MEMOIRS OF THE QUEENSLAND MUSEUM BRISBANE VOLUME 28 31 AUGUST, 1990 PART 2 MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum ANOPLOGNATHUS HILLERI SP. NOV. (COLEOPTERA: SCARABAEIDAE: RUTELINAE) FROM SOUTHEASTERN QUEENSLAND AND NOTES ON A, FLINDERSENSIS CARNE P.G, ALLSOPP Allsopp, P.G. 1990 08 31; Anaplognathus hilleri sp. nov. (Coleoptera: Scarabaeidae: Rutelinae) from southeastern Queensland and notes on A. flinderscnsis Carne. Memoirs of the Queensland Museum 28(2): 377-381. Brisbane. ISSN 0079-8835. Anoplognathus hilleri is described from both sexes collected in Cooloola National Park, SEQ. The adpressed white setae on the dorsal surface, the general shape of the clypeus of both sexes and of the aedeagus, and the black spots on the elytra clearly place A. hilleri with the A. velutinus species-group, A. hilleri differs from both A, velasinus and A. flindersensis in the detailed shape of the aedeagus and in colour, A further specimen of A. flindersensis Carne is also noted. 1] Anoplognathus hilleri, Rutelinae, Scarabactdae, Coleoptera. P.G. Allsopp, Bureau of Sugar Experiment Stations, PO Bax 651, Bundaberg, Queensland 4670, Australia; 18 October, 1989. Anoplognathus Leach was last revised by Carne (1957) who recognised 32 species. A, an- tiquus Arrow has since been transferred to Trioplognaihus Ohaus (junior synonym of T. griseopilosus (Ohaus)) (Carne, 1958) and four further species have been described (Carne, 1981; Allsopp and Carne, 1986). Additional dis- tribution records for Anoplognathus spp. were given by Carne (1958, 1981), Carne and Mon- teith (1971), Allsopp (1975, 1987), Monteith (1986), Allsopp and Lloyd (1987), and De Baar and Hockey (1987), The majority of species occur in coastal and subcoastal eastern Australia with only four species recorded from the arid interior, One species is known also from Papua New Guinea, This paper describes a previously unknown species collected in southeastern Queensland and gives notes on a further specimen of A. flindersensis Carne. The following abbreviations are used for collections: AH = A. Hiller collec- tion, Mt Glorious; ANIC = Australian National Insect Collection, Canberra; PGA = P,G, Allsopp collection, Bundaberg; QDPI = Queensland Department of Primary Industries, Brisbane; QM = Queensland Museum, Brisbane; RIS = R.I. Storey collection, Mareeba. Anoplognathus Leach Anoplagnathus Leach, 1815, p, 43; Carne, 1957, p. 88, 1958, p. 181, 1981, p. 289; Allsopp and Carne, 1986, p. 99. Type species Melolontha viridiaeneus Donovan, L805; designated by Carne, 1957, p. 93. Paranoncea Castelnau, 1840, p. 143; Lansberge, 1873, p So. Anoplognathus hilleri sp. nov. (Figs 4-8) MATERIAL. EXAMINED HoLoryre; OM T11289 3. Coaloola National Park, Queensland (153°5’E, 26°10’S), 15.ix.1988, A. and K. Hiller, to UV light. PARATYPES: 2 d J, same dala as holotype; 14 dd, | 2. same dala as holotype except 28, ix-1.x,1988: 1d, Cooloola, 21.1x.1987, K. and T. Thomas; in AH, ANIC, PGA, ODPI, OM, RIS, DESCRIPTION Male: Total length 23.5-26.5 mm. Head, pronotum, scutellum, pydigium, legs and ventral surface red-brown; elytra red-brown to nearly black with broad longitudinal band of ycllow-brown cither side of sutural interval but narrower and less defined towards apex (Figs 4,6), clytra in paler specimens red-brown; all adpressed setae white. Labrum triangular, apex rounded, surface densely punctate with long selae except scatlered shor! selae on smoother apex, Clypeus with surface of anterior face with micropunctures and scattered long setae arising from larger punctures, anterior face 2.4 times as wide as deep: dorsal surface rugulose with flat- tened udpressed sctac, anterior margin slightly conyex, lateral margins parallel near base then sharply curved and tapering to reflexed anterior margin, 1.7 times as wide across base as mid length; clypeofrontal suture posteriorly sinuate 378 MEMOIRS OF THE QUEENSLAND MUSEUM Fics 1-6. Dorsal view of Anoplognathus spp. 1 - A. velutinus, 3; 2 - A. flindersensis, holotype 3; 3 - A. flindersensis, paratype 2; 4 - A. hilleri, holotype d; 5 - A. hilleri, paratype 2; 6 - A. hilleri, living 3 (photo, A. Hiller). in middle. Frons with large punctures becoming smaller posteriorly, some punctures with single flattened adpressed seta; ocular canthi with dense erect setae. Maxillary palps conspicuous, segment 3 longer than segments 1 and 2 com- bined, segment 3 with large longitudinal con- cavity on upper side. Antennae 10-segmented; club 3-segmented, 2.6 mm long and about as long as segments 2-7 combined, Labium with long dense sctac near base, apex glabrous. Pronotum punctate with sparse irregularly-distributed adpressed setac, with faintly rugose impunctate median stripe; 1.6 times as long as wide; anterior and lateral margins defined by obvious ridges, NEW SPECIES OF ANOPLOGNATHUS posterior margin weakly bilobed in median third where ridge absent; anterior angles slightly ob- tuse, posterior angles distinctly obtuse; long setae arising from beneath posterior margin, denser and longer opposite scutellum. Scutellum with flattened adpressed setae, denser on lateral edges. Elytra with surface conspicuously punctate, coarsely rugose near apices, almost glabrous along sutural interval, elsewhere punc- tures with 1-7 adpressed setae (average 4 to 6); apices contiguous, sutures slightly produced with series of short sharp spinules; epipleurae only visible on posterior half, glabrous. Ventral surface with dense long erect white setae. Post- coxal prosternal process absent. Mesosternal process acute, slightly depressed away from body anteriorly, apex almost glabrous and level with hind margin of fore coxae. Fore tibiae broad, evenly tridentate with teeth at less than right angles to major axis, with white scales dorsally and fine reddish-brown setae on inner margins; mid and hind tibiae with mixture of scattered long reddish or white setae; hind tibial spurs separated by 2 large and | smaller ciliae. Ventrites 1-5 with dense white setae and scat- tered longer white setae; sternite 6 with reduced vestiture, almost bare in middle, slender pale yellow setae across median third of posterior margin. Pygidium finely rugulose, with uniform coating of white setae, longer erect pale yellow sctae near posterior margin; posterior margin with continuous ridge, slightly truncated in mid- dle. Acdeagus similar to that of A. velutinus Boisduval and A. flindersensis Carne but with a more rounded nodule about halfway along outer edge (Figs 7-8). Female (Fig. 5): Total length 26 mm. Head, pronotum, scutellum, pygidium, legs Fics 7-8. Anaplognathus hilleri_ 5 , parametes. Scale line = | mm. 4380 and ventral surface red-brown, pronotum with faint dark spot near mid lateral margin: elytra dark yellow-brown with black spot posterior to humerus. Dorsal surface of clypeus more finely punctate, glabrous, anterior margin almost straight, only very slightly reflexed, lateral mar- gins convex, 2.1 times as wide across base as mid length, Frons more finely punctate, glabrous ex- cept for few adpressed setae across base. Anten- nal club 2.0 mm long. Pronotum with few adpressed selae, mainly on posterior margin. Otherwise as male. COMMENTS The species is named after Anthony Hiller of Mt Glorious who went to great lengths to obtain the two series. The adpressed white setae on the dorsal sur- face, the general shape of the clypeus of both sexes and of the aedeagus, and the black spots on the elytra clearly place A. Ailleri with the A. velutinus species-group (Carne, 1957). A. hilleri differs from both A. velitinus and A. flindersen- sis in the detailed shape of the aedeagus (Figs 7-8) and in colour (Figs 1-6). A. hilleri keys to A. velutinus in Carne’s (1957) ‘key. It and A, /lindersensis can be incorporated into the key by deleting couplet 2 and inserting the following: 2(1). Dorsal surface of body with adpressed white scales (velulinus SPECIES-ZFOUD) reed Dorsal surface of body lacking adpressed white SOAS 20 oj des och dortdongendenoovorenduaachonseeepyosaraqie mses ged 3 2a(2). Head and pronotum red-brown, pronotum of males without black spots near lateral margins; elytra of males red- brown to nearly black with broad longitudinal band of yellow-brown either side of sutural interval, in pale specimens no indication of black humeral spot; southeastern Queensland... cesses eetilleri Allsopp Head and pronotum yellowish-brown, pronotum with black spot near each lateral mar- gin; elytra of males with black humeral spot or longitudinal brownish-black streak extending from anterior margins and becoming progres- BUVELY PAlOT cc cesesserssrsccnerenersssnsesenataesennsnees QO 2b(2a).Elytral punctures with 1-12 scales (average 5-6); elytra of both sexes with black humeral spot, southeastern Australia.....-.s.ccceesve samepengent jeejseatereerey VElulinus Boisduval MEMOIRS OF THE QUEENSLAND MUSEUM Elytral punctures with 0-3 scales; elytra of males with longitudinal brownish-black streak extending from anterior margins and becoming progressively paler; Flinders Ranges, South Austtithia...... cece ene flindersensis Carne The two series were taken in a sandy, dry area with low shrubs and 3-4 Eucalyptus spp. and Casuarina sp. as overstorcy but bordering a Gah- nia (swotdgrass) swamp of at Jeast 0.5 ha. The surrounding cucalypts were searched during the day following the capture of each of the series in an attempt to locate feeding trees but no beetles or tvidence of fveding were seen, When col- lected all specimens had no food in their guts and the female had apparently laid all her eggs. This indicates that the species flics in early Septem- ber. This early fight period appears charac- teristic of the velutinus specics-group; A. velulinus is known from September in south Queensland (Carne, 1957) and A. flindersensis from mid-October in South Australia (Carne, 1981). Most other Anoplognathus spp. fly during summer, The three species of the velutinus species- group have allopatric distributions (A. Aillert oc- curs north of the known northern limit of A. velutinus at Caloundra (Carne, 1957)). Anoplognathus flindersensis Carne (Figs 2-3) Anoplognathus flindersensis Carne, 1981, p. 290. ADDITIONAL MATERIAL EXAMINED ‘South Australia: 1d , Angorichina Hostel, 7 km E of Parachilna, 23.x,1978, E.B, Britton, in ANIC. COMMENTS A. flindersensis was described from Wilpena Pound in the Flinders Ranges. This new specimen comes [rom the western edge of the Flinders Ranges, c. 50 km NNW of the type locality. The black elytral streak is not as well defined as in males of the type series, but there 1s a well-defined black humeral spot. ACKNOWLEDGEMENTS 1 thank Anthony Hiller for access to the specimens and for the photograph of a living specimen (Fig, 6); Phil Campbell for the SEM illustration; and Geoff Monteith for assistance with the layout of the figure. NEW SPECIES OF ANOPLOGNATHUS LITERATURE CITED ALLsopp, P.G. 1975. Scarabaeidae collected in light traps at St. George, Brookstead and Gatton, Queensland. Australian Entomological Magazine 2: 69-71. 1987. New records of scarab beetles (Coleoptera: Scarabaeidae) from central Queensland. Queensland Naturalist 28: 87-8. ALLsopP, P.G, AND CARNE, P.B. 1986, Anoplognathus vietor sp. n. (Coleoptera: Scarabaeidae: Rutelinae) from west Queensland. Journal of the Australian Entomological Society 25: 99-101. ALLSopP, P.G. AND LLOYD, R.J. 1987. New records of Scarabaeidae (Coleoptera) from central Australia. Australian Entomological Magazine 13: 71-6. CARNE, P.B. 1957. A revision of the ruteline genus Anoplognathus Leach (Coleoptera: Scarabae- idae). Australian Journal of Zoology 5: 88-145. 1958, A review of the Australian Rutelinae (Coleoptera: Scarabaeidae). Australian Journal of Zoology 6: 162-240. 1981, Three new species of Anoplognathus Leach, and new distribution records for poorly known species (Coleoptera: Scarabaeidae: Rutelinae). 381 Journal of the Australian Entomological Society 20: 289-94. CARNE, P.B. AND MONTEITH, G.B. 1971. Three little- known species of Christmas beetle. News Bul- letin of the Australian Entomological Society 7: 61-3. CASTELNAU, F.L. 1840. ‘Histoire naturelle des Insec- tes Coléoptéres, avec une introduction renfer- mant l’anatomie et la physiologie des animaux articulés, par M. Brullé. Tome deuxiéme.’ (Dumeénil: Paris). DE Baar, M. AND Hockey, M.J. 1987. Additional records for Queensland Anoplognathus (Coleop- tera: Scarabaeidae: Rutelinae). News Bulletin of the Entomological Society Queensland 14; 153- 4, LANSBERGE, J.G.W. VAN. 1873. Notice sur la Paranonca prasina, Castelnau, Annales de la Société entomologique de Belgique 16: 195-6. LEACH, W.E. 1815. ‘The zoological miscellany; being descriptions of new, or interesting animals’ Vol. 2. (Nodder & Son: London). MONTEITH, G.B. 1986. Insects from Kroombit Tops, Queensland, with some results of a sile survey of Coleoptera. Queensland Naturalist 27: 27-34. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum TWO NEW SPECIES OF TRICHOTICHNUS MORAWITZ FROM NORTH QUEENSLAND (COLEOPTERA: CARABIDAE: HARPALINAE) MARTIN BAEHR Baehr, M. 1990 08 31: Two new species of Trichorichnus Morawitz from north Queensland (Coleoptera: Carabidae: Harpalinae). Memoirs of the Queensland Museum 28(2); 383-388. Brisbane, ISSN 0079- 8835. Two new species of Trichotichnus are described from north Queensland: T. storeyi sp. nov. and T. tolgae sp. nov. Also new records of T. strancoi (Louwerens) {fom north Queensland are communicated. A key is given to all known Australian Tricholichnus. Zwei neue Trichotichnus-Arten aus Nordqueensland werden beschrieben: 7) storeyi sp. nov. und T. telgae sp. nov. AuBerdem werden weilere Funde von 7. straneoi (Louwerens) mitgeteilt. Fir alle ausiralischen Trichotichnus-Arien wird ein Bestimmungsschliissel gegeben. (Coleoptera, Carabidae, Harpalinae, Trichvtichnus, new species, Queensland. Martin Baehr, Zoalogische Staatssammlung, Minchhausenstrape. 21, D- 8000, Miinchen 60, West Germany; 14 January, 1989. Trichotichnus is a genus of Holaretic-Oriental ground beetles that are especially common in the eastern Palearctic Region but also rather numerous as far south as New Guinea (Dar- lington, 1968). Till recently this genus was un- recorded from Australia (Darlington, 1968; Noonan, 1985) but it has since been discovered in northeastern Queensland (Baehr, 1983, 1985) where a widespread Oriental and an endemic Australian species are now known. In Australia, as well as in New Guinea, Trichotichnus are mainly collected at light traps. Single specimens have been found in rain forest litter, which is presumably the true habitat of most Indoaustralian Trichotichnus. The two new species described in this paper were taken at light traps, together with three further specimens of T. straneoi (Louwerens). They were received courtesy of Mr R.1. Storey (Mareeba), They were collected at different localities on the Atherton Tableland and the Windsor Tableland between Mossman and Cooktown, These are areas of upland rainforest. Neither of the new species match any of the described New Guinean or Australian forms. How- ever, both are vaguely associated with different species-groups or subgenera of Noonan (1985). Abbreviations. for collections are CBM, Col- lection of M. Baehr, Miinchen; DPIM, Depart- ment of Primary Industries, Mareeba; and OMB, Queensland Museum, Brisbane. Measurements were made using an ocular micrometer. Length was measured from apex of labrum to apex of elytra. Hence measurements can slightly differ Fic, 1. Trichotichnus stureyi sp, nov,, holatype. Scale: Smm. 384 from those of other authors, especially Dar- lington (1968). Trichotichnus straneoi (Louwerens) (Fig.8) Carbanus straneoi Louwerens, 1962, p. 142, fig. 7. Trichotichnus straneoi Darlinglon 1968, p. 50; Noonan 1985, p. 67; Baehr 1985, p. 21. MATERIAL EXAMINED NEW ReEcorpDs: 1d, Windsor Tableland, Qld, 29- 30, X11, 1984, at light, R.L. Storey (DPIM); 1 2, Tolga, Qld, 22, VIT.1986, light trap, J.D. Brown (CBM), 19, Tully Falls, SF 730m, 18km SSW Ravenshoe, Qld, 18.1.1988, light, Storey and Dickinson (DPIM). REMARKS This specics was recently included in the sub- genus Trichotichnus s. str. by Noonan (1985). It is known from Amboina, New Guinea, New Britain, and New Ireland (Darlington, 1968) and was recently recorded from north Queensland (Buehr, 1985). The new records indicate that this species is rather widely distributed and common in the tablelands of northeastern Queensland, It lives in leaf litter in rainforest (Baehr, 1985). However, the best collecting method seems to be lighting; all three of the records are from light traps. Trichotichnus storeyi sp. nov. (Figs 1,3,5,7,8) MATERIAL EXAMINED HoLoryee: ¢ , Australia, NO, Tully Falls SF 730m, 18km SSW Ravenshoe, 18,1.1988, light, Storey and Dickinson (QMB T.1 1327). PARATYPES: 1d, 12, same data, 2 Jabelled: Trichotichnus demarzi Baehr, det. R.I. Storey 1988 (DPIM, CBM). TYPE LOCALITY Tully Falls, Atherton Tableland, NQ. DIAGNOSIS Easily recognised by yellow border of pronotum, clytra, and abdomen and by wide posterior angles of pronotum. DESCRIPTION Measurements: Length: 7.9-8.2mm, width: 3.3-3.4mm. Width of head/pronotum: 0.78-0.8. Width/length of pronotum: 1.49-1.53. Width of base/apex of pronotum: 1.18-1.2. Width of MEMOIRS OF THE QUEENSLAND MUSEUM pronotum/elytra: 0.76-0.78. Length/width of elytra: 1.55-1.58. Colour, Black with slight metallic lustre. Lateral border of pronotum narrowly yellow. Two outer intervals of elytra yellow, this yellow border widened at apex almost to suture. Lower surface black, lateral parts of thorax brown with scattered yellow spots, abdominal segments with a lateral and a sublateral yellow spot each, brown between, apex of last segment yellow, Labrum brownish, antennae dark yellow, with 2nd and 3rd segments piceous and terminal segments be- coming gradually lighter. Each segment with a distinct dark median stripe on inner and outer Fic, 2, Trichotichnus tolgae sp. nov., holotype. Scale: 5mm. TWO NEW SPECIES OF TRICHOTICHNUS MORAWITZ FROM NORTH QLD | 3 4 Fics 3,4. Clypeo-ocular sulcus. 3. Trickotichnus Storeyi sp. nov.; 4. Trichotichnus tolgae sp. nov, surface. Palpi lighter brown. Femora yellow, tibia yellow at base, infuscate toa apex, consider- ably darker than femora. Tarsi infuscate. Head: Large, with large, rather protruding eyes. Orbits short. Eyes separated from mouth by c.1/6 of eye diameter. Clypeal suture deep. Clypeo-orbital sulcus distinct, clongate, almost attaining eye, though not prolonged inside eye. Clypeus with one seta each, labrum 6-setose. Antennae short, barely surpassing base of pronotum. Median segments wide, square, al- most as wide as long. 3rd segment pilose from middle, Last and fore-last segments of palpi fine- ly sctose. Labium with elongate, unidentate tooth. Microsculpture of upper surface incon- spicuous. Pronotum: Considerably wider than head, c.1.5 times wide as long, widest shortly behind position of lateral seta. Apex very lightly ex- cised, anterior angles rounded off, not projecting. Lateral borders anteriorly convex, behind middle almost straight. Posterior angles wide, obtuse, base laterally rather oblique. Base distinctly bor- dered laterally, medially smooth. Median line well impressed, neither attaining apex nor base. Basal grooves shallow, slightly linear. Base laterally punctate, rather smooth in middle. Microsculpture indistinct, rather irregular. Elytra: Rather parallel, short, lateral borders well sinuate in front of apex. Shoulder rounded off. Striae fine, impunctate, intervals slightly convex, especially near apex. Setiferous punc- ture adjacent to 2nd stria, approximately at posterior 2/3 of elytra. Microsculpture of inter- vals consisting of very indistinct transverse lines. Intervals also with fine punctures, rather glossy, winged. Lower surface: Prosternum, metasternum, and oe) iz <) wn base of abdomen with fine, rather dense pilosity. Pilosity on 4th-6th abdominal sternites extreme- ly short and fine, difficult to see. Lower surface shagrcened. Last abdominal sternite laterally slightly excised. d and 2 with 4 setae at apex of last sternite, Legs: Anterior tibia not much widened at apex. ¢ anterior and median tarsi biseriately squamose beneath. Ist segment of metatarsus rather short, considerably shorter than 2nd and 3rd segments together, Male genitalia: Aedeagus elongate, narrow, basally strongly curved. Apex with minute up- turned denticle. Internal sac and parameres sce Fig. 7. Variation: There is little variation among the few specimens available. DISTRIBUTION Known only from type locality in southern Atherton Tableland, NEQ. HABITAT Not known, but perhaps lives in rainforest lit- ter. This is apparently an upland rainforest species. RELATIONSHIPS It is difficult to assign 7. stereyi to any of the currently distinguished subgenera of Noonan (1985), because it is somewhat intermediate be- 2) 6 Fics 5,6. Posterior tarsus. 5. Trichatichnus stareyi sp, nov.; 4. Trichotichnus toleaé sp. nov, 386 ra 7 (sf es! Fic. 7. d genitalia of Trichotichnus storeyi sp. nov. Scale: 0.5mm, tween Trichotichnus s. str. and Harpaloxenus Schauberger. It has the apex of anterior tibia not conspicuously widened, bul has at the same time the Jower surface of the thorax and abdomen strongly mottled, which is only ascribed to mem- bers of subgenus Harpaloxenus (Noonan, 1985). From descriptions, T. storeyi seems next to T. brandti Darlington and T. obscurus Darlington, respectively, both from New Guinea, However, it iS shghtly smaller than both species and is further distinguished by slightly wider pronotum with lateral borders not at all sinuose, and by wide, conspicuous, yellow, lateral border of elytra. Trichotichnus tolgae sp. nov. (Figs 2,4,6,8) MATERIAL EXAMINED: HovLorype: 2, Australia, NO, Tolga, 3,.X1.1985, J.D. Brown, light trap (OMB T.11326). TYPE LOCALITY Tolga, 5km N Atherton, Atherton Tableland, NEQ. DIAGNOSIS Distinguished by lack of dorsal and ventral pattern and by convex pronotum with short, but accentuate prebasal sinuosity. DESCRIPTION Measurements: Length: ¢.7.Smm, width: ¢.3.2mm. Width of head/pronotum: 0.7. Width/length or pronotum: 1.39. Width of base/apex of pronotum: 1.27, Width of pronolum/clytra: 0.76. Length/width of elytra: c.1.45. Colour: Piccous, rather glossy. Ist antennal scement dark yellow, following segments slight- ly darker, terminal segments yellow again. MEMOIRS OF THE QUEENSLAND MUSEUM Median segments with inconspicuous dark stripe on inner and outer surface. Palpi dirty yellow. Legs reddish, apex of tibia and tarsi slightly darker. Ventral surface piceous, Head: Almost as wide as apex of pronotum. Eyes large, protruding. Orbits distinct, oblique. Eyes separated from mouth by c.1/S of diameter of cye. Clypeal suture deep. Clypeo-orbital sul- cus deep, rather short, almost punctiform, not attaining eye, not prolonged inside median bor- der of eye, Clypeus with one seta each, labrum 6-setose, Antennae medium-sized, attaining base of pronotum. Median segments almost twice as long as wide. 3rd segment pilose from middle. Last and fore-last segments of palpi sparsely setose. Labium with a short, triangular tooth. Microsculpture of upper surface fine, though distinct, consisting of rather regular, slightly transverse meshes. Pronotum: Convex, rather narrow, with wide base. Apex straight, but anterior angles slightly produced. Lateral borders convex throughout, though with a short, distinct sinuation just in front of the rectangular posterior angles. Base almost straight. Pronotum widest at position of lateral setae, slightly in front of middle. Median line superficial, neither reaching apex nor base. Base strongly bordered, also in middle. Basal grooves shallow, circular. Base with very scat- tered punctures, laterally of basal grooves rather smooth. Medially behind apex also with some punctures. Surface glossy, but with fine, regular, slightly transverse microsculpture, Elytra: Rather short and wide, slightly widened behind middle. Shoulders rounded. Lateral borders well sinuate in front of apex. Striae smooth, quite deep. Intervals convex throughout, near apex strongly convex. Setiferous puncture in middle of 3rd interval, though nearer to 2nd stria, far down on apical declivity, at approximately last 1/6 of elytra. Microsculpture distinct, composed of slightly transverse, regular meshes. No additional pune- tures visible. Winged. Lower surface: Prosternum, melasternum, and basal abdominal segment medially with short pubescence. Other abdominal segments virtually smooth. Lower surface fincly microreticulate- Last abdominal sternite laterally not excised, in 2 4- setose. Legs: Anterior tibia not widened at apex, Clothure of anterior and median tarsi of d un- known. Ist segment of metatarsus elongate, ap- proximately us long as 2nd and 3rd segments together. TWO NEW SPECIES OF TRICHOTICHNUS MORAWITZ FROM NORTH QLD Male genitalia: Unknown. DISTRIBUTION Known only from type locality in Atherton Tableland, NEQ. HABITAT Unknown. The type was collected at light. It probably lives in rainforest litter. RELATIONSHIPS Unknown. T. tolgae belongs to subgenus Tricholichnus s. str., though it is a rather atypical species. There is no obvious relationship to Cooktown Innisfail Fic. 8. Distribution of the Australian Trichotichnus- species: @Trichotichnus straneoi (Louwerens); B7richotichnus demarzi Baehr, WTrichotichnus storeyi sp. nov.; ATricholichnus tolgae sp, nov, 387 anyone of the New Guinean species included in Darlington’s (1968) key. Using this key, T. tol- gae comes to 7. medius Darlington which is, however, not closely related to 7. tolgae. As long as the ¢ genitalia and tarsal vestiture is un- known, the relationships will be obscure. DISCUSSION With the two species described here, four Trichotichnus species are now known to occur in northeastern Queensland, all in or near the uplands of Atherton Tableland and Windsor Tableland. All species seem to live in leaf litter of mountain rainforest (Baehr, 1983, 1985) which is the same mode of life thal several of the New Guinean species apparently have (Dar- lington, 1968). The four Australian species belong to different species groups and they are perhaps more closely related to certain New Guinean species than one to another. As was to be expected from the high species diversity in New Guinea, northeastern Australia also has quite a diverse Trichotichnus fauna. Certainly the four species are outliers of the northern, Oriental fauna fram which many members have dispersed to northern Queensland, mainly via New Guinea and Cape York Peninsula (Dar- lington, 1961, 1971; Kikkawa, Monteith and In- gram 1981). This means that north Queensland was subject tu several independent invasions of different Tricholichnus stocks, the most recent apparently being that of 7. straneoi which has conspecific populations in Australia, New Guinea and much of Melanesia. It is to be ex- pected that future collecting in the upland rain- forests of northern Queensland will result in the discovery of yet more Tricholichnus. KEY TO THE AUSTRALIAN SPECIES OF TRICHOTICHNUS 1. Prathorax wide with strongly convex sides, head only 2/3 as wide as prothorax. Surface of elytra- strongly iridescent. Small species, less than 6.5mm long -.-.... T. straneoi(Louwerens) Prothorax less wide, sides posteriorly rather straight oreven sinuate, head almost 3/4 as wide as prothorax. Elytra not or only slightly irides- cent, Larger species, always over 7mm long.. 2 2. Pronotum und elytra distinctly bordered with yel- low. Abdomen with lateral yellow border and sublateral yellow spots, Sides of pronotum near 388 base convex, posterior angles wide, because lateral parts of base clearly oblique oateeTEE TE goontpes send erabhgiap UREA AFTS T, storeyi sp. nov. Pronotum at most inconspicuously bordered with yellow. Abdomen with lateral yellow bor- der or spots or not, but without sublateral spots. Sides of pronotum posteriorly straight or even slightly concave. Posterior angles less wide, DaSe SITATBHE s...seeserecsenconstssnceszoogeserrsrnesetodtestte 3 3. Larger species, 9mm long or longer. Clypeo-ocular sulcus prolonged as a furrow inside eyes. Ab- dominal segments with large, yellow, lateral spots. Femora conspicuously lighter than tibiae. Pronotum heart-shaped, widest well in front of middle, sides near base straight beabiiiebecborndees coateostddnrost cdyebaiest’ T. demarzi Baehr Smaller species. c.7.5mm long. Clypeo-ocular sulcus nol prolonged inside eye. Abdomen without yellow spots. Femora not conspicuously lighter than tibiae. Pronotum widest approximately at middle, sides posteriorly convex, but shortly sinuate just in front of posterior angles Sasa udeduaetco}etp bdosansssheesneesbeneceses T. tolgae sp. nov. ACKNOWLEDGEMENTS I want to express my thanks to Mr R.I. Storey (DPIM, Mareeba) who kindly offered the specimens for examination. MEMOIRS OF THE QUEENSLAND MUSEUM LITERATURE CITED BAER, M. 1983. Trichetichnus demarzi sp. nov, eine weitere Trichotichnus-Art neu fiir Australien (Insecta, Coleoptera, Carabidae). Spixiana 6: 109-112. 1985. Trichotichnus Morawitz, a genus new to Australia (Coleoptera: Carabidae: Harpalinae). Australian Entomological Magazine 12: 21-22. DARLINGTON, P.J. JR. 1961. Australian carabid beetles V. Transition of wet forest faunas from New Guinea to Tasmania. Psyche 68; 1-24. 1968. The carabid beetles of New Guinea III. Har- palinae continued. Perigonini to Pseudomor- phini. Bulletin of the Museum of Comparative Zoology, Harvard 137(1): 1-253. 1971. The carabid beetles of New Guinea IV. General considerations, analysis and history of the fauna, taxonomic supplement. Bulletin of the Museum of Comparative Zoology, Harvard 142(2): 129-337. KIKKAWA, J., MONTEITH, G.B. AND INGRAM, G.J. 1981. Cape York Peninsula: Major region of faunal interchange. pp. 1697- 1742. Jn Keast, A. (ed.), ‘Ecological Biogeography of Australia’. (Junk: The Hague). LOUWERENS, C.J. 1962. New Carabidae from In- donesia, chiefly from Amboina. Tijdschrift voor Entomologie 105: 135-147. Noonan, G.R, 1985. Classification and names of the Selenophori group (Coleoptera: Carabidae: Har- palini) and of nine genera and subgenera placed in incertae sedis within Harpalina. Milwaukee Public Museum. Contributions in Biology and Geology 64: 1-92. HYSTEROTHYLACIUM WARD AND MAGATH, 1917, AND ICHTHYASCARIS WU, 1949, ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISHES NIEL L. BRUCE Bruce, N.L. 1990 08.31: Hysterothylacium Ward and Magath. 1917 jand /chthyascaris Wu, 1949, ascaridoid nematodes from Australian demersal fishes. Memoirs of the Queensland Museum 28(2); 389-426. Brisbane. ISSN 0079-8835. Genera and species of ascaridoid nematodes are recorded fram Australian demersal marine fishes. /chthyascariy Wu is tedefined with two new species and the following new combinations: /. fisheri (Hooper), /. vicentei (Santos), J. lutjani (Olsen), I. mediterraneus (Lebre and Petter). /, chirocentri (Yamaguti) and /. biwakoensis (Fujita). [chthyascaris Jacks flanged lips, any trace of interlabia and intestinal caecum: the excretory system is posteriorly unilateral and filamentar. Descriptions are given for the following species: Hysterothylacium leptaspi n. sp., Hysterothylacium chrysustomi n. sp., H. sebae n. sp., #7. tasmaniense (Johnston and Mawson), H. thalassini n. sp., HH. zenis (Baylis), [chthyascaris gymnocraniae n. sp. and J. sillagoides n, sp. Notes are given on an unidentified species of Goezia from Arius thalassinus. (] Nematoda, Ascarideidea, taxonomy, fish parasites, Indo-West Pacific. Niel L, Bruce, Queensland Museum, PO Box 300, South Brisbane, Queensland 4101, Australia; 27 June, 1989. A brief review of ascaridoid nematodes from Australian marine fishes was given by Bruce and Cannon (1989) recording ascaridoids from Australian pelagic marine fishes. This present work records ascaridoids from demersal and euryhaline fishes. The species treated here in- clude only those species that were represented by adult specimens in good or reasonable condition. Material examined prior to preparation of this work included species of Hysterothylacium and Ichthyascaris from a further 10 host species. Clearly the number of species presently recorded from Australia is but a fraction of the potential total. MATERIAL AND METHODS The follawing demersal fish (or fish stomachs) were examined and, except for those reported in the species description, proved negative for adult ascaridoids. The number in parentheses is the total number of specimens examined. Fish totals of less than 4 are not represented individually. From the Rockhampton coast and central sec- lion Great Barrier Reef, 7 species in total. Ser- tranidae: Plectropoma leopardus (13); Lutjanidae: Lutjanus malabaricus (10) and L. sebae (10). From Heron Island and Wistari Reefs, Capricorn Group, southern Great Barrier Reef, 22 species. Serranidae, 7 species including Plectropomus leopardus (16), Epinephilus fas- ciatus (6) and F. merra (5); Lutjanidae, 3 species including Lutjanus carponotatus (9): Lethrinidac, Lethrinus chrysestomus (30), L. nebulosus (4) and Gymnocranius bitorquatus (34). Labridae, Choerodon venustus (26) and C albigena (6). From southeastern Queensland (Moreton Bay region) Platycephalus fuscus (28), Sillage maculata (60), Pomatomus saltator (34) and Acanthopagrus australis (20). Names are from Grant (1987) and Hutchins and Swainston (1986). Materials and methods are the same as used and detailed by Bruce and Cannon (1989). Meas- urements are given in micrometres except where otherwise indicated, Measurements in mil- limetres were made under binocular microscope, those in micrometres under 4 compound micro- scope, Abbreviations used in the text: AHC - Australian Helminth Collection, at the South Australian Museum; BL- Body length; BM(NH) --British Museum (Natural History), London; CSIRO - Commonwealth Scientific and In- dustrial Research Organisation; ED - eget duct; QM - Qucensland Museum; SAM - South Australian Museum, Adelaide. Goezia Zeder The genus Geezia has been rediagnosed by Sprent (19784) and more recently by Deardorff and Overstreet (1980), In bath diagnoses the lack of interlabia is a character, among others, of 390 generic significance. To date no species of this genus that has been illustrated has shown any trace of interlabia or semi-interlabia (see “Remarks’ for Raphidascaroides for discussion of interlabial structures). The lack of interlabia is clearly illustrated by the SEM’s of Sprent (1978a) and Lébre and Petter (1983). It is of some interest, therefore, to record a species of Goezia (or an undescribed genus closely allied to Goezia) from Arius thalassinus trawled by CSIRO in the Gulf of Carpentaria at Weipa. These samples include one that has a large mature male (31.7 mm long by 1.5 mm wide) and 2 large females (the largest measuring 50.0 mm long by 2.7 mm wide). These specimens (QM G10286) are in very poor condition and cannot be used for description. The remaining LETS S354) [> ===5 ae i == = [| is = PT eas = ss YYW yr A Fic. 1. ? Goezia sp. A, interlabium (scale 100 wm); B, subventral lip (100 jzm); C, oesophageal region (400 pm). MEMOIRS OF THE QUEENSLAND MUSEUM two samples (QM GL10250, GL10285) contain immature specimens (10.7-18.0 mm). All have distinct pseudo- or sessile interlabia (Fig. 1). Additionally the serrations (or cuticular spine tows) fade away completely towards the tail in 2 specimens, and are very weak in the caudal region of the third. These are large worms com- pared to others of the genus, which measure as adults from under 3 mm to about 25 mm. Hysterothylacium Ward and Magath Bruce and Cannon (1989) described those species of the genus to be found is Australian pelagic and oceanic fishes. A further 5 species are recorded here, all from demersal or estuarine hosts. The diagnosis given by Deardorff and a ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH Overstreet (1981) is not in need of modification at present. /ysterothylacium is the largest of the ascaridoid genera, and considerable variation is present with regard to tail morphology, the ex- crelory system, and presence or absence of alae. Tt is still neccessary that further species be described and that 4 detailed description of the type species be available before a critical analysis of the genus can be given. Hysterothylacium leptaspi n. sp. ( Figs 2-4) MATERIAL EXAMINED All from Embley Estuary, Weipa, Gulf of Carpen> laria, Q., from stomachs of Arius leptaspis, coll, CSIRO, 10 males, 11 females, Heinemann’s Creek, middle reaches of Embley Estuary, 20 Feb. 1987 (male HoLorTyre QM GL10244, PaRatyres OM GL10245), 18 males, 7 females, same data as above (OM GL10246, OM GL10247, AHC 18813). Also examined: 2 females, lower reaches of estuary, 15 and 23 Feb, 1987, Arius rhalassinus, co}], CSIRO (OM GL10248, GL 10249), TYPE LOCALITY Embley Kstuary, Weipa, Q.. es 12°13.5'S. 144'57.0°E. Type Host Arius leptaspis (Bleeker), Anidae. DIAGNOSIS Cuticle finely annulated, appearing smooth under light microscopy; posteriorly with transverse folds, Lips with length to width ratio ]: 0.93-1.45 (mean = 1,02, n = 15) with deep postlabial grooves; constricted about one third length from anterior. Alac absent. Caudal papil- lae pairs: precloacal 40-58 in an irregular row; paracloacal 2 pairs; postcloacal 4-5 pairs, usually without doubled papillae; medioventral precloa- cal organ present. Spicules approximately equal in length, 1:1.03-1,10, 3.64-5.74% BL. Tail short, with finely nodulose conical lip, DESCRIPTION Based on 6 mature males and 8 mature females. General. Body reaching greatest width about mid-body. Dorsal lip slightly smaller than sub- ventrals, about as long as wide; flanges posi- tioned abyut half way along Jip, small and triangular, lateral constriction weakly developed. Interlabia prominent, extending 391 anterior to Gesophagus; about half-length of lip, medially expanded, Oesophagus 7,63-10.59% BL. Ventriculus usually narrower than widest level of oesophagus, longer |han wide; posterior- ly narrowing gradually to ventricular appendix. Ventricular appendix very slender, (difficult to observe without dissection) 104.3-261.0% Jength of oesophagus, Cervical papillae con- spicuous, near nerve ring. Nerve ring lying be- tween anterior 15,.0-18,3% of oesophagus. Excretory system H-shaped, anteriorly and posteriorly bilateral, with excretory nucleus about half way between nerve ring and ventriculus, excretory pore immediately posterior to nerve ring. Male. Body 27.5-5S4,2 mm long, 517-1109 maximum width; width at ocsophageo-intestinal junction 414-790; ratio of greatest width to length 1: 45.9-60,.7 (mean = $0.51). Dorsal lip 174-202 long, 179-188 wide (2 specimens); sub- Ventral lips 221-244 long, 216-273 wide (3 specimens), Nerve ring 498-846 from anterior. Excretory pore 611-893 from anterior (3 specimens). Oesophagus 2914-4418 long by 150-244 wide. Ventriculus 141-259 long by 89- 165 wide; ventricular appendix 3948-10058 long by 47-108 wide. Intestinal caecum 1692-2914 long by 169-188 wide; 58.1-75.6% (mean = 65.7%) ocsophageal length (3 specimens). Ejaculatory duct 1880-2350 long, 3.6-5.0% BL. Spicules 1222-2820 long, 112.1-150.0% ED (3 specimens). Caudal papillae pairs 49-66, chang- ing from button to mamillate at about 6th anterior to cloaca, Post cloacal pad absent. Tail 118-188 long, ventrally flexed. Female. Body 48.3-66.8 mm long, 1034-1410 maximum width; width at oesophageo-intestinal junction 470-931; ratio of greatest width to length 1: 45.0-54.7 (mean = 41.22). Dorsal lip 221-230 long by 216-273 wide (3 specimens); subventral lips 226-244 long by 212-282 wide. Nerve ring 658-846 from anterior. Excretory pore 952-912 from anterior. Oesophagus 366A- 5640 long by 216-310 wide. Ventriculus 188-376 long by 197-329 wide (3 specimens); ventricular appendix 5734-8930 long by 113-179 wide (3 specimens). Intestinal caecum 1974-3666 long by 235-282 wide; 51.9-65.0% (mean = 57.5) oesophageal length. Vulva opening 21.7- 30.0 mm or 32.4-48.0% BL from anterior; vulva area nol swollen. Vagina 5.60-8.84 mm (3 specimens); ulerus undivided for 5.84-10.0 mm, divided for further 6,18-7.10 mm. Tail 335-536 long. Variation, Several minor variations were 392 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 2. Hysterothylacium leptaspi n. sp. All figs. d #3 except where indicated. A, anterior end (scale 200 um); B, dorsal lip (100 zm); C, subventral lip (100 jum); D, subventral interlabium (100 jm); E, ventriculus and ventricular appendix (0.50 mm); F, caudal area, d #8 (0.5 mm); G, tail, ventral view, ¢ #3 (100 wm); H, emergent spicule, 6 #1 (200 um). ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 393 Fic. 3. Hysterothylacium leptaspi n. sp. A, tail, laleral view, holotype (scale 100 jzm); B, tail, lateral view, 2 #7 (200 jum); C, vagina and uterus, 9 #6 (3.0 mm), arrow indicates point of division of uterus. Sections (all male): D, nerve ring (100 xm); E, excretory duct (100 jum); F, excretory commissure (100 um); G, excretory nucleus (100 zm); H, ventriculus (100 jum); 1, ventricular appendix (100 zm); J, tail (0.5 mm). 394 MEMOIRS OF THE QUEENSLAND MUSEUM iSKVY S8us BB18 Fic. 4. Hysterothylacium leptaspin. sp. Scanning electron micrographs. A, en face; B, dorsal lip; C, interlabium,; D, postcloacal papillae; E, precloacal papilla, #6; F, precloacal papillae, c. #20. noted. The ventricular appendage is extremely long, and it is likely that the shorter lengths quoted here may be underestimated due to breakage. Lip width varied between being slight- ly wider and slightly shorter than long; this varia- tion is entirely due to perspective when measured as the lateral flanges curve away from the point of view. The smallest male worm (27.5 mm) had the largest number of papillae pairs (66). The largest male was proportionally the most slender. One male had a single double postcloacal papilla (Fig. 4D). Hosts Presently recorded from Arius leptaspis and two single specimens only from Arius thalas- sinus (Rippell). ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 39. DISTRIBUTION. Alt presen! known only fram Embley Estuary, Weipa, Queensland, but see ‘Remarks.* REMARKS Al a species Jevel there are problems posed in the determination of this species. Hysterothylacium arii (Yamaguti, 1954) was taken from an unidentified species of Arius from Sulawesi, There are several points of agreement with Yamaguti’s (1954) description: long caecum, very long ventricular appendage; postlabial grooves; lip shape; tail apex. There are equally several points of distinction, Yamaguti’s species being described as having alae, longer spicules, and fewer caudal papillae. Without ac- cess to Yamaguti’s type material (see Bruce and Cannon, 1989), the recorded differences force the conclusion that the two species are distinct. Amplicaecum indica Srivastava and Gupta, 1975, is a more recently described species from Arius venosus from India. The description con- sists largely of family characters, and the figures do notillustrate in detail the characters needed to assess the correct generic position of the species, or to compare with C, leptaspis, Srivastava and Gupta (1975) state clearly that their single male specimen was ‘without posterior bulb’ and there- fore also without a ventricular appendage. These authors did not state the disposition of their material and further comparisons between these apparently similar species cannot be made. In view of the stated difference (lack of ventriculus), (he two species cannot be con- sidered congeneric. Australia has over 18 species in the catfish family Ariidat (Kailola and Pierce, 1988) and a further 130 species are known world wide (Sands, 1985), With little data on host specificity it is not possible to assume that warms from congencric hosts are the same species or belong to the same genus. In Australia at Jeast four species of ascaridoid have been recorded from Arius species. It should be noted that the ventriculus and Ventricular appendage in this species is difficult to observe, and in all cases dissection was neces- sary. The ventriculus is small and clear, and mer- pes gradually into the ventricular appendage. The ventricular appendage is very flat (Fig. 31) and transparent (in section it appears hollow), and appears to adhere closely to the lateral cord, being only clearly visible where gently separated away, Hysterothylacium leptaspi is readily distin- guished from all other species of the genus by w complete lack of alae, and from those species for which the excretory system has been described by having an distinctly H-shaped excretory sys- tem. Hysterothylacium most commonly have a purely unilateral system, sometimes retaining traces of the left posterior flament. The ex- cretory system of the type species H. brachyurum Ward and Magath, 1917 remains undescribed, Hysterothylacium chrysostomi n. sp. Fig. § MATERIAL EXAMINED Male HOLOTYPE (OM GL10251), [emale PARATYPE (QM GI1.10252) off Dangara, Western Australia, 10 Dec. 1987, slomach of Lethrinus chrysostomus, coll A, Williams, TYPE LOCALITY Dongara, Western Australia TYPE HosT Lethrinus chrysosiomus Richardson, Lethrinidae, DIAGNOSIS Cuticlé annulated. Lips with length to width ratio 1, 1.2-1.7, with deep postlabial grooves; laterally constricted at about 0.3 length from anterior. Alac present, not expanded, originate just posterior to hase of subventral lips, Spicules 4.3-4.5% BL: ratio of 1: 1.02. Caudal papillae: ee peoclocal 30-32, patacloacal 2, postcloa- cal 9-10, DESCRIPTION Based on male holotype and female paratype. General. Body reaching greatest width anterior to mid-body. Dorsal lip smaller than subventrals, shorter than wide (0.63); flanges widest at posterior third of lip. Interlabia large, lateral margin weakly convex, basally wider than Jong. Ocsophagus 8.9-10.8% BL. Ventriculus slightly narrower than widest level of oesophagus, Jonger than wide. Ventricular ap- pendix 33.9-38.8% length of oesophagus. Nerve ring lying between anterior 21.4-22.5% of oesophagus. Excretory system with pore to just posterior to nerve ring, Male. Body 21.4 mm tong, 470 maximum width; width at ocsophageo-intestinal junction 282, Ratio of greatest width to length 1: 45.6, Subventral lip 78 long by 91 wide. Nerve ring 517 from anterior, excretory pore 696 from anterior, Oesephagus 2303 long by 188 wide, 396 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 5. Hysterothylacium chrysostomi n. sp. All figs of male holotype except where indicated. A, anterior (scale 200 jm); B, subventral lip (50 zm); C, dorsal lip, 2 (50 jm); D, interlabium, ? (50 jm); E, ventricular region (200 jm); F, tail, lateral view (100 jm); G, tail, 2 (100 jum); H, caudal region (0.50 mm). ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH Ventriculus 165 long by 188 wide; ventricular appendix 893 long by 85 wide, posterior half wider than anterior. Intestinal caecum 348 long by 141 wide; 15.1% ocsophageal length. Ejaculatory duct 2350 mm long, 11.0% BL. Spicules 940-959 long, 40.0-40,8% ED. Caudal papillae pairs 42-45, becoming mamillate at about 6th or 7th anterior to cloaca. No double papillae discerned in lateral view. Tail 165 long, withweakly nodulose truncated mucron. Female. Body 29.7 mm long, $517 maximum width; width at oesophageo-intestinal junction 385; ratio of greatest width to length 1; 57,4. Dorsal lip 94 long by 150 wide; subventral lip 127 long by 160 wide. Nerve ring 564 from anterior. Excretory pore 658 from anterior. Oesophagus 2632 long by 188 wide. Ventriculus 169 long by 160 wide; ventricular appendix 893 long by 86 wide. Intestinal caecum 423 long by 179 wide; 33.9% oesophageal length. Vulva opening 11,0 mm from anterior or 37.0% BL from anterior extremity; vulva area nat swollen. Vagina about (estimated) 940 long. Tail 400 long, With rounded minutely nodulose tip. Hosts Known only from the type host. REMARKS There are numerous species of Hysterothylacium and related genera described from the Indian and Pacifie Oceans that are in- adequately described (see Bruce and Cannon, 1989, for discussion). Of those that can reason- ably be inferred as belonging to Hysterothylacium only two seem close to the present species. Mysterothylacium epinepheli (Yamaguti, 1941) has a similarly shaped sub- ventral lip, but differs by having more numerous precloacal papillae, fewer postcloacal papillae, 4 Jonger ventricular appendage and a less strongly constricted lip. Similar differences also separate H. pagrosomi (Yamaguti, 1935) from H. chrysastami, ETYMOLOGY The epithet is taken from the host species epithe. Hysterothylacium sebae n. sp. (Figs 6-8) MATERIAL EXAMINED Male, HOLOTYPE, Bundaberg, eastern Queensland, 14 Jun. 1976, intestine of Lutjanus sebae (QM GL10253). PARATYPES 2 males, 2 females. same vila as holotype (QM GI,10254), TYPE LOCALITY Off Bundaberg, Queensland; no more precise locality data was. available, Ty Pr Host Luljanus sebae (Cuvier), Lutjanidae. DIAGNOSIS Cuticle distinctly annulated, Lips with length to Width ratio af 1: 0.72-0.80, with indistinet postlabial groove, Anterior half of lip ap- proximajely rectangular; Manges approximately triangular, anterior margin thickened, Cervical alae not evident; posteriorly alae distinct. Caudal papillae pairs: precloacal 19-20, paracloacal | double, postcloacal 2. Medioventral precloaval papilla present. Postcloacal pad absent. Spicules approximately equal in length, 1,8-2.9% BL. Tail apex acute, unornamented, DESCRIPTION Based on 2 mature males and 2 mature females, General. Body thickest in middle half, anteriorly and posteriorly slender. Dorsal lip slightly shorter than subventrals, shorter than wide (0,72 width); flanges widest at posterior 0,28-0.30 of lips; lateral constriction not strongly developed. Interlabia sessile, short. Oesophagus 9.6-14.9% BL, Ventriculus narrower than widest level of ocsophagus, narrower than long. Ventricular appendix 15.5-17.3% oesophageal length. Nerve ring lying between anterior 14.9- 17.3% of oesophagus, Excretory system with pore opening posterior to nerve ring, otherwise unobserved, Male. Body 38.5-39.2 mm, 771-799 maxi- mum width; width at oesophageo-intestinal junc- tion 508-470; ratio of greatest width to length 1: 49-50. Dorsal lip not measured. Subventral lips 146- 188 long by 189-235 wide. Nerve ring 658 from anterior, Excretory pore 752 from anterior (one specimen). Oesophagus 3807-4277 long by 301-348 maximum width. Ventriculus 282-301 long by 244-273 wide; ventricular appendix 658- 733 long by 56 wide, Intestinal caecum 226-282 long by 197-207 wide, 5.3-7.4% oesophagus length. Ejaculatory duct 2256-2444 tong, 5.9- 6.2% BL. Spicules 658-1128 long, 29.2-45.3% ED. Caudal papillae pairs 22-23, changing to mamillate at about 5th anterior to cloaca, Tail 207-226 long, spex acute, unomamented. 398 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 6. Hysterothylacium sebae n. sp. A, anterior, holotype (scale 200 j1m); B, subventral lip, holotype (100 zm); C, dorsal lip, 2 #1 (100 xm); D, subventral lip, 2 #1 (100 ym); E, subventral interlabium, 2 #1 (100 um); F, ventricular region, holotype (200 jm); G, tail, ventral view, ¢ #2 (100 ym); H, caudal region (0.50 mm). ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 399 Fic, 7. Hysterathylacium sebae n. sp. A, tail, lateral view, holotype (scale 50 um); B, tail, lateral view, 2 #2 (100 jum); C, vagina and uterus, 2 #1 (1.0 mm), arrow indicates point of division of uterus. Female. Body 42,5-45.2 mm long, 752-818 maximum width; width at oesophageo-intestinal junction 564-658; ratio of greatest width to length 1:60.0. Dorsal lip 160 long by 221 wide (one specimen). Subventral lips 165-174 long by 211-226 wide, Nerve ring 658-667 from anterior. Excretory pore not located. Oesophagus 4234- 4418 long by 320-404 maximum width. Ventriculus 329-376 wide by 282- 338 long; ventricular appendix 658-752 long by 66-69 wide. Intestinal caecum 226-329 long by 179- 216 wide, 5.1-7,8% oesophagus length. Vulva opening 13.5-14.8 mm or 31.8-32.8% BL from anterior. Vagina 1320-1500 mm long; uterus 3,67-4.51 mm long, dividing 4.98-6.02 mm from vulva. Eggs not seen. Tail 376- 409 long, apex acute, unornamented. Hosts Known only from type host. DISTRIBUTION Known only from the type locality. REMARKS Distinctive characters shown by this species are the shape of the dorsal lip, lack of obvious cervical alae, and the distinctive tail shape with an unornamented apex. Recent work (Bruce and Cannon, 1989) indicate that the morphology of alae is constant within a species of a sibling complex, such as the species of Maricostula, once the species have been discriminated. At present there is no species of Hysterothylacium that shows any degree of close similarity to this species. To my knowledge no species of Hysterothylacium has been specifically recorded from Lutjanidae. Scanning clectron micrographs show cuticular features not observed by light microscopy. The tail, while having a simple tip has a nodular patch on the ventral side (Fig. 8B). Immediately posterior to this terminal ‘cone’ or ‘cactus’ is an area in which striae form ridges; the dorsal sur- face of the cuticle anterior to cloaca has fine ridges, but these do not constitute crests in the sense of Bruce and Cannon (1989). Unfortunate- ly the material examined was in only adequate condition and it was not possible to examine the anterior of the specimen. Similarly the fixation was not of a standard that would have allowed interpretable sections. 400 ETYMOLOGY Named after the host species. Hysterothylacium tasmaniense (Johnston and Mawson) n. comb. ( Figs 9-11) Contracaecum (Thynnascaris) tasmaniense Johnston and Mawson, 1945: 134, Figs 24-27,.- Korotaeva MEMOIRS OF THE QUEENSLAND MUSEUM and Leont’eva, 1972: 404; Deardorff and Over- street, 1981: 1044, Contracaecum tasmaniense.- Mozgovoi, 1953: 229; Yamaguti, 1961b: 30. Thynnascaris tasmaniense.- Beumer et al., 1982: 22. MATERIAL EXAMINED 8 males, 5 females, one 4th, from Tamar River (estuary presumably), Tasmania, 30 Nov. 1983, from FiG. 8. Hysterothylacium sebae n. sp. Scanning electron micrographs. A, tail apex, cactus; B, cactus lateral view; C, double papilla; D, amphid (positioned dorsal to ala); E, precloacal papilla, #6; F, tail, lateral. ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 401 Tr’ Hh i“ } | ) ted Fic. 9. Hysterothylacium tasmaniense. Figs all ¢ #1 except where indicated. A, anterior 2#1 (scale 100 um); B, dorsal lip, ¢#2 (75 jm); C, subventral lip (100 zm); D, ventral interlabium (100 jm); E, ventricular region (200 um); F, spicule apex, d #4; G, caudal region (300 xm); H, tail, lateral view (100 jz); I, tail, ventral view, d#2 (100 pm). 402 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 10. Hysterothylacium tasmaniense. A, tail, lateral view, 2 #1 (scale 100 xm); B, vagina and uterus, 2 #3 (0.50 m). Sections (scales all 100 xm): C, nerve ring; D, excretory duct; E, excretory nucleus; F, mid oesophagus; G, ventriculus; H, anterior ventricular appendix; I, posterior ventricular appendix. Fic. 11. Hysterothylacium tasmaniense. Scanning electron micrographs. A, anterior and subventral lip; B, dorsal lip; C, interlabium; D, cuticular rings, detail; E, cactus; F, tail; G, postcloacal papillae; H, precloacal papilla #12. ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 403 sng Pseudophycis barbata (AHC 16494, one pair and 41h QM GL10255). TYPES Johnston and Mawson (1945) did not specify types, Their syntype material was reported to beheld at the South Australian Museum, Adelaide, but could not be located, TYPE LOCALITY The material examined by Johnston and Mawson was all from one station. This then is the type locality with the co- ordinates 42°40,0'S, 148°27.5'E. TyPE Host The original description was based on material from two species of host, and neither was designated as type host. DIAGNOSIS Culicle. with prominent cervical rings. Lips with length to width ratio 1: 1.0-1.3 (mean = 1.10), posteriorly pedunculate; without postlabial grooves; weakly constricted about 0.3 length from anterior; flanges forming broadly rounded points. Alae not evident anteriorly; alal grooves run from base of each subventral lip, become evident as alae towards posterior. Caudal papillae pairs: precloacal 18-25, paracloacal 1, postcloacal 4-5 with 2nd doubled. Spicules ap- proximately equal in length, ratio of 1; 1.03-1.08; 4.9-6.4% BL. Tail narrowing evenly, apex covered with fine nodules. DESCRIPTION Based on 6 mature males and 3 malure females. General. Body reaching greatest width about midbody. Dorsal lip slightly smaller than sub- ventrals, usually slightly shorter than wide (0.75- 0,83, 2 specimens); flanges widest at posterior 0,3 of lip; lips laterally constricted about anterior 0.3 of lip, Interlabia triangular, sessile, about twice as wide basally as long, esophagus 7,88- 11.4% BL. Ventriculus narrower than widest level of oesophagus, slightly longer than wide. Ventricular appendix 21.3-47.1% length of oesophagus. Nerve ting lying between anterior 18.3-21.7% of oesophagus. Excretoary system wilh pore immediately posterior to nerve ring, posteriorly unilateral, without anterior filament; excretory nucleus immediately postenor to com- missure. Male. Body 18.2-40.0 mm long, 376-799 max- imum width; width at ocsophageo-intestinal MEMUOIBS OF THE QUEENSLAND MUSEUM junction 282-461; ratio of greatest widih to length 1: 48-79 (mean = 1; 60). Dorsal lip 132 long, 165 wide (1 specimen); subventral lips 193-150 long by 103-165 wide, Oesophagus 2068-3243 long by 122-235 wide. Ventriculus 103-141 long by 103-165 side; ventricular ap- pendix 555-1269 long, 28-103 wide; 21.3-29.5% (mean = 26.8%) oesophageal length. Intestinal caecum 611-799 long, 75-113 wide, 23.7-33.1% (mean= 26.4%) oesophageal length. Ejaculatory duct 5,9-7.9% BL(mean=6.5%). Spicules 1504- 2350 long, 63.3-97.4% ED. Caudal papillae pairs 26-31, changing from button to mamillate at about 9th anterior to cloaca. Tail 127-188 long, narrowing evenly, terminating in minutely nodulose apex. Medioventral papillated precloa- cal organ present; posterior of anus elevated. Female, Body 22,0-35,0 mm long, 376-611 maximum width; width at oesophageo-intestinal junction 282-517; ratio of greatest width to length 1: 55.4-62.1 (mean = $9.4). Dorsal lip 89 long by 118 wide (one specimen); subventral lips 99-141 long by 103-160 wide. Nerve ting 446- 634 from anterior, Excretory pore 517-658 from anterior. Oesophagus 2162-2914 long by 141- 207 wide. Ventriculus 89-235 long by 99-179 wide; Ventricular appendix 611-799 long by 28- 66 wide; 22.5-28.3% (mean = 26.1%) oesophageal length. Intestinal caecum 630-893 long by 75-132 wide; 26.5-30.6% (mean = 28.9%) Oesophageal length. Vulva opening 8.8- 11,7 mm or 31,5-38.6% BL from anterior ex~ lremity; vulva area not swollen, Vagina short, 1-2 mm in length, ovaries not extending beyond vulva; uterus divides 2.7 mm from vulva {smal- lest specimen). Eggs about 56 in diameter. Tail 282-846 long, tapering gradually, apex minutely nodulose. Variation. All but one male had (he spicules shorter than the ejaculatory duct, In the differing male both spicules were about 130% the length of the ejaculatory duct. This male also had an abruptly narrower head than neck. In one specimen the ventricular appendage was markedly longer, 47% oesophageal length. The ventricular appendage and intestinal caecum are of about equal length, with the caecum usually (5 of 9) slightly longer than the ventricular ap- pendage. HosTs Present material is from Pseudophycis bar- bata Giinther (Moridae, Gadiformes). Johnston and Mawson’s (1945) record was fram Coelor- hynehus australis (Richatdson) (Macrouridae, ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH also Gadiformes) and Notopogon lillei Regan (Macrorhamphoridae, Syngnathiformes). Korotaeva and Leont’eva (1972) recorded Macruronus novaezelandiae. DISTRIBUTION Type locality, New Zealand (Korotaeva and Leont’eva, 1972) and now from the Tamar River, Tasmania. REMARKS Johnston and Mawson (1945) described Hysterothylacium tasmaniense from immature females. Their illustrations were few, but there are several points of correspondence between present material and the original description. Lip shape, lack of distinct postlabial grooves, inter- labium shape, the relative proportions of the intestinal caecum and ventricular appendage and the prominent cuticular rings all serve to identify this species. The most similar species to Hysterothylacium tasmaniense appears to be Hysterothylacium aduncum. Descriptive accounts and figures for this species are given by Punt (1941), Petter (1969), Petter and Maillard (1988). Berland (1961) give a detailed account of that species. Points in common that suggest that the two species are closely related are the lip shape, relative proportions of the intestinal caecum and ventricular appendage, papillae details, shape and ornamentation of the tail. Although there are inconsistencies in descriptions of material pre- viously recorded as H. aduncum, H. tasmaniense can consistently be separated by the lack of cer- vical alae, wider interlabia, conspicuous cervical tings, lack of postlabial grooves and more ob- viously constricted lips. The prominent cuticular rings give this species a superficial resemblance to the monotypic genus /heringascaris Peirera. In addition to the generic characters, the cuticular rings are them- selves different between the two species, those of /heringascaris overlapping posteriorly. Hysterothylacium thalassini n. sp. (Fig. 12) MATERIAL EXAMINED All specimens from Albatross Bay, Weipa, Q., Gulf of Carpentaria, March 1987, from stomachs of Arius thalassinus trawled at depth of 36m, coll. CSIRO. Male HoLotyPre, (QM GL10256), 6 females, PARATYPES QM GL10257, GL10258). 405 TYPE LOCALITY Albatross Bay, Weipa, Queensland. Gulf of Carpen- taria, 1240'S, 14142’E. Tyre Host Arius thalassinus (Riippell), Ariidae. DIAGNOSIS Cuticle distinctly annulated; lips with length to width ratio 1:1.05-1.23, with postlabial grooves, constricted a little less than one third (0.29) length from anterior. Alae originate just posterior to subventral lips. Ventricular ap- pendage about 70% length of oesophagus, nearly as wide as oesophagus. Caudal papillae: precloa- cal 16, paracloacal 1, postcloacal 2. Spicules approximately equal in length, ratio of 1:1.11, 5.48-4.96% BL. Tail with conical nodulose apex. DESCRIPTION Based on one mature male and 4 mature females. General. Body slender, about greatest width mid-body. Dorsal lip slightly smaller and basally manifestly wider than subventrals, about as long as wide; flanges widest about half way along length of lip, weakly developed. Interlabia ses- sile, less than half as long as lips. Oesophagus 10.6-15.6% BL. Ventriculus slightly wider than widest level of oesphagus, about as wide as long. Ventricular appendix 67.1-90.0% length of oesophagus. Nerve ring lying between anterior 11.9-16.4% length of oesophagus. Excretory system unilateral, with pore opening distinctly posterior to nerve ring. Male. Body 18.0 mm long, 282 maximum width; width at oesophageo-intestinal junction 212; ratio of greatest width to length 1: 63.8. Dorsal lip 75 long by 71 wide. Subventral lips not measured. Nerve ring 400 from anterior. Ex- cretory pore 494 from anterior. Oesophagus 2444 long by 112 wide. Ventriculus 118 long by 85 wide; ventricular appendix 2115 long by 103 wide. Caecum 1788 long by 94 wide, 73.1% oesophageal length. Ejaculatory duct 1128 long, 6.27% BL. Spicules 893-987 long, 79.2-87.5% ED. Caudal papillae 16-19. Tail 106 long. Female. Body 19.4-27.1 mm long 320-470 maximum width; width at oesophageo-intestinal junction 216-348; ratio of greatest width to length 1: 44.2-72.1 (mean = 1: 58.3). Dorsal lip 66-99 long by 71-94 wide (2 specimens). Sub- ventral lips 85-116 long by 71-94 wide (2 specimens), Nerve ring 329-447 from anterior. Excretory pore 353-517 from anterior. 406 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 12. Hysterothylacium thalassini n. sp. A, anterior, holotype (scale 100 zm); B, dorsal lip, holotype (50 p.m); C, subventral lip, 2 #3 (50 xm); D, ventral interlabium, 2 #2 (50 jm); E, ventricular region holotype (0.5 mm); F, section, posterior to nerve ring (100 zm); G, tail, lateral view, holotype (50 j12m); H, caudal region, holotype (200 xm), ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH Oesophagus 2350-3290 long by 103-132 wide. Ventriculus 94-141 long by 194-146 wide; ventricular appendix 1598-2538 long by 85-113 wide. Caecum 1739-2632 long by 94-169 wide: 74.0-80.0% BL (mean = 76.3%). Vulva opening 7.51- 11.84 mm or 37.6-43.7% BL from anterior; vulya area not swollen, Vagina not measured accurately, approximately 1.3-2.2 mm long. Uterus didelphic, opisthodidelphic. Oviducts not extending anterior to vagina, Tail 235-282 long. Variation. There was notable variation in the length to width ratio, with one specimen being distinctly more slender (1: 71.2) than the others (1:44.2-60.5). One specimen had a subventral lip length to width ratio of 1:0.93, With the limited material at hand no other particular variations were noted. Hosts Known only from the type host. One female specimen from Nemipterus hexadon (Quoy and Gaimard) (Family Nemipteridae) was examined, and although similar, a positive determination could not be made- REMARKS This small species shows no particular affinity with any other member of the genus. The long and, compared to other species, wide ventricular appendage is a noteworthy character which, in combination with lip shape and caecal length, should ensure easy identification. The state of preservation of these specimens was such that detailed histalogy was not success- ful therefore the position of the excretory nucleus remains undescribed, Similarly it was not pos- sible to get accurate measurements of the female reproductive system nor obtain successful SEMs. ETYMOLOGY The epithet is taken from the species name of the type host. Hysterothylacium zenis (Baylis) (Figs 13-15) Contracaecum zenis Baylis, 1929: 547, Fig. 4.- Yamaguli, 1961b: 30. Contracaecum (Cantracaecem) zenis.- Mozgovoi. 1953: 168, Fig. 102. Contracaecum zenopsis Yamaguli, 1941: 366, Fig. 24, plate V. Figs. 27-30.- 1961b: 30. Contracaecum (Contracaecum) zenopsis.- Mozgovoi, 1953: 168, Fig, 103. 407 Hysterothylacium zenis.- Deardorff and Overstreet, 1981; 1042. Hysterothylacium zenopsis. Deardorlf and Over- streel, 1981: 1042. MATERIAL EXAMINED 8 males, 4 females, 3 fourths, off Townsville, Q., na date (but catalogue number implies circa 1982), from Zenopsis nebulosus, coll. J. Stevens, CSIRO (AHC 16332, one pair QM GL10259), Also examined: 4 syntypes. TYPES The syntypes of H. zenis, all female, are held at the BM(NH) 1934.9.29.51-54. The types of H. zenopsis are believed to be at the Meguro Parasitological Museum, Tokyo, where Yamaguti's nematode types were deposiled (see Bruce and Cannon, 1989). TYPE LOCALITY Baylis cited only ‘off South-west Africa, July 8, 1927’. TYPE Host Zeus capensis, Zeidae (Baylis, 1929), DIAGNOSIS Cuticle finely annulated, lips with length to width ratio 1;0,72-0.90: with deep postlabial prooves; constricted about 0.3 length from anterior; dorsal lip flange approximately rectan- gular. Alae originate from and are united with subventral interlabia; expanded to about position of ventricular appendage. Caudal papillae pairs: precloacal: 24-29, paracloacal 1, postcloacal 6-7 af which 4th or 5th from cloaca is doubled, Spicules subequal in length, ratio of 1: 1.05-1.17; 4.67-7.08% BL. Tail narrowing evenly to bluntly rounded apex, provided with small finely nodulose nipple like process. DESCRIPTION Based on 4 mature males and 2 mature females. General. Body reaching greatest width about mid-bady, Dorsal lip slightly larger than sub- ventrals, usually slightly shorter than wide (0.72- 0.81); flanges widest at posterior two thirds of lip; lips laterally constricted about anterior 0.34- 0.37 of lip. Ventral interlabium about 1.3-1.5 jong as basal width, Oesophagus 12,1-16.5% BL. Ventriculus narrower than widest level of oesophagus, wider than long, Ventricular appen- dix 25.0-34,3% length of oesophagus. Nerve ring lying between anterior 7.66-10.18% of 408 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 13. Hysterothylacium zenis. A, anterior, d #1 (scale 100 jum); B, dorsal lip, ¢ #1 (50 jm); C, dorsal lip, 2 #1 (SO am); D, spicule apex, d #2; E, subventral lip, d #1 (50 jm); F, subventral interlabium/ala, 3 #1 (50 jum); G, ventral interlabium, d #2 (50 xm); H, tail, ventral view, d¢ #2 (50 xm); I, tail, lateral view, d #3 (50 um). ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 409 Fic. 14. Hysterothylacium zenis. A, caudal region, d #3 (scale 200 jum); B, tail, 2 #1 (100 ym); C. ventricular area, d #1 (0.5 mm); D, ventriculus, 2 #2 (100 jum); Sections; E, posterior to nerve ring (100 jum); F, mid oesophagus (100 2m). oesophagus. Excretory system initially posteriorly bilateral with right canal terminating anterior to ventriculus, left canal persisting beyond ventriculus; pore opening immediately posterior to nerve ring (one specimen observed). Male. Body 21.5-32.5 mm long, 329 maximum width; width at oesophageo-intestinal junction 251-517; ratio of greatest width to length 1: 44.3- 65.4 (mean = 1: 56.12). Dorsal lip 113-122 long, 146-169 wide (two specimens); subventral lip 103 410 MEMOIRS OF THE QUEENSLAND MUSEUM ASCARIDOID NEMATODES TROM AUSTRALIAN DEMBRSAL. FISH long by 115 wide (one specimen). Oesophagus 46116-5076 long, 160-188 wide. Ventriculus 116- 166 long by 141-188 wide; ventricular appendix 1269-1316 long, 75-94 wide, 25.00-26.17% (2 specimens) oesophageal length. Intestinal caccum 3008-3572 long, 188-216 wide; 62.0- 71.0% (mean = 66.1%) oesophageal length. Eyaculatory duct 5.70-8.74% BL (mean = 7.52%). Spicules 1222-2115 long; 65.0- 87.6% ED (mean = 76.21%). Caudal papillae pairs 24- 32, changing from button to mamillate at about 7th anterior to cloaca. Tail 150-188 long. Female. Body 41,8-43.6 mm Jong, 743-846 maximum width; width at oesophageo-intestinal junction 470-536; ratio of greatest width fo length 49.4-57.9. Dorsal lip 132 long by 165 wide; subventral lip 141 long by 160 wide. Nerve ring 494-517 from anterior. Excretory pore not sighted. Ocsophagus 5075-5452 long by 141- 207 wide. Ventriculus 136-165 long by 141-207 wide; ventricular appendix 1410-1739 long by 66-94 wide; 25.9-34,3% oesophageal length. In- testinal caecum 3572-3854 long by 141-255 wide; 70,4- 70.7% oesophageal length. Vulva opening 21.34-21.76 mm or 49,6-52.1% BL form anterior extremity, vulva area not swollen. Vagina 1974-3729 long; uterus divides 4068 from vulva. Eggs 56-85 in diameter. Tail 282-320 long, apex with small, minutely nodulose nipple like process. Variation, One male had two double papillae on the right side of the tail, all others had only one. The lower figures given for the precloacal papillae may be too low as most specimens were tightly coiled and papillae obscured from view. The lengths for the vagina given for two females of similar body lengths appears dispurate. Ex- amination of further specimens is necessary to place more confidence on the range given, Hosts Recorded only from the Family Zeidae. Present material is from Zenopsis nebulosus (Temminck and Schlegel). Previous records are from Zeus capensis Valenciennes (type host, Baylis, 1929), and Zenopsis nebulosus from Japan (Yamaguti, 1941, 1961b). DISTRIBUTION, Japan (Yamaguti, 1941), South Africa (Baylts, 411 1929), and now eastern Australia. he un- published record of Brunsdon (1956) is not of this species. REMARKS Companson of the present material to the syntypes of Baylis (1929) reveal no points of contradiction. The shape of the dorsal lip in both cases corresponds exactly. There are two other inadequately characterised species of Hysterothylacium that have been recorded only trom zeids, Hysterothylacium baylisi (Yamaguti, 1941) can be separated by having only two postcloacal papillae pairs, fewer precloacal papillae, jonger spicules (potentially 9,.8-13.8% BL) and the vulva positioned at the anterior one third of the body. Hysierothylactum zenopsis (Yamaguti, 1941), the second species, is here placed in synonymy with 1, zenis. Yamaguti's (1941) description agrees in most respects (papillae pairs, spicules, position of vulva, tail apex, Ventricular appendage and intestinal caecum proportions), bul does not have good lip details, and does not mention alae. Yamaguti routinely neglected to mention alae in his desctip- tions, and while I have not been able to obtain his material, lack of mention in Yamaguti's descrip- tion cannot be taken to indicate absence, Several other congeneric species have also been recorded from zeids. These are Hysterothylacium aduncum (Rudolphi), H. clavatum (Rudolphi) and H. fabri (Rudolphi). All of these species lack the long expanded alae and massive subventral interlabia which charac- terise H. zenis, and all have prominently nodulose tail apices. Brunsdon (1956 unpublished) recorded this specics from Zenopsis nebulosus in New Zealand waters. This record is a misidentifica- tion and differs in several respects, most notably in the proportions of the intestinal caecum and ventricular appendage (about equal, versus ventricular appendage 25-40% caecal length) and in lacking prominent alae, lehthyascaris Wu, 1949 Ichthyascaris Wu, 1949: 53. DIAGNOSIS Male, Body elongate, reaching greatest width Fig. 15, Hysterothylacium zenis, Scanning electron micrographs, A, anterior, B, dorsal lip, C, en face; D, lateral interlabium; E, tail, lateral views F, til apex; G, precloacal papilla #10; Ef, postcloacal double papilla. 412 at about anterior one third of body length. Cuticle fincly annulated. Alae run entire length of body, anteriorly united forming flange cunning posterior to subventral lips; forming cordons on tail. Lips with rounded lateral margins, posterior- ly not defined; not pedunculate, without teeth and not laterally constricted; anterolateral angles cach with deep socket; pulp antenorly bilobed. Dorsal lip with 2 lateral double papillae; sub- ventral lips with lateral double papilla and anterolateral papilla with adjacent amphid. Inter- labia entirely absent. Ventriculus shorter than wide, with triradiate lumen; ventriculus in line with oesophagus, Ventricular appendage originates from middle of ventriculus, short (24.6-41.3% ocsophageal length for the species described herein), sac like, with longitudinal sep- tum, Intestinal caecum absent. Excretory system unilateral, filamentar, with excretory nucleus im- mediately posterior to excretory commissure: exerelory pore opens posterior to nerve ring. Gubernaculum absent. Spicules short (2,2-2.9% BL), alate. Medioventral precloacal papilla present. Tail narrows evenly, apex recurved and provided with fine nodules. Female reproductive system, Vulva at about anterior one quarter to one third of body, Vagina not distinctly demarcated from uterus, Uterus didelphic, opisthodidelphic, oviducts extending anteriorly to, bul nol beyond vagina. Tver SPECIES Ichthyascaris laphit Wu, 1949, by monotypy. Wu (1949) did mot state where the material he examined was held or to be deposited. ComPusI TION Ichithyascaris biwakoensis (Fujita, 1928) n, comb, f. chtirocentri (Yamaguti, 1935) n. comb., L fisherit (Hooper, 1983) 0. comb., EL gym- aceranige n. sp., £ lutjani (Olsen, 1952) n comb.,/. meditteraneus (Lébre and Petter, 1983) o. comb., £ siliagaides n, sp. and /. vicentei (Santos, 1970)n, comb. Allexcept /. biwakoensis are from marine hosts. REMARKS The genus Icitipascaris Wu, 1949, has received little attention since its inception, The major compilations of Mozgovoi (1953), and later Yamaguti (1%61b) did not include the genus. Hartwich (1957) placed the genus in synonymy with Raphidascaris, remarking thal the two char- acters Wu (1949) used to distinguish the genus were probably artefacts. Chabaud (1965) fol- MEMOIRS OF THE QUEENSLAND MUSEUM lowed Hartwich(!957), but Hartwich (1975) later did not include /ehthyascaris in the synonymy for Raphidascaris. Although the genus has been overlooked it is distinctive, and differs from Raphidascaris in detail of the lip morphology, a character not considered by Hartwich (1957), Removal of the misplaced species from Raphidasearis to lehthyascaris will allow for a clearer concept of both genera. This genus is readily churacterised by the simple lips, as long as wide, with rounded lateral margins. This is in great contrast to the lip mor- phology shown by the genera Hysterothylacium, Raphidascaroides and Raphidascaris, all of which have flanged lips with a clearly defined posterior border. The lack of an intestinal caecum, short ventriculus, short sac like ventricular appendage, caudal cordons and anterior alal form all serve to further distinguish this genus. Wu (1949) established the genus /ch- thyascaris with a brief illustrated diagnosis. Nonetheless, several critical diagnastic charac- ters arc figured or mentioned. These are; 1, the ventriculus and ventricular appendage; 2, lack of interlabia; 3, alac uniting forming a ventral flange; 4, lack of intestinal caecum; and 5, lips without flanges. While there is little doubt that the genus is valid and clearly distinct from Raphidascaris (with which it had been synonymised by Hartwich, 1957), the descrip- tion of the Lype species is insufficient in detail to allow recognition or clear separation from other species now being placed in /chthyasearis. The anterior alal morphology, with the alae uniting close the subventral lips is a character unique to this genus. It is primarily on the basis of this character that /cAthyascaris chirocentrt is transterred from Raphidascaris, Additional specics of /e/tthyasearis for which there was insufficient material for description were obtained from Macquaria colonorum (Giinther), Solea sp., and Atherinomorus ogilbyi (Whitley). These worm specimens are held in the collections of the Queensland Museum. RELATIONSHIPS The genus most similar to /eAthyascaris is Paraheterotyphlum Johnston und Mawson, 1948 (sce Sprent, 1978b for a detailed description). It differs ftom /ehthyascaris in possessing an intes- tinal caecum, lacking the distinct united anterior alae and lacking caudal cordons. Both genera share the characteristic lip shape, lack of inter- labia and ventricular morphology. Heterotyph- lum Spaul. 1927 (and see Deardorff and ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISIt Overstreet, 1981) js also very similar, but has lips that are much shorter, lacking the characteristic rectangular shape of I/chthyascaris or Paraheterotyphlum, The previously named species now assigned to Ichthyascarts had all (except Rephidascaraides Jiskeri) been placed in the genus Raphidascaris Railliet and Henry, 1915. Smith (1984) gave an excellent redescription of Raphidascaris acus (Bloch) the type species for the genus. In com- paring the lip morphology of A. acus to that of the species now transferred to Ichthyascaris it is evident that the species in question are generical- ly incompatible. This is further supported by differences in alal morphology, excretory system and caudal cordons. There are several other genera that show an affinity to Ichthyascaris. Most of these genera would have been placed in the Heterocheilinae (sensu Sprent, 1983) or the Goeziinae (sensu Gib- son, 1983). Within these groups they appear to form a group apart from the other genera, characterised by the unique morphology of the ventriculus and ventricular appendage. The ventriculus of these penera is about as wide as the preceding oesophagus and in line with the oesophagus. In those genera for which there are appropriate figures, the ventriculus still maintains the triradiate symmetry of the oesophagus. The ventricular ap- pendage originates from the middle of the ventriculus, in contrast to that of Hysterothylacium where the bulb like ventriculus gradually nar- rows to form the posterior ventricular ap- pendage. The genera forming this group are Raphidascaris, Heterotyphlum, Paraheterotyph- lum, Ichthyascaris, Sprentascaris Petter and Cassone, 1984 and probably also Alibagascaris Kalyankar, 1970. None of these genera have interlabia or teeth with dentigerous ridges. Sprentascaris and Raphidascaris have flanged lips. tt is unclear from Yamaguti’s diagnoses (1935, 1961b), or the species contained within the genus, quite where Raphidascaroides should be placed. Most of the genera mentioned above have nol been rediagnosed since their inception. In order to facilitate their clear discrimination from Jch- thyascaris it is necessary to attempt a redefinition that allows comparison to that given for Ich- thyascaris. To this end the genera Raphidascaris, Raphidascaroides, Heterotyphlumn and Paraheterotyphlum are tediagnosed with remarks at the end of the text. 413 Ichthyascaris fisheri (Hooper) n. comb. Raphidascaroides fisheri Hooper, 1983: 8, Fig, 4. MATERIAL EXAM[NED Holotype( AM W157123) and allotype (AM W16039), REMARKS Examination of the type material showed thal discrepancies exist between the material and the figures and description given by Hooper (1983). Hooper (1983) described the species as ‘with interlabia 13 the length of lips’ and “Lips with dentigerous ridges * The figures show distinct interlabia, and also what appear to be narrow lateral flanges on the lips. These character states are not present in the holotype and allotype. The lips are simple, without flanges, dentigerous ridges or interlabia, and conform entirely to the diagnosis piven here for [chthyascaris. The tail is typical of the shape shawn by the genus with the apex tumed up and with the alae forming cordons. The ala¢ are united anteriorly forming, a flange just posterior to the subyentral lips. Accordingly the species is here transferred to ichthyascaris. Ichthyascaris fisheri, recorded only from flathead (Platycephalidae), is easily separated from other Australian species of the genus by the lateral margins of the lips haying a small bulge posterior to the anterolateral sockets and by the very short intestinal caecum. Ichthyascaris gymoocraniae n. sp. ( Figs 16, 17) MATERIAL EXAMINED All taken from the northwestern side of Wistari Reef, Capricorn Group, Queensland, coll. N.L. Bruce and §, Cook, 2 males, 1 female, 21 Apr. 1988, intestine of Gynnecranius hitorguatus (HOLOTYPE mule, QM GLI0260, PARATYPFS QM GL10261), Female, 22 Apr, 1988, intestine of G. bitorquatus (PARATYPE QM GL10262). Female, 21 Apr. 1988, intestine of Lethrinus chrysoxtomus (sectioned, QM GL10263). TYPE LOCALITY Wistari Reef, Capricorn Group, Queensland, 23°26.5'S, 151°54,0°E. Type Host Gymnocranius biterquatus Cockerell, Lethrinidae. 414 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 16. Ichthyascaris gymnocraniae n. sp. Figs. of holotype except where indicated. A, anterior (scale 100 pm); B, dorsal lip ¢ #1 (50 wm); C, subventral lip, d #1 (50 jm); D, caudal region (200 xm); E, tail, lateral view (100 ym); F, ventricular region. d #1 (100 jm); G, ventriculus and appendix (100 zm); H, tail, ventral view, 3d #1 (50 ym). ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 415 Fic. 17. Ichthyascaris gymnocraniae n. sp. A, vagina and uteri, 2 #1 (scale 200 jzm); B, tail, lateral view, 2 #1 (100 xm); Sections (scales all 100 zm): C, nerve ring; D, excretory duct; E, excretory nucleus; F, posterior to excretory nucleus; G, ventriculus; H, mid ventricular appendix. DIAGNOSIS Cuticle finely annulated. Lips as long as, or slightly longer than wide, widest anteriorly. Alae run entire length of body. Caudal papillae pairs: precloacal 24-28; paracloacal 2; postcloacal 7-9, 3rd or 4th from posterior larger than adjacent papillae. Weakly developed medioventral precloacal organ present. Spicules subequal in length, ratio of 1: 1.07-1.08, 2.19-2.91% BL. Tail converging evenly to minutely nodulose apex. DESCRIPTION Based on 2 mature males and 2 mature females. General. Body reaching greatest width at anterior one third to one half of length, of moderately even width. Dorsal lip slightly wider than subventrals. Oesophagus 7.69-9.32% BL. Ventriculus narrower than greatest width of oesophagus, about one half to one third as long as wide (0.45-0.66). Ventricular appendix 24.59- 41.27% length of oesophagus. Nerve ring lying din between anterior 25.2-29.9% of oesophagus. Ex- cretory system with pore opening distinctly posterior to nerve ring. Male. Body 11.3-15.0 mm long by 188-291 maximum width; width at oesophageo-intestinal junction 118-249, ratio of greatest width to length 1: 51,6-60,1. Dorsal lip 61-70 long, 66 wide; subventral lips 66-67 long, 56-67 wide. Nerve ring 241-249 from anterior, Excretory pore 353-494 from anterior. Oesophagus widest at posterior one third, 1053-1194 Jong by 122- 160 wide. Ventriculus 80-85 long, 127-150 wide; ventricular appendix 259-385 long, 71 wide, 24.59-32.25% oesophageal length. Ejaculatory duct 799-1175 long, 5.33-10.4% BL. Spicules 306-353 long, 26.04-44.18% ED (mean = 34.85%). Caudal papillae pairs 33-38, changing from button to mamillate at 9th anterior to cloaca. Tail 127-160 long, apex minutely nodulose. Female. Body 11.6-18.0 mm long by 226-409 maximum width; width at cesophageo-intestinal junction 179-306; ratio of greatest width to length 1; 44,0-51.3. Dorsal lip not measured; subventral lips 56-80 long, 56-80 wide. Nerve ring 249-320) from anterior. Excretory pore 353- 470 from anterior. Ocsophagus 865-1222 long by 132-216 wide. Ventriculus 71-85 long by 108- 188 wide. Ventricular appendix 306-357 long by 75 wide. Vulva opening 2.12-3.57 mm or 18.23- 19.84% BL from anterior extremity; vulva area not swollen. Vagina 235-893 long, uterus divides 1834-2538 from vulva, divided branches short, 271-423 long. Oviducts not extending anterior to vulva. Eggs 38-47 in diameter. Tail 390-423 long, apex minutely nodulose. Variation. The most notable variation is thal of the absolute and relative lengths of the ventricular appendage ranging from 24.5%- 41.3% pvesophageal length. Hosts Presently recorded only from the family Lethrinidae: Gymnocranius bitorquatus and Lethrinus chrysostomus Richardson. DISTRIBUTION At present known only from the type locality. REMARKS This species is readily separated from most others of the genus by the lips being abruptly wider anteriorly. Ichthyascaris lutjani has similar lips, but has a slender ventricular ap- pendage, is much larger in size, has fewer MEMOIRS OF THE QUEENSLAND MUSEUM precloacal papillae (9-11 pairs vs 24-28 pairs) and fewer postcloacal papillae (3 pairs vs 7-9 pairs). The remaining species have the lips il- lustrated as anteriorly rounded, except that the shape of the lips is not known for Ff. chirocentri (Yamaguti). In/. chirocentrithe number of papil- lace (60) clearly separates it from /. gymnocraniae (33-38). For details on how to distinguisn L, gym- nocraniae from L. sillagoides see the “Remarks’ for the latter species. ETYMOLOGY The epithet is taken from the genus name of the type host. Ichthyascaris sillagoides n_ sp. ( Figs 18-20) MATERIAL EXAMINED All from Sillago maculata, Moreton Bay, southeastern Queensland, coll. G. Berry. Male, HototyPr, 17 Feb, 1981, Deception Bay (QM GL9260). PARATYPES: Female (sectioned), 17 June 1978, Deception Bay (QM GL18776); male, 29 Aug. 1982, (QM GL 9448); 2 females (one for SEM), 23 Apr. 1978, Deception Bay (QM GL8768); female, 23 Apr. 1978, Deception Bay (QM GL8769); female, 17 Jun, 1978, Deception Bay (QM GL9106); female, 29 Aug. 1982, (OM GL9447); female, 29 Aug. 1982, (QM GL9450); female, 29 Aug. 1982, (QM GL9451). Two immature specimens (OM GL9449, AHC 18814). TYPE Locality Deception Bay, Moreton Bay, southeastern Queensland. Tyre Host Sillaga maculata Quoy and Gaimard, Sillaganidae. DIAGNOSIS Cuticle distinctly annulated. Lips with length to width ratio of 1,07-1,18, widest anteriorly. Caudal papillae pairs: precloacal 22-26; paracloacal 1; postcloacal 8-10; medioventral precloacal papilla not discerned; no double papillae. Spicules of approximately equal length, ratio af 1: 1.09; 2.27-2.47% BL. Tail with recutved minutely nodulose apex, DESCRIPTION Based on 2 males and 5 females. General. Body of fairly even thickness. Dorsal lip slightly larger than subventrals; widest at ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 417 Fic. 18. Ichthyascaris sillagoides n. sp. Figs of holotype except where indicated. A, atiterior (scale 100 zm); B, dorsal lip, 2 #21 (50 jum); C, subventral lip, 2? #2 (50 wm); D, between lips, 2 #2 (SO wm); E, tail apex; F, caudal region (200 ym); G, tail, ventral view, d #2 (50 wm); H, ventricular area (100 um), 418 MEMOIRS OF THE QUEENSLAND MUSEUM FiG, 19. fchthyascaris sillagoides n. sp. A, tail, lateral view, holotype (scale 50 zm); B, tail, lateral view, 2 #1 (100 jum); C, tail, lateral view, 2 #3 (100 xm); D, tail apex, ° #1; E, tail apex, 9 #2; F, vagina and uterus, 9 #1 (0.5 mm). anterior margin, Oesophagus 5.91-8.16% BL. Ventriculus slightly narrower than widest level of oesophagus, shorter than long. Ventricular appendix 27.59-38.60% length of oesophagus. Nerve ring lying between anterior 24.83- 31.70% of oesophagus. Excretory system with pore opening distinctly posterior to nerve ring. Male. Body 9.5-14.2 mm long, 188-212 max- imum width; width at oesophageo-intestinal junction 155-165; ratio of greatest width to length 1: 51.0-67.0. Dorsal lip not measured. Subventral lips 47-71 long by 42-71 wide. Nerve ring 230-282 from anterior. Excretory pore 352- 376 from anterior. Oesophagus 729-1128 long by 94-113 wide. Ventriculus 52-61 long by 85-94 wide; ventricular appendix 282-447 long by 42- 80 wide. Testes extend anteriorly to vicinity of ventricular appendix, Ejaculatory duct 8.82- 12.89% BL. Caudal papillae pairs 31-37, chang- ing from button to mamillate at about 8th anterior to cloaca. Spicules 282-306 long 17.7-19.1% ED (one specimen). Tail 103-118 long, apex recurved. Female. Body 13.6-30.2 mm long, 216-498 maximum width; width at oesophageo-intestinal junction 179-320; ratio of greatest width to length 1; 48.4-67.8 (mean = 1: 59.7). Dorsal lip 94 long by 80 wide (one specimen); subventral lips 66-108 long by 66-94 wide. Nerve ring 291- 470 from anterior. Excretory pore 611-423 from anterior (3 specimens). Oesophagus 1109-1833 long by 80-188 wide. Ventriculus 66-103 long by 80-174 wide: ventricular appendage 306-611 long by 66-118 wide. Vulva opening 3.50-9.34 mm or 22.8- 29.0% BL from anterior extremity, vulva area not swollen, ovaries extending to vicinity of vulva. Vagina short 329-517, not abruptly differentiated from uterus. Uterus divides 1192-2350 posterior to vagina, branches 658-1880 long (2 specimens). Eggs 19-47 in diameter. Tail 301-479 long, posteriorly Trecurved, apex minutely nodulose. ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 419 Variation. The two largest female specimens Hosts had noticeably thicker tails, while the degree of Known only from the type host. nodule development on the tail apex also varied. Lip shape varied with regard to degree of inden- tation of the anterior margin (e.g. comparing Fig. DIsTRIBUTION 18B to Fig. 18C) but this may be attributable to Known only from localities in Moreton Bay, angle of observation. southeastern Queensland. Bum FiG, 20, Ichthyascaris sillagoides n. sp. Scanning electron micrographs. A, en face; B, dorsal lip; C, between lips; D, subventral ala; E, caudal cordon; F, cactus. REMARKS Ichthyascaris gymnocraniae and J. sillagoides are superficially very similar. There are how- ever numerous differences, especially scxual characters, and these are sufficient to consider the two species as distinct, These differences are in J. sillavoides: spicules 17.7-19.1% ED (vs 26.1-44.2% ED inJ. eymnecraniae); ejaculatory duct 8.8-12.9% BL (vs 5,3-10.4% BL); precloa- cal papillae 22-26 pairs (vs 24-28 pairs); vulva 22.8-29.0% BL from anterior (vs 18.2-19.0% BL); and uterine branches long, 658-1180 (vs short 270-423). Ichthyascaris sillagoides can be distinguished from other species by the Jesser number of precloacal papillae, except for L. weentet and L. mediterraneus from which it differs by having more postcloacal papillae. Khan and Yaseen (1969) described a worm from Sillaginopsis panijus using the combina- tion Raphidascaris panijii Khan and Yaseen, Smith (1984) considered this species as species inguirenda. While the description is not detailed, and the species cannot be reliably assigned to any genus, the very long ventricular appendage sug- gests that it is not conspecific with the material from Sillago maculata described here. ETYMOLOGY The epithet is derived from that of the host genus. REDIAGNOSED GENERA In order to enable comparisons to be made between Jchthyascaris and related genera il is necessary to rediagnose the following genera in as much detail as possible. Without recourse to fresh material, type material, or in some cases a good description, the information contained within these diagnoses is inevitably uneven in detail. Nonetheless these diagnoses are here at- tempted to allow the clear separation of/chthyas- caris from related genéra, A second purpose is to draw attention to these genera and highlight the need for additional descriptive data. In this regard il is Raphidascaroides that is most defi- cient in detail, and will remain so until the types or fresh specimens of the type species are redescribed. Heterotyphlum Spaul Heterotyphlam Spaul, 1927: 634,.- Mozgovoi, 1953) Hartwich, 1957: 238; Yamaguli, 1961b: 31; Dear- dorff and Overstreet, 1981: 4231. MEMOIRS OF THE GUEENSLAND MUSEUM DIAGNaSIS Body elongate, anterior half manifestly nar- rower than posterior half; cuticle finely annu- lated, male with fine crests. anterior to cloaca, Alae narrow, originate. close to subventral lips. run entire length of body. Lips about half as long aS wide, anterior rounded, without anterolateral sockels or denligerous ridges; without Lateral flanges and constrictions, posteriorly not defined. Interlabia entirely absent. Dorsal lip with two lateral double papillae; subventral lips with one double papilla and anterolateral single papilla with adjacent amphid, Ventriculus shorter ihan wide, in line with oesophagus. Ventricular appendage long {(c, 62-72% lengih of ocsophagus) anterior half slender, posterior half expanded, sac like. Intestinal caecum present. Exeretary pore opens near to nerve ring (ex- eretory system otherwise not described). Guber- naculum absent. Spicules short (c. 2.2% BL), alate, Tail broadly rounded, with simple mucron. Female reproductive system. Vulva situated in anterior half of body (38% BL. from anicriar), didelphic, opisthodidelphic. Oviducts not ex- tending anteriorly to Vagina, Tyré SPeciis Heterotyphium himantelaphi Spaul, 1927, by monotypy [BM(NH) 1927.7.22 31-38]. REMARKS The type species of this genus is still known only from the original material. Of the other species placed in the genus only one has been adequately described and that species has been placed in Hysterothylacium by Deardorff and Overstreet (1981), As indicated by Deardorff and Overstreet (1981) the remaining species are in need of redescription before their generic posi- tion can be accurately determined. The morphology of the ventriculus and lips. indicates that Heteroryphlum is most closely re- lated to Paraheterotyphlum and ichthyascaris. Ichthyascaris can be distinguished by the longer Jips and lack of an intestinal caecum. Paraheterotyphlum by having long rectangular lips and a body of even width. The syntypes of H. himantolophi were ex- amined for purposes of the gencric diagnosis, and it was noticed that the male tail is provided with fine transverse crests anterior to the cloaca. These are not prominent as in the species of Maricostula figured by Bruce and Cannon (1989). ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH Species placed in the genus (Yamaguti 1961b), all of which should be regarded as species in- guirendae and incertae sedis are: H. cheni Hsu, 1957; H. multipapillosum (Skrjabin, 1916); and A, obtuscaudatum (Zeder, 1800). Paraheterotyphlum Johnston and Mawson Paraheterotyphlum Johnston and Mawson, 1948: 102.- Hartwich, 1957; 242; Yurnaguti, 1961b: 169; Schmidt and Kunz, 1973: 483; Sprent, 1978b: 164. DIAGNOSIS Body clongate, of even thickness. Cuticular annules not evident. Alae not cervically ex- panded; run entire length of body, originating posterior to subventral lips. Lips longer than wide, rectangular, not pedunculate, posteriorly pot defined; without dentigerous. ridges and lateral flanges, not laterally constricted; anterolateral corner each with socket. Dorsal lip with 2 lateral double papillae; subventral lips each with double papilla and mediolateral single papilla and adjacent amphid. Interlabia entirely absent. Ventriculus shorter than wide, with triradiate lumen; in line with oesophagus. Ventricular appendage long (35-58% oesophageal length). Intestinal caecum present. Excretory system filamentar, unilateral, nucleus near nerve ring; excretory pore opening immedi- ately posterior to nerve ring. Gubernaculum ab- sent. Spicules short (c. 2% BL), alate. Tail straight, narrowly rounded, apex minutely nodulose, Female reproductive system. Vulva in anterior one third of body, didelphic, opisthodidelphic; oviducts not extending anterior ta vagina. TYPE SPECIES Paraheterotyphlum australe Johnston and Mawson, 1948, by monotypy. REMARKS The gonus and its constituent species have been discussed in detail by Sprent (197&b), The most closely allied genus is /chthyascaris which has a near identical lip morphology. Paraheierotyphlum differs in having shallower lip sockets and an intestinal caecum. A further difference is that the two species of Paraheterotyphlum are far larger (61-158 mm) than most /chthyascaris (up to 30 mm) although size is not regarded as a character of generic significance. However /. lutjani is recorded 32- 72 mm which overlaps with that for 421 Paraheterotyphlum. eterotyphlum is distin- guished by its very much shorter lips. Two species from sea snakes are known, the type species and P. ophiophagas Schmidt and Kunz, 1973. Raphidascaris Railliet and Henry Raphidascaris Railliet and Henry, 1915.- Mozgovoi, 1953: 402; Hartwich, 1957; 237; 1974: 9; 1975; 101; Yamaguti, 1961b: 35; Yorke and Maplestone, 1926; 274. Chabaud, 1965: 904. Neogoezia Kreis, 1937: 129, DIAGNOSIS Male ({tom Smith, 1984). Body clongate, fine- ly annulated. Alae originate from between bases of subventral lips and run to middle of tail; ces- vically expanded, without caudal cordons. Lips with Jateral flanges, medially constricted; with postlabial grooves; pulp anteriorly bilobed. Dar- sal lip with two lateral double papillae; sub- ventral lips with double papilla and anterolateral papilla with adjacent amphid. Interlabia absent. Ventriculus shorter than long, in line (= cylindri- cal) with ocsophagus. Ventricular appendage with septum. Intestinal caecum absent. Ex- cretory system filamentar, posteriorly bilateral, with reduced right canal. Excretory nucleus dis- linctly posterior to commissure (Gibson, 1983, fig. 1, Cl). Excretory pore opening posterior to nerve ring. Gubernaculum absent. Spicules short, subequal, alate. Precloacal medioventral papilla present. Tail curving ventrally, apex nar- rowed, unornamented. Female reproductive system. Vulva opening hetween anterior one quarter and one third of body, Uterus didelphic, opisthodidelphic; oviducts not extending anterior to vulva. TYPE SPECIES. Ascaris acus Bloch, 1779, by original designa- tion, REMARKS Smith (1984) redescribed Raphidascaris acts in excellent detail with a full synonymy and also listed the species then placed in the genus, Of a total of LU species that Smith considered poten- tially valid, he regarded five as species inquiren- dae, All of (he remaining species, for which there were adcquate figures, are here transferred to Iehtlryascaris, As discussed for Raphidascarotdes the inter- pretation of interlabia is of greal importance im discriminating these two gencra. Two species of Raphidascarades (R. bishati and R. chilomyc- teri) differ from Raphidascerts only by having distinct (as figured) interlubia. Ye\, examining the SEM’s of Smith (1984) one sees a rudimen- tary interlabial knob (cf. Soleim, 1984) or what is formed by united postlabial grooves, Redescription and reassessment of interlabial morphology is necessary before generic reas- signment of those species can be undertaken. Alt present the type species is the only identifi- able species of the genus. Species of doubtful status are listed by Smith 1984, Raphidascaroides Yamaguti Raphidasceroides Y¥ ammaguii, 1941: 355,- 1961 by 36; Mozgovol, 1953: 418; Chabaud, 1965: 995; Hartwich, 1957: 239; 1974: 9. Ryjikovascaris Mozgovoi, 1950. 1953: 419. DIAGNOSIS (adapted and expanded from Yamaguti, 1941, 196)b). Body clongate, finely annulated. (AJae not described). Lips with lateral flanges, medially constricted, with dentigerous ridges; pulp anteriorly bilobed. Dorsal lip with two lateral double papillae; subventral lips with double papilla and anterolateral single papilla with ad- jacent amphid. Distinct interlabia present. Ventriculus shorter than long, with ventricular appendage. Intestinal caccum absent. Excretory pore opens posterior lo nerve ring (exeretory system otherwise undescribed). Gubemaculum absent. Spicules subequal in length, alate. Tail weakly curved ventrally, apex minutely nodulose, Female reproductive system. Vulva situated about one third of body length from anterior; oviducts not extending anterior to vulva. TYPE SPECIES Raphidascaroides nipponensis Yamaguti, 1941, by original designation. REMARKS Yamaguti (1941) established the genus for one species and a subspecics of the nominate type species, Later (Yamaguti, 1961a) he described a second species, nol mentioning the presence of dentigerous ridges. When redefining the genus (Yamaguti, 1961h), the diagnosis was modified by adding in a footnote ‘|dentigerous ridges] Absent occasionally’. Species were slowly added ta the genus, and by 1988 eleven species MEMOIRS OF THE QUBENSLAND MUSEUM had been assigned to the genus. Two characters are critical in the discussion of Raphidas- caroides: dentigetous ridges on the lips and in- terlabial morphology. Lip shape is a further character that is of significance. Of the species currently placed in the genus only those that Yamaguti (1941) originally placed in the genus and % fisheri Hooper, 1983 have been described as having dentigerous ridges. No species have had the dentigerous ridges figured. Examination of the type material of 2. fisheri failed to reveal dentigerous ridges, and the species has been transferred to Ichthyas- caris (see the species accaunt and ‘Remarks’ for that genus). Presence of a dentigerous ridge is a character that is consistent within genera, and it would seem unacceptable to have both states wilhin a single genus, Therefore those species without dentigerous ridges should be reassigned. Yamaguti's (1941, 19614, 1961b) descriptions clearly indicate that substantial interlabia were present in the species he examined, describing them as ‘conical’ (Yamaguti, 1941) and ‘nearly half as long.as lips’ (Yamaguti, 1961). In at least one species currently placed in the genus, A. africanus Khalil and Oyetayo, 1988, the SEMs show that interlabia, in the sense of Yamaguti (1941, 1961a) or as shown by Hysterothylacium and Maricostula, are absent. There is a problem here of intergradation of characters and of interpretation. Interlabial mor- phology has been discussed by Soleim (1984), who reiterated Berland’s (1961) distinction be- tween interlabia and semi-interlabia. In the genera here under discussion all interlabia are of the latter category. This interlabial form is not clearly defined, and when reduced causes problems in interpretation. In the genus /chthyas- caris, totally lacking postlabial grooves, the lack of interlabia is Unambiguous. In species with defined postlabial grooves, the continuation of the groove clearly gives the impression of an interlabium. This is clearly shown by Hysterothylaciurn tasmaniense where the inter- labial structure is sessile and in great corttrast to the form shown by Hysterothylacium zenis (Fig. 13G) or species of Martcostula Bruce and Can- non, 1989 (SEM’s). While it is pertinent there- fore to allempt differentiate between postlabial grooves, sessile interlabia and scmi-interlabia, it 1s Immediately obvious thal these first two states will intergrade, whereas the free standing ('non- sessile’) form is distinet. Of the species currently placed in Raphidas- caroides four are of uncertain status and those are ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH 423 regarded here as species inquirendae: Raphidas- earoides armatust Gupta and Srivastava, 1984; Raphidascaroides blochii Bilgees and Khanum. 1974; Raphidascaroides jazganathai Gupta and Srivastava, 1984; Raphidascaroides trachino- cephalaust Rajya Lakshmi et al, 1985. No detailed figures have been given for lip shape, and the species descriptions are not in sufficient detail to allow specific determination or genenc placement. Both Raphidascaroides diadonis (Thwaite, 1927) and R. africanus have a similar lip mor- phology. That is flanged lips which are posterior- ly narrowed (or angled) with a medial constriction, and lip papillac positioned anterior to the constriction. The recently described species Sprentascaris hypasiomi Petter and Cas- sone, 1984 has a cephalic morphology entirely similar to that of &. africanus and, contrary to generic diagnosis given for the genus (Petter and Cassone, 1984), is clearly figured with rudimen- tary interlabia (Petter and Cassone, 1984, fig. 6D). While not questioning the validity of Spren- tascaris it seems probably that the species men- tioned above should be placed within a separate genus, defined by their distinctive lip morphol- ogy. Ryjikovascaris Mozgovoi, currently placed in synonymy with Raphidascaroides, is an avail- able name but redescription of the type species R. diadonis is necessary before the validity of the genus can be reassessed. Both Raphidascaroides chilomycteri and R. bishaii are figured with prominent intertabia, and it 18 this character alone which separates them from Raphidascaris. The labial and interlabial morphology of (hese two species also needs to be reassessed. Al present the species are retained within their current combinations and other than those specics listed in the preceeding a8 species tn- quirendae the genus consists of: R. nipponensis, the type species and R. nipponensis lophii Yamaguti 1941, Also within the genus, but here regarded as incertae sedis are: Raphidascaroides bishatt Khalil, 1961, Raphidascaroides chilo- mycteri Yamaguli, 1961b, Raphidascaroides africanus Khalil and Oyetayo, 1988 and Raphidascaroides diadonis (Thwaile, 1927). Resolution of the generic concept of Raphidascaroides, which at present is very loose, and its constituent species can only come about with a detailed redeseription of the type species coupled with re-examination of the type specimens. Unfortunately the types are not avail- able for Joan, and it is not possible to add further to the descriptions of the type species. ACKNOWLEDGEMENTS This study was generously supported by a grant from the Marine Science and Technology Grants Scheme (now the Australian Research Council), together with supplemental funding from the Queensland Museum Trust. Essential material was provided by Dr S, Blaber (CSIRO, Division of Fisheries, Cleveland, Qld), Mr Alan Williams (Murdoch University, WA) and from Professor J.F.A. Sprent’s (Queensland Univer- sity) collection, now housed in the Queensland Museum. Material was borrowed from the col- lections of the South Australian Museum Adelaide for which | thank Dr D.C. Lee and Mrs P.M. Thomas (whom | also thank for their at- tempts to locate the types of H. tasmaniense), and from the British Museum (Natural History) for which ] thank Dr D.I, Gibson. J thank Dr Lawn and Ms M. Prekker and their staff for their assis- jance while collecting at Heron Island, anc Steyen Cook (QM) who ably assisted us tn the field. 1 also thank Mrs C, Lee (QM) for preparing serial sections, and Dr LR.G. Cannon for his comments on the manuscript, REFERENCES BAYLIS, H.A. 1923, Report on a collection of parasitic néematides, mainly from Egypt. Parasitology 15: (-13, 192%, Parasitic Nematoda and Acanthocephala co}- lected in 1925-1927. Discovery Reports 1: 54] - 560. BERLAND, B. 1961. Nematodes from some Norwegian marine fishes. Sursia 2) 1-50, BEUMER, J.P... ASHBURNER, L.D., BuRBURY, M.E., Jerré, E..AND LATHAM, D.J; 1982. A checklist of the parasites of fishes from Australia and its adjacent Antarctic Territories. 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AND PETTER, A.J. 1983 Deux nouvelles espéces d’ascatides (Nematoda) parasites de poissons téléostéens: Raphidascaris mediter- raneus 0. sp. el Goezia artguillae n. sp.; comple- ment de description de Cucullanus micropapillatus Tornquist, 1931 (Nematoda, Cucullanidae). Bulletin du Muséum National d'- ASCARIDOID NEMATODES FROM AUSTRALIAN DEMERSAL FISH Histoire Naturelle, Paris 4th ser. 5, sect A{2): 491-505, Mozcovai, A.A, 1950. On the anisakid (una of fishes and reptiles, Trudi Gel'mintologicheskoi Laboratorii Akademiya Nauk SSSR 3; 102-118 [in Russian]. 1953. Ascaridala of animals and man and the diseases caused by them. Gel'min- tologischeskaya Laboratoriya~ Osnovy Nematologii, Wolume 2, (Akademiya Nauk: Moskva). 617 pp. OLSEN, L.S. 1952. Some nematodes parasitic in marine fishes. Publications of the Institute of Marine Science, University of Texas 2; 173-215, Petter, A.J. 1969. Enquéte sur les nématodes de pois- sans de la region nantaise. Identification des lar- ves d’ Ascarides parasitant les sardines (en rapport avec les pranulomes eosinophiles observés de homme dans la region)Annales de Parasitologie Humaine et Comparée 44; 559- 580, Petrer, A.J. AND CASSONE. J. 1984. Nématodes de poissons du Paraguay; I. Ascaridoides: Spren- tascaris, 0. gen, Revue Suisse de Zoolagie V1: 617-634, PyrTerR, A.J. AND MAILLARD, C, 1988. Larves d’Ascarides parasites de poissons en Mediter- ranee occidentale, Bullerinda Muséum National d'Histoire Naturelle, Paris 4th ser., 10, sect, A(2); 347-369. Punt, A. 1941, Recherghes sur quelques nématodes parasiles de poissons de la Mer du Nord, Memoirs du Musée Royal d'Histoire Naturelle de Belgique 98: 1-110. RAILLIET, A. AND HENRY, A. 1915, Sur les nématodes du genre Goeziu Zeder, Bulletin de la Société de Pathologie Exotique 8: 270-275. Rarya LAKSHMI, 1, HANUMANTHA RAO, K. AND SHYAMASUNDARI, K, 1985, Raphidascaroides trachinocephaluis n, sp. (Nematoda | Heterocheilidae) from marine fishes at Visak- hapatnam, /ndian Journal of Parasitology ¥; 235-238. Rye, L.A. AND BAKER, M.R. 1984, Hysterothylacium analarum n. sp. (Nematoda : Anisakidae) from pumpkinsced, Leponis gibbosus (Linnaeus), in southern Ontarjo. Canadian Journal of Zoology 62: 2307-2312. Sanps, D. 1985. ‘Catfishes of the World, Volume 5, Bagridae and others’. (Dunure Enterprises: Ayr, U.K). 201 pp. Santos, E. Dos. 1970. Nova espécie do género Raphidascaris Railliet and Henry, 1915 (Nematoda, Ascaridoidea). Atas da Sociedade de Biologia do Rio de Janeiro 14; 39-40. 425 Scumipt, G.D. AND Kunz, R.E. 1973. Nematode parasites of Oceanica, XX Paraheterotyphlum ophiophagos n. 8p. (Heterocheilidae), trom the banded yellow-lip sea snake, Laticauda calubrina, American Midland Naturalist 89: 481-484, SKRJABIN, KI, 1916, [Contribution to the study of the helminth fauna of Paraguay], Zoologit Vestnik 1(4): 705-757. [In Russian]. SMITH, J.D. 1984. Taxonomy of Raphidascaris spp. (Nematoda, Anisakidac) of fishes, with a redescriplion of R. acus (Bloch, 1772), Canadian Journal of Zoalogy 62; 685-694, SocLeim, ©, 1984, A synopsis of the genera Thynnas- caris and Contracaecum (Nematoda, As- caridvidea) with an emendation of the generic definitions. Acta Parasitalogica Polonica 29: 85-96. SOLEIM, O, AND BERLAND, B, 198], The morphology of Thynnascaris udunca (Rudolphi) (Nematoda : Ascuridoidea), Zoologica Scripta 10: 147-182. Sraut, B.A, 1927, A new nematode of the subfamily Anisakinae, Annals and Magazine of Natural History (9)19: 627-635. SPRENT, J.F.A. 1978. Ascariduid nematodes of am- phibians and reptiles; Goezla. Journal of Hel- minthology $2; 91-98, 1978b, Ascaridoid nematodes of amphibia and reptiles: Paraheterotyphlum, Journal of Helmin- thology 52: 163-170. 1983. Observations on the systematics of as- caridoid nematodes. pp. 305-319. In ‘Sys- tematics Association Special Volume 22. Concepts |p Nematode Systematics’. Stone, A.R., Platt, H.M, and Khalil, L.F. (eds), (Lon- don; Academic Press). SRivaAsTAvA, A.B. AND GUPTA, S.P. 1975, On twa new species of the genus Amplicaecum Baylis, 1920 ftom marine fishes of Puri, Orissa, /ndian Journal of Helminthology 27: 119-123. THWAITE, J.W, 1927. On a collection of nematodes from Ceylon. The Annals of Tropical Medicine: and Parasitology 21; 225-244, WARD, H.B. AND MAGATH, T.B. 1917. Notes on some nematodes from freshwater fishes. The Journal of Parasitology 3: 57-63, one plate. Wu, H.W. 1949. A note on two parasitic nematodes of fishes. Sinensia, Shanghai 20(1-6): 51-57. YAMAGUTI, 8. 1935, Studies on the helminth fauna of Japan Part 9, Nematodes of fishes, |. Japanese Journal of Zaolagy 6; 337-386. 1941. Studies on the helminth fauna of Japan, Part 33. Nematodes of fishes Il. Japanese Journal of Zoology 9: 343- 408. 1954, Parasitic worms mainly from Celebes Part 9- 426 MEMOIRS OF THE QUEENSLAND MUSEUM Nematodes of fishes. Acta Medica Okayama 9: YORKE, W. AND MAPLESTONE, P.A. 1926. ‘The 112-133. nematode parasites, vertebrates’. (J. & A. Chur- 1961a. Studies on the helminth fauna of Japan part chill: London). 57. Nematodes of fishes, III. Journal of Helmin- ZEDER, J.G.H. 1800. ‘Erster Nachtrag zur Naturge- thology, R.T. Leiper supplement: 217-228. schichte der Eingeweidewiirmer von Johann 1961b. ‘Systema Helminthum. Volume III. The August Ephraim Goeze.’ (Siegfried Lebrecht nematodes of vertebrates. Part I.’ Indian Reprint, Crusius: Leipzig). 320pp. 1985. (International Books and Periodicals Supply Service: New Delhi). 