region; postcervical sulcus not quite reaching the mid-dorsum and curving anteriorly towards the cervical sulcus, but not meeting it; with a low carina in its ventral region. Abdomen dorsally carinate on somites 4-6, each carina ending in a tooth. Mesial length of the optic peduncle/ maximum width at the cornea about 1.0. Tip of stylocerite well short of the disto-lateral spine of the first segment of the antennular peduncle; third segment with a large triangular ventro-lateral expansion ¢ and a rounded bulbous swelling in 2 (Fig. 10A, B). Third maxilliped dimorphic, in the male the dactyl inserted subapically in the propodus, curved and of uniform width, with a dense tuft of 424 subapical setae, Perenpods | & 2 with subdistal spine on the merus. Dorsolateral lobule of petasma expanding distolatcrally before tapering toa bluntly poiited mesial apex. Sternal plate of thoracic somite 7 a large densely setose lanceolate plate, maximum width about 0.65 the length, plate on somite & produced anteriolaterally into a pair of short hoods, REMARKS. Although the genus has been revised (Pérez Farfante, 1987) P sibagae. P kathleenae and P. crassipes have similar ranges and may still be confused. P. sibogue 38 reasonably distinctive (see under 2 sihogae lor a discussion of these features), but the Y ? of P kathleenae and P erassipes are difficult to distinguish. The 4 P. karhleenae is readily separated froin P crussipes by the sexually dimorphic antennule and third maxilliped and the expanded dorsolateral lobule of the petasma. In contrast, in the & only the swollen third segment of the antennular pedunele of 2 kathleenue clearly distinguishes it from P. crussipes and this feature is easily missed. The thelycal plate on thoracic somite 7 appears to be relatively broader in P kathleenae (width/length 0.67-0,75 versus ().43-0.55 in P. crassipes), Otherwise the thelyca are similar. Crosnier (1978) showed that pereopods | & 2 were stouter in 2 sibogae than in P. crassipes and P gracilis by companng the ratios of length/width of the carpi. These ratios for pereopads | & 2 are 8.5-8.9 and 10.5 respectively in PF kethleenae and 8.6-9.5 and 10.1-12.9 in PF crassipes, and 50 these 2 species are indistinguishable by this means. Thus the relative broadness of the thelycal plate is the only feature to assist identification of ° & when the presence or absence of the swelling of the third segment ofthe antennular pedunele ts in doubt. DISTRIBUTION. Australia, Arafura Sea 9°S 131°E, 297m. Known range: Southern India, Indonesia, Arafura Sea, Philippines, 297-|225m. (The Australian location is much shallower than the other records of depth, which start at 549m.) MEMOIRS OF THE QUEENSLAND MUSEUM Pseudaristeus sibogae (de Man, 1911) (Fig. 11A-E) Hemipencers sibogie de Man. $9112 28, 1914. pl. 2. fi, 5. Sa-c; Ramadan, 193%: 48; Anderson & Lindner, 1945; 301, Psrendaristeny sibogoe Crosnier, 1978: 83, figs 27a, 30u-c, 1994q; 353; de Freitas, (985: 12, fig. 0-5: Pérez Varfante, 1987: 332. fig. 18. MATERIAL, AM P41894,22°S 114°, 1158m, 9, 37mm} AMPS55940, 20°S 113°E. 9ldm, @, 24mm, ¥, 36mm: CSIRO SS/V/91, 443, #22, #24. 449, 23-28°S LII-114°R, §54-1305im, 34. 31, 32 33mm, 102, 31-43 mm: OM W24354, 33°S58°S 131°22°E. 1000m, 4. 33mm. DIAGNOSIS. Body finely pubescent. Rostrum normally exceeding the antennular peduncle in both sexes; with 3 dorsal teeth; adrostral carina ending at the first rostral tooth; postrostral carina reaching about 0.9 the length of the carapace. Antennal spine with carina; gastro-orbital carina and orbito-antennal sulcus present. A promineiit branchiostegal spine on the margin of the carapace, with sharp carina reaching the branchiocardiac carina; hepatic sulcus wide; branchiocardiac carina low, but well-defined. the carina and sulcus reaching the posterior margin of the carapace, where they turn down, the sulcus joining the submarginal sulcus. Cervical suleus shallow, but crossing the mid-dorstun, the carina limited to a short prominent shallow are in its ventral region; a posteervical sulcus present, not FIG. VL. A, Pseuduristeus sibogae (de Man, 1911) QMW25354, 33°58'S, 13122 E, 1000m, 2, 33mm; B, dorsal aspect of petasma; C, ventral aspect of petasma; D, appendix masculina; E. propodus and dactyl of maxilliped 3. (Sealebar = linn). AUSTRALIAN ARISTEIDAE AND BENTHESICY MIDAE quite meeting the mid-dorsum; a short auxiliary posthepatic carina and sulcus running in an arc from the cervical sulcus towards the branchiocardiac sulcus. A dorsal carina beginning at the middle of abdominal somite 3, somites 3-6 each ending in a small tooth. Cornea flattened, the mesial length of the peduncle at least 1.4 times the maximum distal width at the cornea. Third maxilliped sexually dimorphic (Fig. 1] E); pereopods 1-3 relatively robust, 4 & 5 much more slender, all covered with a fine pubescence. Distal half of dorsomedian lobule of petasma free and reaching almost as far as the distolateral lobule (Fig. 11 B, C); the latter tapering towards the midline, apex rounded; ventral costa prominent, the distal half free, with truncated tip; ventral surface covered with fine setae. Appendix masculina with ovate outer plate; inner plate (appendix interna) about 2.5 times the length of the outer plate, flexible and tapering (Fig. 11D). Thelycal plate on thoracic somite 8 produced anterolaterally into short hoods, that on somite 7 lanceolate, broad (width 0.6 length), without posterolateral prominences. Colour. Orange. REMARKS. P. sibogae has 4 distinctive features, which while reliable, are not completely definitive: the pubescent pereopods; the relatively long optic peduncle; the well-defined auxiliary carina and sulcus above the posterior hepatic sulcus; the stouter pereopods. The ratio of length/width of the optic peduncle was only 1.45 in these specimens, although Pérez Farfante (1987) recorded 1.5-1.75 (but such measurements are subject to operater variability). Other species also have at least indications of an auxiliary sulcus. Crosnier (1978) showed that the pereopods were stouter than in other species by comparing the ratios of length/width of podomeres of the pereopods, the ratios of the first and second in particular being appreciably smaller in P. sibogae than in other species. In the above specimens examined, the ratios of length/width of the carpus of pereopods 1 & 2 were 5.8-6.7 and 7.0-7.2, respectively, TABLE 3. Distribution of branchiae and epipods on the thoracic somites of the genera of the Benthesicymidae. which are close to those of Crosnier. Thus the only dubious feature in the specimens examined was the length/width ratio of the optic peduncle, but this appears to be a variable characteristic. Pérez Farfante (1987) used features of the genitalia to define species, but again ratios of length/breadth of thelycal plates appear to be variable. The petasma of a mature d (QM W24354, Fig. 11B, C), previously undescribed, is similar to that of P. crassipes, except for the fine setae on the ventral surface. DISTRIBUTION. Western Australia 20-28°S, Great Australian Bight; 854 -1305m. Known range: Off Natal, South Africa, Madagascar, Indonesia, Australia, 834-1305m. Family BENTHESICYMIDAE Wood-Mason, 189] Benthesicvmina Wood-Mason, 1891: 286. Benthesicvmae Bouvier, 1908: 16; Burkenroad, 1936: 15; Anderson & Lindner, 1943: 290: Balss, 1957: 1517; Tirmizi, 1960: 321; Roberts & Pequegnal, 1970: 32. Benthesicyminae Crosnier, 1978: 14; Hayashi, 1983d: 438. Benthesicymidae Pérez Farfante & Kensley, 1987: 56. DIAGNOSIS. Integument thin and soft. Rostrum truncate and blade-like, with the tip usually falling short of the cornea, occasionally not reaching much beyond it; without adrostral carina; dorsally armed only with not more than 3, usually 2 or fewer dorsal teeth. Branchiostegite with a more or less deep antero-ventral emargination, with branchiostegal spine either on or just behind the carapace margin. Hepatic spine present or absent; hepatic carina and sulcus usually well defined; cervical and usually the postcervical sulcus reaching the dorsal mid-line. Telson with 1-4 pairs of movable lateral spines, apex usually acute, sometimes truncate. Eye medium to small, the peduncle with a mesial tubercle; prosartema usually only a tuft of setae; both antennular flagella long and filiform. Exopods on all maxillipeds, present or absent on pereopods. Petasma open, usually broadly lamellar, with the flexible part of the ventrolateral lobule attached to the dorsolateral lobule for most [ Pleurobranchs | Arthrobranchs 2 Arthrobranchs Podobranchs | Epipods | Genus Somites 3-8 Somite | Somites 2-7 Somites 2-6 | Somites 1-7 | | Bentheogennema _ | + _| ] Ft : | + je + : Benthesicvmus | 4 | s | _+ | + | | Benthonectes I + j 1 + | 2—5, 6s _| + \ lL Gennaclas _ + im | r + somite 2 only + | or all of its length; ventral costa entirely attached. Thelycum open or closed with seminal recept- acles formed by paired sternal invaginations at the base of pereopods 3. Arrangement of branchiae and epipods in the genera of the Benthesicymidae is shown in Table 3. KEY TO THE GENERA OF THE BENTHESICYMIDAE 1. Podobranchs on thoracic somites 2-6; telson usually armed with more than one pair of lateral movable spines . . . 2 Podobranchs present on thoracic somite 2 only; telson armed with only one pair of lateral movable spines Pea wed tee he He Gree a oes ed Gennadas 2. Telson apically pointed. At least two abdominal somites dorsally carinate.. 2. 2... ee ee eee 3 Telson apically truncate; only abdominal somite 6 dorsally carinate. 2... eee. Bentheogennema 3. Dactyls of pereopods 4 & 5 not multiarticulate and not markedly elongate; usually a dorsal carina on abdominal somite 4, as well as 5 & 6; dacty| of maxilliped 3 spinous anidsetdsels ua ay KER Benthesicymus Dactyls of pereopods 4 & 5 multiarticulate and markedly elongate; a dorsal carina on abdominal somites 5 & 6 only; dactyl of maxilliped setose but not spinous » ME SE lee 6 adie a eae oo tl Bt Le ford Benthonectes Gennadas is well represented in Australian seas by 8 species, but so far only 2 Benthesicvmus species have been positively identified, with a possible third new species; Bentheogennema has yet to be recorded. NTCR007066 was identified as Benthonectes filipes by Dr A. Crosnier in 1990. However, | found that it is now too badly damaged to be positively identified, is of no value as areference specimen, and a description of this species has therefore not been included (Pérez Farfante & Kensley, 1997 for description and figures). Benthesicymus Bate, 1881 Benthesicymus Bate, 1881: 171, 190, 1888: 326; Alcock, 1901: 42; de Man, 1911:13; Burkenroad, 1936: 23; Anderson & Lindner, 1943: 296; Tirmizi, 1960: 322; Crosnier, 1978: 15; Hayashi, 1983d: 438; Squires, 1990: 21; Kikuchi & Nemoto, 1991: 64; Pérez Farfante & Kensley, 1997: 59. Benthoecetes Smith, 1884: 391, Gennadas Alcock, 1901: 46 [part]. DIAGNOSIS. Rostrum apically acute, often falling short of the cornea, with 0-3 dorsal teeth. Branchiostegal spine on or just behind the margin of the carapace, continuous with hepatic carina; hepatic spine present or absent; cervical, postcervical, hepatic and branchiocardiac sulci well marked. Abdominal somites 5 & 6 dorsally carinate, usually a low carina on the posterior half of the fourth. Telson with 4 pairs of movable lateral spines, apex acute. Eye medium to small, MEMOIRS OF THE QUEENSLAND MUSEUM the cornea slightly wider than the peduncle, brown to black-pigmented; a conical mesial process at about half the length of the peduncle. Dactyli of pereopods 4 & 5 slender and uniarticulate. Petasma with dorsolateral lobule broad, sometimes distally bilobed, longer than the ventrolateral lobule. KEY TO THE INDO-WEST PACIFIC SPECIES OF BENTHESICYMUS |. Branchiostegal spine at margin of carapace, branchiostegal carina not sharp; dactylus of 3rd maxilliped triangular, with only one strong spine at the tip (Group]e Bar pe eR Bee ne of fob 2 Branchiostegal spine just behind margin of carapace, branchiostegal carina very sharp; dactylus of 3rd maxilliped subrectangular, distal margin with more than onestrongspine(Group2)... 0.0... 50.4.5 1 Posterolateral margin ofabdominal somite 4 crenate. . 3 Posterolateral margin of abdominal somite 4 not crenate 4 3. Hepatic spine present; posterolateral margin of abdominal somite 5 without a spine; merus and ischium of maxilliped 3 each with a small acute spine on the median distalpart............-, B. crenatus Hepatic spine absent; posterolateral margin of abdominal somite 5 with a small spine; merus and ischium of maxilliped 3 each without spines on the median distal hm Pat oe caw nese ett anaeeaey B. laciniatus 4. Hepaticspinepresent........-..-..8-2...5 Hepaticspineabsent.......-..-. 10 5, Abdominal somite 3 with a postero-dorsal spine Si ay Pe ers ee, ot B. brasiliensis Abdominal somite 3 without a postero-dorsal spine . . 6 6. Abdominal somite 4 with a postero-dorsal spine cate go ete Ue ng wer ce eee *B. urinator Abdominal somite 4 without a postero-dorsal spine . . 7 7. Merus of 2nd maxilliped more than 3.5 times as long as Brodd®: sy 't Wawnas ce nea W oer i ee a B. strabus Merus of 2nd maxilliped less than 3 times as longas broad. 8 8. Abdominal somites 5 & 6 each with a postero-dorsal spine Abdominal somites 5 & 6 each without a postero-dorsal SPINE, 3) 2 face Sih ay we a 5s 2 B. iridenscens 9. Posterior rostral tooth usually anterior to the level of the orbitalmargin. ©... 2. eee eee B. longipes Posterior rostral tooth well behind the level of the orbital mare te ee eRe *B. sevmouri 10. Abdominal somite 6 more than 2.5 times as long as the FIN SOMME ye es oe ew es B. brevirostris Abdominal somite 6 less than 2.5 times as long as the fifth SOMITE. 6 er ee ee B. carinatus Il. Hepaticspinepresent, 2... 2.2... 00. B. tanneri Hepatic spine absent 12, Abdominal somite 5 with a long spine extending backwatds.5 25 i dood ol Ma gb Eg 13 13. Long spine of abdominal somite 5 extending backwards from the middle ofthe dorsal mesial margin . B. bartletti AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE 427 Long spine of abdominal somite 5 extending backwards from the postero-dorsal margin... ... . B. tirmiziae 14. Postero-dorsal tip of abdominal somite 6 curving upwards’ 22 3248. a Pe oe get wins B. altus Postero-dorsal tip of abdominal somite 6 directed horizontally. 2... ee B. investigatoris *See discussion under B. wrinator howensis sp. nov. Only B. investigatoris, B. urinator and a possible new species have been collected from Australian seas, but probably more species are present as Kikuchi & Nemoto (1991) recorded 8 species from the NW Pacific and Crosnier (1978, 1985) 8 from the W Indian Ocean, 5 of which were common to those of Kikuchi & Nemoto (see Zoogeography of the Aristeidae and Benthes- icymidae). Benthesicymus investigatoris Alcock & Anderson, 1899 (Fig. 12) Benthesicvmus investigatoris Alcock & Anderson, 1899a: 282, 1899b: pl. 41, fig. 2; Alcock 1901: 44; Rathbun, 1906: 906; Balss, 1927: 247, fig. 1; Burkenroad, 1936: 49; Anderson & Lindner, 1945: 298; Crosnier, 1978: 21, pls. 7c-d, 8c-d, 9, 10, 1984: 20, 1985: 857, 1989: 41, 1994a: 351, 1994b: 368; Hayashi, 1983d: 440, fig. 61; Kensley et al., 1987: 276; Kikuchi & Nemoto, 1991: 88, figs 16, 17. Benthesicymus investigatori Borradaile, 1910: 258. Benthesicymus Investigatoris de Man, 1911: 5, 14; 1913: pl. 1, fig.1. FIG. 12. Benthesicymus investigatoris Alcock & Anderson, 1899 AMP39937, 27°59" S, 162°48’E, 1250m, 3, 16.5mm. MATERIAL. AMP39937, 28°S 163°E, 1250m, 60, 15-16.5mm, 52, 12.5-15mm; QMW13483, 17°45’S 148°E, 1115m, 2d, 14, 18mm, 62, 14-21mm; QMW13447, 17°19’S 147°47°E, 1100m, °, 20mm. DIAGNOSIS. Rostrum reaching about half the cornea, with 2 teeth and a minute bump in the position of a third, more posterior tooth; postrostral carina ending at the cervical sulcus. Antennal and hepatic spines absent; orbito-antennal sulcus present; branchiostegal spine prominent, raised above the surface of the carapace, set back just behind its margin and continuous with a prominent hepatic carina, which extends as far as the branchiocardiac carina; both carina and sulcus well defined. Cervical sulcus deep, with a shallow notch where it crosses the mid-dorsum. A posthepatic carina and sulcus present, the sulcus turning towards the dorsum, which it crosses without a notch, at about 0.6 the length of the carapace; the accompanying carina interrupted at this level, but continuing parallel to the branchiocardiac sulcus. A dorsal carina clearly defined on abdominal somites 5 & 6, a barely defined carina on the posterior of somite 4 ; that of somite 5 ending in a small spine. Eye small, with brown pigment; pereopod 1 with a subdistal movable spine on the merus; rudimentary exopods on all pereopods. Petasma with dorsolateral lobule distally entire and semicircular, with thickened rim, ventral costa prominent and reaching almost as far as the dorsolateral lobule. Thelycum with a triangular prominence on the sternum of thoracic somite 6, the sternum of thoracic somite 7 a rounded projection with anterior edge w-shaped, that of somite 8 with two lateral circular indentations. REMARKS. Of the 20 specimens examined, 11 (of both sexes and little different in size) did not have a well-defined dorsal carina on abdominal somite 4, but were otherwise valid B. investigatoris. Thus this appears to be a variable character in this species. DISTRIBUTION. E Australia 10-34°S: Lord Howe Rise 28°S 163°E; WA 29°05’S 113°14°E; Northwest Shelf 13°06’S 122°18°E; 879-1250m. Known range: throughout the Indo-West Pacific E coast of Africa, through the Indian Ocean, Indonesia, Australia, Philippines, Japan, Hawaii, Fiji, Wallis & Futuna Islands, 580-1690m. A common benthesicymid throughout its range; one trawl from the Cidaris’ off NE Queensland included over 150. Benthesicymus urinator Burkenroad, 1936 Benthesicymus urinator Burkenroad, 1936: 29, figs 4, 5, 8, 9, 17, 18, 22, 32, 33, 39, 40, 45. Crosnier, 1985: 843, fig. 2; Kikuchi & Nemoto, 1991: 67, figs 4,5. Benthesicvmus brasilensis Bate, 1881: 191, 1888: 332 [part]. Benthesicymus moratus Rathbun, 1906; 907. DIAGNOSIS. Rostrum reaching about as far as the eye, tapering to the tip, with 2 fairly prominent dorsal teeth, with indications of a third; posterior tooth behind the margin of the 428 FIG. 13. Benthesicymus urinator howensis sp. nov. AM P40648, Lord Howe rise 28°44’S, 161°54’E, 1325m, 2, 22mm. A, carapace; B, thelycum; C, maxilliped 2; D, maxilliped 3 dactyl. (Scalebar = 1mm) carapace. Antennal spine acute, hepatic spine well developed; branchiostegal spine prominent, situated on the margin of the carapace and witha short carina. Cervical sulcus deep, notching the mid-dorsum and with a lower posterior branch which joins the branchiocardiac sulcus, the anterior branch reaching as far as the hepatic spine; branchiocardiac carina prominent; postcervical sulcus obscure; marginal carina well developed. Abdominal somites 4-6 carinate, each ending in a small spine; sixth somite more than twice as long as the fifth. Mandibular palp 2-seg- mented, basal segment 1.5 times the length and much broader than the distal segment, cutting edge of mandible straight with a small anterior tooth; endopod of second maxilla with long apical spine and seven long curved spines on the outer edge of the tip and a row of smaller spines on the inner edge; merus of second maxilliped expanded, length/width ratio 2.5, dactyl with 6 strong marginal spines; dactyl of third maxilliped broadest at its midpoint, tapering distally with 1 large spine at the tip. MEMOIRS OF THE QUEENSLAND MUSEUM Thelycum with inflated sternal plate on thoracic somite 8, a strong groove between sternites 7 & 8, sternite 7 with a prominent triangular projection, the apex directed anteriorly, a conical projection on somite 6. REMARKS, Two 6 d and 2 22 (holotype and paratypes) were collected by the HMS Challenger’ (Station 184) in the eastern approaches to Torres Strait. Kikuchi & Nemoto (1991) note that only 13 specimens have been collected in the Indo-West Pacific, so it appears to be uncommon, although widely distributed. All specimens are lodged in overseas museums and thus were not available (see, however, Fig. 13, B. urinator howensis sp. nov.). As itisa Group I’ benthesicymid it may be easily separated from the relatively common B. investigatoris. DISTRIBUTION. Australia, Torres Strait approaches, 2560m. Known range: Indian Ocean, SW and N Pacific Oceans, 2500-4200m. Benthesicymus urinator howensis sp. nov. (Fig. 13A-D) MATERIAL. AM P40648, Lord Howe Rise, 28°44’S 161°54’E, 1325m, 2°, 22, 24mm. DISCUSSION. The above description for B. urinator (see also Crosnier, 1985, Fig. 2a, Kikuchi & Nemoto, 1991, figs 4, 5a) fits these specimens exactly in all respects except for the armature of the abdomen. They lack a posterior-median spine of abdominal somite 4, which is a diagnostic feature of B. urinator, Crosnier (1985) remarking that this spine is the largest of the 3 abdominal spines. Unfortunately, the larger specimen is in poor condition and appears to lack a spine even on the fifth somite, but the 22mm specimen has a minute, but definite spine in this position, as well as a prominent spine on the sixth somite (both have a carina on the fifth AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE and sixth somites, but only a feeble indication of a carina on the fourth). As such it keys out as B. seymouri, which is obviously incorrect. Since the armature and carination of the abdominal somites are regarded as sufficiently stable to be used as key characters at both the specific and generic levels, these specimens may warrant the status of a new species. However, B. urinator is a rare species and the abdominal armature may be more variable than has been supposed. This plus the limited amount of material (only one relatively undamaged specimen) indicates that the erection of a new species would be unwarranted at this stage. Rather than designate it Benthesicymus sp.’ or Benthesicymus cf. wrinator, in view of its close similarity to B. wrinator, | prefer to assign it provisionally to a new subspecies. Gennadas Bate, 1881 Gennadas Bate, 1881: 171, 191; 1888: 339; Alcock, 1901: 45: de Man, 1911: 15; Kemp, 1913: 60; Calman, 1925: 3; Balss, 1927: 248; Burkenroad. 1936: 59; Anderson & Lindner, 1943: 291; Barnard, 1950: 627 ; Kensley, 1971: 272; Crosnier, 1978: 33; Hayashi, 1984a: 18; Squires, 1990: 26; Pérez Farfante & Kensley, 1997: 63. Amalopenaeus Smith, 1882: 86. Pasiphodes Filhol, 1885: pl 3. DIAGNOSIS. Rostrum short, with a single dorsal tooth, not reaching as far as the cornea of the eye; adrostral carina absent, postrostral carina usually present. Antennal angle acute, narrowly rounded; infra-antennal angle demarcating a deep emargination of the carapace, A branchiostegal spine usually present on the inner edge of this emargination, branchiostegal carina present. Cervical and postcervical sulci present, both reaching the dorsal midline; a weak hepatic carina present. Only abdominal somite 6 with a dorsal carina; telson apically truncate, with a single pair of movable lateral spines. Second and third segments of antennular peduncle slightly expanded; exopod of first maxilliped without distal segmentation. Dactlyi of pereopods 4 & 5 slender, uniarticulate. Petasma with distal margin divided into external, median and internal lobes; accessory lobe always present; thelycum variable, but with small shallow seminal receptacles at the base of pereopod 3, with either paired openings or a common median opening. REMARKS. Seventeen species of Gennadas world wide have been described of which 5 appear to be restricted to the Atlantic and | to the E Pacific, but most of the remainder have been recorded world wide. The following are Indo- West Pacific species: G bouvieri, capensis, crassus, gilchristi, incertus, kempi, parvus, 429 propinquus, scutatus, sordidus, tinayrei. G. parvus Bate, 1881 has not yet been recorded from Australia, but in view of its wide distribution probably occurs here. G crassus Tirmizi, 1960 has been recorded only from Zanzibar; G sordidus Kemp, 1910 appears to be restricted to the NW Indian Ocean: the Arabian Sea, Gulf of Aden and Gulf of Oman. Neither of these species is included in the key below. Gennadas species are soft and mostly < 10 mm CL; the features of the carapace and abdomen are closely similar and they would be extremely difficult to identify were it not for their very distinctive genitalia. The following keys use only features of the latter, with separate keys for males and females. KEY TO THE INDO-WEST PACIFIC SPECIES OF GENNADAS Adult male petasmas: Median lobe undivided... ..--...-...--.. 2 Median lobedivided. .........2-.2.0.,4. 4 External lobe divided by closely approximate blunt Iebuless 65, cm 23 00 iy BY ae te G., tinayrei External lobe undivided or with a small acute medial No PMICESA see te. ben Race Pate +s 3 3. Accessory lobebipartite... 2... ..04;, G. capensis Accessory lobeasingle flap .......... G kempi 4, Lobules of external lobe elongate, subequal, slender ifohe trict par detytdby cds + + mpeduhown pach G, incertus Lobules of external lobe not elongate, subequal, slender 5 5. Lobulesofmedianlobehooked ....... G, bouvieri Lobules of median lobe not hooked 6. Accessory lobeamereridge.......... Accessory lobe welldeveloped.. 2. 2... 0. uk 7 7. Apexofinternallobeacute ......... Apex of internal lobe rounded 8. Inner lobule ofmedian lobe apically acute. G. propinguus Inner lobule of median lobe apically truncate. G scutatus Adult female thelyca: 1. A posteriorly directed tongue-like projection on thoracic sternite 5 G. tinayrei No tongue-like posterior projection on thoracic sternite 5 5) tN No obvious thelycal plate on thoracic somite 8 ob? bored é fof aka > ot eS fe & oct G. capensis Awell-developed thelycal plateonsomite8...... 3 3. Thelycal plate on thoracic somite 8 with broad rounded anterior extension reaching sternite of somite 6 hE pote hp Che ak tod hee G, scutatus Thelycal plate on thoracic somite 8 not reaching the sterniteofsomite6 2... ee ee 4. Thelycal plate on thoracic somite 8 broad, anteriorly notched or indented and overlapping part of somite plate G. incertus Thelycal plate on thoracic somite 8 not indented anteriorly and notoverlappingplate7.........5 wi Thelycal plate on thoracic somite 8 with two slender antero-lateral projections... . 2.2... G. bouveri Thelycal plate on thoracic somite 8 without two slender antero-lateral projections... ............ 6 6. Thelyeal plate on thoracic somite 7 with two medial anterior projections ............ G. gilchristi Thelycal plate on thoracic somite 7 without two anterior projections 7. A leaf-like process arising between the coxae of pereopods 3 & 4and medially directed. . . . G parvus No leaf-like process arising between the coxae of perecpods sitet: |S nat, lees PRS Lie 8 8. Thelycal plate on thoracic somite 7 W-shaped . . . . G. PRODIMGUUS ey ce hte i a Eo te pete fe Thelycal plate on thoracic somite 7 rectangular. G kempi Gennadas bouvieri Kemp, 1909 (Fig. 14A-C) Gennadas bouvieri Kemp, 1909: 726, pl. 74, fig. 1-4, pl 75, fig. 6-7, 1910: 179; Burkenroad, 1936: 80; Anderson & Lindner, 1945: 293; Tirmizi, 1960: 360, figs 40d, 48e, 70-75; Kensley, 1968: 302, 1971: 273, fig. 1, 1972: 12, figs 4c, 6a; Roberts & Pequegnat, 1970: 36, figs 3-2B, 3-3C; Aizawa, 1974: 22, fig. 14; Crosnier, 1978: 34, figs 15a, 18a-b, 1994a: 352; Griffiths & Brandt, 1983: 179; Hayashi, 1984b: 140, fig. 66a-b; Kensley et al., 1987: 276. Gennadas parvus Bate, 1881: 192 [part], 1888; 340 [part], pl. 59; Wood-Mason & Alcock, 1891a: 189 [part], 1891b: 286 [part]; Alcock, 1901: 46 [part]. Gennadas elegans Lenz & Strunck, 1914: 310 [part]. Amalopenaeus Alcocki Balss, 1927: 266, fig. 30. Amalopenaeus Bouveri Balss, 1927: 267. MATERIAL, AM P32892, 33°28’S 152°34’E, 641m, ¢, 8.5 mm, 72, 6.5-8.8mm (49, 6.5-7.2mm immature); AMP32895, 33°20’S 152°32’E, 366m, 2d, 8.0mm, &, 8.2mm. DESCRIPTION. Antennal angle and infra- antennal angles acute, but blunt. Petasma with external lobe divided into 2 broad, bluntly pointed lobules; median lobe broad with wide distal notch, the 2 lobules acute and inwardly A MEMOIRS OF THE QUEENSLAND MUSEUM hooked; inner lobe much shorter than the median lobules, with numerous cincinnuli; accessory lobe large, leaf-shaped. Thelycal plate on thoracic somite 8 with slender processes on its anterolateral corners; the shield-shaped plate on somite 7 attached anteriorly to the plate of the somite 6, with rounded flap opening posteriorly and a small process on either side. Colour, Uniformly red. REMARKS. The hooked lobules of the median petasmal lobe and the slender projections of the 2 8th thoracic sternite readily distinguish this species. Griffiths & Brandt (1983) recorded G bouveri at the edge or outside a warm-core eddy, where it appeared to be mesopelagic, but it was uncommon. DISTRIBUTION. E Australia 17-39°S, 250- 1988m. Known range: Indian Ocean; Indonesia; Arafura Sea; E Australia; Philippines; Japan; northwest Pacific; New Caledonia; Wallis and Futuna Islands, Eastern Pacific, Atlantic Ocean, 0-1115m, mostly 500-650m. Gennadas capensis Calman, 1925 (Fig. 15A-C) Gennadas capensis Calman, 1925: 5, pl. 1, figs 1-2; Burkenroad, 1936: 67, figs 51, 53; Anderson & Lindner, 1945; 292; Barnard, 1950: 630, figs 118e-f; Roberts & Pequegnat, 1970: 34, fig. 3-2A; Kensley, 1971: 277, fig. 3; Abbes & Casanova, 1973: 268, fig. 5; Crosnier, 1978: 36, fig. 18c, 1985: 860, 1994b; Hayashi, 1984b: 141, fig. 66c-d; Kensley et al., 1971: 277, fig. 3, 1987: 277. MATERIAL. AM P32880, 33°19’S 152°25’E, 640m, 5d, 9.0-10.5mm, 72, 8.0-9.5mm; AMP32882, 33°28’S 152°34’E, 641m, 5d, 10.2-13.2mm, 32, 9.1-12.0mm. DIAGNOSIS. Antennal and infra-antennal angles acute. Lobes of the petasma not divided FIG, 14. Gennadas bouvieri Kemp, 1909 AM P32895, 33°20’S 152°32’E, 366m. A, d, 8mm, anterior carapace; B, dorsal right half of petasma; C, 2, 8.5 mm, thelycum. (Scalebar = | mm). AUSTRALIAN ARISTEIDAE AND BENTHESICY MIDAE 431 FIG. 15. Gennadas capensis Calman, 1925 AM P32880, 33°19°S 152°25°E, 640m. A, d, 10.5mm, anterior carapace; B, dorsal right half of petasma; C, 2, 12mm, thelycum. (Scalebar = 1mm). distally, but the external lobe with a small acute process on the medial margin; median lobe truncate and shorter than the adjacent lobes; inner lobe indented distally and covered with cincinnuli; accessory lobe divided, the inner lobule club-shaped, the outer truncate. Thelycal plate of thoracic somite 7 a prominent W-shape, median apex a rounded concave process; coxa of the pereopod 5 expanded and bilobed; coxa of the fourth with a slender elongate process; coxa of the third bluntly lobed; pereopod 3 with a pair of concave spoon-shaped processes posteriorly directed, meeting on midline. Colour. Uniformly red. REMARKS. This is one of the larger Gennadas (up to CL 13.2mm). A d 7.5 mm CL was immature. G. capensis does not appear to be very common in Australian seas. DISTRIBUTION. E Australia 22-39°S, (-1988m, probably mesopelagic. Known range: W Indian Ocean; SE Australia; New Caledonia, Wallis and Futuna Islands, E Pacific, Atlantic Ocean, 0-2000m. Gennadas gilchristi Calman, 1925 (Fig.16 A-C) Gennadas gilchristi Calman, 1925; 6, pl. 1, figs 3, 4; Burkenroad, 1936: 66, fig. 58; Barnard, 1950: 633, fig. 118g,h; Kensley, 1968: 301, 1971: 280, fig. 6; Griffith & Brandt, 1983: 179; Iwasaki & Nemoto, 1987: 5: Kensley et al., 1987: 277. MATERIAL. AMP52815, 34°S 152°E, 950m, 14d, 4.5-7.3 mm, 62, 5-7.3mm DIAGNOSIS. Antennal and infra-antennal angles produced, apically rounded. Petasma with external lobe acute and a smaller lobule at its outer base; median lobe with 2 diverging slender lobules; internal lobe acute; accessory lobe broadly rounded (Fig. 16B). Thelycum with a pair of anterior projections on thoracic 7 thelycal plate; a visible pair of circular seminal receptacles usually containing spermatophores just anterior to it; thoracic 8 thelycal plate apple-shaped, anterior margin extending over thoracic plate 7. Coxa of pereopod 3 expanded, bilobed, more prominent in 2. (Fig.16C) Colour. Uniformly red. REMARKS. The distinctive dark circular spermatophores, red in preserved material, make 2 G gilchristi readily identifiable; the petasma is also distinctive. Kensley et al. (1987) noted that this is the commonest species in NSW collections that they examined. Collections in the Museum of Victoria, made with mid-water trawls off SE Australia confirm this. In some cases they seemed to be associated with trawls consisting largely of salps, being the only decapod crustaceans present. Griffiths & Brandt (1983) recorded G gilchristi as the most abundant Gennadas species associated with the Tasman Sea warm-core eddies that they investigated. G gilchristi was mostly on or outside the edge of the eddy at around 250m depth and thus appears to be mesopelagic. Griffiths observed (F. B. Griffiths, pers. com.) that the gut contents were commonly green algae, and suggested that the Gennadas had been feeding on salp faeces. DISTRIBUTION. SE Australia, 33°-42°S, 200- 1200m, S of Australia, to 45°S 115°-150°E, 0-1050m. Known range: off Cape Penisula and W coast of S. Africa, Argulhas Basin, S$ Indian Ocean, SE Australia, New Caledonia, 0-3400m. MEMOIRS OF THE QUEENSLAND MUSEUM ee | FIG. 16. Gennadas gilchristi Calman, 1925 AM P52815, 34°5’S 151°55’E, 950m, A, 36, 7.3mm, anterior carapace; B, dorsal right half of petasma; C, 2, 6mm, thelycum. (Scalebar = 1mm). A mesopelagic species, preferred depth range appears to be 200-700m, mostly S of 30°S. Gennadas incertus (Balss, 1927) (Fig.17A-C) Amalopenaeus incertus Balss, 1927; 265, figs 24-29. Gennadas incerius Burkenroad, 1936: 66; Anderson & Lindner, 1945: 294; Tirmizi, 1960: 364, fig. 40e, 48f, 76-80; Pearcy & Forss, 1966: 1137; Kensley, 1971: 284, fig. 7, 1972: 12, 14, fig. 41, 5j; Aizawa, 1974: 23, 44, figs 15, 29; Crosnier, 1978: 37, fig. 15b; Griffiths & Brandt, 1983: 179; Hayashi, 1984b: 141, fig. 66e; Kensley et al., 1987: 278. MATERIAL, AM P35740, 33°20’s 152°32’E, 550m, 2¢, 7.2, 7.7mm, 2, 8.0mm; AMP35741, 33°19’S 152°25’E, 640m, 3, 7.4mm DIAGNOSIS. Antennal angle blunt, infra-antennal angle quadrangular. External lobe of the petasma divided into two long tapering lobules, projecting well beyond the other lobes; median lobe divided into 2 rounded lobules, the outer relatively narrow, the inner lobule broad; inner lobe not divided, with cincinnuli along its inner margin; accessory lobe simple and reaching as far as the inner and median lobes. Thelycal plate on thoracic somite 8 indented anteriorly, and overlapping sternite 7 anteriorly; sternite 7 thelycal plate with acute anterolateral corners, posterolateral corners with rounded processes extended anteriorly; somite 6 with large concave plate. Colour. Body generally red, paling anteriorly and posteriorly; thoracic appendages 2-5 dark red with purplish spots; pleopods pale red with dark purple spots. REMARKS. Kensley (1971) described and a small posterior notch on the sternite of thoracic somite 8, which he used in his key to distinguish G. incertus. Liu & Zhong (1986, pl. 24, fig. 5) also showed a small indentation, but no trace of a notch could be found on @ ? from SE Australia. Crosnier (1978, fig.15b) did not show any posterior indentation. It therefore does not appear to be areliable distinguishing feature. Griffiths & Brandt (1983) record this species associated with the edge of a warm-core eddy in the Tasman Sea, probably mesopelagic, but it was not common. DISTRIBUTION. E Australia, 17-34°S, 220- 1406m. Known range: Indian Ocean, Australia, New Caledonia, Japan, NW & E Pacific, SE Atlantic, 100-1406m (diurnal vertical migration). Gennadas kempi Stebbing, 1914 (Fig.18A-C) Gennadas kempi Stebbing, 1914: 283, pl. 27; Calman, 1925: 4; Burkenroad, 1936: 68, figs 52, 54; Barnard, 1950: 630, figs 118a-d; Kensley, 1971: 285, fig. 8; Iwasaki & Nemoto, 1987: 6; Kensley et al., 1987: 278. MATERIAL. AMP32914, 33°31’S 152°20’E, 550m, 3, 9.1mm; P32915, 33°19’S 152°25’E, 640m, 22, 9.2, 10.1mm. DIAGNOSIS. Antennal and infra-antennal angles rounded. Petasma with external, median and internal lobes undivided; external lobe acute; median lobe broad and truncate; internal lobe AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE —— | a | FIG, 17, Gennadas incertus Balss, 1927, AM P35740, 33°20’S 152°32°E, 366m. A, d, 7.7mm, anterior carapace; B, dorsal right half of petasma; C, 2, 8mm, thelycum. (Scalebar = 1mm). truncate, with rounded process bearing cincinnuli; accessory lobe a simple rounded flap not reaching as far as the internal lobe. Thelycal plate on the thoracic somite 8 hexagonal; that on sternite 7 subrectangular, with concave anterior edge, and that on sternite 7 triangular with apex directed anteriorly. DISTRIBUTION. S Australia 32°-42°S, 550- 640m, Antarctic Ocean (8 of Australia, between 115°-150°E) down to 61°27'S, 0-1050m, probably mesopelagic. Known range: SE Indian Ocean, SE Australia, Antarctic Ocean, New Caledonia, SE Atlantic Ocean, 0-3400m; only recorded S of 32°S. A B PrMer caren Pann re drameeme: — thelycum. (Scale bar = 1mm). FIG. 18 Gennadas kempi Stebbing, 1914, AMP32915, carapace; B, AMP32914, 33°31°S 152°20"E 550m, 2. Gennadas propinquus Rathbun, 1906 (Fig.]9A-C) Gennadas propinguus Rathbun, 1906: 907, fig. 6la, b: Burkenroad, 1936:66, 83; Anderson & Lindner, 1945; 295; Pearey & Forss, 1966: 1137: Kensley, 1969: 167. fig. 9; Aizawa. 1974: 24, figs 17-19: Crosnier, 1978: 38, figs l6b, 18d-e, 1989: 41, 1994b: 369; Hayashi, 1984a: 142, fig. 66h; Liu & Zhong, 1986: 60, lig. 25; Kensley et al. 1987: 278, Gennadas clavicarpus de Man, 1907; 144 [part], 1911: 19 [part], 1913: pl. 1, figs 3, 3a-c, pl. 2. fig. 3h.j, 1922: 3, pl. 1, fig. 1; Balss, 1927; Boone, 1930: 129, pl. 45 (9° 2 only); Tirmizi, 1960: 353, figs 40c, 48c, 38-66; Kensley, 197]; 278, fig. 4; Griffiths & Brandt, 1983: 179. Gennadas alvocki Kemp, 1910: 174, pl. 13; 1913: 62, pl. 7 (¥ only). Gennadas seutatus Kemp, 1910: 178, pl. 13. figs 9, 10. Germaday seututus indicus Kemp, 1913; 62 (dd only), 33°19°S 152°25°E, 640m, A, 3, 10,1mm, anterior 9.1mm, dorsal right half of petasma; C, 2, 8mm, 434 FIG. 19. Gennadas propinguus Rathbun, 1906 AM P32904, 33°20°S 152°32° carapace; B,, dorsal right hall! of petasma; C, AM P32903, 33°177S 152°3 (Scealebar — mm), dmalopenqeus scululus indicus Balss, 1927; 259, fig. 13. Amelapendeus clayicarpus Balss, 1927; 267. MATERIAL. AMP32903. 33°17°S 152°32°E, 92m, 3, 6.2mm, 52, 5.1-8.2mm: AMP32904, 33°28 152°32°K, 366m, 52, 5.4-9.5mm. DIAGNOSIS, Antennal and infra-antennal angles blunt. Petasma with the 3 lobes divided apically; the outer lobe with outer lobule acute and slender, inner lobule rounded; median lobe with both lobules prominent, apically acute; in- ner lobe with both lobules rounded, with cinein- nuli; accessory lobe asimple Hap. Thelycum with ovate plate on the sternum of thoracje somite 8, slightly indented anteriorly and posteriorly; thoracic somite 7 with a w-shaped plate, closely applied to the sub-triangular plate of the somite 6; the latter with indented anterior apex, openings of seminal receptacles at its base. Colour. Body generally red, paling towards the anterior and posterior regions; thoracic ap- pendages 2-5 dark red with purplish spots; pleopods pale red with dark purple spots at the base of cach. REMARKS. Griffiths & Brandt (1983) record this species (as G clavicurpus) at the edge and inside a warm core eddy in the Tasman Sea, but it was not abundant. DISTRIBUTION. E Australia, 17°-42°S, 150-950m. Known range: Indian Ocean, Arabian Sea, Gulf of Aden, Philippines, China Sea, NW Pacific, SE Australia, New Caledonia, Wallis and Futuna Islands, E Pacific, SE Atlantic. 0-1200 m; mesopelayic migrating from deeper water at night to 100-200m. MEMOIRS OF THE QUEENSLAND MUSEUM E, 366m. A. 4, 9.5mm, anterior 1*F, 92m, &, 825mm, thelycum, Gennadas scutatus Bouvier, 1906 (Fig. 20A-C) Gennadas scutatis Bouvier, 1906b: 748; 1906d; 3, 9-12 , figs 8, 13, 1908: 42, pl. 8, figs 1-16; Milne Edwards & Bouvier, 1909: 193, figs 10-12; Kemp, 1909: 27, 727_ pl. 75, fig. 2, 1913; 61 (¥ only); de Man, 1911; 6, 15; Lenz & Strunck, 1914: 310, 341; Calman, 1925: 4: Burkenroad, 1936: 83. fig, 34, 1938: 59: Anderson & Lindner, 1943: 295; Barnard, 1950: 634, fig. | [8u-p; Tirmizi, 1960: 342, 346, 357, 358, figs 40¢, 4Xd, A7-H9; Belloc, 1961: 8; Crosnier & Forest, 1969; 349; 1973; 281, Ips 94u, 95a, b, 1973: Roberts & Pequeznat, 1970: 39. fig, 3-1 A; Kensley, 1971; 28K. fig. 10, 1972: 12, 16. fig, 4c Op; Aizawa, 1974: 26, Fig. 20: Crosnier LOTR: 43, fiz. L7a, 19845 20, }994b: 369; Griffiths & Brandt 1983: 179; Kensley et al., L987; 279, Cermudas chivicaypus de Man, 1907: 145 [path POT: 19 [part], 1913: pl. 1, fig, 3f, @ pl. 2, fig. 3k; Boone, 1930; 129 (4 only). Imalopenaeus sculatus Balss, 1927: 25%, fies 11, 12. MATERIAL. AMP32913 33°17°S. 152°317E, 92m. cf. 6.4mm. DIAGNOSIS. Antennal and infra-antennal angles acute. Pelasma with external lobe divided into 2 short rounded lobules: median lobe with slender outer Jobule, the inner lobule broad with a small accessory lobule on ifs euler margin: in- ternal lobe divided into 2 rounded inward-tacing lobules, covered with cincinnuli; accessory lobe a simple triangular Map. Thelycal plate on thoracic somite 8 with a wide rounded anterior flap extending to cover about half sternite 6; sternite @ thelycal plate iiangular, with paired scininal receptacles at its base; sternite plate 7 a wide, fattened triangle; coxae of pereopod 4 with 4-5 inwardly-directed stout setae- Colour. Unitormly red. AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE as88 AAARARA AAR A ACN. FIG. 20. Gennadas scutatus Bouvier, 1906, AMP32913, 33°17’°S 152°31°E, 92m. A, 2, 6.5 mm, anterior carapace; B, 3, 5.3mm, dorsal right half of petasma. C, 2, 6.5mm, thelycum. (Scale bar = Imm). REMARKS. Griffiths & Brandt (1983) recorded G scutatus on the edge of a warm-core eddy in the Tasman Sea, but it was uncommon. DISTRIBUTION. E Australia 17-38°S, 92-1192m. Range: throughout Indo-West Pacific, E Pacific and Atlantic Oceans, 0-3400m, mesopelagic. Gennadas tinayrei Bouvier, 1906 (Fig. 21A-C) Gennadas Tinayrei Bouvier, 1906d: 10, figs 2-4, 14; 1908: 48, pl. 1, fig. 4, pl. 10; 1922: 10; Burkenroad, 1936: 73, fig. 56. Gennadas tinayrei Lenz & Strunck, 1914: 313; Sund, 1920: 29; Anderson & Lindner, 1945: 293; Tirmizi, 1960: 367, figs 40f, 81-83; Kensley, 1971: 290, fig. 12, 1972: 12, figs 4b, 5c; Abbes & Casanova, 1973: 67; Aizawa, 1974: 27, fig. 21; Crosnier, 1978: 44, figs!7b, 19d; Hayashi, 1984b: 143, fig. 66k, 1; Kensley et al., 1987: 279. Amalopenaeus tinavrei Sund, 1920: 29. A ARPAANREROAEAREA BARR ADORE AA, REM BRAD Ane Amalopenaeus Tinayrei Balss, 1927: 252, fig. 2. MATERIAL. AMP32889, 33°31°S 152°20°E, 550m, 6, 6.2mm; AMP52827, 34°0S’S 151°55’E, 950m, 3, 6.3mm; 39, 7.1, 7.2, 7.3. DIAGNOSIS. Antennal and infra-antennal angles acute. Petasma with external lobe slightly indented, edged with minute teeth and with a small lobule at its base; middle lobe broad and slightly convex; inner lobe with 2 rounded projections covered with cincinnuli; accessory lobe flattened and rounded in outline. Thelycal plate on thoracic somite 8 small and subrectangular; plate on the somite 7 a wide triangle with 2 rounded processes on either side of its apex, and just anterior to these a further 2 smaller rounded processes on the somite 6; FIG. 21. Gennadas tinayrei Bouvier, 1906, AM P52827, 34°5’S, 151°55°E, 950m. A, 2, 7.3 mm, anterior carapace. B, ¢, 6.3mm, dorsal right half of petasma. C, 2, 7.3 mm, thelycum. (Scale bar = 1mm). 436 sternite of the somite 5 with a posteriorly directed tongue-like process. Colour. Uniformly red. DISTRIBUTION. SE Australia 33-42°S, 92-950 m. Known range: throughout the Indian Ocean, SE Australia, Japan, NW Pacific, Atlantic Ocean, 92-950, probably mesopelagic and uncommon. ZOOGEOGRAPHY OF THE ARISTEIDAE AND BENTHESICYMIDAE As the Aristeidae and Benthesicymidae inhabit a zone from 0-5000m, to some extent their apparent distribution reflects the collecting methods that have been used in different parts of the world by various expeditions or fishery investigations. Often only benthic collectors have been used (trawls, sledges or dredges) and pelagic or even epibenthic species have been missed or collected adventitiously during shooting or retrieval of the net. Mid-water trawls have often shown that some species hitherto considered rare, are common. The ideal approach is to use various collecting devices as described by Kikuchi & Nemoto (1991), These included an opening-closing net, a mid-water trawl, baited traps and an Agassiz-type trawl. The first 3 were the most successful in collecting Benthesicymus and enabled Kikuchi & Nemoto to distinguish bathy- and meso-pelagic species from predom- inantly benthic species. It is also of interest that of the 8 Benthesicymus species recorded, the trawl collected only B. investigatoris and that only once! The depth range for most of the Aristeidae is within a 200-2000m zone, with a few reaching 5000m or more. All species of Aristeidae have well-developed pleopods and like members of other penaeoid families are probably capable of swimming appreciable distances and may swim up and down in the water column. For example, Crosnier (1978) produced evidence that Aristaeomorpha foliacea undergoes a vertical diurnal migration. The depth range of other species suggests that this may not be an isolated case (e.g. Hepomadus tener, 765-5400m), As shown by the increasing use of mid-water trawls over the last 40 years, the Benthesi- cymidae are largely pelagic and even those classed as benthic probably spend a large part of their time in the water column, All are small, possess thin cuticles, have well-developed pleopods and probably have low density muscle and other tissues, Although some of the species, subsequently shown to be pelagic, have been MEMOIRS OF THE QUEENSLAND MUSEUM collected in bottom trawls, it must have been while the trawl was being shot or retrieved. Pelagic (bathy-, meso- and fully pelagic) genera are Benthonectes, Gennadas and probably all species of Bentheogennema, Benthesicymus brevirostris and B. carinatus are also pelagic (Kikuchi & Nemoto, 1991), Thus the depth range recorded for these species may range from upper levels to 5000m. Longitudinal distribution. Since the Aristeidae are predominantly benthic data collected from bottom trawls probably give a realistic picture of their distribution. In contrast, unless a variety of collecting methods have been used in a given area, the apparent distribution of the Benthesicymidae may not give a true picture. This is certainly the case for Australian seas. Intensive deeper water trawling and later mid-water trawling off SE Australia has collected 8 species of Gennadas, some of which appear to be common or even abundant. None of these species had been collected in other parts of Australia until the Australian Institute of Marine Science ‘Cidaris 1’ expedition, which collected 4 Gennadas species, using various types of gear. As noted previously, of the species of Benthesicymus, only B. investigatoris, B. urinator and a possible third species have been collected from Australian seas, but this is unlikely to be the true picture as Kikuchi & Nemoto (1991) recorded 8 species from the NW Pacific and Crosnier (1978, 1985) 8 from the western and central Indian Ocean, 5 of which were common to those of Kikuchi & Nemoto. Given the provisos regarding collecting methods, it is obvious that the Aristeidae and Benthesicymidae are generally more widely distributed than most of the remaining Penaeoidea. Of 25 species of Aristeidae 8 have been recorded in both the Indo-West Pacific and Atlantic Oceans, with Hemipenaeus carpenteri, H. spinidorsalis, Plesiopenaeus armatus in the E Pacific as well. Of the remaining 17 species, 7 range through the Indo-West Pacific, and 10 appear to be more localised. Among the 37 species of Benthesicymidae, 14 are common to both the Indo- West Pacific and Atlantic Oceans, with 12 of these common to the E Pacific as well; an additional 3 are found through the Indo-West Pacific, and 20 have a more restricted range. A ‘restricted range’ may be due to misident- ification, limited or inappropriate collection methods, or rarity of a species. Latitudinal distribution. The pelagic species of the Benthesicymidae appear to have the widest AUSTRALIAN ARISTEIDAE AND BENTHESICYMIDAE latitudinal range. In the N Pacific the pelagic Bentheogennema borealis has been recorded in the S Bering Sea (around 57°N) where it is a relatively abundant mesopelagic inhabitant (Butler, 1980). Bentheogennema burkenroadi has been collected at 52°N off the coast of British Columbia and again is relatively abundant. No other penaeoids have been recorded in this region at such high latitudes and 45° N appears to be the approximate limit for other pelagic species. In the NW Atlantic G elegans has been collected up to 57°N off the Labrador shelf, while G valens has been recorded at 51°N in the northeastern Atlantic off Ireland (Squires, 1990). No other benthesicymid species have been collected N of about 44°N (Squires, 1990). In the Southern Hemisphere the range is comparable with that of the N Pacific. G kempi was collected S of Aust- ralia as far as 61°27’S in the Antarctic Ocean (Iwasaki & Nemoto, 1987). Various midwater collecting devices were used with oblique tows from 700-1000m to surface. G kempi was collected at 10 stations, from 2-17 per station and thus it appears to be fairly abundant even at the highest latitudes of its range. The same cruise collected G gilchristi around the Sub-Tropical Convergence at 45°S, but none beyond this latitude. It seems, therefore, that the majority of pelagic Benthesicymidae are restricted to a zone between 45°N and 45°S, with a few species adapted to the lower temperatures of higher latitudes. Of the benthic Aristeidae and Benthesicymidae the range appears to be a little less, mostly 40°N - 40°S. The wide longitudinal distribution of many Aristeidae and Benthesicymidae may be explained by lack of geographical barriers at depths of 1000m and more. The reason that the majority are confined to 40°N - 40°S is less obvious. It has been suggested for the Solenoceridae that this is due to the larvae in the upper water column being adapted to higher water temperatures (Dall, 1999). Thus in the Southern Hemisphere as the Subtropical Convergence zone is approached around 40°S, the upper water temperatures begin to fall sharply and so limits the southern distribution of the Solenoceridae. The same mechanism appears to apply to the Aristeidae and Benthesicymidae and perhaps indicates a tropical origin for these families. However, not all species range through tropical or subtropical latitudes. One aristeid species appears to be restricted to higher southern latitudes: Austropenaeus nitidus. It inhabits a zone 26°- 40°S from the S$ Atlantic 437 Ocean, South Africa, across the Indian Ocean (Amsterdam and St. Paul Islands) and across S Australia, where it appears to be common. It is likely that its longitudinal range is even wider. The mesopelagic G gilchristiand G. kempihavea N limit similar to that recorded for Austropenaeus nitidus (around 30°S), except that one G gilchristi has been collected at 21°S by a midwater trawl off E New Caledonia. However, the abundance of G gilchristi south of 33°S in E Australian waters suggests that this may be a stray from higher latitudes. In conclusion, it appears that deep water Penaeoidea are less subject to the geographical barriers which influence the distribution of most of those penaeoid species that inhabit the continental shelf and its outer edges. This particularly applies to the pelagic Benthesicy- midae many of which have a wide longitudinal range with temperature as the major north and south latitudinal limiting factor. Significantly in this regard, 2 species of pelagic Funchalia (Penaeidae) are also cosmopolitan. ACKNOWLEDGEMENTS This work was made possible by a grant from the Australian Biological Resources Survey. My sincere thanks are due to the releyant Curators and/or Collections Managers of the following museums which | visited and which loaned material: The Australian Museum, Sydney, Penny Barents; The Museum of Victoria, Gary Poore and Vicky Barmby; The Museum of Tasmania, Hobart, Roger Buttermore; The ISR Munro Fish Collection, CSIRO Marine Laboratories, Hobart, Alistair Graham; The Queensland Museum, Brisbane, Peter Davie and John Short; The Museum of Tropical Queens- land, Peter Arnold. Gavin Dally, Collections Manager of the Museum of Northern Territory also sent specimens on loan. I thank the Chief, Division of Marine Research, CSIRO for providing facilities at the CSIRO Marine Laboratories, Cleveland. | also thank Alain Crosnier, Muséum National D’ Histoire Naturelle and Ken-Ichi Hayashi, Shimonoseki University of Fisheries for critically reading the manuscript and making a number of helpful suggestions. LITERATURE CITED ABBES, R. & CASANOVA, J. P. 1973. Crustacés Décapodes pélagiques Penaeidea et Caridea récoltés par la “Thalassa’dans 1’ Atlantique eurafricain. Revue des Travaux de I’Institut des Péches maritimes 37: 257-290. 438 AIZAWA, Y. 1974. Ecological studies of micronekton shrimps (Crustacea, Decapoda) in the Western North Pacific, Bulletin of the Ocean Research Institute of the University of Tokyo 6: 1-84. ALCOCK, A. 1899. 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Bulletin of the Biogeagraphical Society of Japan 41(7): 51-60. 1991, Deep-sea shrimps of ithe genus Benthesicvetus (Decapoda: Dendrobranichiata) from the western North Pacific. Journal of’ Crustacean Biology 111): 64-89, KOMAI, T. 1993. Deep-sea shrimps of the family Anstcidae (Decapoda: Dendtobranchiata) from northern Japan, with the description of a new species of the genus dAristeus. Crustacean Research 22: 2)-34, KUBO, 1.1949. Studies on the penagids of Japanese and its adjacent waters. J. Tokyo Coll, Fish. 36: 1-467. LENZ, H. & STRUNCK, K, 1914. Die Dekapoden der Deuischen Stidpolar-Expedition 190}-1903, 1. Brachyuren und Macruren mit Aussehluss der Sergestiden. Wissenschafiliche Ergebnisse der Deutsche Stidpolar-Expedition. 15, Zoologie 7: 259-345. LIU, LY, & ZHONG. Z, 1986, Penaeoid shrimps of the South China Sea, (Agricultural Publishing House; Beijing). MIERS, E.J, 1878. Notes on the Penaeidae in the collection of the British Museum, with MEMOIRS OF THE QUEENSLAND MUSEUM descriptions of same new species. Proceedings of the Zoological Society of Londun 1878: 289-310. MILNE-EDWARDS, H. 1837. Histoire naturelle des Crustacés, comprenant |’anatomie. la physiologie et la classification de ces animaux 2: 1-532, MILNE-EDWARDS, A. & BOLIVIER. E.L, 1909. Les Péneides et Sténopides, Memoirs of Ihe Museum of comparative Zoology at Harvard College 27(3): 177-274. OKADA, Y., SARAMOTO, I., AMANO, R. & TOMINAGA, Y. 1966, Preliminary report of the benthic biological survey in Suruga Bay. Journal of the Faculty of Oceanography of Tokai University 1966(1): 135-155, PEARCY, W.G. & FORSS, C.A. 1966, Depth distribution of oceanic shrimps (Decapoda: Nalantia) off Oregon. Journal of the Visheries Research Board of Canada 23(8): 1135-1143. PEQUEGNAT, W.E. & ROBERTS, T.W, L971. Devapod shrimps of the family Penagidae. 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Peneides and stenopides from the “Michael Sars” North Atlantic Deep-Sea 44] Expedition. Report of the Michael Sars North Atlantic Deep-Sea Expedition 3(2): 1-36. TIRMIZI, N.M. 1960. Crustacea: Penaeidae. Part II. Series Benthesicymae. With a note by I. Gordon. John Murray Exped. 1933-34, scient. Rep. 10: 319-383. WASMER, R.A. 1972. New records for four deep-sea shrimps from the Northeastern Pacific. Pacific Science 26(3):259-263. WOOD-MASON, J. 1891. Phylum Appendiculata. Branch Arthropoda. Class Crustacea. Annals and Magazine of Natural History (series 6)8: 269-286. 1892. Illustrations of the zoology of the Royal Indian Marine Survey Steamer “Investigator”. Crustacea, part 1, pls 1-5. WOOD-MASON, J. & ALCOCK, A. 1891a. Natural History Notes from H.M. Indian Marine Survey Steamer Investigator, Commander R.F. Hoskyn, R.N. commanding. No. 21. Note on the results of last season’s deep-sea dredging. Annals and Magazine of Natural History (series 6)7: 186-202. 1891b. Natural History Notes from H.M. Indian Marine Survey Steamer Investigator, Commander R.F. Hoskyn, R.N. commanding. Ser. II, No. 1. On the results of deep-sea dredging during the season 1890-91. Phylum Appendiculata. Annals and Magazine of Natural History (series 6)8: 269-286. 449 FIRST RECORD OF THE LARGE-TOOTH COOKIE- CUTTER SHARK J/SISTIUS PLUTODUS FROM AUSTRALIAN WATERS. Memoirs of the Queensland Museum 46(2): 442, 2001:- The largetooth cookiecutter shark Tsistius plutodus Garrick & Springer (1964), is an epipelagic and possibly bathypelagic shark known only from the Gulf of Mexico, off Alabama, USA and the western North Pacific, off Okinawa, Japan (Compagno, 1984). /sistius contains /.plutodus and |. brasiliensis (Quoy & Gaimard, 1824). Both species have small cigar-shaped bodies (maximum size about 42cm TL and 50cm TL respectively), a conical snout and two low, spineless dorsal fins, The most obvious differences between the two species are the number of tooth rows in the lower jaw and the distance between the two dorsal fins. In /.p/utodus, there are 19 tooth rows in the lower jaw and the inter-dorsal distance is subequal to the base of the first dorsal fin (D1). In /Arasiliensis there are 25-31 tooth rows, and the inter-dorsal distance is over twice the D1 base (Compagno, 1984). A distinctive dark collar-like marking is found around the branchial region of /. brasiliensis. A 363mm & Jplutodus was recently identified in the Ichthyology Collection of the Australian Museum (AMS 1.28924-001). The specimen was collected by the FV Teresa in 1988 off Newcastle, New South Wales (approximately 33°S 152°E). Full collection details are not available, but discussions with K. Bollinger, the vessel owner, indicate that the shark was probably caught at night as prawn bycatch in an otter trawl at a depth of about 100m. The specimen has 19 tooth rows in the lower jaw and lacks aclearly defined collar around the branchial region. The snout is very short (14.2mm) and the eyes are anteriorly placed. Compagno (1984) suggested that the position of the largetooth cookiecutter’s eyes allows for binocular vision, which aids in precisely locating its victims. The AMS specimen of /.p/utodus was fixed in formalin in 1988 and has been preserved since in 70% ethanol. The pectoral, anal and caudal fins are damaged and the tissue around the snout is distorted. This has affected accurate MEMOIRS OF THE QUEENSLAND MUSEUM measurement of several of the diagnostic characters, however all characters are consistent with the descriptions in Garrick & Springer (1964) and Compagno (1984). The specimen has the following morphometrics (in mm): TL 363, HL 68.6, snout length 14.2, eye length (including posterior notch) 15.0, predorsal length 233, body depth 36.7, DI base 17.5, D1] height 10.6, D2 base 18.2, D2 height 13.2, interdorsal space 22.5, D2 insertion to upper caudal origin 26.2. This major range extension is a new record for Australia and the Southern Hemisphere. The continental shelf in the region of the collection locality is approximately 42km wide. The fish was captured approximately 24km landward of the shelf break (200m isobath). Species of /sistiusare believed to move vertically in the water column from deeper waters. If this is the case then the current fish not only ascended from deep water but also traversed over 20km in shelf (< 200m) waters. Acknowledgements Dr J.D. Stevens (CSIRO Fisheries, Hobart) is acknowledged for confirming the identification and for advice on the manuscript. C. Bento photographed the fish, and S. Morris retrieved the specimen from the collection and did an initial examination. K. Bollinger and the crew of the FV’ Teresa, are thanked for capturing the fish and sending it to the Australian Museum. References COMPAGNO, L.J.V. 1984. FAO species catalogue. Vol.4, Sharks of the world. An annotated and illustrated catalogue of shark species know to date. Part I - Hexanchiformes to Lamniformes: viii, 1-250. FAO Fisheries Synopsis 125: 1-249. LAST, P.R & STEVENS, J.D. 1994. Sharks and rays of Australia. (CSIRO; Canberra). GARRICK, J.A.F & SPRINGER, 8. 1964. /sistius plutodus, a new squaloid shark from the Gulf of Mexico. Copeia 4: 678-682. Mark A. McGrouther, Australian Museum, 6 College Street, Sydney, 2000; 12 April, 2000. FIG, 1. Lsistius plutodus (AMS 1,28924-001). A REVISION OF PROCAMBRIDGEA FORSTER & WILTON, (ARANEAE: AMAUROBIOIDEA: STIPHIDHDAE) VALERIE TODD DAVIES AND CHRISTINE LAMBKIN Davies, V.E. & Lambkin, C. 2001 06 30: A revision of Procambridgea Forster & Wilton, (Araneae: Amaurobioidea: Stiphidiidae). Memoirs of the Queensland Museum 46(2): 443-459. Brisbane. ISSN 0079-8835. Procambridgea rainbowi Forster & Wilton and P. cavernicola Forster & Wilton have been redescribed and 10 new species described. These are P. gravi, kioloa, otwayensis, ourimbah, hunti, carrai, monteithi, lamington, hilleri and montana. The species have been collected from sites in SE Queensland, northern and eastern New South Wales and from the Otway Range in Victoria. Cladistic analysis shows that they form a monophyletic group and infers that Procambridgea is more closely related to the Stiphidiidae than any other group. 4 Procambridgea, new species, Stiphidiidae, Amaurobioidea, cladistics. Valerie E. Davies, Honorary Associate, Queensland Museum, PO Box 3300, South Brisbane 4101; Christine L. Lambkin, Department of Zoology & Entomology, University of Queensland, St Lucia 4072, Australia; Present address: CSIRO, Entomology, PO Box 1700, Canberra 2601, Australia; 28 March 2001. Procambridgea, an Australian genus of cribellate spiders was described by Forster & Wilton (1973: 134) and placed in the Stiphidiidae along with several ecribellate spiders from New Zealand. Comprehensive descriptions of the type species, P. rainbowi and P. cavernicola were given. Examination of further Australian species has shown the importance of several characters which we illustrate for these two species. Procambridgea is a small spider, seldom exceeding 5.0 in length; it has the nondescript pattern of most amaurobioids (Lehtinen, 1967: figs 42-67). Webs are often in the hollows of fallen logs in the form ofa small suspended sheet, on the underside of which the spider sits. In the same way that Stiphidion facetum has been introduced to New Zealand from Australia, recently Procambridgea has been found in Auckland, New Zealand. MATERIAL AND METHODS Spiders from rainforest areas in SE Queens- land, from cave and forest areas in New South Wales (NSW) and 4 mature specimens from the Otway Ranges in Victoria were examined. Most were collected in pitfall (PF) traps. Notation of spines follows Platnick & Shadab (1975). Illustrations were drawn with the aid of a camera lucida; the left male palp is illustrated. The epigyna appear to be very conservative and vary little between species thus there is an emphasis on 3 characters and d ¢ only are used in the Key. Cladistic methods are given under heading ‘Relationships of Procambridgea’. ABBREVIATIONS. Museums: AM, Australian Museum, Sydney; MNZ, Museum of New Zealand, Te Papa Tongarewa, Wellington; QM, Queensland Museum, Brisbane; WAM, Western Australian Museum, Perth. Collectors: CH, C. Horseman; GBM, G.B. Monteith; MRG, M.R. Gray; RJR, R.J. Raven; SRM, S.R. Monteith; VED, V.E. Davies. Location data: SF, State Forest; NP National Park. Anatomical: AL, abdomen length; ALE, anterior lateral eyes; ALS, anterior lateral spinnerets; AME, anterior median eyes; APOPH, apophysis; AW, abdomen width; C, conductor; CB, cymbium; CAL, calamistrum; CH, cheliceral; CL, carapace length; CR, cribellum; CW, carapace width; E, embolic; EG, epigastrial groove; EPIG, epigynal; ID, insemination duct; MAP, major ampullate spigots; mAP, minor ampullate spigot; MT, metatarsal; P, patellal; PCB, paracymbial; PCR, paracribellar spigots; PE, parembolic; PLE, posterior lateral eyes; PLS, posterior lateral spinnerets; PME, posterior median eyes; PMS, posterior median spinnerets; RTA, retrolateral tibial apophysis; T, tarsal; TRICH, trichobothria. Abbreviations on illustrations are explained in the legends. SYSTEMATICS Procambridgea Forster & Wilton Procambridgea gen. nov. Forster & Wilton, 1973: 134 444 TYPE SPECIES. Procambridgea rainbowi Forster & Wilton. DIAGNOSIS. Three-clawed cribellate. Red-brown cephalothorax and legs, darker abdomen with lighter chevron pattern which is often faded or absent. AME smallest; from above posterior row of eyes straight to slightly procurved, anterior row recurved (Fig. 1A); from the front posterior row strongly procurved (Fig. 51). Cheliceral promargin with 2 large teeth and 4-5 minute teeth; retromargin with 7 small contiguous teeth decreasing in size towards the base of the fang (Fig. 1B). Prolateral filamentous seta at base of fang longer than other setae. Labium slightly longer than wide; sternum slightly longer than wide, pointed posteriorly. Legs 1423, without feathery hairs; incomplete pre-distal fracture on tarsi (Raven, in prep.). Trochanters notched. Tarsal trichobothria in single row increasing in length distally, bothrium collariform; tarsal organ slit-like, broadening distally. Cribellum with two spinning fields in female, large broad colulus in male. Small epigynum with median longitudinal ridge, anterior gonopores. Male palp with oval tegulum; course of sperm duct showing clearly. Conductor, a membranous plate often partly sclerotised with distal folds around the spiniform embolus (Fig. IF). Median apophysis reduced or absent. Cymbium with or without long post-alveolar extension; with or without bulge (paracymbium) on proximal retrolateral edge; if present bulge with or without processes. ALS largest with short conical terminal segment; two major ampullate spigots in female. PLS slender with longer terminal segment. KEY TO 6 3d PROCAMBRIDGEA SPP. 1. Palpal tibia with proximal spur (Fig. 5E). . . . monteithi Palpal tibia without proximal spur... 2... 2. 2, 2 2. Paracymbium without processes... ...-....-..3 Paracymbium with processes (Fig.50) 2... 2... 9 3. Medianapophysisabsent.......-....-.... 4 Median apophysis present(Fig. IM). ......... 5 4. RTA with small dorso-retrolateral branch (Fig. 4K) wh, ae Be, cds CRS, teres ee aiae Bie carrai RTA without small dorso-retrolateral branch. . rainbowi 5. Cymbial alveolus as short or shorter than post-alveolus. PosteriorRTA absent... 0... we ee ee 6 Cymbial alveolus longer than post-alveolus. Posterior RTA present (Fig.3E). 2.2. .....--. otwayensis 6, Conductorlargeandmembraneous . 2... 2.06. 7 Conductor small, often partly sclerotised. ... 2... 8 7. RTA with dorso-retrolateral branch, . .. . cavernicola RTA without dorso-retrolateral branch. . 2... . hunti MEMOIRS OF THE QUEENSLAND MUSEUM 8. Alveolus: post-alveolus about equal. Sperm duct with opentoop(Fig.2K) 2... 2. ea ee eee kioloa Alveolus: post-alveolus 1:3. Sperm duct with closed loop Fa et DEE AL oe Ree ELS 6 eR, Swe grayi 9. Paracymbium with retrolateral process only. Loop of spermductsimple .... 0... 2 eee ourimbah Paracymbium with retrolateral and ventral processes. Loop of sperm duct complex (Fig.50)........ 10 10. Embolic region about half length of tegulum. Sperm duct withtransverseloop... 0... ee ee 1] Embolic region about quarter length of tegulum. Sperm duct with longitudinalloop .......0.. montana 11. Ventral paracymbial process pointed. Loop of sperm duct Glosee hie ee eee oe eee lamington Ventral paracymbial process small and blunt. Loop of spermiductopen . - fondue maa EE hilleri Procambridgea rainbowi Forster & Wilton (Figs 1A-I, 8A) P. rainbowi Forster & Wilton, 1973:134, figs 403-406: Brignoli, 1981: 533 (catalogue). MATERIALS. HOLOTYPE; &, rainforest, Jenolan, Blue Mountains, NSW, Australia, 33°30°S, 150°23’E, 18 July 1970, R.R. Forster (AMKS30617), ALLOTYPE: 3, same data as holotype. (AMKS30616). OTHER MATERIAL: NSW. 3, Mt Wilson, Cathedral of Ferns area, 33°30’S, 150°23’E, PF, 14 Aug. 1978, CH (AMKS1677); 2, same locality, PF, 15 Feb. 1978, CH (AMKS1506); 2, Mt Wilson, under log, 28 Oct. 1981, CH, D. Kent (AMKS8412); 2, Mt Wilson, Cathedral of Ferns, under log in rainforest, 17 Apr. 1974, MRG (AMKS32777); 3 @, same locality, in sheet webs in logs, 26 June 1974, MRG (AMKS32778); 2, 18km E Woodford, Blue Mtns NP, 33°44’S, 150°33’E, under log, 17 Apr. 1974, MRG (AMKS32776). DIAGNOSIS. Cymbium with slight sclerotis- ation of posterior retrolateral edge; long cymbial extension; post alveolus is x 1.5 length of alveolus. Without median apophysis; large membranous conductor. ¢ palp with long femoral and tibial segments. DESCRIPTION. See Forster & Wilton (1973:134-135) for description of 2 holotype and ¢ allotype. Female. CL2.1 AL 2.0. Legs: 18.5; 117.1; 116.3; IV 8.1. Epigynum (Fig. 1C-E) with anterior gonopores, simple median insemination ducts to spermathecae (Note: the anterior swellings on insemination ducts (Forster & Wilton fig. 406) were not observed). Females 3.3-5.3 long. Male, CL2.5 AL 2,3, Legs: 112.8; 1110.2; 1119.0; IV 11.3. 3d palp (Fig. 1F-I): embolus short, spiniform arising from distal tegulum; large conductor; no median apophysis, sperm duct with simple loop before entering embolus. Ratio of cymbial alveolus: post-alveolus is 1:1.5. RTA REVISION OF PROCAMBRIDGEA 445 FIG. 1. A-l, Procambridgea ruinbowi Forster & Wilton. A, eyes, dorsal; B, chelicera: C-E, epigynum (veniral, dorsal, lateral); F-I, 3 palp (ventral, retrolateral, dorsal, entire palp). J-P, Pracambridgea cayernicola Forster & Wilton: J-L, epigynum (ventral, dorsal, lateral); M-P, 3 palp (ventral, retrolateral, dorsal, entire palp). drta = dorso-retrolateral branch of tibial apophysis: ma = median apophysis: rta = retrolateral apophysis; vrta = ventro-retrolateral branch of tibial apophysis. 446 with curved ventro-retrolateral and bifid retrolateral branches. Males 4.5-4.8 long. DISTRIBUTION. Jenolan area, Blue Mountains, NSW (Fig. 8A). Procambridgea cavernicola Forster & Wilton (Figs 1J-P, 8A) P. cavernicola Forster & Wilton, 1973: 136, figs 407-410; Brignoli, 1981: 533 (catalogue), MATERIAL. HOLOTYPE: ¢, Wee Jasper Signatore Cave, NSW, 35°09’S, 148°40’E, 2 June 1962, E. Hamilton-Smith (AMKS30614). ALLOTYPE: °, same data as holotype (AMKS30615), OTHER MATERIAL: NSW: 26, 3° , Wee Jasper Punchbowl Cave Bat Chamber, 35°09’S, 148°40°E, 10 Sept. 1977, M. Marx (AMKS32758). DIAGNOSIS. Cave species, less pigmented than P. rainbowi. Long cymbial post-alveolus twice length of alveolus; small slender median apophysis; RTA with extra needle-like dorso-retrolateral branch (ef., P rainbow in all these characters). DESCRIPTION. See Forster & Wilton (1973: 136) for description of 3 holotype and @ allotype. Male. CL2.3 AL2.5. Legs: 113.6; 111.3; 119.9; IV 12.5. d palp (Fig. 1M-P) with small median apophysis. Cymbium with paracymbial bulge; ratio of alveolus: post-alveolus is 1:2.2. RTA with curved ventro-retrolateral, blunt retrolateral and a needle-like dorso-retrolateral branch. Sperm duct with simple open loop (i.e. arms of loop not touching). Males 4.5-4.8 long. Female. CL 2.2 AL 3.1. Legs: I 10.5; 11 9.3; Tf 8.2; [V 10.3. Epigynum (Fig. LJ-L) similar to P. rainbowi with slightly larger gonopores. [Note: The posterior process (Forster & Wilton fig. 409) is thought to be due to damage; it is not present on other females]. Females 5,3-5.5 long. DISTRIBUTION, Wee Jasper, NSW (Fig. 8A). Procambridgea grayi Davies sp. nov. (Figs 2A-G, 8A) ETYMOLOGY. In honour of Michael Gray, arachnologist at the Australian Museum and collector of much of the material for this revision. MATERIAL. HOLOTYPE: 3d, Lane Cove River Park nr Fullers Bridge, NSW, 33°47°S, 151°08’E, under rock Hawkesbury sandstone, 6 Apr. 1974, MRG (AMKS32770). PARATYPES: NSW. 2, same data as holotype (AMKS58083); ¢, Gordon, 33°44’S, 151°09’E, PF, 24 Apr. 1984, CH (AMKS14400); °, Mooney Mooney Ck, 33°31’S, 151°12’E, in rainforest, 13 Jan. MEMOIRS OF THE QUEENSLAND MUSEUM 1977, MRG (AMKS$32771); 3, 2, Bobbin Head, Kuringai Chase NP, 33°39’S, 151°09’E in log, 10 Apr. 1974. MRG (AMKS$32772); d, 9°, same data (AMKS32774); 2 2, same locality, sheet web under logs, 2 Nov. 1974, MRG (AMKS32773); 6, 2, St Ives, 33°44’S, 151°10°E, under log in wet sclerophyll forest, 7 Aug. 1971, MRG GE, Gray (AMKS32775). OTHER MATERIAL: New Zealand. 2, 3, juvs, Air Raid Tunnel, Alten Reserve, Auckland. NZMS 260 R11 685 820, 15 Oct. 1999, M. Hunt (MNZ). DIAGNOSIS. ¢ palp with small membranous conductor with sclerotised tip (cf., P. rainbowi, P. cavernicola). Paracymbial bulge with small posterior projection. Needle-lke median apophysis (cf., P. rainbowi). DESCRIPTION. Male. CL 1.9 AL 2.1. ¢ palp (Fig. 2D-J) with short embolus; small membranous conductor; needle-like median apophysis. Simple sperm duct with arms of loop touching. Ratio of cymbial alveolus: post-alveolus is 1:1.3. Posterior edge of cymbium with small rounded projection. RTA with curved ventro-retrolateral and bifid retrolateral branches. Males 3.9-4.1 long. Female. CL 2.2 AL 2.3. Epigynum (Fig. 2A-C) Females 3.3-4.6 long. DISTRIBUTION. Sydney area, NSW (Fig. 8A) and introduced to Auckland, New Zealand. Procambridgea kioloa Davies sp. nov. (Figs 2H-N, 8A) ETYMOLOGY. From Kioloa SF, N.S.W. MATERIAL. HOLOTYPE: 6, Kioloa SF, Forest Drive, 16km N of Batemans Bay, NSW, 35°37’S, 150°16’E, PF, 28 June 1979, CH (AMKS3834). PARATYPES: NSW. 2, same locality and collector as holotype, 28 Aug. 1978, (AMKS1728); 2, same locality and collector, 30 Apr. 1978 (AMKS1416); @, Boyne SF, Old Highway Rd, 35°38’S, 150°11°E, under rock, moist forest, 15 Aug. 1978, MRG (AMKS82074), DIAGNOSIS. Posterior edge of cymbium with small projection (cf., P rainbowi, P. cavernicola. Ratio of alveolus:post alveolus is 1:1 (cf., previous spp.). Sperm ducts with longitudinal loop with arms not touching (ef., PR. grayi). DESCRIPTION. Male, CL 1.8 AL 1.9. 3 palp (Fig. 2K-N): very short distal embolus, small conductor, slender median apophysis. Arms of loop (longitudinal) in sperm ducts not touching. Cymbium with sclerotised posterolateral edge with small projection. Alveolus and post- alveolus about equal in length. RTA with REVISION OF PROCAMBRIDGEA 447 FIG. 2. A-G. Procambridgea grayi sp. nov. A-C, epigynum (ventral, dorsal, lateral); D-G, d palp (ventral, retrolateral, dorsal, entire palp). H-N, Procambridgea kioloa sp. nov.; H-J, epigynum (ventral, dorsal, lateral); K-N, ¢ palp (ventral, retrolateral. dorsal, entire palp). 448 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. A-G, Procambridgea otwayensis sp. nov. A-C, epigynum (ventral, dorsal, lateral); D-G, ¢ palp (ventral, retrolateral, dorsal, entire palp). H-K, Procambridgea ourimbah sp. nov. 3 palp (ventral, retrolateral, dorsal, entire palp). crp = cymbial retrolateral process; prta = posterior retrolateral tibial apophysis. ventro-retrolateral and pointed retrolateral Procambridgea otwayensis Davies sp. nov. branches. (Figs 3A-G, 8A) Female, CL 1.9 AL 2.3. Epigynum (Fig. 2H-J). Females 3.3-4.5 long. : ar MATERIAL. HOLOTYPE: 6, Erskine Falls, Otway Ra., DISTRIBUTION, Kioloa SE, NSW (Fig. 8A): Victoria, 38°27’S, 143°58’E, in log beech/eucalypt forest, 6 ETYMOLOGY. From Otway Range, Victoria. REVISION OF PROCAMBRIDGEA Apr, 1973, MRG (AMKS32764). Paratypes: Victoria. of, same data as holotype (AMBKSS58082); © , same locality, 6 Apr. 1973, MRG (AMKS32765); 5, Beech Jorest, Otway Ra. 7-Apr. 1973. MEG (AMBS32766): &.Grev R, xing, Olways, 38°39"S, 143°S0°R, 250m, @ Nov. 1977, CiBM, SRM (QMS42196). DIAGNOSIS, Cymbial alveolus is twice as long as postealveolus (ef., all previously described spp.). REA with simple postero-retrolateral branch (ef., all described spp.). DESCRIPTION. Male. CL 2.) AL 2.2. Legs: I 11.45 179.7111 8.0; 1V 10.0. 9 palp ts short (Fig. 31)-G). Embolus spiniform arising from mid- prolateral tegulum; conductor large membranous: median apophysis needle-like. Course of sperm duct with arms of loop coming together and continuing parallel before the anterior arm enters embolus. Ratio of alveolus:post-alyeolus is 1:0.5. Paracymbium a proximal bulge. RTA with ventro- and postero-retrolateral branches. Other ad wasalso 4.3 in length. Female. CL 2.0 AL 2.3. Epigynum (Fig. 3A-C): gonopores open well lateral to the mid-line; spermathecae larger than other species, Females 4.1-4.3 long, DISTRIBUTION, Otway Ra., Victoria (Fig. 8A). Procambridgea ourimbah Davies sp. nov (Figs 3H-K, 8A) ETYMOLOGY. From Ourimbah, NWSW. MATERIAL. HOLOTYPE: 4, Gosford-Ourimbah, NSW, 33°24°S. 1S17°210°1, under lows. rainforest, May 1993 (AMKS35169), DIAGNOSIS. No median apophysis (cf., P. cuvernicola, P. gravi, Pkialea, P. atwavensis), Paracymbium with strong retrolateral process (cf, all previously described spp). DESCRIPTION, Male. CL 2.3 AL 1.9. 3 palp (Pig. 3H-K) with small membranous conductor, sclerotised distally. No median apophysis. Sperm duct simple with closed loop. Paraeymbium with retrolateral process. Ratio of cymbial alveolus: postalveolus is 1:1.4. RTA with small ventro- retrolateral and pointed retrolateral branches. Female. (unknown). DISTRIBUTION, Ourimbah, NSW (Fig. 8A). Procambridgea hunti Davies sp, nov, (Figs 4A-F, 7, 8B) RTYMOLOGY, In honour of the late Glenn Hunt, araghnologist and collector of the holutype. MATERIAL. HOLOTYPE: &, Barrington Tops, NSW. 31°58°S, [S1°28"E, eucalypt forest 1538m, 18 July 1971, CoS. Hunt (AMBS32761). Paratypes: NSW. 3. 4, sane dataas holotype (AMESS5808[): 9, penult. d., Gloucester; 78krn W of Barrington Tops Forest Rd, 32°01'S, 1S 1°09" E, under fog, 19 Mar. 1982, MRG IT. Parnaby (AMKS84 340): ?. Tuglo, 5Ukm NW Singleton, 32°14'S, 1S1°L6°E, srvall sheet webs in lows, 19 Jan, 1977, MRG (AMKS32762), DIAGNOSIS. A larger spider than other species. Very long post-alveolar region of cymbium, more than twice the length of alveolus (cf. all other spp. exeept P cavernicola). RTA without small dorso-retrolateral branch (¢.1., 2 cavernicola), DESCRIPTION. Male. CL 2.8 AL 2.6. d palp (Fiz. 4C-F); embolus, slightly curved; conductor small with sclerotised tip; slender median apophysis; sperm duct with small open loop, Very long cyvmbium with sclerotised posterior edge; ratio of alveolus: post-alveolus is | :224 RTA with ventro-retrolateral and bifid retrolateral branches. Males 5.2-5.8 long, Female,CL2.4 AL2.5. Rpigynum (Fig. 4A, B}. The female from Tuzla was 7.2 long. DISTRIBUTION. Eucalypt forest, Barrington Tops. NSW (Fig, 8B), Procambridgea carrai Davies sp, nov, (Figs 44-M, 7D, 8B) ETYMOLOGY. From Carrai SP, N.S.W. MATERIAL. HOLOTYPE: &. Carrai SF, NSW, 8O0kim W of Kempsey: 31°01°S, 152°20°E, rainforest, 18 July 1971, MRG GE. Gray (AMKS32747). PARALYPES: NSW. ¥, Carrai SF nr Carrai Bat Cave. in Jog, rainforest, 26 Apr. 1974, MRG(AMKS32748); 2 7, Carrai SP nr Carrai Arch Cave nr Kookaburra W ol Kempsey, in cave, | Aug. 1971, L. Henshaw (AMKS$32749); ¢, Carrai SF, start Kookaburra Trail, 31°02°S, 152°20°E, pyrethrum tree fern, 19 Oet. 1992, J, Stanisic, G Ingram (QMS42240); 24a Mt Boss SF (Fenwicks), 31°12"S, 152°24"E, Oct. 1980. GA. Webb, Forestry Commission (AMKS43510). OTHER MATERIAL: 24. Kerewolg SF nr Lorne, 31°36"S, \42°34°B, PF, 15 July 1979, 2. Milledge (AMIKS5405); 2 same data (AMKS5426), & Kerewong SF, 31°345°S, 52°47, PR 29 Aug. 1978, 2, Milledge (AMKS1982), DIAGNOSIS, RTA with dorso-retrolateral branch (cf, all species except P cavernicolc), Without median apophysis (cf, P cavernicola). DESCRIPTION. Male. CL 2.2 AL 2.3. 4 pulp (Figs 4J-M, 7D): short spiniform embolus, small conductor, without median apophysts. Sperm duct with open loop leading to embolus. Cymbium with selerotised posterolateral elves ratio ofalvedlus:postealveolis is 121.8, RTA with ventro-retrolateral, retrolateral and small dorso-retrolateral branches. Males 3.8-4.5 long. Female. CL 1.8 AL 2.1 Epigynum (Fig. 4G-I). Females 3.9-5.0 long. DISTRIBUTION. Rainforest areas W of Kempsey, NSW, (Fig. 8B). Procambridgea monteithi Davies sp. nov. (Figs 5A-G, 8B) ETYMOLOGY. In honour of Geoffrey Monteith, collector extraordinaire. MATERIAL. HOLOTYPE: ¢, Point Lookout, New England NP, NSW, 30°30’S, 152°24’E, 1300m, PF 101, Nothofagus forest, 11 Nov. 1980-16 Mar. 1981, GBM (QMS42206), PARATYPES; NSW. 3 @, Point Lookout (upper), New England NP, 1400m, PF 100, 21 Mar-11 Nov, 1980, GBM (QMS42253). DIAGNOSIS. Cymbial alveolus longer than post-alveolus (cf., all previously described species except P. otwayensis). No median apophysis (cf., P. ofwayensis). Palpal tibia with proximal retrolateral spur (cf., all spp.). DESCRIPTION. Male. CL 1.6 AL 1.3; ¢ palp (Fig. 5D-G) curved embolus; large membranous conductor, sclerotised retrolaterally; without median apophysis. Sperm duct with double loop. Cymbium with flange-shaped paracymbium without processes; cymbial alveolus longer than post-alveolus 1: 0.6. RTA with ventro-retrolateral and non-bifid retrolateral branches. Tibia with proximal retrolateral spur. Other male larger, 3.8. Fenale. CL 1.7 AL 2.1. Epigynum (Fig. 5A-C) Gonopores widely separated; insemination ducts enter spermathecae medially. DISTRIBUTION. In beech forest in New England NP, N NSW (Fig. 8B). Procambridgea lamington Davies sp. nov. (Figs 5H-R, 8B) ETYMOLOGY. From Lamington NP, Queensland. MATERIAL. HOLOTYPE: ¢, Lamington NP, SE Queensland, 28°11°S, 153°11°E, 9-10 Aug. 1977, RJR (QMS42200). PARATYPES: Queensland. d, same data as holotype (QMS42239); 9, 2¢, Lamington NP, notophyll forest, 9 July, 1977, RJR (QMS42199); 9, Nagarigoon, Lamington NP, PF, 27 Mar.-8 Apr. 1976, RJR, VED (QMS42201); 32, ¢, Nagarigoon Ck, 5 Apr. 1976, RJR, VED (QMS42215); ¢, Lamington NP, 9-10 Aug, 1977, RJR (QMS42216): d, Binnaburra, Lamington NP, 1 July 1986, M.S. Harvey, PJ. Vaughan (WAM 98/2095); 2, Daves Ck Country, Lamington NP, 3 Apr. 1976, RJR (QMS42217); 2, Morans Falls, Lamington NP, 28°19°S, 153°05’E, 900m, Berlesate 924, rainforest stick MEMOIRS OF THE QUEENSLAND MUSEUM brushing, 15 Mar.1997, GBM, B. Russell (QMS42218); 2, Albert R, rt Branch, Lamington NP, 244m, PF 65, 8 Sept.-30 Oct. 1976, GBM, SRM (QMS42219). 5d, 28, Springbrook, 28°15’S, 153°16’E, 1000m, rainforest, PF, 28 Oct. 1991, M. De Baar (QMS23032); 2°, Springbrook Repeater Stn, 1000m, stick brushing Berlesate, 27 May 1997, GBM (QMS42198); 2, Upper Tallebudgera Valley, SEQ. 28°14’S, 153°18’E, 530m. PF, Mar.-July, 1985, D. Cook (QMS42220), 23, 22, Mt Tamborine, 27°55’S, 153°10°E, 10 July 1974, VED, RJR, C.L. Wilton (QMS42254). DIAGNOSIS. Paracymbium with retrolateral and ventral processes (cf. previously described spp). DESCRIPTION. Male. CL 2.1 AL 2.0. 3 palp (Fig. 50-R): with curved embolus, large retrolaterally sclerotised conductor, no median apophysis; sperm duct with transverse double loop. Cymbial alveolus about twice as long as post-alveolus 1:0.6, Paracymbium with ventral and retrolateral processes. RTA with ventro-retrolateral and bifid retrolateral branches. Males 4.0-4.6 long. Female. CL 2.0 AL 2.7. Carapace (Fig. 5H) without pattern. Sclerotisation of median epigynal ridge varies (Fig. 5J, K). Dorsal, lateral and posterior views (Fig. SL-N) show the course of the insemination ducts. Females 3.6-4.7 long. DISTRIBUTION. Lamington Plateau and Springbrook area, SE Queensland (Fig. 8B). Procambridgea hilleri Davies sp. nov. (Figs 6A-G, 7A-C, 8B) ETYMOLOGY. In honour of naturalist A. Hiller, collector of the holotype. MATERIAL, HOLOTYPE: 2, Mt Glorious, SE Queensland, 27°20’S, 152°46’E, rainforest, malaise trap, 27 June-18 Oct 1982, A. Hiller (QMS42203). PARATYPES: Queensland 2, Mt Glorious NP, rainforest, 6 Jan. 1977, B.J. and M.J. Marples, RJR, VED (QMS42221); 2, Mt Tenison Woods, 27°19’S, 152°44’E, 750m, rainforest, stick brushing, 15 May 1997, GBM (QMS42222); ¢, 22 Mar. 1979, GBM (QMS42255); d, Mt Mee, via Samford, 27°05’S, 152°41°E, 518m, rainforest, PF 14, 1974-1975, GBM, SRM (QMS42202); 2, Mt Nebo, 27°23’S, 152°47°E, rainforest, 10 Sept. 1973, C, Wallace (QMS42256); 32, 3,2 penult. ¢, Booloumba Ck, Conondale Ra. SEQ, 26°39°S, 152°39’E, 13-18 May 1976, RJR (QMS42234); 3, Casey Ck via Imbil, 26°29’S, 152°38’E, PF 3, 10 Aug.-9 Nov., 1974, GBM, SRM, (QMS42236); 2, Mt Cabinet via Jimna, 26°43’S, 152°34°E, PF, 29 June-23 Aug. 1975, GBM, SRM (QMS42237); 2 Tungi Ck, via Jimna, 26°39’S, 152°28’E, 550m, PF 28, 29 Mar-16 June 1975, GBM, SRM (QMS42238). REVISION OF PROCA MBRIDGEA 451 FIG, 4. A-F, Procambridgea hunti sp. nov. A,B, epigynum (ventral, dorsal); C-F, ¢ palp (ventral, retrolateral, dorsal, entire palp). G-M, Procambridgea carrai sp. nov.: G-1, epigynum (ventral, dorsal, lateral); J-M. ¢ palp (ventral, retrolateral, dorsal, entire palp). drta = dorso-retrolateral tibial apophysis; rta = retrolateral tibial apophysis. DIAGNOSIS. Paracymbium with retrolateral — /amingtom): ventral process small and blunt (cf.. and ventral processes (cf, all species except PP. damington well-defined, pointed). 452 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5, A-G, Procambridgea monteithi sp, nov. A-C. epigynum (ventral, dorsal, posterior); D-G, ¢ palp (ventral, rétrolateral, dorsal, entire palp). H-R, Procembridgea lamington sp. \oy.: H, carapace; 1, Eyes, frontal: J, K, epigyna, ventral; L-N, epigynum (dorsal, lateral, posterior), O-R, ¢ palp (ventral, retrolateral, dorsal, entire palp). erp = cymbial retrolateral process: cvp = cymbial ventral process. REVISION OF PROCAMBRIDGEA ts " ree) FIG. 6, A-G, Procanibridgea hilleri sp. nov. A-C, epigynum (ventral, dorsal, lateral); D-G, d palp (ventral, retrolateral, dorsal, entire palp). H-N, Procembridgea montana sp. nov.; H-J, epigynum (ventral, dorsal, lateral); K-N, ¢ palp (ventral, retrolateral, dorsal. entire palp). 454 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 7. A-C, Procambridgea hilleri sp. nov.: A, epigynum; B, PMS(1.) with large mAP and PCR with 2 shafts; C, 3 palp (ventro-retrolateral). D, Procambridgen carrai sp. nov. 2 palp (ventral). DESCRIPTION. Male. CL 1.9 AL 1.7. d palp DISTRIBUTION. D’Aguilar and Conondale (Figs 6D-G, 7C) with large retrolaterally sclerotised conductor, no median apophysis; sperm duct with transverse double loop, arms of loop not touching. Cymbial alveolus: post-alveolus, 1:0.7. Paracymbium with sharp retrolateral spur and small blunt ventral process. RTA with obliquely angled ventro-retrolateral and bifid retrolateral branches. Males varied in length from 3.4-4.0. Female, CL 1.8 AL 2.3. Epigynum (Figs 6A-C, 7A). ALS with 2 MAP, about 25 piriform spigots. PMS with large anterior mAP, some paracribellar spigots, one with 2 shafts on base (Fig. 7B), one posterior cylindrical spigot and aciniforms. Females 3.5-4.3 long. Ranges, SE Queensland (Fig. 8B). Procambridgea montana Davies sp. nov. (Figs 6H-N, 8B) ETYMOLOGY, From Latin ‘montanus’, of mountains. MATERIAL. HOLOTYPE: ¢, Mistake Mtns (north) 27°58’S, 152°22’E, via Goomburra, SE Queensland, 975m, PF 74, 13 Feb.-13 Oct. 1977, GBM, SRM (QMS42224), PARATYPES: Queensland. ¢, Mistake Mtns (middle) 27°58’S, 152°23’E, via Goomburra, 950m, PF 75, 13 Feb.-13 Oct. 1977, GBM, SRM (QMS42223); 22, Mistake Mts, 16 Oct. 1976, RJR (QMS42226); 3, Bald Mt via Emu Vale, 28°14’S, 152°22’E, 1130m, PF 20, 30 Mar.-2 Aug 1975, GBM, SRM (QMS42205);3 2, Mt Superbus, 28°14’S, 152°28°E, PF, 12 Mar.-13 June 1990, REVISION OF PROCAMBRIDGEA 455 @ PP. rainbowi 1 sP. cavernicola L/ P. grayi OP. kioloa Ley P. otwayensis HEP. ourimbah NEW SOUTH WALES e P, hilleri OP. hunti Z_/ P. monteithi O P. montana LH P. lamington i P. carraj QUEENSLAND NEW SOUTH WALES FIG. 8. A, B, Maps showing distribution of Procambridgea spp. T. Churchill, RJR (QMS15917); 2, Mt Superbus, 30 Oct. 1990, T. Churchill, RJR, K. Williams (QMS25888); ¢, 2, same locality and collectors, 1000m, PF, 13 June-30 Oct. 1990 (QMS26292); 2 Mt Superbus, 1360m. moss on tree trunks, 3 Mar. 1983, D. Yeates (QMS42258); stick brushing, GBM (QMS42259); 2 Spicers Peak, 28°06’S, 152°24’E, 1200m, 30-31 Dec. 1993, GBM (QMS42230); 3, 32, Cunninghams Gap, 28°03°S, 152°23’E, sieved litter, dry forest, 28 June 1991, D. Black (WAM 98/2084-7). NSW. 2d, Nothofagus Mt via Woodenbong, 28°17’S, 152°37°E, 1100m, Berlesate No 416 from stick brushing, 17 June 1982, GBM, G. Thompson (QMS42197); 3, &, Brindle Ck, Wiangaree via Kyogle, 28°22’S, 153°04’E, 740m, PF 42, 22 Mar-2 Aug. 1975, GBM, SRM (QMS42257). DIAGNOSIS. Short embolic area, about a quar- ter length of tegulum (cf., P. hilleri, P.lamington, about half length of tegulum). Sperm duct with longitudinal double loop (cf., P. monteithi, transverse double loop). DESCRIPTION. Male. CL 1.8 AL 1.5. ¢ palp (Fig. 6K-N) with short distal embolus, mem- branous conductor, no median apophysis. Sperm duct in double loop (longitudinal rather than transverse). Ratio of alveolus: post-alveolus 1s 1:0.6. Paracymbium with very small ventral pro- cess and retrolateral process. Tibia short, as long as wide. RTA with curved ventro-retrolateral and bifid retrolateral branches. Males 3.2-3.8 long. Female. CL 2.4 AL 2.4. Epigynum (Fig. 6H-J). Females 4.8-5.2 long. DISTRIBUTION. In the area where the McPherson Range meets the Great Dividing Range (Fig. 8B) on the Queensland/New South Wales border. RELATIONSHIPS OF PROCAMBRIDGEA TERMINAL TAXA. A cladistic analysis ex- amined 52 characters (Table 1) for relationships of the 12 Procambridgea spp. and 27 other Australian taxa as well as Amaurobius from the Northern Hemisphere, and Amphinecta and Matachia from New Zealand (Table 2). Outgroup comparison was with Oecobius navus Blackwall. The data matrix (Table 2) was prepared using MacClade version 3.08 (Maddison & Maddison 1999) and PAUP*. Fig. 9 was prepared using PAUP*. Fig. 10 was prepared using CLADOS version 1.2 (Nixon 1992) with DELTRAN optimisation. DATA ANALYSIS. We analysed the data matrix for the 42 taxa (Table 2) using PAUP* version 4.0b4a (Swofford, 1999) on a Power Macintosh 7100/66. Heuristic searches of the data were completed using 10 random step-wise addition sequences, tree-bisection-reconnection (TBR) branch swapping, MULPARS, and branches having maximum length zero collapsed to yield polytomies. Strict and semistrict (Bremer, 1990) consensus trees of the most parsimonious trees (MPTs) were computed using PAUP*. Analyses 456 TABLE 1. Characters and character states. T AME: as large or larger than ALE (0); smaller (1) 2 CH: normal (0); small (1) 3* Retromarginal CH teeth: no teeth (0); 1 (1); 2 (2); 3 (3); 3+ (4); 5+ (5) 4* Promarginal CH teeth: no teeth (0); 2 (1); 3 (2); 3+ (3); 5+ (4) 5 Long prolateral seta at base of fang: absent (0); present (1) 6 Carapace: round ((); oval (1) 7 Enlarged frontal CH seta: absent (0); present (1) 8 Foveal area highest: absent (0); present (1) 9 @ leg I: shorter than leg [V (0); equal to or longer than leg IV (1) 10 Stridulatory ridges on C coxa I: absent (0); present (1) 11 Enlarged ventral spines on tibia and MT [ and II: absent (0); present (1) 12 Feathery hairs: absent (0); present (1) 13 MT preening comb: absent (0); present (1) 14 MT TRICH: 2+ (0); 1 (1) 15* T TRICH: 0 (0); 2+ (1); double row (2) 16 T rod: absent (0); present (1) 17 Anal tubercle: normal (0); enlarged (1) 18 CR spinning fields: 2 (0); 1 (1); absent (2) 19* CAL: proximal (0); proximo-medial (1); long medial (2); no CAL (3) 20 MAP ALS: 2 (0); 1 and nubbin (1); | (2) 21 MAP 9 ALS: mesal (0); anterior (1) 22* PCR 2 PMS: one shaft per base (0); more than one shaft (1); absent (2); no CR (3) Female characters 23 Medial EPIG atrium: absent (0); present (1) 24* Posterior rim of medial atrium/EG: no medial atrium (0); close (1); well forward (2) 25* ID: simple (0); loosely coiled (1); tightly coiled (2) 26* Posterior EPIG scape: no scape (0); small knob (1); short (2); long (3) 27 Lateral projections on EPIG: absent (0); present (1) 28 Lateral EPIG teeth: absent (0); present (1) Male characters 29 E shape: spiniform (0); broad (1) 30 Direction of E: clockwise (0); anti-clockwise (1) 31 PE APOPH: absent (0); unbranched (1); branched (2) 32 E APOPH with 2 -3 long setae: absent (0); present (1) 33* E APOPH plate-like setae: absent (0); small (1); large (2) 34* Shape of C: irregular (0); rounded (1); plate-like (2); T-shaped (3); s-shaped - falciform (4) 35 C/E: opposite (0); embracing (1) 36 Position of embracing C: retrolateral (0); prolateral (1) 37* C length/ tegulum: quarter (0); half (1); whole (2) 38 Secondary C: absent (0); present (1) 39 Median APOPH: absent (0); present (1) 40* Loops of sperm duct: simple (0); 1 (1); 2 (2); 3 (3) 41 Orientation of CB to bulb: dorsal (0); mesal (1) 42* CB alveolus/ post-alveolus: shorter (0); equal (1); longer (2); twice as long (3) 43* PCB process: absent (1); 1, retrolateral (1); 2, retrolateral and ventral (2) 44 RTA: absent (0); present (1) 45* RTA/CB length: absent (0); quarter or less (1); more than half (2) 46 RTA dorsal branch: no RTA (0); branch absent (1); branch present (2) 47* RTA with small dorso-retrolateral branch : no RTA (0); branch absent (1); branch present (2) 48 RTA extra distal branch: no RTA (0); extra branch absent (1); extra branch present (2) 49* RTA proximal projection: no RTA (0); no proximal projection (1); proximal projection (2) 50 Tibial proximal retrolateral projection: absent (0); present (1) 51 Palpal tibia length/width: shorter to slightly longer than wide (0); much longer than wide (1) 52 Palpal P APOPH: absent (0); present (1) (* Multistate characters treated as unordered] MEMOIRS OF THE QUEENSLAND MUSEUM Oecobius Dictynidae sp. Badumna Amaurobius Paramatachia Matachia Quemusia Jalkaraburra 1 2 Malala 2 Manjala plana 1 Manjala pallida Desis Amphinecta Midgee binnaburra 1 Midgee thompsoni 2 Tasmarubrius 1 Storenosoma Otira 5 Dardurus spinipes Dardurus tamborinensis 4 Kababina 2 Wabua major 2 Wabua halifax 5 Carbinea longiscapa Carbinea breviscapa Malarina monteithi Malarina cardwell 1 Baiami 1 Stiphidion facetum Stiphidion adornatum 1 Procambridgea monteithi Procambridgea lamington i Procambridgea hilleri Procambridgea montana Procambridgea otwayensis Procambridgea rainbowi 1 Procambridgea carrai 1 Procambridgea ourimbah 1 Procambridgea grayi 0 Procambridgea kioloa o Procambridgea cavernicola Procambridgea hunti Kababininae Stiphidiidae FIG, 9. Semistrict consensus of 36 MPT showing Bremer supports above the nodes. were repeated using Hennig&86 version 1.5 (Farris, 1988). The command mh* was used to find initial trees. The trees retained were then passed to the extended branch swapper, bb*. Bremer support (Kallersj6 et al., 1992) to indicate character support for nodes on the cladogram was calculated using the computer program TreeRot (Sorenson, 1999) on the preferred MPT with 20 random step-wise addition sequences, and support indicated above the nodes on the semistrict consensus tree (Fig. 9). RESULTS. Heuristic searches of the 52 characters for the 42 taxa generated 36 MPTs of tree length 185, consistency index (Kluge & Farris, 1969) 0.46, consistency index excluding uninformative characters 0.42, retention index (Farris, 1989) 0.71 and rescaled consistency index 0.33. Hennig&86 finds the same 36 MPTs. Many of the MPTs have terminal polytomies in Procambridgea (Fig. 10) as synapomorphic characters were not able to be scored across all taxa. Thus the strict consensus is less resolved. The MPTs are divided into two groups of 18 cladograms that differ markedly in the placement of Amaurobius in the Amaurobioidea (see REVISION OF PROCAMBRIDGEA HH Oecobius at 1821253441 i Dictynidae sp. 2223244 239 HHH} Badumna 1 HH 3} hind ort 3746 SHHHHHT Amaurobius 3107 oo? [H+ Paramatachia 13.4 6 9 MASIT 44 45 46474849 oI 253942 11034 {HH} Matachia rrsdrPrzrr ais HH} w12 bE: 181922 : 4303638 HH} Quemusia ua f3 2 o072 ft PLT a 1 i+ Jalkaraburra 5 3 2K99 192251 {HH} Malala 3 141520 fH} ae ro 181925 H) { i TT 2. | 523% HH Manjala plana 2 10 Fy oo2 EHH] 42 182039 "11 Lie Manjala pallida ' 1820212537 [HHHt Desis 2291372 4x7 13.28 40.42 ‘ FA Amphinecta 40 35 739 -Midgee binnab. ro EHHHH 2537 er Midgee thomps. HHH} 2027 43 3637 rood {HH} Tasmarubrius {3 132946] 01 i ol 7 0 Fit Storenosoma 16 : Otira 1 3 118224982 7 Dardurus spinip- HH *')22 1) LDardurus tambor. 424548 38% 5) EHH Kababina li fi b t+ 37495) try 2 asa0 HH Wabua major o20 aan) |) bWabua halifax an ° as 26 Malarina montei. asi ou | | 4H Malarina cardw. eH zt 2645 $40 TT Gs1922263151 Carbinea longise. Hi fH} fH ** ri 733729 | Carbinea brevisc. 2324374251 FHHHH} Baiami 12193436 P2201 HHH} oh idi Tia 1 | 1.4 -Stiphidion facet. {HH 25 °2 Lit Stiphidion adorn. ey 50 42 Pro. monteithi Hy ' a. Pra. lamington 4043 fe Pro. hilleri 3 oo |a {+ Pro. montana 52 B cy {-Pra otwayensis 1 ay {+ Pro. rainbowi 1 a 42 HE Pro. carrai Hos . Pro. ourimbah x ti Pro. grayi t » a7 fl {+ Pro. cavernicola 1 2 4 a Pro. kioloa Pro. hunti FIG. 10. Preferred most parsimonious tree showing characters and characters states. 457 TABLE 2, Data matrix. Taxa Oecabius navus Blackwall Dictynidae sp. Badumaa longingua {(Kach) Matachia ranulicola Dalmas Paramatachia Gdecorata Dalmes Desis sp. Quemusia aquilonia Davies Jalkaraburra 4@]ta Davies A!mphinecta milina Forster & Wilton Amaurobius fenestralig (Stroeam) Storenosoma terransa Davies Otirs summa Davies Tesmerubrius miivinwe (Simon) Stiphidion facetwa Simon Stipbhidion adarnatun Devies MEMOIRS OF THE QUEENSLAND MUSEUM Character Number 10 20 30 Aan 5D DLUCGODDON GOOLODIOIZ 0200790000 aoddoz0010 Qood000000 oO 1013010010 OUZSO1IONIN ONODIGWNON 0112100100 O004112010 9001111110 oe 1NZ3010010 OOONLOO12D NNOOBvH1OO OOdDAID1OL? 201111110 oo 80273010010 OODOL9V130 DONNLGOOON 0004101000 CeaLLilllo OL 1024010010 DDbOZUUEaa 1022010010 0000100000 o2no1On101 1022010010 DG00100Z30 UR00LNDIGT GO04TT1100 OG04L1IA11O 19 1024010010 0010100231 G3002D0100 OOOLOPO011 OQZO1111110 19 OO330L0010 0010100000 GO00190110 GONLOF0010 OOd1121i10 O10 LOZLOLOOOL OOLOLOUZT1 0300190110 GOOLOFOOLO ONOLTZ11i0 O0 1021010001 0010110231 LOZLOLOOOD 00101002306 O3001NL01D WOUI0?00TN DO11121110 Co QOZ2L1LIO1T O1001NNN1O 0206100000 0003111002 Q0NL111119 00 GCOZ211001N N1LOOLOO01O 0100300000 0003111001 OHOII1I11N oO ODDLO0DIN 100D3HGNN0 Oodz1I2000 10011111309 00 13O0300L0O 0004102010 DOOLALLI1O 10 Oo04111100 OAOAILITIO 10 0300100010 ON01070010 DNO112i110 CO Baiami volucripes (Simon) 1024110010 0100100010 112100000 0003112002 9101111119 10 Midgee binnaburra Davies 1031111000 LOQDLODZ41 N300100000 OODLONOGG G1G1111110 OO Midgee thompsoni Davies LOF1111000 LOGOLS0AS1 OFMMA0G000 DOOIOFLONN 9091111110 O0 Manjala plane Davies Manjala pallida Davies MNalala Tubinae Davies Dardurus spinipes Davies Dardurus tamborinensis Davies Kababine alta Davies Carbines longistapa Davies Carbines brevistapa Davies Malaring monceithi Davies Malarina cardwell Davies Wabua Major Davies Wabua halifax Davies Procambridgea rainbow? (Forster & Wilton) Procambridgea cavernmicola (Forster & Wilton) Procambridgea grayi sp, nov. Procambridgea kzoloa sp, nov. Procambridgea ofwayensis Sp, nov. Procambridgea curimbal sp, nav. Procambridgea Aunti sp, nay, Procambridqea carrai sp. nev. Procambridgea monteithi sp, nov. Procambridgea lamington sp. nov, 1023111010 LHOGIGAGGL OPLIDOOING 0004232010 0101111110 10 IOZ5LATAIO 1000100231 GILZ1OOLOW HOH4113010 0201111119 10 1043011010 1000190231 OINALOGOGY HOGALAAGOO OLOLIIII19 1h 1N43110010 LOOOLOOLGO OzazLOGOOM YOOLLOIGOA HoNIalii20 12 1093210010 LOOM OO1OO O2i21d000d OUO1LH100L 0600232120 12 1921110110 0100100000 9012100000 9001101001 1021110110 0100100230 0312130010 2001101001 OOHIZ1qiLO AN LW21J1D11G OLO01CO930 0312120019 2O01L01001 0001111116 oO LOZI1101L10 6160100000 0012110010 LIZLLOL001 OOOTIIII90 1D 1021110110 6200100000 6012310019 1011301001 00012111120 10 LOZ1110110 0100100000 0011100000 0001100003 0001111120 tb LOZ1LL10100 9190100000 60612100550 HNN1LNL003 O0O4111110 10 LO54110010 9000100000 0106100005 GONSLOL002 Gadlit1110 Lo ICSALLOOLO OOON1000NO 0200100000 GOoalooo1i O304112110 10 1054110010 OD00100000 0100100600 HoOn1OOOLt O2OLIITII0 10 LOS412G010 ONONLOODOO 8100100006 9007100011 D1IDIIIII10 10 1054119010 (OOOLOOODD 0100100000 5002101011 OO011L1110 10 1954110010 G000100000 0100100000 0002100001 0211111110 10 1054110030 0000100000 0100100000 0002100011 9301111110 10 1054110020 0000100000 5100100000 9002101001 O2H11L1211n 19 1054110010 0000100000 0100100000 0002101001 ON51111211 00 1054110010 9000100999 S1o0100000 0902101002 02012121210 10 0021111110. 00 Procambridgea hiileri sp. noy. 1054110010 GHOOLOOOND G100100000 00021010639 0021111116 O00 Procambridges montana Sp. now. LO44L10010 0000100000 0100100000 0002100002 0521111110 00 discussion). Thus the semistrict consensus of the 36 MPTs (Fig. 9) indicates a basal polytomy that leaves the placement of Amaurvbins ambiguous, Nodes on the semistrict consensus tree (Fig. 9) that receive a Bremer support of 0 do not indicate conflict between the topologies but show that those nodes are unresolyed in some of the MPTs. Figure 10 shows characters and character states onthe preferred MPT with Amaurobius forming a basal clade with Badumna. DISCUSSION Cladistic analyses including many taxa of the Amattobioidea, outlined the difficulty of family placement of the Kababminae (Davies, 1999; Davies & Lambkin, 2000, 2001). Analysis of the Amaurobioidea, including many taxa of Procambridgea, continues to present difficulties with family placement. The Amaurobioidea. Kababininae, and Procamhridgea form well-resolyed clades. However, inclusion of Procaumbridgea with a further |2 taxa, into the analysis causes changes to the basal topology. and results in two alternative resolutions for the placement of Aniurobius Jenesiralis (Stroem). In 8 MPT Amaurohius forms a third, basal clade with Bacdumaa (Fig. 10). Therefore in these MPTs the Amaurobioidea does not form the hwo distinct clades seen in previous analyses (Davies, 1999; Davies & Lambkin, 2000,2001.). In the other 18 MPT Amanrobius ts sister to the large clade including Dardurus. the Kababininae, Stiphidion and Procambrideea. In all MPT one clade continues to contain the type genera of the Desidae and Amphinectidae (Figs 9, 10), REVISION OF PROCAMBRIDGEA Procambridgea forms a separate group, distinct from the Kababininae, and more closely related to Stiphidion and Baiami. The Bremer support for these relationships is poor, never more than I, and based entirely on homoplasious characters (Fig. 10); thus Procambridgea remains in the Stiphidiidae. Procambridgea contains two species-groups: a northern group comprising monteithi, lamington, hilleri, and montana and a more southern group of rainbowi, cavernicola, grayi, kioloa, ourimbah, hunti, carrai, and otwayensis; however support for these groups is poor. ACKNOWLEDGEMENTS We are indebted to the curators and collection managers of the Australian, Western Australian and New Zealand Museums for loans of material and to all collectors of material for this revision. We thank the Council of the Australian Biological Resources Study for funding rainforest surveys in Queensland during which some of this material was collected and for financial support of illustrator and co-author, Christine Lambkin. She also set up the cladistic analyses resulting in the cladograms. We are grateful for the support of other members of the Queensland Museum, particularly Jennifer Cannon and Christine Thai for their help in the preparation of this paper. LITERATURE CITED BREMER, K. 1990. Combinable component consensus. Cladistics 6: 369-372. BRIGNOLI, P.M. 1983. In Merrett, P. (ed.) Catalogue of Araneae described between 1940-1981. (Manchester University Press: Manchester, UK). DAVIES, V.T. 1999. A new spider genus from North Queensland, Australia (Araneae: Amaurobioidea: 459 Kababininae). Journal of Arachnology 27 (1): 25-36. DAVIES, V.T. & LAMBKIN, C. L. 2000. Malarina, a new spider genus (Araneae: Amaurobioidea: Kababininae) from the wet tropics of Queensland, Australia. Memoirs of the Queensland Museum 45(2): 273-283. 2001. Wabua, a new spider genus (Araneae: Amaurobioidea: Kababininae) from north Queensland, Australia. Memoirs of the Queensland Museum 46(1): 231-249. FARRIS, J.S. 1988. Hennig86 Version 1.5. Port Jefferson: New York. 1989, The retention index and the rescaled consistency index. Cladistics 5: 417-419. FORSTER, R.R. & WILTON, C.L. 1973. The spiders of New Zealand. Pt IV. Otago Museum Bulletin 4: 1-309. KALLERSJO, M., FARRIS, J.S., KLUGE, A.G. & BULT, C. 1992. Skewness and permutation. Cladistics 8: 275-287. KLUGE, A.G. & FARRIS, J.S. 1969. Quantitative phyletics and the evolution of Anurans. Systematic Zoology 18: 1-32. LEHTINEN, P.T. 1967. Classification of the cribellate spiders and some allied families, with notes on the evolution of the suborder Araneomorpha. Annales Zoologici Fennici 4: 199-468. MADDISON, W.P. & MADDISON, D.R. 1999 MacClade Version 3.08a. (Sinauer Assoc. Inc.: Sunderland, Massachusetts). NIXON, K.C. 1992. Clados, Version 1.2. (L.H. Bailey Hortorium, Cornell University: Ithaca). PLATNICK, N.I. & SHADAB, M.U. 1975. A revision of the spider genus Gnaphosa (Araneae: Gnaphosidae) in America. Bulletin of the American Museum of Natural History 155: 1-16. SORENSON, M.D. 1999. TreeRot, Version 2. (Boston University: Boston, Massachusetts). SWOFFORD, D.L. 1999. Phylogenetic Analysis Using Parsimony (* and other methods). Version 4.0b4a. (Sinauer Assoc. Inc.: Sunderland, Massachusetts). 460 SOUTHERN RANGE EXTENSION FOR THE DELICATE MOUSE (PSEUDOMYS DELICATULUS). Memoirs of the Queensland Museum 46(2): 460, 2001:- Watts & Aslin (1981) described the delicate mouse (Pseudomvs delicatulus) as a species of the tropical north, from Port Hedland in Western Australia to Bundaberg in Queensland, Despite being widespread, this small cryptic species is rarely encountered across much of its range (Watts & Aslin, 1981). In favourable habitat patches, P. delicatulus populations have been found to undergo dramatic fluctuations, sometimes producing brief erruptions after long periods of continuous breeding, later becoming uncommon (Braithwaite & Brady, 1993). Increased biological survey activity in southeastern Queensland in the past decade has recognised a considerable southward extension of the known range for P. delicatulus. It was reported from Lake Broadwater Conservation Park 25km southwest of Dalby (27°20’S 151°00°E) in August 1996 (QM JM11410) (Mathieson et al., 1999). This note reports a further southerly range extension of 130km for P. delicatulus to a site south of Inglewood following the capture of an individual during a larger project investigating the flow-on effects on biodiversity resulting from rabbit calicivirus disease induced-declines in rabbit numbers. The survey site is located on ‘Whetstone’ (28°31°S 150°55’E), 19km SW of Inglewood. The study site consists of three broad habitat types: 1) cracking clay soils adjacent to McIntyre Brook supporting a partially cleared woodland of forest red gum (Eucalyptus tereticornis), river red gum (E. camaldulensis) and rough-barked apple (Angophora JAloribunda) with an understorey of long grass (Stipa spp.); 2) alluvial plains, including an area of pasture, originally a poplar box (/. populnea) woodland, now grazed by cattle; and 3) a partially-cleared woodland of cypress pine (Callitris glaucophiylla), Eucalyptus spp. and bulloak (Al/ocasuarina luehmannii) on sandy-textured solodic soils, adjacent to an extensive State Forest. Small vertebrate faunas were surveyed at & sites using 4 Elliott trap formations and 4 pitfall lines. There was | trap site of each type in the two woodland communities; one along the river and another adjacent to the State Forest. The remaining 4 sites were located within the pasture habitat, which made up a substantial part of the site. Each Elliott trap site consisted of 49 traps arranged in two cross-arms. Elliott traps were spaced at 10m intervals and baited with a mixture of peanut butter and rolled oats. Pitfall trap sites consisted of a continuous 32m aluminium fly-wire drift fence positioned over 7 evenly spaced pits (PVC pipe 15cm diameter, 50cm deep) buried flush with the ground. Trapping was conducted for three consecutive nights. Whetstone was trapped on nine occasions between October 1996 and June 1999, giving a total trapping effort of 5,145 Elliot trap nights and 588 pitfall nights. A single male P. delicatulus (QM JM12786) was captured in an Elliott trap during May 1997 near the State Forest. The habitat surrounding the trap was mostly bulloak regrowth on sandy soils with several large fallen trees providing considerable cover. No further P. delicatulus were captured despite this site MEMOIRS OF THE QUEENSLAND MUSEUM being sampled on six occasions between August 1997 and June 1999, Extra trapping conducted (100 Elliot trap nights) in bullock regrowth closer to the State Forest boundary in June 1999 and analysis of 70 fox/cat scats collected from the site in winter of 1997 and 1999 (Palmer unpubl. data) also failed to detect this species. Mus domesticus was the most common and widespread mammal species captured. It was most common (33.3 mice per 100 Elliott trap nights) when the P. delicatulus was captured, but numbers were generally low at the site adjacent to the State Forest compared with other sites. Two other native species, Sminthopsis murina and Antechinus flavipes, were captured. These species were rare, but most individuals were trapped in similar habitat (bulloak regrowth) to P. delicatulus. No native mammals were captured at the sites within the pasture. The mouse was kept as a live specimen at the Queensland Museum for almost two years. Initially it was thought to be a Pilliga mouse (P. pilligaensis), Later examination of the mouse’s skull revealed that it was a P, delicatulus or closest to this species. In the meantime, this individual was incorrectly reported by Sandell & Start (1999) to be a P. pilligaensis, as a result it appears in the Rabbit Calicivirus Disease Program Report 4’ as this species. This specimen and several others from SE Queensland and far N Queensland raise a number of taxonomic questions. Lack ofa holotype for P. delicatulus and problems faced with obtaining topotype specimens may make the process of revising this species difficult (S. Van Dyck, pers. com.). Acknowledgements This work was part of a larger study of rabbit calicivirus funded by Land Protection Branch of the Queensland Department of Natural Resources. Thanks go to Steve Van Dyck for identifying the specimen, Technical support was provided by G. Story, D. Moore, W. Gould, A. Gonzalez and S. Pidcock. Literature Cited BRAITHWAITE, R.W, & BRADY, P, 1993, The Delicate Mouse, Pseudomys delicatulus: a continuous breeder waiting for the good times. Australian Mammalogy 16; 93-96, MATHIESON, M., JOHNSON, R. & HOBSON, R. 1999. Additional vertebrate fauna records for Lake Broadwater Conservation Park. Queensland Naturalist 37: 1-3. SANDELL, P.R. & START, A.N. 1999. Rabbit Calicivirus Disease Program Report 4: Implications for Biodiversity. A report of research conducted by participants of the Rabbit Calicivirus Disease Monitoring and Surveillance Program and Epidemiology Research Program. (Bureau of Rural Sciences: Canberra). WATTS, C.H.S. & ASLIN, H.J, 1981, The rodents of Australia. (Angus and Robertson: Sydney). R.A, Palmer, Department of Natural Resources, Robert Wicks Pest Animal Research Centre, PO Box 318, Toowoomba 4350, current address: Department of Zoology and Entomology, University of Queensland, St Lucia 4072; 7 August 2000. THE FOSSIL RECORD OF ELAEOCARPUS L. FRUITS MARY E. DETTMANN AND H. TREVOR CLIFFORD Dettmann, M.E. & Clifford, H.T. 2000 06 30: The fossil record of Elaeocarpus L. fruits. Memoirs of the Queensland Museum 46(2): 461-497. Brisbane. ISSN 0079-8835. Elaeocarpus L. fruit stones from Australian mid-Tertiary sediments are systematically described and their stratigraphic and geographic distributions recorded. The fossil fruit stones comprise 2-9-loculate inner mesocarps, their outer surface with longitudinal sutures and a sculpture ranging from smooth to pitted, baculate/verrucate/echinate/rugulate, or fossulate. The fertile locules have a near apical seed and the seedless locules are usually compressed; thin-walled endocarps dehisce loculicidally and enclose the locules. The seed coat is bitegmic and has a multiplicative tegmen, the outer epidermis of which comprises thick-walled, pitted sclereids. Five types of fruit stones are distinguished on the basis of their surficial sculpture. Types 1 and 2 have verrucate/rugulate surfaces, Type | being distinguished by a higher ratio (>0.1) of sculptural base diameter:transverse diameter of fruit stone than that of (<0.1) Type 2 fruit stones. Types 3, 4, and 5 have pitted, smooth, and fossulate surfaces respectively. A review of fossil fruit stones attributed to Penteune F.Muell., 1874, Pleioclinis F.Muell., 1882, Phymatocaryon F.Muell., 1871, and Rhytidotheca F.Muell., 1871 confirms these categories are congeneric with Elaeocarpus. One new species, E. rozefeldsii, is proposed and type specimens are designated for £. allportii (F.Muell.) comb. nov., E. angularis (F.Muell.) Selling, 1950, E. bivalve (F.Muell.) comb. nov., E. brachyclinis (F.Muell.) comb. nov., E. couchmanii (F.Muell.) comb. nov., £. johnstonii (F.Muell.) comb. nov. (and its junior synonym, £. bassii Ettingsh.), &. /vnchii (F-Muell.) Selling, 1950, E. muelleri Ettingsh., 1886, E. pleioclinis (F.Muell.) comb. nov., and E. trachyclinis (F.Muell.) Selling, 1950. The fossil fruit stone record confirms that Elaeocarpus was represented in the eastern Australian flora as early as the Early Oligocene. Modifications to the distribution range and diversity levels of the genus have occurred in eastern Australia since the Neogene. These involved the loss of taxa with Type 5 stones from Australia and a shift to more northerly areas of eastern Australia of species groups with Types 2, 3, and 4 fruit stones. O E/aeocarpus, fossil fruit stones, Australia, mid-Tertiary. Mary E. Dettmann, Department of Botany, The University of Queensland.St Lucia 4072; H. Trevor Clifford, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia; 8 November 2000. Elaeocarpus L., a genus of ~ 60 or 360 species (Mabberley, 1987) occurring in tropical and warm regions of the Old World excluding A frica has an extensive fossil record from Tertiary sediments of eastern Australia. The fossil taxa are leaf remains that include compressions, impressions and cuticles (Ettingshausen, 1883; 1886; Christophel, 1994; Carpenter et al., 1994) and fruits that occur as permineralised casts and moulds or charcoalified compressions (Mueller, 1871la,b, 1873, 1874a,b, 1878, 1882; Johnston, 1880a,b, 1882; Ettingshausen 1883, 1886; Deane, 1925; Berry, 1926; Kirchheimer, 1935; Selling, 1950; Rozefelds, 1990; Rozefelds & Christophel, 1996a, b, in press; Burrows, 1997). Elaeocarpus-type pollen has been recorded ex- tensively from eastern Australian sediments of late Eocene-Recent age (Luly et al., 1980; Truswell et al., 1987; Kershaw et al., 1994; Blackburn & Sluiter, 1994; Martin, 1998) but the small, psilate, tricolporate pollen are difficult to discriminate from those of certain other Elaeocarpaceae (e.g. S/oanea) and Cunoniaceae, and detailed comparative studies on pollen of the two families are not available. The present paper incorporates a taxonomic account of fossil E/aeocarpus fruit taxa reported from Australia, and documents their known geographic and stratigraphic distributions. Eighteen species of fossil fruits referable to Elaeocarpus have been reported from mid-Late Cainozoic sediments of eastern Australia, and one is newly described herein. STUDIES ON FOSSIL FRUITS OF ELAEOCARPUS In 1883 Ettingshausen described fossil fruits from Tasmania under the name Elaeocarpus bassii and was thus the first to attribute fossil fruits to the genus from the Australian Tertiary. However, the earliest reference to fossil fruits consistent with those of E/aeocarpus dates from Mueller & Smyth’s (1870:390) reference to a ‘five valved capsule of an unknown genus’ recovered from deep lead sediments at Haddon, Victoria. Specimens answering to that description were subsequently attributed to Penteune, which name is ‘an allusion to the five valves in which the seeds are imbedded’ (Mueller, 1874a: 41), Mueller is the author of 10 other fossil fruit species consistent with Elaeo- carpus and instituted (1871a,b, 1874a, 1882) Rhytidotheca, Phymatocaryon, Penteune and Pleioclinis, to accommodate them. Pleioclinis was first used without a description or illustration in a note accompanying the description of Rhytidotheca pleioclinis (Mueller, 1873), and was formally described later (Mueller, 1882). Mueller’s generic diagnoses must be treated with caution for they do not always comply with the characters displayed by the species he attributed to them. For instance, Penteune was proposed for fruits with surface sculpture ‘very slightly rough on the dorsal part’ (Mueller, 1874a: 41), but on the same page, P. trachyclinis, is described as ‘externally very rough, almost verrucular’. For Pleioclinis seed placentation was diagnosed as pendent (Mueller, 1882: 43), but in Rhytidotheca pleioclinis F. Muell., which was later transferred to Pleioclinis (Mueller, 1882), the seeds are said to be erect. Furthermore, fruit-valve number, a character that initially discriminated the genera, was later found to be variable; original diagnoses were not emended. Mueller had difficulty in discriminating between his genera. For instance, he (Mueller, 1874b: 42) emphasised the sculptural similarity between the 5-valved Penteune trachyclinis and the 2-4 valved Phymatocaryon mackayi, but later (Mueller, 1875: 41) reported that fruits of P trachyclinis with 4 valves bore much resemblance to P. mackayi. Subsequently, he (Mueller, 1882: 43) observed that P. trachyclinis MEMOIRS OF THE QUEENSLAND MUSEUM resembled rare 5-valved specimens of Pleioclinis couchmanii, the diagnosis of which specified 7-9, rarely 6 valves. It is also likely that Mueller had doubts that Penteune and Rhytidotheca were discrete genera as he noted for Penteune clarkei that ‘some affinity of these fossils to those of the genus Rhytidotheca is evident (Mueller, 1874b: 41). Thus, over 8 years Mueller directly or in- directly interlinked Rhytidotheca, Penteune, Phymatocaryon and Pleioclinis all of which are here associated with Elaeacarpus (Table 1). Because Mueller possessed so wide a knowledge of the Australian Flora it is of interest to speculate as to why he created these 4 fossil genera. In discussions accompanying the generic descriptions, Mueller made it clear he was of the opinion that it was improper to assign frag- mentary fossil material to extant genera. However, for Penteune, Phymatocaryon and Rhytidotheca he suggested affinities with Sapindaceae amongst living taxa thereby strengthening the view the fossil genera were related. His reasons for selecting the Sapindaceae are spelled out in the discussion accompanying the description of Phymatocaryon (Mueller, 187la: 47). There he noted that the pendent seeds, locular dehiscence, drupaceous fruits (‘with a distinct sarcocarp and putamen’) are all family characters of the Sapindaceae. Furthermore, he observed that the fruits of some members of the family also have surfaces that exhibit ‘tubercular roughness’. He may have been particularly swayed by this character since 5 years earlier he had described Cupaniopsis tomentella (F.Muell. ex Benth.) S.Reyn., fruits of which are described by Reynolds (1985) as ‘valves, thick rugose, deeply wrinkled outside’. Nonetheless Mueller was not entirely satisfied his fossil fruit genera all belonged in the Sapindaceae for a few months later he noted the similarity of Rhytidotheca to the fruits of Flindersia and Chloroxylon (Mueller, 1871b: 39) then placed in the Meliaceae. He discounted an TABLE 1. The three principal characters stressed by Mueller in his original diagnoses of fossil fruit genera now associated with Elaeocarpus. Bracketed valve numbers designate the less common state(s); bracketed shape and sculptural designations comply with terminology used herein. Genus Valve number Shape — Surface eee ; Ovate-globose to broadly ovate (spherical- | Slightly rough-deeply wrinkled (pitted- Penteiner (4) 5 (6) __|__ prolate ellipsoidal) ae i aecrussteruwlntey. at Rhytidotheca 5 Ovate (perprolate ellipsoidal) __ Wrinkled (rugulate) ) Phymatocaryon | 2-3 (4) “sh Spherical to ovate (spherical-prolate) 7 ontecetadeee (baculate/ Pleioclinis —_ (5-6) 7-9 Ovate globular-ovate (spherical-prolate) Tubercular-rough (verrucate-rugulate) | FOSSIL FRUITS OF ELAEOCARPUS affinity with the Sapindaceae because ‘the number of fruit-valves, increased to five, remains exceptional’. Though most Sapindaceae have 3-loculate fruits there are many with fewer or more carpels and so it is surprising Mueller was so concerned on that account. Likewise, in his discussion of the affinities of Penteune he wrote, ‘It belonged, however, most probably to Sapindaceae, although the possibility of it having formed a genus of the meliaceous order is not excluded’ (Mueller, 1874a: 41). Why Mueller failed to recognise the similarity of at least Penteune with fruits of Elaeocarpus is difficult to understand unless it stemmed from him being familiar, as is likely, only with fresh material that had been pressed or stored in preservative. In these circumstances the fruit stones do not disintegrate. They do so after prolonged exposure to wet/dry weathering cycles and/or degradation by fungi and/or insect attack. Under these circumstances disintegration of the stony mesocarp into segments may occur after loss of vascular tissue from the central cylinder and the radial strands that connect the central cylinder to the segment sutures. It is likely that fossil fruits were subjected to similar degradation processes during incorporation into sediments. Moreover, many of those from deep lead sediments have been pyritised, and oxidation after recovery and storage may cause the meso- carp segments to separate. Thus, Mueller was very likely distracted from the real identity of the fossil fruits because many dehisced into segments as a result of burial, fossilisation, and subsequent retrieval and storage. By 1884 Mueller was aware that Rhytidotheca fruits were considered consistent with those of Elaeocarpus because R. johnstonii F.Muell.(in Johnston, 1882) was based on material assigned by Ettingshausen (1883) to E. bassii Ettingsh. Nonetheless, he did not comment on or challenge Ettingshausen’s assignment. Likewise it is surprising that Ettingshausen (1883: 63), having recognised that E. bassii and R. johnstonii were conspecific, relegated other species of Rhytido- theca (R. lynchii, R. pleioclinis) to Incertae sedis without comment. Furthermore, he did not suggest that Penteune and Elaeocarpus might be congeneric and followed Mueller in assigning the former to Sapindaceae (Ettingshausen,1883:16) rather than Tiliaceae in which at that time Elaeocarpus was included. In a lecture to the Royal Society of Tasmania, Mueller (1884) restated his argument (Mueller, 1871a) for not 463 including fossil leaves and other plant organs in extant genera. The likely affinity of Phymatocaryon mackayii F.Muell., 1871 with E/aeocarpus was noted by Deane (1925) and confirmed by Kirchheimer (1935) after detailed comparison of fossils and mesocarps of extant £. angustifolius. Selling (1950) formally transferred several of Mueller’s species to Elaeocarpus without comment. As noted by Selling (1950) transfer of Rhytidotheca lynchii, Penteune clarkei, P. allportii, P. brachyclinis, and P. trachyclinis to Elaeocarpus rendered Penteune and Rhytidotheca superfluous. Rozefelds (1990) and Rozefelds & Christophel (1996a, b; in press) provided evidence for reference of several fossil fruit taxa to Elaeocarpus. These include E. clarkei, E. spackmaniorum, E. cunningii, and E. mackayii. However, they expressed doubts about a relationship, as suggested by Selling (1950), of Rhytidotheca lynchii, Penteune trachyclinis, and Phymatocaryon angulare with Elaeocarpus. MATERIAL Fruits reported upon here include charcoalified material from subsurface sediments near Moranbah and Blackwater in central Queensland and permineralised material from Glencoe (Roz- efelds, 1990; Rozefelds & Christophel, 1996b, in press) that are held in the Queensland Museum (QMF); charcoalified and lignified fruits in the Australian Museum (AMF), the Department of Mines, Geological Survey of New South Wales (MMF), and the Museum of Victoria (NMVP). The last mentioned collection includes most of Mueller’s Victorian material originally housed in the Geological Survey of Victoria. Mueller’s material came from deep lead sediments in Vic- toria and New South Wales and specimens illustrated by Mueller have been identified for most of his Victorian species. Mueller’s New South Wales material may have been destroyed in the Garden Palace fire in 1882 which included ‘the palaeontological specimens of the recently deceased Reverend W.B. Clarke’ (Gilbert, 1986: 107), Clarke collected several of the fruits described by Mueller from New South Wales. The reposit- ory of type material of £. mue//eri Ettingshausen, 1886 is also unknown. Tasmanian material attributed to E. bassii by Ettingshausen (1883) and to Penteune allportii and Rhytidotheca johnstonii by Mueller (in Johnston, 1882) has not been located. Other material not examined includes Rhytidotheca major Deane, 1925 and E. cerebriformis Rozefelds & Christophel, 1996b. 464 LOCALITIES. (Fig. 1). Queensland. a) Picardy Station, near Moranbah 21°5°17.6"S 147°50°34.3”E. Holes RDPD98MA 17, 111-133m and RDPD98MA21, 123-133m. Fruits were recovered from both boreholes from near the base of a thin sequence of sands and silts with interbeds of lignites that occur beneath bas- alts and overlie Permian Coal Measures (Fig. 2A). The basalts are probably related to those of the Nebo Province dated as 30-34 Ma. (Wellman & McDougal, 1974, Sutherland pers. comm.), but stratigraphic relationships between these and those intersected in the boreholes have not been established. Host sediment was not retained for palynological assessment. Thus, the minimum age 1s tentatively suggested as Early Oligocene. b) Near Blackwater 24°1°1.3"S 148°48°50”E. South Blackwater Coal Pty Ltd Hole R8736, 82m. Fruits are from ligneous bands within sands that underlie basalts and overlie Permian sediments (Fig. 2B). Basalts to the SW in the Springsure area are dated at 27-33Ma. (Suther- land et al., 1977), but stratigraphic relationships between dated basalts and those in borehole R8736 are uncertain. Moreover, no sediment was available for palynological analysis and a minimum age of Early-Late Oligocene is tentatively indicated. c) Glencoe Station (23°36’S 148°06’E), near Capella (Rozefelds, 1990; Rozefelds & Christophel, 1996a, in press). These occur in silcretes that overlie undated basalts. However, volcanics near Emerald are dated as 30-32Ma. and those southwest of Capella at 26Ma. imply a maximum Oligocene age, possibly Late Oligo- cene-Early Miocene according to Rozefelds (1990). New South Wales. Elaeocarpus-type fruits are known from a scatter of localities on the western flanks of the eastern highlands of northern and central New South Wales (Fig. 1). a) Newstead near Elsmore (29°47’S, 151°1L7’E), Ettingshausen (1886) described E. muelleri fruits from ironstones overlain by basalts. Pickett et al. (1990) concluded an Early Miocene or younger age based on K-Ar dates of 20.5 +0.2Ma. for a nearby basalt flow. b) At Witherden’s Tunnel, near Emmaville leaf fossils attributed to E. muelleri were recorded from carbonaceous clays and silts beneath basalts (30.4+0.3Ma.); palynological dates confirm a Late Eocene age (Pickett et al., 1990). However, there is no evidence to suggest that the leaf and MEMOIRS OF THE QUEENSLAND MUSEUM a 10°S — Picardy Station Glencoe Station Blackwater Elsmore Gulgong Dubbo Orange . Bathurst . Carcoar Eldorado Talbott Guildford Boola Boola Foster Tanjil Yallourn Haddon Beaconsfield Launceston Geilston Bay Bethany 20°S - 2SGo0artgmMaHN= ere er anarany COaNanaON 30°S — ere 40°S— 140°E 150°E | | FIG. 1. Map of Australia showing localities from which fossil fruits of E/aeocarpus have been reported. mesocarp fossils from the separate localities derived from the same plant taxon. c) Gulgong district (32°12’S, 149°32’E) sites include Home Rule Lead and Black Lead (Mueller, 1876, 1877, 1879). The leads are sediment fills of palaeodrainage channels on Palaeozoic basement and are overlain by basalt flows. Isotope ages (K-Ar) of the basalts range from 13.8+ 1.2Mato 14.8 + 1,2Ma. (Dulhunty, 1971; Meakin & Morgan, 1999). The Home Rule Lead is assigned to the Middle Miocene Triporopollenites bellus Zone (McMinn, 1981) in agreement with isotope dates. d) At Orange (33°17’S, 149°06’E), Carcoar (33°37’S, 149°08’E), and Bathurst (33°25’S, 149°35°E) charcoalified fruits are known from beneath basalts in numerous deep leads and reef mines, As at Gulgong the sediments are overlain POSSIL FRUITS OF FLAEOCARPUS 465 Rio Tinto Exploration South Blackwater Coal Pty Lid A Hole ROPD98MA21, Picardy Station B Hole R8736, near Moranbah near Blackwater 5 gepth im) qo vot im) TERTIARY basic volcanics =e lignite sand with 15 EHS] lanite bands fruit bearleg horizan interval nn fruits sandstone ~=PERMIAN m= ~ PIG, 2, Stratigraphic sequences and sampling horizons in: A, Hole ROPD998MA17, Picardy Station near Moranbah, Queensland; and B, South Blackwater Coal Pty Lid Hole R8736 near Blackwater, Queensland. by basalts that were extruded from Mt. Canabolos 5W of Orange, Basalts that overlie the sediments at Forest Reefs and in the Lucknow Mine have provided dates of |1.2-|3Ma., late Middle-carly Late Miocene (Johnson, 1989). Vietavia. Fossil truiis have been collected from buried placer deposits m the Murray, Gippsland and Oiway Basins. and on the northern Nanks of the Great Divide. There are few data on precise ages of the deep lead sediments. a) The Eldorado deep lead (Ted Ovens G & TM Co, Shalt), Beechworth (37°18'S 146°32°E), in the Murray Basin, (Mueller, 1874e: fig. 2) is al unknown age. Palynological dates on other Mur- ray Basin deep leads near Stawell, W Victoria and in the Woodstock 10008 bore near Bendigo, Vic- toria, indicate Oligocene and late Early Miocene ages based on reference ol the former to the lower Proteacidites riberculatus spore-pollen Zone and of the latter to the upper part of that zone (Parindge & Wilkinson, 1982; Partridge, 1993). b) The bulk of fruits described by Mueller were obtained from deep lead sediments at Smythe’s Creek (Reform Co. Shalt) and Nintingbool (Crucible Co. Shaft), near Haddon (37°18'S 146°32°E), SW of Ballarat. Silty sands of the ‘wash dirt’ overlying basement near the bottom of the shalis (Reform Co, Shati. ~47,.5m (1S56ft); Crucible Co. Shalit, ~23.2m (76tt)) are desig- nated as the source ol the fruits (Mueller, 18740: 29, Map |), and in the Reform Shaft the sedi- ments are beneath basalts. Dates of 2-5Ma. have been obtained from basalts that overtie alluvial sediments in the Ballarat district (Sutherland, 1995) and palynofloras recovered from sedi- ments interealated between two of the flows indivate the Trhuliflaridites pleistocenicus spore-pollen Zone of late Pliocene to Pleistocene age (Partridge, 1995), The minimum age of the subbasaltic sediments is thus Late Miocene age, but Rozelelds & Chnstophel ( 1996b) argue foran Early-Middle Miocene age based on association of the fruit E. mackayii (F.Muell.) Kirehheimer, E. clarkei (F.Muell.) Selling, and Sponedvlo- sirobus symthii F. Muell, (see also Greenwood et al., 2000), Unfortunately, Mueller did not always specily precise depth or mine shafi for localities at Haddon. ¢) Gippsland Basin deep lead sediments al Talbot (37° LOS 146° 14°F), Foster (37°10'S 146°) 4E) and Tanyil (38°01S 146°14°E). Thus far, the ages of these have not been resolved, but as for the Haddon sediments Rozefelds & Christophel (1996b) suggest an Larly-Middle Miocene age. 466 rugose f FIG. 3. Stylised transverse section of Elaeocarpus fruit in which each of the 5 carpels has different surfical sculpture of the inner mesocarp. a, axis; cl, compressed locule; e, endocarp; en, endosperm; em, embryo; ex, exocarp; f, fibres; im, inner mesocarp; om, outer mesocarp; I, locule; r, raphe; s, suture. d) Yallourn Coal Measures, Yallourn (38°10’S 146°21°E) in the Yallourn Formation whithin the Triporopollenites bellus spore-pollen Zone of late Early-Late Miocene age (Rozefelds & Christophel, 1996b). South Australia. Fruits from subsurface sedi- ments near Bethany (34°32’S 139°00’E) in the Barossa Basin (Paterson in Hossfeld, 1949) occur in the Rowland Flat Sand, a lignitic facies over- lain by coarse-grained fluviolacustrine sand and gravel fining upwards to bedded silt and clay. Palynological dates are Early Oligocene-Early Miocene for the basal sediments and Early-Late Miocene for coarser upper facies (Alley, 1995). Tasmania. Deep lead sediments at Brandy Creek, Beaconsfield (41°12’S 148°49°E) and outcrops at Launceston (41°27’S 147°10°E), A mid- Tertiary (Oligocene) age is indicated for these localities based on palynofloras of sub-basaltic sediments in the Tamar Graben (Forsyth, 1989). b) Calcareous fruits from Geilston Bay, Hobart (41°12’S 148°49’E) occur in travertine which is equivalent in age to nearby basalts dated as 22.4 Ma, Early Miocene (Tedford et al., 1975). New Zealand. a) At Mangonui (35°00’S_ 173° 20°24”E), north of Auckland, charcoalfied fruits occur Mangonui Formation lignites (Late Mio- cene) associated with Cocos zeyvlandica Berry (Berry, 1926; Thompson, 1978; Isaac etal. 1994). MEMOIRS OF THE QUEENSLAND MUSEUM b) At Schultz Creek (42°16°48”"S 171° 07°12”E), north of Greymouth, South Island lake sediments deposited during the last interglacial (100Ka.) have yielded fruits comparable to those of extant FE. dentatus (Burrows, 1997). The material illustrated has been deposited at the Queensland Herbarium. METHODS Charcoalified specimens were photographed after whitening with ammonium chloride; internal characters including those of the locules were photographed without whitening. Mueller’s type specimens are figured together with reproduction of the original lithographic illustrations, the latter of which represent mirror images. Available seed coats were examined in transmitted light after clearing in chromium trioxide for several hours, followed by thorough washing in distilled water after which they were mounted on glass microscope slides in glycerine jelly. Fruit stones of extant E. angustifolius Blume and £. reticulatus Smith were collected from trees growing at the Brisbane Botanic Gardens, Mt Coot-tha and the University of Queensland. SYSTEMATIC PALAEOBOTANY Elaeocarpus L.1753 Rhytidotheca F Muell., 1871:39. Phymatocaryon F .Muell,, 1871:41. Penteune F Muell., 1874a:3941. Pleioclinis F Muell., 1882:43 TYPE SPECIES. Elaeocarpus serratus L. FRUITS OF EXTANT ELAEOCARPUS. In extant Elaeocarpus fruits develop from flowers with a superior 2-8 (usually 3-5) loculate ovary that terminates in a single style surmounted by a lobed stigma. There are 2-12 anatropous ovules per locule and these are attached to an axile placenta. When there are few ovules per locule they are attached high up on the axis and so appear to be subapical (Figs 4, 6B,C). Only one ovule per locule develops into a seed and in some species there is only | seed per fruit. Expansion of this 1 seed usually results in compression of the adjoining seedless locules (Figs 4, 5D,G). The fruit is a drupe, a useful term, ill-defined but widely used in the literature (Clifford & Dettmann, in press). In most species the fruit surface is iridescent blue, the colour due ‘not to a blue pigment, but by the structure of the cuticle which reflects blue light’ (Lee, 1991). The outer fleshy mesocarp is usually thin and densely FOSSIL FRUITS OF ELAEOCARPUS 467 Na FIG. 4.Seetions through Alacacurpus wagustifolius Blume fruits: A, transverse; B, vertical. Labels as for big. 3. fibrous, fibres mtertwined with the sculptured surface of, or extending into, the woody inner mesocarp (Fig. 5A). The latter encloses the locules and their surrounding endocarps to form a stone whose outer surface may be smooth, pitted. fossulate, rugulate, baculate, verrucate, or echinate. At maturity the vascular strands of the axis. decay, resulting ina hollow cylinder which extends for ~ two-thirds the length of the mesocarp from its proximal end (Figs 3, 4, 5F). Further decay of radially disposed strands be- neath the segment sutures results in the mesocarp and enclosed endocarps splitting vertically between the septa to produce segments that expose seeds or aborted ovules on their radial walls (Fig. 5G-1), As was recognised by Mueller, such dehiscence ts loculicidal. The lines along which the mesoearp split are usually clearly marked by vertical sutures on the surface of the mesocarp (Fig. 5A-C) and each segment consists of a woody, mner mesocarp, half of 2 adjacent endocarps and associated Jocules (Fig. 5H,1). The inner surface of the endocarp may bear multi- cellular scales ahd hairs (Comer, 1976). Seeds are fusiform, bitegmic with glabrous or hairy surfaces. The testa 1s several cells thick with an outer epidermis of elongate to iso- diametric cells (Figs 61, 7F) that in some species are lignified. The tegmen is multiplicative, the outer epidermis of which has fibriform, bulbous or dumb-bel! shaped, lignilied selereids that are arranged longitudinally (Fig. 5G.H), Both the testa and tegmen are yascularised in the chalaza which forms the woody basal and acute prominence of the seed (Corner, 1976). Shape, structure and surface sculpture of the fruit stones are uselul species discriminators (Rozefelds & Christophel, 1996a). However, such features have been little utilised and remain largely undocumented in current infrageneric classifications, Moreover, litthe attention has been accorded anatomical features of the inner mesocarp wall and the enclosed endocarps. The only record of fossil mesovarps attribut- able to an extant species is that of £. denfanns stones in interglacial (100Ka) sediments from New Zealand (Burrows, 1997), FOSSIL FRUITS. The following fossil taxa ure considered congeneric with Elaeocarpus. PENTEUNE F.Muell., 1874: lype species (designated here) Penteune clarkei F.Muell., 1874a: 41 from Elsmore, NSW; Early Miocene: Neotype (designated Rozetelds & Chrisiophel, (996a: 43). NMVP53913, Fig. LIA, B; Mueller’s figured specimens from Haddon (Smythe’s Creek, ?Reform Co. Shafi, ~47.5m) are believed lost (Rozefelds & Christophel, 1996a: 43). Species included: P af/portij F.Muell., in Johnston 1882: figs 40,41, 44-47.P. hrachvelinis F. Muell., 1874a: 41. pls, figs 1-9, P. trachyelinis F. Muell,, 1874a: 41, pl. 8, figs 10-17. PHYMATOCARYON ¥.Muell., 1871; type speciesPhymatocaryan mackayiF.Muell., 187 la: 41 (by monotypy) from Haddon (Smythe's Creek, Reform Co. Shall, ~47.5m), sediments beneath basalts; ?Early-Middle Miocene: Leetotype (designated Rozefelds & Christophel (in press)) NMV53562, Mueller 187 1a: 41, pl. 2, fig.4: Species included:P angulare F.Muell.. 1874b: 41, pl. 10, figs 1-4. P bivalve F.Muell., 1877: 180, 1878: 39, pl. 15, figs 6-9. 468 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5. Fruit stones of E/aeocarpus angustifolius Blume. A-C, apical, lateral, and basal views showing sculpture of woody inner mesocarp with attached fibre (f) bundles and sutures, x 2. D, photograph of transverse section in Fig. 4A of fresh, mature fruit stone showing sculptured woody inner mesocarp with fibre (f) bundles on the surface, enclosed endocarps, two with fertile locules containing seeds, each showing the raphe, the other three locules compressed, X 2. E, photograph of vertical section in Fig. 4B showing vascular strands in the axis, X 2. F, vertical section portion of inner mesocarp removed to reveal closely adpressed endocarp, x 4. G, internal view of two segments of partially rotted mature fruit stone showing seed in fertile locule (left), compressed sterile locule (right), and hollow axis, x 4. H, I, internal views of dehisced segment comprising inner mesocarp with halves of two adjacent locules and endocarps showing ridges and grooves on their inner surface, x 4. Labels as for Fig. 3. FOSSIL FRUITS OF ELAEOCARPUS 469 FIG. 6. Micrographs of fruit stones of Elaeocarpus angustifolius Blume. A, transverse section of inner mesocarp and endocarp walls, and profile of a concave-crested, arched sculptural element at outer surface of inner mesocarp, X 10. B,C, detail of compressed locule with 2 near-apical, axially attached, aborted ovules (0), and showing coalescence in the apical region of locule of ridges and grooves on inner surface of endocarp, * 10, and x 25 respectively. D, scanning electron micrograph of section of inner mesocarp and endocarp, x 30. E, F, scanning electron micrographs of outer surface and transverse section of inner mesocarp, * 2000 and x 4000 respectively. G, outer epidermis of tegmen showing elongated sclereids with thick, pitted walls, x 750. H, subspherical sclereid from tegmen, X 500. I, outer cuticle of testa, with a sclereid (sc) arrowed derived from outer epidermis of tegmen, x 250, Labels as for Fig. 3. 470 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 7. Fruits, fruit stones and seeds of Elaeocarpus reticulatus Smith. A, apical (upper) and basal (lower) views of fruit stone, x 2. B, lateral view of fruit stone, x 2. C, vertical (left) and transverse (right) sections of immature fruits, x 2. D, transverse section of fruit with outer mesocarp partially detached (left) revealing surface sculpture of inner mesocarp, * 6. E, vertical section of immature fruit showing axile placentation and axial tissue, * 16. F, outer cuticle of testa, x 250. Labels as for Fig. 3. PLEIOCLINIS F.Muell., 1882; type species Pleioclinis couchmanii F.Muell., 1882: 43 (by original designation) from Haddon (Smythe’s Creek or Nintingbool), sediments beneath basalts; ?Early-Middle Miocene; Lectotype (here designated). NMVP53920, Mueller 1882: 41; pl. 19, figs 9, 10; Species included: Rhytidotheca pleioclinis F.Muell., 1873), (=P. shepherdi F.Muell., 1882; junior objective synonym) RHYTIDOTHECA F.Muell., 1871; type species Rhytidotheca lynchii F.Muell.,1871b:39 (by monotypy) from Haddon (Nintingbool, Crucible Co. Shaft, ~23.2m), deep lead sediments; ?Early -Middle Miocene. Lectotype (here designated). NMVP6034, Mueller, 1871b:39, pl. 4, fig. 4 (right hand segment) and NMVP6033, Mueller, 1871b: 39, pl. 4, fig. 1, centre segment broken at apex. Species included:R. johnstonii F.Muell.in Johnston, 1882: 50, fig. 6. R. major Deane, 1925: 491, pl. 60, fig.12 (nomen nudum). R. pleioclinis F.Muell., 1873: 42, pl. 6, figs 1-4. REMARKS. Recognition of fossilised Elaeo- carpus stones has been based on sculpture, shape, locule number, and position of seed attachment (Deane, 1925; Kirchheimer, 1935; Selling, 1950; Rozefelds, 1990; Rozefelds & Christophel, 1996a, b, in press). Woody mesocarps of E/aeo- carpus are distinguished from other Elaeo- carpaceae by: a pitted, smooth, fossulate, verrucate, baculate, echinate or rugose surface with longitudinal sutures that delimit segments; 2-9 carpels; passive loculicidal dehiscence into segments; 1-seeded locules; fertile and sterile locules, the latter often compressed; ovules anatropous, pendulous with a ventral raphe; seeds fusiform with a multiplicative tegmen, the outer epidermis with pitted and lignified longitudinal cells. Further characters with apparent interspecific significance include the internal organisation and structure of the woody inner mesocarp and subjacent locules as revealed by extant E. angustifolius and E. reticulatus (Figs 5-7). In E. reticulatus young fruits have an axial column of elongated vascular strands (Fig. 7E). The inner mesocarp consists of isodiametric stony cells, and the enclosed endocarps comprise lignified cells, elongated and arranged tangent- ially. Fruits of £. angustifolius are similar in their organisation (Figs 5,6). Fruit stones allowed to rot on the ground decompose slowly, but after time they may split into segments. Those of £. angustifolius were little affected after subjecting them to conditions that may be expected in a medium-high energy depositional situation. Stones agitated with sand, gravel and water in a closed container on a rotary shaker for 1 week showed no signs of breakage or abrasion of the FOSSIL FRUITS OF ELAEOCARPUS 47] FIG. 8. Elaeocarpus angularis (F.Muell.) Selling, fruit stone. A-G, holotype E, lateral views of external and internal surfaces of 2 segments and basal part of third segment (NMVP53565), x 2. B, F, lateral views of external and internal surfaces of upper part of third segment (NMVP6017), X 2. C, D, apical and basal views of whole fruit stone, x 2. G holotype as illustrated by Mueller (1874b, pl. X, figs. 1-4), but note his images are reversed and seeds represented in his fig. 4 have not been located. external sculptural elements. However, afier drying some dehisced into segments. Elaeocarpus allportii (F.Muell.) comb. nov. 77 Pentenne aliporti F. Muell. in Johnston, |880a: 27 (nom nud.). Penteune allporti F, Muell. in Johnston, 1880b; 85 (nom. nud. ). Penteune allporti F.Muell. in Johnston, 1882: figs 40, 41, 44-47. MATERIAL. LECTOTYPE (here designated): Whole specimen (Johnston, 1882, figs 40-41) and 4 of its segments (Johnston, |882, figs 44-47) from Early Miocene Geilston Travertine at Geilston Bay, ‘Tasmania. Stony mesocarp prolate (20mm long, 16mm wide), broadly elliptical in longitudinal section and circular in transverse section; 5-loculate (illustrated as 6-loculate but in Johnson’s Explanation of Figure corrected to 5-loculate). External surface punctate and with 5 longitudinal sutures; preserved mesocarp wall 2-3mm thick. Seed cavity fusoid, apically acute, 14mm long, 5mm wide. DESCRIPTION. As for lectotype. REMARKS. No description of calcereous specimens included in the species was provided, but Johnston (1882, Explanation of Figures) illustrated what he believed to be ‘probably PF allporti F. Muell, F.v.M., m.s. Mueller (1874a: 41) noted, under Penteune clarkei F.Muell., that ‘an extremely similar fossil has been discovered in Tasmania by Morton Alport (sic), Esq., at Gerlston-Bay (sic) in tertiary travertine’. Selling (1950) suggested inclusion of P allportii in Elaeocarpus, but did not formally make the transfer. COMPARISON. May be synonymous with, £. clarkei (Johnston, 1880b: Mueller, 1874a: 41). DISTRIBUTION. Geilston Bay, Beaconsfield (Brandy Creek), Tasmania (Johnston 1880a,b, 1882). AGE RANGE. Oligocene-Early Miocene. AFFINITY. Possibly with £. bancroftii F.Muell. & Bailey. Elaeocarpus angularis (F.Muell.) Selling, 1950 (Fig. 8A-E) Phymatocarvon angulare ¥ .Muell. 1874b: 41, pl. 10, figs 1-4. Elaeocarpus angularis (F Muell.) Selling 1950: 558. MATERIAL. HOLOTYPE (by monotypy): NMVP53565 (2 segments plus basal part of third) and NMVP6017 (upper part of third segment) (Fig. 8A-E) from ?Early-Middle Miocene at ~47.5m in Reform Co. Shaft at Smythe’s Creek, Haddon, Victoria. Preserved mesocarp ovoid (22mm long, 19mm wide), oval acuminate in longitudinal section and convexly triangular in transverse section; 3-loculate and with 2 equally developed locules exposed, one of which contained a seed now lost. External surface with 3 longitudinal sutures embedded in protruding ridges that extend from base to apex; intervening surface with low relief striations and pits that may be artefacts of abrasion. Mesocarp wall 2-3mm thick with isodiametric cells; endocarp ~0.5mm thick, composed of tangentially aligned cells. Central axis of mesocarp composed of a hollow cylinder of elongated vascular strands that extend from stalk scar to near the apex. Seed cavity fusoid, the apex acute, 10-12mm long, 3-4mm wide. Seeds fragmented, anatomical detail not determined, DESCRIPTION. As for holotype. REMARKS. Additional material reported from NSW (Mueller, 1883) has not been located. COMPARISON. E. angularis is similar in size and mesocarp sculpture to E. (Penteune) brachyclinis, but was distinguished on its 3 rather than 5 locules (Mueller, 1874b). With 3 locules and position of the seed the taxon was accom- modated in Phymatocaryon (Mueller, 1874b: 42), E. (Phymatocaryon) bivalve was separated from £. angu/are on its 2 segments (valves sensu Mueller), the edges of which are acute and wing- like and in which the sutures are embedded (Mu- eller, 1877, 1878). Moreover, in E. bivalve | of the 2 locules is compressed, unlike E. angularis. DISTRIBUTION. Haddon (Smythe’s Creek, ?Reform Co. Shaft, ~47.5 m), Victoria; Beneree, Dubbo, NSW (Mueller 1874b, 1883). AGE RANGE. ?Early-Late Miocene, MEMOIRS OF THE QUEENSLAND MUSEUM AFFINITY. As suggested by Selling (1950) with Elaeocarpus. However, the lectotype has equally developed locules and the seed coat structure is unknown; an affinity with the Euphorbiaceae is possible (Rozefelds pers. comm.). Elaeocarpus bivalve (F.Muell.) comb. nov. (Fig. 9A-L) Phymatocaryon bivalve F.Muell. 1877: 180. Phymatocaryon bivalve ¥ Muell.1878: 39, pl. 15, figs 6-9. Phymatocaryon bivalve F Muell.1879: 170, pl, 3, fig. 2. Phymatocaryon bivalve F Muell. 1883: 9, pl. 15, figs 6-9. MATERIAL. NEOTYPE (here designated): MMF36220. (Fig. 9A-C) from the late Middle — early Late Miocene Black Lead at Gulgong, NSW. Mesocarp woody, ovoid, vertical axis 17mm, lateral axes 14 and 21mm; with 2 segments delimited by a longitudinal suture and reflexed at their margins; locules 2, one with a near apical seed, the other compressed; dehiscence loculicidal. Mesocarp wall 4mm thick, compact, with a near smooth undulating outer surface. Locules abut the central axis of vascular strands arranged in a hollow cylinder that extends from the stalk scar to near the apex. Endocarp composed of tangentially oriented elements with shortly stalked bifid hairs on the inner surface. Seed cavity ovoid, 10mm long, 4-Smm wide. DESCRIPTION, Fruit stones woody ovoid, vertical axis 17-20mm, lateral axes 13-16mm and 17-22mm; with 2 segments delimited by flanged and reflexed margins adjacent to the longitudinal sutures; locules 2, one of which is usually compressed, the other with a near apical seed. Mesocarp wall 3.5-6mm thick at segment margin, compact, and with a near smooth undulating outer surface. Locules abut the central axis of vascular strands arranged in a hollow cylinder extending from the stalk scar to near the apex. Endocarp of tangentially oriented elements. Seed cavity ovoid, twice as long as wide; seed with near apical attachment. Seed coat near smooth, testa with outer cuticle of rectangular cells up to 30m long, 20 um wide. Tegmen with outer epidermis of elongated, bulbous cells (up to 100um long, 404m wide) with pitted, lignified walls 2-411m thick. Several layers of underlying layers composed of thin-walled ( 0.1 13H E. holopetalus, E. kirtonti, E. obovatus, (baculate, bastionate, verrucate, echinate | E. reticulatus, E. ruminatus, in part) { _ E. williamsianus, E. sp. noy. 1* Type 2 Surface with raised sculpture; basal di- | & /ynchii, E. trachyclinis E. costatus, E. dentatus, &. elliffii, ameter of elorierte (usualy low Rae E. hookerianus, E. grahamii, verrucae, rugulae, grana); transyerse di- -sericghetealiy ameter of fruit stone < 0.1 (echinate in aia La | part, granulate) | ‘a Type 3 Surface with pits or foveolae (punctate) | £. allportii, E. brachyelinis, E. clarkei, EF. baneroftii, E. linsmithii, E.stellaris 7 |E. peterii, E. rozefeldsii(?) L “fy Type 4 Surface smooth or near smooth (smooth) | £. angulare, E. bivalve, E, pleiaclinis E. ferruginiflorus, E. foveolatus, Ran E E. largiflorens, E. johnsonii, E. thelmae | Type 5 | Surface with fossulae E. couchmanii Not known from Australia and New Zealand (Rozefelds & Christophel, 1996b); neither shape of the fruit stone nor locule number was utilized in their classification. Three of the categories — smooth, punctate and granulose — are well characterised, but mild abrasion of granulose fruit stones may reduce the sculpture to smooth and thus far fossil fruit stones having granulose sculpture remain unrecorded. Moreover, sculptural types disting- uished as ‘verrucate’, ‘echinate’, ‘baculate’, and ‘bastionate’ (Rozefelds & Christophel, 1996b, table 2) may be difficult to uphold as exemplified by E. angustifolius whose fruit stone sculpture is said to be bastionate. The sculpture comprises a mix of bacula (straight-sided, flat crested elements), echinae (tapering, pointed processes), and distally expanded processes with flat, pointed or rounded crests, and these elements may be coalesced to form rugulae (Figs 5, 6). Furthermore, they described the fruit stones of E. reticulatus as echinate, but the elements include both round-topped (verrucae) and pointed (echinae) processes (Fig. 7). Sculptural elements, particularly the distal crests of raised elements, may be substantially modified from their original form by abrasion during transportation. Even so, the bases of elements are likely to be preserved in all but extremely abraded fruit stones. The alternative grouping proposed here for stones with raised sculpture is based on the basal diameter of the sculptural elements relative to the transverse diameter of the fruit stone (Table 2). Many of the fossil taxa with raised sculpture included within Types | and 2 have sculptural patterns composed ofamix of bacula, verrucae, echinae and rugulae, and each includes the baculate, bastionate, echinate and verrucate ornamentation classes of Rozefelds & Christophel (1996b, table 2). Types 3 and 4 accommodate taxa with punctate and smooth fruit stones respectively and Type 5 includes taxa with fossulate sculpture, a sculptural type not represented among extant Australian and New Zealand species (Table 2). The fruit stone fossil record confirms that Elaeocarpus in eastern Australia dates to at least the Early Oligocene (Fig. 22) and corroborates evidence from cuticles and leaves (Carpenter, 1994), There are older (Eocene) records of leaves and cuticle of Elaeocarpaceae (Elaeocarpus or Sloanea) and of Elaeocarpus-like pollen from eastern Australia, but these await detailed sys- tematic resolution (Truswell et al., 1987; Rowett, 1991; Rowett & Sparrow, 1994; Christophel, 1994). A Late Paleocene, leat/cuticle record of the family is known from Cambalong Creek, Victoria (Valdala & Greenwood, in press). Thus far, reliable fossil records of fruits/leaves of El- aecocarpaceae indicate a geographic range from southernmost Tasmania north to central Queen- sland and west to southern South Australia. A range to northern South Australia may be implied by Late Paleocene-Early Eocene pollen (Martin, 1998). This distribution varies from its present range insofar as Elaeocarpus is no longer represented in South Australia or Tasmania. In New Zealand the Elaeocarpaceae has a history extending to at least the early Miocene as attested by leaves (Pole, 1993, 1996), but indubitable fossil fruits of Elaeocarpus are thus far unreported from pre-Quaternary sediments. FOSSIL FRUITS OF ELAEOCARPUS PS Ceca ii WALES Ba 493 NEW ZEA 12345 12345/)12345 FIG. 22. Recorded distribution and stratigraphic range of E/aeocarpus fruit stone types as defined in Table 2. Broken lines indicate age uncertainties of sediments; ? indicates uncertain record. Not only has the genus distribution changed in Australia since the mid-Tertiary, so too have the distributions of species groups that shed the individual stone types (Table 2). Today species with Type | fruit stones are distributed from northernmost Tasmania to N Queensland, but during the mid- Tenaty ranged westward into S South Australia (Fig. 22). Type 2 fruit stones are shed by taxa that today occur in NSW (Lord Howe Island), Queensland and NZ in contrast to their more southerly and westerly mid-Tertiary distribution range of Victoria, NSW and South Australia. Extant species that have pitted fruit stones (Type 3) are restricted to N Queensland and NT, whereas during the mid-Tertiary this type had an E Australian distribution from S Tasmania to central Queensland. The mid- Tertiary distribution of smooth stones (Type 4) included Victoria and NSW, whereas today taxa with smooth stones are restricted to Queensland and regions to the north (Fig. 22). The record also implies a former higher species diversity in the region. Type 5 stones, without living counterpart in Australia, are present in Tertiary sediments of Victoria, NSW and South Australia (Fig. 22). ACKNOWLEDGEMENTS We are particularly grateful to Tim Spencer, South Blackwater Coal and Graham Muggeridge, Sandy Menpes and Howard Rewald, CRA for charcoalified fossils and locality details and to Gordon Guymer, Queensland Herbarium, for advice on the taxonomy of Elaeocarpus. Natalie Camilleri, Queensland Museum provided advice and assistance with photography and preparation of the diagrams. We acknowledge helpful discussions with, and comments from, Andrew Rozefelds who also kindly provided illustrations from his Honours thesis and a manuscript copy of Rozefelds & Christophel (in press). Advice on literature was also provided by Miguel Garcia, Sydney Botanical Garden, John Rigby, Queensland University of Technology, and Anthony Valdala, University of Melbourne. We thank Elizabeth Thompson, Museum of Victoria, Robert Jones, Australian Museum, and lan Percival, Geological Survey of NSW for loan of material. Lin Sutherland and Julian Hollis, Australian Museum provided much helpful advice on basalt ages. MED thanks the Director, Queensland Museum for providing facilities and acknowledges support from The Australian Research Council during tenure of a Senior Research Fellowship. LITERATURE CITED ALLEY, N.F. 1995. Barossa Basin. Pp. 173-175. In Drexel, J.F. & Preiss, W.V. (eds) The geology of South Australia, v.2 (Geological Survey of South Australia: Adelaide). 494 BARNARD, C.E, 1881. Notes relating to certain fossil leaves and fruits found in the auriferous drifts of Gulgong, New South Wales. Papers and Proceedings and Report of Royal Society of Tasmania for 1880: 40-43. 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PARTRIDGE, A. & WILKINSON, H.E. 1982. Palynological examination of samples from the Woodstock 10008 bore, Loddon deep lead. Geological Survey of Victoria Report 1982/11] (unpubl.). PICKETT, J.W., SMITH, N., BISHOP, P.M., HILL, R.S., MACPHAIL, M.K. & HOLMES, W.B.K. 1990. A stratigraphic evaluation of Ettings- hausen’s New England Tertiary plant localities. Australian Journal of Earth Sciences 37: 293-303. POLE, M, 1993. Early Miocene flora of the Manuherikia Group, New Zealand. 5. Smilaceae, Polygonaceae, Elaeocarpaceae. Journal of the Royal Society of New Zealand 23: 289-302. 1996. Plant macrofossils from the Foulden Hills Diatomite (Miocene), central Otago, New Zealand. Journal of the Royal Society of New Zealand 26: 1-39. REYNOLDS, S.J. 1985. Sapindaceae. In George, A.S. (ed.) Flora of Australia, v.25. Melianthaceae to Simaroubaceae, (Australian Government Publishing Service: Canberra). ROWETT, A.J. 1991. The dispersed cuticular Noras of South Australian Tertiary coalfields, Part 1: Sedan. Transactions of the Royal Society of South Australia 115: 21-36. ROWETT, A.I. & SPARROW, A.D. 1994, Multivariate analysis of Australian Eocene dispersed cuticle floras: influence of age, geography and taph- onomy on biozonation. Review of Palaeobotany and Palynology 81: 165-184. ROZEFELDS, A.C. 1990. A mid Tertiary rainforest flora from Capella, Central Queensland. Pp. ]23-136. In Douglas, J.G. & Christophel, D.C. (eds) Proceedings of the Third International Organization on Palaeobotany Symposium 1988. (A-Z Printers: Melbourne). ROZEFELDS, A.C. & CHRISTOPHEL, D.C. 1996a. Elaeocarpus (Elaeocarpaceae) fruits from the Oligo-Miocene of eastern Australia. Papers and Proceedings of the Royal Society of Tasmania 130: 41-48. 1996b. Elaeocarpus (Elaeocarpaceae) fruits from the Early to Middle Miocene of eastern Australia. Muelleria 9: 229-237. In press. Elaeocarpus (Elaeocarpaceae) fruits from the Australian Cenozoic, Alcheringa. SELLING, O.H. 1950. Some Tertiary plants from Australia. Svensk Botanisk Tidskrift 44: 551-561. SUTHERLAND, F.L. 1995. The volcanic earth. (University of New South Wales Press: Sydney). SUTHERLAND, F.L., STUBBS, D. & GREEN, D.C. 1977. K-Ar ages of Cainozoic volcanic suites, Bowen-St. Lawrence hinterland, north 496 Queensland (with some implications for petrologic models). Journal of the Geological Society of Australia 24: 447-460. TEDFORD, R.H., BANKS, M.R., KEMP, N.R. & SUTHERLAND, F.L. 1975. Recognition of the oldest known fossil marsupial from Australia. Nature 225: 141-142. THOMPSON, B.N. 1978. Tertiary Stratigraphy. Pp. 443-449. In Suggate, R.P., Stevens, G.R., Te Punga, M.T. (eds) The geology of New Zealand (Government Printer: Wellington). TRUSWELL, E.M., KERSHAW, A.P. & SLUITER, IL.R. 1987. The Australian-south-east Asian connection: evidence from the palaeobotanical MEMOIRS OF THE QUEENSLAND MUSEUM record. Pp. 32-49. In Whitmore, T.C. (ed.) Bio- geographical evolution of the Malay archipelago. (Clarendon Press: Oxford). VALDALA, A.J.V. & GREENWOOD, D.R. (in press). Australian Paleogene vegetation and environments: evidence for palaeo-Gondwanan elements in the fossil records of Lauraceae and Proteaceae. In Metcalfe, I., Smith, J.M.B., Morwood, M. and Davidson, I. (eds) Faunal and floral migrations and evolution in SE Asia-Australasia. (A.A. Balkema: Rotterdam). WELLMAN, P. & McDOUGALL, I. 1974, Cainozoic igneous activity in eastern Australia. Tectonophysics 23: 49-65. FOSSIL FRUITS OF ELAEOCARPUS APPENDIX 1: Register of figured specimens, * denotes nomenclatural type. 497 fius 10, Lectotype * |___Taxon, Fig, No. | Original Figure Specimen Status Locality > _| Repository/ Reg. No. | E angularis BA-F Mueller, 18740c, pl. X, Holotype * Haddon (Smythe’s Ck), Vie | NMVP53565,6017 | E, bivalve SAC | Beats Neatype * Black lead, Gulgong, NSW MMEF36220 | 9D-G_ - Forest reets, Gulgong, NSW _ AMF6669 oe - 7 | Black lead, Gulgong, NSW __MMF36221 | OK 7 : Forest reefs, Gulgong, NSW __AMF6669 £. 1OA-C | Mueller, 1874a, pl, VITI, Lectotype * Haddon (Symthe’s Ck), Vie NMV P6060 brachyelinis | fig. 4 iid are : ~ mie 7 1OE-G.L - | Haddon. Vic NMVP212639 WH = Haddon, Vie NMVP6025 ; WIL . Haddon. Vic NMYVPS53918 EL elarkei }1A,B | Rozefelds & Christophel, = Neotype * Elsmore, NSW AME9281 | 1996, pl. 1, figs A-C See Saree ry HC-l | ’ 7 ¥ “ 4 | Victory Mine, Orange. NSW AMP844U) z= E_ eontehmanii | 12A.€ | eesti 1882, pl. XEX, Lectotype * | Haddon (Nintingboo!), Vic | NMVP53920 - IEG : _|taddon, Vie NMVP53921 E, cunningii | 13A-D_ Rozefelds. 1990, fig. 6D Holotype* ———_| Glengoe, Capella, Qid a QMFI6768 | I3EFG 7 a ee Picardy Station, Moranbah, Qld | QMF30114 13H | Roxefelds 1990, fig, 6E Paratype Glencoe, Capella, Qld __OMPSOLIS eee af = _Gleneoe, Capella, Qld QMESOLI6 | 13K | Glencoe, Capella, Qld QMF50117 _ - [ [a = i: - a ‘Glencoe. Capella. Qld ] | QMFS0118 Fi. lunehii 14A-C | Mueller 1871b, pl LY. figs | Lectotype * (2 segments) | Haddon (Nintingbool), Vic NMVP6034 _ : m Lectotype * (1 seument) Haddon (Nintingbool) Vie _NMVP6035 | dik “Haddon, Vie | NMVP53969 MLO] =i |e ‘Smythsdale. Vie —QMFS01I9 L mackayii ISA\B.1 | Mueller, 1871. ph IL, fig, Lectotype * | Haddon (Smythe’s Creek), Vie | = NMVP53562 ~~ | 4: Rozefelds & “ 5 eT | eae I5C | Christophel. in press, fi: = _| South Blackwater, Qld | OMF 50120 sp 2AB - Picardy Station, Moranbah, Qld | QMPS0121 ISE-G a __| Pieardy Station, Moranbah, Qld _ QMFSU122 i isn === Picardy Station, Morandah, Qld OMPS1078 \E. peterti Rozelelds & Christuphel, Holotype * Glencoe, Capeila, Old OMF | 8088 | 16A-D | 1996b, pl. 3. fuss | AYC.E,G,I _ ! 7 L E, pleioelinis (7A-C | Myeller; 1873. pl. VI. figs Lectotype * Haddon (Nintingbool), Vic NMVPS53747 EG | Haddon, Vie NMVP53741 Hd a Haddon, Vie __NMVP53741 | (Wael | Haddon. Vie NMVP53741 | E rozeteldsii | (BAC | Mueller, 1873, pl. VI, figs | Holotype * _ South Blackwater, Qld QMF350123 isp |? lL | South Blackwater, Old OMPS0124 8D | South Blackwater, Qld OMF50125 a [SEF | __| South Blackwater, Qld QMF30126 E. spackman- 1I9A-C Rozetelds, 1990, figs 4A.B | Holotype * Glencoe, Capelli. Qld QMF15440 vorant | 19D-£,1 | Rozefelds, 1990, fius 4E.F Paratype Glencoe. Capella. Qld |S QMFIS442 19G-1_ | Rozefelds, 1990, fig. 4G Paratype | Glencoe. Capella. (Id QMF 15444 _19K-M 4 |Glencoe, Capella, Old QMF30127 —20A-D ‘Picardy Station, Morandah, Qld | QMFS1076 —_ UEP 7 Picardy Station, Morandah, Qld QMFS1077° | Etrachyelinis IA Haddon, Vic : ——_ _ NMYVP53984 _21B,C = Haddon (Smythe's Creek) Vie _NMVP53922 aip.p | Mueller, 1874a, pl. VIL Haddon, Vic NMVP53758 DUSK CHORUSING BEHAVIOUR IN CICADAS (HOMOPTERA: CICADIDAE) AND A MOLE CRICKET, BRISBANE, QUEENSLAND TONY EWART Ewart, A. 2001 06 30; Dusk ehorusing behaviour in cicadas (Homoptera: Cicadidae) and a mole cricket, Brisbane, Queensland. Memoirs af the Queensland Museunt 46(2): 499-51), Brisbane. ISSN QO79-8835. Diurnal singing behaviour is ducumented for 9 SE Queensland cicadas from Aruaie, Psalioda and Payropsalta, with particular reference to the presence/absence of dawn/dusk chorusing. This ovcurs more within relatively sedentary and also aggregating species. Detailed documentation of dusk chorusing during November 1996 -April 1997 jn janer city St Lucia, Brisbane, is presented lor 5 cicadas Cvsiosama Satindersil Westwood, Glauco- psaltria viridis Godinw & Froggatt. Tamasa tristigma (Germar), Abrieta enrvicoster (Germar). and Pselioda elaripennis Ashton, and the mole cricket (Gryllordlpa plivialis Mjdberg). The choruses closely follow, seasonally. the sunset and civil twilight curves, except for 7 risiemd which systematically changes its pattern duying the season Extensive interspecific synchrony ofchorusing oeeurs al the same loeation, although chorus starvfinish times. vary between species, Fuctors controlling chorusing behaviour, especially ligh| intensity and ambient temperature. and the jmportance of the distinctive interspecific song characteristics, are discussed. [7 Cicadas, mole cricket, chorusing behaviour. Brisbane. al, Rwart, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia; 13 October 2000 Many southeastem Queensland (SEQ) cicadas exhibit both extended daytime singing plus short and intense bursts of dawn and/or dusk chorusing. Certain species are erepuscular, with singing restricted to dusk, the best known local example being the Bladder Cicada, Cystosoma saundersit Westwood. Not all cicadas, however, exhibit dawn/dusk chorusing and observations over 30 years suggesi that the behaviour is best developed in: 1) relatively localised and sed- entary cicada species, and 2) aggregating species, many ol which are also localised, Comparable dawn/dusk chorusing behaviour is documented worldwide, including Malaysia (Govola & Riede, 1995); New Mexico (Crawtord & Dadone. 1979); Tennessee (Sanbom, 2000); Mexico (Moore, 1962); Borneo (Riede, 1996, 1997: Reide & Kroker. 1995); Costa Rica (Young 1976; 1982:102): Thailand (Gogola, 1995); Fiji (Dulfels, 1988): Southern Africa (pers. obs.). Myers (1929: 206-7, 221) quoted further exainples from South America, New Zealand, the Philippines and the Himalaya. The main feature of these choruses are their short, but predictable (day to day) timing and duration, and their intensity. Only severe late atiernoon storms seem to modify their timing. In SEQ, strongly mobile cicada species rarely produce defined dawn/dusk choruses, but instead sing more or less continuously throughout the day from early moming to near dusk. This report outlines observed cicada singing behaviour in a variety of species in SEQ and, in particular, more detailed documentation, over 6 months, of dusk chorusing of 5 cicada and | Orth- opteran species from suburban St Lucia, central Brisbane (27°29.63'°S, 153°00.04'E). All 6 species could be heard singing synchronously within the same localised area. This account starts by reporting more general observations of diurnal singing patterns in SEQ, DIURNAL CICADA SONG PATTERNS Two widespread but relatively sedentary mangrove-inhabiting cicadas, Arunfa lnterc/usa (Walker) and Psulreda plaga (Walker), provide examples of both strong day and dawn/dusk chorusing (Fig. 1). Both species are medium to larger sized cicadas (¢ body lengths [MBL] 27-33 and 27-39mm. respectively) with relatively sedentary behaviour patterns. Both exhibit strong diurnal singing. starting between (600-0700 hours (Eastern Standard Time), with progressive reduction in intensity during mid to Jate afternoon until cessation al approximately 1700 hours. Dusk chorusing occurs immediately following sunset, being less intense for P. plage, Dawn chorusing commences iminediately prior to sunrise. Although dawn chorusing 1s more poorly developed in P. plage, it commences its 500 MEMOIRS OF THE QUEENSLAND MUSEUM Arunta interclusa 0500 0700 0900 1100 D £ B (S) n 2 Sj w = - =9 nig = b=; Psaltoda plaga ® =e) Z 4 = ~ oO n = 3 n = = = = = = 1300 1500 1700 1900 Eastern Standard Time - Based on observations over 12 days between 3 to 18 February, 1981. Mangroves, Caloundra (SEQ) FIG, 1. Daytime singing patterns of two mangrove dwelling cicadas. The graphs are compiled from regular daily observations, at intervals of 15-20 minutes, of times at which songs of each species are heard within a given habitat. The sunrise/sunset times are shown for the period of observation. diurnal singing slightly earlier than A. interclusa. P. plaga normally forms relatively dense singing aggregations with short but frequent flights, this behaviour being less pronounced in A. interclusa. Both species sing from inner and outer mangrove branches which allows micro-habitat selection for variable shade conditions. Mangroves, in fact, are subjected to direct and also indirect solar radiation from reflection off surrounding water and wet mud surfaces. The afternoon decrease in 8p Psaltoda claripennis 12h 6 oa 0 s g 12 Psaltoda harrisii & = 6 i) wn a 0 = S 16 Tamasa tristigma 12 gl 4 0 0500 0700 0900 1100 1300 FIG. 2, Daytime singing patterns of three open woodland/suburban cicadas. Data compiled as in Fig. 1. 1500 song activity is presumably a behavioural response by the insects to avoid superheating (i.e. exceeding thermal tolerances; Sanborn, 1997). Psaltoda claripennis Ashton, Psaltoda harrisii (Leach) and 7amasa tristigma (Germar) (Fig. 2) illustrate further examples of song patterns. Both Psaltoda species are medium to larger sized cicadas (MBL 24-34 and 22-29mm, respectively) and widespread throughout SEQ. P. claripennis forms localised singing aggregations in suburban and open forest habitats while P. harrisii is restricted to open forest and wallum com- munities, again normally in localised aggregations. P. claripennis produces a strong dusk chorus, but no dawn chorus, whereas P. harrisii surprisingly exhibited neither during the observation period. Both are strong diurnal singers, with P. claripennis showing a marked reduction of song intensity during the midday period. As both species sing from exposed tree branches, reduction and later afternoon cessation of song seems to represent behavioural thermoregulation and allow feeding (Sanborn, 1997). T. tristigma is a very 1900 1700 Eastern Standard Time - Based on observations over 19 days between 25 January to 20 February, 1983, Open forest, S.W. Brisbane (SEQ) CHORUSING IN CICADAS AND A MOLE CRICKET 10 Numbers of males calling T Pauropsalta corticinus wun Sunrise 0700 0900 1100 Pauropsalta annulata Pauropsalta fuscata Pauropsalta circumdata WU Sunset 1300 1500 1700 1900 Eastern Standard Time - Based on observations over 19 days between 25 January to 20 February, 1983. Open forest, S.W. Brisbane (SEQ) FIG. 3. Daytime singing patterns of four open woodland Pauropsalta cicadas. Data compiled as in Fig. 1. common, medium sized (MBL 16-23mm), rather sedentary species of open forest and suburban areas. Singing occurs from exposed tree trunks. It exhibits a strong dawn chorus (coinciding with sunrise), followed at about 0700 by the beginning of the extended diurnal song period. This cicada is also notable for the fact that it regularly sings strongly during rain. As shown below, the late afternoon to dusk singing behaviour changes from early summer through to autumn, with discrete dusk chorusing only occurring early and late in a given summer season. Singing patterns of 4 Pauropsalta species (Fig. 3) show further variations. P. fiscata Ewart, P. corticinus Ewart and P. annulata Goding and Froggatt (MBL 12-17, 14-18 and 11-15mm, respectively) are small, cryptic and highly mobile species, common in open forests and wallum habitats (Ewart, 1989). P. annulata is a foliage dweller, while P fuscata and P.corticinus provide good examples of ‘sing and fly’ behaviour (e.g. Sanborn, 1997; Duffels, 1988: 74). Singing occurs from open tree trunks and branches, posts, etc., where their dominantly black coloration facilitates thermoregulation from solar radiation, with additional endogenous heat provided from frequent flight activity (e.g. Sanborn, 1997, 2000). Singing occurs throughout the day, extending to late afternoon, but they do not exhibit discrete dawn/dusk chorusing. This is believed to result from their smaller body size which precludes significant heat retention once solar radiation ceases to be effective (M. Coombs, pers. comm.). Pauropsalta circumdata (Walker) is a slightly larger (MBL 18-21mm) member of the genus, relatively sedentary, singing from open branches of medium to tall eucalypts, usually well exposed to solar radiation. No discrete dawn/dusk chorus is emitted. Diurnal song production extends through the morning, with a clear period of suspended activity during early-mid afternoon, followed by intense singing from late afternoon extending to dusk. This final phase incorporates the ‘dusk chorus’ time interval. During this late afternoon/dusk phase, the insects remain exposed to the sun allowing them to sing until solar radiation effectively ceases. EVENING CHORUS SYNCHRONY IN FIVE CICADA AND ONE MOLE CRICKET SPECIES Start and finish times for the dusk choruses of Glaucopsaltria viridis Goding & Froggatt (Bottle Cicada), C. saundersii (Bladder Cicada), Abricta curvicosta (Germar) (plus P. claripennis), T. tristigma, and the mole cricket Gryllotalpa pluvialis Mj6berg, are illustrated for a 6 month period (Figs 4-6). Each species sang within a localised suburban habitat, the songs each being easily heard from the single observation location. The choruses are compared with sunset and civil twilight times (data from Astronomical Applications Dept., U.S. Naval —— 7 Glaucopsaltria viridis 4 Days of Observation 1800 1900 120 ; a oo Feb ni Civil Twilight 6 look \ 4 a. J oo Cystosoma wz“ AN ra saundersii 4 \ A BS Jan | A + Te] 7 | es 4 60 I aN 7 ise / o / A => Dec / A oO Zz i A | A / A A | 20 > ey J A 2 Nov Ss; F nN re ES —— : 4 ‘ 2000 Time (Eastern Standard Time) - 1996/1997 FIG. 4. Dusk singing patterns, over 4-6 month intervals, of G. viridis and C. saundersii, from St Lucia, Brisbane City. The solid symbols indicate start, and hollow symbols finish, of dusk choruses. ‘St? against a symbol indicates storm. Sunset and civil twilight curves are plotted for comparison. Observatory, Washington). The plots highlight the close correspondence between the seasonally changing dusk chorus timing and light intensity. Sunset is formally defined as the moment the uppermost point of the sun appears to vanish below the horizon and civil twilight as the time between sunset and the moment the sun reaches a point lying 6° below the horizon (Beck, 1980; Nielsen, 1963). Although a similar result could no doubt be obtained with sky light intensity measurements (e.g. Crawford & Dadone, 1979), such measurements do not allow for the differing micro-habitat niches of the cicadas. C. saundersii and G viridis are both cryptic, green, crepuscular species which inhabit dense foliage, whose main song periods are at dusk (see also Doolan & MacNally, 1981; G viridis does produce brief singing and clicking during the day, especially when overcast). C. saundersii and G. viridis are large to medium sized cicadas (MBL 39-53 and MEMOIRS OF THE QUEENSLAND MUSEUM 180 Sa IT ew we April Te VY Finish Gryllotalpa 4 Vy wv . ope 1 Voog¥ pluvialis 3 Ss 140 eV 4 = March v J o v 4 5 * me” % Feb oof v ov 4 2 v id 0 |, y 4 5 Jan : Vv x, 2 60F VW 4 ® ae vay cy 4se wee wy Vv Y WwW e 20 Ml Civil 4 Nov E Vw Twilight | ) ae ere L f 1800 1900 2000 2100 140 ———tg +— 4 March cays = C ; 4 7q Biv es %, B c oY j g 100E SS q G—— | Abricta ne we é & Jan fF curvicosta £ yt \ a s E E ry o 60h # *} 5 4 ES . + wf o @ Dec £8. a JES. 4 2 © —. Pees 20F ¢ 4 BB ~ Civil Twilight 0 4 1 1 [800 900" "2000 Time (Eastern Standard Time) - 1996/1997 FIG. 5. Dusk singing patterns, over 5-6 month intervals, of A. curvicosta and the Common Mole Cricket, G. pluvialis. Filled and hollow symbols indicate start/finish of dusk chorusing. Sunset/civil twilight curves are shown for comparison. 26-35mm, respectively) with abnormally enlarged abdomens, and consequently relatively poor flight. A. curvicosta (MBL 25-3lmm) occurs on tree trunks, normally partly hidden by foliage, while P. claripennis and T. tristigma sing from more open tree trunks and branches. The mole cricket occurs in shallow soil burrows. Absolute light intensities will vary within these different microhabitats at any given time. Nevertheless, the mole cricket data are less regular than the corresponding cicada data, possibly due to variable diurnal and seasonal shadow effects over their fixed burrow positions. Rain flooding of their burrows presumably explains ‘non-singing’ during and following afternoon storms. The dusk chorusing of P. claripennis overlapped extensively with, and was largely masked by the singing of 4. curvicosta. Although the 2 species commenced dusk chorusing nearly simultaneously, P. claripennis stopped earlier. CHORUSING IN CICADAS AND A MOLE CRICKET e Dusk chorus g Marked increase in song intensity towards sunset start Days of Observation 1800 Time (Eastern Standard Time) - 1996/1997 FIG. 6. Dusk singing patterns, over 6 month interval, of 7. tristigma showing the seasonal changes of afternoon/dusk singing behaviour, noting that singing cessation is consistent throughout. The solid and dotted lines represent isolated days of anomalous singing behaviour relative to each behavioural segment. True dusk chorusing is restricted to early and late in season. Sunset/civil twilight curves are shown for comparison. See also Fig. 8D. 1700 T. tristigma shows a systematic change of singing patterns from early summer through to autumn (Fig. 6). Discrete dusk chorusing occurs in early and late summer season (November and April), with a very brief additional occurrence in early February. In mid-summer (mostly January), singing is continuous at relatively high intensities through until near civil twilight. The periods either side of mid-summer (December and February/March) have continuous singing through to civil twilight, but with marked increases in intensity in late afternoon/dusk (1.e. intermediate behaviour to discrete dusk chorusing). At all times during summer, however, singing consistently ends close to the civil twilight curve, irrespective of whether a discrete dusk chorus, or continuous late afternoon to dusk singing occur. Although clearly correlated with fading light, triggering mechanisms of dusk chorusing may be more complex. These include (Crawford & Dadone, 1979; Riede & Kroker, 1995): i) total light intensity; ii) rate of change of light intensity; iil) changing spectral composition of evening light, such as the suppression of middle wavelengths of visible light (Endler, 1992); and iv) barometric pressure changes prior to dusk. © Singing stops Separate and clearly defined dusk chorus. Continuous song to dusk: Marked increase of intensity towards sunset (grey circles). Continuous song to dusk: Intensity remains at a high and constant level. Continuous song to dusk: Marked increase of intensity towards sunset (grey circles). Separate and clearly defined dusk chorus. 1900 Some support for a more complex triggering mechanism, for start of dusk chorus, is provided by the lack of statistically significant differences between fine to cloudy/overcast days (Table 1), although the means do show systematic shifts. Only severe late afternoon storms caused the early onset of dusk chorusing (points labeled ‘st’ in Fig. 4). Another relevant aspect is the small seasonal variation of twilight period (i.e. between sunset and civil twilight), 0.40-0.47 hour for the total observation period. The close correlations between start and finish of dusk choruses and sunset/civil twilight curves, for each species, are illustrated further by Pearson correlation coefficients (r) derived from the linear plots between these variables (e.g. Fig. 7D). The coefficients between sunset and chorus start (finish) times or G . viridis, C. saundersii, T. tristigma, A. curvicosta and G. pluvialis are, respectively, 0.96 (0.99, n=121); 0.95 (0.88, n=84); (0.99, finish only, n=121); 0.85 (0.88, n=76); and 0.89 (0.86, n=117). Identical values are calculated using civil twilight instead of sunset times. The correlation coefficients are high, especially those for 7. ¢ristigma (chorus finish), G. viridis, and chorus onset of C. saundersii, pointing to absolute light intensity as TABLE 1. Dusk chorus starting times, relative to sunset, MEMOIRS OF THE QUEENSLAND MUSEUM under clear and cloudy conditions, chorus duration’s, reference CREP data, ambient temperatures (T,) and temperature gradients during dusk chorusing. Data presented as means and standard deviations (in parentheses). * Chorus finish only. = a — = —— ——————— = - — | > T. tristigma. G. viridis C. scundersii _A. curvicosta | _G. pluvialis Ag beecig stare = Panset) its 0,32 (0.07)* 0.030 (0.11) 0.26 (0,06) 0.24 (0.09) 0,29 (0.21) } <- - —_ - — -— - ~ - | (Chorus start — sunset) (hrs). 0.075 (0.071) 0.28 (0.052) 0.26 (0.086) 0.33 (0.16) | | Clear to <50% cloud cover 7 (n=77) (n= 46) (n= 39) (n= 74) _ | (Chorus start — sunset) (hrs), -0,031 (0.12) 0,23 (0.068) 0.22 (0.086) 0.23 (0.26) | | Overcast to >50% cloud cover 7 7 (n= 44) (n= 38) : (n= 37) I (n=43) | Chorus duration (hrs). All data = - | 0.33 (0.09) 0.61 (0.16) | | 0.29(0.14) | (0.87 (0.34) (CREP; chorus start. Alldata = 0.087 (0.25) _0.59.(0.15) 0.54 (0.20) _ 0.68 (0.49) | CREP: chorus finish, All data 0,74 (0.16) | 0.86 (0.13) _ 1.97 (0.35) 1.19 (0.27) | 2.69 (0.56) T,atchorus start(°C). Alldata | 24.2 (1.9) 23.8 (1.7) —23.6(18) | 24OCLS) | 23.5 (1.7) T,at chorus finish. (°C), Alldata | 23.5(1.7) | 23.5 (1.7) _ 23.3 (1.8) 23.8(1:5) | __23.0(1-8) __| Tesipeasiye sseeientaintng dusk 0.8 (0.8) 0.3 (0.4) 0.3 (0.6) 0.2 (0.3) 0.9 (0.8) q Seat - = _—t —— 4. - as } = _— Total T, range during dusk of . | chorus CC) 8 | 18-30 | 19-29 18-29 227 18-29 n (number of data) 121 121 84 76 “7 the dominant control on the timing of dusk chorusing. The coefficients for C. saundersii (finish), G pluvialis and to a less extent A. curvicosta, are numerically lower, reflecting the fact that chorusing ceases well after twilight, indicating additional controlling factor(s). Even for G. viridis, there is a discernible deviation of the chorus starting time, relative to sunset, early and late during the summer season (Fig. 7D). This also suggests secondary controlling factors, possibly seasonally changing spectral properties of the evening light. An alternative time parameter that has been advocated for such studies is the crepuscular time unit (CREP; Nielsen, 1963; Beck, 1980). Although noted in Table |, this parameter was not found to be as useful in this study as simpler comparative plots using standard sunset and twilight data. The possible influence of ambient temperature (T,) and temperature gradients on chorusing patterns are evaluated from temperature data, taken every 30 minutes during the whole period of observation, from records for Brisbane from the Australian Bureau of Meteorology. These data were linearly extrapolated to the seasonal starting and finishing times of chorusing for each species. T, ranged between 18-30° during chorusing, averaging between 23-24° (Tables | and 2) during the 6 month observation period. These temperature ranges suggest that T, is not a controlling factor in triggering or controlling the duration of dusk (or dawn) chorusing. This is confirmed, for all 5 species, by the lack of significant correlation’s between the start and finish of dusk choruses and T, (r values range between —0.22 and 0.42) or between temperature gradients during chorusing and the start/finish timing of the choruses (r values —0.24 to 0.35). It is recognised, however, that for singing to occur, body temperature needs to exceed some critical minimum value for each species (Sanborn, 1997, 2000), although this is in part controlled by T,. Josephson & Young (1979) indicated that C. saundersii 3 3 are ectothermic. It is unknown whether this also applies to the other cicadas considered here. Nevertheless, the patterns of singing behaviour and habitats of G viridis and T. tristigma males suggest possible ectothermy. Chorus duration (Fig. 7A; Table 2) for C. saundersii increases during the summer season, but sharply decreases at the end of its season, believed to reflect a combination of slightly shorter twilight period and especially the diminishing and aging population. G viridis, and A, curvicosta exhibit shorter chorus lengths both early and late in the season (Fig. 7B,C; Table 2), thought to also reflect smaller population and twilight length effects. G pluvialis shows no systematic seasonal change of chorus length. Reference to the seasonal variation of T,’s for dusk chorus finish (=start) times for G. viridis (Fig. 8A) suggest that seasonal T, variations could have a significant effect in determining seasonal changes in chorus duration. This is negated, however, by: 1) the lack of significant correlation’s, for all five species, between chorus CHORUSING IN CICADAS AND A MOLE CRICKET 120 -— at he 7 ' — ad A | 100 $A 4 c C 4 a ah J 2 f at a] © 80- bha | ic [ : aa A ] o t Cystosoma a“ n J 3 60. Saundersii AMA ac Qo" ws ert 1 ‘On ae . ‘ cs ri 1 g.°40F r 5 q o a ay a a Q é x 20¢ a or. + [ aw, 4A ot pe gt Aa A | 0.0 0.2 0.4 0.6 0.8 1.0 1.2 Dusk Chorus Duration (hours) 140g TO TT 1 E ¢@ Abricta curvicosta | - 20% o 1 2 c My r é ° : @ 100F +e pe 4 e E oe ° : B sot “yet E es 80F $ a . o : e) E % ars 4 eo 4 % 60b ot - L ¢ 4 o [ ¢ > E ® 4oF ° aes :° 4 ja) E * ° ¢¢ 20F ¢° 4 E Cj OL ! fl i i I | 1 J 0.0 0.1 0.2 0.3 0.4 0.5 0.6 Dusk Chorus Duration (hours) 505 180 [a * “Ee da ] 160- fm on Glaucopsaltria —~ Poe viridis 1 140; Sm a « =< & 120+ ial 4 $ ne LS 100} Hy = oF & . 8 80 a ha r 4 ‘O 60F i saul La | o r all @ 40- es = Hy + Qa L ag ZF aos Lad J 20+ "Zana B { [ a” fas = | 0 \ 5 ae a 0.0 0.1 0.2 03 04 05 06 07 Dusk Chorus Duration (hours) 1854. + Glaucopsaltria . a viridis | 1830 - + ra) L = L J - = ra 2 L © 1800+ | a B r 1 1730 > a L 1 i L L l 4 1700 1800 1900 Eastern Standard Time FIG. 7. A-C, dusk chorus duration in relation to progress of season (days of observation starting on | Nov, 1996, as in Figs 4 to 6). D, linear plot of sunset and twilight times in relation to the start/finish (filled/hollow symbols) of the dusk chorus of G. viridis over 6 month interval (as in Fig. 4). duration and either start/finish T,’s (r values of —0.20 to 0.28), or temperature gradients during choruses (r values 0.06 to 0.43); ii) although A. curvicosta and C. saundersii populations collapsed before those of the other 3 species, plots of days of observation (= season) versus T,’s of chorus finish (=start) for both of these species show no corresponding late season decrease in T, (e.g. Fig. 8B, illustrating the data for C. saundersii); 111) T, data for G pluvialis show late season decreases (Fig. 8C), yet no corresponding decrease in dusk chorus duration is found (although in this case, the subsurface micro-habitat is a complicating factor). Synchrony: A high degree of synchrony occurs between the 6 insect species during dusk chorusing (Figs 3-6). A general time progression occurs from 7. tristigma to G. viridis, A. curvicosta (+ P. claripennis) to G. pluvialis, (Fig. 9). Mean chorus duration (Table 1) ranges from 0.29 (A. curvicosta) to 0.87 hour (G pluvialis), whjch are consistent with dusk chorusing observations in Malayasian and Bornean rainforests (Gogola & Riede, 1995). Only the pre-sunset part of the 7. tristigma song and the later part of the G p/uvialis chorus do not overlap with competing insect songs. The late afternoon to early evening period is an acoustically ‘busy’ period of the day, from spring through to autumn, particularly as birds are also acoustically active during the same time period. This implies that the temporal structures and frequencies of the songs of the respective insects are sufficiently species specific to enable mate recognition to occur, as further outlined below. 506 18057 7 ai i E . iy . H La = A - < 1406 ey ges q 2 Ff fe, 4 L = = 100F ‘ » h 4 g eet | v 4 oO 7" H . a = 6 605 . ; a L a = il gE vo. ie _ 4 OQ Ep a . | 205 i + F . : ' glu 1 jo ny pe TT, PT 18 19 20 21 22 23 24 25 26 27 28 T (°C) Dusk Chorus Finish - G viridis 180 —-TJo I YT T TT T T 71 [ vy vy J L Vv 4 [- v ti] v Cc q E tal yey | c 140F wa * 7 rot E y = se ¥Y v 7 4 Oo E vy = 1005 shites ce 4 g E w ' bari j 5 E Ty 3’ im 4 5 60F pte Ss : o Mi w 1 > E ¥ 7 4 oO [ vy 4 a E a v } q 20- v vi¥ v J [ ¥ vy v Ow. 1 gh FL | pet a oo tl 18 19 20 21 22 23 24 25 26 27 28 T (°C) Dusk Chorus Finish - G pluvialis MEMOIRS OF THE QUEENSLAND MUSEUM 120 = Salt oT a ee ae a a 4 4 [ B | 100- 4 * . f 4 [ at FI i 44 1 80+ 7 a4 a [ a Sree ; t 6ob a j t ; A 44 1 4ob li 4 4 [ ek 4 | L. a a 4 20- a 4 ‘ A 7 PY Mek, 4 Ober. ee Bg ee ee hd 18 19 20 21 22 23 24 25 26 27 28 T (°C) Dusk Chorus Finish - C. saundersti 18067717 7 wy T T T T TF EI eet 4 Cte ee ee pt 1405 inn, 4 Pu 4 ole “er E oe J L ¥ my 3 ee 100 eames eeeeeeceroe Toe area " vr. , Il e of . 4 Se ONE ae We: Og OPE UP 60F « iat " . 4 r OT : a . ° 20 Fea TSear aera sas ae 5 are t eh eS bo = ee ] QOLr.te if § tg 18 19 20 21 22 23 24 25 26 27 28 T (°C) Dusk Chorus Finish - T tristigma FIG. 8. Seasonal (days of observation) variation of ambient temperatures (T,,) at the cessation of dusk chorusing for : A, G. viridis (Fig. 4); B, C. saundersii (Fig. 4); C, G. pluvialis (Fig. 5); and D, T. tristigma. In (D) the phases of changing singing behaviour are shown, based on Fig. 6, as follows: I, discrete dusk chorusing; II, continuous afternoon singing through to dusk, with marked intensity increase near sunset; II], continuous song through to dusk, with no change in song intensity. DISCUSSION Henwood & Fabrick (1979) highlighted, with particular reference to vertebrates, the optimal acoustic environment provided within the dawn chorus window, e.g. reduced temperature gradients, low wind and wind gradients, and low abiotic noise. This is expected to lead to more efficient broadcast coverage and therefore significant advantages to individuals selecting the early morning calling environment. Young (1981) pointed out that the same data indicate that dusk is also a time of low background noise and relatively stable wind and temperature gradients and again may lead to an adaptive advantage towards dusk chorusing. The underlying reasons for dusk/dawn chorus- ing in many cicadas are still unknown, but the following points are relevant to the phenomenon: i) Not all cicadas exhibit such behaviour, which seems to be most prevalent in more localised (static) species with continuous/monotonous calls. Dusk/dawn chorusing may not occur in smaller species (e.g. some Pauropsalta) due to more rapid heat loss once solar radiation ceases, thereby causing body temperatures (T;, ) to drop below the critical levels needed for song production (Sanborn, 1977). 11) Excepting crepuscular species, dawn/dusk chorusing CHORUSING IN CICADAS AND A MOLE CRICKET 507 TABLE 2. Monthly mean ambient temperatures (T,,), temperature gradients during dusk chorus (start to finish), chorus duration’s and chorus starts relative to sunset (SS). Data presented as means and standard deviations (in parentheses). iv — | November 1996 | December 1996 | January 1997 | February 1997 | | March 1997 | April 1997 | 2, trastieme 23.6 (3.0) 24.1 (14 24.2 (1.3) 25.8 (1.1 24.8 (0.8 22.8 (13 |Start T, °C), 23.6 (3, 24.1 (1.4) 2(1.3 25.8 (1.1) 24.8 (0.8) 22.8 (1,3) Finish T, (°C) 22.5 (2.6) 23.7 (1.2) 23.3 (1.3) 25.0 (1.2) 24.0 (0.8) 22.214) Temperature gradient 1.0 (1.4) 0.4 (0.5) 0.8 (0.6) 0.9 (0,6) 0.8 (0.5) 0.6 (0.6) Chorus duration (hr) 0.53 (0.17) 0.75 (0.14) 1.79 (0.51) 1.49 (0.57) 0.77 (0.37) 0.44 (0.10) (Chorus finish - SS) (hr) (),28 (0.08) 0.30 (0.08) 01,33 (0.05) 0.33 (0.06) 01.36 (0.02) (1.30 (0.06) ‘ 20 24 24 [ 20 7 16 | A. curvicosta Start T, (°C) 23.3 (2.3) 23.7 (1.3) 3.5 (1.2) 25.0 (1.2) 24.5 (0.9) - Finish T, (°C) 23,2 (2.4) 23.5 (1.2) 3.2 (1.2) 24.9 (1.2) 24.4 (0.9) - | ‘Temperature gradient | 0.5 (0.1) 0.2 (0.3) 0,3 (0.5) (0.2) (0.1) | Chorus duration (hr) 0.20 (0.09) 0.31 (0.08) 0.35 (0.17) 0.24(0.10) 0.04 (0.02) - (Chorus start — SS) (hr) 0.31 (0.05) 0.23 (0,09) 0.21 (0.07) 0.26 (0.10) | 0.20(0.09) 2 | 6 24 24 19 3 - G. viridis Start T,, (°C) 23.2 (2.7) 23.8 (1.3) 23.5 (1.2) 25.1 (1.2) 24.1 (1.3) 22.7 (1.3) Finish T, (@C) 22.6 (2.5) 23.6 (1.2) 23,3 (1.2) 24,9 (1.2) 23.7 (1.2) 22.6 (1.3) | Temperature gradient 0,6 (0.7) 0.2 (0.3) 0.2 (0.2) 0,2 (0.3) 0.4 (0,3) 0,2 (0.2) Chorus duration 0.33 (0.09) 0.39 (0,07) 0.38 (0.09) 0.34 (0.05) 0.30 (0.06) (1.20 (0.05) (Chorus start — SS) (hr) 0.04 (0.11) -0.03 (0.11) 0.00 (0.11) 0,01 (0.07) 0.09 (0.04) 0.16 (0.04) | we 21 25 24 20 18 13 C. saundersii Start T, (°C) 22.8 (2.6) 23.7 (1.3) 23.4 (1.2) 24.9(1.2) | - - | Finish T, OC) 22,3 (2.4) 23.3 (1.3) 23.1 (1.1) 24.8(1.3) | - - | Temperature gradient 0.5 (0.8) 0.4 (0.6) 0.3 (0.5) 0.1 (0.2) - - Chorus duration | 0.57 (0.14) 0.60 (0.15) 0.70 (0.13) 0.54 ((0,17) - - (Chorus start — SS) (hr) (1.26 (0.06) 0.27 (0.07) 0.26 (0.05) 0.26 (0.08) - - t 20 24 24 16 E : r, pluvialis Sift ace 23.1 (2.7) 23,7 (1.3) 23.4 (1.3) 24.8 (1.2) 23.9 (0.8) 22.1 (1,5) Finish T, (°C) 22.0 (2.4) 23.1 (1.4) 23.0 (1.2) 24.6 (1.2) 23.5 (0.8) 21.5 (1.5) Temperature gradient 1.1 (1.7) 0.6 (0.6) 0.4 (0.5) 0.2 (0.5) 0,4 (0.3) 0.7 (0.6) Chorus duration 1.07 (0.37) 1.00 (0,43) 0.81 (0.24) 0.71 (0.27) 0.81 (0.23) 0.75 (0.29) (Chorus start — SS) (hr) 0.13(0.29) 0.24 (0.24) 0.34 (0.16) ().32 (0.17) 0.37 (0.07) 0.38 (0.08) ha 20 23 23 20 15 16 species produce their main singing periods during the day. iii) During dusk chorusing, cicadas commonly become active, undertaking frequent localised flights to nearby branches or trees. This is especially notable in 29 of C. saundersii, corresponding to the period of crepuscular mating activity (Daws et al., 1997). iv) Although overlap of chorusing occurs, species differ in chorus start and finish times. This suggests differing response levels to critical external stimuli (e.g. light intensity) for each species, but these may be modified by differing micro-habitat niches (e.g. dense vs open foliage). v) C. saundersii (and apparently other crepuscular species) do not exhibit dawn chorusing. vi) The dusk song of the 7. rristigma is indistinguishable from the day song, while in G viridis the dusk song is an extended version of the very brief ‘whistle’ song sporadically emitted during the day. A. curvicosta has a continuous coarse dusk song, which is the extended equivalent of the longer phrase emitted as part of the day song. P. claripennis produces a continuous rattling song (which forms a major, but not continuous part of the day song). The dusk choruses of all species are therefore emitted as continuous songs, irrespective of the structure of the day songs. vii) T tristigma systematically changes its late afternoon to dusk calling behaviour through the season, which seems most plausibly related to seasonal temperature patterns. A very general correspondence does exist between the T, and singing patterns (Fig.8D), as for example, the occurrence of discrete dusk chorusing behaviour during the slightly cooler November and April periods. In contrast, the brief re-occurrence of this behaviour in early February seems to follow a sharp shift to higher T,. Overall, however, the T,’s occurring during and between the periods of changing behaviour extensively overlap, as reflected in the very similar monthly average T,’s for chorusing (Table 2). Available data are therefore inconclusive as to the role of T, in explaining the changing behaviour patterns, and imply influence of additional factors. Twilight = Gryllotalpa ‘ pluvialis Cystosoma saundersiti Abricta curvicosta Glaucopsaltria viridis << Sunset —~> Tamasa (ristigma ‘ Song Duration Measured as Hours after Sunset: Mean plus 10 FIG. 9. Summary of dusk chorus duration, relative to sunset, of the five cicada and the mole cricket species. Twilight represents time between the sunset and civil twilight lines, the width of the civil twilight line marking the restricted seasonal variation of twilight through the observation period. Staicer et al. (1996) listed 12 hypotheses in 3 categories to explain dawn/dusk chorusing in birds: 1) intrinsic to internal state, e.g. hormonal levels; 2) social function; 3) dawn preference resulting from daily timing of environmental selective pressures. Within these categories, the following aspects, in combination, are considered potentially relevant to cicadas: 1) Self-stimulation. Dawn chorusing perhaps represents a ‘warming-up’ strategy necessary for relatively static species (cf. Josephson & Young, 1979). A possible cue for dawn/dusk song could be the changing xylem-flow pressure within vegetation which stops at dusk and resumes at dawn, presumably responding to transpiration rate (e.g. Dolling, 1991:8). As noted, however, not all cicadas sing at dawn/dusk. 2) Mate attraction, certainly critical for dusk singing crepuscular insects. In other species, however, mating occurs throughout the day, with no observational evidence for unusually high mating activity at dawn/dusk. 3) Mate stimulation, The intensity of dawn/dusk chorusing, plus the optimal acoustic conditions, may facilitate female location of males at a time of lowered predation pressure. 4) Social dynamics. Chorusing may represent an effective mechanism, by signaling, of adjusting spatial distributions between calling males, after MEMOIRS OF THE QUEENSLAND MUSEUM dispersion during the day (e.g. by predation). Doolan & MacNally (1981) have shown that although aggregation is important in C. saundersii for increasing mating success, individuals space themselves at ~1-1.5m apart, with females selecting males only on the basis of their acoustic display. Doolan (1981) further suggests that the spatial distribution results from the interplay between selective pressures to aggregate (ensuring greater numbers of females), and selective pressure to space (enhancing individual success in mating). The short and intense dawn/dusk chorusing provides a mechanism for this to happen and explains the increased, but localised activity of individuals. It also may explain why chorusing is important for relatively localised and also for aggregating species. 5) Lowered predation rates from birds, reptiles, arachnids, and predatory insects. Doolan & MacNally (1981) note the more intense bird predation early in the dusk chorus of C. saundersii. 6) Acoustic transmission enhanced at dawn/dusk, as previously noted. 7) Energy reserves conserved/accumulated during night, stimulating burst of singing at dawn. This does not, however, explain the dusk chorus. UNIQUENESS OF THE CICADA AND MOLE CRICKET SONGS. As noted, the extensive temporal overlap of songs during the acoustically ‘busy’ dusk window requires that the songs are species specific. A detailed presentation of the song characteristics is beyond the scope of this paper, but the essential temporal structures of the cicada songs are presented (as oscillograms/ waveform plots) in Young (1972a,b; 1980), Simmons & Young (1978), Young & Josephson (1983) and Ewart (1995). The G plivialis call is documented by Otte & Alexander (1983). These data clearly show the distinctive temporal song structures of each species, seen in their pulse and phrase structures, and pulse repetition rates. The differences are also reflected in the dominant frequencies of the songs. For G viridis, C. saundersti, T. tristigma, A. curvicosta, P. claripennis and G. pluvialis, these frequencies (author data) are, respectively (kHz): 1.8-1.9; 0.8-0.9; 6.1-8.4; 9.5-9.6; 5.9-6.8; and 2.1. Further differences are seen in the detailed structures of the frequency bands, i.e. whether broad or narrow, indicating the ‘purity’ of the emitted tones. These are represented as bandwidths, based on the sound energy emitted between the lower (25%) and upper (75%) quartiles determined from song spectra. Respective values are: 0.14: 0.28; 1.0; 2.3; 2.1; and 0.27kHz. The G CHORUSING IN CICADAS AND A MOLE CRICKET viridis song closely approaches a modulated pure tone (see also Young & Josephson, 1983), while the songs of G viridis and G. pluvialis exhibit well defined harmonics (to 5f), giving increased transmission flexibility to the songs. Notwithstanding the extensive synchrony during dusk chorusing between species, the interspecific songs each have their uniquely defined acoustic properties. ACKNOWLEDGEMENTS I thank Geoff Monteith, Queensland Museum, for making numerous improvements to the manuscript and Dr M. Coombs for emphasising the potential importance of temperature in interpreting the data. LITERATURE CITED BECK, S.D. 1980. Insect photoperiodism. 2nd edition. (Academic Press: New York, London). CRAWFORD, C.S, & DADONE, M.M. 1979, Onset of evening chorus in Tihbicen marginalis (Homoptera: Cicadidae), Environmental Entomology 8(6): 1157-1160. DAWS, A.G, HENNIG, R.M. & YOUNG, D, 1997. Phonotaxis in the cicadas Cystosoma saundersii and Cyclochila australasiae, Bioacoustics 7(3): 173-188. DOLLING, W.R. 1991. The Hemiptera. (Oxford University Press: Oxford). DOOLAN, J.M. 1981, Male spacing and the influence of female courtship behaviour in the bladder cicada, Cystosoma saundersii Westwood. Behavioral Ecology and Sociobiology 9: 269-276. DOOLAN, J.M. & MacNALLY, R.C. 1981. Spatial dynamics and breeding ecology in the cicada Cystosoma saundersii: The interaction between distributions of resources and intraspecific behaviour. Journal of Animal Ecology 50: 925-940. DUFFELS, J.P. 1988. The cicadas of the Fiji, Samoa and Tonga Islands, their taxonomy and biogeography (Homoptera, Cicadoidea). Entomonograph 10, (E.J, Brill/Scandinavian Science Press). ENDLER, J.A. 1992. Signals, signal conditions, and the direction of evolution. The American Naturalist 139 (Supplement): $125-S153. EWART, A. 1989, Revisionary notes on the genus Pauropsalta Goding and Froggatt (Homoptera: Cicadidae) with special reference to Queensland. Memoirs of the Queensland Museum 27(2): 289-375, 1995. Cicadas. Pp. 79-88. In Ryan, M. (ed.) Wildlife of Greater Brisbane. (Queensland Museum: Brisbane), GOGALA, M. 1995. Songs of four cicada species from Thailand. Bioacoustics 6; 101-116. 509 GOGALA, M. & RIEDE, K. 1995. Time sharing of song activity by cicadas in Temengor Forest Reserve, Hulu Perak, and in Sabah, Malaysia. Malayan Nature Journal 48: 297-305. HENWOOD, K. & FABRICK, A. 1979. A quantitative analysis of the dawn chorus: Temporal selection for communicatory optimization, The American Naturalist 114(2); 260-274. JOSEPHSON, R.K. & YOUNG, D. 1979. Body temperature and singing in the Bladder Cicada, Cystosoma saundersii. Journal of Experimental Biology 80: 69-81. MOORE, T.E. 1962. Accoustical behavior of the cicada Fidicina pronoe (Walker) (Homoptera: Cicadidae), Ohio Journal of Science 62: 113-119. MYERS, J.G. 1929. Insect singers. A natural history of the cicadas. (George Routledge & Sons: London). NIELSEN, E.T. 1963. Illumination at twilight. Oikos 14(1): 9-21. OTTE, D. & ALEXANDER, R,.D. 1983. The Aus- tralian crickets (Orthoptera: Gryllidae). Academy of Natural Sciences of Philadelphia Monograph 22 RIEDE, K 1996. Diversity of sound-producing insects in a Bornean lowland rain forest. Pp. 77-84, In Edwards, D.S. et al. (editors) Tropical Rainforest Research — Current Issues. (Kluwer Academic Publishers: Netherlands). 1997. Bioacoustic monitoring of insect com- munities in a Bornean rainforest canopy. Pp. 442-452. In Stork, N.E., Adis, J. & Didham, R.K. (eds) Canopy Arthropods. (Chapman & Hall: London), RIEDE, K. & KROKER, A. 1995. Bioacoustics and niche differentiation in two cicada species from Bornean lowland forest. Zoologischer Anzeiger 234: 43-51, SANBORN, A.F. 1997. Thermal biology of cicadas (Homoptera: Cicadoidea). Trends in Entomology 1: 89-104. 2000. Comparative thermoregulation of sympatric endothermic and ectothermic cicadas (Homoptera: Cicadididae: Tibicen winnemanna and Tibicen chloromerus). Journal of Comparative Physiology A 186: 551-556. SIMMONS, P & YOUNG, D. 1978. The tymbal mechanism and song patterns of the bladder cicada, Cystosoma saundersii. Journal of Experimental Biology 76: 27-45. STAICER, C.A., SPECTOR, D.A. & HORN, A.G. 1996. The dawn chorus and other diel patterns in acoustic signaling. Pp. 426-453. In Kroodsma, D.E. & Miller, E.H. (eds) Ecology and evolution of acoustic communication in birds. (Cornell University Press: Ithaca, London). YOUNG, A.M. 1976. Notes on the faunistic complexity of cicadas (Homoptera: Cicadidae) in northern Costa Rica. Revista de Biologia Tropical 24: 267-279, 1981. Temporal selection for communicatory optimization: The dawn-dusk chorus as an 510 MEMOIRS OF THE QUEENSLAND MUSEUM adaptation in tropical cicadas. The American 1980. The calling song of the Bladder Cicada Naturalist 117: 826-829. : an ; 1982. Population Biology of Tropical Insects. CHSTOSOMIE: SiNGEESE. B. POMP tet ANALYSTS: im (Plenum: po York, ee ‘ ; Journal of Experimental Biology 88: 407-411. OUNG, D. 1972a. Neuromuscular mechanism o . sound production in Australian cicadas. Journal of is D. beheeour ders oe ee es tee Comparative Physiology 79: 343-302 ue ES ie pe reer 1972b. Analysis of songs of some Australian Physiology 152: 183-195. cicadas (Homoptera: Cicadidae). Journal of the Australian Entomological Society 11: 237-243. PLATYHELMINTHES FROM SALT MARSHES OF COOMERA RIVER, SOUTHEASTERN QUEENSLAND, AUSTRALIA ANNO FAUBEL AND BRONWYN CAMERON Faubel, A. & Cameron, B. 2001 06 30: Turbellaria from salt marshes of Coomera River, southeastern Queensland, Australia. Memoirs of the Queensland Museum 46(2): 511-519. Brisbane. ISSN 0079-8835. Three new species, Childianea coomerensis gen. et sp. nov. (Acoela), Macrostomum greenwoodi sp. nov. and Macrostomum coomerensis sp. nov. (Macrostomida) are described from the brackish water estuarine area of the Coomera river, Gold Coast, Queensland, Australia. 1 Taxonomy, morphology, flatworms, Acoela, Macrostomida. Dr A. Faubel, Institut fiir Hydrobiologie und Fischereiwissenschajt, Universitét Hamburg, Zeiseweg 9, D-22765 Hamburg, Germany (e-mail: faubel(@uni-hamburg.de); B. Cameron, The University of Queensland, St Lucia 4072, Australia; 14 March 2000. The Australian fauna of Acoela and Macrostomida of limnetic, brackish and marine habitats is not very well known. Over a long time the only study known on Acoela was carried out by Haswell (1905) on Heterochaerus australis. More recently, however, more intensive studies have been started by Trench & Winsor 1987 on Amphiscolops sp. and Haplodiscus sp. (in part Waminoa litus Winsor, 1990), Winsor 1988 on Wulguru cuspidata, Winsor 1990 on Convolutriloba hastifera, Waminoa litus, W. sp. 1, and M sp. 2. A first census of macrostomid species of Australia by Faubel et al. (1994) listed 6 species. Of these Macrostomum tuba v. Graff, 1882 and Promacrostomum palum Sluys, 1986 are known from freshwater environments. Dunwichia arenosa Faubel, Blome & Cannon, 1994, Bradburia australiensis Faubel, Blome & Cannon, 1994, Macrostomum australiense Faubel, Blome & Cannon, 1994, and Macro- stomum sp. Faubel, Blome & Cannon, 1994, however, are all known from eulittoral habitats of sandy beaches of Stradbroke Island, South Queensland. In the present paper one acoelous species and two macrostomid species from brackish water habitats are described. METHODS For studies on meiobenthos, sediment was qualitatively collected from the Coomera salt-marsh pools. The freshly collected sediment was transferred into a glass tank in a constant temperature tom approximating ambient conditions (Salinity ca 30 ppt, 27°C). Habitat water was added to the sediment tank, to a depth of about 15cm. A series of 60 watt light bulbs were suspended above the tank to attract organisms into the water column. Every hour, the tank was siphoned through a 63m sieve to collect emerged fauna. The defaunated water was then returned to the tank and the extraction process repeated until the majority of the organisms had been collected. Sexually mature specimens of Turbellaria were studied alive and in squash preparation, i.e. flattened under the increasing pressure of the coverslip as the preparation dried. Measurements of living organisms (m.l.s.) were made from squashed ones. These measurements are given in parentheses in the running text. All other measurements were made from sections of the holotype. For histological observation specimens were relaxed in 7% MgCl, and fixed in Bouin’s fixative. Specimens were embedded in Paraplast plus (Reichert & Junk) and cut sagittally at 3.0,1m and stained with haematoxilin-eosin according to Mayer. Types are deposited in the Queensland Museum, Brisbane. SITE DESCRIPTION Coomera salt-marsh (27°54’S, 153°17°E) was chosen as a site for turbellarian collection. For the past 4 years this site had been excluded from the broad-scale mosquito control measures that typically occur in southeast Queensland. It was therefore considered to be a relatively pristine site. Coomera salt-marsh is only inundated by tides of 2.4m or more above datum. Salinity ranges between 25-38 ppt. The salt-marsh pools are 2-7m in area and are bordered by dense clumps of Sporobolus virginicus (Kunth.) the dominant marsh-grass, and Sarcocornia quinqueflora Bunge (ex. Ung. Stern) a sprawling herbaceous halophyte. The sediment consists of much highly organic mud with negligable amounts of sand. SYSTEMATICS Order ACOELA Uljanin, 1870 Family CHILDIIDAE Darjes, 1968 Childianea gen. nov. DIAGNOSIS. Childiidae with frontal organ. Body wall musculature with outer circular and inner longitudinal muscle fibers. Separate genital openings. Seminal bursa with vagina and cellular bursa mouth directed anteriad. Paired ovaries and testes. Male copulatory apparatus without seminal vesicle; proximal ejaculatory duct with sclerotised layer working as penial stylet; distal ejaculatory duct with prostatic glands. False seminal vesicles present. TYPE SPECIES. Childianea coomerensis sp. nov. ETYMOLOGY. Derived from the generic name Childia. KEY TO THE GENERA OF THE CHILDIIDAE 1. withsingle male copulatory organ, . 2... 0.00. 3 with several male copulatory organs... ....... 2 Ne with 2 male copulatory organs . . Childiav. Graff, 1910 with 4 male copulatory organs deena ¢ My Ae er Tetraposthia An Der Lan, 1936 3(1). . . with single male copulatory organ; without seminal Dursaw yp ORS Re met Ta RY et withseminal bursa. 6... 6. ee et ee ee es 6 4. without male atrium orvery short ........... 5 withlong maleatrium ... . .. Atriofronta Dérjes, 1968 5. copulatory organ with sclerotised spines... . 2. . td ce aphes Noetiat ahd Actinoposthia An Der Lan, 1936 copulatory organ with rosette like muscular elements encasing the seminal vesicle eS ee en Paraproporus Westblad, 1945 6(3). with seminal bursa; body muscle wall with outer circular and inner longitudinal muscle fibers... 2... 0... 7 body muscle wall with inverse muscle layers porbha wis a ees Paraphanostoma Steinbick, 1931 without bursanozzle . Pseudactinoposthia Dirjes, 1968 8. bursanozzle directed anteriadorcaudad. . 2... .. 9 bursa nozzle invaginated in the seminal bursa ale et yee pe ee Pelophila Dérjes, 1968 bursa nozzle sclerotised . . Philactinoposthia Dérjes, 1968 10, male complex with prostatic glands, without seminal WOSICION. ae 6 5 eases 54 a Childianea gen. nov. male complex with seminal vesicle, without prostatic glands. ........4¢ Archactinoposthia Dorjes, 1968 MEMOIRS OF THE QUEENSLAND MUSEUM Childianea coomerensis sp.nov. (Figs 1A-B, 2 ETYMOLOGY. From the Coomera River estuary on the Gold Coast of Queensland. MATERIAL, HOLOTYPE. Coomera salt-marsh (27° 4’S, 153°17°E), Gold Coast, southeast Queensland, QM G217362, 2 specimens, May 1998 leg. B. Cameron. DESCRIPTION. Length of body of living sexually mature specimens up to 0.9mm, when extended and gliding; maximum width up to 0.18mm in second half of body, area of growing oocytes. Body spindle-shaped with rounded frontal and rear body end, dorso-ventrally flattened. Posterior body end with a small vacuole being horseshoe-like (Fig. 1A). In transmitted light, colour of body yellowish but digestive parenchyme and lumen bright yellow through ingested diatoms. Glandular ducts of frontal organ open to exterior through several pores at the frontal end, the glands of which lie posterior to the statocyst. Secretion of the glands coarsely granulated. Statocyst embedded within the medio-frontal brain, about 55um (m.1.s.) distant from anterior body end. Epidermal cells 2.0 - 2.5m high, with intraepithelial nuclei; rhabdoids lacking. Epidermis completely covered with cilia 3.5 - 4.0um in length. Body wall musculature with faint outer circular and inner longitudinal fibers. Dorsal and ventral subepidermal hyaline glands open to the exterior throughout body surface, more abundant lateral. Digestive lumen bordered by nucleus-rich parenchymal tissue which starts immediately at level of frontal glands and fills the whole mid-body covering the ovaries dorsal. Gut lumen filled with diatoms and detritus. Mouth ventral behind mid-body, about 400m (m.l.s.) distant from frontal body end. Reproductive system (Fig. 2). Lateral testes situated about 1 80um (m.l.s.) from anterior body end. Spermatids penetrate the parenchyma on both sides lateral of the ovaries. Anterior to the transverse level of the male complex the spermatids accumulate in two false seminal vesicles which enter the common proximal ejaculatory duct in common. True seminal vesicle lacking. Ejaculatory duct is closed by a sphincter at junction of the vasa deferentia. The ventral male pore (Fig. 1B,2), 32um distant from rear body end, leads to the bulbous pyriform copulatory organ. Muscle bulb a highly muscular complex 43m in length with central ejaculatory duct (Fig. 2). The muscle fibers run more or less PLATYHELMINTHES FROM THE COOMERA RIVER 513 FIG. |. Childianea coomerensis, gen.et sp.noy. A, dorsal view in squash preparation. B, diagrammatic sagittal reconstruction after serial sections. Scale: 200um. (b = brain, bs = seminal bursa, bm = mouth of bursa, ca = copulatory apparatus, d!= digestive lumen, f= frontal organ, fs = false seminal vesicle, m = mouth, o = ovary, oc = oacyte, s = statocyst, t = testis, v= vacuole, d = male gonopore, ° = female gonopore). parallel, inclining toward the ejaculatory duct, and obviously function as retractors during copulation. Ejaculatory duct tripartite; proximal part of duct (Fig. 2) about 10um long, highly sclerotised forming a seal controlling sperm ejaculation; central one, 31m long, lined by a sclerotised epithelial layer looking stylet-like; distal part is 12um long, not ciliated, and of prostatic function, Cell bodies of epithelial layer very elongated, enclose copulatory complex (Fig. 2). LOS\WAUAC NA 20 MAI ee rR n bs fs MEMOIRS OF THE QUEENSLAND MUSEUM re ej FIG. 2. Childianea coomerensis, gen. et sp. nov.; sagittal reconstruction of the male and female complex. Scale: 50um. (bs = seminal bursa, bm = mouth of bursa, c = circular muscles, dl = digestive lumen, ej = ejaculatory duct, fs = false seminal vesicle, | = longitudinal muscles, n = nucleus, oc = oocyte, po = prostatoid glands, re = retractor muscles, se = seal, sg = subepidermal gland, v= vacuole, d = male gonopore, ° = female gonopore). Female system with paired ovaries and a seminal bursa. Oogonia of each ovary lie ventro-lateral, 190-200um behind statocyst (Fig. 1B). A line of growing oocytes runs latero- caudad. Hindmost oocytes are about 170um (m.l.s.) distant from posterior body end. Seminal bursa with a distal ciliated atrium (7.5m long), a central vesicle filled with sperm (20m in diameter), and a cellular mouth piece 13m long. The atrium rises dorsad, 65u:m anterior to male pore, widening proximad to form a vesicle filled with sperm. The cellular bursa mouth piece is directed anteriad, reaching central the hindmost part of the oocytes. Aperture between atrium and vesicle of bursa is closed by a sphincter. DIAGNOSIS. With characters of the genus. DISCUSSION. The family Childiidae was established and discussed in detail by Dérjes 1968. Up to now the family contains 10 genera. D6rjes included in this family all these genera of Acoela which have species with a sclerotised (cuticular according to Dérjes 1968), muscular, or cellular penis plug which is never invaginated in the seminal vesicle. The new species Childianea coomerensis is characterized by a sclerotised ejaculatory duct obviously working as a pointed stylet. On the basis of this character the species belongs to the Childiidae. Coomera coomerensis, however, differs from all species of the known genera of the family Childiidae in the presence of prostatic glands entering the distal ejaculatory duct. In combination with the characters of presence of a seminal bursa with a cellular spermiducal duct directed frontad and absence of a true seminal vesicle, establishment of monotypic Childianea and its species C. coomerensis sp. nov. is justified within the family Childiidae. Order MACROSTOMIDA, Meixner, 1924 Family MACROSTOMIDAE Van Beneden, 1870 Macrostomum greenwoodi sp. nov. (Fig. 3) ETYMOLOGY. Named in honour of Prof. J. Greenwood, University of Queensland. MATERIAL. HOLOTYPE. Coomera salt-marsh (27°54’S, 153°17’E), Gold Coast, southeast Queensland, QM G217363, 2 specimens, April 1998 leg. B. Cameron. DESCRIPTION. Length of body of living sexually mature specimens up to 1.3mm, when PLATYHELMINTHES FROM THE COOMERA RIVER 515 FIG, 3. Macrostomum greenwoodi, sp. nov. A, dorsal view in squash preparation. B, male stylet. C, sagittal reconstruction of the female organ. Scales: A, 200m; B, 20pm; C, 50m, (ad= adhesive glands, b = brain, bs = seminal bursa, c= circular muscles, ce= cement glands, d= diatoms, ds = spermiducal duct, e= eyes, fa= female atrium, i= intestine, | = longitudinal muscles, m = mouth, n= nucleus, 0 = ovary, pg = pharyngeal glands, r= rhabdite, ra = rhammites, t = testis, vg = vesicula granulorum, vs = seminal vesicle, ¢ = male gonopore. 9 = female gonopore). 516 extended and gliding; maximum width up to 0.18mm in mid-body. Outline of body with characteristic macrostomid like anterior end; posterior end rounded, provided with strong adhesive glands (Fig. 3A). Around margin weak sensory hairs of varying length and stiffness present. In incident light body brownish with darker contrasting oocytes in the posterior part of body; gut intensively coloured yellow based on the uptake and consumption of bright yellow diatoms. Eyes present, 175ym (m.Ls.) far from anterior margin of the body. Crescentic brain 169um (m.l.s.) distant from anterior end. Rhammite glands open ventro-frontal through the anterior margin of body, their cells extending posteriad up to the level of the pharynx simplex. In mid-body cells of epidermis 4.6m high covered with 4.0 to 4.6m (ventral) and 3.4 to 3.9um (dorsal) long cilia. Intraepithelial nuclei present; rhabdites about 9.011m long, the cell bodies of which subepidermal, up to 11.3um long. Rhabdites distributed in longitudinal rows over the dorsal and ventral body surface. Body muscle wall with outer circular and inner longitudinal muscle fibers. Basal membrane not observed, Digestive system with pharynx simplex pierced by extrapharyngeal glands and large intestine but less numerous ciliated than pharynx. The intestine fills the median parts over the ovaries, oocytes, and seminal bursa and extends caudad up to the seminal vesicle. Mouth behind brain, 210,1m (m.l.s.) distant from anterior end. The species feeds on diatoms and detritus particles. Ingested sand grains covered with detritus are abundant. Excretory system not observed, probably absent. Male and female gonopores separate, 185,1m (m.l.s.) distant from each other, Reproductive system (Fig. 3). The male system is typically macrostomid like. It consists of bilateral testes, vasa deferentia running caudad to either side of the intestine and ovaries, a common vas deferens, a seminal vesicle, a prostatic vesicle, and a male stylet which projects into the short male antrum. The stylet depicted in Fig. 3A, is 98.3,1m long; the proximal opening is 12.0j1m in diameter; the distal opening is subterminal.The male pore is 105um (m.l.s.) distant from rear body end. The antrum rising dorsad is 10.8um long. The female system consists of bilateral ovaries and a seminal bursa. The ovaries forming lateral fields of oogonia, lie behind the germinative zone of the testes in mid-body (Fig. 3A), generating caudad large oocytes. The seminal bursa consists MEMOIRS OF THE QUEENSLAND MUSEUM of a ciliated atrium (11.4um long), a bursal vesicle (37.9um long), and a spermatic duct (12.8um long). The bursal vesicle is totally lined with elongated cells. The proper epithelium of the bursa looks syncytially in which the surrounding cells (up to 35,1m) excrete obviously coarsely granulated secretion. Cement glands surrounding the female pore, discharge their secretion into the female atrium. DIAGNOSIS. The species is diagnosed on the outline of the male stylet (Fig. 3B). Macrostomum coomerensis sp. nov. (Fig. 4) ETYMOLOGY. From the Coomera river of the Gold Coast of Queensland. MATERIAL. HOLOTYPE. Coomera salt-marsh (27°54’S, 153°17°E), Gold Coast, southeast Queensland, QM G217364, 2 PARATYPES. Same data, QM G217365-366, numerous specimens, April 1998 leg. B. Cameron. DESCRIPTION. Length of body of living sexually mature specimens up to 1.4mm, when extended and gliding; maximum width up to ().25mm in anterior part of second body-half, area of oocytes (Fig. 4A). Anterior and posterior body end rounded. Marginal hairs or spines absent. Frontal glands present, not reaching anterior level of brain. Eyes present immediately behind crescentic brain, 125m (m.l|.s.) distant from anterior margin of body. In incident light colour brownish yellow with contrasting darker lateral ovaries. In transmitted light body greyish translucent with yellowish digestive system. Mouth opening immediately caudal of eyes. From mouth the pharynx simplex rises dorso- caudad and receives openings of digestive glands. Intestine with less numerous cilia. The gut fills the median parts of the body extending caudad over ovaries and oocytes, ending up at the anterior level of the last oocytes. Rhabdites evenly distributed in serial lines over the body. Ventral rhabdites 7.5um and dorsal ones 9.4um long. Rear dorsal body end with dense aggregates of large rhabdites (10.7um long). Ventral epidermis is 5.7um thick and dorsal one is 2.8m thick, entirely covered with 5.7um long cilia; epithelial nuclei subepidermal. Basal membrane not observed. Body muscle wall with outer circular and inner longitudinal muscle fibers. Excretory system not observed, probably absent. Gonopores separate, 65um distant from each other. Bilateral testes anterior to ovaries; last in mid-body. PLATYHELMINTHES FROM THE COOMERA RIVER 317 A c r Eu! 7 ce SS le ee \ .] eet bs Hg Wore FIG. 4. Macrostomum coomerensis, sp. nov.; A, dorsal view in squash preparation. B, male stylet. C, sagittal reconstruction of the female organ. Scales: A, 200um; B,C, 50pm. (b = brain, bs = seminal bursa, c = circular muscles, ce =cement glands, ds = spermiducal duct, e = eyes, f= frontal organ, fa= female atrium, i= intestine, | = longitudinal muscles, m = mouth, n= nucleus, o = ovary, pg = pharyngeal glands, r= rhabdite, t= testis, va = vagina, vg = vesicula granulorum, vs = seminal vesicle, ¢ = male gonopore, ° = female gonopore). Reproductive system (Fig. 4). The male system is testes, vasa deferentia running caudad to either typically macrostomid like. It consists ofbilateral side of the intestine and ovaries, a common vas deferens, a seminal vesicle, a prostatic vesicle, and a male stylet which projects into the male antrum. The sac-like testes are located 100 to 110m (m.Ls.) behind the pharynx. Latero- caudal of the female bursa the vasa deferentia unite to a common vas deferens on the left side before entering the seminal vesicle. The distal part of the seminal vesicle, the prostatic vesicle, and the proximal part of the penial sheath in which the stylet rests, are covered with a complicated system of strong muscle fibers. These fibers are responsible for protrusion and retraction of the stylet and ejaculation of sperm into the female system. The distal part of the prostatic vesicle is encased in the proximal portion of the stylet bemg slightly funnel-like extended, The stylet is 125,1m (m.l.s.) long; the distal two thirds are spiralled as outlined in Fig. 4B. The distal tip of the stylet is broadend, the opening of which lies subterminally. The male pore, 42.7u1m distant from the rear body end leads antero-dorsad to a rather long male antrum (15.2)1m). The female system consists of bilateral ovaries and a seminal bursa. The oogonia generate laterally immediately behind the testes and are growing up caudad forming oocytes on each side of the body. The female genital pore is 280,1m (m.l.s.) distant from the caudal body end. The female atrium (43.5.4m long) entered by cement glands, is outlined with cilia and rises dorsad. The transition to the vagina is closed by a sphincter. The vagina, up to 31.5um long, ciliated, extends frontad to form a bursal vesicle filled with sperm. The cilia of the vagina up to 13.0um long. Anteriad of the bursa a spermiducal duct (24m long) is present. The spermiducal duct reaches up mediad to the level of the last oocytes. The epithelia of the vagina, the bursal vesicle, and the spermiducal duct are cellular with intraepithelial nuclei. DIAGNOSIS. The species is diagnosed on the outline of the male stylet (Fig. 4). DISCUSSION, Macrostomidae currently contains 11 genera. The dominant taxonomic characters concern the male copulatory apparatus: a distally armed ejaculatory duct (stylet), or an unarmed cirrus. Genera having a penial stylet are Macro- stomum Schmidt, 1848, Omalostomum Van Beneden, 1870, Promacrostomum An Der Lan, 1939, Archimacrostomum Ferguson, 1954, Bradynectes Rieger, 1971, and Bradburia Faubel, Blome & Cannon, 1994. Of these genera, Macrostomum has the most species (127). MEMOIRS OF THE QUEENSLAND MUSEUM Beklemishev (1951) has assigned the species of Macrostomum to 3 taxonomic groups as follows: Macrostomum hystricinum group (stylets being funnel-like proximally, with tapered hook-like shape distally), Macrostomum orthostylum group (stylets being straight or curved, distad evenly tapering), and Macrostomum tuba group (stylets having a distal characteristic Macrostomum tuba-like enlargement, with the opening of the ejaculatory duct in the centre of the tip, or subterminally). Although these groupings are useful in a taxonomic sense, they should not be regarded as phyletic groupings because there are different lines of the development of the female genital organs within each group. A revision of the taxon Macrostomida is in preparation by the first author. Based on the distal features of the stylet, Macrostomum coomerensis belongs to the Macrostomum tuba group and Macrostomum greenwoodi to the Macrostomum orthostylum group. Macrostomum coomerensis is characterised by having the male stylet formed like a cork-screw with a distal enlargement, and a subterminal opening of the ejaculatory duct. On the basis of these features the species belongs to the Macrostomum tuba group. Macrostomum species with a spiral stylet are: MZ. /eptos An Der Lan, 1939, M. lewisi Ferguson, 1939, M. reynoldsi Ferguson, 1939, M. riedeli Ferguson, 1940, M. delphax Marcus, 1946, M. contortum Beklemishey, 1951, M. phytophilum Beklemishev, 1951, M. phocorum Marcus, 1954, M. spirale Ax, 1956, M. poznaniense Kolasa, 1973, M. bicurvistvla Armonies & Hellwig, 1987, and M. extraculum Ax & Armonies, 1990. Of these species, however, only M. leptos, M. reynoldsi, and M. extraculum have the characteristic distal enlargement of the stylet which assigns them to the Macrostomum tuba group. The other species listed above belong to the Macrostomum orthostylum group. Macrostomum coomerensis, M. reynoldsi, M. extraculum, and M. leptos are differentiated by the different length and outline of their stylets. The stylet of M. coomerensis (125\1m in length) is essentially longer than that ofboth M. reynoldsi (72um) and M. extraculum (68-72\m). For M. leptos, only the length of the body (0.8mm) and the outline of the stylet are known, The stylet of M. leptos differs greatly from that of M. coomerensis both in the nature of the distal swelling with subterminal pore and in having the proximal and median part of the stylet greatly PLATYHELMINTHES FROM THE COOMERA RIVER curved. Distally, the stylet is only weakly corkscrew-like. Macrostomum greenwoodi belongs to the Macrostomum orthostylum group based on the curved and evenly tapered distal region of the stylet. The main distinguishing character for this species concerns the outline of the distal termination of the stylet. The distal tip in M. greenwoodi is obtuse, with a subterminal ejaculatory duct opening. In this respect, and in not having a cork-screw like stylet, the following 9 species resemble with M. greenwoodi: M. curvituba Luther, 1947, M. infundibuliferum Plotnikov, 1905, M. johni Young, 1972, M. longituba Papi, 1953, M. lutheri Beklemishev, 1927, M. mediterraneum Ax, 1956, M. magnacurvituba Ax, 1994, M. mosquense Beklemishev, 1951, and M. tenuicauda Luther, 1947. With the exception of M. /utheri and M. greenwoodi all other species have stylets with the ejaculatory duct opening subterminally on the concave side of the stylet. The stylets of M. greenwoodi and M. lutheri have ejaculatory openings subterminally on the convex side of the stylet. M. lutheri was established by Beklemishev (1927) only on the characteristic termination of the stylet. Other morphological characters are unknown. Therefore, the only main differential character resides in the complicate enlargement of the distal termination of the stylet. ACKNOWLEDGEMENTS Financial support was given by the Deutsche Forschungsgemeinschaft to Dr A. Faubel. We thank Prof. G. Grigg, Head of Zoology 519 Department, Prof. Jack Greenwood and staff for facilities and help, University of Queensland, Brisbane. Dr L.R.G. Cannon, Queensland Museum, Brisbane and Prof. K. Rohde, University of New England, Armidale, NSW, assisted with help and arrangements. LITERATURE CITED BEKLEMISHEYV, V.N. 1927. Uber die Fauna des Aralsees. Zugleich ein Beitrag zur Morphology und zur Systematik der Dalyellioidea. Zoologisches Jahrbuch, Systematik 54: 87-138. BEKLEMISHEYV, V.N. 1951. The species of the genus Macrostomum (Turbellaria, Rhabdocoela) of the USSR [in Russian]. Bulletin of the natural Society .. of Moscow (Biology, new series) 56, 31-40. DORJES, J. 1968. Die Acoela (Turbellaria) der deutschen Nordseekiiste und ein neues System der Ordnung. Zeitschrift fiir zoologische Systematik und Evolutionsforschung 6: 56-452. FAUBEL, A., BLOME, D. & CANNON, L.R.G. 1994, Sandy beach meiofauna of eastern Australia (southern Queensland and New South Wales). I. Introduction and Macrostomida (Platyhelminthes). Invertebrate Taxonomy 8: 989-1007. HASWELL, W.A. 1905. Studies on Turbellaria. Quarterly Journal of microscopical Science 49: 425-467. TRENCH, R.K. & WINSOR, H. 1987. Symbiosis with dinoflagellates in two pelagic flatworms, Amphiscolops sp. and Haplodiscus sp. Symbiosis 3: 1-22. WINSOR, L. 1988. A new acoel (Convolutidae) from the north Queensland coast, Australia. Fortschritte der Zoologie/Progress in Zoology 36: 391-394. 1990. Marine Turbellaria (Acoela) from north Queensland. Memoirs of the Queensland Museum 28: 785-800. BOWERBIRD (PTILONORHYNCHIDAE) BIOMETRICS, WITH OBSERVATIONS ON SEXUAL DIMORPHISM AND INTRASPECIFIC VARIATION CLIFFORD B. FRITH AND DAWN W. FRITH Frith, C. B. & Frith, D. W. 2001 06 30: Bowerbird (Ptilonorhynchidae) biometrics, with observations on sexual dimorphism and intraspecific variation. Memoirs of the Queensland Museum 46(2): 521-542. Brisbane. ISSN0079-8835. Australasian bowerbirds (Ptilonorhynchidae) belong to 18 or 19 species made up of 35 subspecies. Comprehensive biometrical data are presented for the Ptilonorhynchidae. Mean values and ranges of measurements of each species and subspecies are given, and interspecific and intraspecific variation in size and body proportions discussed. These and plumage traits are used to assess species and subspecies. We accept 20 species including 31 subspecies. Ai/uroedus crassirostris, Sericulus ardens and Chlamydera guttata are maintained as species, while C. nuchalis subspecies are reduced from four to two. Ai/uroedus melanotis joanae and Chlamydera guttata carteri are considered valid and Sericulus chrysocephalus rothschildi invalid. Bowerbird, biometrics, sexual dimorphism, variation. Clifford B. Frith and Dawn W. Frith, ‘Prionodura’, P.O. Box 581, Malanda 4885, Australia; 8 May 2000. Gilliard (1969) considered the Australasian bowerbirds (Ptilonorhynchidae) consist of 8 endemic genera, containing 18 species, with 32 subspecies. With subsequent revisions the present consensus has 18 or 19 species, and 35 subspecies (Beehler & Finch, 1985; Christidis & Boles, 1994; Frith et al., 1995; Frith & Frith, 1997a,b, 1999; Schodde & Mason, 1999). Bowerbirds attract attention particularly because of the elaborate bowers dd build and decorate. These are related to their polygynous mating system and associated ¢ promiscuity (Gilliard, 1963; Borgia, 1986). Adult ¢ ¢ show remarkable interspecific diversity of ornate plumages, bower structures and decorations, courtship, and vocalizations. They use these in complex and ritualised displays at traditional sites (Diamond, 1986; Borgia, 1986, 1995; Frith, 1970; Frith & Frith, 1989, 1990, 1993, 1994, 1995a, 2000a,b,c,d; Johnsgard, 1994). Size differences between the sexes (dd typically larger than 2 ° ) are particularly pertinent in most polygynous vertebrates but no comprehensive review of size variation among, bowerbirds has been published. Gilliard (1969) provided size ranges for basic traits without indicating either sample sizes or subspecific groupings. Cooper & Forshaw (1977) gave measurements for “five or more specimens” of species. Measurements for some subspecies appear in Schodde & Mason (1999), without sample sizes or means. Biometrical data are available for Amblyornis (Schodde & McKean, 1973; Frith & Frith, 1997b, 1998), Sericulus (Diamond, 1969; Lenz, 1999), Ptilonorhynchus (Schodde & Mason, 1999) and Chlamydera (Frith & Frith, 1997a, 1999), Some bowerbird subspecies have been distinguished on geographical plumage and size variation (Gilliard, 1969; Schodde & McKean, 1973; Diamond, 1969, 1972; Coates, 1990; Lenz, 1999; Frith & Frith, 1997a,b; Schodde & Mason, 1999). We examine our biometrical data with respect to subspecies defined by these authors. Differences in average population sizes has limitations, given that size is a continuously varying trait. While this is not a formal taxonomic revision, we record key broad plumage characters where appropriate. Genera and their sequence herein are a combination of those of Beehler & Finch (1985) and Christidis & Boles (1994). We largely follow Gilliard (1969) at the species and subspecies levels. That is, as taxa are acknowledged in Gilliard’s species accounts. Since Gilliard’s (1969) text was written Ai/uroedus buccoides oorti (Mees, 1964), Se. chrysocephalus rothschildi (Schodde & Mason, 1999), C. nuchalis oweni and C. n. yorki (Frith & Frith, 1999) have been rejected. Al. b. cinnamomeus (Mees, 1964), Al. melanotis joanae (Ford, 1977), Am. macgregoriae kombok, Am. m. nubicola (Schodde & McKean, 1973), Am. m. amati (Pratt, 1982), Am. m. lecroyae (Frith & Frith, 1997b) and C. guttata carteri (Frith & Frith, 1997a) have been erected or resurrected, C. 7. guttata and Se. aureus ardens have been raised to species level (Schodde, 1982; Lenz, 1999). We agree with TABLE |. Measurements (mm) and weights (g) for Ailuracdus buccoides. Wing Tail farsus ‘Total Sill Bill Sill Weight lengin lenelhy lencth) fewd —lencth = owedlh depth length A, }. buccoees Moles adult Mean 1346 so. 443 «6631 a4 wr \72 sD 77 06988 2a) pg gus oF Mot lag a Ma 24 27.6 45 Mae ag as 4)6 «86835 «6353 «(ON o 16 14 14 z Wat 3 | Females, adutt Mears 130 85 369 582 295 95 iat sD Tle 62B 197) Beh 1S O86 4 Min {fe 78 346 4550 Ga 83 150 Max ag 34 470) COBH 22 OT 152 f 5s 1& 14 7 15 15 2 AL o sheorit Males, adult Mean 136 2 we SY WS 35 142 ha 344 a3 115 164 146 «049 5,2 Mir, 130 56 3683 S76 260 89 is Max 14g 96 Any &26 33.4 ‘3 150, f ie 16 16 a] \s 16 7 Females, adult Mewar) (32 6 <3W2 576 28.6 af 140 sD 1 26! 112 214 132 OM 9.05 Min 128 a2 354 558 269 B4 6 Mee 136 30 381 si2 NT $5 150 i \2 12 12 7 a] te 7 4. & gersioronin Males: adult Mean {34 90 S77 564 WS4 OF 126 199 sD 416 a4 {sT 36 106 «0.80 078 12,26 Mir VPS as 22S Sea 276 6s 20 No Max 143 $8 406 BS 3425 7 ‘at 15g n ae S) 42 36 42 4} 2 3 Femades, adull Mean 129 Q5 38f StH ast M1 417) (128 sp AST 6330 131 152 O91 OS) O55 I82 Min 119 m 33 S59 265 4) itr 100 Man 138 83 faz 6BnY) 6S m3 123 150 “ 46 45 a5 a0 ad ad 3 30 4. & onnameametis Malwa: atiuil Mean var a gat sa) 306 on 150 so $44 279 146 183 133 o8) Min 130 85 370 S64 277 a2 Mie My 35 447 G24 326 {te n 13 48 3 " 8 1B ' Femabes: axdull Mean 13a ay 370 S70 292 96 130 so ae7 224 tis o78 141 O82 Min t2g a4 354 56.) 27,1 7 Mv iat 9 383 543 316 105 n 17 7 16 a 7 16 ' All subspecies: Males. adult Meaty 196 o) 382 S98 305 98 126 mt sp 510 a75 17s tee 128 OSD «OFA 1227 Min 123 6 329 565 ett 65 420 110 Mae 143 ao 47 S&S Sag we ist \72 n 9 7 st 6 sul 84 < 3 Forales adull Mean iy f S65 Si wi a2 117 129 50 $9 375 135 [68 (20 OSS OSM 15,06 Mitr Wa 7 Sd S39) BBS By we yon Maw 141 9% 410 822 B22 107 Ra 152 n 0 as or LP 6? 7 4 40 these decisions. 4/, crassirvosiris was considered a subspecies of d/. melanotis by Schodde & Mason (1999) but see our discussion herein. We MEMOIRS. OF THE QUEENSLAND MUSEUM use vernacular names of Beehler & Finch (1985) and Christidis & Boles (1994) except Spotted Catbird for 4/. melanotis and Yellow-breasted Bowerbird for C. lauterbachi; we retain the traditional Black-eared Cathird and Lauterbach’s Bowerbird, respectively. METHODS We examined all sexed skin specimens with a locality of origin in 32 collections (see Acknowledgements). The catalogue numbers of all specimens used and the measurements taken to produce Tables 1-21 are tabulated and lodged in the Library of the Queensland Museum: copies may be obtained trom the authors or front the library. Standardized measurements were taken by CBF with the same instruments. * Wing length’ 1s the flattened and straightened chord, measured witl a stopped steel decimal rule. “Tail centrals’1s the maximum length of the longest of the central pair of rectrices, from point of feather entry into skin to its terminal tip, and ‘tail length’ likewise but to the tip of the longest feather ather than the central pair, Tails were measured with an unslopped narrow steel decimal rule. When central and outer tail rectrices were the same length. or <3mm dilferent. only the longer (usually ‘tail length’) is given. Where there 1s intrageneric variation. both are given. Other measurements were taken with new tine-pointed stee! electronic digital vernier eallipers (checked/zeroed daily) to the nearest whole decimal point, ‘Bill length’ is from the union of bill and fore skull (cranio-manillary hinge) to the upper mandible tip. We measured bill lengths only [rom specimens with a complete upper mandible and on which the undamaged cranio-maaillary hinge could be confidently located. ‘Bill width’ and ‘bill depth’ (of fully closed bills only) were taken at the anterior nostril edge. *Total head length’ is the maximum distance lrom rear skull to the upper mandible tip, measured only from specimens retaining sufficient skull bone. Some specimens may appear complete in this respect but are not-so total head length figures are minimums. “Tarsus length’ is from the intertarsal joint to the lower edge of the last undivided scute (scale) before the toes diverge. Our measurements of live wild birds were generally similar to those from museum specimens, but we only include live bird measurements where samples of museum specimens are relatively small. Measurements of live birds included are for 10 immature/subadult 3 Sc, dentirosiris; 4 adult and 2 immature 4 and BOWERBIRD BIOMETRICS 52 2 2 Archboldia papuensis sandfordi; 7 subadult and 42 immature d Prionodura newtoniana; 3 adult, 1 subadult and 17 immature 6 Pr. violaceus minor. As a large majority of Schodde & Mason (1999) wing and tail length ranges fall well within the limits of our ranges we conclude that their samples were smaller. From our samples we evaluate previous assessments of several size-related characters. Gilliard (who used both “culmen” and “culmen from base”), Cooper & Forshaw (1977) and Schodde & Mason (1999) present “‘culmen length” or “exposed culmen”’, as opposed to bill length. Culmen length is the same as exposed culmen, and is measured from where anterior forehead feathers no longer cover the culmen to the bill tip, and is not of an entire structure. Thus culmen measurements are shorter, often exclusively, than our bill lengths. Culmen length appears more subjective and variable than bill length and may mislead as it is not structural bill length (Frith & Frith, 1997c: 173; Frith & Beehler, 1998; P?. violaceus below). Some 6 ¢ have a nuchal or fuller crest, which was measured from the posterior crest base to its tip (Schodde & McKean, 1973). We also measured crest length as viewed from above (Frith & Frith, 1997b, 1998). All measurements are in mm, Weights (in grams) were noted from specimen labels. We include additional weights obtained from live birds as follows: 27 adult ¢ and 36 2 Al m, maculosus; 46 adult and 10 immature ¢d and 1 @ Sc. dentirostris; 3 adult and 2immature ¢ and2 2 Ar. p. sanfordi; 79 adult, 7 subadult, 42 immature ¢ and 12 adult @ Pr newtoniana; 3 adult, | subadult, and 32 immature 3 Pt. v. minor; 14 adult, 4 subadult dd, and | adult 2 C. maculata; 5 adult 3 C. nuchalis. Differences in weight between sexes are commented upon only where samples are adequate, because body weight is subject to bias with respect to time of day/year recorded. Data are reported for adult samples unless otherwise stated. Data for markedly sexually dimorphic species in which dd may have an adult, subadult (i.e. trace to almost complete adult 3 plumage intruding into -type), and immature (purely 2 -type) plumage are presented separately for each age group. For monomorphic species, in which adult ¢ 3 and 2 @ are similar but ¢¢ have a discernibly different immature plumage, data are given separately. Data for 2 are of adults, but some samples might inadvertently include the odd younger Lod individual, given the similarities of 2 plumages. We exclude conspicuously smaller individuals (i.e. presumed juveniles-immatures). We do not describe plumages of monotypic species or those of nominate subspecies as these are widely available (Rand & Gilliard, 1967; Gilliard, 1969; Cooper & Forshaw, 1977; Beehler et al., 1986; Coates, 1990; Donaghey, 1996; Schodde & Mason, 1999). In a few instances we refer to numbered colours of Smithe (1975), with his nomenclature indicated by capitalisation. Geographical ranges indicated do not include controversial or vagrant records. West Papua (WP) was Irian Jaya. SYSTEMATIC NOTES Family PTILONORH YNCHIDAE Ailuroedus buccoides (Temminck in Temminck & Laugier, 1835) White-eared Catbird (Tables | and 22) 2 © average 5% smaller than 6 ¢ in tail, tarsus and bill lengths, 6% in bill width, 7% in bill depth and 9% in weight. Tail/wing ratio is 66%, tarsus/wing 28% and bill/wing 22%. Measurements and body size ratios are near uniform across subspecies, except in geislerorum which has fractionally shorter wings. We follow Mees (1964) in synonymising oorti with A. 6. buccoides. Ailuroedus buccoides buccoides (Temminck in Temminck & Laugier, 1835) Al, b, aorti Rothschild & Hartert, 1913. ? 