679pp. REDESCRIPTION OF THE ASCARIDOID NEMATODE AYSTEROTHYLACIUM SCOMBEROMORI (¥Y AMAGUTT) FROM AUSTRALIAN SPANISH MACKEREL SCOMBEROMORUS COMMERSON (LACEPEDE) NIEL L. BRUCE Bruce, N.L, 1990 08 31: Redescription of the ascaridoid nematode Hysterethylacium scomberomori (Yamagutt) from Australian spanish mackerel Scomberamarus cammerson (Lacepéde). Memoirs of the Queensland Museum 28(2); 427-434 Brisbane. ISSN 0079- 8835, Stomachs of 5 species of mackerel were examined. Hysterothylaciurt were obtained only from Scamberomorus commerson, Hysterothylacium scomberomeri is redescribed, together with comments on another undescribed species, Goezia aspinulosa Arya is placed in synonymy with H. scomberomori, The other ascaridoids recorded from mackerets are discussed, Notes are included on Hysterothylacium sp. which occurs concurrently with 7. scomberomori in stomachs of Scomberomorus commerson, DAscarididoid nematode, Hysterathylacium, mackerel, Scomberomarus commerson, Niel L.. Bruce, Queensland Museum, PO Box 30), South Brisbane, Queensland 4101, Australia; 31 January, 1989. Ascaridoid nematodes from Australian pelagic marine fishes were reviewed by Bruce and Can- non (1989). Since the completion of that work a small number of specimens has been collected from Scomberomorus cammersen (Lacepéde) from Queensland waters. The specimens proved to belong to two species of Hysterothylacium Ward and Magath: H, scomberomori (Yamaguti, 1941) and an undescribed species. The un- described species is represented by four mature females, but no males. This record of H. scom- beromori brings the total number of adult marine species of the genus recorded from Australia {Bruce,1990; Bruce and Cannon, 1989) to nine. Five species of mackere] were examined for worms. With the exception of one worm from the Solomon Islands, all were collected from eastern Queensland. The species examined were: Scom- beromorus commerson (Lacepéde), Lizard Is- land (northern Great Barrier Reef) (19 examined), Heron Island (southern Great Barrier Reef) (14), southeastern Queensland (10); Scom- beromorus munroi Collette and Russo, southeastern Queensland (35); Scomberomorus queenslandicus (Munro), Heron Island (1), southeastern Queensland (34); Scomberomorus semifasciaius (Lacepéde), Lizard Island (5); Grammatorcynus bicarinatus (Quoy and Gaimard), Lizard Island (7). Nomenclature for the hosts follows Collette and Russo (1984), All material is housed in the Lower Inver- tebrates collections of the Queensland Museum (QM). Abbreviations used are BL - body length; ED - ejaculatory duct. All measurements are in microns (jm) unless. otherwise indicated and were obtained using a calibrated micrometer eyepiece. Numbers in the figure caption tefer to specimens numbered in the collection, SYSTEMATICS OF ASCARIDOIDS FROM MACKERELS Mackerels of the genera Scomber and Scoim- beromorus are important food fishes in many parts of the world. Larval ascaridoids, commonly of the genera Anisakis Dujardin, Contracaecum Railliet and Henry, Hysterathylacium Ward and Magath and Terranova Leiper and Atkinson, have been widely reported in mackerel hosts (é.g., Beumer er al, 1982; Cannon, 1977; Korotaeva, 1974; Oshima, 1972). Not surpris- ingly there are many nominal species. Most of these are unidentifiable by contemporary stand- ards, with brief and sometimes unillustrated descriptions. For many of these species there is no information on the whereabouts of the type material or, indeed, if there was any, A survey of the literature revealed 9 names which have been established for ascaridoids with species of Scom- ber ot Scomberomorus as type host. Of these names | regard four as valid. It is important to recognize that although some of these names are here regarded as nomina dubia and are unlikely ever to be resolved, they still remain available. The species involved are listed in Table 1 together with remarks on their status. All names based on larval stages are here regarded as nomina dubia, 428 MEMOIRS OF THE QUEENSLAND MUSEUM SPECIES HOST REFERENCE |TYPES | REMARKS Goezia aspinulosa Arya, L980 Dujardinascaris cybii Arya and Johnston, 1978 S. brasiliensis and spp. Hysterothylacium fortalazae (Klein, 1973) Porrocaecum paiva S. cavalla Silva Molta and Gomes, 1968 Ascaris papilligerum Scomber scomber Creplin, 1846 Ascaris pedum Scomber scomber Deslongschamps, 1824 Scomber japonicus and spp. Hysterothylacium saba (Yamaguti, 1941) S. sinensis, 5. commerson Hysterothylacium scomberomori (Yamaguti, 1941) Ascaris scombromorum Scomber colias Stossich, 1892 Hysterothylacium sp. S. commerson present work Synonym of H. scomberamori larva of Terranova ? Deardorff and valid Overstreet, 1981 valid, incerta cedis Stossich, 1896 valid, nomen dubium Stossich, 1896 nomen dubium valid present work valid Stossich, 1896 larva, nomen dubium known only from females present work TABLE 1. Ascaridoid species described from type hosts of the genera Scomber and Scomberomorus (S. = Scomberomorus; OCT = Oslwaldo Cruz Institute; MPM = Meguro Parasitological Museum). There are several other species, such as Hysterothylacium fabri (Rudolphi), H. aduncum (Rudolphi) and H. incurvum (Rudolphi) (see Mozgovoi, 1953; Yamaguti, 1961; Zhukov, 1960), that have been widely recorded from a variety of hosts including scombroid fishes. These species were not established on the basis of type material from Scomber or Scom- beromorus hosts. In many cases the identities of these species are uncertain or, when reassessed, are shown to be misidentifications. Examples of such misidentifications include records of Maricostula incurva (in various combinations) from hosts other than Xiphias and records of Hysterothylacium cornutum (Stossich) from hosts other than tunas (Bruce and Cannon, 1989). Without examination of the material on which the decisions were made, it is not possible to corroborate or refute many of the older deter- minations. To tabulate the numerous unsub- staniated records of ascaridoids from scombroid hosts is a task beyond the scope of this work and such records are not included in Table 1. Hysterothylacium scomberomori (Yamagutt) (Figs 1, 2) Contracaecum meanest Yamaguti, 1941: 362, pl.V, figs 21, 22, text figures 18, 19.— 1962: 30, Contracaecum (Erschovicaecum) scomberomori— Mozgovoi, 1953; 208, fig. 127. Goezia aspinulosa Arya, 1980: 96, fig. 1. Hysterothylacium scomberomori.— Deardorff and Overstreet, 1981: 1042. MATERIAL EXAMINED All from stomachs of Scomberomorus com- merson (Lacepéde). Male, Lizard Island, 5.xi.1988, coll. L.R.G. Cannon and N.L. Bruce. (OM GL10264). Male, 10 females, Heron [sland Reef, 26.iv.1988, coll. N.L. Bruce and S. Cook (QM GLI0147). 2 females, several immature, HYSTEROTHYLACIUM SCOMBEROMORI northern side of Wistari Reef, 21.iv.1988, coll. N.L. Bruce and 8, Cook (QM GLI10148). 2 damaged females, 2 immature, Heron Island Reef, 25.iv.1988, coll. N.L. Brice and S. Cook (OM GL10146). 2 females (frozen sample), off Moreton Island, iv.1988. coll. S. Watson and N,L. Bruce (OM GL10265). 2 males, Moreton Bay, 12.11.1989, coll. T.H. Cribb (QM GL10343), Female, Vana Vana, Solomon Is- lands, viii. 1988, coll. CSTRO (QM’'GL10266). TYPES Believed to be held at the Meguro Parasitologi- cal Museum, Tokyo, Japan. S. Kamegai (in fit.) informs me that the collection was received in poor condition, uncatalogued, and therefore type specimens cannot be identified. TYPE LOCALITY Yamaguti (1941) merely stated ‘Pacific’. Col- tette and Russo (1984) give the distribution of the type host from Japan to Cambodia and Victnam. The probable type locality can therefore be restricted to north western Pacific, Tyre Host Yamaguti (1941) gave Scomberomorus chinensis as the host for his material. The cur- rently accepted name and spelling (Collette and Russo, 1984) is Scomberomus sinensis (Lacepéde). DIAGNOSIS Cuticle with distinct cuticular rings. Lips with length to width ratio 1; 1,08-1,33 (mean = 1.14, n=5), with deep postlabial grooves; lateral con- striction wide, positioned slightly less than one third length from anterior. Alae commence posterior to ventral postlabial groove of sub- ventral lips, Caudal papillae pairs: precloacal 28-33; paracloacal 2-3; postcloacal 7-11, without doubled papillae; medioventral precloa- cal organ present, Postcloacal pad present. Spicules approximately equal in length, 1; 1.02- 1,09, 3.01-4.60% BL. Tail apex with 5-7 small acute spines. DESCRIPTION Based on 3 mature males, 1 mature female and 4 immature females. General. Body reaching greatest width about midbody. Dorsal lip slightly shorter than sub- ventrals, flanges moderately developed, trian- gular, Interlabia prominent, not extending beyond anterior of oesophagus, less than half 429 length of lips. Oesophagus 6.97-7.83% BL, Ventriculus narrower than widest level of oesophagus, longer than wide. Yentricular ap- pendage 27.9- 37.5% length of oesophagus, In- testinal caccum 18.2-25.0% length of oesophagus. Nerve ring lying between anterior 25.6-29.4% of oesophagus. Excretory system filamentar, unilateral, with canal extending slightly anterior {o commissure, posteriorly beyond ventriculus. Male. Body 19.2-43.8 mm long, 353-818 max- imum width; width at intestinal/oesophageal juschion 273-395; ratio of pean width to ength 1:42,1-54.4, Dorsal lip 71 long by 94 wide (one specimen); subventral lips 85-141 long by 94-150 wide. Nerve ring 442-630 from anterior. Excretory pore 470-705 from anterior, Oesophagus 1504-2632 long by 115-179 wide, Ventriculus 71-118 long by 82-141 wide; ventricular appendage 564-799 long by 47-66 maximum width. Intestinal caecum 273-658 long by 108-188 wide. Ejacutatory duct 1,692- 3,854 long, 8,8- 10,6% BL, Spicules 884-1,363 long, 37.9-52.2% ED. Caudal papillae pairs 40- 43, changing from button to mamillate at about Sth anterior to cloaca, Tail tapering evenly, 155- 329 long. Female. Body 28.3 mm long, 705 maximum width; width at oesophageal/intestinal junction not observed; ratio of maximum width to length 1; 40.1. Dorsal lip 118 long by 127 wide. Sub- ventral lip 122 long by 136 wide. Nerve ring and excretory pore not observed. Oesophagus 1974 long by 188 wide. Ventriculus 235 long by 160 wide; ventricular appendage lost in dissection, Intestinal caecum 432 long by 235 wide, Vulva opening 10.0mm or 35.3% BL from anterior, vulva areca not swollen. Vagina 3290 long; uterus undivided for 846, divided part 3478. Eggs about 47. Tail 564 long. Immature females. Body length 12.5-22.5 mm. Oesophagus 1105-1739 long by 84-122 wide; 7.73-9.30% BL. Ventricular appendage 376-564 long by 33-71 wide; 31.85-39.29 length of oesophagus. Intestinal caccum 165-371 long by 66-118 wide; 14.57-21,33% length of oesophagus. Vulva 4.57-8.34 mm or 28.90- 37.04% BL from anterior. Variation, The largest males (QM GL10342) have the tail more abruptly narrowed than shorter males and also have a greater number of paostcloa- cal papillae (11) than the smaller males (7-8). 430 MEMOIRS OF THE QUEENSLAND MUSEUM TeRNyTURUNReNUHOUUD UUMULAWUELENIVIELUEE! as = G Fic. 1. Hysterothylacium scomberomori. A, anterior end, ¢ #1 (scale bar 200,.m); B, dorsal lip, 2 #5 (100m); C, subventral lip, ¢#2 (100j.m); D, interlabium, d #2 (100j.m); E, ventricular region, 3 #2 (200m); F, tail, lateral view, 6 #2 (100,.m); G, caudal area, ¢#2 (400m). HYSTEROTHYLACIUM SCOMBEROMORI 431 Fic. 2. Hysterothylacium scomberomori. A, tail, ventral view, #1; arrows indicate deemed paracloacal papillae (scale bar 50j.m); B, tail, lateral view, 9#2(200,m); C, reproductive tract, 2 #5; small arrow indicates end of vagina, large arrow indicates point of division of uterus (1.0mm). Sections (from immature female, scale bars all 50,.m): D, through nerve ring; E, excretory commissure; F, about 80,4.m posterior to nerve ring; G, anterior of intestinal caecum; H, posterior of ventriculus; 1, mid-ventricular appendage. 432 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 3. Hysterothylaciumscomberomori, Scanning electron micrographs. A, en face; B, subventral lip; C, dorsal lip; D, postcloacal papillae. Hosts Australian material recorded only from Scom- beromorus commerson;, also Scomberomorus sinensis (type host) and Scomberomorus guttatus (Bloch and Schneider) by Arya (1980, see synonymy). DISTRIBUTION In Australia, at least in Queensland, presumab- ly throughout the range of Scomberomorus com- merson. One specimen recorded from Solomon Islands. Also India (Arya, 1980) and within the tange of Scomberomorus sinensis, that is, southeast Asia to Japan (Collette and Russo, 1984) if the type host was correctly identified. REMARKS The adult male material at hand agrees well with the brief description and figures of Yamaguti (1941). Particular points of correspon- dence include the proportions of the oesophagus, intestinal caecum and ventriculus, the caudal papillae and spicule details and the description of the tail, Other ascaridoid species described from Scomberomorus hosts are Hystero- thylacium fortalazae (Klein) (Deardorff and Overstreet, 1981), Porrocaecum (P.) paivai Silva Motta and Gomes, 1968, Dujardinascaris cybii Arya and Johnston, 1978 and Goezia aspinulosa Arya, 1980. Hysterothylacium fortalazae is easily distin- guished by the prominent cervical alae, more spinose tail apex and fewer precloacal papillae. Of the remaining species, Porrocaecum (P.) paivai should be treated as incertae sedis until more fully described. However, it differs from H. scomberomori in having wider alae and, most notably, in lacking a ventricular appendage. From the brief description and rudimentary figures, Dujardinascaris cybii cannot be as- IfYSTEROTHYLACIUM SCOMBEROMORI signed to a genus. It was listed as incertae cedis by Soota (1983) Goezia aspinulosa is obviously not a species of Goezia and, as figured, has the characteristics of an Aysterothylacium. Soota (1983) com- mented that the species lacked cuticular spines. The description and figures given by Arya (1978), consisting largely of family or generic characters, are not adequate to allow absolute resolution of the identity of the species. The general correspondence of lip size, prominent cuticular rings, tail ornamentation, spicule size, and morphology of the ventricular appendage and intestinal caecum strongly suggests that G. aspinulosa is a junior synonym of H. scom- beromori and is so included in the synonymy. Data taken from immature females are in- cluded to illustrate the constancy of proportions of the characters listed. Hysterothylacium sp. A second species of Hysterothylacium (represented by four mature females QM GL10267) was collected from the stomach of ene specimen of Scomberomorus commerson fram Heron Island together with immature specimens of H. scomberomeri, This worm is easily distinguished from H. seomberomori (including synonyms and related species) by the following: alae starting immediately posterior to postlabial groove; long intestinal eaccum (75.7-78.4% length of oesophagus), longer ventricular appendix (S2,7-58.1% length of oesophagus}, a bluntly rounded tail with conical nodulose apex, oviducts extend- ing 1.5- 2.5mm anterior to the vagina and the ulerus with the undivided portion (3854) ap- proximately equal in length (93.2%) to the un- divided portion (4136) in the dissected female. This latter character is very different in H. scevn- beromori, which has the undivided part 24.3% the length of the divided part. Without adult males, it is mot possible to give a full and diagnostic description of this species or to be sure if it has previously been described. The specimens are recorded here lo draw attention to the presence of twa species of Hysterathylacium that can be found in Scomberamorus hosts in Australia and that can occur together within the single host stomach, Additionally two immature specimens from Scaomberamorus queenslandicus were 433 tentatively identified as belong to this species rather than H. scomberomari, ACKNOWLEDGEMENTS This study was generously funded by the Australian Research Council. The staff of Heron Island Research Station and Lizard Island Re- search Station are thanked for their assistance, [ also thank the following for assistance with field work; Dr L.R.G. Cannon, Dr T. Cribb, Mr S, Cook and Mr R.T. Springthorpe. I also thank Mr D, Steel, Mr S. Watson (Brisbane Sportsfishing Club) and Mr J. Smith (CSIRO - Cleveland) for their efforts in continuing to supply me with host stomachs. Mrs Christine Lee is thanked for her careful preparation of sections. LITERATURE CITED ARYA, S.N. 1980. A new nematode of the genus Goezia Zeder, 1800, from marine fish of India. Indian Journal of Helminthalogy 30(for 1978): 96-99, ARYA, S.N. AND JOHNSON, S. 1978. A new species of the anisakid genus Dujerdinascaris from the fish Cybium gutratum, with a key to the Indian species of the genus Dujardinascaris. Veytnik Ceskoslavernske Spolecnoski Zoologicke 42: 81- 84. BEUMER, J.P. ASHBURNER, L.D., BURBURY, M.E,, Jerry, B. Anp LATHAM, D.J. 1982. A checklist of the parasites of fishes from Australia and its adjacent Antarclic Territories. Commonwealllt Institute of Parasitology, Technical Com- munication 48; i-viii, 1-99. Beuck, NL. 1990, Mysterathylacium Ward and Magath, 1917 and /ehthyascaris Wu, 1949, as- caridoid nematodes from Australian Demersal Fishes. Memoirs of the Queensland Museum 28(2): 389-426, Bruce, N.L. AND Cannon, L.R,G. 1989, Hysterothylacium, Theringascaris and Maricos- tila new genus, nematodes (Ascaridoidea) from Australian pelagic marine fishes, Journal ef Natural History 23:1397-1441. CANNON, L.R.G., 1977. Some larval ascuridoids from south-eustern Queensland murine fishes. /nter- national Journal of Parasitology 7: 233-243. CoLLetTe, B.B. AND Russo, J.L. 1984. Morphology, sysiemalies, and blology of the spanish mack- erels (Scomberomorus, Scoambridae), Fishery Bulletin $2: 545-692. DEARDORFF, TL, AND OVERSTREET, R.M, 198f, Review of Mysrerothvlacium and [heringasearis 434 (both previously = Thynnascaris) (Nematoda : Anisakidae) from the northern Gulf of Mexico. Proceedings of the Biological Society of Washington 93(for 1980): 1035-1079. DOLLFUS, R.P. 1933. Thynnascaris legendrei, n, gen., n. sp., de l’estomac du germon, Germo alalonga (Gmel.). Bulletin de la Société Zoologique de France 58: 7-13. KoROTAEVA, V.D. 1974. Helminths of some marine fishes of the suborder Scombroidei from the Australian region. /zesfaya Tikhookeanskogo Naucho-Issledovafel’skogo Instituta Rybnogo Khozyaistue : Okeanografii, Vladivostok 88: 61- 66. Mozcovol, A.A. 1953. Ascaridata of animals and man and the diseases caused by them. Gel’mintologischeskaya Laboratoriya. Osnovy Nematologii, Volume 2, (Akademiya Nauk: Moskva). 617 pp. OsHIMA, T. 1972. Anisakis and anisakiasis in Japan and adjacent area. Jn Morishita, K., Komiya, Y. and Matsubayashi, H. (eds), ‘Progress of Medi- MEMOIRS OF THE QUEENSLAND MUSEUM cal Parasitology in Japan’. 4: 310-393 (Meguro Parasilogy Museum: Tokyo). SILVA MOTTA, C. AND GOMES, D.C. 1968, Sébre uma nova espécie do género <> Rail- lie. and Henry, 1912 (Nematoda, Anisakidae). Atas da Sociedade de Biologia do Rio de Janeiro 11: 163-164. STossicH, M. 1896. Il Genere Ascaris Linné, Lavoro Monografico. Bolletina della Societa Adriatica di Scienze Naturali in Trieste 17: 9-120. YAMAGUTI, S., 1941. Studies on the helminth fauna of Japan. Part 33. Nematodes of fishes II, Japanese Journal of Zoology 9: 343-408. 1961b. ‘Systema Helminthum. Volume III. The nematodes of vertebrates. Part I.’ Indian Reprint, 1985. (International Books and Periodicals Supply Service: New Delhi). 679pp. ZHUKOV, E.V,, 1960. Endoparasitic worms of fishes in the Sea of Japan and shallow water of South Kurile Islands. Trudy Zoological Institute, Akademiya Nauk, SSR 28: 3-146 (in Russian). APIDIOPLANA APLUDA N. SP., A TURBELLARIAN SYMBIOTE OF GORGONIAN CORALS FROM THE GREAT BARRIER REEF, WITH A REVIEW OF THE FAMILY APIDIOPLANIDAE (POLYCLADIDA:ACOTYLEA) L.R.G, CANNON Cannon, L.R.G. 1990 08 31: Apidioplana apluda n. sp., a turbellarian symbiote of gorgonian corals from the Great Barrrier Reef, with a review of the family Apidioplanidae (Polycladida: Acolylea). Memoirs of the Queensland Museum 28(2); 435-442. Brisbane, ISSN 0079-8835. Apidioplana apluda n. sp. is from gorgonian corals, Melithea sp., from the Great Barrier Reef, The worms are small, red like their host, and live closely applied to it. The worms are characterized by a slender, tapering penis with a fine, terminal sclerotic tube, which may be recurved. The genus has been found only with gorgonians of the genus Melithea and is known from Fiji and Japan. This is the first report from Australia. Apidioplana, Polycladida, Turbellarian, Symbiote, Gorgonian, Coral, Great Barrier Reef. L.R.G, Cannon, Queensland Museum, PO Box 300, South Brisbane, Queensland 4101, Australia; 20 March, 1989 During his expedition to the South Pacific in 1917-18, Dr Sixten Bock recovered some small polyclad turbellarians from gorgonians taken from the surf zone from reefs in Fiji. Their orange to red colour matched well that of their host, Melithea (=Melitodes). Furthermore, he found they had numerous pyriform, glandular struc- tures. on the ventral surface, each with a hard, conical or tubular mouthpiece. Bock (1926) called these organs apioids and created a new genus, species and family within the Acotylea for the worms: Apidioplana mira, Apidioplanidae. Subsequently Bock (1927) published a much fuller account of these animals and erected a second species, A. similis, which he had earlier overlooked among the specimens. Kato (1944) described a third species, A. okadai, found on Melithea (=Melitodes) flabellifera at Simoda, Izu Japan. Although briefly mentioned by Bresslau (1928-33), Stummer- Traunfels (1933), Hyman (1951), Faubel (1983), Prudhoe (1985) and Cannon (1986), Apidioplana has been un- recorded for nearly 50 years. The discovery of a new species of the genus prompts my review of the family. Polycladida: ACOTYLEA Superfamily PLANOCEROIDEA Poche, 1926 Family APIDIOPLANIDAE Bock, 1926 Genus Apidioplana Bock, 1926 Apidioplana apluda n. sp. Fic. 1. Apidioplana apluda n. sp. whole warm (Scale; (Figs 1,2,3a-e) = 500m) (B = bursa, G = gut, Od = oviduct, Oc = oocyte, Ph = pharynx, T = tesis, t= tentacle, V=vas © MATERIAL EXAMINED deferens) HOLOTYPE: ex Melithea sp, 3m on Milln Reef, GBR, P. Alderslade, March 1978, G1.10337 (whole mount: stained with Mayer's Haemalum, mounted in Canada Balsam). PARATYPES: data as for Holotype, GL10338 (whole mount: treated as for holotype) and GL10339-10342 (sections; serial sections cul at 7m and prepared as follows ~ longitudinal sagital sections stained with Haematoxylin and cosin GL10340 and GL10342, lon- gitudinal facial sections stained wilh Papanicolou's GL10339 and transverse sections stained with Heidenhain’s iron haematoxylin GL10341), All material is in the Queensland Museum. DESCRIPTION The whole worms are about 2 by Imm, quite russet red dorsally and grey ventrally, with a short, tubular pharynx opening via a mid-ventral mouth about one third from the anterior margin. They have a bilobed brain and four anterior- lateral cyes of unequal dimensions. The largest (about 15ym in diam.) and most posterior pair abut the ventral basal membrane, slightly anterior and in the midbody are the next largest pair. The two smaller, more median pairs of eyes are closer ta the dorsal surface. A rudimentary pair of tentacles are found on either side of the anterior just lateral to the brain. They are repre- sented in fixed material by a pair of low bosses (Fig. 3a). The dorsal and ventral epithelia are weakly ciliated, and the dorsal epithelium (but not the ventral) is filled with prominant rhabdoid MEMOIRS OF THE QUEENSLAND MUSEUM bundles, often appearing to be arranged in small rings anterior to the pharynx, but more irregular- ly in the posterior. In section, small blisters from the basal membrane of the dorsal epithelium can be seen and may account for this arrangement. Subepidermal muscles consist of outer lon- gitudinal fibres underlain by transverse then diagonal layers. All three muscle layers are more strongly developed ventrally than dorsally. Strong dorso-ventral muscles run through the parenchyma. Unicellular basiphilic glands are scattered through the mid-body, Larger, granular cells lie in the parenchyma more dorsally. The brain is bilobed, with a scattering of anterior nuclei. Thick lateral cords tun posteriorly lateral to the pharynx and form a ventral plexus. The mouth opens on the ventral surface about 300 - 350,.m from the anterior margin. The anterior buccal region has a ciliated pouch (about 100..m long by 60-70,.m wide) extending into the pharyngeal chamber in which lies the short, tubular and strongly muscled pharynx (3b). The gut opens behind the pharynx and proceeds posteriorly as a more or less tubular canal: the gut is virtually saccate (3c). The male reproductive system consists of numerous testes (about 40-S0,.m in diam.) scat- tered laterally to the centrally placed gut and stretching from the region of the phanynx posteriorly to just in front of the penis. They lie predominantly ventrally or ventro-laterally- From the posterior of this field on each side a Fic, 2, Apidioplana apluda n. sp. reconstruction of copulatory siructures (Scale: = 100,.m) (B = bursa. g = accessory glands of \he vagina media, Od = oviduct, p = penis, S = stylet, V = vas deferens) APIDIOPLANA FROM THE GREAT BARRIER REEF 437 Fic. 3. Apidioplana apluda n. sp. (a) tentacle (r = rhabdoids), (b) pharynx and buccal pouch (g = brain ganglion), (c) section through intestine, (d) tangential section through gonopores and revealing the penis stylet (s), (€) bursa (b) without hook (Scales: a = 251m, b = 100pm, c-e = 50j.m) 438 weak-walled and much coiled duct, the vas deferens, runs posteriorly to near the gonopore then bends and runs forward along the penis to curve again and enter the penis stem. There is no evidence of a distinct seminal vesicle or of pros- tate tissue, The penis is a long, tapering muscular organ 400) - 500,.m long within a penis cavity. Internally there is a narrow ejaculatory duct, The penis ends in a very fine tip covered with a thin, tubular, sclerotic stylet (Pig. 3d). This fine tip may be strongly reflexed within the antrum, The male gonopore opens separately to the ventral surface immediately anterior to the female pore (Pig. 3d) i.e., 100 - 200m from the posterior margin which is somewhat indented in the mid- posterior region. The female system consists of numerous ovaries (about 60 - 100..m in diam.) developing laterally and slightly more dorsal io the testes. There is a tendency for gonads to interdigitate. Dorsally on each side oviducts. run sinuously posteriorly from the hind most quarter of the ovaries. In the region of the stem of the penis the ducts curve medially, join and a common duct (vagina media) then runs anteriorly for a short distance. Numerous acidophilic glands dis- charge into it. The duct loops ventrally and then runs posteriorly above the penis. At the level of the gonopores it joins the duct running from the bursa copulatrix and this short common duct opens into a female antrum and then to the female gonopore. The bursa (about 150 by 40j.m) lies medially and is a short blind sac running forward over the top of the penis. It has smooth walls lined with tall cilia and is without evidence of any sclerotic boss or hook (Fig. 3e). Characteristic of the worms 1s the presence, on the ventral surface, of numerous ‘apioid’ organs or prostatoids (Fig. 3d). These range from 15 - 60..m in diameter and are scattered around the lateral regions of the body, being smaller and less numerous (or absent?) anteriorly than posterior- ly. The organs are prominent in the posterior lateral body. Each organ consists of a strong muscular capsule through which the necks of glands pass. Inside the organ there are several large cells filled with a granular secretion, Each organ opens to the exterior via a short, strong sclerotic cone which projects from a small MEMOIRS OF THE QUEENSLAND MUSEUM depression of the ven(ral epithelium. Within the organ the sceretion granules failed to stain with the stains used, but become increasingly basiphilic towards the exterior of the cone, ETYMOLOGY The specitic name apluda L. = scale, chaff. It pertains to the habit of these animals to be closely applied to their host. REMARKS The presence of only one type of apioid, thase with conical mouthpieces, and their presence scattered about the body, place the new species close to A, mira with which it also shares an unarmed bursa. The gonopores are well separated in A. mira and the penis armature is quite different. A. mira has a short, wide, sclerotic ring with scalloped distal rim, like a coronct, but A, apluda has a long, finely tapering stylet that may be reflexed. Purther, A. mira has a large, thin-walled seminal vesicle which passes via a small pore into a voluminous penis cavity with a well developed interpolated prostate, Apidioplana mira Bock, 1926 (Figs 4a-b, Sa and e)} Bock (1926) 133, 3 figs,: (1927) 6, 17 tigs.; Bresslau (1928-33) 89, fig.; Hyman (1951) 116, fig., Faubel (1983) 96; Prudhoe (1985) 115, fig.; Cannon (1986) 72, lig.; on Melithea (=Melitodes) surf zone Mbau, Fiji Islands. MATERIAL EXAMINED 13 slides of serial sections from Fiji of Sixten Bock’s material from the collections of the Natural History Museum, Stockholm, REMARKS Bock (1927) gave a thorough but discursive account of A. mira, Although the prostatoids (apioids) (Fig. 5e) can be found anteriorly in some cases, their number can be small. The gonopores are well separated by about 100u.m. The bursa copulatrix is strongly muscled with a thick homogenous lining (Fig. 5a). The male copulatory apparatus (Fig. 4a) has a seminal vesicle with a thin muscular wall about 10m Fic. 4. Male copulatory organs: (a) Apidioplana mira ~ longitudinal section showing seminal vesicle and commodious interpuluted prostate (v = vagina and s= stylet), (b) same Section as (a) but at greater magnification showing the ring-like sclerotic stylet, (c) A. similis - longitudinal section showing heavy musculature of seminal vesicle and penis (p) which lacks a sclerotic stylet (v = vagina) (Scales: a and c= 100m, b = 25pm), APIDIOPLANA FROM THE GREAT BARRIER REEF 439 440 MEMOIRS OF THE QUEENSLAND MUSEUM *% SOLE A 3 ee a Fi. 5. Bursae and prostatoids: (a) A. mira - longitudinal section of bursa showing thick wall, but no hook, (b) A. similis - longitudinal section of bursa as figured by (Bock, 1927) showing a blunt hook, (c) A. similis - longitudinal section of bursa showing spinous hook, (d) A. simils - transverse section showing tubular hook (v = vagina), (e) A. mira - postatoid, (f) A. simils - a large Type-A prostatoid and a small Type-B prostatoid (B) (Scales: a,c and e = SOjum, d = 100pum, f = 254m) APIDIOPLANA FROM THE GREAT BARRIFR REEF thick, ‘sehr stark muskulds" according to Bock; the interpolated prostate is extensive and the terminal armature (Manschettc) is a small sclerotic ting bearing distal scalloping like a coronet (Fig, 4b). The mouth opens into a short buccal pouch (Mundrohr) which is provided with a high ciliated epithelium and is protrusible. The gut is virtually saccate. Apidjoplana similis Bock, 1927 Bock (1927) 87, 97, 2 figs, MATERIAL EXAMINEDY & slides of serial sections: details as for A. mira REMARKS Bock's (1927) description of A. similis is very rnuch less detailed than that of A, mira, He high- lighted the major differences he perceived be- tween the species. He recognized a second, much smaller prostatoid (apioid type -B) (Fig. Sf) withoul a sclerotic mouthpiece. In some of Buck’s sections these small prostatoids are strongly stained and readily seen. In other sec- tions, however, they stain weakly and easily could be overlooked, Many of the type A apioids (Pig. St) are larger and more strongly muscled than those of A. mira. The gut is’ similar to that of A. mira in having both the buccal pouch and little lateral branching. The seminal vesicle is extremely muscular with walls about 25m or more thick. A narrow strongly muscled sphincter leads inta the ejaculatory duct with only modetate prostate development (Fig. 4c). The penis is muscular, but completely unarmed. Bock described the bursa (his fig. 19) with a rounded sclerotic boss (Zapfen) (Fig. 5b). From his material, this structure is quite variable being also with spinous projections (Fig. 5c) or merely a long thin spine (Fig. Sd). The gonopores are well separated. Apidioplana okadai Kato, 1944 Kato (1944) 287, figs. REMARKS Kato’s material was lost during the allied bombing of Japan during the World War U (Kawakatsu, pers. comm.). This specics was characterized by having a single large apioid with a very muscular wall immediately posteriar to a common gonopore, having a muscular copulatory organ containing Atl a seminal vesicle with a narrow duct leading to an ejaculatory duct with moderate prostale development and an unarmed penis, Also there is a long fine sclerotic spine in the bursa. Kato made no mention of a buccal pouch and described the gul as ‘provided with numerous lateral branches which form a network’. DISCUSSION Bock (1927) gave a thorough account of the ‘apiotd’ organs and of the many terms used to describe them. Since then, similar organs have been recorded several times most notably within the triclads where they are known as adenodac- tyls. Myman(1951) suggested the term ‘prostatoid’ be adopted as a convenient term fur all of them. Within the polyclads, prostatoids are found in the Discocelididae, Polyposthiidae as well as the Apidioplanidae within the Acotylea and also in the Boniniidae (Cotylea), They may be clustered about the male antrum and open into it (Coronadena, Adenoplana, Cryptocelides (Discocelididac) and Boninia (Boniniidae)), be on the walls of the penis (Discocelis (Dis- cocelididae)) or open onto the ventral surface either clustered around and radiating fram the gonopore (Paraboninia (Boniniidae), Polypas- thia, Polypkalloplana, Metaposthia (Polypos- thiidae)), forming a line of mid-ventral rosettes but alsa scattered about the ventral periphery (Polyposthides (Polyposthiidae)), simply scal- tered ventrally aboul the periphery, mainly posteriorly Apidioplana mira, restricted to the posterior A. similts or reduced to one large organ opening ventrally just posterior ta the gonopores in A. okadai as Well as Traunfelsia (Boniniidae). In most cases the organ is as described above, i.e, a pyriform muscular capsule filled with secretion and extracapsular glands feeding into it. There is a short conical or tubular sclerotic mouthpiece opening to the exterior. Sometimes ihe organ is nol armed and sometimes vasa deferentia, or just teminant ducts, Jead to the organs. Marcus and Marcus (1968) state regard- ing Boninia antillarum (p.63) ‘The prostatoids of the original worm have no stylets, because the animal is growing’. Bock (1927) described Apidioplana similis with small, unarmed pros- tatoids as well as the larger armed type. He discounted the possibilty of these being from young individuals since he found the prostatoids (8 in all) in one quite juvenile individual, The function of these glands is unclear, bul some stimulatory role in facilitating copulation 442 and/or fertilization is possible, All species of Apidioplana are reported from gorgonians of the genus Melithea (=Melitodes), Boninia mirabilis was recorded also from the same coral. It may be tempting to speculate that these glands in some cases have becn modified to assist with adhesion to the host. Gorgonians live in turbulent waters and Bock (1926) collected his specimens from the surf zone. Similar glands are found, however, in several species, both Acotylean and Cotylean, not known to be associated with hosts nor known to live in areas of exceptional turbulence. A comparison of the principal characters of the species in the genus is given in Table 1. Re-ex- amination of Bock’s original material suggests little change in details for A, mira, the type species of the type genus of the family, However, the details of the bursal hook in A. similis reveal it to be more variable than Bock indicated. The presence of a specimen with a long fine bursal spine, together with the unarmed penis indicates A, similis is quite close to A. okadai. It is regret- fable that the type material of the latter species is ost. A. okadai is reported to have a common gonopore, in the other species the pores are separate, though in A. apluda they are close together. A. okadai has only one large posterior apioid, and while A. similis has. both large and small prostatoids, these are all confined to the posterior. Further, the large apioids of A. similis are more prominent than those in A, mira. The small (Type B) ones are quite tiny and may be ees [ac PA eae Prostatoids armed + + unarmed number many many arrangement | all round |all round|posterior | posterior Penis fine stylet = | + sclerotic ring + unarmed + + Genital Pores | separate |separate separate [common unddomed junadomed|sclerotic | sclerotic boss spine or spine efter ben fae [ein | TABLE 1, Comparison of the Species of Apidioplana MEMOIRS OF THE QUEENSLAND MUSEUM easily overlooked if the staining does not high- light them, Insufficient is known of the develop- ment of these organs or of the variability of their occurrence: in both A, mira and A, apluda anterior prostatoids may not be comman. The two species A. similis and A. okadai also share a similar internal arrangement of the penis with a long muscular duct between seminal vesicle and ejaculatory duct. While these two species must remain distinct, suffice to say they appear closely related. Kato’s (1944) report of the gut anatomy of A. okadaz is at variance with the other species in the genus. Apidioplana mira and A. apluda appear close, but the differences in the penis and its armature make them distinct. LITERATURE CITED Bock, S. 1926. Eine Polyclade mit muskulésen Driisenorganen rings um det Korper. Zoologis- cher Anzeiger 6: 133-138. 1927. Apidioplana, eine Polycladgattung mit muskuldsen Drisenorganen. Gdteburgs Kunglingen Vetenskaps-och Vitterhets- Samenhdlles Handlingar. Fjdrde faljden 30: 1- 115. BRESSLAU, EB, 1933. Turbellaria, pp. 52-320. In Kiikenthal, W. and Krumbach, T. (eds), “Hand- buch der Zoologie’ (Walter de Gruyter & Co.: Berlin und Leipzig). CANNON, L.R.G. 1986, ‘Turbellania of the World.’ (Queensland Museum, Brisbane). 136pp. FaAuBeL, A. 1983, The Polycladida, Turbellaria, Proposal and establishment ofa new system. Part I, The Acotylea. Mitteilungen aus dem hambur- gischen zoologischen Museum und Institut 80: 17-121. Hyman, L.H. 1951. “The Invertebrates: Platyhelmin- thes and Rhynchocoela. The Acoelomate Bilateria. U.’ (McGraw-Hill, London). 550pp. KATO, K. 1944. Polycladida of Japan. The Journal of the Sigenkagaku Kenkyusyo 1: 257-322. Marcus, E. AnD Marcus, E, DU Bojs-REYMOND. 1968. Polycladida from Curagao and faunistical- ly related regions, Studies on the Fauna of Curagao and other Caribbean Islands 25: 1- 106. PRUDHOE, 8. 1985. *A Monograph on Polyclad Tur- bellaria.’ (Trustees of the British Museum (Natural History): London), 259pp. STUMMER-TRAUNFELS, R. von. 1933, Polycladida (continued), pp. 3485-3596. Jn Bronn, H.G, (ed,), ‘Klassen und Ordnungen des Tier-Reichs IV. (Vermes).’ (Leipzig). ROCK PAINTINGS OF THE YINGALARRI REGION: PRELIMINARY RESULTS AND IMPLICATIONS FOR AN ARCHAEOLOGY OF INTER-REGIONAL RELATIONS IN NORTHERN AUSTRALIA BRUNO DAVID, MAREE DAVID, JOSEPHINE FLOOD AND ROBIN FROST David, B., David, M., Flood, J. and Frost, R. 1990 08 31: Rack paintings of the Yingalarri region, preliminary results and implications for an archaeology of inler-regional relations in northern Australia, Memoirs of the Queensland Museum 28(2): 443-462. Brisbane, ISSN 0079-8835. Recent research in Wardaman country, Northern Territory, has revealed an extremely varied body of cave paintings. Preliminary investigations of painting techniques from 48 sites located around the Yingalarri waterhole has failed to identify any significant changes in the distribution of rock paintings from the region through time. Given that intensive interactions between Wardaman people and their westerly and southerly neighbours have been tepeuledly documented in the anthropological literature for the recent past, and that the distribution of rack art does not indicate any significant changes in the pattern of inter- regional relations, we conclude that these contacts have considerable antiquity. (JReck art, cave paintings, Wardaman, Northern Territory, intensification, Australian prehistory. Bruna David, Department of Anthropology and Sacialagy, University of Queensland, Queensland 4072, Australia; Maree David, 3 Seaman Avenue, Villawood, New South Wales 2163, Australia; Josephine Flood, Australian Heritage Commission, P.O. Box 1567, Canberra, Australian Capital Territory 2601, Australia; Robin Frost, 464 Greenwich Street, New York, U.S.A.; 24 November, 1989. It was once customary to view the Australian Aboriginal past as consisting of a series of dis- tinct episodes of prehistory, each observable ar- chacologically as made up of a distinct and relatively discrete set of material artefacts. For example, the early ‘Core Tool and Scraper Tradition’ has been seen to consist of large ar- tefacts often charactetised by steep edges and percussion flaking, and followed by the ‘Small Tool Tradition’ which, amongst other things, saw the beginnings of pressure flaking and blade technologies. Each Tradition was seen as an in- ternally coherent system of artefactual produc- tion (and use), and each implied a system In steady-state equilibrium (cf. McCarthy, 1967; Mulvaney, 1975). In the late 1970s and early 1980s a dissatisfac- tion with culture-historical approaches to Aboriginal prehistory began to appear in the literature (e.g. Lourandos, 1983). These dissatis- factions stemmed from the failure of such ap- proaches to address issues which concern social processes. For this reason, archaeology has been, until recently, more concerned with document- ing material aspects of Aboriginal life, and ar- dering the resultant classifications in structural- functionalist terms, than in trying to understand how these structures. came to be. Cultures were seen as made up of interacting parts, each part at once functionally and structurally contributing to the formation of an integrated whole. This is the case not only for the archaeological ‘traditions’ which investigators claimed to have identified, but also for the alliance and trading systems documented ethnographically (and which by im- plication operated in the past) (e.g. McCarthy, 1939), This, to a large extent, is testimony to the influence of Radcliffe-Brown’s work on Australian archaeology (and Australian anthropology in general). It alsa highlights a tendency which has prevailed amongst ar chaeologists to treat prehistoric Aboriginal sys- tems as ccologically adaptive ones, whilst ignoring the role that socio-cultural relations have played in socio-cultural change and stability. Humans interact with each other, create and re-create social and individual identities (creations of a symbolised self in relation to ‘the other’) observe and break social rules, and in the process create the dynamic environments which we study. It is not just in “the norm’, in people frozen in time and space, that we can hope to understand the past, but by inquiring into the social forces which influence people’s be- haviour, and consequently structure social change. It is only thus that we may arrive at an understanding of those forces which have shaped the prehistoric past (and, consequently, the ane present). Yet it is the very dynamism of Aboriginal peoples which for a long time has been neglected, even denied, in early works. In this paper we therefore wish to argue that, in spite of the difficulties involved in addressing questions of social relations and social change in archaeology, such issues should not only be ad- dressed, but should be central in our inquiries of the past. We illustrate this issue by reference to our own research in Wardaman country, North- ern Territory, by looking at the rock paintings of the Yingalarri region. Here, a rich and varied body of rock art is-analysed with respect to the information it can supply on inter-group rela- tions, both past and present. It is concluded that not only have extensive alliance networks been established in the region, but also that such net- works have probably been operating for a very long time, although the particular configuration of alliances may or may not have changed sig- nificantly during this time. WARDAMAN COUNTRY The Yingalarri waterhole is located towards the heart of Wardaman country, approximately 130km southwest of Katherine, Northern Ter- titory. Wardaman country currently extends from the Victoria River to the west, the southern Flora River to the northwest, Scott Creek in the northeast, and Romula Knob in the east (Merlan, 1988, p.3). Their country is made up of a con- figuration of territories to which people are af- filiated in numerous ways, but where patri- and cognatic affiliation are of primary importance. It is in relation to the Dreaming that these affilia- tions to the land are given meaning and identity. Although identified with the region delimited above, Wardaman people have wide-ranging as- sociations with peoples sometimes hundreds of kilometres from their homelands. For instance, Stanner (1959-63) reports seeing a Wardaman man visiting the Port Keats region, 250km from Wardaman country, early this century. In this meeting, he records a Wardaman version of the Rainbow Serpent story, and shows its great similarity to versions recorded west-northwest of Wardaman country. In effect the story itself links Wardaman places with regions all the way to Port Keats, incorporating places such as Garnawala (Mt Hogarth), Murning and Nimji (both near the Yingalarri waterhole) into the story line. Some of the actors in the Rainbow Serpent story go on to participate in other Dreaming stories, and al- though some stories appear unrelated to others, MEMOIRS OF THE QUEENSLAND MUSEUM it is in the Dreaming itself that the whole landscape is linked into a broad configuration of sites each expressing a broader ideological sys- tem. It is this organising of the landscape, via the identification of localised Dreaming beings and events, which articulates the relation of people to the land. As a result, the landscape cannot just be seen as comprising autonomous units in space, but must be understood also as expressing a broader ideological system in continuous social space. Extensive interactions of Wardaman with peoples to the west and south have been docu- mented by Merlan (1988), Davidson (1935) and Spencer (1914). Davidson has noted recent trad- ing relations with people to the west, whilst Merlan (1988) notes that there is much evidence for extensive inter-marriage between War- daman, Mudburra, Bilinara and Garranga peoples to the south, and Ngariynman, Ngaliwurru, Jaminjung and Nungali peoples to the west and southwest (see Fig. 1). These inter- regional links echo favourably the patterning of Fic. 1, Distribution of linguistic groups mentioned in text. After Tindale (1974). YINGALARRI CAVE PAINTINGS linked Dreaming places, such as the Rainbow Serpent and Lightning Brothers stories, which show an extensive network of relatedness be- tween Wardaman country and places to the west and south. In the case of the Lightning Brothers, although the main participants in the story, Jabir- tinggi and Yagjagbula, are said to be from Yiwarlarlay itself (and not to have come from anywhere else), the frogs and rain (wiyan) as- sociated with the story are said to have come from the south, watching the brothers fight. In this case, the Yiwarlarlay site and its associated story are extremely significant to both Mudburra and Warlpiri peoples of the south (Walsh, 1988). Further links with the south are implied in that Warlpiri, Mudburra and Wardaman peoples, amongst others, all share a similar 8-class kin- ship system (along with a similar terminology), as well as the practice of subincision. The pat- terning of both the 8-class system and subin- cision not only links Wardaman social practice with peoples to the south and west, but also sets them apart from peoples to the north and north- east, where both subincision and the 8-class sys- tem are not practised. With respect to trade, little information is available. It is assumed that given the widespread interactions between Wardaman people and those to the south and west, both material goods and ideas, and with them stylistic conventions, have been extensively shared (see below). Items traded include reed/bamboo spear shafts from the Daly River to the northwest of Wardaman country, in return for stone spear points (Riley Birdun, Wardaman man, pers. comm., 1989). Ground stone axes made from greenstone have been located on the surface of Wardaman sites. Visual inspection of these axes points to an original Western Australian source, although more detailed petrological analysis, planned for the near future, may pinpoint the source more precisely. In short, the available evidence,as limited as it is, points to widespread contacts with peoples to the west and south. As will be argued below, these interactions have resulted in the diversification of rock art styles in specific locations, whilst at the same time standardising artistic conventions within the broader interacting sphere. The region is set apart in terms of rock art from northern regions, where other forms of rock art (especially x-ray art) are common, and we believe that this geographical separation of artistic conventions reflects broader issues centring upon the struc- 445 ture of alliance networks and patterns of inter- regional interaction. YINGALARRI ROCK PAINTINGS The rock art of the Yingalarri region analysed below was recorded in 1988 during the first season of the Lightning Brothers Earthwatch Project. In the field, the rock art was extensively photographed and sketched with annotated drawings of each picture being undertaken. The ensuing preliminary classification presented in this paper was constructed by allocating each picture a technique category (Y-axis) and a for- mal category (X-axis) (Fig..2). The current paper is concerned purely with a preliminary examina- tion of the technique characteristics of the rock paintings from the region. The formal characteristics of the paintings of the Yingalarri region are extremely varied. By far the most numerous are anthropomorphs (43.2%), with zoomorphs (28.9%), abstract designs (23.9%), boomerang-forms (0.5%), track-forms (0.7%), and images of contact (European) objects (0.8%) also present. In this preliminary analysis, 45 motif forms have been identified (see Appendix 1), 33 of these consist- ing of biomorphic forms (anthropomorphs and zoomorphs). These can be divided into eleven painting technique types (Fig. 2): A: linear. These include ‘stick figures’ and other, non-area paintings. These are rare and very dif- ferent from the ‘stick figures’ documented from the Hammersley Ranges to the south-west (Walsh, 1988) and those of the earlier periods of the Arnhem Land regions to the north (Chaloupka, 1984; Lewis, 1988; Brandl, 1973) (Fig. 3); B: enclosed linear, defined as linear motifs (Type A) enclosed by an outline. Extremely rare and not known from elsewhere (Fig. 4); C: outlined; very common and widespread throughout Wardaman country and beyond, especially to the west and south, including the Victoria River region all the way to Keep River and further west in Western Australia (Lewis and Rose, 1988); D: infilled; very common, and found commonly throughout most of Australia, and especially the northern half of Queensland, the Northern Territory and Western Australia, Chaloupka (1984) has ar- gued that similar paintings as some infilled biomorphs from the Yingalarri region have been found in late Pleistocene rock art in the Alligator Rivers region (but see Lewis, 1988) (Figs 5 and 6). E: striped. These include a range of longitudinally MEMOIRS OF THE QUEENSLAND MUSEUM 446 So avea-x c xaTIdwoD ‘ant/1n0 ‘9380 /4N1 ‘o30/1n0 a1d0wis oadistlis WAN anming uvann x Fic. 6. Type D (infilled) contact image and Type G (outlined and internally decorated) zoomorphic painting. MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 7. Type E (striped) paintings, YINGALARRI CAVE PAINTINGS figures (type E) appear much more commonly under other paintings (especially outline/infilled ones) than over them (Table 2). These results, however, should be further explored by increas- ing the sample size, a project planned by the authors for the near future. In short, four sites, Yingalarri 2, 3, 21 and 22 separate out as significantly different from the other Yingalarri sites on the basis of their paint- ings. These differences can be best explained by the fact that these sites contain more paintings than the other sites (with the exception of site 4), although sample size itself is not a contributing factor as all variables from each site were given equal weight. An examination of superimposi- tions in these sites has failed to locate any clear change in painting technique through time, al- though it is possible that longitudinally striped biomorphs are earlier depictions, whilst the x-ray and outlined-infilled paintings are relatively recent, The recent beginnings of x-ray paintings is supported by their fresh appearance, and this may reflect recent interactions with peoples from the north, where very similar x-ray (fish) repre- 455 sentations are common. Such paintings are not found to the south or west of Wardaman country. In summary, we emphasise 5 points: 1) the Yingalarri paintings are dominated by biomorphic paintings, undertaken in outline, in- fill, outline/infill, outline/infill/internal decora- tion, or striped longitudinally. The latter in particular are found only in Wardaman country and to the south and west, with some very rare examples found in Jawoyn country. All other painting techniques are mostly found to the west, with the exception of some extremely rare x-ray paintings (mainly fish), which are charac- teristically northern influences. 2) Yingalarri sites 2-49 cannot be divided into distinct groups on the basis of the techniques used to make the paintings which decorate these sites. 3) Four sites separate-out as significantly dif- ferent. These differences can be best explained by reference to site size: a) the four sites, having more paintings than any other site in the sample, may have witnessed more painting episodes than the other sites, and as such a more heterogeneous painting assemblage has resulted in these sites, Fic, 8. Type F (sectioned) painting, 456 orb) this may purely be attributable to significant differences in site function. 4) The surviving paintings do not show any conclusive evidence of changing techniques through time. All techniques are roughly con- temporaneous, although some techniques may indicate recent influences from the north (x-ray paintings) (see also Mulvaney, 1975, p.273). 5) From 4) above, we suggest that the paintings observed reflect largely contemporaneous in- fluences from numerous sources by local popula- tions. The art may reflect more the particular painting styles of individuals who’s styles have differentially developed via influences from numerous places, rather than representing a strict cultural convention that can be specifically iden- tified as ‘The’ Wardaman rock painting style (as distinct from neighbouring ‘styles’). In a later paper, these issues will be further addressed by us in a more detailed analysis of the paintings. This analysis includes both an inves- tigation of the distribution of motif forms (see Appendix 1) and particular stylistic elements MEMOIRS OF THE QUEENSLAND MUSEUM (e.g. type of internal decoration). We present below a preliminary discussion of the above points in relation to documented Wardaman inter- regional relations. DISCUSSION Perhaps the most influential debate in the his- tory of Australian prehistory has fallen under the guise of the ‘Intensification Debate’ (Lourandos, 1983; Beaton, 1983). It has recently been the subject of focused attention, and represents a critical stage in Australian prehistory as it has addressed a number of questions which have re-oriented the nature of archaeological enquiry in this country. It has not only brought the issue of intensification, for long a major source of discourse in international circles, to Australia, but more importantly, it has brought it out of its traditional “hunter-gatherer to farmer’ frame of reference to one addressing hunter-gatherers as capable of ‘intensifying’ without venturing out of the hunter-gatherer mode of production. In Fic. 9. Type H (infilled and internally decorated) zoomorphic painting (uppermost painting) and Type J (outlined, infilled and internally decorated) zoomorph (lowermost painting). YINGALARRI CAVE PAINTINGS 457 Fic. 10. Type J (outlined, infilled and internally decorated painting). 