2 average 6% smaller in wing, tarsus and bill lengths and bill width than dd. Sexual size dimorphism is little different in the other subspecies. Characterised as “Under surface pale cinnamon with large black dots; crown brown, usually tinged with greenish.” (Mees, 1964: 127). Range: W Papuan Islands (Is) and Vogelkop lowlands, coastal W and central Geelvink Bay to Siriwo River (R) in north New Guinea. Triton Bay E to upper Fly R in south New Guinea(NG). Ailuroedus buccoides stonii Sharpe, 1876 Underparts deep cinnamon, black spots smaller than other subspecies. Crown dark blackish-brown, sometimes tinged greenish, distinctly blacker than in other subspecies. Range: SE coastal Papua New Guinea (PNG) from Amazon Bay to upper Purari R, including 524 TABLE 2. Measurements (mm) and weights (g) for Ailuroedus crassirostris. Wing Tail Tarsus Total Bill Bill Weight length length length head length width length Males: adult Mean 166 124 471 647 34.0 10.0 215 sD 7.38 5.60 1.85 2.50 1,68 056 35.41 Min 151 WW 40.4 59.6 31.2 8.4 167 Max 179 139 50.0 69.6 38.0 10.8 289 n 38 36 37 25 37 36 13 Females: adult Mean 161 120 458 631 332 38 193 SD 6.03 5.35 2.16 2.15 1.64 0.51 15.20 Min 150 108 416 604 305 8.9 169 Max 172 127 43.3 67.3 36.9 10.8 2it n 25 25 26 1 26 26 7 the Karimui, Bomai, Soliabeda areas, E Highlands (Diamond, 1972). Ailuroedus buccoides geislerorum Meyer, 1892 Al. b, molestus Rothschild & Hartert, 1929. Underparts as in nominate form but, unlike it and all other subspecies, crown light tan brown. The pure white ear covert feathering extends forward onto the lower lores. Range: Japen Is and north NG from Mamberamo R to Collingwood Bay, PNG. Ailuroedus buccoides cinnamomeus Mees, 1964 Underparts save throat deep cinnamon, as in stonii, with large black spots. Crown as nominate buccoides but on average tinged more green. We found, as did Mees (1964), that at the eastern end of its range 5 of 6 individuals had a darker crown colour, thus approaching sfonii further east. Range: south WP from Mimika R, E to upper Fly R and Lake Kutubu, PNG. Al. b. geislerorum is distinctive in its paler tan crown and white ear covert feathering extending forward to include the lower lores. Stomii is equally distinctive in its black crown and lores, contrasting pure white ear coverts, and deep cinnamon ventral colouration. 4. 6. buccoides and cinnamomeus, have brown crowns (variably washed greenish). Both have heavy black spotting on the white ear coverts, but cinnamomeus has considerably darker cinnamon underparts and usually more white (as opposed to dark brown in buccoides) lower lores. These differences are greater than the “northern form, brown cap” and “western and southern form, blackish cap” defined for field use (Beehler et al., MEMOIRS OF THE QUEENSLAND MUSEUM 1986), which may give the erroneous impression of only two subspecies. Ailuroedus crassirostris (Paykull, 1815) Green Catbird (Tables 2 and 22) 29 average 10% lighter than dd. Adult tail/wing length ratio 74%, tarsus/wing length 28% and bill/wing length 20%. Thus this species is proportionately longer-tailed than A/. m. maculosus (Table 3), to its N within Australia Mean bill/tarsus length is 72.3% compared to 77.5% in Al. m. maculosus. Range: Coastal Australia from Dawes Range (Ra), Qld to E of Canberra in NSW. Ailuroedus crassirostris has been considered the southern subspecies of A/. melanotis (Pizzey, 1980,1997; Simpson, 1984,1999; Schodde, 1976 a,b,1986) or a separate species (Mack, 1953; Slater, 1974; Slater et al., 1989). Schodde & Mason (1999) argue that “macu/osus cannot be combined with melanotis without bringing in crassirostris as well”, an argument we do not accept. While the 10 subspecies of A/. melanotis demonstrate considerable geographical variation, they all share the traits of a dark crown spotted with conspicuous white to buff markings, strongly contrasting black chin feathering and ear coverts that give a ““black-eared” appearance, and broad dusky edging to predominantly whitish throat and chest feathering to give a scalloped appearance, As Al. crassirostris (a) lacks the first three characters, (b) has a green throat and chest finely streaked by white central feather lines, (c) is longer tailed and shorter billed than Al m. maculosus and (d) is geographically isolated by more than 600 km, we treat it as a distinct species. We find the acceptance of AL. crassirostris as a species, on morphological and zoogeographical grounds, consistent with similar treatment of C. guttata as distinct from C. maculata (Christidis & Boles, 1994:74; Schodde & Mason, 1999). The latter are separated by 250 km. Al. crassirostris evolved in isolation from A/. me/anotis stock, the issue being to what level of taxonomic significance it has differentiated. Equivocal (Schodde & Mason, 1999) allele frequency data (Christidis & Schodde, 1992) supported separation of crassirostris as a species. Given the foregoing, and that tissue samples remain unavailable for the Aru Is population of Al. m, melanotis, for Al. m. arfakianus, and for other montane populations (Schodde & Mason,1999), we concur with Christidis & Boles (1994) in BOWERBIRD BIOMETRICS TABLE 3. Measurements (!nm) and weights (g) for Ailuroedus melanotis. Wing Tail Tarsus Total — Bill Bill Bill = Wenght length length length head length width depth Males: adult Mean 166 «119 457 70.0 38.0 13 221 SD 613 414 237 222 168 O72 21.0) Min 156 12 38 666 3249 101 200 Max 77 125 4686 754 411 126 242 " 24 24 25 15 23 23 3 Females: adutt Mean 158 145 436 656 356 106 SD B85 417 154 313 1.44 O7B Min 147 10 ©4090 «614 © «334 a0 Max 166 22 457 #6890 379 #114 n 10 g g 4 3 9 Am. maculesus Males: adult Mean 143, 103 438 63.2 340 98 141 173 SD 438 439 181 157 1.36 052 O77 1404 Min 140 34 #399 B04 29.6 B87 128 145 Max 1ST Wi2 479 «#6660 376 190 47 205 n 57 40 4) 23 45 42 5 40 Females; adult Mean 146 98 431 625 33.3 $3 133 169 sD £00 434 182 225 134 055 O47 14.07 Min 136 30 402 579 309 63 129 140 Max 189 106 470 B74 359 {07 141 199 n 36 30 ww \9 30 30 5 50 A. m. arlekianus Maies: adult Mean 162 «123 467 «69.0 374 108 247 SD 51) 528 185 185 177) O94 5.77 Min 156 17 440 665 351 90 240 Max 71 1339 503 721 404 120 250 n " 12 12 8 W a 3 Females: adult Mean 155 Vs 461) 653 355 106 216 SD 270 252 106 460 1.02 0.62 173 Min 151 i116 452 588 341 aT 215 Max 158 123 478 #668 366 N12 218 n 5 5 5 4 5 5 3 Am, melanocephatus Males: adult Mean 156 3 467 6BA2 361 104 223 SD 410 435 187 207 #147 O58 2oi14 Min 147 «116 «6421 6386 3536 92 204 Max 163, 29 515 15 407 #116 285 " v 30 34 24 31 4 7 4 7 7 A. m. misoliensis Males: adult Mean 165 = 140 a6 233 SD Min Max n i} i} i] ‘ Females; adult Mean 169 1255 457 707 36,8 11.36 18) 424 071 316 212 148 Min 166 125 43.4 35.2 10.3 Max 72 126 «ATA 3 (124 n 2 2 2 1 2 2 Al m@m joanae Males: adult Mean 147 «100 429 619 339 QT 178 sD 283 212 198 233 0.07 Min 145 g8 415 322 96 Max 149 yor 44,3 355 37 n 2 2 2 { 2 2 1 Females: adult Mean 137 94 40.2 596 S22 89 150 SD 286 435 104 158 O59 029 {2.02 Min 133 90 391 S72 NT 484 4) Max 147 100 417 612 332 38) 158 n 5 4 5 5 5 5 2 All subspemes. Males: adult Mean 156 116 456 674 36.1 105 141 198 SD 786 968 235 3.37 229 O86 O77 3025 Min 140 34° 36.6 6 «6=BO4A 629.6 a7 128 145, Max vw 140 S15 754 413 126 14,7 285 " 151 139-156 Ba 137 134 5 65 Females: adult Meany 1st tio 442 645 344 100 134 182 so 796 1006 216 314 168 O84 O45 2737 Min 133 90 38.) S572 309 Ba 129 140 Max 172 126 489 7Oo7 392 124 141 261 f 102 34 95 58 96 95 6 69 Ee ww th ao maintaining A/. crassirostris as a separate species. Ailuroedus melanotis G.R. Gray, 1858 Black-eared Catbird (Tables 3 and 22) Body ratios are similar to those of A/. crassirostris. 2 2 average 5% smaller than dd in bill width and 8% in weight. Proportions of size sexual dimorphism among the various characters vary slightly between subspecies, but no pattern is evident. Subspecies are generally similar in leg/wing and bill to wing length ratios, but there is some variation in tail/wing length. Ailuroedus melanotis melanotis (Gray, 1858). Darker areas of head plumage and dark edging to the upper breast feathers are blackish to black. Range: Aru Is and Trans-Fly lowlands of south NG. Ailuroedus melanotis arfakianus Meyer, 1874 Tail/wing length ratio is 68%. The tail is thus proportionately short, as is that of joanae (at 69%). Throat much darker than in me/anotis melanotis, being black spotted with dirty white to pale buff down to the chest. The latter is not marked blackish as in me/anotis melanotis, but is darker green marked with narrow and pointed pale buff feather centres. Pale patch behind black ear coverts large and clear white; large spotting on the black crown paler (white or almost so) than in melanotis melanotis. Range: Arfak and Tamrau Mountains (Mts), Vogelkop, possibly also Kumawa Mts, WP (Diamond, 1985). Ailuroedus melanotis maculosus Ramsay, 1874 Head, nape and upper back less black and far paler than in nominate form, with dark brownish feather edges. Upper back spotting very small, pale buff and feathers lacking blackish edging. Green of back and upper tail paler and slightly more yellowish than in nominate form. Underparts far paler buffy-white, more green, than in melanotis melanotis and feathers with broad brown-green edges. Chest far paler and less heavily marked than melanotis melanotis. Galbraith (in Mees, 1982) found 4 ¢ d and2 2 2 had mutually exclusive maxilla depths (11-12.5 versus 10-10.5, respectively) which Mees (1982) thought separated the sexes. In measuring bill depth of 5 of each sex, we found considerable MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 4. Measurements (mm) and weights (g) for Scenopoeetes dentirostris. Wing Tail Tarsus Total Bill Bill Bill Weight length fength length head length width depth length Males: adult Mean 149 1046 317 576 30.7 103 135 158 sD 386 226 109 177 #115 O47 O43 11,12 Min 141 100 294 521 283 94 126 132 Max 156 108 340 598 329 112 144 199 n 35 35 34 23 35 35 20 5 immature/subadult Mean 150 103 315 587 300 104 14 164 sD 281 155 181 100 150 O31 O81 12.48 Min 146 101 280 574 272 100 133 151 Max 155 106 33.0 597 314 107 149 185 10 8 7 5 7 ¥ 7 10 n Females: adult Mean 146 #101 316 581 310 107 #136 169 sD 316 3.08 142 137 116 O39 O61 12.62 Min 138 96 294 540 2681 99 127 157 Max 152 107 345 597 330 113 146 182 n 25 25 25 16 24 26 8 3 overlap. Range: Australian Wet Tropics, from Mt Amos to Seaview-Paluma Ra, Queensland (Qld). Ailuroedus melanotis melanocephalus Ramsay, 1883 Tail/wing length ratio is 78%. Like nominate form but underparts generally darker, blacker on chest and throat, and more rufous below. Crown blacker, as the buff spots are slightly smaller and/or sparser (more so in two birds from ‘Yule I.’). Range: mountains of SE NG, west to Herzog Mts in north and Mt Karimui in south, including Owen Stanley, Hydrographer, Astrolabe, Wharton and possibly Kratke Mts. Ailuroedus melanotis jobiensis Rothschild, 1895 Dorsally like melanotis melanotis but spotting on the blackish crown less pure white, more buff. Ventrally quite different in that the chin, throat and upper chest are blackish with only fine buff spotting thereon. Remaining underparts similar to, but darker than me/anotis melanotis. This subspecies is more like gu/ffaticollis ventrally, the throat to upper chest being darker due to smaller pale spotting. White patch behind black ear coverts more obvious than in most subspecies. Range: Bewani, Torricelli, Prince Alexander Mts, middle Idenburg R, and (tentatively) Adelbert Mts, NG. Ailuroedus melanotis guttaticollis Stresemann, 1922 Like nominate form but spotting of crown to upper back more rufous, less buffy. Throat and BOWERBIRD BIOMETRICS chin much darker and underparts generally more rufous. In colour and markings indistinct from Jobiensis. Range: Sepik and (tentatively) Jimi R, PNG. Ailuroedus melanotis astigmaticus Mayr, 1931 Tail/wing length ratio is 81%, this being the longest-tailed subspecies. Like melanotis melanotis but crown blacker, with fewer, finer, and paler spots. Range: mountains of Huon Peninsula, PNG. Ailuroedus melanotis facialis Mayr, 1936 Tail/wing length ratio is 71%. Like nominate form but throat darker, less white and more buff. Spotting of crown and upper back much darker, more cinnamon (123A to 123). This is not a “poorly differentiated subspecies” (Cooper & Forshaw, 1977) when compared with its nearest conspecific population of A/. m. jobiensis, as the latter has a much darker throat, breast, and crown. Range: Nassau and Oranje Mts, WP. Ailuroedus melanotis misoliensis Mayr & de Schaunsee, 1939 Like arfakianus but blacker at base of throat, and averaging much larger but with some overlap (Mees, 1965). Range: Misool I, WP. Ailuroedus melanotis joanae Mathews, 1941 Tail/wing length is 69%, the tail being similar in length to the other Australian subspecies maculosus (68%). Compared to maculosus: smaller, blacker on crown, nape, and mantle, less black on face and chin, throat whiter, lower breast, flanks, belly, ventrals and under tail coverts far cleaner (less marked with buff-green feather edging) and thus clean yellowish on these underparts. Differs from other Australian catbirds in having all underwing coverts pure white (save a few exceptions involving only the outermost greater primary coverts). In A/. crassirostris and Al. m. maculosus these feathers are centrally heavily pigmented with blackish-grey and their edges dirty whitish with the faintest of greenish wash in places. Scalloping on chest notably blacker than on the other Australian subspecies, but less black than on the nominate form in New Guinea to which joanae is overall more similar than it is to maculosus. Range: QLD, Australia. Pascoe R and Iron Ra areas to Rocky R, Mellwraith Ra, Cape York Peninsula. Wn bho ~J Ford (1977) resurrected joanae and defined some plumage characters (enlarged upon by Schodde & Mason, 1999) distinctive from maculosus. Ironically, the only character Mathews (1941) used in naming joanae, that it is smaller than maculosus, is invalid. To test this we compared sizes of joanae (2 dd and 5 2°, given that we agree with Ford (1977) that the holotype is 2) with northernmost maculosus. from Mt Finnegan (4 6 ¢ ), Gap Creek, 7 km N of Bloomfield (3 ¢¢, 1 2), Granite Creek, Bloomfield R (3 ¢d,4 @ 2), and Big Tableland (2 4,1 2). Results show joanae no smaller than closest maculosus. Scenopoeetes dentirostris (Ramsay, 1876) Tooth-billed Bowerbird (Tables 4 and 22) 2 2 7% heavier than d d, but this may be due to time of year few weights were taken. This species has by far the shortest proportional tarsus length and is arboreal except when, rarely and briefly, on their forest floor courts (Frith & Frith, 1994). Immature/subadult ¢ d, of Table 4, were all live court-attending birds identifiable as such by their pale mouth (black in adultd ¢; Frith & Frith, 1995a). Measurements of wing length by Cooper & Forshaw (1977) are near the higher limit of our sample, while most of their other measurements are outside our ranges. As this is the only potentially lekking bowerbird (Frith & Frith, 1995a) in which males form an exploded (vocal rather than visual contact (Johnsgard, 1994)) lek it is surprising to find adult d ¢ little larger than ° 2 (and less so than in monogamous Ailuroedus). While Scenopoeetes has long been considered closest to monogamous Ailuroedus, on similarities in plumage and egg colour, behavioural and molecular data indicate its closer relationship to typical, polygynous, genera particularly Amblyornis (Frith & Frith, 1993; Kusmierski et al., 1993, 1997; Schodde & Mason, 1999). Range: Australian Wet Tropics uplands, from Mt Amos to Mt Elliot, Qld. Archboldia papuensis Rand, 1940 Archbold’s Bowerbird (Tables 5 and 22) 2 2 8%, 22% and 9% shorter in wing, tail and tarsus lengths, respectively, than dd but the same in bill length. With a tail/wing length ratio of 96% (103% for dd only) this is the longest-tailed bowerbird. We assume Gilliard (1969) and Cooper & Forshaw (1977) included some younger individuals in their adult dd TABLE 5. Measurements (mm) and weights (g) of Archboldia papuensis. Wing Tail Tail Tarsus Total Bill Bill Bill ~Weight length length central length head length width depth length length A. p. papuensis Males: subadult Mean 159 152 137 39.6 652 33.0 8.4 173, SD 21 5.7 5.7 og 15 0.2 a3 35 Min 157 148 133 369 641 328 82 170 Max 160 156 141° 402 662 331 86 175 n 2 2 2 2 2 2 2 2 immature Mean 158 «135° «125 «422 «64.0 328° 82 sD 3.50 966 727 128 #%4101 145 036 Min 156 127 120 408 629 307 77 Max 163 148 «#136 439 649 3414 65 n 4 4 4 4 3 4 4 Females: adult Mean 148 126 16 38.1 640 33.4 LB) sD 439 365 635 173 142 160 0.33 Min 144 122 12 366 626 31.0 65 Max 155 130 123 407 661 353 92 n 5 5 3 5 5 5 5 A. p. sanforat Males: adult Mean 168 173 146 434 663 33.5 85 10 184 SD 339 859 7289 #170 130 O91 056 043 3.86 Mint 161 155 133 394 646 318 75 87 180 Max 174 189 165 459 698 359 94 10.3 190 n 2 20 18 21 15 20 20 9 6 immature Mean 165 152 137 43.4 663 336 683 99 186 sD 1.69 672 650 134 1.03 126 039 038 963 Min 163 145 132 417 644 B15 7.8 96 170 Max 167 162 150 454 677 357 68 103 195 n 8 7 7 8 7 8 8 3 5 Females: adult Mean 158 140 130 404 653 33.5 8.7 10.0 176 SD 474 480 433 165 102 131 O70 018 9.42 Min 148 132 125 371 64.0 315 7.8 98 163 Max. 163 149 139 427 665 356 97 10.2 185 n 9 9 8 3 6 9 9 4 4 All subspecies Males:adult Mean 166 #175 #148 433 666 336 84 96 184 SD 345 825 662 157 132 093 058 016 3.68 Min 161 155 136 394 650 316 75 94 180 Max 174 169 165 455 698 359 g4 98 190 n 17 16 4 17 W 16 16 5 6 subadult Mean 159-1520 «137-396 465.2 330 84 173 SO 24 57 57 O89 15 02 03 3.6 Min 157 148 133 389 641 328 8.2 170 Max 160 156 141° 40.2 66.2 33.1 86 175 n 2 2 a 2 2 2 2 2 immature Mean 163 1460 «(133 43.0 656 334 83 99 186 sD 423 11.59 869 139 148 133 036 038 9.63 Min 156 127 120 408 629 307 TF 9.6 170 Max 167 162 «4150 454 677 357 88 103 195 n 12 " i" 12 10 12 12 3 5 Females: adult Mean 154 135 126 39.5 647 335 88 100 176 SD 670 837 610 197 1,32 136 O58 O18 942 Min 144 122 12 366 626 31.0 78 98 163 Max 163 149 139 427 #665 356 97 102 185 n 14 4 a0] 14 " 14 14 4 4 because their wing length ranges of 155-170 and 155-162 are small versus our 161-174. The two geographically isolated forms of this bird are considered separate species by some (Gilliard, 1951, 1958, 1959; Peckover & Filewood, 1976; Collar, 1986) and indistinct, or invalid, subspecies by others (Frith et al., 1995). While lack of adult d specimens of papuensis MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 6, Measurements (mm) and weights (g) of Amblyornis inornatus. Wing Tail Tail Tarsus Total Bill Bill Weight length length central length head length width length length Males: adult Mean 134 95 93 364 607 30.1 8.4 128 sD 3.37 377 3.84 1.37 1.70 1.43 051 15.51 Min 129 88 B4 34.0 57.2 27.4 73 105 Max 140 104 100 39.20 «6380 06334 97 155 n 28 28 26 28 19 27 27 12 Fematles: adult Mean 130 a1 a1 35.2 60.2 30.2 B6 121 SD 31 277 3.00 1.18 1.89 1.37 0.87 15.21 Min 125 85 84 3340 «678 «6279 69 105 Max 136 96 95 37.8 63.7 32.1 10.0 146 n 18 18 ¥ 18 4 18 18 9 papuensis hampers comparisons, adult ? ° (Table 5) are smaller than those of sanfordi. Archboldia papuensis papuensis Rand, 1940 The smaller subspecies, averaging 6% smaller in wing and tarsus length, and 10% in tail length (22 only), but similar in bill size. This difference in tail length is exclusive, the range in this form being 122-130 but in sanfordi 132-149. The adult tail/wing length ratio averages 96% in 33 and 85% in 2 2. Plumage is generally less black, more grey, than that of sanfordi. Range: Ibele R (Balim Valley) to Lake Habbema region, Oranje Mts, Nassau Ra and Wissel Lakes, Weyland Mts area, WP. Archboldia papuensis sanfordi Mayr & Gilliard, 1950 Larger than papuensis papuensis and plumages blacker. Tail/wing length ratio averages 103% in 36 and 89% in 2¢. Adult dd fully crested. Mean exposed crest length of 20 dd is 91 (77-98, SD 5.41). Range: Mt Hagen, Giluwe, Tari Gap and S Karius Ra (unconfirmed on Kubor Ra), PNG (Coates, 1990). Amblyornis inornatus (von Rosenberg in Schlegel, 1873) Vogelkop Bowerbird (Tables 6 and 22) 2 2 average fractionally smaller than dd in body sizes and weight. Both sexes in this and other Amblvornis are identical or similar in tail/wing length ratios (Table 21), a trait shared by Scenopoeetes among polygynous genera (and also by monogamous Ai/uroedus), Range: Arfak, Tamrau, Wandammen, Kumawa, and Fakfak Mts, WP (Diamond, 1987; Uy & Borgia, 2000). TABLE 7. Measurements (mm) and weights (gz) of Amblvornis macgregoriae, BOWERBIRD BIOMETRICS 529 Wing Tall Tail Tarsus Total — Bill Bill Bill Weight length langth central length head lenoth width depth length tenath Alm, macegonas Malan) adult Meare 135 a7 eS 374 5879 286.0 76 120 SD 322 335 31/9 108 095 085 066 15.47 Min 129 al 79 355 554 25 67 104 Max 142 92 91 400 593 21 Bg 142 n 22 22 20 22 "3 22 22 7 immature Mean 133 aT 86 373 573 275 74 120 5D 384 955 949 143 116 #123 O48 822 min 126 80 60 MB 556 252 65 110 Max ia. 92 5} 406 601 299 82 198 v 22 2 19 22 16 22 22 14 Females: adult Mean 13) B5 834 36) 582 265 40 B89 (|23 5D 342 3/2 3.02 110 154 O82 O50 $71 Min 23 74 7 #333 S4t 270 68 5 Max 138 90 90 380 S99 296 a3 130 a 26 26 22 26 is 26 25 7 Am, germans Males adult Mean 130 87 84 963 S578 274 7.3 96 112 so 3yf 182 218 O68 (46 O86 O38 Bea Min 24 BS 8) 349 550 260 66 104 Maw 135 ‘90 8 378 #597 265 73 \27 " VW ” Q N 6 wt Th 1 5 imenestuire: Mean 126 BE 4 386 559 287 77 4 so 256 226 #333 088 #%136 #118 #O2F 287 Mire 122 83 76 353 534 246 73 V2 Max 130 ag a7 378 570 283 B2 We n a 8 6 3 6 8 7 4 Females: adult Mean (27 85 64 °360 STS 285 BS 5 sD 57° 351 3599 O78 (68 O85 0.65 7.88 Mite 125 a 7a 352 581 275 72 ine Mak 129 g2 9) 373 SAS 29.6 g2 128 n 7 7 7 7 3 7 7 4 Alm may Males: adult Mean 139 92 695 377 598 293 B82 95 145 sD 299 #306 287 186 210 O88 O84 45d Min 1348 a7 B39 338 564 284 7.0 140 Maw 145 38 55 403 6524 296 o4 146 n 22 21 20 22 10 21 2 1 2 immature Mearr 136 69 876 376 S84 289 74 1e7 sb 337) 470 580 100 (76 #110 O80 643 Min 134 Bt 78 3659 S67 era 6H 120 Max 143 Es) 98 389 607 B16 LE} 132 " 15 14 13 15 7 15 15 3 Females: adult Mean (54 89 866 364 59) 302 86 sb 316 383 356 O91 211 197 O61 Mir 127 B3 so 344 S49 273° «70 Maor 142 i 94 381) G24 386 ve nm 26 26 25 26 13 26 26 Am. hambok Males adult Mean 137 ay 84 374 5B) 281 74 BO 125 SD 368 320 969 1191 167 O90 O52 Us eT Mine 129 7a 72°«279 «4554 265 62 84 W0 Max 144 92 so 400 607 ah) 53 95 140 n 7 32 27 36 7 a7 6 5 7 immature Mean 135 85 63378 S79 278 75 9.0 124 5D 366 260 254 O87 126 O79 O8F O27! 335 Min 128 80 80 362 56) 263 57 ao V0 Maox 14) a0 a8 392 599 296 B87 Gr 134 n 7 7 7 7 10 7 vw Zz 7 Females: adult Mean iat 84 6 361 574 288 83 92 130 50 313 2982 291 194 23) UBO O74 Haw aid tin \27 7a 7 37 300 2a TG fe iat Max 138. 88 B7 398 S97 22 98 IMI 140 a 19 1@ " 19 15 {9 19 7 2 Al im mitweokis Males: adult Mean 135 aI 50 371 582 276 67 128 SDB 173 344 388 090 O71 095 O25 5.69 Min {33 a9 87 362 S76 268 64 122 Mata 188 95 s4 380 S590 287 69 183 n 8 x) 3 a Ki = | 3 3 imate: Mean 14 3) 69 36.3 S587 28) 73 126 §D (72 251 207 130 156 O92 O30 395 Min 132 a7 85 348 S560 268 70 125 Max 137 95 92 35) BOS 295 78 135 " 7 T 7 7 7 7 T 7 Females: ariutt Mean 131 89 66 355 579 26) 76 130 SD: 265 299 252 (2) O82 040 026 Ta Min 7 128, BB 65 337 870 2745 73 wg Max {37 95. 92 38| 605 295 Te Is " 3 4 3 4 4 4 a 4 Al im amatt Males: adull Mean 128 62 801 357 262 72 sp 144 035 1.84 Min 127 a2 a44 Maw 125 «825 37.0 n r J 2 | 2 ' 1 Alan fecrayse Males: adult Mean 132 82 60 368 544 2759 80 33 SD 187 256 356 095 1¢@ 116 O57 Min 130 rc) 75 383 «534 #259 75 Maw 105 Ba 84377) «6553 287 86 n 6 4 5s 5 2 s 5 i] immature Mean 1a 82 Bi 36.5 276 TA SD 200 120 225 G46 os9 42 Min 123 80 79 «#434b 272 A) Max 133, 83. B35 385 26.3 73 n 3 4 3 2 a z Femutes adult Mean 129 aI 81 379 265 84 n ' 1 ' 1 i] 1 All subspecies Males: adult Mea 136 86 BE 372 S82 282 TE 3.2 123 SD. 436 402 416 162 (|.83 (07 O67 O39 |233 Min \2a 78 73 «273 S34 259 82 as 104 Max 145 3B 95 403 824 308 a4 a6 145 a 102 o6 85 10) 51 joo 38 4 24 trnireature: Mean 133 aT 65 373 S76 278 75 90 22 sD 438 390 416 129 #155 #122 O84 O21 810 Min 122 au 7a 380 544 246 65 e8 m0 Max 143 39 5B 406 6807 GIB 58 3) 138 a 72 70 65 7 46 72 7 2 35, Females: adult Mean, 132 86 64 362 581 291 83 92 124 SD 364 404 377 1.27 #106 |.48 O65 O46 897 Min 123 7a 7 #307 S00 266 68 a6 108 Max 142 38 94398 624 WE 36 16) 140 n 82 B2 69 x] 54 63 a 8 17 Amblyornis macgregoriae De Vis, 1890 Maegregor’s Bowerbird (Tables 7 and 22) 2 2 average 9% wider in bill width than dd, both sexes having all bul the same bill length. Mean crest length from posterior base of 47 6 is 64 (46-75, SD 6.56) and exposed crest length of 96 3 4, 82 (52-105, SD 10,26). Amblyornis macgregoriae macgregoriae De Vis, 1890 Most similar in size to kombok and in proportions, for both sexes, to pubicola and kombok ? @ average 5% wider in bill width than TABLE 8. Measurements (rum) und weizhts (g) of Amblvornis subalaris. Wing Tail Tal Tatsus Tolal Sill Bil Weight length length centrak length. bead length width lenath Males: adult Mean 125 as 65 386 So4 259 73 96 8D 26) BaF S325 157? 144 126 O37 Min {21 80 7 #30) S46 235 oF Meor 120 3a 92 367 600 248 80 n au 28 a 2g iz 23 2s 1 subadult Mean 123 a7 BE 342 565 25.4 7I 85 SD i00 6115 6645 14) Me a S50 Miry 122 65 665 329 SB2 251 66 Maw \2a 68 a? 357? S665 257 75 n 3 3 3 3 2 3 3 1 smmealure Mean 124 89 a6 347 35 259 75 108 SD 280 2355 339 O76 $99 100 O82 556 Min 12 84 82 G35 528 244 66 104 Max (28 93 9% 29 S85 276 85 126 aa] ‘a 10 1 =} W W 5 Females adult Mean 124 87 85 34) 5668 265 76 108 SoD 374 274 236 157 (15 '18 Usa, 966 Miri WT 8 GO 3207 S52 248 G67 35 Max 13) 92 68 184 585 296 85 122 n 22 22 zz rd 7 2 2i 5 34. Mean crest length from posterior base of | dé 65 and exposed crest length of 22 dd §1 (35-92, SD 9,50), Range: W Kukukuku and Herzog Ra to W Owen Stanley Ra, PNG, excluding range of 4m. im. nubicola to the E (Schodde & McKean, 1973). Amblyornis macgregoriae germanus Rothschild, 1910 Bath sexes average 7% lighter and fractionally smaller in wing length than jmacerezoriae macgregoriae, but almost identical to it in tail length and similar in tarsus and bill lenyths and bill width. Mean crest length from posterior base of 3 do 48 (46- 30, SD 2.08) and exposed crest length of 10 dd 68 (56-79. SD 6.51). Range: mountains of Huon Peninsula, PNG. Amblyorois macgregoriae mayri Hartert, 1930 The largest subspecies, °° 5'% wider in bill width than ¢ d. Mean crest length from posterior base of 2 dd 72 (71-73, SD 1.63) and exposed crest length of 22 ¢4 90 (79-105, SD 7.79). Range: Weyland Mts, WP to E Star/W Hidenbure Mis, extreme western PNG, Amblyornis macgregoriae kombok Schodde & McKean, 1973 Similar in size and proportions to the nominate form. In bill width, 9 2 average 12% larger than MEMOIRS OF THE QUEENSLAND MUSEUM 24. Mean crest length from posterior base of 28 a5 4 63(51 -71,8D4) 9?) and exposed crestlength of 32 oo 84 (72-95, SD 5.59), Crest rather dense. Throat and upper breast pale-hrownish olive; lower breast, abdomen, and under taul-coverts rather hright light buffy brown (Schodde & McKean,1973). Range: Kubor, Hagen and Bismarek Ra, central PNG, probably W fo at least the Strickland R or Hindenberg Ra, and Eto Kratke Ra (Schadde & McKean, 1973). Amblyornis maceregoriae nubicola Schodde & McKean, 1973 The tail/wing length ratio fractionally longer than average for the species (as in ger manus), In bill width, &% average 13% wider than 3. Mean exposed crest length of 3.¢.¢ 83 (75-90, SD 7.75). & erest length trom posterior base 39-65 (n=3, SD=3.0) and rather densely feathered, entire under surface uniformly dull coffee-brown. Range: Mt Simpson-Dayman area, E Owen Stanley Ra, probably W to Mt Suckling, PNG (Schodde & McKean, 1973). Amblyornis macgregoriae amati Pratt, 1982 specimens indicate this is the smallest subspecies, with wing length closest to gerimantis and tail as in lecrayue. Mean crest length from pasterior base of] ¢ 71 and exposed crest Jength of 2 4d 54 (52- 57, SD 3.18). Chin, throat and upper breast dark olive brown, only slightly paler than side of head and forehead: breast medium cotlee brown. Range: Adelbert Mts, PNG. ya = Amblyornis macgregoriae lecroyae Frith & Frith, 1997 Asmall and particularly short-tailed form, sunilar to amet? in size. Tail/wing length averages fractionally shorter than lor the species. OFS 34, mean crest length from posterior base 64 (62-66, SD 1.64) and exposed crest length 76 (67-83, SD 6.64). Darker and more brown-orange than Lombok, Range: NNW slopes of Mt Bosavi, PNG. Amblyornis subalaris Sharpe, 1884 Streaked Bowerbird (Tables 8 and 22 Smallest, and relatively shortest winged, Amblvornis species. °? ¢ average fractionally smaller than ¢@. Mean crest length from posterior base oft 4d 50; the exposed crest Jength of 29 24 al (50- 72, 8D 5 79), and that of a crested: 2 30. The species is known to have BOWERBIRD BIOMETRICS 531 TABLE 9. Measurements (mm) and weights (g) of TABLE 10. Measurements (mm) and weights (g) of Amblyornis flavifrons. Wing Tail Tail Tarsus Total Bill Bill length length central length head length width length length Males: adult Mean 137 85 82 342 584 308 73 SD 2.65 1.15 3.46 0.65 1.41 5.78 064 Min 135 84 680 334 574 27.2 75 Max 140 86 86 35.1 594 375 86 n 3 3 3 3 2 3 3 hybridised at least once with Am. macgregoriae. Range: Mountains of extreme SE of NG, from upper Angabunga R to Mts Suckling, Simpson and Moiba, PNG (Schodde & McKean, 1973). Amblyornis flavifrons Rothschild, 1895 Golden-fronted Bowerbird (Tables 9 and 22) do wing length similar to Am. macgregoriae. Short-tailed relative to wing length, and long-winged relative to tarsus length, compared with Am. macgregoriae. Mean crest length from posterior base of 2 d d 54 (53-54, SD 0.71) and exposed crest length of 3 dd 94 (89-97, SD 4.16). No 2 collected. Range: Foya (or Gauttier) Mts of WP (Diamond, 1982). Prionodura newtoniana De Vis, 1883 Golden Bowerbird (Tables 10 and 22) Immature ( 2 -plumaged) d ¢ all-but identical in body measurements to adult 2 ¢ but, like adult 3 6, average slightly smaller in tarsus and bill lengths and notably more so in bill width and depth. The great disparity between d and @ tail length, and the relative proportions of these, reflects sexual selection upon 6d for arboreal/flight courtship display traits (Frith & Frith, 2000b,d). Other than this, 2@ are fractionally larger, averaging 15% heavier, than 3d but influences of sampling bias upon the latter are unknown. Range: Australian Wet Tropics uplands, from Thornton Ra and Mt Windsor Tableland to Seaview-Paluma Ra, Qld (Nix & Switzer, 1991). Sericulus aureus (Linnaeus, 1758) Masked Bowerbird (Tables 11 and 22) Both sexes average almost the same wing length. 2 2 average 5% and 7% larger in tail length and bill width but 7% and 6% smaller in Prionodura newtoniania. Wing Tail Tail Tarsus Total Bill Bill Bill Weight length {length central length head length width depth length length Males: adult Mean 122 110 95 303 509 23.1 6.0 7.0 73 sD 153 3.50 2.24 120 O74 054 029 O16 5.13 Min 119 88 85 231 49.0 220 54 67 62 Max 1260 «115 98 323 524 244 7.0 74 86 n 74 60 57 57 41 49 55 47 80 subadult Mean 119 87 84 299 507 23.3 66 6.9 71 sD 1.28 372 151 O89 030 101 046 062 267 Min 118 79 82 290 503 217 62 59 68 Max 122 91 86313 SIT 243 70 TA 76 n 8 8 7 5 5 5 4 5 7 immature Mean 119 68 851 309 500 23.1 64 7.18 75 sD 176 217 210 O67 062 O86 040 028 377 Min 115 84 76 292 488 213° 54 64 69 Max 124 99 90 321 SI5 242 74 8.1 82 n 56 55 52 41 14 42 43 33 42 Females: adult Mean 119 88 85 308 509 237 68 7.5 84 sD 254 191 194 O79 211 O70 052 022 836 Min m2 85 80 287 474 #223 56 7 62 Max 124 92 68 323 «#4564 251 78 #78 96 n 30 30 27 28 15 30 29 W 17 bill depth and weight than 3, respectively. With tail/wing length ratio of 60%, this bowerbird has the third shortest proportionate tail. Gilliard’s culmen “from base” should be compatible with our bill length, but his 23 for d and 24 for 2 ° are shorter and therefore probably of exposed culmen. Moreover, his 2 wing length of 137-140 is 2mm shorter than our shortest. Range: Mountains of western and northern NG almost to the Sepik R in the E. TABLE 11. Measurements (mm) and weights (g) of Sericulus aureus. Wing Tail Tail Tarsus Total Bill Bill Bill Weight lenath length centrale length head length width depth length Males: adult 143 84 82 41.4 61.2 30.6 7.5 8.5 178 Mean 3.37 372 355 123 198 161 034 026 3.54 sD 136 7 7 #397 «S74 259 68 79 175 Min 151 90 68 #441 648 328 83 88 180 Max 22 2! 15 18 18 22 22 W 2 n subadult Mean 147 87 86 42.3 62.9 31.5 75 86 71 sD 286 295 340 144 122 125 O52 O30 778 Min 142 81 81 403 598 295 6.6 8.2 165 Max 151 a1 92 448 #644 331 82 89 176 n W W 9 W n W W 4 2 immature Mean 139 90 89 420 608 32.0 75 149 sD 263 519 608 O73 265 179 068 gat Min 135 83 so 410 573 30.2 67 135 Max 141 94 94 426 642 342 6.3 156 n 4 4 3 4 6 4 4 4 Females: adult Mean 142 88 87 #419 618 315 6.0 TS 168 sD 207 481 604 1.41 100 108 068 O14 577 Min 139 62 79 401 604 298 68 78 165 Max 146 97 97 447 63.3 33.3 69 8.0 175 n W 10 7 W 10 10 W 2 3 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 12. Measurements (mm) and weights (g) of TABLE 14. Measurements (mm) and weights (g) of Sericulus ardens. Wing Tail Tail Tarsus Total Bill Bill length length central length head length width length Males: adult Mean 138 65 64 42.2 60.7 296 7.4 sD 412 3.02 333 264 128 047 0.49 Min 135 62 60 36.0 58.7 288 6.5 Max 148 72 70 446 62.0 303 8.1 n 93 | 8 g 6 8 8 subadult Mean 141 70 67 426 59.1 29.0 TA sD 299 310 379 1.72 0.70 040 058 Min 138 67 63 401 584 284 6.9 Max 145 74 70 43.7 598 29.3 8.2 n 4 4 3 4 3 4 4 Females: adult Mean 143 74 73 «414.8 #599 289 7.8 sD 495 1414 212 007 014 O57 0.07 Min 139 73 71 641.7 598 285 77 Max 146 75 74 418 600 293 7.8 n 2 2 2 2 2 2 2 Sericulus ardens (D’Albertis & Salvadori, 1879) Flame Bowerbird (Tables 12 and 22 Sericulus aureus has a tail longer than Se. ardens and averages a slightly longer (and pale) bill. This results in strikingly different tail/wing length ratios of 60% and 50%, respectively, the TABLE 13. Measurements (mm) and weights (g) of Sericulus bakeri. Wing Tail Tail Tarsus Total Bill Bill Weight length length centrals length head length width length Male: adult Mean 138 79 76 416 601 300 80 182 SD 130 235 265 063 102 058 O39 212 Min 136 7 73 «411 583 29.0 76 180 Max 139 83 80 428 607 304 85 183 n 5 5 5 5 5 5 5 z subadult Mean 143 85 83 425 606 306 85 178 SD O71 212 141 148 O07 O21 0.14 Min 142 83 82 414 60.7 304 8.4 Max 143 86 84 435 608 307 8.6 n 2 2 2 2 2 2 2 1 immature Mean 139 83 82 418 595 297 7.7 170 SD 495 566 7.07 049 O07 O78 049 Min 135 79 7 414 #594 291 73 Max 143 87 87 43.5 608 30.7 8.6 n 2 2 2 2 2 2 2 1 Females: adult Mean 139 85 84 405 606 31.5 86 173 SD 310 413 385 199 147 076 048 838 Min 133 80 80 387 583 306 79 = 164 Max 142 91 91 432 624 329 92 184 n 6 6 6 6 5 6 6 5 Sericulus chrysocephalus. Wing Tail Tail Tarsus Total Bill Bill Bill Weight length length centrals length head length width depth length Males: adult Mean 130 85 79 349 585 30.9 59 69 390 SD 256 219 282 126 090 O73 030 0.28 12.44 Min 125 82 74 325 566 293 5.5 64 76 Max 135 31 86 368 601 323 6.5 7.3 110 n 23 23 22 21 37 21 2 12 5 subadult Mean 134 92 85 346 564 306 5.9 7.2 86 sD 228 465 802 116 195 054 022 0.26 Min 131 87 7 334 53.6 30.0 5.6 6.9 Max 137 98 96 365 580 313 62 74 n 5 4 4 5 4 4 5 3 1 immature Mean 133 99 95 346 573 30.0 61 69 30 SD 268 258 310 102 108 O87 O24 O39 390 Min 129 94 91 329 557 268 5.9 6.3 85 Max 140 105 102 362 590 311 6.8 8.0 97 n 14 14 14 14 14 14 14 14 13 Females: adult Mean 138 106. 102 356 578 304 7.0 77 Wt sD 385 336 329 103 113 096 O37 O29 1213 Min 131 96 93° (340 S57 279 6.0 7 35 Max 148 Ww 107) 382 «597 321 75 82 134 n 26 26 ral 26 26 26 26 23 20 latter having a shorter tail than other bowerbirds (Table 21). Sexual size dimorphism is also quite different in the two species: 2° ardens average 14% and 5% larger in tail length and bill width but fractionally smaller in tarsus and bill lengths than 3 3 . Our bill width figures do not agree well with Lenz’s (1999: 53) statement that ardens has a “much narrower bill than aureus”. The bill width figures of Lenz (1999, table 3.8) do not support his statement. They do, however, indicate ¢ ardens have a slightly narrower bill than 3d aureus and that aureus has a deeper bill than ardens. In size differences between the sexes, and in body proportion ratios of the 3 Sericulus species in New Guinea, aureus differs more from ardens than it does from bakeri. Range: Patchy in lowlands-foothills from Wataikwa-Mimika, upper Noord-Endrich R of WP to upper Fly R, Strickland-Nomad R and Mt Bosavi, PNG (Mackay, 1984). Se. a. aureus and Se. a. ardens have different 2 plumages and bill colouration (Cooper & Forshaw, 1977; Beehler et al., 1986). They are all but allopatric, in being isolated by the central ranges, and are restricted to predominantly different altitudes, but do meet and hybridise on the Wataikwa R, south WP as indicated by | or 2 36 specimens (Gilliard, 1969). Thus, aureus and ardens are considered separate species (Lenz, 1999), TABLE 15. Measurements (mm) and weights (g) of Ptilonorhynchus violaceus. BOWERBIRD BIOMETRICS Wing Tail Tail Tarsus Total Bill Bill Weight length length central length head length width length length P. ¥. olaceus Males: adult Mean 172 «110 «©6103 493 666 354 98 238 SD 3.68 5.20 545 198 138 170 O34 195 Min 162 103 9 443 637 315 90 208 Max 182 125 14° 530 694 403 104 278 n 66 46 27 47 30 46 46 13 subadult Mean 1730 («1150115 48.3 66.5 357 97 205 SD 3.77 492 2.37 151 067 0.46 Min 168 109 453 648 353 94 Max WF (lad 50.2 677 365 10.2 n 4 4 i; 4 3 3 3 1 intermediate-plumaged immature Mean 177i 119-114 49.3 66.1 349 10.1 227 SD 3.98 534 435 238 152 106 029 2597 Min 161 106 107 43.7 617 326 95 176 Max 178 131 125 53.7 677 375 106 284 n 26 26 18 26 17 26 26 14 female-plumaged immature Mean 169 118 «6 111 498 653 343 99 225 SD 3.90 455 550 184 153 1.08 034 6879 Min 162 107) 103 46.7 620 325 92 208 Max 178 #126 120 542 678 359 104 234 n 21 20 14 21 18 21 21 12 Females: adult Mean 165 117) «113 46.1 641 346 102 209 sD 474 467 472 215 150 150 033 18.80 Min 156 108 102 415 601 31.0 96 170 Max 176 128 123 516 672 382 109 250 n 60 59 36 60 43 59 58 30 Pv minor Males: adult Mean 153 96 89 473 639 33.2 9.3 187 sD 2.74 280 234 184 120 086 0.81 16.70 Min 150 92 87 443 620 32.0 84 173 Max 158 100 93 496 651 344 113 205 n 10 10 ¢ 10 7 3 10 4 subadult Mean 156 = 102 97 479 645 334 98 206 n 1 1 1 1 1 1 1 1 intermediate-plumaged immature Mean 153-101 97 459 624 323 968 193 sD 485 321 325 235 129 076 0.37 23.08 Min 144 933 90 414 605 309 9.0 164 Max 165 110 104 489 642 334 103 232 n 18 18 17 18 7 18 18 18 female-plumaged immature Mean 152 102 98 474 623 333 88 197.8 sD 473° 3.79 2.31 1.18 0.75 0.81 15.20 Min 148 99 95 467 32.6 79 «187 Max 157 106 99 48.8 341 95 209 n 3 3 3 3 1 3 3 2 Females: adult Mean 150 102 95 450 615 317 9.5 174 SD 465 332 330 167 021 O50 0.32 Min 145 98 91 #436 613 313 92 Max 156 106 99 470 617 324 99 n 4 4 4 4 3 4 4 1 All subspecies Males: adult Mean 163 108 100 489 66.1 35.0 97 226 SD 735 746 763 208 %1.70 1.78 0.49 2865 Min 150 92 87 443 620 315 B4 173 Max 1820«125°0C—i114 «533.0 69.4 «6403 «113 ©«278 n 76 56 34 57 37 55 56 17 subadult Mean 169 «#113 106 48.2 66 35.2 97 206 sD 817 7.30 1273 206 159 129 038 O71 Min 156 = 102 97 453 645 334 94 205 Max 177 121 15 502 67.7 365 10.2 206 5 5 2 5 4 ee Se J 533 intermediate-plumaged immature Mean 164 it 106 47.9 643 33.8 100 208 SD 10.22 988 942 288 232 159 035 29.57 Min 144 93 90 414 605 309 3.0 164 Max 178 131 125 53.7 677 375 106 284 n a4 44 35 44 34 44 44 32 female-plumaged immature Mean 167 116 109 495 65.1 34.2 9.8 221.1 SD 7.07 725 7.36 193 1.64 109 055 13.44 Min 148 39 95 467 62 32.5 79 187 Max 178 126 120 542 678 359 104 234 n 24 23 i7 24 19 24 24 14 Females: adult Mean 164 116 111 460 639 344 102 208 SD 6.1 6.0 7.0 21 1.6 1.6 04 19.5 Min 145 98 91 415 60.1 31.0 92 170 Max 176 128 123 516 672 382 109 250 n 64 63 40 54 46 63 62 31 Sericulus bakeri (Chapin, 1929) Adelbert Bowerbird (Tables 13 and 22) Its tail/wing length ratio is 59%, the second shortest for the family. 2 2 average 8%, 5% and 8% larger than 6 ¢ in tail and bill length and bill width, respectively, but 5% lighter in weight. In most ¢/¢ ratios, as well as tail/wing length for both sexes combined, this (dark-billed) species is more like dark-billed Se. aureus than pale billed Se. ardens. Gilliard (1969) gave the short bill length “from base” of 23 for dd and of 24 for 2° (cf. Table 13) but these are probably of exposed culmen. Range: Adelbert Ra, PNG. Sericulus chrysocephalus (Lewin, 1808) Regent Bowerbird (Tables 14 and 22) Tail/wing length ratio 71%. 2 ¢ average 6%, 24%, 19% and 12% larger than d d in wing and tail lengths and bill width and depth, respectively and average 23% heavier. With increasing age, 3d wings and tails become relatively shorter and their mean central tail feather length gets shorter relative to tail length. As a result, the tail/wing length ratio in ¢ d is 66% but in & 2 is 77%. Lenz (1999:38) gave bill length of 26.0 for 36 and 25.6 for 22 . We concur with Hartert (1929) and Schodde & Mason (1999) in considering the doubtful northern Se. c. rothschildi, Mathews, 1912 invalid. Range: From N of Sydney to Connors and Clarke Ra, inland of Mackay with a gap at the Fitzroy R valley inland of Rockhampton, Australia. Ptilonorhynchus violaceus (Vicillot, 1816) Satin Bowerbird (Tables 15, 16 and 22) Tail/wing length ratio 67%, tarsus/wing length 28%, and bill/wing length 21%. 2 ° average 6% Wn re) rs TABLE 16. Mean measurements of Ptilonorhynchus v. violaceus, from N to § of its range in latitudinally bounded subdivisions. Degress of latitude S. Wing Tail Tail/wing Sexage group length (n, SD) length (n. SD) ratio 26-30 Males: adult 170 (20, 3.07) 107 (20. 4.05) 0.63 immature " 169 (12. 4.40) 114 (12. 4,77) 0.67 Females: adult 164 (9, 4.55) 114 (9, 3.71) 0,7 All birds 169 (41, 4.40) 114 (41. 5.42) 0.66 30 - 32 Males: adult 172 (22, 3.21) 110 (3, 3.21) 0.64 immature 168 (1) ~- - Females: adult 166 (4, 7,14) 117 (4, 5.85) 07 All birds 171 (27. 4.30) 114 (7, 6.08) 067 32-34 Males: adult 174 (6. 6.63) 113 (6. 4.72) 0.65 immature * 172 (16, 3,40) 120 (16, 3.82) 07 Females: adult 163 (17, 3.17) 115 (16. 3.66) 0.71 All birds 188 (39. 6.17) 117 (38, 4,78) 0.7 34 - 36 Males: adult 171 (2, 0.71) 112 (2, 3.54) 0.65 immature 169 (8, 4,34) 117 (8. 4.50) 0.69 Females: adult 166 (2, 7.07) 116 (2, 5.66) 07 All birds 169 (12. 4.30) 116 (12, 4.54) 0.69 36 - 39 Males: adult 174 (10, 4,56) 115 (9, 5.52) 0.66 immature ~ 171 (13, 3.69) 121 (13, 4.64) O71 Females: adult 167 (23, 4.82) 120 (23, 3.97) 0.72 All birds 170 (46..5.14) 119 (45. 4.85) 07 “Samples for 26-30 degrees include two subadullts (i.e. immature showing some signs of adult plumage) and for 32-34 and 36-39 degrees one subadult each and 8% smaller than 34 in tarsus length and weight, but 7% and 5% larger in tail length and bill width. Thus dd tails become relatively shorter with increasing age. Mean bill length averages 2% shorter in ° 2 than 3d d, contrary to comparison of the exposed culmen (Schodde & Mason, 1999). Males in their first to third year have ° 2-like plumage, but with whiter underparts. [n their fourth to sixth year, or prior to attaining subadult characters, their plumage becomes indicative of that of ¢ 3 by increasingly solid green breast and throat plumage (Disney, 1970). o lacking adult or subadult plumage are *©-plumaged immature 6’. Birds with any sign of the more solid green breast/throat are ‘intermediate-plumaged immature 3 3’, Ptilonorhynchus violaceus violaceus (Vieillot, 1816) Based on the surgical sexing of 19 birds, and 3 on plumage, caught and measured at Healesville, MEMOIRS OF THE QUEENSLAND MUSEUM Victoria, 1t was concluded that birds with a tarsus length >57.5 and wing length>161 are 3d and less are 22 (Miller, 1995). Tarsus length measurements of Miller are “tarsus with foot length” of Lowe (1989), thus his are 18% larger than ours. One ¢ of our sample had a wing length of 161, and a tarsus length of 48.9. While our 60 ? tarsus lengths are shorter than 58, and thus agree with Miller (1995), 45 9 ¢ had a wing length of 162 or more. Range: coastal Australia in a <250 km band, narrowing to the N, from the Otway Ra W of Melbourne to Dawes Ra, Qld. Schodde & Mason (1999) described a ‘step’ in size and tail proportions of the adult (I. Mason pers. comm.) populations in the centre of the range. They wrote that “Within the southern form, there is evidence of incipient divergence at the Hunter River (- Sydney) Barrier, NSW” and go on to observe that “Populations of nominotypical violaceus north of the Hunter to northern limits in the Dawes Range (Kroombit plateau), Qld, are relatively small with short tails (wing: o¢ ¢ 167-173, 2c 157-165 mm; tail/wing ratio: d 3d c 0.58-0.64, 2 2c 0.66-0.70; n= 76). Those § of the Illawarra (Wollongong- Nowra) region are all larger and longer tailed (wing: od ¢ 170-178, 2 Pe 163-170 mm; tail/wing ratio: d dc 0,63-0.67, ° 2 c 0.68-0.72; n = 43). In between, from the Hunter to the Illawarra, populations appear to grade from one size morph to the other (n = 65).” Schodde & Mason (1999) claimed that birds N of the Hunter River (or of 32°) are “relatively small with short tails”. While our data support this in broadest terms (Table 16) the differences are at best slight, and are less so in adult 22 than in all dd. Schodde & Mason also found birds S of 34° larger and longer tailed, but we find this true of only adult 2 wing length, as there is little difference in tail/wing length ratio (Table 16). Mean values for ¢ and ° wing length and tail length are compared using Student’s two-tailed t-test between more pertinent pairs of the five geographical samples (Table 16). Results are as follows: there is no significance in d or in & wing length between the sample pairs of 30-32° and 32-34°, 32-34° and 34-36°, 30-32° and 34-36° S (all being P > 0.1). There is also no significance in d and tail length between these same sample pairs (P > 0.1). Wing length between 26-30° and 36-39° S were significant (P <0).02), but 2 wing length between them was not (P > 0.1). Differences in ¢ and in @ tail length between these extremes are, however, more significant (P< 0.001). The latter differences are BOWERBIRD BIOMETRICS TABLE I7. Measurements (mm) and weights (g) of Chlannydera maculata. Wing Tail Tarsus Total Bil) Bil fength jength length head jenath wath tength Weight Males aduft Mean 151 409 407 597 W5 6.0 133 SD B29 412 156 106 147 O46 6.25 Min 145 io 6376 «S77 2B6 70 125 Max 157 17) «485 6B A 389 fo 150 n 30 30 30 2) 23 30 13 subadutt Mean 150 2 404 586 U7 al 144 sp 26 42 4 10 4 05 155 Min 145 104 376 S69 296 7H 127 Max 154120 2S OD 84H 8B oi] 22 22 22 aI 22 22 12 immature Mean 82 7 405 Sh2 arg TS 7 so 2.63 148 26 113 Min 150 394 2265 8687 Max 154 4s a3 63 7 2 2 2 ' 2 2 t Females adull Mean 146 10 369 S85 B44 82 41 50 344 410 190 147 447 O39 1171 Mire 141 103 «36.7 554 294 77 124 Mar 156 117 408 616 354 89 I62 n 30 30 2g 26 29 30 13 not surprising, however, given they are between populations at each end of a distribution. As itis possible that small diffcrences apparent in some sizes for sex/uge classes in Table |6 are due to small samples (e.g. for adult od of the 32-34° and 34-36°_ and adult 4 2 ol the 30-32° and 34-36") it is worth comparing figures for all birds combined for each zone, These show little difference in wing length for the subspecies, with notevena discernable N-S cline. Our data do not indicate a cline, or step, is in tail/wing length ratio with latitude. Tails are slightly longer in the $ thanin the N, but even between the N-and S-most vones the difference is minimal (4%) and clinal, with no obvious step (Table 16). Ptilonorhynchus violaceus minor A.J. Campbell, 1912 All measurements are about 10% less than in violaceus Violaceus but body proportions are similar Plumages are similar but the green of plumage has a dull bluish-grey cast (Schodde & Mason, 1999). Range: Australian Wet Tropics. from Mi Amas & of Cooktown toa Seaview- Paluma Ra, Qld. Table 15 shows full overlap in wing and tarsus lengths of 2 @within the ¢¢ range. Our bill length data (21% of wing length in violaceus minor and 20% in violaceus violaceus) do no ye \pa at support the observation that the bill of vielacens vialaceus is “long and thick” (Schodde & Mason, 1999). This diflerence trom Schodde & Mason (1999) may be due to method of measurement. Their measurement can be misleading, particularly in dimorphic species in which only one sex has modified and/or longer plumage al the base of the upper mandible (Frith & Frith, 1997e: 173). Adult d 4 show atinurto have a bill width 5% narrower (han vie/aceus violaceus. and adult 2 minor have a bill width 7% narrower than violaceus violaceus. For the sexes combined bill widih inuninoer is 9.6 and in violaceus violaceus 9.9 which, viven the difference in their overall sizes, is not striking, The bill of wiolacens violaceus is not proportionately thicker than that of minor. In view of this, we assume Schodde & Mason (1999) meant that the bill of violaceus violaceus is long and deep (not thick) and thus “appears slumpier in shape” bul, like us. they provide no bill depth measurements. Bill width/length is the same in 3 4 (28%) and 2 2 (30%) of both subspecies, Chlamydera maculata (Gould. 1837) Spotted Bowerbird (Tables | 7 and 22) Gilhard treated wullate as a subspecies of C. maculata, bul others consider tla distinct species (Frith & Frith, 1997a; Christidis & Boles, 1994: Schodde & Mason, 1999), The tail/wing length ratio is 73%, tarsus/wing length 27%, and bill/wing length 21%. °° are fractionally smaller than 2d, but fractionally larger in bill width and in weight. The species is thus less sexually dimorphic in size than congeners, while being similar in major proportions (Tables 17-21) Immature 4 lack crest feathers, while subadult dé haye some to many, Range: Interior of Old § of 20°S, exceplextreme W and SW, and interior of nomhem and central New South Wales (NSW) (except extreme W border country). Alsa extreme NW corner of Victoria and just into South Australia (SA), on the Murtay R, Once confirmed to have hybridised where tt meets C. nuchalis some 100 km SSE of Charters Towers, Qld (Frith & Frith, 1995b), Chiamydera guttata Gould, 1862 Western Bowerbird (Tables 18 and 22 2 average fractionally smaller than dé in wing and tarsus lengths, but 5% larger in tail length and bill width. the sexes being the same in total head and bill lengths. Tail/wing length ratio nt us ° TABLE 18, Measurements (mm) and weights (g) of Chlamvdera guitaia. Wing Tail Tarsus Total Bill Bill ~=Weight length length length head length width length Cg guitala Males: adult Mean 149 $2 389 579 304 74 134 sD 247 330 103 095 O99 O31 517 Min 144 86 361 557 281 68 128 Max 157 100 «416 #593 326 72 \42 " “4 33 “4 3 a 4 8 subadult Mean 148 97 #391 S78 308 WW 133 f=18] 147 449 152 O7F 034 034 495 Min 146 90 376 566 202 67 129 Max 150 102 41.9 566 312 77 136 n 6 6 6 8 6 6 2 Females: adult Mean 147 $8 38.1 562 302 tT 137 5D 236 270 121 124 417 O36 820 Min 143 94 357 #561 277 69 122 Max 152 104 4140 599 319 84 148 n 21 21 21 17 20 19 g C g carter Males-adult mean 135 93 37.3 267 T n ' i} i} i} 1 Females: adult Mean 137 31 #365 57.) 30.5 77 sD. 288 333 057 O81 O38 Min 135 87 «355 297 TA Max 141 95 372 32.0 B2 n 6 6 6 ! 5 6 All subspecies Males: adult Mean 148 82 389 579 304 73 134 SD 336 326 #105 ag5 [02 O81 SIT Min 135 86 361 557 281 68 128 Max 157 100 416 593 326 7a 142 n 36 34 35 3) 36 36 8 subadult Mean 148 97 391 578 308 71 133. SD 147 449 152 O71 O34 O34 495 Min 146 90 376 566 302 67 129 Max 150 102 449 S86 312 77 138 n 6 6 6 6 6 6 2 Females: adult Mean 145 37 378 581 303 7.7 137 5D 490 430 %!3/ ¢t23 109 O36 820 Min 135 87 355 561 277 69 122 Max 152 194 410 S589 320 a4 148 " 27 27 27 16 26 25 a 65%, tarsus/wing length 26%, and bill/wing length 21%, the first of these confirming the short tail (8% shorter than C. mereidlare). This Australian taxon was treated as a separate species until Mathews (1912) considered it a subspecies of C. maculata. We agree with Schodde (1982) that guftafa is a separate species, For diagnosis see Schodde & Mason (1999), MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 19, Measurements (mm) and weights (g) of Chlamydera nuchalis. Wing Tail Tarsus Total Bill Bil =~Werght length fength length head length andth length C2, nuchals Mates adult Mean 182 144° 4491 #690 357 30 216 so $73 683 190 154 1.30 055 27,94 Min 171 133 456 656 360 73 160 Max 193 157 546 7IB 408 106 265 " 35 36 36 26 m4 35 3 subadult Mean iso (144 «484 695 391 390 222 sD 480 593 (38 %1§02 {28 O45 12.36 Min 170 «(1350 «AA 681 GB 7D 201 Max eS 1S7 498 «720 410 35 280 n 7 17 17 12 16 7 Q immature Meary 79 «15) 492 689 390 85 213 sD 783 75) 218 212 153 O49 1730 Min 163 #138 444 652 365 74 195 Mas 196 '65 S40 732 416 93 242 n 3t wu a 18 28 30 10 Females: adult Mean 73 «(1430 «(4570 «665 «37506 «6B OB 8D 689 749 180 148 155 O47 1415 Min 158 127 423 BAZ 322 76 164 Mas 191 160 505 697 415 98 215 " 54 50 st 35 50 50 17 Cn, onentatis Males: adult Mean 175 #13) 4269 685 384 89 210 sD a49 450 2039 148 #%1(69 O49 1353 Min 167 122 437 554 345 82 \87 Max 82 139 526 716 Ald 101 236 n 26 26 26 15 26 26 10 subadult Mean 77 134 474 G76 378 BB 240 sD 521 508 |54 156 178 O55 5646 Min 168 «124 «44606 «659 6352067 Max 185 145° «502 703 417 OF OMT 15 15 15 6 15 14 5 immature Mean 16g «134 46.1 668 373 BB 200 SD 698 736 192 122 154 O51 10.67 Min 156 «1211S 350 666188 Max 166 «149«s50.7) «6681 399 «9B 23 fn 32 32 32 " 32 32 5 Females. adult Mean 167 131 4368 656 3659 67 1686 sD 555 526 173 174 #%(1I47 053 15.58 Min 65 122 407 629 326 77 153 Max vw 142 469 B76 386 a9 205 0 x 33. 33 4 32 32 13 All subspeces. Males, adult Mean 79 «1138 «6482 6BB 386 30 212 so 608 891 224 152 147 O53 2hI7T Min 1670122, 43) 54 OS OTS (BO Max 133 157 S46 718 414 106 265 n =) 6 81 4) 60 a1 19 subadult Mean 178 «#139 «479 «688 385 8S 228 so 510 726 1.51 150 165 049 34.04 Min 168 «6124 «4430 «(65.9 «352 79 «1a7 Max {a9 «1570 S020 720 «641700 87 OH? ” 32 32 32 18 3 31 4 immature Mean 174 «#142 476 681 361 86 209 sD Ber 1109 256 206 175 O50 16.36 Min 158 121 415 $635 350 74 168 Max 196 «165 «4580 732 «416 9B 242 n 63 63 is] 29 50 62 15 Females: adult Mean \7o 138 449 666 374 87 187 5D 586 B71 20) 165 157 O48 1456 Min 158 122 807 6829 322 7.6 153 Mas 19) 160 50S 697 415 99 215 n a4 aa 84 49 82 62 1) BOWERBIRD BIOMETRICS TABLE 20, Measurements (mm) and weights (g) of Chlamydera cerviniventris, Wino Tail Tal Tarsus Total Bill fu = Weight feneyth ength central tenath fread engi witty lene fenath Austraha Males, adult Mean 140 ie 4°94 614 306 76 175 sD S30 336 0,85 48 1,08 o44 479 Min 14) 105 34 Savy 264 be 70 Max 154 WE 429° 628 32) BY ae al 24 4 24 5 22 23 7 Femates, adult Mean \a4 We 104 39) B04 305 a4 sv 50 3960 (57! 142 1\O2 103 Oar 978 Min sa 0 365 594 BAA 76 144 Mae 180 8 ats ole 7 a5 187 n 8 17 ' 16 a 17 15 5 New Guiness Males: adult Meaty $49 mW 106 «410 Sh 306 Bz 159 SDB a5 t59 aor 145 5,61 115 14 1066 Min ica) ww? oo 3h4 0 356 BB 74 145 Man 157 WwW? 8 432 642 aa? gt 180 n a7 5 6 a5 a2 36 OT a7 Females: adult Mean 146 WH 96 «983 S05 309 Ba jag so 402 297 248 442 144 196 DRA 13.85 Min ag 195 4 M7 S72 380 7a V7 Mae 155 19 iio 422 85 Mg 105 170 n ae ue 4 3B 23 x) 33 16 All turds Males: adult Mean 48 62 «6106 «412 60) 3207 Bo Me so a6) 299 207 (25 520 '12 DSO 1146 Mir 14) 105 104 “S44 396 781 fa 45 Max sv 118 wS 439 baa 397 a1 182, " 5 60 6 5p x 58 oa a4 Ferrales. adtull Mean 145 Wi 106 302 605 206 a2 151 sp 4i0 322 240 ral rs 10 «058 1219 Nant 138 «(04 104 347 BT ee 70 WW Nave 156 ng 110 42 68:5 349 ws rs) Chlamydera guttata guttata Gould, 1862 The larger subspecies, blacker dorsally and with a longer and broader nuchal crest. d d have crests and 2 S vary from no crest to 4 complete one. Range: Western Australia (WA), from the base of North West Cape through the Pilbara and into the central interior, Thence FE in a narrowing band, across the Northern Territory: (NT) border into the S of NT to c. 200-300 km N and E of Alice Springs, and extending 100 km S of the NT/SA border and E to due S$ of Alice Springs. Chlamydera guttata carteri Mathews, 1920 Averages 8% smaller in wing length and 5% in tarsus length than the nearest populations of gnitata guttuta (Hammersley R, Onslow & Sherlock R areas) but only fractionally so in tail and bill lengths. Thus carter: has a long (and broad) bill, in addition to being richer and more russet coloured than guittuta guitatu (Frith & Frith, 1997a), Five of six 2 2 examined have a full ¢-hke crest, Range: Restricted to North West Cape, WA. Chlamydera nuchalis (Jardine & Selby, 1830) Great Bowerbird (Tables 19 and 22) Adults have iail/wing length ratio 79%, tarsus/wWing length 27%, bill/wing Jength 22%. 22 average 5%, 7% and 12% smaller than 2 4 in Wing and tarsus lengths and weight, respectively. Alladull ¢ ¢ have a full crest. and the 17 subaduli do have between 10 crest feathers to three-quarters of'a full 3 -like crest, OF 84 2 2 none havea full d-like crest while 4 have only 4-10 crest feathers, 2 a quarter-developed crest, 3 a third-, | a half-, and | a three-quarter developed crest (Frith & Frith, 1999), Chlamydera nuchalis nuchalis (Jardine & Selby, 1830) Chiamydera nuchalis aweni Mathews, 1912, The larger subspecies, ? & ayerage 5% smaller than dd in wing length, 7% smaller in tarsus length and 14% lighter, Mean ratios for sexes combined are: tail/wing length &1%, tarsus/wing, 27%, bill/wing 22%. We agree with Mayr & Jennings (1952), Gilliard (1969) and Hall (1974) that this subspecies has paler and less contrasting, more uniform and greyish, upperparts and underparts usually darker than orfentalis. immatures exhibit shahtand faint ventral (mostly flank) barring (stronger in some more W individuals), but adults less so ornetat all. Adults typically lack silky silyery-white feather tipping, or spotting, throughout the crown (and when present only aboutthe pink nuchalcrest feathers). Range: Kimberley, NT. extreme NW Qld, islands off N coast, to no further 8 than 20° S. Chlamydera nuchahis orientalis Gould, 1879 Chlampdera michealis york Mayr & Jennys, 1952. The smaller subspecies, averaging 9% smaller in tail length, and fractionally so in tarsus and bill lengths and bill width than nuchalis nuchalis. Similar to nuechalis nuchalis in 2 size as proportion of. Mean combined sexes tarsus/wing length and bill/wing length ratios the same as nuchalis muchalis, but tail/wing 77% (Frith & Frith, 1999). Dorsal colouration and markings (Mayr & Jennings, 1952) are darker and more variegated or contrasting in pattern. Ventral 538 TABLE 21. Measurements (mm) and weights (g) of Chlamvdera lauterbachi. Wing Tail Tail Tarsus Total Bill Bill Weight length length central length head length width length length C Lf fauterbachi Males: adult Mean 133 104 39.7 57.0 27.5 69 135 SD 1.41 2.83 0.21 1.70 0 Min 132-102 39.5 26.3 6.9 Max 134 106 39.8 28.7 69 n 2 2 2 1 2 2 1 Cf uniformis Males: adult lean i34 = 106 $9 386 56.3 28.1 639 i30 sD 267 3.39 1,59 1.51 087 029 3.89 Min 130 =—:100 348 526 263 64 128 Max 139 We 414 578 29.6 7TH 133 n 21 20 1 22 10 22 2i 2 Females: adult Mean 131 106 102 375 566 282 7.2 W5 sD 2862 425 1.73 147 129 069 034 416 Min 125 99 100 353 539 266 6.2 112 Max 137 116 103 «44.1 582 295 78 120 n 26 26 3 26 12 26 26 3 All subspecies Males: adult Mean 134 ©6106 99 387 563 280 69 132 sD 258 3.34 155 145 092 027 3.88 Min 130 = 100 34.8 526 263 6.4 128 Max 139 112 414 578 296 7.5 135 n 23 22 i 24 W 24 23 3 Females: adult Mean 131 106 102 37.5 56.6 262 7.20 «115 SD 282 425 173 147 1.29 0.69 4.16 Min 125 99 «©6100 353 539 26.6 6.2 112 Max 137 N16 103 41.1 582 295 78 §=120 n 26 26 3 26 12 26 26 3 plumage, particularly flanks, usually strongly barred. Birds in the NE (formerly yorki) average slightly smaller and paler, more so ventrally and particularly on the throat, than those to the S (Frith & Frith, 1999). Range: Qld N of 20° S, a little further S in the E. Absent on Cardwell- Tully coast and E watershed of the Wet Tropics ranges. Locally sympatric with C. cerviniventris on NE Cape York Peninsula, but hybrids unknown. Gilliard (1969) followed Mayr & Jennings (1952) in accepting 4 subspecies. Now oweni is considered a synonym of nuchalis and yorki is synonymous with orientalis (Frith & Frith, 1999; Schodde & Mason, 1999). Colouration and pattern of the crown are important discriminators in Ailuroedus, Amblyornis and Chlamydera (Gilliard, 1969; Frith & Frith, 1995b, 1997a,b, 1998). The marked difference in crown and dorsal plumage colouration and pattern separate nuchalis, in WA and the NT east to the Gulf of Carpentaria in W Qld from orientalis inthe E and MEMOIRS OF THE QUEENSLAND MUSEUM NE of Australia (Storr, 1967, 1973, 1977, 1980, 1984; Ford, 1974, 1987; Schodde, 1986). See C. maculata (above) for details of hybridisation. Chlamydera cerviniventris Gould, 1850 Fawn-breasted Bowerbird (Tables 20 and 22) There is negligible difference between NG and Qld populations (Table 20). Tail/wing length ratio is 76%, tarsus/wing length 27%, and bill/wing length 21%. 2 2 average 5% smaller in tarsus length and are 7% lighter than ¢ 3. Range: PNG coast to 500 m asl, to Humboldt Bay in N and Marauke in the S of WP. Jimi Valley, PNG at 1700 m (Frith, 1987). Locally sympatric with Lauterbach’s but hybrids unknown. Isolated populations in Ransiki and Kebor Valleys of E Vogelkop; along eastern coastal zone of Cape York Peninsula to Coen, Qld where locally sympatric with Great Bowerbird but hybrids unknown. Chlamydera lauterbachi Reichenow, 1897 Lauterbach’s Bowerbird (Tables 21 and 22) Tail/wing ratio of 80%, tarsus/wing 29%, and bill/Awing 21%; 2 2 13% lighter than dd. Chlamydera lauterbachi lauterbachi Reichenow, 1897 Known from two specimens, with coppery coloured crown and cheek feathering having a Sericulus-like metallic rose-golden sheen to it. Range: Aiome area of upper Ramu R, PNG, possibly also near Bogadjim, Finisterre Mts, PNG (Gilliard & LeCroy, 1968). Chlamydera lauterbachi uniformis Rothschild, 1931 No differences between this and lauterbachi lauterbachi in average sizes or body proportions. Crown yellowish olive-green (50). Range: NG lowlands tol 800m asl, from Geelvink Bay, WP to Okapa-Aiyura area of E Highlands, PNG. Patchy in Snow and Star Mts to upper Ramu R. Sympatric with C. cerviniventris SE of Hupai, Ramu Valley, and in Aiyura Valley (Bailey, 1992; Doyle et al., 1981), but hybrids unknown. CONCLUSIONS It has been stated that sexual size dimorphism, expressed by the 6/2 wing length, is no greater in polygynous bowerbirds than in monogamous Ailuroedus species (Payne, 1984). Polygynous Se. dentirostris is, however, less sexually BOWERBIRD BIOMETRICS nh 1e9) ‘© TABLE 22. Mean size and some mean body proportions of adult 2 vs ¢ bowerbirds, expressed as % & are smaller than d or as a % of mean size the former trait is of the latter. Values of less than 5% are not indicated. Tables 1-21 give sample sizes. MWL = mean wing length; MTL = mean tail length; MLL = mean tarsus length; MBL = mean bill length; MBW = mean bill width; MBD = mean bill depth; MW = mean weight. Species Al buecordes Al crassirostis Al melanotis Sc. dentirostis Ar. papuensis Am. inomatus Am. maogregorae Am. subalaris Am. flavitrons Pr. newtoniana Se, aureus Se. ardens Se. bakers Se. chrysocephalus Pt. wolaceus C maculata C guttata C nuchalis C cermniventis C lauterbach/ Mean of mean ratios dimorphic in size than Ailuroedus (Tables 1-4, 22). Our wing length data agree with Payne (1984) in demonstrating Ar. papuensis to be the most sexually dimorphic bowerbird in this trait. They also agree with Lenz (1999) in showing Se. chrysocephalus to be the second most sexually dimorphic (but reversed, in that 2 2 are larger than 3 3) and not the least dimorphic bowerbird (pace Payne, 1984). In all polygynous bowerbird genera except Sericulus 2 2 are typically shorter in wing and tarsus lengths than dd. In all bill dimensions 22 Ailuroedus average slightly smaller than dd, whereas in the majority of polygynous species ¢ 2 have bills the same size or larger than d d. We consider the morphology and geographical isolation of Al crassirostris supportive of its distinct status from A/. melanotis, this being in harmony with the separation of Se. ardens and C. guttata. MTUMWL (male, female) =MLL/MWL MBL/MWL 66 (67. 66) 28 22 74 (74, 74) 28 20 73 (73, 73) 29 23 70 (70, 69) al a1 96 (103, 88) 26 21 70 (71, 70) 27 23 65 (65. 65) 27 21 70 (70. 70) 28 24 62 (62, -) 25 22 82 (90. 74) 25 19 60 (59. 62) 29 22 50 (47. 52) 30 a 59 (57. 61) 30 22 71 (66, 77) 26 23 67 (64. 71) 28 21 73 (72.74) 27 21 65 (62. 67) 26 21 79 (77, 81) 27 22 76 (76. 77) 27 24 80 (79. 81) 29 21 ACKNOWLEDGEMENTS For access to collections in their care we thank the staff, too numerous to mention, of the mainland Australia State Museums and the following institutions: Australian National Wildlife Collection, CSIRO, Canberra; Australian Bird and Bat Banding Scheme, Canberra; American Museum of Natural History; The Natural History Museum, Tring; National Museums and Galleries on Merseyside, Liverpool; Museum of Vertebrate Zoology, University of California, Berkeley; The Field Museum, Chicago; Museum of Comparative Zoology, Cambridge; Bishop Museum, Honolulu; Royal Ontario Museum; Academy of Natural Sciences, Philadelphia; Carnegie Museum of Natural History, Pittsburgh; Peabody Museum, Yale University; Delaware Museum of Natural History, Wilmington; Museum Zoologicum Bogoriense, Bogor; National SA() Museum and Art Gallery of Papua New Guinea, Port Moresby; Museo Nacional de Ciencias Naturales. Madrid: Museo Civico, Genova; Nationaal Natuurhistorisch Museum, Leiden; Swedish Museum of Natural History, Stockholm; Zoologisk Museum Kebenhavns Universitet, Kobenhaven: Museum National d?Histoire Naturelle, Paris; Staatliches Museum fiir Trerkunde, Dresden: Zoologische Staats- sammlung, Minchen, Zoologisch Museum, Berlin; Museum Alexander Koemg, Bonn: Staatliches Museum fiir Naturkunde, Stuttgart: Forschungsinstitut und Naturmuseum senckenberg, Frankfurt; Zoologisches Institut und Zoologishes Museum, Hambure. People who kindly provided help in other ways include Joan Airey, Brian and Del Coates, JefY and Barbara Dayies, Paul and Luisa Frith, Peier and Daphne Fullagar, Andrew and Trish Gillison and Peter and Janet Marsack. Sincere thanks to the Chapman Fund Committe for support, We thank fan McAllan, Walter Boles, Norbert Lenz, Wayne Longmore, and an anonymous referee, Opinions expressed are ours alone. 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(Am- erican Museum of Natural History: New York). STORR GM. 1967. List of Northern Territory birds. Special Publication of the West Australian Museum 4: 1-90. 1973. List of Queensland birds. Special Publication of the West Australian Museum 5; 1-177. 1977. Birds of the Northern Territory. West Aust- ralian Museum Special Publication 7: 1-130. 1980. Birds of the Kimberley Division, Western Australia. West Australian Museum Special Publication 11: 1-71. 1984. Revised list of Queensland birds. Records of the West Australian Museum, Supplement 19: 1-189, UY, J.A.C. & BORGIA, G 2000. Sexual selection drives rapid diverence in bowerbird display traits. Evolution 54: 273-278. WESTERN PACIFIC TINGIDAE (HETEROPTERA): NEW SPECIES AND NEW RECORDS ERIC GUILBERT Guilbert, F. 2001 06 30; Western Pacifie ‘lingidae (Heteroptera); new species and new records. Memoirs of the Queensland Museum 46(2)-543-554. Brisbane. ISSNOQ079-8835. Nine new species are deseribed from the Western Pacific, namely, Crsveochila kraussi, Leptoptyx varians, L. longispina, Nesoeypselas simples, N- straphii from Solomon Islands, Eleoneus samoaensis from American Samoa, &. palauens/s froin Palau Islands, Jediocvsta vanudna from Fiji, Qmaplay majorcarinae Fram Bonin Islands. Twenty other species are reported from Western Pacific islands (Bonin, Fiji, Marianas, Palau, Samoa, Society. Solomons, Tonga Islands) with comments about their distribulion. 9 7ingidae, Heleroplera, Western Pacific islands, taxeriomy, Museum National d'Histare Naturelle. ESA 8043, Laberataire d'Entamolagie,43 rue Button, F-75231 Paris cedex 05 France (e-mail: siulhertqunahnJdr); 31 January 2000, This paper deals with lave bugs (leteroptera: Tingidae) from Bonin, Fiji, Marianas, Palau, Samoa, Society, Solomons and Tonga Islands. The first species deseribed from the region was Tingis irregularis (Montrouzier, 1861) from New Caledonia. By 1956 Tingidae of Micronesia (Bonin, Voleano, Mariana, Caroline Marshall and Gilbert Islands) had reached 20 species (Drake, 1965). Since then it has risen to 8&1 species of which about 70% were described by Drake, Poor & Ruhoff between 1943 and 1966.1 added 9 species (Guilbert 1997a, 1997b, 1998a, }998b) to this list and 9 for Vanuatu (Guilbert, 1999). Herein, 29 species are recorded trom Bonin, Fiji, Marianas, Palau, Samoa, Society, Solonions and Tonga. Nine are new, namely, Cysteachila kraussi, Nesovypselas simplex. N. Strophil, Leptoptyy varians, L. longispine trom Salomon Islands. Efeonews samouensis trom American Samoa, FE. palavensis Irom Palau, Idiveysta vanuane from Fiji, Oniwplax major- carinae {rom Bonin. Repositories mentioned are Bernice P. Bishop Museum, Honolulu (BPBM), Natural History Museum, London (NHM), Muséum National d*Histoire Naturelle. Paris (MNHN) and National Museum of Natural History, Washington (NMNH). Abbreviations for colleetors are: J.L. Gressitt (IG); N.H.L. Krauss (NK); T.-C. Maa (TM); C.W, O’Brien (CO); C.W. Sabrosky (CS): R. Straatman (RS); G.A, & $.L. Samuelson (GSS); C.M. Yoshimoto (CY); B.LL. Gagne (BG); W-C. Cagne (WG): E.C, Zimmerman (EZ); J.M. Sedlacek (JS): E.S. Brown (BB); S.N. Lal (SL); C.H. Swezey (CII); WR. Kallen (WK); JA. Listinger (JL): M.K. Kamath (MR); R.G, Oakley (RO): JW, Beardsley (JB); F.M. Snyer (FS); W. Mitchell (WM). SYSTEMATICS Order HEMIPTERA Suborder HETEROPTERA Family TINGIDAE Cysteochila Stal, 1873 Cysieuchila comprises around [35 species, and has 4 widespread distribution; 12 species are known Irom the Indo-Pacific region: 4 are known fron the Solomon Islands: C. kraussi, C. idonea Drake,1956, C. consanguinea (Distant,1909), and C. preta Drake & Ruhotf!965, Cysteochila kraussi sp, nov. (Fig. 1) HOLOTYPE. SOLOMON ISLANDS: ©, Poitete, Kolombangara, 0-60m, 29.xi.1976, Nik, BPBM. ETYMOLOGY. For the collector, DESCRIPTION. Head. antennae, abdomen black, legs brawn fusecous, pronotum and hemelytra brown yellowish, Body 3.70mm long, |,08nim wide, Head short, with 2 long, inwardly curved occipital sptnes, 2 short straight frontal spines and stout, tubercle-like median spine. Antennae slender, I: 0.15, 1b: 0.14, Hh: 0.79, TV: ().46, longer than first 2 together, pilose, Bucculae broad, with 3 rows of areolae, closed anteriorly: labium short, reaching middle of mesosternum;: labial sulcus sinuate. narrow, open posteriorly. Pronotum broader than hemelytra, gibbose, deeply punctate, tricarinate, carinae straight. uniseriate: arcolae small. Collar moderately large, 6 arcolae wide transyersely on top, sligthly 544 4s o oe > tre < 7 4 om ate sre ee ane te PacomGac? io, a 2 i s ere 222 > aa FIG. 1. Cysteochila kraussi sp. nov., habitus. raised to form a tectiform hood not covering head. Paranota large, 8 areolae broad, reflexed inwards, resting on the pronotum, not joining on top but covering lateral carinae and not reaching median carina; areolae small, round. Hemelytra narrow, longer than body; costal area narrow, straight, margins reflexed upwards, uniseriate, areolae small; subcostal area narrow, mostly biseriate, areolae small; discoidal area large, 6-7 areolae across, areolae small but larger than on costal and subcostal areas; sutural area 11 areolae wide, areolae moderately large, larger at apex; hypocostal laminae uniseriate. REMARKS. C. kraussi differs from C. nativa Drake,1960 by being smaller, and by having paranota not joining dorsally and subcostal area mostly biseriate. It differs from C. jimmina Drake,1960 (New Britain and New Guinea) by the paranota not meeting on top, costal area width and the number of areolae on the costal area, MEMOIRS OF THE QUEENSLAND MUSEUM which is 1-2 areolae wide for C. jimmina. It differs from C. brunnea Hacker,1928 (Queens- land) by the shorter labium (reaching the meso-metasternal suture in C. brunnea), paranota not meeting dorsally, labium length and coloration. Cysteochila prata Drake & Ruhoff, 1965 NEW RECORDS. SOLOMON ISLANDS: 1¢, Kolombangara, Pepele, 30m, 15.1[1.1964, PS, BPBM. REMARKS. C. prata is known from New Guinea. This is the first record of this species from the Solomon Islands. Cysteochila idonea Drake, 1956 NEW RECORDS. SOLOMON ISLANDS: 22 ,2¢ Guadalcanal, Honiara, 0-100m, NK, BPBM1985.163; 12, Kolombangara, Gizo, 0-140m, XII.1980, NK, BPBM1981.79. REMARKS. C. idonea is known from NE New Guinea, Kusaie (the Caroline Islands) and the Solomon Islands. Cysteochila vitilevuana Drake & Poor, 1943 NEW RECORDS. FIJI: 3 Viti Levu, Namosi Rd, 6km N Queens Hwy, 250m, 3-7.X1.1981, BG BPBM1981.601; 1d, 12, 90km E of Tavua, 28.VII.1967, JS, BPBM; 1¢ 52, Vanua Levu, Nakawanga, 9.X.1955, JG; 1? Vanua Levu, Nakorocau, 23.111.1966, NK, BPBM. REMARKS. C. vifilevuana is known only from Fiji, but is recorded here for the first time from Vanua Levu. Leptoptyx Drake & Ruhoff, 1965 Previously known from atopia and icelia from Solomon Islands and New Britain and following Drake & Ruhoff (1965), Leptoptyx is allied to Leptopharsa Stal 1873, but separated by its reflexed paranota, and longer and tectiform hood. Leptoptyx generally resembles Trachypeplus Horvath, 1926. Both have reflexed paranota, but Trachypeplus has spines on the pronotum and hemelytra, and wider hemelytra. Leptoptyx varians sp. nov. (Fig. 2) HOLOTYPE. SOLOMON ISLANDS:<¢, Honiara, Guadalcanal, 0-200 m, xi.1976, NK, BPBM1977.29. DESCRIPTION. Head, pronotum, legs and antennae beige to yellowish; abdomen dark brown; femora with a dark brown transverse medial band. Body long, with sparse minute pubescence, 2mm long, 0.74mm wide. WESTERN PACIFIC TINGIDAE 545 FIG. 2. Leptoptvx varians sp. nov., habitus. Head small, with 5 slender, suberect, mod- erately long spines; occipital and median spines parallel; median and frontal spines slightly stouter than occipital spines; frontal spines crossed; antenniferous process short; bucculae broad, mostly triseriate, closed in front; labium reaching middle of mesosternum, labial channel moderately wide. Antennae long, slender, I st and 2nd segments of equal length, 1: 0.08, 1: 0.08, IIL: 0.54, IV: 0.25; 4th segment shgthly stouter, pilose. Legs short, slender; tarsi slender. Pronotum gibbose, punctate; areolate on hind process, tricarinate; carinae subparallel, uni- seriate; areolae tiny, indistinct; lateral carinae terminating anteriorly at calli; median carina slightly higher than lateral carinae, contiguous with collar: collar 4 areolae broad, erected medially as a tectiform hood higher than top of pronotum: hood extending back between calli to pronotal disc; paranota narrow, reflexed, resting on pronotum, 3 areolae broad: areolae sub- quadrate; 2 inner rows forming a ridge with last ouler row, reaching median carinae anteriorly. Hemelytra moderately broad and broadened at base, constricted in the middle, flat: costal area mostly biseriate, triseriale al widest part; sub- costal area almost vertical, slightly sinuate. biseriate; discoidal area > 1/2 hemelytral length, 5 FIG. 3. Leptoptyx longispina sp. nov., habitus. areolae broad; junction between RM and Cu at apex of discoidal area slightly tumid; sutural area broad at apex, 6 areolae broad; areolae larger than in other areas. REMARKS. L. varians differs trom L. utopia Drake & Ruhoff 1965 in having paranota 3 areolae broad and a discoidal area 5 areolae broad, while £. afopia has paranota 4 areolae broad and a discoidal area 7 to 8 areolae broad. L. varians differs from L. icelia Drake & Ruholf 1963 by the costal area biseriate while it is uniseriate in L. icelia. Leptoptyx longispina sp. nov. (Fig. 3) HOLOTYPE. SOLOMON ISLANDS: &, Honiara, Guadalcanal, 0-200 m, xi.1980, NK. BPBM1981.79. PARATYPE; 1 © same data as holotype, BPBM. ETYMOLOGY. For the collector. DESCRIPTION. Head dark brown; cephalic spines yellow;, pronotum and hemelytra beige: hemelytra with some veinlets fuscous, legs and antennae beige: tarsi and first antennal segment sligthly darker. Body long, narrow, glabrous. Body 2.4mm long, 0.71mm wide. Head small, slightly pilose on top, with 5 long, slender spines; occipital and median spines parallel, adpressed; frontal spines subereel, crossed; bucculae broad, biseriate: antennae long, slender, I: 0.12, TE: 0.09, 111:0.79, TV: 0.37; 4th segment slightly stouter, more pilose; labium reaching middle of mesosternum; Jabial channel slightly enlarged posteriorly, Pronotum narrow, long, gibbose, punctate, areolute on hind process, Uri- carinate: carinae distinctly uniseriate; areolae small; lateral carinae not reaching calli anteriorly but interrupted before top of pronotum; median carina higher than lateral carinae: callar 5 areolae broad, elevated medially as a tectiform hood extending backwards to pronotal disc; paranota reflexed, resting on the pronotum, almost touching hase of lateral carinae, 4-5 arcolae broad; 3-4 outer rows of areolae forming a ridve With the timer row, Hemelyira long, narrow, flat, slightly constricted posteriorly at 2/3 length; costal area beni dorsally, narrow, uniseriate, arcolae moderately large, subquadrate; subcostal area straight, bent downwards, narrow, biseriate, areolae small; discoidal area >1/2 hemelytra, 5 areolae broad at widest pari, areolae small: junction of RM and Cu at apex of discoidal area slightly tumid; sutural area large al apex. 7 areolae broad at widest part, arcolac small at base and large al apex. REMARKS. L. /ongispine is easily distinguish- able from 1, varians by its narfower hemelytra and costal area, broader paranota and lateral carinae ending before the top of the pronotwm, It differs from L. atopia which has a biseriate costal area and a sutural area 7-8 areolac wide. Very similar to L. ieedia, it differs by its broader (4-5 areolae) paranota (2-3 in L. ivelia), and longer cephalic spines, L. afopia and L. icelia have short cephalic spines while LZ. varians and L- langispina have long spines. The lateral carinae MEMOIRS OF THE QUEENSLAND MUSKUM FIG, 4. Nesvevpselas strephii sp. nov. A, habitus; B, profile. of L, atopie are partly covered by the paranota unlike varices, longispina and icelia, Nesocypselas Kirkaldy. 1908 Nesoeypselas includes 12 species (including these 2 new species). Half are known from Pijt. and the others from neigbouring islands (Vanuatu, New Guinea, New Britain, New Ireland and now the Solomon Islands). Nesocypselas strophit sp. noy. (Fig. 4) HOLOTYPE. SOLOMON ISLANDS: 9 Popomanasiu, Guadaleanal, 4,400 m; 9-10,x1.1965, Hunuvalekama, Roy. Soc. Exped, Brit. Mus. 1966. 1. malaise trap, NHM, ETYMOLOGY. For the collector. DESCRIPTION, Head, pronotum and abdomen dark brown, Hemelytra and paranota hyaline. veins beige to yellowish. Legs and antennae yellowish, except tarsi dark brown. Body length: WESTERN PACIFIC TINGIDAE 3.45, body width: 2.25, hood length: 0.83, hood width: 0.46. Head small, sparsely pubescent, with 2 long, slender frontal spines. Bucculae small, short, mostly triseriate, slightly open in front. Labium long, reaching middle of meta- sternum. Labial channel broad, closed behind. Antenniferous process short, acute. Antennae long, slender, sparsely pubescent; 1: 0.31, I: 0.11, HT: 0.68, IV: 0.65. Legs long, slender; tarsi short, stout, pilose beneath. Pronotum short, wide, tri- carinate, slightly punctate, sparsely pilose; hind process rounded, not covering base of abdomen, with a transverse row of 6 rounded areolae on posterior margin. Carinae moderately narrow, raised, foliated, without areolae, extending from calli to posterior margin. Collar short, without hood, but anterior margin raised to form a small collarette. Paranota hyaline, large, reflexed but not resting on pronotum, outer margins not meeting dorsally but bent downwards, partly covering head but not covering pronotum, slightly serrate, 7 areolae wide, areolae large. Hemelytra hyaline, much larger than abdomen, sharply widened at base. Outer margins anteriorly bent upwards, then bent downwards. Costal area wide, 6 areolae broad at widest part, areolae large. Subcostal area sinuate, bent downwards, uniseriate. Discoidal area small, biseriate, 7 areolae, inner areolae larger. Sutural area triseriate, areolae large. REMARKS. N. strophii is distinguishable within the genus by its collarette and transverse row of areolae on the pronotum. It is very close to N. muiri Drake & Poor,1943 (Fiji) which has a costal area 6 areolae deep, clouded black transverse bands on the hemelytra and 2 transverse rows of areolae on the posterior margin of the pronotum. 547 FIG. 5. Nesocypselas simplex sp. nov. A, habitus; B, profile. Nesocypselas simplex sp. nov. (Fig. 5) HOLOTYPE. SOLOMON ISLANDS: @ Popomanasiu, Guadalcanal, 4,400 m; 10.xi.1965, Hunuvalekama, Roy. Soc. Exped. Brit. Mus. 1966.1, low vegetation in camp, NHM. ETYMOLOGY. For the collector. DESCRIPTION. Head, pronotum and abdomen dark brown. Paranota hyaline, hemelytra hyaline, with slightly clouded spots, veins brown. Legs and antennae yellowish, except tarsi and 4th antennal segment dark brown. Body 4.84 long, 3.78 wide; hood 2.40 long, 1.75 wide. Head small, spineless. Bucculae very small, short, biseriate, widely open in front. Labium reaching meso-metasternal suture. Labial channel broad, closed behind, ostiolar canal small. Antenniferous process short. Antennae long, slender, slightly pubescent; I: 0.35, II: 0.15, Ill: 1.23, IV: 0.92. Legs long, slender; tarsi short, stout, pilose beneath. Pronotum short, wide, tricarinate, slightly punctate, sparsely pilose; hind process rounded, 548 not extending over base of g abdomen, without areolae on posterior margin. Carinae moderately narrow, raised, foliate, without areolae, not reaching posterior margin. Collar short, without hood, triseriate. Paranota hyaline, large, raised, reflexed, not resting on pronotum, outer margins almost meeting in front and below but not meeting dorsally, partly covering head and pronotum, sligthly serrate, 8 areolae wide, areolae moderately large. Hemelytra hyaline, much larger than abdomen, sharply widened at base. Outer margins anteriorly bent upwards, then bent downwards. Costal area wide, 6 areolae broad at widest part, areolae large. Subcostal area sinuate, bent downwards, uniseriate, areolae larger than on costal area. Discoidal area small, barely distinct from the subcostal area, of 2 very large areolae. Sutural area biseriate, areolae large. REMARKS. JN. simplex resembles N. evansi Drake,1953 generally, but differs in its non areolate posterior process of the pronotum and uniseriate subcostal area. It is distinguished within the genus by its paranota. Nesocypselas dicysta Kirkaldy, 1908 NEW RECORDS. FIJI:¢ Viti Levu: Nausori, 11.1951, NK; 16 2 sex undet., Rewa, Muir, XII.1905, BPBM. REMARKS. WV. dicysta is known only from Fiji. Idiocysta China, 1930 Five of the 6 /diocysta species are restricted to Fiji. L. hackeri is only known from Samoa. Idiocysta vanuana sp. noy. (Fig. 6) HOLOTYPE. FIJI: 2, Wainigata Res. Stn, Vanua Levu, 0-100m, 5.X.1979, SL &GSS, BPBM1979,387. DESCRIPTION. Head and body beneath black; pronotum, hemelytra, antennae and legs beige; MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 6. /diocysta vanuana sp. nov. A, habitus; B, profile. posterior part of hemelytra and top of pronotum darker. Body 2.68 long, 0.89 wide. Head small, short, armed with a short median spine; bucculae small, narrow, mostly triseriate, closed in front; labium almost extending beyond mesometasternal suture; labial channel widened and closed posteriorly; antennae long, slender, I: 0.12, Il: 0.08, If: 0.55, TV: 0.43, 4th segment slightly pilose. Legs and tarsi long, slender. Pronotum gibbose, punctate, areolate on hind process, tricarinate, densely pilose between carinae; carinae raised, uniseriate, areolae subquadrate, moderately large; collar narrow, biseriate, raised dorsally to form a tectiform hood, lower than median carina but higher than top of pronotum, slightly extending forwards but not covering head; paranota large, raised, reflexed but not resting on pronotum, not meeting dorsally, but covering part of pronotum, also covering lateral carinae but not median carina, 6 areolae wide; areolae deep, moderately large, polygonal. Hemelytra narrow, a little wider than pronotum; principal veins slightly raised; costal area bent upwards, uniseriate, areolae large and WESTERN PACIFIC TINGIDAE quadrate; subcostal area same width as costal area, straight, almost vertical, biseriate, areolae small and rounded; discoidal area narrow, >1/2 length of hemelytra, 4 areolae wide, areolae small and rounded; sutural area 5 areolae at widest part, areolae small anteriorly, large posteriorly. REMARKS. /. vanuana is close to 1 bicolor Drake & Poor,1943 in hemelytral structure, however the paranota of /. bicolor are smaller (5 areolae wide) and meeting on top. It is also close to 1. dryadis Drake & Poor,1943 but its paranota are wider (6 areolae deep while /. dryadis 4). Its subcostal area is biseriate, while that of /. dryvadis is bi- to triseriate. Also similar to /. fijiana Drake & Poor, 1943, it differs by its wider subcostal area (biseriate while that of /. fijiana is uni- to biseriate), and by its paranota (larger by one row of areolae) which do not meet dorsally. Idiocysta hackeri China, 1930 NEW RECORDS. SAMOA: 2¢, Upolu, Afiamalu, 8.VI.1940, 2200m, Eugenia sp., CH; 12, Upolu, Afiamalu, 24.VII.1940, 800m, beating, EZ; 12, Upolu, Afiamalu, 13.VII.1940, 1000m, beating dead branches, EZ. 12, Upolu, Malololei road, 8.VII.1940, 1600-1800m, beating shrubs, EZ. 19, Tutuila, 12.1.1958, WK, BPBM. Idiocysta fijiana Drake & Poor, 1943 NEW RECORDS. FIJI: 12, Viti Levu, Nukurua Forest, logged area, 60-130m, 15.X.1979. MK, GSS, BPBM 1979.260. REMARKS. This specimen is slightly different from the type in that its paranota are almost closed dorsally but not as closed as in the type and the labium reaches the meso-metasternal suture, while in the type it extends to near the base of mesosternum. Idiocysta floris Drake & Poor, 1943 NEW RECORDS. FIJI: 2 42, Viti Levu, Namosi Rd, 16 km N Queen’s Hwy, 330m, 3-7.X1.1981, on Alpinia boia leaves, WG BPBM1981.601. Idiocysta dryadis Drake & Poor, 1943 NEW RECORDS. FIJI: 1d, Viti Levu, Namosi Rd, 3km N Queen’s Hwy, 100m, Myristica sp. leaves, BPBM1981.601; 9¢ 3° 3 fifth instar larvae, Viti Levu, Foster’s Pk, 10km N of Suva, 1100m, Mvistica sp. BPBM1981.601. Eteoneus Distant, 1903 Eteoneus comprises 20 species; 5 in the Ethiopian region, | in China; 14 in the Oriental FIG, 7. Eteoneus samoaensis sp. nov., habitus. region from west India to the Bismarck Archipelago and Palau Islands. This is the first description of an Eteoneus species from Samoa and 2nd from Palau. Eteoneus samoaensis sp. nov. (Fig. 7) HOLOTYPE.: AMERICAN SAMOA: & Mulinuu, Tutuila, 8.xii.1963, TM, BPBM. DESCRIPTION. All body, antennae and legs brown fuscous. Body 4.39 long, 1.54 wide. Head large, with sparse minute pubescence on top, armed with 4 spines; occipital spines short, slender; frontal spines small, tubercle-like; eyes very large, with post-ocular plate behind; bucculae small, narrow, triseriate, closed in front; labium long, reaching meso-metasternal suture; labial channel enlarged posteriorly, closed behind; | st segment of antennae stouter than 2nd, I: 0.25, Il: 0.15, 3rd and 4th segments lacking. Pronotum strongly gibbose, long, clothed with short pubescence, deeply punctate, areolate on hind process, tricarinate; median carina distinct all pronotum length; lateral carinae almost indistinct, present only on hind process, not reaching top of pronotum; calli wide, collar narrow, biseriate, not raised to form a hood: paranota almost indistinct, reduced to a slight ridge all pronotum length. Hemelytra Mat, not widened, same width as pronotum, covered with short pubescence: costal aréa straight, natrow, moderately bent upwards, uniseriate; areolae rounded, varying in size from small to moderately large; subenstal area narrow, slightly bent downwards, triseriale, ureolac rounded and small: diseoidal area >1/2 hemelytral length, 8 areolae wide at widest part, areolae small and rounded; sutural area large and short, 6 areolae wide at widest part, areolae small to large. REMARKS. E. samociensis is easily distinguish- able within the genus except for E. sigillatus Drake & Poor, 1956 by the uniseriate costal area, It differs from F. sigi//atus in its lack of median cephalic spine. Eteoneus palauensis sp. nov. (Fig. 8) HOLOTYPE. PALAU: 3, Imeliik Netkeng, Sabelthuap, 6.V1,1957, CS. BPBM, DESCRIPTION. Head, pronotum, fuscous, hemelytra fuscous with a yellowish spot through costal and subcostal areas, at level of apex of discoidal area, body beneath brown; legs and antennae yellowish, except tarsi and 4th antennal segment brown. Body 2.62 long, 0.92 wide. Head short, wide, slightly pilose on top, without spines: eyes large: bucculae short, broad, biseriale, closed in front; labium reaching meso-metasternal suture, labial channel wide, narrowed at apex, open behind; antenniferous process short; antennae Jong, slender, pilose; b 0.14, U: 0.11, Hk 0.94, 1V: 0.54, 4th segment slightly stouter than others; legs slender, slightly pilose; tarsi long, slender, pilose, Pronotum strongly gibbose, long, broad, punctate, clothed by short pubescence, unicarinate; carinae distinct, not raised all pronotum length: collar large, triseriate; paranota almost indistinct, reduced to a small ridge. Hemelytra moderalely longer and broader than body, the same width as prouotum, flat, constricted posteriorly at level of apex of discoidal area; costal area narrow, bent upwards, uniseriate, with areolae rounded and moderately large; subcostal area narrow, triseriate, with MEMOIRS OF THE QUEENSLAND MUSEUM FIG. &. Eteaneus palavensis sp. nov., habitus, areolae rounded and sinall; diseoidal area >1/2 length of hemelytra, 5-6 areolae wide at widest part, areolae small, same size as subcostal area; sutural area wide, 7 areolae wide at widest part, areolae small to large posteriorly. REMARKS. Like F. samoaensis, E. palauensis differs from the other species by its uniseriate costal area. It differs from EF. samoaensis. by lacking cephalic spines, smaller size and discoidal area 6-7 areolae wide. It differs from EF. sigillatus by lacking cephalic spines and its triseriate subcostal area (pentaseriate in E. sigillatus), Eteoneus lectus Drake, 1960 NEW RECORDS. SOLOMON ISLANDS: 22. New Georgia, Gizo, 100m, 16.VIL1964. JS. 12, New Georgia, Munda, 0-200m, XT.1972. NK. 1, New Georgia, Munda, 0-L00m, XIL1975. NK. 14, New Georgia. Munda, 0-100m, X1.1980. NK, BPBM1981.79. 9, New Georgia, Munda, 0-100m, X1.1976. NR, 1984-299, BPBM.2° 24, Guadalcanal, Honiara, 0-200m, 1.1984. NK, BPBM1984.168. 1d 22, Santa Ysabel, Kolotuve. IS.VI1960. CO. 1d, Kolombangara, Gizo, 0-140m, XIT.1980, NK. BPBM1981.79. REMARKS, This species is known trom New Britain, Bismarck Archipelago, Philippines, NW New Guinea and the Solomon Islands. Eteoneus esakii Drake, 1939 NEW RECORDS. PALAU: 1c, Malakal. 2.V.1957, CS. 60 3%, Koror, 19.1V,1957, sweeping Premme sp.. CS, BPBM. WESTERN PACIFIC TINGIDAE 5 —o 5 STS ls > O°, Oy FIG. 9. Omoplax majorcarinae sp. nov. A, habitus; B, profile. REMARKS. Known only from Palau Islands (Koror, Malakal, Peleliu). Omoplax Horvath, 1912 Originally a subgenus of Stephanitis, was raised to full generic rank by Takeya (1962). It now contains two species from the same islands. Omoplax majorcarinae sp.nov. (Fig. 9) HOLOTYPE. BONIN ISLANDS: 6, Chuo san, Chichi Jima, 300 m, 23-25.VIII.1980, Cinnamomum sp., JG, BPBM1980.377. PARATYPES: all from Chuo san, Chichi Jima. 12, 300m, 23-25. VII1.1980, Ligustrum sp., JG, BPBM1980.377; 1d, 300 m, 15.VIII.1980, JG BPBM1980.377; 12, 300 m, 23-25.VHI.1980, JG, BPBM1980.377. ETYMOLOGY. For the collector. DESCRIPTION. Head, body, legs and antennae brown to yellowish; hemelytra with a dark spot on costal area at anterior third; body beneath darker. Body 3.45 long, 1.85 wide. Head short, small, without cephalic spines; bucculae small, narrow, much narrower in front, mostly biseriate, closed in front; labium reaching middle of mesosternum; labial channel very w — wide, wider and closed behind; antennae long, slender, I: 0.26, II: 0.12, N Ill: 1.31, TV: 0.71,4th segment slightly pilose. Pronotum gibbose, punctate, areolate on hind process, unicarinate; carina all pronotum length, raised, uniseriate, with large areolae, subquadrate and hyaline, collar biseriate, inflated dorsally to form a hood cyst-like, narrow but sharply raised, extending forwards to cover part of the head, 4-5 areolae high, 8 areolae long, areolae small; a paranota narrow, present all pronotum length, a small hyaline surface just i0/ near calli, ridge-like \ posteriorly, wider just opposite humeri, there raised, of 5-6 small areolae, outer margins slightly serrate. Hemelytra sharply widened at base, wider at base than posteriorly, hyaline, with tumid area, outer margins slightly serrate, raised anteriorly and posteriorly bent downwards; costal area broad, 4-5 areolae broad at widest part, areolae large, angular; subcostal area wide anteriorly, narrow posteriorly, sinuate, 4 areolae wide at widest part, areolae small, tumid as to form a vesicle with discoidal area; discoidal area wide, tumid, 4 areolae wide at widest part, areolea small; sutural area moderately wide, 3-4 areolae wide, areolae large, angular. REMARKS. Omoplax majorcarinae is similar to O. desecta, and differs only in its labium extending to middle of mesosternum (beyond metasternum in O. desecta )and in its paranota having 5-6 (vs 3-5) areolae, and uniseriate median carina (vs biseriate). Omoplax desecta Horvath, 1912 (Fig. 10) NEW RECORDS. BONIN ISLANDS: 3¢ 52, Haha Jima, Okimura, 26.1V-9.V1.1958, FSr. 2d, Chichi Jima, Sakai-ura, Bull beach, 12-31.V.1958 FS & WM. 12, Haha Jima, Chibusa Yama, 200-462 m, 17-20. VIII.1980, Cimamonum sp., JG BPBM1980.377. 2d 12, Haha Jima, Chibusa Yama, 200-462 m, 17-20. VIII.1980, 2S 7. ee ONo, o s eve. FIG. 10. Omoplax desecta Horvath, 1912, profile. Terminalia sp., JG. BPBM1980.377. 1d, Haha Jima, Chibusa Yama, 200-462 m, 17-20. VIII.1980, beating, JG BPBM1980.377, 1d 22, Chichi Jima, Chuo san, 300 m, 23-25.VII1.1980, Ligustrum sp., JG BPBM#1980.377. 12, Is., Chichi Jima, Chuo san, 300 m, 23-25. VIII.1980, Cinnamomum sp., JG, BPBM1980.377. 2d 12, Chichi Jima, Chuo san, 300 m, 23-25. VIII.1980, Ardisia sp., JG BPBM1980.377. 1°, Chichi Jima, Chuo san, 300m, 23-25. VIII.1980, JG) BPBM1980.377. 22, Chichi Jima, foot of Mt Shigure, 100 m, 16.VIII.1980, dead branches, JG, BPBM1980.377. 1d 22, Chichi Jima, 13-16. VIII.1980, host #3, JG BPBM1980.377, BPBM. REMARKS. O. desecta is known only from the Bonin Islands (Chichi Jima and Haha Jima). Tingis Fabricius, 1803 Tingis parvoroe Guilbert, 1999 NEW RECORDS. SOLOMON ISLANDS: 1° 12, San Cristoval, Maniate, 6. VIII.1960, light trap, CO. 2¢, San Cristoval, Maniate, 5.VIII.1960, light trap, CO. 12, Guadalcanal, Roroni, 35km E of Honiara, 10m, 12.V.1964, light trap, RS. 1°, Guadalcanal, Roroni, 35km E of Honiara, 10m, 9.V.1964, light trap, RS. 1¢, Santa Cruz, Graciosa Bay, 0-50m, 1.1977, NK. 1 2, Mono, 150-300m, 6-11.X1.1980, Bidens sp., JG, BPBM1980.484. 12°, New Georgia, Gizo, 30m, 13.VII.1964, light trap, JS. 12, San Cristoval, Wugiroga, 7. VIII.1960, light trap, CO, BPBM. MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS. This species was described from Vanuatu, and is recorded for the first time from the Solomon Islands. Perissonemia Drake & Poor, 1937 Perissonemia torquata Drake & Poor, 1937 NEW RECORDS. SOLOMON ISLANDS: 1° 13 1 sex undet., Choiseul, Sasamongga, 0-100m, 11.1984. NK, BPBM1984.168. REMARKS. P. torquata is known from New Guinea, the Philippines and Solomon Islands. Minor differences exist between specimens from these localities. This species was described from Mindanao and has a labium reaching slightly beyond the midde of the metasternum, a triseriate subcostal area and paranota with 2 inner areolae opposite to calli, The outer row of areolae is tiny and without distinct veins on membranous mar- gins. In addition, the outer row has 3 areolae, while some specimens from New Guinea (NMNH) are slightly different; the outer row of the paranota has >3 areolae with a minute outer vein; and the subcostal area is mostly biseriate. These specimens from the Solomons have the labium almost reaching the meso-metasternal suture, a triseriate subcostal area, and narrow paranota, with | row of small areolae and 2 extra inner areolae opposite the calli. Eritingis Drake & Ruhoff, 1962 Eritingis includes 11 species; 9 from Australia (Qld, NSW and Sth Aust)and 2 from the Indo -Pacific region, The genus was erected for Tingis -like species with narrow paranota, uniseriate, erect and reflexed against the pronotum. Eritingis recens (Drake & Poor, 1937) NEW RECORDS. SOLOMON ISLANDS: 1d, Guadal canal, Kukum, 18.X.1953. EB, N °3923, Press. By Com. Inst. Ent., B.M. 1958-79, NHM. REMARKS. E. recens is known from Singapore, Vietnam, North Borneo, New Ireland and Palawan and now the Solomon Islands. Eritingis pacifica (Kirkaldy, 1908) NEW RECORDS. FIJI: 1d 12, Viti Levu, Korotongo, 0-100m, HI.1981, NK, BPBM1981.131. 19, Viti Levu, Nandi, 0-50m, IV.1981, NK, BPBM1981.131. REMARKS.Known from Fiji and New Britain. WESTERN PACIFIC TINGIDAE Teleonemia Costa, 1864 Teleonemia scrupulosa Stal, 1873 NEW RECORDS. FIJI: Viti Levu: Nandarivatu 850m-950m; Suva; Lami; 40km E of Nadi; Rakiraki 0-50m; Korotongo 0-100m; 90km E of Tavua; Tacirua. Ovalau: Levuka 0-200m. Vanua Levu: Nakawanga, Savusavu 0-100m. SOCIETY ISLANDS: Tahiti: Papeete 0-200m; Vallée de Papenoo 0-100m; Vaiufaufa 500m; Punaauia 0-150m; Punaauta 0-50m; Fare Rau Ape-Aorai trail 600-1400m; Arue 90-150m.TONGA: Tongatapu: Nuku'alofa 0-100m; Kolovai 0-100m; Mu’a 0-100m; Houma 0-50m; Haamonga. Vavau: Neiafu 0-100m. Eua: Hafu 100-200m; Pangai 0-100m. SAMOA: Savaii: Salelologa 0-50m. REMARKS. T. scrupulosa isa biocontrol species for Lantana sp. It is widely distributed in neotropical and oriental regions, also in the Indian islands and Australia. It occurs in Hawaii, Vanuatu, New Caledonia, Fiji, Tonga, Carolines, Solomon, Society Islands, Bismarck Archi- pelago and now Samoa. Phatnoma Fieber, 1844 Phatnoma pacifica Kirkaldy, 1908 NEW RECORDS. FIJI: 1d, Vanua Levu, Nakawanga, 9.X.1955, JG 1d, Vanua Levu, trans-insular road above summit, 500-550m, 6-9.X.1979, dry forest, GSS, BPBM1979.387. REMARKS. Known only from Fiji. Holophygdon Kirkaldy, 1908 The genus contains only H. melanesica from Fiji and H. nishidae Guilbert (1999) from Vanuatu. It was included in the tribe Litadeini (Drake & Ruhoff, 1965), due to the greatly swollen and pilose 2nd tarsal segment, like Litadea China, 1924. However, both genera share many different character states with other genera. In addition, Litadea is monotypic and known only from Madagascar. A revision of these genera would probably not support the validity of this tribe, Holophygdon melanesica Kirkaldy, 1908 NEW RECORDS. FUL: 1.4, Viti Levu, Namosi, rd, 8km N Queen’s Hwy, 320m, 3-7.X1.1981, MV light, BG & WG BPBM1981.601. 1d, Viti Levu, Namosi Rd, 14km N Queen’s Hwy, 300m, 3-7.X1.1981, on climbing aroid, BG & WG BPBM1981.601. 1d, Viti Levu, Colo-i-suva, 3-6.111.1963, Malaise trap, CY. 1d, Rewa, Muir, ?.1908. 1, Vanua Levu, Tabia (Thakaudrove), 0-2m, 5.X.1979, SL & GSS, BPBM 1979.387. Ww in aS) Berotingis Drake, 1956 Berotingis includes B. yvapensis from Yap (Caroline Islands) and B. guamensis and B. rugiana from the Marianas (Guam and Rota, respectively). These last two species are the only Tingidae known from those islands. This genus was separated from 7ingis (Drake, 1956) by the large eyes, narrow and scarcely reflexed paranota, absence of hood and the indistinct lateral carinae. Itis allied to Eteoneus (B.yapensis was formerly included in this genus) by the large eyes, the wide paranota, and the indistict lateral carinae. Berotingis rugiana Drake, 1956 NEW RECORDS. MARIANAS: 7¢ 62, Rugi, Rota, 29.V1.1946, RO, BPBM925, Stephanitis Stal, 1873 Stephanitis subfasciata Horvath, 1912 NEW RECORDS. PALAU: I, Angaur Is., 1.V.1954, Hernandia sp., JB, BPBM REMARKS. This species is known from China, Taiwan, Java, Burma, India, New Guinea and Palau. DISCUSSION Species richness in the western Pacific varies greatly among the islands. Small islands are species poor, but endemicity is high. The known tingid faunas of Hawaii, Tonga, and Society Islands are only Teleonemia scupulosa, which is an introduced species. Excluding 7. scrupulosa, the tingid fauna of Bonin, Marianas, and Samoa islands is represented by 2 endemic species each. 5 species are known from Palau, and 2 of them are endemic. Only | of 4 species known from the Carolines is endemic (to Yap); 10 species occur in Vanuatu, and 4 of them are endemic. New Caledonia has 14 species, all endemic; 20 of 21 species in Fiji are endemic, and 12 of the 21 species in the Solomons are endemic. Before 1999, only 2 species were known from Vanuatu, and 9 (including 7. scrupulosa) were added to its fauna this year. A recent study of the New Caledonian fauna (Guilbert, in prep.) added 19 species (12 endemic). Thus that many species remain undiscovered on Pacific islands. Many genera are restricted to the Pacific region. Omoplax has 2 species, both endemic to Bonin. Nobarnus Distant, 1920, including 5 species, is endemic to New Caledonia. Nesocypselas has 11 species distributed from Fiji to New Guinea. Leptoptyx, with 4 species, is known only from the Solomons and New Britain. The 7 species in /diocysta are restricted to Fiji, Samoa and Vanuatu. Holophygdon has | species in the Solomons and | in Vanuatu. Agaotingis Montrouzier, 1861, Cephalidiosus (Guilbert, 1998) and Corinthus Distant, 1920, each with 2 species, are endemic to New Caledonia. Berontingis 1s restricted to the Carolines (Yap) and Marianas. However, high generic endemism in this region may be due to oversplitting at the generic level. Many monotypic endemic genera could be grouped with related genera. Aulotingis Drake & Poor, 1943, related to Leptoypha Stal, 1873, is monotypic and known only from Fiji. The long tubular hood-like lateral carinae separate it from Leptoypha. Monotypic, Fijian Corythotingis Drake & Poor, 1943, differs from Physatocheila Fieber, 1844 in its pronotal structure. Monotypic, Fijian Nesocysta Kirkaldy, 1908 is allied to Nesocypselas but differs in paranotal shape. Monotypic Oeocharis Drake & Ruhoff, 1965from the Solomons, is distinguished by the globular hood concealing the pronotum. ACKNOWLEDGEMENTS 1 am grateful to Gordon Nishida, Bishop Museum, Hawaii, for the loan of specimens in his care, to Thomas J. Henry, National Museum of Natural History, Whashington D.C. and Mike Webb, Natural History Museum, London, for access to collections. This study was supported in part by the Large Scale Facility, TMR program. LITERATURE CITED CHINA, W. E. 1930. Heteroptera. In Insects of Samoa and other Samoan terrestrial Arthropoda, part II. Hemiptera, 3: 81-162. British Museum (Natural History). DISTANT, W.L. 1920. Rhynchota from New Caledonia, Annals and Magazine of Natural History series. 9, 6: 143-164, DRAKE, C.J. 1939. A new tingitid from Palau Islands (Hemiptera), Mushi 12: 102-103. 1941. New American Tingitidae (Hemiptera). Journal of the Washington Academy of Sciences 31: 141-145, 1946. A new tingid from Yap Island (Hemiptera). Musht 17: 27-28. 1953. Synonymic data and description of new genera and species of Tingidae (Hemiptera), Great Basin Naturalist 13: 91-99, 1956. Hemiptera: Tingidae. In Insects of Micronesia vol. 7. Bernice P. Bishop Museum. Pp. 101-116. 1960. Tingidae of New Guinea. Pacific Insects 2: 339-380. MEMOIRS OF THE QUEENSLAND MUSEUM 1965. Insects of Micronesia. Hemiptera: Tingidae. Occasional Paper of the Bernice P. Bishop Museum 7: 101-116. DRAKE, C.J. & POOR, M.E. 1936. New Indian Tingitidae (Hemiptera). Indian Forest Records 2: 141-149. 1937. Tingitidae from Malaysia and Madagascar (Hemiptera). Philippine Journal of Sciences 62:1-18. 1943. Fijian Tingitidae (Hemiptera). Occasional Paper of the Bernice P. Bishop Museum 17: 191-205. DRAKE, C.J. & RUHOFF, F.A. 1962. Synonymic notes and descriptions of new Tingidae (Hemiptera). Studia Entomologica 5: 489-506, 1965. Lacebugs from New Guinea, Borneo, Solomons, and other islands of the Western Pacific and Indian oceans (Hemiptera: Tingidae). Pacific Insects 7: 243-290. FIEBER, F.X. 1844. Entomologische Monographien, Leipzig. 138 pp. Tingideae pp. 20-111. GUILBERT, E. 1997a. Two new species of Dicysta from New Caledonia (Hemiptera, Tingidae). Zoosystema 19 (2-3): 515-521, 1997b. Nouveaux Tingidae (Hemiptera) de Nouvelle-Calédonie. Bulletin de la Société Entomologique de France 102 (3): 293-298. 1998a, Stenotrachelus : a new genus and two new species of Tingidae (Hemiptera) from New Caledonia. Australian Journal of Entomology 37 (1): 17-21. 1998b. Revision of the New Caledonian genus Nobarnus (Hemiptera: Tingidae) with description of three new species. European Journal of Entomology 95: 395-406. 1999, Tingidae (Hemiptera : Heteroptera) of Vanuatu (New Hebrides): new species and new records. European Journal of Entomology 96: 419-426, HACKER, H. 1928. New species and records of Australian Tingitoidea (Hemiptera). Memoirs of the Queensland Museum 9: 174-188. HORVATH, G. 1912. Species generis Tingitidarum Stephanitis, Annales Musei Nationalis Hungarici 10: 319-339. 1926. Hemipterologische Notizen Neiderlandisch-Indien. Treubia 8: 327-330. KIRKALDY, G. W. 1908. A catalogue of the Hemiptera of Fiji, Proceedings of the Linnean Society of New South Wales 33: 345-391. MONTROUZIER, P. 1861. In Perroud & Montrouzier. Essai sur la faune entomologique de Kanala (Nouvelle-Calédonie) et description de quelques espéces nouvelles ou peu connue. Annales de la Société Linnéenne de Lyon 11: 46-256. STAL, C, 1873. Enumeration Hemipterorum, vol. 3. Kongliga Svenska Vetenskaps-Akademiens Fér Handlingar 11: 1-163. TAKEYA, C. 1963.Taxonomic revision of the Tingidae of Japan, Korea, the Ryukyus and Formosa. Mushi 36 : 41-75. aus PARASITE FAUNA OF AUSTRALIAN MARINE OLIGOCHAETES SASCHA L. HALLETT, CHRISTER ERSEUS, PETER J. 0’ DONOGHUE AND ROBERT J.G. LESTER Hallett, S.L., Erséus, C., O’ Donoghue, P.J. & Lester, R.J.G. 2001 06 30: Parasite fauna of Australian marine oligochaetes. Memoirs of the Queensland Museum 46(2): 555-576. Brisbane. ISSN 0079-8835. A survey of 7,200 marine oligochaetes from Queensland (Moreton Bay, Brisbane, Heron Island and Lizard Island), New South Wales (Georges River, Sydney) and the Northern Territory (Darwin Harbour) revealed infections by 5 major parasite groups. Tubificid oligochaetes of the Limnodriloidinae, Phallodrilinae and Rhyacodrilinae were host to: 10 actinosporeans (Myxozoa) namely Sphaeractinomyxon ersei, S. leptocapsula, Endocapsa rosulata, E. stepheni, Endocapsa type | nov., Tetraspora discoidea, T. rotundum, Triactinomyxon of Roubal et al., 1997, Triactinomyxon type | nov. and Triactinomyxon type 2 nov.; an aseptate eugregarine (Apicomplexa) Oligochaetocystis sp.; an astomate ciliate (Ciliophora); a peritrichous ciliate Seyphidia sp. (Ciliophora); mermithid nematodes (Nematoda); a haplosporidian (Haplosporidia); and a coccidian (Apicomplexa). A single enchytraeid specimen, Grania sp., harboured astomate ciliates. Australia, actinosporeans, marine oligochaetes, parasites, protozoans. Sascha L. Hallett (sascha@wildfire.com.au), Peter J. O'Donoghue & Robert J.G. Lester, Department of Microbiology and Parasitology, The University of Queensland, St Lucia, Queensland 4072, Australia; Christer Erséus, Department of Invertebrate Zoology, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden; 22 September 2000. The literature suggests that parasites of marine oligochaetes are common (Giere & Pfannkuche, 1982; Raftos & Cooper, 1990), yet few specific examples can be found. Documented parasites belong principally to the protozoan orders Astomata (Ciliophora: Holotricha) and Gregarinida (Apicomplexa: Telosporidia) with at least 13 and 5 species described, respectively (Giere & Pfannkuche, 1982). Six actinosporeans (Myxozoa), Sphaeractinomyxon stolci Caullery & Mesnil, 1904, Sphaeractinomyxon type 1 & 2 Hallett et al., 1997a and Aurantiactinomyxon type 1, 2 & 3 Hallett et al., 1997a, also have been recorded. Prior to this study, no parasites had been recorded from an Australian marine oligochaete. Freshwater oligochaetes are reported as hosts to these parasite groups as well as to cestodes (e.g. Archigetes iowensis and Hunterella nodulosa), nematodes (e.g. Eustrongylides sp., Dioctophyma renale) (Raftos & Cooper, 1990) and rotifers A/bertia spp. (Koste, 1970; Erséus, 1976). They also are host to about 116 actinosporeans (McGeorge et al., 1997; Lom et al., 1997; Xiao & Desser, 1998a, b; El-Mansy et al., 1998b,c); some of these parasites are alternate stages in the life cycles of myxosporeans (Myxozoa) in fish (see Kent et al., 1994a; Lom et al., 1997). To date, some 24 life cycles have been elucidated and they have all involved freshwater species for which no other form of life cycle is known. No complete life cycle has been determined for any marine myxozoan. Myxosporeans are common in marine fish in Australia; this study was undertaken to determine if actinosporeans were present in Australian marine oligochaetes. Some of our early findings have been published elsewhere. These records are of the actinosporeans Sphaeractinomyxon ersei Hallett, O’Donoghue & Lester, 1998, S. leptocapsula Hallett, Erséus & Lester, 1999, Endocapsa rosulata Hallett, Erséus & Lester, 1999, E. stepheni Hallett, Erséus & Lester, 1999, Tetraspora discoidea Hallett & Lester, 1999, 7. rotundum Hallett & Lester, 1999, and Triactinomyxon of Roubal et al., 1997. Here, we present a review of all parasite types so far encountered in Australian marine oligochaetes, including published records. MATERIALS AND METHODS Sediment samples were collected to a depth of 15cm from the intertidal zone in Moreton Bay (27°15-25’S), and from Heron (23°27°S, 151°55’E) and Lizard (14°40’S, 145°28’E) Islands during 1995-1997 (Fig. 1). Infected oligochaetes were also obtained from Sydney, NSW (33°53’S, 151°10°E) and Darwin, NT (12°25’S, 130°51°E) (Fig. 1B). Sediment was collected in 500ml jars, each emptied into a 556 Mi} Hays inlet” North @ Redcliffe Point a Clontarf @ Scott Point Point a Woody Point @ Sandgate Moreton Cabbage Tree Bay Creek Brisbanie River 0. North : \ yg Lizard : Island s Heron Island Ul AUSTRALIA. ee . Brisbane /Sydney wo B FIG. 1. A, Moreton Bay collecting localities. B, Map of Australia showing other collecting localities. Brisbane incorporates Moreton Bay. 0 500km tt bucket containing about 4 litres of seawater, thoroughly mixed and the supernatant poured through a 0.4mm sieve. The contents of the sieve were washed into a petri dish and the supernatant returned to the bucket. This process was repeated about 5 times per sample. The material in the petri dish was then examined under a dissection microscope and any oligochaetes present were recovered for detailed examination under a compound microscope. The worms were placed on glass slides and a drop of 25% ethanol in MEMOIRS OF THE QUEENSLAND MUSEUM seawater was added to inhibit movement. Parasites (with the exception of those obtained from Lizard Island, Darwin Harbour and Sydney) were photographed, sketched and measured prior to fixation in either Bouin’s fixative for 24 hours for host identification or 3% gluteraldehyde in 0.066M cacodylate buffer for 24 hours for electron microscopy. For host identification, infected oligochaetes were transferred to 70% ethanol, stained in alcoholic paracarmine, mounted whole in Canada Balsam and examined under a compound microscope. Nematodes were fixed within the oligochaete host and several were cleared and stained with chlorlactophenol and Mayer’s haemotoxylin. Actinosporean descriptions follow the guidelines presented by Lom et al. (1997) except that ‘germ cell’ is used for ‘daughter cells’ and we give an additional measurement ‘basal width’ which is maximum width of the spore in apical view. New forms are identified in accordance with Kent et al. (1994) and Lom et al. (1997). Reference oligochaete host specimens containing parasites are lodged in the Queensland Museum (QM), Brisbane. RESULTS A total of 5,200 oligochaetes were examined from Moreton Bay and a further 2,000 oligochaetes from Heron Island. The density of worms sampled in Brisbane was up to 0.8 worms per cm’. Six parasitic/commensal groups were identified: actinosporeans (Myxozoa), aseptate eugregarines (Apicomplexa), astomate and peritrichous ciliates (Ciliophora), mermithid nematodes (Nematoda), haplosporidians (Haplosporidia) and coccidians (Apicomplexa) (Tables 1, 2). Phylum MYXOZOA Grassé, 1970 Class MY XOSPOREA Biitschli, 1881 ACTINOSPOREAN FORMS Kent et al., 1994 Infections by actinosporeans were detected in 196 (3.8%) of 5,200 worms from Moreton Bay, 25 (1.3%) of 2,000 worms from Heron Is. and in 8 worms from Lizard Is. One worm from Moreton Bay harboured a double infection of Sphaer- actinomyxon ersei and Tetraspora discoidea. Three worms, Duridrilus sp.(QMG463613), Limnodriloides sp. (QMG463615) and Doliodrilus diverticulatus (QMG463614) from Darwin Harbour and an unidentified oligochaete from Georges River, Sydney, harboured un- identified actinosporeans (all fixed samples). Ten actinosporeans belonging to 4 collective groups were found in marine oligochaetes (Table 1). PARASITES OF MARINE OLIGOCHAETES TABLE 1. Actinosporeans from Australian marine oligochaetes. ___ Actinosporean No. Infected Hosts a Host Sd Sphaeractinomyxon ersei 2 Doliadrilus diverticulatus Moreton Bay | ] Limnodriloides cf. victoriensis Moreton Bay | 44 Tubificidae sp./spp. Moreton Bay | 5 Tubificidae sp./spp. H Island ~ Thalassodrilides cf. gurwitsehi er On AST l Limnodriloides lateroporus Lizard Island 2 Bathydrilus sp. Lizard Island 7 - 1 _ Lizard Island I Sphaeractinomyxon spp- 24 — Tubificidae sp./spp. it Moreton Bay _S. leptocapsula 2 Heronidrilus sp. | Lizard Island _ | Endocapsa rosulata 3 Heterodrilus cf. keenani | Heron Island | Thalassodrilides cf. gurwitschi Lizard Island | 1 Heronidrilus sp. Lizard Island ; 9 Tubificidae sp./spp. Moreton Bay UE cf, rosulata : 6 Tubificidae sp./spp. | Heron Island E. stepheni 1 Heterodrilus cf, keenani Heron Island 2 Heterodrilus queenslandicus Heron Island - 1 oh! lubificidae sp. Heron Island | Endocapsa type | _ 8 __Tubificidae spp. Moreton Bay she rangiasin Av unnitcte, 2 Doliodrilus diverticulatus Moreton Bay Tetraspora discoidea 2 Tubiticidae sp./spp. Moreton Bay | Tetraspora rotundum 3 Tubificidae sp./spp. Moreton Bay wl 36 , 1 Limnodriloides cf. victoriensis Moreton Ba . Tide tinomyxon al ; 24 ‘Tubificidae sp./spp. x Maret: Bay Triactinomyxon type | 1 | Limnodriloidinae sp. Moreton Bay | L sia yP Z | : _ i | | Triactinomyxon type 2 | 1 Tubificidae sp. Moreton Bay ] I Unidentified: Pahific] Heron Island Sphaeractinomyxid 10 sepia seLabe. Moreton Ba: coelomic infection 53 pO OTe Be AD APD! kp intestinal infection 12 Tubificidae sp/spp. Moreton Bay caglomicintektion 1 Duridrilus sp. Darwin Harbour colors infsction \ Limnodriloides sp. Darwin Harbour chelomitankection | : Rolearites diverticulatus Darwin Harbour infection 1 CRT EIOAS SR Sydney SPHAERACTINOMYXON FORMS Sphaeractinomyxon ersei Hallett, O’Donoghue & Lester, 1998 (Fig. 2A) TYPE HOST. Doliodrilus diverticulatus Erséus, 1985 (Tubificidae: Limnodriloidinae). SITE IN HOST. Immature stages located within the coelom and mature spores present in the intestinal lumen. TYPE LOCALITY. Boggy Creek, Moreton Bay, 27°24’S, 153°09°E. SPECIMENS LODGED. QM G462452 (#110), G462453 (#185), G462465 (#210), G463601 (LI95-4), G463602 (L195-24/1), G463603 (LI95-24a), G463604 (LI95-24b). DESCRIPTION. Triradially symmetrical spores packed in groups of eight in the pansporocyst. Spores triangular in apical view, diameter 17-34um, basal width 17-33ym; ellipsoidal in side view, length 17-33ym. Polar capsules, 3, round to pyriform, centrally located, diameter (width) 3-5.5um, length 3-7pm. Sporoplasm, rounded triangular in apical view, single binucleate, about 46 germ cells, almost fills the spore cavity. REMARKS. S. ersei was detected in 47 (0.9%) of 5,200 oligochaetes examined from Moreton Bay; the number may be higher because 77 un- identified (immature) coelomic actinosporeans were also recorded (see Table 1). It also infected Limnodriloides cf. victoriensis Brinkhurst & Baker, 1979 from Boggy Creek, tubificid species from Heron Is. and Thalassodrilides cf. gurwitschi (Hrabe, 1971) (Limnodriloidinae), Limnodriloides lateroporus Erséus, 1997 and Bathydrilus sp. (immature) (Tubificidae: Phallodrilinae) from Lizard Is. Sphaeractinomyxon leptocapsula Hallett, Erséus & Lester, 1999 (Fig. 2B) TYPE HOST. Heronidrilus sp. (2 immature specimens infected) (Tubificidae: Rhyacodrilinae). 