458 other words Aboriginal society is not seen as a static socio-economic network, but rather is referred to as a dynamic socio-ideological sys- tem, subject to the same kinds of influences characterising farming or agricultural societies. The questions asked of prehistoric Aboriginal life are now more directed at understanding the nature of inter-personal relations (i.e. at the power structures which mediate the maintenance of social traits and changes there-in), questions which traditionally have been the realm of ar- chaeologists studying more complex (structural- ly differentiated) societies. It is in questioning the nature of inter-personal relations that we may best attain information on the social forces which serve to both re-direct and maintain trajectories of change. Lourandos (1983, 1984) has recently taken up these issues with reference to Australian prehis- tory. He has argued that in many parts of Australia, and in the Victorian southwest in par- ticular, the very structure of alliance networks MEMOIRS OF THE QUEENSLAND MUSEUM has witnessed a fundamental shift during mid- to late-Holocene times. These shifts, he believes, are visible archaeologically in the following ways: 1] there is an increase in use of individual sites; 2] there is an increase in the rate of estab- lishment of new sites; 3] there is a diversification and increase in the complexity of resource management strategies; 4] there is an increase in the use of marginal environments (see also Flood et al., 1987; David, 1987; Walters, 1989); 5] the development of widespread trading net- works takes place. Together, Lourandos sees these changes as reflecting an intensification of socio-economic variables, where these can refer to ‘economic as well as social variables which may themselves bear directly or indirectly upon economy’ (Lourandos, 1983,p. 81). Yet in spite of the far-reaching implications of 1315 14 3923 33 12354142 44 494847 452746 7 291836 1734 6 4020433231 3722 5 211125383 4 2 Fic, 11. Average Linkeage Cluster Analysis, showing site numbers. YINGALARRI CAVE PAINTINGS Lourandos’s work, it is surprising that little at- tempt has yet been made to investigate the dis- tribution of, and changes in, particular stylistic conventions, and this is especially so of rock art. One notable exception has been Lewis (1988), who has argued that during the height of the last glaciation, an extensive interaction network ex- isted throughout the Bonaparte catchment, which extended from the Kimberleys to the Vic- toria River, Arnhem Land and beyond (including Wardaman country). In relation to climatic chan- ges (especially the development of wetlands) and population increases during the late Holocene, Lewis (1988) sees the rock art of northern Australia becoming more regionalised during the last 2000 years or so. Although we agree with the directions of Lewis’ (1988) work, we have ar- gued in this paper that the patterning of stylistic conventions in Wardaman rock paintings does not reflect any significant changes in this region, and we therefore argue that there have been no major changes in inter-regional alliance net- works for the period documented by the region’s surviving rock paintings. We briefly discuss the probable antiquity of this body of rock art below, situating it in a broader temporal sequence. We 459 conclude below with some implications for the Intensification question. CONCLUSION A number of factors lead us to believe that most, if not all, of the surviving rock paintings from the Yingalarri region are of relatively recent age (late Holocene): 1) The sandstones of the region are soft and friable, unlike those of the Kombolgie Formation to the north (where much of the Kakadu rock art is located). Chaloupka (1984, p. 53) notes, for instance, that sandstones found on the Amhem Land Plateau are far more stable than those found in Wardaman country. The conditions for preser- vation of long-standing paintings are not widespread in Wardaman country. 2) Despite the elaborate, figurative nature of Wardaman rock paintings, no evidence of extinct fauna has yet been found in the region. This is so despite the fact that 109 sites and 3379 paintings have so far been systematically recorded. This is in direct contrast to the situation to the north, where numerous examples of extinct fauna (i.e. Thylacines) have been found (Lewis, 1977). Fic. 12. PCA on sites Yingalarri 2-49, showing 0.5 level of significance (boxed area). 3) Compounding |) above, dramatic increases in precipitation, and increases in conditions promoting exfoliation of rack surfaces, before 3000 years ago, would mitigate against the stabilisation of rock surfaces in friable sandstone outcrops. 4) Excavations of two sites at Yiwarlarlay have shown: a] dramatic increases in occupational intensity, along with presence of ochre, in tate Holocene levels; b] the presence of exfoliated, painted wall cor- lex only in very recent times, 5) Excavations at Mennge-ya (near Yingalarri} have revealed dramatic increases in occupational intensity and in amounts of ochre in late Holocene Jayers. 6) Paintings always overlie patinated peckings when superimpositions occur. No example of peckings over paintings are known to us despite the fact that 18,354 pictures (engravings, paint- ings, stencils, prints) have been recorded from Wardaman country, 7) The creation of engravings have been dated Stratigraphically to over 5000 years by Mulvaney (1975) at the Yingalarri 1 (Ingaladdi) site. How- ever, it is likely that some of these (abraded grooves especially) have been created con- tinkously for long periods of time, as both Mer- lan (1988) and Flood er al. (in press) document the recent making of abraded grooves by War- daman people, No examples of paintings under- lying abraded art have been observed. 8) The fragility and therefore instability of paintings is re-enforced by the fact that numerous clear paintings [rom Yingalarri which were photographed in the 1960s have since disap- pearcd to the point of total disintegration (mainly fading), As a result of these observations, we suggest that the paintings from the Yingalarri region are mostly, if not entirely, relatively recent (late Holocene). Furthermore, as is shown in the dis- tribution of superimpositions {Table 2}, no major temporal changes in painting techniques have been noted for sites 2-49, As a result of these observations, we suggest that {here is no evidenee for any major shift in the distribution of painting conventions in the Yingalarri region during jhe late Holocene, By implication the alliance networks observed ethnohistorically from the broader region (which have been central to the dispersal of painting conventions throughout the broader region) have been operat- ing perhaps throughout the late Holocene period MEMOIRS OF THE QUEENSLAND MUSEUM (and, as previously argued by Lewis [1988]. linked the Wardaman region with the Victoria River and western areas). In effect, we suggest alse that the seeds to the alliance networks docu- mented during ethnohistoric times were already in place during or hefore the mid-Holocene. We say this as 4 result of the great similarities be- tween the patinated engravings of whal is today Wardaman country (some of which have been dated stratigraphically at Yingalarri 1 to over 5000 years BP), and those to the south and to the west. The similarities in the carly engravings may even document a system of alliance net- works far broader than that documented eth- nohistorically, as the ‘early’ engravings are stylistically similar to those of Keep River and beyond to the west, and to those of numerous parts of South Australia (e.g. Meadow’s Bluff) to the south. Engravings are. considerably rarer in the north and, contra Lewis (1988), this may imply considerable antiquity for an Arnhem Land-Victoria River cultural discontinuity. Al- though these issues will form the subject of a separate study, We suggest that early Holocene: socio-organisational systems were more open than those observed ethnohistorically, perhaps stressing the maintenance of social relatedness, a feature of arid and semi-arid life which Myers (1986, p, 164) has argued is central to survival and everyday life in central Australia. Purther assessment of these questions will, however, have to aWail a more detailed analysis of the rock arnt found in both what is today Wardaman country und beyand. ACKNOWLEDGEMENTS We would first and foremostly like to thank the many Wardaman people who showed us and allowed us to tecord theirsites. Special thanks go to Ruby Alison, Riley Birdun, July Blutcher, June Blutcher, Daisy Gimin, Lily Gingina, Raymond. We would also like to thank the numerous Earthwatchers from the 1988 season for con- tributing so much to this project, and to Frances- ca Merlan for sharing so much of her anthropological knowledge with us, For useful comments and discussions, bur thanks go to lan McNiven, Mike Morwood, Harry Lourandos, Darrell Lewts, Francesca Mer- lan, Su Higgins, Bryce Barker, and Jobn Clege, YINGALARRI CAVE PAINTINGS whilst we are extremely grateful to Richard Wright for making his VOTRANS program available to us. LITERATURE CITED BEATON, J. 1983. Does intensification account for changes in the Australian Holocene archaeologi- cal record? Archaeology tn Oceania 18: 94-97. BRANDL, E. 1973. ‘Australian Aboriginal Paintings in Western and Central Arnhem Land’. (A,1.A.S.; Canberra), CHALOUPKA, G, 1984. ‘From Palaeoart to Casual Paintings’, (Northern Territory Museum of Art and Sciences: Darwin). Davin, B. 1987. *Chillagoe: from Archaeology to Prehistory. Contributions to a late Holocene prehistory of the Chillagoe region’. (Un- published M.A, thesis, A.N.U,, Canberra), Davipson, D.S. 1935. Archaeological problems of Northern Australia, Journal of the Royal Anthropological Institute 65: 145-83. FLoop, J., Davip. B., AND Frost, R. in press. Dream- ing into Art: Aboriginal interpretations of rock engravings, Yingalarri, Northern Territory (Australia). / Morwood, M. (ed.), “Ethnography and Art.’ (preliminary title). (A.LA.S.; Canber- Ta). FLoop, J., DAVID, B., MAGEE, J. AND ENGLISH, B. 1987. Birrigai: a Plesitocene site in the southeastern highlands. Archaeology in Oceania 22; 9-26. Hucnet, B. 1988. Striped anthropamorphs from Laura. (Unpublished paper presented at the A.A.A. conference, Armidale). Lewis, D. 1977. More striped designs in Arnhem Land tock paintings. Archaeology and Physical Anthropology in Oceania 12 (2): 98-111. 1988. “The Rock Paintings of Ammhem Land, Australia: social, ecological and material culture change in the post- glacial period’. (BAR Inter- national Series 415; Oxford), Lewis, D. AND ROSE, D. 1988. ‘The Shape of the Dreaming: the Cultural Significance of Victoria River Rock Art’. (A.1,A.8.: Canberra). LaurANbos, H. 1983. Intensification: a late Pleis- tocene-Holocene archaeological sequence fram southwest Vicloria. Archaeology in Oceania 18: 81-94. 1984. ‘Changing perspectives in Australian prehis- tory: a reply to Beaton. Archaeology in Oceania 19 (1): 29-32. McCartny, F. 1939. *Trade’ in Aboriginal Australia, and ‘trade’ relationships with Torres Strait, New 461 Guinew und Malaya. Oceania ¥ (4): 405-38, 1: 80-104, 171-95. 1967, ‘Australian Aboriginal Stone Implements’. (Australian Museum: Sydney), MERLAN, F. 1988. The interpretive framework of War- daman rock art: a preliminary report. (Paper presented atthe Ist AURA conference, Darwin), MULVANEY. D.J. 1975. ‘The Prehistory of Australia’. (Penguin: Ringwood). Myers, F. 1986. ‘Pintupi Country, Pintupi Self’ (A.1.A.S,: Canbetra). SPENCER, W.B, 1914. ‘Native Tribes of the Northern Territory of Australia’. (Macmillan: London). STANNER, W.E.H. 1959-63, ‘On Aboriginal Religion’. (Oceania Monograph: Sydney). WALSH, G. 1988. ‘Australia’s Greatest Rock Art’, (EJ. Brill/R. Brown & Ass.: Bathurst), WALTERS, |. 1989. Intensified fishery production at Moreton Bay, southeast Queensland, in the late Holocene. Antiquiny 63: 215-224. APPENDIX 1 Description of the 45 Motif Forms identified from sites Yingatarn 2-49 in this preliminary classification system (see Fig. 2). In the descriptions below, the number corresponds to the Motif Form number as is shown on Fig, 2, and this is follawed by a key word description (e.g. 1] elongated anthropomarph). 1] elongated anthropomorph, where length of painting is al least twice its width. 2] split anthropomorph, where the image is either split along the entire length of the bady, or atits junction with the arms (giving the impression of an absence of torso). 3] elongated limbed anthrapomorph, where arms/legs aré elongated, often showing presence of joints (elbows, knees). 4| detailed anthropomorph, where facial and body features are detailed (e.g. eyes and mouth especial- ly). 5] split face anthropomorph, where the face is split into two halves. 6] dynamic anthropomorph, where body features are standardised into geometric elements (e.g. square head, oval breasts, circular elbows), and there is a intal absence of any detail of internal features (such as eyes, mouths, etc.). 7] generalised anthropomorph, where only the basic body shapes are shown, 8] other anthropomorph. This category refers to one- off anthropomorphs thal do not conform to 1)-8] above. Further surveys will, hopefully, increase the sample sizes of these pictures, and each form will 462 be treated separately and further described in our final report. 9) indeterminate anthropomorph, where the picture can be identified as an anthropomorph but is too faded or damaged to further identify. 10] macropod, whete the picture has the formal char- acteristics of a mammalian quadruped, and where the fore-limbs are noticeably shorter than the hind- limbs. 11] canid, where the picture has the formal charac- teristics of a mammalian quadruped, where the limbs are all of approximately equal size, and where ears and tail resemble those of dogs. 12] other quadruped, where the picture does not con- form fo either 10] or 11] above. 13] emu, where the body is extremely large relative to the head, and where the formal characteristics of the image resemble |hose of birds, 14] long-beaked bird, where the beak-length is pronounced, and the image has the formal charac- teristics of a bird. 15] generalised bird, where the image has the formal characteristics of a bird, but where no other distinct feature can be identified. 16] owl, where the picture consists of a generalised oval body with the head above it, and where the head is divided into two equal halves. 17] short bird, where the image is of a biped whose length is approximately equal to its width. where the picture is relatively small (SOcm or less in length), and where the legs emanate {rom the side of the body (rather than from underneath it), 18] other bird, ane-olf pictures where the image resembles a bird, and where specific features distin- guish it from 13] to 17] above. 19] snake, being elongated images with identifiable heads and/or tails. 20] eared snake. As with 19] above, with the addition of one or two ears above the head. 21] turtle, being four-limbed zoomorphs with head and sometimes tail emanating from a circular body, 22] echidna, having the formal characteristics of echid- nas. There is a linear extension protruding from a roundish head when in plan view, whilst the linear extension emanated from the body when in profile. 23] indeterminate echidna/turtle, being pictures which are loo faded or damaged to determine whether they belong to category 21] or 22], 24] crocodile, having the formal characteristics of crocodiles, and where head is pictured as relatively circular and followed by an elongated ‘snout’. MEMOIRS GF THE QUEENSLAND MUSEUM 25| lizard, similar to 24] above, but where the head is elongated, 26] indeterminate cracodile/lizard, where the picture resembles 24] and 25] above, but where it cannot be identified as cither one or the other. 27| indeterminate crocodile/lizard/fish, where the image cannot be differentiated between 23}, 24] above or 28] below, 28] fisk, where the picture resembles a fish (presence of fins, tail and head), 29] bat, where image resembles a bat/flying fox, These are either upright ar up-side down, These are small, generalised bipedal badies with heads and relative- ly large ears. 30) unidentified quadruped, being a four-legged and tailed zoomorph whose formal characteristics do not resemble those of any known animal. 31) unidentified biped, being a two-legged biped whose formal! characteristics do not resemble those of any known animal, 32] star zoomorph, being a tailed-image with a star- shape in head position, and where two circles occur in the position of the eyes. 33] indeterminate zoomorpk, where degree of fading. and/or damage precludes it from further identifica- tion, 34] foot, being a toed track-form resembling the shape of a human foot. 35] macropad track. being a track-form which resembles (either singly or in pairs) the tracks of macropods. Side toe(s) must be present. 36) bird track, being (hree- or four-pronged pictures. resembling the shape of bird tracks, and where the meeting point of the prongs must be angular. 37] concentric circles. including single circles. 38] geometric non-figurative, these being one-off geometric non- figurative piclures which do not conform to 39], 40,41] or 42] below. 39] tally-forms, being series of dots or parallel lines. 40] grids. 41) matchsticks, being geometric linear shapes with circular ‘head’ above them. These sometimes have tadiating lines emanating from the ‘heads’. 42) half-suns, being geometric semi-circles with radiating lines. 43] other non-figurative, being amorphous non- figurative designs. 44] boomerang-shapes, being, curvilinear pictures re- sembling boomerangs, or hooked curvilinear pictures. 45] contact images, being pictures having formal char- acteristics of post-contact items (e.g. guns, person on horse). NEW AND RARE CRABS OF THE SUBFAMILY DOTILLINAE (CRUSTACEA: OCYPODIDAE) FROM NORTHERN AUSTRALIA AND PAPUA NEW GUINEA P.J.P. DAVIE Davie, P.J.F. 1988 05 31: New and rare crabs of the subfamily Dotillinae (Crustacea : Ocypodidae) from northern Australia and Papua New Guinea. Memoirs of the Queensland Museunr 28(2):463-473, Brisbane. ISSN 0079-8835, Two new species of the previously monolypic genus Tmethypocoelis, T. koelbeli and T. oduntadactylus, are described. They are separated from each other and from 7. ceratophora (Koelbel) by their distinctive male pleopods, and cheliped characters. //yoplax strigicarpus is also described, and is separated from its closest ally. /. orientalis, by the shape of the lower orbit and the male first pleopod. /lyaplax dentains is discussed and the male and female abdomens and male pleopod are figured. [1 Crustacea, Ocypodidae, Dotillinae, Tmethypocoelis, Hyoplax, new spectes, Australia, Papua New Guinea, P.J.F. Davie, Queensland Museum, PO Box 300, South Brisbane, Queensland 4101, Australia; 14 August, 1989, The ocypodid subfamily Dotillinac has long been referred to 4s the Scopimerinae, but Man- ning and Holthuis (1981; 192) point out that Dotillidae Stimpson, 1858, is older than Scopimerinae Alcock, 1900, and therefore has priority. The dotillines are poorly represented in Australia; only Scopimera inflata A. Milne Ed- wards and /lyeplax dentatus Ward have been previously recorded. This paper records two new species bringing the total number to four. All appear to be endemic, Only Scopimera kochi Roux, 1927, has been previously recorded from New Guinea, All specimens examined are in the collections of the Queensland Museum (QM), the Australian Museum (AM), or the Wesiern Australian Museum (WAM), Genus Tmethypocoelis Koelbel, 1897 Tmethypacoelis Koelbel, 1897, p. 715.* (Type species: Dioxippe (Tmethypocoelis) ceratophora Koelbel, 1897, by vriginal designation, sub- sequently elevated by Shen, 1935). DIAGNOSIS Scopimeninae with ocular peduncle prolonged beyond the cornea as a long styliform projection. Sub-orbital margin cut into two parts by a deep groove which runs obliquely and medially downwards; the inner part has twa transverse granular ridges separated by a groove. The en- dopod of the second maxilliped has an ovate palp. Chelipeds subequal. Second maxilliped * Nol seen, pagination follows Shen (1935) and athers, although Tesch (1918) gives p. 573. with the penultimate segment not expanded and with the ultimate segment attached terminally. Carapace with upper surface and lateral walls nol conspicuously sculptured. Ambulatory legs with large tympana. REMARKS Within the Dotillinae, Tmethypocoelis shows closest affinities with the genus Jlyoplax and, indeed, if was originally considered a subgenus of Ilyoplax (then referred to by the preoccupied name Dioxippe de Man), Tmethypacoelis shares in common with /lyeplax: subequal chelipeds; a similar type of second maxilliped that does not have the penultimate segment expanded, and has the ultimate segment attached terminally; and, the upper surface and lateral walls of the carapace are not conspicuously sculptured. The most unusual character of Tmethypocoelis is the long styliform projection on the cornea, This. character is not unique to Tmethypocoelis as it alsa occurs in a number of species of Ocypode and Uca (see Barnes, 1968; Hagen, 1970). In itself the ocular projection would not be sufficient to delimit the genus, however, com- bined with the grooves of the sub-orbital margin and the extremely consistent overall appearance of the chelae and carapace, the three species now described must be considered generically dis- lincl. Shen's (1935) diagnosis gave eight chatac- ters to separate Tmethypocoelis from Tlyaplax. My diagnosis is somewhat shorter because, in the hight of the new species described here, U decided that several of his characters were useful only at the specific and not the generic level. 464 —— Imm MEMOIRS OF THE QUEENSLAND MUSEUM a Imm Fic. 1. Tmethypocoelis koelbeli sp. nov. A, male abdomen (paratype, QM W7982); B, male first pleopod (paratype, QM W7982); C, female abdomen (paratype, QM W7983). Tmethypocoelis koelbeli sp. nov. (Figs 1,2,6A) MATERIAL EXAMINED HOLOTYPE: QM W7981, d (6.9 mm), South Alligator R., Northern Territory, P. Davie, 11.5,1979. PARATYPES: QM W7982, 6d ¢ (7.3, 7.1, 7.0, 6.8, 5.9, 5.7 mm), same data as Holotype. QM W7983, 1d (5.5 mm), 322 (6.0, 5.8, 4.6 mm), East Alligator R., Northern Territory, P. Davie, 30.4.1979. QM. W15085, 22.¢ 6 (5,3-8.2 mm), 1 2 (6.7 mm), Magela Ck, East Alligator River, Kakadu National Park, Northern Territory, 13.vi.1981, P. Davie. DESCRIPTION Carapace: Approximately pentagonal, some- what flattened. Convex along mid-dorsal line, slightly convex laterally. About 0.6 as long as broad. Regions semi-defined; epigastric lobes slightly swollen, joined in a depressed arc behind front but separated behind. Ovoid mesogastric region prolonged anteriorly between epigastric lobes; cervical groove dividing gastric and car- diac regions, short and distinct, narrow medially, widening laterally. Cardiac region with a slight central depression. Orbital, hepatic and branchial regions not distinctly separated; with undulating surface. Branchial region sloping, with irregular small setiferous tubercles. Sub-branchial region bulging, regularly hairy, separated from branchial region by sinuous lateral border which is hairy and minutely granular in its anterior half and with closely spaced short hairs posteriorly. Front at base about one-fifth distance between external orbital angles. Side borders slightly con- vex, converging; frontal angles rounded. Anterior borders with sides oblique and shallow- ly concave; a central blunt prominence. Supra- orbital borders sinuous, sloping backward, microscopically beaded particularly towards ex- ternal orbital angles. External orbital angle, with edges more or less serrulate; posteriorly fol- lowed by a broad U-shaped sinus which con- tinues as a depressed oblique channel! onto dorsal surface behind — supra-orbital = margin. Epibranchial angle capped by small tubercle. NEW AND RARE DOTILLINE CRABS Fis. 2. photographs of apex of male first pleopod (paratype, QM W7982). Scale line = 0.1 mm. Tmethypocoelis. koelbeli sp. nov. S.E.M, Distance between epibranchial angles subequal or slightly less than between external orbital angles. Hind margin slightly concave and about two-thirds distance between external orbital angles; a fine ridge parallel with the hind margin forms a broad tim. Infra-orbital border projects beyond supra- orbital, and is of two parts separated by a notch, from which on the pterygostome, runs an inward- ly directed oblique channel. This channel forms a tilted Z-shape and terminates above the base of the chela. Inner part of infra-orbital border about equal in length to outer part and consists of two tows of large granules separated by a concavity. Anterior row follows the arc of the arbit, posterior row is slightly itregular, almost straight, and converges on but does not intercept the anterior row at the notch. Just behind this second row is a line of closely spaced long feathery hairs which extend from the base of the eyestalk and are longest behind the notch. Outer 465 part of infra-orbital border also granular; ter- minates just below external orbital angle such that a broad notch is formed. Undersurface of the external orbital angle with a brush of feathery hairs. Side walls perpendicular anteriorly but sloping outwards at the sub-branchials; regions not separated: setiferous tubercles over whole area. Eyestalks: Widen distally; cornea bulging; medial thickening gives twisted appearance; usually reach level of external orbital angle. Eyestalk projects beyond the cornea in the form of a long style, which in adult males is equal to, or longer than cornea, but shorter in younger specimens. Female style short and fine, almost like a stout hair. Tip with two or three stiff hairs disposed in single file. Always a long stout hair placed just before cornea and projecting up- wards. External maxillipeds: Do not close buccal cavern, slightly vaulted. Ischium subquadrate, although antero-internal angle produced as an obtuse lobe. Dense fine hair longest on postero- internal curve and extending dorsally up inner side, Anterior margins slightly concave. A line of fine hair runs obliquely from anterior margin near antero-internal angle and then down outer margin, increasing in Jength so that are longest at the outer margin. Merus slightly larger than ischium, Lateral margins converge distally; inner margin straight with long feathered hairs, outer margin slightly convex with short pile. Merus and ischium covered with very short scattered hairs. Carpus occupies the narrow anterior mar- gin of merus as with 7. cerataphora. Dactylus slender and twice length of propodus. Long feathered hairs apically. Abdomen: Terminal segment rounded. Penul- timate segment slightly longer than fifth seg- ment, and with slightly concave, parallel sides. Fifth segment with straight sides, narrowest at base. Fourth segment expanded, Female ab- domen as figured. Chelipeds: Massive; long bui not remarkably; subequal. Merus is sharply three faced with seru- late borders; tympanum on inner surface, broadly oval; feathery hairs. on whole inner surface in- cluding tympanum. Outer face granulate distally and with tympanum smaller and elongate. Car- pus 4 little elongated in large males, but not so in smaller males and in females; distal internal angle is extended to form tooth-like articulation point with the palm; unarmed except for fine serrulation of inner and outer borders; inner face in large males with scattered, pointed granules proximo-dorsally and a short near vertical ridge of large rounded granules ventrally, Palm bulky, length approximately three-quarters distance be- tween ouler orbital angles, height about half length, and about equal to length of immovable finger; outer surface granulate dorsally, granules largest distally; smooth ventrally. Large granules down edge of gape between fingers, on both outer and inner surfaces. Inner surface evenly and finely granulate on upper half. These granules extend over the dorsal curve to the sharply cut superior border of the outer surface. Both upper and lower borders finely granulate. Lower border extends onto, and is obvious, for about half length of immovable finger. Small group of long hairs on upper surface just behind the articulation joint of dactylus, Fingers gaping at base. Both fingers curved inwards, expanded distally to form spooned tip. Cutting margins with even rows of tecth, however on moveable finger of smaller males, a raised platform of teeth is differentiated in proximal half and some trace of this is usually evident in fully mature chelae. Inside surface of moveable finger with short irregular line of large tubercles just above cutting margin near base. Band of finc granules on dorsal surface tukes the same form as palm. Superior border straight, extending three-quarters length of finger; terminates in an overhanging “shelf because of sharp inward turning of tip, Outer surface with two subregular lines of granules originating at base; development of granules and length of line variable. Superior one may extend three-quarters of length to lip, lower one a little less. Inner margin at tip of both fingers with a short row of 6-10 stout hairs. Spooned tips with comcous edge. Chelipeds of females of small and simple “ocypodid’ type, Ambulatory legs: Meri approximately two and a half times as long as broad, as long as the next two joints together. Upper and lower margins convex, Upper margin slightly crenulate distally, otherwise smooth. Provided both sides with oval tympana. Tympana large and of same relative size on anterior surface of all meri; on posterior surface, become progressively smaller from Ist lo 4th legs. Second pair of legs the longest, second and third pairs both Jonger than firsi; fourth pair the shortest. First pair a little longer than distance between external orbital angles. Carpo- and propodites with fine bristles. Dactyli nearly straight, flattened dorso-ventrally, pointed, shorter than propodites. Closely spaced fine hairs laterally, not extending to tip. Length of hairs tapering distally, MEMOIRS OF THE QUEENSLAND MUSEUM Hairy edged pouch present between bases of first and second, and second and third walking legs, Hair tults are long, thick, and conspicuous and extend to bases of meri. First male pleopod: As figured. Colour: Chesinut brown to grey with white chelae. HABITAT Burrow in soft moist mud banks in upper es- tuary, low salinity, mangrove situations. REMARKS This species is only known from the Alligator Rivers System, Northern Territory. It differs most conspicuously from 7, ceratophera, by being not quite as broad (c. 0.6 as long as broad, as Opposed to c. 0.5 in T. ceratophora); having a distinctive first male pleopod; and having the carpus of cheliped not as elongated in adult males. Jt differs from 7. odontodactylus by the shape of the first male pleopad and the form of the chelae. Tmethypocoelis odontodactytus sp. nov. (Figs 3,6B) MATERIAL EXAMINED HovotyPr: WAM 953-88, 3 (7.7 x 4.5mm), Mouth of Gogol River, South of Madang, Papua New Guinea, sand, intertidal, 1.vil.1987, GJ, Morgan. PARATYPES: WAM 218-88, 8c d (6.1-9.3mm), 52 9 (5,6-7,6mm), data as for holotype, OM W15385, 2d 9 (6.2, 8.6mni), 12 (6.6mm), data as tor holotype. DESCRIPTION Carapace: Approximately pentagonal, evenly convex longitudinally, slightly convex laterally; about 0.6 (0.57-0.61) limes as long as broad; regions semi-defined. Frontal region furrowed; epigastric lobes small, not joined anteriorly; mesogastric region ovoid and slightly swollen; cervical groove separating gastric and cardiac regions is quite wide and widens into depressed areas laterally bordering the cardiac region; in- testinal region separated by a shallow furrow. Orbital, hepatic, and branchial regions not dis- tinctly separated from each other, marked by several short lateral, setiferous ridges and the whole arca laterally with short strong setae. Sub- branchial region bulging, regularly hairy, separated from branchial region by a sinuous lateral border of short stout setae. Front at base about one-fifth distance between NEW AND RARE DOTILLINE CRABS Imm 467 0.2mm ol | — Imm Fic. 3. Tmethypocoelis odontedactylus sp. nov. A, paratype WAM 218-88 (7.6 mm), female abdomen; B-D, paratype male, WAM 218-88 (9.1 mm); B, third maxilliped (denuded); C, left chela; D, abdomen; E-G, holotype male, WAM 953-88 (7.7 mm), first pleopod, and magnifications of apex. external orbital angles; lateral borders slightly convex, converging; anteriorly concave either side of a central blunt point. Supra-orbital bor- ders sinuous, sloping backward, microscopically beaded. External orbital angle bluntly pointed, granulate; followed by U- shaped sinus which forms a depressed oblique furrow onto dorsal surface behind outer half of orbit. Epibranchial angle blunt and at the same level as external orbital angle although may be slightly more protruding on small specimens. Hind margin slightly concave, about two-thirds of the distance between the external orbital angles; with a broad smooth rim. Infra-orbital border projects beyond supra- orbital and consists of outer and inner sections of about equal length, and separated by a notch; inner part consists of two rows of granules separated by a deep furrow, outer part with small granules on border, terminating below the exter- nal orbital angle such that a broad notch is formed; ventral margin of external orbital angle with a brush of feathery hairs. Side walls perpen- dicular anteriorly but sloping outwards at the sub-branchials; covered in short stout setae. Eyestalks: Widen distally; slight medial thick- ening, cornea bulging; usually reach level of external orbital angle. Style projects beyond cor- nea in males; length variable but usually slightly more than length of cornea; tipped with two or three long bristles. A stout hair usually placed on stalk just before the cornea, and projecting up- wards, External maxillipeds: Do not close buccal 468 cavern; slightly vaulted; internal margins and palp with thick lining of fine setae; outer margins with short sparse setae. Ischium subquadrate, a little broader than long, and with the internal superior angle produced as an obtuse lobe. Merus distinctly larger than ischium, lateral margins convergent distally; palp occupies the narrow anterior margin. Abdomen: Telson rounded, about the same length as fifth segment; sixth segment longest; fifth segment basally constricted; fourth segment expanded; second segment a thin strip. Female abdomen a wide flap as figured. Chelipeds: Massive in males, subequal; merus trihedral with serrulate borders; broadly oval tympanum on proximal two-thirds of inner face; outer face with a smaller elongate tympanum near lower border. Carpus elongated (length c. 1.5 times breadth), sub-rectangular, granulate borders. Palm bulky, height about half total length, and equal to length of moveable finger; outer surface finely granulate mid-dorsally, granules largest distally, smooth ventrally, larger granules along edge of gape. Lower border granulate except for distal portion of immovable finger. Upper half of inner surface evenly and finely granulate, granules extending over the dorsal curve to the sharply defined superior bor- der of the outer face. Both fingers curved inwards distally, and expanded distally to form spooned corneous tips; cutting margins are evenly toothed although on the proximal half of the moveable finger the teeth are a little more elevated. Move- able finger with a medial granulate ridge running the whole length and another granulate crest on the superior margin which terminates subdistally in a strong upturned tooth. Chelipeds of females of small and simple ‘ocypodid’ type. Ambulatory legs: Similar to T. koelbeli; tym- pana on anterior surface of meri are large and of similar relative size, on posterior surface becom- ing progressively smaller from first to fourth legs. Second pair of legs longest. Hairy edged pouch present between bases of first and second, and second and third walking legs. Colour: Chestnut brown, with lighter chelae becoming white on the fingers. Some specimens have large lateral cream patches extending over the orbital floor, and the hepatic and anterior branchial regions but not onto the frontal and protogastric regions. First male pleopod: As figured. HABITAT Found on intertidal sand at the mouth of the MEMOIRS OF THE QUEENSLAND MUSEUM Gogol River and so would appear to be more tolerant of high salinity than 7. koelbeli. REMARKS This species is only known from the type locality. It is distinguished from both the other species by the remarkable subdistal tooth on the anterior margin of the moveable finger of the chela, and by the distinctive tip of the first male pleopod. Both T. odontodactylus and T. koelbeli differ from 7. ceratophora in having the carpus of the cheliped rather short and not remarkably elongated. Ilyoplax strigicarpus sp. nov. (Figs 4,6D) MATERIAL EXAMINED HoLotyPe: QM W14944, ¢ (7.1 mm), near the Australian Institute of Marine Science, Cape Fer- guson, Townsville, NEQ, Nov. 1980, N. Zucker. PARATYPES: QM W11258, 26 ¢ (6.4, 6.4 mm), data as for Holotype. QM W2996, 2d d (4.9, 5.0 mm), Pioneer River, Mackay, MEQ, B. Campbell, 4.i1.1965. QM W4584, 2d 6 (5.0, 5.3 mm), 22 @ (4.9,5.9 mm), Redbank Ck, Trinity Inlet, Cairns, NEQ, R. Timmins, 11.x1i,1974. QM W4625, 1d (4.2 mm), Bogimbah Ck, Fraser Island, SEQ, Australian Littoral Society, 3.1.1973. QM W4776, 33d (4.6, 5.9, 6.5 mm), Bogimbah Ck, Fraser Island, SEQ, Australian Littoral Society, 1.1.1974. QM W5383, 1d (5.2 mm), Pulgul Ck, Hervey Bay, SEQ, P. Davie, 19.vii.1975. QM W15083, 1d (4.7mm), 3% 2 (3.8, 4.1, 5.3 mm), Point Farewell, East Alligator River, Kakadu National Park, Northern Territory, 1 1.vi.1981, P. Davie. WAM 185- 80, 1d (5.8 mm), Main Channel, Broome, N.W. Australia, between Broome Pearls Ck and Roebuck Hotel Ck, 25.ix.1980, D.S.Jones and R.W. George. WAM 219-80, 1d (6.0 mm) 12 (5.4 mm), Main Channel, Broome, N.W.A., 2.x.1980, D.S. Jones and R.W. George. OTHER MATERIAL: AM Unreg. 16 (5.3 mm), mouth of Nungbalgari Ck, NT, mudflat burrows, 22.viii.1975, D. Grace (Messel). AM Unreg, 1d (4.2 mm), Hutchinson Strait, 10 km W/B, mudfloor, Rhizophora forest, 25.ix.1975, D. Grace/Green. AM Unreg. 3d ¢ (5.5, 5.9, 5.0 mm) Wurugois Ck, NT, 19.viii.75, mudbank, burrow, D. Grace. DESCRIPTION Carapace: Sub-rectangular, regions poorly defined; smooth or microscopically granular, with sparscly scattered small hairs. Front rela- tively broad (approx. 0.29 x distance between external orbital angles), slightly deflexed; lateral NEW AND RARE DOTILLINE CRABS borders converging, angular; a blunt median prominence; distinctly concave dorsally. Gastro- cardiac groove well defined; cardiac region with a low swelling either side of the mid-line; branchial regions with three short, low, horizon- tal crests posterolaterally above the articulation of the last walking leg, each with a row of short hairs. Posterior border slightly concave, and with a very broad rim. Side walls divergent; greatest carapace width at about the first walking leg. Supra-orbital borders, sinuous, sloping back- ward, minutely granular on central convexity otherwise smooth. External orbital angles pointed although not sharply, posteriorly fol- lowed by broad U-shaped notch. Lateral borders slightly divergent, straight or slightly concave, sharp and clearly defined in anterior half then becoming indistinct. Infra-orbital border projects beyond supra- orbital; smooth; slightly sinuous, and is continuous with the lower edge of the external A B ———— eal Inu —EEEE | Bam Ynint 469 orbital angle laterally; the anterolateral edge has a tooth capped by a molar-like pectination and from this tooth a ridge extends almost to the outer lobe of the epistome so forming a horizontal, triangular, concave shelf anterior to the orbit. Eyestalks; c.0.36 X distance between external orbital angles, cornea bulging. External maxillipeds: Metus longer than is- chium(c. 1.3 *). Antero-internal angle of ishium produced along edge of merus. Anterior half of ishium with a line of microscopic granules slant- ing down toward external border and equipped with feathery hairs longest near external border. Maxilliped slightly bulging and completely closing the buccal cavity. Abdomen: Terminal segment rounded trian- gular, slightly shorter than penultimate. Penul- timate with sub-parallel margins. Fifth segment subequal in length to penultimate, charac- teristically constricted near base. Fourth and third segments divergent, fourth longer than 02mm 1 Fic. 4. [lyeplax strigicarpus sp. nov. A,C-G, holotype male; A, male abdomen, B, female abdomen, paratype (OM W4564; 5.9 mm c_b_); C, male first pleapod; D,E, magnifications of apex; F, left chela; G, suborbital border showing chitinous peg on tip of outer orbital tooth: H, suborbital margin of //yoplax orientalis (Zool. Ref. Coll., University of Singapore; Cat. No. 1965,7.19.192-196; ¢ (4.2 mm c.b.), Pandan Forest Reserve Singapore, coll, Sept. 1934), 470 third. Second segment constricted. First with horizontal kee] subparallel to anterior margin. Female abdomen as figured. Chelipeds: Massive, very long (c.3 * distance between external orbital angles); equal; merus trihedral, inner margin granulate, outer margin serrated; posterior border rounded. Posterior face with squamiform markings except for a smooth band down outer edge. Carpus greatly elongated (c, 2X as long as wide); and about as long as length of carapace; widest proximally, tapering distally. This clongation is typical in males over 6 mm carapace breadth however length is vari- able with age being not nearly so long in smaller males; in females it is short and quadrate, The internal proximal angle possessess an obtuse tooth which on its inner edge bears a series of ridges which extend in a band along the proximal lower edge of the carpus. These ridges, in as- sociation with the tooth on the inferior orbital border would appear to be a stridulatory ap- paratus, Above the stridulatory ridges is a row of fine hairs, longest distally, Chela considerably elongated (length c. 2.6% height). Dorsal margin of palm rounded, marked by a line of fine granules which continue down about two-thirds of the inside face. Upper surtace of outside face slightly roughened by a serics of Jow wrinkles, ridges or flattened tubercles, other- wise outer face is smooth, Ventral margin formed by a fine granulate line arising from the lip of the immovable finger and extending c. two-thirds distance toward articulation. An ac- cessory line of granules also arises from the tip of the finger and extends the whole length of the lower outer face. The lower surface of the im- movable finger between these two lines is also finely granulate. A granulate ridge also arises near the tip on the inside of the finger and curves upwards behind the gape for about one-third the length of the palm. This forms one side of an approximately triangular smooth area directly behind the gape. Fingers hollowed but with a brush of hairs on inside edge near tip; poinied; immovable finger slightly down-turned; cutting margin canvex. armed with an even row of teeth that range from large and rounded proximally to tiny points dis- tally. Upper margin of movable finger granulate; cutting margin with similar dentition to fixed finger except for the differentiation of a medial protruberance. Walking legs: Meri c. 4X as long as wide; about as long as the last three segments. There MEMOIRS OF THE QUEENSLAND MUSEUM appear to be faint signs of large oval tympana on both sides of the meri but these are not as distinct us are those of other members of this genus. The second leg slightly the longest. Dense hair is presenton the carpi and propodi of the first and second pairs of walking legs however the extent of coverage is variable. The propedi may be completely covered except for a small bare area on the postero-ventral edge behind the dactylar joint, and the carpi similarly, may be well covered except for the dorsal proximal third and the ventral surface. There does not appear to be any relationship between extent of coverage and size Or SCX. First male pleopod: As figured. REMARKS In overall appearance /, strigicarpus is almost identical with //yoplax orientalis and cannot be distinguished from the type description and il- Justration. The differences are: the presence of a pectinate tooth on the projecting lobe of the outer orbital border; the ‘stridulatory ridges on the proximal inner tooth of the carpus; and the form of the tirst male pleopod. ft falls into the Group l species of Seréne and Lundoer (1974), which includes /. orientalis, 1. tansuiensis, I. gangeticus and /. longicarpus, The species of this group have long ambulatory legs without obvious tym- pana. /. orientalis and I. longicarpus share with f, strivicarpus a similarly formed first male pleopod characterised by an apex distally divided into several short lobes. The gonopods of the other two species are still unknown. HABITAT Common on very soft mud flats at the mouths of estuaries, and sheltered bays. DISTRIBUTION Northern Australia from Hervey Bay in Queensland, north and westwards to Broome in NW Australia. Nlyoplax denlatus Ward, 1933 (Figs 5,6C) Myuplax dentaia Ward, 1933. p, 391, pl. xxii, figs 5,6. Tweedie, 1935, p, 53; 1937, p, 148 (in key), Hyoplay dematus, Serene and Lunduer, 1974, pp. 4,5. MATERIAL EXAMINED HMororyrr AM P10638, 3 (6,0 mm), Port Curtis, Queensland, M, Ward. June 1929, OTHER MATERIAL OM W7428, Lo (5, lL mm). Trinity NEW AND RARE DOTILLINE CRABS Imm 471 0.5mm Fic. 5. Ilyeplax dentatus Ward; A, male abdomen (QM W7428); B, female abdomen (QM W8565); C, male first pleopod (QM W12964); D,E, magnifications of tip. Inlet, Cairns, NEQ, R. Timmins, 14.xi1.1975. QM W8565, 192 (5.2 mm), Murray River, north of Cardwell, NEQ, P. Davie, 19.v.1978. QM W8566, 1 2 (3.8 mm), same data as QM W8565. QM W12964, 184 (3.44.8 mm), 89 9 (3.84.7 mm), Murray River, NEQ, upstream of Tates Landing, exposed bank, P. Davie and J. Short, 19.11.1987. QM W8567, 329 (3.9, 4.5, 6.5 mm), 1d (4.1 mm), Calliope R,, Gladstone, SEQ, P. Saenger, July 1979. QM W8570, 13 (3.4mm), same data as OM W8567. QM W8568, 19 (5.8 mm), Oct. 1975, same data as QM W8567, QM W8569, 1 2 (4.5 mm), 2 juveniles (2.6, 2.8 mm), May 1977. same data as OM W8567. REMARKS Although Ward’s description is short, this species is quite distinctive; the large spine on the inner angle of the carpus of the chela 1s diagnas- tic. Some further description and notes on variability are warranted. Lateral margins of carapace sinuous, bifid anteriorly such that a straight edge continues ta the base of the third ambulatory leg and clearly separates the sub-branchia] region. External or- bital angles rounded, the distance between them less than between epibranchial angles, which are also rounded. The margin between these angles continuous, depressed and shallowly concave. The underside of the external orbital angle forms a sub-acute tooth produced into the orbit lateral- ly. A supplementary row of granules is present on the lower portion of the chela. This is not men- tioned by Ward but is vaguely indicated as a ridge on the right chela in his figure. It arises about midway along the outer surface of the immovable finger and is prominent for only a short distance (about one-quarter length of chela) before fading to an indistinct ridge proximally. This, however, ts variable as the granulate row may extend the full length of the smaller chelae of females and juvenile males. The first two pairs of walking legs may have a thick short fur on the dorsal anterior surfaces of the carpi and propodi. When present this fur extends from the joint but varies in extent of cover. In well preserved specimens the tympana are large and obvious on both the inner and anterior surface of the merus of the cheliped, and al- though not as distinctive on the other legs never- theless occupy most of the width and about MEMOIRS OF THE QUEENSLAND MUSEUM 472 WU UT aur] 2R9S “(PhEPTAA WO) ae adAjojoy “aou “ds snduvaisiais xojdodyy *q (87PL AA WO) 18 ‘prey SMaDjUap xvjdoaq] ‘5 {(98-81Z AVA) Aleut adAyered *aou ‘ds snjAjavpojuopo syaosoddyjawy ‘gy (SROSTAA WO) afew adAqeied “sou ‘ds yaqjaoy syaosoddyjamy ‘yo 9 “O14 ee ee ee ee NEW AND RARE DOTILLINE CRABS two-thirds the length on both sides of the meri of the first to third pairs of legs and about a half the width and length of the last pair. Tlyoplax dentatus is a little difficult to place into one of the groups of Scréne and Lundoer (1974). The presence of the tympana on the legs must place it in their ‘Group TIT’ but the form of the first male pleapod is not particularly like those so far figured for other species in this group, except perhaps for that of /. formosensis. According to Serene and Lundoer (1974), ‘Group III” species have a male first pleopod characterised by a narrow tongue-like apex, and a stem with a longitudinal row of setae on one side, and a subdistal lobe with long setae on the other side. /. dentatus does not have an obvious subdistal lobe. HABITAT Most common on firm mud banks of mid- to upper-estuary, DISTRIBUTION Eastern Australia between Port Curtis and Cairns. ACKNOWLEDGEMENTS I wish to thank the Australian Littoral Society, particularly its Director, Mr Ed Hegerl, for invit- ing me to take part in the Alligator Rivers Field Survey, and also the Australian National Parks and Wildlife Service who allocated the money for this survey to be done. Dr Gary Morgan of the Western Australian Museum kindly loaned me the specimens of Tmethypocoelis adontadac- tylus that he collected in Papua New Guinea while in receipt of a Christiansen Fellowship, and allowed me to describe it. Dr Michael Turkay of the Forschungsinstitut Senckenberg kindly gave me comparative maternal of 7. ceratophora and he is especially thanked. Mrs Yang Chang Man of the Zoological Reference Collection, University of Singapore also kindly loaned me comparative material of /. orientalis. Mr Bruce Campbell of the Queensland Museum is thanked for his valuable comments which have improved the manuscript, Mrs. Peta Woodgate typed the manuscript, and Mr J. Short 473 photographed the specimens, and to both I am grateful. LITERATURE CITED Barnes, R.S.K, 1968, On the evolution of elongate ocular peduncles by the Brachyura. Systematic Zoalogy 17: 182-7, HaGen, H.O0. von. 1970, On the significance of elon- gaied and horned eyes in ocypodid crabs (Decapoda, Brachyura). Forma et Functio 2: 13-57. Kemp, S. 1919, Notes on the Crustacea Decapoda in the Indian Museum. XIT, Scopimerinae. Records of the Indian Museum 16(5): 305-48. KOELBEL, K. 1897. Beschreibung der Krebse. Wiss. Ergebn. der Reise des Grafen Béla Széchenyi in Ostasien. Bd. 2: 709-18, | PL (not seen), MAN, J.G.bE. 1888. Report on the podophthalmous Crustacea of the Mergui Archipelago, collected for the Trustees of the Indian Museum, Calcutta, by Dr John Anderson, F.R.S., Superintendent of the Museum. Journal of the Linnean Society (Zoology) Landan 22; 1-312. MANNING, R.B. AND HOLTHUIS, L.B. 1981, West African brachyuran crabs (Crustacea : Decapoda). Smithsonian Contributions te Zaol- agy 306: 1-379. SERENE, R. AND LUNDOER, S. 1974. Observations on the male pleopod of the species of Ilyeplax Stimpson with a key to the identification of the species. Phuket Marine Biological Cenier (Phuket, Thailand) Research Bulletin 3; 1-10. SHEN, C.J, 1935. On some new and rare crabs of the families Pinnotheridae, Grapsidae and Ocypadidae. Chinese Journal of Zoology 1: 19- 40. Tescu, LJ, 1918. The Decapoda Brachyura of the Siboga- Expedition. I], Goneplacidae and Pin- notheridae. Siboga-Expedition, Monograph 39c. Leiden. pp. 149-295. TWEEDIE. M.W.F. 1935. Notes on the Genus /lyoplax Stimpson (Brachyura, Ocypodidae). Bulletin of the Raffles Museum 10: 53-61, 1937, The crabs of the Family Ocypodidae in the collection of the Raffles Museum. Bulletin of the Raffles Museum U3: 140-70. Warn, M. 1933. New genera and species of marine Decapoda Brachyura. The Australian Zoologist 7: 377-94. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum LITORIA ELECTRICA: A NEW TREEFROG FROM WESTERN QUEENSLAND GLEN INGRAM AND CHRIS CORBEN Ingram, G, and Corben, C. 1990 08 31: Litoria electrica: a new treefrog from western Queensland. Memoirs af the Queensland Museum 28(2); 475-478. Brisbane, ISSN 0079- 8835. Litoria electrica sp. nov. is a member of the L. rubella complex. Morphologically, itis very similar to L, rzbella but readily distinguished by its distinctive colour-pattern and mating call. L. electrica inhabits semi-arid country in northwest and west central Queensland, It is sympatric with L. rubelia. A lectotype for Hyla rubella Gray, 1842, is designated, [Litoria electrica, Litoria rubella, Hylidae, Queensland, Glen Ingram, Queensland Museum, P.O. Box 100, South Brishane, Queensland 4101, Australia; Chris Corben, Queensland Forest Service, 80 Meters Rd, Indooroapilly, Queensland 4068, Australia; 12 October, 1989. In 1975, at Polygammon Creck, west central Queensland (WCQ), one of us (CC) heard a frog's call with which he was unfamiliar. When the frog was located and captured, he was con- vinced it was an undescribed species, The frog was very similar in morphology to Litoria rubel- Ia, which was also common at the locality. In 198], Ingram found the frog again near Cloncur- ry, WCQ. Like Corben, he was impressed with the differences between the calls of the frog and nearby Litoria rubella. In this paper, we describe the frog as a new species. Although it is generally very similat to L, rubella, we consider that the differences in the mating calls indicate the presence of specific- mate recognition systems (sensu Paterson, 1985) A maintaining the genetic isolation of two species in sympatry. In addition, the new species differs consistently, though subtly, in colouration and body form. To check which taxon the name ‘rubella’ designated, we examined two of the three syn- types of Hyla rubella Gray, 1942 (British Museum (Natural History) numbers 1947.2,.24.7 and 1947,2.24.9: we select the latteras lectotype) from Port Essington, Northern Territory. The syntypes are typical frogs of the taxon tradition- ally called Litoria rubella (sensu Copland. 1957), hence the name has been correctly ap- plied. Despite fading, they lack the brown dorsal bars and the brown blotching on the posterior of the thighs of the new taxon. FiG. 1. Litoria electrica sp. nov., holotype, J38963. A. Lateral view of head, B. Dorsal view of head. A76 The following abbreviations are used in the text: SV - snout-vent length; TL - tibial length; HW - width of head at broadest part: ED - diameter of eye opening between anterior and posterior borders; EN - distance between the external nostril and anterior border of eye open- ing; IN ~ distance between the two external nostrils. Specimens with registralion numbers prefixed by ‘J’ and *R’ are housed in the Queensland and Australian Museums respec- tively. Measurements are in millimetres and ratios are expressed as percentages. The sonograms were made on a Kay Sonograph. Litoria electrica sp, nov. (Figs 1-4) MATERIAL EXAMINED HoLoryre, Adult ¢, J38963, 25.1km E of Cloncurry on Julia Creek-Cloneurry road, WCQ (20°43°S, 140°39'E). Collected by G.J. Ingram and G.V. Czechura on 21 January, 1981. PaRATYPeEs: Floraville, NWO (R129391); Floraville Crossing, NWQ (Rj 29393-396, 129407-16); Lawn Hill Station, NWQ (J49227-8): 13.7km Eot Cloncurry on Julia Creek-Cloncurry road, WCQ (J38964); 20km E of Cloncurry on Julia Creek-Cloncurry road, WCO (J38973-4); 25.1km E of Cloncurry on Julia Creek- Cloncurry road, WCQ (J38976-7), Polygamman Creek, on Middleton-Hamilton Hotel road, WCO (J27240-3). DESCRIPTION OF HOLOTYPE SV 38, Tl 11, TL/SV 28.9, HW 9, HW/SYV 23.7, HWY/TL 81.8, ED 2.5, ED/HW 27.8, EN 3.0), IN 2.0, EN/IN 150. Eye small. Snout pointed in lateral view; blunt in dorsal view, Canthus rostralis poorly defined, curving in then out to the nostril, Loreal region concave, Tympanic annulus prominent. Supratympanic fold present, poorly defined. Neck slightly elongate. Subarticular tubercles on hand rounded, one each on first and second fingers and two each on third and fourth fingers; one outer metacarpal tubercle, Subarticular tubercles on feet rounded, one each on first and second toes, two each on third and fifth toes, three on fourth toe: two metatarsal tubercles, outer small, inner elon- gated. Hands and toes poorly webbed. Finger disks large, bigger than toe disks. Nuptial pads present, covering three-quarters of dorsal and lateral surfaces of first finger. Dorsal surface of skin finely granular, ventral surfaces coarscly granular. Vocal sac distended. MEMOIRS OF THE QUEENSLAND MUSEUM Fic, 2, Lituria electrica sp. nov,, holotype, J38963, A. Underside of foot. B. Underside of hand. Ground colour of dorsum yellow-brown, with an indistinct chocolate mark across the upper back; a chocolate forward- pointing chevron across the lower back; also a chocolate blotch above the cloaca, Sides speckled and blotched with chocolate markings that tend to coalesce above and form a distinct dark line running from nostril to eye, beginning again behind eye and continuing lo hind leg. Back of thighs with brown and yellow (white in preservative) blotch- ing. Skin of vocal sac dark grey. Nuptial pads purple-brown. VARIATION IN THE PARATYPES There are 26 paratypes. SV 26-38 (mean 31.1), TL9-12 (mean 10,2), TL/SV 30-35 (mean 33.1), HW 7-10 (mean 8.6), HW/SY 26-31 (mean 27.9), HW/TL 78-91 (mean 84.4), ED 2.4-3.4 (mean 2.77), ED/HW 28-35 (mean 32.1), EN 2,6-3.4 (mean 3.14), IN 2.0-2.9 (mean 2.47), EN/IN 100-149 (mean 127.9). The two bars across the dorsum can vary from bold and weil- defined to indistinct. The brown blotching on the back of the thighs can be faint. DIAGNOSIS L. electrica closely resembles.L. rubella. It can be readily distinguished by the banded dorsal markings and the pattern on the posterior surface of the thigh. In L. rubella, this area is unicalourous or finely dusted with brown, whereas in L. electrica the area is patterned with well-defined dark blotches, In life, L. electrica A NEW TREE FROG 477 hi 0.5 0.4 r om sie ar aa vu wav a SM on wit “ei . ae . i Ms nasi ‘ sal one ser mi winner os = or asian hee gle am o ahaa vn ltt tlm pid ru cy yater . te hegi na 12) SECONDS 0.2 0.3 0.4 0.5 0.6 01 large dark patches at c. 4.5KHz are representations of the call of Ranidella deserticola. B. L. rubella, recorded at 8km E of Toowoomba, SEQ, 22 FIG, 3, Sonograms ot calls. A. L. electrica sp. nov., recorded at Polygammon Creek, on Middleton-Hamilton Hotel road, WCQ, 27 October, 1975; the 478 Fic. 4, Distribution of L, electrica sp. nov. further differs from L. rubella by its longer neck- ed appearance, darker irides, yellower coloura- tion and distinctive mating call. CALL The call of L. electrica has a wavering quality that suggests the sound of a high voltage, long duration, electric arc. This appears to be due to irregular variations in amplitude between pulses of a call. Moreover, some pulses may be left out altogether (Fig. 3A) L. electrica has a higher pitched call compared with that of L. rubella (3.1 vs 2.0-2.7KHz respec- tively. See Fig, 3). As well, the pitch remains MEMOIRS OF THE QUEENSLAND MUSEUM much the same throughout the call while that of L. rubella rises. The pulse repetition rate is higher than in L. rubella (70 vs 55Hz respective- ly) and the duration is longer (585 vs 510 mil- liseconds respectively). Thus, the number of pulses per call is much greater in L. electrica (40+ vs 30). DISTRIBUTION Known only from the semi-arid northwest and central west of Queensland (Fig. 4) in the Gulf drainage: Gregory, Leichardt and Flinders Rivers; and in the Lake Eyre drainage : Hamilton River. REMARKS During breeding, L. electrica is usually found calling from the ground next to, or from low emergent vegetation in, temporary water. ACKNOWLEDGEMENTS We thank Ester Lawless and Clare Bremner for their assistance and Kate Couper for the draw- ings. We also thank Anita Smyth, Gregory Czechura and Gregory Roberts for their help with field work. For specimens, we thank Ross Sadlier, Australian Museum, and Keith Mc- Donald, Queensland National Parks and Wildlife Service. Finally, we thank Jeanette Covacevich and Steve Donnellan. LITERATURE CITED COPLAND, S.J. 1957. Australian tree frogs of the genus Hyla. Proceedings of the Linnean Society afNew South Wales 82: 9-108. GRAY, J.E. 1842. Description of some hitherto un- recorded species of Australian reptiles and batrachians. pp.51-57. In Gray, J.E. (ed.), ‘Zoological Miscellany’. Pt 2, pp.41-80. (Treut- tel, Wurtz and Co.; G.B. Sowerby; W. Wood: London). PATERSON, H.E.H. 1985. The recognition concept of species. pp.21- 29. Jn Vrba, E.S. (ed.), ‘Species and speciation.’ Transvaal Museum Monograph No. 4. ‘YES, | WELL REMEMBER THAT MRS W.C.C. WRIGHT...’ THE ELSIE WRIGHT NEEDLEWORK COLLECTION KERRY KLEINSCHMIDT Kleinschmidt, K. 1990 08 31; ‘Yes, I well remember that Mrs W.C.C. Wright...’ The Elsie Wright Needlework Collection, Memoirs of the Queensland Museum 28(2): 479-497. Brisbane. [SSN 0079-8835. Mrs Elsie Wright (1898-1986) of Nambour, Queensland, exhibited needlework and hand- icrafts from the 1920s to the 1970s at agricultura! shows and Country Women’s Association compelitions throughout Ausiralia. In 1980, she donated a collection of her work to the Queensland Museum, The Elsie Wright Collection consists of 129 pieces of her needlework and handicrafts. As well, ithas valuable documentation such as prize certificates and sashes, her embroidery needles, a needlework encyclopaedia and photocopies of her scrapbook of undated press clippings. Elsie Wright was no ordinary embroiderer, but a significant figure in the history of women’s needlework in Queensland. Throughout her successful exhibiting career, Elsie achieved a national reputation and won some 10,363 prizes and awards. ClElsie Wright, Mrs W.C.C. Wright, needlework, embroidery, handicrafts, Queensland, Kerry Kleinschmidt, 3] Jacaranda Street, Fast Ipswich, Queensland 4305, Australia, 15 February, 1989. The Elsie Wright Collection was donated to the Queensland Museum in 1980 by Mrs Elsie Wright (1898-1986) of Nambour. The collection comprises 129 pieces of needlework and hand- icrafts, These were worked by her during the fifty years in which she actively exhibited at agricul- tural shows and Country Women’s Association competitions throughout Australia from the 1920s ta the 1970s. The collection also has valu- able documentation in the form of prize certifi- cates and sashes, her embroidery needles, a needlework encyclopaedia and photocopies of her sctapbook of undated press clippings (see Appendix). Elsie Wright (Pig. |)was no ordinary embroiderer, She was a significant figure in the history of women’s needlework in Queensland. Throughout her successful exhibiting career, she achieved a national reputation and won some 10,363 prizes and awards. Sadly, fame for Elsie was fleeting. After a brief return to the spotlight in 1976 with a one-woman fund-raising exhibi- tion at Nambour, Maroochydore and Buderim, she returned to ananmynity. The achievements of women in the domestic and decorative arts have been largely overlooked by mainstream art history. However, with the emergence of feminist art history and the revival of interest in traditional handicrafts, women like Elsie Wright are being rediscovered. The Elsie Wright Collection is significant as a document of high quality Australian needlework from the mid-1920s to the mid- 1970s. It provides much information about embroidery styles, techniques and design. For example, the fine hand-made and hand-embroidered garments of the 1920s and 1930s, are of a standard of technical virtuosity that can not be reproduced in modem domestic needlework; the materials needed to produce them are no longer commer- cially available. The Collection is an important source of historical needlework. In it, the trends and changing styles in embroidery, from Vic- torian white work to the cruder, more interpretive crafts of the 1960s and 1970s, can be traced in the production of one woman. In addition to its artistic and technical sig- nificance, the Collection has social sig- nificance as the work of a woman whose life experiences were typical of many Australian women of her time. She had an isolated country childhood and a limited public educa- lion, She experienced the strictures of social expectations for women in the early decades of this-century. She was a war-bride in the First World War. She ‘made do’ during the Depres- sion. She coped with the rapidly changing technology and mores of post World War Two socicty. And, finally, there were illness, ageing and death. In the following, | discuss Elsie Wright’s Collection and her career (1), My major sources of evidence include interviews with her family and friends, show catalogues and newspaper reports. 480 MEMOIRS OF THE QUEENSLAND MUSEUM ELSIE WRIGHT NEEDLEWORK COLLECTION FROM CHILDHOOD TO MOTHERHOOD On 17 February L898, Elsic Sallaway was bom on her parents’ dairy farm at Alstonville in the Richmond River area of New South Wales. She was the fifth child and first daughter of Ada May and Edward Sallaway, Needlework Was an im- portant part of Elsie’s life from the age of four (2). Her first sewing lessons were taken at her mother’s knee, There followed some formal in- struction in needlework during her schooling at the Rous Public School (3), In New South Wales, the Public Instructions Act of 1880 made elementary cducation compulsory for children between the ages of six to fourteen years of age. Needlework probably dominated Elsie's school life, from plain sewing with coloured threads in her first year as an infant to fancy needlework as a teenager, as prescribed in the school curriculum for girls (4). Unlike countless Australian school- girls, Elsie loved needlework. Through it, Elsie expressed an artistic crealivily, which was shared by all of her six siblings, Her brother Arthur took up painting. Her sister Ethe! in later life won numerous awards in the needlework section at the Brisbane, Sydney and Melbourne Royal Shows, Their mother Ada continued sewing and doing fancywork into her seventies (5). Elsie was still a young girl when her family Jeti New South Wales and took up a farming property near Pomona in Queensland, During the suc- ceeding ycars Elsic’s needlework became ‘her preaccupation, her means of self-expression and her recreation’ (13), As a teenager she travelled long distances to country shows and {o “the big one’ in Brisbane, primarily to visit the embroidery exhibits, “‘Lused to study closely any piece (hal look my eye to work out how it was done... then | would pay a penny (o go into the ladies’ toilet and sit in there reproducing the different stitches on the tap of my stockings. This way I could carry the new sliiches home with me and put them into my embroidery’ (7), In this way Elsic taught herself new stitches and techniques, developing her skills by obser- vation and trial and error. Life for Elsie entered » new phase when, in 1916 at the age of cighteen, she married William Fic. 1, Elsie Wright nee Sallaway ()891-1986), c. 1940s, Elyic Wright designed and executed the panched hole decoration on her hal, Photo courtesy Mr B. Wright. 48] Charles Ceci] Wright al Cooroy in Queensland. At twenty-five, William had spent.a number of years in the Sunshine Coast area of Queensland, He was a good-looking young man with an eye for the ladies and a great sense of humour. Soon after the marriage William enlisted for active service in the Great War and Elsie returned to her parents’ property near Pomona. In 1917 she gave birth to their first and only child, Edward Charles. AFTER THE GREAT WAR Although the sequence and details of events are sometimes unclear, and the memories of those limes have a timeless quality of family legend, the story of Elsie’s life in the decades following the waremerges vividly from her pub- lished reminiscences and from the recollections of Edward. Upon William's return from the war, probably late in 1919 or early the following year, the young family moved to Tweed Heads on the northern coast of New South Wales, where they stayed for at least twelve months. They then moved to the hinterland of the Sunshine Coast to their first farm in White's Road, near Landsborough, As a returned soldier, William reccived a government subsidised loan to buy the property. Life was not easy. The homestead was little more (han a slab hut, Amenities were primi- tive; lighting was by kerosene lamp and there was neither running walter, nora hand- pump al the back of the house. Both Elsie and William worked long and hard. During the day William worked three miles away on another man’s farm at Bald Knob Mountain, while Elsie did the housework and her shave of the farm Work, During the season, William cut bananas in the evening and Elsie packed them, He then took them the ten to twelve kilometres into Landsborough, returning home again at 2.30am, At seven o'clock thal morning it was time to leave again for his day-time job. Elsie’s experience in ‘making do’ as a farmer’s daughter staod her in good stead. She coped with the physical labourof farm work and keeping house in primitive conditions, She met the challenge of providing for her family’s comfort and well- being on limited resources in the isolation of their bush farm, Elsie's son, Edward, recalls that she was an cxcellent cook and a good shot with a rifle, She shot parrots and made them into parrot pies. She also made her own bread and grew vegetables. Their houschold furniture was also 482 ae * Binns: LETT UT MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 2. Diamond Jubilee medal, Brisbane Exhibition, 1935. Awarded to Mrs W.C.C. (Elsie) Wright for the best piece of needlework in the show. Scale in mm. Photo courtesy Mrs J. Fawke. improvised. Edward’s first bed was two corn sacks suspended on poles. In the following years, William and Elsie owned several properties in the Landsborough district. After some twelve months of share- farming and dairying at Maleny, they moved to Caboolture. William found work on a farm at Mount Mellum, a soldier settlement near Landsborough. Elsie looked after her home and family, ran a fruit stall on the Caboolture railway station and still found time to do her needlework. In those days Caboolture was an important rail- way junction with big refreshment rooms (8). Elsie held the franchise on the fruit stall for twelve months. THE BEGINNING OF A PRIZE-WINNING CAREER During this period, Elsie’s national show career was launched. Prior to 1927, she had ex- hibited successfully at small local shows like Woombye and Landsborough (9). In 1926, she also exhibited eight articles, in the open classes of the needlework section at the Brisbane Show, but without success (10). In 1927, Mrs Maude Egan, herself an exhibitor and needlework judge at country shows, saw some of Elsie’s work and persuaded her to enter it in the next important show on the calendar - at Toowoomba. Mrs Egan provided Elsie with a prospectus and entry form and she entered twelve pieces of needlework (11). Maude Egan’s confidence in Elsie’s skill was well founded. Elsie did extremely well against stiff competition from some of the best Queensland and interstate needlewomen. She won one first and four second prizes. The first prize was awarded for an example of fibrone embroidery. Hers was the sole entry in that class but this did not diminish her achievement. The judge was not obliged to issue the prize (12). Her white table centre, hand-embroidered pillow sham, article of lazy daisy work and guest towel attracted the second prizes. In some of these classes she was beaten into second place only by veteran prize-winners of national standing, such as Miss Roma Field of Sydney and Mrs A. M. Price. Encouraged by this success, Elsie later the same year entered seven pieces in the Brisbane Show. On her second attempt at cracking ‘the big one’, Elsie was more successful than in the pre- vious year. She won second prizes in the classes for Oriental embroidery and white applique (13). ELSIE WRIGHT NEEDLEWORK COLLECTION MOUNT MELLUM In late 1927 or carly 1928, William and Elsie purchased the fatm at Mount Mellum on which William had worked for the last twelve months, Elsie recalled: ‘Lean still remember seeing the farm for the first time. I saw this small slab hut and asked my husband if that was the packing shed. He told me it was where we would be living, Well it was pretty rough I can tell you. Two rooms and a dirt floor’ (14). They grew bananas and small crops on the farm, Elsic working alongside her husband, Only when the house and farm work were done did Elsic have time to indulge her passion for needle- work, AL night, she worked by kerosene lamp to complete a collection for the show circuit. Wil- liam recounted in a newspaper interview: "She slashed bananas with those hands you know, she hoed and dug and even picked vegetables. Bul even when she was getting up at 2.10am in the morning to milk the cows she'd often be doing her needlework until midnight sometimes. Ud be lying there in bed and the needle would be coming backwards and for- wards in front of my nose’ (15), Hardwork took its toll on Elsic’s hands. Clean hands were important toa needlewoman because washed needlework was disqualified at shows and a soiled piece had no chance of winning a prize (16). Rough hands could also snag fine fabrics, like lawn and silk crepe de Chine, which she used for handkerchiefs, lingerie and baby’s clothes. Elsie’s remedy for ingrained dirt and work-roughened hands was a home-made cold cream of equal parts of glycerine, lemon juice and methylated spirits. Towards the end of 1928, Elsie and William felt secure enough to take on the expense of building a homestead fo replace the slab hut. Elsie designed the new weatherboard bungalow and took the measurements hersell, Her sketch was passed onto an architect who drew up the plans late in October that year. William and Elsie ordered pre-cut materials from Toowoomba at a cost of 450 pounds. Then, when they were heavily committed financially, disaster struck, In the 1928 season, the market for bananas and small crops col- lapsed. Terrible ycars followed as the Depression approached, Along with countless other Australians, Elsie and William were badly af- fected, During the day they laboured on the farm, After dark, Elsie worked on her embroidery which had suddenly achieved a new importance, Edward recalls that it was only Elsie’s prize money that pulled them through the Depression years and spared them fron) selling the farm, One yearshe made SO! puunds, a huge addition to the family’s income (17). THE UPS AND DOWNS OF EXHIBITING (19308 TO 19405) During the 1930s, Elsic’s exhibiting carcer flourished, From 1926-40 she exhibited annually at the Brisbane Show, and al the Toowoomba Show from 1927-41, and al scores of smaller shows from Cairns to Wagga Wagga. She was on the mailing lists of many show societics, When she discovered a show new to hér, she wrote asking to be added to their mailing list. Catalogues and entry forms were constantly ar- riving at their home, William looked after the clerical side of exhibiting. He wrote the entry forms and dispatched the precious boxes of embroidery by train. ff the exhibits were for local shows such as Nambour, he took (her in person, Elsie herself rarely went to the country shows, although she liked to visit the Brisbane Exhibi- tion (18). Elsic’s growing reputation can be traced through award lists and newspaper articles. These give fuscinating ghmpses of the fierce competition thut raged bencath the refined and lady-like face of the needlework section. In 1928, Elsie made her media debut, when her collection of white fancywork at the Brisbane Show was among those complimented in The Queenslander (19). Prom. then on, throughout the 1930s, glowing reports of Elsic’s latest achievements ut the Brisbane Show were rarely missing from the women’s pages of The Queenslander and The Courier-Mail, For ex- ample in 1932 in ‘Needlework at the Show’, it was reported enthusiastically, if slightly inac- curately; ‘Mrs W.C.C, Wright .,. another needlewoman who has had many successes in previous Shows, was again represented by some beautiful work, and five first, seven second, and one third prize tickets bore her name’ (20). Elsie reached the peak of her success at the Brisbane Show in 1935, when she won twenty lirst prizes and three second prizes oul of a total of twenty-five entrics. She also was awarded the fubilee Medal (Fig. 2) for the best pivee of needlework in the show, for a ‘three-piece trous- seau set in pink silk, which Is adorned with x4 exquisite French embroidery and tiny bars of hand veining, and also has i dainty edging’ (21), In 1933 Elsie felt confident enough to enter her work against the best Australian needlewomen in the Melbourne Show. She did very well, gain- ing two first prizes for her buttonholes on linen and on tweed (22). She continued to exhibit at Melbourne throughout the 1930s, winning the prize for (he most successful exhibitor (that is, the exhibitor with the highest aggregate points) in the faney needlework section in [935, 1936 and from 1938 to 1940. Her exhibits were often singled out for praise in The Age's columns ad- ding to her growing national reputation. For ex- ample in 1936, it was reported: ‘Another veteran Who retains her title is Mrs W.C.C, Wright, who has once again scored the highest aggregule of points in the fancy needle work. Mrs Wright submitted 25 entries, and took 8 firsts, 7 seconds, a third and 4 high commen- dation certificates, Her work is amazing in its delicacy and her hand-made trousscau set of white crepe Je Chine with its minute scalloping and hemstitching, its exquisite eyelet and satin stitch embroidery, is as dainty and charming as itisexpertly worked, whyle the child’s frock that won her the prize for plain hand sewing is the perfection of fine stitchery’ (23), In 1939, a Women’s Industries section was reintroduced al the Royal Baster Show, Sydney, following a long suspension because of World War I. Elsie submitted twenty-three entries, and won ton first prizes, four seconds, and the pres- ligious award for champion picce of needlework (24). Elsie’s successes in the big metropolitan shows Were reflected by her results in the smaller Queenslind, New South Wales and Victorian country shows, At Toowoomba, she won the points prize for the highest aggregate of points in the needlework section in 1930, 1931, 1935, 1936, 1938 and 1941. Al Wagga Wagga, New South Wales, she won the cham- pion prize for best piece of hand made under- wear in the shows in 1934, 1936, 1938 and 1939. fn 1934 und 1939, she also won a cham- pion prize for the best piece of a fancywork, and in 1934 and 1939 won the grand champion- ship for ‘chumpion piece of work in Class C, needlework’. Many of her surviving prize cer- tificaltes and sashes in the Queensland Museum's collection were awarded for Cham- pion piece of fancy needlework at rural shows, such as Cairns (1937-1941), Dalby (1939- 1941) and Killarney (1937-1938, 1941), There MEMOIRS OF THE QUEENSLAND MUSEUM were many more awards far which prize certifi- cates have not survived. With this propensity to ‘scoop the pool’, a number of country show societies politely re- quested that she abstain from exhibiting with them in future years. Few other women cared to compete against such a formidable opponent, in the classes that she monopolised. Many com- petitors must have shared the feeling of frustra- lion and acimiration, expressed recently by Mrs Lyn McKay (née Skerman), formerly of Millmerran, in a letter to her daughter. ‘Yes, | well remember that Mrs W.C.C. Wright, She used to win all (he needlework prizes at country shows- as well as city shows. I would have had several first prizes at Millmerran shows, only for her! -She used to “scoop the pool” in every section- absolutely perfect work’ (25). Early in the (940s, World War Two halted Elsie’s winning streak. Many shows, including the large metropolitan ones, were curtailed and eventually suspended during the later years of the war. During those years, Elsie, like many other Australian women, directed her energics to the war effort. When agricultural shows resumed after the war, so did Elsic’s prize winning career. In the Toowoomba Show of 1946, she Won sixteen first and twenty-two second prizes out of forty-four entries. At the Melbourne Show in the same year, she won special prizes for the best picce of fancy needlework and forthe most successful exhibitor in the fancywork section (26), She exhibited annually in the Toowoomba Show until 1950, but restricted her participation in the shows in Brisbane (1946) and Melbourne (1946,1947 and 1950), Elsic was not unchallenged throughout those years. Stiff competition was provided by a num- ber of expert needlewomen, including Miss Roma Field, Mrs C.J, Dwyer, Miss Isla Me- Conachie and Mrs J.A. Anlezark. Of these, the competitor who matched Elsie most closely in skill and techniques was Mrs Dwyer of Toowoomba, who also came to the fore in the late 1920s (27), idward recalls that his mother hated Mrs Dwyer with a passion, although they. had never met. The day after judging at the Brisbane Show, Elsie waited impatiently for her father to ring from Brisbane to read the prize list from the newspaper, When Mrs Dwyer beat Elsie there would he frowns, but when Elsie won over her rival, then... the delight! Elsic would go to great lengths to beal Mrs ELSIE WRIGHT NEEDLEWORK COLLECTION Dwyer. One day, frustrated by her rival’s consis- tent victories jn the coloured afternoon tea or supper cloth class, Elsie broke with her normal tule of not visiting country shows. She went to see whal gave Mrs Dwyer’s work the winning edge. On her return home, she announced with mutch satisfaction lo her family, “I’ve gotit!’, and then set to work embroidering. Needless to say, at the nextshow it was Elsie’s supper cloth which carried off first prize (28). Her show caréer gave her other high moments, such as the time when a Melbourne bank manager sent a telegram seck- ing to buy all her articles at the Royal Melbourne Show for his fiancee. There were sad incidents too, such as the time in the 1930s when a fire at the Toowoomba Show pavilion destroyed an entire collection of thirty-twa pieces of needle- work (29). ‘PREMIER NEEDLEWOMAN'’ In the 1950s, Elsie continued to enjoy her national reputation. She exhibited successfully at metropolitan and country shows in Queensland, New South wales and Victoria: at Brisbane (1955). Townsville (1951-52), Ayr (1950, 1952- $4); Sydney (1953), Wagga Wagga (1951-52): Melbourne (1950-53), Bairnsdale (1951-52); and Mirboo North (1951, 1954). In 1953, she won the Central Agency (Aust.) Ltd's special trophy for best exhibit in the needlework section at the Royal Easter Show in Sydney (30). By 19S], a national women’s magazine is said ta have run a story on the ‘mysterious Queensland necdlewoman’ from the ‘backwoods’, revealing the personality behind the awards (31), Flsie dic not welcome such exposure. She preferred her long periods of seclusion on the farm, After many years of struggle, the early 1950s also brought material comfort to Elsie and Wil- liam. In c.1952. their Mount Mellum home was described as “a comfortable weatherboard cot- tage overlooking the sea and surrounded by the 14U-acre plantation of bananas, oranges, bush nuts, paw paws and other tropical fruits’ (32). In 1953, they sold the farm to Edward, and moved to Perwillowen Road, Nambour, Here for the first time they had the luxury of clectricity . although Wilham held it in deep distrust and Elsic continued to cook on an internal combus- tion stove for the rest of her lite. As their prosperity inercased, Elsie began to suffer the legucy of those early years of heavy work. She wis Constantly nt and out of hospital during the Jatter half of her life. However. she continued to 485 pursue her love of needlework. Moreaver, her contacts at the local hospital proved a valuable source of materials for her, growing interest in handicrafts. The celluloid of X-ray film, when soaked to remove the light-sensitive coating, was pertect for making decorative containers, such as work boxes and waste-paper baskets (33). Elsic joined the local branch of the Country Women's Association, when she moved to Nam- bour in 1953. Soon she participated in both their state and national handicraft competitions. Local CWA members urged Elsie to enter a special piece of work im the Association’s national needlework competition for the Clifton Joseph Rose Bowl. Elsie won it with a waffle-ette stitched article she designed herself. She wenton fo win this prestigious award for five successive years, each time for a different type of embroidery: binca work, huckaback, drawn thread work, counted thread work and Richelieu embroidery (34)- In 1955, at the urging of her grandchildren, Elsie entered the 2000 pound Embroidery Con- test, run by The Australian Women’s Weekly, It attracted over 4000 entries from Australasia, Elsie entered an article in Section Two: ‘Best hand-embroidered tray-cloth, runner,table- centre, duchess set,orsetof4d’oyleys’. She won the third prize of 35 pounds with a blue table centre. Together with the other prize-winning entries it was exhibited at the Parmer department store’s Blaxland Gallery in Sydney in February 1956, and later in stores throughout Australia (35). LATER LIFE Despite illness and deteriorating eye-sight and co-ordination, Elsie continued to design and embroider well into her seventies. Bul instead of the fine needlework of her carlier years, Elsie now exceuted the coarser types of counted thread work, such as Swedish darning and blackwork (36) which were in vogue in the 1960s and carly 1970s, Flsic exhibited at the Brisbane Show from 1963 to 196¥, and from 1974 ta 1975, Her new interest in handicrafts is reflected in entries in the classes fur artificial flowers and in one year, in bark painting (37). Her beautifully hand-tinted bouquets of fabric fuchias and nasturtiums, are now in the possession of her family. Throughout the sixties and seventies, Elsic participated in CWA competitions, winning first prize for a Swedish daming pillow cover in the Queensland NBG CWA handicraft contest in 1971, at the age of seventy-three (38). Sadly, in the mid 1970s, a stroke pul an end to Elsie’s needlework, There- after her creativily was expressed through a wide variety of handicrafts. As part of her recuperation therapy in hospital, she learnt to decorate ceramic plaques and to make poker Nowers, She also made scratched pictures on painted metal foil and decorative pot-plant holders from large tins, In 1976, Elsie was asked to display her work to raise money to aid the Sundale appeal to build a new wing on the Sundale Nursing Home al Nambour. Throughout her life, Elsie had used her talents for charity, donating many of her trophies to aid worthy causes. On the 29th February, accompanied by consid- erable publicity in the local press, a one-woman exhibition of Elsic’s work was held in the CWA Hall at Nambour. A local newspaper reported that: ‘hundreds of people came to sec it. The or- ganisers were amazed at the interest the exhibi- tion created and at the wide cross section of people who came to admire and marvel’ (39). On (his occasion, $300 was raised for the Sun- dale appeal, and the success of the exhibition was such that jt was later shown at Maroochydore and al Buderim, on the Sunshine Coast. Apparently, at this time the Queensland Muscum was offered its choice of Elsie’s work (40). However, it was not until November 1980, that the Museum (ook custody of the collection, just prior to her admission to the James Grimes Nursing Home, Sundale Garden Village at Nam- hour on New Years Day 1981. Sadly, Elsic was never Lo see her work displayed in ils new home, She died on 20th May, 1986 at the age of cighly- eight. The end of a remarkable life of courage and creativity. ELSIE: NEEDLEWOMAN AND 'DESIGN ARTIST’ Elsic Wright's achievement is remarkable. Apart from needlework instruction atschool, she was entirely sell-taught, Unlike more privileged women, who had access to art classes and tech- fical education, Elsie had to rely on her own hatural abilities, on observation and on trial and error, An indication of her lack of basic needle- work lraining was her inability to knit or crachet (41). She made up for this deficiency with inven- live and painstaking neediclace edging, Some time in the 1930s or 1940s, she bought herself an MEMOIRS UF THE QUEENSLAND MUSEUM encyclopacdia of needlework. I! is difficult to trace the influence of this manual on Elsie’s work. In one instance she copied a monogram from the book and used it on a handkerchief (42), Elsic usually designed her own work for ex- hibition. She gained inspiration from many sour- ces, Such as shows and shop displays, memorising what she saw and adapting it in her own work. Occasionally, she used commercially produced designs, She won prizes in a section at {he Toowoomba Show for a coloured alternoon lea or supper cloth traced with ‘Semco’ designs only (43), ‘The output and range of Elsic’s creative ac- tivity throughout her life was enormous and varicd. She had to continually produce new pieces. Prize-winning exhibits oflen could not be re-entered jn the same show in following years. In addition to her embroidery which included calceolaria work, punch work, richelicu work, lace stitch, binca Work, fibrone embroidery, broderic anglaise, and Mount Mellick work, she produced a wide variety of handicrafts. These crafts included, artificial flowers, shell work, burk painting, decorations of painted and threaded pine cones and gum nuts and cut, rolled and painted jam-tin dahlias. She made her own hats (Fig. 1), even out of hessian bags (44). The best of Elsie’s needlework is remarkable forits exquisite fineness, achieved by using very fine No. 12 embroidery needles, made by H. Milward and Sons. For her most delicate work, she would unravel the finest cotton she could buy into three strands, All her sewing supplies, such as needles, cotton and silk thread and fine Irish linen came from MeWhirters in Brisbane, where she bought them by the dozen (45), ACKNOWLEDGEMENTS The following gave generous assistance with my research; Elsie Wright's family, Mr Edward and Mrs Vera Wright and Mrs Joan Fawke; the staff of the Queensland Museum, especially Judith McKay; Mrs Joan Selnes of the Embroiderers’ Guild of Queensland; Mrs Ma Nipperess; the Royal National Agricultural and Industrial Association of Queensland and the Royal Agricultural Society of Queensland, REFERENCES (1) This paper is based on the major project | underlook in partial fulfilment of the require- ments lar the Diploma of Museum Studies, ELSIE WRIGHT NEEDLEWORK COLLECTION 487 University of Sydney, 1988. The project followed work experience at the Queensland Museum in August 1988, when the Elsie Wright Collection was accessioned. (2) Wright, Elsie (comp.) Scrapbook of press clip- pings (unpublished): ‘She has the Art All Sewn Up’ ¢.1952. (3) Robinson, Dr D., Notes from interview with Elsie Wright 5 November 198{) (unpublished: Queensland Museum). (4) Kyle, Noeline ‘Her Natural Destiny. The Educa- tion of Women in New South Wales’ (Sydney c.1986) pp, 44, 46-47. (5) Wright, Elsie, Scrapbook: ‘6000 First Prizes for Needlework’ c.1939, (6) Sunshine Coast Datly 24 January 1981. p.29. (7) ibid, (8) Wright, Elsie, Scrapbook: ‘Delight in Fine Needlework’ c.1976. (9) ibid.: ‘The Prizewinning Work of Elsie Wright’ c.1976 (Woombye); Queenslander 26 August 1926, p.42 (Landsborough). (10) Royal National Agricultural and Industrial As- sociation of Queensland Awards Ledger 1926, pp.470- 477. (11) Wright, Elsie loc. cit, and ‘Delight in Fine Needlework’ c,1976; Sunshine Coast Daily 24 January 1981, p.29; Royal Agricultural Society of Queensland Official Catalogue 1927, pp.179- 185. (12 R.A.S.Q. Prize Schedule 1941, p.18. (13) R.N,A, Awards Ledger 1927, pp.491-498. (14) Sunshine Coast Daily 24 January 1981, p.29. (15) ibid. (16) Wright, Edward and Vera, pers. comm. 1988; R.A,S.Q. Official Catalogue 1941, p.220. (17) Wright, Edward and Vera, pers. comm. 1988; (18) Queenslander 16 August 1928, p.S0. (19) Queenslander 18 August 1932, pp.405-412. (20) Courier-Mail 19 August 1935, p.19, (21) The Age 20 September 1933, p.15. (22) The Age 15 September 1936, p.9 (23) The following data was obtained from prize cerlificales in the Elsie Wright Collection (H.i9781). (24) McKay. Lyn, Letter to Judith McKay 28 August 1988, (25) R.A,S.0. Official Catalogue 1946, pp.219-228 (Toowoomba): The Age 20 September 1946, p.5 (Melbourne). (26) For example: Queenslander 16 August 1928, p.50. (27) Wright, Edward and Vera, pers. comm. 1988 (28) Wright, Elsie, Scrapbook: ‘The Prizewinning Work of Elsie Wright” c.1976. (29) Sydney Morning Herald 25 March 1953, p.12. (30) Sunshine Coast Daily 24 January 1981, p.29. (31) Wright, Elsie, Scrapbook: ‘She has the Art All Sewn Up’ c.1952, (32) For example: A.19777. (33) Sunshine Coast Daily 24 January 1981, p.29. (34) Probably H.19688. Australian Women's Week- 1y 6 July 1955, p.12, and 15 February 1956, p.22: Robinson, Interview 1980. (35) For example: H.19775 and H.19721, respec- lively. (36) R.N.A. Awards Ledger 1967, pp.13-21, (37) Courter Mail 21 May 1971, p.11. (38) Wright, Elsie, Scrapbook: ‘Delight in Fine Needlework’ c.1976. (39) Wright, Elsie, Scrap- book: ‘Wonderful Display of Fine Needlework’ c. 1976. (40) De Dillmont, Therese ‘Encyclopedia of Needlework” (France nd.) p.66, fig, 121. H.19789, cf, H.19706. (41) Forexample: R.A.S.Q. Official Catalogue 1934 p.207. Other exceptions: H.19682/1-3, which is in the “Semco’ coloured doyley tradition. (42) Fawke, Joan pers, comm, 29 September 1988. (43) H.19785. (44) Wright, Elsie (comp.) Scrapbook; ‘The Prizewinning Work of Elsie Wright’ c.1976, (45) Swift, Gay, ‘The Batsford Encyclopedia of Embroidery Techniques’ (London 1984) pp.138-9. (46) ibid., pp.31-33; Thomas, M. “Mary Thomas's Embroidery Book’ (London 1943) pp.24-27. (47) Swift, op. cit. p.186; Thomas, op. cit., pp.247- 248, (48) Women's Domestic Needlework Group ‘The Doyley Show. An Exhibition of Women’s Domestic Fancywork’ (Sydney 1979) p.54; Swill, Joe. cif,, pp.18-19. (49) ibid., pp.75-74. (50) ibid., p.58; Women's Domestic Needlework Group. loc; cit., p.47; Isaacs, J. ‘The Gentle Arts. 200 Years-of Australian Women's Domestic and Decorative Arts’ (Sydney !988) p.106; Thomas, loc, cit., pp.75, 84. (51) ibid. p.84. (52) ibid., pp.75,77-80, (53) ihid., pp-75, 81, (54) Isaacs, foc. cir, p.105. (55) Swill, Joc, cit., p.61; Thomas, loc. cit,, pp.96- 97, (56) Selnes, J, pers, comm, (57) Swift, loc. cit., pp.66- (S58) ibid., pp.69-71; Isaacs, loc. cit, p.106; Wamen’s Domestic Needlework Group, lac. cit, p.54. dss (59) thid., pp.54-55. (60) See H.1)9792; Seines pers. comm (61) Institute of Domestic Arts, ‘Dressmaking. Decorative Stitches and Trimmings’ 2nd edition (England 1967) p.41. (62) Swifl, loc, cil., pp.28-30; Isaacs, loc. cit, p, 137, (63) Swill, foe. cif, pp.37. 193; Courier-Mail 21 May 1971, p.11. APPENDIX CATALOGUE OF THE ELSIE WRIGHT COLLECTION Because Elsie Wright died before the Collection was accessioned in 198K, it was nol possible to assign specific dates and awards to individual items. Ap- proximate dates were assigned on stylistic grounds by Mrs Joan Selnes of The Embroiderers’ Guild of Queensland. She also provided valuable assistance in identifying and describing embroidery techniques and stitches. The following sections in this catalogue are arranged in approximate chronological sequence, based on when the particular needlework techniques were populur in Australia, Within each section, the items are arranged by accession number. The descrip- tive information is taken from the Queensland Museum Accession Register for the History and Tech- nology collections. MOUNT MELLICK WORK (45) 19718 Table Centre, while linen, poppies of fine Mount Mellick embroidery wilh detached buttonhole jace fillings, commercial lace edging, circular shape §2.0cm diameter. ¢. 1920s. (Fig.3) H.19720 Dayley, white linen, flowers oF fine Mount Mellick embroidery, with detached buttonhole lace fillings, scalloped edge, butionhule lace edging with details in sealjops, 36.1 x 53.9¢m, © 1920s, The examples of Mount Mellick wark in the Elsie Wright Collection are fairly typical in colour and design. However, they are much finer in texture than usual, Elsie Wright has used some unusual stilches, such as fly stitch, seeding slitch and detached button- hole lace fillings. The edgings of buttonhole stitch and buttonhole lace details are typical of her work. These pieces probably date to the 1920s. BRODERIE ANGLAISE [46) FINE SCALE H.19709 Handkerchief, yellow lawn, with broderie anglaise, Moral design, with stem stich stems and satin stitch leaves, buttonholed edging. o.1930s. 1.19710 Handkerchief, yellow lawn, with broderie anglaise, floral design with stem Stitch stems and satin stitch leaves, buttanhaled edging, c. 1930s, MEMOIRS OF THE QUEENSLAND MUSEUM 1.19711 Handkerchief, while lawn, with broderie anglaise and running stitches in yellow thread, floral design with stem stilch stems and satin stitch leaves, hemmed edyes. c. 1930s. 1.19712 Doyley, white linen, with broderie anglaise, floral design with satin stitch surface embroidery, buttonholed lace edging, circular shape 14.6cm diameter, ¢,1930s. H.19714 Handkerchiel, white lawn, with broderie anglaise, Moral design with stem stich stems and satin stitch leaves, bultonholed edging with eyelets. ©. 1930s, LARGE SCALE H.L971S Doyley, while linen, with large scale broderie unglaise, clustered vine design with satin stitch surface embroidery, scalloped edge, bullonhale lace edging with details in scallops, oval shape 36.2 x 53-bem. c.1920s, 1.19716 Doyley, while linen, with large scale broderie anglaise, {oral design with satin stitch and seed stitch surface embroidery, buttonhole Jace edving, oval shape 34.3 x 50.0cm, ¢. 1920s. BRONERIE ANGLAISE WITH CUT WORK 1.19713 Handkerchief, white lawn, with broderie anglaise, and cutwork diamonds with buttonholed lace f\llings al each corner, floral design with satin stitch and stem stitch surface embroidery, buttonhole edging. c. 1930s, 1.19768 Woman's Apron, pink organdie broderie anglaise wilh culwork, commercial lace edwing. c, late 1930s. BRODERIE ANGLAISE WITH DRAWN THREAD WORK 19,19767/1,11.19767/2 Baby’s Dress and Matching Bonnet, white lawn, braderie anglaise and drawn thread work. commercial lace edging. H.19767/1 - dress. H.19767/2 - Bonnet, broderie anglaise with drawn thread work, underlaid with cream net, com- mercial lace edging and medallion. c.194(1s. Of the examples of broderie anglaise in the Collec- lion, mostare on & very Tine scale with tiny punched and overcast eyelets. Two dayleys, however, are done in large scale broderje anglaise, Elsie Wright used this form of embroidery on doyleys, lady’s handkerchiefs, bahy’s garments and lingerie. Sometimes i! appears with other forms of embroidery, such as.cut work and drawn thread work Scalloped buttanholed edges are typical of these pieces, as is buttonhale lace edging and details. They date (o ¢c.1920s-1930s. SHADOW WORK (47) 1.19729 Duchess Doyley, while organdie, with 3- colour (yellow, green and pink) embroidery including shadow Work, buttonhole lace edging, oval shape 42.1 x 63.4em. c. 1935-19405, ELSIE WRIGHT NEEDLEWORK COLLECTION H.19730/1, H.19730/2, H.19730/3 Duchess Set, white organdie, with 4-colour (yellow, green, pink and blue) embroidery including shadow work, scalloped, with buttonhole lace edging. H.19730/1 - centre, diamond shape 45.7 x 61.1cm. H.19730/2-3 - side mats, circular shape 17.3cm diameter. c.1930s-1940s. Elsie Wright’s duchess set and doyley in this tech- nique appear to be typical of coloured shadow work in fabric, design and stitchery. The buttonhole lace edging is typical of her work. These pieces probably date to the 1930s-1940s. 489 APPLIQUE (48) H.19727 Child’s Apron, unbleached linen; applique embroidery with buttonhole stitch; toadstools, rabbits and elf design, pocket with buttonhole lace edge; buttonholed edging with needle lace details in scal- lops. c.1953. Won a first prize in the Melbourne Show, 1953. H.19728 Child’s Apron, unbleached linen; applique embroidery with buttonhole stitch; ducks and flowers Fic. 3. White linen table centre in Mount Mellick work, c.1920s. (H.19718) 4901 design, duck pocket, buttonholed edging with needle lace details in scallops. c. 1950s, H.19755 Tea Cosy, cream organdie, with applique flowers (?hollyhocks) and fence; yellow organdie iinet cosy, padded, bultonhole lace edging, c. 1930s. APPLIQUE WITH CUT WORK H,19725 Doyley, cream linen, border of applique flowers and leaves and cutwork loops, satin stitch, stem stitch, french knots and feather stich surface embroidery, oval shape 27.2 % 41.6cm, ¢.1940s, 11.19726 Doyley, cream linen, border of applique flowers and leaves and cutwork loops, oval shaped 32.8 x 46cm, ¢.1940s, Elsie Wright used applique on « wide range of articles from delicate objects such as organdie baby's bonnets and rayon lingerie to sturdy unbleached linen child's aprons, All her applique work is characterised by the use of bultonhole stitch to apply the design to the ground fabric, and by added interior and exterior details in satin stitch, stem stitch and running stitch, Elsie Wright's individual touch can be seen in details, such as the unusual filling of feather stitch within the cut work loops in H,19725, and the interior details of the appliqued flowers in H,19726, where on one flower they are dark, on the next they are light coloured and on the third one they alternate light and dark, petal by petal. The completely buttonholed edges on the aprons are typical of Elsie Wright's edging and are a marathon feal of embroidery. In addition lo surface embroidery. Elsie Wrightalso used applique in combination with other types of embroidery, such .as cut work; cut work and net inser- tion; cut work and drawn thread work; cut work, applique, net inserlion and drawn thread work; and richelieu work and drawn thread work, SIMPLE CUT WORK (49) In Elsie Wright's work, cul work (50) is often found combined with other techniques such as broderie anglaise, applique, or applique and netinsertion, Other combinations are cut work with applique and drawn thread work; cut work with applique, drawn thread work and net insertion; cut work with drawn thread work, cut work with net insertion and drawn thread work; cut work with net/lace insertion; and cut work and embroidery, H.19691 is the only piece in the Collection with a design based on Australian flora or fauna (Fig.4). The design of H.19692, which uses the traditional English daffodil motif, has a strong feel of William Morris and the Arts and Crafts Movement, and it is possible that Elsie adapted i! from an illustration, These pieces are dated to ¢.1930s, H.19691 Doyley, white linen, cul work embroidery, padded satin stitch details, flannel flowers design, 28,1 x 43.40m. 0.19308. (Fig. 4) 1.19692 Doyley, while linen, cut work embroidery, MEMOIRS OF THE QUEENSLAND MUSEUM padded satin stitch details, daffodils design, oval shape 33.6 x 49,6cem, ¢. 1930s. 11..19693 Doyley, while linen, cut work embroidery, ivy design, buttonhole lace edging, circular 16.1em dtameter, c, 1930s, H,19706 Handkerchief, white handkerchief linen, cul work embroidery, ivy design, embroidered monogram, buttonholed edging. ¢.1930s, The design for the monogram was taken From the new revised and enlarged edition of ‘The En- cyclopedia af Needlework’ by Therese De Dillmont. CUT WORK WITH APPLIOUR AND NET INSERTION H.19722 Handkerchief, white lawn, flowers in cut work with applique, net insertion with satin stitch and stem stitch embroidery, scalloped buttonholed edging with buttonhole lace details in the scallops. ¢.1930s. H.19723 Handkerchief, while lawn, flowers and leaves in cul work with applique and net insertion at corners, stem stitch and satin stitch surface embroidery, scalloped bultonholed edging with needle lace details in the scallops. ¢. 1930s. H.19724 Doyley, yellow organdie, poinsettia flowers and leaves in cut work with applique and net insertion, sulin stitch, stem stitch und buttonholed eyelets, oval shape 36.5 x 51.2em. ¢, 1930s. H.19773 Baby's Bonnet, pink organdie, cut work with applique, lace insertion and drawn thread work, bultonhole lace edging. c.1930s. CUT WORK WITH APPLIQUE AND DRAWN THREAD WORK 11.19769 Woman's Apron, yellow ofgandie, cut work and applique, with drawn thread work, vine leaf and grape design, edging bullonhole lace details in scallops, ¢ late 1930s, CUT WORK WITH APPLIQUE, NET INSERTION AND DRAWN. THREAD WORK H.19772 Baby's Bonnet, pink organdie, cut work, net insertion and applique with drawn thread work, bullonhole edging, ribbon work rosettes. «1930s. Cur Work Wittt DRAWN THREAD Work H.19770 Baby's Bonnel, yellow crepe de Chine, cut work, with drawn thread work buttonhole edge with bullonhole lace details, vine leaf design, ribbon work rosette trim, ¢, 1930s, Cur Work WITT! NET INSERTION AND DRAWN THREAD WORK 11.1977) Baby's Bonnet, pink crepe de Chine, cut work with net inserlions with drawn thread work, floral and heart design, ribbon work trim. c,1930s, ELSIE WRIGHT NEEDLEWORK COLLECTION Fic. 4. White linen cut work doyley with a design of flannel flowers, c.1930s. (H.19691) 492 CuT WoRK WITH NET/LACE INSERTION H.19717 Doyley, white linen, border of cut work flowers and leaves with lace insertion, buttonhole lace edging, circular shape 44.5cm diameter. c.1930s. H.19719 Tray Cloth, white linen, centre medallion of cut work deer and leaves with lace insertion, satin stitch berries, commercial lace edging, 45.3 x 60cm. c.1930s. CuT WORK WITH EMBROIDERY H.19694/1, H.19694/2, H.19694/3 Duchess Set, cream linen; cut work and satin and seed stitch embroidery; floral and scroll design, cream and green threads. H.19694/1 - centre, oval shape 27.6 x 41.1cm. H.19694/2-3 - side mats, circular shape approx. 19.6cm diameter. c.1950s. VENETIAN CUT WORK (51) H.19689 Doyley, white linen, Venetian cut work, floral design, buttonholed edge, oval shape 31.1 x 46.4cm. c.1930s. H.19690 Doyley, white linen, Venetian cut work, border of roses design, circular 17.0cm diameter. c.1930s. ROMAN CUT WORK (52) Roman cut work occurs in the Elsie Wright Collec- tion on garments, in combination with some other embroidery technique. For instance, with embroidery; with smocking; and with drawn thread work. ROMAN CUT WORK WITH EMBROIDERY H.19754 Cushion Cover, unbleached linen with lining of yellow satin, Roman cut work with padded satin stitch, deer and foliage design in medallion. c.1950s. ROMAN CUT WORK WITH SMOCKING H.19766 Child’s Dress, yellow crepe de Chine, Roman cut work with smocked front, buttonhole edge with edging of buttonhole lace details. c.1930s. ROMAN CUT WORK WITH DRAWN THREAD WORK H.19679 Guest Towel, white textured linen, com- posite embroidery (including Roman cut work, drawn thread work, padded satin stitch), butterflies and floral design. c.1930s. H.19680/1, H.19680/2, H.19680/3 Duchess Set, white linen, Roman cut work, drawn thread and padded satin stitch, floral design, buttonhole lace edging. H.19680/1 - centre 32.3 x 46.5cm. H.19680/2- 3 - side mats 20.5 x 20.5cm. c.1950s. H.19758 Woman's Nightdress, pink crepe de Chine, MEMOIRS OF THE QUEENSLAND MUSEUM Roman cut work with drawn thread work, buttonhole edge with buttonhole lace details in scallops, tie belt with overcast eylets. c.1930s? H.19765 Child’s Dress, pink crepe de Chine, Roman cut work and drawn thread work, ruching, edging in plain buttonhole with buttonhole lace edging. c.1930s. H.19761/1, H.19761/2, H.19761/3 Woman’s Lingerie (‘Trousseau’) Set, pink crepe de Chine, Roman cut work with drawn thread work. H.19761/1 - nightdress, ruched waist and self tie. H.19761/2 - petticoat. H.19761/3 - scanties. ? Prize-winner at Brisbane R.N.A. Exhibition of 1935. RICHELIEU EMBROIDERY (53) The Collection contains traditional white richelieu, as well as self-colour and coloured richelieu pieces. The traditional white pieces probably date to c.1930s and 1940s, while the coloured work has been dated to c.1940s. In a number of pieces, richelieu embroidery is combined with surface embroidery. These were probably produced c.1930s to 1940s. Again there is extensive use of buttonhole edgings, both simple buttonhole and buttonhole lace. Some- times, as in H.19696, the use of buttonhole lace edging and details is excessive and mars the design qualities of the article. Richelieu embroidery often appears on articles com- bined with other forms of embroidery, for example, applique and drawn thread work, drawn thread work or with net insertion. WHITE OR ECRU RICHELIEU H.19698 Doyley, white linen, richelieu embroidery, floral design, buttonhole lace edging, square shape 14.7cm sides. c.1930s- 1940s. H.19699 Doyley, white linen, richelieu embroidery, star shape S0.S5cm diameter. c.1930s-1940s, H.19700 Doyley, white linen, richelieu embroidery floral and scroll design, buttonhole lace edging, cir- cular shape 17cm diameter. c.1930s-1940s. H.19701 Doyley, white linen, richelieu embroidery floral and scroll design, square shape 14.5 x 13.5cm. c.1930s-1940s. H.19703 Handkerchief, white lawn, richelieu embroidery, floral and ribbon design, scalloped but- tonhole edging with buttonhole lace details in scallops. c.1930s. H.19704 Handkerchief, white lawn, richelieu embroidery, floral (?blossom) design, buttonhole edging. c.1930s. H.19705 Doyley, white linen, border of richelieu embroidery, floral design, buttonhole lace edging, cir- cular shape 16.0cm diameter. c.1930s-1940s, H.19707 Doyley, white linen, richelieu embroidery, art nouveau floral (?lily) design, star shape 48.7cm diameter. c.1930s- 1940s. H.19708 Doyley, white linen, richelieu embroidery, ELSIE WRIGHT NEEDLEWORK COLLECTION floral design, oval shape 29.9 x 43,9cm, ©,1930s- 1940s, COLOURED RICHBLIEU H.19695 Doyley, cream linen: coloured richelieu embroidery with needle lace flowers in brown, orange and yellow threads; bultonhole lace cdying with details, square shape 42.1em sides, c.1940s. H.19696 Dayley, cream linen, coloured richelieu embroidery with needle lace flawers in brown, orange and yellow threads, oval shape 33.8 x 50,3cm. ¢. 1940s, H.19702 Doyley, cream linen, coloured richelicu embroidery, in brown, orange, green and yellow threads, floral design, diamond shape 40 x 42,9cm c.1940s. H.19756 Tea Cosy, cream linen, coloured righelieu embroidery in blues, blue taffeta and net lining; cream net inner cosy, padded, c.late 19305-19405, H.19757 Tea Cosy, cream linen, coloured richeliew embroidery in blues, blue net lining; cream satin cation inner cosy, padded, c.Jate 1930s-1940s, RICHELIEU EMBROIDERY WITH SURFACE EMBROIDERY H.19686 Doyley. cream linen, composite embroidery (including richelieu and padded satin stitch); in brown thread, buttonhole lace edging with many details, square shape 42.0cm sides. c.1950s, H,19687 Doyley, cream linen, richelieu embroidery and some hedebo filling, with coloured padded button- hole stitch flowers, in yellow thread, oval shape 30.7 ¥ 45.6cm. c.1940s. H.19688 Doyley, cream linen; coloured richelieu, padded satin stitch and seed stitch embroidery, in blue threads, 30.5 x 41.1cm, c.1950s, RICHELIEU EMBROIDERY WITH APPLIQUE AND DRAWN THREAD WORK H.19759 Woman's Nightdress, pink rayon, satin applique, tichelieu and drawn thread work, ruched shoulders, buttomhole edge with buttonhole lace details in scallops, tie belt. c.1930s? RiIcHeLi-U EMBROTDERY WITH DRAWN THREAD Work H,1967) Lady's Apron, cream linen, composite embroidery (with coloured richelieu, drawn thread work, pudded satin stitch), floral design in blue threads, buttonhole edging with buttonhole lace details in scallops, two pockets. c. 1940s. H.19672 Tablecloth, cream linen, composite embroidery (with coloured richelicu, drawn thread work, padded satin stitch), floral and ribbon design in blue threads, butionhole lace edging, 84,2 x 85,5cm, ¢,1940s-1950s, H.19673 Tablecloth, cream linen, composite embroidery (with coloured richelieu, drawn thread 493 work, padded sulin stitch), floral design in blue threads, buttonhole lace edging, 85.6 x $7.3cm. c.1940s-1950s. 1.19674 Tablecloth, cream linen, composite embroidery (with coloured richelieu, drawn thread work, padded satin stitch and buttonhole), floral and ribbon design in orange and brown threads, bultonhole lace edging, 86.8 x &4.7em. c.1940s- 1950s. H.19675 Doyley, cream linen, composite embroidery (including richelieu, drawn thread work, padded satin stitch), in ted, brown and black threads, buttonhole lace edging, circular shape. 49cm diameter. €.1950s, (Fig.4) H.19676 Guest Towel, while textured linen, com- posite embroidery (with richelicu, drawn thread work, padded satin suitch), scroll design with monogram. c,1930s, H.19677 Guest Towel, white textured linen, com- posite embroidery (with richeliew, drawn thread work, padded satin stitch), floral design, c. 1930s. H.19678 Guest Towel, pink textured linen, com- posite embroidery (with richelieu, padded satin stitch, drawn thread work), butterflies and floral design, fringed ends, c.1930s, H.19685 Doyley, cream linen, composite embroidery (including richelieu and drawn thread work); in yellow, green and brown threads; butionhole lace edging with many details, 34.1 x 49cm. c.1950s. H.19760 Woman's Bed Jacket, pink crepe de Chine, richelieu embroidery with drawn thread work, ruching on shoulders, edging buttonhole lace details in scal- lops ribbon work rosettes, c. 1930s? H.19762 Child’s Dress, yellow crepe de Chine, richelieu embroidery and drawn thread work edging of buttomhole and buttonhole lace details. c.1930s. H.19763/1, H.19763/2 Child's Dress and Matching Jackel, cream silk, richelieu embroidery wilh drawn thread work, applied waist band, buttonhole edge. H.19763/1 - dress. H.19763/2 - Jacket..c.1930s, RICHELIEU EMBROIDERY WITH NET INSERTION H.19764 Child’s Dress, pink crepe de Chine, tichelieu embroidery with net insertion, buttonhole lace edgings of details. ruching. c.1930s. PLAIN SEWING Elsie Wright was very expert in the art of plain Sewing, H.19753 Series of five samples of buttonholes: two of linen: three of woollen fabric. no date SURFACE EMBROIDERY (54) White WorK H.19666 Handkerchief, while linen, hemmed on al? sides, embroidered monogram. ¢. 1930s. H.19667 Handkerchief, white linen, hemmed on all sides, embroidered monogram. c, 1930s, 494 This monogram appears to be a modified version of the one illustrated in Elsie’s “Encyclopedia of Needle- work’ by Therese De Dillmont which was also used in H.19706. H.19668 Handkerchief, white linen, hemmed on all sides, embroidered monogram. c.1930’s, COLOURED EMBROIDERY H.19681 Child’s Apron, unbleached linen with TEORER AES) t a} FRE CRMOCO RDO CHEE EEE f MEMOIRS OF THE QUEENSLAND MUSEUM green bias binding, transfer design of birds and flowers, coloured embroidery. c.1950s. H.19682/1, H.19682/2, H.19682/3 Duchess Set, white linen, coloured embroidery with birds, possibly worked from a transfer design, buttonhole lace edging. H.19682/1 - centre 30.8 x 48.9em. H.19682/2-3 - side mats, circular shape 17.5cm diameter. c.1950s, The black and red birds in H.19681 and H.19682 are fanciful, no known species, H.19683/1, H.19683/2, H.19683/3 Duchess Set, - * « s Twn CETEEEELEEEIE PTH EERE {UTE tt i) . craven Supeee enn erence eee CELE pene QUIETER "wh sos oe SEES Fic. 5. Cream linen doyley combining richelieu embroidery, drawn thread work and padded satin stitch, c.1950s. (H.19675) ELSIE WRIGHT NEEDLEWORK COLLECTION white linen, composite coloured embroidery, with blue birds and flowers, buttonhole lace edging. H.19683/1 - centre 39.3 x 59.6cm, H.19683/2-3 - side mats, circular shape 21.7cm diameter. c. 1950s. H.19684 Doyley, fawn cotton, coloured embroidery with parrots and flowering creeper on trellis, commer- cially prepared edging with buttonhole lace, worked from a transfer design, 37.9 x 53.7cm. c.1940s. i OO IO ed -v-F- —- a ee ee en ee ee eee Tare ‘fle -<<- eg ee = = o ~ a, 7 7 S oc Ly 495 Elsie Wright usually used her own designs for her exhibition pieces. However, there are a number of pieces in the Collection whose designs are very dif- ferent from her usual style. Their motifs, in fact, are strongly suggestive of commercial transfer designs. On some pieces, such as H.19681, the printed design can still be seen under the embroidery threads, while other pieces (H.19682 and H.19684) have a commer- 29 SRY -, CaS se cho Ade 30) 4 r ' ' e t ' ‘ ‘ ' ‘ ‘ t ' ‘ Fic. 6. Cream linen mat in Dorset feather stitchery in pink, orange and browns, c.1960s. (H.19697) 4up cially made spoke stitch edging, These pieces were probably produced inc, 1940s and 1950s. HUCKABACK DARNING (55) The Collection, contains a number of guest lowels decorated by Elsie Wright in this technique. They have been dated to ¢. 1940s and 1950s. Generally, Elsie has used fairly typical geometric designs, with some (H,19740 und H.19744) displaying particularly care- ful shading. H.19741, however, has a most unusual design (56) H.19740 Guest Towel, white huckaback cotton, huckaback darning in red, yellow and orange threads, hemmed sides, fringed ends. c.1940s-1950s. H.19741 Guesi Towel, white huckaback cotton wilh huckaback damning in yellow and orange threads, hemmed sides and [ringed ends. c. 19405-19505, 4.19742 Guest Towel, white huckaback cotton with huckuback durning in yellow and orange threads, hemmed sides and fringed ends. c, 19405-1950), 11,19743 Guest Towel, white huckuhack colton with huckaback darning in yellow and arange threads, hemmed sides and fringed ends, ¢, 1940s-1950s, 31.19744 Guest Towel, while huckaback colton with huckaback darning in yellow and orange threads, hemmed sides and fringed ends, c, 19405-1950s, NORSET FEATHER STITCHERY (57) The one example (Fig, 6) of Dorset feather stitchery in the Collection seems to be typical of this technique. In addition to feather stitch, it includes satin stitch, plain and whipped buttonhole stitch and wheat ear stitch. It was probably produced in c¢, 1960s, §1.19697 Mat, cream linen, with Dorset feather stitchery, in orange, browns and pink threads, border of ric rac braid attached with buttonhale stitch; button- hole lace edging, 39.3 x 45.7em. c. 1960s. (Pig.6) DRAWN THREAD WORK (58) Of counted thread work (59), There are two examples of pure drawn threadl work in the Collection. The appearance of a pattern in her exercise hook (H.19792), indicates that Elsie counted and sketched at least the more complex drawn thread work designs, The coarse- ness of the work and lack of precision in technique indicates a date of ¢.1960s-1970s for this work. A bonnet of drawn thread work with some broderie anglaise (H.19774) is a much earlier and finer piece. Drawn thread work often appears in Elsie Wright's embroidery as a subsidiary decoration, for example with broderie anglaise on children's garments; cul work and applique; cul work, applique and nel inser- tion; cut work; cut work and net insertion; richelieu embroidery and applique: richelieu embroidery; or ramian cut work, MEMOIRS OF THE QUEENSLAND MUSEUM H.19669 Mat, cream even weave cotton, drawn thread work, in ecru thread, hemmed un all sides, 46.5 x 48em. c. 19605-1970. U.19670 Mat, cream even weave cotton, drawn thread work, in eeru thread, hemmed on all sides, 31.8 x &5.5en. c. 19608-1970. DRAWN THREAD WORK WITH BRODERIE ANGLAISE, H.19774 Baby's Bonnet, pink organdie, drawn thread work and broderie anglaise, commercial braid edging, ribbon work. ¢. 1930s. CONTEMPORARY COUNTED THREAD WORK The Collection contains a number of pieces of counted thread work in 4 style that [have not been able lo identify. This style of contemporary embroidery has a peasant feel (0 its designs, They were designed by Elsie and seem to utilise several motifs, which reoceur with varlations, Elsie appears to have worked up these motifs, first sketching them on graph paper and then reproducing them on aida cloth using a basic repertory af stitches, These stitches include herringbone stitch, satin stitch, fly stitch, delached chain stitch, back stitch, chevron stitch, leather stitch, sheaf stitch and stem stitch (40). In some cases (H.19731 to H.19733), counted thread work has been combined with free stitchery, Occasionally, as on H.19732, 4 buttonhale lace edging has been added thal does not marry happily with the coarse fabric and stitchery or the geometric design, The technique is sometimes lacking in precision, which, together with the modern style of the fabric and embroidery, would seem to indicate a date later in her life, ¢, 1960s, 0.19731 Mat, blue aida cloth, contemporary counted ihread embraidery in red thread, hemmed on all sides,, buttunhole lace edging, 39.5 x 37cm. c.196(s. H.19732 Mat, blue aida cloth, contemporary counted thread embroidery in red thread, hemmed on all sides, buttonhole lace edging, 43.4 xX 38.7em, cl 96t)s, 1.19733 Mul, blue aida cloth, contemporary counted thread embroidery in red thread, hemmed on all sides, buttonhole lace edging, 39.4 x 38.8cm, ¢. 19605, H.19734 Mat, uchre aida cloth, counted thread embroidery, in brown, green and red threads, fringed on all sides, 36 x 35.1em. c.1960s. H.19735 Mat, ochre aida cloth, counted thread embroidery, in brown. green and orange threads, hemmed on all sides, 69 x 34.4cm_c.1960s, H.19736 Mat, ochre aida cloth, counted thread embroidery, in blue and red threads, hemmed on all sides, buttonhole lace edging, 44.8 x 34.8cm. c. 1960s. HE.19737 Mat, lemon aida cloth, counted thread embroidery, in red and blue threads, hemmed on all sides, 45 x 35.2em. c. 1960s. H.19738 Mat, lemon aida cloth, counted thread embroidery, in brawn and purple threads, hemmed on all sides, 91.8 x 37,2cm, ¢, 1960s. ELSIE WRIGHT NEEDLEWORK COLLECTION 1.19739 Tablecloth, yellow aida cloth, counted thread embroidery, in brown, green and orange threads, hemmed on all sides, 88.7 x 87.Scem. c.1960s. CROSS STITCH (61) H.19745 Mat, ecru even weave cotton cloth, cross stitch embroidery in brown, orange and green threads; hem-stilched and [ringed borders, 41.2 x 39.7em. ©. 1950s-1960s. H.19746 Mat, ecru even weave cotton cloth, cross stitch embroidery in brown, orange and green threads, hem stitched and fringed borders, 43.4 x 41.5em. e, 1950s-1960s. 1.19747 Tablecloth, black and white gingham, cross stitched in white, border of white ric rac braid and buttonhole lace, 87 x 85,3cm. c.1960s-1970s, 1.19748 Woman's Apron, black and white gin- gham, cross stitvhed in white, border of white ric rac braid. c. 1960s-1970s. H.19749 Woman's Apron, black and while gin- gham, cross stitched in red and white, border of white ric rac braid..c, 196()s- 1970s. H.19750 Tablectoth, red and white gingham, cross stitched in black, border of red ric rac braid, 87.8 x 85.3cem. ¢.1960s- 1971s. H.L9751 Tablecloth, black and while gingham, cross stitched in yellow, brown and orange, border of yellow ric rac braid, 87 x 83.5em.c.1960s-1970s. H.19752 Tablecjoth, green and white gingham, cross stitched in black, border of black rie rac braid, 89,2 x 83,2em. c. 1960s-1970s. Cross stitched pieces in the Elsie Wright Callection range from quile Sine stitchery on linen lo coarse work on gingham tablecloths and aprons. The finer cross Stilched pieces (H.19745, H.19746) have a multi- colored floral design, and a drawn thread work border. Elsie designed these, skelching the design first an graph paper. The coarse work on gingham uses geometric designs and applied ric rac braid. They are dated to c.1950s to 1960s, and ¢.1960s to 1970s, respectively. BLACK WORK (62) H.19721 Mal, cream even weave calton, with black- work embroidery, fringed on all sides, 45.8 x 40.5em. 1960s-70s. Only one piece of blackwork is found in the Collec- tion. In il, blackwork is combined with a border of drawn thread work. Elsie designed this piece and the pauiern, counted out and sketched on graph paper, survives. Lack of precision in the stitching suggesis thal this piece was execuled by Elsie Wright laler in life. It has therefore heen dated to c, 1960s to 1970s. CANVAS WORK: SWEDISH DARNING (63) H.19775 Cushion Cover, top of yellow canvas stitched in Swedish darning in red raffia, back of red satin, cearly 1970s. 497 A reference and photograph in The Courier-Mail (64) identifies the cushion cover (H.19775) with this technique with a fair degree of certainty, Another cushion cover in the same technique remains with the family. SMOCKING (65) There are two examples of this technique. One is applied to a child's dress (H.19766) with Roman cui work, which dates Io c. 1930s; the other is a gingham cushion cover (H.19776) smocked in snow-flake smocking, which was popular in ¢, 1960s, H,19776 Cushion cover, brown and white gingham, top is snow-flake smocked. c.1 960s. HANDICRAFTS The Elsie Wright Collection contains a wide variety of handicrafts made by Elsie from the 1950s to the 1970s. H.19777 Waste Paper Basket, made of cardboard: celluloid and wallpaper sides and base; joined by green blanket stitch, sides decorated with @ collage of floral wrapping paper, plastic ferns and ribbon bows; 18.0cm high. 1950s H.19778 Posy of flowers made with painted shells, buttons and wire; plastic ferns, undated, H,19779 Ladys Bag, nylon embroidery on plastic, lined with taffeta, metal clasp. undated. H.19785 Lady's Hat, made from a hessian sugar bag, machine stitched brim, band of petals of the same hessian, undated. H.19786 Posy of French flowers, hand-made, syn- thetic fabric, rose, bud and leaves, one white, one pink, carnation flower, bud and leaves, shades of pink. c, 1960s, H.19787 Lady's Brooch, comprising a pink imita- Lion pearl bution mounted in a commercial imitation gold setting, decorated with sequins, imitation pearls and beads threaded onto sewing pins. undated. 1.19788 Lady's Brooch, comprising a cream imita- tion pearl button mounted as above. undated. H.19791 Work Box, made of cardboard, celluloid and paper sides and base, jormed by green blanket stitch with buttonhole lace edging. c. 1950s. TOOLS OF THE TRADE 11.19782 Sewing needles. made by H. Milward and Sons of Redditch, England, steel, sharps no.12, 2 packets. H.19789 ‘Encyclopedia of Needlework’ by Therese De Diilmont, new edition, Mulhouse, France, not dated. H.9792 Exercise book, ‘The Shell Geometry Book’, containing pencil drawn patterns for counted thread work, undated. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum A REVIEW OF THE POLYRHACHIS VIEHMEYERI SPECIES-GROUP (HYMENOPTERA: FORMICIDAE: FORMICINAE) RUDOLF J. KOHOUT Kohout. RJ. 1990 08 31: A review of the Palyrhachiy viehmeyert species-group (Hymenoptera: Formicidae: Formicinae). Memoirs of the Queensland Museum 28(2): 499-508. Brisbane. ISSN (079-8835, Nine species of the Polyrhachis viehmeyeri species-group sre recognised, meluding P. davydovi Karawajew, P. hirta Viehmeyer and P. vichmeyeri Emery and six new species: P. bamaga, P. eremita, P. greensladei, P. laweryi, P. rustica and P. stigmatifera, Lectoly pes are desipnated for P. davydovi and P, hirta A key to the species-group is provided. LFormicidae, Polyrhachis, viehmeyerl species-yroup, systematics, distribution. Rudol/ I, Roheut, Queensland Museum, PO Box 300, South Brisbane, Queensland 4101, Australia; 20 June, 1989, The Polyrhachis viehmeyeri species-zroup was delimited by Emery (1925) within the sub- genus Myrmhopla Foret for two of ils more un- usual constituents: P. firfa Viehmeyer and P. wehmeyert Emery. A third species, P. davydovi, was added by Karawajew in 1927 and, since (hen, the composition of the specics-group has remained unchanged. Subsequent systematic work has been hampered by the scarcity of re- search material because specimens of this rather distinct group are seldom collected and only a few have been taken in addition to the types. My study hus been possible because of recent collections, particularly those gathered by Dr P.J.M. Greenslade in the Solomon Islands, and Rev. B.B. Lowery in Australia and Papua New Guinea. Their collections ate lodged in the Australian National Insect Collection and provided three of the new species, namely P. greensladei, P. loweryi and P. rustica, The material collected by Rev. Lowery also included the only known specimens of P. viehmeyeri apart from the holotype. My collecting in northern Australia and Papua New Guinea has produced two more new species, P. bamaga and P. eremita, together with additional specimens of P. greensladei and P. hirta. Finally, two specimens located by Burry Bolton in the collections of the British Muscum (Natural History) provided the unique holotype of P. stigmatifera sp.noy. and an additional paratype of P. rustica sp.nov. The P, viehmeyeri species-group ranges from Moluceas and Papua New Guinea to Guadal- canal in the Solomon Islands (09/159) and north- ern Australia, south to southern Queensland (lal. 26"), In the following, Australian and some Melanesian records are given using 1-degree coordinates as initiated by Taylor (1987). The illustrations were prepared using a Zeiss (Oberkochen) SR Stereomicroscope with camera lucida. All figures depict the primary types. The measurements (in mm) and indices follow those of Kohout (1988): HL - maximum head length, measured from the anterior clypeal border to the oceipitul margin; HW - width of the head, measured immediately in front of the eyes: Cl - cephalic index (HW x 100/HL); SL - length of the antennal scape, excluding the condyla; SI - scape index (SL x 100/HW), P'W - width of the pronotal dorsum, measured at the bases of the pronotal spines; and MTL - maximum measurable length of the tibia of the hind leg. Acronyms for museums and depositories are: ANIC - Australian National Insect Collection, CSIRO Division of Entomology, Canberra; BMNH - British Museum (Natural History), London, U.K.; BPBM - Bernice P, Bishop Museum, Honolulu, Hawaii, U.S.A.; IZAS - In- stitutc of Zoology, Academy of Sciences, Kiev, U.S.S.R.; MCSN - Musco Civico di Storia Naturale ‘Giacomo Doria’, Genoa, Italy; MCZC - Museum of Comparative Zoology, Harvard University, Cambridge, Mass., U.S.A.; MHNG- Muséum d'Histoire Naturelle, Geneva, Switzer- land; MNHU - Museum fiir Naturkunde, Hum- boldt-Universitat, Berlin, D.D.R.; NMNH - National Museum of Natural History, Smith- sonian Institution, Washington, D.C., U.S.A.; QMBA - Queensland Museum, South Brisbane: RJK - Rudolf J. Kohout, Brisbane (author's ac- cessions and private collection dita) S00 CHARACTERS OF THE P. VIEHMEYERI SPECIES-GROUP The P. viehmeyeri species-group can be char- acterised within the genus Polyrhachis by the following combination of characters; 1. All dorsal surfaces of the body with bristle- like hairs, which are distinctly shorter than the maximum diameter of the eye. 2. Dorsa of head, mesosoma and petiole with characteristic vermiculate-rugose sculpturation. 3.Mesosomal dorsum bluntly marginale on each side along its entire length. 4, Pronotum and propodeum each armed with a pair of spines. $.Pronotal spines flattened dorsally, with anterior and lateral margins acute; their length, direction and degrce of elevation usually highly variable within species. 6.Pronotal and propodeal dorsa almost flat, mesonotal dorsum transversely convex with rounded lateral margins. 7.Node of petiole with more or less flat dor- sum, bearing a pair of widely separated, diverg- ing spines, and without intercalary spines or teeth. 8. Eyes strongly convex, almost hemispherical, with numerous short, erect hairs. 9.Mandibles very finely longitudinally striate. 10.Clypeus with anterior margin medially truncated; posterior margin usually deeply im- pressed, 11.Antennal carinae rather flat, widely separated. KEY TO MEMBERS OF THE P. VIEHMEYERI SPECIES-GROUP BASED ON WORKER CASTE 1. Australian species with base of first gastral tergite finely micro-reticulate and more or less shiny; mandibles. with 4 distinct teeth ... Indonesian, Melanesian or Australian species with base of first gastral tergite more or less closely, transversely striate and opaque; man- dibles with 5 teeth, of which the basal tooth is Often vestigial sees epeeceeeenn 4 2, Larger species (HL>1. a" median ocellus well developed, diSHINCE woe 3 Smaller species (HL<1,78); median ocellus rather small, indistinct... P. rustica 3, Body bicoloured, medium reddish-brown withmost of the head, pronotal collar and antero-median MEMOIRS OF THE QUEENSLAND MUSEUM patch on mesosomal dorsum light yellowish- brown: antennal scapes shorter (SI<144) auvesvarvvesyersepisvarsnecqerassen seoquevesveasenoney P, eremita Body more or Jess uniformly coloured, dark reddish-brown wilh only the mandibles, spines and subpetiolar pracess slightly lighter; anten- nal scapes longer (SI>146) .......... P, loweryi 4. Inner surfaces of hind femora without erect hairs, or with only a few hairs at their proximal and distal ends; body dark reddish-brown .....,.:cs 5 All surfaces of hind femora with numerous erect hairs; Dow. Tught to medium reddish- brown .. Lak halesettetsetsbedsee 5. Metathoracie spiracles prominent, situated on laterally projecting tubercles .. P. stigmatifera Meltathoracic spiracles not prominent, more or Tes, Teal, sls stssctetasdewsattasevactoasvettoaccsarsmatcarsvassats Gh 6, Sides of head between eyes and mandibular bases with mumerous projecting short hairs lune PF, greensladet Sides of head between eyes and mandibular bases without i prcpcting a short hairs (Fig, 4) 4 . P, bamaga 7. Smallerspecies (AL 1.93); antennal scapes relative- ly short (ST 130).u..cccceereeeee P. davydovi Larger species CHL >2. eE 4 antennal mapes longer (SI >135) .., ire seieioes 8. Pronotal, propodeal and petiolar spines well eleyaled (Fig. 7); truncated median portion of anterior clypeal margin distinctly dentate laterally (Fig. 3).ccsctsssereeeee PL vleAneyeri Pronotal, propodeal and petiolar spines scarce- ly elevated (Fig, 5); truncated median portion of anterior clypeal margin bluntly angulate laterally (Fig. 1) c.cjupcsence ee. PL Aleta Polyrhachis bamaga sp.nov. (Fig, 4) MATERIAL EXAMINED HOLOTYPE: AUSTRALIA: Queensland, Cape York Peninsula, Bamaga, 10°53'S,142°23’E, 18 March 1987, RIK acc. 87.4 (worker). PARATYPES: data as for holotype (10 workers). Type deposition: Holotype in OMBA (type no. T 11123); 2 paratypes each in ANIC and RJK; | paralype each in BMNH, BPBM, MHNG, MCSN, MCZC and NMNH. POL YRHACHIS VIEHMEYERI SPECIES-GROUP WORKER Dimensions (holotype cited first): TL 7.41, 7.11-8.01; HL.1.90, 1,.84-1.96; HW 1,53, 1.50-1.42; CI 81,79-83; SL 2.12, 2.03-2.25; ST 139, 134-142; PW 0.75, 0.72- 0.81; MTL 2.74, 2.62-2.90 (11 measured). Clypeus in profile distinctly sinuate, with the base deeply impressed; median carina feebly marked anteriorly, indistinct posteriorly; trun- cated median portion of anterior margin dentate laterally. Oce if lacking. Pronotum with anterior- ly converging lateral margins, bearing a pair of antero-lateral, more or less horizontal spines. Propodcal spines well clevated, moderately divergent, slightly sinuate in lateral view. Dor- sum of petiole Mat, anterior and posterior mar- gins well defined, spines well elevated, widely divergent. Clypeus with fine, V-shaped tugae, extending to the frontal area of head: sculptural intensity increasing posteriorly, so that dorsa of head and mesosomu are coatscly vetmiculate-rugose; propodeal declivily and dorsum of petiole transversely rugose, Sculptural intensity dis- tinctly less coarse laterally, with sides of mesosoma and petiole only weakly rugose. Propodeal spines smooth and shiny, with only microscopic reticulation. Basal half of first gastral tergite very Cinely and regularly, transver- scly striate, opaque, Brown. bristle-like hairs longest and most dense on gaster, only slightly shorter and more dilute on dorsa of head and mesosoma, Hairs almast completely absent from sides of head between cyes and mandibular bases. dorsal sur~ faces of front and middle femora and tibiae, and dorsal and inner surfaces of hind femora. Golden, relatively short, appressed pubescence very Spotadic on darsum Uibody except gaster, where it is rather abundant. Very dark reddish-brown; sides of mesosoma and petiole, spines and appendages a shade lighter. Sexuals and immature stages unknown. REMARKS The eleven specimens of the PF. bamaga type- series were collected on the edge of riverine lowland rainforest at Bamaga (Grid cell 10/142), near the tip of Cape York Peninsula. This species shares characteristic features with P. vieluneyeri- group species from the Solomons, New Guinea and Indonesia, including 5-dentate mandibles and an opaque first gastral tergite. In contrast, the more southern Australian members of ihe group (P. erentita, P. laweryi and P. rustica) possess be 4-dentale mandibles and have more or less shiny gasters. At the lime of collection the site was inundated following torrential rain and the ants were run- hing in disarray over low. vegetation and flood debris in company with Polyrhachis pexilla Fr. Smith, a species of similar appearance and with almost identical vermiculate-rugose sculptura- tion. Despite repeated visits over following days no other P. bamaga specimens could be found. Polyrhachis dayydovi Karawajew, 1927 (Figs 2, 6) Palyrhachiy (Myrmbopla) davydovi Karawajew, 1927:24, Syntype workers. Type locality: IN- DONESIA, Aru [s., Wammar (= Wamarl., Kepulavan Aru) (05/134), 19 iff 1913, Karawajew, Nr.2746, IZAS (1 synlype examined), LECTOTYPE DESIGNATION | have examined one of two synlypes compris- ing the P. davydovi type series, kindly loaned by Dr A.G. Radchenko of the Zoological Institute, Ukrainian Academy of Sciences, Kiev. The Specimen is in good condition and, besides the orange lay reading davidovi (sic), which is glued directly to the card triangle with the specimen, it bears three additional labels of which two are apparently in Karawajew’'s handwriting and read as follows: “Wammar, Aru. 2746. Karavaiev’; ‘Polyrhachis (Myrmhopla) davydovi Karav. Typus’. The third label (on a ted tag) reads: ‘Holotypus Polyrhachis (Myrmkopla) davydovt Katawajew’. Despile the specimen being labelled ‘Holotype’, Karawajew's original description clearly indicates that both specimens are of equal! value and thus syntypes. | designate ihe specimen | have examined as lectotype and, consequently, the second specimen is a paralec- totype. Dimensions of lectotype: TL 7.91; HL 1.93; HW 1.56, C181; SL 2.03; $1130; PW 0.81; MTL 2.90. WORKER Clypeus in profile almost straight, with moderately impressed basal margin; median curina Vague; truncated median portion of anterior margin obtuse laterally. Ocelli lacking (a shallow depression in the cephalic sculptura- tion indicates the relative position of the median ocellus). Pronotum with anteriorly converging lateral margins, bearing a pair of antero-laterally projecting, well elevated spines. Propodeal bal spines well clevated, subparallel, almost straight in lateral view, Dorsum of petiole convex, anterior and posterior margins ill defined, spines moderately elevated, widely divergent. Fine, mostly V-shaped rugac, on clypeus and fromtal area of head. Sculptural intensity increas- ing dorsally and posteriorly, with dorsum of head and mesosoma vermiculate-tugose; the pattern tends to be less coarse laterally, with sides of mesosoma only weakly rugose. Dorsum of petiole with rather fine, somewhat transverse, but mostly irregular rugulations. Base of first gastral tergite very finely, mostly transversely striate, opaque. Yellow and reddish-brown, bristle-like hairs Jense on head and gaster, but rather sporadic on dorsa of mesosoma and petiole, Mosily silvery, appressed pubescence fairly sparse everywhere, except the gustral dorsum, where it is longer and golden-yellow, with a distinct reddish-tint. Medium reddish-brown; dorsum of mesosoma and sides of mesonotum and propodeum a shade darker. Mundibles, clypeus, antennal carinac and posterior margins of gastral tergites bordered dark brown, Sexuals and immature stages unknown. REMARKS P. davydavi closely resembles P. Aira and P. viehmeyeri, and is undoubtedly closely related to both. Besides the characters given in the key, it differs from viehmeyeri in having the truncated portion of the anterior clypeal margin bluntly terminated laterally, and the propodeal and petiolar spines distinctly shorter. From Aira it differs in having the propodeal spines distinctly shorter and well elevated, and the petiolar spines more Widely divergent. The anterior margin of the petiolar dorsum in davydovi is blunt and indistinct (Fig, @), while it is clearly defined or even dorsally produced in the other two species (Figs 5, 7), Also, the bristle-like hairs in P. davydovi are much shorter and more dilute than in P, kirta and P, viehmeyert. Polyrhachis eremita sp.nov. MATFRIAL EXAMINED HOLOTYPE, AUSTRALIA, Queensland, ¢. 4-10) km Nol Marlhureugh, 22°45'S,149°54 Eo April 198t, RIK nee, 81.5) (worker). PARATYPES data as for holotype (® workers). ‘lype deposition: Holotype in QMBA (type no, T 11124); 2 paratypes each in ANIC and RIK, | paratype each in BMNU, MUNG, MCSN and MOZO, MEMOIRS OF THE QUEENSLAND MUSEUM WORKER Dimensions (holotype cited first): TL 7.66, 7.66-8.72,, HL. 1.92, 1.87-2.03; HW 1.61, 1.56-1.70; C184, 82-85; SL 2.28, 2.21-2.43; ST 142, 141-144; PW 0.87, 0.81- 0.91; MTL. 3.00, 2-97-3,22 (9 measured), Clypeus in profile almost straight, with posterior margin moderately impressed; median carina rather smooth and shiny for most of its length; truncated median portion of anterior mar- gin obtuse laterally. Median ocellus distinct; lateral ocelli Jacking (their relative location marked by shallow depressions in cephalic sculpturation), Pronotal dorsum narrowed anteriorly, bearing a pair of slender, well elevated, short to medium long, often asym- metrical spines (see below under remarks on P. rustica). Propodeum flat, bearing a pair of straight, well elevated, moderately divergent spines. Petiole with posteriorly sloping dorsum, whieh is more or less concave between diver- gent, well elevated spines, Clypeus and front of head with fine, more or less longitudinal rugaec, sculptural intensity in- creasing posteriorly to vermiculate-rugose on dorsum of head and occipital border. Dorsa of mesosoma and petiole vermiculate-rugose; Sculptural intensity decreasing laterally to weak- ly rugose. Propodeal spines, besides a few piliferous pits at their bases, highly polished. First gastral lergite fincly, microscopically reticulate, more or less shiny. Short, yellowish to reddish-brown, bristle-like hairs most dense on head and gaster, rather dilute on dorsa of mesosoma and petiole. Short, ap- pressed pubescence very sporadic everywhere, save for the gaster, Where it is rather abundant, ranging from reddish- golden dorsally to silvery on lateral and ventral surfaces, Distinetly bicoloured; head mostly light red- dish-brown with mandibles, clypeus, antennal carinac, median ocellus and the lateral ocellar depressions, narrowly bordered very dark brown, occipital border dark brown. Dorsa of mesosoma and petiole dark reddish-brown, ex- cept the pronoral collar, antero-median patch on mesonotal dorsum, propodeal spines, and sides Of mesosoma and petiole, which are yellowish to light reddish-brown, Guster reddish-brawn, its base widely diffused yellowish-brown, Posterior margins of tergites and sternites widely bordered very dark brown. Appendages reddish- brown. Sexuals and immature stages unknown, POLY RHACHIS VIEHMEYERI SPECIES-GROUP REMARKS The type-series of P, eremita was collected in open sclerophyll forest at the base of Pine Moun- tain, near Marlborough, CQ (Grid cell 22/149). The ground had been disturbed, possibly by recent logging activitics, and the immediate sur- face of the forest floor was badly ravaged. A few ants were observed running in disarray, together with a similarly coloured unidentified species of Rhytidoponera Mayr (Ponerinae), Some specimens were found dead and damaged. Sub- sequent visits lo the area in following years failed to produce further specimens of P. eremita. Polyrhachis greensladei sp.nov. MATERIAL EXAMINED Hovwotyee: SOLOMON IS.: Guadalcanal Prov... MI Austen, 10 ¥-28 vi 1965, P.J.M, Greenslade (worker). PARATYPES: dala as for holotype, 5 x 1965, PIM. Greenslade (1 dealate female); data as for holotype, 5 vii 1984, RIK aee. 84.1 (1 worker); Central Prov., Savo 1, 5 viti 1963, P.M. Greenslade (1 worker). PAPUA NEW GUINEA (PNG): New Ireland Prov.. Lelet Plateau, 800-1000 m, e. 03°20°S,15 1°56’ EB, 19-24 vii 1984, RIK ace. 84.93 (worker). Type deposition: Holotype in ANIC (type no. 7732); 1 paratype (dealate female) in ANIC; | paratype (worker) in BMNH; 2 paratypes (Workers) m RIK. WORKER Dimensions (holotype cited first): TL 7.86, 7,.81-8.77; HL.2.00, 1.96-2.15; HW 1.61, 1.59-1.73; C181, 80-82; SL 2.15, 2.15-2.34; SL 134, 133-135; PW 0.84, 0.87- 0.94; MTL 2.97, 2.97-3.28 (4 measured). Clypeus of holotype in profile almost straight, but rather distinctly sinuate in some paratypes, with posterior margin deeply impressed; median longitudinal carina feebly marked anteriorly, in- distinct posteriorly; truncated median portion of anterior margin distinctly dentate laterally. Ocel- li lacking. Pronotal dorsum with sides subparal- lel; spines moderately long, scarcely elevated. Propodeal suture more or less marked by a shal- low transverse depression. Dorsum of propodeum straight in profile, spines well clevated, moderately divergent, with tips gently turned outwards in dorsal view, Dorsum of petiole flat, sloping posteriorly, with distinet anterior margin; spines relatively long, well elevated, widely divergent. Clypeus, front and sides of head with irregular, mostly longitudinal rugae; sculptural intensity in- creasing posteriorly, so that the dorsum of head is rather coarsely vermiculate-tugose. Dorsa of 503 mesosomad and petiole vermiculate-rugose; sculpturation markedly more fine laterally, Propodeal spines smooth and polished, petiolar spines witha few longitudinal rugac. First gastral tergite basally with more of less regular iransverse striations, opaque. Brown, bristle-like hairs abundant on head and gaster, only slightly less dense on dorsa of mesosoma and petiole, The rather dilute, whitish pubescence has a somewhat reddish tint on the dorsal aspect of gaster. Very datk brown; clypeus, sides of head, mesosoma, petiole and appendages a shade lighter. Mandibles reddish-brown, bordered very dark brown. FEMALE Dimensions: TL 9,17; HL 2.05; HW 1.62; CL 80; SL 2.28; S1 141; PW 1.71; MTL 2,97 (1 measured). The female differs from the worker in the usual characters identifying full sexuality, including three ocelli, complete thoracic structure and wings. The sculpturation, pilosity and colour is essentially that of the worker, and only the con- figuration of spines js different, The pronotal spines are reduced to pair of minute denticles; the propodeal spines are distinctly shorter than in the worker. almost horizontal in lateral view, and divergent. The petiolar spines are similar to those of worker, but markedly shorter. Male and immature stages unknown, REMARKS The known distribution of P. greensladez is from the Bismarck Archipelago in Papua New Guinea to Guadalcanal in the Solomon Islands (Grid cells 3/151, 9/159, 9/160). The locality data label under the holotype states that the specimen was taken in a carrion trap. The single specimen from New Ireland was collected dead onanarrow path winding through low dense fem thickel, which is typical of parts of the Lelet Plateau. Polyrhachis hirta Viehmeyer, 1913 (Figs 1,5) Palyrhachis hirta Viehmeyer, 1913:59. Syntype workers. Type locality; NEW GUINEA, Wareo (Madang Prov,, PNG) (06/147), MNHU (1 syntype examined), LECTOTYPE DESIGNATION Ihave examined one syntype of P. hirta, kindly loaned by Dr Frank Koch of the Museum fir 504 MEMOIRS OF THE QUEENSLAND MUSEUM Fics | - 4, Head in full face view (right antenna omitted): | - P. hirta; 2 - davydovi; 3 - viehmeyert, 4 - bamaga. Fics 5 - 7. Lateral view (antennae, legs and gaster omitted): 5 - P. Airta; 6 - davydovi; 7 - viehmeyeri, Naturkunde, Humboldt-Universitat, Berlin. The specimen is in a fair condition and bears six labels as follows: ‘Wareo, D. Neuguinea’, ‘Typus’ (on red tag), ‘Coll. Viehm.’, ‘Zool. Mus. Berlin’, ‘Syntype’ (round label) and ‘hirta Viehm., det. B. Bolton, 1973’. This specimen is here designated lectotype of P. Airta, and has been so labelled. Dimensions of lectotype: TL 8.16; HL 2.03; HW 1.65; CI 81; SL 2.28; $1138; PW 0.87; MTL 3.22. ADDITIONAL MATERIAL EXAMINED PAPUA NEW GUINEA: Northern Prov,, Managalese Plateau, c. 09°05'S, 148°26'E, S of Popondetta, July 1964, R. Pullen (1 worker); Pongani Riv., c. 500 m, Boikiki Plantation, c. 8 km NNE Afore, 09°06’S, 148°25’E, 29- 30 viii 1984, RIK acc. 84.386 (4 workers): Morobe Prov., Bulolo (07/146), 2300 ft, 3 January 1968, B.B. Lowery (7 workers, 1 dealate female), WORKER Dimensions: TL 7.76-8.92; HL 1.96-2.18; HW 1.59- 1,75; Cl 78-81; SL 2.15-2.43; ST 135-139; PW 0.79- 0.90; MTL. 3.02-3.38 (12 measured). Clypeus in profile almost straight, with basal margin deeply impressed; median longitudinal carina vague; truncated median portion of antcrior margin bluntly angulate laterally. Ocelli lacking. Pronotal dorsum with lateral margins subparallel, bearing a pair of barely elevated, POLYRHACHIS VIFHMEYER! SPECIES-GROUP often asymmetrical spines (including those of leclotype). Propodeal spines only weakly elevated, somewhat divergent. Petiolar dorsum sloping posteriorly, almost flat, with anterior margin distinct or even dorsally produced in some specimens; spines only weakly elevated, divergent. Clypeus with fine, mostly V-shaped mugae, ex- tending to sides and frontal areas of head; sculptural intensity increasing dorsally and posteriorly, with dorsa of head and mesosoma vermiculate- rugose, and decreasing laterally, with sides of mesosoma and petiole less coarsely, somewhat irregularly, rugose. Buse of first gastral lergite finely, mostly transversely, striate, opaque, Short, reddish-brown or yellowjsh, bristle-like hairs on all dorsal surfaces of body, most dense on head and guster. White to silvery, appressed pubescence very sparse, except on gastral dar- sum, where itis more abundant and yellow, with a distinct reddish tint. Medium reddish-brown, mandibles, propodeal and petiolar spines a shade lighter. Mandibles, antennal catinae and lateral margins of mesosoma narrowly and posterior margins of gastral tergites more widely, bordered dark brown. Appendages reddish-brown. FEMALE Dimensions: TL 9.22; HL 2.03; HW 1.57; CI 77; SL 2.28; SI. 145; PW 1.72: MTL 3.17 (1 measured), The single available female closely resembles the worker and, besides the usual characters identifying full sexuality, differs only in the con- figuration of the spines. The pronotal spines are reduced to minute denticles. The propodeal spines are relatively short with tips curved gently outwards: in lateral view the spines are horizon- tal at their bases and then gently downturned- Pctiolarspines are rather short, widely divergent. Male and immature stages unknown. REMARKS I have directly compared the lectotype of P. hirta Viehmeyer with the holotype of P. vieh- meyert Emery and other available material. Despite cerlain similarities of the specimens, | am confident that each name designates separate and valid species. Polyrhachis loweryi sp-nov- MATERIAL EXAMINED HOLOTYPE. AUSTRALIA: Queensland, Miles, 20 viii 1975, B.B, Lawery (wurker). PARATY?ES: data as for holotype (4 workers). Type deposition: Hololype in ANIC (type no. 7733); 1 paralype each in BMNH, MCZC. OMBA and RJK. WORKER Dimensions (holotype cited first): TL8.26, 8.26-8.77; HL 2.03, 1.93-2.00; HW 1.65, 1.56-1.65; CI81, 81-84; SL 2.46, 2.31-2,46: SE 149, 145-152; PW 0.94, 0.87- 0,94; MTL 3.28, 3.12-3,33 (5 measured). Clypeus in profile almost straight with the base moderately impressed; median carina rather blunt, frequently interrupted throughout its length; truncated median portion of anterior mar- gin obtuse laterally. Median ocellus distinct, lateral ocelli lacking (their relative position marked by shallow depressions in the cephalic sculpturation). Pronotal dorsum with sides con- Verging anteriorly; spines well elevated, long and slender. Propodeal suture marked by a shallow transverse depression. Propodeal spines only moderately elevated, subparallel. Dorsum of petiole widely concave between well elevated, relatively long, divergent spines. Clypeus and front of head finely, mostly lon- giludinally, striate-rugose; sides irregularly rugose. Dorsa of head, mesosoma and petiole vermiculute-rugose, with sides distinctly less rugose than dorsum; propodeal declivity transversely rugose, First gastral tergite very shallowly micro-reticulate, more or less shiny. Light to dark brown, bristle-like hairs, most dense on head and gaster, rather dilute on dorsa of mesosomu and petiole. Silvery, appressed pubes- cence, very scafce everywhere except on gaster, where jt is abundant with distinctly reddish tint dorsally. Very dark reddish-brown, with only mandibles (except their masticatory borders), spines and petiole belew spiracles, a shade lighter. Sexuals and immature stages unknown- REMARKS The type-scries was collected in dry sandy sclerophyll forest with Ca/litris, near Miles in southern Queensland (Grid cell 26/150). Lowery’s original data label states that the specimens Were ‘found only in galleries of com- mon large species of Rhytidoponera’, Polyrhachis rustica sp.nov. MATERIAL EXAMINED HOLOTYPE: AUSTRALIA: Queensland, 4 km N of Col- linsville (20/147), savannah Woodland, 24 v 1981, BB, Lowery (worker) 506 PAKATYPES; data as for holotype (1 dealate female). AUSTRALIA. Queensland, Mareeba, Chico(?) Rd. (17/145), 17 vi 1961, LH, Weatherill (BM 1960- 163) (worker). Type deposition: Holotype (type no, 7735) and | paratype (dealate female) in ANIC; 1 paratype (worker) i BMNH, WORKER Dimensions (hlotype cited first); TL 7.16, 7,00; HL 1.78, 1.75; HW 1.47, l.43; CI 83, 82; SL 2.15, 2,00; SI 146, 140; PW 0.81, 0.80; MTL 2.84, 2.68 (2 measured). Clypeus in profile almost straight with rather shallowly impressed posterior margin; median longitudinal carina poorly marked, except for a short, clearly defined anterior section; truncated median portion of anterior margin obtuse lateral- ly, Median ocellus vestigial; lateral ocelli lack- ing. Pronotal dorsum narrowed anteriorly, bearing a puir of well elevated spines, which are rather short and triangular in the holotype, and distinctly longer and more slender in the paratype. Propodeal suture distinct laterally, rather obsolete medially, Propodeal spincs scar- ecly ¢levated, subparallel. Dorsum of petiole with more or less distinct anterior margin, bear- ing # pair of Samewhat divergent, gently curved spines, Clypcus and front of head with fine, more or less longitudinal rugosity. Sculptural intensity increasing dorsally and posteriorly, so that the dorsa of head, mesosoma and petiole are mostly vermiculate-rugose. Sculpturation distinctly less intense: laterally, with sides of mesosoma and peliole somewhal relculate-rugose. Dorsum of first gastral tergite very finely, microscopically reticulate, shiny. Very short brown to yellowish bristle-like hairs rather scarce on dorsa of mesosoma and petiole, more numerous on head and gaster. Short, appressed, silvery pubescence sporadic over most of the body, except the gastral dorsum, where it is more abundant, with a distinct, red- dish-golden tint. Dark reddish-brown with dorsum and sides of mesosoma and petiole infuscated medium red- dish-brown. Dorsum of gaster medium reddish- brown with posterior margins of tergites widely bordered dark brown. FEMALE Dimensions: (LS.97; HL 1.90; HW 1,56; €) 82; SL 2.37; SI152; PW 1.76; MTL 3.22 (1 measured). Very similar lo worker and, besides the ab- vious characters identifying full sexuality, with MEMOIRS OF THE QUEENSLAND MUSEUM the following differences: pronolal spines reduced to triangular, somewhat dorso-ventrally flattened teeth which, in direct comparison to those in other known viehmeyeri species-group females, are distinctly longer; propodeal spines relatively short, their length equal to about half the distance between their bases; petiolar dorsum concave between short, widely divergent, well elevated spines, the anterior petiolar margin rather blint, posterior margin indistinct, Colour brownish-black; mandibles medium reddish- brown with masticatory borders narrowly bor- dered very dark brown. Sides of head at mandibular bases, sides of mesosoma, petiole and gastral (ergites and sternites infuscated medium reddish-brown. Appendages medium reddish-brown, tarsi a shade lighter. Male and immature stages unknown. REMARKS P. rustica is Similar to P. eremita, WW shares with that species a number of characteristics, includ- ing an almost identical clypeal outline, with dis- tinct median carina, The colour scheme of the mesosoma and gaster is similar, Besides the characters given in the key, it differs from eremita in the colour of the head which is uniformly dark reddish-brown in P, rustica, bul conspicuously bicoloured in P, eremita, Also, the petiole in lateral view is distinctly higher and more slender in P. rustica. The holotype and paratype worker of P. rustica were collected from relatively distant localities but, besides the marked differences in the length of pronotal spines, they are closely comparable and undoubtedly conspecific. The variability in the length, elevation and orientation of the pronotal spines ts a peculiar character exhibited to some degree by all known viehyneyeri-group species. | examined many individuals (including types) with the pronatal spines asymmetrical, Although morphologically interesting, this variability seems to be random, and is apparently of no taxonomic value, Like some other species of the group (eremita, loweryi and viehmeyeri), P. rustica seems to be closely associated in the field with ants of the genus Rhytidoponera. Lowery's original label states thal the specimens (holotype and paratype female) were collected ‘under same rock as large Rhytidoponera’. Also the specimen sent by Barry Bolton was apparently collected in as- sociation With ants of that genus, since the GBMNH register for 1966-163 states ‘315 Rhytidoponera Australia... POLYRHACHIS VIEHMEYERI SPECIES-GROUP Polyrhachis stigmatifera sp.nov, MATERIAL EXAMINED HOLOTYPE: INDONESIA; Seram, Solea, viii 1987, M.C. Day (worker). Type deposition: Unique holotype in BMNH. WORKER Dimensions: TL 8.06; HL 2.00; HW 1.59; CI 7% SL. 2.25; SI 141; PW 0.78; MTL 2.96. Clypeus in profile straight, with basal margin deeply impressed; medial longitudinal carina rather acute anteriorly, somewhat less distinct posteriorly; truncated median portion of anterior margin dentate laterally. Ocelli lacking. Pronotal dorsum narrowed anteriorly, bearing a pair of weakly elevated, asymmetrical spines (right spine on the unique holotype is long and slender, while the lefi spine is distinctly shorter and more (riangular). Metathoracic spiracles situated on prominent, laterally projecting tubercles. Propodeum bearing a pair of moderately elevated spines. Petiolar spines scarcely elevated, widely divergent. Clypeus with fine, more or less. Y-shaped Tugae, continuous over frontal area of head. Sculptural intensity increasing posteriorly, with dorsa of head, mesosoma and petiole vermicu- late-rugose, and decreasing laterally with sides only irregularly. rugose. Propodeal and petiolar spines smooth and shiny, save for a few transverse rugae al their bases. First gastral ter- gile opaque; the base somewhat striate-rugase laterally, with sculpturation distinctly less regular dorsally and posteriorly. Rather short yellow to reddish-brown, bristle- like hairs very sparse everywhere, save on the gaster, where they are longer and more abundant, Short silvery to golden appressed pubescence very dilute over most of body, except gaster, where the hairs are longer, with adistinct reddish tint on the dorsal aspect. Dark reddish-brown, lateral portions of head a shade lighter. Mandibles reddish-brown, with masticatory borders narrowly bordered dark hrown. Appendages and tips of spines medium reddish-brown. Sexuals and immature stages unknown, REMARKS P. stigmatifera is only the second species of the P. viehmeyeri species-group known from In- donesia (the other is P. davydovt). Besides the prominent metathorucic spiracles, it is easily separable from davydavwi by the anterior section S07 of the median clypeal carina which forms rather acute ridge; the petiolar darsum with well defined anterior margin; and the generally dark colour of the body. In contrast, the clypeus in P. davydovi is almost flat with the median carina vague; the anterior margin of the petiolat dorsum is ill defined: and general coloration distinctly lighter. Polyrhachis viehmeyeri Emery, 1921 (Figs 3, 7) Polyrhachis (Myrmhopla) viehmeyeri Emery, 1921-19. Holatype worker. NE NEw GUINEA, MCSN (Examined). Dimensions of holotype: TL 8.77; HL 2.06; HW 1.65; CI 80: SL 2.31; SI 140; PW 0.86; MTL 3.22. ADDITIONAL MATERIAL EXAMINED PAPUA New GUINEA: Morobe Prov., Bupu Riy., Lae (06/146), 6 January 1968, B.B. Lowery (47 workers, 2 alate lemules, 4 males); Northern Prov., Kokoda (08/147), 1100", 17 January 1971, B.B. Lowery (1 worker). WORKER Dimensions: TL 7.46-8.52; HL 1.90-2.15; HW 1.57- 1.78: Cl 81-84: SL 2.18-2.34; SI 130-141; PW 0.75- 0.94; MTL 2.92-3.24 (21 measured). Clypceus in profile almost straight, with well impressed posterior margin; median longitudinal carina vague; truncated median portion of anterior margin distinctly dentate laterally. Ocel- li generally lacking, but in some specimens a vestigial median ocellus is evident. Pronotal dor- sum narrowed anteriorly, bearing a pair of well elevated, long, slender spines. Propodeal spines well elevated, subparallel, somewhat divergent in some specimens, Dorsum of petiole with clearly defined anterior margin and well elevated, long, slender spines. Clypeus with fine, somewhat longitudinal Tugae, extending to the frontal areas and sides of head. Sculptural intensity increasing dorsally and posteriorly, so thal the dorsa of head, mesosoma and petiole are vermiculate-rugose. Sculpturation is more: fine laterally, with the sides of the mesosoma and petiole only reticu- late-rugose. Base of first gastral tergite finely, more or less transversely striate, opaque, Mostly reddish-brown, bristle-like hairs abun- dant over most of body. Silvery appressed pubes- cence yery sparse, save for the gastral dorsum, where itis more abundant and somewhat yellow with a reddish tink. Light to medium reddish-brown, dorsa of head 508 and mesosoma a shade darker. Mandibles, anterior clypeal margin and antennal carinae nar- rowly bordered dark brown. FEMALE Dimensions: TL 8.47-8.87; HL 1.93-2.06; HW 1,53~ 1.62; CL 79: SL 2.06-2,15; S1 153-135; PW 1.75-1.81; MTL 2.72-2.92 (2 measured). The female is very similar to the worker, with almost identical sculpluration and colour. Be- sides the characters identifying full sexuality it differs only in the length of the mesosomal spines: pronotal spines reduced to minute den- ticles; propodeals short, subparallel, and almost horizontal in lateral view, The petiolar spines are similar to those of the worker, except shorter, The female of P. viehmeyeri is distinguishable from that of the closely related P. Airfa, and from other known females of the viehmeyert species- group, by its relatively wide mesoscutum, which is as wide, or even wider than long. In com- parison, the mesoscutum of hirta, greensladei and rustica is distinctly more narrow, being longer than wide, The mesoscutum in P, vieh- meyeri is also somewhat concave posteriorly, when viewcd from the side; il is almost straight in all other species. Males and immature stages in ANIC. REMARKS I have examined and directly compared the unique holotype of P. viehmeyeri with numerous specimens collected by Rev. Lowery. They matched the holotype closely. The main charac- ters separating this species from the closely re- lated P. hirta and other species of the group are given in the key and in the discussion under P. davydovi and P. hirta. ACKNOWLEDGEMENTS [am grateful to Dr Robert W. Taylor, Division of Entomology CSIRO, Canberra, for access to ANIC material, to Barry Bolton, for providing specimens from the British Museum (Natural History), London, U.K., and to both for their MEMOIRS OF THE QUEENSLAND MUSEUM comments. received during the course of this study. Thanks for the loan of types is due to Dr Frank Koch, Museum fiir Naturkunde, Hum- boldt-Universitat, Berlin, D.D.R.; Drs Roberto Poggi and Valter Raineri, Museo Civico di Storia Naturale ‘Giacomo Doria’, Genoa, lialy; and Dr A.G, Radchenko, Zoological Institute, Ukrainian Academy of Sciences, Kiev, U.S,S.R. 1 am also indebted to Dr Geoff B. Monteith, Queensland Museum, Brisbane, and to Dr Robert W. Taylor, for their comments on the draft manuscript. Finally, to Mr Edward C. Dahms, Queensland Museum, I owe my thanks for his support in various aspects related to this study. LITERATURE CITED Emery, C. 1921. Le genre Polyrhachis. Classification; espéces nouvelles ou critiques. Bulletin de la Suciété vaudoise des. sctences naturelles. Lausanne 54: 17-25. 1925. Hymenoptera, Pam, Formicidae, Subfam. Formicinae in Wytsman (Ed,) Genera Insec- torum fase. 183, 302 pp., 4 pls, Bruxelles. FoReEL, A. 1915. Results of Dr BE, MjGbergs Swedish Scientific Expeditions to Australia 1910-1913. 2. Ameisen. Arkiv fér Zovlogt 91-119, pls 1-3. KARAWAJEW, W. 1927. Ameisen aus dem Indo- Ausiralischen Gebiet. Travaux du Musée zovlozique de UAcadémie dey sciences de |’- Ukraine, Kiev 3:3-52, 21 figs. Konourt, RJ. 1988. A new species of Polyrhachis (Polyrhachis) from Papua New Guinea with a review of the New Guinean and Australian species. Memoirs of the Queensland Museum 25(2); 417-427, TAYLOR, R.W, 1987. A Checklist of the Ants of Australia, New Caledonia and New Zealand (Hym.; Formicidae). CSIRO Australia, Division of Entomology Report No.41;1-92. TAYLOR, R.W. AND Brown, D.R. 1985. Hymenop- tera: Formicoidea in Zoological Catalogue of Australia 2:1-149, 306-348. VIEHMEYER, H. 1913, Neue und unvollstindig bekan- nte Ameisen der Alten Welt. Archiv fiir Natur- geschichte, Berlin 79A:24-60, NOTES ON AUSTRALIAN ANTS OF THE GENUS POLYRHACHIS FR.SMITH, WITH A SYNONYMIC LIST OF THE SPECIES (HYMENOPTERA: FORMICIDAE: FORMICINAE) RUDOLF J. KOHOUT AND ROBERT W. TAYLOR Kohout, R.J. and Taylor, R.W. 1990 08 31. Notes on Australian ants of the genus Polyrhachis Fr.Smith, with 2 synonymic list of the species (Hymenoptera: Formicidae: Formicinae). Memwirs of the Queensland Museum 28(2); 509-522. Brisbane. [SSN (1079- 8835, The named Australian species of the ant genus Polyrhachisate reviewed. Eleven subspecies are raised lo Species: P. guerini lata Emery, P. hoakeri lownei Forel, P. hookeri obscura Forel, P. aurea obtusa Emery,P. guerinipallescens Mayr, P. rastellata pilosa Donisthorpe, P. sexspinosa reclinata Emery, P. appendiculata schoopae Forel, P. guerini vernliculosa Mayr, P. lombokensis varrabahkensis Forel and P, rastellaia yorkang Forel. P. australis Mayr, formerly a junior synonym of P. levior Roger, is declared a valid species. Nine new synonyms are proposed (senior names cited first): P. australis Mayt = P. nox Donisthorpe, P, creusa Emery = P. creusa chlorizans Forel = P. hecuba Forel, P. heinlethii Forel = P. heinlethit sapkiae Forel, P. hookert Lowne = P. hookeri aerea Forel, P. lata Emery = P. gab aegra Forel, P. pallescens Mayr =P. aurea depilis Emery, P. phryne Forel =P. sidnica perthensis Crawley, P. sexspinosa (Latreille) = P. barnardi Clark. Seven Papuasian species are recorded for the first time from Australia: P. argenteosignata Emery, P. atropos FrSmith, ?. denticulata Karawajew, P. mucronata Fr.Smith, P. obtusa Emery, P. reclinata Emery and P. semiobscura Donisthorpe. P. guerini Roger, P, levior Roger and P. lomboken- sis Emery, which were previously reported from Australia, apparently do not occur there. P. guerini is known only trom New Caledonia and is apparently endemic there. A current synonymic listof the 114 valid named Australian Polyrhachis species is provided, and (he position of all names in the traditional subgeneric arrangement is indicated. O Formicidae, Polyrhachis, Australia, synonymy, distribution, checklist. Rudolf J. Kohout, Queensland Museum, PO Rox 300, South Brisbane, Queetsland 4101, Australia; Rabert W. Taylor, Australian National Insect Collection, CSIRO Division of Entomology, GPO Box 1700, Canberra, A.C.T. 2601, Ausiralia; 10 May, 1989, This study culminates a series to which both authors have contributed (Kohout, 1988a, 1988b, 1988c, 1989, 1990; Taylor, 1989), With these studics, we have aimed to review the iden- titics and nomenclature of all described Australian species of the large and diverse for- micine ant genus Polyrhachis Fr.Smith, In this paper, a number of species-group taxa are dis- cussed in detail. They are the last remaining for consideration in this project. Dunng the 210 years preceding 1985, 20 authors generated a nomenclature comprising 137 available specific and subspecific names for Australian Polyrhachis. As well, several in- frasubspecific names, which have no formal status in zoological nomenclature (Taylor, 1986), were proposed. The arrangement of species thal resulted was summarised by Taylor and Brown (1985) and Taylor (1987). Because ho comprehensive revision of the genus was available, these lists ranked the available names according to the their then most recent taxonomic assignment. In Taylor’s checklist (1987), for example, 86 names were ranked as species, and 33 as subspecies, At that time, only 5 names had previously been assigned to junior synonymy and 3 tojunior homonymy. However, because this arrangement was assembled piecemeal and neither subjected to critical review nor revision, any impression of sys- tematic comprehensiveness in the resulting nomenclature is illusory. Following Our studies, we are now able to recognise 114 validly named species of Polyr- hachis trom Australia, and 33 junior synonyms or homonyms. We find no justification for con- tinued nomenclatural recognition of subspecies. Tn our recent papers, 20 of the subspecies listed by Taylor (1987) have been elevated to specific rank, and 13 consigned to junior synonymy, Of the latter, 5 are treated as junior synonyms of their species, 2 as junior synonyms of anelevated name that was previously treated as a subspecies of the same species, and 6 as juninr synonyms of species with which they were not previously associated, Six of the 28 names cited below as senior synonyms were originally proposed for Papuasian species. Four of these (P. creusa Emery, P. dives Fr.Smith, &. fervens Fr.Smith, and P. insularis Emery) have not been used before for Australian ants. Five names of South- east Asian or Melanesian taxa (P. everini Roger, P. ithona VrSmith, P. leviar Roger, P. lom- bokerisis Emery and P. relucens (Latreille)), which were previously treated as senior synonyms of Australian names of were con- sidered to be nominotypical subspecies of Australian taxa, no longerapply to the Australian fauna. Finally, the first Australian records are noted of LO species originally described from lands to the north (P. argenteasignata Emery, P. atrapos Fr.Smith, P. consimilis Fr.Smith, P. den- ticulata Karawajew, P. mucronata Pr.Smith, ohiusa Emery, P, paxilla Fr.Smith, P, reclinata Emery, P. rufofemorata Fr.Smith and P. semi- obscura Donisthorpe), Some of the species listed below have not been discussed in detail in our papers. Nonctheless, the species have been studied, found to be valid, and do not require further comment at present We cnvisage that few changes wil) be required in the nomenclature proposed below, However, there 18 (he possibilily thal we may have oVver- looked sibling species concealed by variability that we have interpreted here as infraspecific (a problem always prominent in taxonomic work on Australian ants). All of the species listed are represented in the Australian National Inseet Collection (ANIC), Canberra, and most are in the Ouvensland Muscum (QMBA). Brisbane. The ANIC has type material of a number of taxa, including syntypes or paralectotypes of several of the species or Subspecies described by Auguste Forel, These were generously donated in 1968 by the Muséum d Histoire Naturelfe, Geneva, Swit- zetland. In addition the ANIC contains many specifically labelled voucher specimens iden- tified by one of us following direct comparison with type matenal on loan from foreign or other Australian collections. Between us, we have seen types of every relevant available name for which types are known to exist, We also believe that any specimen bearing one of qur determination labels writlen during ar since 1987 can be con- sidered to have been identified with confidence. We have also designated and used a few par- ticular ANIC vouchers as nomenclatural MEMOIRS OF THE OURFNSLAND MUSEUM paradigms for species of which the types are deemed to have been lost (see P. guerini), Asa result, we believe that Australian workers using our published notes, and the resources of either collection should have little difficulty in iden- tifying material of named Polyrhachis species or in recognising specimens representing un- described species. The undescribed species known to us will at least double the Australian species tally. The next phase of our project will altend toTheir description and ultimately to the provision of keys enabling the ready identifica- tion of all relevant species. Although we do not make formal use of the Polyrhachis subgenera of the Emery/Wheeler classification (Wheeler, 1922; Emery, 1925), we consider that they do provide useful foci for the sorting of species. The collections in our care are arranged following this system, generally as simplified by Hung (1967). For this reason a second list of species is given below, arranged in subgeneric groups. We do not support the direct use of species-group names to replace those of the subgenera. It seems unlikely that all of these subgenera are monophyletic. As well, a number of apparently monophyletic species-groups, which do not correlate with the present sub- genera, can be discerned, We expect the present subgenera to be replaced by a larger set of infor- mally-named species-groups bit, prior to an overall formal taxonomic analysis, we will not initiate such a classification, No apology is made for this apparent ambivalence; the categorical subvision of Polyrhachis cannot be resolved until all of the specics now known are nomenclaturally and taxonomically accessible, and have heen subjected to classificatory analysis. We trust (hat this paper will contribute meaningfully to the achievement of such a clas- sification. Abbreviations for institutions (with the names of co-operating curators) are: ANIC, Australian National Insect Collection, Canberra; BMNH, British Museum (Natural History), London (Barry Balton); BPBM, Bernice P, Bishop Museum. Honolulu, Hawai’i, U.S.A. (Dr G, Nishida); HNHM, Hungarian Natural History Museum, Budapest (Dr J. Papp); [ZAS, Institute of Zoology, Academy of Sciences, Kiev, U,S,S.R. (Dr AG. Radchenko); MCSN, Museo Civico di Storia Naturale ‘Giacomo Doria’, Genoa, llaly (Drs R. Poggi and V. Raineri); MCZC., Mustum of Comparative Zoology, Har- vard LIniversity, Cambridge, Mass., U.S.A. (Dr A.V, Newton, Jr); MHNG,. Muséum d'Histoire ANT GENUS POLYRHACHIS Naturelle, Geneva, Switzerland (Dr C. Be- suchet); MVMA, Museum of Victoria, Mel- bourne (Dr A. Neboiss and K. Walker); NHMW, Naturhistorisches Museum, Vienna, Austria (Dr M. Fischer); NHRS, Naturhistoriska Riksmusvet, Stockholm, Sweden (Dr K-J. Hed- quist); OXUM, University Museum, Oxford, U.K. (Drs G. McGavin and C, O'Toole); QMBA, Queensland Museum, Brisbane (E.C. Dahms and Dr G.B. Monteith). SYNONYMIC LIST OF AUSTRALIAN POLYRHACHIS SPECIES The following list includes all described Australian Polyrhachis species known to the authors. Non-Australian junior synonyms are ex- cluded. Names of the unresolved P. (sidnica) complex (Taylor. 