558 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 2. Parasites and commensals identified from Australian marine oligochaetes. #NR = not recorded. Lh Parasite No. Infected Hosts Host — Site a APICOMPLEXA a Giacaigeonystir se. | I Tubificidae sp. , Moreton Bay Unidentified coccidian species NR Limnodriloidinae sp./spp. _ Moreton Bay CILIOPHORA con, Radiophrya sp. 43+ Tubificidae spp. Heron Island Unidentified astome | Grania sp. | Heron Island Seyphidia sp. NR Limnodriloidinae sp./spp. Moreton Bay HAPLOSPORIDIA Haplosporidium sp. l Heterodrilus sp. Heron Island 15 Tubificidae sp./spp. _ Heron Island _ NEMATODA Mermithid nematode Heterodrilus cf. keenani | Tubificidae sp./spp. Heron Island Heron Island SITE IN HOST. Coelom. TYPE LOCALITY. Intertidal sand, Lizard Is., 14°40°S, 145°28°E. SPECIMENS LODGED. QMG462459 (LI95-16b), G462460 (LI95S-16e). DESCRIPTION. Spores triangular in apical view, diameter 20-24,1m, basal width 20-23m, length 17-22um. Outer valve cell membrane follows contours of inner valve cell membrane. Polar capsules ~5um long, slender, pyriform, orientated with pointed ends facing centre of spore, each positioned opposite a corner of spore about midway along spore radius, each contain polar filament with at least two turns. In side view, spore ellipsoidal to broad pyriform. Suture lines not discernible. Pansporocysts each with eight spores. REMARKS. S. leptocapsula was observed on only in 2 oligochaetes from Lizard Is. ENDOCAPSA FORMS Endocapsa rosulata Hallett, Erséus & Lester, 1999 (Fig. 2C-F) TYPE HOST. Heterodrilus cf. keenani Erséus, 1981 (Tubificidae: Rhyacodrilinae), SITE IN HOST. Immature stages in coelom; mature spores in intestinal lumen (Fig. 2E-F). TYPE LOCALITY. Heron Is., 23°27’S, 151°55’E. SPECIMENS LODGED, QM G462454 (H122), G462455 (H146), G463605 (LI95-24c), G463606 (LI95-16a), G462723 (#200). DESCRIPTION. Spore diameter 25-271, basal width 25-28um, length 20-23um, resembles rosette in apical view, compressed dorsoventrally in side view (Fig. 2C); develops in groups of 8 within pansporocysts (55-75um). Valve cells inner membrane subspherical; outer membrane follows contours of dorsal and ventral surfaces of this but exhibits swellings laterally in 3 regions equidistantly apart; inner and outer membranes close at each valve junction; suture lines prom- inent. Valve cell swellings formed within host, little or no further expansion on contact with seawater (Fig. 2D), Cavity diameter 19-23um, ellipsoidal in side view. Polar capsules ellipsoidal, 4-5um long and wide, located beneath a suture line anteriorly in, but not extruding from, spore, adjacent to remains of capsulogenic cell and perpendicular to processes, embedded in sporoplasm in side view, with polar filament with at least 3 turns. Sporoplasm granular, remaining areas of spore clear. REMARKS. Thalassodrilides cf. gurwitschi and Heronidrilus sp. (immature) from Lizard Is. and tubificids from Moreton Bay also harboured this parasite. Nine tubificids of the 5,200 oligochaetes from Boggy Creek, Moreton Bay, were infected. Spores were observed only in the coelom; none were found in the gut lumen as were Heron Is. infections. Spores were smaller FIG, 2. Actinosporeans from marine oligochaetes. A, mature Sphaeractinomyxon ersei spores (each with 3 polar capsules) in the intestinal lumen ofa limnodriloidine oligochaete from Moreton Bay. Fresh, unstained material. Scale = 25um; B, Sphaeractinomyxon leptocapsula pansporocysts with eight spores in the coelom of an oligochaete. Preserved material from Lizard Island. Scale = 50um. C-F, Endocapsa rosulata, fresh unstained material. C, Moreton Bay; D-F, Heron Island. C, spore in sea water, Scale = 20um; D, spores emerging from host. Scale = 251m; E, developing stages of E. cf. rosulata in coelom. Scale = 150um; F, spores of E. cf. rosulata free in intestinal lumen. Scale = 150um. PARASITES QF MARINE OLIGOCIIAETES (S i i A | te meee 7 AY Ad 560 than those from Heron Is. being 20-25um in diameter and 25-28um in basal width. Endocapsa stepheni Hallett, Erséus & Lester, 1999 TYPE HOST. Heterodrilus cf. keenani. SITE IN HOST. Coelom. TYPE LOCALITY. Heron Is., 23°27°S, 151°55’E. SPECIMENS LODGED. QM G462456 (H132), G462457 (H135), G462458 (H148). DESCRIPTION. Spore, diameter 25-28um, basal width 23-25y1m, length ~20um, irregularly shaped in apical view. Spore cavity diameter ~23um. Valve cells follow shape of roughly triangular spore cavity but with a single lobe-like swelling at one corner. Sutures, detectable, from corners to middle of spore. Sporoplasm roughly triangular in apical view, almost fills spore cavity, depressed in side view where polar capsules positioned. Polar capsules round in apical view, diameter 4-5um, pyriform in side view, length 4-5um, centrally located in spore, close to each other, beneath suture lines. Spore appearing ellipsoidal in side view, except that valve cells form an extension at one side. REMARKS. One Heferodrilus cf. keenani, 2 H. queenslandicus Jamieson, 1997 and | unidentified tubificid were infected with this actinosporean. Endocapsa type 1 nov. (Fig. 3) HOST. Immature tubificids. SITE IN HOST. Developing stages in peritoneum, mature spores in coelom and intestinal lumen. LOCALITIES. Hays Inlet and Boggy Creek, Moreton Bay, 27°16’S, 153°04’E and 27°24’S, 153°09’E . DESCRIPTION. Spores subtriangular in apical view, diameter 17-30um (22m, n=8), basal width 19-3lum (25um, n=2) (Fig. 3A). Spore body (inner valve cell membrane) basically round but corner formed at 3-way valve junction (Fig. 3A, C). Valve cells, upon contact with sea water, form 3 equally-sized biconcave processes (swellings) which join at their narrowest part at each 3-way valve junction (Fig. 3A, C). Swellings frequently present prior to spore contact with seawater. Polar capsules round, oval to pyriform in side view, diameter 3-5um (4um, n=4), located centrally in spore, proximal to one MEMOIRS OF THE QUEENSLAND MUSEUM other, each situated beneath a suture (and therefore opposite a spore corner) (Fig. 3A), embedded in anterior part of sporoplasm, do not form an apex (Fig. 3B). Spore round in side view, length 16-28 um (22,1m, n=7), diameter 19-31 um (25um, n=6) (Fig. 3B). REMARKS. Some spores had smooth rather than pinched corners. The non-protrusive polar capsules and reduced swellings, at times present within the host, place this actinosporean in the Endocapsa. Two other forms of Endocapsa have been recorded: E. rosulata; and E. stepheni. Endocapsa type | differs from both these in the shape of the spore and swellings. Its valve swellings encompass fully the spore body (visible in both apical and side view), whereas those of E. rosulata do not, and it possesses 3 swellings whereas FE. stepheni forms just one. The Aurantiactinomyxon types described from Hong Kong marine oligochaetes possess valve projections rather than swellings and the spores are considerably smaller, being less than 20j1m (Hallett et al., 1997). The principal difference between the Neoactinomyxon types described by El-Mansy et al. (1998b) and Endocapsa type | is that the valve cells of the former form triangular shaped extensions whereas they are curved in the valve cells of the latter. Endocapsa type 1 was recorded from 8 tubificid specimens. TETRASPORA FORMS Tetraspora discoidea Hallett & Lester, 1999 (Fig. 4A) TYPE HOST. Doliodrilus diverticulatus Erséus, 1985 (Tubificidae: Limnodriloidinae). SITE IN HOST. Coelom. TYPE LOCALITY. Boggy Creek, Moreton Bay, 27°24’S, 153°09°E. SPECIMENS LODGED. QM G462461 (#104), G462462 (#114), G462463 (#96). DESCRIPTION. Spore diameter 33-52,1m, basal width 33-38um, length 14-22um, disc-like, almost round, valve junctions form corner in apical view, dorsoventrally compressed in side view. Valve cell processes absent. Polar capsules, 3, subspherical in apical view, each within a pyriform capsulogenic cell, situated beneath a spore suture line, opposite a spore corner, bases proximal; pyriform in side view, located midway across spore at dorsal surface of sporoplasm. Polar filament oblique, at least 7 turns. Sporoplasm, shape similar to spore, unisporal, PARASITES OF MARINE OLIGOCHAETES FIG. 3. Endocapsa type | nov. from fresh material from Moreton Bay. A, drawing of apical view and B, side view. Scale = 5um. C, spores in seawater. Scale = 50um. contains at least one somatic nucleus and >100 germ cells. Pansporocysts, freely floating within host coelom, irregularly shaped, 47-70um, each containing 4 developing spores. Spores do not alter in size or shape following release from host. Development between pansporocysts asynchronous, but within synchronous. REMARKS. Twelve immature unidentifiable Tubificidae from Boggy Creek and 2 specimens of D. diverticulatus from Hays Inlet were infected with this parasite. Tetraspora rotundum Hallett & Lester, 1999 (Fig. 4B) TYPE HOST. Immature Tubificidae sp. SITE IN HOST. Coelom. TYPE LOCALITY. Boggy Creek, Moreton Bay, 27°24’S, 153°09°E. SPECIMENS LODGED. QM G462464 (#113, #186). DESCRIPTION. Triradially symmetrical spores packed in groups of eight in the pansporocyst. Spores triangular in apical view, diameter 17-34um, basal width 17-33um; ellipsoidal in side view, length 17-33um. Polar capsules, three, round to pyriform, centrally located, diameter (width) 3-5.5um, length 3-7um. Sporoplasm, rounded triangular in apical view, single binucleate, about 46 germ cells, almost fills the spore cavity. REMARKS. Three tubificids from Boggy Creek harboured 7. rotundum. FIG. 4. Tetraspora spores; fresh unstained material. A, T. discoidea in seawater. Scale = 10pm. B, T. rotundum pansporocysts (arrows) with four spores. Scale = 20um. TRIACTINOMYXON FORMS Triactinomyxon of Roubal et al., 1997 (Figs 5, 6) HOST. Limnodriloides cf. victoriensis. SITE IN HOST. Immature stages in intestinal epithelium (Fig. 6B, D) and mature spores present also in intestinal lumen. LOCALITIES. Hays Inlet, Clontarf Point, and Boggy Creek, Moreton Bay, 27°15-25’S DESCRIPTION (expanded from Roubal et al., 1997). Spore anchor-shaped, total length 96-142um (125m, n=6) (Fig. 5A). Valve cells inflate upon contact with seawater to form 3 anteriorly curved projections (caudal processes) from spore stylus at 90-100° angle (a) (Figs. 5A, 6A). Projections equal length 94-185um (138um, n=6) and width, at end taper to a point, equidistantly apart, arm base not wider than stylus base. Polar capsules, three, 3-5um (4um, n=5) long, located at anterior end of stylus, MEMOIRS OF THE QUEENSLAND MUSEUM pyriform, protrude at apex, bases abut, anterior end of each at slight angle (c.45°) facing away from other polar capsules (Fig.5 inset). Stylus widens gradually from tip to base (to ~22um). Sporoplasm, single, 17m long, with indis- cernible number of germ cells, irregularly in the stylus. REMARKS. This Triactinomyxon most closely resembles 7. /egeri Mackinnon & Adam, 1924 and T. ignotum Stolc, 1899, but spore dimensions are most like 7: /egeri (style 90-140um, arms 150um, sporoplasm 15-20um [Marques, 1984]). T. legeri and T. ignotum differ in size and number of germ cells in the sporoplasm (7: legeri= 24; T. ignotum = 8) and both develop in the intestinal epithelium of Tubifex freshwater oligochaetes. Triactinomyxon differs from 7: /egeri in its host and environment but these characters are con- sidered insufficient to establish a new species. The number of germ cells was indeterminable by light microscopy in the marine triactinomyxon but the size of the sporoplasm suggests about 32. The orientation of the processes varied between spores emitted from a single host and were directed out, up or down; spores were always observed under a coverslip and this may have influenced the orientation of the projections. The range in spore length and arm length appears initially large but these are comparable with other species (range of 50 and 100um respectively) (Marques, 1984; Lom & Dykova, 1992b). This was the first Triactinomyxon recorded from the marine environment. At least 25 oligochaetes harboured the parasite. Other hosts than Limnodriloides cf. victoriensis were ?Thalassodrilides sp. (Limnodrilinae) and Duridrilus sp. (Phallodrilinae). Infections were difficult to detect without squashing and killing the hosts. Clusters of 3 polar capsules in the intestinal epithelium or a distended intestine are indicative of an infection (Fig. 6B, D). Another 12 oligochaetes had developing Triactinomyxon- like stages in the intestine but no free spores were seen (QM G463607 #203). Triactinomyxon type 1 nov. (Figs 5, 6) HOST. Immature limnodriloidine oligochaete. SITE IN HOST. Posterior gut distended with pansporocysts. LOCALITY. Boggy Creek, 27°24’S, 153°09°E. DESCRIPTION. Ovoid spores in pansporocyst become anchor-shaped when exposed to PARASITES OF MARINE OLIGOCHAETES 563 FIG. 5. Drawing of Triactinomyxon types. Side view of spores fully expanded in seawater. A, Triactinomyxon of Roubal et al., 1997. Scale = 50um. [Inset: Spore in intestinal lumen before contact with seawater and valve expansion. Scale = 10um.] B, Triactinomyxon type | nov. Scale = 100um. seawater (Fig. 5B). Polar capsules (8-11 1m long) remain at anterior end of spore; valve cells form elongated stylus and 3 caudal processes. Spore body (polar capsules plus sporoplasm) 32-44um long (344m, n=20) (Fig. 6C) and total spore length 208-268um (236um, n=20). Caudal processes, 240-360um long (296m, n=46), curve slightly anteriorly, taper to point, a 90°. Suture lines visible, germ cells indiscernible. REMARKS. Triactinomyxon type | closely resembles Triactinomyxon of Roubal et al., 1997, except it is twice the size. When compared to other types (Marques, 1984 [8 types]; McGeorge et al., 1997 [1 type]; Xiao & Desser, 1998a [6 types]; El-Mansy et al., 1998b, c [9 types]), including those involved in a myxosporean life cycle (El-Matbouli & Hoffmann, 1989, 1993, 1998; Kent et al., 1993; El-Mansy & Molnar, 1997a, b; El-Mansy et al., 1998a; Székely et al., 1999; Eszterbauer et al., 2000), Triactinomyxon type | most closely resembles Triactinomyxon type 4 of El-Mansy et al., 1998c and Triactinomyxon ‘E’ of Xiao & Desser, 1998a. However, although the length of the processes and polar capsules are similar in type 4 of El-Mansy et al. and our type 1, the average length of the spore body (451m) is greater and the length of the style (149m) less for the former than that of the latter. Similarly, the process length of Triactinomyxon ‘E’ is within the range of Triactinomyxon type |, but the polar capsules of the former are smaller (Spm) as is the total spore length (spore axis 190-210um). Triactinomyxon type 2 nov. (Fig. 7) HOST. Unidentified tubificid oligochaete. SITE IN HOST. Not possible to determine if infection is coelomic or intestinal; intestine distended with pansporocyst within its boundary, but the latter may be above rather than within intestine. TYPE LOCALITY. Boggy Creek, 27°24’S, 153°09°E. DESCRIPTION. Pansporocyst ~139um across, with 8 subspherical spores (Fig. 7A). Spores fill out into characteristic triactinomyxon anchor- shape after contact with seawater (Fig. 7C). Polar capsules 3, pyriform, ~8 X 6um, at anterior end of stylus (Fig. 7B). Spore ~346um long, ~38um wide, narrowed anteriorly, posterior end dividing into 3 caudal processes directed posteriorly which taper at ends, ~517pm long, a 130°. Sporoplasm within stylus, ~130um long. REMARKS. The thick stylus and arms are reminiscent of Siedleckiella Janiszewska, 1955 but spores of Triactinomyxon type 2 did not appear interconnected either in the pansporocyst or in seawater and arms of Siedleckiella are blunt rather than pointed. Triactinomyxon type 2 is larger than any known Triactinomyxon. The arms of 7 magnum Granata, 1923 are >500um but the stylus is only 25-30um long. The size of the sporoplasm suggests it contains numerous germ cells. Development is asynchronous. Spores were liberated from pansporocysts under pressure. This form was observed on only one occasion and the host worm disintegrated during observation under the cover slip. Phylum APICOMPLEXA Levine, 1970 Order EUGREGARINORIDA Léger, 1900 Merogony absent; gametogony and sporogony present; typically parasites of annelids and arthropods, but some in other invertebrates. 564 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 6, Triactinomyxon spores, Fresh unstained material. A, Triactinomyxon of Roubal et al., 1997 spores in sea water. Scale = 125um. B, prominent polar capsules of Triactinomyxon of Roubal et al., 1997 spores within pansporocysts developing in the intestine of an oligochaete. Scale = 20m, C, anterior end of Triactinomyxon type | nov. spore in seawater showing polar capsules and sporoplasm. Scale = 20um, D, intestine of limnodriloidine oligochaete distended with developing stages of Triactinomyxon of Roubal et al., 1997. Left side uninfected, right side is filled with pansporoblasts. Scale = 200j1m, Family MONOCYSTIDAE Biitschli, 1882 Gamonts spherical to cylindrical, with anterior end little differentiated ifat all: oocysts biconical or navicular; mostly coelomic; the great majority are parasites of oligochaetes. Genus Oligochaetocystis Meier, 1956 Gamonts club-shaped, solitary or in syzygy; syzygy head-to-head [type-species: O. pachydrili (Lankester, 1863) Meier, 1956 emend. Leyine. 1977). Oligochaetocystis sp. (Fig. 8) HOST. Immature tubificid oligochaete. SITE IN HOST. All stages coelomic. Several gamonts appeared attached to the oligochaete intestine by their anterior ends. Infection extended from anterior region to middle of oligochaete, LOCALITY. Boggy Creek, 27°24’S, 153°U9"E. SPECIMEN LODGED. QM G462725 (#147), PARASITES OF MARINE OLIGOCHAETES DESCRIPTION. Gamonts (trophozoites) club-shaped and aseptate 148-185 (167) x 31-40 (35)um (Table 3). Posterior region paddle-like, narrows into a ‘neck’ that forms a bulb-like ant- erior region (Fig. 8A,B). Mucron inconspicuous. Nucleus clear, oval to circular, diameter 6-12 (10)um, in mid-posterior region. Gamonts solitary, granular, syzygy not observed. Gametocysts round (diameter 77-96um), con- taining numerous (100+) gametes (Fig. 8C, D). REMARKS. This is the only record of Oligo- chaetocystis sp. among the oligochaetes examined. The worm also harboured a coelomic actinosporean, Sphaeractinomyxon ersei. Oligochaetocystis contains 3 species: mesenchytraei, pachydrili and saenuridis in the coelom of European freshwater oligochaetes; Mesenchytraeus flavidus, Lumbricillus spp. (Enchytraeidae) and Tubifex tubifex (Tubificidae) respectively (Levine, 1977,1988). O. saenuridis also occurs in the seminal vesicles. We 365 “4 J44 4 aaa Cees ita FIG. 7. Triactinomyxon type 2 nov. Fresh unstained material. A, Pansporocyst (bound by arrows) in intestine of a limnodriloidine oligochaete, Scale = 50um. B, polar capsules and partial sporoplasm of a spore in seawater. Scale = 40um. C, drawing of side view. Scale = 250um. Phylum CILIOPHORA Dolflein, 1901 Internal ciliates were most prevalent in worms from Heron Is.; infected worms were rare in Moreton Bay and were represented usually by a single ciliate. In contrast, ectocommensal ciliates, represented by Scyphidia sp., were found only on Moreton Bay oligochaetes. Class OLIGOHYMENOPHORA de Puytorac et al., 1974 Order ASTOMATIDA Schewiakoff, 1896 Large body; uniformly ciliated; mouthless; endosymbiotic in oligochaetes, polychaetes, leeches, free-living flatworms and molluscs. Family RADIOPHRYIDAE de Puytorac, 1972 Body flattened; V-shaped apical cytoskeletal organelle; dense somatic ciliation. Radiophrya sp. (Fig. 9A-C) HOST. Tubificid oligochaetes. SITE IN HOST. Intestinal lumen, near clitellar region (Fig. 9A). SPECIMENS LODGED. QM G463608 (H116), G463609 (H133). LOCALITY. Heron Is., 23°27’S,151°55’E. 566 FIG. 8. Oligochaetocystis sp. in the coelom of an immature tubificid oligochaete. Fresh unstained material. A, drawing ofa gamont. Scale =20um. B, Gamonts. Scale = 50m. C, early gametocysts (arrows). Scale = 50um. D, Gametocyst (arrow) containing numerous gametes. Scale = 50m. DESCRIPTION. Body elongate, straight to curved, 140-180.1m long, tapering posteriorly and anteriorly, V-shaped attachment structure anteriorly, arms of V of about equal length, longitudinal kineties converge at each end; macronucleus large; much of one surface reinforced by cortical fibres extending almost whole length of body (Fig. 9B, C). MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS. Endocommensal, up to 30 per host. Cilia beating. Unidentified astomate ciliates (Fig. 9D-F) HOST. Grania sp. (Enchytraeidae) LOCALITY. Heron Is., 23°27’S, 151°5S°E. SITE IN HOST. Intestinal lumen (Fig. 9D). PARASITES OF MARINE OLIGOCHAETES SPECIMEN LODGED. QMG463610(H153). DESCRIPTION. Ciliates elongate, TABLE 3. Morphometric characters of the gregarine Oligo- chaetocystis sp. 177-257um long, 654m wide, | Mean | Minimum | Maximum No. posteriorly tapered, anteriorly blunt, — ——~—— _| um) =} (tm) __| observations oral structures (vestibulum, cytostome, | }AMONT duel | Maes Hest : trichites) absent, no prominent anterior — = asi SR ned FO == attachment region or structures such as!" ————+— eae 2 ee ee suckers, hooks or spines; macronucleus Nee width |__ 10.3 1 13,9 8 elongate ribbon-like; kineties |Bulbwidth 24.7 24.7 24.7 3 longitudinal, ~10 rows (Fig. 9E, F), |Bulblength 25.7 | 216 TR 3 Budding not observed. Nucleus diameter | 10.3 | 6.2 124 | 3 REMARKS. All characters conform to oc oat | gra | 173 86.5 3 | those of the Astomatida but none [ae = | 958 T =, al conforms exactly to any of the 8 families of astomate ciliates known from aquatic and terrestrial annelids. Apparent lack of an anterior attachment structure is consistent with the Anoplophryidae but most species in this family are large (>300um) and have about 40-100 kineties, whereas the present ciliates are up to 260um long and have only 10 longitudinal kineties. Species belonging to all other families have distinctive attachment organelles. At least 6 ciliates inhabited the worm with only | per segment. Order PERITRICHIDA Stein, 1859 Body goblet-shaped; conspicuous oral ciliature winding counter-clockwise to cytostome; scopula antapical; widespread throughout aquatic habitats; many free-living or symphorionts on diverse hosts; some commensals or parasites. Family SCYPHIDIIDAE Kahl, 1933 Solitary zooids; stalkless; disc-like scopula; generally found as epibionts mainly on invertebrates. Scyphidia sp. (Fig. 10) HOST. Limnodriloidine oligochaetes. SITE IN HOST. Attached to posterior integument (Fig. 10A). LOCALITY. Boggy Creek, Moreton Bay, 27°24’S, 153°09°E. DESCRIPTION. Zooid bell-shaped, ~40-50um long, aloricate, stalk absent; scopula sessile, solitary (but can occur in close proximity to each other), macronucleus U-shaped (Fig. 10A). REMARKS. This ectocommensal ciliate was found attached to the posterior region of oligochaetes from muddy habitats but was never associated with oligochaetes from the coral reef. About 10 ciliates were present simultaneously on a worm either singly or in small groups of 2-5 zooids, Scyphidia spp. attach to both vertebrates and invertebrates as well as to submerged objects; about 19 species have been recorded. Phylum HAPLOSPORIDIA Caullery & Mesnil, 1889 Histozoic, coelozoic unicellular parasites which form unicellular, typically uninucleated distinctive propagules, ‘spores’ without polar capsules or polar filaments (Perkins, 1990). Haplosporidium sp. (Fig. 10) HOST. Tubificid oligochaetes (including Heterodrilus sp.). SITE IN HOST. Intestinal epithelium, free spores in intestinal lumen. LOCALITY. Heron Is., 23° 27°S, 151° 55°E. SPECIMENS LODGED. QM G463611 (H97), QM G463612 (H147). DESCRIPTION. Unicellular spore without polar capsule or polar filament. Spores irregularly shaped, round to oval, 8-10 * 13-15um, groups of about eight enclosed by membrane (pansporoblast ~23um) (Fig. 10B). Spores possess characteristic operculum-like caps. REMARKS. Haplosporidia were first believed to be actinosporeans because of the arrangement of the spores in groups of eight in the intestine; however, they do not contain polar capsules, Sixteen of the 2,000 oligochaetes examined from Heron Is. harboured Haplosporidium sp. 568 MEMOIRS OF THE QUEENSLAND MUSEUM me st age ioe 2, at, et ~~ FIG. 9, Astomate ciliates. A-C, Radiophyvra sp. D-F, unidentified, from the intestinal lumen of oligochaetes at Heron Island. Fresh unstained material, A, ciliates in intestinal lumen of a tubificid oligochaete (arrowheads). Scale= 150m; B, curved ciliate in seawater, Scale = 50m; C, elongate ciliates in sea water. Scale= 100m; D, ciliate (arrow) in the intestinal lumen of an enchytraeid oligochaete. Scale = 100um; E, ciliate in seawater with macronuclei in focus. Seale = 100um; F, same ciliates with somatic kineties in focus. Scale = 100um. PARASITES OF MARINE OLIGOCHAETES FIG. 10. Protozoans from marine oliguchaetes. A, Seyp/icdia sp. contracted zooids attached to posterior region of limnedriloidine oligochaete. Fresh unstained material. Scale = 30pm, B, aplosporidinm sp. pansporoblast containing eight spores, Scale = Sym. Phylum APICOMPLEXA Levine. 1970 Class COCCIDEA Leuckart, 1879 Lite cycle with merogony, gamogony and sporogony; producing small intracellular gamonts, single macrogamete; monoxenous or heteroxenous parasites in vertebrates and invertebrates. Unidentified coccidian (Fiz. 11) HOST. Tubificid oligochaetes meluding Linnodriloidinae sp. SITE IN HOST. Primarily coelomic but the parasites appear to ‘bud off from intestine (Fig. 11 A). LOCALITY. Moreton Bay, 27°15-25°S DESCRIPTION, Gocysts nol seen or not present, sporocysts spherical, 12-l6,um (14um, n=8) tn diameter: contain ~16 sporozoites, roughly spherical, ~3pum across (Fig. 1]A-C). Meronts and gamonits not observed. REMARKS, An accurate record of prevalence was not maintained. Parasites were recorded also from the posterior part of the gut. Some sporocysts were observed to ‘excyst’ in seawater (Fig. 11D). The cysts resemble those of the protococcidian Gre/lia Levine, 1973 which has ellipsoidal sporocysts 12-!4um Jong. contain 5-14 sporozoites and which inhabits the coelom of archiannelids and polychaetes. The specimens we found had neither ellipsoidal oocysts nor large gamonts. The sporocysts may representa hitherto unidentified eucoceidian genus. Phylum NEMATODA Class ENOPLEA Order MERMITHIDA Family MERMITHIDAE Adult worms free-living; juveniles parasitic in body cavity of various invertebrates, primarily insects; no functional gut at any stage. Unidentified mermithid nematodes (Fig. 12) HOST. Hererodrilus cf. keenani and possibly other tubificid taxa. SITE IN HOST. Coelom, in posterior part of host. LOCALITY. Heron Is... 23° 27°S. 151° 55°E, SPECIMEN LODGED. QM G318270 (11145). DESCRIPTION. Two types of mermithid nematodes were observed; both juvemle (no gonads) and coiled in the host (Fig. 12). Width uniform along length. Anterior end blunt, posterior end tapered to a point. Buccal cavity, short, narrow, anterior end rounded, reminiscent of ascarophid nematodes. Pharynx, not well defined; lips absent. One nematode type long with green intestine, four specimens, length 1.400-2,064um, width 30-41lum (Fig. 12A). Second type short, withoul pigmentation, two specimens, length 244-1,080um, width 10-35 um (Fig. 12B). REMARKS. Four oligochaetes each harboured | nematode and a fifth oligochaete held 2 nematodes, one of each type. The nematodes were fixed inside the oligochaete host. Some 570 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 11. Unidentified coccidian from limnodriloidine oligochaete. Fresh unstained material. A, coccidian sporoblasts ‘budding-off intestinal epithelium into coelom, Scale = 30j1m; B, numerous sporocysts in coelom. Scale = 50m; C, sporocysts in seawater. Scale = 25pm; D, excysted sporocyst with sporozoites netted in seawater. Scale = 254m. PARASITES OF MARINE OLIGOCHAETES were then cleared and stained with chlorlacto- phenol and Mayer’s haemotoxylin but this did not facilitate identification of them. DISCUSSION Australian marine oligochaetes were found to be infected with a range of metazoan and protozoan parasites/commensals (Tables 1, 2). Tubificids dominated the oligochaete samples and although enchytraeids comprised at least 11% of the Heron Is. collections, only one worm was infected (Grania sp. with an astomate ciliate). Members of the tubificid Limno- driloidinae, Phallodrilinae and Rhyacodrilinae were infected with actinosporeans, ciliates, haplosporidians, coccidians, nematodes and gregarines. While double infections with species from the same parasite group were rare, infections by two parasites belonging to different groups were not uncommon; e.g. actinosporeans and ciliates or actinosporeans and gregarines. MYXOZOA. The actinosporeans we described (except possibly Triactinomyxon of Roubal et al., 1997) have been found only in Australia. Marine actinosporeans do not exhibit the same degree of host specificity as do freshwater actinosporeans and most (>60%) infect 2 or more marine oligochaetes e.g. Sphaeractinomyxon ersei was recorded from Doliodrilus, Limnodriloides, Thalassodrilides and Bathydrilus. Conversely, a species of marine oligochaete may be host to several actinosporean types. e.g. Thalasso- drilides is infected by both S. ersei and Endo- capsa rosulata but, as far as we have observed, not simultaneously. In contrast, 90% of freshwater actinosporeans occur in only one oligochaete species (cf. Marqués, 1984; Xiao & Desser, 1998c). One oligochaete species, however, may be host to several actinosporeans e.g. Tubifex tubifex is host to at least 12 types represented by 7 collective groups: Neo- actinomyxum, Guyenotia, Echinactinomyxon, Raabeia, Triactinomyxon, Hexactinomyxon and Synactinomyxon. The role of actinosporeans in marine oligochaetes is not yet fully understood but both are implicated in the life cycle of marine myxosporeans. Of particular importance are members of the myxosporean genus Kudoa that dwell in the skeletal muscles of marine fish and cause economic losses in mariculture around the world (Ireland, USA, Canada and Australia) (see Kent et al., 1994b; Palmer, 1995; Hallett et al., 1997b). Knowledge of the biology of the 57] FIG, 12. Mermithid nematodes in posterior coelom of oligochaetes from Heron Island. Fresh unstained material. A, green mermithid nematode. Scale = 100um. B, clear mermithid nematode. Scale = 100um. parasite, including its life cycle, is required. to control and alleviate these problems. Despite evidence that at least 24 freshwater myxosporeans alternate with an actinosporean stage in an oligochaete, similar connections (or any others) have not been established for any marine myxosporeans or actinosporeans. Indeed, Diamant (1997) provides experimental evidence for direct fish-to-fish transmission of at least | species, Myxidium leei, which would theoretically eliminate the need for an alternate invertebrate host (Diamant, 1997); though this direct transmission may alternate with transmission via actinospores (Lom & Dykova, 1995). Significant changes were proposed for the taxonomic treatment of actinosporeans during the course of our studies. The result of conclusions made by Kent et al. (1994a) that alternate myxosporean development probably occurs in all actinosporean families and genera, was the proposal that nominal actinosporean generic names should not be distinguished from 572 myxosporean genera and, consequently, all actinosporean genera and species be declared invalid (except Tetractinomyxon) thereby treating their nominal generic names as collective group names, The majority of actinosporean descriptions postdating these proposals adopt them without comment (McGeorge et al, 1997; Xiao & Desser, 1998; El-Mansy et al., 1998b,c). Various researchers over the past two decades have proposed the redundancy, either partly or fully, of every taxonomic level within the Myxozoa, notwithstanding the phylum itself (Kent et al., 1994a; Siddall et al., 1996), The Myxozoa are now recognised generally as being Metazoa, although their exact placement within this group is still unclear, Because the position of most actinosporeans within the phylum is currently uncertain, and for the sake of consistency, we employ the identification system recommended by Kent et al. (1994a) and Lom et al. (1997), although we would prefer to ascribe a binomial identity to actinosporean forms that do not have clear links to myxosporean genera. We have added ‘nov.’ at the start of descriptions of previously undescribed forms to avoid confusion between new forms and those already described (cf. El-Mansy et al., 1998c). Prior to this study, marine actinosporeans had been described from France (S. s/olci from the oligochaete Clitellio arenarius and Hemitubifex henedeni (= Tubificoides henedii) (Caullery & Mesnil, 1904; Marques, 1984)), Romania (S. stolci from Tubifex sp. (Radulescu & Motilicia, 1957)), England (Tetractinomyxon intermedium and 7. irregulare from the sipunculid worm Petalostoma minutum (Ikeda, 1912)) and Hong Kong (Aurantiactinomyxon type 1 & 2 from Pacifidrilus vanus, Aurantiactinomyxon type 2 from P. darvelli and Limnodriloides toloensis, Sphaeractinomyxon type | from Aktedrilus mortoni and Sphaeractinomyxon type 2 from Ainudrilus geminus (Hallett et al., 1997a)). We have now observed marine actinosporeans not only in Australian tubificid oligochaetes but also marine oligochaetes from: near Honiara, Solomon Islands (Tubificidae sp. (unidentified), Limnodriloidinae sp. and Heterodrilus sp.); Jiaozhou Bay, near Qingdao, China (Doliodrilus tener); Florida, USA (Tectidrilus squalidus); and Ascension Island, South Atlantic (Thalasso- drilides gurwitschi) (all material retrieved from second author’s collection; unpubl. data). This study demonstrates that actinosporeans occur in marine oligochaetes, and these findings MEMOIRS OF THE QUEENSLAND MUSEUM will facilitate experimental and molecular studies into the life cycle and systematics of this group. Infections in oligochaetes could not be detected on the basis of differences in worm motility, colour, size or shape. Instead, coelomic infections were readily discerned by the presence of iridescent spheres in the coelom when examined using incident light under a dissection microscope (the refractile bodies representing pansporocystic stages of the actinosporeans). The sporogonic stages varied considerably in size and internal composition and were more difficult to recognise, but the presence of numerous mature spores, monomorphic in appearance, was indicative of infection. Light microscopy generally revealed the coelom to be packed with pansporocysts in coelomic infections or the intestine to be distended with pansporocysts in gut infections. Similar distension has been reported also by Janiszewska (1955), Wolf et al. (1986), however, found the intestinal (freshwater) actinosporean Triactinomyxon gyrosalmo to be abundant in worms that were pale, had generalised anterior swellings and displayed an opaque outer layer. Similarly, Molnar et al. (1999a) could distinguish tubificids heavily infected with raabeia to be pale in colour and move sluggishly, El-Matbouli & Hoffmann (1993) recognised triactinomyxon infected tubificids by their whitish discolouration and Molnar et al. (1999b) noted that intestinal segments infected with neoactinospores appeared darker in colour and had thickened walls compared to uninfected areas. In contrast, Yokoyama et al. (1991) found that Raabeia sp. infections, apparently in the body cavity of the freshwater tubificid Branchiura sowerbyi, were not visible externally. We found coelomic infections to develop anteriorly in an oligochaete with more advanced stages located more posteriorly as the infection developed. El-Mansy et al. (1998a) observed triactinospores in the centrally located intestinal segments in moderate infections but in most segments when severe. The findings presented in this review support the literature that natural actinosporean infections have a low prevalence in oligochaetes being 0.1-9.5% (Mackinnon & Adam, 1924; Hamilton & Canning, 1987; Yokoyama et al., 1991, 1993a, 1993b; McGeorge et al., 1997; Hallett et al., 1998; Xiao & Desser, 1998c; Ozer & Wootten, 2000). An exception is the findings of El-Mansy et al. (1998c) who recorded a significantly higher prevalence of up to 43% which they attributed to their examination PARASITES OF MARINE OLIGOCHAETES 373 technique. Mixed infections are reported to be rare (Yokoyama et al., 1991; Xiao & Desser, 1998c) and indeed, only one of the 222 infected worms of the present study harboured two actinosporean species. In experimental infections, prevalences of 3 to almost 100 percent have been observed (Wolfet al., 1986; Kent et al., 1993; Yokoyama et al., 1993a; Uspenskaya, 1995; El-Mansy et al., 1997b; El-Mansy et al. 1998a; Molnar et al. 1999a,b; Ozer & Wootten, 2000). Coelomic actinosporeans accounted for most infections (>80%) and Sphaeractinomyxon ersei represented 23.9% of all infected worms from Moreton Bay. In contrast, all 25 actino- sporeans recorded by Xiao & Desser (1998c) parasitised the intestinal epithelium. Gut- inhabiting actinosporeans like Neoactinomyxum, Guyenotia and Hexactinamyxon spp., are common in freshwater oligochaetes, however the only gut-inhabiting actinosporeans we identified were forms of Triactinomyxon. There was no obvious seasonal influence on infections; the highest prevalence (12.1%) was recorded in June at Boggy Creek and the lowest (0.47%) in May at Scott Point. A number of other studies, however, have reported temporal patterns (see Yokoyama et al., 1993; Xiao & Desser, 1998c; El-Mansy et al., 1998c; Ozer & Wootten, 2000). Most infected oligochaetes were sexually immature, which hindered identification of the host. It is not known whether the presence of actinosporeans and other parasites may adversely affect the maturation of oligochaetes. Sexually mature worms, however, constitute only a small part of populations of marine oligochaetes at most times (Erséus, 1994). Different fixatives (including ethylalcohol, Bouin’s, Trump’s, glutaraldehyde and Karmovsky’s) were used to preserve infected worms, depending upon their intended use. These chemicals had varying effects on actinosporean spores when combined with the stain paracarmine and the mounting process which were necessary to identify the host worms. Karnovsky’s fixative preserved spores best even though they had ‘shrunk’ by 1-2um (3.5%); all other fixatives resulted in greater shrinkage of the spores. Endocapsa specimens were affected to a greater extent than Sphaeractinomyxon ot Tetraspora types, related perhaps to the valve cell properties of these groups; Endocapsa species have valve cells that swell whereas those of Sphaeractinomyxon and Tetraspora types do not (see Hallett et al., 1998; Hallett et al., 1999; Hallett & Lester, 1999). Detailed drawings accompanied with a range of micrographs are therefore recommended (see also Lom et al., 1997). Material fixed in Bouin’s or Trump’s fixative, but not processed further, appear to be representative but spores become deformed and distorted when stained and mounted. A complete taxonomic description requires information about: mature spores observed both in the host and free in seawater (or freshwater) to monitor changes in size and shape; the host stained in alcoholic paracarmine and mounted whole in Canada Balsam preferably after fixation in Bouin’s solution; and developing stages and where they are located in the host and their appearance. The anterior part of the oligochaete fixed separately in Bouin’s solution is needed to check host identity regardless of the intended use of the infected worm for either histology, TEM or DNA studies. APICOMPLEXA. Gregarines are widespread, common parasites of invertebrates, particularly arthropods. New hosts ingest gametocysts and the oocysts they contain, to become infected. The low prevalence of gregarines in marine oligochaetes suggests they may be an atypical host group; nevertheless gregarines formed gametocysts and developed. Apparently, oocysts were ingested and sporozoites migrated across the gut epithelium into the body cavity of the host worm where they underwent gamogony, but neither oocysts containing sporozoites nor syzygy were observed. We believe this to be the first record of a marine tubificid oligochaete infected with a gregarine, but enchytraeids infected by monocystid gregarines dominate the records (Giere & Pfannkuche, 1982). CILIOPHORA. In this study, most ciliates were fixed in sifu within worms so that the hosts could be identified. Subsequent dissections of infected worms yielded few intact ciliates and silver impregnation studies were uninformative. None of the ciliates were therefore identified to species level. Nonetheless, the ciliates were clearly astomate and peritrichous species as determined by their morphological characters. Similar groups occur as endo- or ecto- commensals in oligochaetes (Giere & Pfannkuche, 1982). Ciliates should be carefully extracted from host tissues and observed live to note colour, rigidity, motility, contractile vacuole, location, etc. Ciliates should then be fixed in Bouin’s, Stieve’s or Dafano’s fluid prior to silver impregnation to reveal patterns of ciliation, attachment structures, nuclear arrangement, etc. Regrettably, the best 574 silver stain to use for any particular ciliate group can vary considerably so it is advisable to use multiple stains including silver nitrate, silver carbonate and silver proteinate. The prevalences and intensities of infection by internal ciliates were lower in Moreton Bay than at Heron Island. External ciliates, however, were observed only on worms from Moreton Bay, particularly those collected at Boggy Creek with substantial silt loads compared to the pristine coral cay of Heron Island; the parasite fauna reflects this difference. Similar external ciliates (order Sessilida) were reported from Limnodriloides biforis Erséus, 1990 in muddy sediments associated with estuarine habitats in Hong Kong (Erséus, 1990). Grania spp. (Enchytraeidae) for some reason seem untouched by the parasites which were relatively prevalent in marine tubificids. NEMATODA. Oligochaetes are phoretic, paratenic, intermediate and definitive hosts for nematodes (Poinar, 1978; Smith, 1985). Juvenile mermithid nematodes primarily infect insects but also molluscs, crustaceans, arachnids and other invertebrates; mermithid adults are free-living (Poinar, 1976). The juvenile emerges from the egg, penetrates into the body cavity of an invertebrate host, develops for a period to emerge finally into the environment where it moults to the adult stage. Mermithids almost always kill their host (Poinar, 1976); the large size of the nematodes relative to their oligochaete host observed in the current study imply a similar life cycle for the mermithids we found. The majority of associations of nematodes and oligochaetes are with earthworms (Poinar, 1978). Only | of the 83 species cited in Poinar (1978) was a tubificid (freshwater) oligochaete and its nematode was listed as unidentified. Smith (1985) subsequently documented at least 3 microdrile families as hosts to this group; these were Lumbriculidae, Naididae and Tubificidae to members of Dioctophymatidae, Rhabditidae, Anisakidae and Mermithidae. Smith (1985) found Mermithidae usually in the anterior half of the naidid oligochaetes examined. All six nematodes we found were located posteriorly in the tubificid oligochaetes. Smith (1985) considered migration to be unlikely but rather assumed that the larvae hatched quickly from ingested eggs, penetrated the gut and then resided in the anterior portion of the worm. No more than two nematodes were observed per worm in either study and Smith (1985) recorded that the total MEMOIRS OF THE QUEENSLAND MUSEUM prevalence of infection was low and concluded that naidid infections were probably incidental. Parasitology of marine oligochaetes is fertile ground for studies and we made many new records of infection. The diverse range of organisms detected in oligochaetes indicate that worms are susceptible to external, intestinal and coelomic infections by both commensal and parasitic species. The host range, geographic distribution, habitat requirements and specificity of infection for parasites of marine oligochaetes remain to be determined. ACKNOWLEDGEMENTS We thank Ms Kylie Jennings and Mr Stephen Atkinson for collecting assistance in Moreton Bay and Heron Island and Dr Rob Adlard for providing the NSW sample; and Ms Barbro Léfnertz and Mrs Anna Hedstrém for staining and mounting oligochaete specimens. Assistance with digital imaging from Mr Atkinson is appreciated also. Financial support came from a University of Queensland Postgraduate Research Scholarship to SLH, an Australian Research Council grant (# A195065) to RJGL and Swedish Natural Science Research Council to CE. LITERATURE CITED CAULLERY, M. & MESNIL, F. 1904. Sur un type nouveau (Sphaeractinomyxon stolci n.g, n.sp.) d’Actinomyxidies, et son développement. Comptes rendus des séances de la Societé de biologie Paris 56: 408-410. DIAMANT, A. 1997. Fish-to-fish transmission of a marine myxosporean. Diseases of Aquatic Organisms 30; 99-105. EL-MANSY, A. & MOLNAR, K. 1997a. 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Some biological characteristics of actino- sporeans from the oligochaete Branchiura sow- erbyi, Diseases of Aquatic Organisms 17:223- 228. THE *‘WATER-TRAP’ SPINY OYSTER, SPONDYLUS VARIUS G.B. SOWERBY I, 1827 (MOLLUSCA: BIVALVIA: SPONDYLIDAE) FROM AUSTRALIA JOHN M. HEALY, KEVIN LAMPRELL AND JENNIFER L. KEYS Healy, J.M., Lamprell, K. & Keys, J.L., 2001 06 30: The ‘water-trap’ spiny oyster, Spondylus varius G.B. Sowerby I, 1827 (Mollusca: Bivalvia: Spondylidae) from Australia. Memoirs of the Queensland Museum 46(2): 577-588. Brisbane. ISSN 0079-8835. Spondylus varius G.B. Sowerby I, 1827, the largest member of the pectinoidean family Spondylidae, is recorded for the first time from Australian waters. Previously known from the Philippines, the Solomons and New Caledonia, it is now recorded from Orpheus and Lizard Islands in northern Queensland. A neotype is established for S. varius because: 1) no type material appears ever to have been deposited and no illustrations accompany Sowerby’s original description; 2) validity of the species has been questioned; 3) juveniles of S. varius can potentially be confused with other species of Spondylus (especially S. echinatus and S. castus). Aside from its exceptionally large size (rv height up to 400mm), S. varius is also unique among the Spondylidae in often producing water/gas-filled chambers (defined by thin, shell septa) in one or both of the valves in mature specimens (> 100mm, sometimes occurring in subadult specimens) especially those living in coral overhangs or threatened by epibiont smothering. These chambers possibly play a role in maintaining the position of the animal in relation to the shell margin, thereby optimising water current flow (and therefore feeding and respiration). Given the extremely odorous quality of the enclosed liquid (an acidic, saline solution) the chambers could also act as a last resort deterrent to predation. A total of 55 species of Spondylidae are now recorded from Australia. O Spondylidae, Pectinoidea, spiny oysters, bivalves, Australian fauna, new record, neotype. John M. Healy and Jennifer Keys, Department of Zoology and Entomology, University of Queensland, StLucia 4072 (e-mail: jhealy@zoology.ug.edu.au; jkevs@zoology.uq.edu.au); Kevin Lamprell, Queensland Museum, PO Box 3300, South Brisbane, 4101, Australia (e-mail: k. lamprell@uq.net.au); 14 April 2000 The waters around the Australian coastline are prolific in bivalve molluscs but especially so in tropical and subtropical areas which share several species with adjacent regions of the Indo-Pacific (Lamprell & Whitehead, 1992; Lamprell & Healy, 1998). Featuring prominently within the Australian molluscan fauna are the Spondylidae (‘spiny oysters’), with 54 previously recorded species (Lamprell & Healy, 1998). Although several species are known from the southern states, including some endemics (Lamprell, 1992), the Spondylidae are more speciose off the subtropical to northern Australian coastline. During November 1998, one of us (JK) collected a very large spondylid from shallow water off Orpheus Island, northern Queensland. After cleaning and close inspection, the species was determined as Spondylus varius G.B. Sowerby I, 1827 — a new record for the Australian bivalve fauna. A subsequent search of the collections of the Australian Museum (Sydney) revealed other specimens of this species. Spondylus varius is probably the largest spondylid to have ever existed, reaching a maximum of 400mm in shell height (umbones to ventral margin — measurement based on in situ measurements of Solomon Islands specimens, P. Clarkson, pers. comm.). Only the tropical Atlantic species S. limbatus GB. Sowerby II, 1897, rivals the dimensions of S. varius (up to 267mm maximum height according to Eisenberg (1981)). Together with the giant clams (Tridacnidae) and pen shells (Pinnidae), S. varius and S. /imbatus rank among the largest of the extant Bivalvia. In addition to its exceptionally large size, S. varius is also unique among the Spondylidae for its habit of often producing fluid and gas-filled chambers sometimes referred to as water-traps’ (Lamprell, 1986) in one or both valves. This phenomenon was investigated by Sir Richard Owen (1837, 1838) who concluded that chamber production in S. varits was probably a response of the animal to overgrowth from encrusting life such as corals. Juvenile §. varius differ from adults in colour and, usually, in the absence of water-filled chambers, leading to difficulty in identifying immature specimens and confusion with other species such as S. echinatus Schreibers, 1793 and 578 S. castus Reeve, 1856. Tomlin (1937: 350) considered Sowerby’s (1827b) original des- cription as vague and possibly based on material of more than one species. The purposes of the present account are to: 1) record Spondylus varius in Australian waters; 2) establish a neotype; 3) clarify the publication date; 4) provide a revised diagnosis based on material from the entire known range and 5) discuss possible functions of the fluid and gas-filled chambers. MATERIAL AND METHODS The living specimen of Spondylus varius GB. Sowerby I, 1827 was collected at depth of 10m, attached to a slight coral overhang, off the point between Pioneer Bay and Hazard Bay, Orpheus Island (18°367S,146°29°E) during November, 1998. The specimen was photographed in the aquarium at the Orpheus Island Research Station to record the distinctive mantle pattern. After removal of the animal (for later gonad studies) the shell was partially cleaned of excessive coral encrustations by overnight immersion in 10% w/v sodium hypochorite solution. The shell is now deposited at the Queensland Museum (QMM067048), For comparison with the Australian specimen, material from the Solomon Islands and the Philippines (all Lamprell Collection), and the Natural History Museum (London) (the specimen figured by GB. Sowerby II (1847) and Reeve (1856)) are also figured. Height measurements are from umbones to ventral margin; width measurements are from anterior to posterior margins. Abbreviations: AMSC = Australian Museum, Sydney; BMNH = Natural History Museum, London; I = Island; KL= Lamprell Collection; lv = left (or top) valve; pv = paired or conjoined valves; Qld = Queensland; rv = right (or lower) valve. SYSTEMATICS FAMILY SPONDYLIDAE Gray, 1826 Spondylus Linnaeus, 1758. TYPE SPECIES. Spondvlus gaederopus Linnaeus, 1758, by subsequent designation of Schmidt, 1818: 61. Spondylus varius G.B. Sowerby I, 1827 (Figs 1-6) Spondylus varius Sowerby, 1827; G.B. Sowerby 1, 1827b:1-2; Tomlin, 1937: 350; 1943: 143. Spondylus varius Sowerby, 1829; [sic] Fulton, 1915: 358, sp. 71; Lamprell, 1986: 68, pl. 25, fig. 2. MEMOIRS OF THE QUEENSLAND MUSEUM Spondylus varians Sowerby, [sic, no date specified]; G.B. Sowerby II, 1847: 426, pl. 86, figs 21, 22; Reeve, 1856: sp. 3, pl. 1, fig. 3: Hanley 1842-56: 293 Spondylus varians Sowerby, 1829 [sic]; Springsteen & Leobrera 1986: 325, pl, 92, fig. 16. Spondylus delessertii Chenu, 1845: 5, TYPE LOCALITY. Not stated by Sowerby (1827b); type locality of neotype (here designated; specimen BMNH 1952.10.30.3): ‘Pacific Islands’ (locality as stated on label). TYPE MATERIAL. NEOTYPE (here designated; Figs 1A-E, 6B): | py BMNH 1952.10.30.3, ‘Pacific Islands’ (locality as stated on label), Figured by Sowerby, 1847, pl. 86, fig. 21 and Reeve, 1856, pl. 1, sp. 3. Dimensions of neotype (excluding spines): lv height 154mm, width 137mm; rv height 169mm, width 140mm; pv height 169mm, width 140mm, depth 70mm; (measurements including spines) - lv height 163mm, width 143mm; rv height 174mm, width 155mm, pv height 179mm, width 155mm, depth 75mm. OTHER MATERIAL. (all measurements excluding spines; Table 1) AUSTRALIA: AMSC104588 Watson Bay, Lizard I., Qld, 14°40” S, 145°27°E, 2.5m among corals and sand lv height 195mm, rv height 255mm, pv depth 100mm; AMSCI50016 Orpheus I., Qld, 18°36’S, 146°29’E, 10m base of bommie, | py, lv height 155mm, rv height 200mm, pv depth 80mm, water chambers on lv; I pv QMM067048 approximately 10 m depth attached to a slight coral overhang off the point between Pioneer Bay and Hazard Bay, Orpheus I., Qld, 18°37°S, 146°30’E, rv height 170mm, Iv height 138mm, pv depth 94mm, water chambers in both valves (see Figs 2A-D, 3A,B). NEW CALEDONIA: AMSC100790 Noumea, ex- aquarium, lv height 175mm, rv height 220mm, pv depth 90mm, water chamber on ly; one rv with water chamber (no data); Sud nouvelle Caledonie 2pv Grotte Merlet, 22°42.4’S, 166°41.2’E, 20-30 m, 21.1.1993, lv height 190mm (for figures see Lamprell & Healy, 2000). SOLOMON ISLANDS: AMSC303014, | pv, off Bonegi I shipwreck (lv height 115mm, rv height 135mm, pv depth 80mm, water chambers in both valves) (specimen figured by Lamprell, 1986); KL, 4 py, off Bonegi I. shipwreck, 43m: spec | lv height 85mm, rv height 93.2 small water chamber in lv; spec 2 lv height 62mm, rv height 72.2mm, large water chamber in Iv; spec. 3 lv height 56mm, rv height 65mm, water chamber in lv; spec. 4 lv height 36mm, rv height 42mm, water chambers absent (Fig. 3C,F); PHILIPPINES: KL, off Cebu L., spec 1 lv height 94mm, rv height 113mm, water chambers absent; spec 2 lv height 99mm, rv height 113mm, water chambers in both valves (Fig. 3D,E); NO DATA: KL, ex Rombouts collection, | py, no locality data lv height 210mm, rv height 240mm, water chamber (empty) in both valves (senescent specimen; Fig. SA-C). DIAGNOSIS. Shell height of py to 400mm; elongate-ovate; inequivalve, top valve (lv) usually depressed to slightly inflated; lower (rv) appreciably deeper than lv. Sculpture of lv with numerous, low, irregular, radial ribs ornamented with dense, slightly flattened, raised or depressed SPINY OYSTER, SPONDYLUS VARIUS 579 FIG. 1. Spondylus varius Sowerby, 1827. Neotype (here designated) 1 py BMNH 1952.10.30.3, ‘Pacific Islands’ (locality as stated on labels). Figured by GB. Sowerby II, (1847) in the Thesaurus Conchyliorum: pl. 86, fig. 21 (see Fig. 6B herein) and Reeve, (1856) in the Conchologia Iconica: pl. 1, sp. 3. A, External view of |v. B, Internal view of lv showing visible (but empty) water chamber (arrow heads). C, External view of rv. D, Internal view of rv showing visible (but empty) water chamber (arrow heads). E, Profile view of pv. Scale bars = 20mm. 580 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 2. Spondylus varius Sowerby, 1827. A-D, Specimen from 10m, between Pioneer Bay and Hazard Bay. Orpheus I., Queensland, 18°37°S, 146°30°E (QMMO67048). A, External view of py from aspect of lv. B, Internal view ofrv showing the visible water chamber (arrow heads). C, External view of pv from aspect of ry, D, Internal view of rv showing two visible water chambers. Note broken uppermost water chamber overlying most of lower (intact) water chamber (the latter indicated by arrow heads). Scale bars = 20mm. SPINY OYSTLE, SPONDYLOUS FARIUS 381 nibs in juvenile specimens (< 100mm ry height), becoming longer wwards ventral margin. {n mature specimens (100-400mimn ev height) spines sometimes long but usually short, blunt and appressed (extremely stunted in specimens over 200mm rcv height); interstices in uneroded specimens have fine, dense appressed spines. Rv with similar radial ribs jo those of Iv; omamentation of foliations extending out from umbonal (fixation) area; remainder of rv similarly spined to lv. Hinge line broad, straight, showing typical spondylid dentiiion of two, large, isodont teeth on ry, fitting into sockets on ligament and ligament pit between teeth/sockets. Auricles broad. Externally, shell usually white, almost always with a clearly defined orange-red-purple areca umbonally which occasionally extends into the spines or over entire shell length (especially in small juveniles < SOmn rv height), Internally, glossy white with yellow-orange margin. Radiating ribs visible internally, Stronger marginally forming a strong, wide crenulated margin. Mature and senescent specimens (height of rv 100-400mm, excluding spines) oflen exhibiting Muid and gas-llled, vertically-stacked chambers within one or both valves. Chamber septa composed of smooth, translucent shell, associated with or completely surrounding the adductor scar (rarely passing partially under scar). Chambers sometimes also oceurring in juveniles (of ry height 60-80mm, excluding spines) bul absent in smaller specimens (rv height < 60mm), Fluid from ebambers usually lost through evaporation over an extended period of dry storaze (or rapidly if septum is cracked), Table | lists the incidence af externally detectable water chambers in material examined herein, For material =lO0mim in shell height the absence of chambers represents a verifiable absence of these structures, whereas in thicker, more mature specimens, the absence of visible, surface chambers (that is, ones which are fluid/gas-filled in fresh material) in one or both valves does not necessarily preclude the presence of one or more lower (empty) chambers. OBSERVATIONS ON LIVE-COLLECTED SPECIMENS. The live-collecled specimen from Orpheus 1. (Fis 1 A-D,2A-D,3A,B, 44,B) has lv 170mm in height, and 130mm in width (from anterior ta pestetiog margin in the lv), Although considerably encrusted with calcareous material, treatmemt with commercial hydroehloric acid soon revealed the external colour lo be white with some traces of red-purple umbonally. The rv contains twa visible witer chambers, one overlying the other (Figs 2B,D, 4A4,B) whereas the lv exhibils only a single visible chamber. The resence af Muid within the chambers is clearly indicated by the large gas bubbles which move around when the shell is tilted, During the initial stage of shell cleaning afler removal of the animal, the uppermost of the two water chambers lining the inside of the rv was accidentally broken, thereby allowing release of the clear, faintly yellowish fluid contents, This fluid was slightly more viscous than water and extremely foul-smelling (? partly due to the gas within the traps) — both charucteristics suggesting decomposing proleimaceous material, The septum of the broken chamber consists of iranslucent-white, glossy, shell material and has va thickness of approximately 0.3mm (lamella thickness), Internally bath valves are white with the excepuion of the crenulated, brown-purple margins and the regions. associated with the water chambers which appear brownish owing to the discoloured Muid contained therein. Live Spondvius varius has an extensive and gaping mantle which exhibits large, irregularly-shaped motilings (Fig, 3; white ona black background in Orpheus |. specimen, this study; orange on green background in Soloman Islands specimens, P. Clarkson. pers comm; white on yellow background (Slack-Smith, 1998, pl. 12.5). Variation in mantle colour is possibly associated with age, sex of the animal or represents some degree of geographical variation. L:dges of the mantle are red-orange in colour and ornamented with numerous blue ovelli, The ocelli are associated with rapid closure of the valyes during disturbances including the approach of a diver, Large specimens of S. varivy may weigh several lalograms, and live to be at least ten years of age (P. Clarkson, pers. comm.), HABITAT AND DISTRIBUTION. Spondylus varius is usually found cemented umbonally to venily sloping reefs, vertical prope ike, shipwrecks and concrete pylons, im depths from 3m to more than 69m water in prominent positions exposed to nutrient laden currents. Spechnens with particularly well-developed spines occur in sheltered positions within shipwrecks, where they are attached to a variety of surlaces and obscured by silt or sponges rather than calcareous enerusting organisms. They commonly provide habitat ‘for other molluses, including other cemented bivalves (other Spondylidae, Chamidae, Ostreoidea) and gastropods such as cowries and muricids. 5 582 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. Spondylus varius Sowerby, 1827. A, Profile of py from 10m, between Pioneer and Hazard Bays, Orpheus I., Queensland (QMMO 67048). B, Same specimen photographed alive in aquarium at Orpheus I. Research Station. Clearly visible is the black and white mottled mantle. C, Pv of juvenile from 43m off, Bonegi [sland wrecks, Solomon Islands showing long spines (KL). D, External of lv of juvenile specimen from off Cebu L., Philippines — note darker shell colour (purple in this case) of younger portion of shell (KL). E, Rv of same specimen as Fig. 3D showing water chamber (arrow heads), F, Internal views of left valve (at right) and right valve (at left) of juvenile specimen from 43m, off Bonegi I. wrecks, Solomon Islands, showing water chamber in lv (KL). Scale bars = 20mm. SPINY OYSTER, SPONDYLUS VARIUS adductor scar water chamber ‘lower' water chamber ‘upper’ water chamber FIG. 4. Spondylus varius Sowerby,1827. Sketches showing extent of the water chambers (dotted lines) in QM M067048. A, Lv; B, Rv — note two visible water chambers, the upper one broken. Scale bars = 20mm. varius is recorded from the Philippines (Lamprell, 1986; Springsteen & Leobrera, 1986), Solomon Islands(Lamprell, 1986), northern Queensland and New Caledonia (Lamprell & Healy, 2001). The species has yet to be taken in the eastern Pacific, and we know of no Hawaiian records. DISCUSSION. General remarks and comparisons. This is the first published record of this large and unusual bivalve from Australian waters, bringing the number of spondylid species known to occur in the region to 55. Sowerby (1827b) offered no precise locality data for S. varius other than to say that the material offered in the auction catalogue were ‘shells collected by Mr. Samuel Stutchbury on the coast of some islands of the Australian and Polynesian groups’. Tomlin (1943) located a complete copy of the Sowerby auction catalogue which records in one annotation that catalogue specimens of S. varius were from ‘Bow Island’ (‘one of the Paumoto group according to Tomlin (1943) = Hao Atoll, Archipel des Tuamoto, 9°22’S, 171°14, French Polynesia). Although we have not examined material from French Polynesia, there is no compelling reason to believe that S. varius does not occur there (possibly all of Stutchbury’s material was collected at ‘Bow Island’). Reeve (1856) gave the collection locality of the Cuming Collection specimen (designated herein as the neotype of S. varius) as ‘Pacific Islands’. Possibly because of its preference for semi-secluded habitats such as sunken ships or coral overhang, S. varius may have been observed previously by divers in Australian waters but not identified as such. Specimens of S. varius were also found in the collections of the Australian Museum, but somehow these have remained overlooked and undocumented until now. The exceptionally large size reached by S. varius (maximum height 400mm — measurement based on in situ measurements of Solomon Islands specimens — P. Clarkson, pers. comm.) could have precluded collection of specimens prior to this study as divers are generally hesitant to collect or even disturb large, attached molluscs such as these. The measurement of 450mm quoted by Lamprell & Healy (1998) for the Spondylidae (not stated by them but indirectly alluding to S. varius) was based on andecotal sources only. While diving on the reefs between Pioneer and Hazard Bays at Orpheus Island, one of us (JK) noted very large spondylids (estimated pv height of 250+ mm) in the vicinity of the S. varius specimen collected for this study. Almost certainly these were additional specimens of S. varius because no other spondylid occurring in the Western Pacific reaches this size. 584 FIG. 5. Spondylus varius Sowerby,1827. A-C, Aged specimen showing stunted nature of spines (KL; no locality data), height from umbones to ventral margin 257mm; A, External view of pv from lv aspect; B, External view of pv from rv aspect; C, Lateral view of pv. Scale bars = 20mm. In its juvenile state (and before any water chambers have developed), S. varius can potentially be confused with S. echinatus Schreibers, 1793 and S. castus Reeve, 1856. However, S. varius can usually be distinguished from these species by its umbonal colouration (white, or often orange-red-purple in S. varius; white with black speckling in S. echinatus and S. castus). Spondylus wrightianus Crosse, 1872 and S. tenuitas Garrard, 1966 also exhibit orange-red umbonal colouration like S. varius, but the former is smaller, solid and equivalved and the latter is a relatively small (65mm length), equivalved temperate Australian species with delicate spines. A review of the Spondylidae from New Caledonia (Lamprell & Healy, 2001) uncovered several new species and a significant number of species previously unrecorded from that region MEMOIRS OF THE QUEENSLAND MUSEUM including S. varius. It is therefore not surprising that S. varius should also occur on the Great Barrier Reef, and indeed it is likely that other Indo-West Pacific spondylids known from the Solomon Islands and New Caledonia will eventually be recorded from Queensland. Taxonomic history of Spondylus varius. Confusion concerning the date of publication of the description of Spondylus varius stems from the fact that Sowerby I chose to publish this text in an appendix to the auction catalogue of Samuel Stutchbury’s collection (principally of Polynesian shells and artefacts). Tomlin (1937) located incomplete copies of the catalogue, each with the date ‘July 1827’ added to a MS version of the missing title page and noted the discrepancy between this date and the tentative date *? 1829’ written on the Natural History Museum’s (then only) copy of the catalogue. Later a complete copy of the catalogue was sighted by Tomlin (1943) allowing him to clarify the date of the auction as July 26th, 1827. Tomlin concluded that this complete copy of the catalogue was probably Stutchbury’s, for it is annotated and bound into the back of a copy of J.G. Children’s Lamarck’s Genera of Shells bearing Stutchbury’s signature). Although it is impossible to state the exact date of publication of the auction catalogue, it seems almost certain that this occurred in the first few months of 1827, and not in 1826. For some reason, however, the year 1829 has regularly been associated with the name Spondylus varius Sowerby and, at present, the labels of the Natural History Museum specimen figured by G.B. Sowerby II (1847) and Reeve (1856) still bear the date 1829. Fulton (1915) appears to be the first author to cite 1829 as the authority year for S. varius Sowerby. Conceivably however, the first usage of this erroneous date may have been earlier than 1915, perhaps arising in dealers’ listings and subsequently repeated by Fulton. SPINY OYSTER, SPONDYLUS VARIUS 58 Ww FIG. 6. Spondvlus varius Sowerby, 1827. A, Illustration from Owen (1838, fig. 21) of a longitudinal section through pv of S. varius showing the multiple, vertically stacked, slightly offset septa of water chambers (note bifurcation of each septum into two lamellate layers). B, Illustration from Sowerby (1847, fig. 21, pl. 86) of S. varius here chosen as neotype (BMNH 1952.10.30.3): external view of pv from aspect of rv. To our knowledge, alteration of the name 5S. varius to S. varians was first made by Sowerby (1847) and unaccompanied by any explanation. This name change appears deliberate and not a lapsus calami as it occurs not only in Sowerby’s (1847) main text but also in his List of Figures and Alphabetical Index. Tomlin (1937: 350) considered the original description of S. varius (Sowerby, 1827) to be ‘rather vague, and one rather suspects that it may have been taken from a series comprising more than one species’. Undoubtedly this view was influenced not only by Sowerby’s choice of name (varius) but also his admission that ‘Under all other circumstances their form is exceedingly varied’ (Sowerby, 1827b: 2). Sowerby (1827a, b) did not illustrate S. varius nor did he mention deposition of any type material. It is clear from Sowerby’s original description that he was familiar with his new species outside of the series of specimens included in the auction (‘Many of the specimens of this shell are extremely beautiful, and there is in the Sale an interesting series of specimens of different sizes and variously circumstanced’, Sowerby, 1827b: 2). The neotype, figured by Sowerby (1847, pl. 86, fig. 21) and Reeve (1856, pl. 1, fig. 3.) (BMNH 1952.10.30.3.), has obvious historical significance, but to our knowledge, there is no surviving documentation to prove that it originated from the material offered in the auction catalogue or from other material used by Sowerby in his 1827 description (or from material collected after 1827). Associated with this specimen in the Natural History Museum isa note stating that it has no type status. Reeve (1856) referred to this shell as being from Cuming’s collection (“Cuming Mus.’) but as to the ultimate source of this specimen nothing is known, other than it was collected in the ‘Pacific Islands’. Our reasons for nominating BMNH 1952. 10.30.3. as neotype are: 1) this shell was accepted both by G.B. Sowerby II (1847) and Reeve (1856) (and presumably also Sowerby I who died in 1854), as characterising the species, and is accurately illustrated in colour by both authors; 2) the specimen is in an excellent state of preservation, even though the liquid contents of the water chambers have been lost through gradual evaporation; 3) the locality data associated with this specimen (‘Pacific Islands’) reasonably accurately reflects the known distribution of S. varius, although the species has yet to be collected from the eastern Pacific or the Indian Ocean. Nomination of a neotype is here deemed necessary to eliminate any doubts concerning the validity of S. varius (e.g. Tomlin’s (1937) comment that the species may have been based on specimens of more than one species). Structure and possible function(s) of the water chambers. The function of the water chambers in S. varius remains unclear. Over 150 years ago, Sir Richard Owen investigated the structure of the fluid/gas chambers in this species in two brief, but highly informative, papers (1837, 1838). He demonstrated a sequence of 14 vertically- stacked, slightly off-set chambers (each defined 586 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1. Comparison of Spondylus varius specimens examined. Collection Reference Locality (ncitie cue} tentitdlag Spinks} (exatucine spies) | QMMO67048 Orpheus I., Qld In both valves — 170mm | 138mm | AMSC150016 __ Orpheus f., Qld In lv only 200mm 155mm | AMSC104588 Lizard I.,Qld In lv only 255mm 195mm | AMSC100790 a New Caledonia In Iv only 220mm 175mm AMSC303014° _ Solomon Is In both valves 135mm — smm | KL Solomon Is In Iv only 65mm S6mm KL Solomon Is ___Inlv only 72mm 62mm KL Solomon Is __Chambers absent —__ 42mm | 36mm _ KL Solomon Is In lv only 93mm | 85mm [KL Philippines _ Chambers absent 113mm | 94mm KL __ Philippines Inboth valves 113mm | 99mm | KL —_ Nodata sn both valves 240mm 7 | 210mm ae Pacitic Islands In both valves 169mm 154mm by shell septa) in the rv (lower valve) of a large specimen which he had sectioned longitudinally from umbones to ventral margin (Fig. 6A). The upper valve of his specimen contained fewer, more lenticular chambers, which Owen (1837) originally ascribed to a reduced shell-secreting capability of the lv (upper valve) mantle but later (1838) interpreted as a result of reduced demand for such a capability in this region of the mantle (that is, both the upper and lower valve mantle margins probably have equal shell-secreting abilities). Septa of the chambers in the rv were shown by Owen to bifurcate away from the attachment area of the valve, so that strictly each septum is composed of two thin lamellae for most of its length. After discussing the phenomenon of septal production in other attached bivalves (ostreids and gryphaeids), in attached gastropods (vermetids and certain muricids) and in chambered cepahalopods, Owen (1837, 1838) hypothesised that the presence of septa in S. varius was probably a continuing response of the animal to impending overgrowth by corals. Hence the attached animal could effectively counteract overgrowth, and therefore maintain an unimpeded water flow for respiration and filter feeding, by raising the mantle edge on a series of successive shell platforms (= the septa defining the chambers). In support of this idea, Owen (1838) cited a personal communication from Stutchbury claiming that chambers only occurred in specimens associated with coral overhangs or other situations where over-growth by corals seemed imminent. In this species, water chambers may occur on the upper valve, the lower valve or commonly on both (Owen, 1837, 1838; Lamprell, 1986; Springsteen & Leobrera, 1986; present study), and even though it has been suggested that the fluid/gas chambers are only a feature of older specimens (Sowerby, 1827, 1848; Lamprell, 1986) we have sometimes observed them in relatively young shells (rv 72mm height) from the Solomons and the Philippines (Fig. SC-F). We cannot definitively assess Owen’s ‘overgrowth’ theory, but admit that his explanation is well argued and probably correct. However the chambers in some sub-adult S. varius — animals not as yet threatened by epibiont smothering — and the absence of such chambers in other spondylid species (including all other large species, Lamprell & Clarkson, unpubl. data), suggest that there may be additional functions for these structures. The foul-smelling odour of the fluid contained within the water chambers may offer another clue as to the function of these structures. Owen (1837,1838) had the fluid analysed by a colleague, Dr Bostock, who reported that ‘it was turbid, had an acid-saline taste, and a rank disagreeable odour’. After allowing the fluid to settle for 24 hours he determined that the clear supernatant was essentially a saline (sodium chloride) solution, with some hydrochloric acid and a little sulphuric acid also present. Dr Bostock made a point of emphasising that the fluid differed in its composition from sea water, raising the question as to its origin. If Owen’s (1837, 1838) proposed sequence of events for septal secretion is accepted, then sea water seems SPINY OYSTER, SPONDYLUS VARIUS the most likely source of the water chamber fluid. Possibly the mantle has an ability to chemically modify the contents of the fluid immediately before it is sealed off from the environment. More likely perhaps, chemical changes occur within the water once it is enclosed resulting in the precipitation of certain dissolved salts leaving a predominantly sodium chloride solution within the water chamber. In addition to the above results Dr Bostock isolated ‘a little brown matter’ which he tentatively concluded ‘gave the fluid its peculiar flayour and odour’, We can certainly confirm the repugnant odour of the fluid (? and gas) in S. varius and given this property, and the brittle nature of the septa, it is possible that fluid/gas-filled chambers may act as a last-resort defence against predators once maturity has been reached. The use of repugnatorial chemicals for defence against predators is widely recorded in the animal kingdom, although in most cases species adopting such a strategy advertise their inedibility (or even poisonous nature) through the use of bright colours and patterns (for example many nudibranchs, xanthid crabs). ACKNOWLEDGEMENTS We thank the Director of the Orpheus | Research Station, Mr M. Fogg, for allowing us access to the station’s facilities during November, 1998. Miss N. Wilson and Mr. B. O’Kane (Department of Zoology, University of Queensland) acted as dive buddies and volunteer assistants to one of us (JK) during the course of the field work. We extend our thanks to Mrs J. Pickering and Ms K. Way (Natural History Museum, London) for providing information on the status of the specimen of Spondylus varius figured by Sowerby and by Reeve and for a photocopy of the 1827 auction catalogue. Mrs Pickering and Ms Way are also thanked for facilitating our access to the Natural History collections for photography of spondylid material during our visit. Dr T. Waller (Malacology Section, Smithsonian Institution) kindly checked the Smithsonian malacological collection for possible type material of S. varius. We also thank Mr P. Clarkson (Port Lincoln, South Australia) for sharing his observations on living S. varius from the Solomon Islands. The referees are thanked for their constructive comments on the original manuscript. We are especially grateful to Ms Alison Miller (Australian Museum, Sydney) and Mrs Victoria Harrison (Queensland Museum Library) for obtaining copies of some old literature. This study was supported financially by a research grant and Senior Research Fellowship both from the Australian Research Council (to JH), a grant from the Malacological Society of Australasia (to KL) and University of Queensland Research Grants (to JH and JK). LITERATURE CITED CHENU, J.C. 1845. Illustrations conchyliologiques Vol. 3. Bivalves, Part 2. EISENBERG, J. M. 1981. A collector’s guide to seashells of the world. (McGraw-Hill Book Company: New York & Maidenhead). FULTON, 1915. List of the Recent species of Spondvlus Linne with some notes and descriptions of six new forms. Journal of Conchology 14: 331-360. HANLEY, S. 1842-1856. An illustrated and descriptive catalogue of Recent bivalve shells. with 960 figures by Wood and Sowerby (Williams & Norgate: London). LAMPRELL, K. 1986. Spondylus, spiny oyster shells of the World. (Robert Brown & Associates: Bathurst). 1992. Notes on the genus Spondy/us Linnaeus with descriptions of two new species from Western Australia, Memoirs of the Queensland Museum 32: 189-194. LAMPRELL, K. & WHITEHEAD, T. 1992. Bivalves of Australia. Vol. 1. (Crawford House Publications: Bathurst). LAMPRELL, K. & HEALY, J. 1998. Bivalves of Australia. Vol. 2. (Backhuys Publishers: Leiden). 2001. The Recent Spondylidae from New Caledonia and adjacent waters, with the descriptions of eight new species, and some taxonomic notes on the species Spondyvlus anacanthus (Mawe) (Mollusca, Bivalvia, Spondylidae). Bulletin du Muséum National d’Histoire Naturelle, Paris (Zoologie) 22. OWEN, R. 1837. Structure of water clam shell. Proceedings of the Zoological Society, 5: 63-66. 1838. Observations upon the camerated structure in the valves of the water clam (Spondylus varius, Sow.). Annals and Magazine of Natural History 2: 407-412. REEVE, L.A. 1856. Monograph on the genus Spondylus. Pls 1-18. In Conchologica Iconica or illustrations of the shells of molluscous Animals. Vol. 9. (Reeve: London). SLACK-SMITH, S.M. 1998. Ostreoida. Pp. 268-282. In Beesley, P.L., Ross, GJ.B, & A. Wells (eds) Mollusca the southern synthesis. Fauna of Australia, Vol. 5 Part A. (CSIRO Publishing: Melbourne). SOWERBY, GB. | 1827a. Catalogue of the whole of that splendid collection of rare shells, madrepores, carved paddles and other curiosities formed by Mr Samuel Stutchbury (principally among the Polynesian group of the south seas) and brought home by the Pacific Pearl Company’s ship the Sir George Osbourne. [Shell sale catalogue; 588 MEMOIRS OF THE QUEENSLAND MUSEUM authorship almost certainly GB. Sowerby I, the auctioneer; catalogue printed sometime prior to the auction date, July 26, 1827, but presumably within the early months of 1827). Pp 1-10, plus two pages with hand-written notes (one identifying the copy as belonging to Stutchbury and one giving locality data for three species, including Spondylus varius] 1827b. Observations on a few of the most remarkable shells collected by Mr. Samuel Stutchbury on the coast of some of the islands of the Australasian and Polynesian groups, together with descriptions of a few new species. Appendix to July 26th, 1827 Sales Catalogue, pp 1-4 with | late. SOWERBY, G.B. I 1847. Thesaurus Conchyliorum Vol. 1. Pp. 417-433, pls 84-89. (Sowerby: London). SPRINGSTEEN, F.J. & LEOBRERA, F.M.1986. Shells of the Philippines. (Carfel Shell Museum: Manila). TOMLIN, J.R. Le B. 1937. Book notes. 6. Stutchbury’s catalogue. Proceedings of the Malacological Society of London 22: 350. 1943. Book notes. Addendum to book note no. 6. Stutchbury’s catalogue. Proceedings of the Malacological Society of London, 25: 143. SIX SPECIES OF MEGASCOLECINAE (MEGASCOLECIDAE: OLIGOCHAETA) FROM NEW SOUTH WALES AND THE AUSTRALIAN CAPITAL TERRITORY B.G.M. JAMIESON Jamieson, B.G.M. 2001 06 30: Six species of Megascolecinae (Megascolecidae: Oligochaeta) from New South Wales and the Australian Capital Territory. Memoirs of the Queensland Museum 46(2): 589-602. Brisbane. ISSN 0079-8835. A small collection of earthworms from New South Wales and the Australian Capital Territory which are the subject of agricultural studies is shown to consist of five species: Notoscolex bakeri sp. nov.; Spenceriella bywongensis sp. nov.; 8. hamiltoni (Fletcher, 1887), S. macleayi (Fletcher, 1889) and S. nevillensis sp. nov. Notoscolex sensu stricto is known from New South Wales and Victoria, with one species, doubtfully included, from south-western Australia. N. bakeri appears to be the closest known relative of NV. montiskosciuskoi Jamieson, 1973, but differs from the latter in lacking calciferous glands. Spenceriella occurs in the Kosciuskan Division and Darling Basin province of Australia, Lord Howe Island, and Norfolk Island. S. macleayi (Fletcher), S. bywongensis and S. nevillensis spp. noy., all with two pairs of spermathecae, appear closely related but differ, among other respects, in having respectively two, three and four pairs of calciferous glands. S. hamiltoni, with three pairs of spermathecae and of calciferous glands, is considered to be the senior synonym of Anisochaeta chani Blakemore, 2000. Material from the Upper Manning River, New South Wales, previously referred to S. raymondiana (Fletcher 1887) is distinguished as Spenceriella manningi sp. nov. O Notoscolex, Spenceriella, new species, Megascolecinae. B.G.M. Jamieson, Zoology Department, University of Queensland, Brisbane 4072, Australia; received 11 May 2001, A collection of earthworms from New South Wales and the Australian Capital Territory which are the subject of agricultural studies by Geoffrey Baker, C.S,LR.O. Entomology, is shown in the present paper to consist of: Notoscolex bakeri sp. nov.; Spenceriella bywongensis sp. nov.; S. hamiltoni (Fletcher, 1887), S. macleayi (Fletcher, 1889) and S. nevillensis sp. nov. In addition, material from the Upper Manning River, New South Wales, previously referred (Jamieson, 2000) to S. raymondiana Fletcher (1887) is distinguished as Spenceriella manningi sp. nov. Notoscolex Fletcher, 1886 emend Jamieson, 2000 Notoscolex Fletcher, 1886a: 546. Notoscolex(emend.); Jamieson, 2000: 858-860. DIAGNOSIS. Setae 8 per segment. Combined 3 and prostatic porophores a pair on XVIII. A single gizzard, in V and/or VI. Meronephric, with exonephric stomate nephridium median to exonephric astomate micromeronephridia caudally (the notoscolecin condition), Prostates racemose, exceptionally tubuloracemose. REMARKS. Notoscolex occurs in NSW and Victoria; one species, doubtfully included, from SW Western Australia. The genus contains some species which form what is undoubtedly a monophyletic core and other species which can, with varying confidence, be associated with or placed in this monophylum. Members of the core (Notoscolex s. s., including the type-species N. camdenensis Fletcher, 1886a) are associated, inter alia, by 3 pairs of extramural calciferous glands, in XIV-XVI, a clear synapomorphy (Jamieson, 2000). Calciferous glands are absent in N. bakeri but it nevertheless shows close similarities to N. montiskosciuskoi which has 3 pairs of calciferous glands, albeit in X-XIT. Notoscolex bakeri sp. nov. (Figs 1, 2) MATERIAL EXAMINED. HOLOTYPE QMG218232. PARATYPES QMG218233-—218234, All from N.S.W., 36°10°S.149°20’E., in a black basaltic soil on a flat near Rock Flat Creek, ‘Rosebrook’ approx. 14 km NE of Cooma, L. Robinson; 3 macerated clitellate specimens. DESCRIPTION. Length 225 (P2), 250 (P1), >255mm (H). Width (midclitellar) 8mm (H, P1,2). Segments ca 185 (H) (posterior amputee), ca 200 (P1). Form moderately stout, anterior end club-shaped though apically tapering, clitellar region wider; segments III-XHI strongly biannulate, clitellar and more posterior segments weakly biannulate. Pigmented reddish brown; pale ventrally. Prostomium pro-epilobous but Sug FIG..1. Natascaley bukeri sp. nov., holotype. Ventral view of fore- and mid-body, MEMOIRS OF THE QUEENSLAND MUSEUM broken up by longitudinal and transverse Hssures extending the length of peristomium. First dorsal pore 4/5. Setae small, discernible with difficulty, clearest ventrally on clitellum; in 8 Jongitudinal rows, commencing on Il,.c and ¢ caudally with pale epidermal areolae; ce and 4 absent in X VIL; in XE (X11 not elear), aa: ab: be: ed: da = 8.5: S: 19: G@: 50%; caudally setal lines ¢ and « and occasionally / irregular; d lines becoming irregular shortly behind clitellum. Nephropores not externally visible. Clitellum annular, well developed, thick and rigid, embracing XIV-XIX; interrupted ventrally in XNVII-XIX: inter- segmental furrows and dorsal pores visible though weak; setae retained. Male pores on inconspicuous small circular papillae median to a lines in XVII; each papilla lyme near inner posterior border of a flat, ear-shaped genital marking with slightly raised margin, the two markings conjoined across midline anteriorly and filling XVIII longitudinally. A further genital marking, also flattened with raised margin, but forming a single midventral transverse ellipse, presetally in XIX; both sets of markings extending laterally into ah, Female pores paired anteromedian lo setae a of XIV. Spermathecal pores 4 pairs in 5/6-8/9, minute points median to a lines (Holotype). Some anterior septa strongly thickened; 9/10-11/12 very thick. Dorsal blood vessel paired seomentally, unpaired at septa, in VII-XVI and possibly further posteriorly, the two slender halves widely divergent so as to form a diamond patie in each segment; last hearts in XI, those in X-XII (all very slender) with their major connection to Supra-oesophageal vessel which is at Jeast partly double. Gizzard very large but flaccid in V, enclosed in septa 3/6 and 6/7: preceded by an equally wide pharyngeal mass. Oesophagus lacking calciferous glands. Intestine commencing in XVII (tragmentary). Meronephric; forebody segments with forests of minute (astomate?) parietal micromeroneph- ridia, Caudally with transverse rows of few astomate, integumentary micromeronephridia with, on each side of nerve cord, a median stomate exonephric megameronephridium, funnel with long preseptal neck. Large iridescent sperm funnels and copious sperm masses [ree in X and XI; small, compact slightly lobulated seminal vesicles in LX and XII, on their posterior and anterior septa, respectively, but also a pair on posterior septum of VIL. Small ovaries in NIT. Prostates elongate, flattened racemose {tubuloracemose?), extending laterally, NSW AND ACT MEGASCOLECID OLIGOQCHAERTES prostate gland spermatheral pore multiloculate spermathecal divarticulur B ventral nerve card spearmatheca) ampulla 8/9 Timm FIG. 2. Nofascoley bakeri sp. novi. paratype 1. A, dorsal view of right prostate. B, dorsa) view of spermatheeae of VIL, 277 site. restricted to XVITL tongue-like with lobulated surface, each gland with single ental bend; duet short, narrow and flaccid, lacking a muscular sheen. Penial setae absent. Spermathecae 4 pairs opening, anteriorly in their segments, each with subspheroidal ampulla, a wide, somewhat shorter ectally tapering duct; and a large, sessile, multiloculate iridescent diverticulum, usually with three lobes, which joins ectal end of duct; size unilorm (Paratype 1), ETYMOLOGY. For Dr Geotlrey Baker, who provided this. collection, in recognition of his contributionsto earthworm ecology in Australia. REMARKS. N. montiskosciuskoi Jamieson, 1973, isthe only other species of Notoscolex with four pairs of spermathecal pores and, like N. hakeri, has a double dorsal vessel (double also in N. cameroni), Wt differs from N. bakeri in the configuration of genital markings which form ill-detined transverse slightly tumid strips one in front of, the other behind the equators of segments XVIT-XXIL; those in XVII between the male papillae. N. baker? further differs in the absence of extramural caleiferous glands; in lacking, penial setae. in the auxiliary seminal vesicles, in VIII, and in the multiloculate, not simple clavate form of the spermathecal diverticula. Despite the absence of valeiferous glands in NV. hakeri, similarities with NV. montiskoscinxkot and the geographical proximity suggest that they are sister-species. Spenceriella Michaelsen, 1907, emend. Jamieson, 2000 Spenceriella Michaelsen, 1907: 161, Spenceriella (emend., ): tamieson, 2000, 1123, DIAGNOSIS. Perichaelin: setae 16 or more per segment. A pair of combined ¢@ and prostatic pores in scument XVIL Spermathecal pores intersegmental. Gizzard in V well-developed or rudimentary. Calciferous glands 3 or 4 pairs in X-XIT, XU, or absent. Intestine acaceate. Meronephric; bucca-pharyngeal tufts. present or absent. oesophageal nephridia astomate or stomate; caudal nephridia stomate (and astamite’’); nephridia in regions in which they are stomate (always?) with a median preseptal funnel and multiple intrasegmental funnels, Caudal enteronephry present or, more commonly, absent. Rarely with nephridial bladders, No setae median to the male pores. Prostates racemose or tubuloracemose. Spermathecae diverticulate. REMARKS. Spenceriella occurs in the Easter Subregion, Kosciuskan Division of Australia (sersu Kikkawa & Pearse, 1969): Lord Howe Island and Norfolk Island and the Western Subregion Darling Basin province (Jamieson. 2000). Spenceriella as redefined by Jamieson (2000), differs from Anisochaeta Beddard, 1890, in having multiple intrasegmental (not preseptal) nephrostomes, a less extensive series of ealciferous glands (if present), and more numerous setae which are not in the anisochaetin arrangement. It dilfers from Genraseolex in having segmental not intersegmental genital markings and multiple imtrasegmental (not preseptal) nephrostomes. The four species of Spenceriella described here are referable to the 8. rosabilis-group (Spenceriella s. sy.) of Jamieson (2000). Spenceriella bywongensis sp. nov. (Figs 3-8) MATERIAL. HOLOTYPE. (cliltellae) QM G 218235, PARATYPES. 1-3 (weakly clitellate) OMG2182346- 218240), All from A.C,T., 35°10°S.149°.20'E,, 'Bywony'. Sutton, ca 20 km NF of Canberra, improved pasture, July 2000. CSIRO sample ‘BR’. PARATYPES 6-8 (QMG218241-218243.'Gold Creek'. NE of Gungahlin, Canherra, J. Seown, July 2001). CSIRO sample *D*; faintly clitellate: vil) “) Reread spermathecal pore 2 genital markings spermathecal RX! drm MEMOIRS OF THE QUEENSLAND MUSEUM Xill xe Nv fernale pores XV ae ‘clitellurrr AVE Po: genral marking xvii XVII male pore FIG, 3. Spenceriella bywongensis sp, noy., holotype. Ventral view, A, forebody; B, midbody. DESCRIPTION. Length (clitellate specimens) 65-86mm, mean 75mm (FL, P1-5). Width 3mm. Segments 97 (P3 is shortest)-106 (Holotype is longest), Form (as preserved) slender and elongate, cylindrical, tapering at each end, with no clubbing; slightly wider at clitellum than elsewhere. Postclitellar segments weakly triannulate. Pigmentless buff in ethanol. Prostomium epilobous 2/3 or tanylobous. Peristomium bisected ventrally. First dorsal pore 5/6, Setac per segment 20 in XII and caudally; in forebody aa about 4 ab but ab smaller than be; = less than twice adjacent intervals. All rows regular. Clitellum XIV-XVIL (=4 segments): annular but interrupted ventrally, in ae, in XVII: intersegmental furrows obscured dorsally, dorsal pores and setae clearly visible. Male pores minute but distinct, ina lines of XVII, each near median aspect of a large circular porophore which fills XVII [longitudinally and extends slightly median of aand well laterally, beyond c; porophore more sharply defined medianly than laterally, on transversely oval porophores; each male pore on smal! circular papilla on porophore. Genital markings: prominent circular eminences, in [X-X1; unilateral left in IX in H; right in P2, PS: absent from IX in others; paired in X and XI (constant); each with conspicuous immediately presetal pore-like centre but prominence extending both pre- and post-setally. Posterior markings: midventral oval pads filling their segments longitudinally and extending laterally of b lines, in XVI, XX and XXI (constant in the 3 segments), or XX1L also (P1, P2, ‘pore’ unilateral left; P5, ‘pore’ paired) each with 2 tranversely elongated ‘pores’ immediately anterior to and including setal arc, excepting P| in which in XVI and XXII pore-like marking is unpaired midventral and is hardly bifid in XX and XXI:a slightly posterior crescentic swelling in XIX (H) and a midventral pad in XVIL(P3, P5); in only P2 a pair of widely separated porelike markings in NSW AND ACT MEGASCOLECID OLIGOCHAETES 59 vasa deferentia N sp ernathecal diverticulum sp emathecal ampulla FIG. 4. Spenceriella bywongensis sp. nov., Bywong material, holotype. A, dorsal view of right prostate; B, dorsal view of right spermatheca of VIII. XIX. Female pores paired on XIV, just anterior to setal arc, about one-third aa apart, in common glandular field. Spermathecal pores 2 pairs of large eye-like papillae, in 7/8 and 8/9, centred approximately in 4 lines. Several anterior septa moderately thickened; 8/9 strongest. Dorsal blood vessel continuous onto pharynx; hearts in X-XII latero-oesophageal with chief origin from calciferous vessel, near origin of latter from supra-oesophageal vessel; commissurals of IX anterior dorsoventral only. Gizzard large, barrel-shaped in V, with muscular sheen but readily depressed; extending to level of intersegment 8/9 posteriorly, septa 5/6 and 6/7 funnel-shaped around it. Oesophagus with 3 pairs of very large almost spherical calciferous glands, in X, XI and XII, each with its ventromedian aspect narrowly attached to oesophagus. Intestinal origin XVI, superficially appearing to commence in XV but septum 15/16 adherent to its anterior limit; acaecate; a well-developed dorsal typhlosole commencing though there rudimentary in XVIII. Meronephric with pairs of small tufted nephridia ventrally in IT], 1V and V; first 2 pairs small, those in V very large; all apparently exonephric; reducing to parietal but not numerous micromeronephridia by clitellar region. A large median preseptal funnel and post-septal (intrasegmental) funnels demonstrated for caudal nephridia, confirming placement in Spenceriella; all exonephric. Ovaries in XIII, an exceptionally large palmate LoS) prostomium peristomium first setiger Vi first dorsal pore mouth prostomium peristomium 1mm FIG. 5. Spenceriella bywongensis sp. nov. Bywong material. Dorsal view of prostomial region. A, holotype; B, paratype 1; C, paratype 3. pair with numerous strings of large oocytes, and paired thick funnels; conspicuous ovisacs on anterior wall of XIV, only slightly smaller than ovaries and with several oocytes projecting. Holandric, a pair of large testes and funnels in each of X and XI (only those in X notably iridescent), embedded in large free sperm masses; large racemose seminal vesicles in IX and XII, posterior pair larger. Prostates a pair of large, racemose glands, in XVIII-XX, but resolvable into a flattened S-shape the anterior half of which is enlarged and forms a square outline; vasa deferentia joining ental end of duct [as in S. macleayi]; duct thick and with muscular sheen, shorter than width of square portion of gland, extending directly median. Penial setae absent. Spermathecae 2 pairs, in VIL and LX with 594 MEMOIRS OF THE QUEENSLAND MUSEUM male pore FIG, 6. Spenceriella bywaengensis sp. nov. Bywong material. Variation in configuration of posterior genital markings. A, paratype |; B, paratype 2; C, paratype 3, large ovoid or polex-shaped ampullae tapering to ducts each bearing near its ectal end a thickly digitiform diverticulum which is a little shorter than, and (on right VIII) may be almost as wide, as ampulla; common duct of ampulla and diverticulum scarcely developed. The 3 Gold Creek specimens (P6-8) are closely similar to those from Bywong but differ in having paired pore-like markings in XVII (P6, P8) and large markings of this type in XIX (constant); furthermore, one specimen has a marking (unpaired, midventral) on XXII. In detail, there are paired (sometimes unilateral) pore-like markings in 1X (P6 only, unilateral left); X and XI (constant but unilateral left in P7); XVI (constant); XVII (P6,P8, absent P7); XLX-XXI (constant); and midventral marking in XXII in P7. Paired markings in XIX-XXI are progressively closer, posteriad, to the midventral line, those in XXI being on common pad. ETYMOLOGY, From the type locality, Bywong. REMARKS. Other species of Spenceriella with 2 pairs of spermathecal pores and 3 pairs of calciferous glands, in X-XII, are S. australis (Fletcher, 1886) from Burrawang and Mt Wilson, NSW, S. indissimilis (Fletcher, 1889), from Lake Alexandrina, S. Australia, and S. montanus (Spencer, 1900), from Mt Baw Baw, Victoria. S. australis differs from S. bywongensis in the wide separation of the spermathecal pores and a very different configuration of the genital markings. The little known S. montanus appears to differ in having unpaired, not paired genital markings in X and XI, together with those in XIX and XX. The geographically distant S. indissimilis is the most similar to S. bywongensis but differs, among other respects, in restriction of its anterior genital markings (in some or all of VII-X) to the presetal parts of their segments; in absence of any indication of pairing in the posterior genital markings; the closer pairing of the spermathecal and male pores; the smaller gizzard; and the bipartite condition of each prostate. The wide geographic separation of S. bywongensis, S. montanus, and S. indissimilis further supports specific distinction. Spenceriella hamiltoni (Fletcher, 1887) (Figs 9, 10) Perichaeta hamiltoni Fletcher, 1887: 399-400 Megascolex hamiltoni; Beddard, 1895: 373. Anisochaeta hamiltoni; Blakemore, 2000: 4. Spenceriella hamiltoni; Jamieson, 2000: 1225-1227, Fig. 4) ,28. Anisochaeta chani Blakemore, 2000: 18-19. New Synonymy. Spenceriella chant; Jamieson, 2000; 1193-1195, NEW RECORD. N.S.W., ca 5km S of Neville (33°43’S.149°13’E.), near Blayney and Bathurst, from improved pasture (CSIRO sample ‘E - dark head’); 2 weakly clitellate, QMG218244—-218245, and 2 (unregistered) aclitellate, macerated specimens. DESCRIPTION. Length 134-150mm (spec- imens | and 2 respectively). Width ca 6.4mm. Segments 128-145mm. Form (as preserved) NSW AND ACT MEGASCOLECID OLIGOCHAETES sp ermathecal FIG. 7. Spenceriella bywongensis sp. nov. Gold Creek material. Paratype 6. Ventral view of fore- and mid-body. un Oo wn vasa deferentia prostate gland prostate duct spermathecal pore. B spermathecal ampulla spermathecal diverticulum 1mm FIG. 8. Spenceriella bywongensis sp. noy. Gold Creek material. Paratype 6. A, dorsal view of right prostate; B, dorsal view of right spermatheca of VIL. slender, elongate, cylindrical, tapering at each end, slightly club-shaped anteriorly; clitellum about as wide as club-shaped region. Segments simple, lacking secondary annulation. Darkly pigmented, pale ventrally, in ethanol. Prostomium narrow epilobous 2/3, open. Peristomium weakly bisected ventrally. First dorsal pore 5/6. Setae per segment: 18 in XII, 20 in XX; about 28 caudally; in forebody aa ca 3.5 ab but ab smaller than cd (cd: ab = 1.5); dorsal break large, zz = about 4 zy. All rows regular, with occasional slight divergence. Clitellum weakly developed, from redder coloration, XIV-XVII (= 4 segments); annular?, intersegmental furrows and setae clearly visible; dorsal pores obscured (but minute elsewhere). Male pores in XVIII, each projecting mediad from a large mound; minute but distinct, very slightly median of a lines of XVIII, each on small but distinct circular porophore which fills about two-thirds of length of XVIII, is continuous laterally with long- itudinal prominence which occupies whole length of segment and extends laterally, beyond ¢; prominence more sharply defined medianly than laterally. Genital markings: pre- and post-setal paired pore-like markings faintly indicated on common transverse pad, on X. Paired pore-like markings close to midventrum immediately postsetal in XVI and XVII and prostomium wide dorsal setal interval (zz) pore 3 female pore ditellum, FIG. 9. Spenceriella hamiltoni (Fletcher, 1887). A, dorsal view of prostomial region; B, ventral view of fore- and mid-body of specimen 2. presetally and slightly more laterally, though anteromedial to setae a in XIX. Female pores paired on XIV, just anterior to setal arc, close together in a common glandular field. Spermathecal pores 3 pairs of small but sharply defined open slits, in 6/7, 7/8 and 8/9, in setal lines b. Several anterior septa thickened; 8/9-11/12 strongly. Dorsal blood vessel single, continuous onto pharynx; hearts in X-XII latero-oesophageal with chief origin from calciferous vessel, near origin of latter from supra-oesophageal vessel; commissurals of [X anterior originating from dorsal vessel only. Gizzard large, barrel-shaped in V, firmly muscular; extending to level of MEMOIRS OF THE QUEENSLAND MUSEUM vasa deferentia trilobed prostate duct 1mm FIG. 10. Spenceriella hamiltoni (Fletcher, 1887). Dorsal view of (immature) right prostate of specimen " 2. intersegment 8/9 posteriorly, septa 5/6 and 6/7 funnel-shaped around it. Oesophagus with 3 pairs of large almost spherical calciferous glands, in X, XI and XII, each broadly attached to oesophagus but separated by a deep trench middorsally. Intestinal origin XVI; acaecate; a low ridge but no true dorsal typhlosole present. Intestinal contents unusual in containing very little fine grit, but with sparse (siliceous?) grains and filled with dense masses of what appear to be strips of grass blades. Meronephric, nephridia commencing in II; forming large masses in II-VI but not united as tufts; all apparently exonephric; reducing to parietal but not numerous micromeronephridia by XII or XIII. A single median preseptal funnel (not multiple funnels) demonstrated on each side of nerve cord for caudal nephridia, confirming placement in Spenceriella; all exonephric. Sparse ovaries with visible oocytes and small funnels, in XIII; ovisacs not found. Holandric, seminal funnels in each of X and XI, lacking spermatozoal iridescence; racemose seminal vesicles in LX and XII, posterior pair tortuous, vermiform (incompletely mature). Prostates a pair of racemose glands, in X VILL, incompletely mature; divided from ental end of duct into three major, elongate lobes; vasa deferentia joining anterior lobe near its base; duct wide and fusiform. Penial setae absent. Spermathecae 3 pairs, small and immature, in VII, VIII and [IX with ovoid ampullae tapering to ducts each bearing near its ectal end a small polex-shaped diverticulum; common duct of ampulla and diverticulum scarcely developed. REMARKS. The clearly phytophagous nature of this species, demonstrated in the new material, correlates with the dark pigmentation of the body NSW AND ACT MEGASCOLECID OLIGOCHAETES as it presumably emerges above ground to remove portions of grass blades. This material conforms very closely to the description of Spenceriella (=Anisochaeta) chani (Blakemore, 2000), reported from Neville and Cowra, even to the multilobed prostates. However, there seems no valid reason to separate it, or chani, from the prior S. hamiltoni (Fletcher, 1887) which was collected from Cowra and Oberon by Easton, as reported in Jamieson (2000). These localities are respectively 51 km W and 59 km E of Neville. The type locality of S. hamiltoni, Guntawang, is about 150 km N of Neville. The ¢ pore on the right side in the putative syntype of S. hamiltoni ( Jamieson, 2000) is near a line, the disposition of genital markings, albeit absent in XVI, resembles that in S. chani and the prostates are again multilobed. Location of each of the small ¢ porophores, in the new material, median to a longitudinal prominence is not here considered a significant difference from hamiltoni and chani as in both of these there is a tendency to lateral enlargement of the prostate porophores. The possibility that S. hamiltoni is a junior synonym of S. austrina (Fletcher, 1886) remains to be investigated. Spenceriella macleayi (Fletcher, 1889) (Figs 11) Perichaeta macleayi Fletcher, 1889: 1556-1558. Perichaeta macleayi yar. a, b and c; Fletcher, 1890: 1004-1007. Megascolex macleayi (Fletcher); Beddard, 1895: 376, Spenceriella macleayi (Fletcher); Blakemore & Elton, 1994: 251-254, fig. 1; Jamieson, 2000: 1254-1258, fig. 41.43-41.47. Anisochaeta macleayi (Fletcher); Blakemore, 2000; 4. ? Anisochaeta filix Blakemore, 2000: 21-22, Fig. 11. NEW RECORDS: A.C.T., 35°10’S.149°.20°E., 'Bywong’, Sutton, ca 20 km NE of Canberra, in improved pasture, July 2000. 8 clitellate specimens. CSIRO sample ‘A, Dark head’. QMG218246-218253. 'Gold Creek’, native pasture just NE of Gungahlin, Canberra, J. Scown, 2 July 2000. CSIRO sample ‘C. Native 1’; 3 clitellate; 3 (unregistered) aclitellate specimens. QMG218254-218256. DESCRIPTION. Length of clitellate specimens from both localities 55-95 (mean 70) mm. Width (midclitellar) 3.8mm; segments 87 (A, illustrated specimen). Colour in ethanol dark purplish grey-brown dorsally and laterally, especially anterior to clitellum, setal areolae and ventrum pale. Prostomium epilobous 3/4, closed, wedge-shaped. Peristomium bisected ventrally. First dorsal pore 5/6. Setae per segment 22 in XII. Clitellum annular, XHI-X VII. Genital markings (constant in 8 A and 3 C clitellate specimens): a 597 Vi vil sp ermathecal Vill pore 1 IX spemathecal pore 2 xX genital | markings Xil Xill XIV female pores XV clitellum XVI genital XVII markings XVIII male pore L— genital XIX markings 1mm aK ap pee wae FIG. 11. Spenceriella macleayi (Fletcher, 1889). Ventral view of fore- and mid-body female pores Gitellum genital marking? FIG. 12. Spenceriella nevillensis sp. nov. Holotype. Ventral view of fore- and mid-body of holotype. MEMOIRS OF THE QUEENSLAND MUSEUM transverse pad on each of X and XI, extending laterally to about c lines and filling segment longitudinally, each with pair of pore like presetal circular markings in ab; a pair of presetal and postsetal circular papillae with pore-like centre in each of segments XVII and XIX, most anterior and posterior pair of four, slightly more median than other two and all slightly median of line of male pores; an additional, midventral, eye-like marking equatorial in XVIII between male porophores. Male pores in bc lines on small elliptical papillae which are situated on larger porophores which do not, however, reach anterior and posterior borders of XVIII. Female pores paired on XIV, presetally and about 1/3 aa apart, ina common glandular field. Spermathecal pores 2 minute pairs, in 7/8 and 8/9, in or slightly ventral of c lines. Dorsal blood vessel single, continuous onto pharynx. Last hearts in XII. Gizzard large, a posteriorly slightly tapering cylinder in V, moderately muscular. Prostates large, racemose bipartite glands (C) or tortuous racemose, in XVII to as far as XXI(A). Spermathecae 2 pairs, in VII and IX with large subspherical or elongate ampullae tapering to ducts each bearing near ectal end a large, clavate diverticulum (A, C). REMARKS. The combination of calciferous glands limited to XI and XII with the particular arrangement of genital markings is diagnostic of S. macleayi. Fletcher (1889) noted as a variant a midventral marking in XVIII, also seen here. Specimens from Manning River, NSW, identified as S. raymondiana Fletcher (1887) by Easton (unpubl.) with similar restriction of calciferous glands to XI and XU are referred below to a new species which differs, among other respects, from S. macleayi in having 3 pairs of spermathecae, They were tentatively described as S. raymondiana by Jamieson (2000). The only other species of Spenceriella with 2 pairs of calciferous glands is S. filix (Blakemore, 2000), collected by Easton from Tree Fern Valley, NSW. It appears probable, from its description, that it is conspecific with S. macleayi. S. macleayi is widespread in NSW, being known from Elizabeth Bay, Sydney (the type locality); Mt Wilson; Mt Lawson; Burrawang.; Mt Victoria; Raymond Terrace; Morpeth; Richmond; Queenscliff; Mt Tomah; Bunadoon, Moss Vale District; Upper Manning River; and Old Newington (details in Jamieson, 2000). NSW AND ACT MEGASCOLECID OLIGOCHAETES Spenceriella nevillensis sp. nov. (Figs 12, 13) MATERIAL EXAMINED. HOLOTYPE QMG218257. PARATYPES 1 & 2, QMG218258-218259. All from ca 5 km S of Neville (33°43°S.149°13’E.), near Blayney and Bathurst, NSW, from improved pasture (CSIRO sample ‘F’). Two weakly clitellate; one (excluded from type series) aclitellate. DESCRIPTION. Length (clitellate specimens) 80-115mm (H, P;). Width (forebody) 4mm. Segments 91-129. Form (as preserved) slender and elongate, cylindrical, tapering at each end, slightly club-shaped in forebody; clitellum forming a narrower cylinder. Segments weakly triannulate. Pigmentless buff in ethanol. Prostomium tanylobous, its lateral borders gradually converging posteriad but posteriorly so weakly defined that it appears epilobous. Peristomium bisected ventrally. First dorsal pore 5/6. Setae often difficult to discern, about 18-20 in XIL and caudally; in forebody aa about 2.6 ab but ab smaller than bc; zz in forebody less than twice adjacent intervals, not an appreciable break in setal circlet behind clitellum; setal rows mostly regular. Clitellum XHI-X VII (=5 segments); less developed in XIII; annular but interrupted ventrally, to approximately c lines, in XVI and XVII; intersegmental furrows and dorsal pores, except 13/14, obscured dorsally, setae clearly visible. Male pores minute but distinct, in a lines of XVIII, each on a very small transversely elliptical papilla which is bordered laterally by a longitudinal prominence which extends length of segment. Genital markings a weakly visible circular disc on right side only (H) or paired (P)) almost filling segment longitudinally and bearing a pore-like presetal marking in cac line. Posterior genital markings consisting of faintly visible midventral pads with raised rims in each of XX-XXIII, first 3 of them with suggestions of paired pore-like markings. A suggestion of ventral glandularity in XVI is probably not a true genital marking; a marking in XIX is also doubtful (H). Posterior genital markings in the paratypes are too poorly defined for certain determination of their distribution. Female pores paired on XIV, just anterior to setal arc, about one third aa apart, in a common glandular field. Spermathecal pores 2 pairs of large eye-like papillae, in 7/8 and 8/9, centred approximately in b lines. Several anterior septa moderately thickened; 8/9-10/11 strongest. Dorsal blood vessel con- tinuous onto pharynx; dorsoventral com- missurals in X-XII forming large hearts. Gizzard 599 spermathecal diverticulum spermathecal duct a fn f A Yn spermathecal ampulla vasa deferentia prostate uct FIG, 13. Spenceriella nevillensis sp. nov. Holotype. A, dorsal view of right spermatheca of IX; B, dorsal view of left prostate. in V, large, wider anteriorly than posteriorly, with muscular sheen but readily depressed; extending to level of intersegment 8/9 posteriorly, septa 5/6 and 6/7 funnel-shaped around it. Oesophagus with 4 pairs of large reniform calciferous glands, in X-XII] (holotype and paratype), a short duct from hilus connecting to dorsolateral aspect of oesophagus; each pair of glands supplied by bifurcation of supra-oesophageal vessel present anteriorly in segment but does not continue posteriorly of pair of glands and originates anteriorly from dorsal vessel. Intestinal origin XVI; acaecate; a well-developed dorsal typhlo- sole commencing in XVIII-XIX though there rudimentary. Meronephric with at least 3 pairs of tufted nephridia in buccopharyngeal region with thick anteriorly running (enteronephric?) ducts; parietal micromeronephridia, about as numerous as setae in clitellar region. Ovaries not visible (H) or small, bushy (P)). Holandric, a pair of free sperm masses and funnels (with negligible spermatozoal iridescence) seen in each of X and XI; large laterally extensive racemose seminal vesicles in IX and XII, posterior pair larger. Prostates S-shaped tubuloracemose; a stout muscular duct of moderate length arising from anterior limit where it is joined basally by combined vasa deferentia. Penial setae absent. 600 pore 1 genital marking genital marking male pore genital marking FIG. 14. Spenceriella manningi sp. nov. Holotype. A, prostomium; B, ventral view of fore- and mid-body. Spermathecae 2 pairs, in VIII and IX with large ovoid ampullae tapering to ducts each bearing near its ectal end a clavate diverticulum whichis a little shorter than ampulla; common duct of ampulla and diverticulum scarcely developed. ETYMOLOGY. From near Neville. REMARKS. Two pairs of spermathecae and 4 pairs of calciferous glands, as in S. nevillensis, are also seen in S. monticola (Fletcher, 1887) and S. calpetana (Blakemore, 2000). S. monticola differs from S. nevillensis in being larger, pig- mented reddish brown, with more setae (16-50); anda different configuration of genital markings. MEMOIRS OF THE QUEENSLAND MUSEUM S. calpetana differs, among other respects, in having about 60 setae per segment, a markedly different configuration of genital markings, seminal vesicles in XI and XII; and no typhlosole. Spenceriella manningi sp. nov. (Fig. 14,15) Spenceriella raymondiana (part.) Jamieson, 2000; 1293-1297, fig. 41.70, 41.71. MATERIAL. HOLOTYPE AM W197644 from 31°49°S.151°56’E., Upper Manning River, NSW, coll. and ident. as S. rayvmondiana by E. Easton, 1983. DESCRIPTION. Length 165mm. Width (mid- clitellar) 71mm. Segments 117. Prostomium epilobous, almost tanylobous, with wide, deep lateral grooves, slightly convergent posteriad. Prostomium bisected ventrally. First dorsal pore 4/5. Setae per segment: in XII 26; caudally 32. In XID aa: ab: zz = 14: 4.5: 6.0; dorsal and ventral breaks clearly visible preclitellar; not apparent behind clitellum but aa becoming wide caudally; no evident irregularity. Clitellum annular, limits indistinct but dorsally from setal zone of XIII to posterior XVII; dorsal pores of 14/15-16/17 occluded. Male pores in a lines on small rounded papillae surrounded by dark glandular border with narrow connection across ventral midline with that of other side. Genital markings a pair of small pore-like markings presetally in bc and post-setally in 6 lines of X; 2 ‘pores’ of a side lying on a common raised glandular pad which is well defined laterally but not medianly. A pair of pore-like markings immediately presetal and median to a lines of XVII, each surrounded by a dark glandular border which is broadly confluent with that of other side. Similar markings in XIX with a narrower connecting zone. Female pores shortly presetal, well median of a lines, in a common dark oval field with a narrow, tumid, pale margin. Spermathecal pores 3 pairs of min- ute orifices apparent only on opening up inter- segmental grooves, in 6/7, 7/8 and 8/9, shortly below 6 lines. Septa 7/8-13/14, strong. Dorsal blood vessel single, continuous onto pharynx; last hearts in XII; those in XI and XII, only, latero- oesophageal. Supra-oesophageal recognizable in XI and XII. Gizzard in V, broad, cylindrical and strongly muscular, preceded by a short, almost equally broad proventriculus. Oesophagus with a pair of large, extramural calciferous glands, each with a narrow posterior connection to gut, in each of segments XI and XI]; glands with numerous NSW AND ACT MEGASCOLECID OLIGOCHAETES A vasa deferentia prostate duct imm spermathecal duct spermathecal diverticulum spermathecal ampulla 1mm FIG. 15. Spenceriella manningi sp. nov. Holotype. A, right prostate; B, right spermatheca of IX. narrow internal longitudinal septa. Intestine commencing XVI but pushing septum 15/16 close to 14/15; no definite typhlosole seen. Nephridia tufted in III and I'V and, less compact, in V. Transverse bands of apparently astomate meronephridia posterior in succeeding segments of forebody. Caudally with dense bands of meronephridia filling segments; no preseptal funnels seen; some intrasegmental funnels tentatively identified. Large sperm masses in X and XI invest iridescent sperm funnels which lie posteriorly in each segment; the masses possibly enclosed in thin, membranous testis-sacs. Seminal vesicles large, racemose, in [X and XII. Ovaries webs of numerous large oocytes in XII. Large morula-like ovisacs on anterior walls of XIV. Prostates appearing to be compactly tubuloracemose but not resolvable into distinct tubes, therefore racemose; restricted to XVIII, each with a short, muscular duct which is joined, near its junction with gland, by a thick vas deferens. Spermathecae 3 pairs, each with somewhat flattened subspheroidal ampulla, narrowing with no distinct duct to body wall and joined ectally by clavate uniloculate 601 diverticulum of about equal length (1.4mm), with spermatozoal iridescence. ETYMOLOGY. From the type locality. REMARKS. Jamieson (2000) stated that in the absence of type material, it was uncertain that the material identified by Easton as S. raymondiana, including that from the Upper Manning River (W197644 ex 1515), was referable to this species, in which it was provisionally placed. The Manning River material is here distinguished as S. manningi. The remaining Easton material is not available for examination. Only S. macleayi Fletcher, 1889, and the probable junior synonym of the latter, 4. filix Blakemore, 2000, resemble S. manningi in having calciferous glands restricted to 2 pairs, in all 3 taxa being in segments XI and XIl. S. manningi differs in having 3 pairs of sperm- athecal pores, the other 2 taxa having 2 pairs. The genital field of S. manningi differs in important respects from that of S. macleayi. The posterior genital field in the Manning River material is also similar to that of S. jenolan- ensis, and may indicate close relationship though that species differs in its 4 pairs of spermathecae and 4 pairs of calciferous glands, in X-XIII. Molecular studies are required to aid elucidation of the phylogenetic relationships of these and other Spenceriellas. ACKNOWLEDGEMENTS The author is grateful to Dr. Geoffrey Baker, CSIRO, Entomology, Canberra, for providing the CSIRO material described in this paper. Dr. Pat Hutchings and Dr. Penny Barents are thanked for providing facilities in the Australian Museum. All illustrations (by camera lucida) are by the author. LITERATURE CITED BEDDARD, F.E. 1890. Observations upon an American species of Perichaeta, and upon some other members of the genus. Proceedings of the Zoological Society of London 1890: 52-69. BEDDARD, F.E. 1895. A Monograph of the Order of Oligochaeta. Clarendon Press, Oxford. BLAKEMORE, R.J. 2000. New species of the earthworm genus Anisochaeta (Oligochaeta: Megascolecidae) from New South Wales. Records of the Australian Museum 52: 1-40, BLAKEMORE, R.J. & ELTON, K.L. 1994. A hundred-year-old worm? Australian Zoologist 29: 251-254. FLETCHER, J.J. 1886a. Notes on Australian Earthworms-Part I. Proceedings of the Linnean Society of New South Wales 1(2); 523-574. 1886b. Notes on Australian Earthworms-Part II. Proceedings of the Linnean Society of New South Wales 1(2): 943-973. 1887. Notes on Australian earthworms-Part III. Proceedings of the Linnean Society of New South Wales 2(2): 375-402. 1889, Notes on Australian earthworms-Part V. Proceedings of the Linnean Society of New South Wales 3(2): 1521-1558. 1890. notes on Australian earthworms-Part VI. Proceedings of the Linnean Society of New South Wales 4(2): 987-1019. JAMIESON, B.G.M. 1973. Earthworms (Megascolecidae: Oligochaeta) from Mount MEMOIRS OF THE QUEENSLAND MUSEUM Kosciusko, Australia. Records of the Australian Museum 28: 215-252. 