1989) - comprising P. sidnica, P. cedarensis, P. leae, and P. tambourinensis - are listed as species. For publication details see Taylor and Brown (1985) or Taylor (1987), and Emery (1925). Synonyms are in italics. ammon (Fabricius, 1775) ammon angustata Forel, 1902 ammonocides Roger, 1863 andromache Roger, 1863 connectens australiae Emery, 1887 angusta Forel, 1902 appendiculata Emery, 1893 arcuata (Le Guillou, 1841) argentosa Forel, 1902 argenteosignata Emery. 1900 atropos Fr.Smith, 1860 aurea Mayr, [876 australis Mayr, 1470 nox Donisthorpe, 1938 syn.nov. bamaga Kohout, 1990 barretti Clark, [928 bedoti Forel, |} 902 bellicosa Fr.Smith, 1859 bicolor Fr.Smith, 1858 cataulacoidea Stitz, 1911 cedarensis Forel, 1915 chalchas Forel, 1907 cleopatra Forel, 1902 clio Forel, 1902 clotho Forel, 1902 constricta Emery, 1897 consimilis Fr.Smith, 1858 contemta Mayr, 1876 crawleyi Forel, 1916 creusa Emery, 1897 hecuba Forel, 1902 syn.nov. cupreata Emery, 1895 daemeli exlex Forel, 1915 daemeli Mayr, 1876 denticulata Karawajew, 1927 dives Fr.Smith, 1857 exulans Clark, 1941 doddi Donisthorpe, 1938 erato Forel, 1902 aeschyle Forel, 1915 eremita Kohout, 1990 cuterpe Forel, 1902 femorata Fr.Smith, 1858 emeryi (Forel, 1880) fervens Fr.Smith, 1860 kershawi Clark, 1930. flavibasis Clark, 1930 foreli Kohout, 1989 fuscipes Mayr, 1862 semipolita hestia Forel, 1911 gab Forel, 1880 glabrinota Clark, 1930 gravis Clark, 1930 heinlethii Forel, 1895 heinlethii sophiae Forel, 1902 syn.nov. hermione Emery, 1895 hexacantha (Erichson, 1842) froggatti Forel, 1910 hirsuta Mayr, 1876 hirsuta quinquedentata Viehmeyer, 1925 hookeri Lowne, 1865 hookeri aerea Forel, 1902 syn.nov. inconspicua Emery, 1887 thalia Forel, 1902 insularis Emery, 1887 inconspicua subnitens Emery, 1895 inusitata Kohout, 1989 io Forel, 1915 jacksoniana Roger, 1863 lachesis Forel, in Emery 1897 lata Emery, 1895 stal.nov, gah aegra Forel, 1915 syn.nov. latreillii (Guérin-Méneville, 1838) leae Forel, 1913 loweryi Kohout, 1990 lownei Forel, 1895 stal.nov. lydiae Forel, 1902 lysistrata Santschi, 1920 machaon Santschi, 1920 mackayi Donisthorpe, 1938 macropus Wheeler, 1916 longipes Wheeler, 1915 maculata Forel, 1915 micans Mayr, 1876 mjobergi Forel, 1915 anguliceps Viehmeyer, 1925 mucronata Fr.Smith, 1859 obscura Forel, 1595 stat.nov. obtusa Emery, 1897 stat.nov. 312 MEMOIRS OF THE QUEENSLAND MUSEUM ops Forel, 1907 ornata Mayr, 1876 humerosa Emery, 1921 chrysothorax Viehmeyer, 1925 pallescens Mayr, 1876 stat.nov. aurea depilis Emery, 1897 syn.nov. patiens Santschi, 1920 paxilla Fr.Smith, 1863 penelope Forel, [895 phryne Forel, 1907 sempronia Forel, 1907 perthensis Crawley, 1922 syn.nov. pilosa Donisthorpe, 1938 stat.nov. polymnia Forel, 1902 prometheus Sanischi, 1920 pseudothrinax Hung, 1967 punctiventris Mayr, 1876 pyrrhus Forel, 1910 queenslandica Emery, 1895 delicata Crawley, 1915 rastellata (Lalreille, 1802) reclinata Emery, 1887 stal.novy, rowlandi Forel. 1910 rufifemur Forel, 1907 terpsichore elegans Forel, 1910 rufofemorata Fr.Smith, 1859 rustica Kohout, 1990 schenkii Forel, 1886 schoopae Forel. 1902 slal.nov. schwiedlandi Forel, 1902 semiaurata Mayr, 1876 semiobscura Donisthorpe, 1944 semipolita André, 1896 senilis Forel, 1902 gab tripellis Forel, 1915 comata Crawley, 1915 erawleyella Santschi, 1916 sexspinosa (Latreille, 1802) barnardi Clark, 1928 syn.nov. sidnica Mayr, 1866 quadricuspis Mayr, 1870 sokolova Forel, 1902 sokalava degener Forel, 1910 tambourinensis Forel, 1915 templi Forel, 1902 terpsichore Forel, 1893 thais Forel, 1910 thusnelda Forel, 1902 townsvillei Donisthurpe, 1938 trapezoidea Mayr, 1876 tubifera Forel, 1902 turneri Forel, 1895 urania Forel, 1902 yermiculosa Mayr, 1876 slat.nov. yarrabahensis Forel, 1915 stat.nov. yorkana Forel, 1915 stat:nov. zimmerae Clark, 194i SUBGENERIC ARRANGEMENT OF THE SPECIES The following list, which basically follows the subgeneric arrangement of the Emery/Wheeler classification, is presented with the qualifica- tions previously mentioned. Polyrhachis (Campomyrma): cedarensis, creusa, fémorata, flavibasis, fuscipes, gravis, hexacantha, hirsuta, inconspicua, insularis, to, jacksoniana, leae, macropus, maculata, micans, ops, patiens, phryne, polymnia, prometheus, pseudothrinax, pyrrhus, schwiedlandi, semi- polita, sidnica, tambourinensis, templi, zim- merae. P, (Chariomyrma): appendiculata, arcuata, argenteosignata, aurea, bedoti, cataulacoidea, constricta, cantemta, gab, heinlethii, hookeri, lata, latreillii, lownei, obscura, obtusa, palles- cens, punctiventris, rawlandi, schoopae, senilis, urania, vermiculosa. P. (Cyrtomyrma): australis, doddi, mackayi, pilosa, rastellata, townsvillei, yorkana. P. (Hagiomyrma): ammon, ammonoeides, an- gusta, chalchas, crawleyi, denticulata, lachesis, lydiae, paxilla, penelope, schenkii, semiaurata, semiobscura, sokolova, thusnelda, trapezoidea, tubifera. P. (Hedomyrma): argentosa, atropos, barret- ti, cleapatra, clio, clatho, consimilis, cupreata, daemeli, erato, euterpe, fervens, hermione, machaon, mjobergi, ornata, rufifemur, terpsichore, thais, turneri. P. (Myrma): andromache, foreli, inusitata, rufofemorata. P. (Myrmatopa): yarrabahensis. P. (Myrmhopla): bamaga, bicolor, dives, eremila, glabrinata, loweryi, mucronata, reclinata, rustica, sexspinosa. P. (Myrmothrinax): lysistrata, queenslan- dica. P. (Polyrhachis): bellicosa. NOMENCLATURAL NOTES ON SOME RECENT POLYRHACHIS RECORDS Voucher specimens supplied by Dr A.N. Andersen enable the following allocation of the names he used in his ecological papers (Ander- sen, 1986a, 1986b, 1988). Specimens of: (1) P. femorata wete vatiously identified as ‘femorata’,‘femorata sp A’ or ‘femorata sp B’; (2) P. fuscipes as *hexacantha’ or ‘hexacantha sp A’; (3) P. patiens as ‘micans’, ‘micans sp B’ ANT GENUS POLYRRACHIS 413 or ‘micans sp C'; and (4) P. (sidnica) as ‘hexacantha sp B’. Specimens identified by Imai, Crozier and Taylor (1977) using the voucher designation ‘Polyrhachis sp no 1 (ANIC) tepresent an un- named ‘Campomyrma™ species, which will be described elsewhere. DISCUSSION OF INDIVIDUAL SPECIES In the following, distributions are summarised using ‘short’ coordinates to specify 1-degree mupping grid-cells; all latitudes are for the southern hemisphere. Unless otherwise indi- cated, all specimens discussed are workers and all nomenclatural changes have been based on direct comparison of relevant type specimens. Polyrhachis atropos Fr.Smith, 1860 Palyrhachis atropoy Fr.Smith, 1860: 100, pl.t,fig.22. Type Jocality: New Guinea, Dory (= Manokwari, Irian Jaya), (hololype examined, OXUM). Palyrhachis (fledamyrma) eucharis Karawajew, 1927:22, fig.12. Type locality: Aru, Wammiar 1., Dobo (= Kepulauan Aru, Indonesia) (syntype ex- amined, IZAS). syn.nov, P. atrapas has not previously been recorded from Australia. Specimens from Maer (= Mur- ray) fsland (09/144) in Torres Strait (29,vii1974, H. Heatwole and E, Cameron) and Bamaga (10/142) on Cape York Peninsula (18,24. i.1987, R.J. Kohout (RIK)) are in the ANIC and QMBA. The species is known from the Moluccas through West Irian to Papua New Guinea and northern Cape York Peninsula, Queensland (Qld). Recent New Guinean records are from Oransbari, S of Manokwari (01/134), Aitape (03/142), Lumi (03/142), Umboi [. (05/147), Wampit nr Lac (06/146), Kokoda (08/147), Rouna (09/147) and Pongani R. (09/148), This species. nests in small hollow twigs on standing low trees and shrubs. Polyrhachis argenteosignata Emery, 1900 Polyrhachis argenteosignata Emery, 1900;335, Type locality: New Guinea (synlype examined, MCSN). This New Guinean species is recorded for the first time from Australia where it is known only from lowland rainforests on northern Cape York Peninsula, at Lockerbie Scrub (10/142), Bamaga (10/142) and Iron Range (12/143). New Guinean records are from at or near the following localities: Oransbari, S of Manokwari (01/134); Kar Kar I., Kurum (04/145); and Wampit, nr Lae (06/146), P. argenteosignata inhabits lowland and gallery rainforests where it mostly frequents the edges, rather than deeper in the forest or in the surrounding savannah woodJand. Polyrhachis australis Mayr, (870 Polyrhachis australis Mayr, 1870:945, Type locality: Port Mackay, Ostaustralien (= Mackay, Qld) (type presumed lost). Polyrhachis levior Roger, 1863; Mayr, 1876:71, (spurious synonymy of australis under P. levior), Polyrhachis (Cyriomyrma) nox Donisthorpe, 1938:249. Type locality: Mackay, Queensland (2 syntypes examined, BMNH). syn.noy. The holotype of P. australis cannot be found in the Mayr collection and appears to have been last. However, details given in the original description, and stated in support of the sub- sequent inappropriate (see below) synonymy of australis under P. levior Roger (= laevissimus Fr.Smith, 1859) (Mayr, 1876:71), enabled us to identify P. australis with reasonable confidence. In this species the pronotal humeri are angulate and the propodeum is armed with a pair of spines. The same characters were used by Donisthorpe to characterise his P. nox, the types of which were evidently collected at Mackay (Qld), as were those of P. australis. We are confident that these names apply to a single species and thatnox is thus a junior synonym of australis. Qn the other hand, examination of the holotype of P. levior shows il cleatly to be representative of another species. Its head is missing, but the mesosomal structure is yery distinctive. The pronotal humeri are distinctly bilobed, and the propodeal spines relatively long and thick, some- what dorsoventrally flattened and rather bluntly terminated. In contrasl, the pronota! humeri of P. australis (as represented by the P. nox syntypes and other available material) are only bluntly angulate, and the propodeal spines distinctly shorter, more siender and more acute than in levior. We have not seen confidently identified Australian specimens of P. levier, and thus con- sider it no longer to be referable to the Australian fauna. P. australis is known from just north of Mack- ay (20/148), south to Rundle Range (23/150). It is an arboreal species that builds its nests by a4 joining the leaves of various trees and shrubs with silk. Polyrhachis creusa Emery, 1897 Polyrhachis creusa Emery, 1897:577. Type locality: New Guinea, Ighibirei (holotype examined, MCSN). Polyrhachis creusa var. chlorizans Forel, 1901:30. Type locality: Bismarck Archipelago, Ralum (04/152) (Papua New Guinea (PNG), East New Brilain Prov.) (4 syntypes examined, MHNG). syn.nov. Polyrhachis kecuba Forel, 1902:527. Type locality: Qld, Mackay (21/149) (2 syntypes examined, ANIC). syn.oov. The creasa holotype has somewhat less clearly expressed gastral shagreening than the chlorizans syntypes. This difference is slight, however, and does not justify separate specific status, When describing P. creusa Emery noted its similarity to the then unpublished P. hecuba Forel (of which Forel had sent him specimens). Comparison of the types shows that (hese names are synonyms. In fact, were it not for modern material collected in both New Guinea and Australia, we would suspect that one of the type series had been mislabelled, so closely do they match. Jt is historically of intcrest to note that Forel had evidently distributed identified specimens of P. hecuba by 1897, five years before he formally published the name. Modern type-compared ANIC vouchers are from Mingende (05/144), Papua New Guinea (collected in grassland at SOO elevation, B.B. Lowery, 13.13.1968), and Finch Hatton Gorge, Eungella Nat. Park (21/148), Qld (7-13 April 1975, RJK). The creusa ehlorizans types were taken ‘im Graslande und am Strande’ (Forel, 1901), This species is widely distributed both geographically and altitudinally and, like other species discussed here that are found both in Australia and New Guinea, it was probably vicariantly isolated when the two lands were last separated, Polyrhachis denticulata Karawajew, 1927 Polyrhachis (Hagiomyrma) denticulata Karawajew, 1927:13, Type locality; Amboina (= Ambon), {n- donesia)(syntype examined IZAS). P. denticulata has been reported previously MEMOIRS OF THE QUEENSLAND MUSEUM only from the Molucceas and New Guinea. The first Australian record is based on a single worker collected on Mabuiag Island (09/142), Torres Strait (1974, H. Heatwole and E.Cameron: ANIC). Recent Papua New Guinea records are from at or near Lac (06/147), Buna (08/148), Oro Bay (08/148), and Milne Bay (10/150). Polyrhachis guerini Roger, 1863 Although its holotype cannot be located in any relevant European collection and appears to have been lost, P. gveriné may be recognised as the only Polyrhachis species known from New Caledonia, where it has been abundantly col- lected, In lieu of the type we recognise several ANIC youcher specimens from Mt Panié (20°34’S, 164°46'E) (Melaleuca scrub, 120m, 15 February 1977, P.S. Ward acc. 2218) as the nomenclatural paradigms of the name gwerini. Designation of a neotype would not be ap- propriate here, The Australian species P. lata, P. pallescens and P. vermiculosa were first described as sub- species of P. guerini (see below). We have seen no Australian specimens that are conspecific with the New Caledonian species and consider P. guerini to be endemic to New Caledonia. It was originally described from ‘Neuhalland’, but this citation scems to have been an error (Emery, 1897:588-589; Emery, 1914:428). Polyrhachis heinlethii Forel, 1895 Polyrhachis heinlethii Forel, 1895:47. Type locality: Qld, Mackay (21/149) (6 syntypes examined, MHNG, ANIC). Polyrhachis heinlethii var. sophiae Forel, 1902:521. Type locality: Old, Mackay (21/149) (5 syntypes examined, MING, ANIC), syn.noy, P. heinlethii sophiae was. putatively distin- guished from the nominotypical subspecies by its smaller size, the shape and length of its prona- tal and propodeal spines, and its more regularly striate pronotal dorsum. These characters are now known to vary infraspecifically, Examina- tion of numerous specimens from throughout the range of P. heinlethit has revealed no other taxonomically significant variation. This is a ground nesting species which inhabits open forests and woodlands. It is known from Mt Ossa, NQ, south to Campbelltown, New South Wales(NSW),. (Grid cells 20/148, 20/149, ANT GENUS POLYRHACHIS 25/153, 26/153, 27/152, 27/153, 34/150). 28/153 Js Polyrhachis hookeri Lowne, 1865 Polyrhachis hookeri Lowne, 1865;334, Type locality: NSW, vicinity of Sydney (as Sidney) (33/151) (lypes presumed Jost), Polyrhachis hookeri var. aerea Forel, 1992:521. Type locality: Qld, Mackay (21/149) (2 syntype workers, | alate queen examined, MHNG), Syn.nov. Lowne’'s type material of P. hookeri cannot be found in the collections of the BMNH or OXUM and must be presumed to have been lost (Bolion, pers.comm.). The original description, however, enables reasonably confident jdentification of this species. In lieu of a type we recognise a topotypical worker (ANIC) from Manly Reser- voir, Sydney (33/151), NSW (sandstone scrub, 250ft. 19.v.1960, B.B. Lowery) as the nomenclatural paradigm of the name hookert. Designation of a neoltype would not be ap- propriate here, because the type(s) might still be extant. Comparison of syntypes of P. hookert aerea with this paradigm, and with other specimens collected near Sydney, shows them Lo be conspecific. P. hookeri is acommon ground-nesting ant which inhabits open forests and woodlands. [t ranges from NOQ to central coastal NSW (Grid cells 19/146, 20/148, 20/149, 21/148, 21/149, 22/149, 23/149, 23/150, 25/153, 27/150, 27/152, 27/153, 28/151, 28/152, 33/151), Specimens from N and CQ tend generally to be relatively large, with more acute pronotal spines than those of other series, and with the colour of the mesosomal dorsum ranging from bright metallic green to various. shades of blue and purple, Southern specimens are generally somewhat smaller, with the pronotal spines less strongly projecting and the mesosomal dorsum more uniformly green. These forms, however, intergrade, and we perceive a single, somewhat variable, species, Polyrhachis lata Emery, 1895 stat.nov. Polyrhachis guerini ssp. lata Emery, 1895;357. Type locality: Old. Somerset (10/142) (synlype ex- amined, MHNG). Palyrhachis (Chariomyrma) gab var, aegra Forel, 1915:109, Type locality: Old, Atherton (17/145) (6 synlypes examined, MHNG) (Raised to species by Kohout, 1988:49). syn.nov, Comparison of the P. guerini lata syntype with the AN[C nomenclatural paradigms of P. guerini (see above) and consideration of other Australian and New Caledonian material shows P. guerini and P. lata to be separate species. Comparison of the lata and aegra syntypes shows that they are conspecific. P. lata is known from the Torres ‘Strait (Horn 1.) and Cape York Peninsula, south to Rundle Range near Gladstone, Qld (Grid cells 10/142,.16/146, 17/145, 17/146, 19/147, 20/148, 21/148, 23/150). It is an inhabitant of open sclerophyll forests and savannah woodlands. Polyrhachis lownei Forel, [895 stat.nov. Polyrhachis hoekeri r. lownei Forel, 1895:44. Type locality: Old, Mackay (21/149) (10 syntype workers, | alale queen examined, MHNG, ANIC), The most obvious difference between syntypes of P. hookeri lownei and specimens considered here to be conspecific with the ANIC nomenclatural paradigm of P hookeri (sec above) is the colour of the mesasomal dorsum. In hookeri this ranges from metallic green to blue and purple, while in /ownei it is uniformly very dark brown, at times almost black. Also, the propodeal spines in Haekeri are relatively short (distinctly shorter than the distance between their bases), straight and divergent, while in lownei they are longer (with length equal to or exceeding the distance between their bases), and are gently curved, so that the extreme apices project posteriorly. We consider these types to represent separate species, P, lownei appears to be uncom- mon, and is perhaps confined to the higher al- titude sclerophyll forests of NO, from Kuranda south to Eungella, inland from the type locality (Grid cells 16/145, 18/145, 18/146, 21/148). Polyrhachis mucronata Fr.Smith, 1859 Polyrhachis mucronatus Fr.Smith,1$59:140, Type locality: Indonesia, Aru (= Kepulauan Aru, 06/134) (holotype examined, OXUM). P. mucronata has been reported previously only from Kepulauan Aru (Moluccas) and New Guinea. It has recently been taken ata number of localities on or adjacent lo Cape York Peninsula. Examination of the holotype and of all available Australian and New Guinean specimens shows mucronata to be somewhat variable. However, the observed differences between specimens Sift seem not to be taxonomically significant, They include relatively ‘soft’ characteristics, such as the degree of elevation of the propodeal spines and the colour of the appendages. In Australia mucronata ranges from Lockerbie Scrub, south to Hinchinbrook Island, Old (Grid cells 10/142, 12/143, 16/145, 17/145, 17/146, 18/146). Moluccan and New Guinean records are from Vogelkop, Sururai (01/133); Biak [. (01/136); Lumi, Torricelli Mts (03/142); Naru, SW of Madang (05/145); Wampit, nr Lae (06/146); and or Sogeri (09/147). This species builds arboreal nests by joining leaves with silk, or by constructing small, pocket-like structures of silk and assorted plant fragments against the underside of living Jeaves on low trees and shrubs, It inhabits lowland tropical rainforest. Polyrhachis obscura Forel, 1895 stat.nov. Polyrhachis hookeri t. obscura Forel, |895;44. Type locality; Qld, Mackay (21/149) (8 syniype workers, Z alate queens examined, MHNG, ANIC). Comparison of P. Aookeri obscura syntypes with the ANIC nomenclatural paradigm of P. hookeri (see above), and consideration of other material, demonstraics that obscura almost cer- tainly constitutes a species separate from P. hookeri. The most obvious differences involve the colour of the mesosomal dorsum, which in hookeri is mostly metallic green, blue ar purple, but dark brown in obscura, with a faint but distinct coppery metallescence. Also, the gaster in hookert as brownish-black, usually with a metallic green sheen, and has a dark brawnish- maroon coloured median Jongitudinal dorsal patch, The gaster in P. obscura is reddish-brown (lighter and more reddish than the mesosoma), with an equivalent dorsal patch much less dis- linctly visible. The propodeal spines of P. obscura are relatively short, dorsa-ventrally flat- tened and unusually wide at the base; cach has a conspicuous, brightly reflective smooth dorsal patch near its base; the petiolar spines are rela- tively shart compared ta those of hookeri. P. obscura is apparently rare; the only specimens known to us are the types. Polyrhachis obtusa Emery, 1597 stat-nov. Polyrhachis aurea vat. obtusa Emery, 1897; 589, Original localities; New Guinea: Haveri (09/147), Kapa Kapa (09/147), Humboldt Bay (02/140) (sya- type from Haveri examined, MNHG). MEMOIRS OF THE QUEENSLAND MUSEUM The syntypes of P. aurea (original localities: Qid, Rockhampton (23/150) and Gayndah (25/151), workers, NHMW, MHNG) and P. aurea obtusa clearly represent separale species. The pronotal hameri of aurea are distinctly spinose, while those of obtusa are at most obtuse- ly angulate or subdentate, The head in aurea is relatively narrow, and the Scape Index (SI = Scape Length x 100/Head Width, measured im- mediately in front of the eyes) relatively high (>125). SI in P. obtusa is <117. Available specimens of P. obtusa are generally smalier than those of P. avrea; Head Length (HL) of the obtusa syntype is 1.65 mm; other New Guinean worker specimens are consistently smaller; the smallest examples we have seen are from the Torres Strait Islands, where HL ranges from 1.37 to 1.53 mm, HL in available P. aurea specimens exceeds 1.70 mm, The first Australian records of P. ob/usa are from the Torres Strait Islands: Boigu (09/142), Darnley (09/143), Moa (= Banks, 10/142), and “Prince of Wales (10/142)(all 1974, H. Heatwole and E. Cameron; ANTC). Recent West Irian and Papua New Guinea records are from Nabire, S of Geelvink Bay (03/135); Lumi, Torricelli Mountains (03/142); Madang, Amele (05/145); near Lae (06/147); and Obo, Fly River (07/141) (ANIC, BPBM, OMBA), Polyrhachis pallescens Mayr, 1876 stat.nov. Polyrhachis guerini var. pallescens Mayr, 1876.74, Type locality: Old, Rockhampton (23/150) (type presumed lost). Polyrhachis aurea var. pallesceny Mayr, Emery, 1897:584, Palyrhachis aurea vat. depilis Emery, |1897:589 (foat- note), Type locality; Queensland (syatype ¢x- amined, MHNG). syn,nov, The unique holotype of P. guerini pallescens cannot be located in the Mayr collection (NHMW) and must be presumed to have been lost. Syntypes of P. aurea (see above under P. ab- fusa) and P. aurea depilis, have been compared and matched against the ANIC nomenclatural paradigms of P. guerini (established above), and other material. We conclude that P. aurea, P. guerint and P. depilis are almost certainly separate Species, The mesosomal dorsum of P. aurea is covered with dense, golden pubescence and abundant, ANT GENUS POLYRHACHIS long, somewhat undulating hairs. The pubes- cence of the gaster is Very dense and golden, often with u distinct reddish tint mid-dorsally. The mesosomal dorsum of P, depilis has relative- ly sparse, whitish appressed pubesecnee, with fewer scattered hairs than in aurea (\hese ate lacking in some of the specimens cxamined, apparently due to wear in life). The relatively long, appressed pubescence on the gaster is sil- very-white, wilh at most only a very slight yel- lowish tint. The original descriptions of depilis and palles- cens show these taxa to be closely similar, and we conclude with reasonable confidence that they are conspecific. P. pallescens is evidently not common. It is known from the types, col- lected in central coastal Old, and from a single additional series taken further south, from wal- lum community (a coastal heathland formation with dominant Banksia aemula, which extends from the Tropic of Capricorn south to northern NSW) at Lake Cootharaba (26/153) (near Boreen Point, SEQ, 13.x1.1976, RIK ace, 76.86). Polyrhachis phryne Forel, 1907 Polyrhachis phryae Forel, 1907:41. Type locality: NSW, Mt Victoria (33/150), Polyrhachis sidnica vat. perthensis Crawley, 1922:30, Type locality; WA,. Perth (31/115). syn.noy Taylor (1989) discussed affinities between the eastern Australian P. phryne (= P. sempronur Forel) and the southwestern P. sidnica perthen- sés, and raised perthensis to species rank. Sub- sequent examination of ANIC material from bulk spirit storage, and consideration of pre- viously unstudied QMBA, Western Australian Museum and South Australian Museum specimens, along with the ecological voucher material of Dr P.J.M. Greenslade (CSIRO Division of Soils and ANIC), shows that these taxa should now be considered conspecific, P. phryne is ane of the most widespread of all southern Australian ants. [tis known frome. 100 km N of Geraldton (28/114), Western Australia, southcastwards to the Great Australian Bight, across the southern part of the continent to southeastern South Australia and Victoria, thence northwards along the Great Dividing range and its eastern flanks to the Mackay district (21/148,149), NQ. An apparently outlying population is present further north at Forty Mile Scrub (18/144), where the species has been taken in deciduous monsoon tainforest. P. phryne ts S17 unrepresented in available collections from Tas- mania, though its presence there seems likely. P. phryne is generally collected from nests in the soil under stones, or taken abroad at night. usually in dry scilcrophyll forests or woodlands, including mallee and subalpine woodland. It is commonly encountered foraging nocturnally on tree trunks in tall mallee near Poochera (32/134), SA, at well studied sites where Nothomyrmecia macrops Clark is present; it has only rarely been seen abroad there during daytime; nests are ex. cavated in very fine calcareous soil, often without covering stones, P. phryne is not a particularly variable species when compared to some others in ‘Campomyrma’. Detailed examination of scores of mounted specimens from all parts of the known range has not revealed churacters imply- ing the presence of more than a single species, and there is no clear evidence of consistent geographical variation. Relevant specimens have been examined from at or near the following localities: QUEENSLAND: Forty Mile Serut (18/144); Britton Range, Eungella Nat Fk, Finch Hatton Gorge (21/148); Blackdown Tableland (23/149), Cammoy Caves (23/150); Rundle Range (23/151), Biloela (24/150); Kroombit Tops (24/151); Bunya Mountains (26/151); D’Aguilar Range, Mt Coot-tha (27/152); Cedar Creek Nat Pk, North ‘Tamborine, Tamborine Mountain, Thunderbird Park (27/153); Goondiwindi (28/150); Fletcher (28/151); Mt Devlin (28/152); Binna Burra Lodge, Lamington Nat Pk (28/153). NEW SoutTy Watés: Lismore, Mt Nullum (28/153); Ar- midule, Ben Lomond Hill, Moonbi Range (30/151); Wauchope (31/152); Bulga, Fassilern, Heaton State Forest (32/1/51); Bulahdelah (42/152); Hillston (33/145); Grenfell (23/148): Blaxland, Kanangra Creek, Mount Victoria (33/150); Bucketty, Lane Cove, Swunsew (33/151): Griffith (34/142): Eusten (34/143); Tabbita (34/145); Boorowa, Cootamundra, Yass (34/144), Gunning, Wombeyan Caves (34/149); MI Flora (34/150); Como (34/151); Finley (35/145); Tumut (35/148); Bungendore, Jerrabomberra Hill, Queanbeyan (35/149); Rosedale Beach (35/150, Khancoban (36/148): Tathra (36/149), AUSTRALIAN CAMTAL TERRITORY: Cotter Dam, Gibraltar Falls, Or- roral Crossing, Uriatra Forest (35/148); Black Moun- tain, Mt Ainslic, Mt Majuta, Mt Pleasant, Stirling Park, Tharwa, Yarralumla (35/149), VicTorRun: Linga (35/141); Nyah (35/143); Kiata (36/141), Heathcote (36/144); Ovens Valley (36/146); Chewton, El- phinstone (37/144), Sour AUSTRALIA; Nullarbor (31/130), 100 mi W of Penong (41/132): Wilpena Pound (31/138); Candada, Ponchera, Streaky Bay (32/134), Alligator Creek (32/138); Koonamore (32/139); Flinders [stand (33/144); Arno Bay, Cur tinye Hill, Kintba (33/136); Blyth, Jamestown, Leighton, Sevenhill (33/138); Worlds End (33/139), Boston Bay. Part Lincoln (34/135); Wurouka (34/137). Port Parham (34/138), Cambrai, Morgan (34/139), Cape Borda, Sandy Creek Buy, West Bay (35/136); D’ Estrees Bay, Marion Bay (35/137): Belair (35/138); Murray Bridge (35/139). WESTERN AUSTRALIA: Billubong Roadhouse (26/1 14); Mullewa (28/115); Dongira (29/114); lo0km S of Eneabba (30/115); Bungalbin Hill (30/119); Karonie (30/122); Perth (41/115), Coomalling (31/116); Merredin (A/S); Madura (31/127); Euelia(3 1/128); Armadale (32/116); Narrogin (32/117); 120km W of Balladonia (32/122); Booyana Rock (32/123); Cacklebiddy (32/126); Ongerup, Pingrup (33/118); Newdepate (33/119); Esperance, Telegraph Till (33/121); Goora Rock, Jununa Rock, Mt Ragged, Thomas River (33/123), Mt Barker (34/117); MUTn0 (34/118), Salis~ bury Island (34/123). Polyrhachis pilosa Donisthorpe, 1938 stat.nov, Polyrhachis rastellata vr. levior var. pilosa Forel, 1902:527, workers trom author's study series ex- amined, MHNG, NHRS, ANIC (an inadmissible infrasubspecific name), Polyrhachts rastellata var, pilosa Donisthorpe, 1938;256 The epithet pilosa was introduced by Forel (1902) as the fourth name in a quadrinomen. It was later excluded, as an infrasubspecilic name (Taylor, 1986), from the catalogues of Taylor und Brown (1985) and Taylor (1987). These actions overlooked Donisthorpe’s 1938 nomenclatural act, in which ‘pilosa’ was used in a trinominal context for a subspecies, with clear reference to the carlier Fore! name. As uv result, authorship of the available species-group name pilasa in Polyrhachis must be formally attributed to Domisthorpe, under the Iaiernational Code of Zoological Nomenclature, and the year of publi- cation for purposes of priority is 1938. We take existing specimens from Forel's 1902 series to be the types of PF. rastellata pilosa Donisthorpe, 1938, The subgenus Cyriamyrma, to which this {axon isussigned, has a number of Australian and New Guinean species, and the differences separating them can be subtle. Nonetheless, we recognise ?. pilosa as a good species, in con- sideration of ils very distinctive habitus. It uni- quely possesses the following combination of MEMOIRS OF TRE QUEENSLAND MUSEUM characters: pronotal humeri evenly rounded in dorsal view, lacking projections; propodeum without spines or other armament; and (most distinctively) entire dorsal surfaces of mesosoma and gaster with numerous long, erect hairs. The only other Known (apparently undescribed) Australian species with long gastral hairs occurs in coastal areas centred on Cairns (16/145), NQ, but it hus distinct, relatively long propodeal spines, We have seen 2. pilosa in Qld from Mt Ossa, NW of Mackay (20/148), Rundle Range, nr Gladstone (23/150); several sites near Bun- Jaberg (24/152): Seary Scrub, Cooloola (25/153): Blackall Range (26/152); Peregian Beach (26/153). and in NSW from near Lismore (28/153) and Taree (31/152), 1 has been cal- lected most frequently towards the southern limits of this range. [tis a relatively common ant in parks and suburban gardens in Brisbane (27/153). The (ypes are putatively from Wol- longbar (28/153) and Richmond River (29/153), NSW (Forel, 1902), and this accords with the known distribution, However, two syntypes (ANIC, donated by MHNG) on a single pin labelled ‘COTYPUS? carry a Forel autograph label reading ‘fevior var pilosa, Mackay, Turner, no 7 Polyrhachis reclinata Emery, 1887 stat.nov. Polyrhachis sexspinasa var. reelinata Emery, 1887:236. Type locality: New Guinea, Fly River (O7/141) (3 syntypes examined, MCSN). Polyrhachis sexspinesa (Latreille, 1802); Bolton, 1975:12 (in part). P. sexspinosa reclinata syntypes have been compared with specimens identified by Bolton (1975) as P. sexspinesa, and with the ANIC nomenclatural paradigm of sexspinasa (see below). We consider reclinata to represent a Species separate from sexspinosa. The head in sexspinosa is Strongly tapered immediately be- hind the eyes, and the somewhat Mange-like oe- cipital margin forms prominent, well developed lateral lobes, which are usually clearly visible in full face view. In addition, the front of the head behind the eyes is coarsely rugose, with rather dilute, mostly while or silvery pubescence. In the P. reclinata \ypes the head is generally shorter behind the eyes and less strongly tapered, and the lateral occipital lobes are relatively weakly developed und barcly visible in full face view; the sculpturing on the back of the frons is a very ANT GENUS POLYRHACHIS faint, somewhat polished, fine puncturation, which is usually largely masked by rich, brassy, appressed pubescence. The propodeal spines in sexspinosa are generally vertical relative to the main axis of the body, or even inclined forwards when viewed from the side. In reclinata they are posteriorly inclined, more curved in side view, and somewhat more divergent in dorsal view. P. sexspinosa is relatively common and widespread on the mainland of New Guinea (Grid cells 03/135, 03/142, 05/145, 07/146, 08/147), but P. reclinata is known only from the savannas of the Port Moresby area, and from the Fly River delta in the south of the island, in areas gencrally opposite the tip of Cape York Penin- sula (Grid cells 08/143, 09/143, 09/147). In Australia both species are known only from northern Cape York Peninsula, where sexspinosa ranges from Lockerbie Scrub south to Rocky River, ENE of Coen (Grid cells 10/142, 12/143, 13/143), and reclinata has been taken only at Iron Range (12/143) (RJK acces 81.185, 209, 210), where it nests in the hollow internodes of the bamboo Bambusa forbesti. Bamboo-inter- node nesting is common to most Australian species of the sexspinosa-group. The exceptions are P. sexspinosa itself, which builds camouflaged pockct nests of silk and bark frag- ments attached to the trunks of rainforest trees, and P. glabrinota, which uscs silk to build nests by joining the Icaves of various rainforest trees and shrubs. There are, incidentally, 5 species of the sexspinosa-group present on N Cape York Peninsula (P. reclinata, P. sexspinosa, P. glabrinota and 2 species yet to be described). Polyrhachis schoopae Forel, 1902 stat.nov. Polyrhachis appendiculata 1. schoopae Forel, 1902:520. Type locality: Qld, Mackay (21/149) (8 syntypes examined, MHNG, ANIC). The P. appendiculata and P. appendiculata schoopae syntypes, and specimens considered conspecific with each, are consistently separable. We consider them to represent separate species. P. appendiculata is generally smaller than P. schoopae, with the relatively fine sculpturing of the promesonotal dorsum partly to largely obscured by silvery pubescence. The pronotal dorsum in profile is more strongly con- vex than in schoopae, and the mesonotal and propodeal dorsa almost straight. P. schoopae is relatively large, with the promesonotal dorsum mostly longitudinally striate and shining, with 519 appressed pubescence virtually lacking. The en- tire mesosomal dorsum in profile forms a single, strongly convex, almost semicircular arc. The known range of P. schoopae is from Cape Tribulation, Qld, south to Mackay (Grid cells 16/145, 20/148, 20/149, 21/148, 21/149). P. ap- pendiculata is known only from the Torres Strait islands (Yam I., Stephens I., Murray I.) (Grid cells 09/142, 09/143, 09/144), except for a single record from mid Cape York Peninsula (Jane Table Hill, Princess Charlotte Bay, 28 June 1980, RJK acc. 80.47) (Grid cell 14/144). Polyrhachis semiobscura Donisthorpe, 1944 Polyrhachis (Hagiomyrma) semiobscura Donis- thorpe, 1944:65. Type locality: New Ireland, Kavieng (02/150) (holotype examined, BMNH). P. semiobscura has been previously reported only from mainland PNG, and adjacent islands. It was recently collected for the first time in Australia, on Cape York Peninsula, Qld, at Bamaga (10/142) (RJK acc. 87.76) and Iron Range (12/143) (RJK accs 81.176, 179, 203). A colony at Iron Range occupied two adjacent woody galls attached to the thin branch of a low shrub in open forest. Melanesian records are from at or near the following localities: Bubia, nr Lae (06/146); Bulolo (07/146); and nr Sogeri (09/147), This is an open forest and savannah-in- habiting species. Polyrhachis sexspinosa (Latreille, 1802) Formica sexspinosa Latreille, 1802:126, pl.4,fig.21. Type locality: East Indies (type presumed lost). Polyrhachis (Myrmhopla) barnardi Clark, 1928:39, pl.l,figs 37,38. Type locality: Qld, Cape York (10/142) (syntypes examined, MVMA, ANIC, MCZC). syn.nov, Five nominal specimens (all labelled ‘Type’, one on a red tag, the others on blue tags) of P. barnardi have been examined, and the series is considered to include 2 distinct species. Three specimens match Clark’s original description and illustrations, and these we take authentically to represent P. barnardi. A lectotype (the specimen bearing a red tag, MVMA) has been selected, and the other 2 specimens (MVMA) are designated paralectotypes. Further comparison of these specimens with others identified as P. sexspinosa by Bolton (1975), shows sexspinosa and barnardi (as restricted here) to be con- 320 specific. The 2 remaining original barnardi syn- types (ANIC, MCZC, the last examined by Bol- ton, 1975) are to be included in the type series of a new species for description by RIK. P. sexspinosa was originally described from a queen collected in the “East Indics’. Like Bolton (1975) we have been unable to locate the type. In lieu we recognise a Specimen in the ANIC from a long series collected at Pes Mission nr Aitape (03/142), West Sepik Province, PNG (RJK ace. 84.207) as the nomenclatural paradigm of P. sexspinosa. Designation of a neotype would not be appropriate here. Other details are given above under P. reclinata, Polyrhachis yermiculosa Mayr, 1876 statinov, Polyrhachis guerini var, vermiculosa Mayr, 1876:74. Onginal localities: Old, Rockhampton (23/150) and Peak Downs (22/148), NSW, Sydney (as Sid- ney) (33/151), (3 syntype workers, 1 alate queen examined, NHMW, MHNG). Polyrhachis aurea var, vermiculasa Mayr, Emery, [KYTSR4, The compared syntypes of P. guerini ver- miculosa and P. aurea (specimen details given under P. pallescens) are similar but, following examination of other relevant specimens, we consider them 0 represent separate specics. P, aurea has the mesosoma strongly narrowed posteriorly, with the dorsum of the propodeum distinctly mdented across the bases of the propodeal spines; the sculpturing of its pronotal dorsum consists of fine, more or less generally Jongitudinal, fragmented rugulation. P. ver- miculasa is more stoutly built, with the propodeal dorsum at the base of the spines rela- lively Wide, and the pronotal dorsum irregularly and rather vermiculosely rugose. The antennal scapes ure relatively long in aurea, with Scape Index (sce above under P. obtusa) >125, versus <115 in vermiculosa. We have seen P, vermiculosa specimens from many localities, ranging from Lakefield on Cape York Peninsula, Old, to just south of Brisbane (Grid cells 14/144, 20/148, 21/148, 22/149, 23/150, 26/153, 27/152, 27/153, 28/152). This species inhabits open forest and woodland, and nests mostly in the soil between grass roots, or under suituble covering objects, such as stones or logs. MEMOIRS OF THE QUEENSLAND MUSEUM Polyrhachis yarrabahensis Forel, 1915 stal.nov. Polyrhachis (Myrmatopa) lombokensis var. yar- rabahensis Forel, 1915:115. Type locality; Qld, Malanda (17/145) (for ihe queen), Yarrabah (16/145) (for the worker) (syntype workers ex- amined, ANIC). Workersyntypes of P. lombokensis yarrabahen- sis have been compared to a syntype of P. lom- bokensis (Emery, 1898; 239; worker (MCSN); Indoncsia, Lombok). Withsomce hesitation we con- sider them lo represent separate species. We are hesitant to synonymize the names because of the great distance separating the known ranges of these faxa and because no material is known from areas in between, One syntype was unfortunately the only specimen of P. lambokensis available to us, and we have been unable to match it with any of the numerous Indonesian and Melanesian “Myrmatapa’ specimens we have seen, The char- acters differentiating these taxa are relatively slight, but consistent in the material available, The eyes of ?. yarrabahensis are tather strong- ly convex, extending promimently beyond the outline of the head in frontal view. In P. Jom- bokensis the cyes are Jess strongly convex, and exceed the outline of the head less strongly. The humeral margins of the pronotal dorsum in yar- rabahensis are slightly angled in dorsal view, and the propodeal declivity descends trom the dorsum as a relatively gentle slope (the accurate- ly goniometer-measured angle between the basal and declivitous propodeal faces in side view ranges fram 37 to 45° in 10 yarrabahensis specimens examined). The humeti in lomboken- sis are broadly and smoothly rounded in dorsal view, and the propodeal declivity descends more abruptly in the available syntype, at an angle of 50°. P. yarrabaheasis is an arboreal species which uses silk to build complex, often polydomous nests by joining together the leaves of various lowland rainforest trees, shrubs and vines. Known records (NQ) are from Massy Spur Creek, near Silver Plains (13/143) and from Kamerunga and Yarrabah, in the Cairms area (16/145). We would expect that this species is limited to altitudes below about 350 m in the coastal strip of the Base-of-Peninsula area (as are many ant species of Papuasian affinity, including jhe prominent green weaver ant Oecophylla ANT GENUS POLYRRACHIS smaragdina (Fabricius)), and that the alate type- queen from Malanda had probably flown to elevations beyond the usual nesting range. Polyrhachis yorkana Forel, 1915 stat.nov. Patyrhachis (Cyrtomyrma) rastellaia yar, yorkana Forel. 1915:110. Type locality: Old, Cape York Feninsula (syatypes examined, ANIC), P. yarkana is the oldest available name based on Australian type-material that is applicable to a widespread species (or complex of species) represented throughoul the range of the subgenus “Cyriomyrma’ in eastern Australia (i.c. from Cupe York south to near 30°S in coastal eastern NSW). These ants are relatively nondescript, lacking any of the features discriminating more easily characterised Cyriamyrma species (such u8 propodeal spines, abundant gastral or mcsosomal/gustral pilosity, dense body pubcs-: cence), except for a tuft of fewer than 10 long hairs on the summit of the strongly arched mesosoma. Similar ants are present in SE Asia and New Guinea, so that senior names based on foreign specimens could be applicable to the Australian species, All available names of the specics-group have identical status in nomenclature regardless of their rank, and the assigning of species rank to the names of unresolved taxa of the species- proup seems the best course to follow. For these reasons we here attribute full specific rank to P. yvorkana. We suspect that it will survive as a good species following revision of Polyrhachis. ACKNOWLEDGEMENTS We thank Philip S. Ward and Kazuo Ogata for useful comments, and the institutions and curators listed in the introduction for the loan of lypes and other material in their care. Sound taxonomic work would not be possible in Australia without such collaboration, and We are deeply appreciative of their help. Much of the matcrial pertinent to the project of which this paper is a part was collected by Rev. B.B. Lowery SJ, or Geoff B. Monteith. We take this opportunity especially to acknowledge (heir merils 4s ant collectors. LITERATURE CITED ANDERSEN, A.N, 1986a. Diversity, seasonality and community organization of ants al adjacent $21 heath and woodland sites in south-eastern Australia, Australian Journal of Zoology 34: 53-64. 1986b. Patterns of ant community organization in mesic southeastern Australia. Australian Jour- nal of Ecology (1: 87-97. 1988) Immediate and longer-term effects of fire on sced predation by ants in scleraphyllous vegeta- lion in south-eastern Australia. 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Ber- liner entomologische Zeitschrift 7: 129-214. SMITH, F. 1859. Catalogue of hymenopterous insects collected by Mr A.R. Wallace at the islands of Aru and Key. Journal of the Linnean Society. Zoology 3:132-178. 1860. Catalogue of hymenopterous insects col- lected by Mr A.R. Wallace in the islands of Bachian, Kaisaa, Amboyna, Gilolo, and at Dory in New Guinea. Journal of the Linnean Society. Zoology 4 Suppl.: 93-143,pl.1. TAYLoR, R.W. 1986. The quadrinominal in- frasubspecific names of Australian ants (Hymenoptera: Formicidae). General and Ap- plied Entomology 18: 33-37. 1987. A Checklist of the Ants of Australia, New Caledonia and New Zealand (Hymenoptera: Formicidae). CSIRO Australia. Division of En- tomology Report 41: 1-92. 1989, The nomenclature and distribution of some Australian ants of the genus Polyrhachis Fr.Smith (Hymenoptera: Formicidae: For- micinae). Journal of the Australian entomologi- cal Society 28(1): 23-27. TAYLOR, R.W. AND BROWN, D.R. 1985. Hymenop- tera: Formicoidea in Zoological Catalogue of Australia 2: 1-149, 306-348. WHEELER, W.M. 1922. Ants of the Belgian Congo. Part 7. Keys to the genera and subgenera of Ants. Bulletin of the American Museum of Natural History 45: 631-710. A NEW CYRTOMAIA, C. GRIFFINI, FROM AUSTRALIA (CRUSTACEA: DECAPODA: BRACHYURA) BERTRAND RICHER DE FORGES AND DANIELE GUINOT Richer de Forges, B. and Guinot, D. 1990 08 31: A new Cyrtomaia, C. griffini, from Australia (Crustacea: Decapoda: Brachyura). Memoirs of the Queensland Museum 28(2): §23-530. Brisbane. ISSN 0079-8835. A new species of crab of the genus Cyrtomaia is described from deepwater off the eastern coast of Australia: C. griffini sp. nov. This species is compared with its closest relatives, C. suhmii and C. curviceros, and remarks on the geographic distribution of the six species in this genus with large protogastric spines are given. Une nouvelle espéce de crabe de profondeur du genre Cyrtomaia est décrite de la céte est de |’ Australie: C. griffini sp. nov. Aprés la description de l’espéce et une comparaison avec les espéce proches, C. suhmii et C. curviceros, des remarques sur la distribution géographique des six espéces de ce genre a grandes épines protogastriques sont faites. (1Crustacea, Decapoda, Brachyura, Cyrtomaia, new species. Bertrand Richer de Forges, Australian Institute of Marine Science, PMB No.3, Townsville M.C., Queensland 4810, Australia; Daniéle Guinot, Muséum National d'Histoire Naturelle, 61 rue de Buffon, Paris cedex 05, France; 14 August,1989. In earlier works (Guinot and Richer de Forges, 1982a: 1097, under Cyrtomaia gaillardi; 1982b: 24, under C. suhmi; 1986: 119, under C. suhmi), we raised the question of the identity of some Australian material that was attributed to Cyr- tomaia suhmii' Miers, 1886 (p. 18, pl. 3, fig. 2, 2a-2c) by Griffin and Brown (1976: 252, fig. 6). All the specimens from the East Australian coast have a strong intercalated orbital spine but on the holotype of C. suhmii Miers (from the Tulur islands to the north of the Molucca Sea) the supra-orbital edge bears only a small granule. The holotype was examined and this feature figured by Guinot and Richer de Forges (1982b, fig. 10, 11a-b). Another species, Cyrtomaia cur- viceros Bouvier, 1915 (pp. 9-15, pl. 1), originally described from Japan and similar to C. suhmii (cf. Guinot and Richer de Forges, 1986: 118, fig. lla, under C. suhmi) is typified by a completely smooth, inermous supra-orbital edge. In their important work on the Majidae gathered by the Siboga, Griffin and Tranter (1986a: 30, 31) put C. curviceros Bouvier in synonymy with C. suh- mii and questioned the taxonomic value of the intercalated orbital spine: ‘The spine in the orbit of Cyrtomaia and similarly Platymaia doesn’t constitute a more useful distinguishing feature than the other small carapace spines’. When they wrote this, the two Australians were not aware of 1 The specific spelling of Miers’ species should in fact be C. suhmii to respect the original formulation, as Davie and Short (1989) have pointed out. our work, published the same year (Guinot and Richer de Forges, 1986), in which we reported the discovery of the Philippines topotypical Cyr- tomaia suhmii. In all our material, juvenile and adult at various stages, from the Philippines, the supra-orbital edge uniformly bears a small, but distinct, intercalated granule. The fine fresh material collected during the Cidaris I cruise and also found by the vessels Soela, Southern! ntruder and Iron Summer on the east Australian coasts, does not belong either to C. suhmii or to C. curviceros, and the description of a new species, C. griffini sp. nov., is therefore required. List of abreviations: MP, Museum national d’Histoire naturelle, Paris (B - Brachyoures); QM, Queensland Museum. Length measure- ments exclude pseudorostral spines. Cyrtomaia griffini sp. nov. (Figs 1, 2, 3,4) Cyrtomaia suhmi: Griffin and Brown (not Miers, 1886), 1976: 252, fig. 6. Griffin and Tranter, 1986a: 30, fig. 91g. not Cyrtomaia suhmi: Griffin and Tranter, 1986b: 351, fig. 1. Cyrtomaia suhmii: Davie and Short, 1989: 27. MATERIAL EXAMINED HOLOTYPE: QM W15362, 6 (78.8 x 83.3mm), Soela 524 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 1. Cyrtomaia griffini sp. nov., paratype 2 (55 x 60.6mm), R.V. Franklin, Cidaris I, Stn 47-3, 505m, 17°51.35'S—143°07.83’E, 17.v.1986, beam trawl (QM W16067). A, dorsal view of fronto-orbital region. B, ventral view. C, D, detail of the antenna from two different angles. NEW CYRTOMAIA FROM AUSTRALIA Cro, stn. 79, 6x11 1985, 800m, 17°O1'S, 151°20'E, P, Davie. Pagaryies: MP-B 20555, 2 (58x 65mm),Soela Cro, stn, 78, 6.xi) 1985, 880m, 16°55'S- 151°34'B, P, Davie. OM W15360, 62 (57.4 x 62,,58.8 x 65.6, 61.9 x 65.7, 55.5 x 58.6, 60.5 x 65.7, 56.1 x 60.6), 50 (75.9 x B18, 46.1 x 50.1, 47.8 x 52.6, 47.9 x 52.2, 48.5 x 50.7mm), Cr6, Stn 78, 6.xii,1985, 880m, 16°55'S, 151°34'E, P. Davie, OM W16067, 2 (55 x 60.6mm), Cidaris 1, Sta 47-3, 15.¥.1986, 17°51,35'S, 143°07,83'E, 505m. QM W16068, ovig. 2 (50.5 x 63mm), Cidarix J, Sm 48-3, 17.v, 1984, 17°52,00'S, 147°08,00'R, 740-680m, OM W16069, 2 (43.8 x 495m), Cidaris 1, Stn 1-4, 6.v.1986, 18°08.69'S, 147°33,97'E, 966m, OTHER MATERIAL: R.V. Soela: QM W15373, 6 @ (40.2 x 42.2, 40.6 x 43, 25.5 x 27.4, 24.4 x 25,7, 26.8 x 28,2, 24.4 x 24.6mm), | juv, d (20 x 20.8mm), Cr6, Sty 78, 6,¥.1985, 880m, 16°55’S, 197°14'E, P. Davie. QM W15375, 36 (27.2 x 27, 20.3 x 20.9, 17,3 x 17.9mm), Cr6, Sin 79, 6.xii.1985, 800m, 17°01'S, 151°20'E, QM W15361, 1¢ (74,2 » 78.9mm), | crushed specimen (74mm width), Cré, Stn 80, 6.8iL 1985, 700m, 17°02'S, 151°03'E, P. Davie. QM W15932, crushed juv. od (20.8 x 21. 5mmj, Cré, Sin 59, 2.xii.1985, 900-908m, 17°30'S, L49°00'F, P-. Davie. M.Y, fron Summer: QM W14908, 2d (60.2 % 62.7, 58.1 x 62.1mm), Shot 2, 10.v.1983,600m, 28°19,91'S, 153°53.47'E. OM W10611, 1 2 (64.9x 72.5mm), Shot 1, 3.x.1982, 500-54()m, 27°22'S, 153°E, M. Holmes. QM W10612, 1d (61.3 x 66.3mm), Shot 6, 25,iii.1983, 520m, 27°13'S, 153°22'E, R. Morton. OM W10609, 10 (68.4 x 73.2mm), Shot 5, 13.viii 1982, 54m, 27°18'S, 193°S4'E, G, Smith and J. Burke. OM W14909, 1d’ (63.2 x 61.9mm), Shot I, 10.v.1983, 590m, 27°13,00'S, 153°52.53'E, R. Morton, OM W14916, 12 (63.5 x 72.7mm), Shot 3, 10.¥.1983, 555m, 27°12.83'S, 153°52.87'E, R. Morton. OM W10610, 12 (67x 75mm), 1d (62.3x 66.9mm), Shot 4. 10.v.1983, 540m, 27°34'S, 153°56’E, R, Morton. MLV, Southern Intruder, OM W)1233, 12 (64.4 x 43.8mm), Shot 40, 30.xii.1983, 460m, 23°17'S, 152°56’E, P. Davie. QM W11232, 19 (60.6 x 46.2mm), Shot 37, 29.xii.1983, 550m, 23°45'S, 1535°07'E, P. Davie.. DESCRIPTION A large specics, that may measure as much as 80mm across the cephalothorax. Carapace is smooth and shiny, with a few bristles in front of the protogastric spines. Protogastric spines dis- tinctly longer than the others, in the form of arallel horns, pointing forward and very slightly incurved; the spines appear smooth to the naked wn tw “ eye and to the touch, but in fact are very finely granular. A single, pointed gastric spine with a very slight granule in front. Two pointed and raised cardiac spines. Very sharp, slanting anterior branchial spine, similar in size to the single gastric spine. A very short, but also sharp, posterior branchial spine, with a tubercle behind it. A branchial spine, pointed, but shorter than the abovementioned. On the latero-posterior edge of the branchial area is a line of five spinules, The intestinal area is almost imperceptible. Long pointed pseudo- rostral spines, subparallel on the paratype (Fig. 1A) but diverging on other specimens (Fig. 4A); the pointed rostral spine extends from the front; there is a broad supraocular hood with smooth edges. A very distinct intercalated orbital spine, fine and sharp, pointing obliquely towards the stocular spine. Strong postecular spine, point- ing forwards (Fig, 4B), Dorsal facet with marked relief; clear branchio-cardiac groaves (Fig. 4A), very fine and diffuse granulation, visible only under a binocular microscope, Short ocular peduncle with a single horn, (Fig, 1A), Antennal basal article with three large spines, plus a small one on the lower facet (Fig- 1C). The meri of P4 and P5 are smooth, as are the carpi and propodi; in the female, the P2 merus is distinctly longer than that on P1; in the male, on the other hand, the P2 merus is much shorter than that of P1, and the P1 propodus is broader. The live animal is orange-pink all over; the eggs of the ovigerous female are navy blue. DISCUSSION The differences between Cyrtamaia griffin sp. nov. and C. sukmii Miers, 1886 are as follows: an intercalated orbital spine is not found in C. sufynit but is found in C. griffini, both in males and in females, whatever their size; (he number of spines on the antennal basal article is 3 in C, griffini and 4 or S in C. suAmii. The study of numerous species of the genus Cyrtomaia shows. a stability in the shape and spinulation of the supra-orbital edge, The presence of an intercalated orbital spine con- stitules a constant character whatever the size or sex. Guinot and Richer de Forges (1986: 117-19, fig. 11a) reported on many specimens of C. suh- mit {rom the Philippines Islands and confirmed that, as on the holotype, the supra-orbital edge was alWays smooth, Cyrtomaia curviceros Bouvier, 1915, is another lutge species wilh very MEMOIRS OF THE QUEENSLAND MUSEUM “pINUsOD “QL ‘syo10s ‘Dy ladpiapood “DEH ‘ipavjjivd “De affiaS “DH NyDO][NIINUE “DW sso4ao1ANd *D pue MWYNS “DE ‘9QRT ‘SATA Mayns “-D Oj parejas DIBWUO4D JO saroads yo dnosd ay) jo uonnqiisiq *Z “Old YA Yr Ss ‘et MME NEW CYRTOMAIA FROM AUSTRALIA Fic, 3. Cyrtomaia griffini sp. nov., holotype d (78.8 x 83.3mm), Queensland, Soela, Cr.6, Stn 79, 800m, 6.xii.1985, P. Davie (QM W15362), dorsal view. well-developed protogastric spines but is known only from the holotype, a very large specimen from Japan. It also has a smooth supra-orbital edge and this is the reason why some authors think it is a synonym of C. suhmii. It will be necessary to examine topotypical material of C. curviceros before this problem can be finally resolyed. C.curviceros (see Guinot and Richer de Forges, 1982b: 24, fig. 12) like C. suhmit is different from C. griffini by having the supra-or- bital edge smooth and the basal antennular article with five spines. C. gaillardi Guinot and Richer de Forges, 1982, found in the waters of Madagascar can easily be distinguished from C. griffini by its divergent pseudorostral spines, the granular carapace (in C. griffini it is smooth), and the possession of a small intercalated orbital spine (in C. griffini this is large and pointed). The other Australian species, C. maccullochi Rathbun, 1918, from the Great Australian Bight in southern Australia, is smaller than C. griffini. It also has long protogastric spines but has no intercalated orbital spine and its carapace is granular. It would appear that the specimens of C. suhmii reported by Griffin and Tranter (1986b), from the Molucca Passage and from the continental slope of northwestern Australia (Port Hedland) are very different from those of the eastern coast. Indeed, the excellent photographs illustrating this work show that the supra- orbital edge is smooth, with no spine or granule; this species should therefore be related to the C. suAmii - C. curviceros group of species. Itseems, therefore, that of the species that have very long protogastric spines, there are three that inhabit the Australian coasts, C. maecullochi in the south, C. suhmii in the northwest and C. griffini in the east (Fig. 2). One other species of Cyrtomaia, C. horrida also occurs off the east coast, Richer de Forges and Guinot (1988) have just described three new species of the genus Cyr- tomaia, gathered in New Caledonia and in the Chesterfield Islands. Two of them, C. cornuta and C. coriolisi, are relatively large in size and 4) tw ~ 528 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 4. Cyrtomaia griffini sp. nov., holotype & (78.8 x 83.3mm), Queensland, Soela, Cr.6, Stn 79, 800m, 6,xii,1985, P. Davie (QM W15362). A, enlargement of the carapace; B, lateral view. NEW CYRTOMAIA FROM AUSTRALIA the fact that their longest spines are the protogastric spines places them in the same group of species as C. griffini. C. coriolisi, taken at a depth of 650m in the Chesterfield Islands and on the Norfolk Ridge, is very similar to C. griffini, but it is smaller in size; its carapace is finely granular (it is smooth in C, griffini); and its protogastric spines are longer. C. cornuta, found in the south of New Caledonia (at depths of 270 to 535m), has granular legs and carapace, and very long, curved protogastric spines. DISTRIBUTION Northeastern coast of Australia between 35°S and 17°S, on sandy-mud bottoms situated at depths of 360 to 980m. The present geographical distribution of the large species of Cyrtomaia with long protogastric spines (C. suhmii, C. cur- viceros, C. maccullochi, C. griffini, C. gaillardi, C. goodridgei) indicates that they are quite dis- tinct entities (Fig. 2). It appears that the group C. suhmii/C. cur- viceros is limited to the western Pacific and East Indian Ocean zone between Japan and the northwest of Australia, with several spots in the Philippines and the Sunda Islands; it would be impossible for these bathyal species to pass through the Torres Strait on account of the very shallow shelf between Papua New Guinea and northern Australia. These species have not been reported from the Marianas despite all the trap fishing that has been done in this area by L. Eldredge. It would be most interesting to obtain some Cyrtomaia from the Solomon Islands to see if there is a relationship between them and C. suhmii or C. griffini. The species C. maccullochi has been found only in southern Australia, at a very low latitude (33°S). Griffin and Tranter (1986a) quote a fine male in the South China Sea, which they relate, with reservations, to C. maccullochi. The distribution of the species C. goodridgei and C. gaillardi in the Indian Ocean appears very scattered (Guinot, 1985); this is probably be- cause there has been so little sampling. To under- stand the distribution of these species, it would be necessary to collect them from the different types of sea- bed relief (e.g. sea mounts, ridges) which have served as bridges during the geologi- cal eras. The genus Cyrtomaia Miers is typically bathyal (200-2000m), and the geographical spread of the species seems to be limited by the 529 immense abyssal zones; their distribution is thus naturally discontinuous. ETYMOLOGY We dedicate this species to Dr D.J.G. Griffin, who has done a great deal of work on the Majidae and especially on bathyal fauna. ACKNOWLEDGEMENTS Dr D.J.G. Griffin received one of us (Richer de Forges) at the Australian Museum in Sydney and it gives us much pleasure to pay him a tribute in dedicating this species to him. We extend sincere thanks to Mr P. Davie who placed his C. suhmii material at our disposal at the Queensland Museum and who also suggested we describe Cyrtomaia griffini sp. nov. in the Memoirs of the Queensland Museum. We also thank Ms V. Sery for the drawings, J. Rebiére for the photographs and J. Semblat for preparing the manuscript. LITERATURE CITED Bouvier, E.