2000. Native Earthworms of Australia (Megascolecidae, Megascolecinae). (Science Publishers Inc.: Enfield, New Hamsphire). (CD ROM). KIKKAWA, J. & PEARSE, K. 1969. Geographical distribution of land birds in Australia — a numerical analysis. Australian Journal of Zoology 17: 821-840. MICHAELSEN, W, 1907. Oligochaeta. Pp117-232. In W. Michaelsen & R. Hartmeyer (eds) Die Fauna Siidwest-Australiens (Gustav Fischer: Jena). SPENCER, W. B. 1900. Further descriptions of Australian earthworms, Part I. Proceedings of the Royal Society of Victoria new series 13: 29-67. STATUS OF ESTUARINE CROCODILES IN THE POPULATED COAST OF NORTHEAST QUEENSLAND CHRISTOPHER P. KOFRON AND RUSTY SMITH Kofron, C.P. & Smith, R. 2001 06 30: Status of Estuarine Crocodiles in the populated coast of northeast Queensland. Memoirs of the Queensland Museum 46(2): 603-610. Brisbane. ISSN 0079-8835. Tully to Cooktown encompasses 325km of coastline in northeast Queensland. During the past 50 years this area has undergone tremendous urban, rural residential and agricultural development. North of Cooktown human population density is low. Since 1990, Estuarine Crocodiles (Crocodvlus porosus) have attacked six people in Queensland resulting in one death and five serious injuries. Two attacks occurred in the study area in Cairns during 1997 and 1998. Consequently public perception is that numbers of crocodiles have increased greatly since cessation of commercial hunting in 1974 and elimination of removal zones around cities and towns in 1991, and public concern for human safety from crocodile attack is high, We surveyed the major waterways between Tully and Cooktown for Estuarine Crocodiles from June 1996 to May 1998: Hull R, Maria Ck, Moresby R, Johnstone R, Russell/Mulgrave R, Trinity Inlet, Barron R, Daintree R, Annan R and Endeavour R. These waterways comprise most of the habitat occupied by Estuarine Crocodiles between Tully and Cooktown. We surveyed 346km of waterway, sighting 146 crocodiles at densities 0.11/km to 1.00/km. Overall density was 0.34/km, which relative to most waterways in Cape York Peninsula is low. Contrary to public perception, the crocodile population between Tully and Cooktown is of low density. Certain human activities such as urban, rural residential and agricultural development, clearing of riparian vegetation, disturbance by motor boats, commercial gill netting in estuaries, and removal of crocodiles appear to be keeping crocodile numbers low in this area. 0 Crocodiles, north Queensland, estuarine. Christopher P. Kofron Queensland Parks and Wildlife Service, Northern Regional Centre, PO Box 2066, Cairns 4870, (e-mail: Chris.Kofron@env.qld.gov.au); Rusty Smith Queensland Parks and Wildlife Service, Innisfail District Office, PO Box 44, Innisfail 4860, (e-mail: Rusty.Smith@env.qld.gov.au), Australia; 24 May 2001, The Estuarine Crocodile, Crocodylus porous, is the world’s largest living crocodile, with total lengths up to 9m (Groombridge, 1987). Its large size and predatory habits cause special management problems because most people do not want to live or recreate near large, dangerous, predatory wildlife. The geographic range is greater than for any other crocodile, extending from Vanuatu and Solomon Islands in the east, across northern Australia, New Guinea, Indonesia, Borneo, Philippine Islands and Southeast Asia to eastern India and Sri Lanka, Despite this enormous geographic range, its populations in most countries are very severely depleted with continuing decline (Groombridge, 1987). Prospects for survival appear to be poor except in Indonesia, Papua New Guinea and Australia (Ross, 1998). The geographic range in Queensland extends from Fitzroy R system (23°27’S), along the eastern coast to islands of Torres Strait, along the western coast of Cape York Peninsula, to the Northern Territory. In Queensland, Estuarine Crocodiles inhabit coastal waterways, and freshwater sections of rivers where they may travel several hundred kilometres inland (Taplin, 1987). Estuarine crocodiles are listed as ‘vulnerable’, if threatening processes continue (Department of Environment and Heritage, 1994). Since 1990 Estuarine Crocodiles have attacked 6 people in Queensland resulting in 1 death and 5 serious injuries. Two recent attacks occurred in Cairns. On 28 November 1997, a man swimming at Yorkey’s Knob Beach was attacked by a 2.6m crocodile, suffering puncture wounds on head, shoulder and back. On 6 February 1998, a teenage girl swimming in a mangrove-lined drain of Chinaman’s Ck was attacked by a 3.1m crocodile, suffering deep puncture wounds in both legs and a broken ankle. Public perception is that crocodile numbers have increased since cessation, in 1974, of commercial hunting, and elimination, in 1991, of removal zones around cities and towns. Consequently public concern for human safety from crocodile attack is 604 especially high. The purpose of this study, therefore, was to determine the current status of Estuarine Crocodiles in the populated coast of northeast Queensland. Some rivers between Tully and Cooktown were surveyed previously by Queensland National Parks and Wildlife Service from 1984-1986 (Taplin, 1989) and in 1991 (Kreiger & Fell, 1991). Although no data exist on crocodile numbers prior to commercial hunting in Queensland, crocodile numbers were apparently reduced to low levels relative to pre-hunting abundance (Taplin, 1987). It seems probable that the combination of commercial hunting and removal zones in late 1980s to early 1990s reduced crocodile numbers to very low levels within the study area. However, some people argue that these tidal waterways, mostly short and with headwaters in mountains to the west, were never a major crocodile breeding area and never contained high numbers, cooler water temperatures being a determining factor. STUDY AREA Cairns (16°55°S) is 119km N of Tully (17° 56’°S) and 169km S of Cooktown (15°28’S) on the eastern coastal plain in the Tropical Rainforest Bioregion. Rivers on the NE coast between Tully and Cooktown are not extensive. Tully to Cooktown encompasses 325km of coast, and here the coastal plain does not extend beyond 30km inland where it is bounded by mountain ranges, including Bellenden Ker range with Queensland’s highest peak (Bartle Frere 1,622m). During the past 50 years the coastal plain between Tully and Cooktown has undergone major urban, rural residential and agricultural development. Total human population between Tully and Cooktown is 168,855 (Cook et al., 1997), 76% (128,022 people) living in Cairns (Centre for Applied Economic Research and Analysis, 1997). In addition to Tully, Cairns and Cooktown, other human population centres are Mission Beach/Innisfail (Johnstone Shire) and Port Douglas/Mossman areas (Douglas Shire). North of Cooktown, population density is low and centres are sparse, There are 10 major waterways between Tully and Cooktown (south to north): Hull R, Maria Ck, Moresby R, Johnstone R, Russell/Mulgrave R, Trinity Inlet, Barron R, Daintree R, Annan R and Endeavour R. Only one species of crocodile (C. porosus) inhabits this area, and tidal MEMOIRS OF THE QUEENSLAND MUSEUM waterways comprise most of its habitat. Six flow through towns or cities: Hull R (Tully); Johnstone R (Innisfail); Russell/Mulgrave R, Trinity Inlet, Barron R (Cairns); and Endeavour R (Cooktown). METHODS We surveyed the major waterways between Tully and Cooktown for Estuarine Crocodiles from June 1996 to May 1998. Surveys were conducted by spotlighting from a small motorboat at night within 3 hours each side of low tide. Tidal reaches of waterways were surveyed, including adjacent freshwater sections passable by small motorboat. Total length ofeach crocodile was estimated and recorded in size classes: 0,2-0.4m (hatchling/yearling), 0.5-1.0m, 1.1-2.0m, 2.1-3.0m, 3.1-4.0m and ‘eyes only’ (crocodile sighted only by eye shine). Locations of crocodiles were determined by global positioning systems and topographic maps. We compare our data to surveys by Taplin (1987) and Kreiger & Fell (1991). However, we are unable to make statistical comparison. Surveys were conducted to international standards (Bayliss, 1987; Kofron, 1992) with recognition of inherent weaknesses (Webb & Smith, 1987). For example, wariness in crocodiles is a function of size (Webb & Messel, 1979), older and larger crocodiles being more ‘wary’ and thus difficult to detect. However, we surveyed tidal rivers, which relative to other aquatic habitats (e.g. vegetated swamps) are easy’ to survey. In such areas a higher proportion of total number of crocodiles present 1s seen (Webb et al., 1987). Following convention, we define density as number of crocodiles observed per kilometre of waterway, excluding hatchlings and yearlings. Distance of a waterway from Cairns is measured from the central business district for this study. RESULTS AND DISCUSSION We sighted 146 crocodiles in 346km of waterway (Table |): 0.2-0.4m (7 = 29), 0.5-1.0m (23), L.1-2.0m (41), 2.1-3.0m (19), 3.1-4.0m (10) and eyes only (24). Crocodile densities in river systems were 0.11/km to 1.00/km. Overall crocodile density in waterways between Tully and Cooktown was 0.34/km. HULL RIVER. Hull R system is east of Tully with a catchment area of 12,996ha (Russell & Hales, 1997), originating in Walter Hill Range to the west (highest peak 1,068m) and draining a ESTUARINE CROCODILES IN NORTHEAST QUEENSLAND TABLE 1. Results of spotlight surveys of Estuarine Crocodiles (Crocodylus porosus) in major waterways between Tully and Cooktown from June 1996 to May 1998. Density is number of crocodiles sighted per kilometre of waterway, excluding hatchlings and yearlings (0.2-0.4m total length). [_—- — = | fr Estimated Lengths | ihe: Surveyed | “Gross |_0.2-0.4m [ Q5-1.0m | 1.1-2.0m | 21-3.0m | 3.1-4.0m | YoY | Density ‘Hull River | 38 2a | 2 | o | 3 [| 3 | 3 1 | 024 Maria Creek _ 4 2. oO 1 | 1a | o | o | 0 | on | | Moresby River 44 | 6 0 0 he oO 0 3 0. 14 Johnstone River a 7 | 3 | 3 0 o | 9 0.15 /Russell/MulgraveR | 63_ 9 2 2. 4. L 0 0.11 | Trinity Inlet 40 2 | oo 4 10 3 0 es 0.50 Barron River 22 7 | 07 Oo | 2 [ 3 2h 0 0.32 Daintree River 25 21 6 8 4) _0 0 3 0.60 Annan River | 12 12 oO 4 2 a [oe [3 1.00 Endeavour River 44 41 8 4 | 9 6 3 i 0.75 ; _ Total 346 146 29 3 | 41 19 10, 24 0.34 narrow coastal plain. Hull R and North Hull R are the major waterways, converging 2km from the coast into an estuary. Hull R flows through Hull R National Park (3,070ha) immediately adjacent to the coast. The catchment is 76% forested, almost all in Wet Tropics World Heritage Area, and 14% under agriculture (sugarcane). Mangrove (1,326 ha), other swamp forest (8lha, predominantly Melaleuca ) and rainforest comprise the existing natural vegetation. Although the only urban centre is the township Hull Heads (Russell & Hales, 1997), the Hull R system is heavily used for recreational boating and fishing. Commercial gill netting occurs at low to moderate level. We surveyed 38km on 5-7 April 1997, sighting 21 crocodiles at 0.24/km: 0.2-0.4m (n = 12), 1.1-2.0m (3), 2.1-3.0m (3), 3.1-4.0m (2) and eyes only (1). Two nests were sighted. MARIA CREEK. Maria Ck system is a group of small coastal streams 100km S of Cairns originating in Walter Hill Range to the west and draining a narrow coastal plain. It flows through Maria Ck National Park (749ha) immediately adjacent to the coast and in Wet Tropics World Heritage Area. Catchment area is 24,642ha, 49% cleared predominantly for agriculture (sugarcane; Russell & Hales, 1997). The main waterway is 25km long with a relatively large estuary, sheltered and lengthened by a sandspit extending south from Kurrimine Beach. Half of the national park is mangrove (350ha), the remainder other swamp forest and rainforest (Australian Nature Conservation Agency, 1996). Maria Ck National Park is an important wetland in Australia. Maria Ck system is heavily used for recreational boating and fishing, and commercial gill netting occurs at low level. High levels of nutrients and sediments in runoff from adjacent agricultural lands may be impacting on the wetlands (Australian Nature Conservation Agency, 1996). Catchment area contains E| Arish, Mission Beach and Kurrimine townships. We surveyed 17km on 4-5 April 1997, sighting two crocodiles: 0.5-1.0m (7 = 1) and 1.1-2.0m (1). Density was 0.12/km. MORESBY RIVER. Moresby R system is 80km south of Cairns, a wetland 24km long and 12km wide, mostly in Wet Tropics World Heritage Area. Catchment area is 126km’, 48% remaining as natural vegetation, and with a population of 300 (Eyre & Davies, 1996). Moresby R communicates with the Coral Sea through a restricted passage in Moresby Range, with a broad sheltered estuary west of the range. The southern system is bounded to the east by a series of sand dunes. The tidal reach of Moresby R is 20km long with headwaters in Basilisk Range, a low range in the western coastal lowlands. Natural vegetation is mangrove, other swamp forest and rainforest, and the estuary contains significant seagrass beds. Moresby R system is an important wetland in Australia (Australian Nature Conservation Agency, 1996). Coastal lowlands north, south and west of Moresby R system have undergone extensive clearing for agriculture (sugarcane). Mourilyan Harbour, at northern end of the estuary and sheltered by Moresby Range, loads sugar onto UG sea-20ing vessels. Moresby R is heavily used for recreational fishing and boating, and commercial gill netting occurs al low level We surveyed 44kmon 31 January 104 February 1997, sighting. six crocodiles: 1,1-2.0m (a2 = 3) and eyes only (3). Density was 0.14/km. JOHNSTONE RIVER. North Johnstone R (125km) and South Johnstone R (9Okin) are the main waterways of Johnstone R system 64k south of Caims, originating on Atherton Tableland and Howing through vast rainforest on the eastern escarpmem of Bellenden Ker Range. Catchment area is 163,050ha (population 17.860: Ryre & Davies, 1996): 34% rainforest, predominantly on slopes; 38% under agriculture (cattle, sugar cane, bananas). rural residential and urban uses, predominantly in coastal lowlands and Atherton Tableland; and 8% mangrove and other swamp forest (Pitts, 1993), Along the lower river system much clearing has occurred to very edge of the stream bank. North Johnstone R is 43km long in the coastal plain, and South Johnstone R 3tkm, The two rivers converge at Innisfail forming Johnstone R estuary Skm long. Johnstone R Hows through centre af Innisfail, population 8,987 (Cook et al., 1997), Innisfail is home fora commercial fishing eet, and the river is also heavily used for recreational boating and while-water rafing. Johnstone R has. been closed to commercial gill netting for 30-40 years, We surveyed 4}km on 10-12 January 1997, sighting 7 crocodiles at 0.) 5/lune 0.2-0,4i (a = 1);.0,5-1.0m (3); 1.1-2.0(3). Johnstone R system (1 [kni) was surveyed previously in 1986 (Taplin, 1989),.6 crocodiles sighted at 0.55/km: 0.5-1.8m (m2), 1.9-3-0m (3) and eves only (1). Densities are low and show no increase, RUSSELL/MULGRAVE RIVERS. Russell R and Mulgrave R flow through agricultural and rural residential lands of south Cairns, headwaters in mountains to the west. The lower river system is in a narrow plain between Bel- lenden Ker Range and Ciraham Range/Malbon Thompson Range to the east coustally, Russell R flows northeast and Mulgrave R southeast. The two rivers converge [ki from the sea to form Mutchero Inlet, a constricted passage throuwzh submerged parts of Graham Range {o the south and Malbon ‘Thompson Range to the north. Catchment area of Russell R is 57,000ha, and Mulgrave R 145,000ba (Australian Nature Conservation Agency, 1996), MEMOIRS OF THE QUEENSLAND MUSLUM Surrounding natural vegetation 18 mangrove, other swamp forest and raintorest, although most floodplain in coastal lowland has been cleared. Forty-eight percent of the catchment area remains in natural state and is Wet Tropics World Heritage Area; and 27% is under agriculture (sugarcane). Russell R flows through Russell R National Park (4,1 00ha), an important wetland in Australia (Australian Nature Conservation Agency, 1996), Tidal scetions of the Russell/ Mulgrave Ro are heavily used lor recreational fishing and boating, and adjacent freshwater seclions for swimming and rafting. The rivers have been closed to commercial gill netting for more thaty 10 years, We surveyed 63km (Russell R S38km, Mulgrave R 25km) 18-21 January 1997. sighting 9 crocodiles at 0.11/km: 0.2-0.4m (7 = 2), 0.5-1.0m (2), 1. 1-2,0n (4) and 3, 1-4.0m (1). One problem crocodile (1.9 m) was removed trom Mulgrave R during the study because it was in a freshwater swimming area. Russell/Mulgrave R (27kin) Were suryeyed previously in [991] (Kreiger & Fell, 1991), seven crocodiles sighted at 0.26/km: 0.5-1.0m (7 = 1), 1.1-1.8m (2) and eyes only (4). Also, 24km were surveyed in 1984 (Taplin, 1989), 2 crocodiles sighted at 0,04/kin; ).2-U.4m (7 =L) and eyes only (1). Densities are consistently low and show no increase. TRINITY INLET. Trinity Inlet is 18km long by 6km wide in north-south direction, comprising 6,475ha: mangrove 3,600ha, seagrass 1,000ha, open water 940ha, Udal fats 900ha, salt Mats 20ha, freshwater swamp [Oha and sand ridge Sha. Cutchment area ts 30,000ha. ‘Trinity Inletas a blind estuary, previously the mouth of Mulgrave R whose communication with the sca has shifted south (Australian Nature Conservation Ageney, 1996), Caims surrounds Trinity Inlet: Cams Port ataits mouth; urban development and industry on its western side; and rural residential and agriculture om its southern and eastern sides, Most of the previously fringing freshwater swamp forest has been cleared. The system is heavily used for recreational fishing and boating, and also by large sea-going vessels; commercial gill netting occurs at extremely high level, Despite proximity to a major population centre, the ecosystem is intact and overall water quality good (Australian Nature Conseryation Agency, 1996). We surveyed 40kim on 7-9 and 20 May 1998. sighting 20 crocodiles at 0.50/km: 0.5-1.0m (= A), 1,-2.0 (10), 21-3,0 (3) and eyes only (3), ESTUARINE CROCODILES IN NORTHEAST QUEENSLAND Also, we previously surveyed 40km of Trinity Inlet on 18 April and 19 May 1997, sighting 13 crocodiles at 0.33/km: 0.5-1.0m (7=2), 1.1-2.0m (4), 2.1-3.0m (5), 3.1-4.0m (1) and eyes only (1). Although we sighted no hatchlings, Keith Cook (Cairns Crocodile Farm, pers. comm) observed one in 1997. In addition, courtesy Mark Read (Queensland Parks and Wildlife Service), we searched for nests by helicopter on 21 February 1998 but sighted none. Two problem crocodiles were removed during the study: one (3.1m) from Chinaman’s Ck (February 1998) that attacked a teenage girl, and a second (1.5m) from the same drain in April. Trinity Inlet (55km) was surveyed previously in 1991] (Kreiger & Fell, 1991), seven crocodiles sighted at 0.13/km: 0.5-1.0m (7 = 1), 1.1-1.8m (2) and eyes only (4). Densities are consistently low and show no increase. BARRON RIVER. Barron R is one of the longest easterly-flowing rivers (165km) in Cape York Peninsula, with headwaters in Atherton Tableland at 1234m elevation (North Queensland Joint Board, 1997). It breaches Macalister/Lamb Ranges through a major gorge (6km length) then forms a delta (50km? ) in the narrow coastal plain (8km wide). Barron R flows through Cairns just north of the urban centre, Machan’s Beach community at its mouth. Catchment is 200,000ha, population 42,000. Natural vegetation along lower Barron R is mangrove, other swamp forest and rainforest, most cleared for sugarcane and residential land use. A significant area of mangrove (6km*) remains at the mouth. Queensland Government (Department of Primary Industries, 1993) is concerned about catchment condition, particularly erosion, weeds, urban expansion, nutrient enrichment, impacts on wildlife and loss of wetlands. Tidal section of the river is heavily used for recreational fishing and boating, and adjacent freshwater section for swimming and rafting. Commercial gill netting occurs at low level. We surveyed 22km (tidal reaches) on 4-6 May 1998, sighting seven crocodiles at 0.32/km: 1.1-2.0m (7 = 2), 2.1-3.0m (3) and 3.1-4.0m (2). Also, we surveyed the same tidal reaches previously 28-29 June 1996, sighting seven crocodiles at 0.32/km: 0.5-1.0m (#7 =1), 1.1-2.0m (1) and 2.1-3.0m (5). Densities are consistently low. Two problem crocodiles (2.6m each) were removed during the study, one from Richters Ck (29 January 1997) frequenting swimming beaches and one from Thomatis Ck (9 December 607 1997) that attacked a swimmer at Yorkey’s Knob Beach. DAINTREE RIVER. Daintree R (120km length) is 78km northwest of Cairns, headwaters in Great Dividing Range. Catchment is 2125km*, 90% in Wet Tropics World Heritage Area and most protected in Daintree National Park and Dagmar Range National Park, population 1000. Rainforest comprises 74% of catchment, originally 90%. The estuary is 5km long, tidal influence extending 24km upstream. The lower 13km of Daintree R, up to 10km wide with mangrove and other swamp forest, is an important wetland (Australian Nature Conservation Agency, 1996; Eyre & Davies, 1996). Daintree R in the coastal lowlands occupies a narrow valley between Thornton Range to the north and Dagmar Range to the south. One-half (16,400ha) of this floodplain is cleared for sugarcane (12,000ha) and cattle (4,400ha), the other half (17,300ha) remaining as mangrove, other swamp forest and rainforest (Burrows, 1998). Mangrove is most extensive at river mouth to 6km upstream. Daintree R is heavily used for recreational boating, and also commercial boating for wildlife tours. The river was closed to commercial gill netting about 12 years ago. We surveyed 25km (starting 7km above river mouth at ferry crossing) on 5 September 1997, sighting 21 crocodiles: 0.2-0.4m (# = 6), 0.5-1.0m (8), 1.1-2.0m (4) and eyes only (3). Density was 0.60/km, however fog made visibility poor. One crocodile (2.4m) that attacked a dog was removed during the study (8 May 1997). Daintree R (31km) was surveyed previously by Kreiger & Fell (1991), 27 crocodiles sighted at 0.85/km: 0.2-0.4m (7 = 1), 0.5-1.0m (17), 1.1-1.8m (3), 1.8-4.0m (3) and eyes only (3). Also, Taplin (1989) surveyed 21km in 1984, sighting 16 crocodiles at 0.75/km: 0.5-1.8m (7 = 9), 1.9-3.0m (2) and eyes only (5). Densities are consistently low and show no increase, Also in Douglas Shire during the study, | problem crocodile (1.8m) was removed at Wonga Beach (7km south of Daintree R); 2 (3.4m each) from Mossman R (17km south of Daintree R; 29 January and 23 March 1998) that took dogs; and | (2.1m) from Bloomfield R (41km north of Daintree R, 10 February 1998). ANNAN RIVER. Annan R is 6km south of Cooktown and is least disturbed of the major 608 waterways. Catchment is 750km*, with population 300 (Eyre & Davies, 1996). Ninety percent of catchment retains natural vegetation (40% rainforest, predominantly upper half; 40% dry Eucalyptus woodland, predominantly lower half), and 10% cleared for cattle. The tidal section is bounded immediately by mangrove, other swamp forest and rainforest. The estuary is 6km length. Recreational boating occurs at low to moderate level in the tidal section, and commercial gill netting at moderate level. We surveyed 12km (tidal reaches) on 9-10 October 1997, sighting 12 crocodiles at 1.00/km: 0).5-1.0 (7= 1), 1.1-2.0(2), 2.1-3.0 (4), 3.1-4.0 (2) and eyes only (3). Annan R (1 1km) was surveyed previously by Kreiger & Fell (1991), sighting six crocodiles at 0.55/km: 1.9-4.0m (7 = 1) and eyes only (5). Densities are low but may indicate a slight increase. ENDEAVOUR RIVER. Endeavour R system has two major channels: Endeavour R flowing west to east, and Endeavour R Right Branch north to south, Cooktown is on the southern banks of Endeavour R at its mouth, population 1411 (Cook et al., 1997). Cooktown Mclvor River Rd parallels Endeavour R for 30km near Cooktown, and adjacent land is under rural residential use and agriculture (peanuts, corn, bananas); however only little clearing has encroached to the banks of Endeavour R. Vegetation along Endeavour R Right Branch remains predom- inantly natural. The estuary is 5km long, its upper half in Endeavour R National Park (2,170ha). Endeavour R is 500m wide at its mouth, narrowing to 100m at junction with Endeavour R Right Branch 11km upriver. Vegetation in the tidal area is mangrove; along freshwater sections of the river, rainforest; and along freshwater creeks, Melaleuca swamp. Recreational boating occurs at moderate level, The river was closed to commercial gill netting about 30 years ago. We surveyed 44km on 14-16 September 1997, sighting 41 crocodiles at 0.75/km: 0.2-0.4m (n= 8), 0.5-1.0m (4), 1.1-2.0m (9), 2.1-3.0m (6), 3.1-4.0m (3) and eyes only (11). Two crocodiles (3.7m, 3.8 m) frequenting Cooktown wharf were removed during the study. Endeavour R system (28km) was surveyed previously by Kreiger & Fell (1991), sighting 28 crocodiles at 0.96/km: 0.2-0.4m (7 = 5), 0.5-1.0m (16), 1.1-1.8m (1), 1.9-4.0m (2) and eyes only (4). Densities are consistently low and show no increase. MEMOIRS OF THE QUEENSLAND MUSEUM MANAGEMENT CONSIDERATIONS Densities of Estuarine Crocodiles in waterways between Tully and Cooktown range from 0.11/km to 1.00/km. In total, we surveyed 346km of waterway between Tully and Cooktown, sighting 146 crocodiles at overall density 0.34/km. Previously Kreiger & Fell (1991) surveyed five waterways, sighting crocodiles at overall density 0.45/km. In these five waterways we sighted crocodiles at density 0.47/km. Also, Taplin (1989) surveyed 3 waterways in 1984-1986, sighting crocodiles at overall density 0.41/km. In these 3 waterways we sighted crocodiles at overall density 0.22/km. The densities are consistently low and show no increase. Overall crocodile density between Tully and Cooktown (0,34/km) is low relative to most waterways on Cape York Peninsula: 0.7/km, rivers in Lakefield National Park (165km northwest of Cooktown; Read & Miller, 1998); 5.9/km and 10.5/km, Wenlock R and Tentpole Ck (northwest tip of Cape York Peninsula), respectively (Read, 1998); and 0.3/km to 3.6/km, other rivers on western Cape York Peninsula (Read, 1998). In the Northern Territory, Estuarine Crocodile populations are recovering, with mean annual rate of increase 2-3% reported in several rivers (Webb et al., 1987) and overall annual rate of population increase 8% (Bayliss, 1987). Estuarine Crocodiles nest in the wet season, and flooding is a major threat to egg survival. In the Northern Territory, mean egg survivorship is 25%, inundation of nests the major cause of mortality (Webb et al., 1987). In some areas flooding accounted for 100% egg mortality. No effort was made to locate nests except for one helicopter search of Trinity Inlet, but none was seen. However, 2 nests were observed near the Hull R, and | near the Bloomfield R subsequently inundated (Tony Frisby, Queensland Parks and Wildlife Service, pers. comm.). In addition, hatchlings/yearlings were seen in several rivers: Hull R (12), Johnstone R (1), Russell/Mulgrave R (2), Trinity Inlet (1, Keith Cook, Cairns Crocodile Farm, pers. comm.), Daintree R (6) and Endeavour R (8). There was no evidence of nesting in Maria Ck, Moresby R, Barron R, Mossman R or Annan R, In total, 30 hatchlings/yearlings were observed in 346km of waterway, which is a low number. Crocodile densities in tidal rivers are partly a function of proximity to successful nesting areas. ESTUARINE CROCODILES IN NORTHEAST QUEENSLAND For example in the Northern Territory, Webb et al. (1987) observed crocodiles at density 3.2/km in tidal rivers with successful nesting areas, and 0.7/km without. We attribute the low number of hatchlings/yearlings between Tully and Cooktown to negative impacts of human activities, largely preventing successful nesting. Extensive deforestation and development in the catchment of some rivers probably effect greater rises in water levels than occurred previously, partially accounting for the paucity of successful nesting. In addition, many river banks previously available for nesting are now cleared to water’s edge, and disturbance by motor boats may also discourage nesting. The human population and agricultural development were identified previously as causing significant degradation of crocodile habitat between Tully and Cooktown (Taplin, 1987). Further, Taplin (1987) referred to this crocodile population as depleted’, with only small numbers at low densities. There are 16 protected areas with crocodile habitat between Tully and Cooktown. They range from 6ha to 76,000ha. Daintree National Park is the largest, however it encompasses predominantly Great Dividing Range and lowland rainforest, with relatively little crocodile habitat. The next 3 largest are Cedar Bay National Park (5,650ha), Russell R National Park (4,100ha) and Ella Bay National Park (3,71 0ha). The fifth largest is Hull R National Park (3,070ha) and almost all crocodile habitat (waterway, mangrove, swamp forest). Crocodiles are at low denisty here too, 0.24/km. Estuarine Crocodiles are highly mobile, and probably no protected area between Tully and Cooktown contains sufficient habitat for the life cycle. Taplin (1987) believed adult crocodiles moved in and out of protected areas, juveniles and subadults dispersing into surrounding non-protected areas. Commercial gill netting in tidal rivers was identified as a major threatening factor to crocodiles (Taplin, 1987), but the impact remains to be quantified. This activity continues in six major waterways between Tully and Cooktown. Johnstone, Russell/Mulgrave, Daintree and Endeavour Rivers are closed to this fishing. Despite low densities, problem crocodiles are not infrequent in the study area, often as a result of human behaviour, such as discarding fish scraps at public facilities (boat ramp, wharf, jetty). Two crocodile attacks occurred in Cairns 609 during the study, and public perception is now that large numbers of crocodiles occur between Tully and Cooktown. However many people use these waterways daily, usually without incident, and consequently some crocodiles may be coming less wary and also more visible. Twelve problem crocodiles were removed during the study: 2 had attacked humans, 2 from swimming areas, 3 had attacked dogs, 2 frequented a wharf and 3 were otherwise threats to humans. Removed crocodiles measured 1|.1-2.0m (#7 =3), 2.1-3.0m (4) and 3.1-4.0m (5). In the interests of public safety, the Queensland Parks and Wildlife Service initiated a Trial Intensive Management Area for Crocodiles in May 1998. This 3-year trial program targets all crocodiles for removal in designated areas near Cairns, Port Douglas and Mossman. Also, an education program provides information on appropriate and safe behaviour in crocodile habitat. Research is focussed on surveys and on-going monitoring. The survey data presented here precede implementation of the crocodile removal zone and comprise baseline data for comparison against future surveys. CONCLUSIONS Contrary to public perception, the crocodile population between Tully and Cooktown exists at low density, showing no increase over 14 years. Human activities such as urban, rural residential and agricultural development, clearing of riparian vegetation, disturbance by motor boats, commercial gill netting in estuaries, and removal of crocodiles appear to be keeping crocodile numbers low. ACKNOWLEDGEMENTS The following people rendered assistance: Angela Chapman, Tony Frisby, lan King, Gordon LaPrak, Jeffrey Miller, Frank Nissan, Richard Orchard, Mark Read, Rupert Russell, Clay Smith, Willie Smith, Laurie Taplin and Brent Vincent. LITERATURE CITED AUSTRALIAN NATURE CONSERVATION AGENCY 1996. A directory of important wetlands in Australia. 2nd edn. (ANCA: Canberra). BAYLISS, P. 1987, Survey methods and monitoring within crocodile management programmes. Pp. 157-175. In Webb, GJ.W., Manolis, S.C. & Whitehead, P.J. (eds) Wildlife management: crocodiles and alligators. (Surrey Beatty & Sons: Sydney). 610 BUREAU OF METEOROLOGY 1975. Climatic averages: Australia. (Australian Government Publishing Service: Canberra). 1977. Rainfall statistics: Australia. (Australian Government Publishing Service: Canberra). BURROWS, D.W. 1998. Far North Queensland 2010 Regional Environment Strategy: Key Waterways Report. (James Cook University of North Queensland: Townsville). [Draft report]. CENTRE FOR APPLIED ECONOMIC RESEARCH AND ANALYSIS 1997. Report on the Cairns Regional Economy. (Australian Economic Consultants Group: Townsville). COOK, T., GILLAM, E., BARKER, R. & WARD, G. 1997. Recent Population and Housing Trends in Queensland 1997. (Queensland Department of Local Government and Planning: Brisbane). DEPARTMENT OF ENVIRONMENT AND HERITAGE 1994. Nature Conservation Act 1992. (Government Printer: Brisbane). DEPARTMENT OF PRIMARY INDUSTRIES 1993, The Condition of River Catchments in Queensland: A Broad Overview of Catchment Management Issues. (Department of Primary Industries [ICM]: Brisbane). EYRE, B. & DAVIES, P. 1996. A preliminary assessment of suspended sediment and nutrient concentrations in three Far North Queensland catchments. Pp. 57-64. In Hunter, H.M., Eyles, A.G. & Rayment, GE. (eds) Downstream effects of land use. (Department of Natural Resources: Brisbane). GROOMBRIDGE, A. 1987. The distribution and status of world crocoditians. Pp. 9-21. In Webb, G.J.W., Manolis, S.C, & Whitehead, P J. (eds) Wildlife management: crocodiles and alligators. (Surrey Beatty & Sons: Sydney). HERBERT, B. & PEETERS, J. 1995. Freshwater fishes of Far North Queensland. (Department of Primary Industries: Brisbane). KOFRON, C.P., 1992. Status and habitats of the three African crocodiles in Liberia. Journal of Tropical Ecology 8: 265-273. KREIGER, G. & FELL, D. 1991. Results from Population Survey of Crocodiles on the North, East Coast Of Queensland — February and June 1991. (Queensland Parks and Wildlife Service: Cairns). [Unpublished internal report]. MESSEL, H., VORLICEK, GC., WELLS, A.G & GREEN, W.J. 1981. Surveys of tidal river systems in the Northern Territory of Australia and their crocodile populations. Monograph No. 1. (Pergamon Press: Sydney). NORTH QUEENSLAND JOINT BOARD 1997, Barron River Catchment Rehabilitation Plan: Technical Report on Rehabilitation Needs. (North Queensland Joint Board: Cairns). MEMOIRS OF THE QUEENSLAND MUSEUM PITTS, D. 1973. Water quality management in the Johnstone River catchment. Pp. 1-43. In Resource Assessment Commission (commissioned by) Water quality management: five case studies. (Resource Assessment Commission: Canberra). READ, M. 1998. Report on Distribution and Abundance of the Estuarine Crocodile (Crocodvlus porosus Schneider) in Queensland. 3. Western and Northwestern Cape York Peninsula. Research Conducted October- November 1997. (Department of Environment and Heritage: Townsville). [Unpublished internal report]. READ, M. & MILLER, J.D. 1998. Report on Distribution and Abundance of the Estuarine Crocodile (Crocodylus porosus Schneider) in Queensland. 2. Lakefield National Park. Research Conducted September 1997. (Department of Environment: Townsville). [Unpublished internal report]. ROSS, J. P. (ed.) 1998. Crocodiles. Status Survey and Conservation Action Plan, 2nd edn. (IUCN: Gland). RUSSELL, D.J. & HALES, P.W. 1997. Fish Resources and Stream Habitat of the Liverpool, Maria and Hull Catchments. (Queensland Department of Primary Industries: Brisbane). TAPLIN, L.E. 1987. The management of crocodiles in Queensland, Australia. Pp. 129-140. In Webb, GJ.W., Manolis, S.C. & Whitehead, P.J. (eds) Wildlife management: crocodiles and alligators. (Surrey Beatty & Sons: Sydney). 1989, Crocodile Management Plan 1989. (Department of Environment and Heritage: Townsville). [Unpublished internal report]. WALSH, B. & WHITEHEAD, P.J. 1993, Problem crocodiles, Crocodylus porosus, at Nhulunbuy, Northern Territory: an assessment of relocation as a management strategy. Wildlife Research 20: 127-135. WEBB, GJ.W. & MESSEL, H. 1979. Wariness in Crocodylus porosus. Australian Wildlife Research 6: 227-234. WEBB, GJ.W. & SMITH, M.A. 1987. Life history parameters, population dynamics and the management of crocodilians. Pp. 199-210, In Webb, GJ.W., Manolis, $.C. & Whitehead, P.J. (eds) Wildlife management: crocodiles and alligators. (Surrey Beatty & Sons: Sydney). WEBB, GJ.W., WHITEHEAD, PJ. & MANOLIS, S.C. 1987. Crocodile management in the Northern Territory of Australia. Pp. 107-124. In Webb, GJ.W., Manolis, S.C. & Whitehead, P.J. (eds) Wildlife management: crocodiles and alligators. (Surrey Beatty & Sons: Sydney). SPONDYLIDS FROM THE MEDITERRANEAN SEA AND ATLANTIC OCEAN (MOLLUSCA: BIVALVIA: SPONDYLIDAE) KEVIN LAMPRELL, JOHN STANISIC AND PETER CLARKSON Lamprell, K., Stanisic, J. & Clarkson, P.2001 06 30: Spondylids from the Mediterranean Sea and Atlantic Ocean (Mollusca: Bivalvia: Spondylidae). Memoirs of the Queensland Museum 46(2): 611-622. Brisbane. ISSN 0079-8835, A new spondylid from the Ligurian Sea, Spondylus cevikeri sp. noy., is described. A spondylid from the Cape Verde Islands, possibly new and very similar to the Indo-Pacific Spondyvlus linguafelis Sowerby, G.B. II, 1847 is recorded and figured. The numerous synonyms of Spondylus gaederopus Linnaeus, 1758, are discussed and additional taxonomic notes presented. Spondylus reevei Fulton, 1915 is shown to be valid; Spondylus limbatus Sowerby, GB. I, 1847 is shown to be a prior name for Spondylus calcifer Carpenter, 1857; Spondylus powelli Smith, 1892 is shown to be a junior synonym of Spondylus senegalensis Schreibers, 1793; Spondylus multisetosus Reeve, 1856 previously known only from the Indo-Pacific is now recorded from the Mediterranean Sea; Spondvlus lamarcki Chenu, 1845, is placed in synonymy with Spondylus spinosus Schreibers, 1793; and variations of S. spinosus, including an all-brown specimen, are discussed and figured. O Spondylus, new species, Mediterranean Sea, Atlantic Ocean. Kevin Lamprell & John Stanisic, Queensland Museum, PO Box 3300, South Brisbane 4101; Peter Clarkson, PO Box 713, Port Lincoln 5606; 8 May 2000. Spondylids are an extremely difficult group to identify by shell characters because of the high level of intraspecific variability related to growth pattern. In part of their life cycle all spondylids are attached to the substrate by the lower (right valve), at least vestigially. In some species, such as Spondylus imperialis Chenu, 1845, S. pratti Parth, 1998 or S. regius Linnaeus, 1758, the right and lys are virtually mirror images of each other in terms of spination, because the right valve grows free of the substrate in adulthood. However, in the majority of species the area of attachment of the right valve is widely variable and, to some extent, the shell will mould itselfto the shape of its anchorage. If they become dislodged due to wave action or some other reason, the ornamentation of the right valve will be further affected. As such, there may be little consistency in the overall shape and ornamentation of the right valve. This leaves only the top (left) valve and internal features of the shell as useful characters for field identifications. Because of the intraspecific variability of many species, such as S. gaederopus Linnaeus, 1758, a wide range of specimens may be necessary in order to establish accurate identification. Reasonably reliable taxonomic characters appear to be: width of ears (auricles); inflation of left (top) valve; inflation of right (lower) valve: number and structure of ribs; ornamentation (sculpture) of ribs; sculpture of interstitial areas; external colour (in some species); internal shell colour particularly the colour of the internal margin crenulations. Problems of identification could be solved by molecular studies, sperm ultrastructure or scanning electron microscopy, and some species may yet be reduced to subordinate rank by the use of such methods. However, these tests are laboratory-based and do not help the field worker with identification of species. The authors have had the opportunity to study spondylids from the Mediterranean Sea, the Senegal coast, Cape Verde and Canary Islands including several variations of the extremely variable S. gaederopus. The various synonyms (forms) of this species are discussed and a new species, S. cevikeri, is described. Additional taxonomic notes are also presented on several other spondylids. ABBREVIATIONS. AMS, Australian Museum, Sydney: QM, Queensland Museum, Brisbane; DC coll, Dogan Ceviker collection; FS coll, Frank Swinnen collection; KL coll, Kevin Lamprell collection; NSW, New South Wales; Qld, Queensland; pv, paired valves; rv, right or upper valve; lv, left or lower valve. MEASUREMENT DETAILS. Height= greatest vertical distance between the centre of the umbo and the lowest part of the ventral margin of lv, excluding spines; width = greatest horizontal distance between the anterior and dorsal margins of ly, excluding spines; depth of pv = greatest 612 distance between the external surfaces of the left and tight valves excluding ribs and spines, Family SPONDYLIDAL Gray, 1826 Spondylus gaederopus Linnagus, 1758 (Fig. 1) Spomidvins geredeveymas Linnaeus, (758: 1136: Schreibers 1795; 152 (based an Cherie, 1784! 4497: Chenu 1845; pl. b fig. 1; pl. 2. igs 1-24) Sowerby 147, figs 24, 47) Reeve VeSo: My. 13; Fultar (975; 331, sp. is Lucas 1978, 0-4, Abbott & Dance (982: 317; Lamprell 1986: pl. 1. fy. ba, Spondvins madierruneus Heonam, C7812 10, Spondvits spinesas Marivo,1784 non Sebrethers. 1793. Spondvlis iments Monterosata, IR7S: 64. Spondylus clbiaus Monterosato, ATS: 64. Spondias coralinus Manterosata, |R75; 64, Spanevins faliovuy Montenrsato, 18757 04, Spondviis aculeatus Philippi. 1884): 74. Spemdvius tomellesuy Pallary, 1904: 52, 244. Spandus mixtes Koch and Pallary, 1900; 48, 371 Spondvius wnicus Jousseame, (927: 33, 307. TYPE MATERIAL. LECTOTYPE: Linnean Collection, London (Dodge, 1952), Mediterranean Sea, DESCRIPTION. Shell ovate, almost equivalve, height to 90mm, rv deeper than ly. Seulpture variable, both valves usually with &-9 ribs omamented with irregular shaped spines Lhal are hollow undermeath, varying from spatulate to fat or shurp spines; interstices with dense small prickles. Colour of tv usually purple, ry white; some orange or all white specimens: are also known. Internally the crenulations are moder- ately fine, purple on the lv and usually white on the rv. DISTRIBUTION AND HABITAT. Mediter- ranean Sea; northwest African coast; attached to dead coral or rock amongst alvae to at least 30m, Usually covered in a distinvlive orange-red sponge that appears lo grow only on Spordylus (P. Clarkson, pers. obs.). REMARKS. In both the 1758 and 1767 editions of the “Systema Naturae’, Linnaeus gave the same brief description of Spanedvius gaederopus: ‘S. testa aubaurita spmosa ... Natum altera MEMOIRS OF THE QLULLNSLAND MUSEUM longior. hine plana, ac si arte aut abrasa fuisset’ with the locality “M. Mediterraneo, arce adhuerens scopulis’. Twenty-eight figures from 9 different authors were quoted for.5. gaederopus in the 12th edition of Linnaeus’ ‘Systema Naturae’, It has been shown that many of these figures referto a number of spondylids other than S. gaederopus so that the diagnosis of the species by Linnaeus must be considered a composite and therefore undefined (Dodge, 1952). Dodge (1952), however, recognised a specimen of § gaevderopuy lodged in the Linnaean Collection, London as the Linnaean type specimen. This specimen, while unidentified, not only closely agrees with the few details of the original description, but also was one of only two spondylids present in the collection. Numerous names have been used Jor the different forms of 5. gaederopus. S. spinosus Murtyn,1784 non Sehreibers, 1793 was introduced for specimens possessing sparse, lon spines; 5, dermis Monterosato, 1875 for specimens with sparse sculpture without strongly developed spines or other projections; &. a/binus Monterosato, 1875 for white shelled forms; S. coralinus Momerosato, 1875 for coral red forms; §. foliosus Monterosato, 1875 for specimens with omamentation of large, leaf-shaped radiating lamellae; S. /amellosus Pallary, 1904 for, specimens with numerous spatulate lamellae: and §. mixtus Koch & Pallary. 1900 for specimens with numerous mixed, spatulate lamellae and spines, These names, while strictly synonyms of 8. gaederopus have nevertheless proven useful when referring to the various forms ol 8, geederapus. Spondylus cevikeri sp, nov. (Figs 2F-K, 3A-C) PTYMOLOGY. Named for Dogan Ceviker (Istanbul). MATERIAL. HOLOTYPE: AMS C204238, | py, Ligurian Sea, height 80.7mm, width 76.5mm, depth of conjoined valves 43. Jinm, PARATYPES: OMMO66960 FIG_\_A-L. Spondylus gaederopus (Linnaeus). A, external view, lv, form spinayus, DC coll 209: height 713mm, width 60.0nm, depth of py 32.0inin, B, external view, ly, form erivivs, FS coll; height 59.Simm, width 59. 7mm, depth of py 24.0mm. C, external view, lv, form spinasus, KL coll, Antibes 15.0m: height 40.02om, width 34.5mm, depthofpy 22.5mm. D, external view, ly, form mernzix, KL, coll, Ligurian Sea, 45,.0m: height 66,9mm, width 61.9mm. depth of py 38.2mm. E, external view, ly, form a/birws, KL call, Majorea, Spain: height 49,Snim, width 42.5imm, depth of pv 32, 5mm. V', external view, lv, form corafinus, DC coll 265: height §4.2mm, width 83,4mm, depth of lv 23.0mm. G, external view, rv, form a/binus, DC coll, Karatas. Adana, 60-90.0m: beight 44.2mm, width 36.6mm, depth of rv 14.8mm, 1, external view, lv, typical form, KL coll, Mediterranean Sear height 796mm, width 65.8inm, depth of py 40.0mim. I, L, KL coll, Bodrum, southern Aegean Sea. 1. external view, ly, L, ventral view, pv. J, K, AMSC303118, | py Guvenile), Hilat, Israel. J. internal view, rv. RK. external view, Iv; height 29.5mm, width 31.3mm, depth of py (4.1mm. 613 MEDITERRANEAN AND ATLANTIC SPONDYLIDS 6l4 MEMOIRS OF THE QUEENSLAND MUSEUM MEDITERRANEAN AND ATLANTIC SPONDYLIDS | py, same data as holotype; AMSC204279 1 py, South side of Zaborgad Island, Red Sea, P. Clarkson, x 1994; AMSC99484, 1 pv (juvenile) Bay of Stari Grad, Bosnia, 2-10m, attached to rocks. DESCRIPTION. Shell elongate-ovate to pear-shaped. Height to 80.7mm, approximately 1.74 times greater than the auricle width. Approximately equivalve, lv moderately convex; interior slightly excavated under hinge plate, with a strong, raised, coloured crenulated margin. Sculpture consists of numerous, strong, raised radial ribs; interstices narrow with a minor riblet centrally. Major ribs bear numerous spines varying from long and slightly spatulate to short or long and sharp. Lower (right) valve equally convex as the lv; ornamentation of unattached areas more densely spined than lv; cardinal area triangular; internally with a deep excavation under the hinge plate and a strong, coloured, raised crenulated margin. Fixation area large, with foliations supporting fixed area. Colour red-purple, lighter umbonally with indistinct black lines and markings at the umbonal region; internally blue-white with dark red-purple crenulated margin, external colour visible centrally. Based on 4 specimens. TYPE LOCALITY. Ligurian Sea. DISTRIBUTION AND HABITAT. Ligurian Sea - Red Sea; attached to corals or rock, to 25m. REMARKS. Spondylus cevikeri sp. nov. most closely resembles S. gaederopus Linnaeus, 1758 in shape and colouration. S. cevikeri can be readily separated by the purple-coloured right valve (mostly white or orange in gaederopus), the numerous, equally sized radial ribs and smooth interstices (8-9 ribs ornamented with irregular shaped spines varying from spatulate to flat or sharp spines and interstices with dense small prickles in gaederopus), dark coloured lines and marks umbonally (absent in gaederopus) and purple internal crenulated margins in both valves (lv purple, rv usually white in gaederopus). Spondylus reevei Fulton, 1915 Spondylus hystryx Reeve, 1856; 12, 42. non Réding, 1798. 615 Spondvlus reevei Fulton, 1915: 332, sp, 7. Spondylus cuneus Lamprell 1986; pl. 8, figs 1A-D, non Reeve, 1856. TYPE MATERIAL. HOLOTYPE: BMNH1984252/1, Philippine Islands. DESCRIPTION. Shell elongately ovate, height to 90mm. Sculpture of many irregular ridges, ornamented with numerous, regular, strong, slightly depressed, blunt spines of varying lengths; interstices usually smooth but minor spines do occur in some specimens. Colour purple-red to brown, spines usually purple, internally white with a purple crenulated margin. Area of attachment variable. DISTRIBUTION AND HABITAT. West Indies (Reeve, 1856), Mozambique, South Africa; and widely distributed throughout the Indo- and central Pacific; on and under dead coral, to 30m. Fine specimens have been collected from shipwrecks in the Solomon Islands and Palau. REMARKS. This species was figured by Lamprell (1986) as a synonym of S. cuneus Reeve, 1856. Examination of numerous specimens of S. americanus Hermann,1781 obtained by diving off Florida by Peter Clarkson, has, in our opinion, confirmed that S. cuneus is a junior synonym of that species. Spondylus limbatus Sowerby, G.B. II, 1847 Spondylus limbatus Sowerby, G.B. IH, 1847: 427, fig. 51; Lamprell, 1986: pl. 16, fig. 2; Lamprell, 1998: pl. 2, figs 8,10. Spondylus radula Reeve, 1856: pl. 14, sp. 52; Spondylus calcifer Carpenter, 1857: 152; Fulton, 1915; 357, sp. 68; Eisenberg, 1981: pl. 145, sp. 4: Lamprell, 1986: pl. 20, fig, 1: Skoglund & Mulliner, 1996: 102. Spondylus smithi Fulton, 1915: 357, sp. 66. TYPE MATERIAL, HOLOTYPE: BMNH 1846.12.4.1, Persian Gulf (sic). DESCRIPTION. Shell ovate to elongately ovate, equivalve, height to 200mm. Sculpture of 6 principal radial ribs on top (lv) with spatulate appressed spines, stronger marginally, upright umbonally; interstices with numerous radial riblets with several stronger than the others and ornamented with similar but smaller spines than on the principal ribs; rv with numerous radial ribs FIG. 2. A-D, Spondylus sp. 1 pv, FS coll, St Vincents, Cape Verde Islands. A, external view, lv. B, ventral view, pv. C, internal view, rv. D, internal view, lv: height 58.5mm, width 54.0mm, depth of pv 23.2mm. E, Spondylus linguafelis Sowerby, 1847, KL coll, Herald Prong Reef, Swain Reefs, external view, Iv: height 41.5mm, width 39.0mm, depth of pv 19.7mm., F-K. Spondylus cevikeri sp. nov. F-J, holotype, AMS C204238, Ligurian Sea, | pv; F, external view, lv. G, external view, rv. H, ventral view, pv. I, internal view, rv. J, internal view, lv: height 80.7mm, width 76.5mm, depth of conjoined valves 43.1 mm. K, paratype, AMSC99484, | py (juvenile), Bosnia; external view, lv: height 27.3mm. 616 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. A-C, Spondylus cevikeri sp. nov. A, paratype, AMSC204279, | pv, south side of Zaborgad Island, Red Sea, P. Clarkson, Oct 1994, external view, lv: height 49.1mm, width 40.7mm, depth of conjoined valves 29.1mm. B-C, paratype, QMMO66960, | pv, same data as holotype. B, external view, rv. C, internal view, lv. height 60.7mm, width 51.5mm, depth of conjoined valves 29.1mm. D-K, Spondylus senegalensis Schreibers, 1793. D, lectotype (illustration from Adanson, 1757). E-I, FS coll, Canary Islands. E, 1 pv, juvenile, external view, lv: height 24.2mm, width 26.0mm, depth of conjoined valves 13.3mm. F, | pv, juvenile, external view, lv: height 37.7mm, width 44.5mm, depth of conjoined valves 30.7mm. G, | pv, juvenile, external view, lv: height 41.0mm, width 47.0mm, depth of conjoined valves 27.8imm. H-I, | pv. H, internal view, rv; I, external view, lv: height 71.0mm, width 63.8mm, depth of conjoined valves 36.3mm. J-K, KL coll, | py, Canary Islands. J, external view, lv, K, external view, rv: height 50.0mm, width 56.0mm, depth of conjoined valves 29.1mm. MEDITERRANEAN AND ATLANTIC SPONDY LIDS and dense appressed, spatulate spines, mterstices natrow with nine minor radial ribs and smaller, similar spines. Colour purple, maroon, orange, or brown with some yellow or white areas- Attachmentarea variable, but in larger specimens rv i$ usually entirely cemented to the substrate, DISTRIBUTION AND HABITAT. Persian Gulf (Sowerby, 1847), Sea of Cortez, Mexico and Ecuador. Attached to coral rubble, dead shell debris, or solid rock. at 1-30m. Frequently covered with a greyish maroon sponge, which usually preserves the omamentavion ofthe shell. REMARKS. Sowerby (1547) desertbed 5S. Jimbatus trom the Persian Gulf, however examination of numerous spondylids from that area has failed to identily 8. Jinbafis among them. A companson of S) caleifer Carpenter, La57 collected in the Sea of Cortez (by Peter Clarkson) and the holotype ol S. /inhatus show these are the same, Flence, until there is evidence ol SJimbetus frant the Persian Gull) this type locality should be regarded as erroneous. Spondylus sp, (Fig, 24-B) MATERIAL. L pv, St Virwents. Cape Verde Islaneds. FS voll, DESCRIPTION. Shell: ovate, heightto 58.Simm, approximately 1,73 times greater than the auricle width: both valves moderately shallow. Sculpture of numerous fine radial ribs densely spined with short to long spatulate and sharp spines: interslices narrow with dense. short, minute spines, Colour of both valves purple, white at umbonal area ; internally white, lv with purple crenulated inner margin, ry crenulated inner margin white. outer edge mauve, Area of attachment large. DISTRIBUTION AND HABITAT, Cape Verde Islands: attached to coral, at 15-20m. REMARKS. The equi-distant and close radial ribs. fine dense spines and purple nght valve colouration distinguish the Cape Verde specimen from typical forms of 8. gaederopus. Spanedylus ap. is most similar to short spined §. /ingua/eliy G.8. Sowerby U1, 1847 from the Indo-Pacilic and Australia and appears remarkably similar to a purple coloured, short spined form of this species fromthe Swain Reefs, Queensland (Fig. 26), The lovality, while previously considered yery improbable for a predominantly Indo-Pacific gpecies, is now acceptable in the hight of known ol) ingress into the Medilerranean and Red Seas of other Indo-Pacific species of Spondylys. In the absence Of additional material itis diffeult to be certain of its correc! identification. Spundylus senegalensis Schrethers, 1793 (Figs 3D-K, 4A-1) Spondvins senegalensis Schreithers. (793 PURO! PL VF, fig, 3 Spondi tie. powell: Smith, | 892; 36, Spondivins haeidis Quuraenbere, (895 pon Belbirdi, |B52 (fossil) MATERIAL. 3 pv. Puerta del Carmen. 1998, 2(-30n7 by diver, PS coll; 7 py, ix-1999, 20-30m, wrowdh series, Los Caneajos, La Palma Island. FS coll: | py, Lide. Funchal, Madeira, 20m by diver, /S coll; | py, Sardina, FS coll; Canary Islands, KL coll. DESCRIPTION. Shell pear- to fan-shaped: height io 150.0mm, approximately 1.88 times greater than the auricle width (based on measurements of 4 speeimens). Approximately equivalye, Ly with 4-11 principal radial ribs, ornamented with numerous short to moderately long, depressed, blunt and sharp spines; interstices with 5 or 6 minor radial ribs; minot ribs and interstives with dense, short overlapping minor spines or imbrications: rv sculpture where visible of strong, close, radial ribs with numerous overlappiny spatulute Lo sharp spines. Speeimens examined from Senegal usually have 4-5 radial nibs with strong overlapping spatulate spines, interstices devoid of spination. Colow” orange red, bright red or brown with off-white spines, internally white with red or dark brown, wide crenulated margins in boil valves. TYPE LOCALITY. Senegal, DISTRIBUTION AND HABITAT. Madeira, Porto Santo Island, Canary [slands, Cape Verde Islands, Senegal, Liberia, Ivory Coast. Principe Islands, Cameroons and Gabon; attached to roek or dead coral. REMARKS. Sponedyluy powell! Smith, 1892 has been placed in the synonymy ofS. senegalensis (Schreibers, 1793). However. until specimens including a growth series were recently obtained (rom Mr Frank Swinnen (Belgium), both species were considered valid by several authors. Mr Swinnen’s specimens have enabled a thorough examination and camparison of humeraus variable forms of the species which show that although the typical fan-shaped Senegal specimens differ from the larger and ofien pear-shaped specimens from other localities, there is msulficient variation to separate the lwo (42: Lamprell 618 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Spondylus senegalensis Schreibers. 1793, Guinea, FS coll. A-B, | py, A, external view, lv, B, dorsal view, py; height 59.5mm. width $7.5mim, depth of conjoined valves 40.6mm. C, F’, 1 pv; C, dorsal view, pv. F, external view, lv: height 89.0mm, width §3.2mm, depth of conjoined valves 50.2mm. D-E, | py. D, internal view rv. E, external view of ly: height 118.7mm, width 98.3mm, depth of conjoined valves 59.5min. G, | pv, external view of ly: height 115.8mm; width 119.7mm; depth of py 71.2mm. H-I, 1 pv. H, internal view of rv. 1, external view, lv: height 4.5mm; width §1.9mm; depth of py 42.0mm, MEDITERRANEAN AND ATLANTIC SPONDYLIDS 619 FIG. 5. Spondylus nuultisetosus Reeve, 1856. A, BMNH1998094, syntype, external view, lv. B-E, Mediterranean Sea, DC coli No. 281. B, external view, lv. C, internal view, lv. D, external view, rv. E, dorsal view, pv: height 70.2mm, width 75.2mm, depth of conjoined valves 36.6mm. F, Cebu I., Philippine Islands, KL coll, external view of lv: height 59.8mm, width 52.2mm, depth of conjoined valves 31.0mm. G-I, Mediterranean Sea, DC coll, G, No. 269, external view, lv: height 72.5mm, width 67.9mm, depth of conjoined valves 44.3mm. H, No. 270, external view, lv: height 85.0mm, width 77.0mm, depth of conjoined valves 47.5mm. I, No. 233, external view, lv: height 73.0mm, width 65.5mm, depth of conjoined valves 43.3mm. J, Capricorn Channel, Qld, 127m, KL coll, external view of lv: height 71.8mm, width 64.6mm, depth of conjoined valves 40.2mm. 620 forms. As a growth series of this unusual species has not been previously figured, a plate showing shape, size and sculpture variations 1s included. Spondylus multisetosus Reeve, 1856 (Fig. 5A-J) Spondvlus multisetosus Reeve, 1856: pl. 3, fig. 11; Fulton, 1915: 353, sp. 48; Lumprell, 1986; pl, 13, lig. 1, MATERIAL. 4 specimens, DC coll,: nos 233,270, Tasucu, Turkey; nos 269, 281, Iskenderun, Turkey. DESCRIPTION. Shell ovate, height to 68mm; approximately 2.48 times greater than the auricle width (based on measurements of 4 specimens). Ly usually more inflated than the ry. Sculpture of numerous radial ribs and striae with numerous upright spines, hollowed underneath with some remote, slightly spatulate spines; interstices densely ribbed, ornamented with small sharp spines. Colour mauve or brown with some yellow on ribs and spines, sometimes yellow white, dark coloured umbonally; internally blue white with a moderately wide, dark purple-brown or variegated yellow and brown crenulated margin. TYPE LOCALITY. Philippine Islands. DISTRIBUTION AND HABITAT. Indo-Pacific (Philippines); Mediterranean Sea; attached to Ilammer Oysters, shell debris, corals or rock in shallow, turbid water. REMARKS. These specimens confirm this Indo-Pacific species in the Mediterranean Sea. The brown or mauve base colour and yellow upright spines distinguish S. multisetosus from any other Mediterranean species. Spondylus spinosus Schreibers, 1793 (Fig. 6A-J) Spondylus spinasus Schreibers, 1793; 154 (based on Chemnitz, 1784: fig. 460); Oyama & Takemura, 1960: 97, fig. 2; Habe 1977: 93; Lamprell, 1986: pl. 14, fig, 2a-b; Mienis et al., 1993, Spondylus proboscideus Schreibers, 1793: 468, pl. 145, sp. 11. Spondvlus aculeatus Schreibers, 1793: 476. Spondylus marisrubri Réding,, 1798: 460. Spondvlus dentatus Chenu, 1845: pls 25, 27. Spondyvlus lamarcki Chenu, 1845; pl. 9, figs 3-4; Lamprell, 1986: pl. 1S, fig. 3. Spondvlus actleatus Sowerby, 1847; figs 11-13. MEMOIRS OF THE QUEENSLAND MUSEUM TYPE LOCALITY. Red Sea. DISTRIBUTION AND HABITAT. Mediterranean Sea, Red Sea, Indo-Pacific (Japan, Philippines, Mauritius, Solomon Islands, north Western Australia); attached to dead coral or debris in 3m or more of water. A species frequently found within protected lagoonal environments, where it grows amongst delicate Acropora corals. REMARKS. Some authors have considered Spondylus lamarcki Chenu, 1845 to be a variety of S. squamosus auct., non Schreibers, 1793 (=S. sinensis Schreibers, 1793) however, in S. lamarcki the interstitial areas are quite narrow and the shell more elongate. After examining numerous specimens we consider S. /amarcki to be a form of S. spinosus Schreibers. We have figured a wide variety of the species from the typical form with dark coloured base, white ribs and spines to the all-brown and long spined forms. Small specimens can bear long spines, but these erode as the shell grows, leaving the shell rather poorly sculptured in its adult state. ACKNOWLEDGEMENTS We would like to thank the following museum staff and collectors for their assistance; Ms Kathie Way and Mrs Joan Pickering of the Malacology Section of The Natural History Museum, London for the loan of specimens and supplying information on the status of specimens in the collection; Dr Philippe Bouchet, Muséum National d’ Histoire Naturelle, Paris for the loan of type material; Dr Claude Vaucher and Dr Yves Finet, Muséum d’ Histoire Naturelle, Geneva; Dr John Healy, University of Queensland, Brisbane for helpful advice on the presentation of this paper; Mr lan Loch, Malacology Section, Australian Museum, Sydney for the loan of specimens; Dr Michele Dardano, Mr Frank Swinnen, and Mr Dogan Ceviker donated material. Thanks are also due to Darryl Potter, Malacology Section, Queensland Museum for his helpful comments on the manuscript. The Malacological Society of Australasia provided tavel assistance. FIG. 6. Spondylus spinosus Schreibers, 1793. A, Mediterranean Sea (off coast of Turkey), DC coll nos 239-240, external view of two joined pvs, one all brown, one (largest) typically brown with white ribs and spines: height 89.5mm, width 76.5mm, depth of conjoined valves 56.4mm. B, external view of a lv: height 58.0mm, width 53.4mm, depth of lv 19,6mm, C-D, 1 pv; C, internal view, rv. D, external view, Iv: height 78.2mm, width 73.9mm, depth of conjoined valves 45.8mm. E, external view ofa lv: height 53.1mm, width 47.3mm, F-G, 1 pv, No. 246. F, external view, lv; G, internal view, rv: height 92.6mm, width 91.6mm, depth of conjoined valves 47.0mm, H, external view ofarv: height 68.0mm, width 64.5mm, depth of rv 20.0mm. I-J, 1 pv; 1, external view, lv. J, internal view of rv: height 62.2mm, width 53.5mm, depth of conjoined valves 35.3mm. 621 MEDITERRANEAN AND ATLANTIC SPONDYLIDS 622 LITERATURE CITED ABBOTT, R.T. & DANCE, S.P. 1982. Compendium of seashells. (E.P. Dutton: New York). ADANSON, M. 1759. Histoire naturelle du Sénégal. Coquillages avec la relation abregée du voyage fait en ce pays, etc. Illust. (London). CHEMNITZ, J.H. 1784. Neues systematisches conchylien-cabinet. Vol 7. G.N. (Raspe: Niimnberg). DODGE, H. 1952. A historical review of the mollusks of Linnaeus. Bulletin of the American Museum of Natural History 100: 242-260. FULTON, H.C. 1915. List of the Recent species of Spondylus Linne with some notes and descriptions of six new forms. Journal of Conchology 14: 331-360. LAMPRELL, K.L. 1986. Spondylus, spiny oysters of the world. (E.J. Brill-Dr W. Backhuys: Leiden). 1992a. Notes on Spondylus Linnaeus with descriptions of two new species from Western Australia (Mollusca: Bivalvia: Spondylidae). Memoirs of the Queensland Museum 32: 189-194. 1992b. On some species of Spondylus. La Conchiglia 23: 40-41. 1998. Recent Spondylus from the Middle East and adjacent regions, with the description of two new species (Mollusca, Bivalvia, Pectinoidea). Vita Marina 45: 41-60. LINNAEUS, C. 1758. Systema Naturae, per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 10th edition. (Holmiae: Laurentii Salvii). MEMOIRS OF THE QUEENSLAND MUSEUM LUCAS, M. 1978. Pectinoidea from the European coasts. La Conchiglia 116-7: 6-8. MIENIS, H.K., GALILI, E. & RAPOPORT, J. 1993. The Spiny Oyster, Spondylus spinosus, a well-established Indo-Pacific bivalve in the Eastern Mediterranean off Israel (Mollusca, Bivalvia, Spondylidae). Zoology in the Middle East 9: 83-91. MONTEROSATO, T.A. di 1875. Nuova revista delle conchiglie Mediterranee. Atti della Accademia Palermitana sienze e lettre. new series, 5: 1-50. PALLARY, P. 1904. Coquilles marines du littoral du départment D’Oran. Memoires du Muséum national d’ Histoire Naturelle de Paris. Journal de Conchyliologie. 48: 211-434. REEVE, L.A. 1856. Monograph of the genus Spondylus. Conchologia Iconica or illustrations of the shells of molluscous animals, Vol. 9. (Reeve: London). RODING, P.F. 1798. Museum Boltenianum sive Catalogus cimeliorum e tribus regnis naturae. Pars secunda, Conchylia sive Testacea univalvia, bivalvia & multivalvia. (J.C. Trappii: Hamburg). SCHREIBERS, K. 1793. Versuch einer Vollstandigen Conchylienkenntniss nach Linné’s System. (Vienna). SMITH, E.A. 1892. Description of a new species of Spondylus and a new Helix. Journal of Conchology 7: 71. SOWERBY, G.B. II 1847. Descriptions of several new species of Spondylus. Proceedings of the Zoological Society of London 1847: 86-88. SPRINGSTEEN, F.J. & LEOBRERA, F.M. 1986. Shells of the Philippines. (Carfel Seashell Museum: Manila). SOME SPONDYLIDS FROM THE PACIFIC OCEAN (MOLLUSCA: BIVALVIA: SPONDYLIDAE) KEVIN LAMPRELL, JOHN STANISIC AND PETER CLARKSON Lamprell, K., Stanisic, J. & Clarkson, P. 2001 06 30: Some spondylids from the Pacific Ocean (Mollusca: Bivalvia: Spondylidae), Memoirs of the Queensland Museum 46(2): 623-628. Brisbane. ISSN 0079-8835. Spondylus swinneni sp. noy. is described and figured; the type of Spondylus multimuricatus Reeve, 1856 is figured and redescribed; Spondylus hawaiensis Dall, Bartsch & Rehder, 1938 is placed in the synonymy of Spondylus candidus Lamarck, 1819; Spondylus punicus Bernard, Cai & Morton, 1993 (nom. nov. for Spondylus coccineus Lamarck, 1819 non Schreibers, 1793) is discussed. O Spondvlus, new species, Pacific Ocean. Kevin Lamprell & John Stanisic, Queensland Museum, PO Box 3300, South Brisbane 4101, Peter Clarkson, PO Box 713, Port Lincaln 5606; 8 May 2000. Since publication of ‘Spondylus, Spiny Oyster Shells of the World’ (Lamprell, 1986), several important taxonomic changes have been effected within the Spondylidae (Lamprell, 1992a; Lamprell, 1992b; Lamprell & Kilburn, 1995; Skoglund & Mulliner, 1996; Lamprell, 1998; Lamprell & Dekker, in press; Lamprell & Willan, 2000; Lamprell & Healy, 2001). Thirteen new species have been described and several species listed in the book have been placed in synonymy. These changes were largely based on the results of extensive deep-water exploration off the coast of New Caledonia by the ORSTOM expeditions (Lamprell & Healy, 2001) and diving off the American coast and Solomon Islands. As a result there has been renewed interest shown in the group by collectors in Europe, Australia and other Indo-Pacific countries and consequently much more material from private collections has become available for scrutiny. The present study aims to rectify several additional taxonomic problems that have come to notice following examination of this material. ABBREVIATIONS. QM, Queensland Museum, Brisbane; AMS, Australian Museum, Sydney; BMNH, The Museum of Natural History, London; MNHN, Muséum National d’ Histoire Naturelle, Paris; MHNG, Muséum d’Histoire Naturelle, Geneva; USNM, United States National Museum, Washington; KL coll, Kevin Lamprell collection; PC coll, Peter Clarkson collection; lv, left valve; rv, right valve; pv, paired valve; NSW, New South Wales; WA, Western Australia; Qld, Queensland. MEASUREMENT DETAILS. Height = greatest vertical distance between the centre of the umbo and the lowest part of the ventral margin of lv, excluding spines. Width = greatest horizontal distance between the anterior and dorsal margins of lv, excluding spines. Depth of pv = greatest distance between the external surfaces of the left and right valves excluding the ribs and spines. Family SPONDYLIDAE Gray, 1826 Spondylus swinneni sp. nov. (Fig. 1G-K) Spondylus multimuricatus Reeve, 1856: Lamprell, 1986; 32; pl. 9, fig. 2a-b, non Reeve, 1856. ETYMOLOGY. For Mr Frank Swinnen. TYPE MATERIAL. HOLOTYPE: AMSC303105, Philippine Islands. PARATYPES: AMSC303116, northern Qld; QMM066961, Finger Reef, Swains Reef Qld, off corals, 18m, P. Clarkson. OTHER MATERIAL. KL coll, 1 pv, Bonegi Wreck, Solomon Islands, 28.viii.85; KL coll, 1 py, West of Honiaro, Solomon Islands, dived, P.Clarkson; KL coll, 2pv, Gneering Shoals, south Qld, dived 6-30m, P Clarkson; PC coll, 9pv, West of Honiara, Solomons, dived 7-50m, shipwrecks and reefs, P. Clarkson; PC coll, 1 pv, Santo, Vanuatu, 8m; PC coll, | py, North West Reef, Exmouth, WA, 24m, P. Clarkson. DESCRIPTION. Shell elongate-ovate to ovate, height to 125mm; lv moderately convex to flat, width of the auricles approximately half the height of the shell. Sculpture consisting of numerous radial ribs; interstices wider than the ribs with one or two minor riblets. Major ribs bearing numerous overlapping spines varying from short to moderately long, sharp, blunted or slightly spatulate, frequently curved inward and hook-like. Internally slightly excavated under the hinge; pearl white centrally, deep brown outer crenulated margin, orange to yellow-brown at the crenulations and lighter brown inner margin. Rv slightly more convex than the ly; ornamentation 624 MEMOIRS OF THE QUEENSLAND MUSEUM SPONDYLIDS FROM THE NORTH AND SOUTH PACIFIC of unattached areas consisting of evenly spaced radial ribs and dense overlapping spines, sharp and hook-like, rarely if ever blunt or spatulate. Cardinal area triangular; internally with a shallow excavation under the hinge plate and a strong, crenulated margin similarly coloured to the lv. Area of attachment variable with limited foliations supporting fixed area. Colour orange-brown with indistinct black lines and markings at the umbonal region; some specimens have black in the interstitial areas extending to the lower margin, TYPE LOCALITY. Philippine Islands. DISTRIBUTION AND HABITAT. Indo-Pacific, Solomon Islands, and northern Australia; cemented to ledges, overhangs, and cave walls, frequently along drop-offs festooned with gorgonian corals and subject to moderate currents, in 10 to more than 50m. Delicately sculptured specimens have been collected from sheltered positions within shipwrecks at similar depths. REMARKS. Specimens of Spondylus swinneni sp. noy. have been confused with S. multimuricatus Reeve, 1856 in the past (Lamprell, 1986). However, compared with the latter species the hinge teeth of S. swinneni are finer and tinged with brown, the spines overlap and are not arranged concentrically, the ribs are more numerous and less prominent. The ornamentation of the lv of S. swinneni is of short to moderate length spines, deposited evenly on the ribs, and consistently sharp and hook-like (on the lv of S. multimuricatus the major spines are relatively broad and blunt while the species has a limited number of prominent ribs). Spondylus swinneni is also superficially similar to the sympatric S. linguafelis Sowerby, G. B. I, 1847. However, the shorter, spatulate or hook-like spines (longer and needle-like in S. /inguafelis) readily distinguishes the new species. Spondylus multimuricatus Reeve,1856 (Fig. 1A-F) Spondylus multimuricatus Reeve, 1856: pl. 4, fig. 15; Fulton, 1915: 353, sp. 49; Abbott & Dance, 1986 (3rd ed.): 316. 625 TYPE MATERIAL. Specimen figured by Reeve, BMNH1998093/1 lectotype herein selected, Philippine Islands; paralectotype, BMNH1998093/2, same data as lectotype. Dimensions of lectotype: height 70.5mm, width 64.0mm, depth of py 35.0mm, OTHER MATERIAL. KL coll, | pv, Koh Samui, southeast Thailand, Gulf of Siam; KL coll, | pv, Cebu Islands, Phillipines, by dealer; KL coll, 1 pv, Heron Island, Qld, L. Newman; KL coll, 1 pv, Exmouth Gulf, north W.A, P. Clarkson. DESCRIPTION. Shell elongate-ovate, moderately gibbous, height to 100mm. Sculpture of numerous weakly formed ribs, 5-6 of which are ornamented with irregular squamate, short to long spines; interstices with numerous minor radial ribs that are densely spined with minor spines; 2-3 interstitial ribs bear slightly larger sharp spines of varying lengths. Colour red-orange, orange or mauve with some purple at the umbonal area; internally white; crenulated margins deep-orange. Area of attachment variable. TYPE LOCALITY. Philippine Islands. DISTRIBUTION AND HABITAT. Philippine Islands; habitat unknown, REMARKS. Spondylus multimuricatus was incorrectly figured by Lamprell (1986) and Springsteen & Leobrera (1986). The species figured by Lamprell, 1986 is 8. swinneni sp. nov. (see above). The species figured by Springsteen & Leobrera (1986) is a specimen of S. reesianus Sowerby, 1903. Spondylus candidus Lamarck, 1819 (Fig. 2A-H) Spondylus candidus Lamarck, 1819: 188. Spondylus candidus Lamarck: Chenu, 1845: pl. 12, fig. 4; Sowerby I], 1848: figs 3-5; Reeve, 1856: sp. 2; Fulton, 1915: 358, sp. 74; Springsteen & Leobrera, 1986: pl. 92, fig. 6; Lamprell, 1986: pl. 25, fig. 3a-b; Lamprell & Kilburn, 1995; 91, figs d-e (as flabellum); Lamprell, 1998: pl. 3, figs 9-11. Spondylus hawaiiensis Dall, Bartsch & Rehder, 1938: 100, TYPE MATERIAL. HOLOTYPE: MNHN, Lamarck Collection. (Lamprell, 1986: pl. 3, fig 9); Holotype of Spondylus hawaiensis, USNM337515. OTHER MATERIAL. KL coll, | pv, Koko Crater, Hawaii, attached under boulder at 22m, 1996, Chris Takahashi; KL coll, 1 FIG. 1. A-F. Spondylus multimuricatus Reeve, 1856. A-C, lectotype herein selected, BMNH1998093/1, Philippine Islands. A, external view lv. B, internal view rv. C, ventral view pv: height 69.3mm, width 63mm, depth of pv 36.3mm. D-F, paralectotype herein selected BMNH1998093/2, Philippine Islands. D, external view lv. E, internal view rv. F, ventral view pv: height 74.0mm, width 66.0mm, depth of pv 48.0mm. G-L, Spondylus swinneni sp. nov. G, holotype, AMSC303105, Philippine Islands, external view lv. H-J, KL coll, 1 pv, Bonegi Wreck, Solomon Islands. H, internal view rv. I, ventral view pv. J, external view lv: height 27,0m. K-L, paratype, QMM066961, 1 pv, off corals, Finger Reef, Swains Reef, Qld, 18m. K, internal view rv. L, external view ly. 626 MEMOIRS OF THE QUEENSLAND MUSEUM n 2 haere Spe wd ylas Coccintdth fucabli Qnanchvee 8 Sct A FIG. 2. A-H, Spondylus candidus Lamarck, 1819, A,B, KL coll, 1 pv, SW side Cassin I., N of Broome, WA, 13m. A, external view lv. B, internal view rv. C-E, KL coll, 1 pv, Koko Crater, Hawaii, 17m. C, external view lv. D, internal view rv. E, ventral view pv. F-H, KL coll, 1 pv, Oahu, attached under ledge on ceiling, | 8m. F, internal view ly. G, internal view rv. H, ventral view pv. I,J, 1 py, MHNG1088/96, specimen labelled ‘Spondylus coccineus Lk Type locality Amérique? Chenu 15/3 No 11’. 1, external view lv. J, internal view rv. K, | pv, MNHN, specimen labelled ‘Spondylus coccineus Lam nomme par Lamarck’, external view lv and view of hinge of rv. SPONDYLIDS FROM THE NORTH AND SOUTH PACIFIC py, Oahu, attached under ledge on ceiling, 20m, 1999, Chris Takahashi; KL coll, 1 py + attached pv juvenile, Hakeiwa, Oahu, attached under ledge at c.20m, 1999, Chris Takahashi; KL coll, | pv, inside lagoon, Pt Havau, Moorea, Tahiti; KL coll, 1 py, SW side Cassini |., N of Broome, 13m, WA, P. Clarkson. TYPE LOCALITY. Nouvelle-Hollande [= Australia|. DISTRIBUTION AND HABITAT. Mozambique- South Africa, Japan (as S. flabellum) and Indo-West Pacific, Hawaii, Qld, north WA; common under dead coral slabs or on the under faces of huge living Porites coral colonies also occurs in marine caves or on the walls of shipwrecks in 1-40m. REMARKS. While we have been unable to obtain the holotype of Spondylus hawaiensis, Kay (1979) synonymised it with S. tenebrosus Reeve, 1856 (= S. violaceus Reeve, 1856). Specimens of S. hawaiensis sent by Mr Chris Takahashi for examination, after cleaning, proved to be inseparable from S. candidus. Further examination of specimens of S. hawaiensis in the collection of one of the authors (KL) has led us to the conclude that these two species are the same. Spondylus punicus Bernard, Cai & Morton, 1993 (Fig. 21-K) Spondylus punicus Bernard, Cai & Morton, 1993; 55 (nom. nov. for Spondylus coecineus Lamarck, 1819, non Schreibers, 1793). Spondylus coccineus Lamarck, 1819: 190, non Schreibers, 1793. Spondylus coccineus Lamarck: Chenu, 1845; pls 14, 15 (not pl. 25, fig. 2); Reeve, 1856: sp. 44; Fulton, 1915: 336, sp. 31; Lamprell, 1986: 70, fig. 1. Spondylus coccineus var. vaillanti Jousseaume (in Lamy), 1927: 301, DISTRIBUTION AND HABITAT, Indian Ocean, Philippines, South and east China to Okinawa (Bernard, Cai & Morton, 1993); attached to corals in moderately deep water. REMARKS. Spondylus coccineus Lamarck (non Schreibers, 1793) was briefly described by Lamarck (188: 11) without a stated locality. Examination of the Lamarck specimen marked ‘Spondylus coccineus (pl. 2, fig. 11)’ in the MNHN showed a pear-shaped spondylid with narrow, oblique auricles and dense radial ribs devoid of ornamentation, probably due to age and deterioration; internally off-white with a raised crenulate, dark coloured margin. A specimen marked ‘Spondylus coccineus Lk Type’ (pl. 2, fig. 9-10) with the locality ‘Amérique?’ Chenu 627 15/3 No 11, registration number 1088/96 is in the collections of the MHNG. This specimen is ovate, with wide interstices and strong, sparce spines on the 5 or 6 principal ribs. The specimen is tan-brown in colour with a light orange coloured internal crenulated margin. The specimen bears a strong resemblance to S. gilvus Reeve, 1856. In our opinion both S. coccineus type specimens are of doubtful status. Bernard, Cai & Morton (1993) introduced the replacement name S. punicus for this species. They recorded the species as occurring in China but gave no reference to the specimen(s) on which they based their identification. ACKNOWLEDGEMENTS We would like to thank the following, museum staff and collectors for their assistance. Ms Kathie Way, malacology section of The Natural History Museum, London for the loan of specimens and access to the type collection; Dr Philippe Bouchet, Muséum National d’Histoire Naturelle, Paris for permission to access the type collection and loan of type material; Dr Claude Vaucher and Dr Yves Finet, Muséum d’ Histoire Naturelle, Geneva for access to the type collection; Mr Ian Loch, Malacology Section of the Australian Museum, Sydney for the loan of specimens. Mr Chris Takahashi, Hawaii donated Hawaiian Spondylus specimens. The Malacol- ogical Society of Australasia provided travel assistance to KL. LITERATURE CITED ABBOTT, R.T. & DANCE, S.P. 1982. Compendium of seashells. (E.P. Dutton: New York). BERNARD, F.R., CAI, Y. & MORTON, B. 1993. Catalogue of the living marine bivalve molluscs of China. (Hong Kong University Press: Hong Kong). CHEMNITZ, J.H. 1784. Neues systematisches conchylien-cabinet. Vol 7. (G.N. Raspe: Niirnberg). CHENU, J.C. 1845. Illustrations conchyliologiques Vol. 3. Bivalves. Part 2. DALL, W.H., BARTSCH, P. & REHDER, H.A. 1938. A manual of the Recent and fossil marine pelecypod mollusks of the Hawaiian Islands. Bulletin of the Bishop Museum 153: 1-233. FULTON, H.C, 1915, List of the Recent species of Spondylus Linné with some notes and descriptions of six new forms. Journal of Conchology 14: 331-360. KAY, E.A. 1979. Hawaiian marine shells. Reef and shore fauna of Hawaii. Section 4, Mollusca. B.P. Bishop Museum Special Publication 64 (4): 1-653. 628 LAMPRELL, K.L. 1986. Spondylus, spiny oysters of the world. (E.J. Brill-Dr W. Backhuys: Leiden). 1992a. Notes on Spondylus Linnaeus with descriptions of two new species from Western Australia (Mollusca: Bivalvia: Spondylidae). Memoirs of the Queensland Museum 32: 189-194, 1992b. On some species of Spondylus. La Conchiglia 23: 40-41. LAMPRELL, K.L. & DEKKER, H. In press. Rectification of nomenclature and notes on species of Spondylus Linnaeus, from the Chemnitz Cabinet and held in the Zoological Museum, University of Copenhagen (Mollusca: Bivalvia: Pectinoidea: Spondylidae). LAMPRELL, K.L. & KILBURN, R.N. 1995. The recent Spondylidae of South Africa and Mozambique, with the description of a new species (Mollusca, Bivalvia, Pectinoida). Molluscan Research 16: 81-95. LAMPRELL, K.L. & WILLAN, R.C. 2000. Rectification of nomenclature for three species of Spondylus Linnaeus (Bivalvia: Pectinoidea: Spondylidae) from the Indo-Pacific based on re-examination of type specimens. Vita Marina 47: 1-7. LAMPRELL, K.L. & HEALY, J.M. 2001. The Recent Spondylidae from New Caledonian and adjacent waters, with the descriptions of 8 new species, and MEMOIRS OF THE QUEENSLAND MUSEUM some taxonomic notes on the species Spondylus anacanthus (Mawe) (Mollusca, Bivalvia, Spondylidae). Bulletin du Muséum National d’Histoire Naturelle. Paris (Zoologie) 22. LAMY, E. 1927. Les Spondyles de la Mer Rouge (d’apreés les matériaux recueillis par le Dr Jousseaume). Bulletin du Museum National d’Histoire Naturelle 33: 259-266; 301-308. LINNAEUS, C. 1758. Systema Naturae, per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 10th edition. (Holmiae: Laurentii Salvii). REEVE, L.A. 1856. Monograph of the genus Spondylus. Conchologia Iconica or illustrations of the shells of molluscous animals, Vol. 9. (Reeve: London). SCHREIBERS, K. 1793. Versuch einer Vollstandigen Conchylienkenntniss nach Linné’s System. (Vienna). SKOGLUND, C. & MULLINER, D.K. 1996. The genus Spondylus (Bivalvia: Spondylidae) of the Panamic Province. The Festivus 28: 93-107. SOWERBY, G.B. IT 1847. Descriptions of several new species of Spondylus. Proceedings of the Zoological Society of London 1847: 86-88. SPRINGSTEEN, F.J. & LEOBRERA, F.M. 1986. Shells of the Philippines. (Carfel Seashell Museum: Manila). A BREEDING POPULATION OF THE YELLOW-BELLIED SEA SNAKE, PELAMIS PLATURUS, IN THE GULF OF CARPENTARIA COLIN J. LIMPUS Limpus, C.J. 2001 06 30: A breeding population of the Yellow-bellied sea-snake, Pelamis platurus, in the Gulf of Carpentaria. Memoirs of the Queensland Museum 46(2): 629-630. Brisbane. ISSN0079-8835, A surface aggregation of Pelamis platurus was observed on 13 July 1992, along a 99.4km line between 14.59°S, 140.73°E and 15.34°S, 140.25°E within the Gulf of Carpentaria. Recently born, immature and adult sized snakes provide the first evidence of a resident breeding population in northern Australian waters. 1 Sea snake, Gulf of Carpentaria, breeding, Pelamis. Colin J. Limpus, P.O. Box 541, Capalaba 4147, Australia; 10 April 2000. The Yellow-bellied Sea Snake, Pelamis platurus, is widely distributed in Australia as beach-washed specimens but is poorly documented in its marine habitat (Cogger, 1975; Guinea, 1992). Shuntov (1972) implied that P platurus in northern Australian waters were transported from the Coral Sea, whereas Guinea (1992) suggested they could be ‘part of a geographically discrete, yet seldom encountered, breeding population’. METHODS Observations of sea snakes were made from the bow of the One and All, a twin masted, square rigged, 42.7m brigantine under sail en route from Weipa to the Wellesley Group. A bow watch during daylight hours recorded marine reptiles, mammals and birds. With each sighting, a latitude, longitude and water depth (corrected for depth of sensor below the waterline) were recorded from the ship’s navigational instruments. No animals were captured. Surface water was sampled by casting a bucket over the side and hauling it in quickly, for an immediate recording of water temperature using a quick reading, mercury in glass thermometer (+0.1°C). All species identifications were made by the author while crew members assisted in locating wildlife. The uniquely conspicuous colouration of P. platurus (Heatwole, 1975a) contributed to the ease of identification. RESULTS A total of 112 sea snakes were seen in the 6 days. Of these 84 (75.0%) were P. platurus. All P. platurus were observed on 13 July 1992, along a 99.4km line between 14.59°S, 140.73°E and 15.34°S, 140.25°E (Table 1). All sightings occurred between 0936-1730hr. TL estimates for 32 P. platurus that passed close to the bow ranged from 25-60cm with 88% <45cm. At locations where P. platurus was recorded, mean water depth was 47.6m (SD=1.04, range=43-49m, n=35) and mean surface water temperature was 24.7°C (SD=0.05, range=24.7-24.8°C, n=6). Most P. platurus did not dive on close approach of the ship. Other species of sea snakes were more difficult to identify, often because they dived on the ship’s approach. Of the 28 non-Pelamis sea snakes observed (Table 1), only 3 Lapemis hardwickii, including a copulating pair, 6 Hydrophis elegans and | Disteira kingii were identified. On the transect with P platurus were post hatchling marine turtles Chelonia mydas and Natator depressus with carapace length ~15cm. While it is well documented that P. platurus and post hatchling marine turtles aggregate along drift lines where currents converge (Kropach, 1975; Carr, 1987), there were no signs (floating debris) in this case, of a convergence zone. DISCUSSION P. platurus is the most widely distributed sea snake species globally, with records extending from Possiet Bay in southern Siberia to New Zealand and Tasmania and from the Cape of Good Hope to Panama (Minton, 1975). However, the species rarely inhabits waters <20°C and its upper lethal temperature for indefinite survival is ~33°C (Dunson & Ehlert, 1971); while it is tolerant of low salinities, it does not occur in coastal waters subject to high variability in salinity (Dunson & Ehlert 1971). Cogger (1975) postulated a permanent breeding population off central NSW coast where surface waters reach 17°C in winter. However, Hecht et al. (1974) identified the 26°C isotherm as delimiting the 630 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE I. Description of transects over which searches for sea snakes were conducted from the tall ship One and All in the Gulf of Carpentaria during 12-22 July 1992. Date Start Finish | Sea snakes sighted 7 a Transeet > 1992 time latitude longitude | depth time latitude ongitude | depth length | p. platurus cose 12 July 1105hr 12.67°S 141,77°E | 123m | 3759hr | 13.15°S 141.62 °E 64km 0 4 13 July | O71Shr | 14.40°S 140.87°E | 48m 1800hr | 15.40°S | 140.23°E | 43m 130km 84 16 14 July | 0930hr 16.62°S_| 139.83°E | 23m 1100hr | 16.50°S 39.84°E | 23m ISkm 0 0 17 July | 1440hr 16.08°S_ | 139,.20°E | 27m 1855hr_ | 15.56°S | 139.13 °E | 48m 63km 0 3 18 July | 0730hr 14.43°S 138,33°E 58m 1850hr | 13.62°S | 136.93°R | 28m (77kan 0 __ 4 22 July | 1031hr 12.07°S_| 136.77°E | 20m 1200hr | 11.87°S 36.75°E | 35m 22km | 0 0 distribution of permanent breeding colonies. P platurus is a pelagic sea snake that aggregates in association with slicks and drift lines (Dunson, 1975), Aggregations are not seasonal although calm weather favours their formation (Kropach, 1975). P. platurus is piscivorous, in the top 2m rather than at the bottom like most sea snakes (Kropach, 1975); has been recorded diving to c. 20m (Kropach, 1975) and can adjust buoyancy to suit activities (Graham et al. 1975); cannot regain the sea if beach-washed (Dunson, 1975). In the Gulf of Panama P platurus has: TL at birth = 22-26cm; dd sexually mature at TL ~50cm; 2 2 sexually mature at TL ~ 64cm; non- seasonal breeding (Kropach, 1975). By analogy, the eastern Gulf of Carpentaria population encompassed all size ranges from recently born to adult, with the majority being immature, thus demonstrating a breeding population, the first recorded for northern Australia. Although sea snakes have been well studied in the Gulf of Carpentaria P. platurus has rarely been reported (Guinea, 1992; Shuntov, 1972; Heatwole, 1975b; Wassenberg et al., 1994). The Gulf of Carpentaria is in a region that Hecht et al. (1974) predicted should support a permanent breeding colony, although its water was slightly cooler than identified as limiting permanent breeding populations. The extent and habits of this population are yet to be determined. ACKNOWLEDGMENTS Penny van Oosterzee and Noel Preece of Discovery Ecotours provided the opportunity to participate in the expedition and provided logistical support. The crew and guests assisted in data collection. This assistance is appreciated. LITERATURE CITED CARR, A. 1987. Impact of non degradable marine debris on the ecology and survival outlook of sea turtles. Marine Pollution Bulletin 18(6B): 352-6. COGGER, H.G. 1975. Sea snakes of Australia and New Guinea. Pp. 59-140. In Dunson, W.A. (ed.) The biology of sea snakes. (University Park Press: Baltimore), DUNSON, W.A. 1975. Adaptations of sea snakes. Pp. 3-20. In Dunson, W.A. (ed.) The biology of sea snakes. (University Park Press: Baltimore). DUNSON, W.A. & EHLERT, G.W. 1971. Effects of temperature, salinity and surface water flow on the distribution of the sea snake Pelamis. Limnology and Oceanography 16: 845-53. GRAHAM, J.B., RUBINOFF, I. & HECHT, M.K. 1975. Temperature physiology of the sea snake Pelamis platurus. An index of its colonisation potential in the Atlantic Ocean. Proceedings of the National Academy of Sciences 68: 1360-3. GUINEA, M.L. 1992. The yellow-bellied sea snake Pelamis platurus in the Northern Territory. Northern Territory Naturalist 13: 37-39. HEATWOLE, H, 1975a. Predation on sea snakes. Pp. 233-250. In Dunson, W.A. (ed.) The biology of sea snakes. (University Park Press: Baltimore). 1975b. Sea snakes of the Gulf of Carpentaria. Pp. 145-50. In Dunson, W.A. (ed.) The biology of sea snakes. (University Park Press: Baltimore). HECHT, M.K., KROPACH, C. & HECHT, B.M. 1974. Distribution of the yellow-bellied sea snake, Pelamis platurus, and its significance in relation to the fossil record. Herpetologica 30: 387-396. KROPACH, C. 1975. The yellow-bellied sea snake, Pelamis, in the eastern Pacific. Pp. 185-216. In Dunson, W.A. (ed.) The biology of sea snakes. (University Park Press: Baltimore). MINTON, 8.A. 1975. Geographic distribution of sea snakes. Pp. 21-32. In Dunson, W.A. (ed.) The biology of sea snakes. (University Park Press: Baltimore). SHUNTOV, V.P. 1972. Sea snakes of the North Australian Shelf. Soviet Journal of Ecology, New York 2(4); 338-44. Translation of 1971 paper in Ekologiya 4: 65-72. WASSENBERG, T.J., SALINI, J.P., HEATWOLE, H. & KERR, J.D. 1994. Incidental capture of sea snakes (Hydrophiidae) by prawn trawlers in the Gulf of Carpentaria, Australia. Australian Journal of Marine and Freshwater Research 45; 429-43, THE LOGGERHEAD TURTLE, CARETTA CARETTA IN QUEENSLAND: FEEDING ECOLOGY IN WARM TEMPERATE WATERS COLIN J. LIMPUS, DEIDRE L. DE VILLIERS, MICHELE A. DE VILLIERS, DUNCAN J. LIMPUS AND MARK A. READ Limpus, C.J., de Villiers, D.L., de Villiers, M.A., Limpus, D.J. & Read, M.A. 2001 06 30: The loggerhead turtle, Caretta caretta in Queensland: feeding ecology in warm temperate waters. Memoirs of the Queensland Museum 46(2): 631-645. Brisbane. ISSN 0079-8835. Gut contents and faecal samples from 53 loggerhead turtles, Caretta caretta, from the Bundaberg coast, Hervey Bay, Sandy Straits, Moreton Bay and Gold Coast regions of southern Queensland continental shelf waters were examined. C. caretta in these coastal waters are carnivorous, consuming at least 94 taxa of benthic and near benthic organisms. Large immature and adult C. caretta are specialised for feeding on slow moving, hard bodied invertebrate prey with molluscs and crustaceans being the most commonly consumed taxa. Four feeding methods were identified for these C. caretta. The specific prey species selected was a function of the turtle’s feeding area rather than its sex or size. As individuals their diet is unpredictable with some variability in the diet being attributed to individual preference. 4 Loggerhead turtle, Caretta, feeding ecology, southeast Queensland. Colin J. Limpus, Michele A. De Villiers, Duncan J. Limpus, Queensland Parks and Wildlife Service, P.O, Box 155 Brisbane, 4002; Deidré L. De Villiers, Queensland Parks and Wildlife Service, P.O. Box 5116, Daisy Hill, 4127; Mark A. Read, Queensland Parks and Wildlife Service, P.O, Box 5391, Townsville M.C., 4810, Australia; 8 May 2000. The loggerhead turtle, Caretta caretta, is carnivorous and feeds on a very wide range of prey species. While it preys primarily on benthic invertebrates such as crustaceans, bivalves and gastropods, it consumes many other taxa including jellyfish, sea pens, sea urchins, holothurians, tunicates, and fish (Bleakney, 1967; Burke & Standora, 1993; Ernst & Barbour, 1989; Gudynas, 1980; Plotkin, 1996; Porter, 1972; Preen, 1996). Some of this diversity in diet is a function of the life history phase of the turtles. C. caretta, typical of cheloniid turtles, occupies a wide range of habitats throughout its life history (Carr, 1986; Dodd, 1988; Limpus, 1985, 1994), Eggs are laid in sandy tropical and warm temperate beaches. The hatchlings leave the beaches and disperse within days into deep water from where they enter a pelagic phase, being dispersed by ocean currents. For the first years of their life they occupy open ocean surface waters before recruiting to live in widely dispersed feeding areas over the continental shelf. Breeding adults migrate from their feeding areas to their traditional breeding sites and occupy courtship and internesting habitats within the waters adjacent to the nesting beaches for some months during each breeding season. At the completion of the breeding season they return to their respective feeding areas. Feeding by C. carefta is largely restricted to the pelagic phase where the young turtles utilise planktonic prey at or near the ocean surface (Plotkin, 1996; van Nierop & den Hartog, 1984) and inshore shallow waters where the larger sized turtles feed predominantly on benthic prey (Conway, 1994; Plotkin et al., 1993; Moodie, 1979). The breeding adult does not feed while ashore for egg laying. Similarly, while in the courtship and internesting habitats, the breeding female does not feed or substantially reduces her feeding, while she is in this egg production phase (CJL unpubl. data). The hatchlings do not feed while in the nest, while crossing the beach or in the inshore waters as they disperse from the nesting beach. The eastern Australian C. caretta stock (Bowen et al., 1994) is endangered with a declining breeding population (Limpus & Reimer, 1994). As part of general studies to understand their biology, we describe the diet of adult and large immature C. caretta feeding in inshore warm temperate waters of southeastern Queensland. METHODS Gut contents were obtained opportunistically from C. caretta from southeastern Queensland during 1989-1998. The study area extended from the Kolan River near Bundaberg (24°35’S, 152°07’E) to the Gold Coast (28°02’S, 153°26’E). Habitats utilised by C. caretta encompass rocky reefs, bays, estuaries and coastal open waters. 632 Free ranging (. earentee were capoired durmy mark-recaplure studies (Limpus. 1978, 1985: Limpus ct al. 1994), Dead and moribund C. caretia were obtained through the Queensland Parks and Wildlite Service (QPWS) marine wildlife stranding program. Faecal samples were obtained from the live turtles, Digestive tract contents were oblained during necropsy of’ the turtles from the stranding program. The amount of gut contents collected was dependent on the state of decomposition of the carvass but. where possible, the entire alimentary tract was sampled. For each sample, the turile’s sex, age class, breeding status, midline curved carapace length (CCL), injuries, cause of death, date, and location were recorded, Carcasses were selected only if the body organs were intact. While the turtles were not feeding at the point of stranding, given the limiton state of decomposition, itis presumed that they would have fed in the adjacent waters. For case of description, gut contents are identified by the (ag number of (he turtles or the museum specimen numbers. All except one sample were from non breeding turtles in or adjacent to their presumed feeding areas. No suh-sampling was taken of large samples. Samples with fleshy biomass were fixed in 5$-10% formalin Solution. Samples of predominantly hard skeletal remains, including molluse and crustacean exoskeletons, were dried und stored. Prior to sorting, samples which had been stored in formalin were rinsed in freshwater and spread on sorting trays to remove most ofthe moisture, Prey items from the Brachiopoda, Mollusca, Crustacea and QOsteichthyes were identified to species or genus level where possible. Prey items from the Povifera, Cnidaria and Echinodermats were identified at higher laxononic levels. No attempt was made to identify the algae and seaprass. As each prey taxon was identified, the number of individuals present was counted using identifying features of fragments that temained intact through the feeding and digestive process, Decapod crustaceans were counted by the number, size, and orientation (leti/rizht) of chelae, and toa lesser extent, mandibles. The number of hinges and the orientation of valves assisted with bivalve counts, With gastropods, intact spires were the primary indicator of the number of individuals consumed, bul at times, counts of opercula (e.g. Yurbo spp., Sitrombus spp.) were more appropriate, Sand, stone, coral fragments, dead shell (identitied by eroded and dull inner shell surfaces), charcoal and tree bark were treated.as MEMOIRS OF THE QUEENSLAND MUSI\UM imcidentally ingesi¢d debris, Queensland Museum staf!’ and private shell collectors assisted with the identification of prey items, Por unalysis, the sumples were grouped by geographic origin: Bundaberg coast, [lervey Bay, Sandy Straits, Moreton Bay and Gold Coast. For the purposes of this analysis, a dominant prey speoies was delined as one comprising =5'% ofa sample by either the number of individual prey ilems present or by volume. Volume was used in the analysis of'six samples where there were very large differences in the sizes of prey species (see Tables 2.4). RESULTS Fifly-three Samples included 6 faecal samples and 47 digestive tract contents, The geographic origins of samples were the Bundaberg Coast (el 6), Hervey Bay (Woodgate + Burrum Heads (3); Sandy Cape (2), Sandy Straits (2), Moreton Bay (22), and Gold Coast - Jumpinpin (8). Prey from at least 94 taxa representing § Phyla were identified (Table |): 36 species of crustaceans, 43 molluses (18 gastropod, 23 bivalve, | scaphopod, | cephalopod), | poriferan, 2 cnidarians, | brachiopod, 3 echinoderms, | urochordate, and at least 7 Osivichthyes, Most food items were crushed and the soft tissues and some or all of the skeletal fragments ingested. A small range of items were myested whole, including some fish, héche-de-mer and small molluses. The largest intact prey was a porcupine fish (Family Diodontidae, 2)em), lodged in the oesophagus, and which probably caused the death of the turtle (lag=Z1819), The smallest intact prey were the gastropod Olive caldania (dem) and the scaphopod Denfalium sp. (2.4em), both in the same turtle, TS9145. [t is presumed that these shells had been ingested incidentally and not crushed when the turtles had targeted larger prey, Other taxa that were considered to be ingested accidentally were seagrass and barnacles: there were only isolated blades of seagrass in the gut contents from 4 C. cured, Some barnacles were attached to other largeted prey species such as portunid crabs. The greatest abundance of prey from an entire put content was 670 individuals and the largest number of a single species was 565 Sole vagenoides (Chinese fingernail shell). Where there were >400 prey items occurred ina sample, they primarily consisted ofa single species: 36'%o of samples comprised =90% of a single species and 15% of samples were made up of a single species, LOGGERHEAD TURTLE IN QUEENSLAND 633 TABLE 1. Caretta caretta prey species by regions within subtropical Queensland. The value in each cell denotes the number of turtles recorded with each prey species. The percentage range that each prey species contributed to a turtle’s diet sample is shown in parenthesis. * indicates material which was considered incidental and not included in the analysis of prey species; # identified fish included: porcupine fish (Diodon), flathead (Platycephalus), bigeye (Priacanthus), flounder (Pseudorhombus), whiting (Si//ago) and wrasse (Labridae). Sandy Family Goneplacidae Family Leucosiidae Family Parthenopidae Family Pinnotheridae Family Portunidae Family Raninidae Family Xanthidae Eucrate dorsalis Galene bispinosa Galene cf bispinosa Leucosia sp. Myra affinis Myra sp. Cryptopodia queenslandi Parthenope nodosa Parthenope (?) valida Xenophthalmus pinnotheroides Charybdis natator Portunus pelagicus Portunus sanguinolentus Thalamita sima portunid crab, unidentified 1 portunid crab, unidentified 2 portunid crab, unidentified 3 portunid crab, unidentified 4 portunid crab, unidentified 5 Ranina ranina Halimede ochtodes crab, unidentified 6 crab, unidentified 7 crab, unidentified 8 crab, unidentified 9 unidentified (fragment) 1(0.2%) 8(0.2-7.1%) 4(0.6-6.9%) 1(1.7%) 5(0.1-4.5%) 8(0.2-31.8%) 1(0.2%) 3(0.5-6.9%) 1(2.89%) 4(1.4-25%) 1(3.8%) 1(0.5%) 1(1.72%) 1(0.5%) | 2(0.5-5.2%) 1(0.5%) 1(1.72%) 1(1.4%) 1(33%) 2(7.7-8.3%) 1(10%) 1(1.3%) 1(25%) 1(1.3%) 2(25-90%) | | 12(0.6-100%) 8(0.6-55%) 1(1.6%) | 2(12.5-16.7%) 1(0.6%) 4(3.5-100%) 11(0.2-50%) 3(0.2-48.4%) 3(2.38-50%) 1(17.5%) 1(5%) 2(1-50%) Taxa Genus/Species/Common name | Bundaberg Hervey Bay | Straits Moreton Bay | Gold Coast No. turtles in sample n=16 n=5 | n=2 n=22 n=8 Phylum ARTHROPODA | Class Cirripedia * barnacle, unidentified 2(1.1-1.3%) | 2(0.3-4.3%) 11.4%) | 7(0.2-1.4%) 1(<1%) Class Malacostraca | cn. Order Decapoda | Infra-order Anomura | Family Diogenidae Clibanarius sp. 9(2.7-31.8%) Clibanarius taeniatus | 1(0.4%) Dardanus imbricatus 12(5-90.6%) 1(16.7%) | _ | 116.7%) | Infra-order Brachyura | | Family Calappidae Calappa (?) hepatica 1(2.2%) Calappa sp. 2(2.7-4.5%) | Matuta sp. 1(8.3%) | 1(20%) 3(2.5-20%) 2(2.5-5%) 3(2.5-20%) 1(5%) 4(1.7-40%) Infra-order Penaeidae penaeid prawn — 1(<5%) Order Stomatopoda Squilla sp. 3(3.1-10%) Phylum BRACHIOPODA Family Lingulidae Lingula murphiana 1(90%) TABLE 1 (cont.). MEMOIRS OF THE QUEENSLAND MUSEUM Family Turbinidae Turbo haynesi 1(3.1) Taxa Genus/Species/Common name | Bundaberg Hervey Bay aie Moreton Bay | Gold Coast Phylum CHORDATA Class Ascidiacea ascidian, unidentified 2(14.2-22.2%) | Class Osteichthyes fish, multiple species # 1(100%) 1(25%) 3(2.4-50%) 8(20-95%) Phylum CNIDARIA Class Anthozoa Order Gorgonacea sea whip, unidentified 1(20%) Order Actiniaria anemone, unidentified 1(0.6%) Phylum ECHINODERMATA Class Echinoidea sea urchin, unidentified | 1(100%) 2(0.6-2.2%) Class Holothuroidea béche-de-mer, unidentified 1(5%) Class Stelleroidea starfish, unidentified 1(90%) | Phylum MOLLUSCA Class Bivalvia Order Arcoida Family Arcidae Anadara trapezium 3(0.2-18.4) 1(6.3) 1(100) 2(60.7-100) Family Glycymeridae | Glycymeris holsericus 1(2.6) Order Mytiloida Family Anomiidae Patro australis 1(25) Family Mytilidae Botula sp. (elong. shiny mussel) 1(4.5) Modiolus ostentus 1(100) Stavelia horrida 1(2.6) Trichomya hirsuta 2(18.8-100) Family Ostreidae Crassostrea commercialis 1(10.7) Ostrea bresia 1(0,2) Saccostrea commercialis 1(0.7) Family Pectinidae Annachlamus flabellata 3(1.3-33) 1(4.5) Amusium balloti 4(0.5-30.4) Family Pinnidae Atrina pectinata 6(0.2-63) 1(87) 2(3.6-33.3) Family Pteriidae Pinctada albina sugillata 1(2.6) Pinctada fucata 2(1.3-8.3) | Family Spondylidae — | Spondylus wrightianus 1(1.4) | - oyster fragment 1(5) Order Veneroida Family Carditidae Cardita incrassata 1(10.7) Family Mactridae Mactra abbreviata 1(2.6) Family Solenidae Solen vaginoides 1(99.5) Family Tridacnidae Tridacna maxima Family Veneridae Antigona lamellaris 2(1.4-2.6) 1(2.6) 3(3.6-75) Family Crassatellidae | Eucrassatella cumingii 1(2.6) 1(2.6) 1(16.7) - unidentified (fragment) 1(0.6) 2(0.2-0.6) Class Cephalopoda octopus, unidentified 2(1.7-2.5) Class Gastropoda Order Archaeogastropoda Family Trochidae Calthalotia indistincta 1(0.7) Monilea callifera 1(90) LOGGERHEAD TURTLE IN QUEENSLAND 635 TABLE 1 (cont.). Taxa Genus/Species/Common name | Bundaberg Hervey Bay gandy Moreton Bay | Gold Coast Order Mesogastropoda _ = Family Cerithidae | Pyrazus ebeninus 1(5) Family Naticidae Polinices conicus 1(0,.2) Polinices didyma 1(0.6) Family Potamididae Velacumantus australis 1(98.6) Family Strombidae Strombus campbelli 6(3.3-85.6) 1(91.1) Family Tonnidae Tonna tessellata | 1(20) Order Neogastropoda Family Buccinidae Dolicholatirus thesaurus 2(0.5-1,8) | Family Fasciolarinae | Fusinus colus (0.6) Family Muricidae Lataxiena fimbriata (1.2) Family Olividae Oliva caldania (0,2) | Family Volutidae Amoria maculata (0.2) | | { Cymbiolacca complexa (0.2) | | | Melo amphora 1(95) | Melo sp. (<5) __| unidentified fragment (0.1) 1(5) Class Scaphopoda | — 7 = | _ Family Dentaliidae | Dentalium sp. 10.1) Phylum PORIFERA | sponge, unidentified 1(2.6) 3(7-50) Other |* algae <1) 1(0.3) 7 : * seagrass 2(<1) —_— 4(<1) A faecal sample comprising both valves of the hairy mussel, Modiolus ostentatus, was obtained from a nesting ¢ C. caretta during oviposition at Mon Repos near Bundaberg (X37114). This was the only identifiable item obtained as faecal material from the many thousands of nesting females observed. This prey species was not found in any other turtle sample. Given that Bun- daberg is outside the geographical distribution of this tropical bivalve species (Lamprell, 1998) and that breeding female marine turtles do not feed, or substantially reduce their food uptake, while away from their home feeding areas during their breeding migrations (C. Limpus, unpubl. data), it is highly unlikely that this food item originated from the Bundaberg coast. Hence this sample was excluded from the following analysis. Bundaberg coast. Fifty two species were identified in 14 C. caretta samples from the Bundaberg region (Table 1). The mean number of prey species per sample was 8.4 (SD=4.7, range=2-16). Of the 52 prey species only 14 (26.9%) were dominant prey items (Table 2). These C. caretta fed mostly on hermit crabs (12/14 individuals), gastropods (5/14 individuals), bivalves (3/14 individuals) and small brachyuran crabs (5/14 individuals). For nine of these turtles the sample comprised many specimens of multiple species of prey. In contrast, for three individuals, a single large item (gastropod [Melo amphora], sea urchin and starfish, respectively) dominated the sample. One turtle had many specimens of the hermit crab, D. imbricatus, comprising >90% of the gut content. The dominant prey items from the Bundaberg Coast (Table 2) were benthic species that live on or superficially burrow into the substrate. These dominant prey species were slow moving with the exception of the more agile saucer scallop, A. balloti. Hervey Bay. Sixteen species were identified in the gut contents of five C. caretta from the Hervey Bay region (Table 1). The mean number of prey species per sample was 3.8 (SD=1.0, range=2-5). Of the 16 prey species, 12 (75%) were dominant prey items (Table 3). These C. caretta fed mostly on bivalves (4/5 individuals), crabs (4/5 individuals) or hermit crab (1/5 individuals) With one exception, this geographic group consumed 636 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 2. Relative abundance by number of individuals of prey items identified in the gut contents of beach washed Caretta caretta (n=14) from the Bundaberg Coast. All samples were obtained from turtles that had been feeding immediately prior to their death. Relative abundance values are summarised only for those species comprising >5% of the total sample for the turtle. y’ denotes that the species was present in the gut sample at less than the 5% level. Other species present at less than 5% ofa gut content within all samples are not listed. * Where there was a very large difference in the size of prey items the abundance has been adjusted to reflect the relative volume of the prey species. Tag number - N39890 N39891 N39925 N39944 N39961 N39970* N39975, | Date 14.02.94 18.02.94 07.02.94 04.02.94 _ 15.01.94 25.01.93 12.01.93 Sex Q g g g | g g | g | Maturity Adult Immature | Adult Immature Immature Immature Adult |Carapace length (cm) 94.7 83.5 93.5 75.5 82.0 83.0 104.5 Latitude 24°52’S 24°50’S 24°49’S 24°52’S 24°47’S 24°58’S 24°49’S Longitude 152°29°E 152°28°E 152°27°E 152°28°E 152°26’E 152°29"E 152°28°E Prey Items Mollusc, gastropod Strombus campbelli | 86% y 10% Melo amphora Mollusc, bivalve Ambusium balloti 30% y 51% Anadara trapezium y | __ Atrina pectinata y = - y 5 Crustacea, hermit crab Dardanus imbricatus 22% 5% 18% 70% 50% 5% 17% Clibanarius sp. 29% | 8% 32% y 5% 5% 10% Crustacea, brachyuran crab Charybdis natator y 7% Halimede ochtodes y 5% Leucosia sp. y y y 5% 7% Myra affinis y y T% Parthenope nodosa y y 32% _ T% 12% y | Echinoderm, starfish 90% Echinoderm, sea urchin | = z Tag number N39979* N39985 N39993 T85156 T89145 T89184 Z2029 Date 06.01.93 15.12.92 28.11.92 15.01.95 06.01.96 01.06.96 26.11.96 Sex = g 3 3 io} 3 3 | Maturity Immature Immature Immature Immature Immature Immature Immature _Carapace length (cm) | 83.0 78.0 85.5 82.7 91.5 91.5 95.1 Latitude 24°48’S 24°48’S 24°47’S 24°58’S 24°40’S | —_-24°40°S 24°43’S Longitude 152°26°E 152°27°E 152°26°E 152°29°E 152°13°E 152°13°E 152°17°E Prey Items Mollusc, gastropod Strombus campbelli 41% 30% | 81% | Melo amphora 95% Mollusc, bivalve Ambusium balloti | y Anadara trapezium | y 18% Atrina pectinata = y | y 63% Crustacea, hermit crab | Dardanus imbricatus 5% 43% 91% 27% 11% Clibanarius sp. 9% 31% Crustacea, brachyuran crab | Charybdis natator y | Halimede ochtodes Leucosia sp. y y 5% Myra affinis y |__Parthenope nodosa y y Echinoderm, starfish Echinoderm, sea urchin 100% LOGGERHEAD TURTLE IN QUEENSLAND few prey species, usually with only one species dominating the sample. For example, a single prey species constituted ~90% of the gut content (A. pectinata, P. sanguinolentus, or Lingula murphiana). The prey for this region represented a range of habitats: within the water column (fish), on or superficially burrowed within the substrate surface (A. pectinata and crabs) or burrowed deep within the substrate (L. murphiana). That 72 L. murphiana were the only species present in | sample indicates that the turtle was intentionally feeding on this cryptic species to the exclusion of other species. Sandy Straits. Four species were identified in the 2 gut con- tents from the Sandy Straits region (Table 1). The number of prey species per sample ranged 1-3, Two species (50%) were dominant prey items (Table 3). The 2 C. caretta had fed almost exclus- ively on either the gastropod Velacumantus aust- ralis, or the bivalve A. trapezium. Both prey species are epibenthic or superficially burrow. Moreton Bay. Thirty-four prey species were identified in 22 C. caretta samples from the Moreton Bay region (Table 1), The mean number of prey species per sample was 3.8 (SD=2.2, range=1-7). Of the 34 prey species, 27 (79%) were dominant prey items making up >5% of any one gut content (Table 4). The C. caretta in Moreton Bay had been feeding mostly on crabs (14/21 individuals), especially portunid crabs. When molluses occurred in the sample, they were the major component of the prey (8/21 individuals) and a single species of gastropod or bivalve dominated, Other benthic animals (sea cucumber, sea urchin, sea whip, ascidian and sponge) were taken commonly and many C. caretta consumed a wide range of prey species. However, 8 of the 21 turtles ingested a very limited range. Six had gut contents approaching 100% ola single species of bivalve or crab; two had gut contents with a single gastopod species accounting for ~90%. The majority of the prey items were benthic species that live on or superficially burrow within the substrate, except for the bivalve S. vagenoides which burrows well below the surface. Most of these prey species are slow moving, although the fish and stomatopod are active species. Gold Coast - Jumpinpin. Ten species groups were identified in the gut contents of eight C. caretta from the Gold Coast - Jumpinpin region (Table 1). Of these, six (75%) were dominant prey items (Table 5). The C. caretta sampled fed predominantly on small fish (8/8 individuals) with six of eight turtles consuming a diet consisting of >90% fish. While some species of fish could be recognised (Table 5) most material could not be identified and is grouped as all fish. Crabs contributed significant amounts to the overall volume of prey ingested by 2/8 individuals. A single specimen of the gastropod Jonna tessellata was a major component of the diet of one turtle. The principal prey items in this area were highly mobile species that live close to the substrate and in the adjacent water column (fish). A small amount of the slow moving benthic species that live on the substrate surface or superficially burrow were eaten also. There was circumstantial evidence (unpublished data, QPWS stranding database) that at least several of these turtles had been killed during fish trawling activities off this coastline. The prey items identified from these turtles are common among the ‘trash fish’ discarded from this trawl fishery. There is a high probability that these turtles were scavenging the water column or the substrate for discarded dead bycatch. COMPARISON AMONG REGIONS. Molluscs and crustaceans dominated the diet of C. caretta in most areas but not inthe Gold Coast region. However, the dominant species in the diet varied spatially. For example, on the Bundaberg coast the diet included a range of hermit crabs, a gastropod (S. campbelli) and a bivalve (A. balloti) for the major part of the diet (Table 2). The diet in Moreton Bay was dominated by a range of brachyuran crabs (not hermit crabs) and bivalves (not A. halloti) (Table 4). Within a localised area, many turtles displayed idiosyncratic feeding patterns, choosing to feed on individually unique suites of prey. Over half (54.7%) of C. caretta in this study had consumed < 5 species of prey. The turtles from the Bundaberg region contained more prey species per sample (8.4 prey) while the turtles from Moreton Bay and Hervey Bay had similar numbers of prey (3.8 prey species per sample) (1 way ANOVA: Fi3g = 9.57; p<0.001). The comparison excluded Sandy Straits because of the small sample size and the Gold Coast - Jumpinpin samples because it was impossible to count the masses of fish bone in the gut contents. The size, sex and maturity of the sampled turtles are listed in Tables 2-5. There were 27 males and 21 females and two unsexed turtles in the study group. The prey ingested was unrelated to sex or size because there was no significant 638 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 3. Relative abundance by number of individuals of prey items from Caretta caretta gut contents (G) of beach washed turtles or faecal samples (F) from captured wild turtles from Hervey Bay (n=5) and Sandy Straits (n=2) areas. All samples were obtained from turtles that had been feeding immediately prior to their death. Relative abundance values are summarised only for those species comprising >5% of the total sample for the turtle. The sample from specimen T79317 was a faecal sample. The remainder were from gut contents. Species present at less than 5% of a gut content are not listed. Hervey Bay 7 Sandy Strait ‘Tag number Q18573 T57853 Q22481 2322 179317 122833 Date 11.01.96 | 25.11.95 27.01.98 19.12.96 24.09.92 13.06.95 | 11.06.95 | Sex ; 3 3 é é. |e 9 Q | Maturity Immature Immature Immature Immature Immature Immature Adult | Carapace length (cm) 79.4 80.9 95.2 83.4 104.0 71.0 94.1 Latitude 25°11'°S 25°09°S 25° 08'S 24°43’S_ | 25°01"S__ | __25°42°S 25°45’S Longitude 152°37E | 152°37"E 152°35°E [53°27 EK 153°2 VE 15°55°E 152°S7°7E Prey Items | Mollusc, bivalve | Anadara trapezium 6% Atrina pectinata Patro australis 25% Pinctada fucata y Mollusc, gastropod | Velacumantis australis 87% 100% 8% 99% Crustacea, hermit crab Dardanus imbricatus Crustacea, brachyuran crab Charybdis natator 25% Mutata sp. Myra affinis Portunus sanguinolentus Brachiopod Lingula murphiana _ | 90% Leucosia sp. B% 75% 8% 8% 10% 90% Fish , unidentified 25% difference in the number of prey species per gut sample between the sexes (t=0.357, d.f.=47, p>0.05) or by size of these large immature and adult-sized turtles (regression analysis: F\ 4g=0.005, p>0.25; r’=0.0001, df=48, p>0.05). FEEDING OBSERVATIONS. C. caretta in eastern Australia used four modes of behaviour to locate and obtain their prey. Mining. Some C. caretta in shallow soft-bottom habitats of Moreton Bay located buried infaunal prey items by ‘mining’ (Limpus et al., 1994; Preen, 1996). The turtles use sweeping motions of their front flippers to dig shallow meandering trenches ~1.5m wide with the advancing edge 0.3-0.45m deep. This mining action resembles the front flipper actions used during nesting be- haviour when the @ is digging or filling in the body pit (Bustard et al., 1975). Thick and thin shelled bivalves and polychaetes (Preen, 1996) of exposed infauna are then crushed and ingested by the turtles. While this feeding behaviour has been regularly observed in sea grass meadows of Moreton Bay, we have not observed it with C. caretta in sandy lagoons of the southern Great Barrier Reef. To be effective, mining requires a substrate that will not readily collapse as trenches are dug. The seagrass root-mass provides this short term stability in Moreton Bay. Biting into substrate surface. In the southern Great Barrier Reef, some C. caretta feed on molluscs living within the top few centimetres of sand of coral-reef lagoon habitat (Limpus, 1978; Moodie, 1979). ‘The turtle walks across the bottom biting up mouthfuls of the molluscs and sand, blowing out the latter with water before LOGGERHEAD TURTLE IN QUEENSLAND 639 TABLE 4. Relative abundance by number of individuals of prey items from Caretta caretta gut contents of beach washed turtles or faecal samples from captured wild turtles from Moreton Bay (n=21). All samples were obtained from turtles that had been feeding immediately prior to their death. Relative abundance values are summarised only for those species comprising >5% of the total sample for the turtle. y’ denotes that the species was present in the gut sample at less than the 5% level. Samples from specimens K7489, T51210 and T53780 were faecal samples. The remainder were from gut contents. Not all species present in all samples are listed. * Where there was a very large difference in the size of prey items the abundance has been adjusted to reflect the relative volume of the species. Tag number J49809 JS1131- J51658 553275 * J53276 K7489 Q10419_ Date | 12.09.89 4.09.90 18.01.91 22.06.91 | 24.06.91 29.05.97 29.09.91 Sex 3 g g 3 é g 2 Maturity Immature Immature Immature Immature | Immature Adult _ Immature — Carapace length (cm) 90.9 625 _ 771A SOT 86.2 91.5 74.6 2711'S 27°18’S 27°22’°S 27°12’S 27°18’S 27°31’8 | Latitude Longitude 153°2’E 153°04E 153°23°E _| 153°22’E _153°22°E 153°23’E Prey Items Mollusc, gastropod Monilea callifera Pyrazus ebeninus Strombus campbelli Mollusc, bivalve Anadara trapezium Antigona lamellaris Atrina pectinata Cardita incrassata Eucrassatella cumingii Solen vaginoides Trichomya hirsuta Crustacea, hermit crab Dardanus imbricatus | Crustacea, brachyuran crab | Eucrate dorsalis Galene bispinosa Leucosia sp. Portunus pelagicus Portunus sanguinolentus Thalamita sima | Portunid Crab, unidentified | Portunid Crab, unidentified 4 | | Crab, unidentified 9 “Crustacea, Squilla sp. 12% 12% 38% 38% 50% 50% 100% 30% 10% 20% 16% 12% 100% 100% Echinoderm, sea urchin Echinoderm, béche-de-mer Cniderian, sea whip Porifera, sponge Urochordate, ascidian | Fish, porcupine fish |Fish, unidentified 640 TABLE 4. (cont.). MEMOIRS OF THE QUEENSLAND MUSEUM Tag number Q10460 | T51210 T53780 | 170057 | _T83099 X10789* | _X10791 Date 23.05.92 10.11.90 _ 30.08.90 19.09.95 | 30.05.95 7.10.96 17.05.96 Sex _ : = $ fo} 3 Q $ 3 Maturity — : - __ Immature Immature Immature | Immature Adult Immature Carapace length (cm) = - 81.4 92.5 86.0 78.9 | 105.8 aw | | Latitude 2708'S | 27°21’S_| |___27°26’S __27°15°S 27°28 27°25°S 2721'S _ | + | Longitude 153°22"E | 153°24’E | 153°21’E | 153°04VE | 153°07E_ | 153°31"E_ | _153°04°E Prey Items | Molluse, gastropod Monilea callifera Pyrazus ebeninus Strombus campbelli | Mollusc, bivalve Anadara trapezium Antigona lamellaris Atrina pectinata Cardita incrassata Eucrassatella cumingii Solen vaginoides 99% Trichomya hirsuta Crustacea, hermit crab | Dardanus imbricatus Crustacea, brachyuran crab Eucrate dorsalis Galene bispinosa Leucosia sp. Portunus pelagicus | Portunus sanguinolentus Thalamita sima Portunid Crab, unidentified 1 Portunid Crab, unidentified 4 Crab, unidentified 9 Crustacea, Squilla sp. _ “< 50% Echinoderm, sea urchin | Echinoderm, béche-de-mer | Cniderian, sea whip | Porifera, sponge | Urochordate, ascidian 100% | 90% 5% 33% 14% 57% | 29% 10% y 48% “< Fish, porcupine fish Fish, unidentified crushing and swallowing the shells’ (Limpus, 1978). Picking off the substrate surface. C. caretta will feed on visible prey items on the substrate surface. Limpus (1973) described C. caretta grasping, crushing and tearing a 19cm long clam, Tridacna maxima, from the substrate surface of a coral reef. An adult male loggerhead (CCL= 94.2cm) was observed pulling a large distended anemone (Stichodactvla haddoni) from the surface of the sandy substrate on the Moreton Banks of Moreton Bay (Limpus et al., 1994). Several large immature C. caretta which were not engaged in ‘mining’ at the time have been captured while feeding on portunid crabs, P. pelagicus, on the bottom over the Moreton Banks (Limpus et al. LOGGERHEAD TURTLE IN QUEENSLAND 641 TABLE 4. (cont.). Tag number a | Z107_ | 21806 © Z1819 * _Z3014* Z3057 - : Date = __ 18.04.93 28.06.96 | 20.07.96 | 12.03.98 | __ 22.01.98 2.01.92 29.01.91 | Sex é g | 3 F |! 3 3 an) - Immature | _Immature | trninature Adult? | Carapace length (cm) = a) 955°} 2 F8A | 9G el 91.0 | : - - 86.3 | ae" | Latitude 26°S7TS | 2657S | 27'S 27°14" | 27°23°8 27°12°S | Longitude _ 153°09°E 153°00E | 153°22E | 1S3°IVE | 1S3°IVE | 153°12"E_ | 153°07°E us Band | Adult | Immature’ Adult Maturity | Prey Items Mollusc, gastropod Monilea callifera Pyrazus ebeninus Strombus campbelli 91% | | Mollusc, bivalve Anadara trapezium | 61% | Antigona lamellaris | 75% 5% y Atrina pectinata 33% Mi Cardita incrassata 11% Eucrassatella cumingii 17% Solen vaginoides | | Trichomya hirsuta | 100% | : - _ 12% Crustacea, hermit crab Dardanus imbricatus | 17% Crustacea, brachyuran crab | Eucrate dorsalis 35% y 55% Galene bispinosa 35% y 15% Leucosia sp. 17% | Portunus pelagicus | y Portunus sanguinolentus 17% y y y 15% Thalamita sima Portunid Crab, unidentified 1 | Portunid Crab, unidentified 4 | Crab, unidentified 9 | Crustacea, Squilla sp. x . 10% Echinoderm, béche-de-mer Cniderian, sea whip | Porifera, sponge _ 4 z - | t _Urochordate, ascidian | Fish, porcupine fish = ‘Fish, unidentified _ ; : | y | 1994; CJL, DJL, MAR unpubl. obs.). Individual narrow for the turtle’s jaws to reach the mollusc. turtles may be persistent in their attempts to The turtle repeatedly circled and nudged the obtain an individual food item. For example, an adult C. caretta was observed for ~15 minutes as it attempted to bite a large Zrochus sp. from a crevice in a coral boulder on Wistari Reef in the southern Great Barrier Reef. The crevice was too captured by amateur anglers in Moreton Bay boulder apparently searching for an alternative access to the food and regularly returned to push into the crevice. C. caretta are occasionally MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 5. Relative abundance by number of individuals of prey items identified in the gut contents of beach washed Caretta caretta from the Gold Coast - Jumpinpin area (n=8). All samples were obtained from gut contents of turtles that had been feeding immediately prior to their death. Relative abundance values are summarised only for those species comprising >5% of the total sample for the turtle. y’ denotes that the species was present in the gut sample at less than the 5% level. Not all species present in all gut contents are listed. | Tag number UQ91/1533 Z3051 23052 | 23053 23054 Z3116 Z3117 | Date | 24.10.91 | 13.12.91 | 141.98 | 14.1.98 14,1.98 | 14.1.98 | 19.03.98 | 19.03.98 Sex an a 8 é | 8 o 3 Maturity | Adult Adult =| Adult Adult | Adult Adult Adult Adult ae | L | Carapace length (cm) 9 97 100.5 CLe i | 99.5 92.7 95.8 93.6 | Latitude __|_28e02s_| 27es4's_|_27esa's | 27°5s's_ | 27esa's | 27ess's | 27e42's | 27°43's._ | | Longitude 153°26°E | 153°24°E | 153°25°E | 1593°25°E_ | 1593°25°E | 153°2S"E | 153°27E | 153°27°E Prey Items | | Molluse, gastropod Tonna tessellata 20% 1 = -_ —— | Crustacea, brachyuran crab Charvhdis natator 20% y y Matuta sp. 20% Portunus sanguinolentus 20% y y Ranina ranina_ 40% 20% | y y 7 J Fish, mixed species 20% 40% >90% >95% >90% >95% >95% >90% | including flathead | (Platycephalus sp.), ‘whiting (Si//ago sp.), | flounder (Pseudorhombu | sp.), wrasse (Labridae), | bigeye (Priacanthus sp.) | 7 r | when the turtles ingest hooks baited with fish that lie on the bottom. Plucking from the water column. C. caretta will feed within the water column (Limpus, 1978; CJL, MAR unpubl. obs.). C. carefta living on the reef edge at Heron Island in the southern Great Barrier Reef grazed on clusters of goose-neck barnacles, Lepas sp., growing on floating timber. When swarms of the jellyfish, Pelagia noctiluca, drifted over the coral reefs near Heron Island, adult and large immature C. caretta of all sizes present cease feeding on their normal food of benthic molluscs (Moodie, 1979) and rise to feed at or near the surface on these jellyfish. In contrast, C. caretta in Moreton Bay has not been observed to feed on the abundant Catostylus mosaicus within their close proximity. In the Queensland Shark Control Program, drumlines (large hooks suspended near the surface from a float) are baited with dead fish to catch sharks (Kidston et al., 1992). C. caretta, especially off Point Lookout (27°26’S, 153°32’E) and the Gold Coast, regularly eat the fish baits on drumlines and are hooked. Many of these turtles have been tagged on release. Tag recoveries indicate that C. caretta learn to seek food from such artificial sources and return regularly to the hooks. In an extreme case, an adult male C. caretta (tag T74407) was hooked at least 18 times on the baited drumlines off Point Lookout between 06 December 1993 and 09 September 1995. These turtles demonstrate that C. caretta will scavenge on food items floating at or near the water surface in addition to taking live food. These four feeding modes appear to be employed by the turtles examined in the present study. The brachiopod, L. murphiana (Table 3), and the bivalve, S. vaginoides (Table 4), burrow deeply within the substrate and can occur in dense aggregations (de Villiers & Hodgson, 1993). These species can be obtained in quantity only by mining. The scallop, A. balloti (Table 2) and the many hermit crabs (Tables 2-4) which do not burrow would have been taken from the substrate surface. The species that burrow to only a limited extent (A. trapezium, A. pectinata, S. campebelli) would be obtained from beneath the surface by biting into the substrate. When a turtle scavenges discarded bycatch from trawlers (e.g. Table 5), itis simply feeding on prey items on the substrate surface or in the water column. Dif- ferences in prey encounter rate and processing may influence a turtle’s choice in procuring food. LOGGERHEAD TURTLE [IN QUEENSLAND When burrowing bivalves and brachiopods were consumed in large numbers, presumably by Mitting, few gastropods or crabs were consumed, It would involve ‘biting’ as well as intaunal mining to. obtain such variety, When crabs were consumed, very few, ifany, bivalves were found inthe yul contents, When bivalves did occur with crustuccuns, jt was usually superficially burrowing bivalves like the scallop 4. ballori or razor clams 4. pecrinafa that were also targeted, Large fragments of prey cxoskeleton can cause blockage of the intestine with resulling death of the turtle: £22833 (Table 3) was found floating and moribund with a necrotic large intestine blocked wilh a compacted mass of A. trapeziun. DISCUSSION Extant turtles do not have teeth. Rather caretta has keratinised sheaths to its jaws that are specialised for grasping and fragmenting hard-bodied food items (Thompson, 1980). With murine turtles, the food is pushed from the mouth through the oesophagus to the stomach using water and w hyoid pump (Bjornadal, 1985; C. Limpus, unpubl. data), The ocsophagus is lined with backwardly projecting keratinised spines that act as a filter system to allow the foad 10 be pushed easily down the throat while impeding tts return back from the stomach but not impeding the return Flow of walter (Thompson, 1980). The back flow of water is also used to flush sand and other small particles from the buceal cavity when the turtle bites up a mouthful of prey and sediment (Limpus, 1978). The thick, keratinised epidermal surlaces of the buccal cavity and throat provide protection to the turtles during Ingestion of the sharp and abrasive surfaces of the exoskeletal ragments of molluses and crusl- aceans. This thick epidermal lining to the mouth and throat also provides protection trom enven- omation during ingestion of venomous prev such as Physalia and jellyfish hike P noviiluca. Once the food bolus has passed beyond the aesoph- agus, acid seeretions by the stomach would inhibit discharge of cnidarian stinging cells (Sutherland, 1983). During the pelagic lite history phase, the young C. curete feed in surface waters (Carr, 1986). Those in later life history phases that live in the shallower inshore waters, have changed their feeding behaviour to utilise prey items on the sea tHoor. The C. caretta in this study area were carnivorous, consuming a wide variety of benthic organisms, Prey items ranged from a single to hundreds of animals per gut sample. Quantifying has prey abundance was highly problematic as our methods give a minimum estimate of the number ofeach speeres per sample. Preen (1996) observed abundant broken molluse shells in feeding scars lefi from infaunal mining, suggesting that much shell, including the identifying components, 1s not swallowed. Alternatively, identifiable features af prey items such as thin-toothed molluse radula may accumulate in the gut, resulting in exaggerated numbers. hor example. a count of the radulae of S. campbell? in sample TS89I45 indicated 541 individuals were consumed, yet only 2) spires were recorded from the shells retrieved. Seyphomedusag (jellytish), hydromedusae, siphonuphores and other solt- bodied maeroplankton are significant prey of pelagic post-hatchling C. Caretta (Bleakney, 1967; van Nierop & den Hartog, 1984; Plotkin ct al,, 1993). These soft budied prey without skeletons are rapidly digested (Plotkin et al. 1993; van Nierop & den Hartop, 1984) and therefore, it may be difficult to identify them in gut content studies, [fowever, many of these turtles were treshly dead and such prey items would be recognised in the anterior of stomachs if fed upon immediately prior to death, This was not the ease, While the present study underestimates the occurrence of soft bodied prey, it is concluded that C, caretia in their non-pelagie phase does nol feed extensively on large, soft-bodied planktonic organisms, Even so, the meestion of charcoal ane tree bark in some turtles suggests at least some surface feeding, It appears that when lane swarms of some species of jellytish are available, large immature und adult C caretfa will temp- orarily abandon benthic foraging in favour of plucking these jlems from the water column, a reversion to the feeding behaviour used during the previous pelagic phase (Prick et al,, 1999). Given the finite time for food passage through the gut, aguleontent orsingle faeces) sample provides only a ‘snapshot’ of the diet and probably retains fewer species than were actually consumed. C curetta iS Queensland have uipredictable and opportunistic diels that result from selective foraging modes and chanee encounters with prey patches. Our study was based primarily on ereriies C. caretta and so foraging areas could only be approximated to broad regions. Large immature and adult C. carerta are specialised for feeding on slow moying, hard bodied invert- ebrate prey (Conway, 1994; Dodd, 1988; Plotkin et al, 1993; Moodie, 1979). As a generalised carnivore, difterent prey species can be expected to dominate C. earena diet in different regions, 644 Turtles from the same locality have a common set of potential prey species yet individuals do not necessarily consume the same prey. Moodie (1979), and Conway (1994) determined in studies of the loggerheads from the southern Great Barrier Reef and the Northern Territory, respectively, that prey availability at feeding sites was different to the relative frequency of those species in the turtle gut contents. The present study also shows that individuals may feed on a very limited range of species and use specialised methods such as infaunal mining to expose the prey. Thus some variability of the diet may be attributed to individual food preference. As in the Northern Territory and GBR (Conway, 1994; Moodie, 1979) we find no correlations in diet for sex, size, Season, or year. Furthermore, Moodie (1979) found that turtles foraging over the same section of coral reef were each selecting a different species of mollusc. Yet the same turtles would rise to the surface to consume jellyfish that sporadically drifted over this reef in large numbers (CJL, pers. obs.). The extent to which C- caretta is obtaining nutritional components via a strategy of foraging on diverse taxa or is merely optimising available food sources that can be gathered with minimum energy expenditure is not addressed herein.Brachiopods are here ident- ified for the first time in the diet of C. carefta. Fish were present in samples other than those from the Gold Coast - Jumpinpin region. Frick (1997) suggested that fish are captured alive but it is more likely that they are scavenged, especially from discarded trawler bycatch (Shoop & Ruckdeschel, 1982; Plotkin et al., 1993). With regular trawling, there is the potential for aggregating C. caretta that scavenge discarded bycatch in the trawled area and hence increasing their risk of capture and death. Once the turtle is associated with a chosen area such as an inter- nesting refuge (Limpus & Reed, 1985) or feeding onareliable food source (feeding off baited shark hooks, this study), they can be persistent with the association despite other disturbances or human- related perturbations. C. caretta, through their diet, has additional unquantified interactions with a number of coastal fisheries. Commercially fished species such as the sand crabs P. pelagicus and P. sanguinolentus, the spanner crab R. ranina and the scallop A. balloti, are targeted as food by some C. caretta. Potentially more significant to fisheries is the role of C. caretta in the life cycle of the ascaridoid nematode Sulcascaris sulcata, MEMOIRS OF THE QUEENSLAND MUSEUM a parasite of commercial scallops. C. caretta is the definitive host of S. suwlcata (Sprent, 1977) with the adult worms inhabiting the stomach and intestine while eggs are shed to sea with faecal material. Molluscs are the secondary host to the larval worms and immature S. su/cata are frequently found in the adductor muscles of large bivalves, especially 4. balloti (Cannon, 1978). The parasite completes its life cycle when the mollusc is eaten by C. caretta. This parasite can cause loss of fisheries production in areas where there is coincidence of large numbers of C. caretta and scallops (Lester, 1980). The occurrence of C. caretta off the Bundaberg coast where scallops are abundant enough to form a significant part of their diet, has the potential for maintaining locally elevated levels of infection of scallops by S. sulcata. ACKNOWLEDGEMENTS This study was a part of the Queensland Turtle Research project of the Queensland Parks and Wildlife Service. Patrick Couper (Queensland Museum), numerous QP WS staff and volunteers assisted in obtaining the diet samples. Thora Whitehead, Alan Limpus, Beverley Heidke and Kevin Lamprell provided assistance in identification and advice for molluscs, Dr Peter Davie (Queensland Museum) associated in identification of crustaceans, Two unidentified reviewers made helpful editorial comments. This assistance is gratefully acknowledged. LITERATURE CITED BJORNDAL, K.A. 1985. Nutritional ecology of sea turtles. Copeia 1985: 736-751. BLEAKNEY, J.S. 1967. Food items in two loggerhead sea turtles Caretta caretta (Linnaeus) from Nova Scotia. Canadian Field Naturalist 81: 269-272. BOWEN, B.W., KAMEZAKI, N., LIMPUS, C.I., HUGHES, G.R., MEYLAN, A.B. & AVISE, J.C. 1994, Global phylogeography of the loggerhead turtle (Caretta caretta) as indicated by mitochondrial DNA haplotypes. Evolution 48: 1820-1828. BURKE, V.J. & STANDORA, E.A. 1993. Diet of juv- enile Kemp’s Ridley and Loggerhead sea turtles from Long Island, New York. Copeia 1993: 1176- 1180, BUSTARD, H.R., GREENHAM, P. & LIMPUS, C. 1975. Nesting behaviour of loggerhead and flatback turtles in Queensland, Australia. Proceedings Koninkl. Nederlands Akademie van Wetenschappen, Series C 78: 111-121. CARR, A. 1986. New perspectives on the pelagic stage of sea turtle development. National Oceanic and Atmospheric Administration Technical LOGGERHEAD TURTLE IN QUEENSLAND Memorandum, National Marine Fisheries Service Southeast Fisheries Center 190: 1-36. CANNON, L.R.G. 1978. A larval acsaridoid nematode from Queensland scallops. International Journal of Parasitology 8: 75-80. CONWAY, S.P. 1994. Diets and feeding biology of adult Olive Ridley (Lepidochelys olivacea) and Loggerhead (Caretta caretta) sea turtles in Fog Bay (N.T.). Unpubl. thesis, Northern Territory University, Darwin. DE VILLIERS, C.J. & HODGSON, A.N. 1993. The filtration and feeding physiology of the infaunal estuarine bivalve Solen cylindraceus Hanley 1843. Journal of Experimental Marine Biology and Ecology 167: 127-142. DODD, K. 1988. Synopsis of the biological data on the loggerhead sea turtle Caretta caretta (Linnaeus 1758). Fish and Wildlife Service, U.S. Depart- ment of Interior Biological Report 88(14): 1-110. ERNST, C.H, & BARBOUR, R.W. 1989. Turtles of the World. (Smithsonian Press: Washington DC.) FRICK, M. 1997. Lepidochelys kempi (Kemp’s Ridley) Caretta caretta (Loggerhead) and Malaclemys terrapin centrata (Carolina Diamondback Terrapin). Diet and predation. Herpetological Review 28: 149. FRICK, M.G, QUINN, C.A, & SLAY, C.K. 1999, Dermochelys coriacea (leatherback sea turtle), Lepidochelys kempi (Kemp’s ridley sea turtle), and Caretta caretta (loggerhead sea turtle). Pelagic feeding. Herpetological Review 30: 165. GUDYNAS, E. 1980. Notes on the sea turtles of Uruguay. ASRA Journal 1: 69-76. KENNELLY, S.J. & WATKINS, D. 1994. Fecundity and reproductive period, and their relationship to catch rates of spanner crabs, Ranina ranina, off the east coast of Australia. Journal of Crustacean Biology 14: 146-150. KIDSTON, W., DWYER, K., BUHK, A., WALDOCK, J. & ANDERSON, G. 1992. Review of the Operation and Maintenance of Shark Meshing Equipment in Queensland Waters: Report of the Committee of Enquiry. (Queensland Department of Primary Industries: Brisbane). LAMPRELL, K. 1998. Bivalves of Australia. Vol. 2. (Backhuis: Leiden). LESTER, R.J.G. 1980. Marine parasitology research aiding industry. Australian Fisheries October 1980: 13-14. LIMPUS, C.J. 1973. Loggerhead turtles (Caretta caretta) in Australia: food sources while nesting. Herpetologica 29: 42-45. 1978. The reef. Pp.187-222. In Lavery, H.J. (ed.) Exploration North: Australia’s Wildlife from desert to reef. (Richmond Hill Press: Richmond). 1985. A study of the loggerhead turtle, Caretta caretta, in eastern Australia. Unpubl. PhD thesis, Zoology Department, University of Queensland. 1994. The loggerhead turtle, Caretta caretta, in Queensland: feeding ground selection following 645 her first nesting season. National Oceanic and Atmospheric Administration Technical Memorandum National Marine Fisheries Service Southeast Fisheries Centre 351: 78-81. LIMPUS, C.J., COUPER, P.J. & READ, M.A. 1994. The loggerhead turtle Caretta caretta, in Queensland: population structure in a warm temperate feeding area. Memoirs of the Queensland Museum 37: 195-204. LIMPUS, C.J. & REED, P.C, 1985. The loggerhead turtle, Caretta caretta, in Queensland: observations on internesting behaviour. Australian Wildlife Research 12: 535-40. LIMPUS, C.J. & REIMER, D. 1994, The loggerhead turtle Caretta caretta, in Queensland: a population in decline. Pp. 39-59. In James, R. (compiler) Proceedings of the Australian Marine Turtle Conservation Workshop (Australian Nature Conservation Agency: Canberra). MOODIE, E.G. 1979. Aspects of the feeding biology of the loggerhead turtle (Caretta caretta). Unpubl. BSc Honours thesis, James Cook University of North Queensland, Townsville. PLOTKIN, P.T. 1996. Occurrence and diet of juvenile loggerhead sea turtle Caretta caretta in the North West Gulf of Mexico. Chelonian Conservation Biology 2(1): 78-80. PLOTKIN, P.T., WICKSTEN, M.K. & AMOS, A.F. 1993. Feeding ecology of the loggerhead sea turtle Caretta caretta in the Northwestern Gulf of Mexico. Marine Biology 115: 1-15. PORTER, K.R. 1972. Herpetology. (Saunders: Philadelphia). PREEN, A.R. 1996. Infaunal mining: a novel foraging method of loggerhead turtles. Journal of Herpetology 30: 94-96. SHOOP, C.R, & RUCKDESCHEL, C. 1982, Increasing turtle strandings in the South-east United States: a complicating factor. Biological Conservation 23: 213-215. SPRENT, J.F.A. 1977. Ascaroid nematodes of amphibians and reptiles: Su/cascaris. Journal of Helminthology 5: 379-387. SUTHERLAND, S.K. 1983. Australian Animal Toxins. (Oxford University Press: Oxford). THOMPSON, S.M. 1980. A comparative study of the anatomy and the histology of the oral cavity and alimentary canal of two sea turtles: the herbivorous green turtle Chelonia mydas and the carnivorous loggerhead turtle Caretta caretta (includes discussion of diet and digestive physiology). Unpubl. MSc thesis, James Cook University of North Queensland, Townsville. VAN NIEROP, M. M. & DEN HARTOG J.C. 1984, A study on the gut contents of five juvenile loggerhead turtles Caretta caretta (Linnaeus) (Reptilia, Chelonidae), from the south-eastern part of the North Atlantic coast, with emphasis on coelenterate identification. Zoologische Mededelingen 59(4): 35-54. 646 FIRST RECORD OF ELSMAN’S WHIPNOSE ANGLERFISH GIGANTACTIS ELSMANI (LOPHIFORME GIGANTACTINIDAE), FROM AUSTRALIAN WATERS. Memoirs of the Queensland Museum 46(2); 646, 2007-- klsman’s Whipnose Anglerfish Gigantactis elsmani Bertelsen, Pietsch & Layenberg (1981), isa meso- and bathypelavic species previously known from five specimens, The 384mm SL holotype (ISH 1360/71) was trawled in the central Atlantic and the 283mm SL paratype (LACM 10687-1) was caught in the eastern South Pacific. Other specimens have been recorded trom Japan (Amaoka, 1984), the Sea of Okhotsk (Fedorov, 1994), and one specimen tentatively (dentified as this species from the South Atlantic (Bertelsen, Pietsch & Layvenberg, 1981) Giventactis contains 21 species (Bertelsen, Pietsch & Lavenberg, 1981), and these show extreme dimorphism, The females have an elongate body, head length less than 35" SL, caudal peduncle length greater than 20% SL, and a very long illicrum, greater than half the SL. The largest female grows to 40em, whereas the largest male is only 2.2em long. Males have highly developed sense organs that are presumably used to find females, Dermal denticles on the snout allow the male to attach to the female, Unlike some other ceratioid anglerfishes, male/female attachment in whipnose anglerfishes is not parasitic (Pietsch, 1999), A 310mm SL lemale Gelsmrani (AMS 1.28742-001) was collected by Australian Museum stall on the HAMAS Cook in 1989 in the Tasman Sea off Sydney, NSW (trawl! start and finish courdinates; 33°52°30"S 152"39°00"E to 33°53°09"S }52"05°54"E). The specimen was collected by noidwater trawl ata depth between the surface and | 800m over a bottom depth from 1700m to 4856m, and is a new record for both Australia and the SW Pacific, This is the second species of Gigantactis recorded from Australian waters, the first bem CG puxtoni (Bertelsen & Pietsch, 1983) The AMS specimen of Gef/smueni was fixed in Formalin in 1989 and has been preserved since in 70% ethanol, The specimen has the following characters which, in combination, diagnose the species: illicial length 315mm (102% SL): escal papillae absent; dentary teeth relatively short, 9.0mm (longest 2.9% SL) in approximately 3 rows; longest caudal fin ray 90mm (29.0% SL). These characters are in agreement with the original description of the species, with the minor exception that the length of the longest dentary tooth is 2-8 vs. 2.9% SL. The characters of the esca also 2¢ with. the original description, With specimens previously collected [rom the Atlantic, SI Pacific and Japan it was not surprising that this species was found in Australian waters. The known distribution of this species is here extended into the SW Pacific, increasing the likelihood that its distribution 18 eireumplobal c sexual Acknowledgements J. Leis is acknowledged for giving helpful advice on the manuscript. C. Bento and T. Trnski photographed the fish, J Trski compared characters of the fish with those in the original description Literature Cited AMAOKA, K. 1984. Fumily Cigantactinidae. Whipnoses. Pp 106-L08. In Mastida, H., Amaoka, Is... Araga, C.. Uyeno. T. & Yoshino, T. (eds) Fis of the Japanese Archipelago. (Toke University Press; Tokyo) & PIETSCH, TW. 1983. The Cerauoid sof Australia, Records of the Australian Museum ,E PIETSCH, TW, & LAVENBERG, RJ. 1981 Ceratioid anglerfishes of the family Gigantactinidae morpholovy, systematics, and distribution. Contribributions in Science (los Anweles) 332: 1-74 MEMOIRS OF THE QUEENSLAND MUSEUN A PIC. 1.A,310mm SL temale & e/smani (AMS L28742-001) collected in the Tasman Sea off Sydney, NSW: B, head {nole (he origin of the illicium at the tip of the snout); C, esca. FEDOROV, VV. | 4b4 the (rst report of a epecies Of the family Gigantactinidae (Lophitormes) trom the Sea of Okhotsk, Voprosy Ikhtiolowii. 34(3);414-415, [English translation in Journal of k vyolowy 34 (RK); | 32-133] PIETSCH, TW rantaclinidas, Whipnose anglerfishes {deepsea anylerfishes). P2036. In Carpenter, KE & Niem, VE. (eds) PAO species identification puide for Fishery platy he Living Marine Resources of the Western Central Pacific Vol. 7 (FAQ: Rome) Ciganractis elsmeani 1009, G Mark A, MeCGrouther, Division of Vertebrate Zoology, tusiralian Museum, 6 College Street, Svdriev 20h, 15 June JNO) AN ADDITION TO THE RAINBOWFITSH (MELANOTAENIIDAE) FAUNA OF NORTH QUEENSLAND KATRINA L. MeGUIGAN McGuigan, K.L, 2000 06 30; An addition to the rainbowfish (Melanotaeniidae) fauna of North Queensland. Memoiry of the Queensland Museum 46(2); 647-655. Brisbane. ISSN 0079-8835. Anew Melanotaentid species is described from 28 specimens vollected from the Johnstone River, north Queensland, Melanotaenia uicheensis sp. nov was found in sites with moderate to high water flow over cobbjes and boulders and all sites are in close proximity to major agricultural activity and have moderately to highly disturbed riparian vegetation. Melanotaenia utcheensis sp. nov. has a distinctive colour pattern with a blue-black mid-lateral band and orange margins on vertical scale rows. It is morphologically distinct from the broadly sympatric Melanatdenia eachamensis (Allen & Cross, 1982) and Melinatwenia splendida splendida (Peters, 1866), as well as from its sister species from southern Queensland/northern New South Wales, Me/unatuenia duboulayi (Castelnau, (878). In particular, M. uteheensis sp, nov. has more first dorsal spines and fewer vertical scale rows and anal rays than Af. §. splendida, and fewer soft second dorsal rays and more pectoral rays than either Af. eachamensis or M. duboulay!. The new species is alse generally smaller than vither Ad. 5. splendida or M. cachamensis and intermediate between them in eye diameter, predorsal length, head depth and body depth. O Melanoreeniidae, Melunotaenica, ralibawfsish, freshwater, North Queensland, Johnstone River. KL. MeGuigan, Department of Zovlogy and Entomology, The University of Queensland, St Lucia 4072, Australia (e-mail: kmeguigan@isen ug.edu.au); 10 May 2001, The family Melanotaeniidae is endemic to freshwaters of Australia and New Guinea. These small rainbowfish (usually less than |2em standard length) tend to be locally abundant, representing a major coinponent of the fresh- water fauna of the region. Rainbowfish are also popular in (he aquarium trade both in Australia and overseas. There are currently 68 described Melanotaeniidae species in seven genera (Allen & Renyaan, 1998), Recent changes to taxonomy have primarily resulted from surveys in New Guinea, where fifleen species und one genus have been described since 1990 (see Allen & Renyaan, 1998). In Australia there are 4 genera with 13 species, a number that has remained static since Crowley et al. (1986) reassessed the status of southern Me/anotaenia. Melanotaenia is the numerically dominant genus in Australia where it is represented by ten described species and 4 subspecies, Some Melanotueniu species are geographic- ally restricted while others are widespread, occupying a range of habitats, Widespread taxa often display interpopulation variation in morphology and colouration, making classification difficult. Intraspecific phenotypic variation is recognised in the aquarium trade where rainbowfish are sold as types, usually tamed for the collection locality. Melanotaenia splencdida Splendida (Peters, 1866) is one such widespread taxon (Cape York Peninsula to Gladstone) that is sold as several types due to variation among populations in both colour and morphology. The high level of intraspecific variation in M. s. splenedida has caused con(usion over the status of the geographically restricted and poorly characterised Me/anolaenia eachamensis (Allen & Cross, 1982) (Crowley & Ivantsoff, 1991; Zhu et al,, 1994, 1998; Pusey et al., 1997), Pusey ct al.’s (1997) study of morphological variation of rainbowfish in the region supported separate Species status for ML eachamensis. Mapping the distribution of and determining the relationship among mtDNA lineages on the Atherton Tablelands confirmed the species status of M. eachamensis (Zhu et al., 1994, 1998), Several of the At, eathamensis populations identified by Pusey et al. (1997) represented new lower altitudinal limits for the species (Rankin, Fisher and Utehee Creeks), However, subsequent mtDNA analysis of fish from those sites indicated that they represented a distinct lineage, more closely related lo the southern species, Melanotaema duboulayi (Castelnau, 1878) and M. fluviatilis (Castelnau, 1878) than to M, eachamensiv (McGuigan et al, 2000; fig. 1), MIDNA analysis of high altitude populations nat 648 previously characterised for morphology revealed another unique lineage, closely related to that observed in Rankin, Fisher and Utchee Creeks (McGuigan et al., 2000: fig. 1). These lineages exhibit divergences from described species consistent with a cessation of gene flow between one and two myr ago (McGuigan et al., 2000). The status of the Utchee Creek population has been debated previously. It is sold in the aquarium trade as the Utchee Creek Type. Leggett & Merrick (1987) considered the population to be banded rainbowfish (Melanotaenia trifasciata Rendahl, 1922), which they resemble in colour pattern. Allen & Cross (1982) identified fish from Utchee Creek as a population of M. s. splendida with unusual colouration but conceded the possibility that they represented an undescribed species. In his rainbowfish classification scheme Schmida (1997) distinguished the Utchee Creek Type from all described species and, as did McGuigan et al. (2000), placed it in a group containing M. eachamensis, M. duboulayi and M. fluviatilis, along with M. s. australis populations from Western Australia. This paper describes M. utcheensis sp. nov. from Utchee, Fisher and Short Creeks in the Johnstone River, north Queensland. The new species is compared to the broadly sympatric M. eachamensis and M. s. splendida and also to its sister species, M. duboulayi. MATERIALS AND METHODS SPECIMEN COLLECTION. Melanotaenia eachamensis, M. s. splendida and M. utcheensis sp. nov. were collected during 1998 and 1999 from sites in northeast Queensland (Fig. 2) using dip nets and traps. Melanotaenia duboulayi were collected in the same manner from Kholo Creek (Brisbane River) and Amamoor Creek (Mary River) in southeast Queensland (Fig. 2). Inform- ation on land use, riparian vegetation, substrate, and channel characteristics was collected at each site. Fish were transported to The University of Queensland, Brisbane and held in 72L tanks at 26°C until processing. MORPHOLOGICAL CHARACTERISATION. The definition of characters and the format of the description follows Allen & Renyaan (1998). Morphological characterisation was performed on anaesthetised live fish (1:10000 MS222, Sigma Chemical Company). Fin ray and scale row counts were made using a light microscope. Morphometric measurements were made on microscope images using Video Trace (Leading MEMOIRS OF THE QUEENSLAND MUSEUM Edge Pty Ltd, 1994); a program that facilitates calibrated measurement directly from a live video feed. Type specimens were then euthanased by anaesthetic overdose and deposited at the Queensland Museum. Data analysis was conducted using SPSS for Windows v. 9 (SPSS Inc., 1999). Meristic data was non-normally distributed, and could not be normalised through standard transformations. A Kruskall-Wallace test was conducted on all meristic variables and the sequential Bonferroni technique (Rice, 1989: «=0.05, k=7) used to control for group-wide type-I error. Variables that remained significant after correction were subject to a non-parametric multiple comparison test with unequal sample sizes (Zar, 1984) to determine whether the significant result was due to M. utcheensis sp. nov. The same technique was used to assess the meristic similarity of M. eachamensis and M. s. splendida. Morphometric data was natural log trans- formed. To allow comparison of shape without the confounding effect of size (as indexed by standard length) data were size corrected using the formula: scaled variable = set standard length y observed variable x ( observed standard length where b was the slope of the regression (specific for sex within population) of the observed variable on standard length; set standard length was 4.0 (=In 55mm) as this value fell within the range of all species. A discriminant functions analysis (DFA) was conducted on the size-corrected data with species as the discriminator. A one-way ANOVA with planned comparisons was used to compare the discriminant scores of M. utcheensis sp. nov. with all other species. A one-way ANOVA with planned comparisons was also used to compare M. utcheensis sp. nov. with all other species for univariate morphometric variables. Again, the morphological similarity between M. eacha- mensis and M. s. splendida was determined in the same way. Melanotaenia utcheensis sp. nov. (Fig. 3) ETYMOLOGY. Named for the type locality, Utchee Creek, and in recognition of the history in the aquarium trade of the Utchee Creek Type. NEW NORTH QUEENSLAND RAINBOWFISH MATERIAL. HOLOTYPE: QM 132159, 3, Utchee Creek, North Johnstone R. (17°38'30"S. 145°56’20”E). PARATYPES: Utchee Creek, North Johnstone R. (17°38°30"S 145°56°20”E), 5 females, QM 132160-32164 inclusive; Fisher Creek, North Johnstone R. (17°34°55”S 145°53°55”E), 2 males, QM 132165 and 32166, 5 females, QM 132167-32171 inclusive; Short Creek, North Johnstone R. (17°23’00"S 145°40°00"E), 10 males, QM 132172 and 5 females, QM 132173. DESCRIPTION. The value of the holotype is presented with the observed paratypic range in parentheses. Dorsal rays VII-1I,12 (V to VII-I, 10 to 12); anal rays 1,19 (I, 16 to 20); pectoral rays 12 (11 to 15); horizontal scale rows 10 (9 to 11); vertical scale rows 34 (32 to 35); predorsal scales 13 (13-16). Greatest body depth 34mm (31-38), head length 36mm (34-42) both as a proportion of standard length. Greatest body width 24mm (17-28) as a proportion of body depth. Snout length 32mm (34-43), eye diameter 28mm (24-32), interorbital width 22mm (19-28), depth of caudal peduncle 25mm (20-30), length of caudal peduncle 17mm (11-18) as proportions of head length. Upper and lower jaws are of approximately equal length, oblique with a typically abrupt bend in the premaxilla between the anterior horizontal and lateral portions; the maxilla ends in front of the anterior edge of the eye; lips are thin. Scales are arranged in regular horizontal rows and their posterior edge is slightly crenulate; predorsal scales extend to the posterior end of the interorbital; preopercle has 2 scale rows from the posterior angle to the edge of the eye. The origin of the first dorsal fin is anterior to the anal fin origin. The depressed longest first dorsal ray (second or third from origin) reaches half way between the insertion of the first dorsal and the origin of the second dorsal in females and ranges in males from the spine to the fourth soft ray of the second dorsal. The second dorsal fin origin is posterior to the anal fin origin. When depressed the longest rays of the second dorsal fin (usually second or third from insertion) extend just past the point of insertion in females, but reach almost to the caudal fin in males. The longest rays of the anal fin (usually second or third from insertion) have a depressed length the same as those of the second dorsal. In males both the second dorsal and anal fins are elongated and show a boxy outline when extended, whereas in females the fins are rounded, giving them a more ovaloid outline when extended. Length of pelvic fins is 18mm (15-26), pectoral fins 21mm 649 TABLE 1. Structure matrix from the discriminant functions analysis. Superscripts denote loading rank for the 3 variables that contribute most to that function, ranging from highest loading (1) to third highest (3). ee | Snout Length 0.026 0.544" 0.749! | Eye Diameter F 0.3717 | 0.019 0.338" Head Length -0.079 : 0.266 : 0.149 | | Predorsal Length | 0.289 0.672! 0.177%. | | Head Depth “a -0.618! 0.268 0.061 __—| | Body Depth hh 0.4697 ve 0.589 | -0.104 (13-19) and caudal fin 17mm (8-19), all as a proportion of head length. The pelvic fin extends half to two-thirds of the way to the anal fin origin in females and smaller males, but reaches to the third anal soft ray in large males. Pectoral fins are rounded. The caudal fin is moderately forked. Colour in Life. The overall body colour is silver, often with an orange cast near the midline. The head and gill region is silver to pink and fish often have an obvious reddish cheek patch. Scales tend towards purple iridescence, especially on the upper half of the body. There is an obvious orange stripe between horizontal scale rows. A typical Melanotaenia mid-lateral band starts dark at the tail and fades forwards, tending towards blue in males and black in females. Anal, dorsal, pelvic and caudal fins range from translucent to deep red, being most strongly pigmented in males. Anal, second dorsal and pelvic fins often have black margins, especially in males. Pectoral fins are translucent, Colour in Alcohol. The underside is pale, generally tawny in colour. Above the mid-lateral line scales are outlined in grey, with blue tones in some specimens. Fins retain black margins, but the red in anal and dorsal fins fades to pink. Dark mid-lateral bands are retained. Sexual Dimorphism. 33 and 2 are easily distinguished on the basis of external character- istics. d d tend to be more strongly pigmented than 22. 36 are also deeper bodied. As described above, ¢ ¢ have longer pelvic, dorsal and anal fins. The outline of extended second dorsal and anal fins is distinctively boxy in d d and oyaloid in 2°, COMPARISONS. Melanotaenia utcheensis can be discriminated from M. eachamensis, M. s. splendida and M. duboulayi on the basis of multivariate morphology (Fig. 4). .M. utcheensis differs from all other species on Discriminant 650) MI. Punotitls Ml dubBouloyt A — 5 thames Af. s, tatel Ms. splendida Ad 5. splendida Se MM. trifasciata tthica Creek Fisher Creek Ufchée Creek AT esachamensis Mi & Quairalls Ws salanaca Aq. $, rubrostriara. M7 5. inarnata Mo meaecullochi MEMOIRS OF THE QUEENSLAND MUSEUM Ms. splendida M. eachamensis — 0.005 changer FIG. J. A, Maximum parsimony phylogeny of mtDNA cytochrome b (351 bp) and control region (331 bp) sequence, adapted from McGuigan et al. (2000). Bootstrap support for nodes are indicated above branches. M!. utcheensis sp. noy. represented by sequences from [thica, Utchee and Fisher Creeks. B, Neighbour-joining network of mtDNA control region sequence (331 bp) (McGuigan & Moritz, unpubl.). Geographic locations of the nine M. utcheensis alleles are: 1, North Johnstone R. below the Malanda Falls; 2, Bromfield Swamp, North Johnstone R. below Malanda Falls and Ithica R.; 3, Ithica R.; 4, Ithica R.: 5, Gillies Creek; 6, Utchee Creek: 7, unnamed tributary of North Johnstone R.. near Glenn Allyn; 8, Rankin, Fisher, Tregothanana and Utchee Creeks; 9, Utchee Creek. Functions 1 (DF1) and 3 (DF3), but is distinct from only M. duboulayi on DF2 (one-way ANOVA with planned comparisons: p<0.001 for all significant comparisons) (Fig. 4). Factor loadings indicate that negative contributions from depth variables and positive contributions from length variables dominate DF1 (Table 1). M. eachamensis has low scores on DF 1 due to its deep body and head, and small eyes and across DF 1 species are progressively shallower in head and body, and larger eyed, with M.s. splendida at the extreme of this trend (Fig. 4; Table 2). DF2 is determined by positive contributions from all traits (Table 1) and M. duboulavi is distinct on DF2 because it is generally shorter and shallower than other species (Fig. 4; Table 2). DF3 is dominated by positive contributions from snout length and eye diameter, with M. utcheensis hav- ing short snouts and small eyes (Table 2; Fig. 4). Melanotaenia utcheensis was observed to be morphologically most similar to the southern species M. dyboulayi (Table 2). This supports the mtDNA sequence data, which indicates that they are sister species (Fig. |). Melanataenia utcheensis differs from M. duboulayi in having fewer soft second dorsal rays and more pectoral rays, as well as a longer predorsal distance and a deeper maximum body depth (Tables 2 and 3), Melanotaenia utcheensis is more distinct from its sympatric congeners IM. eachamensis and M. s. splendida, differing in most of the traits measured (Tables 2 and 3). Melanotaenia utcheensis has fewer vertical scale rows, anal rays, and first dorsal spines than Af. 5. splendida; fewer second dorsal rays and more pectoral rays than At, eachamensis (Tables 2 and 3). Standard length, snout length, bead length and depth, and body depth of Mf. uicheensis are less than that of M. eachamensis, but eye diameter is. greater (Table 2). Compared to M. 8, splendida, M. utcheensis has a shorter standard length, snout length, eye diameter, predorsal length and head depth, but a deeper body (Table 2). Melanotaenia eachamensis differs from M. s. splendida in having fewer vertical scale rows and anal rays, more first dorsal spines and second dorsal rays, and being shorter in standard length and predorsal length, having smaller eyes and a deeper head and body (Tables 2, 3). The intermediate position of Af. uteheensis between M. eachamensis and M. s, splendida in both multivariate morphospace (Fig. 4) and in univariate traits (Table 2) probably contributed to the lack of previous recognition of species status, Morphometric analyses of rainbow fish species by McGuigan et al. (2000) suggested that some characters particularly reflect phylogenetic history, while others reflect local adaptation, NEW NORTH QUEENSLAND RAINBOWFISH 651 TABLE 2. Mean meristic and morphometric measurement (mm) + standard error. All morphometric measurements (except standard length) are corrected for standard length (see equation in text). Asterix (*) indicates a significant difference between M. utcheensis and the asterixed species; hash (#) inicates a significant difference between M. eachamensis and M. s. splendia at p<0.05 significance level (from one-way ANOVA with planned comparisons for size-corrected morphometric data and from nonparametric multiple comparisons for merisite data; see text for details). M.utcheensis (53) | _ M. eachamensis (90) M. s. splendida (25) M. duboulayi (40) Vertical Scale Rows 31,724 0.27 31.67 £0,154, 33.00 £0.13" _ 32.95 + 0.19 Horizontal Scale Rows 9.87 = 0.08 10.17 + 0.06 10,00 + 0.06 10.25 + 0.08 | | Anal Rays : 18.79 + 0.14 «18.80 + 0.144 20.12 + 0.22* 18.73 + 0.17 lst Dorsal Spines 5.40 + 0.11 5.42 + 0.08# 4.88 + 0.12* 4.95 + 0,12 | 2nd Dorsal Rays 10.79 + 0.09 | WN.91+0.12*# | 10.72+0.17 | 11.43 + 0.24* Pelvic Rays 5.96 + 0,04 | 6.02 + 0,04 5.00 + 0.00 6.00 + 0.00 | Pectoral Rays 12.62 + 0.14 | 1190+ 0.08* 12.00 + 0.1] 148+ 011" | Standard Length _ 46.02 + 0.74 SL2840.76%# | 56.364 1.90" 46.18 + 0.88 Snout Length 3.24 + 0.05 3.62+0.05* 3.73 + 0.11% 3.10 + 0.08 Eye Diameter 456+ 0.05 437+ 0.04*# 5.06 + 0.08% _ | 448 +004 Head Length 13.364 0.10 13.68+0.10* — 13.40 + 0.14 : 13,124+0.12 | Predorsal Length 24,05 + 0.15 23.62 + 0.18# —25,50£0,22* 22.13 + 0.15* ‘Head Depth 12.63 + 0.16 14.27420.13%4 1137+ 0.20* 12.90 + 0.17 Body Depth __ 14.924 0.15 16.48 + 0.16*# 13.49 + 0.31* 14.10 + 0.19% plasticity, or the effect of random genetic drift. Traits that contributed strongly to discrimination among clades in McGuigan et al.’s (2000) study (i.e. those with strong phylogenetic signal) showed the greatest differences among species in this study, strongly supporting the species status of M. utcheensis. Additionally, all populations included in the morphological analyses came from similar habitats (fast flowing streams in closed forest), reducing the possibility that observed differences are due to local adaptation, or phenotypic plasticity. As documented above, historically there has been considerable confusion over species assignments in the Wet Tropics. In this study, all 3 species of the region are distinct in morphology. However, the differences between them are not pronounced and the level of variation within species suggests that none of the traits are diagnostic (Tables 2 and 3). Allen & Cross (1982) described M. eachamensis and indicated that it differs from M. s. splendida in a number of traits, including having a consistently shallower body. The opposite was observed in this study, with M. eachamensis being consistently deeper in the body than M. s. splendida (Table 2). Given the evidence of a long-term lack of gene flow (McGuigan et al., 2000), the morphological divergence between species is surprisingly limited. The lack of morphological specialisation of Australian freshwater fish has been noted previously (McDowall, 1981), and similarity of rainbowfish in the Wet Tropics may be a related phenomenon. Despite the inconvenience such an approach would cause, | recommend that assignment of fish to species be based on multivariate morphological analyses of multiple populations, preferrably with supporting molecular data (see Zhu et al., 1998). DISTRIBUTION AND HABITAT, Melano- taenia utcheensis was discovered through the identification of a unique mtDNA lineage, more closely related to southern species than to other north Queensland species (McGuigan et al., 2000: fig. 1). The above analyses confirmed M. utcheensis as a discrete species by demonstrating that the unique mtDNA lineage correlates with a unique morphology. Morphological data for north Queensland rainbowfish is limited, such that the distribution of M. utcheensis might be better determined through examination of the distribution of mtDNA lineages (McGuigan & Moritz, unpubl. data; Fig. 1). The M. utcheensis mtDNA lineage has been observed allopatrically in a tributary of the lower South Johnstone River (Utchee Creek) and in lower (Fisher and Rankin Creeks) and upper (Short Creek and an unnamed creek near Malanda) tributaries of the North Johnstone R (McGuigan & Moritz, unpubl. data; Fig. 2). The M. utcheensis mtDNA lineage was also observed MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 3. The percentage of the surveyed fish that are observed to have each values of the meristic traits vertical scale rows, pectoral fin rays, first dorsal spines and second dorsal and anal soft fin rays. These traits are those observed to differ among species (Table 2). | Species _ _ Vertical Scale Rows 7 ee Pectoral Rays : 2» | 30 | 31 32 33 34 35 mo | u | 2 | 43 14 15 utcheensis z | 47 | 8 4 9 21 ay. | 6 53 24 | 8 9 eachamensis 7 | 3B 27 24 | 19 9 | l | 29 | 49 21 splendid 28 | 44 28 16 68 | 16 _duboulayt = 12 25. 28 25 10 7 40 | 50 ) - Species 7 First Dorsal Fin Spines | Second Dorsal Soft Rays —— _. wih “2 4 5 6 | 7 5 9 10 iW} 12 13 | 14 15 | utcheensis 7 | a8 55 | 28 | 9 a 36 49 15 eachamensis | ou 41 42 6 8 | 33 29 | 26 2 2 splendida 24 _ 64 12 | 7 4 40 36 20 | 7 duboulayi | 3 | 17 | 60 | 20 [2 3 is | 28 | 30 | 2 | 2 Species J __ Soft Anal Fin Rays | _ | 16 17 | 18 19 20 21 22 23 utcheensis _| 8 30 45 9 8 | eachamensis | 8 7 ll 28 26 «|= (25 3 3 splendida _ | s_ | 16 | 40 32 | | 4 | | duboulayi 12, | 33 30, | 20 5 7 al at several sites in the main channel, upland (Ithica, Gillies and Williams Creeks) and lowland (Tregothanana) tributaries of the North Johnstone R (Fig. 2), admixed at various proportions with M. s. splendida mtDNA (Zhu et al., 1998; McGuigan & Moritz, unpubl. data). Melanotaenia utcheensis co-occurs, at a low tre- quency, with the more common ™. eachamensis mtDNA lineage at one site, Bromfield Swamp, the headwaters of the North Johnstone River (McGuigan & Moritz, unpubl. data). The South Johnstone River has been surveyed extensively, making it unlikely there are undisc- overed populations in this drainage. However, the M. utcheensis lineage in so many sites in the North Johnstone River, along with the many unsampled tributaries, raises the possibility of undiscovered populations in that catchment. Analyses of mtDNA sequence suggest that, while M. utcheensis and: M. eachamensis are old species that evolved in situ, M. s. splendida is a young, species and has colonised the region recently (McGuigan et al., 2000; Hurwood & Hughes, 2001). It is not yet clear whether the admixture of mtDNA lineages in the North Johnstone River is the due to the occurrence of M. utcheensis and M. s. splendida in sympatry, or to either current or historical hybridisation. The co-occurrence of M. eachamensis and M. utcheensis at only one site, despite frequent co-occurrence of each with M. s. splendida, suggests the old endemics are characterised by barriers to dispersal and gene flow; lack of geographic discontinuity suggest the barriers may be ecological. If M. utcheensis and M. s. splendida evolved allopatrically, only recently coming into contact, they may not have evolved any mechanisms that would prevent hybridisation. Freshwater fish show unusually high levels of hybridisation and introgression (Turner, 1999). A documented threat to the conservation of freshwater fish is loss of genetic identity through introgressive hybridisation with introduced taxa (Berrebi, 1997). In many cases hybridisation is facilitated by modification of habitat and human-mediated species intro- ductions; conditions that are met in N Qld. Within M. utcheensis there are 2 MtDNA lineages, | on the Atherton Tablelands (e.g., Short Creek) and 1 primarily in the lowlands (e.g., Utchee Creek) (McGuigan et al., 2000: fig. 1). The presence of these sister lineages with a highly structured geographic distribution sug- gests long-term barriers to gene flow, and consid- erable antiquity of the endemic lineage. With additional information on ecology and inter- actions of rainbowfish in the Wet Tropics, it may become appropriate to accord species status to these 2 lineages, which are differentiated morph- ologically (McGuigan, unpubl. data). Lacking NEW NORTH QUEENSLAND RAINBOWFISH 653 COE. rf \ co ON pod 15 Mary River er VWEHrishane River t ¥ Barron \ ae River Malanda , a 7 ya vi) 5 \ 4 o> 6 ~40km 7 8 14 Le Millaa Millaa Innisfail e th Johnstone River 4 10 12 : Sth 13 Johnstone River FIG. 2. Distribution of Melanotaenia species as determined from the distribution of mtDNA lineages (McGuigan & Moritz, unpubl. data). Sites sampled in this study indicated with *: 1, Tinaroo Dam (M. s. splendida: 17°09°30"S 145°35°10"E); 2, Lake Euramoo (M. eachamensis: 17°09°30"S 145°37°40”E); 3, Upper North Johnstone R. (M. s. splendida: 17°30’30”S 145°37°20”E); 4, Ithica R. (MA utcheensis: 17°24’25"S 145°37’10”E); 5, * Short Ck (M. utcheensis: 17°23’00”S 145°40°00”E); 6, unnamed tributary (M. utcheensis: 17°23°50"S 145°39°20"E); 7, Gillies Ck (M. utcheensis: 17°25’40"S 145°36715”E); 8, * Dirran Ck (M. eachamensis: 17°28°30"S 145°32°53”E); 9, * Upper South Johnstone R. (M. eachamensis: 17°39°25”S 145°42°55”E); 10, Rankin Ck (M. utcheensis: 17°34°15"S 145°53’55”E); 11, * Fisher Ck (M. utcheensis: 17°34°55”S 145°53"55”E); 12, * Lower South Johnstone R. (M. s. splendida: 17°43’50"S 145°56’00”E); 13, * Utchee Ck (M. utcheensis: 17°38°30"S 145°56°20”E); 14, Tregothanana Ck (M. utcheensis: 17°31720"S 145°57°30”E); 15, * Amamoor Ck (M. duboulayi: 26°21’S 152°40°E) and; 16, * Kholo Ck (M. duboulayi ). such data, I have taken the conservative approach of recognising both lineages as a single species. Melanotaenia utcheensis is found in moderate to fast flowing water, in sections of stream consisting of deep pools separated by short runs. The substrate consists of cobbles and boulders with little fine sediment. Most sites have good visibility, but Rankin Creek and the lower North Johnstone main channel have very poor visibility due to large amounts of suspended solids. Visibility varies substantially across time, probably due to seasonal changes in land use and rainfall. Utchee Creek is the most structurally complex site with exposed root masses and overhanging vegetation. Other sites have undercut banks or grass beds. Sampled sites in Fisher, Utchee and Ithica Creeks have a riparian buffer zone dominated by native vegetation. Other sites have highly disturbed riparian veg- etation (completely absent or dominated by exotic species). These sites are located in agri- cultural lands such as banana and tea plantations. Melanotaenia utcheensis commonly co-occurs with purple spotted gudgeons (Mogurnda adsp- ersa), long finned eels (Anguilla reinhardtii) and less commonly with blue-eyes (Psuedomugil signifer), roman nosed gobbies (Awaous acri- tosis), swamp eels (Ophisternon sp.), grunters (Hephaestus sp.) and exotic guppies (Poecilia reticulata) (pers. obs.; B. Pusey, pers. comm.). The Wet Tropics of north Queensland are listed 654 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 3. Melanotaenia utcheensis, &, Utchee Creek, South Johnstone R., northeast Queensland. as a World Heritage area, partly in recognition of the high level of endemism. As yet, there is little known about this most recent addition to the endemic fauna, M. ufcheensis. The geographic separation of populations of M. eachamensis and M. utcheensis, and the structure of genetic diversity within the latter, suggest a distribution that has been stable over along period. It is not yet clear what the mixed populations of M. s. splendida and M. utcheensis represent. Pure M. utcheensis populations are currently known from only five sites (Fig. 2), with stream structure suggesting a restricted area of occupancy at those sites. Observations on population size during sampling suggested that population size was fewer than 1,000 mature individuals per site. If the mixed populations of M. s. splendida and M. utcheensis represent different stages in an ongoing exclusion of M. utcheensis through competition, predation or introgressive hybridisation, M. utcheensis would qualify as vulnerable under the criteria of the World Conservation Union. M. eachamensis is infamous as the first Australian freshwater fish to be declared extinct (in the wild). Although it has been rediscovered in the wild (Zhu et al., 1998), its disappearance from Lake Eacham is a strong warning for management of freshwater fish. Circumstantial evidence suggests that the cause of the demise of M. eachamensis in Lake Eacham was the introduction of 4 non-endemic _ pisci- vorous species (Barlow et al., 1987). In addition to the threat of predation, the potential for loss of genetic identity through hybridisation, at the population and species levels, should be con- sidered in formulation of management strategies. Translocation of fish, either deliberately, or accidentally as a by-product of general water movement, or as live bait for fishing, has the potential to impact severely on diversity. ACKNOWLEDGEMENTS DPI Walkamin, Brad Pusey, Geoff Carey, David Hurwood, Brian Stumer, Adam Fletcher, Laura Paetkau, Kym Collins, Lisa Pope, Craig Franklin, Craig Moritz and Culum Brown assisted in collection. This paper benefited from the input of Walter Ivantsoff, Culum Brown, Craig Moritz, Craig Franklin, Mark Blows, Elizabeth O’Brien and Lisa Pope. This paper is an outcome of a research project on rainbowfish funded by an ARC grant to Craig Moritz. LITERATURE CITED ALLEN, GR. & CROSS, N.J. 1982. Rainbowfishes of Australia and Papua New Guinea. (Angus & Robertson: Sydney). ALLEN, GR. & RENYAAN, S.J. 1998. Three new species of rainbowfishes (Melanotaeniidae) from Irian Jaya, Indonesia. aqua, Journal of Ichthyology and Aquatic Biology 3: 69-80. NEW NORTH QUEENSLAND RAINBOWFISH Function 2 Function 1 Function 3 Function 1 FIG. 4. Mean score for ds and @s of each population on: A, functions | and 2; and B, functions | and 3 of the Discriminant Functions Analysis. (0 M. eachamensis, (| M. s. splendida, A M. duboulayi, @ M. utcheensis, + group centroids. BARLOW, C.G, HOGAN, A.E. & RODGERS, L.J. 1987, Implication of translocated fishes in the apparent extinction in the wild of the Lake Eacham’ rainbowfish, Melanotaenia eachamensis. Australian Journal of Marine and Freshwater Research 38: 897-902. BERREBI, P. 1997. Species introduction in freshwater environments: Genetic impacts. Bulletin Francais de la Peche et de la Pisciculture 344-45: 471-487. CASTELNAU, F.L. 1878. On several new Australian (chiefly) freshwater fishes. Proceedings of the Linnean Society of New South Wales 3: 140-144. CROWLEY, L.E.L.M., IVANTSOFF, W. & ALLEN, G.R. 1986. Taxonomic position of two crimson-spotted rainbowfish, Melanotaenia duboulayi and Melanotaenia fluviatilis (Pisces: Melanotaeniidae) from eastern Australia, with special reference to their early life-history stages. Australian Journal of Marine and Freshwater Research 37: 385-398, CROWLEY, L.E.L.M. & IVANTSOFF, W. 1991. Genetic similarity among populations of 655 rainbowfishes (Pisces: Melanotaeniidae) from Atherton Tableland, Northern Queensland. Icthyological Exploration of Freshwaters 2: 129-137. HURWOOD, D. & HUGHES, J. 2001. Interdrainage dispersal of eastern rainbowfish from the Atherton Tableland, north-eastern Australia. Journal of Fish Biology 58: 1125-1136. LEADING EDGE PTY. LTD. 1994. Video Trace: Image Measurement System. (Leading Edge Pty. Ltd.: Marion, South Australia). LEGGETT, R. & MERRICK, J.R. 1987. Australian Native Fishes For Aquariums. (Artarmon: NSW). McDOWALL, R.M. 1981. The relationships of Australian freshwater fishes. Pp. 1253-1273. In Keast, A. (ed.) Ecological Biogeography of Australia. (Dr W. Junk Publishers: The Hague). McGUIGAN, K., ZHU, D., ALLEN, GR. & MORITZ, C. 2000. Phylogenetic relationships and historical biogeography of Melanotaeniid fishes in Australia and New Guinea. Marine and Freshwater Research 51: 713-723. PETERS, W.C.H. 1866. Mittheilun uber Fische. Monatsberichte der KAniglichen Preussischen Akademie der Wissenschaften zu Berlin 1866: 509-526. PUSEY, B.J., BIRD, J.. KENNARD, M.J. & ARTHINGTON, A.H. 1997. The distribution of the Lake Eacham rainbowfish in the Wet Tropics Region, North Queensland. Australian Journal of Zoology 45: 75-84. RENDAHL, H. 1922. Acontribution to the ichthyology of north-west Australia. Meddeleser Fra det Zoologiske Museum, Kristiania 5: 163-197. RICE, W.R. 1989. Analysing tables of statistical tests. Evolution 43; 223-225. SCHMIDA, G. 1997. What’s in a name anyway. Fishes of Sahul 11: 481-500. SPSS INC. 1999. SPSS for Windows v. 9. (SPSS Inc.: Chicago, Illinois). TURNER, GF. 1999. What is a fish species? Reviews in Fish Biology and Fisheries 9: 281-297. ZAR, J.H. 1984. Biostatistical Analysis. (Prentice-Hall International, Inc.: New Jersey). ZHU, D., JAMIESON, B.G.M., HUGALL, A. & MOR- ITZ, C. 1994, Sequence evolution and phylo- genetic signal in control-region and cytochrome b sequences of rainbowfishes (Melanotaeniidae). Molecular Biology and Evolution 11: 672-683. ZHU, D., DEGNAN, S. & MORITZ, C. 1998. Evol- utionary distinctiveness and status of the en- dangered Lake Eacham_ rainbowfish (Melanotaenia eachamensis), Conservation Biology 12: 80-93. 656 SEA SPIDERS (PYCNOGONIDA) FROM THE GREAT BARRIER REEF, AUSTRALIA, FEED ON FIRE CORALS AND ZOANTHIDS. Memoirs of the Queensland Museum 46(2); 656, 2001;-Tropical sea spiders have been scarcely studied and associations between pycnogonids and reef organisms are rarely reported. The possibility of sea spiders feeding on scleractinian corals had been suggested once (Child, 1998) but no evidence had been presented for either fire corals or zoanthids as prey of sea spiders before. This note reports the occurrence and feeding activity of the pycnogonid /ndeis on the fire coral Mi/lepora exaesa (Class Hydrozoa) and species of Zoanthidea (Class Anthozoa) from Rib Reef, Pandora Reef and Goold Island in the central section of the Great Barrier Reef. This is the first report on coral reef sea spiders feeding activity, and is based on both field and laboratory observations. Adults of Endeis mollis (32 individuals) were found on different colonies of Millepora exaesa (Fig. 1) at Rib Reef (6-9m depth). On several occasions, the sea spiders were observed feeding on the coral inserting the tip of the proboscis into a coral polyp and remaining attached for about 60 sec. Individuals of £. mollis were also observed feeding on Palythoa caesia and crawling on a Parazoanthus sp, at Rib Reef. Related Endeis biseriata, was found in the reef flat of Goold Island and Pandora Reef (2-4m depth). Twenty-two adults of E. biseriata were found on Protopalythoa sp. Similar feeding activity to that of £. mollis was observed in some individuals of £. biseriata at both sites. The relative high number of £. mollis on the coral Millepora exaesa and of E. hiseriata on the zoanthid Protopalythoa sp. suggest a preference by the Endeis spp. for MEMOIRS OF THE QUEENSLAND MUSEUM these hosts, at least at the reefs visited. Observations made under captive conditions in the laboratory with the aid ofa low light video camera corroborated the sequence of movements observed in the field and more importantly, the ingestion of cnidarian tissue by the sea spiders. Some pycnogonids are known to sequester and accumulate metabolites from prey, apparently to be used as defence (Sheerwood et al., 1998), It remains to be studied whether this association between Endeis spp. and cnidarians from the Great Barrier Reef could be chemically mediated, since Millepora and zoanthids in general, are known for their high concentration of toxins (Fossa & Nilsen, 1998). Acknowledgments Thanks to G. Diaz-Pulido for help with collecting and photography. I thank the Australian Institute of Marine Science (AIMS) for providing access to the Great Barrier Reef on board the RV Harry Messel. Literature Cited CHILD, A. 1998. Nymphon torulum, new species and other Pycnogonida associated with the coral Oculina varicosa on the east coast of Florida. Bulletin of Marine Science 63: 595-604, FOSSA, S.A. & NILSEN, A.J, 1998. The modern coral reef aquarium. (Birgit Schmettkamp Verlag: Bornheim). SHEERWOOD, J., WALLS, J.T. & RITZ, D.A, 1998, Amathamide alkaloids in the pyenogonid, Sty/opallene longicauda, epizoic on the chemically defended bryozoan, Amathia wilsoni. Papers and Proceedings of the Royal Society of Tasmania 132: 65-70 Claudia P. Arango, Department of Zoology and Tropical , Ecology, James Cook University, Townsville 4811; 8 November 2000. FIG, 1, Underwater photograph of Endeis mollis (Pycnogonida) on the fire coral Millepora exaesa. DIAGRAMMA MELANACRUM NEW SPECIES OF HAEMULID FISH FROM INDONESIA, BORNEO AND THE PHILIPPINES WITH A GENERIC REVIEW JEFFREY W. JOHNSON, JOHN E. RANDALL AND STEPHEN F. CHENOWETH Johnson, J.W., Randall, J.E. & Chenoweth, S.F. 2001 06 30: Diagramma melanacrum, new species of haemulid fish from Indonesia, Borneo and the Philippines with a generic review. Memoirs of the Queensland 46(2): 657-676. Brisbane. ISSN 0079-8835. The haemulid fish Diagramma melanacrum is described from 5 specimens: 1 S of Lombok, | off Bali, 1 off Halmahera, and 2 from Bonebetang Is., Sulawesi. The species is known also from underwater photographs at the islands of Moyo and Romang, S Indonesia, Sipadan and Mabul off NE Borneo, Tomini Bay in Sulawesi, Bohol in the Philippines and Ashmore Reef, Timor Sea. It is distinct among haemulids in having IX-X, 22-24 dorsal rays; 55-57 lateral-line scales; swimbladder with 3 small anterior projections; dorsal spine 3 or 4 longest, 1.8-2.9 in head; first dorsal spine about half length of second; pelvic fins reaching vent or slightly beyond, 1.2-1.3 in head; and in colour: pale yellow dorsally, shading through grey to whitish ventrally, with numerous small dark brown spots on head, body and dorsal fin, and caudal fin with lower fourth black; anal and pelvic fins mainly black. Diagramma pictum is divided into 5 geographically separate subspecies: D. pictum pictum (Thunberg), from the western Pacific and Indo-Malay Archipelago, excluding Australia and S New Guinea; D. pictum labiosum Macleay, from N Australia and S New Guinea; D. pictum cinerascens Cuvier, from the Bay of Bengal to the Persian Gulf; D. pictwm punctatum Cuvier, from the Red Sea; and D. pictum centurio Cuvier, from East Africa. O Diagramma, Haemulidae, Indonesia, Philippines, generic review. Jeffrey W. Johnson, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia (e-mail: jeffi@qm.qld.gov.au); John E. Randall, Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817; Stephen F. Chenoweth, Genetics Laboratory, Faculty of Environmental Sciences, Griffith University, Nathan 4111, Australia; 18 May 2001. Haemulid fishes of Plectorhinchus and Dia- gramma (commonly Thicklips, Sweetlips, Rubberlips or Slateys) are important components of the fish fauna of coral reefs and adjacent habitats of the Indo-Pacific region. Indo-Pacific species of Plectorhinchus (23) and Diagramma (2) are are strikingly different in colour as juveniles, and some have intermediate phases that differ from either the juvenile or the adult. Association of juveniles to adults is still not complete for all species. Like other haemulids, Plectorhinchus and Diagramma are primarily nocturnal. They tend to form small, nearly stationary aggregations on coral reefs by day and disperse at night for feeding on a wide variety of small benthic invertebrates. In 1988, while diving in 36m on a reef off NW Moyo Island (8°13’N, 117°30°E) in S Indonesia, one of us (JER) encountered a small group of an unknown haemulid fish that was believed to be a species of Plectorhinchus Lacepede. A photograph (Fig. 3D) was sent to Roland J. McKay at the Queensland Museum who reported that it appeared to be undescribed. The Queensland Museum has a specimen of this species, taken by trawl S of Lombok in the Lesser Sunda Islands. It had been misidentified as P. picus and was reported and illustrated as such (Gloerfelt-Tarp & Kailola, 1984). Burgess et al. (1988, pl. 221) followed Gloerfelt-Tarp & Kailola with their artist’s impression. Re-examination of the QM specimen revealed that it has X dorsal spines (XI-XIV in species of Plectorhinchus) and a swimbladder with 3 small anterior projections (not known in Plecto- rhinchus). These are features of Diagramma Cuvier to which it is now assigned. Additional specimens are known from Denpasar, Bali and Halmahera, Indonesia and Bonebetang Is., Sulawesi. Underwater photographs by JER at Romang Island, Indonesia (7°34’S, 127°27’E); by Klaus E. Fiedler at Sipadan Island, Sabah; by Dieter Eichler at Bohol Island, Philippines; by Rudie Kuiter at Mabul, Borneo (QMNR 44-45) and the S edge of Tomini Bay, central-eastern Sulawesi (QMNR 46-47, QMNP 200); and by J. Barry Hutchins at Ashmore Reef, Timor Sea indicate that the species is widespread in the East Indian region. Diagramma has generally been regarded as monotypic (Smith, 1962; McKay, 1983), although Kuiter & Debelius (e.g.1994) recognised 2 closely related species. Apart from the species described here, 5 apparently allopatric colour forms of D. pictum (Thunberg) occur, with the SE Asian form most distinctive. These variants are most readily recognisable in the large juvenile to subadult colour phases. All forms have previously been assigned specific names; however, with the exception of D. labiosum Macleay from Australia, these species have generally been regarded as junior synonyms of D. pictum. Attempts were made using colour and morphometric and meristic data, as well as through analysis of mitochondrial DNA, to determine whether some of these forms represent distinct species. The results of both methods were inconclusive. DNA analysis suggested that there may be almost as much genetic divergence among widely separated Australian populations of D. ‘abiosum’ as there is between some of these populations and D. pictum from Singapore. From these comparisons, as well as non-genetic examination of other colour forms from the Indian Ocean, it was concluded that more detailed genetic study would be necessary. All colour varieties appear to form geographically separate populations and should be atforded interim recognition as subspecies of D. pictum. METHODS The following measurements were taken: Standard length (SL) = tip of snout (not upper lip) to base of caudal fin (hypural plate flexure); body depth = maximum depth from base of dorsal spines; body width = maximum width just posterior to gill opening; head length (HL) = tip of snout (not including upper lip) to posterior margin of opercular membrane; snout length = tip of snout (not including lip) to fleshy margin of orbit; interorbital space = least fleshy width; preorbital depth = least depth from lower edge of orbit to above end of maxilla; caudal peduncle length =rear of anal fin base to base of caudal fin; spine and ray lengths were measured to the base of these elements; diameter of spots on the body were taken from spots of most common width on the upper body. The last ray of the dorsal and anal fins was counted as one, although divided to base; pectoral ray counts include the upper rudimentary ray; lateral-line scale counts were made to the caudal flexure and include tubed scales only; gill-raker counts were made on the first arch and include all rudiments that are at least long as wide; the upper-limb gill-raker count was given first; the lower-limb count MEMOIRS OF THE QUEENSLAND MUSEUM includes the raker at the angle. Figures for the holotype are given in parentheses. Total length (TL) is quoted where size was estimated from photographs of fish not collected, and in the key, to assist in field identification of fresh or live material. Institutional abbreviations: Australian Museum, Sydney(AM); The Natural History Museum, London (BMNH); Bernice P Bishop Museum, Honolulu (BPBM); CSIRO Marine Laboratories, Hobart (CSIRO); Museum National d’Histoire Naturelle, Paris (MNHN); Miyazaki University, Miyazaki (MUFS); Northern Territory Museum, Darwin (NTM); Phuket Marine Biological Centre, Phuket (PMBC); Queensland Museum, Brisbane (QM); JLB Smith Institute of Ichthyology, Grahamstown (RUSI); Museum fur Naturkunde an der Universitat Humbolt zu Berlin (ZMB). DNA EXTRACTION, PCR AND SEQUENCING. The genetic analyses and its statistical treatment were carried out by SC, Total genomic DNA was isolated from muscle tissue using sequential phenol/chloroform extractions following digestion with Proteinase Kk (Sambrook et al., 1989), Two oligoprimers were used to amplify a 420bp fragment of the mitochondrial Cytochrome B gene, L 1475 (Meyer, 1991) and CB2H (Palumbi et al., 1991). PCR mixtures contained 5u1 of 1.25mM dNTP’s’(Promega), 51 of Promega Polymerase reaction buffer, | unit of Promega tag DNA polymerase (#1662), 5 ul of MgCl2 , 2.5u1 of each primer (10uM), 1 yl of genomic template DNA, made up to a volume of 50u1 with autoclaved distilled water and overlaid with 301 of light mineral oil. The mixture was subjected to 35 cycles of the following thermal profile: Smin at 94°C (only on initial cycle), 30sec at 94°C, 30sec at 55°C, Imin at 72°C and S5min at 72°C (on final cycle only). PCR-amplified DNA was then purified for sequencing using an agarose gel purification kit (QIAGEN). Light strand sequences were generated for each individual using Automated DNA Sequencing Technology (Applied Biosystems Inc.) following the manufacturers directions. Nucleotide sequences were aligned by eye and identical sequences were assigned the same haplotype number for subsequent statistical analysis. Nucleotide diversity and nucleotide divergence statistics (Nei, 1987) were calculated within and among each pair of samples using REAP (McElroy et al., 1991). The phylogenetic relationships among haplotypes was represented NEW SPECIES OF HAEMULID FISH as a minimum spanning network. Minimum spanning trees were reconstructed with the aid of a program supplied by L. Excoffier. SYSTEMATICS Diagramma melanacrum Johnson & Randall, sp. nov. (Common name: Blackfin Slatey) (Figs 1A-F, 8) Plectorhinchus picus (non Valenciennes) Gloerfelt-Tarp & Kailola, 1984; 198 (fifth fig.), 199, 340. Plectorhinchus picus (non Valenciennes) Burgess, Axelrod & Hunziker, 1988: pl. 221, lower right fig. ETYMOLOGY. Greek me/anos, black; and akros, tip or at the end; refers to black outer part ofanal and pelvic fins and lower part of caudal fin. MATERIAL, HOLOTYPE: QMI20291, 410mm SL, Indonesia, S of Lombok, 8°24’S, 116°01’E, trawl, T. Gloerfelt-Tarp, 1982. PARATYPES: QMI30725, 187mm SL, Indonesia, Kampung Loleba, Wasile district, Halmahera, 00°58’N, 127°56’E, trawl, H. Singou, 1/2/1979; QMI20285, 133mm SL, Indonesia, Denpasar, Bali, 8°46°S, 115°10°E, trawl, T. Gloerfelt-Tarp, 1982; BPBM 26719, (2) 219-254mm SL, Sulawesi, W side Bonebetang Is., spear, 20m, sand at reefedge, J.E. Randall, 8/9/1978. (Fig. 8). DIAGNOSIS. Dorsal rays X-X1, 22-24; tubed lateral-line scales 55-57; outer row of teeth in jaws slightly enlarged, movable; swimbladder with 3 short anterior projections; third or fourth dorsal spine longest, 1.8-2.9 in head; first dorsal spine 1.7-2.3 in length of second; interspinous membranes of dorsal fin not incised; pelvic fins reaching vent or slightly beyond, |.2-1.3 in head; body light yellow dorsally, shading to pale silvery grey on sides and ventrally, with numerous small dark brown spots, which are smaller and form oblique rows on body below lateral-line; head light purplish grey with numerous small brown spots, opercular membrane black; dorsal fin and upper 3/4 of caudal fin with small dark spots; lower 1/4 of caudal fin black; anal and pelvic fins mainly black. Juveniles from 133mm SL with spots about size of pupil on head, body, dorsal and caudal fins becoming progressively smaller and more numerous with age. Juveniles and subadults up to 254mm SL with narrow discontinuous wavy lines on operculum and cheeks, extending to about level of anterior margin of eye; these lines breaking up into spots in larger specimens. DES NN (Table 1) Dorsal rays X-XI, 22-24 (X, 22); anal rays III, 6-7 (III, 6); pectoral rays 17 (first ray rudimentary, second unbranched); pelvic rays I, 5; tubed lateral-line scales 55-57 (56); scales above lateral-line to origin of dorsal fin 14-15(14); gill rakers 6-7+13-14 (6+14). Body oblong and moderately deep, depth 38.0- 41.5% SL (41.5%); body compressed, width 31.6-40.8% depth (39.4%); caudal peduncle depth 10.7-11.8% SL (10.7%); caudal peduncle length 22.6-25.2% SL (23.4%); head length 29.5-33.2% SL (31.2%); snout length 21.6-28.7% HL (27. 3%); "orbit diameter 21.1-29.3% HL (21.1%); interorbital space convex, width 25.8-33.6% HL (33.6%); preorbital depth 22.2-27,.3% HL (27.3%). Mouth small, slightly inferior, maxilla reaching a vertical from just beyond anterior margin to nearly the anterior third of eye, upper- jaw length 25.6-35.5% HL (35.5%); numerous rows of small slightly curved conical teeth at front of jaws, narrowing to a single row posteriorly, outer row slightly enlarged and somewhat movable; lips fleshy; chin with 3 pairs of prominent pores. Scales small and ctenoid; lateral line gently curved, following dorsal contour of body; head scaled except snout just forward of anterior nostrils (to anterior nostrils in holotype), and a naked band to eye; small scales basally on median fins, those on dorsal extending about 1/4 distance to margin; opercle witha single flat blunt spine; margin of preopercle very finely serrate, serrae slightly larger at corner. Dorsal fin scarcely notched, interspinous membranes not incised; base of spinous portion of dorsal fin 26.9-3 1.2% SL (27.8%); base of soft portion of dorsal fin 36.6-41.1% SL (36.6%); first dorsal spine about half length of second, 43.2-60.4% (49.3%); longest dorsal spine the third or fourth, 34.0-55.9% HL (third, 34.0%); longest dorsal soft ray 44.5-61.5% HL (44.5%); base of anal fin 28.9-35.7% HL (28.9%); first anal spine very short, 7.0-12.9% HL (7.0%); second anal spine 25.8-45,1% HL (25.8%); third anal spine 28.1-45.4% HL (28.1%); longest anal ray 53,1-69.6% HL (53.1%), reaching when adpressed more than half distance to caudal-fin base; caudal fin truncate to slightly emarginate, its length 20.0-23.8% SL (20.7%); longest pectoral ray 21.1-21.9% SL (21.5%); pelvic fins reaching to or slightly beyond vent, 24.1-27.4% SL (24.1%). Colour in alcohol of holotype: light brown with numerous small dark brown spots on head, dorsally on body, dorsal fin, and upper 3/4 of 660 TABLE 1. Proportional measurements of the type specimens of Diagramma melanacrum expressed as a percentage of standard length. MEMOIRS OF THE QUEENSLAND MUSEUM (specimens not collected): snout, interorbital, nape, body Holotype mostly above lateral-line, | Eatetypes ——| dorsal fin, and upper 3/4 of | QMi20291 | QM130725 | QMiz0285 | BEBM | BPN’ caudal fin pale yellow with IL Ie ihe a 19 : |Standard length(mm) | 410.0 187.0 133.0 | 219.0 954.9 | Humerous small dark brown BEA Eta ree Lea cay oh abs 3 spots; body below lateral-line * ff : ae 2°: on ares ers light grey with numerous faint me wie igs 138 a — a close-set orange brown spots, Head length 32 33.2 30.8 29.5 | 31.7 smaller than those dorsally on | Snout length __ 8.5 9.5 719 64 | 83 body, and arranged in oblique Upper jaw length VL 9.4 79 9.0 | 9.4 rows; head light purplish grey [Orbit diameter 6.6 80 9.0 82 | 79 with numerous small dark | Preorbital depth | BS TS 6.8 6.8 1.7 orange brown spots; ventral Interorbital width | 10.5 86 | 84 9.2 9.8 part of head and body whitish; Caudal peduncle depth | 10.7 11.4 11.8 11.7 11.7 opercular membrane black; ; f i lower 1/4 of caudal fin black; Caudal peduncle length 23.4 22.6 25.2 24.0 23.4 . “ik '§ isions dorsal base 27,8 29.4 29,3 31,2 26.9 anal and pelvic fins whitish =a - =a =" “2: = =: basally, soon shading to black i ss lorsa ase 36.6 36.7 36.6 38.8 41.1 distally, the leading edge of First dorsal spine 4.3 7.5 10.2 6.1 6.0 pelvics narrowly whitish; | Second dorsal spine 8.7 15.2 16.9 13.9 13.7 | pectoral fins pale with dusky |Third dorsal spine 10.6 16.7 16.8 14.3 14.7 | rays and a narrow dark reddish Fourth dorsal spine | 10.0 16.8 172, 13.6 14.2 brown bar at base. Subadults of Fifth dorsal spine 9.9 15.8 15.6 13.0 13.4 about 250mm TL with narrow Longest dorsal ray 13.9 20.3 15.8 18.2 16.1 discontinuous wavy long- Anal fin base _\|e: 9.0 12 10.9 10.5 10.9 itudinal dark brown lines on power Es | head, extending from about irst anal spine 2:2. 43 4.0 3.3 | 2.5 ' : : . _ anterior margin of eye to Second anal spine 8.0 13.0 13.9 12.1 11.6 | h f | Third anal spine 8.8 14.2 140 | 12.8 126 _| apeacles; The eolour patter: P : == wa = small juveniles is unknown. Longest anal ray 16.6 20.9 19.5 20.5 19.7 Longes . a) Caudal fin length 20.7 23.8 23.3 21,5 20.0 REMARKS, Kuiter (1993; Longest pectoral ray 21.5 21.9 21.1 21.8 21.7 aor! Sie oSRiee | Longest pelvic ray 24,1 27.4 25.9 25,3 25.1 ( ) an ICHUEE EES caudal fin; spots below lateral-line smaller, not as dark, very close-set, following centres of oblique scale rows; spots on snout and interorbital smaller and more close set than those on remainder of head and upper body; opercular membrane dark brown; lower 1/4 of caudal fin and all but basal part of anal and pelvic fins dark brown; pectoral fins dusky, the upper half of base with a dark brown bar that curves over dorsal edge of fin base to axilla. Paratypes, 133-254mm SL: Similar to holotype except spots on head, body and caudal fins progressively smaller and greater in number with increasing size. Oper- culum and cheeks with narrow discontinuous wavy longitudinal lines extending to about level of anterior of eye. Anteriorly, on snout, lines re- placed by small dark brown spots as in holotype. Colour in life from underwater photographs (1997) recognised 2 species of Diagramma, D. pictum (Thunberg), widespread in the Indo-west Pacific excluding Australia and S PNG, and D. labiosum Macleay from Australia and S PNG. Sightings of both species from Flores, Indonesia were reported by Rudie H. Kuiter (pers. comm., 1996), but the presence of D. ‘abiosum’ in this area has not been substantiated by specimens or photographs. J. Barry Hutchins reported (pers. comm., 2000) that D. ‘labiosum' was absent from Ashmore Reef. This is not surprising as the fish fauna of this reef has close affinities to Indonesia and includes species typically Indo-Malaysian, some of which are not known from Australian inshore waters (Allen, 1993). Another haemulid, Plectorhinchus vittatus (Linnaeus) is recorded from offshore reefs at Ashmore, Hibernia and Rowley Shoals, but not from other Australian waters. NEW SPECIES OF HAEMULID FISH 661 FIG, 1, A, Diagramma melanacrum Holotype, QM 1.20291, 410mm SL, Lombok, Indonesia, T. Gloerfelt-Tarp. B, Diagramma melanacrum Paratype, BPBM 26719, 254mm SL, Off Ujung Pandang, Sulawesi, J.E. Randall. C, Diagramma melanacrum ~350mm TL, Romang Island, Indonesia, J.E. Randall. D, Diagramma melanacrum ~330mm TL, Moyo Island, Indonesia, J.E, Randall. E, Diagramma melanacrum ~350mm TL, Tomini Bay, Sulawesi, R.H. Kuiter. F, Diagramma melanacrum ~ 250mm TL, Mabul, Borneo, R.H. Kuiter. G, Diagramma pictum labiosum, 300mm TL, Lizard Is., Australia, J.E. Randall. H, Diagramma pictwn labiosum, ~130mm TL, Sydney Harbour, Australia, R.H. Kuiter. 662 TABLE 2. Meristic data of Diagramma species. MEMOIRS OF THE QUEENSLAND MUSEUM Tubed Lateral-tine Scales 55 56 S7 S58 59 60 61 G2 63 64 65 66 67 6R 69 70 71 72 #73 74 ~+%TS 76 77 7B melunacrim % -f.2 a ee ad - = eowwr™ . lee ee. + pictum lahiasum - -~ + = F A 1 4 ! Lo ¢ | & FF WD FT B@ & 2 BYE = |) pictum pictum 4 3 5 $3 3 1 s&s Ff Fey F FS VD + ae ~ = pictum cinerascens | 5 Lak. *3 43 i 2 | | - : = - = _ - : + 4 - - - pletum centurio 2 i} 3 8 1 | 2 2 3 - I - - - - - - - . 2 = = picum punctataum z | i] ee 1 | - - - - - = - = ~ = — = 2 s e Gill Rakers Upper Lower Total 5 6 7 8 9 1 o6420C« 13 45 TT 708 OH melanacrum - 4 ! - - - - 4 1 = C - - 3 2 = < . pictum labiosum 3 330023 3 - | 19 «37 4 r E | 4M 24 616 3 \ pictum pictum | 1s 26 7 1 - 9 36 6 i j - 6 \4 4 id 5 pictum cinerascens—- & {is 3 ] 5 I 7 - ! i 6 x § - pictum centuriv - | 9 16 2 - - - 12 16 - - - - 6 q 3 pictum punctatum 6 2 - - - 2 3 3 - 2 3 ] z Dorsal Spines Dorsal Rays Pectoral Rays 1x x XL 20 21 22 23 24 2s 26 16 v 18 melanacrum 2 3 - - - | - 4 - - - 5 pictim labiosum 12 SO. - - 3 16 «29 |2 x 3 16 13 piclam pietum G 43 - | 6 7 13 4 5 - 2 Au 6 pictum cinerascens 2 2) l + 3 al i of - 20 | pletum centurio 7 22 € - | 5 I 10 2 - - 25 3 plot punectaruin = 8 . - 2 4 2 - - - 5 3 Diagramma pictum and D. ‘labiosum’ are distinguished on few characters other than colouration, with all meristic and morphometric values strongly overlapping (Tables 2, 3). Small juveniles are difficult if not impossible to separate. Given the similarity between pictiim and ‘labiosum’ there was some doubt as to whether they should be separated at the specific or subspecilic level (as geographically separate colour forms). With no firm evidence of sympatry, which would lend support to the premise of separate species status, it was decided that some tests would be conducted to determine the genetic variability among and between Australian and SE Asian populations, and to see if these indicated more conclusive separation. DNA ANALYSIS. Fresh frozen specimens of D. ‘Tabiosum' were obtained from the following widely separated Australian localities: 1) Comboyuro Point, Moreton Bay, Qld (4 fish); 2) North West Island, Capricorn Group, Qld (4 fish); 3) Gulf of Carpentaria, Qld (3 fish); 4) off Port Headland, WA (3 fish). Specimens of D. pictum were obtained from Senoko Fishery Port, Singapore (4 fish). One specimen of Plectorhinchus flavomaculatus (Ehrenberg, 1830) from off Mackay, Queensland, Australia was used lo gain an outgroup sample. Frozen muscle tissue from the lower caudal peduncle was removed from each specimen for analysis. TABLE 3. Selected morphometric data of Diagramma melanacrum, D, pictum pictum and D. pictur labiosum. | D, melinaerum | D, p. pictur | D. p. labiosun | Number of i gnecifoare 5 33 40 Standardlengtly | 433.410 130-503 129-620 Percentage of standard length “Depth 38.0415 | 358-406 | 313413 | “Head length(HL) 295-332 | 268315 | 257326 | Orbit diameter 66-90 | 549.5 | AS -8.7 | Fauceleeivel| 226252 | 229.279 | 242089 GERD | 17-1L8 | 96-119 8.7-12.3 Pelvic fin length | 24.1-27.4 18,7-23.8 170-243 Ratio CPL:HL 1215 | 10-13 09-13 “Ratio CPD-CPL 2.0.2.8 22-31 24-22 NEW SPECIES OF HAEMULID FISH TABLE 4. Cytochrome B nucleotide diversity within (bold) and among pairs of samples of D. pictum (5) and D. ‘labiosum’ (1-4). - veel Se ie eet «is 1 | (loteion Bay 0.000 | 0.000 | 0.003) 0.002 0.005 | 2 NorthWestl, 9,000 | 0.000 | 0.003 | 0.002 | 0.005 3 fata (ne aP | 0.005 | 0.005 | 0.003} 0.001 | 0.003 | 4 Boe erent 0.005 | 0.005 | 0.006 aia 5 |Singe a 0.005 | 0.005 | 0.005 | 0.003 | 0.000 | Our aim with the mitochondrial DNA analysis was to determine if each of the D. ‘labiosum’ and D. pictum forms formed monophyletic mtDNA clades. Such a pattern would have allowed us to conclude that the 2 colour forms have existed allopatrically for a considerable amount of time (at least 4N generations (Avise et al., 1984)) and may have provided an insight to the 2 forms being separate species. The results are however inconclusive as mtDNA haplotypes do not cluster by colour form or location (Tables 4, 5). That haplotype 4 is shared between forms in the North west Shelf and Singapore provides evidence of gene flow between the 2 forms (Fig. 2). It should be noted however that several Indo-Pacific species of butterflyfishes Chaetodon are paraphyletic for Cytochrome B sequences despite being considered good species (MeMil- lan & Palumbi, 1995). This may reflect the rapid diversification of the Chaetodon species com- plex and suggests that there has not been sufficient time since speciation for complete mtDNA lineage sorting. Such a situation cannot be rejected for D. ‘labiosum’and D. pictum on the basis of our cytochrome B data set. Thus mitochondrial DNA analysis alone does not provide a perfect test of species status due to its haploid non-recombining nature (Avise, 1994). An assay of nuclear genes from areas where the two forms are sympatric would have provided a more powerful test of the taxonomic status of the two forms. This luxury was not afforded in the present study for two reasons. First, reports of the two forms existing in sympatry are anecdotal at best. The one report from southern Indonesia is based only on sightings of large adults, in which colour is a less reliable distinguishing feature. No specimens of D. ‘labiosum’ from this region have been located. Despite extensive collecting efforts off northern 663 TABLE 5. Cytochrome B haplotype frequencies for samples of D. pictum and D. ‘labiosum’. | ] | iia Haplotype Frequencies 2 3 4 | l 4 - : - - North West Is. 4 - _- | - | a Gulf of i | Carpentaria | ; - North West Shelf * f | }& 1 | Singapore = 7 . 4 uy Location Moreton Bay Australia, no specimens of the D. pictum form have been found. Second, tissue material could not be transported back to the laboratory from SE Asian sites in good enough condition to permit allozyme electrophoresis to be carried out. On this basis, and results enunciated below, it was decided that the separate colour forms be recognised as subspecies of D. pictum Thunberg. Diagramma pictum pictum (Thunberg, 1792) (Common name: Yellow-spotted Slatey) (Figs 3A-H, 8) Perca picta Thunberg, 1792 (Japan). Perca pertusa Thunberg, 1793 (Japan). Holocentrus radjaban Lacepéde, 1802 (East Indies). Diagramma balteatum Cuvier, 1830 (Java). REMARKS. D. pictum pictum is known from the W Pacific and Indo-Malay Archipelago, N to Japan and S to New Caledonia, but excluding Australia and S New Guinea. Various stages are figured in colour, from Japan (Burgess & Axelrod, 1972, pls 392-393; Masuda et al., 1984; Masuda & Kobayashi, 1994:162, plsl-3, 5-6 (note fish in pl. 4 are D. pictum labiosum from the Great Barrier Reef, Australia); Okamura et al., 1997: 352, 12 pls of specimens 2.5 to 65cm; Masuda, 2000: 96), through Taiwan (Shen, 1984: FIG, 2. Minimum spanning network of the relation- ships among Cytochrome B sequences. Each dash represents a single nucleotide substitution. 664 pl. 65 no. 327-6a,b,c, pl. 66 no. 327-8b&c; Burgess & Axelrod, 1974: pls 271, 275-276), the Philippines (JER photo QM NR63), Palau (Myers, 1999, pl. 67, a&b), Indonesia (Gloerfelt-Tarp & Kailola, 1984; Kuiter & Debelius, 1994; photos JER (QM NR62) and R.H. Kuiter (QM NRS52)), Rabaul, New Britain (Allen & Steene, 1987, pls 50-8), New Caledonia (Fourmanoir & Laboute, 1976: 99; JER photo QM NX793-795) and Santo, Vanuatu (Burgess & Axelrod, 1975, pl. 222-223). Fowler (1931) illustrated in black and white a range of 12 juvenile colour variations, one of which (fig. 22, lower left) is similar to a juvenile paratype of D. melanacrum (QMI20284), but lacks the width and intensity of black pigmentation to the lower caudal lobe. D. pictum pictum is distinguished by relatively large, close-set, yellow to burnt orange spots on head, body and unpaired fins in juveniles from about 160mm TL, subadults, and at least smaller adults. This subspecies progresses from the striped to fully spotted phase at 160-200mm TL. Diameter of the spots varies from about 1-3 times in the pupil or 2-5.3 times in the eye diameter, Short bars and wavy lines of the same colour are often present, mostly on the suborbital and operculum, in specimens larger than 160mm TL, but usually break up into spots in large specimens. Specimens to about 600mm TL usually retain conspicuous, although relatively smaller, spots on the body and particularly the cheeks. In fish above 800mm TL, spots on the body have often faded into a generally plain slate or silver-grey ground colouration, but there are usually at least some yellowish spots remaining on the lower head. We have no records of specimens with scattered dusky blotches, as is common in other large D. pictum subspecies. This subspecies also lacks the rows of bronze centres to individual scales present in D. pictum punctatum. The only other subspecies to share yellowish spots, or yellow pigmentation on the unpaired fins of all but small juveniles, is D. pictum cinerascens. However, in the latter the spots fade and disappear from the body at a much earlier stage, at about 300 to 400mm TL. Other subspecies have darker, usually brownish spots. The median tubed lateral-line scale count of 65 (range 57-74) is lower than the median of 69 recorded for D. pictum labiosum (range 59-78), but is higher than that for all other pictum subspecies (median 58-59, range 55-66). The median dorsal ray count (23) is lower than for D. pictum labiosum (24), but higher than that of D. MEMOIRS OF THE QUEENSLAND MUSEUM pictum cinerascens (22). The median total gill raker count (21) is higher than that for D. pictum labiosum and D. pictum cinerascens (20) and lower than that for D. pictum centurio (22). Diagramma pictum labiosum Macleay (Common name: Australian Slatey) (Figs 1G-H, 4A-H, 8) Diagramma labiosum Macleay, 1883 (Wide Bay, Queensland). REMARKS. D. pictum labiosum is known from S New Guinea and N Australia, from Houtman Abrolhos, WA, E to Lord Howe Island and Sydney, NSW. It is figured in Burgess & Axelrod (1976: pls 345 & 350-354), Coleman (1980: 160), Grant (1982: pl. 190; 1987: pl. 456), Sainsbury et al. (1985: 215), Allen & Steene (1987: pls 51-2), Randall et al. (1990 & 1997; 191), Kuiter (1993, 1996) and Masuda & Kobayashi (1994: 162, pl. 4). Macleay (1883) described D, /abiosum from a single 300mm specimen. The type is no longer extant (Eschmeyer, 1998). Macleay made reference to dense spotting on the soft dorsal and caudal fins, but not to any spots on the head and body. He stated that ‘the general colour is a dark silvery bluish grey’, This is consistent with fresh specimens from Australian waters, some of which had lost all spots on the head and body by 250mm TL, and all by 300-350mm TL, It is usual for other subspecies, except for D. pictum cen- turio from E Africa, to exhibit spots at 300mm. D. pictum labiosum has relatively small, close-set dark brown to bronze spots on the head and body in specimens from about 150-300mm TL. These spots vary in diameter from 2.6-6.5 times in pupil or 4.7-11.6 times in eye, rapidly diminishing in size on fish from 150-200mm TL. The stripes of the juvenile stage break up into a larger number of much finer spots than in any other subspecies. Golden-brown to dusky spots and short bars may be present on the suborbital and operculum of juveniles, but fade and disappear by about 200mm TL. Small dark brown spots gradually fade and disappear from the anterior toward the posterior part of the body with age. From 200-300mm TL spots on the body have reduced to a cluster of spots less than 1/4 pupil diameter, peppered on the upper half of the caudal peduncle. Beyond 350mm TL, all spots have disappeared and the head and body are plain slate to sooty silver-grey, often with some violet reflections on the cheeks. Adults, especially those in excess of 500mm TL, often develop scattered diffuse dusky blotches, In most NEW SPECIES OF HAEMULID FISH 665 FIG. 3. Diagramma pictum pictum; A, ~800mm TL, Maumere, Flores, R.H. Kuiter; B, 800mm TL, Menjangan Is., Bali, RLH. Kuiter; C, ~S50mm TL and ~250mm, Flores, Indonesia, R.H. Kuiter; D,~350mm TL, Tomini Bay, Sulawesi, R.H. Kuiter; E, ~300mm TL, New Caledonia, J-E. Randall; F, 236mm TL, Dumaguete, Philippines, J.E. Randall; G, ~200mm TL, Amed, Bali, R.H. Kuiter; H, ~150mm TL, Bitung, Sulawesi, R-H. Kuiter. individuals numerous small spots persist on the sizes, spots on these fins are greater in number caudal and rear of the soft dorsal fin, even in and smaller in diameter than in D. pictum pictum. specimens of 1000mm TL. At all comparable In specimens 350-500mm TL, a maximum of 666 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Diagramma pictum labiosum; A, 392mm TL, Moreton Bay, Australia, J.W. Johnson; B, 335mm TL, North West Is., Great Barrier Reef, Australia, J.W. Johnson; C, ~220mm TL, Shark Bay, Western Australia, R.J. McKay; D, 160mm TL, Gulf of Carpentaria, Australia, J.W. Johnson; E, 146mm TL, Gulf of Carpentaria, Australia J.W. Johnson; F, 136mm TL, Gulf of Carpentaria, Australia, J.W. Johnson; G, 126mm TL, Gulf of Carpentaria, Australia, J.W. Johnson; H, 108mm TL, Gulf of Carpentaria, Australia, J.W. Johnson. 9-14 rows of spots on the soft dorsal fin membrane and 12-20 rows of spots on the caudal fin membrane parallel to the fin rays were recorded, versus a maximum of 4-6 and 6-10 respectively for D. pictum pictum. This species lacks yellow or orange spots at any size and has no bars or wavy lines on the head, except occasionally in small juveniles. Adults have a scarlet to crimson-red throat, from the rear of the palate to the interior of the operculum and branchiostegal membranes, as well as the outer NEW SPECIES OF HAEMULID FISH gill arch membranes, broadly in common with other subspecies. D. pictum labiosum progresses from a striped to a fully spotted stage at a smaller size than other subspecies of Diagramma (130-160mm TL); and spots, while present, are smaller and more close-set at any given size than other subspecies. Subadults lose all markings on head and body ata smaller size (300-350mm TL) than all other subspecies, except some D. pictum centurio from E Africa. It differs from other subspecies by a higher median lateral-line scale count (69 versus 57-65) and, more importantly, the upper range of lateral line scale counts in all other Diagramma (except D. pictum pictum with 74), is below the median for D. pictum labiosum, The median total gill raker count (20) is lower than for all other D. pictum subspecies (21 or 22), except D. pictum cinerascens, and the median soft dorsal ray count (24) is higher than all other D. pictum subspecies (22 or 25). Diagramma pictum cinerascens Cuvier (Common name: Fork-striped Slatey) (Figs SA-H, 8) Diagramma cinerascens Cuvier, 1830 (Trincomalee, Sri Lanka), Diagramma hlochii Cuvier, 1830 (Trincomalee, Sri Lanka). Diagramma poecilopterum Cuvier, 1830 (Pondicherry, India). REMARKS. Diagramma pictum cinerascens occurs in the Indian Ocean from the Bay of Bengal to the Persian Gulf. It is figured in colour photographs taken by R.H. Kuiter (QM NR48-49), Kuiter (1998:104), JER (QM NL742) and H. Voigtmann (in Debelius, 1999: 100, top left) from the Maldives; from Sri Lanka (Debelius, 1999: 100); from the Persian Gulf by Randall (1992, pl. 130a) and J. Hoover (in Randall, 1995, pls 528-530) and from the Gulf of Oman by P. Woodhead (in: Debelius, 1993: 127 top left, p. 128 centre, and Debelius, 1998: 88 centre) from Ras al Hamra, Oman. The brief original description of D, cinerascens is from a 256mm SL specimen, MNHN 7803 (Bauchot et al., 1983). The holotype has a uniform brown body and small darker brown spots on the rays of the dorsal and anal fins. Cuvier (1830) gave a dorsal ray count of ‘XII/16?? for D. blochii, which suggests a species of Plectorhinchus, however his description was based on a drawing by Raynaud and the query given after the count suggests he had difficulty in discerning the true fin formula from the drawing. His dorsal, anal and pectoral 667 fin formulae tor D. poecilopterum are also shown to be erroneous by Smith (1962). Smith examined the holotype and paratype (MNHN 7811 from Pondicherry and 7810 from Trincomalee, respectively) and obtained counts consistent for this form (Table 2). Stripes on the 155mm SL holotype, as depicted in Smith (1962: fig. 22), are usual for specimens of this size. This subspecies differs from others in the configuration of the body stripes of juveniles, the size of the spots, and nature of the lines on the cheeks and operculum of large juveniles to small adults. Juveniles progress from a striped to a fully spotted phase at 180-240mm TL. Juveniles of about 150mm TL often have broad, clearly defined body stripes, not yet beginning to break into broken lines or numerous spots, as is usual for other Diagramma of this size. As mentioned by Day (1878), unlike juvenile stages of any other form of Diagramma, the second and third stripes on the body merge to form a single stripe above the posterior half of the pectoral fin (Day, 1878, pl. 21, fig. 3; Debelius, 1993, fig. p. 127; Randall, 1992, pl. 130a; Smith, 1962, fig. 22). At the point where the stripes merge, there is often a small break or up-curved interruption in the stripe. The front of the head, including the snout to the interorbital and suborbital, as well as much of the dorsal and caudal fin membrane, may be bright yellow in fish of up to about 200mm TL. In specimens from about 200-350mm TL, there are usually wavy yellowish lines or spots on the operculum and cheeks. The lines are narrow in larger specimens, but vary in width in small specimens. Spots on the body are tan to orange- brown, in longitudinal rows. In specimens about 180-300mm TL, the spots range from 3.1-6 in eye diameter. The spots are generally smaller and more close-set than D. pictum pictum, larger and more close-set than D. pictum centurio and larger and more sparse than in D. pictum labiosum. Unlike the latter, they do not fade and disappear first from the dorso-anterior region, producing a life-stage with a cluster of spots on the upper part of the caudal peduncle only. The only other subspecies to have lines or bars on the head of specimens above 180mm TLis D. pictum pictun, however, in the latter, these are usually broader and accompanied by yellow or orange spots on the body. Specimens in excess of 400mm TL are generally plain slate to silver-grey, often with scattered irregular dusky blotches on the head and body, similar to large D. pictum labiosum. However, unlike the latter, narrow wavy yellowish lines or small spots may still be evident 668 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5. Diagramma pictum cinerascens; A, ~800mm TL, Oman, J.P. Hoover; B,~500mm TL, Great Basses, Sri Lanka, R.C. Anderson; C, ~450mm TL, Maldives, R.H. Kuiter; D, ~300mm TL, Oman, J.P. Hoover; E, ~240mm TL, Trincomalee, Sri Lanka, R.C. Anderson; F, ~200mm TL, Trincomalee, Sri Lanka, R.C. Anderson; G, 172mm TL, off Bahrain, Persian Gulf, J.E. Randall; H, ~150mm TL, Oman, P. Woodhead. NEW SPECIES OF HAEMULID FISH from the suborbital to the rear of the operculum in some larger specimens. This subspecies is best distinguished by the unique striped pattern of the juvenile and by the narrow yellowish wavy lines or spots on the head in small adults. It has a lower median lateral line scale count (58) than D. pictum pictum (65) and D, pictum labiosum (69); fewer median total gill rakers (20) than other D. pictum subspecies (21 or 22), except D. pictum labiosum; and fewer median dorsal fin rays (22) than all other subspecies (23 or 24). Diagramma pictum punctatum Cuvier (Common name: Red Sea Slatey) (Figs 6A-H, 8) Diagramma punctatum Cuvier, 1830 (Red Sea). Diagramma punctatum Riippell, 1830 (Northern Red Sea). Diagramma cinerascens (non Cuvier) Riippell, 1830 (Red Sea) - name preoccupied by D. cinerascens Cuvier. REMARKS. Diagramma pictum punctatum is known only from the Red Sea. It is figured in colour photographs by GR. Allen at Jeddah (QMNRS54-55); JER at Sudan and Nuweiba, Egypt (McKay, in Fischer & Bianchi, 1984: vol. II, pl. 2; Randall, 1983: pl. 111; Randall, 1992: pl. 130b); JER photos at Gulf of Aqaba (QMNX806) and Dahab, Egypt (QMNX807); R. Kuiter photo at Egypt (QM NX800); D. Eichler photo (QMNX798) and H. Debelius photos at Sinai, Egypt (Debelius, 1998: 88 upper left (adult) and upper right (large juvenile)), QMNX799. Cuvier (1830) in his account of D. punctatun, initially referred to 3 specimens collected by Ehrenberg. A description and meristic formulae for this material was quoted separately and a single specimen 9 inches in length was listed. These specimens were from the Red Sea and are represented by one unregistered specimen in the Natural History Museum, Humbolt University, Berlin (Eschmeyer, 1998). Cuvier went on to mention other material of Diagramma sourced by Kuhl and van Hasselt from Java (RMNH D 2172); Stadhouder from the Indian Archipelago (MNHN 7836 (1), MNHN A.7832 - latter specimen is the holotype of D. radjabau Lacepéde, 1802); Raynaud from Batavia (MNHN 7801 (1)); and Quoy & Gaimard from Vanicolo (MNHN 7802 (2)). The museum catalog numbers for these specimens were listed by Bauchot et al. (1983) and Eschmeyer (1998). Cuvier recognised all the material from the East Indies as D. radjabau of Lacepede (1802), and surmised that the Ehrenberg material from the Red Sea was ‘the same species, or very near’ to it. However, Cuvier’s description of D. punctatum was not a compilation of the Red Sea and East Indian forms of Diagramma. The Ehrenberg material was treated separately in the initial 669 stages of the paper, followed by an account of other specimens from the east, including a redescription of the type material of Holocentrus radjabau, Also, Cuvier, by his equivocal comments above, cast doubt on whether the other specimens were conspecific with the Red Sea material. The unregistered ZMB specimen from the Ehrenberg collection is implicitly the holotype and the rest constitute other material, rather than part of a syntypic series. Riippell (1830) also proposed D. punctatum for specimens from the N Red Sea, catalogued SMF 2215 (2), but appears to have been pre-empted by Cuvier (Eschmeyer, 1998). D. pictum punctatum has dark brown, orange-brown to golden tan spots on the head and body of most specimens in excess of 140mm TL. In smaller specimens the spots are usually slightly elongate, and individual spots along the rows may be slightly inclined obliquely. Juveniles transform at about 135-145mm TL from a colour pattern including 3 broad dark stripes, to a phase that is uniformly marked with spots of approximately equal size. The spots on the body of large juveniles and subadults (180-350mm TL) are smaller than in D. pictum pictum, similar to D. pictum cinerascens, but larger than other Diagramma subspecies. They are fewer than in D. pictum labiosum and usually greater than in D. pictum centurio. From about 140-350mm TL spots generally decrease from about 1/2 to 1/4 pupil diameter, or 4.1-6.6 in eye diameter. At this size, there is no bright yellow pigmentation to the head, body or fin membranes, as in some D. pictum cinerascens and D. pictum pictum. The ground colour of adults is plain silver-grey to grey-bronze. Adults usually have small round dark brown spots on the upper head and along the upper body to the caudal peduncle, above the lateral-line. The only other subspecies to have distinct spots on the body in adults is D. pictum pictum, however in the latter the spots (when present) are larger, usually lighter in colour and not confined to the upper margin of the body. In adults there are bronze centres to individual scales, forming rows, a feature absent or very ill-defined in other forms. Few specimens were available for examination; however the lateral-line scale counts obtained (55-61) are at the low end of the range for the genus and modally well below that for D. pictum labiosum. 670 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 6. Diagramma pictum punctanun; A, 594mm TL, Sudan, Red Sea, J.-E. Randall; B, ~500mm TL, Ras Muhammad, Sinai, Red Sea, H. Debelius: C, ~400mm TL, Daheb, Sinai, Red Sea, J.E. Randall; D,~300mm TL, Jeddah, Red Sea, GR. Allen; E, 222mm TL, Nuweiba, Sinai, Red Sea, J.E. Randall; F,~200mm TL, Egypt, Red Sea, R. H. Kuiter; G, 180mm TL, Red Sea, J.E. Randall; H, 132mm TL, Eilat, Gulf of Aqaba, Red Sea, J.E. Randall. Diagramma pictum centurio Cuvier REMARKS. D. pictum centurio is known from (Common name: East African Slatey) the Seychelles to E Africa and § to Durban. It has (Figs 7A-F, 8) been photographed by JWJ (QM NRS33, Diagramme centurio Cuvier, 1830 (Seychelles), subadult), G.R. Allen (QM NRS56, large juvenile), N. Coleman (subadult) and H. Debelius NEW SPECIES OF HAEMULID FISH 671 FIG. 7. Diagramma pictum centurio; A, ~500mm TL, Mahe, Seychelles, H. Debelius; B, ~350mm TL, Pemba, Kenya, D. Eichler; C,~250mm TL, Pemba, Kenya, D. Eichler; D, ~300mm TL, Mahe, Seychelles, N. Coleman; E, ~230mm TL, Mahe, Seychelles, GR.Allen; F, ~200mm TL, Mahe, Seychelles, H. Debelius. (Debelius, 1999: 99 (adult and juvenile) from Mahe, Seychelles; by D. Eichler (Eichler & Lieske, 1994: 132, from Pemba, Kenya; and a 340mm specimen from Inhaca, Mozambique and a 750mm specimen from Shimoni, Kenya are illustrated (Smith, 1949, pl. 42, fig. 688; Smith, 1962, figs 1, 2). Cuvier’s description of 257mm SL holotype (MNNH8526) of D. centurio mentioned scattered small brown spots on the nape, upper body and caudal region. Playfair & Giinther (1867) reported a specimen 16 inches from the Seychelles, with ‘head, back, sides, upper part of tail, dorsal and sometimes caudal with brownish-yellow spots’. No small juveniles from the Seychelles were available for examination, however Debelius (1999: 99 left) illustrated a specimen estimated to be about 200mm TL, with numerous small dark brown spots peppered on the head and upper body. Subadults from this area have slightly larger, sparsely distributed, orange-brown spots on the upper head, body mostly above the lateral line, and on the caudal peduncle. The spots are Al2 smaller ihan im similar sized D. pict piciini, 2. picnim cineraseens. or D, pictum punefatum, and are more sparse than in D, pletimi febiastan. Adulls are slate to silver-grey on the head and body, and may retain a few seattered small spots onthe upper body, near the base of the dorsal fin. The sott dorsal and caudal fins may have numerous small close-sel datk brown spots, similar to D. pictum labiosum. There appears to be some variability 1 colour- alion between individuals from the Seychelles and those from E_Africa. Preserved specimens of in excess of 170mm TL fram coastal E Atrica lack spots on the head and body, and spots on the dorsal and caudal fins, if present, are faint. Fish of this size allustrated in underwater photographs were unvormly silvergrey with no distinctive markings, however few photographs from this region Were available for examination. Juveniles are striped ia about 150mm TL, but the stripes rapidly regress, forming profuse rows of small dark spots by about (50-160mm TL. Smith's (1949, 1962) figures of a 340mm subadult from Mozambique, however, clearly illustrate fine dark spots on the body and unpaired fins, similar 10 those in the photograph of Debelius (1999) from Mahe. These spots appear to have been somewhat artist-enhaneed in Smith's latter reproduction, bal prior io bis expedition to the Seychelles in 1954, Smith (1949) alluded briefly lo the spots in adults, Identification of this subspecies is complicated by some variability between regions, however large juveniles (beyond the striped phase) may be peppered with tiny dark brawn spots, and the subadult stage may best be distinguished by the presence of small, sparse orange-brown spots on the body. Alternatively, large juveniles and subadults lacking any distinctive markings (Some E. Altican specimens) are separable in that all other subspecies, except some D. protmm labiosuni, have spots ofsome type at 160-350mm TL. Also, the median upper, lower and total vill raker counts are highest for all Qiagen; the median dorsal! ray count (23) is higher than in D plerum cinerascens (22) but lower than in D, pictun labiosum (24); and the median lateral-line seule count (59) is lawer than in D. prenen pinttan (65) and D. pictum labiasum (69). DISCUSSION D. mekimacrunris the most distinctive member ofthe genus, with its low scale count, more robust body and unique colouration, The subspecies of D. pictum, although differing considerably in MEMOIRS OF THE QUEENSLAND MUSEUM colouration between one another at certain onto- genetic siqzes, are remarkably similar as adults. Only D. pietum pietim retains obyious spots on the body as an adult. D. melanacrim differs from all .D. pictun Subspecies in having the third or fourth dorsal spine longest (second dorsal spine usually longest in the latter. fourth always shorter than second); a taller first dorsal spine, 1.7-2,3 in length of second (2,3-3.9 m the latter), a stoufer caudal peduncle, depth of pedunele in iis length 2-2.2 (2.2-3.1 in speenmens greater than 130mm SL of the latter) and in having pelvie tins that at least reach the vent in all life stages (pelvic fins reach lo or slightly beyond the vent in most juvemiles up to 200mm SL, but become increasingly remote with growth, large adults fallme short by over half length of tin in J, pictun subspecies). The tubed lateral-line scale count Of 55-37 is lower than that of D. pierun lahiasuim (59-78, usually higher than 65), bur overlaps the extreme lower end of the range for olher D. piclum subspecies (cunmulative ranges 55-74). Seale rows above the lateral line to the origin ofthe dorsal fin, at 14-15, are lower than D. pictum labiosum (17-19) and usually lower than other D_ pienmesubspecies (15-18). The range of scale counts for D, melanuerum, however, could reasonably be expected to expand when further specimens are avadable for examination. There are also noteworthy differences in colour. Adults of D pictym pictint have orange to yellow spots on the head and usually also on the body; adults of D. pictuny piiicratum have brown spots on the upper head and body, but they are smaller, more evenly rounded and move sparsely distributed; adults of other D, picrun subspecies are slate or silvery-grey, Sometimes with large irregular scatiered dark blotches. They all lack numerous smal! close-set dark brown spots on a yellowish background (bronze spots Lo centres of scales in D, pichim punctatum are Vague. smaller, lighter in colour and generally lollow along the scale rows), and ihe prominently black lower part of the caudal lin. Other subspecies also usually lack the intensity of black pizmentation to outer part of the anal and pelvic fins, although some juveriics may have dusky fins. D. melancerum is heht yellow dorsally on the body,.as well as. the dorsal 4in and upper 3/4 of the caudal fin, and the dark spots on the upper pan of the body and on these fins are larger, darker and nat arranged one per scale; also there are no large dark blotches on the body. Ditgrommnia melanacrum appears to be rare compared to most other Indo-Pacific haemulids, NEW SPECIES JER has observed it only 3 titnes in Indonesia and not in the Philippines, Borneo or New Guinea. in spite of many hours underwater af numerous localities in the East Indies region. Nor was it encountered in fish markets, Pieter Bleeker apparently failed to find this species during his many productive years of research on the fishes of Indonesia. KEY TO DI4GRAMMA SPECIES N.B. Differences in colouration between most subspecies of D. pietwm is limited mainly ta certain ontogenetic stages. Small initial stage juveniles are generally patterned with broad Jongitudinal black stripes on a cream to ereamishi yellow background, and are generally nol separable using this key. Transtormation from striped to spotted colour phases oceurs ar various lengths between subspecies. Large adults of labiasum, cineraseens and centurio are all generally silver-grey to slate-grey, often with scattered dusky blotches, but differ medially in several meristic features, The colouration of small D. melanacrum juveniles is unknown. {. Lower 1/4-1/3 of caudal fin black: upper body and Heal With numendus sinall Close-set dark brown spots; thirdor fourth dorsul spine lonuest: first dorsal spine | 723 in length of second; pelvic fins reaching to orbeyond anus: vaudal-pedinele depth 2.9-2.2 tn its lenwth: tube luteral-line scales about $§-37 “cA: a5 Diomtelitniae runt (Philippines, Borneo, Sulawesiand southern (ndonesiu) Lower 1/4-1/3 of caudal fin not black (often dusky ts black Near lower margin in juveniles and subadults); upper body und head of adults wathoul small Glascescl dark brown spots, second dorsal spine Wsually longest, first dorsal spine 2.3 or more yn length of second, pelvic fins not reaching anus in adults (may extend tr anus in juvemlos), cuudul-poduncle depth usually ied thar 2,2 inits length; Wubed liatersal-line scales 535-78, 2 Lemon yellow, orange to orange-tun spots on head and body of large juveniles and subadulls. spots lane, 2-6 in eye diameter; adulls cilher lacking spots on the head and body. ur with large yellow to orange spots. af least on head: short burs or wavy lines usually on head of lane juveniles and subadults; ground colour af median. fins often bright yellow in jpventlts (6 subudules: puvenles transform from siriped to fully spolted phase al 160-240mM TL ee ' 3 Dark brown, bronze or orange-brown spats on head and body of lame javeniles and stubadults, spats usually smaller, 4-I] in cye diameter; adulls ecnerally silverprey to slate prey, often with scattered dusky blotches, sometimes with small brown spots, but never with large yellow or orange spots; no short hars or wavy lines on head of juveniles and sobadulls, ground colour of median fins never brivht yellow (except occasionally in initial slage suveniles of up to about 100mm TL), juveniles transform trom striped to fully apt phuse at 130-L60imm TL , : 4 1 Tubed hueral-Line velo a7- 7 ried 5): doarun! atl third body stripes afjuveniles not meqging near peetort t2 § OF HAEMULID FISH 073 fin; largo-yellow (o bum orange spots usually presentan head und body of adults (if not, hen at least ya checks); no scattered wvevuluy dusky blutehes an head and body of udults; head. of subadults und adults asually wath combination of spots, short bars and broad wavy lines: spots present on head and body of large juveniles to adults, 2-5 in eye diameter; tolal gill takers [9-23 (median 21) : LD pact peri (New Caledonia north to southern Japan wnd west to the tndo-Matay Vechipelage, BrHleding Australia and southern Now Guinca) Ot Tubed lateral-line suules 55-04 {moclian 58): pane, auil (Hird body stripes of juvenites merging above posterior half of pectoral Ain din specimens of about [50mm TL), Hi Spols on bady of adults: scattered imegular dusky blotches often present on head and body of aciilis; head of subadulis and adults with narrow wavy lines, small spots or unmarked; spots presen! on head aid body of large juveniles tosubadults, 3-0 in eve chameter; total yall rakers 17 2) (inedian 20) . 2D, pista cinerasee ns (Northern Walia Quean, from Bay of Bengal to the Persian Gulf) 4, Spots on head and body of large juventles ta aubadults (TS8C-350min TL) larger, 4. 1-6,01n eye diameter, olten shutily elongate and obliques adults usually with small scultered round spots on upper head and along body Wear dorsal fin base and caudal peduncle, adults with small bronze centres lo mdividial scales, fonming rows ( 2D. prenem punerarum (Red Sea) Spats on heud and betty of large juveniles to subsdults (1K0-350mI6 TL) smaller, 65-116 i eye diameter, always round: adulls either lackime or wilt only few! spots on head and body; adults without small bronze centresta individual seals. . _ 5 5 Fubed jateral-line scales 59-7# (median 69): tual 2 will rakers usually [9-21 (median 20), spols on body (when present) bronze tu dark brown, always close-set, gradually disappearing trom anténor toward posterior pert of body With growth, ultimately leaving @ srnall patch of spots al upper part of caudal pedunele, before fading completely 5, pictum labiasun (northern Australia, from Hounman Abralhos, Wi te Sydacy Harbour, NSW; southern New Guinea) - Tubed lateraltine scales 56-06 (median 59); total pill rakers 21-23 (median 22): spots on body (when present) usually orunge-brown, peppered finely in some laree juveniies, but sparsely distributed in subadulis, not eridually disappearing from anterior toward posterior part of body with growth, not leaving wsmall pateh of spots al upper part of caudal Meneses before tading completely, . Oo plemmeenturia (East Anica lo Seychelles} OTHER MATERIAL EXAMINED (Fig. 8) {Nurnbers in brackets are lengths of specimens in mim, Catalogue numbers refer fg single specimens unless otherwise indicated by number in italics) D. pictim lahiosum QUEENSLAND: QMI337L (148); QMI3442 (124): QMI3946 (203)> QMIH079 (187); QMI6678 (168): QMI7094 (261); QMI7S79 (24%): QMI7804 (283); OMI78S3 (115); OMII1555 (266); QM112535 (129); OMI12536 (127): QMLII71 (20K): QM) 2676 (292), QMI12724-30 7 (224-337), MIT 2908 674 60° 60° @ D. melanacrum te D. pictum pictum WY D. pictum labiosum @ DP. pictum cinerascens A D. pictum punctatum BD. pictum centurio HAY OF BENGAL ANDAMAN Is 7 4, ARABIAN SEA fh = Socotra LACCADIVE! 1st SAI LANKA , NICOBAR Maldives e is INDIAN OCEAN CHAGOS ARCH * Dwgo Garcia MSEYCHELLES we Cocos-Keeling © Mauntius Reunion MADAGASCAR + Amsterdam * Saint-Paul Chetan MEMOIRS OF THE QUEENSLAND MUSEUM 180° BERING = SEA SEA OF OKHUTSK At ae R ‘e #, e Ls OGASAWARA + ARCH > Minami: Ton Shima (Marcus) Drainan » MARIANAIS SOUTH CHINA Qonines | SEA “2 Guam 2 LA yao % PALAU, © 1S CAROLINE IS Nau Banaba * SOLOMON 1S (Gceuny . ic ; TUVALU **. WALLIS & | FUTUNA, Fidi ; VANUATU ore: CORAL SEA * a New ee LOYALTY Caledonia |S +Nortolk Lord Howe NEW ZEALAND | ; TASMAN ‘WS SEA TASMANIA Q 50° FIG. 8. Confirmed distributional records of examined Diagramma species 2 (117-119); QMI15058 (156); QMI15097 (118); QMI15965 (98); QMI16467 (116); QMI16852 2 (96-127); QMI20132 (192); QMI20850 (107); QMI20884 (123); QMI21223 (110); QMI21334 (151); QMI23229 2 84-126); QMI30757 (401); QMI30758 (434); QMI30759 (370); QMI30760 (380); QMI30761 (601); QMI30762 (620); BPBM14335 (349); BPBM14465 (138). Northern Territory: NTMS10577-001 (222); NTMS11613-028 (356); NTMS13318-002 3 (175-213). WESTERN AUSTRALIA: QMI10225 2 (83-105); QMI14230 (112); QMI30726 2 (123-196); QMI30727 (255); QMI30728 (232); QMI30729 (238); QMI31110 (468); QMI31111 (519); QMI31112 (511); NTMS10987-002 (436); CSIROH3834-02 (341). D. pictum pictum. JAPAN: MUFS12534 (192); MUFS12847 (245); MUFS11816-7 2 (136-156.5); MUFS12162 (142); MUFS12226 (114.5); QMI3 1403-4 2 (372-377). INDONESIA: BMNH1858.4.21:364 (85); QMI20383 (304); QMI20284 (136); QMI20286 (100); QMI20287 (186); QMI20288 (503); QMI20289 (302); QMI20304 (61); BPBM36675 (38); BPBM18593 (99); BPBM20678 (239); BPBM30100 2 (60-61); NTMS11037-001 (332); NTMS11127-016 (95). THAILAND: QMI21687 /2 (58-212); PMBC14425 (152); PMBCS5884 (190), PMBC5885 (185), PMBC5890 (175) and PMBCS891 (187). MALAYSIA: AMIA3343 (94); QMI30873 (311); QMI30874 (325); QMI31076 3, 136-188); QMI31113 (224). PHILIPPINES: AMI10568 (118); AMI10503 (120); BPBM28550 (95); BPBM22143 (187). TAIWAN: BPBM18687 (200). VANUATU: AMI17142-015 (173). NEW CALEDONIA: AMIB2410 (275). SOLOMON ISLANDS: BPBM17371 2 (172-245). PAPUA NEW GUINEA: BPBM15709 (104). D. pictum cinerascens. NDIA: AMI15599-006 (125); AMB8320 (141); BMNH1888.11.6:7-8 2 (147-181); BMNH1889.2.1:2961 (45); BMNH1847.11.22:134-136 2 (74-90); BPBM20666 2 (67-86); PMBC5886 (181), PMBC5887 (181), PMBC5892 (167), PMBC5893 (169) and PMBC5894 (197). GULF OF OMAN: BMNH1888.12.29:48 (172). PERSIAN GULF: BMNH1904.5.25:184 (78); BMNH2000.4.19:1113 (155); BPBM33174 (68); BPBM21181 3 (63-132); BPBM29495 3, 117-144). D. pictum punctatum. RED SEA: BMNH 1860.11.9:93 (157); BMNH1960.3.15:763 (36); BMNH1871.4.13:9 2, NEW SPECIES OF HAEMULID Fish! (193-737); RUSIT9R9 2 (113-128); BPBM 19856 (180); BPBM31867 (100). D. picnim venturio. SEYCHELLES: AMI32067-001 (232), KENYA; RUSI41682 (168), RUSI41677 2, (50-61). TANZANIA! BMNH1985,7,.9:193 (203), MOZAMBIQUE: RUSI41656 4, (59-125), RUSH 1681 (161); RUSI41680 (141), RUSI4227 (127); RUSI41679 (122); RUSIS63R6 2, (98), RUSI41678 (76). MADAGASCAR: RUSI52829 10, 73-105), SOUTH AFRICA: RUSI3846 (68), RUSII2754 (46); BMNH1919.9,12:24 (193), ACKNOWLEDGEMENTS We are grateful to Klaus FE. Fiedler, Dieter Eichler, Rudie H. Kutter and J, Barry TMutechins for providing underwater photographs of D, melunacrum, Rudie WW. Kuiter, R. Charles Anderson, Dieter Eichler, Helmut Debelius, Gerald R, Allen, Philip Woodhead, Neville Coleman and John Hoover provided photographs of various phases of D, pietum, Phillip C. Heemstra and Andrew Bentley of the JLB Smith Institute of Ichthyology, Chatatsee Angtanya of the Phuket Marine Biological Centre, Anthony C, Gill of the Natural History Museum, Yukio Iwatsuki and Miroyuki Molomura of the Miyazaki University, Helen K. Larson of the Northern Territory Museum and Mark A. McGrouther of the Australian Museum made available comparative material of Diagramme under their care, Alastair Graham ofthe Division of Fisheries of CSIRO, Hobart arranged for fresh specimens to be sent from WA and sent a duplicate colour slide of the holotype of D. melanacrum taken by Thomas Giloerfelt-Tarp. Patricia J, Kailola kindly gave us permission to reproduce this photograph herein, Peter K.L, Ng and Thomas Tan Han Tong sent fresh specimens from Singapore and Michael McDade collected specimens from North West Island. Special thanks are due to Roland J. McKay of the Queensland Museum for assistance in the description of D. melanacriym. LITERATURE CITED ALLEN, GR, 1993, Part 7, Fishes of Ashmore Reel‘and Cartier Island. Records of the Western Australian Museum Supplement 44; 67-91. ALLEN, GR. & STEENE, R.C, 1987, Pacific marine fishes, Book 10, Reef lishes of the Indian Ocean, (UF, Publications: Neptune City, New Jersey), AVISE, F.C, 1994, Molecular markers, natural history and evolution, (Chapman and Hall: New York), AVISE, LC,, BERMINGHAM, E., KESSLER, L.G. & SAUNDERS, N.C 1984. Characterization of mitochondrial DNA variability ina hybrid swarm 67s berween subspecies of bluegill suntish (Leponiis macrochirus). Evolution 38(5); 941-941, BALCHOT. 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Pl. 1-370. In The fishes of the Japanese Archipelago. (Tokai University Press: Tokyo). MASUDA, H. & KOBAYASHI, Y. 1994. Grand atlas of fish life modes: color variation in Japanese fish. (Tokai University Press, Tokyo). MYERS, R.F. 1999. Micronesian reef fishes. (Coral Graphics: Guam). MEMOIRS OF THE QUEENSLAND MUSEUM NEI, M. 1987. Molecular evolutionary genetics. (Columbia University Press: New York). OKAMURA, O., AMAOKA, K. & OKATA, Y. 1997. Sea fishes of Japan. (Yama-kei Publishers: Tokyo). PALUMBI, S.R., MARTIN, A., ROMANO, S., McMILLAN, W.O., STICE, L. & GRABOWSKI, G. 1991. The simple fool’s guide to PCR. (Department. of Zoology, University of Hawaii: Honolulu). PLAYFAIR, R.L. & GUNTHER, A.C.L.G. 1867. The fishes of Zanzibar. (John Van Voorst: London). RANDALL, J.E. 1983. Red Sea reef fishes. (Immel Publishing: London). 1992, Divers guide to fishes of Maldives. (Immel Publishing: London). 1995. Coastal fishes of Oman. (Crawford House Press: Bathurst, Australia). RANDALL, J.E., ALLEN, GR. & STEENE, R.C. 1990. Fishes of the Great Barrier Reef and Coral Sea. (Crawford House Press: Bathurst). 1997, Fishes of the Great Barrier Reef and Coral Sea. (Crawford House Press: Bathurst). RUPPELL, W.P.E.S. 1830. Atlas zu der reise im nordlichen Afrika, fische. (Fische des Rothen Meeres: Frankfurt). SAINSBURY, K.J., KAILOLA, PJ. & LEYLAND, GG, 1985. Continental shelf fishes of northern and north-western Australia. (Clouston & Hall & Peter Pownall Fisheries Information Service: Canberra). SAMBROOK, J., FRITSCH, E.F. & MANIATIS, T. 1989. Molecular cloning: a laboratory manual. (Cold Spring Harbour Laboratory Press: New York). SHEN, S.C. 1984. Coastal fishes of Taiwan. (Shih- Chieh Shen, National Taiwan University: Taipei). SMITH, J.L.B. 1949. Sea fishes of southern Africa. (Central News Agency: South Africa). 1962. Fishes of the family Gaterinidae of the western Indian Ocean and the Red Sea with a resume of all known Indo Pacific species. Ichthyological Bulletin of the Rhodes University 25: 469-502. NEW OCTOPUS SPECIES FROM QUEENSLAND MARK D. NORMAN Norman, M.D. 2001 06 30: New octopus species from Queensland. Memoirs of the Queensland Museum 46(2): 677-690. Brisbane. ISSN 0079-8835. Three new octopus species are added to the growing list of Australian octopods. All are known solely on the basis of trawl material from habitats not easily surveyed using standard diving techniques. Octopus harpedon sp. noy. attains arm spans in excess of 2m. It occurs in shallow muddy waters in the Gulf of Carpentaria. Octopus bulbus sp. noy. is also a long-armed species, which possesses a swollen bulbous ligula and a skin ridge around the lateral mantle. Octopus micros sp. nov. is tiny (mature at <25mm mantle length), the first pygmy species to be reported with a lateral mantle ridge. The latter 2 species occur on the continental shelf off southern Queensland in water depths of 18-195m and 166-195m, respectively. The phylogenetic affinities and potential life styles of these octopuses are discussed in light of their morphological attributes. Octopus, taxonomy, morphology, Queensland, Cephalopoda. Department of Zoology, University of Melbourne, Parkville 3052, Australia; Present address: Marine Invertebrates, Melbourne Museum, GPO Box 666E, Melbourne, 3001, Australia (e-mail: m.norman@unimelb.edu.au); 17 March 2000. Research into benthic octopuses of Australia and the Indo-West Pacific region over the past decade has revealed more than 70 new species of octopus in Australian waters (Stranks, 1988a-b, 1990; Norman, 1992, 1993a-d, 1998; Stranks & Norman, 1993; Norman et al., 1997; Norman, unpubl. data), the bulk of which await formal description. Within this fauna are 3 distinctive and very different octopus species from Queensland waters. The ‘Spaghetti Octopus’, Octopus harpedon sp. nov. is reported from the shallow waters of the Gulf of Carpentaria. This octopus has extremely elongate arms (up to 10 times mantle length) and would attain arm spans of >2m when foraging over its soft sediment habitat. The other two species were collected from continental shelf waters (>150m) off the south coast of Queensland. The ‘Swell-Club Octopus’, Octopus bulbus sp. nov., is also long-armed (arms to 5 times mantle length) with a distinctive swollen ligula. Arm spans of live animals would be up to 50cm. This species possesses a distinctive lateral mantle ridge. The ‘Pygmy Keeled Octopus’, Octopus micros sp. nov., 1s tiny with an arm span of <12cm and a weight of <6g. It is the first pygmy species to be reported with a lateral mantle ridge. There are few specimens available for these species, all originating from trawl surveys. None of these new species have been observed live. Based on the available material, the taxonomic affinities and potential habits of these little- known creatures are discussed. SYSTEMATICS FAMILY OCTOPODIDAE Octopus harpedon sp. nov. (Figs 1-2, 9A-B, 10A-C) MATERIAL. HOLOTYPE: 16,: 56.9mm ML, AMC30411, Albatross Bay, near Weipa, Gulf of Carpentaria. PARATYPE: 12, 96.lmm ML, AM C304112, SE Gulf of Carpentaria, 16°27°40"S, 141°15°25”E, 2m. TYPE LOCALITY. Albatross Bay, near Weipa, Gulf of Carpentaria, Australia ETYMOLOGY. Greek harpedon, thread-like; referring to elongate, thread-like arms.”Spaghetti Octopus” is proposed as a common name. DIAGNOSIS. Moderate-sized (ML to 96mm) with spindle-shaped mantle and bulbous eyes. Arms greatly elongate, up to 9 times mantle length. Second arm pair appears longest (arm formula approximately 2>1=3>4). Webs very shallow, less than 5% of arm length. Approximately 270 suckers on intact arms of mature animals. Third right arm of male hectocotylised, very short (only 20% of length of opposite arm) and bearing 49 suckers. 10-11 gill lamellae per demibranch. Eggs large-type. Skin largely unpigmented, dark blue subdermal pigment around eyes produces ‘bruised’ appearance. Skin smooth. Lateral ridge absent. GTB PIG. 1. Oetapus harpedon sp, nov. (¢ holotype, AM C30411). A, dorsal view (scale bar = 30mm): B, ovantle and arm crown (scale bar = 30mm); C, copulatory organ (seale bar = 2mm). DESCRIPTION. Counts and measurements were taken offa single specimen of each sex, the only known specimens of this distinctive species. Raw morphological data are presented in Table 1. Moderate sized species with extremely long arms (Figs. LA~B, YA-B); mantle lengths (ML) 56.9mm for male holotype and 96.lmm for female paratype. Total length of @ 394mm and 3 997mm: weight to at least 104¢. Mantle elongate to spindle-shaped, much longer than wide (¢; width 35.1% of ML; ¢: width 24.9% of ML), mantle walls thin to moderately muscular. Head narrower than mantle (¢ 226.40 of ML. 75.0% of mantle width; “+ 15.8% af ML, 63.6% of mantle width). Eyes moderate to small, only slightly pronounced. Stylets absent. Mantle aperture of moderate width, approximately half circumference of body at level of opening. Funnel narrow and elongate, approximately 50% of mantle length (451.5% of ML; 2: 47.9% of ML), iree portion short, approximately one third of funnel length (4 : 23.9% of funnel length; 2: MEMOIRS OF THE QUEENSLAND MUSEUM 33.5% of fuel length), Funnel organ not evident in either specimen, Arms extremely long, longest >5 times mantle length (¢: 5.9 =~ ML; 9:91 * ML), Arms narrow relative to mantle length (d': 9.3% of ML; ® [ex frozen]: 5.9% of ML), widest midway along arm, roughly square in cross section. Arms unequal in length, ventral pair distinctly shortest, second pair appears longest (arm formula; <: ~2>)>3>4, 9: ~2>1=3>4). Suckers in two rows and of moderate size (a: 7.0% of ML, 2: 5.0% of ML), slightly elevated with only slight flare, No enlarged suckers evident in either sex, Approximately 280 suckers on intact arms of mature animals (4 : 272 suckers on lefi third arm; &: 291 suckers on left third arm). Webs tiny, shortest relative to arm length reported for any octopus (4 : deepest web 3.9% of longest ann; 9 : deepest web 2.4% of longest arm). Web sectors approximately equal in length. Web margins along arms absent. Third right arm of males hectocotylised. Modified arm very short, less than mantle length (79.1% of ML), and around 20% of length of opposite arm (20.4% of opposite arm). Ligula moderate-size (7.3% of arm length, 5.8% of mantle length), in the form ofa small thick-lipped spoon with transverse creases across the open ligula groove (Fig. 1C). Calamus distinct and sharp, approximately one half of ligula length (54.6% of ligula). Spermatophore groove well developed and wide with fine transverse creases, Spermatophore guide distinct with a ridge of elevated square papillae. Forty-nine suckers on hectocotylised arm of single male. Gills with 10-11 lamellae on both inner and outer demibranchs, plus terminal lamella. Digestive tract (Fig. 2A). Anterior salivary glands extend along approximately one third of buccal mass from posterior margin on dorsal surface. Posterior salivary glands elongate and of moderate length (similar in length to buccal mass, approximately 30% of digestive gland length). Crop diverticulum present, long and narrow. Stomach bipartite. Caecum coiled in approximately 1.5 whorls with distinct striations. Digestive gland long and narrow, not bound in an iridescent membrane. Narrow intestine reflexed several times in proximal third, Ink sac well developed, embedded in ventral surface of digestive gland. Anal flaps absent. Upper beak with a hooked rostrum, concave and notched on the cutting edge, and a small hood (Fig. 2B). Lower beak with narrow short rostrum, hood NEW OCTOPUSES FROM QUEENSLAND narrow, widely spread wings and slightly flared lateral walls (Fig. 2C-D). Radula with 7 teeth and 2 marginal plates in each transverse row (Fig. 10A-C). Rhachidian tooth with 2-3 lateral cusps, on each side of moderately long medial cone (Fig. 10A). Lateral cusps in asymmetrical seriation, migrating from lateral to medial position over 7-8 transverse rows (Fig. 10B). First lateral teeth unicuspidate with cusp towards lateral edge. Second lateral teeth unicuspidate and long with curved base. Lateral marginal teeth straight. Marginal plates oblong and plain (Fig. 10C). Male genital tract not fully formed. Submature terminal organ (‘penis’) T-shaped with diverticulum slightly longer than distal portion. Female genital tract not fully formed but eggs in tiny ovary already large and low in numbers (<100), This species would produce large eggs and the young are likely to be benthic on hatching. Oviducts elongate, opening posterior to the narrow short septum. Colour in life unknown. Preserved specimens uniformly cream (¢) and pink (3d) with dark subdermal pigmentation around the eyes of both specimens giving a ‘bruised’ appearance (Fig. 1B). Dorsal White Spots (sensu Packard & Sanders, 1971) absent. Skin smooth. Lateral mantle ridge absent. Nothing is known of the behaviour and general biology. The short hecto- cotylised arm of the d suggests mounted copulation in this species as opposed to other octopus species where a long hectocotylised arm enables copulation from a distance (as found in other long-armed species such as O. aculeatus, Norman & Finn, 2001). Numerous spiral parasites were present along sections of the digestive tract, particularly adjacent to the crop and intestine. TAXONOMIC REMARKS. Only one other Australian octopus has arms of comparable relative length. Ameloctopus litoralis Norman, 1992 is an intertidal, smaller species (ML to 30mm) with arms up to 10 times the mantle length. It is distinguished from O. harpedon in that it has a much lower gill count (5-6 versus 10-11 lamellae per demibranch), a linear terminal organ which lacks a diverticulum, distinctive bands along the arms (compared with little pigmentation in O. harpedon), and it lacks an ink sac. An undescribed octopus from Hong Kong, China and Taiwan also shows similarities to this new species, sharing similar arm lengths and gill 679 FIG. 2. Octopus harpedon sp. nov. A, digestive tract (2, AM C304112): as = anterior salivary glands, bm = buccal mass, c= caecum, cd=crop diverticulum, er =crop, dg = digestive gland, i= intestine, is = ink sac, o = oesophagus, ps = posterior salivary gland, s = stomach (scale bar = 10mm). B-D, beaks (d holotype, AM C30411) (scale bar = 3mm); B, upper beak, lateral view; C, lower beak, lateral view: D, lower beak, ventral view. counts. This Asian taxon has been treated under a number of names by various authors: Octopus sp. B. Voss & Williamson, 1972; Octopus fusiformis (non Brock, 1887) in Dong, 1987; Octopus sp. | Norman & Hochberg, 1994. It can be distinguished from O. harpedon on the basis of differences in arm formula (1>2>3>4 versus second pair longest in O. harpedon) and hectocotylised arm length (~50% versus 20% of opposite arm length in O. harpedon). Additional mature material of both taxa is required to further resolve the relationship between these octopuses. As for higher taxonomic affinities, O. harpedon shares a number of characters with poorly-known Ezaxoctopus Voss, 1971. Euax- octopus contains 2 quite different species from 680 FIG. 3. Octopus bulbus sp. nov. (do holotype, MV F87067); A, dorsal view (scale bar = 20mm); A, mantle (scale bar = 10mm); B, funnel organ (scale bar = 5mm); C, copulatory organ (scale bar = 5mm). either side of the Panama Isthmus. E. pillsburyae Voss, 1971 and E. panamensis Voss, 1975 share long arms of which the second pair are longest, shallow webs, paired semi-circular ocelli on the mantle, a large crop, a rhachidian tooth of the radula with 1-2 lateral cusps on each side (typically 1), a blunt linear terminal organ with a diverticulum longer than the distal free portion, and distinctive spermatophores with flattened coils in the oral tip. The two member species differ in that £. panamensis has a hectocotylised left arm, a gill count of 11-13 lamellae, and a VV-shaped funnel organ, while £. pillsburyae has the right third arm hectocotylised, a gill count of 7, and a W-shaped funnel organ. Octopus harpedon shares the same arm formula (second pair longest) and blunt linear terminal organ with a diverticulum longer than the distal free portion. It differs in lacking the semicircular ocelli on the mantle and possesses a radula with a higher number of lateral cusps on the rhachidian tooth MEMOIRS OF THE QUEENSLAND MUSEUM (2-3 per side). In the absence of mature material of QO. harpedon spermatophores can not be compared. The disparate composition of Euwax- octopus and the absence of replicate well- preserved material for both this genus and O. harpedon prevent further resolution of their affinities. Based on available material, I place this species in Octopus. Octopus harpedon shares several morphological characters with the ‘Octopus macropus group’ (Norman, 1993a). This group 1s characterised by an arm formula of 1>2>3>4, high gill counts (10+ lamellae per demibranch) and a rhachidian tooth of the radula with 2-3 cusps on each side of the medial tooth, migrating from medial to lateral positions over 7-8 transverse rows. The radula and gill count of O. harpedon match those of the Octopus macropus group but arm length, arm formula and ligula shape differ. Until more material becomes available (including fresh tissue for molecular analyses), the higher affinities of this distinctive octopus remain unknown. Octopus bulbus sp. nov. (Figs 3-5, 9C, 10D-F) MATERIAL. HOLOTYPE: 1d; 41.lmm ML, MV 87067, east of Mooloolaba, 16-20 miles north of Cape Moreton, southern Queensland, 90-106fim (166-195m), trawl, FV ‘Debie-Marie’, 11-13 Aug 1981, coll. G Smith, Queensland Fisheries. PARATYPES: 12: 53.0mm ML, MV F87068, East of Noosa, southern Queensland, 63fm (116 m), trawl, FV ‘Rhonda Lane’, 12 Dec 1980, coll. M. Potter, Queensland Fisheries; 12: 49.2mm ML, MV F87069, off Mooloolaba, southern Queensland, 10fm (18.4m), trawl, 1500-1600hrs, FV ‘Rhonda Lane’, 14 Dec 1980, coll. M. Potter, Queensland Fisheries. TYPE LOCALITY. East of Moololaba, southern Queensland, Australia. ETYMOLOGY. Latin bu/bus, fleshy swelling; referring to the distinctive swollen ligula. ‘Bulb-tip Octopus’ is proposed as a common name, DIAGNOSIS. Small (ML to 50mm) with relatively long arms, approximately 5 times mantle length. Dorsal arms longest, receding to ventral arm pair (arm formula 1>2>3>4). 7-8 gill lamellae per demibranch. Approximately 200 suckers on intact arms. Third right arm of dd hectocotylised, bearing approximately 90 suckers. Posterior salivary glands large and elongate (almost twice length of buccal mass, approximately 60% of digestive gland length). Ligula large (8% of hectocotylised arm length) NEW OCTOPUSES FROM QUEENSLAND with greatly swollen lips to the ligula groove. Spermatophores thick and short, approximately halfmantle length. Dorsal mantle and arm crown sculptured with scattered pink-red raised patches. Lateral ridge present. DESCRIPTION. Counts and measurements were taken off the only known specimens, 2 mature dd and a submature 2. Morphological data are presented in Table 1. Moderate-sized elongate (Fig. 3A); mantle length to around 50mm (¢: to 53.0mm ML, ?: 49.2mm ML). Total length to 341mm; weight to at least 47g. Mantle elongate to spindle-shaped, much longer than wide (width 53.5% of ML in holotype, other material distorted from freezing), mantle walls moderately muscular. Head approximately same width as mantle (53.3% of ML, 99.5% of mantle width in holotype, other material distorted from freezing). Eyes large and pronounced. Stylets absent. Mantle aperture of moderate width, approximately half circum- ference of body at level of opening. Funnel broad-based, approximately one half of mantle length (41.8-62.2% of ML), free portion elongate, 32.1-56.0% of funnel length. Funnel organ W-shaped with broad limbs (Fig. 3A). Outer limbs slightly shorter in length than median limbs (outer limbs 86.4% of median limbs in holotype). Funnel organ occupies approximately two thirds of funnel length (59.8% of funnel length). Arms long, approximately 5 times mantle length (longest 4.9-5.6 x ML). Arms of moderate width relative to mantle length (13.9% of ML), widest at one third of arm length from base then tapering evenly to tip, roughly square in cross section. Arms unequal in length, dorsal pair longest (arm formula: 1>2>3>4). Suckers in 2 rows and of moderate size (6.7-9.8% of ML), slightly elevated with low flare and a deep cup. No enlarged suckers evident in either sex, Approximately 190 suckers on intact arms of mature animals (196 suckers on fourth right arm of holotype). Webs shallow and thin (deepest web 10.9-11.5% of longest arm in intact spec- imens). Dorsal and lateral web sectors approx- imately equal in length, ventral webs shallower (web formula A=B=C>D>E). Web margins extend along both dorsal and ventral aboral edges ofarms for less than one third of the arm length. Third right arm of dd hectocotylised. Modified arm relatively short, almost halfas long as normal arms (2.0-2.5 * ML, 56.3% of opposite arm). Ligula large (8.0-8.6% of 681 FIG. 4. Octopus bulbus sp. nov. digestive system (2 paratype, MV F87068); A, digestive tract: symbols as in Fig, 2 (scale bar = 5mm). B-D, beaks; B, upper beak, lateral view; C, lower beak, lateral view; D, lower beak, ventral view (scale bar = 3mm). hectocotylised arm length, 16.1-21.9% of mantle length), in the form of an elongate oval with a deep closed groove (Fig. 3C-D). Calamus small and sharp, <20% of ligula length (17.2-18.2% of ligula). Spermatophore groove well developed, of moderate width with fine transverse creases. Spermatophore guide not obvious. Approx- imately 90 suckers on hectocotylised arm (91, 94). Gills with 7-8 lamellae on both inner and outer demibranchs, plus terminal lamella. Digestive tract (Fig. 4A). Anterior salivary glands extend along approximately one third of buccal mass from posterior margin on dorsal surface. Posterior salivary glands large and elongate (almost twice length of buccal mass, approximately 60% of digestive gland length). Crop diverticulum present, moderately small. Stomach bipartite. Caecum coiled in 1.5 whorls, with distinct striations. Digestive gland approx- imately ovoid. Muscular intestine reflexed approximately one third along length from proximal end. Ink sac wel-developed, embedded in ventral surface of digestive gland. Anal flaps present, small. Upper beak with a hooked rostrum, concave on cutting edge, and moderate 682 FIG. 5. Octopus bulbus sp. nov. reproductive system (3 holotype, MV F87067). A, reproductive tract: mg = mucilagenous gland, ns = Needham’s sac, sg] = spermatophoric gland, sg2 = accessory spermatophoric gland, t = testis, to = terminal organ (‘penis’), vd = vas deferens (scale bar = 5mm); B, spermatophore: ea = ejaculatory apparatus; sr = sperm reservoir (scale bar = 5mm). hood (Fig. 4B). Lower beak with pointed rostrum, narrow hood, widely-spread wings and slightly flared lateral walls (Fig. 4C-D). Radula with 7 teeth and 2 marginal plates in each transverse row (Fig. 10D-F). Rhachidian tooth with 2-3 lateral cusps, typically 2, on each side of long thin medial cone (Fig. 10D). Lateral cusps in asymmetrical seriation, migrating from lateral to medial position over 7-8 transverse rows (Fig. 10E). First lateral teeth unicuspidate with cusp towards lateral edge. Second lateral teeth unicuspidate with curved base. Lateral marginal teeth robust and curved. Marginal plates oblong and plain (Fig. 10F). Male genitalia (Fig. 5A). Terminal organ (‘penis’) robust and roughly linear with a diverticulum of similar length as the free distal portion. Distorted by spermatophore in holotype (Fig. 5A). Spermatophores (Fig. 5B) short, around half mantle length (22mm, 53.5% of ML in holotype), and thick (1.1mm, 2.7% of ML), produced in low numbers (1 in spermatophore storage sac, | in terminal organ in holotype). Oral MEMOIRS OF THE QUEENSLAND MUSEUM cap contains thick coils of ejaculatory apparatus and bears a thick cap thread. Sperm reservoir long, 54% of spermatophore length in holotype, containing a thick sperm cord forming coiled in approximately 24 regular whorl. Only ¢ spec- imen is submature. Submature eggs large-type and produced in low numbers (<100). The large eggs indicate that juveniles are likely to adopt a benthic habit on hatching. Colour in life unknown. Preserved specimens cream with pink-red raised patches scattered on dorsal mantle and upper arm crown. Pink low small papillae on dorsal skin between larger patches. Regular fine papillae on ventral mantle, Dark blue subdermal pigmentation around eyes with superficial red brown chromatophores (Figs 3A, 9C). Webs cream in contrast to pink brown arms. Dorsal White Spots (sensu Packard & Sanders, 1971) absent. Scattering of low papillae pronounced around eyes. Lateral mantle ridge present. Small regular low papillae on ventral mantle within lateral ridges. Nothing known of behaviour or general biology. DISTRIBUTION. Octopus bulbus sp. nov. is known from only 3 specimens, collected off southern Queensland, in 18-195m. TAXONOMIC REMARKS. Octopus bulbus shares a number of attributes with O. australis Stranks & Norman, 1993, also found in shallower waters in the region. Both species share a lateral ridge, bulbous ligula and similar gill counts (7-8 in O. bulbus versus 7-9 in O. australis). However, these taxa have very different floorplans and reproductive characters, suggesting separate evolutionary origins. Octopus bulbus has arm and web formulae in which dorsal arms and webs are longer/deeper (AF 1>2>3>4, WF A=B=C >D>E), whereas O. australis has longer/deeper lateral arms and webs (AF 3>2>4>1, WF typically D>C>B>E>A). Octopus bulbus also has longer arms (4.9-5.6 versus 2.7-4.3 x ML), shallower webs (10-12% versus 20-30% of longest arm), a proportionally shorter hecto- cotylised arm (56% versus 66-86% length of opposite arm) with a higher sucker count (91, 94 versus 62-77), absence of enlarged suckers in mature dod (suckers 6.7-9.8% versus 12.6-15.3% ML), and spermatophores with far fewer sperm cord whorls (24 versus >60). The longer dorsal arms (arm formula ]>2>3>4) and a multicuspid radula (2-3 cusps on each side of the rhachidian tooth) are similar to NEW OCTOPUSES FROM QUEENSLAND those of the ‘Octopus macrapus group’ (Norman, 1993a). However, the gill count of 7-8 is lower than any previously reported for the group all of which possess 10-15 lamellae per demibranch. Until more material becomes avatlable (including fresh tissue for molecular analyses), the higher affinities remain unknown. Octopus micros sp. nov. (Figs 6-8, 9D, 10G-I1) MATERIAL, HOLOTYPE: I¢: L8.8mm ML, MV F87070, east of Mooloolaba, 16-20 miles north of Cape Moreton, southern Queensland, 90-106fm (166-195m). trawl, FV ‘Debie-Marie’, 11-13 Aug 1981, coll. G Smith, Queensland Fisheries. Paratypes: | ¢: 20.7mm ML, ] 2: 34.5mm ML, MY F78815, off Mooloolaba, southern Queensland, trawl, FV ‘Debie-Marie’, 11-13 Aug 1981, coll, G Smith, Queensland Fisheries (no depth data). TYPE LOCALITY. East of Moololaba, southern (Queensland. ETYMOLOGY. Greek mikros, small referring to its small size. ‘Pygmy Keeled Octopus’ is proposed as a common name. DIAGNOSIS. Small species (ML to 25mm) with short arms (2-3 » ML) of approximately equal length, dorsal pair slightly shorter. Lateral webs slightly deeper than dorsal web, Enlarged suckers absent in both sexes. Gills with 6 lamellae per demibranch. Hectocotylised arm (third mght) approximately 80% of length of opposite arm. Ligula of moderate size (~6% of arm length) with open groove. 85-93 suckers on hectocotylised arm. Terminal organ (penis) robust and linear with simple rounded diverticulum. Spermatophores approximately equal in length with mantle. Lateral mantle ridge present. DESCRIPTION. Counts and measurements are from the known specimens, 2 mature dd anda submature 7 (Table 1). Robust pyginy species (Fig. 6A); mantle length to 20.7mm for dd, 24.5mm for ¢. Length to 91mm: weight to at least 5.82. Mantle ovoid to spherical, slightly longer than wide (width 63.3-76.1% of ML), mantle walls moderately muscular. Head width similar to mantle (56.3-76,1% of ML, 89.0-100% of mantle width). Eyes moderate to large and moderately pronounced. Stylets present (Fig. 6B). non- mineralised, 4.4mm in holotype, 23.4% of ML. Mantle aperture of moderate width, approx- imately half circumference of body at level of opening. Funnel broad and short, approximately one third of mantle lenvth (33.0-41.1% of ML), 653 FIG. 6. Octopus micros sp. nov, (¢ holotype, MV F87070). A, dorsal view; B, stylet (scale bar=2mtn). C, funnel organ (2 paratype, MV F78815) (scale bar = 3mm). D, copulatory organ (¢ paratype, MY F78S815) (seale bar = 2mm). free portion 44.7-63.5% of funnel length. Funnel organ W-shaped with broad limbs (Fig. 6C). Outer limbs similar in length to median limbs (outer limbs 92.9-105.2% of median limbs), Funnel organ occupies approximately two thirds of funnel length (60.4-67,7% of funnel length). Arms of moderate length, longest approx- imately 2.5 times mantle length (2.3-2.7 * ML). Arms moderately robust relative to mantle length (16,9-19.7% of ML) tapering eyenly to fine tips, rounded in cross section. Arms roughly equal im length, dorsal pair slightly shorter (arm formula typically 4=3=2>1). Suckers in 2 rows and of moderate size (9.8-11.7% of ML), slightly eley- ated with moderate flare and a thin rim. Enlarged suckers absent in both sexes. Approximately 150 suckers on intact arms of mature animals. Webs of moderate depth (deepest web 22.3-27.1% of 684 FIG. 7, Octopus micros sp. nov. digestive system (2 paratype, MV F78815); A, digestive tract, symbolsas in Pig. 2 (scale bar = Smm), B-D, beaks (scale bar = jinm), B, upper beak. lateral view; C, lower beak, lateral view; D, lower beak, ventral view. longest arm). Lateral web sectors slightly deeper than other webs (web formula B=C=D>E=>A). Web margins extend as thin ridges for a short distance along ventral edge of arms. Third right arm of @d@ hectocotylised. Modified arm relatively long, almost as long as normal arms (2.0-2.2 x ML, 86% of opposite arm in intact male). Ligula moderate-size (6.4% of arm length, [4.4% of mantle length), in the form of an elongate pointed leaf with fine trans- verse creases across the open ligula groove (Fig. 6D). Calamus distinct and sharp, approximately one half of ligula length (44.4% of ligula). Spermatophore groove well-developed and wide with fine transverse creases. Spermatophore guide shallow with no obvious papillae. 85-93 suckers on hectocotylised arm, MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 8. Octopus micros sp. nov. reproductive system (¢ holotype, MV F87070), symbols as in Fig, 5; A, teproductive tract (scale bar = 5mm); B, spermatophore (scale bar = 3mm), Gills with 6 lamellae on both inner and outer demibranchs, plus terminal lamella. Digestive tract (Fig. 7A), Anterior salivary glands extend along approximately 20%. of buccal mass trom posterior margin on dorsal surface, Posterior salivary glands large (slightly longer than buccal mass, approximately 80% of digestive gland length), Crop diverticulum present, small. Stomach bipartite. Caecum coiled in single whorl, with striations. Digestive gland approximately ovoid. Muscular intestine rellexed approximately one third along length from proximal end. Ink sac well developed, embedded in ventral surface of digestive gland. Anal flaps present. Upper beak with slightly hooked rostrum and small hood (Fig. 7B). Lower beak with rounded rostrum, hood narrow, widely spread wings and nearly parallel lateral walls (Fig. 7C-D). Radula with 7 teeth and 2 marginal plates in each transverse row (Fig. 10G-H). PIG.9, Type material. A, Octopus harpedansp. nov. (4 holotype, AM C3041 1) dorsal view; 1-4: arms numbered from dorsal to ventral pair (scale bar = 50mm). B, Octopus harpedan sp, nov. (¥ holotype, AM C30412) dorsal view; I-4: as in A (scale bar = 50mm). C, Qetopus bilbus sp. nov. (d holotype. MV F87067), dorso-lateral view; lr= lateral mantle ridge (scale bar = 20mm), D, Octopus micras sp. noy. (S holotype, MV P87070) dorsal view (scale bar = 10mm). NEW OCTOPUSES FROM QUEENSLAND 685 686 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 10. Radulae. A-C, Octopus harpedon sp. nov. radula (2 paratype, AM C304112). A, dorsal view showing multicuspid rhachidian tooth; B, lateral view showing serial progression of cusps on rhachidian tooth; C, lateral teeth and marginal plates. D-F, Octopus bulbus sp. nov. radula (2 paratype, MV F87069); D, dorsal view showing multicuspid rhachidian tooth; E, lateral view showing serial progression of cusps on rhachidian tooth; F, lateral teeth and marginal plates. G-H, Octopus mticros sp. nov. radula (2 paratype, MV F78815); G dorsal view; H, lateral view. NEW OCTOPUSES FROM QUEENSLAND Rhachidian tooth with 1-2 lateral cusps, typically 1, on each side of short robust medial cone (Fig. 10G). Lateral cusps in asymmetrical seriation, migrating from lateral to medial position over 4-5 transverse rows (Fig. 10H). First lateral teeth unicuspidate with cusp towards lateral edge. Second lateral teeth unicuspidate and long with curved base. Lateral marginal teeth long and straight. Marginal plates square, plain (Fig. 10H). Male genitalia (Fig. 8A). Terminal organ (‘penis’) in mature dd short and robust with simple swollen diverticulum. Spermatophores (Fig. 8B) approximately equal in length with mantle length (20.1, 22.8mm, 97.1, 121.3% ML), and of moderate width (0.5mm [n=2], 2.3, 2.5 % of spermatophore length), produced in low numbers (2, 4 in spermatophore storage sac). Ejaculatory apparatus linear with slight coils at oral end. Oral cap simple, bearing long cap thread. Sperm reservoir 37.3% of total spermato- phore length in dd, containing robust sperm cord, most of which forms regular whorls with some bunching to produce a plaited appearance. Submature ° with eggs in undeveloped ovary already large (~2mm) and few in number (<100). This species would produce large eggs and the young are likely to be benthic on hatching. Colour in life unknown. Preserved specimens red-brown formed by fine uniform chromato- phores on dorsal surfaces and lateral arm crown to midline of third arm pair. Darker purple-brown pigmentation around eyes creates a ‘bruised’ appearance to the eyes (Fig. 6A). Dorsal White Spots (sensu Packard & Sanders, 1971) present. Skin relatively soft (preservation artefact?) with a single distinct primary papilla over each eye and 4 papillae in a diamond on the dorsal mantle. Large primary papillae on posterior tip of mantle. Lateral mantle ridge present. Nothing known of behaviour or general biology. Its depth range (>150m) and small size make it unlikely ever to be observed in the wild. TAXONOMIC REMARKS. O. micros sp. nov. is the first pygmy species reported with a lateral mantle ridge. Two other larger species in the area also possess this ridge, O. bul/bus described here and O, australis Stranks & Norman, 1993, Octopus micros is clearly distinguished from QO. bulbus by the arm formula (4=3=2>1 versus |>2>3>4), shorter arms (2.3-2.7 versus 4.9-5.6 x ML), lower gill count (6 versus 7-8) and the presence of ‘dorsal white spots’ (sensu Packard & Sanders, 1971) and the diamond of primary 687 papillae on the dorsal mantle (latter two attributes absent in O. bulbus). O. micros is distinguished from O. australis by a higher sucker count on the hectocotylised arm (85-93 versus 62-77), shorter arms (2.3-2.7 versus 2.7-4.3 = ML) and a lower gill count (6 versus 7-9). These 2 species share several characters, namely a lateral mantle ridge, similar arm formulae, ‘dorsal white spots’ and a diamond of primary papillae on the dorsal mantle. These species may share common ancestry. Only 2 pygmy species have been reported from Australian waters: O. warringa Stranks, 1990 and O. superciliosus Quoy & Gaimard, 1832 (see Stranks, 1988a). Both are restricted to temperate southern Australian waters. They are easily distinguished from O, micros in that they both lack a lateral mantle ridge. DISCUSSION Due to the nature of certain marine habitats, our only knowledge of some sea creatures comes from dead material collected by fishing or research trawls. For such animals our capacity to interpret their lives is restricted to deductions based on morphology, stomach contents and habitat associations. The new species described here are all from environments where direct observation is difficult or impossible due to poor water clarity or excessive depths. Combined with the cryptic and/or nocturnal behaviour typical of octopuses, it is likely that none of these species will be observed in their natural environs. Attributes of their morphology, however, may provide some clues to their lifestyles. O. harpedon is a long-armed species from shallow coastal waters typically clouded with suspended silt over soft sediment substrates. This area is also the regular haunt of abundant tiger sharks and crocodiles, effectively deterring any attempt to find and observe this species in sitv. The long arms of O. harpedon are similar in scale to those of Ameloctopus litoralis Norman, 1992, a small intertidal mudflat species found across northern Australia. Ameloctopus litoralis lacks significant webs between the arms and feeds by probing its long and thin arms individually down holes and burrows to capture small crustaceans and fish (Norman, 1992). The long arms and shallow webs of O. harpedon suggest that it is similarly using long arms to probe subterranean burrows for crustacean and fish prey. The absence of complex colour patterns 688 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE |. Counts and measurements (mm) for O. harpedon, O. bulbus and O. micros spp. nov. d= damaged; fr = frozen distorted specimen; H =hectocotylised arm; InD = indistinct; t= very tip ofarm damaged; TO = terminal organ; A-E = web sectors from dorsal sector. Species O. harpedon | O. harpedon | _O. bulbus O. bulbus | O. bulbus O. micros O. micros O. micros | “Museum AMS AMS | MV MV | MV MV MV MV_ Reg.No. | C304111 C304112 F87067 F87068 F87069 F87070 | F78815 F78815 ‘Status a Holotype Paratype Holotype Paratype Paratype Holotype Paratype | Paratype [Sex Male | Female | —_—Male _ Male Female _Male | Male Female | Maturity | submature_ | submature mature mature submature | mature mature | _submature Mantle length __ 56.9 { 96.1 41.1 | 53.0 49.2 18.8 20.7_ 24.5 Total length _ 304 997 251 318 67 67 nm | 91 Weight (g) 453 | 1043 | 27.5 47.2 30.1 4.5 4.0 5.8 Mantle width 20.0 23.9 22.0 23d 22d “143 13.1 15,5 Head width 15.0 152 | 219 17d 22d 143 | 12.3 13.8 Funnel length 29.3 46.0 20.9 33.0 24.3 6.2 8.5 | 9.6 Free funnel length 7.0 Isa) |) $7 13.0 13.6 37 | 38 | 6 Funnel organ limb (medial). InD iD | 128 InD InD 42 | ID 5.8 Funnel organ limb (lateral) In | ms 108 InD | InD 3.9 ind ae etal Sl ae E: 15 E: 18 E: 26 A: 19f A: 10 A: 10 A: 10 CCF ihe | . = ben sat ia BCE: 13 C:21 BC: 23 B: 34 D: 30fr CD: 13 C213 C: 14.5 ie] - veers ee a ae) ie = : _ 7 (meas 246d 292 | 5431591 200d dd 275 239d 3741 391 43 55d i 2 290d?334_ | d 877° 171 167 2431262 233.253 48.50 45 49 dd 3. _-|_22145H_ =| 628495d | 106d 82H | 240135H | 176199 391 42H 48 41H dd - 4 143 148 | 298t 308t 125 144 187 173 159d 47 48 4748 60t 65 Arm width — 6.1 a ES fr fr 37 | 38 | 45 Sucker diameter 4.0 | 4.8 3.0 5.2 [33 2.2 2.3 2.4 Sucker count: R3 _49H d IH 94H fr 93H 85H ds | L3 272 291 |__ 196 (R4) fr fr 156 (R4) 143 153 (R4) | 7 if - a 7 Gill lamellae court 10-10 11-10 Ae 8-8 8-8 66 | 66 6-7 L 10-10 10-1 1-1 8-8 8-8 6-6 6-6 6-7 | | Ligula length 3.3 7 oS 11.6 - 27 36 = Calamus length 1.8 as | Ten 2.0 - he eet) - fi aa ge : : 1(+1inTO)} — InD ’ 3 (+1 in TO) 2 : ed le : 2 In| 23 20 R Pepin : : Ll InD 3 0.6 0.5 A LY — = 3. = a - 7 pee seco : : 12 InD ; 8.5 75 - Egg number > <100 - - <100 . - - <100 — Egg length | - large - - large | - - large | Egg width - = - - . > Y: -_ -_ and skin sculpture suggest that this species may known in a handful of other described octopods, be nocturnally active. namely the Australian shallow water O. australis Hoyle, 1885 and O. berrima Stranks & Norman, The other 2 species described here both possess 1993, and the deeper water Benthoctopus leio- a lateral mantle ridge. This skin structure is only derma (Berry, 1911), Eledone palari Lu & NEW OCTOPUSES FROM QUEENSLAND Stranks, 1992, Megaleledone senoi Taki, 196) and members af Bathypolyprs Grimpe. |921, Pareledane Robson, 1932, Scaeurgus Troschel, 1857 and Tetracheledone Voss, 1955, A partial or broken Jateral mantle ridge also occurs in two other shallow water species: O. hunurong of the ‘Ocrepus macropus group’ (Norman, 1993a) and QO. fungsiva of the ‘Octopus aegina group’ (Norman, 1993h). Presence of this skin ridge in such disparate taxa suggests either independent origins for this structure or that it is a common primitive State lost in many groups. All the species listed are primarily associated with soft sediment substrates and it is possible that this ridge may relate to their capacity to bury in sand ur mud. Octopus micros 1s only known from the continental shelf off southern Queensland, This species shows some similarities with Q. australis and QO. berrima, from eastern and southern Australia, respeetively. All 3 species are unlike any other known shallow-water species and they may represent older Palaco-austral lineages, the product of the long and isolated northward drift of the Australian continent following the break up of Gondwana around 200 million years ago. The small size at maturity of O. micros is similar to that found in other pygmy species. Such dwarfism appears linked with specialised microhabitats. O. micropyrsus Berry, 1953 of California (Lang. 1997) and a species from Tasmania (unpubl. daia) occur primanly w kelp holdfasts (Meeroeystis spp.) while Q. hocks Adam, 1941 of the tropical Indo-West Pacitic lives in coral heads (pers. obs.). Juveniles of many octopus species seek refuge in such porous structures (pers. obs.). Jt is possible that selective pressures against outgrowing the protection of such micro-refuges (and their resident prey species) may have lead to neotenous dwartism in certain groups, The small size of O. micros may reflect a similar specialization for particular microhabitats, ACKNOWLEDGEMENTS This research was supported by the Australian Biological Resources Study. Sincere thanks to the following people for assistance with various aspects of this study: Drs Mike Vecchione and Louise Allcock for their constructive comments on the manuseript. David Paul for photography, Joan Clark for SEM, and lan Loch and Phil Colman (Australian Museum) and Chris Rowley (Museum Vietoria) for curatorial assistance. 68Y LITERATURE CITED DONC, 4. 1987, Fauna Sittica, Phylum Mollusca. class Cephalopoda, (Science Press: Beijing, China). NORMAN, M.D. 1992. “imelectopuy tifaraliy gen. & , Cj e Scare a i a) yf o fe % \ \“Y ' : ) Vv S FIG. 1. Cepidognathus adonis sp. nov., 3. A, idiosoma, dorsal view; B, detail of anterior epimeral plate and posterior pair of setae; C, idiosma, ventral view; D, gnathosoma, ventral view; E-G, lelt ocular plate in 3 specimens; H, 4 genitoanal plate. Abbreviations: aa, anterior areola; AD, anterior dorsal plate: AE, anterior epimeral plate; ar, areola; ca, canaliculi; ds-| to ds-6, dorsal setae | to 6; ep, epimeral pore; yb, gnathosomal base; glp-! to glp-4, gland pores | to 4; Ja, lateral areola: Ic, lateral costa; ma, medial areola: me, medial costa: ms, maxillary setae; OC, ocular plate; pa, posterior areola; pe, pore canaliculus; PD, posterior dorsal plate; PE. posterior epimeral plate; pgs, perigenital seta; pi. pits; P-1 to P-4, palp segments | to 4; ro. rostrum; rp, rosette pore; sgs, subgenital seta, Scale bars: A, C, D-H = SOum; B= 251m. band of shallow pits in the anterior half from 27-30 perigenital setae. Gnathosoma as depicted about the inner-most seta, this group not in Fig. 1D. Ventrolateral lamellae on telofemora contiguous with anterolateral areola; posterior(o — elaborate (Fig. 2A-D), with smooth or slightly leg IV insertions without canaliculi. Male with undulate edge. Ventral margin of tibia | lacking a 720 MEMOIRS OF THE QUEENSLAND MUSEUM G — FIG. 2. Copidognathus adonis sp. nov., 3; A, leg 1, medial view; B, leg IT, medial view; C, leg III, medial view; D, leg IV, medial view; E, claws of tarsus II, medial view. Abbreviations: ba, basifemur; bp, bipectinate seta; ds, dorsal setae; d-pas, doubled parambulacral seta; ec, empodial claw; ge, genu; pas, parambulacral seta; pcl, paired claw; pec, pecten; ta, tarsus; te, telofemur; ti, tibia; tr, trochanter; vs, ventral setae; vll, ventrolateral lamella; @, solenidion. Scale bars: A-D = 50m, E = 25m. conspicuous cuticular protuberance. Tibia IV with bipectinate seta. Bipectinate seta on tibia I as finely pectinate as those on other tibiae and not distinctly heavier than these. Tarsi II] and IV each with 4 dorsal setae. Paired claws on tarsi II-IV with relatively fine pecten extending along the inside of the entire shaft (Fig. 2E, compare with Fig. 12E). Empodial claws on tarsi [I-IV not clearly seen. REMARKS. Other species of the ornatus group that have 4 setae on each of tarsi II] and IV are barrierensis, emblematus, orarius, ornatus and prideauxae, all of which are described below. Copidognathus barrierensis sp. nov. (Figs 3,4) ETYMOLOGY. From the Great Barrier Reef. MATERIAL. HOLOTYPE: QMS105731, 3, GBR, No Name Reef, ca. 14°39’S 145°40’E, 9 Oct. 1998, medium coarse sand at 6m. PARATYPES: GBR: QMS105732 (1 3), data as for holotype; QMS105692 (1 2), QMS105695 (1 &), QMS105693/ 105694 (2 ds), Loadstone Reef, 18°41.91°S 147°06.49’E, 12 Apr. 1998, sand & rubble at 2m; ANIC (1 2), ZMH (1 &), Loadstone Reef, 18°42.03°S_ 147°06.54’E, 12 Apr 1998, coarse sand & rubble at 12-15m; ZMH (1 3), ANIC (1 ¢&), Loadstone Reef, 18°42.05’S 147°05.98’E, 12 Apr. 1998, coarse sand & rubble at 8m; QMS105696-105699 (4 9s), Faraday Reef, 18°25.93’S 147°21.11’E, 13 Apr. 1998, coarse sand HALACARIDAE OF THE GREAT BARRIER REEF AND CORAL SEA 721 = , By Bay: = rn a th, ~ 4 * cs oa FIG, 3. Copidegnathus barrierensis sp. nov. A, 2. idiosoma, dorsal view: B, ©, idiosoma, ventral view; C, gnathosoma, 2. ventral view; D,E, ocular plate of 2 2s; T, d, genitoanal plate. Scale bars: A-F = 50m. ke rubble at 6-9m; QMS105700 (1 2), QMS105701 (1 ¥), Bramble Reef, 18°25.25°S 146°40.65°F, 10 Apr. 1998, medium coarse sand at 3-6m: QMS105702 (1 <2), OMS105703 (1 2). Bramble Reef, 18°26.36°S 146°42.24E, 9 Apr. 1998, coarse sand at 5m; QM§ 105704 (1 3), Turmer Cay, NE, ca. 21°43°S 152°33°E, 8 Dec. 1998, medium coarse sand at 3m; QMS105705 (1 ¥), Howard Patch, ca. 22°23.5'S 152°37°E, 6 July 1998, D. Fenner, sand at 6m; QMS105706 (L 2), 23°12.22'S 151°S58.49E, 27 Aug. 1999, coarse sand at 60m, I. Zagorskis; QMS105707 (1 2). Myrmidon Reel, 18°16.69°S 147°23,21'E, 14 Apr 1998, coarse sand at 12m; QMS105708 (1 ¢), Elizabeth Reel, 19°20.12'S 149°02.85’°E. 25 Dee. 1997, coarse sand & rubble at 3m; QMS105709 (1 4), Faraday Reef, 18°24.87°S 142°20.79°E, 12 Apr, 1998, on sponge at 10m; QMS105710-105711 (2 Ss). Lavers Cay, 21°I3°S 151°59°E, 20 Apr. 1999, chunks of coral rubble just above low tide mark, sediment depth 0-10em; QMS105712 (1 3), Boulder Reef, ca, 15°24°S. 145°27°E, § Oct. 1998, A. Thompson, coarse sand at 2m; QMS105713 (1 &), Lizard Island, Coconut Beach, ca, 14°40°S 145°28°E, 15 Oct. 1998, medium coarse sand at 0.5m, 722 MEMOIRS OF TIE QUEENSLAND MUSEUM FIG. 4. Capidognathus barrierensis sp. nov., °. A, leg L ventromedial view; 3, claws on tarsus II, ventromedial view; C, leg IL, ventromedial view: D, leg Il, medial view; K, leg ['V, medial view, Scale bars: A, C-D=50,u.m;B =25um- DESCRIPTION. Male and Female, Male idiosoma 313-326j.m (holotype 326,.m), female idiosoma 298-3264.m long. AD with anterior areola more slender than the 2 posterior ones; posterior areolae not extending to glp-| (Fig. 3A). Medial areola (Fig. 3A,D,E) on OC with usually 2-3, maximally 5 rosette pores; between lateral and medial areola faintly reticulate. PD with pair of porous medial costae (2-3 rosette pores wide), and pair of narrow non-porous lateral costae in form ofa narrow ridge; glp-3 and glp-4 associated with medial costa; ds-4 on anterior half of plate; between costae with conspicuous reticulated ornamentation that becomes fainter towards posterior. AE with concave posterior margin (Fig. 3B). PE in anterior halfclose to inner margin with a group or band of shallow pits, this group or band not contiguous with the anterolateral areola. Male with 23-30 perigenital setae. Ventrolateral lamella on telofemora elaborate, with smooth or slightly undulate edges (Fig. 4A,C-E). Tibia IV with bipectinate seta, Bipectinate seta on tibia Las finely pectinate as those of other tibiae and not heavier than these. Tarsi Hl and 1V with 4 dorsal setae, Patred claws of tarsi II-[V finely pectinate along most of shaft (Fig. 4F), Empodial claw on tarsi [I-IV barely visible. REMARKS. Copidognathus barrierensis can be distinguished from all other species in the arnatus group by having the lateral costae on the PD transformed into narrow non-porous ridges. In all other species in the group the lateral costae are furnished with rosette pores. A further HALACARIDAE OF THE GREAT BARRIER REEF AND CORAL SEA 723 oes Xe Q % 0° b ¢ 2 4 23o a £ 6 0° #3. % Q a 1° G 0 o @% 0 bide) cs 90 Sas SN 5 8 ° o yz o. s ro oe S35 Piha 2 Bieory car se FIG. 5. Copidognathus emblematus sp. nov. A, 3, idiosoma, dorsal view; B, d, idiosoma, ventral view; C, 2, genitoanal plate; D,E, ocular plates in 2 ds; F, 9, gnathosoma, ventral view. Scale bars: A-F = 50m. character by which it can be identified is the relatively short medial areola on the OC that has fewer rosette pores than any other species in the ornatus goup. Copidognathus emblematus sp. nov. (Figs 5,6) ETYMOLOGY; Greek, emAb/lematus = ornament, MATERIAL. HOLOTYPE: QMS105714, ¢. GBR, Lizard Island, Coconut Beach, ca. 14°40°S 145°28"E, 13 Oct. 1998, medium coarse sand at 0.5m. PARATYPES: Great Barrier Reet: QMS105715 (1 ¢), ANIC (1 2), ZMH (1 &), data as for holotype; QMS105716 (1 3), Lizard Island, Coconut Beach, va. 14°40°S 145°28"E, 13 Oct. 1998, medium coarse sand at mid-tide level, sediment depth 10cm; QMS105717 (1 &), Elizabeth Reef, 19°20,12’S 149°02.85°E, 25 Dec. 1997, coarse sand & MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 6. Copidognathus emblematus sp. nov., 3. A, leg 1, ventromedial view; B, leg Il, ventromedial view; C, leg II, medial view; D, leg IV, medial view. Scale bars: A-D = 50u.m. rubble at 3m; QMS105718 (1 3), John Brewer Reef, 18°38.25°S 147°04.42’°E, 11 Apr. 1998, coarse sand at 15m; ANIC (1 @), ZMH (1 2), QMS105719-105721 (3 ds), East Cay, 21°29°S 152°33’E, 18 Apr. 1999, reef flat, coarse sand; QMS105722-105725 (4 gs), QMS105726/105727 (2 2s), Boulder Reef, ca. 15°24’S 145°27°E, 8 Oct. 1998, A. Thompson, coarse sand at 2m; QMS105728 (1 2), Reef 22-101, 21°02.5’S 151°30°E, 16 Apr. 1999, reef flat at 1m; QMS105729 (1 d), Turner Cay, 21°43’S 152°33’E, reef flat, 17 Apr. 1999, coarse sand at 2m. DESCRIPTION. Male and Female. Male idiosoma 326-374m long (holotype 352m), female idiosoma 346-368um long. AD with anterior areola more slender than the 2 posterior ones; posterior areolae not extending to glp-l. OC (Fig. 5A,D,E) with medial areola consisting of at least 15 rosette pores; both areolae in some specimens barely separated (Fig. 5D); between areolae with faint reticulate ornamentation. PD with pair of medial and pair of lateral costae, medial costae 2-3 rosette pore wide, lateral costae 1-2 rosette pore wide; medial costae either connected anteriorly by a transverse band of rosette pores (variable width between specimens, in some specimens only one rosette pore, in others 2-3 rosette pores wide) or both costae completely separate; between costae with reticulated ornamentation, gradually becoming fainter towards posterior; glp-3 and glp-4 associated with medial costae; ds-4 on anterior half of plate. AE with slightly concave posterior margin. PE in anterior half close to inner margin with a band of pits and underlying canaliculi that is contiguous with the anterolateral rosette pore areola (Fig. 5B); further groups of canaliculi posterior to insertions of leg IV. Male with 26-29 perigenital setae. Ventrolateral lamella on telofemora elaborate, with smooth or slightly undulate edges (Fig. 6A-D). Tibia IV with HALACARIDAE OF THE GREAT BARRIER REEF AND CORAL SEA 725 bipectinate seta. Bipectinate seta on tibia I as finely pectinate as those on other tibiae and not heavier than these. Tarsi LI] and IV with 4 dorsal setae. Paired claws of legs II-IV finely pectinate. Empodial claws on tarsi II-IV barely visible. REMARKS. The combination of 4 dorsal setae on tarsi II] and IV and 4 porous costae on the PD is present also in C. adonis, C. prideauxae and C. ornatus, all of which are described below. Copidognathus emblematus can be distinguished from C. adonis by having groups of canaliculi on the PE posterior to leg IV insertions and by having a broad band of pits and underlying canaliculi arising from anteromedial areola on the PE. A group or band of shallow pits in the anterior half of the PE is also present in C. adonis. However, in the latter species, these are not contiguous with the anterior rosette pore areola on the PE, as is the case in C. emblematus. A further difference between both species is in the ornamentation along the anterior margin of the PD. In C. emblematus a transverse band of rosette pores connecting both medial costae anteriorly was found in some but not all specimens, while in C. adonis it was absent in all specimens. It follows that presence of a transverse band identifies C. emblematus while absence of a transverse band is inconclusive. It is interesting to note that the transverse band was absent only in C. emblematus specimens specimens collected on the most southerly sites, south of 21°S (East Cay, Reef 22-101, Turner Cay). Differences between C. emblematus, C. orarius and C. prideauxae are discussed under the remarks to the latter species. Copidognathus hawaiiensis Bartsch, 1989 (Figs 7,8) Copidognathus hawaiiensis Bartsch, 1989: 141, MATERIAL. Australia, GBR: QMS105742 (1 2), Morris Island, 13°29°S 143°44°E, 18 Aug. 1999, C. Bastidas, K. Fabricius & S, Uthicke, fine- medium coarse sand; QMS 105743 (1 3), Whitsunday Islands, Long Island, ca. 20°23’8 148°52’E, 28 Feb, 1997, sand at 0.5 m. Indonesia, Bali: QMS105744 (1 3), QMS105745 (1 2), Menjangan Island, 9 Mar. 1999, J. Benzie, coarse-fine sand in 0.3-0.5m water depth. DESCRIPTION. Male and Female (listed material only). Idiosoma in Australian and Indonesian males 352m and 314m, respectively, in Australian and Indonesian females 314m and 310m, respectively. AD with posterior areolae extending to glp-1; along posterior margin with a band of rosette pores (Fig. 7A). Medial areola on OC with at least 11 rosette pores; remainder of plate with scattered shallow pits. PD with 2-4 rosette pore wide medial and lateral costae, all conntected anteriorly; no reticulation seen between costae. Pores glp-3 in lateral costa, glp-4 in medial costa; ds-4 in anterior half of plate. AE. with straight or concave posterior margin (Fig. 7B); PE with rosette pores posterior to leg IV insertions. Male with ca. 20-24 perigenital setae. Telofemur I with ventrolateral lamella transformed into 2 conspicuous protuberances (Fig. 8A), on one side of a specimen with an additional protuberance (Fig. 8E); a cuticular spine distally on medial flank. Tibia I with one ventral and one smaller distomedial protuberance, each associated with a seta. Telofemur I conspicuously pitted on lateral flank (Fig. 8F). Tibia IV with bipectinate seta. Bipectinate seta on tibia I more robust than those on other segments. Tarsi HI and 1V with 3 dorsal setae. Empodial claw on tarsi I-IV clearly visible. Paired claws of tarsus I smooth, those on tarsi II-[V with moderately coarse pecten over most of the shaft. REMARKS. I have been unable to find any taxo- nomically important differences between the Hawaiian type and the Australian or Indonesian material listed above, and conclude that these are conspecific. Thus, C. hawaiiensis is the third halacarid species that is known to occur in Hawaii as well as the Great Barrier Reef. The other 2 are Acarochelopodia biunguis Bartsch (Otto, 2000b) and Scaptognathus kunzi Bartsch (Otto, 2000a). The common occurrence of these species on both the Hawaiian archipelago and the Great Barrier Reef is noteworthy, as Halacaridae in general have poor dispersal ability and areas that are separated by large and deep bodies of water therefore usually have different halacarid faunas. C. hawatiensis is similar to C. acanthoscelus Bartsch, 1992, in having the ventrolateral lamella on telofemur | transformed into spines and in having a conspicuous ventral protuberance on tibia I that is associated with a seta. C. acanthoscelus differs from C. hawatiensis in that the spines on telofemur I are more numerous and in the glp-3 being associated with the medial instead of the lateral costae on the PD. Copidognathus orarius sp. nov. (Figs 9,10) ETYMOLOGY. Latin, orarius = of the coast. MATERIAL, HOLOTYPE: QMS105735, 3, Great Barrier Reef. Taylors Beach, near Lucinda, ca.18°37’S 726 ) aay Seen oo fal Fd enh, OS ch fay ie Ree i ee a7 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 7. Capidognathus hawaiiensis Bartsch; A, @, idiosoma, dorsolateral view; B, . idiosoma, ventrolateral view; C, 2, gnathosoma, ventral view; D. d. genitoanal plate. Scale bars: A-D = 50m, 146°20°B, 14 Dec 1997, medium coarse sand just above low tide mark, sediment depth 0-Sem. PARATYPES: ANIC (1 ¥), ZMH (1 9), QMS105733/105734 (2 &s), ANIC (1 3),ZMH(1 2),QMS105736-05740(5 3s), data as for holotype. OTHER MATERIAL: QMS105741 (1 3), Indonesia, Bali, Menjangan Island, 9 Mar., 1999, J. Benzie, coarse-fine sand in 0,3-0.5m water depth. DESCRIPTION. Male and Female. Idiosoma in Australian and Indonesian males 288-307um (holotype 307m) long and 278um long, respectively, in Australian females 282-304m long. AD with distinetly blunt nose (Fig. 9A); posterior areolae extending to glp-l: along posterior margin with band of rosette pores; culicle between areolae with scattered pits. OC with medial areola consisting ofat least 15 rosette pores and extending to lateral margin of plate; remainder of plate with scattered pits. PD with medial and lateral costae, all connected anteriorly; lateral areola on average 2 rosette pores wide; medial areola increasing in width HALACARIDAE OF THE GREAT BARRIER REEF AND CORAL SEA FIG. 8. Copidognathus hawaiiensis Bartsch, 2; A, leg I, ventromedial view; B, leg II, medial view; C, leg II, ventromedial view; D, leg IV ventromedial view; E, telofemur I, medial view; F, telofemur I, lateral view. Scale bars: A-F = 50pm. towards posterior, at level of glp-3 ca. 4-5 rosette pores wide; reticulation between costae barely visible; glp-3 and glp-4 associated with medial costae; ds-4 in posterior half of plate. AE with concave posterior margin (Fig. 9B). PE in anterior half with broad band of pits and underlying canaliculi extending from about the inner-most seta to the anterior-most seta or the anterolateral rosette pore areola; posterior to leg IV insertions with canaliculi. Male with ca. 24-27 perigenital setae. Female GA as in Fig. 9D. Gnathosoma as depicted in Fig. 9C. Ventrolateral lamellae on telofemora poorly developed, with smooth edges (Fig. 10A-C,E). Tibia IV with one bipectinate seta. Bipectinate seta on tibia | distinctly more robust than those on the other tibiae, and with coarser pectination; ventral seta on tibia I distinctly thickened and spine-like. Empodial claw on all tarsi clearly visible. Paired claws of tarsus I smooth, those on tarsi H-I'V with a short cluster of about 5-7 teeth that increase in size towards the distal end of the claw; cluster of teeth discontinuous with accessory process (Fig. 10D); paired claws I-IV conspicuously slender. REMARKS. Among the species of the ornatus group C. orarius resembles most closely C. umbonatus. Both species can be distinguished by the morphology of the tarsal claws. Copi- dognathus orarius has maximally 7 tines on each claw II-IV, and this short row of tines does not MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 9. Copidognathus orarius sp. nov.; A, °, idiosoma, dorsal view; B, 3, idiosoma, ventral view; C, 2, gnathosoma, ventral view; D, °, genitoanal plate. Scale bars: A-D = 50m. connect to the inconspicuous accessory process. By contrast, in C. wumbonatus each claw I-IV has at least 12 tines and this row of tines extends along the entire apical half of the claw and merges into the accessory process. Copidognathus ornatus Bartsch, 1981 Copidognathus ornatus Bartsch, 1981: 58. MATERIAL: QMS105730 (1 ¢), Great Barrier Reef, Whitsunday Islands, Long Island, ca. 20°23’S 148°52’E, 28 Feb. 1997, sand & coral rubble at 0.5m The 352\.m long specimen agrees in all aspects under examination with the holotype of C. ornatus from the Mocgambique Channel. C. ornatus can be distinguished from all other species in the ornatus group by having 2 glabrous ventral setae on tibiae IV, instead of one bipectinate and one glabrous seta that are present in all other species. HALACARIDAE OF THE GREAT BARRIER REEF AND CORAL SEA 729 PIG. 10. Copidognathus orarius sp. nov.; A, 2. leg 1, ventromedial view; B, ©, leg 11, ventromedal view: C, °°, leg UL. medial view: D. 3, detail of claws on tarsus 1V, ventromedial view: FE, 2, leg IV medial view. Abbreviation; ac, accessory process. Scale bars; A-C\E = 50um;D = 25pm. Copidognathus prideauxae sp. nov. (Figs 11,12) ETYMOLOGY. For my friend, the late Anne Teresa Prideaux Payne. MATERIAL. HOLOTYPE: QMS105746, ¢, Great Barrier Reef, No Name Reef, ca. 14°39"S 145°40°E, 9 Oct. 1998, medium coarse sand at 6m. PARATYPE: OMS105747 (1), data as for holotype. DESCRIPTION, Male. Idiosoma 320-3524.m (holotype 320m) long. AD with posterior areolae not extending to glp-1 (Fig. 11A); along posterior margin with band of rosette pores. OC with medial areola consisting ofat least | 3 rosette pores which in holotype and paratype show a gap posteriorly. PD with both medial costae appearing clearly separated when focussing onto the canaliculi in deeper cutucular layers but appearing connected through a band of shallow pits on the surface of the plate; medial costa about 2-3, lateral costa |-2 rosette pores wide; between costae with reticulated ornamentation becoming fainter towards posterior; glp-3 and glp-4 associated with the medial costa; ds-4 about half way along plate. AE with concave posterior margin (Fig. 118). PE with extensive and conspicuous band of canaliculi extending from near the inner-most seta to the anterior areola; similar canalicul posterior to insertions of leg [V. GA with 27-33 perigenital setae. Gnathosoma (Fig. 11C). Ventrolateral lamella on telofemora with smooth or slightly undulate edge (Fig. 12A-D). Tibia [ without ventrolateral cuticular protrusion, Tibia TV with bipectinate seta. Bipectinate seta on tarsus | as finely pectinate as those on other tibiae. Tarsi IIT and TV with 4 dorsal setae. Paired claws legs II-TV coarsely pectinate (Fig. 12E), Empodial claws on tarsi II-IV barely visible under oil immersion. 730 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 11. Copidognathus prideauxae sp. nov., d; A, idiosoma, dorsal view; B, idiosoma, ventral view; C, gnathosoma, ventral view. Scale bars: A-C = 50m. REMARKS. The only other species of the ornatus group with 4 setae on tarsi IIT and IV and similarly coarse pecten on tarsal claws [I-IV is C. ornatus (my observation on the holotype). C. prideauxae differs from C. ornatus by having a bipectinate seta on tibia IV. KEY TO SPECIES OF THE COPIDOGNATHUS ORNATUS GROUP 1. . Ventrolateral lamella on telofemur I transformed into at least 2 large spines (Fig. 8A,F); tibia | with a ventral protuberance that is associated witha seta (Fig. 8a) . . 2 Ventrolateral lamella on telofemur I either smooth or slightly undulated (Fig. 2A), not transformed into 2 conspicuous spines; tibia | without a ventral pro- tuberance that is associated witha seta Ventrolateral lamella on telofemur I transformed into 2 large spines, or 2 large and one much smaller spine; glp-3 in lateral areolae, more lateral than ds-5 SA Te Cie ee C. hawaiiensis Bartsch Nm Ventrolateral lamella on telofemur | transformed into more than 3 spines; glp-3 on outer edge of medial areola and directly anterior to ds-5 Tarsi [Land IV each with 4 dorsal setae 4. Paired claws on tarsi [I-IV each with a short row of 5-7 teeth that is discontinuous with the accessory process a Ste Se Ree AMES me. 8 le, C. orarius sp. nov. Paired claws on tarsi II-IV with a row of at least 12 teeth that is continuous with the accessory process NAS ued gt eye, Me C. umbonatus Bartsch an PD without lateral costae bearing rosette pores; medial areola on OC consisting of 2-5 rosette pores safer: ee beh east Ness C. barrierensis sp. nov. PD with lateral costae bearing rosette pores; medial areola on OC consisting of at least eight rosette pores . 6 6. TibialV with abipectinate seta(Fig.2D),....... 7 Tibia IV without a bipectinate seta . . C. ornatus Bartsch 7. Paired claws on tarsi II-IV with teeth that increase in size towards the distal end of the claw. C. prideauxae sp. nov. Paired claws on tarsi II-IV finely pectinate throughout, pectines not increasing in size towards distal end of claw 8. PE with a broad band of canaliculi arising from the anterolateral areola on the PE and reaching to about the innermost seta; posterior to leg |V insertion with several groups ofcanaliculi. .. 2... C. emblematus sp. nov. PE with canaliculi near the innermost seta but not in form of a band that reaches the anterolateral areola; posterior tolegIV withoutcanaliculi. .. 2. . C. adonis sp. nov. ACKNOWLEDGEMENTS I thank the Australian Biological Resources Study for funding, the Australian Institute of Marine Science for providing infrastructure support, the Great Barrier Reef Marine Park Authority for permission to collect mites in the Marine Park, and Mark Judson, Ilse Bartsch and Gordon Nishida for loan of specimens. I also acknowledge Carolina Bastidas, John Benzie, Guillermo Diaz-Pulido, Katharina Fabricius, Doug Fenner, Angus Thompson, Sven Uthicke HALACARIDAE OF THE GREAT BARRIER REEF AND CORAL SEA 731 FIG. 12. Copidognathus prideauxae sp. noy., 3; A, leg 1, medial view: B, leg I, medial view; C, leg LI. medial view; D, leg |V, medial view; E, detail of claws on tarsus IT, ventral view. Scale bars: A-D =50um; E = 25j.m. and Irena Zagaroskis for collecting some of the material. LITERATURE CITED BARTSCIL, I. 1981. Halacaridae (Acari) aus dem Kanal von Mocambique. Cahiers de Biologie Marine 22: 35-63, 1989. New species of Copidognathus (Acari: Halacaridae) from Hawaiian Islands. Bishop Museum Occasional Papers 29; 138-148. 1992. Halacaridae (Acari) from Hong Kong. Three new species of Copidognathus, Entomologische Mitteilungen aus dem zoologischen Museum Hamburg 10: 229-24], 1997, Copidognathinae (Halacaridae, Acari) from northern Australia; description of four new species. Pp. 231-243. In Hanley, J.R., Caswell, G., Megirian, D. & Larson, HK. (eds) Proceedings of the Sixth International Marine Biological Workshop. The marine flora and fauna of Darwin Harbour, Northern Territory, Australia. (Museums and Art Galleries of the Northern Territory & Australian Marine Sciences Association; Darwin). NEWELL, 1.M. 1947. A systematic and ecological study of the Halacaridae of eastern North America, Bulletin of the Bingham Oceanographic Collection 10: 1-232, 1984, Antarctic Halacaroidea. Antarctic Research Series 40: 1-284. OTTO, J.C. 2000a. Halacaridae from the Great Barrier Reef and Coral Sea: the genera Lohmannella, Scaptognathides and Seaptognathus (Acarina: Halacaridae: Lohmannellinae). Memoirs of the Queensland Museum 45: 535-555. 2000b. Acarechelopodia and Actacarus species (Acari: Halacaridae) from Australia, with remarks on 4. pacificus and A. orthoiectus, Species Diversity 5: 111-127. ANEW SPECIES OF TORRENT-DWELLING FROG (ANURA: HYLIDAE: LITORIA) FROM THE MOUNTAINS OF INDONESIAN NEW GUINEA (WEST PAPUA) STEPHEN J. RICHARDS Richards, S.J. 2001 06 30: A new species of torrent-dwelling frog (Anura: Hylidae: Litoria) from the mountains of Indonesian New Guinea (West Papua). Memoirs of the Queensland Museum 46(2): 733-739. Brisbane. ISSN 0079-8835. Litoria macki sp. nov. from the mountains of West Papua, Indonesia, is a torrent-dwelling species characterised by medium size (adult 6 d 42.1-45.7mm SVL), unwebbed fingers, and prominent conical tubercles on the dorsum and limbs. The new species most closely resembles L. spinifera (Tyler), from which it can be distinguished by its Jarger size and different advertisement call. The calls of Z. macki and L. spinifera are described and compared, and natural history observations on L. spinifera are presented. 0 Torrent-dwelling frog, West Papua, new species. Stephen Richards, School of Tropical Biology, James Cook University, Townsville 4811 (present address, Vertebrate Department, South Australian Museum, North Terrace, Adelaide 5000); 11 August 2000. Torrent-dwelling hylid frogs within Liforia are a diverse assemblage reaching their greatest div- ersity in the mountains of New Guinea. All species for which life history features are doc- umented lay unpigmented, macrolecithal eggs and have tadpoles with dorsoventrally flattened bodies and large, ventral suctorial mouthparts (Haas & Richards, 1998; Tyler & Davies, 1978). The L. becki species-group (Tyler & Davies, 1978) contains 5 predominantly montane torrent- dwelling frogs (L. becki, L. micromembrana, L. modica, L. pratti and L, spinifera) from mount- ains of New Guinea. These species are most easily distinguished from other torrent-dwelling Litoria by their medium size (6 ¢ and 2 to about 42mm and 53mm, respectively) and long, unwebbed fingers (Tyler & Davies, 1978). Litoria spinifera (Tyler, 1968) is distinctive within the L. becki group with a sharply pointed snout and greatly enlarged tubercles on the dorsum, eyelids, tarsus and foot (Tyler, 1968). It was described from a large series of frogs collected by Mr Fred Parker at altitudes of 1000 - 1500m in dense rainforest S of Kundiawa in the central mountains of PNG (Tyler, 1968). The type series was collected ‘in the vicinity of small waterways’, where they were found among leaves near the streams during the day and on leaves overhanging the water at night (Tyler, 1968). Nothing else is known about its biology. During 1997 and 1998, I recorded its advertisement call and made observations on its general natural history in the Crater Mountain Wildlife Management Area (CMWMA), in Eastern Highlands Province, PNG. Conservation International’s 1998 biodiversity survey of the rugged Wapoga River headwaters region of West Papua (Mack & Alonso, 2000) accumulated a significant collection of torrent- dwelling hylid frogs (Richards, S.J., Iskandar, D. & Allison, A. in Mack & Alonso, 2000:54-57). Among these is an undescribed Litoria that resembles L. spinifera in its possession of large conical tubercles on the dorsum and limbs, but differs from that species in its larger size and different advertisement call. In this paper I describe the new species from West Papua, and analyse and compare its vocalisations with those of L. spinifera. I also present brief observations on the natural history of L. spinifera in the CMWMA. MATERIALS AND METHODS Specimens are deposited in the Museum Zoologie Bogor (MZB), Indonesia, the Queensland Museum (QM), Australia, and the South Australian Museum (SAMA), Australia. Measurements (to the nearest 0.1mm) were taken with dial callipers and a stereomicroscope fitted with an ocular micrometer, and follow Menzies (1993). They are: SVL (snout-vent length), TL (tibia length), HW (head width at tympanum), HL (head length from tip of snout to posterior edge of tympanum), EYE (horizontal eye diameter), TYM (horizontal tympanum diameter), IN (inter-narial distance), EN (distance between anterior edge of eye and posterior edge of naris), 3FD (width of 3rd finger disc at right angle to digital axis) and 3FP (width of penultimate V4 cabal VON as Pa a 4 oe Neyer MEMOIRS OF THE QUEENSLAND MUSEUM FIG, |, Literia macki sp. noy, (QMJ75810). A, lateral view of head: B, palmar view of hand; C, plantar view of fool. Scale bars = 2mm. phalanx of 3rd finger), 47D and 4TP (4th toe dise and 4th toe phalanx, as for 3rd finger). Calls were recorded with a Sony Professional Walkman tape recorder and Sony SMZ-200 microphone, and wete analysed with the sound analysis program Avisofi SAS-Lab Pro. Note structure was highly variable, both within and between call sequences and individual males. Variation involved the degree of pulse definition, and the presence or absence of minor pulses in terminal portions of the note. In an attempt to attain consistency only well defined pulses and pulses that contributed substantial energy to the note, were calculated for comparisons among species. SYSTEMATICS Litoria macki sp. nov. (Figs 1-6) ETYMOLOGY. Named for Dr Andy Mack in ap- preciation of his unfailing encouragement and support and in recognition of his passion for conservation and science education in New Guinea. MATERIAL, HOLOTYPE: MZB Amp.3870, adult Y, collected by S.J, Richards and M, Moore, Wapoga Alpha mineral exploration camp, West Papua, Indonesia, 3°08.687°S, 136°34.423°F, 1070m,, [7-iv,98. PARATYPES: MZB Amp.3871. 3872, QM J75810. SAMA R55363 same collection data as holotype, SAMA R55364 collected by M. Moore at Lagori Landing Site 21 (LS-21). West Papua, Indonesia, 3°00,348"S, 136"33.412°E, 275in, 26, iy.98, DIAGNOSIS. A medium-sized Litoria, 32 42.1-45.7mm SVL, distinguished from all known New Guinea species except L. spinifera by the combination of: 1, blotched green and brown dorsally in life; 2, canthus rostralis well-defined, moderately curved; 3, fingers long, unwebbed; 4, large conical tubercles on tarsus and foot, and on dorsum (concentrated on head and eyelids). Distinguished from L. spinifera by its larger size (SVL of male L. spinifera 35.3-42.3) and different advertisement call, which consists of a rapidly-repeated series of loud, bell-like notes in which note repetition rate, note length, and pulses/note increase during the call sequence, DESCRIPTION OF HOLOTYPE. An adult ¢ measuring: SVL 45.7; TL 29.0; WW 15.6; HL 16; EYE §.5; EN 4.0; IN 5.1: TYM 2.0; 3rd finger 1.2; 3rd dise 2.9; 4th toe 1.2; 4th dise 2.5, Body slender, limbs long (TL/SVL 0.634). Head slightly longer than broad (HL/HW 1.04), more than one third of snout-vent length (HL/SVL 0.35); snout slightly pointed in dorsal view, pointed in lateral view, projecting slightly beyond lower jaw. Canthus rostralis well- defined, moderately curved, loreal region strongly concave. Nostrils close to tip of snout, internarial distance greater than distance between snout and naris (EN/IN 0.784). Eyes large, prominent (EYE/SVL 0.12), pupil horizontal, eyelid without reticulations. Vomerine teeth in two prominent oblique ridges between the choanae, yocal slits present. Tongue cordiform, Tympanum clearly visible, but dorsal edge obscured by prominent, slightly curved supratympanic fold. Fingers long, unwebbed, TORRENT-DWELLING FROG FROM WES FIG. 2. A, adult Litoria macki sp. nov. trom Wapoga River headwaters, West Papua. B, adult_L. spinifera from Maimafu, Eastern Highlands Province, PNG. relative lengths 3>4>2>1, terminal discs large (3FP/3FD 0.41); A brown nuptial rugosity on first finger extends distally penultimate tubercle. Toes 2/3 webbed, web reaching to just below penultimate tubercle on 4th toe, and nearly to disc on toes 2, 3 and 5, relative lengths 4>5=3>2>1, terminal discs large (4TP/4TD 0.48). Conical tubercles on limbs (including heel) and dorsum, concentrated on head and eyelids, and on outer edge of tarsus and foot. In life, mottled with large patches of brown and green dorsally, including dorsal surfaces of legs. Yellow in groin and axilla. L aterally pale y ellow grading to white anteriorly and ventrally. ‘Venter white, mottled with faint brown pigmentation Iris gold with purple reticulations. Tubercles beneath vent white. In preservative brown dorsally, with extensive blue (green in life) patches. Pale yellow lateral and mguinal regions have faded to white. to level of T PAPUA 73 ih VARIATION, The paratypes are adult males (SVL42.1-45.3) and the colour pattern is uniform in the type series: in life all were blotched with brown and green dorsally. The size and distribution of tubercles on the dorsum and legs are variable but in all specimens tubercles on the eyelids, tarsus and heel are prominent. Measurements are summarised and compared with C, spinifera in Table 1, ADVERTISEMENT CALL. The advertisement call is a series of 14-44 bell-like notes lasting about 7-50 seconds. Dominant frequency is 2606-3144Hz. A consistent and distinctive feature of the call is an increase in note repetition rate and pulses/note in terminal sequences (Fig. 5; Table 2). Individual notes are about 0,02s in length at the beginning of a call sequence and contain a single pulse. During the call sequence note length increases to 0.04-0.06s and the number of pulses/note increases to 6-8. A call sequence is illustrated in Fig. 5 and structural features are summarised in Table 2 FIG 3. Torrential stream habitat of L. macki in lower montane rainforest, West Papua. 736 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 1. Morphological comparison of 3d types of Litoria macki sp. nov. and 6 L. spinifera (Tyler). Measurements involving finger and toe discs only recorded where discs are well-preserved. from small trees adjacent to and hanging over small (<5m wide) torrential streams in closed- canopy rainforest (Fig. 3). They were never observed in swampy or slow-flowing aquatic habitats. Perch sites were high above or adjacent to the water, normally exceeding 2m high, but two frogs were observed on perches about 1m above the water. Several species of suctorial tadpoles were observed in streams occupied by this species, but it is not possible to associate any of these with the new species. DISTRIBUTION. Known only from 2 localities in the mountainous headwaters region of the Wapoga River in NW West Papua, Indonesian New Guinea (Fig. 4). COMPARISON WITH OTHER SPECIES AND DISCUSSION. Lack of webbing between the fingers and presence of prominent conical tubercles on the dorsum and limbs will distinguish L. macki from all New Guinea Litoria except L. spinifera. | have examined 7 paratypes (SAMA R6295-6301) and 5 additional specimens (SAMA R9167, SAMA R9108A-D) of Litoria spinifera collected in the vicinity of the type locality, and compared them with 8 specimens (SAMA R55357-62; UPNG 9963-4) from the Crater Mountain Wildlife Management Area (55-75km SE of Kundiawa). The frogs from CMWMA have prominent tubercles along the edge of the tarsus and foot (Fig. 2) and agree in all other respects except minimum adult size with the type series (Table 1), The minimum SVL reported for the type series is 38.4mm (Tyler 1968) but the minimum for frogs from CMWMA is 36.7mm. However re-measurement of the seven L. spinifera paratypes shows that six Litoria mackt (n= 6) Litoria spinifera oe CMWMA (see | Litoria sie rhea ra vicinity i = Mean (SD) Range Mean (SD) Range | Mean (SD) Range [SVL | 43,93 (1.36) _ 42.1-45.7 38.98 (1.42) 374-414 37.26 (1.29) 353-394 TL/SVL | 0,629 (0,016) 0.604-0.650 0.632 (0.017) (0.609-0.658 0.657 (0.020) | —_(0.630-0.691 | BYE/SVL 0.125 (0.009) 0.114-0.142 0.130 (0.003). 0,126-0.135 0.116 (0.009) 0,107-0.137 ENN | __0.664 (0.075) 0.592-0.784 0.630 (0.032) 0.592-0.680 0.698 (0.052) | _0.615-0.829 HW/SVL | __0.340(0.005) | —_ 0.335-0.349 0.348 (0.005) 0,341-0.355 | _0.352(0.010) | __(0.337-0.371 3EPBFD | __0.402 (0.029) 0.370-0.448 0.456 (0.036) 0.416-0.500 0.449 (0.022) —_-0.409-0.476 | |ATP/4TD __|_0.503 (0.015) | __0.480-0.520 0.567 (0.058) | _0.523-0.666 0.512 (0.032) | 0.473-0.588 | 3FD/SVL —_|_0.064 (0.004) 0.061-0.073 | _-0.057 (0.002) 0.053-0.059 0.058 (0.002) 0.053-0.061 _4TDISVL 0.055 (0.005) 0.050-0.064 0.051 (0.002) 0,048-0.054 0,050 (0.003). 0.047-0.059__| NATURAL HISTORY. Males called at night specimens have an SVLless than 38.4mm and the minimum SVL of this sample is 35.5mm. The smallest adult male L. spinifera that I have examined has an SVL of 35.3mm and the mean SVL of frogs from the vicinity of the type locality and from CMWMA are 37.26 and 38.98mm respectively (Table 1). There is therefore complete overlap between the type series and specimens from the CMWMaA in all characters examined and | have no hesitation in referring the CMWMA specimens to this species. In contrast there is no overlap in size between the type series of L. macki and the eight L. spinifera from the CMWMA (SVL 42.1-45.3 vs 36.7-41.4 in spinifera; Table 1), and only marginal overlap between L. macki and the large type series of L. spinifera (29 males from Oruge and vicinity measuring 35.5-42.3mm; upper size limit based on Tyler, 1968) (Table 1). Size of finger discs is another character that is useful for distinguishing between these two species. The disc of the third finger is larger in proportion to SVL in L. macki than in L. spinifera (3FD/SVL 0.061-0.073 vs 0.053-0.061; Table 1). Only one of the L. spinifera examined had a 3FD/SVL ratio that was as large as the smallest ratio recorded for L. macki. Difference in size alone would be insufficient to warrant recognition of the Wapoga specimens and L. spinifera as distinct taxa, given that the populations are allopatric and separated by nearly 1000km (Fig. 4). However L. macki can be further distinguished from L. spinifera by its advertisement call, which is a short sequence of 14-44 explosive notes (see above). In contrast the calls of L. spinifera are distinctly pulsed musical notes uttered singly at regular or irregular intervals of 1.5-8 seconds (Fig. 5). The call of L. TORRENT-DWELLING FROG FROM WEST PAPUA 737 TABLE 2. Advertisement call characteristics of Litoria macki sp. nov. and L. spinifera (Tyler). | structural features of notes based on two call series only, * only a short portion of a much longer sequence was recorded, excluding territorial calls. Data are presented as mean (SD) range. __Litoriamacki _ 5 Litoria spinifera es ica QM J75810 SAMA R 55363 Herowana SAMA R55361 | Maimafu SAMA R55357 (cal sequences) _ (n=3)! * |b (n=3) (n= 1y (n= 2) Date _ 17.iv.98 : 17.iv.98 a 3x98 | Xi. 97 Snout-vent length (mm) 42.1 43.7 - 41.4 39.5 Temperature “¢ : 20.8 225 LOH 20 4 |Fullcall sequence | 7 : a! a Total notes in sequence 18-44 14-19 6 25-49 | Call duration (s) 10.3-48.7 ; ; 7.2-15.4 22.75 __97,2-125.3 Note repetition rate (nates/s 0.9-1.66 1.22-1.82 : J 0.22 Ci 0.25-0.39 Note duration (s)° 0.04 (0.01) 0.015-0.084 | 0.03 (0.01) 0.015-0.066) noe ass 0.03 (0.01) 0.013-0.095 Pulses per note* 32 Q. 4) 1 9 3.0 Q. 0) 1-8 3.5 (1.22) 3-6 | 4.04 (15)2-9 Single-pulse notes? Yes Yes No No Dominant frequency (Hz) | 2606-2842 2961-3144 2885-3015 1335-2821 | Introductory notes a Note repetition rate (notes/s 0.78-1.16 0.84-0.92 - | ‘Terminal notes 7 Note repetition rate | (notes/s) ae 1.9-2.39 2.56-2.57 - . Territorial calls ee 7 Type | om =6) Length (s) “ - | : - ___|_ 0.358 (0.09) 0.21-0.44 _Number ee = * - | - - _ | 34.83 (9.5) 19-44 Ty Pin ion oF sits per 5-6 sequence 7 7 a 7 7 z Note repetition rate |_(notes/: js). =a - . : ; & a jal 32) 2 2 2021 i 200 0 200 400 600 800 1000 1200 Kilometers SS ee FIG. 4. Distribution of L. macki (@) and L. spinifera (@: Crater Mountain sites only) specimens examined during this study. 738 > 9 MEMOIRS OF THE QUEENSLAND MUSEUM 34 = 3 | | : 2 z= 4 ne ) T | | T T 1 2 3 4 5 6 7 B c d a | | adie ] ] bike fH ot 4 de g 2 = 1 0 4 2 3 4 5 6 + Time (s) Amplitude 80 ms Time FIG. 5. Advertisement calls of A, Litoria spinifera (SAMA R55361) from Herowana, PNG, two consecutive notes from a long series at an air temperature of 20.6 °C; B, Litoria macki (SAMA R55363) at an air temperature of 22.5°C, audiospectrogram of full call sequence showing increasing note repetition rate during call sequence; C, wave forms of notes indicated, showing distinctly pulsed calls of L. spinifera and change in pulse number during call sequence of L. macki. spinifera also differs from that of L. macki in the following features: in L. spinifera all notes are distinctly pulsed (vs single-pulse introductory notes in L. macki; Table 2), note repetition rate is much slower than that of L. macki at similar temperatures, and note repetition rate and pulses/note do not increase dramatically during a sequence of calls. The call of L. spinifera is described in more detail below and structural features are presented and compared with L. macki in Table 2. DISTRIBUTION AND NATURAL HISTORY OF LITORIA SPINIFERA Litoria spinifera was previously known from several localities SW of Gumine in Chimbu Province. Although precise coordinates are not available, all of these sites are clustered in an area extending from approximately 6°12’S, 144°57’E in the north to approximately 6°43’S, 144°43’E in the south. L. spinifera was a common species in the Crater Mountain Wildlife Management Area (Herowana 6°39714.5"S, 145°11’49.8"E; Maimafu 6°30°06.0"S, 145°01°59.1"E) during TORRENT-DWELLING FROG FROM WEST PAPUA kHz 5 4 i—| = WTTTITTT IIIT 1a) 1s sesasadadee demesne g ° ; ty 0.05 0.1 Time FIG. 6. ‘Territorial’ call of Litoria spinifera from Maimafu, given during vocal interaction with another calling frog. November-December 1997 and 1998, and in January 2000 adults were heard calling, and several immature specimens were observed, at the Wara Sera Research Station (6°43.423’S, 145°05.693’E) (Fig. 4). The CMWMaA records extend the known distribution of this species about 40km to the east. At Herowana and Maimafu males called from leaves one to 3 metres above the ground, above or adjacent to small torrential streams. At Herowana males were common along streams running through disturbed rainforest and old gardens, suggesting that this species is tolerant of significant human disturbance. Some of these streams had beds that were heavily silted due to erosion from the surrounding denuded hill slopes. ADVERTISEMENT CALL OF LITORIA SPINIFERA. The advertisement call of L. spinifera is a single distinctly pulsed note. Each note consists of 2-9 pulses (Fig. 6; Table 2), and notes are repeated at intervals of about 1.5-8 seconds for very long periods. Two complete call sequences recorded at Maimafu lasted for 97.2 seconds and 125.3 seconds (Table 2). Two other call-types, tentatively identified as territorial calls, were heard but were much less common and only produced by one male in response to calls of anearby conspecific. Territorial call type 1 consisted of a single drawn out note with subs- tantially more pulses than ‘normal’ calls (Table 2). Territorial call type 2 involved the production of 5-6 notes in rapid succession, with a repetition rate substantially higher than that of the whole call sequence (Table 2). Both of these territorial call types were uttered intermittently during long sequences of ‘normal’ (advertisement) calls. All calls are finely tuned, with energy concentrated in narrow bands giving them a musical quality. In calls from Herowana, energy 015 02 0.25 03 035 04 045 § 739 is concentrated in a single band, but in calls from Maimafu energy is con- centrated in 2 distinct, harmonically related bands. In some notes the lower frequ ency is dominant; in others the higher frequency dominates. ACKNOWLEDGEMENTS I am extremely grateful to Dr Arie Budiman of Puslitbang Zoology, and the head of Museum Zoologicum Bogoriense - LIPI, Ibu Liliek Prijono, for their assistance in Jakarta and Bogor. Dr Djoko Iskandar assisted in many ways in the field, and I am most grateful for his input to this project. This research was part of Conservation International’s Rapid Assessment Program, and | greatly appreciate their assistance. Funding support was provided by the CI-USAID Cooperative Agreement # PCE-5554- A-00-4028-00. I am especially grateful to Andy Mack, Yatna Supriatna, and Burke Burnett of CI for support during the survey, and to Mike Moore who helped collect. P.T. Freeport Indonesia provided logistical support for which I am most grateful. Mike Tyler commented on the manuscript and Bronwen Scott produced Fig. 1. Andy Mack helped produce the map. LITERATURE CITED HAAS, A. & RICHARDS, S.J. 1998. The correlation of cranial morphology, ecology and evolution in suctorial tadpoles of the Australian genera Litoria and Nyctimystes (Amphibia: Anura: Hylidae: Pelodryadinae) Journal of Morphology 238: 109-141, MACK, A. & ALONSO, L. (eds) 2000. A biological assessment of the Wapoga river area of northwestern Irian Jaya, Indonesia. RAP Bulletin of BiologicalAssessment 14. (Conservation International: Washington). MENZIES, J.I. 1993. Systematics of Litoria iris (Anura: Hylidae) and its allies in New Guinea and a note on sexual dimorphism in the group. Australian Journal of Zoology 41: 225-255. TYLER, M.J. 1968. Papuan hylid frogs of the genus Ayla. Zoologische Verhandelingen 96: 1-203. TYLER, M.J. & DAVIES, M. 1978. Species-groups within the Australopapuan hylid frog genus Litoria Tschudi. Australian Journal of Zoology (supplement 63): 1-47. THE HELICEINID LAND SNAIL PLEUROPOMA ENTINCTA (ODHNER, 1917) AS AN ENVIRONMENTAL INDICATOR IN ARCHAEOLOGY CASSANDRA ROWE, JOHN STANISIC, BRUNO DAVID AND HARRY LOURANDOS Rowe, C., Stanisic. J., David, B. & Lourandos, 11, 2001 06 30: The helicinid land snail Pleuropoma extineta (Odhinet, 1917) as an environmental indicator in archaeology. Menrairs af the Queensland Museum 46(2); 741-770, Brisbane. ISSN 0079-8835, Land snails have lone been used as palacoenvironmental indicators in some parts of the world, butin Australia they have received Jittle attention, Here we present new data on the geographical distribution and ecological characteristics ofthe helicinid P/eurapoma extineta (Odhner, 1917). The species is confined to ihe limestone oulcrops of the Chillagoe Formation where it oecurs abundanily both as living populations on the limestone rock and associated yegetation, and as dead shells in the litter deposits. After analysing 1,100 modern snail shells from 24 collection localities (from two major limestone regions within the formation), we argue that the sensilivity of shell growth (measured as size-correlated differences in whorl counts) to environmental moisture variation in P extineta makes this species particularly suitable for palaeoenviranmental research, Differences in whorl counts af sub-fossil 2 exfincra shells from archaeological excavations ait Hay Cave, in tropical northeastern Australia, are then investigated. We conclude with observations about the implications of the land snail shell sequence for the nature and timing of changes in rainfall levels during the last 19,000 years in north Queensland, and how these changes compare and contrast with existing palynological trends for the Atherton Tableland nearby, [7 Lend sreily, Helicinidae, Pleurapoma extincta, palaevenviroyments, yrchdeolagy, narth Queensland. Australia. Cassandra Rawe and Bruno David (e-meil: brune,dayidlaiarts.nonash edict, Department of Geography & Environmental Science, Monash University, Wellington Read. Clayton 3800; John Stanisic, Queensland Museum, PO Box 3300, Santh Brisbane 4101s Harry Laurandas, Department of Anthropology & Saciology, University ef Queensland, St Lucia 4072, Australia; 10 November 2000, In Australia, the use of archaeologically obtained land snails for palacoenvironmental reconstructions is in its infancy. This is despite their frequent presence jn excavated sites. Some of the problems associated with the utility of land snails in archaeological analysis were outlined by Evans (1972) and highlighted by other authors since, especially in Europe (e.g, Carter. 1990: Connah & McMillan, 1964: Sparks, 1969). Difficulties arise in two major areas: the level of knowledge about the land snail species to be used, including not only their taxonomic status but also their behavioural and ecologic traits: and ithe taphonomic status of the excavated samples. These difficulties are equally, if not more, applicable in the Ausiralian sphere. This paper presents new data from north Queensland. iuldressing these issues 50 as. to investigate late Quaternary palacoenvironments alony the western slopes of the Great Dividing Range, an area hitherto poorly researched. QUATERNARY LAND SNAILS; HISTORY OF RESEARCH Study of gastropod molluses from Pleistocene archacological deposits began in the 19th century, and the possibility of using snails as indicators of past climates, local environments and as chronological indicators has been recognised since at least this time (Kennard, 1897: Lowe & Walker, 1984). As in other branches of palaeontology, much of the early work was concerned with taxonomy; relatively little attention was paid to palaeoecological considerations. By the late L9th and early 20th centuries, however. researchers in the United Kingdom began to use molluscs as palaeo- climatic indicators and as a means ol dating geological events (Kennard, 1897; Woodward, 1908). During 1919 to 1939, numerous investigations followed their lead. These studies were usually published as appendices to archaeological reports. One of the earliest studies concermed the Neolithic (4,000-2,400 BC) flint mines of Grimes Graves in Norfolk, where land snails were used not only as indicators of past 742 landscapes, but also as evidence for the post-ulacial antiquity of the mines —at that ome an issue in considerable dispute (Pvans, 1972). Despite such promising pioneering works emanating from the UK (and to some degree from central Rurope, particularly Czechoslovakia), to this day the analysis of land snatls from archacological deposits continues to be largely ignored in many parts of the world, Yet the argument for using land snails in pulaco- ecological reconstructions seems irresistible. Allernations.of cool and warm periods, spanning various lengths of time from decades to thousands of years, ure represented by soil horizons which, when not deealeitied, asually contain molluses of yarious species. Such changing climatic conditions during the Quaternary have been successfully identified by land snail analysis in Czechoslovakia, in much the same way that pollen has been used to define various interglacial periods. Not only can the distinction between interglacial, interstadial and wlactal stages be recognised via land snails, butin many cases the fauna are sufficiently diagnostic to make relatively fine-grained chmatic determinalions for sume periods of time (ef. Fvans, 1972). In a study with particular relevance to the current investigation, Solent (1972) documented land snails from deposils recovered in the Aq Kupruk Caves. Afghanistan. He used the general abundance af taxa in the land snail assemblage as well as size differences in the enid Sufzehrinus eremitus (Benson, 1849), a pulmonale gastropod, to postulate prehistoric changes in moisture patterns. Hlowever, he readily acknowledged that his conclusions were restricted by the absence of knowledge about the ecology and distribution of (he contemporary land snail fauna of the region and a lack of data on environmental vanation within local species, In spite of these short- comings Solem’s (1972) analyses demonstrated the principle that changes in the shell-size of land snails could be used to determine past changes in environmental moisture levels. Briel discussions of research in central Rurope, the Middle Last, Affica and North America can be found in Evans (1969) and Goodtriend (1992), and will not be recounted here, However, in central urope and North America, land snail analysis is of considerable importance in various lines of Quaternary research, This is due in large part to their widespread occurrence in areas of calcareous loess (Evans, 1972). MEMOIRS OF THE QUPPNSLAND MUSPUM LAND SNAIL RESEARCHES THI UK: EMERGENCE OF ATECHNIOLIE In Britain, numerous regional studies have Jooked to and snails lo myestizate the timing and extent of human-induced Jandscape clearances from the Neolithic to the Bronze Age (Wainwright & Davies, 1995). Where land snails were Once used fo look at general charges in land-use patterns (e.g. Evans 197Ja, 1Y7Tb Lor southem England), they are now being used tor much higher resolution mapping of archacological landscapes, typically at seules of ahout 35-80kim’ (eg, Allen 1997, in press). Several factars have contributed to thrs. Land snails are abundant in many types of depositional vontexts (including oxidised sediments) these are usually identifiable to species; and they are visible in the field, thus allowing preliminary assessments of local depositional environments (Porch, in prep.). Moreover, their appeal is also directly related to a fundamental limitation of pollyn research in the UK, where an absenee of peat and liminie deposits trom the large tracts of chalk and limestone in southern, central and eastern England renders pollen analysis inappropriate (Evans, 1972), Pollen analysis until recently has been able to reveal virtually nothing about the environmental and ecological history of the chalklands (but see for example Seaile, 995; Sharples, 1991). This is particularly unfortunate given the importance of such environmental settings to settlement and avricullure during prehistoric und Roman times (Evans, 1972). In contrast, palacoenvironmental research based on land snails has enabled conclusions to be drawn on the nature of people-land relations for various periods in prehistory, including the timing of landscape clearances, the onset of ayricullural practices, and the emergence of intensified settlement systems (Fwans, 1972), The thrustof vans’ own work shows that ecalogical information can be gained trom land snails with which patierns of land-use change can be addressed. Criven these research foci, i is common to find in the archacological literature discussions of land snails and the presentation of data included under cuptions such as “Evidence for the environment and farming ccanomy’, This ia particularly so when sites such as barrows (burial mounds), pits, hillforts, pastureland and ploughzones are considered (Allen, 1995), It has also been recognised for some time, largely from tbe results of pollen analysis. that LAND SNAIL AS AN ENVIRONMENTAL INDICATOR 743 many parts of Britain which are today open country were once forested. The East Anglican Breakland and the North York Moors are classic examples, the former haying been cleared by Neolithic people some 3,000 BC, the latter during the Bronze Age by around 1,550 BC (Evans, 1972). In view of this, it has generally been assumed that the chalk and limestone upland regions were likewise once forested, having suffered a similar history of forest clearance through the actions of prehistoric farmers. By undertaking land snail analysis, landscape clearance dating as far back as the 4th millennium BC has been confirmed. As a result, it is now thought that few, if any, parts of the British Isles can today be thought of as entirely natural (Evans, 1972). The Buckskin Barrow site at Basingstoke, Hampshire, provides a good example of how changing frequencies of molluscan faunas with distinctive ecological affiliation can be used to identify changes in patterns and intensities of land use. An excavated section through a series of silts and mixed deposits suggested the clearance of deciduous woodlands during the Late Neo- lithic to Early Bronze Age. This clearance was associated with pre-barrow features and a cremation pit. Subsequently, short-grazed grasslands became established during the Middle to Late Bronze Age. By the Iron Age, the site had become overgrown by long grasses and shrubs (indicating very little or low pressure land-use) followed by intermittent arable and pastoral activity, resulting in erosion of the barrow monument (Allen et al., 1995). Consequently, molluscan evidence indicated to the excavators that major changes had taken place in systems of land use at Basingstoke since the late Neolithic, and that these changes could be divided into a number of well-defined demographic, settlement and economic systems. LAND SNAIL ANALYSIS IN AUSTRALIAN ARCHAEOLOGY Australian archaeological site reports typically include site descriptions, vertebrate faunal lists and a detailed evaluation of stone artefacts, while rarely making even a passing note of land snails. They are rarely identified even when their presence is noted, and are rarely incorporated in ensuing discussions (David & Stanisic, 1991). To illustrate this point: in SE Australia, almost all non-marine molluscan analyses have been qualitative and peripheral to the main objectives which are generally geomorphological or palynological in emphasis (Porch, in prep.). This is somewhat surprising given the attention often paid to palaeoenvironmental indicators such as pollen from archaeological sites. It is doubly so given the high profile of land snail research in archaeology in the UK, where most of Australia’s first generation of professional archaeologists were trained in the 1960s. BIOGEOGRAPHY OF AUSTRALIAN LAND SNAILS AND THEIR SUITABILITY AS PALAEOENVIRONMENTAL INDICATORS. A key biological feature of terrestrial molluscs that makes them useful in climatic reconstruction is their sensitivity to environmental moisture regimes, They have not evolved totally effective structural means to prevent water loss and desiccation is their greatest enemy. This characteristic makes them especially relevant for such studies in Australia where climatic drying (with shorter intermittent wet periods) since the Miocene has been an overriding force in determining the distribution of the biota. Such climatic changes were particularly prevalent in the Quaternary when they occurred in greater rapidity (Galloway & Kemp, 1981; Kemp, 1981; Nix & Kalma, 1972). Biogeographic analyses of Australian land snails (Bishop, 1981; Stanisic, 1994a; Solem, 1992a, b; Pokryszko, 1996) stress the post- Miocene drying of the continent as a basic factor in the evolutionary processes that have led to contemporary distribution patterns. Reasons for this have a broader ecological basis than purely snails ‘drying out’. Moisture, to a large extent, also determines the distribution of vegetation systems (Webb & Tracey, 1981) so it is not surprising to find strong correlations between distribution of land snails and certain vegetation communities; particularly rainforests in eastern and northern Australia where the majority of land snail species occur (Solem, 1991; Stanisic et al., 1994; Stanisic, 1998). Such relationships with vegetation are evident not only at the community (=guild) level but also in the geographical distribution of individual species (Solem, 1991; Stanisic, 1997c, 1998). These trends are perhaps best illustrated by the current distribution of the Charopidae, a Gondwanan family that established a close relationship with rainforests during the late Cretaceous when mesic forests were more widely spread (Stanisic, 1990). The ecological bond between land snails and rainforest is predicated on convergent habitat preference since ecological conditions that 744 favour the growth and persistence of rainforest, particularly high and stable moisture availability, and eutrophic soils, are also those favoured by terrestrial molluscs. For land snails, rainforests also provide shelter and food. In most cases, the present-day restriction of land snails to suitable patches of rainforest vegetation is absolute (Stanisic & Ingram, 1998) though in a historical sense these distribution patterns should be viewed as temporally dynamic (Stanisic, 1999), Hence from a biogeographical perspective land snails are useful for determining the vicariant shattering and isolation of mesic habitats when conditions were dry and hostile. They are also useful indicators of historic dispersal routes that became available to a wide array of organisms when conditions were moister, and thus more favourable to the movement of wet-adapted organisms (Stanisic & Ingram, 1998). From an archaeological viewpoint, they would appear to be ideal candidates for palaeoenvironmental reconstruction. ADVANCE AT A SNAIL’S PACE. So why have land snails been so rarely used in Australian archaeological studies? Kershaw et al. (in press) suggest that palaeoenvironmental research based on land snails in Australia to date, has been limited by inadequate ecological and systematic knowledge. The shortcomings in taxonomic knowledge seem to be borne out by preliminary analyses of land snail diversity in eastern Australia (Stanisic, 1994a). Since the late 1970s considerable descriptive work has been completed on land snails of the Kimberley region, central Australia and South Australia (see Solem, 1992a and accompanying references), but much remains to be done elsewhere. Several detailed regional surveys of land snails by Stanisic (1994a, 1997c, 1998) in eastern Australia demonstrate more fully the extent of these shortcomings. In southeastern Australia, Smith & Kershaw’s (1979, 1981) checklists of Victorian and Tasmanian land snails form the basis of knowledge for this region, however, even these are not based on comprehensive survey work. Nevertheless pioneering studies using these animals in palaeoenvironmental reconstruction have been attempted. David & Stanisic (1991) reported a Holocene (and possibly terminal Pleistocene) land snail sequence from Echidna’s Rest, an archaeological site in north Queensland and interpreted variations in the relative MEMOIRS OF THE QUEENSLAND MUSEUM abundance of mesic and dry-adapted taxa in terms of changes in regional moisture regimes, Ina study of land snail remains relative to local environments for the past 35,000 years at Warreen Cave, southwest Tasmania, N. Porch & J. Allen (pers. comm.) argued the snails indicate environmental changes reflecting regional climatic change rather than exclusively local factors. In a second Tasmanian study (N. Porch pers. comm.) a long sequence of land and freshwater molluscs from Pulbeena Swamp, northwestern Tasmania suggests that palaeoenvironmental trends based on pollen and ostracod sequences are supported by the molluscan sequence. Notwithstanding these studies, the limited use of land snails in Australian archaeology is regrettable when it is realised that in many parts of Australia, other palaeoenvironmental data do not appear suited to addressing archaeological questions. Many palaeoenvironmental records are founded on pollen sequences obtained from places tens or even hundreds of km from archaeological sites. OTHER CONSIDERATIONS. There are two major differences between the UK and Australia in this type or research. Firstly, in the UK people have been directly responsible for massive changes in vegetation communities since the Neolithic, principally through agricultural practices. Human impacts on the landscape were extreme to the point that most climate-induced changes were dwarfed by comparison. Consequently, major changes in molluscan assemblages (Allen, 1992, 1995a) can be directly attributed to human activities. This is not the case in Australia, where climate still is a major controlling force behind both vegetation and land snail changes. While anthropogenic fires in Australia could be considered to have affected environments in a similar way as human clearance in the UK, the protected nature of limestone outcrops in Australia has significantly limited the impact of fire in these environments. Secondly, high levels of molluscan diversity in Australia over-rides abundance. This is well illustrated by the 222 Wet Tropic species known in northeastern Queensland (Stanisic et al., 1994), as against 200 for the British region as a whole. Stanisic (1994a) recorded individual southeastern Queensland sites yielding in excess of 40 species. This allows for an organisation and interpretation of species based on their environmental associations (e.g. wet vs dry LAND SNAIL AS AN ENVIRONMENTAL INDICATOR adapted [David & Stanisic, 1991; Stanisic, 1994a]) in Australia. However, given limitations in sampling methodologies as practised in Australia, where sediment samples are sieved through wire mesh of various sizes and only the fractions caught in the mesh are kept and hand sorted for land snail shells, most very small species are by-passed and therefore ignored. Procuring and processing land snail samples from sieved excavated sediments will result in the non-collection of snails smaller than the wire-mesh (David & Stanisic, 1991) which may be critical to the level of interpretation placed on analyses. As this paper aims to show, variation in shell size of particular species, no matter how small, can provide palaeoclimatic information that may over-ride issues based on biodiversity. RESEARCH METHODOLOGIES. Two major analytical methods have been applied in palaeoenvironmental research based on fossil land snails. The first, a qualitative approach, uses a taxon’s modern ecological requirements to make statements about the environment of sub-fossil assemblages (Goodfriend, 1992), In general, shells can be extracted from ancient deposits and identified to species or genus. By attributing ancient shell assemblages to their nearest present-day ecological analogues, where species composition is known to be governed by climate and local habitats, some idea of past environments can be obtained (Lowe & Walker, 1984). This methodology follows procedures outlined by Evans (1972), and has recently been characterised as standard (Cleal & Allen, 1994; Allen & Wyles, 1993), The second, a quantitative approach, uses absolute numbers or the relative abundance of particular taxa to interpret palaeoenvironments (Goodfriend, 1992). The total number of molluscan species (living and extinct) in Britain is not more than 200, and fewer than 50 of these can generally be expected in terrestrial sites (Evans, 1972). When changing proportions of represented taxa are investigated through time, it is common to divide species into ecological categories: woodland vs. shade-loving, intermediate vs. catholic, open-country vs. closed woodland, disturbance-sensitive vs. disturbance-tolerant species (Evans, 1972; Lowe & Walker, 1984). O’Conner (1988) suggested the addition of a fifth category — wet terrestrial — after studies of molluscs as indicators of slums and ditches highlighted their benefits to the history of settlement sites in wetland areas, although Robinson (1988) is more specific in taxonomic and ecological terms. In these contexts, the usefulness of such ecological categories to the study of past people-land relations is well illustrated by the chalkland block of Malling, Cliff and Caburn. Here, the molluscan sequence has been sub-divided into phases of mature deciduous woodland with dense leaf litter, to woodland clearance and localised ground disturbance, and finally to fluctuating pressures through a grazed, dry grassland (Allen, 199Sa, 1995b). CONCEPTS AND DIFFICULTIES. Evans (1972) noted that practical difficulties fall into two main categories: stratigraphic and ecological. In the former, variations in the horizontal and vertical distribution of land snails in soil or sediment profiles may skew death assemblages in a way not truly reflective of past populations. For example, snails may occur on rock surfaces, among trees and leaf litter, or at varying depths underground. This results in the presence of different but contemporaneous species of land snails at different vertical levels, a potential problem often accentuated by the activities of worms and other invertebrates known to significantly disturb soil profiles (David & Stanisic, 1991). At death, snails on or above the ground accumulate on the ground surface through gravity, while those buried under the ground remain in situ. The physical, chemical and biological soil processes acting on this signify that the ‘subfossil’ assemblage (including that extracted for analysis) is distinct from the ‘death’ assemblage, and one further stage removed from the ‘living’ population (Evans, 1972). Studies from southern Britain have highlighted processes where sediment profiles have been altered, eroded or obscured to such an extent that ‘stratified’ data may no longer reflect real prehistoric settlement patterns (Allen, 1994). For example, the erosion of fine soil particles by sheetwash and small rills has often resulted in the regular removal of the silt, fine sand and small stone fractions of soil profiles. The effect of this in areas of soil loss has been to artificially concentrate evidence of specific types of human activity along the upper slopes. Areas of deposition tend to show a reduction in the density of such evidence on or near the ground surface, especially where total burial has taken place. Under more energetic erosion regimes, the removal, travel and physical smashing of evidence is possible (Allen, 1991, 1994). 746 Previous alternating, perhaps rotational farming regimes and subsequent erosional events through valley systems have been suggested as one explanation for a paucity of Neolithic and early Bronze Age settlement in Wiltshire (Evans, 1972). Interpretations of erosion events through land snail research have been possible given that flushes of terrestrial sediments into river and/or lake systems can be identified by the presence of non-aquatic shells and sediments within a soil profile (Evans, 1972). The relative abundance of individual species of land snails within a community is partly controlled by hereditary factors congenital to those species. Some taxa, such as Vertigo pygmaea, are seldom abundant, irrespective of the suitability of the habitat; others, such as Carychium tridentalum and Discus rotundatus, ave often prolific. Thus, ratios between species are to some extent a function of patterns of reproduction and survival (Evans, 1972). The shells of particular species of snail also vary in their ability to resist physical and chemical destruction. Hence thin-shelled taxa, such as Oxychilus, tend to be more readily damaged or destroyed than more robust shells, such as Cepaea. Some shell apices, notably those of Pomatias elegans and Clausilia, become enlarged by the accretion of calcium carbonate, and often remain in the soil for very long periods of time. The ability of some shell species to withstand both chemical and mechanical pressures during and following burial may thus cause artificial stratification down a soil profile (Evans, 1972). These issues highlight the need to include both an adequate assessment of snail behaviour and a discussion of the taphonomic status of excavated samples for each species analysed. Taking this into account, palaeoenvironmental reconstructions based on species composition or abundance in fossil assemblages depend ultimately on a knowledge of ecological factors controlling species distribution and abundance among modern land snails (Goodfriend, 1992). ENVIRONMENTAL SETTING OF STUDY AREA Northeastern Australia is perhaps best known for the Wet Tropics World Heritage Area, a region described as possessing great biological significance in terms of species diversity and endemicity. Unlike this humid belt, however, most ecosystems in the region are seasonally stressed by drought. Their research profile, MEMOIRS OF THE QUEENSLAND MUSEUM however, does not match their geographic prominence. As Gillison (1987) noted, the distribution and ecology of vegetation surrounding the Wet Tropics World Heritage Area are poorly known. Rainforest, as defined in Australia, covers a whole range of floristically related, closed canopied communities (Webb & Tracey, 1981). These communities include not only the floristically diverse and structurally complex types requiring high rainfall, but also small patches of semi-evergreen and deciduous monsoon forests and vine thickets (Kahn & Lawrie, 1983; Winter et al., 1987). In wet coastal areas complex mesophyll or mesophyll vine forest dominate lowlands and lower mountain slopes. Mesophyll vine forests also occur at altitudes of up to 800m on basalt soils where soil fertility is particularly favourable. However, patches of sclerophyll vegetation can occur within these rainforest massifs, generally but not exclusively where rainforest development is inhibited by poor or excessive drainage and regular disturbances (Kershaw, 1994). These vegetation patterns are replaced progressively by notophyll and microphyll vine forests, and eventually simple microphyll vine-fern thickets with increasing altitude and by notophyll vine forest with decreasing rainfall. West of the ranges there is a rapid change from rainforest to open forest as the rainfall declines sharply due to the rain-shadow effect of the mountains. This usually occurs above 600m with a transition band of tall open forest extending continuously for about 360km from Mount Halifax in the south to Mount Windsor Tableland in the north. Further west towards the Gulf of Carpentaria, eucalypt forests are drier and stunted. In this area, semi-deciduous vine thickets with relatively small and restricted distributions occur. Generally they are found in locally moist and fire shadow niches. The vine thickets that occur on the limestone outcrops of the Chillagoe Formation which encompass two major karst areas (Mitchell-Palmer and Chillagoe-Mungana) are typical. Dry vine thickets can also occur in environ- ments very similar to surrounding sclerophyll vegetation (Hiscock & Kershaw, 1992) and in the south, a few small patches of low, semi- deciduous, dry monsoon forest endure, scattered in an arc extending southeast from Chillagoe (Bell et al., 1987). The Mitchell-Palmer and Chillagoe-Mungana limestones, towards the northern end of this LAND SNAIL, AS AN RNVIRONMENTAL INDICATOR TAT TABLE 1. Rainfall figures for Mitchell-Palmer (mean rainfall readings taken at Palmerville for years [N89-1998, and at Bellevue for 1957-1998), ‘ | Jun) Feb Mar | Agr Mat Jun Jul Aug Sep Ovi Nov | Leer Palmeryille tear (tytn) 7AU 259. | 183 | 49 16 | 2 a | & 1 oom | 1 | mw) oP Ss Belles he mean (rim) | 225° | iy 16l . Ww \7 | | a | li 6 | 6 { Is. § a List Palmerville mean ni. rain days 7 O18 1s 7 3 | 3 R- { Zz % | 6 WW limestone arc, are geographically isolated from each other by the Walsh River and extend from just north of the Palmer River near Palmerville Station, to northwest of Almaden in the south (Fig. 1), In both areas the climate is basically semi-atid but highly seasonal. Vegetation on these limestone outcrops, where there 1s shelter from fire, is deciduous, microphyll vine thicket. Fensham (1995, 1996) denoted these vine thickets asa floristically distinct group restricted mainly to limestone karst. The relatively closed canopy of the vegetation cover and the highly dissected micro-topography of the limestone karst provide a wide range of moist, shady micro-enVironments favoured by land sails. Within each of the two areas individual outcrops are isolated by eucalypt woodland communities which are devoid of limestone, The vine thickets on these limestone outcrops and their associated pediments are largely lire-protected, Fire appears to have been a feature of the Australian dry tropics landscape for a considerable period of time (Kershaw, 1985), but its influence has increased in intensity with the arrival of Aboriginal people more than 40,000 years ago (Singh et al., 1981) and Europeans in the last 150 years. This has complicated imerpretations of environmental change based on the analyses of vegetation assemblages from pollen studies (Kershaw, 1985, 1986). However, the limestone substrate forms a natural barrier and protects the fire-sensilive vine thicket vegetation and the associated land snails. Hence climatic interpretation based on analyses of variation in shell characteristics from these environments should be relatively free of the influences of fire. Limestone outcrops in each of these locations has been mapped by the Chillagoe Caving Club (1982. 1988, 1990) and the followiny brief assessments Of local environments are based on these publications, Mitehell-Palmer. The Mitchell-Palmer limestones outcrop ina series of towers running roughly in a north-south line from north of the Palmer River to the Mitchell River in the south (Pig 1). Tower height varies, but expasures up to 150m above the surrounding landscape have been recorded (Fig, 2), Mean annual rainfall at Palmerville in the north is 1034imm; at Belleyuc Station at the southern end it is 9<0[mm, in beth cases with more than 90% of the rainfall coming in the period November-April. Mean annual rain days at Palmeryville = 88 (Table |). Evaporation and transpiration exceed precipitation for eight months of the year, with the area experiencing a marked winter deficiency of water for plant growth. Towers are higher than those at Chillagoe-Mungana and include mussive scree slopes (Chillagoe Caying Club, 1988). Chillaugae-Mungena. The limestone bluffs of the Chillagoe-Mungana region are situated in u rough northwest to southeast line extending from Rookwood Station in the north, through Mungana and Chillagoe to slightly northwest of Almaden in the south (Fig, 3A). The individual towers vary from narrowly circumscribed outcrops to expansive, relatively high limestone masses (Fig. 3B). Raintall at Chillagoe-Mungana is Jess than m the Mitchell-Palmer region, with the annual ayerage around 850mm, most of which occurs between November and April (Table 2). The entire Chillagoe-Mungana region, from its northwestem to southeastem ends, can TABLE 2. Rainfall figures for Chillagoe-Mungana region (from north to south) (Chillagoe mean mm from 1902-1998; Rookwood mean mm from 1961-1997; Almaden mean mm from 1972-1998; Chillazoe mean rain days for 1979-1982). Jan Feb 0 | ay | yal 30 195 37 Ss | 15 | Rookwood mean (mm) pat Chillagoe mean (im) Almaden mean (mim) -Chillagoe mean no, rain days Mar Ane | May | Jua (30 21 13 a Ju), Aue Sep Oe | ae Dee raps db ow 2 | 64) dak | | is 1b ,o# | a ' 5s In Ao | daa | _i6 | 8 | 5 41 | & | 1s [| 67 | 132 40) ei a) LP Be 748 MEMOIRS OF THE QUEENSLAND MUSEUM bl Ried Horse Mushragm Ro 7 1 ] at \ Giant Horse, Tt Sar Early Man , ~Sandy Creek 182 | | ‘Fed Blut =] Magnificent Gallery ® Yam Camp ns yay Cave 1, Mitchell- 4) Palmer ;. Limestone 1,', Belt waist River a> ay da ; ss, Se Fon Gaye Chillagoe. Mungana Ye x Limestone 4} Bell. Ft) . J Waikunder °° arch Cave Pillar Cave FIG. |. Map of northeastern Australia showing areas >600m ASL; and distribution of limestone karst towers of the Mitchell-Palmer and Chillagoe-Mungana regions indicating locations of archaeological sites and other places mentioned in text. LAND SNAIL AS AN ENVIRONMENTAL INDICATOR be treated as a single climatic region, with variation through space being minimal. This is well illustrated by the similarities in mean annual rainfall levels for Rookwood (850mm), Chillagoe (854mm) and Almaden (852mm). Similar patterns also exist for mean monthtly rainfall levels (Table 2). The number of annual rain days in the Chillagoe-Mungana region is also less than in the Mitchell-Palmer (e.g. average for 1979-1982 = 65). A number of the limestone towers contain caves which are today the focal points of local tourism (Chillagoe Caving Club, 1982; Robinson, 1982). PALAEOENVIRONMENTS IN NE QUEENS- LAND. Most of what we know of Quaternary palaeoenvironments in northeast Queensland comes from pollen research along the eastern seaboard and the convection-affected Great Dividing Range a few kilometres to the west. Few investigations have been made in the rainshadow area to the west of the Range, an area which covers most of north Queensland where the magnitude of environmental change can be expected to be significantly less than on the Atherton Tableland and coastal regions where coastal processes are strong. Kershaw’s work at various sites from the Atherton Tableland, some 150km to the southeast of Mitchell-Palmer and Chillagoe-Mungana — at Lake Euramoo, Quincan Crater, Bromfield Swamp, Strenekoff’s Crater and Lynch’s Crater — has revealed a series of pollen sequences covering the last 215,000 years. More recent work at the ODP 820 offshore site (Kershaw et al., 1993), Isabella Swamp, Kings Plains Lake, Garden Creek Swamp, Napabina Swamp and Barwon Swamp in the Laura region some 100km to the northeast of Mitchell-Palmer (Stephens & Head, 1995), Lake Carpentaria 500km to the west (Torgensen et al., 1988) and Negarrabullgan some 75km to the southeast (Butler, 1998) have revealed further pollen data for palaeoenvironmental recon- structions, especially for the Holocene period. Other relevent research from tropical northern Australia and the near Pacific include works on past sea levels (Hopley, 1983; Chappell et al., 1983), oxygen isotopes on corals (Aharon, 1983), ancient charcoals and woods (Hopkins et al., 1993), cheniers (Chappell, 1982; Chappell et al., 1983), lake levels (Bowler, 1983; Torgensen et al., 1988), dunefields (Galloway et al., 1970), cave sediments (Hiscock, 1984; Hughes, 1983) and simulation studies (Nix & Kalma, 1972). Together, these studies enable us to present the 749 following palaeoenvironmental reconstructions for the last 40,000 years of northeastern Australia. 40,000-38,000 BP. This was a relatively wet phase, although both absolute and relative rainfall levels were probably lower than they are today. Records from Lynch’s Crater indicate that the araucarian vine forests that had dominated much of the last glacial period as well as the previous glacial, were displaced by pyric sclerophyll forests dominated by Eucalyptus. An order of magnitude increase in charcoal peaks at Lynch’s Crater coincides with this change. Given that there is little evidence for substantial climatic change around this time, Kershaw (1985, 1986) has used this change as indicating the onset of human activity, particularly burning of the landscape. The regional nature of the Araucaria decline is indicated by a similar sequence of events at both Lynch’s Crater and the offshore ODP 820 record (Kershaw et al., 1993). 38,000-25,000 BP. Temperatures were low, but increased slightly from c.36,000 to 30,000 BP, then decreased to their lowest value sometime after 25,000 BP. Geomorphological investigations indicate that lake levels were higher than they were before c.26,000 BP. Bowler (1983) argued that this lake-full stage indicates greater rainfall (a humid expansion) around 28,000 to 26,000 BP. This is probably best explained by increases in summer rainfall. Lakes then began to contract c.25,000 BP, indicating the advent of maximum glacial activity (Bowler, 1983). 25,000-17,000 BP. The time of the Last Glacial Maximum was characterised by low effective and absolute precipitation, described as about one-third of today’s levels. This was probably caused by a mixture of exposed continental shelves (drier air masses), decreased oceanic temperatures (decreased convection), and a severance of warm ocean currents from northern Australia (decreased convection) (Hopley, 1983). 17,000-8,500 BP, Effective and absolute precipitation remained low. After the height of the last glacial maximum sometime between 13,000 and 10,000 BP, however, conditions were slowly reversed. Levels of atmospheric humidity and precipitation increased with the initial flooding of the Gulf of Carpentaria c.11,000 BP, but it was not until c.8,000 BP that warm ocean currents flowed across Torres Strait (Hopley, 1983). The spatial distribution of charcoal fragments in and around present-day rainforests 750 MEMOIRS OF THE QUEENSLAND MUSEUM Mitchell Palmer limestones. A, pul eh vine thicket; B. eucalypt woodlands surroundiny the limestone Lowers. FIG. 2 suggests that the Fucalypins expansion and rainforest displacement reached their maximum extents during the period 13,000-8,000 BP (Hopkins et al., 1993). By c.8.500 BP, both effective and absolute precipitation had inmereased to levels similar to today's: temperatures were probably also similar to those of today. 8,500-3,000 BP. This is a period of high rainfall and high temperatures. By 8,500 BP. both effective and absolute precipitation had increased to levels higher than today. Temperatures were probably similar to today. By 6.500-5,900 BP, rainfall attained levels nearly five-times those of the terminal Pleistocene on ihe Atherton Tableland. Explanations include an opening-up of the northern and eastern continental shelves, the creation of an eastern high energy window, the presence of convection as a result of warmer ocean currents, increased air and oceanic limestone tower and temperatures and higher seas levels. Around 6,000 to 5,000 BP, sea levels were similar to today’s (and possibly slightly higher), and micro-atoll and reef formations began to appear — much of the eastem continental shelf experienced reef growth and the formation of micro-atolls by around 5.000 ta 4,000 BP (Hopley, 1983). The latter closed the high-energy window, slightly decreasing rainfall levels and thus effective precipitation. Palynological and sedimentary evidence from the crater lakes suggest that fire frequencies had declined and rainforest species recolonised certain areas, with rainforest achieving its maximum extent around 6,500-5,900 BP. This coincided with an increase in Elqeocarpus relative to Cunoniaceae and Rapanea pollen values. From surface sample data, Elaeocarpus pollen has tts major representation in lower altitude forests. Bioclimatic analyses of taxa ftom Lake Euramoo suggest that summer temperatures may have been higher than today’s, while rainfall continued to be high during winter (Hiscock & Kershaw, 1992). 3,000 BP-present. Following a lowering of absolute and effective levels of precipitation around 3,000 BP. northern Australia attained its present climatic regime, There is also an indication from Ngarrabullgan that vegetation changed shortly afler 2,000 BP from forests with a relatively dense overstorey to more open forest with a grassy understorey, and that charcoal levels in sediments increased, perhaps in response to the increased lire susceptibility of that forest type. Dry rainforest was probably present, al Jeast in patches, due to the oecasional representation of Araicaria pollen. From this interpretation it is speculated that the change to more open communities facilitated the onset of swampy conditions or lake formation due to increased erosion and sediment input from more exposed soil surfaces (Butler, 1998). This is in accord with the suggestion of Stephans & Head (1995) that swamp formation was widespread in southern Cape York Peninsula after 2,700 BP (but see Hiscock & Kershaw, 1992), LAND SNAIL AS AN ENVIRONMENTAL INDICATOR There is some evidence for the onset of El Nino/Southern Oscillation (ENSO) after ~7,000 BP. The evidence for changing ENSO influences in Australia is evident in past fire, drought and erosion regimes and in the stress- and disturbance-tolerant vegetation that indicate more variable climates (McGlone et al., 1992: 435). This evidence has been argued to indicate colder, drier summers than previously between 5,000 and 3,000 BP, approximating present conditions. McGlone et al. (1992) conclude that early Holocene climates were more stable than for the mid and late Holocene, pointing to ‘either a much reduced amplitude of the ENSO fluctuations, or to a change in the extra-tropical expression of ENSO due to different climatic boundary conditions. It is unlikely that typical ENSO cycles were a major factor in Australasian and South American climates before about 7,000 BP, and they only began to exercise their present strong influence beginning at 5,000 and fully developed by 3,000 BP’. In short, the record for the last few thousand years emerges as problematic. This is particularly 751 > FIG. 3. Chillagoe-Mungana limestones. A, small outcrops; B, large tower; C, litter zone. evident given that Stephans & Head (1995) have proposed that water levels have been more reliable in the last 2,700 years, while Hiscock & Kershaw (1992) nominate driest Holocene conditions occurred between 2,600 and 1,400 BP. The disturbance of upper sediments within pollen sites is suggested as one possible explanation (Stephans & Head, 1995). However, such discrepancy more likely implies not a single discrete period of changed environmental conditions but increased climatic variability during the last 3,000 years, with short periods of decreased wet season rainfall (Lees et al., 1992). As noted above, some studies (McGlone et al., 1992; Shulmeister & Lees, 1995) have gone further, arguing for the onset of ENSO from the mid Holocene. Australian droughts generally coincide with El Nifio events, a relationship most consistent over eastern and northern Australia, while prolonged, heavy rainfall and consequent floods accompany the reverse cycle known as La Nina. The general effect of these events is to increase rainfall variability across the continent and through time (McGlone et al., 1992). INVESTIGATING PALAEOENVIRONMENTS IN NORTH QUEENSLAND VIA LAND SNAILS While some land snail taxa, such as Vertigo spp. in Europe (Evans, 1972) and Subzebrinus eremitus in Afghanistan have been used as indicators of past changes in moisture levels, comparatively few detailed studies have ever been made of the response of particular shell characteristics to specific climatic or environ- mental factors (see Goodfriend, 1986 for summary). David & Stanisic (1991) used the total land snail assemblage from an excavation at Echidna’s Rest, tropical northeastern Queensland (Fig. 1), to make broad-scale regional interpretations of environmental change in the Chillagoe region, and also emphasised that individual species may yield useful micro-environmental data. Much of our understanding of the alternation of wet and dry phases since the late Pleistocene has been gained from pollen analyses of sample cores from lacustrine environments relatively close to the eastern seaboard (e.g. Kershaw, 1970, 1971, 1975, 1976, 1985). However, while this pollen work has revealed extremely long and more or less continuous sequences, some spanning hundreds of thousands of years, palaeoclimatic and palaeoenvironmental conditions for some periods of time and for some modern biogeographic zones are better understood than others, One deficiency that is particularly problematic for archaeologists is a paucity of such data for the period 17,000 to 10,000 BP. This poorly understood period of time is compacted in the pollen sequences studied so far, particularly at the major pollen site of Lynch’s Crater (Kershaw, 1974, 1976, 1994). Elsewhere in Australia, the palaeoenvironmental evidence is clear and consistent, indicating sustained rainfall rises after the last glacial maximum at various times after 17,000 BP. In northeastern Queensland, however, the existing pollen evidence for this period, while of poor temporal resolution and dubious stratigraphic quality, implies that annual rainfall levels may not have increased until ~13,000, and perhaps even as late as 10,000 BP. Indeed, terminal glacial aridity may have been greatest around 13,000 BP (Kershaw, 1994). A determination of the timing of increased rainfall in this region has major implications for archaeological reconstructions. For example, Lamb (1996) and David & Chant (1995) argued MEMOIRS OF THE QUEENSLAND MUSEUM that people were restricted to a few sources of surface water during the dry season in the Chillagoe region during the last glacial maximum, broadening their territories and altering systems of land use as conditions became wetter and biomass increased after ~17,000 BP. Evidence for such changes in cultural behaviour is evident at Fern Cave near Chillagoe, and possibly at Hay Cave in the Mitchell-Palmer limestone zone to the north where large, permanent and apparently long-standing springs can be found. However, if rainfall levels did not begin to increase until well after people had already abandoned these places and/or broadened their lived horizons, alternative models of change in settlement behaviour — specifically site and regional land use — and cultural dynamics relative to environmental conditions would have to be considered, Unfortunately, appropriate pollen sites are rare in these regions, and sedimentation rates across the landscape are generally very slow, rendering suitable palaeoenvironmental indicators outside of caves hard to find. The shells of land snails from archaeological excavations within this region, particularly those which span the period from the last glacial maximum to present, would appear to be capable of providing the information necessary to address such questions — especially if sufficient numbers could be recovered for size analyses. Their presence within caves and rockshelters, where sedimentation is usually varied but more or less continuous, affords us the possibility of obtaining stratified sub-fossil assemblages covering relatively long periods of time. Land snails could thus act as an independent test for existing palaeoenvironmental trends, as well as perhaps also providing a finer level of detail than pollen analyses are capable of achieving. Pollen sequences are often limited to evidence of genus or higher-order changes in community composition. Vegetational changes affecting the species or sub-species often fail to register in the pollen record. This is particularly problematic in Australian semi-arid conditions, where Eucalyptus, Acacia and Melaleuca tend to predominate the upper storeys. Under such circumstances, changes in species composition are rarely identifiable from the pollen record, unless a genus or higher-order taxonomic change also takes place. However, some environ- mentally sensitive species of land snails may be able to overcome this limitation. The Echidna’s Rest study identified the helicinid Plewropoma LAND SNAIL AS AN ENVIRONMENTAL INDICATOR 753 extincta (Odhner, 1917), which is a prosobranch gastropod, as a likely candidate for such analyses, Prosobranch gastropod land snails, such as P. extincta, are more appropriate environmental indicators than pulmonates. They have an open pallial cavity rather than a closed lung, suggesting that these species are much more liable to respond to environmental moisture changes than is the case with pulmonates (Solem, 1991), In the latter the lung can be used as a ‘water-bag’ for storing moisture, thus potentially ameliorating the effects of dry climatic phases, both spatially and temporally. This greater sensitivity to changes in environmental moisture regimes could be expected to reflect in more easily detectable size changes in the shells of these animals. The underlying rationale is that as rainfall patterns and microenvironmental humidity levels vary, so will activity periods which determine growth rates. Hence the usefulness of P. extincta in palaeo- environmental reconstruction is dependent ultimately on two main factors: firstly, the capacity to show that this species does in fact respond to environmental moisture change (in time and space) in a way that can be easily measured and analysed; and secondly, the ability to recover sufficient undamaged material from an archaeological or palaeontological excavation with an extended and finely calibrated temporal sequence. The latter problem involves not only questions of site preservation, but also of excavation methods. PLEUROPOMA EXTINCTA: ECOLOGY AND DISTRIBUTION. In many land snails moisture-related responses are reflected in changes in shell patterns which can be measured. These include size, whorl number and shell thickness, although in the latter the correlation has yet to be thoroughly studied (Goodfriend, 1986). While other factors — such as temperature, insolation, the availability of calctum and population density — are also known to affect shell size, moisture appears to be the best documented factor (Goodfriend, 1986) and has been the basis of several detailed studies (Tillier, 1981; Gould, 1984) which suggest that it perhaps is a critical factor in determining both size and shape differences within species and between populations. Considering the acute moisture sensitivity of land snails, such a conclusion would seem appropriate. However, given the relative lack of studies in this sphere and the inconclusive nature of some, it is not possible to accept the universality of the moisture-size relationship and each case must be assessed separately. It also needs to be kept in mind that though the size of shells is determined by whorling pattern, shells can become large in two ways: few large-sized whorls or many small-sized whorls (Gould, 1984). However, a strong positive correlation between size and whorl count may also indicate that size increase is purely a function of whorl count increment without any appreciable change in whorl size. Hence, determining the shell parameter which will be most useful as an index of moisture levels also becomes a crucial consideration. LAND SNAILS IN SEASONAL ENVIRON- MENTS. Environmental moisture-related variation in the shell size of Australian land snails has been reported by Solem & Christensen (1984) for camaenids from the vine thickets of the Kimberley and inferred for a number of non-camaenids from other seasonal environ- ments in tropical northern and northwestern Australia (Solem, 1982; 1984). In each of these cases the implication was that variations in shell size correlated with differences in whorl numbers of adult shells and that they were not merely due to variations in overall size wherein large shells result from larger animals with larger whorls and vice versa. Furthermore, Solem (1991) suggested that shell-size variation in populations of Pleuropoma walkeri (Smith, 1894), a helicinid from the vine thicket patches of northwestern Australia, may also be a reflection of differences in local moisture conditions, although no analysis was undertaken. This latter inference has significant implications for using P. extincta in fine-grain palaeoenvironmental reconstruction. Reasons for the size differences in the shells of the land snails from these monsoonal (= seasonal) environments relate directly to the nature of the climate and the specific adaptation of the local snails to this climate (Solem & Christensen, 1984: Solem, 1988). These land snails have the capacity to vary in adult shell size because of variations in the length of feeding activity available during growth from juvenile to adult (Smith & Stanisic, 1998), This feeding activity is related to intensity and duration of summer rainfall (‘the wet’), and since these snails appear to reach adulthood synchronously at the end of their second wet season (Solem, 1982; 1984), maturity can lead to adults with different whorl numbers depending on the quality of the growing 754 seasons. However, it also needs to be noted that while rainfall results in growing activity, this response is not always a simple function of annual rainfall or annual number of rain days. Rather, land snail activity periods during and after significant ‘activating’ rain events vary according to the complexity of local environmental conditions, which include such factors as vegetation cover, topography and geology (Solem, 1988). The Chillagoe-Mungana and Mitchell-Palmer limestone outcrops (Fig. 1) are part of this archipelago of seasonal environments and receive most of their rainfall in a relatively short time during summer (see above). The land snail faunas of limestone areas in eastern Australia are comparatively rich when compared with the snail fauna of the surrounding non-limestone areas (Stanisic, 1994; 1997a,b). However, while the diverse nature of the fauna in some of these areas has been documented (Stanisic, 1997c), many of the taxa have yet to be formally described. The land snails of the limestones of the Chillagoe Formation are no exception, Odhner (1917) described the first species from Chillagoe and since then only a few extra species have been added (Iredale, 1937, 1938). Their classification and distribution are summarised in Smith (1992). Stanisic (1997d) briefly detailed and illustrated the land snail fauna of the Chillagoe-Mungana limestones, including undescribed species, and drew attention to its diverse make-up of widespread and locally endemic taxa. The equally prolific Mitchell-Palmer land snails are largely undocumented, though they are well represented in the collections of the Queensland Museum. These two limestone areas share a number of taxa, including P. extincta. This species displays shell size variation across the range of its occurrence (Stanisic, unpubl.). P extincta has also been recovered from archaeological excavations at Hay Cave, towards the northern end of the Mitchell-Palmer limestones (Fig. 7). Radiocarbon dates associated with cultural remains from this cave suggest a temporal sequence covering the last ~19,000 years at least, a time period which spans a number of major wet/dry climatic phases in the region, These circumstances provide an opportunity to examine the extent of shell variations in recent P. extincta and to see whether these have palaeo- environmental implications. MEMOIRS OF THE QUEENSLAND MUSEUM Pleuropoma extincta (Odhner, 1917). The shell of P. extincta is small (adult size: diameter 2.3-6.6mm, height 2.0-5.4mm), top-shaped and sculptured with numerous spiral cords; the periphery is angulate, there is no umbilcus and the shell is relatively thick (Fig. 4). Its small size and relatively thick walls make P. extincta shells capable of withstanding relatively high levels of chemical and mechanical weathering, an important consideration when investigating buried sequences. P. extincta is comparatively widely distributed (N-S approximately 175km) in the Einasleigh Uplands Biogeographic Region, but it is environmentally restricted (Stanisic, unpubl.). It appears to live exclusively on the limestones of the Chillagoe Formation (Fig. 1). The species was mistakenly recorded from a tiny patch of vine thicket in the Mt. Mulligan area, northeastern Queensland by Stanisic & Ingram (1998). While the Mt Mulligan population is superficially similar to P extincta, recently obtained anatomical details show that it is a different species (Stanisic, unpubl.). P extincta is an obligate vine thicket dweller (Fig. 2A) and has not been recorded from the more expansive eucalypt forest/woodland communities which dominate much of this region (Fig. 2B). It is also absent from the vine thickets growing on the massive lava flows which occur in the surrounding countryside, such as at Undara and Kinrara to the south. The latter habitats are occupied by another, undescribed species of Pleuropoma (Stanisic, unpubl.). Present-day distribution of P extincta is the result of long-term, climatic changes which have also affected the distribution of vine thicket communities (Stanisic, unpubl.). Because of their enhanced ability to retain ground moisture, vine thickets associated with the limestone pediments act as refugia for P. extincta and other micro-organisms during dry phases. P. extincta’s current restriction to these habitats is indicative of its past and present sensitivity to regional and local moisture regimes. P. extincta lives chiefly under strewn limestone talus, on leaves and bits of timber in the litter and on the trees which grow on these outcrops. It may also occur on the limestone rock where it seals itself to the karst surface in the many cracks and crevices. Compared with most other land snails on these limestones, it is present in very high numbers. P. extincta is not subterranean but may occur around the entrance of caves. It does not burrow, an important consideration when analysing subsurface, sub-fossil assemblages. Its LAND SNAIL AS AN ENVIRONMENTAL INDICATOR 75 NIG. 4, Shell of Pieurpome exrineta.[A, Francis). presence in great numbers along drip points within caves is probably a result of the removal of finer, surface sediment particles by percolating water, concentrating larger particles (including snail shells) in exposed pockets. Because of their large numbers in living populations and the alkaline nature of the environment, dead shells also accumulate in the litter in great abundance (Fig. 3C), In MATERIALS AND METHODS Recent specimens of P. extincta came from the collections of the Queensland Museum (QMMO). Methods comprised hand-collecting specimens and specimen-containing leaf litter from various limestone outcrops, including the ground surfaces of cave entrances and interstitial spaces between rocks and boulders on talus slopes. The shells were subsequently separated from the leaf litter by hand-sorting in the laboratory. Shells retrieved from leaf litter represented the accumulation of several years of dead individuals. The ability to sample land snails pos!-morien greatly enhances their utility in environmental interpretation. Preliminary analysis of samples involved separation of individuals into three categories; adult or juvenile, based on the thickness of the outer-lip (perisiome), and unclear according to the degree of coating of calcium carbonate. Each sample category was counted, totalled and labelled, To facilitate measurement 10 adult P extincta shells were placed at a lime on the stage of a calibrated dissecting microscope (Zeiss 475052-9901) and observed by reflected light under * 12 magnilication, The parameters measured were whorl count, height and diameter. Of those thal were partially broken or damaved, it was only possible to measure one or two of the three variables. These damaged specimens were also counted and labelled, Shell height was measured from the apex to the base of the aperture, and diameter covered the lateral span of the body whorl. These observations were recorded to the nearest 0,05mm, Measurements beyond 5.00mm were performed with a dial calliper (Mitutoyo 505-667 D20F), accurate to 0.0] mm. Calculation of whorl number was coordinated by aligning the origin of the whorl pattern at the apex in the vertical position and counting to the nearest eighth rotation (Solem, 1976). STATISTICAL ANALYSIS. The mean value for each parameter within each oulcrop unit sample was calculated to a single standard deviation. In this way, statistical comparisons of parameters from different sites were possible. In each analyses the range of measurements was noted for each character. For convenience of data presentation and interpretation, Variations within shell characteristics are presented as scatter-graphs of diameter vs. whorl count, height vs. whor! count and height vs. diameter. Such statistical analyses 756 Mitchell-Palmer > 7 6.5 + = 6 E 55 — 5 + + 5 45 ts : + s cA y =0.9996x + 0.8938 3% R? = 0.2227 2 3 35 4 45 5 55 6 Whorl Count Chillagoe-Mungana y = 0.7477x + 1.3932 R? = 0.0735 E £45 Ss 4 o 53s o 3 25 2 3 3.5 4 45 5 Whorl Count MEMOIRS OF THE QUEENSLAND MUSEUM Mitchell-Palmer Height (mm) » y = 1,4084x - 2.0885 R? = 0.4352 3 3.5 4 5.5 6 4.5 5 Whorl Count Chillagoe-Mungana 45 y = 0.8628x - 0.3298 4 R? = 0.153 E 3.5 = e ¢ 3 a ‘3 =x 3 3.5 4 Whorl Count 45 5 FIG, 5. Surface collections (control samples), P. extincta statistics (with simple regressions). A, whorl count versus diameter (mm), Mitchell-Palmer; B, whorl count versus height (mm), Mitchell-Palmer; C, whorl count versus diameter (mm), Chillagoe-Mungana; D, whorl count versus height (mm), Chillagoe-Mungana. were first undertaken for each individual sample set (i.e. samples from individual limestone outcrops). In order to investigate spatial patterning in shell height, diameter and whorl count within a regional framework, adult data were pooled into a combined Chillagoe- Mungana dataset, and into a Mitchell-Palmer dataset. To describe and quantify the relationship between the parameters, and determine the strength of any possible correlation, a simple regression analysis and associated equation is provided for each graph. In addition, to allow for an understanding of the relationships between the parameters for the entire study and determine the correlation strengths at this scale, all regional data were combined on a single graph. This technique therefore used all measured variables in the one analysis. Here, the regressions were second- order polynomial to account for the presence of two regional populations (Chillagoe-Mungana and Mitchell-Palmer). RECENT PLEUROPOMA EXTINCTA Analysis of shell variation (Tables 3-5). Average largest shells and highest average whorl counts occur among the Mitchell-Palmer material (Fig. 5A,B). When the Mitchell-Palmer and Chillagoe- Mungana areas are treated together, whorl count depicts strong positive correlation with height and diameter (correlation coefficient ~0.72-0.83; Fig. 6A,B). When each region is treated in isolation, however, the strength of this relationship is not as evident (Fig. 5A-D), with the correlation coefficient failing to exceed 0.44. This latter poor correlation is also evident when each collection area was analysed as a separate | Diameter No. of | (mm) | whorls | sem and range). | salty | No. of Height Locality | | Specimens | (mm) _| ‘Chillagoe- 602 | 3:075+0.224 | 4.34440.280) 3.947+0.101 _ Mungana | 9? | (2.05-3.70) | (2.30-5.30) | (3.625-4.250) Mitchell- gy | 4.205+0.353 | 5.361+0.350) 4.46840.165 (Palmer =| > | (3.00-5.35) | (4.00-6.55) | (3.375-5.000) LAND SNAIL AS AN ENVIRONMENTAL INDICATOR 757 TABLE 4. Local variation in P. extincta from Mitchell-Palmer area (mean, sem and range). | QM Reg. no. No. of Mean height | Height range | Mean diameter Diameter range Mean whorl Whor! count hc _ specimens | (mm) _ (mm) (mm) (mm) count range MO229988 | 5 3.110+0.119 3.00-3.30 | 4.15040,087 4.00-4.20 | 3.92540.112 | 3.875-4.125 | MO23375 i 2 errem 3.45-4.20 | 5,179+0,226 E 4.75-5.80 4.37740.159 | 4.000-4.750 | MO23388 | a2 | 3.950+0.515 | 3.20-4.70 | 4.85040.420 4.45-5.60 | 4.46420.310 | _4.125-4.875_ | MO23584 | __29 | 4.03640.202 | 3.65-4.45 | 4.98440.234 | 4.55-5.35 4.40140.158 | 4.125-4.750 | -MO23592, 7 | -4.15040.367 | 3.65-4.65 5.52940.438 | 4.90-6.10 | 4.482+0,.152_ | 4.250-4.625 MO61156 208 | 4.148+0.249 3.60-4.85 | $.28540.243 | 4.70-5.95 | 4.46840.134 | 4.125-4.875 | }MO61172, | 24 | 438140210 | 3.85-4.70 5.42740,251 5.00-5.85 | 4.53140.129 | 4.250-4.750 — _MO61191 87, 4.3420.262 | 3.80-5.05 5.45240.222 | 4.95-5.90 | 4.476£0.180 | 3.375-5.000 MO61205 _ 1 103 | 4.529+40.317 | 3.95-5.35 5.73540.294 | 5.00-6.55 4.557+0.123 | 4,250-4.875 | outcrop unit. In terms of variation, the whorl counts, diameters, and heights of shells from any one region, single limestone outcrop unit or individual collection unit appear smaller; the spread between the values becoming narrower, forming a cluster, rather than distinct linear spread, on the graph. Relating this to analytical scale, the positive correlation between whorl count and size of the shells is thus illustrated at the inter-regional level and does not arise at a more local scale. This patterning is also visible in the geographical pattern of rainfall variation, which is fairly marked at the inter-regional level, but not at a local scale. This correlation implies that the relationship between whorl count and diameter or height is likely to be extrinsic rather than intrinsic. It implies that the relationship between whorl count and diameter or height requires variation in environmental influence(s) on these variables. This is consistent with findings of strong environmental, and in particular humidity, influences on whorl growth (see above). Highest whorl counts, calculated as % of total numbers of individuals occupying various whorl count classes, shows the Mitchell- Palmer specimens to be consistently higher than those at Chillagoe-Mungana (Table 3). ENVIRONMENTAL INTERPRETATION. Differences in mean shell size between the Mitchell-Palmer and Chillagoe-Mungana material correlate positively with differences in local rainfall regimes. On average the Mitchell- Palmer specimens are larger then Chillagoe- Mungana ones, which has lower average annual rainfall. Although larger shell size can result from a number of other factors, such as temperature, insolation, calcium availability, population density and soil pH, their effects were not measured. Studies elsewhere have linked some of these factors with differences in whorl expansion rates, shell weight, diameter and height (Bengston et al., 1979; Owen, 1965). In most instances, differences in shell size were related to differences in whorl cross-section. Goodfriend (1986) presented several hypotheses to explain the relationship between these environmental variables and adult shell size. However, in most cases the differences were in connection to the initial size of the whorls and the rate at which the whorls expanded — that is, the aperture and whorl cross-sectional area (=whorl volume). In this case, variation in aperture and whorl cross-sectional area do not represent a similar function to the number of whorls, but they are generally interpreted as representing either an adaptation for regulation of water loss, to reduce predation, or both. They are, therefore, not as dependent on the surrounding habitat (Goodfriend, 1986). In the context of this study, the strong positive correlation between whorl count and both diameter and height in P. extincta indicates that size differences in this species have an affinity to variations in whorl ‘count’ rather than whorl ‘size’. These findings coincide with the camaenid land snail growth patterns in semi-arid areas of northwestern Australia as discussed by Solem & Christensen (1984), Adult shell size in their study is seen to depend upon moisture conditions near the end of the snail’s second wet season. An early interruption of moisture supply causes growth to stop at a lower whorl count and produces smaller adults. Favourable moisture conditions allow extended growth time, with cessation occurring at a higher whorl! count, and producing a larger adult size (Solem & Christensen, 1984). What Solem & Christensen (1984) implied is that the regular and predictable wet season in the Kimberley region is the key to understanding the maturation pattern ofits land snails. The length of 758 A Chillagoe-Mungana + Mitchell-Palmer Diameter (mm) 3 y = -0.1102x" + 2.6638x - 4.3984 o R= 0.7244 3 ae 4 45 5 55 Whorl Count MEMOIRS OF THE QUEENSLAND MUSEUM B Chillagoe-Mungana + Mitchell-Palmer y = 0,3436x° - 0.938x + 1.4721 5 R?=0,831 Height (mm) 3 3.5 4 4.5 5 55 Whorl Count FIG. 6. Mitchell-Palmer + Chillagoe-Mungana. A, whorl count versus diameter (mm); B, whorl count versus height (mm). Regressions are second-order polynomial, to take into account the presence of two regional populations. the wet season may vary from as little as two months on the southern fringes, to as long as six months in the Mitchell Plateau. The median wet season rainfall is only 500mm near Fitzroy Crossing and Halls Creek, but nearly 1600mm on the Mitchell Plateau. These parameters define the potential maximum length of the activity period for the snails (Solem & Christensen, 1984). The lower number of rain days at Chillagoe- Mungana compared to the Mitchell-Palmer appears to at least partially account for the smaller-sized (= lower whorl count) snails. Given this, whorl count in P extincta appears to be a useful indicator of differences in regional rainfall levels, which in turn suggests that this shell feature may also be a useful index of change in past environmental moisture levels along the limestone belt of the Chillagoe Formation. Evidence indicating the sensitivity of P. extincta to environmental moisture levels is strengthened through the presentation of initial findings highlighting little intra-regional variation, where each outcrop has a fairly uniform rainfall pattern. However, the most significant correlation between size and whorl count is between regions with perceptible differences in mean annual rainfall levels. In contrast to the above, Solem & Christensen (1984) concluded that the pattern of size variation in land snails in the Kimberley region was very much a local phenomenon dependent on both moisture regimes and microhabitat. Populations of Westraltrachia at Wombarrella Gap differed by up to 3mm in mean adult diameter over distances of less than 100m. The small shells were collected near a single isolated boulder, and the larger specimens were taken in shaded talus that would retain moisture for a longer period of time (Solem & Christensen, 1984). Although the north Queensland results do not show the strong patterns of local variation seen in some Kimberley species it should be borne in mind that the recent material was mostly collected from large karst towers whose micro-topography was generally considered to be similar. There are however, tantalising indications that local influences would be worth further investigation and that outcrop size and aspect may also play a mitigating role in final shell size of P. extincta. For instance, material from Mitchell-Palmer sample QMMO22998 was collected from a small, low relief outcrop and has comparatively smaller whorl counts and size than others from the region (Table 4). Specimens from Royal Arch Caves, Chillagoe-Mungana (QMMO53436) displayed the largest size and whorl counts for specimens from this region (Table 5). While this limestone outcrop does not demonstrate any apparent physical differences to others from the region, the land snail assemblage does suggest that it has intrinsic differences. The large camaenid land snail, Spurlingia praehadra (Odhner, 1917), occurs almost exclusively on this outcrop indicating that it probably is some- how environmentally distinctive. Further research on these aspects appears warranted because if these casual perceptions about local differences in P. extincta are related to local habitat differences, then the role of P. extincta as a useful indicator species would be enhanced. APPLICATION TO AN ARCHAEOLOGICAL ASSEMBLAGE: HAY CAVE Hay Cave is located towards the northern end of the Mitchell-Palmer limestone belt, near LAND SNAIL AS AN ENVIRONMENTAL INDICATOR 759 TABLE 5. Local variation in P. extincta from Chillagoe-Mungana area (mean, sem, range). QM Reg. no. ; No. of Mean height Height range | Mean diameter Diameter range | Mean whorl Whorl count 1 ~ |_ specimens (mm) (mm) (mm) (mm) ___ count : range | _ MO1S955 9 3.067+0,224 2.70-3.40 4.41140.204 | 4.00-4.65 3.93140.110 | 3.750-4.125 | MO19527 85 2.968+0.220 | 2.05-3.40 | 4.18940,208 | 3.65-4.60 3.89140.107 | 3.625-4.250__| | MO025996 40 —3.11140.195 _2.55-3.55 4.32840.186 4.00-4.75 | 3.93140,109 | 3,750-4.125 | | MO38499 6 —___3.10040.148 2.90-3.35 __3.97540.829 | 2.30-4.45 | 3.97940.094 | 3.875-4.125 | MO50229 15 | 2.82340.231 2.50-3.40 | 4.17340.171 3.90-4.45 3.908+0,088 | 3.750-4.000 MO50231 19 —3.32440,.197 | 2.95-3.65 4.61640,242 | 4.00-4.95 3.96140,094 | 3,750-4.125 |_MO53325 31 | 3.14820.166 | 2.80-3.45 4.574£0.237 4.20-5.15 | 3.948+0,090 | 3.750-4.125 | __M053335 19 3.005+0.222 | 2.70-3.50 | 4.36340.309 | 3.90-4.95 3.888+0.109 | 3,750-4.125 MOS53436 25 | 3.452+0.154 3.20-3.70 4.910+0.194 4.60-5.30 4.070+0.063 | 4.000-4.125 | MO53448 20 | 2.860+0.240 2.15-3.20 | 4,05040.145 | — 3.65-4.25 3.94440.137 | 3,625-4.125 MOS3551 21 | 3.06040.119 | 2.90-3.30 | 4.400+0.152 | 4.15-4.65 3.92940.075 | 3.875-4.125 | __MO53564 20, 3.06340.143 | 2.85-3.40 4.31040,136 | 4.05-4.55 _-4.01340,099 | 3.875-4.125 | MOS8485 220 | 3.116+0.184 2.05-3.40 | 4.38640.216 3.80-5.00 | 3.953+0.089 | 3.750-4.125 MOS8540 | 31 | 2.94740,158 2.65-3.35 | 4.07440.145 | 3.70-4.40_——3.948+0.063 | _3.875-4.000 | _MO58547 41 | 2.985+0.174 | 2.60-3.45 4.23740.164 3.90-4.55 _3.976+0.098 | 3.750-4.125 Palmerville Station. Mean annual rainfall is estimated to approximate 1,034mm given its proximity to the station. It is a medium- to large-sized cave positioned at the base of a large limestone tower. The site’s entrance is located at an estimated 2m above the surrounding plains. The area outside the dripline displays evidence of roof collapse in the form of numerous boulders. Boulders caused by roof-fall are also present inside the dripline in some cave chambers, but never in the central area of human occupation, where the excavation was undertaken. The cave entrance is itselfnot visible from the outside, as it is hidden by a thick web of vine thickets. These circumstances lead to a lack of direct sunlight reaching the cave, and consequently a relatively constant, cooler and humid internal climate. Evidence of human activity includes a massive boulder whose edges have been flaked and numerous paintings that occur from the entrance to the very back of the cave, a distance of approximately 17m. The central part of the cave possesses soft, ashy floor deposits, signalling the presence of ancient fireplaces below ground. FIELD AND LABORATORY METHODS. Hay Cave was excavated by BD and HL as part of an Earthwatch-funded project in mid 1996 (Fig. 7). The reasons for the excavation were multiple, focusing on human responses to environmental change in north Queensland during the last glacial maximum (in particular the period 21,000 to 13,000 BP). Four juxtaposed 50 50cm squares (test pits), forming a | x lm square matrix, were excavated. Excavation commenced near the central zone of soft ashy surface deposits where sediments appeared to be deepest (and potentially oldest) and least disturbed by animals and water. The 4 test pits were labelled M30, M31, N30 and N31. Square M30 proved to be the deepest square excavated and the focus of subsequent investigations. Excavation methods employed ~2cm excavation units (XU) within stratigraphic units (SU). This method simply recognises that sediments have in the past accumulated progressively, at times resulting in the formation of distinct strata. Excavation units are employed during excavation because sediments of similar composition may have accumulated over long periods of time, so that a single stratigraphic unit may represent many hundreds or even thousands of years of sediment accumulation. Excavation units thus enable the excavator to sample different parts of a given stratigraphic unit, enabling an investigation of its internal structure and contents. Major changes in sediment colour, texture and observable content at the time of excavation and section description were deemed separate SU: minor stratigraphic changes were identified as changes in sub-layers. The SU were numbered sequentially from top to bottom, with sub-layers characterised by an alphabetical letter after the SU number (e.g. SU2a and SU2b). The location of all cultural material measuring over 2cm maximum diameter and observed in situ was 760 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 7. Hay Cave. recorded in three dimensions, plotted on recording forms and bagged separately. The remaining sediments were weighed, dry-sieved in 3mm-mesh wire sieves and bagged for subsequent sorting. In the laboratory, this material was later wet-sieved, air-dried and sorted. Sorted sediments thus resulted in various components for each XU, including stone artefacts, bone, mussel shells, land snails and calcium carbonate (CaCQ;) concretions. Sediment samples from the <3mm residue were collected from each XU from each square along with pH readings. Pollen and oriented sediment micromorphology samples were also collected TABLE 6. Radiocarbon dates from Hay Cave. Age (BP) _ Lab no. XU, Depth(cm) | || “xU2 | 44 350255 Wk-6053 XU6 8.0 66075 OZD-006 | XUI2 17,1 | 870+65 OZD-007 | xU26 460 | 2.590280 OZD-008 | xU34 64.0 3.100260 | —OZzD-009 |XU45 9L0 13, 4504150 [ OZD-011 xua9 |__ 102.8 13,6004180 | 7-422 | XUS6 118.2 | 193004140 = OZD-012 from the pit walls upon completion of the excavation. Dating. Eight samples for radiocarbon dating were extracted from square M30. Seven of these dates were derived from charcoal: the remaining OZD-012 determination was on freshwater mussel shell. All samples were collected in situ from the upper two SU where sediments were relatively easy to dig. These radiocarbon dates were obtained from various parts of the square, covering much of the sequence. The charcoal and mussel shell were pre-treated for possible rootlet and carbonate contamination by the radiocarbon laboratories before dating. Sample Wk-6053 was sent to the University of Waikato Radiocarbon Laboratory (Hamilton, New Zealand). All other samples were forwarded to ANSTO (Physics Division, Lucas Heights Research Laboratories, Australia). The former date is a conventional radiocarbon determination, the latter by accelerator mass spectrometry (AMS) (Table 6). Microscopic and statistical analysis of excavated materials. The segregation of materials from the excavated sediments provided a useful opportunity to identify changes in the relative proportions of components. This allowed the LAND SNAIL AS AN ENVIRONMENTAL INDICATOR 76) quantity of deposited land snail shell to i be graphed according to excavation a unit. Asimilar method wasemployedto _ °° graph the frequency of CaCO; 2 5° concretions. In turn, the P extincta ®& 4° shells were separated in an equivalent = °° manner to the modern collections, 200 enabling the number of specimens tuo deposited per unit time to be calculated. 0 Shell measurements and statistical analysis were undertaken in the same manner as employed in the modern samples. The mean whorl count, height, and diameter from each excavation unit were outlined on three graphs. Low P. extincta counts in some XU highlighted a need for caution in interpretation, and consequently neighbouring XU with under 10 specimens were combined to increase sample size. A third order polynomial regression was super-imposed on the graphs. This choice of regression type reflects the aim of investigating long-term environmental change, the nature of variation within the data and the possible representation of three major climatic regimes for the period in question, as informed by pre-existing palaeoenvironmental research. Trends in shell characteristics were then identified through time. RESULTS. Radiocarbon determinations (Table 6) of age range from c.19,300 to c.350 years BP, and a precision of one standard deviation is given for each. All dates are in sequence. Deposition of sediments within Hay Cave. There is a high incidence of land snail shells by weight (all species combined, but consisting mainly of large Hadra aff. bipartita and Xanthomelon pachystylum) from 13,600 BP to around or shortly before 3,100 BP (XU49-XU38) (Fig. 8). This incorporates a peak of 688.0g at XU40. Prior to this time, total shell weights are comparatively low. Above XU38, a pronounced decrease to almost negligible levels is evident, encompassing the period 3,100 BP-present (XU33-XU1). The highest incidence of CaCO; concretions in Fig. 9 corresponds in timing with the period of peak land snail shell deposition, between 13,450 BP and 3,100 BP(XU45-XU34). Deposition rates of P. extincta shells per 100 years (Table 7) reveal two major depositional regimes: one before approximately 3,100 BP (XU56-XU35) characterised by significantly low shell frequencies, and the other since 3,100 BP (XU34-XU1), when numbers are some 20-fold higher. S Se SAD PPP dA YP SP XU FIG. 8. Land snail shell excavated from square M30, Hay Cave: total land snail shells by XU, Pleuropoma extincta at Hay Cave. A total of 295 P. extincta shells were recovered from excavation square M30 at Hay Cave. Most (236) are adult specimens, with 59 at undetermined stages of maturity given the presence of CaCO, encrustation or breakages effectively removing or hiding characteristic features. Of the adult specimens, 17 were partially broken. This left 219 specimens where each of the three measurements — height, diameter and whorl count — could be made. Mean whorl counts, heights and diameters by excavation unit at Hay Cave are represented in Fig. 10. A description of relationships is based on the shapes of the regression curves, which display similar trends but are not identical to each other. Of the three represented graphs, mean whorl counts demonstrate minor variation across the time span, indicating slight decreases from an under-pronounced peak at 13,600-13,450 BP (XU49-XU45). Values change little from about 3,100 BP (XU34) until 660 BP, after which they decline further to 350 BP (XU2) and present levels. However, variability between individual samples through the sequence is high. TABLE 7. Deposition rates of P. extincta numbers per 100 years at Hay Cave (note that XU45 to XU49 were not separated as dates were within one standard deviation of each other). Duration in | No. shells/100 years | years _ | 350 leewnrt =| i | XU range No. of shells _xu-2 | oa XU3-6 33 | 310. | 10.64 jes Ea | 20 15,24 | XUI13 - 26 _ 89 1720 S70 2 | XU27-34 | $9 S10 11.57 XU35-49 | 53 10,500 0.50 XUS0- 56 1 $700, 0.02 762 ( nN [s} i=] Oo CaCO3 concretions (g ro) oa oO 0 foarererwere mt NO OB bog BP op mh PO wo XU 2 FIG. 9, Total weight of calcium carbonate concretions excavated from square M30, Hay Cave, by XU. Mean heights show more extreme variation of the trends evident in whorl counts, expressing a greater decrease through the last 660 years (XU6-XU2) and indicating an obvious peak at 13,600 BP (XU49). This peak is followed by a marked decrease through the period 13,450- 2,590 BP (XU45-XU26) and a rise to a lower peak at 660 BP. The trend for mean diameter is similar to that for mean height. This tendency was generally expected given that height showed strong correlation with diameter in the Mitchell-Palmer and Chillagoe-Mungana control study. The P. extincta shells dating to 13,600 BP (XU49) correspond to peak levels, decreasing to relatively small measurements from 3,100 to 2,590 BP (XU34-XU26). Diameters rise to a second, almost equal peak through the period 870-600 BP (XU12-XU6), after which they decrease to low levels. As with mean height, variability between individual samples is high. DISCUSSION, Major temporal phases can be identified from the Hay Cave fossil molluscan records via an analysis of shell sizes and whorl counts. These quantifications in turn can be linked to trends in relative frequency of land snail shells through the dated sequence. However, such comparisons also raise questions about the broader geographical applicability of existing palaeoenvironmental trends based on pollen. The general shapes of the regression curves are likely to be reliable, given the stratified nature of the sequences which appear to possess good internal integrity. However, the degree of stretch (as opposed to the shape) of the regression curves — that is the length of each section of each curve — is dependent on the period of time covered rather than by the depth of deposits. Currently these curves have been presented by excavation unit, which basically represents depth below MEMOIRS OF THE QUEENSLAND MUSEUM ground. The last 3,100 years are over-emphasised in such curves, given peak sediment deposition rates during that time. The early to mid Holocene period appears to be present in the Hay Cave sequence despite an absence of radiocarbon dates from this period, given the high land snail shell and So ¢ CaCO; concretion frequencies from XU45 to XU34 (Figs 8,9). However, their exact timing Is uncertain, given a total absence of radiocarbon determinations for this part of the sequence. The frequency of redeposited CaCO; can be taken as an indicator of changing moisture regimes, given the susceptibility of limestone to solution. The greater the solubility of a substance, the greater its liability to come out of solution. Many minerals are deposited in caves as chemical precipitates, and calcite is the mineral form of CaCO;. Although reactions are reversible, calcite is chiefly precipitated through the diffusion or ‘degassing’ of carbon dioxide (CO,) from water into the atmosphere. To describe a general reaction, seepage water, initially enriched in CO» given the presence of soil and vegetation, becomes saturated in calcite when passing through joints in the limestone. Upon entering the cave environment with lower partial pressure of CO) in its atmosphere than in the soil above, CO, diffuses from dripping and flowing water and causes calcite to be precipitated out of solution and deposited. In essence, CaCO; deposition in caves implies a supply of water and vegetation growth sufficient to enrich soil water with carbon dioxide for limestone solution. From this, calcium carbonate is observed as an inorganic indicator of high moisture levels (Jennings, 1985) The calcium carbonate curve is matched by other proxy indicators of environmental humidity. As a biological index, the incidence of land snail shell (Fig. 8) mimics the relative frequency of CaCO, concretions between 13,450 and 3,100 BP, an alliance related to the notion of Evans (1969) that essentially all molluscs are aquatic animals. As discussed, the habitats and behaviour patterns of terrestrial snails are largely controlled by the humidity of the environment. For example, Hadra aff. bipartita, a dominant taxon between XU45 and XU34, requires humid conditions. Observations on species in north LAND SNAIL AS AN ENVIRONMENTAL INDICATOR > y @ 4E05e 1 OTT. -O 02934 + 4 1632 PATI I7+18 24425 § Excavation Unit 31+32 3/+38 47448189 £ 7O35 —__——————————————— SS. w 4, y =-,00020*+ 7 OOTas" - DGB 1857 se however, there is no simple relationship between amount of charcoal in a site and the number of radicearbon dates obtained. In 1utm, 260BP ri the number or presence of a radiocarbon dates may not be aber indicative of the magnitude of humun occupation, as biases in the creation, recovery, and preservation of eaten charcoal and radiocarbon dates in a sequence or in a region are more 34b0EF likely to critically affect results. 2 4508 Biases may skew the date curve, and 3.60087 ag 34 14 a4 36 #37 a8 ag 4 4 Mean Height (mm 2) ur DOSGSt + ya TE dx 7 0 Ud 1 « 7034 thus potentially falsify the common 42 43 gssumplion thal human occupation will produce more deposited carbon, even if the relationship is not strictly aaoae = Proportional. At Hay Caye this 1s well illustrated by the continued presence of burnt bone, shell and BRIBP oe =nypp _ Carth in the purported early Holocene 5 isasia . levels Where no radiocarbon dates Baha have yet been obtained and chareoal ¥ er is very sparse, We argue that such an a zssoeP = absence of charcoal in these layers 1s 312 sioee = OL due to its original absence, as 37438 _ evident in the ample cases of burn! i a tsoep Materials in those layers. Rather, GG. dae Gi Middac uc aD 9 Be ars charcoal! has probably been removed Mean Diameter (rom FIG. 10. P. extineta measurements by XL. Hay Cave. A, mean whor! count: B, mean height: C. mean diameter. Queensland suggest that land snail activity is largely conlined to the wet season. and hence there may be a relationship between the length of the wet season and number of individuals. The high shell accumulation, together with CaCO, concretions could indicate conditions were wettest during the period 13,450 to 3,100 BP in the Hay Cave sequence, independent of P. extincia shell measurements. This would be consistent with high rainfall regimes during the early to mid Holocene, as informed by existing post-depositionally, cither as a result of high humidity levels rendering the charcoal fragile and susceptible to mechanical damage, or through other, as yet undetermined means. This issue ts currently the subject of further invesuigalions by BD and HL. However. increases in deposition rates of all cultural materials during the late Holocene indicate that broader factors. are also at work. Land snail shel] deposition rates, as calculated by numbers of 2 exfineta per unit time, show that shell frequencies were low within this early Holocene period, in comparison to the Jate Holocene. This may mean that terminal Pleistocene/early Holocene conditions were dry 764 rather than wet. If this was the case, this trend would question the validity of the above interpretations of CaCO; and total land snail concentrations as evidence of a wet phase. However, we would rather argue that the low frequency of P. extincta (and other small) shells during the early Holocene is due to their very small size and their inability to frequent soggy ground surface conditions, as was probably the case at the time (see above). Ifrainfall levels were significantly higher during the early to mid Holocene as it was during the terminal Pleistocene, which appears to be the case from the pollen evidence to the east, moisture regimes may have breached the upper threshholds for very small land snails (<6mm in size). This is consistent with the predominance of large species of shells at Hay Cave during this time, including the wet-adapted species Hadra aff. bipartita. While the issue of continuous vs. intermittent deposition of sediments at Hay Cave is beyond the ability of this paper to resolve, its resolution may be important for a better understanding of palaeoenvironmental trends in the region as a whole. If in fact the sediment accumulation rate at Hay Cave is a function of the presence of Aboriginal people in the environment rather than of natural phenomena, compared to the late Holocene, the formation of only 27cm of deposit between 13,4504£150 and 3,100+60 BP would be consistent with lower levels of human occupation than subsequently. For Hay Cave, what these late Holocene sediment increases mean in terms of human behaviour remains uncertain, but it is possible that, like Mitchell River Cave, Hearth Cave and Mordor Cave also in the Mitchell- Palmer region (David & Chant, 1995), intensive occupation did not begin until the late Holocene, and before this time the caves may not have been extensively used as living sites. CONCLUSIONS This paper has focused on techniques of land snail shel] analysis and their usefulness to palaeoenvironmental reconstructions for the period spanning the Last Glacial Maximum to present in north Queensland. In the process, a wide range of themes have been covered, including: history of Aboriginal land use (archaeology); plant biogeography; geological and climatic processes; and land snail ecology and taxonomy, [rom historical as well as modern day perspectives. [tis salutary to recall here that the presence and value of molluscs in archaeological deposits was MEMOIRS OF THE QUEENSLAND MUSEUM recognised more than a century ago, and yet their detailed investigation has only recently begun in Australia. Considering the rapid development of palynology in the last 30 years, and together with the great potential of Mollusca to inform us on historical trends in both climatic and environmental change, it is perhaps rather surprising that such a state of affairs should have arisen (Evans, 1969). At present, nonetheless, most Australian studies of fossil land snails require taxonomic and ecological investigations of modern land snails to be undertaken before palaeoenvironmental research can be conducted. Once this necessity is accepted, a wealth of potential palaeoenvironmental information is available from fossil land snails. As is apparent from their abundance in buried deposits, their ability to be identified to species level, their sensitivity to moisture regimes, and their strong affinity with vegetation type, P. extincta snails have demonstrated their value as proxy indicators of palaeoenvironments in semi-arid landscapes. It is likely that both P. extincta and other species of land snails can be used for palaeoenvironmental research in other environmental zones in various parts of this continent. An existing body of regional palaeoenviron- mental evidence has provided a context for the Hay Cave land snail data. Although the quality of these data is variable, marked but variable community and taxon responses to glacial/interglacial cycling is apparent. The northeastern Queensland environment has been a dynamic and diverse system (or rather, interconnected set of systems) where the gross composition of the vegetation fluctuates in relation to cyclical changes in climate and sea level. The humid tropics region of northeastern Queensland has provided a substantial and detailed history of rainforest and rainforest- sclerophyll interactions via pollen analysis at a number of sites on the volcanic Atherton Tableland. These pollen sequences indicate that during the late Quaternary, the Tableland was subject to significant environmental variability. It has been proposed that similar changes in climate have been experienced throughout northern Queensland, although changes may not have been entirely synchronous nor of equal degree in all places, Complementing these established palaeo- environmental studies, we argue that P. extincta shell whorl counts are also dependent on moisture regimes and thus reflect fluctuations in LAND SNAIL AS AN ENVIRONMENTAL INDICATOR mean annual rainfall.Based largely on pollen research, last glacial maximum has been interpreted, independently of land snail research, as being a period of dry conditions. This was followed by a period of increased (to peak) precipitation during the early to mid Holocene. In its turn, this period of time was itself followed by an effective drying. These major changes in moisture regimes can be expected to have affected the spread, floristic structure and complexity of semi-deciduous vine thicket in north Queensland. Given P. extincta’s restricted distribution to such vegetational communities and their sensitivity to moisture, the broader applicability of our extant knowledge of palaeoenvironmental (including palaeoclimatic) trends can be tested and refined. Thus, the most important changes in the P. extincta record since 19,300 years BP are the relative increases in precipitation starting at 13,000 BP and a relative lack of whorl count variation at Hay Cave. Results at Hay Cave, to some degree, differ from those of the Atherton Tableland models based on pollen analyses, which have so far largely failed to address their relevance for regions to the west of the Great Divide. Far from constituting negative evidence, in contrast to the Atherton curves, the Hay Cave data suggests a more stable rainfall regime through time, although directions of change are consistent with the Atherton trends. As Head & Stuart (1980) concluded, this type of pattern has the potential to help quantify the limits within which climate systems operate. The lower magnitude of environmental change west of the Great Dividing Range suggested by this study now warrants further investigation. Given the current status of land snail research, multidisciplinary studies would aid in assessing the extent of this phenomenon, as well as testing the accuracy of land snail models in palaeo- environmental reconstructions. The need for the application of additional tests in such research was briefly demonstrated in this paper by a consideration of the formation of CaCO, concretions in cave sediments, and of depositional frequencies of all land snail species. A more extensive investigation of these issues may provide a better impression of past moisture levels and palaeoenvironmental factors than a consideration of P. extincta alone has been able to. Further insights into the ecology of other land snail species may also enhance our ability to model palaeoenvironments via excavated fossil land snail shells. Although good long-term data from ecological and biological information is available on P. extincta shells, there is still a need to refine our understanding of this species. In particular, P. extincta deposition rates before and after 3,100 BP raise more questions than answered here; further sediment analyses may clarify some of these questions. Similarly, the changes in P extincta shell characteristics documented here may indicate changes in human and climatic impacts on sedimentation rates within caves. Further attempts should be made to determine the influence of human activities on ecosystems within the Mitchell-Palmer region, and how people may have adjusted to environmental change in the past. In recognising the ability of vegetation to accommodate to changes in certain environmental variables, it is probable that the Chillagoe Formation vine thickets are now remnants of a more extensive Pleistocene-early Holocene flora, now represented as permanent refugia. In this sense, P. extincta highlights the relevance of, and promotes the need for, knowledge of scales of biotic changes in the recent past and for ‘dynamic’ conservation strategies to accommodate these. Tn short, the main value of this work is that the land snail data point to a reasonable but limited degree of variability in palaeoenvironmental change for hitherto unstudied parts of north Queensland. In particular, it questions the exact applicability of the Atherton curves to inland regions, while at the same time supporting established trend directions. The use of P. extincta in this project indicates that land snails represent a significant and new potential source of data for Australian palaeoecological studies. ACKNOWLEDGEMENTS We thank Kylie Stumkat for SEM work; the Queensland Museum for photography and for a Summer Scholarship for CR; Monash University for a Logan Fellowship to BD; Gary Swinton of the Department of Geography and Environ- mental Science, Monash University, for Figure |; Lana Little for assistance with fieldwork, data collection and for comments on an earlier draft of this paper; Prof. Peter Kershaw for comments on aspects of this work; Chris Clarkson for helping with the Hay Cave excavation; Earthwatch and the 1996 Earthcorps for helping with all aspects of the fieldwork; Angela Holden for sorting much of the Hay Cave material; AINSE for a radiocarbon grant to HL; Ewan Lawson and 766 ANSTO'S ANTARIS AMS laboratory for undertaking the radiocarbon dates: and Mrs Ethel Ahlers for kindly supplying the rainfall data for Bellevue Station, LITERATURE CITED AWARON, P. 1983. 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Unpublished report to the Queensland Department of Environment (Queensland Museum: Brisbane). 1999, Land snails and dry vine thickets in Queensland: using museum invertebrate collections in conservation, Pp. 257-263. In Ponder, W. & Lunney, D. (eds) The other 99%, The conservation and biodiversity of invertebrates. Transactions of the Royal Zoological Society of New South Wales. (Surrey Beatty & Sons: Chipping Norton), STANISIC, J. & INGRAM, G.J. 1998. Sandstone, snails and slaters: Pp, 112-118. In David, B. (ed.) Ngarrabullgan: geographical investigations in Djungan Country, Cape York Peninsula’ Monash Publications in Geography and Environmental Science, Number 51. (Monash University: Melbourne). STANISIC, J., EDDIE, C., HILL, A. & POTTER, D. 1994. A preliminary report on the distribution of land snails occurring within the Wet Tropics area. Unpubl. report to the Wet Tropics Management Authority. (Queensland Museum: Brisbane), STEPHANS, K. & L. HEAD, L. 1995. Palaeoecology of archaeological and swamp sites in SE Cape York Peninsula. Pp. 18-32. In Morwood, M. & Hobbs, D.R. (eds) Quinkan prehistory: the archaeology of Aboriginal art in southeast Cape York Peninsula, Australia. (Tempus 3. University of Queensland: St. Lucia). TILLIER, S, 1981. Clines, convergence and character displacement in New Caledonian diplommatinids (land prosobranchs). Malacologia 21: 177-208. TORGENSEN, T., LULY, J., DE DECKKER, P., JONES, M.R., SEARLE, D.E., CHIVAS, A.R. & ULLMAN, W.J. 1988. Late Quaternary environments of the Carpentaria Basin, Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 67: 245-261. WAINWRIGHT, GJ. & DAVIES, S.M. 1995. Balksbury Camp, Hampshire: Excavations 1973 770 and 1981, English Heritage Archaeological Report 4. (English Heritage: London). WEBB, LJ. & TRACEY. J.G. 1981. Australian rainforests: patterns and change. Pp. 605-694, In A. Keast, A. (ed.) Ecological biogeography of Australia. (Dr W. Junk: The Hague). WINTER, J.W,, BELL, F.C., PAHL, L.L. & ATHERTON, R.G. 1987. The distribution of MEMOIRS OF THE QUEENSLAND MUSEUM rainforest in north-eastern Queensland. Pp 223-227. In Werren, G. & Kershaw, A.P. (eds) The Rainforest Legacy: Australian National Rainforest Survey, Vol. 1, (Australian Government Publishing Service: Canberra). WOODWARD, B.B. 1908. Notes on the drift and underlying deposits at Newquay, Cornwall. Geology Magazine V (5): 10-18, 80-87. APPENDIX 1 Locality data tor P. extincfa from Mitchell- Palmer area. (QM reg. no., locality and habitat data, latitude/longitude). MO22998, Laura-Palmerville Rd, N of Palmerville, 1S5°48'45"S, 144°03°46"E M0O23375, Laura-Palmerville Rd, N of Palmerville, 15°53°20"S, 144°0S°20"E MOQ61172, Limestone Ck Rd, SSE 16°02°367S, 144°07°32"E MO61191, Limestone Ck Rd, SSE of Palmerville, 16°02°517S, 144°08°28"E MOQ23592, Limestone Ck Rd, SSE 16°02°53"S, 144°07°23°E MOQ61205, Limestone Ck Rd. SSE 16°03" 13S, 144°08'247°E MO61156, Limestone Ck Rd, SSE of Palmerville, 16°03722"S, 144°07°39"E MO23584, Limestone Ck Rd, SSE 16°07733"S, 144°08°47”" E MOQ23388, Limestone Ck Rd, SSE 16°08758"S, 44°09 077 E of Palmerville, of Palmerville, of Palmerville, of Palmerville, of Palmerville. Locality data for P. extincta from Chillagoe- Mungana area. (QM reg. no., locality and habitat data, latitude/longitude). MO15955, Marachoo Cave, 17°05739"S, 144°23°28"E MO25996, Carpentaria Cave, 17°05746’S, 144°23°56"E MO50231, Capricorn Cave, 17°06°15°'S, 144°24°360"E MQ53325, Chillagoe, c.13km WNW on Chillagoe- Mungana Rd. 17°06°15"S, 144°23°40"E MOS58485, Chillagoe, c.l0km WNW on Chillagoe- Mungana Rd, 17°06°45"S, 144°24’55"E MO53335, Chillagoe, c.l0km WNW on Chillagoe- Mungana Rd, 17°06"45"S, 144°24°55°E MO53551, Chillagoe, ¢.3km N on road to Metal Hills, 17°07°20°S, 144°3 P10" E MO19527, Spring Cave, 17°08703"S, 144°25°30"E MO50229, Tea Tree Cave, 17°10°33"S, 144°087 18"E MO53448, Chillagoe, c.3km SW on road to Royal Arch Caves, 17°10°35"S, 144°00 1S"E MO058540, Chillagoe, c.3km SW on road to Royal Arch Caves, 17°10°35"S, 144°00°15"E MO53436, Chillagoe, c.6km SW at Royal Arch Caves, 17117 15"S, 144°30°00"E MO038499, Chillagoe, c.8km SE on road to Almaden, 17°11?20"S, 144°33°05"E MO053564, Chillagoe, ¢.8km SE on road to Almaden, 17°11720"S, 144°33°08"E MO58547, Chillagoe, c.8km SE on road to Almaden, 17°11720"S, 144°33'0S"E OBSERVATIONS ON THE POSTCRANIAL MORPHOLOGY, ONTOGENY AND PALAEOBIOLOGY OF SCLEROCEPHALUS HAEUSERI (AMPHIBIA: ACTINODONTIDAE) FROM THE LOWER PERMIAN OF SOUTHWEST GERMANY ULLA LOHMANN AND SVEN SACHS Lohmann, U. & Sachs, 8S. 2001 06 30: Observations on the postcranial morphology, ontogeny and palaeobiology of Sclerocephalns hagusey (Amphibia: Actinodontidae) trom the Lower Permian of Southwest Germany, Memoirs of the Queensland Museum A6(2): 771-781, Brisbane. [SSN 0079-8835. The temnospondyl amphibian Sclerncephalus, in particular its postcranial anatomy, is deseribed in 4 ontogenetic stages: larval, juvenile, adult and late adult. Some specimens preserve stomach contents. consisting of paramblypterid fishes and smal) amphibians (Micromelerpeion, Aputean). In one specimen, the remains of a smal) Sclerocephalus were found. Larval and juvenile individuals probably lived in a different habitat than adult and late adult ones. Inthe juvenile, adult and lute adult stages, Sclerocephalus was the top predator ia iis environment. DO Amphibia, Linmarchia, Actinodontidue, Scleracephalus haeuseri posterapial morphology, ohtegeny, Lower Permian, Germany. Ulla Lehmann, Institure af Tropical Enviranmemal Studies and Geography, Janes Cook University, Townsville 48/1, (e-mail: ulle(@ullalahmann.cde); Sven Sachs, Queensland Museum, PO Box 3300, South Brishane 4101, dusrralia: 28 Mare 2001. Selerocephalus is one of the best-known limnarchian amphibians in the Lower Permian of Central Europe. The type species, 8. haenseri, was described by Goldfuss (1847), but the holotype is lost (Boy, 1988), Scleracephalus was described or referred to by several researchers (Ammon, 1889; Boy, 1976, 1988; Branco, 1887; Broili, 1908, 1926; Burmeister, 1850; Credner, 1893; Fritsch, 1901: Heyler, 1975; Meyer, 1857, 1858; Romer, 1947, Wemeburg, 1983, 1989, 1992), Three known species are: S. fweuseri Goldfuss, 1847.8. bavaricus Braneo, 1887 and S. jogischneideri Werneburg, 1992, Other species referred to Sclerovephalus do not belong to that taxon, e.g. ‘8.’ /atirosrris Jordan, 1849 (= Cheliderpeton Jatirastre sensu Boy, 1993), ‘8.’ labvrinthicus Geinitz, 186) (= Onchivdon labvrinthicus sensu e.g. Watson, 1919) and “S.° eredneri Fritseh, 1901 (= Onehioden labyrinthicus sensu Werneburg, 1993 and Capetus. palustris sensu Sequeira & Milner, 1993), On the other hand some larval individuals of Scleracephalus where first misinterpreted and therefore deserihed as or referred to other taxa, Romer (1939) first pointed out that Branchiosaurus ainblystomus described by Credner (1882,1885, 1886, 1893) and Ammon (1889) belonged to Sclerecephealus (Credner’s specimens are today assigned Lo Onchiodun (sensu Boy, 1990)). Later Boy (1972) recog- nised that Leptorophus levis Bulman, 1928 (= Branchiosaurus levis in Watson, 1963) and Pelosaurus longisentains Theobald, 1958 may represent larval individuals of Se/erocephalus. Selerocephalus had a wide palaeogeographical distribution (Wemeburg, 1988) and is recorded throughout the Autunian. The oldest known species is S. bayvaricus from the Altenglan Formation (Lower Autunian) of Ohmbach (Rheinland Pflalz, SW Germany), lt is represented by only one incomplete skeleton (MB Am_.442) with a well-preserved skull, The type species §. Aaeuser/ is best known and is represented by a large number of articulated skeletons, from the Saar-Nahe area in SW Germany (Rheinland Pfalz and the Saarland). Two subspecies were recognised by Boy (1988); S.h. haeuseri Goldfuss, 1847 and S. A. jecken- bachensis Boy, 1988, It is also the only species for which all ontogenetic stages are known. New specimens, showing different ontogenctic stages, have been found during the last decade, They also provide new data concerning diet. This paper characterises the stages of ontogeny of S. haeuseri and discusses its autecology. MATERIAL AND METHODS MATERIAL. PMNB uncataloged, PMNB 395, larval individuals, complete skeleton with sott issue preservation, length 8.5 and 9.5em; GMS 307, larval individual, skull and pectoral girdle, skull length |6mm; PMNB 85, juvenile, com- plete skeleton with soft tissue preservation, len- eth 13cm: PMNB 308, 177 juveniles, complete 772 skeleton with soft tissue preservation, length |1 and 12.5cm; PMNB 179, juvenile, skull and parts of the pectoral girdle, skull length 18mm; GMS 24, juvenile, skull and parts of the pectoral girdle, skull length 25mm; GMS 228, 395, juveniles, complete skeleton with soft tissue preservation, length 14 and 24cm; GMS 394, juvenile, skull and anterior half of the body with soft tissue preservation, skull length 35mm; GMS 52, juv- enile, complete skeleton with soft tissue preserv- ation, length 30cm; PMNB 174, juvenile, almost complete skeleton with soft tissue preservation, length 22cm; GMS 396, PMNB 93, juvenile, complete skeleton with soft tissue preservation, length 26cm; PMNB 103, GMS 348, PMNB 415, GMS226, PMNB GRE-1, juveniles, comp- lete skeleton with soft tissue preservation, length 25, 28, 29, 30, 48cm, respectively; PMNB uncat- alogued, juvenile, complete skeleton with soft tissue preservation, length 26cm; PMNB PDC 327, adult, complete skeleton, length 74cm; PMNB BGC 69, adult, complete skeleton, length 72cm; PMNB BGC 112, adult, complete skel- eton, length 79cm; BSPHG-1981 199, adult, skull and partial skeleton, skull length 16cm, PMNB BGC 112, late adult, complete skeleton, length 182cm. GMS P/70, adult, isolated pelvis, length ca. 10cm. Private collection SKO, adult, complete skeleton, length 82cm. REFERRED MATERIAL. About 800 coprolites from PMNB, GMS, GPIM and SKO. LOCALITIES AND AGE. All larval and most of the juvenile specimens are from Riimmelbach/ Gresaubach (Top L-O 10). One juvenile is from Gresaubach (PMNB GRE-1, Top L-O 10) and one from Worsbach (PMNB uncataloged, Top L-O 10). Adults are from Niederhausen an der Appel (PMNB PDC 327, L-O 8), Jeckenbach (PMNB BGC 69, L-O 6), Odernheim (SKO uncataloged, L-O 6) and Raumbach (SKO no Nr., L-O 6). The oldest adult was found in Jeckenbach (PMNB BGC 112, L-O 6). The isolated pelvis was found in St. Wendel (GMS P/70, L-O 5). The coprolites are from horizons Q 1 to L-O 10. In general the collecting horizons are in the Lower Rotliegendes (Autunian of European strat- igraphy sensu Boy & Fichter or Gzehlian/Assel- ian in global stratigraphy sensu Deitze, 2000). METHODS. All specimens were drawn prop- ortionally from radiographs. For smaller individuals a WILD M-3 binocular with camera lucida was used. MEMOIRS OF THE QUEENSLAND MUSEUM REPOSITORIES. BSPHG, Bayerische Staat- ssammlung fiir Palaéontologie und historische Geologie, Munich; PMNB, Pfalzmuseum fiir Naturkunde, Bad Diirkheim; GMS, Geologisches Museum der Saarberge AG, Saarbriicken; GPIM, Geologisch- Palaontologisches Institut, University of Mainz; MB, Museum fiir Naturkunde, Berlin; SKO, private collection Kratschmer, Odernheim. ABBREVIATIONS. Geological, L-O, Lauterecken-Odernheim-Formation (current stratigraphy Lauterecken-Formation L-O | + L-O 2, Jeckenbach-Odernheim-Formation L-O 3 — L-O 10 (3 = lowermost/10 = uppermost). Q Quirbach-Formation (Q1 = lower / Q2 = upper) (Dietze, 2000, fig. 1) Anatomical. Cl, clavicle; Cr, caudal rib; Ct, cleithrum; F, femur; Fi, fibula; H, humerus; I, interclavicle; Il, ilium; Is, ischium; Mc, metacarpus; P, pelvis; Ph, phalanges; Pu, processus uncinatus; R, radius; Sc, scapula; St, stomach contents; Ti, tibia; U, ulna; Va, ventral armour. SYSTEMATIC PALAEONTOLOGY AMPHIBIA Linnaeus, 1758 TEMNOSPONDYLI Zittel, 1888 LIMNARCHIA Yates & Warren, 2000 STEREOSPONDYLOMORPHA Yates & Warren, 2000 Superfamily ARCHEGOSAUROIDEA Meyer, 1857 FamilyACTINODONTIDAE Lydekker, 1885 Sclerocephalus Goldfuss, 1847 Sclerocephalus haeuseri Goldfuss, 1847 ADULT POSTCRANIAL. Vertebral column of 37-39 rhachitomous vertebrae (25 presacrals and 12-14 caudals). Neural arches high, robust, with those of the atlas-axis-complex only visible in older adults. Neural spine of the fourth cervical vertebra somewhat shorter; feature of other temnospondyls such as Eryvops (Moulton, 1974) and Balanerpeton (Milner & Sequeira, 1994), Transverse processes short but prominent, post- eroaterally directed, with large, vertical diapoph- yses. Pre- and postzygapophyses well-developed and about equal in size. Proximal articulation surface of ribs relatively broad, because of coossification of the capitulum and tuberculum. Ribs 2-14 with a prominent processus uncinatus close to their posterior margin (Fig. 5). These processus hook-like, contacting the anterior margin of the following rib. Ribs 2-4 longest. Following ribs taper ONTOGENY OF SCLEROCEPHALUS HAEUSERI 773 098 NOR, SSD. FR 77 he005 . x y i te a io a Ww y a . FIG. 1. Larval individual (PMNB uncataloged) in dorsal view. Scale-bar = lcm. slightly in length back to 14-15 vertebra and then rapidly to the pelvis. Shoulder girdle of inter- clavicle (which is the central ventral element) and paired clavicles, scapulocoracoid and cleithrum. Interclavicle a relatively flat bone of rhombic shape; ventral surface with numerous cristae and furrows, running from the edges to the centre. Clavicles flat, triangular, medially curved in their ventral section, broadly contact the interclavicle; dorsal process with posteriorly directed tip contacting the ventral process of the cleithra; ventrolateral margin also with some furrows. Scapulocoracoids the most prominent bones of the shoulder girdle. Shoulder blade for- med as a broad, posteromedially directed, dorsal process, articulating with the clavicle antero- ventrally and with the cleithrum anterodorsally. Coracoid part of the scapulocoracoid ventrally on the posteroventral margin. Prominent glenoid surface in the posterior margin. Large supraglenoid foramen above the glenoid surface. Cleithra flat, arch-shaped, of triangular outline, FIG. 2. Gill-teeth (“Kiemenzahne’), which were attached at the ceratobranchial filaments. Scale-bar = Imm contacting the scapula blade with their postero- medial margin, running ventrally out in a sharp process meeting the dorsal process of the clavicle in about the anterior mid-section of the scapula blade. Humerus robust, moderately elongated, with tetrahedal shape (Meckert, 1993 sensu Romer, 1939) similar to the humerus of Ervops (Miner, 1925);.proximal head prominently convex, above the processus latissiumus dorsi and the crista pectoralis posteriorly and the crista dorsalis anteriorly; distal surface anteroposteriorly expanded, bearing a relatively small processus supinator and somewhat larger ectepicondylus anteriorly, as well as a very prominent entepicondylus posteriorly. Radius and ulna relatively short elements; radius more robust, with slightly widened ends, with concave shaft; ulna more slender, with proximal part bearing the olecranom widened somewhat medially directed; olecranom very prominent, only fully ossified in older adults. Manus well ossified in older adults, described in detail by Meckert, 1993. Phalangeal formula 2/2/3/3. On the ventral part of the body, between the interclavicle and the pelvis, there is a compact ventral armour of epidermal scales. Boy (1988) and Broili (1926) described this armour in detail. Scales generally long and sharp in the midsection and oval or circular laterally. At the level of the forelimbs, there is a prominent recess on each side of the armour. Ossification of these scales had already started in the late larval stage, in individuals with a skull length of approximately 20mm (Boy & Sues, 2000). Pelvis very robust; ilium its most prominent element; ventral part fan-shaped anteropost- eriorly bearing most of the acetabulum; dorsal part elongate-rectangular, curved posteriorly. 774 FIG. 3. Juvenile individual (PMNB uncataloged) in ventral view. In this specimen, scales of paramblypterids are visible as stomach contents (St). Scale-bar=lcm. The ventral section of the pelvic-girdle consists of the pubis and ischium, which together represent a somewhat triangular shaped plate. Hind-limbs well-developed in adults. Femur massive, rectangular, clearly the largest element of the limbs; proximal end broadened and somewhat more prominent than the distal one. Fourth trochanter weakly developed, situated proximally, with distal half of shaft still robust, terminating in a relatively straight articulation surface. Tibia and fibula much shorter than the femur; fibula somewhat shorter than tibia. Tibia less robust, with slender distal portion, contacting MEMOIRS OF THE QUEENSLAND MUSEUM the tibiale ventrally and the intermedium medially. Fibula massive, with expanded distal end with a medially directed tip. This tip touching the intermedium, while the medial margin contacts the fibulare. Metatarsals only visible in older adults. Tibiale, intermedium and fibulare very well-developed. Tibiale triang- ular, situated laterally, smallest of proximal metacarpals. Inter- medium oval, between tibiale and fibulare; fibulare situated med- ially, very prominent, roughly tri- angular. Phalanges rod-like, terminate in a claw-shaped phalanx. Phalangeal formula varies between 2/2/3/3/3 and 2/2/2/3/3. ONTOGENY Four ontogenetic stages (larval, juvenile, adult and late adult) are characterised by features of the postcranial skeleton. The pectoral girdle has already been described by Meckert (1993) and the cranium by Boy (1988). LARVAL STAGE (skull length 0.8-2.5cm). The basis for larval anatomy is an 8cm long specimen (PMNB uncataloged) (Fig.1) with well-preserved body outlines and carbonaceous imprints of the external gill. The basibranchiale was probably cartilaginous. It ossifies in the juvenile stage and is prominently developed in adults (Boy, 1972, 1988; Boy & Sues, 2000). The ceratobranchials were arranged in up to 4 rows and situated laterally at the level of the shoulder girdle. They probably also consisted of cartilage, but had attached to them small bony plates, so-called gill-teeth (‘Kiemenzihne’ after Boy, 1972) (Fig. 2). Each gill-tooth had approx- imately 5 spine-like denticles, which vary by 1-2 spines. Similar structures are known in Micromelerpeton (Boy, 1995) and Gerrothorax (Nilsson, 1946). The hypobranchials, hypohyals and ceratohyals were not preserved. The limbs are conspicuous but only slightly ossified in the Bad Diirkheim specimen. The ONTOGENY OF SCLEROCEPHALUS HAEUSERI humerus is very short and nearly quadratic; the radius and ulna are weakly developed, but the radius is longer. Elements of the manus are not recorded and were probably cartilaginous. The femur is rectangular in lateral view, relatively short, very robust and longer than the less developed tibia and fibula. Elements of the pes are badly preserved. The centra are rhachitomous and bear paired neural arches with low neural spines (Boy, 1972). Ribs of vertebrae 1-14 are ossified, rod-like and slightly broadened proximally. Capitulum and tuberculum not clearly developed. The sacral rib is slender. Caudal ribs have not been recorded. The pelvis is relatively weak and not as robust as the well-developed shoulder-girdle (Meckert, 1993). The tail occupies approximately 50% of body length. No stomach contents could be found. JUVENILE STAGE (skull length 3.5-7.5cem). Representative juvenile isa 26cm long specimen with clearly visible body-outlines (PMNB uncataloged) (Fig. 3). The external gill has been lost and limbs are more ossified. The humerus shows no major differences to that in the larval stage. Radius and ulna are about as long as the humerus. Radius and ulna are approximately equal in length, but the radius is somewhat more robust. The femur is elongate-rectangular with slightly expanded proximal and distal ends, It is clearly more robust and about twice as large as the tibia and fibula. The tibia is rod-shaped and slightly expanded distally. The metapodials are not ossified; phalanges are clearly visible. Claw- like terminal phalanges are developed. GMS 394 shows that the vertebrae are well-ossified and bear lower neural arches and spines as in adults. The ribs are more robust than in the larval individual. The 2nd to 14th presacral ribs are club-like and broadened laterally. Between |2 and 14 caudal ribs are presented. The second and third ribs, distal to the sacral rib, are longest. The 775 FIG, 4. Adult individual (PMNB PDC-327) in dorsal view. Scale-bar = Sem. tail occupies about 45% of body length, but is shorter than in larval individuals. ADULT STAGE (skull length 9.5-16cm). The representative adult is a well-preserved skeleton 74cm long (PMNB-PDC 327) (Fig. 4). In general, the body has become somewhat more compact. The humerus is robust, but not as stout as in juveniles. Its proximal and distal ends are still cartilaginous (Broili, 1926; Boy, 1988). Ulna and radius are about equal in length, but the radius is more robust. In both, the shaft is inwardly curved, with the ulna showing a more prominent inflexion.Phalanges are broader than 776 MEMOIRS OF THE QUEENSLAND MUSEUM Cl +I? FIG, 5. Late adult individual (PMNB BGC-1 12) in lateral view. In this stage the ventral armour (Va) is clearly visible. Scale-bar = 20cm. in juveniles and still not completely ossified. Vertebrae are well-developed. The pleuro- Also the metacarpus is not completely ossified. centra are slender and of rhachitomous shape The hind-limb elements are similar in shape to (Schoch, 1999: 107). Boy (1988) described the those in the juvenile specimens, but are more _ intercentra as unpaired with a low semicircular robust. Metatarsals are present. shape and the neural arches as robust and ONTOGENY OF SCLEROCEPHALUS HAEUSERI < Insects Zooplankton \\ 777 an Aeduella aa win exe Paramblypterus SA Nanoplankton “——____* Larval — Juvenile a el Adult / Late Adult “as Malacostraca Ne” Micromelerpeton —— Apateon FIG. 6. Proposed food chain of Sclerocephalus in the larval, juvenile and adult/late adult stage. relatively high. This could also be observed on PMNB-PDC 327. All ribs are strongly developed and have a co-ossified capitulum and tuberculum. This is especially visible in ribs 2-14 where the proximal ends are broadened into a club-like shape. These ribs bear a hook-like processus uncinatus posteriorly. In large individuals, there are lateral depressions on ribs 2-6. Pelvic- and shoulder- girdles are robust and the pelvis is now completely ossified. The tail is shorter than in juveniles. It occupies about 40% of body-length. LATE ADULT STAGE (skull length >18cm). The late adult stage is based on PMNB-BGC 112 (Fig. 5). In general the body is shorter than in earlier stages and all bones are well ossified. Humeri are robust and slightly elongated. Metapodials are completely ossified. Of the atlas-axis-complex only the neural arches are visible. Tuberosities on the proximal ends of the ribs change from being club- to bow-shaped and can be observed to rib 7 (but possibly extended to 9). Also the lateral depressions, mentioned for the adults, can be recorded to ribs 7-9, and are deeper. The pelvis is extremely robust. The tail is shorter then in earlier stages, but does not differ much from that of adults (it also occupies 40% of body-length). HABITAT RECONSTRUCTION During the Lower Permian, the Saar-Nahe Basin in SW Germany was traversed by several river systems. In some areas water was trapped in low relief, forming lakes (Dietze, 1999). These lakes (Table 1) normally existed only for a relatively short geological period, but contained a well-balanced ecosystem (palaeocommunities after Boy, 1998 sensu Jarvinnen et al., 1986), composed of large amphibians (e.g. the limnachian Archegosaurus), small amphibians (e.g. the branchiosaurs Micromelerpeton and Apateon), large fishes (e.g. the xenacanthodian freshwater sharks Xenacanthus and Orthacanthus), small fishes (e.g. the amblypterid Paramblypterus or the acanthodian Acanthodes), as well as a variety of different invertebrates (e.g. ostracods, bivalves, shrimps) (Boy, 1998). One of the larger lakes was Lake Humberg (L-O 10) that extended over an area of 3,400km* and deposited nearly the whole Saar-Nahe Basin (Stapf, 1990). Boy (1994) mentioned, that lake deposits from Lake Humberg can be divided into 4 phases. TABLE 1. Distribution of the ontogenetic stages of Sclerocephalus haeuseri in different lake localities (primary based on unpublished excavation reports of the Pfalzmuseum fiir Naturkunde, Bad Diirkheim). a ea a sw & | Lake-size r. , Late Layer | Lake | (km?) [pars | Tuy. | Adult | Adult | | |Riimmelbach | |. | | | 2 10/SHumberg. c= 3400) 14 |ca100) 0 o- L-O 9 | Ruthweiler cal0 | 21 | 0 | 0 07 L-O 8 | Odernheim ca. 760 | 31 (ca.l00, 2 | 0 5 Jeckenbach- sak (Beak 7 Heimkirchen | ° 230 0 | 4 ca.50) | 8 ee st — — i 06 |Niederkirchen, ca.40 | 0 | 2, 2. 2 coz i | wees thee, = -_ [Q2 Quirnbach _ | ca.500 | 0 | 0 0 1 (QL |StWendel_| ca. 40 go | 2 | 20 | 3 778 Sclerocephalus is found (together with Para- mblypterus uvernoyi and Apateon pedestris) only in the first phase, when the Lake was deepest. In other lakes, such as in Lake Odernheim (L-O 8), Sclerocephalus was found together with the large branchiosaur Micromelerpeton (Boy, 1994) or with the small freshwater shark Triodius (Lake Klauswald, L-O 9, after Dietze, 1999). Boy (1988) concluded that juvenile Sc/ero- cephalus individuals might have lived in shallow lakes. We suggest that juvenile Sc/erocephalus lived in large, relatively deep lakes of the Rtimmelbach-Humberg type. Recent extensive excavations by the Pfalzmuseum fiir Naturkunde in Bad Diirkheim in the corresponding localities Gresaubach/Lebach and Humberg/Odernheim (Top L-O 10) support this suggestion as only larval and juvenile individuals have been found. It therefore is suggested that these lakes were being used as spawning grounds whilst adult individuals lived in another environment. Also, this could explain why, in lakes where adult and late adult individuals are common, as in Lake Jeckenbach (L-O 6) (Stapf, 1990), larval and juvenile individuals are rare or absent. DIET DURING ONTOGENY As mentioned above, no stomach contents could be associated with the larval specimens. Most probably their diet consisted of plankton and malacostracans (as the common Lronectes) or maybe small insects. Boy (1993) suggested that larval Sc/erocephalus, similar to some recent amphibians could have caught its prey with a suck-snap method (Bramble & Wake, 1985). Juveniles caught large prey. As mentioned above, a complete, large Paramblypterus with its head folded backwards was found in the stomach of PMNB (uncataloged). Altogether 6 fish-bearing specimens were observed (only one in a public collection, PMNB, uncataloged, fig. 3); another one (GPIM-N 1166) was mentioned by Boy (1988). We therefore conclude that Sclero- cephalus swallowed these fishes whole. In numerous specimens, scales of paramblypterids, and also traces of small amphibians such as Apateon and Micromelerpeton, have been found. As a rarity, the stomach of 1 specimen (GMS, uncataloged) contained remains of a smaller Sclerocephalus. This is the first record of cannibalism in this taxon. Adult and late adult individuals were primary piscivores, in which Paramblypterus represents the common prey. Observations of stomach contents and a large number of coprolites showed MEMOIRS OF THE QUEENSLAND MUSEUM that smaller amphibians and probably smaller Sclerocephalus individuals also belonged to the prey. Acanthodians were never present as stomach contents or in the coprolites. These fishes were very common in the environment of Sclerocephalus and of moderate size, but had large spines anterior to their fins. It is possible that the more heavily built adult and late adult Sclerocephalus individuals mostly did not hunt actively, but watched for prey next to the shore and caught them with the suck-snap method. If we accept this premise, Paramblypterus and small amphibians would have been an easier prey than the spine-bearing acanthodians. CONCLUSIONS In the first ontogenetic phases, Sc/erocephalus haeuseri was well adapted to its aquatic habit, FIG. 7. ldealised reconstruction of the four ontogenetic stages: A, larval; B, juvenile; C, adult; and D, late adult. The right side of the body shows the dorsal bones, the left side shows the ventral elements. ONTOGENY OF SCLEROCEPHALUS HAEUSERI which is especially perceptible in the slender shape of the body, the weakly ossified limbs and the long rudder-tail. Larval individuals are up to 10cm in length. Their jaws were not very strong and bore only slightly developed teeth (Boy, 1988); therefore their diet must have consisted of plankton and probably insects or malacostracans. In the juvenile stage, individuals could reach to 50cm long. They still had a slender body-shape and a long tail and probably were active hunters. Stomach contents and coprolites contain remains of paramblypterids and small amphibians, including smaller individuals of Sclerocephalus. The prey was swallowed entirely. It is conspicuous that some lakes, especially those of the ‘Riimmelbach-Humberg-Lake’ type, nearly exclusively yielded larval and juvenile individuals (up to the late juvenile stage (= late ‘metamorphic’ stage after Boy & Sues, 2000) or subadult stage (Table 1). We therefore conclude that adult S. haewuseri might have changed their habit from fully aquatic to amphibious. Adults probably visited the habitat of the younger ones to spawn; therefore they are rarely recorded in these layers. According to Boy (1998) both juvenile and adult/ late adult individuals were the so-called top-predators in their environment (Fig. 6). In Lake Niederkirchen Sclerocephalus shared this position with the freshwater shark Orthacanthus senckenbergensis. In Lake Humberg, as mentioned above, Sclerocephalus was only recorded in the first of 4 phases. In the second phase the position as top-predator was held by the freshwater shark Xenacanthus meisenheimensis (Boy, 1994), while in the fourth phase the top-predator position was shared by the archegosaurids Archegosaurus and Cheliderpeton. The latter is the closest relative to Sclerocephalus (Yates & Warren, 2000) and is also known in different ontogenetic stages (Boy, 1993; Steyer, 2000). In the adult/late adult stage, Sclerocephalus specialised on paramblypterids, but also caught smaller amphibians such as Apateon or Micromelerpeton, Although Acanthodes was very common in the lakes, it was not recorded in the stomach contents or coprolites and therefore probably was not preyed upon. The limbs were well ossified in the adult stage, so that these individuals also could have been terrestrial from time to time. ACKNOWLEDGEMENTS UL is grateful to Bernd Graumann (formerly Pfalzmuseum fiir Naturkunde, Bad Diirkheim), 779 Klaus Kratschmer (Odernheim), Rainer Schoch (Staatliches Museum fiir Naturkunde, Stuttgart), and Jiirgen Boy, Karl Stapf and Jahn Hornung (University of Mainz) for fruitful discussions and access to the specimens under their care. SS thanks Scott Hocknull (Queensland Museum) and Anne Warren (La Trobe University, Melb- ourne) for helpful discussions during writing and for reviewing and commenting upon the manuscript. He is also grateful to James Cox (Brisbane) for making the drawings of Fig. 6. LITERATURE CITED AMMON, L. v. 1889. Die permischen Amphibien der Rheinpfalz. (Straub Verlag: Munich), BOY, J. A. 1972. Die Branchiosaurier (Amphibia) des saarpfilzischen Rotliegenden (Perm, SW-Deutschland). Abhandlungen des hessischen Landesamtes fiir Bodenforschung 65: |-137. 1974, Die Larven der rhachitomen Amphibien (Amphibia: Temnospondyli; Karbon-Trias). Paldontologische Zeitschrift 48: 236-268. 1976. Uberblick tiber die Fauna des saarpfalzischen Rotliegenden (Unter-Perm). Mainzer geowissenschaftliche Mitteilungen 5: 13-85. 1988. Uber einige Vertreter der Eryopidea (Amphibia: Temnospondyli) aus dem europaischen Rotliegend (?héchstes Karbon — Perm) 1. Sclerocephalus. Palaiontologische Zeitschrift 62(1/2): 107-132. 1990. Uber einige Vertreter der Eryopidea (Amphibia: Temnospondyli) aus dem europaischen Rotliegend (?héchstes Karbon — Perm) 3. Onchiodon. Palaontologische Zeitschrift 64(3/4): 287-312. 1993. Uber einige Vertreter der Eryopidea (Amphibia: Temnospondyli) aus dem europdischen Rotliegend (?héchstes Karbon — Perm) 4. Cheliderpeton latirostre. Palaiont- ologische Zeitschrift 62(1/2): 123-143. 1994. Seen der Rotliegend-Zeit — Ein Lebensraum vor rund 300 Millionen Jahren in der Pfalz. Pp. 107-116. In Koenigswald, W.v. & Meyer, W. (eds) Erdgeschichte im Rheinland. (Verlag Dr Pfeil: Munich). 1995, Uber die Micromelerpetontidae (Amphibia: Temnospondyli). 1. Morphologie und Paliiodkologie des Micromelerpeton credneri (Unter-Perm; SW-Deutschland). Palaontologische Zeitschaft 69: 429-457, . 1998. Méglichkeiten und Grenzen einer Okosystem- Rekonstruktion am Beispiel des spatpalaozoischen lakustrinen Paldo-Okosystems. Paléontologische Zeitschrift 72(1/2): 207-240 BOY, J.A. & FICHTER, J. 1982. Zur Straigraphie des saarpfalzischen Rotliegenden (?Ober-Karbon — Unter-Perm; SW-Deutschland), Zeitschrift der deutschen geologischen Gesellschaft 133: 607-642. BOY, J.A. & SUES, H.-D. 2000. Branchiosaurs: Larvae, metamorphosis, and heterochrony in 780 temnospondy! and seymouriamorph amphibians. Pp. 1150-1197, In Heatwole, H. & Carroll, R.L. (eds) Amphibian Biology Vol. 4 (Surrey Beatty & Sons: Chipping Norton). BRAMBLE, D.M. & WAKE, D.B. 1985. Feeding mechanics of lower tetrapods. Pp. 230-261. In Hildebrand, M., Bramble, D.M., Liem, K.F. & Wake, D.B. (eds) Functional Vertebrate Morphology (Belknap Press: Cambridge). BRANCO, W. 1887. Weissia bavarica g. n. sp. n., ein neuer Stegocephale aus dem unteren Rothliegenden. Jahrbuch der kéniglisch Preussischen geologischen Landesanstalt und Bergakademie Berlin 1886: 22-39. BROILI, F. 1908. Ueber Sclerocephalus aus der Gaskohle von Ntirschan und das Alter dieser Ablagerungen, Jahrbuch der kaiserlisch kéniglischen geologischen Reichsanstalt Wien 58: 49-70. 1926. Uber Selerocephalus Hduseri GOLDFUSS. Sitzungsberichte der Bayerischen Akademie der Wissenschaften, Mathematisch-naturwissen- schaftliche Abteilung 1926: 199-222. BULMAN, O.M. 1928. Additional notes on some branchiosaurs from Odernheim. Annals and Magazine of Natural History 10: 250-255. BURMEISTER, H. 1850. Die Labyrinthodonten aus dem Saarbriicker Steinkohlengebirge. (Berlin) CREDNER, H. 1882. Die Stegocephalen und Saurier aus dem Rothliegenden des Plauen’ schen Grundes bei Dresden. III. Theil, Zeitschrift der deutschen geologischen Gesellschaft 34: 213-237. 1885. Die Stegocephalen und Saurier aus dem Rothliegenden des Plauen’ schen Grundes bei Dresden. V. Theil. Zeitschrift der deutschen geologischen Gesellschaft 37: 694-736. 1886. Die Stegocephalen und Saurier aus dem Rothliegenden des Plauen’ schen Grundes bei Dresden, VI. Theil. Zeitschrift der deutschen geologischen Gesellschaft 38: 576-632. 1893. Die Stegocephalen und Saurier aus dem Rothliegenden des Plauen’ schen Grundes bei Dresden. X. Theil. Zeitschrift der deutschen geologischen Gesellschaft 45: 639-704. DIETZE, K. 1999. 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Bulletin Trimestriel de la Société d’Histoire Naturelle et des Amis du Museum __d@’Autun 75: 15-27. JARVINNEN, O., BABIN, C., BAMBACH, R., FLUGEL, E., FURISCH, F., FUTUYMA, D., NIKLAS, K., PANCHEN, A., SIMBERLOFF, D., UNDERWOOD, A. & WEIDICH, K. 1986. The neontologico-paleontological interface of community evolution: How do the pieces in the kaleidoscopic biosphere move? Pp, 331-350. In Raup. D.M. & Jablonski, D. (eds) Patterns and processes in the history of life (Springer-Verlag: Berlin). JORDAN, H. 1849. Ergiinzende Beobachtungen zu der Abhandlung yon GOLDFUSS iiber die Gattung Archegosaurus. Verhandlungen des naturhistorischen Vereins fiir Rheinlande und Westphalen 6: 76-81. MECKERT, D. 1993. Der Schultergiirtel des Sclerocephalus haeuseri GOLDFUSS, 1847 im Vergleich mit Eryops COPE, 1877 (Eryopidea, Amphibia, Perm). Palaeontographica (A)229: 113-140, MEYER, H. v. 1857. Reptilien aus der Steinkohlen- Formation in Deutschland. Palaeontographica 6: 59-220. 1858. Nachtrag zu den Reptilien aus der Steinkohlen-Formation in Deutschland, insbesondere zu Archegosaurus latirostris. Palaeontographica 6: 219-220. MINER, R.W. 1925. The pectoral limb of Eryops and other primitive tetrapods. Bulletin of the American Museum of Natural History 51: 145-312. MILNER, A.R. & SEQUEIRA, S.E.K. 1994, The temnospondy! amphibians from the Viséan of East Kirkton, West Lothian, Scotland. Transactions of the Royal Society of Edinburgh: Earth Sciences 84: 331-361. MOULTON, J.M. 1974. A description of the vertebral column of Eryops based on the notes and drawings of A.S. Romer. Breviora, 428: 1-44. NILSSON, T. 1946. A new find of Gerrothorax rhaeticus Nilsson, a plagiosaurid from the Rhaetic of Scania. Lunds Universitet Arsskrift, N.F. 42: 1-42. ROMER, A.S. 1939. Notes on branchiosaurs. American Journal of Sciences 237: 748-761. 1947, Review of the Labyrinthodontia. Bulletin of the Museum of Comparative Zoology, Geological Series 99; 1-368. SEQUEIRA, S.E. & MILNER, A. 1993. The temnospondly amphibian Capetus from the Upper Carboniferous of Czech Republic. Palaeontology 36(3): 657-680. ONTOGENY OF SCLEROCEPHALUS HAEUSERI SCHOCH, R.R. 1999. Comparative osteology of Mastodonsaurus giganteus (JAEGER, 1828) from the Middle Triassic (Lettenkeuper: Longobardian) of Germany (Baden- Wiirttemberg, Bayern, Thiiringen). Stuttgarter Beitrige zur Naturkunde (B) 278: 1-175. STAPF, K.R. 1990. Fazies und Verbreitung lakustriner Systeme im Rotliegend des Saar-Nahe-Beckens (SW-Deutschland). Mainzer geowissen- schaftliche Mitteilungen 19: 213-234. STEYER, J.-S. 2000. Ontogeny and phylogeny in temnospondyls: a new method of analysis. Zoological Journal of the Linnean Society 130: 449-467. THEOBALD, N. 1958. Contribution a l’étude des stégocéphaes du Permien inférieur de la Sarre. Annales Universitatis Saraviensis, Natur- wissenschaftliche Rethe (7)2: 192-210. WATSON, D.MLS, 1919. The structure, evolution and origin of the Amphibia. — The ‘order’ Rhachitomi and Stereospondyli. Philosophical Transactions of the Royal Society of London (B)209; 1-73. 1963. On growth stages in branchiosaurs. Palaeontology 6: 540-553, WERNEBURG R. 1983. Stegocephalen (Rhachitomi, Amphibia) aus dem hohen Unterrotliegenden 78] (Oberhéfer Schichten) des Thiirmger Waldes. Freiberger Forschungshette (C)384: 111-129. 1988. Paléobiogeographie der labyrinthodonten Amphibien im Oberkarbon und Rotliegenden Mitteleuropas. Zeitschrift der geologischen Wissenschaften 16: 929-932. 1989, Labyrinthodontier (Amphibia) aus dem Oberkarbon und Unterperm Mitteleuropas — Systematik, Phylogenie und Biostratigraphie. Freiberger Forschungshefte, Reihe (C)436: 7-57. 1992, Sclerocephalus jogischneideri n. sp. (Eryopidea, Amphibia) aus dem Unter- rotliegenden (Unterperm) des Thiiringer Waldes, Freiberger Forschungshefte (C)445; 29-48. 1993. Onchiodon (Eryopidae, Amphibia) aus dem Rotliegend des Innersudetischen Beckens (Béhmen). Paléontologische Zeitschrift 67(3/4): 343-355, YATES, A.M. & WARREN, A.A. 2000. The phylogeny of the ‘higher’ temnospondyls (Vertebrata: Choanata) and its implications for the monophyly and origins of the Stereospondyli. Zoological Journal of the Linnean Society 128: 77-121. A NEW SPECIES OF CALAMOECIA (COPEPODA: CALANOIDA) FROM ARID AUSTRALIA, WITH COMMENTS ON THE CALANOID COPEPODS OF THE PAROO, NORTHWESTERN MURRAY-DARLING BASIN BRIAN V. TIMMS Timms, B.V. 2001 06 30: A new species of Calamoecia (Copepoda: Calanoida) from arid Australia, with comments on the calanoid copepods of the Paroo, northwestern Murray-Darling Basin. Memoirs of the Queensland Museum 46(2): 783-790. Brisbane. ISSN 0079-8835. Calamoecia baylyi sp.nov. is described from claypans in the Paroo catchment in southwestern Qld and northwestern NSW. It also occurs in WA where it was previously known as the Cue form of C. /ucasi. Ten species of calanoid copepod occur in the Paroo with Boeckella triarticulata, C. canberra and C. lucasi common, B. fluvialis and C. zeidleri present and B. robusta robusta, B. timmsi, C. baylvi, Diaptomus lumholtzi and Gladioferens spinosus uncommon. Co-occurrences are common, especially in claypans and riverine waterholes. 1 New species, Calamoecia, Calanoid copepods, environmental ecology, biogeography. Brian V. Timms, School of Geosciences, University of Newcastle, Callaghan 2308, Australia (e-mail: ggbvt(@alinga.newcastle.edu.au); 28 March 2001. Calamoecia Brady contains 16 species (Bayly, 1992, 1998; Halse & McRae, 2001). Two of the most recent additions, C. zeidleri Bayly and C. halsei Bayly live in waters in remote parts of arid Australia. Also occupying some of these waters is a form similar to C. /ucasi Brady but different enough to be recognised as the Cue form (Bayly, 1984). It has been reported from Cue (Bayly, 1984), near Port Hedland (Timms & Morton, 1988), and the Gascoyne-Murchison (Bayly, 1998), all in NW WA. Although Bayly (1998) thought this western form of C. /ucasi was anew species, he was reticient to proceed partly because it is allopatric with C. /ucasi s. s. (LLA.E. Bayly, pers. comm.). C. /ucasi s.s. is widespread in eastern and central Australia (Maly & Bayly, 1991), but records of C. /ucasi in WA in Maly & Bayly (1991) and in Maly et al. (1997) refer to its western form (alias Cue form) (Maly & Bayly, 1991, and S. Halse, pers. comm.). However C. lucasi s.s. occurs at Lake Gregory, S Kimberley (Halse et al., 1998) and another variant of C. lucasi, the ‘Lake Grace form’ lives near Lake Grace in SW WA. Discovery of populations similar to the Cue form in the Paroo, NW NSW and SW Qld, and sympatric with C. /ucasi s.s. confirmed that the Cue form is a separate species. Not only are there many differences between it and C. lucasi s.s., but the Cue form has maintained its distinct- iveness despite chances for interbreeeding. The Paroo area contains 9 other calanoid cope- pods including many inland forms for which few ecological data are available. Ecological and biogeographic studies on Australian calanoid copepods (Bayly, 1996; Maly, 1984; Maly & Bayly, 1991; Maly & Maly, 1997; Maly et al., 1997; Timms & Morton, 1988; Walsh & Tyler, 1998) have not included data from the eastern arid zone. The aims of this paper are to describe the Cue form of Calamoecia lucasi s./. and to comment on ecological biogeography of calanoid cope- pods in the Paroo catchment of the NW Murray- Darling Basin. METHODS Specimens were measured under a Wild M3C stereomicroscope fitted with an eyepiece micrometer, stained with Chorazol Black, and dissected with tungsten needles in DePeX mountant on a microslide under the same microscope. Appendages were studied and drawn using a Olympus BHA microscope fitted with Olympus drawing attachment. Terminology and abbreviations used in describing the appendages follow Bayly (1992). Ecological data on the copepods of the Paroo are gleaned from my monitoring studies of 100+ wetland sites over 13 years (Timms 1993, 1997a, 1997b, 1998; unpubl. data). The area covered ranges from Currawinya National Park in the N (centred on 144°25’E, 28°50’S) to Lake Peery in the Overflow lakes area (143°37’E, 30°45’S) and encompassing about two-thirds of the Paroo’s catchment of 73, 600km’. 784 SYSTEMATICS Class CRUSTACEA Order COPEPODA Family CENTROPAGIDAE Giesbrecht Calamoecia Brady Calamoecia baylyi sp. nov. (Figs 1, 2) Calamoecia lucas; Brady; Bayly, 1984 (in part, the Cue population, fig. 4A-D). ETYMOLOGY. In honour of Dr Ian A.E. Bayly, to whom so much is owed on the taxonomy and ecology of centropagid copepods. MATERIAL, HOLOTYPE ¢ QMW25483, length 0.77mm prosome, 1.19mm total, ALLOTYPE 2 QM W25484, length 1.10mm prosome, 1.52mm total, both mounted on microslides in DePeX., PARATYPES 30 ds, 20 2s QMW25485, Queensland Museum. Collected by the author on 17 May, 1996. OTHER MATERIAL. 30 ds, 15 9s QM W25487, Turkey Pan, 29°33'S, 144°49°E, Bloodwood Station, 130km NW of Bourke, NSW.collected by the author, 19 May, 1998; 5 ds, 39s, Melaleuca Pan, 29°33’S, 144°48°E, Bloodwood Station, NSW, collected by author 19 May 1998; a claypan near S end of Lake Wyara, Currawinya National Park, 28°48’S, 144°15°E, Qld, collected by author 14 May, 1996; dissected ¢ WAM C24994 and vial of 10 ds and 10 9s WAM C24996 from an unnamed claypan, 24°47°35"S, 114°09°14”"E (CB 58e of Halse et al., 2000), collected S.A. Halse 25 July 1995; dissected 2 WAM C24995 from Tirigie Claypan, 24°38°34"S, 113°59°29”E, collected by S. Halse 17 August, 1994. TYPE LOCALITY. A claypan 1.5km NE of Coomburra Waterhole, Currawinya National Park, 28°47°E, 144°22°E, SW Queensland. DESCRIPTION, Male. On first legs, outer edge spines on exopodite promixal segment (Rel ) and distal segment (Re3) with abnormally large secondary spinules (same in females [Fig. 1A]). On fifth leg (Fig. 1B,C) distal basidopodite segment (B2) with a distinct space between insertion of exopodite and endopodite, which in the right limb is wider and a little indented. Proximal segment of right exopodite (Rel) somewhat cuboidal with a spine on its outer distal corner and a small rounded protrusion on its inner distal corner. Middle distal segment (Re2) witha prominent rounded projection midway on its inner surface. Claw (Re3) strongly bent about one-quarter way along its length, and lacking an inner spine. Right endopodite (Ri) 3-segmented, proximal (Ril) and middle segment (Ri2) round-oval shaped, but distal segment (Ri3) more elongated and bearing 2 long setae subterminally. Left exopod (Re)1-segmented with a small spine MEMOIRS OF THE QUEENSLAND MUSEUM two-thirds along outer margin and tip expanded into rounded spade-like structure with denticles on and towards its margin. Base of this structure with short spine on a protuberance on the inner side and longer spine next to it. Left endopodite (Ri)l-segmented, a little longer than the first 2 segments of the right exopodite (1.e. exopodite without its claw), with almost parallel sides. This segment with 5 spines (2 terminal, 3 subterminal) decreasing almost uniformally in length from the inner spine to outermost one. Female. On first legs (Fig. 1A), spine on outer edge of proximal segment (Rel) and 2 spines on outer edge of distal segment (Re3) of exopodite with abnormally large secondary spinules (as in 3 ). These outer spines absent on exopods of legs 2-4, On fifth leg (Fig. 1D), proximal exopodite segment (Rel) with an outer spine with secondary spinules. Middle exopodite segment (Re2) also with such a spine and a large curved slender outgrowth on its distal inner corner. This outgrowth with 2 rows of spinules. Distal exo- podite segment (Re3) with 5 or 6 spines, outer- most with secondary spinules, next inner one longest but still a little shorter than the segment. Curved outgrowth on Re2 subequal to Re3 plus its spines. Endopodite (Ri) 1-segmented, with 7 or 8 long setae (4 or 5 on inner side, | terminal, and 2 on outer edge.). Prosome terminating in elongated lobes on either side of prosome- urosome junction (Fig. 1E,F). Urosome (Fig. 1E,F) with genital segment about 1.5 x as long as wide. Genital operculum with a distinct nipple and on a large mound. Left side of genital segment with a bulge almost halfway along its length, but no indication ofan associated furrow. REMARKS. Variation between populations. Specimens from the Carnarvon region of WA are slightly different from the type material in a number of features. On the 6 fifth leg, the spine of the outer distal corner of the first exopodite segment (Rel) is much longer (Fig. 2B). The claw of the right exopodite (Re3) is markedly bent further along its length than in type material (ca one-third to halfway along its length). The right endopodite is curved inwards and the first and second segment (Ril, Ri2) even more bulbous so that especially the second segment presents a semicircular outer edge (Fig 2B). In the ° the most noticeable difference is on the third segment of the exopodite (Re3) where spines (6 instead of 5) are longer than in the type material, so that they exceed the length of the segment and of the curved inner process of the NEW CALAMOECIA FROM THE MURRAY-DARLING BASIN second segment (Re2) (Fig. 2A). Bayly (1998) thought different material he examined from_ the Carnarvon region ‘agreed exactly with the Cue population’, i.e. was the form now known as C. baylyi. The Cue population of C. lucasi s.1. studied by Bayly (1984) is almost identical to the type material from the Paroo. Like the Carnarvon population the second segment of the right endopodite (Ri2) in d is enlarged and with a semicircular outer edge, and in © the proportions and lengths of the spines on the third exopodite segment are the same.(Fig.2C; Bayly, 1984, fig. 4). The Cue population is similar to the Paroo type material in other features, including the short spine at the outer distal corner of the first right exopodite segment (Rel) and the marked bend in the right exopod claw (Re3) being about a quarter to a third along its length. Both these western populations of C. /ucasi s.l. are considered as belonging to C. baylyi. The Lake Grace population of C. /ucasi s.1. studied by Bayly (1984) exhibits most of the features of C. baylyi including most of the minor modifications seen in the Carnarvon and Cue material. It has however some curious features unique to it, including an outgrowth on the inner distal corner of the proximal segment of the right exopodite (Rel). This is much more prominent than the usual small rounded protrusion in typical C. baylyi, but not as long as in C. elongata, the only other species with an outgrowth in this position. Also unique to this population is the variable setation on the distal segment of the right endopodite (Ri3) so that generally the 2 setae are of very unequal lengths and usually much reduced. In addition some of the features typical of C. baylyi are either not as well developed (e.g. 785 FIG. 1. Calamoecia baylyi sp. nov.; A, 2 first leg; B & C, anterior and posterior aspects, respectively, of ¢ fifth legs; D, 2 fifth leg; E, ventral aspect of genital segment with lobes of last prosomal segment; F, left lateral aspect of 2 genital segment with lobes of last prosomal segment. B and C from holotype, A, D, E, F from allotype. Scale bars = 0.1mm. the projection midway along the inner surface of the middle right exopodite (Re2) is hardly developed) (Fig. 2D), or more developed (e.g. the outgrowth on the left side of the genital segment is more pronounced) (Bayly, 1984, fig. 5E,F). This Lake Grace material should be considered as C. baylyi, though a little more abberant than other western populations. For 5 populations of C. baylyi, prosomal lengths are a little less than 1mm, with d lengths ca 10% less than 2(Table 1). Differential Diagnosis. C. baylyi is easily recognisable by the rounded projection on the middle inner surface of the second right exopodite segment (Re2) of the male (Fig.1B,C). The only other species with a remotely similar structure is C. australica, in which the projection IMIG. 2. Caleamneeia buylyi sp. nov. (see Material for details); B, posterior aspects of ¢ specimen from Carnarvon basin; C, posterior aspect of population (after Bayly, 1984, fig. +4); D, posterior aspect of 4 fifth legs, Lake Grace population (afer Bayly, 1984, fig-SA). Seale bars = (Imm. is about a third along the inner edge and moreover sharp pomted. Other notable features are variously shared with a limited range of other species. These include: 1) Outer edges of the exopods of the first pair of legs in both sexes have 3 spines each with a few large stout secondary spinules (Fig. 1A), Of other Calamoecia species examined for these spines, C. halsei has them and significantly C, /ueusi s,s, does not (Bayly, 1998), 2) The claw of the rightexopodite of the 7 fifth leg(Re3), lacks an inner spine (Fig. [B.C)asinc. australica, C. cunberra, C. halsei and C. zeidleri (Bayly, 1998). C. /ycasis.s, alsa lacks this spine but all other species have it (Bayly, 1992), 3) The lefLendopodite ofthe ¢ fitth leg (Ril) is l-scgmented, very elongated, exceeding the length of the lefi exopod and bears 5 terminal and subterminal spines. This feature is shared with the same 4 species listed above, but usually not TABLE 1. from WA; A, second and third segment of exopodite of 2 filth leg , specimen from Carnarvon basin a filth legs, Cue Prosomal lengths ii mim of 10 as and 10) 2S from 5 boculartoes oF C. bens i. * Only 6dsand4 ? MEMOIRS OF THE QUEENSLAND MUSEUM broad sense, this feature 1s shared only by C. aanpulla and C. lucast s.4,. but the latter has no terminal denticles. In addition, C. luceasi lacks an inner subterminal spine that is present in CL baviyi and overall the length of the Jett exopodite is shorter relative tothe left endopodite (a Inttle greater than half in C. lueasi ss. cf three-quarters in C. baylyi). 5) On the ? fifth leg, the curved outgrowth on the inner distal corner of the middle exopodite seginent (Re2) is relatively long in C. bayvlyi reaching about to the end of the distal exopodite segment plus its spines (Fig, 1D), In many other species, including C. lucasi s.s., this outgrowth terminates well short of the distal segment plus its spines (Bayly, 1961). Only C, halsei exhibits a similar condition (Bayly, 1998). 6) Again onthe & fifth legs, some of the 5 or 6 spines of the distal exopod segment (Re3) are only a little longer than the length of the segment in C. haviys (Fig, 1D), as tor C. Aalsei (5 spines) and C, salina also (but C. salina has only 2 terminal spines mstead of 6). By contrast in most other species of Calamoecia, including C. luecsi, the spines are more than twice the length of the segment (Bayly, 1961) and siynilivantly in C. lucast §.. there are only 5 spines, though some species have 6 (e.g. C. #rifide). Only C. zeidleri has shorter terminal spines (which number 5) (Bayly, 1984). In the dichotomous key provided by Bayly (1992). C. baylyi would key out through couplets 1.2 8.9 and Il to C. /ueas7, but as pointed out above is easily distinguished from this species, Also, as shown by Bayly (1984) the Cue form of right fifth lex, 5 measured. with C. ducasi s.s.. which although of | sa ; Pdpuluticn Mean,3 a | are ofd Mean ¥ | Rangeof? Source of | similar construction. has 2-5, usually | *"? lengxti tenths | length | lengths dita 4 terminal/subterminal spines, (Bayly, |.CNT, Paroo U7 078-092 0.94 0.91) 1.88 Original 9? sodw 1961,1992, 1998). Bloodwaoch Oy) | 0856094 | 0,96 892-).0) Grizinal 4) The left exopodite of the 3 fifth wae 089 | 086-090) 088 087-100 Onainal leg 1s l-segmented and with a aa ; . ‘ . Cue, WA 0.86 0.96 Bayly, 1984 | spade-like terminal protrusion