-L. 1915. Etude sur un Cyrtomaia suhmi du Musée de Marseille. Ann. Mus. Hist. Nat. Marseille 15: 9-15, pl. 1. DAVIE, P. AND SHORT, J. 1989. Deepwater Brachyura (Crustacea : Decapoda) from Southern Queensland, Australia; with descriptions of four new species. Mem. Qd Mus, 27(2): 157-187. GRIFFIN, D.J.G. AND BROWN, D.E. 1976. Deep water Decapod Crustacea from Eastern Australia : Brachyura Crabs. Rec. Aus. Mus, 30: 248-271, figs 1-10. GRIFFIN, D.J.G. AND TRANTER, M.A. 1986a. The Decapoda Brachyura of the Siboga Expedition. Part VIII : Majidae; Siboga Expedition Monograph 39 (C4), 148: 1-335, text. figs 1-112, pls 1-22, 1986b. Some Majid Crabs from the deep Indo- west Pacific. Rec. Aus. Mus. 38: 351-371, figs 1-14, Guinot, D. 1985. Crabes bathyaux de l’ile de la réunion; description de Cyrtomaia guillei sp. nov., de Platypilumnus inermis sp. nov. et de Psopheticus vocans sp. nov. (Crustacea, Decapoda, Brachyura) CNFRA, no 55, 1984 (1985): 7-31; figs 1-5, 1 fig. n.n., pl. 1-4. GuInoT, D. AND RICHER DE FORGES, B. 1982a. Nouvelles récoltes des genres Cyriomaia Miers et Pleistacantha Miers (Crustacea, Decapoda, Brachyura). Bull. Mus. natn. Hist. nat., Paris 530 MEMOIRS OF THE QUEENSLAND MUSEUM nat., sér. A. Zool., 1985(1986), 133: 83-178, figs 1-21, pls 1-11. MIERS, E.J. 1886. Report on the Brachyura collected tomaia Miers, 1886 : campagnes océano- by H.M.S. “Challenger” during the years 1873- graphiques du “Challenger”, de I’"Albatross", 76. Rep. Sci. Res. Voy.‘Challenger’, Zool. du “Siboga” et du “Vauban” (Crustacea, 17(2):i-1, 1-362, pls 1-29. Decapoda, Brachyura). Annies Inst. océanogr., RICHER DE FORGES, B., AND GUINOT, D. 1988. Monaco 58(1): 5-88, figs 1-55, 1 tabl. Description de trois espéces de Cyrtomaia 1986. Crustacés Décapodes : Majidae (genres Miers, 1886, de Nouvelle Calédonie et des iles Platymaia, Cyrtomaia, Pleistacantha, Spheno- Chesterfield (Crustacea, Decapoda, Brachyura). carcinus et Naxioides) In Rés. Camp. Bull. Mus. natn. Hist. nat., Paris,(4), 10, sect. MusorstToo I et II. 2, 4. Mém. Mus. natn. Hist. A(1): 39-55, figs 1-4, pls 1-2. (4),3, 1981 (1982), sect. A (4): 1087-1124, figs 1-8, pls 1-4. 1982b. Révision du genre indo-pacifique Cyr- A REVISION OF THE AUSTRALIAN PARCOBLATTINI (BLATTARIA: BLATTELLIDAE: BLATTELLINAE) LOUTS M. ROTH Roth, L.M. 1990 08 31: A revision of the Australian Parcublattini (Blattaria: Blattellidae: Blatiellinae), Memoirs of the Queensland Museum, 28(2)) 531-596, ISSN 0079-8835, Diagnostic characters and descriptions of species helonging to 4 genera of Australian Blattellinae, tribe Parcoblattini are presented. Gislenia Princis issynonymised with Neorem- nopterys Princis. Shawella Princis and Franwalkeria Princis are synonymised with Paratemnopteryx Saussure. Neotemnopteryx ferruginea (Tepper), Gislenia apicalix (Walker), and Gislenia australica (Brunner) are synonymised with Neotemnopteryx fiulva (Saussure), Paratemnopteryx blattoides Tepper is synonymised with P. couloniana (Saus- sure), P. 2evfzi Tepper is synonymised with P, rufa (Tepper). Symploce centralensis Roth, previously known only from New Guinea, occurs in Australia, and is transferred to Parateninopteryx. There are LO species of Neotemnopteryx, 3 of which are new combinations; fulva (Saussure), australis (Saussure), and douglasi (Princis); the remaining 7 are new taxa: gloriousa, bifurcata, glossa, concava, elliptica, nana and styliparedra, A neotype is designated for Neatemnopteryx fulva (Saussure). Paratemnopteryx contains 10 taxa of which 3 were correctly placed; australis (Saussure), rufa (Tepper), and afra Princis: 3 are new combina- tions! couloniana (Saussure), glauerti (Princis), centralensis (Roth): 4 are new taxa stortel, howarthi, broomehillensis and suffuscula. One species af Neotemropteryx and 3 species of Poratemnopteryx ate cavernicolous. Trogloblatiella Mackerras contains 1 Australian cavernicolous species, nullarborensis. The new genus Keyella comains 2 new taxa, gaye and armidalensis. Keys are provided to separate the genera and to distinguish between males Of the species, DBlattaria, Blattellidae, Parcoblatiini, Australia, cockroaches, cave insects, Louis M. Roth, Museum of Comparative Zoology, Harvard University, Cambridge, MA, 02138, USA; Correspondence; P.O, Box 540, Sherbarn, MA, 01770, USA; 25 March, 1989, According to Rentz and Cameron (1983) the Blattellidac of Australia are among the poorest Rydalmere, New South Wales, Australia, Dr G.R. Brown; MCZH - Museum of Comparative known cockroaches found on that continent, They suggest that the specics belonging to this family may be twice the number actually recorded, From my studics thus far, they may be lov conservative in their estimate of undescribed taxa. This paper includes the following 7 closely related blattellid genera belonging to the tribe Parcoblatlini: Neotemnopleryx (including Gis- lenia), Trogloblattella, Paratemnopteryx (in- cluding Shawella and Franwalkeria) and Keyella, The specimens were louned to me by the following museums and their curators or collec- tion managers: ANIC - Australian National In- sect Collection, CSIRO, Canberra, ACT, Australia; Dr D.C.F. Rentz and Mr John Balder- son; ANSP - Academy of Natural Sciences of Philadelphia, Philadelphia, USA, Mr Donald Azuma; BMNH - British Muscum (Natural His- tory), London, England, Mrs Judith Marshall, BPBM - Bernice P, Bishop Museum, Honolulu, Hawaii, Mr Gordon M. Nishida; DARA - Biological und Chemical Research Institute, Zoology, Harvard University, Cambridge, MA, USA; MHGN - Musée d'Histoire Naturelle, Neuchatel, Switzerland, Dr Jean-Paul Haenni; MHNG - Museum d'Histoire Naturelle, Geneva, Switzerland, Dr V. Aellen and Dr B. Hauser, NMYM - Museum of Victoria, Melbourne, Vic- toria, Australia, Dr A. Neboiss; NMWA< - Natur- historisches Museum Wien, Vienna, Austria, Dr A. Kaltenbach; QMBA - Queensland Museum, Brisbane, Queensland, Australia, Dr G.B. Mon- teith, SAMA - South Australian Museum, Adelaide, South Australia, Dr G.F. Gross; WAMP - Western Australian Museum, Perth, Western Australia; Dr T.F. Houston, ZILS - Zoological Institute, Lund, Sweden, Dr Roy Danielsson. | have examined some of Saussure’s specimens borrowed from the Geneva and Neuchatel muscums. According to Dr Bernd Hauser (pers. comm., 1986), Saussure was an independent scientist and his private collection was transterred to the Geneva Museum after his death, He did not designate (ypes, some 532 MEMOIRS OF THE QUEENSLAND MUSEUM specimens had no labels on them, and he never indicated how many specimens he used in his descriptions, A minimum of 2 specimens was certain only if he described, or gave measure- ments of both sexes of a species. He made many exchanges with friends and colleagues so that his syntypes, if any, may be widespread in major world collections. The fact that Saussure indi- cated ‘Mus. Neuchatel’ in his publications does not mean that he did not have specimens of the same species in his private collection. Dr Hauser believes that Saussure’s specimens in the Geneva Museum are syntypes, even though he listed only Neuchatel as. the location of the specimens, in his publications. DIAGNOSTIC CHARACTERS The characters that have been used to distin- guish between the genera discussed in this paper are shown in Table 1. A diagnostic character is one that is useful in distinguishing a species or genus from its relatives, but as Key (1970, p.144) points out the ‘difficulty here is to know which characters may prove to be needed when all related species have been discovered.” In this Paper | attempt to evaluate the generic: impor- tance of the characters shown in Table 1, and redefine those genera which I believe to be valid. My conclusions are based for the most part on examination of the type species (including type specimens wherever possible) as well as the known and new taxa. Wings. ‘The fully developed, reduced or absent organs of flight, in one or both sexes, is often Gislenia ' Fully Developed Fully Developed Neotemnopteryx *! Reduced Reduced Shawella | Reduced Reduced Franwalkeria | Fully Developed Fully Developed Parutemnopteryx ') Reduced Reduced Keyella * Fully Developed Fully Developed Trogloblattella * | Reduced Reduced specific and sometimes attributable to nothing more than mere individual variation. Such dif- ferences may occasionally be given generic recognition, bul only in conjunction with other differential characters of much greater usual stability,” (Hebard, 1929, p.4). Regarding Australian cockroaches, Hebard (1943, p.4, foot- note 4) stated ‘Different degrees of reduction in the organs of flight have obviously been used as major generic factors, although it is now well known thal such characteristics rarely have any generic significance whatever, and indeed such dissimilarity between the sexes 1s now recog- nised as characteristics of many species.’ The wing venation of those species with fully developed flight organs (Neotemnopleryx and Paratemnopteryx) are essentially similar. Whether or not the tegmina and wings are reduced may be intraspecifically variable in some species of a genus, and is of questionable generic value. Princis (1954, p.35) claimed that Shawella (=Paratemnopteryx) ditfered trom Gislenia (=Neotemnopteryx) and Franwalkeria (=Paratemnopteryx) by its reduced wings which do not reach beyond the first abdominal tergum. Paratemnopteryx couloniana (Saussure) has brachypterous tegmina and micropterous wings, but the degree of reduction varies, and there are males whose flight organs are fully developed and extend beyond the end of the abdomen. Neotemnapteryx fulva (Saussure) males are mac- topterous and the females may be macropterous or brachypterous. Male tergal specialisations. ‘In males the dor- sal surface of the abdomen may always be un- Front Femur Type Pulvilli Arolia Presen| Present Present Present Absent Absent Present | Present Absent | Present Present or Absent 'From Princis (1954). “From Princis (1951), based on female only. “This paper. ‘From Mackerras (1967), TABLE. 1. Characters that have been used to distinguish between 7 genera of Parcoblatinni. AUSTRALIAN PARCOBLATTINI specialised in certain genera and show a distinc- live type of specialisation in alhers, but various differences in specialisation or rarcly both types may be found to exist in species of the same genus.” (Hebard, 1929, p.5), According to Princis, Gislenia males have a gland on Ti and T7. as compured to Shawella and Franwalkeria that have only TL modified. The abdominal tergum(a) on which the sexual gland(s) is/are located may be # geod generic character (Roth, 1969), Bul as pointed out by Hebard, the number of modified segments, and their position and morphology may vary in the same genus. Some examples of this are: in Blat- tella spp., the tergal glands may occur on T7 only, or on T? and TS, and their morphology differs between different species-groups (Rath, 1985a), The Australian species discussed here belong in the Parcoblattini.and of special interest is the American genus Parcablatta Hebard whose males have glands on TL only (6 spp.), Tl and ‘T2 (4 spp.), or they are absent (2 spp.) (Hebard, 1917), A striking example of unifor- mity in position dnd morphology of male tergul glands on T7 and TS is found among species of Ischnoptera Burmeister (Roth, 1969, figs 47-53), The degree of development and morphology of the glund may differ between species and may be diagnosue (i.c., deep or shallow fossae, with or without assoctated setae, or whether setal patches are dense or sparse). Neotemnopteryx spp. males always have T1 modified, but based on new (axa, [7 may or may not be modified, In Neatemnapicryx fulva the first abdominal. ter- gum has relatively few setac along the anteromedial margin of Tl, and sometimes these are difficult to see because they are hidden by the hind margin of the metanotum, In his original redescription of Shawe/la couloniana (=N, fulva) Princis (1951, p.61) stated that the male’s ab- domen is unspecialised, but later ( ac ee he corrected (his, A similar tergal gland 1s found on TL of Paratemnopteryx. The gland on TT ol Neotemnoptery* often covers a larger arca, and the group of setae ts denser, exposed, and located in about the middle of the segment, Legs, Hebard (1917) defined 2 kinds of spines on the anteroventral margin of the front femur: those armed witha row of spines which decrease gradually in size und length distad (Type A) (p.12) and those armed with a row of heavy proximal spines, succeeded by a row of more slender, shorter distal spines (Type B) (p.11). Later (Hebard, 1929, p,5) stated that the margin mity have an elongate row of minute piliform spines (‘type A’) or a row of heavy spines which decrease gradually in size distad (‘type B*), twa or three apical spines furnishing an additional feature of high importance. These definitions are not consistent with his earlier ones. | follow his (1917) original definitions as do recent workers (wg. Bruijning, 1948. p.33; Bey-Bienko, 1950, pp.15-L7, fig. 5) and recognise the following types of front femurs (numerals denote the num- ber of terminal spines, exclusive of the genicular spines); hairlike sctac are not considered in the classification: 1. Types Ao, Aa, Ad: long, robust spines which gradually orsometimes suddenly become shorter towards the apex of the femur, the 2 to 4 terminal spines of Unequal length, the most distad longest; the first of the terminal spines may be very short and not much longer than the one preceding il, 2. Types G2, B3: one or more large proximal spines succeeded by a row of minute piliform spinules, terminating in 2 or 3 large spines. In some instances the row of spines may be uniform in length and the size of but stouter than piliform spinules: or there may be a few piliforn spinules among the small heavy spines, These forms may be intermediate between Types A and B, bul usually are considered Type A, 3. Types Cz, C3: row of piliform spinuies only. 4. Types Dy, Da: robust spines and piltform spinules absent, Sometimes a genus may have 2 types of front femurs. Balta spp. have Type B or C (Hebard, 1943), The Parcoblaltim’ discussed here have Types A or B. Type D femurs are found in Choristina Tepper, Choristimodes Hebard, Ee- toneura Shelford, and Stenectoneura Hebard. “The number and type of pulvilli, armament of the ventral surface of the tarsi and development of the arolium has proved to be useful in many ways, whilethe symmetry orspecialisation of the tarsal claws tre features of importance.” (Hebard, 1920. p.3). Princis (1974, p.518) quoted (his sentence und so strongly beleved in the importance of the presence or absence of pulvilli and arolia that he used them as a principal diag- nostic character for certain genera. Bruijsing (1948, p.33) also stated that whether or not the arolia are well developed has “high generic Value.” This may well be true for some genera, but apparently is of less significance in a number of taxa. including some closely related to Paratenimopleryx Saussure, discussed in this paper. Shelford (19U8a, p34), and Rehn (1922, p.A5) indicated that in Tha spp, (Polyphazidae), che 334 presence or absence of tarsal arolia varies, and does not have as much taxonomic importance in the polyphagids as in the Blattidae. But even species of Tryenicus Shaw (Roth, 1987a, p.152) and Neostylopyga Shelford (Rath, 1988, p.316), both Blattidae, have some species without arolia, and with some or all pulvilli absent, whereas most of the species in these genera have the Structures. There ure several closely related Australian genera in the Blattellinae (Parcoblattini) in which the presence or absence of pulvilli and arolia has been ased as an important character to distinguish between them. Pulvilh and arolia are present in Neofemmopierye and absent in Paratemnopteryx (original diagnosis) and Trogloblatiella, (pn Shawella (=Paratemnop- feryx) pulvilli are present of absent, and in Fran- walkeria (=Paratemnopteryx) pulvilli are present, arolia absent. In his original diagnosis of Shawella (based only on coulodiana), Princis (1951, p.61) stated that small pulvilli are present on the 4 proximal tarsomeres. However, pulvilli are absent in Shewella douglest Princis (=Neotemnopteryx douglast) (Princis, 1963, p.11). Sometimes it is difficult to determine if pulvilli are absent. In Trogloblatrella the pulyil- lar regions on mast tarsal segments are heavily sclerotised and bear small spines, but some spines form a line limiting a clear apical area of the tarsomere. A ‘normal’ pulvillus is convex and generally is not completely surrounded by short stoutspines. Princis and Mackertas probab- ly considered pulvilli absent if, in lateral view, the apex of the tarsomeres dg not show small, colourless convex swellings that protrude beyond the margins of the segments, Perhups the presence of clear, non-protruding structures at (he apex of the tarsomeres (viewed in ventral view) should be considered as subobsolete pul- villi; probably these are nonfunctional and can- not be used in climbing. Mackerras (1967, p.43) stated “Why the tarsi should fack pulvilli and arolia is not obvious, bul they are so frequently lost in cavernicolous cock- roaches that they must be disadvantageous in some way; possibly mud adheres to (hem and retards progress. Among their relatives living on the surface neither Shawella couloniana nor Gis~ lenia australica (Brunner), nor G. fulva (Saus- sure) have pulvilli in the nymphal stages. These structures are only acquired al the last ecdysis. IL is noticeable (hat nymphs cannet run up the sides of a glass container in the way that adults can, so that T.Jrogloblattella| nullarborensis would not MEMOIRS OF THE QUEENSLAND MUSELIM be able (o negotiate a smooth vertical surface, Shawella douglasi which has so far only been found in caves, lacks pulvilli in nymphs and adults, but small arolia are present in both, In another related genus, Paratemmnopleryx, lack of pulvilli and arolia is cited as a generic character.’ Although she didn't say so, Mackerras may have. been questioning the generic importance of the presence or absence of pulvilli and arolia in this group of cockroaches. Although absence of pul- villi and arolia may be associated with strictly Cave species, these characteristics may occur in truly epigean species as well, During climbing, the tarsal claws function on rough surfaces whereas the arolia and pulvilli are important in climbing smooth surfaces (Roth and Willis, 1952; Arnold, 1974). Christiansen (1965) found that the more elongated tarsal claws in cave collembola reduced adhesion by the surface tension of water in moist caves, and this facilitated locomotion on wet surfaces. In a study of some domiciliary pest cockroaches, Roth and Willis (1952, p.499) found that al least two species, Blatta arientalis Linn. and Periplaneta australasiae (Fab.) have great difficulty walking on a smooth horizontal surface like glass, be- cause their first and sometimes second pulvilli {euplantulae) adhere to the substrate. The insect walks readily if its body is raised high above the surface, Bul if the body is held close to the substrate and its legs are extended so that the first pulvilli of the mid and/or hind legs touch the surface the cockroaches have difficulty pulling or lifting the tarsi away from the smooth sub- strate, In some instances the insect will actually pull itself free leaving the tarsi adhering to the glass (Roth and Willis, 1952, fig. 5), Whether or not the presence or absence of pulvilli and arolia influences the locomotor ability of cockroaches on different cave surfaces remains to be determined, Pulvilli and arolia are adaptive characters (Arnold, 1974), although their loss is not restricted to cave habitats, and they ure insufliciently stable to be used as a principal diagnostic generic character in the encra discussed here. /schnoplera peckorum oth (Blattellidae) and Neostylopyzga jam- busanensis Roth (Blattidac), both cavernicolous species, differ principally from other epigean members of their respective genera, by the ab- sence or reduction of eyes, absence of some or all pulvilli, and absence of arolia. There is na need to erect new genera based on these differen- ces ulone (Roth, 1988), Mebard (1917, p.9) stated that “It must be AUSTRALIAN PARCOBLATTINI 53 remembered that until the Blattidae of the world are monographed, the relative values of all diag- nostic features can not be fully determined— Mackerras (1967, p.39) stated that Trogloblatrel- la has the characters of Gislenia but is modified for a cave existence. In Franwalkeria, Shawella and Paratemnopieryx, we find intermediate stages towards a complete loss of pulvilli and arolia, and the species in these genera generally have reduced eyes. My study of these three genera indicates that they are congeneric, Except for specific determinations, Princis tended to ignore similarities in male characters (if males were known) when he diagnosed genera. | believe that male styles, genital phallomeres, tergal glands, which tend to link the Parcoblattini together, are more important than the presence or absence of pulvilli and/or arolia, although in some instances the latter may be used for species differences, Size. The Parcoblattini contains same of the largest species of Blattellidae (e.g. Neotemnoap- teryx gloriousa 1. sp., Paratemnopteryx broomehillensts n. sp., Trogloblattella nullar- borensis Mackerras), Considerable size varia- {ion occurs in some species and the difference between the smallest and largest specimens is so great thal they appear to be different species (c.g. Paratemnopteryx ceniralensis (Roth)). How- ever, if all important male characters (supraanal and subgenital plates, styles, and genitalia) are similar between the different sized specimens, I have considered them to be one taxon, Measure- ments are given in millimetres. Colouration. The Patcoblattini are more or less unicolourous and for the most part yellowish brown (Neoftemnopteryx), or dark-, reddish- or blackish-brown (Paratemnopteryx). Head, pronotum, and tegmina do not have distinctive markings, KEY TO AUSTRALIAN GENERA OF PARCOBLATTINI based mainly on male characters 1. Anteroventral margin of front femur Type A . 2 ree Anteroventral margin of front femur Type B 2. Epigean species: Eyes extending below level of antennal sockets (Figs 2B,5C,13C). Tegmina and wings fully developed. Styles usually asymmetrically placed on hind margin of sub- in genital plate (Fig. 2F), rarely clase tagethernear mid-jine (Fig, 136). Intercercal ridge usually presenton undersurface of supraanal plate (Figs 1D,13B), rarely subobsolete (Fig. 12B), or ab- sent. Pulvilli presenton 4 proximal tarsameres, arolia present. First abdominal tergum specialised, the setae located in a group medial- ly; seventh abdominal tergum with or without a selal specialisation. Cavernicolovs species: Eyes reduced, not extending below level of antennal sockets (Fig, 14B-D), Tegmina reduced, width normal (Fig, 144), hind wings vestigial, Pulvilli absent from all tarsomeres, or with a subobsolete pulvillus on the fourth seg- MEML ONLY «.scecresegerecesenners eaten Neotemnopieryx Epigean and cavernicolous species: Eyes reduced, usually not extending below level of antennal sockets (Figs 16B,20D,228B), rarely well below level of sockets (Figs 29A,31A). Tegmina often reduced in length, rarely fully developed (Figs 16C,E,24A), Hind wings usually reduced or vestigial (Figs L6C,E,25A). Styles usually symmetrically placed, wide apart an hind margin of subgenital plate (Figs 17D,E,18D,24D). Intercercal ridge absent. First abdominal tergum specialised, the setae arranged or dispersed along the anterior region of segment (Figs 17B,20C,27C), or with asmall group of setae Jocated anteromedially partly or sometimes completely hidden under hind mar- gin of metanotum; seventh abdominal tergum usually unspecialised, rarely with a large, dense, medial group of setae (Figs 18B,27A). Pulvilli present, subobsolete, or absent, aralia small or absent ...,.....00.5+, Paralemnopteryx 3. Eyes present. Tegmina and wings fully developed. First abdominal tergum with a pair of medial setal tufts (Figs 32A,33A) wu... Keyella Eyes absent. Tegmina and wings reduced to small, coriaceous, lateral pads (Fig, 1SA). First abdominal tergum with an elliptical depression without any or only a few minute setae (Fig. ISD) cep ctei ener, Drogtoblatiella Neotemnopteryx Princis Neotemnopteryx Princis. 1951, p. G1. Type species: Temnopteryx ferruginea Tepper, by monotypy. Gislenia Princis, 1954, p. 33. Type species: Ischnap- lera fulva Saussure, by selection, N. Syn, 536 REMARKS It is impossible to separate Neolemnopteryx from Gislenia using Princis’ characters. The holotype of ferruginea is an unusually small female of Gislenia fulva with reduced tegmina that do not reach to hind margin of T7, and wings only slightly smaller; in spite of wing reduction the cubitus has 6 complete and 4 incomplete branches, similar to specimens from a colony established under the name Gislenia australica MEMOIRS OF THE QUEENSLAND MUSEUM at CSIRO, Canberra. I have not seen the type of Ischnoptera fulva but | have examined specimens which Princis determined as Gislenia fulva and which I find is conspecific with Gis- lenia australica and Neotemnopteryx ferruginea both of which I consider synonyms of fulva. Princis (1969, p.732) listed 1 species of Neotemnopteryx and 6 species of Gislenia (1969, p-729). The former (ferruginea) and 3 Australian species of Gislenia belong in Neotemnopteryx Fic. 1. Neoleninopteryx fulva (Saussure), A, B, D-H, J, d 3d: A, Pronotum; B, Eye (lateral view); C, 2, eye: D, Supraanal plate and paraprocis (ventral view); E, Supraanal and subgenital plates, styles (rear view); F, Setal gland on seventh abdominal tergum; G, Subgenital plate (ventral view; styles reflexed dorsad and not visible); H, Hind wing; 1, 2, hind wing; J, Subgenital plate and genitalia (dorsal view). Abbreviations: a, left paraproct; b, right paraproct; c, intercercal ridge; d, left hooklike genital phallomere; e, median phallomere; f, right phallomere. Localities: A, D, J, lectotype of /schnoptera australica Brunner, Tarangower, N.S.W.; B, C, H, 1, Canberra, A.C.T.; E-G, Black Mt., A.C.T. AUSTRALIAN PARCOBLATTINI which is restricted to that continent. The two Indian species, Gislenia brevipes (Walker) and G. himalayaca (Brunner), are species of Episymploce (Roth, 1987b). Gislenia indica (Brunner) from Malacca, West Malaysia is a Symploce (Roth, 1986). [recognise 10 species of Neotemnopteryx of which 6 are new and 3 are 537 new combinations. One species is cavernicolous. the others epigean. The distribution of species of Neotemnopteryx is shown in Fig 34. DIAGNOSIS Epigean species. Eyes well developed usually 2mm Fic. 2. Neotemnopteryx fulva (Saussure). 3 d: A, Pronotum; B, C, eye (lateral view); D, Supraanal plate and paraprocts (ventral view); E, Distal region of subgenital plate, and styles (dorsal view); F, Subgenital plate and styles (ventral view); G, Genital phallomeres (dorsal view), Localities: A, D, E, G, Australia (no exact locality), holotype of Periplaneta apicalis Walker; B, Perth, W.A.; C, D, Margaret River, W.A, 538 extending below level of antennal sockets (Figs 1B,5C,6D). Tegmina and wings usually fully developed extending beyond end of abdomen; in one species (fulva) some females have reduced tegmina not reaching end of abdomen, hind wings only slightly smaller, their veins distinct. Cubitus vein of hind wing with 2-6 (usually 4-6) complete and 2-7 (usually 3 or more) incomplete branches, apical triangle absent or subobsolete (Fig. 1H,I). Anteroventral margin of front femur Type A3; pulvilli present on 4 proximal tar- someres, tarsal claws simple, symmetrical, arolia present. Male. First abdominal tergum specialised, the dense group of setae exposed medially on segment. Seventh abdominal tergum with or without specialisation; if present may vary from dense to sparse (Fig. 3D-G). Supraanal plate usually with ridge or shelf on ventral sur- face between cerci (intercercal ridge); this may be darkly sclerotised extending ventrad into genital chamber (Figs 5B,8C) but sometimes is teduced (Fig. 1D), subobsolete (Fig. 12B), or absent (Fig. 6B). Paraprocts dissimilar, right one larger, often with spinelike processes (Figs 1D,8D-F). Styles present, partially covered by small dark spines, usually asymmetrically placed, right one at or near apex of plate, the other to its left (Fig. 2E,F), rarely both symmetri- cally located at apex (Fig. 13E). Genital hooklike phallomere on left side. Ootheca rotated 90 prior to deposition. Cavernicolous species. In the one cave-dwell- ing taxon (douglasi) eyes are reduced and do not extend below level of antennal sockets (Fig. 14B-D). Pulvilli are lacking from all tarsomeres or a subobsolete pulvillus occurs on fourth tar- somere only. KEY TO MALES OF NEOTEMNOPTERYX Neotemnopteryx fulva appears twice in the key because its tergal gland on T7 is variable and some individuals apparently lack this specialisa- tion. 1. Eyes reduced, not extending below level of antennal sockets (Fig. 14B-D). Tegmina reduced reach- ing to TS or T7 (Fig. 14A). Hind wings ves- tigial. (CaverniCOlOuS) ......:cseeeeeeeees douglasi Eyes well developed, extending below level of antennal sockets (Figs 2B,5C). Tegmina and wings fully developed reaching beyond end of abdomen. (€pigean) oc cecseeeseeseeeeeeseeeens 2 MEMOIRS OF THE QUEENSLAND MUSEUM 2. Styles close together at or near apex of subgenital plate (Fig. 13E) ......ccceeeeeeeeeee styliparedra Styles not aS ADOVE ...... ee eeeeseeeeeseeeeseeeeeeeeneeenes 3 3. Posterior halves of lateral margins of pronotum, straight, weakly oblique (Figs 6A,7A)........... 4 Posterior halves of lateral margins of pronotum MOLSHAIBHE 56h Zrraersrscenrrpeserocanonsrabersederstenateene 5 4. Supraanal plate subtrapezoidal (Fig. 6B). Right style decidedly more robust than the left one (Fig. GE )icisscsteohsccsccssnssastoptenctussonwiescepdesee seen australis Supraanal plate produced, tongue-shaped (Fig. 7C). Right and left styles about same size (Fig. LEED) ducers ryote syenpsee ders tepienkteyaved he ceretoedebes glossa 5. Hind margin of supraanal plate distinctly concavely excavated (Fig. 10A) ........:scccceseeeee concava Hind margin of supraanal plate not concavely SXCAV ALC cares consnddtsanssiossetsvidsebevitasoatsasvaysentees 6 6. Abdominal terga 1 and 7 with setal specialisation Abdominal tergum 1 specialised, T7 un- SPO CHAISE sccrtcsecssecsnpdostsecennescesneonsevoounsscnancteg 8 7. Right style distinctly more robust than left one (Fig. 5F,G). Paraprocts as in Fig. 5B. Median genital phallomere with square-shaped preapical ex- tension (Fig. SE) .........cssescseeeseseeee gloriousa Right and left styles similar (Figs 1J,2E). Paraprocts as in Figs 1D,2D. Median genital phallomere without preapical extension (Figs LU PAT) cestssvessersvistintreshndernrhsteineeheproeaacenthy fulva 8. Pronotum subelliptical, greatest width at about middle (Fig. 11A) oc. cscseseeeeseeseee elliptica Pronotum with greatest width below middle 9. Median genital phallomere apically bifurcate (Fig. BG) asliecdnadessetsneapecadogareastgnostt fa peatocscees bifurcata Median genital phallomere not apically PIPE CRS 50 5ceinprorpindavonscegnian cguvancnpoeapenstsaeseat 10 10. Pronotum subparabolic (Fig. 12A). Intercercal AUSTRALIAN PARCOBLATTINI ridge subobsolete (Fig. 12B), Genital phal- lomeres 48 1 Fie. LAE ec esseeesteeeenee nana Pronotum suboval (Figs |A,2A). Tntercercal ridge distinct etPIE. 3D) Genital erat in Fig. 1... iy Patna Lal .. fulva Neotemnopteryx fulva (Saussure) n.comb.(Figs 1A-J,2A-G,34) Ischnoptera fulva Saussure, 1863, p. 156, pl, |, fig. 18 (3); Walker, 1868, p, 119; Saussure, 1869, p, 251; 1872, p. 102; Tepper, 1893, p. 55; 1894, p. 172; Kirby, 1904, p. 81; Shelford, 1908b, p. 7. Gislenia fulva (Saussure): Princis, 1954, p. 33, fig, 31 (d and ©); 1959, p, 125. N. Comb, Ischnoptera australica Brunner, 1865, p. 131 (a); 1862, p95 (/schnoplera australica sp. nov., nom, nud, till 1865); Walker, 1869, p, 145; 1871, p, 31, Tepper, 1893, p. 50; Kirby, 1904, p. 81 (incorrectly synonymised under /schaoptera australis Saus- sure), Gislenia australica (Brunner): Richards, 1967, pp. 37-39; Roth, 1968, pp. 84, 110, fig. 111 (ootheca); Princis, 1969, p. 730, Periplaneta apicalis Walker, 1468, p 129 (d), Tep- per, 1893, p. 105; Kirby, 1904, p. 81 (incorrectly synonymised under /schnaplera australis Saus- sure), Gistenia apicalis (Walker): Princis, 1959, p, 125 (synonymised under Gislenia fulya); 1969, p. 730, WN. Syn. Temnopteryx ferruginea Tepper, 1895b, p, 148 (2), Ceratinoprera ferruginea (Vepper): Shelford, 1908b, p. 20. Neatemnopteryx ferruginea (Tepper); Princis, 1951, p. 61, N. Syn, HOLOTYPE (nol examined) d, Australia. (According to Saussure, the specimen wus in the Mus. Neuchatel. However, Jean-Paul Huen- ni (pers. comm., 1956) informed me that it is not there. It is not in the Geneva Museum (Dr B, Hauser, pers. comm., 1986)). MATERIAL EXAMINED NEOTYPE (here designated); d, Canberra, ACT, Australia, light trap, 7.xiii, 1950, P,B. Carne; in ANIC. New SoutH WALES. NMWA; d (terminalia slide 45) lectotype (selected by Prineis in 1965), of /sehnoprera australica Brunner, Tarangower; a 2 ‘paratype’, from Qld, ‘Boucard ded.” also was selected by Princis, but itis not conspecific with fulva and is Neoterinepterys gloriousa n. sp. This specimen does not have type status because Brunner did nol describe the fernale of vi eo = australica and did not indicate Queensland as one of the localities, listing only Tarangower, and Sydney. ANIC; Armidale, 19. 26,1.1960, A. Stock, 16, 12.xi.1959, 1d (terminalia slide 59), 26.xi11.1959, 1d, 5.1.1960, C.W, Frazier; NE National Park, Pt Lookout, 5200ft. 1d, 7.xii.1962, A. Stock; Armidale (U.N,B, Campus), 670m, 12, 30.xii.1978, S. Harrington; Yass, F.C.T., 19 (cattying partially rotated oatheca), 10.17.1929, K, English; Jervis Bay, 1¢ (reared), 19.ix.1951, H.M, Cane. ANIC: Arrawarra, north coast, al light, 1d (terminalia slide 75) 7.xii.1961, 1¢, 9x196), 18, 5.xi.1961, C.W. Frazier; Armidale, 19, at light, 27.1.1963, CW. Frazier. SOUTH AUSTRALIA, MHNG: L® paralectotype (here designated) of /schnopterd australis Saussure (misidentification). This female is not australis be- cause the posterior halves of the lateral margin of the pronotum are rounded, not straight (see Fig. 4A). Tis fegmina and wings are fully developed and extend beyond end of abdomen; cubitus of hind wing with 6 complete and 5 incomplete branches. Based on Dr. Hauser’s comments that Saussure did not indicate ‘synlypes’ (see introduction) I have designated this specimen a paralectotype, but I believe it is fulva, The male lectotype of australis is in the Neuchatel Museum (MHGN). ANIC: 25km E by N of Kimba, 33.058 136.41E, 22 2, 3.ix.1981, D.C.F. Rentz. Victoria. NMVM: Fernshaw, 2 holotype of Tert- Nopteryx ferruginea Tepper, iji.1958, type no. 4446; same data as holotype of T. ferruginea, 1 (abdomen missing). ANIC; &km SW of Moyston, Id, 16.xi. 1959, E.F, Riek. AUSTRALIAN CaPITAL TERRITORY. ANIC: Canberra, in house, 17, 16,ii.1963 (killed 8.iv.1963, ‘progeny in jar 33°, 20 d, 12 nymph (det. Gislenia fulva (Saus- sure) by Mackerras); Canberra, C.S.1.R.0., colony 146 (started from 1 2), 200 3 (1 with terminalia slide 43), 1722 (1 with genitalia slide 44), 14¢¢ and 929 nymphs (det, as G. fulva by Muckerras); Canberra, 1, 18.11.1965, 19 (carrying partially rotated ootheca), 11.1952, D.F. Waterhouse; Canberra, light trap, CS.LR.O,, 34 od, 4.1.1953, 5d dF, 23.xi1.1953, 1d, 14,811, 1950, PB. Common, 20d, 25,1.1954, PJ, Sinclair, 1d, 29,xi, 1950, 204, 18.xij,1950, P.B. Came; Canberra (Turner), 1¢ (terminalia slide 72), 22.1.1969, R.W. Kerr; Black MU, (some taken in light trap), 1d (terminalia slide 46), 23,xi1,1966, 1d (ter- minalia slide 71), 14.1963, 1d, 29,xii,1961, 3d 3, 17.1,1962, 10, 18.xi1.1962, 1d, 5-6,x1,1966, 14, 9. xi), 1968, 1d, 7.1,1969, LFLB. Common. 1d, 7.xit. L950, 2d od, 18.xii.1950, 1, 26.11.1950, 20 3, (31.1951, 10, 134.1953, PLB. Carne, 1d, 1.xii,1948, RW, Kerr, WESTERN AUSTRALIA, ANIC: Denmark, Id, 23,4,.1935, K.R. Norris; Dandalup, |d (terminalia iM slide 74), 19.i,1948, D. Nicholson; Augusta, 1°, i.1968, R.F. and E.M, Watson; Victoria Park, 1d, 111.1950, J.A. Mahon: Perth, ex colony 79 (CSIRO), 68,49 % (alt with oothecae), 30d and 329 nymphs, xli.i964, BPBM; Skm W of ‘Lake Cave’, )- 40m, 1¢ (terminalia slide 454), 7-8,x1, 1963, J, Sedlacek. MCZH: The following were collected by P.J. Darlingion or W.M, Wheeler, on the Harvard Australian Expedition 1931, Pemberton, 2d, 19, 10.xi,, 12, 13.xi.; Margaret River, 3d d (1 with ter- minalia-slide 239), 29 2, 3.xi., 1d, 30.x. ZILS: The following were collected by T. Gislén and reported as Gislenda fulva (Sauss,) by Princgis (1954, p.33): Mar- garet River, Mammoth Cave, Bd, S29, 10 xin, 1951; S Karridale, Hamelin Bay, 20d, Vwi, (981; C, Naturalisie, Bunker's Bay, 36d, 19, Y.ni1.1951; Nornalup, | 2, 20,i,1952; Denmark, Karri forest. 12, 261.1952, WAMP; Manjimup, | &; Yul- lingup, 1d, 19 (labelled Gislenia fulva by Princis, 1954); Cottesloe, 1d, iv.1916; Apple Cross, 1d. L.xii, 1963, G.M. Riley; Margaret River, 22%, Kalamunda, 1d, 15,1971, P. Shaw; Rottnest, 1d, 1.1931 (labelled G. falva by Princis, 1954); Mandurah, 1d, 23,1,1979, R. Ruston: Highbury, 19%, 641.1965, W.H. Butler; nt Devils Lair Cave, 3 miles from sea, 19, 20.11.1973, A. Baynes; Denmark, 12, 27.1.1979, R.P. MeMillan, AUSTRALIA (fo exact locality), BMNH; 3 (not 2 as indicaled) (lerminalia slide 25()) holotype of Periplaneta apicalis Walker, presented by C,G, Grey DESCRIPTION Male. Eyes well developed extending slightly below antennal sockets (Figs 1B,2B.C), Inter- ocular space about same or stightly greater than space between antennal sockels, Pronolum widest below middle (Figs 1!A,2A). Tegminaand wings fully developed extending beyand end of abdomen; cubitus vein of hind wing with 3-6 complete and 2-6 incomplete branches, apical triangle absent (Fig. 1H). Front femur Type Aa; pulvilli on 4 proximal tarsomeres, tarsal claws symmetrical, stmple, arolia small. First ab- dominal tergum specialised, setae dense and numerous. Seventh abdominal tergum specialised, with sctac as dense as on Tl (Fig. LF), or sometimes sparse and difficult to see. Supraanal plate with hind margin (when flat- tened) with small median protrusion (Fig. 2D), (runcale (Fig, 1D), or rounded; intercereal ridge present but variably developed (Figs 1D,2D); paraprocts dissimilar, right one with U-shaped sclerotization (Figs |D.2D), Subgenital plate weakly asymmetrical, styles small located to left of midline, right one slightly more robust (Fig. MEMOIRS OF THE QUEENSLAND MUSEUM 1E,J); in pinned specimens hind margin may be teflexed dorsad (Fig. 1E) so that styles are not visible in ventral view; if margin is not reflexed, styles are visible from below (Fig. 2F). Genitalia as in Figs U,2G: hook on left side, with a subapi- cal incision, median phallomere rodlike, apically acute. Jolauration. Yellowish to reddish brown. Hind wings with anterior ticld weakly yellowish, sometimes darker apically. Terminal abdominal segments sometimes.darkly infuscated, Measurements, Length, 17.0-23,0; pronotum length x width, 4.5-5.8 x 5.9-7,5; tegmen length, 14,5-24.0, Female. Teymina fully developed extending beyond end of abdomen, or reduced reaching to about T5-T8, rounded apically; in brachypterous individuals, wing venation is distinct (Fig. LI), and eyes (Fig. 1B) are similar to those of males and macropterous females. Supraanal plate trigonal, reaching to hind margin of subgenital plate or slightly beyond. Ootheca, Dark teddish brown, 4.1-4.3mm high, 6.4-10.4mm long, with 20-33 small keel serrations and 20-30 egg chambers; oothecal wall nol covered by a removable membrane. Measurements. Length, 16.4-22.8; pronotum length ¥ Width, 4.9-6.0 x 6.1-7.7; tegmen length, 11,0-14,2 (brachyplerous), 165-21.8 (macro- pterous). VARIANTS (Figs 34-H,4A-D,34) MATERIAL EXAMINED QUEENSLAND. OMBA; Braemar, 5.F., via Kogan, SE Old, 1d, 15-19 %,1979, GB, Monteith and OM; Hinchinbrook Is., NE Qld, Gayundah Ck, 10m, Id, 19, 7°15.xi.1984, Monteith, Cook and Thompson; ‘Morey’, 120km W of Windorah, 2¢ d, 24.ix.1983, G.B, Monteith. ANIC: The Boulders, Skm W of Babinda, 17,218 145,52E, 1d (terminalia slide 65), 14-15.xi.1981, J. Balderson; 24 miles SW by W of Kingaroy, at light, 1d (terminalia slide 66), 21.xi. 1968, Britton and Misko; Bidsvold, 1d. New Sout WALes, ANIC; 31km WNW of Tenter- field, 28.588 [5.43E [sie) probably 151.438], 1d, 23.x1.1983, Stap 64; Lake Cowal, 1d (lerminaliastide 62), (6.xi.1971, W. Vestjens; Willandra Bridge, iIkm N of Mossgiel, dry swamp, al light, 33.168 144,34E, 1¢ (terminalia slide 68), 21.xii,1970, Brit- ton, Misko, and Pullen, Arrawarra, north coast, 1d (terminalia slide 75), al light, 7.xii.1961, C.W. Frazier. SOUTH AUSTRALIA, ANIC: 2km SW of Mt Barr, SSE of Abminga, 26.208. 134.565, at light, 1d (terminalia AUSTRALIAN PARCOBLATTINI 541 Fig. 3. Neotemnopteryx ?fulva (Saussure), variant. dd: A, Pronotum; B, Eye (lateral view); C, Supraanal plate and paraprocts (ventral view); D-G, Setal glandular area on seventh abdominal tergum; H, Subgenital plate and genitalia (dorsal view). Localities: A, B, Tenterfield, N-S.W.; C, H, Lake Cowal, N.S.W.; D, The Boulders, Qld; E, 24 miles SW. by W. of Kingaroy, Qld; F, G, exact localities unknown. MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 4, Neotemnopteryx ?fulva (Saussure), variant. 6 d: A, Pronotum: B. Supraanal plate and paraprocts (dorsal view); C, Bye (lateral view); D. Subgenital plate and genitalia (dorsal view). Localities: A, C, Braemer, Qld; B, D, The Boulders, Qld. slide 69), 25.ix.1972, K.ALL, Key ef a/, WAMP: Etadunna Stn, 1d. x.1972, K. and B, Oldfield. DESCRIPTION A number of small specimens appear to be different from fulva, but in spite of their size 1 am provisionally considering them to be variants because their paraprocts and genital phallomeres are similar to what I believe to be that species. The hind margin of the supraanal plate may have a small medial protuberance (Fig. 3C, ep. Fig. 2D), or is rounded (Fig. 4B). The intercercal ridge may be divided (Fig. 4B, cp, Fig. 2D), or undivided and concave (Fig. 3C). The tergal gland on T7 varies from dense and distinct to sparse and subobsolete (Fig. 3D-G), or apparent- ly absent. Because characters such as shape of hind margin of supraanal plate, intercercal ridge, and presence or absence of a gland on T7, which in some species are diagnostic, but here are so variable in the large number of specimens I have examined, J do not feel it advisable to describe a AUSTRALIAN PARCOBLATTINI $3 new species. However, biological studies might show that more than one taxon is present among this material. Measurements (33). Length, 13,5-18.4; pronotum length x width, 3.4-4.7 x 4.7-6.0; teg- men length, 15.0)-19.7. REMARKS Neotemnopteryx apicalis was synonymised by Kirby under australis, and by Princis under fulva. it ts clearly not australis, as indicated by Princis (1969, p,.730). Princis (1969) gave only Western Australia as the locality for fiedva, and all his records (Princis, 1954, p.33) are from that slate. Both Saussure and Walker simply indicated that their species, fulva and gpicalis, came from Australia. The differences between the types of apicalis and australica (cp. Figs 1,2) are small and, based on examination of a large number of individuals, I consider them synonyms of fulva. As pointed out under remarks of the generic diagnosis, Neatemnopteryx ferruginea is based on a brachypterous specimen of NV. fulva. Collection data indicates that this is acommon and widespread species occurring in ACT, NSW, Vict., SA, and WA. Neotemnopteryx gloriousa n. sp. (Figs SA-G,34) MATERIAL EXAMINED Hovotyer: 3, Mi Glorious, southeast Queensland, 21.1974, A. Hiller, in OMBA. PARATYPES; QUEENSLAND. QMBA: Mt Gloridus. 2d, 211974, 1d, 1S, 3.xi1.1973, A. Hiller; ML Misery, Shiptons Flat, via Cooktown, 1d, 29 9, 1982, L.Roberts; Mt Chinghee, 12km southeast of Rathdowney,] 9, 17.xii.1982 (Rf, 720m), Monteith, Yeules and Thompson; Kroombit Tops (Upper Kroombit Ck), SSW Calliope, 12, open forest. 9 19. xi. 1983, G. Monteith and G. Thompson; Black- down Tableland via Duaringa, ©. Old, 1°, 28-29.x.1980, G.B, Monteith; Mt Cannon summit via West Burleigh, SE Old, rainforest, 1d, 1 2, 18.1.1987, G, Monteith and D, Cook} Mt Moffat, N.P., Top shelter shed, 1000m, 1, 10-12.x.1987, Top Mofial Camp, 1d, 13+1S.xii,1987, Monteith, Thompson. Yeates. BPBM: Mt Glorious, | Y terminalia slide 453), 29,41 1968, R. Rice. The following were collected by J.Loand M, Gressitt: Mt Glorivus, Malaise trap, 6d d (1 with terminalia slide 452), 39 9, S-8.4i.1961, 1d, SYR, 13.01.1961, selerophyll forest, 1d, 13- 1641, 1961, rain forest, |}, 24-2801, 1961 (1d, 12, relained al MCZE). ANIC: Brisbane, 1d, 13.11.1960, C.F. Ashby; 2km NW of Mt Mowbullan, Bunya Mins, we light irap in Nothofagus forest, 36 d, 6.11970, Brit- ton, Holloway, and Misko; Binna Burra, Lamington Nat. Park, 1¢ (terminalia slide 86), 19, 12.11.1964. New SouTH WALES, BPBM; Ourimbah, 1 9, 1.1904. ANIC; Armidale, 1d, 26.xi1.1959, 12, at light, ).17, 1960, CW, Frazier; Allyn River, Chichester, S.F., 32.088 151.27E, Ld, L0-11.81.1981, T. Weir; Muka N. Res. 4km Nol luku, 12, 21.xi.1982, J.T. Doyen; Plot Forest Road, 3km NNE of Minyon Falls, Whian Whian State Forest, NNE of Lismore, 28,355 153.23E, 2d (1 with terminalia slide 52), 17.x4, 1982, D.C.F. Rentz and C.D. MacNeill. The following were collected by D.C.F. Reniz and M.S. Harvey: Sawpit Ck, 23km E of Woodenbong, 28.228 IS2.S1E, Sd cd, 22.71.1983, Condong Falls, Whian Whian Staite Forest, NNE of Lismore 28.375 153.23E, 1d,1 2, 18.xi.1983; Big Bend Flors Reserve, Gibber- gunyah Rd, Whian Whian State Forest NNE of Lis- more, 28.378 153. 19E, 12, 19.81.1983; Moore Park, Richmond R,, 27km E by S of Woodenbong, 28.278 1§2.54E, 1d, 21.xi.1983; The Bird Tree, Middle Brother State Forest, near Kendall, 31.418 152.418, 13,19, 16.41.1983. The following were collected by J.T. Dayen; Lake Cathie, 1d, 10-11.41.1982; Cobcrafi Camp, Werrikimbe Nat, Park, 1d, 12.41.1982. The following were collected by H.M. Cameron: Mt Warn- ing Nat. Park. near Murwillumbah, 1, 10.vili.1979; Victoria Park near Alstonville, 1d, 6.vii, 1981, 19 (with ootheca, feared from nymph, Feb, 1981). The following were collected by A, Calder; Gloucester R, Barrington Tops Nat. Pk, 32.048 151. 41E.20d,19, 12-14.8).1981, Wiangaree S.F.. 28,228 153,.05B, 150m, 1d, 10- 12.11.1983 (coll, with T. Weir). DARA; Lune Cove. 52 2, 14.xi7.1977, G.R. Brown. AUSTRALIAN CAPITAL TERRITORY, ANIC: Canberra, CSIRO hight trap, 14, 4.i1.1953,1.F,.B, Common. DESCRIPTION Male. Eyes extending betow level of antennal sockets (Fig, SC); interocular and interocellar distances similar; last palpal segment elongate. Pronotum as in Fig. SA. Cubitus vein of hind wing wilh S-6 complete and 4-7 incomplete branches, apical triangle subobsolete. Front femur Type Aa; pulvilli present on 4 proximal tarsomeres. tarsal claws symmetrical, simple. arolia present. First, and seventh (Fig. 5D) ab- dominal terga specialised. Supraanal plate with posteromedial region deflexed, hind margin with small mesal convexity (Fig. SB), or broadly, smoothly convex; intercereal ridge on ventral surface large, darkly sclerotised, divided, covered with minute setae; right and left paraprocts with clongated spinclike processes, tight one covered with small setae (Fig. SB). 544 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 5. Neotemnopteryx gloriousa, n. sp. 3 paratypes: A, Pronotum; B, Supraanal plate and paraprocts (ventral view); C, Eye (lateral view); D, Setal gland on seventh abdominal tergum; E, Genitalia (dorsal view) F, Distal region of subgenital plate showing styles (dorsal view); G, Styles (ventral view). Localities: A-F, Mt Glorious, Qld; G, Condong Falls, N.S.W. AUSTRALIAN PARCOBLATTINI Right style at apex of subgenital plate, much more robust than left cylindrical one, both covered with small spines (Fig. 5F,G), Genitalia us in Fig. 5E: hook on left side with subapical ingiston, median phallomere, apically acute, with preapical square-shaped extension on onc side, Colouration. Chestnul brown, Antennae uniformly pale or first two segments light, remainder dark brown. Pronotum with posterolateral and hind border regions ranging from practically opaque to variably translucent. Supraanal plate with dark brown medial region on posterior half. Cerci dark brown. Hind wing yellowish brown, distal region may be darker. Specimens from New South Wales may have broad dark brown lateral border on venter, with a dark subgenital plate. Female. Supraanal plate broadly trigonal, apex acute or subacute. Supraanal plate may be darkly infuscated in New South Wales specimens; dark brown lateral borders of abdominal sterna, and dark subgenital plate are more extensive and intense than in males.. Measurements (2 in parentheses), Length, 19.0-25,5 (19.0-26.2); pronotum length x width, 5.0-6.3 x 6.3-8.3 (5.1-6.4 x 6.0-8.2); tegmen length, 22.3-27,8 (19.0-26.6), REMARKS There is considerable variation in size in this species, Specimens trom Mt Glorious tend lo be larger than those from New South Wales. Neotemnopteryx australis (Saussure) n. comb. (Figs 6A-E,34) isctnoptera australis Saussure, 1863, p. 155, pli, tig. 17 (dc). Symploce fulva (nec Saussure, 1863): Princis, 1951, p. 59, pl. 6, figs 61, 62(¢d and 2) (misidentification). Gislenia australis (Saussure): Princis,. 1954, p. 34; 1969, p. 730. MATERIAL EXAMINED Lectotyre. (here designated): d, Australia; in MHGN. (I received 4 specimens, supposedly of Ischnoptera dustralis Saussure, from the Neuchiitel Museum. These were Saussure's specimens bul were not individually labelled and were arranged in the collection under the above name, One was ‘Ischnoptera’ australis which Lam designating the lectotype of Neotemnopteryx australis, Of the other 3 specimens, one is émale of an /sehmoptera, having the typical specialisations on abdominal terga 7 and 8; the 545 other 2 specimens are females and probably belong in Ischnomera based on their Wing venation and Type B3 front femurs. /schnopteéra js found principally in South America and does not occur in Australia. Saussure deserihed several species from South America, and the 3 specimens probably were incorrectly arranged in the Museum collection), AUSTRALIAN CAPITAL TERRITORY. ANIC: Canberra, CSIRO light trap, 2d d, 12.iy.1953, PJ. Sinclair; Black ML, light trap, 1¢ (terminalia slide 51), 15, xii, 1968, 19,.8.1,1969, L.F.B, Common. Niw Sourtt WALES. ANIC: 16 miles B of Bungen- dore, 1d, 18x. 1951, H.M. Cane: 13km ENE of Kyogle, 28.35S 153.08E, 19, 20.xi.1983, DCF. Rentz and M.S. Harvey: Durras, North, nt Batemans Bay, | 2 (with detached oothees), 16,),1985, H.M, Cameron, DESCRIPTION Male. Eyes extending below level of antennal sockets (Fig, 6D), interocular width about same as distance between ocellar spots, less than space between antennal sockets. Pronotum with pair of shallow oblique impressions on distal half, lateral margins truncate, oblique, on posterior half, find margin weakly convex, posterior region mostly hyaline (Fig. 6A). Abdominal lerga | and 7 with large dense medial tufts of setae, those on the latter segment (Fig, 6C) partly hidden by T6. Supraanal plate subtrapezoidal (Fig. 6B); intercercal ridge absent, represented by few, robust, minule spines; some small to minute spicules along hind margin and posterior halves of lateral margins of plate; left paraproct with slender surface setac, posterodistal corner prolonged into robust sclerotised spine, right paraproct with large, single.curved, spine-like process, robustand slender setae on other regions of sclerite (Fig, 6B). Subgenital plate with ex- posed region trigonal, convex, large right style at its apex, the smaller left style separated from it by more than its length, both with surface spines (Fig. 6E). Genital phallomeres as in Fig. 6E: genital hook on left side, hardly any neck be- tween hook portion and broad basal arm, former uniformly slender, apex slightly enlarged without subapical incision; median phallomere todlike, apex acute, right phallomere with cleft sclerite. Colouration. Yellowish brawn. Female. Supraanal plate trigonal, sides not concave, apex rounded, reaching about hind margin of subgenital plate, Pronotum as in male, Ootheca black, microscopically roughened, in- dividual egg cells not discernible, keel with 24 S46 MEMOIRS OF THE QUEENSLAND MUSEUM Fic, 6, Neotemnopteryx australis (Saussure). od: A, Pronotum; B, Supraanal plate and paraprocts (ventral view); C, Setal gland on seventh abdominal tergum; D, Eye (lateral view); E, Subgenital plate and genitalia (dorsal yiew). Localities; A- C, E, Black ML, A.C.T.; D, Canberra, A.C.T. Fig. 7. Neotemmopteryx glossa, n.sp. A-D, F, 3 Holotype: A, Pronotum; B, Subgenital plate and genitalia (dorsal view); C, Supraanal plate and paraprocts (ventral view); D, Subgenital plate and styles (ventral view); E, Female paralypefrom ‘Baren Grounds’ , southeast of Robertson, N.S.W., supraanal plate (dofsal view); F, Eye (lateral view). AUSTRALIAN PARCOBLATTINI 547 shallow, undulate teeth; 4.0mm high, 7.6mm long, Measuremenis (2 in parentheses): Length, 20.5-23.0 (19.6-21.4); pronotum length x width, 4,8-5.6 * 6.3-7.2 (5.2-6.1 x 6.7-7.3); tegmen length, 24.0-25.0 (22.4). Neotemnopteryx glossa n, sp. (Figs 7A-F,34) MATERIAL EXAMINED HOvoryee: & (terminalia slide 61), Jolly Flora Res., 17km N of Dorriga, New South Wales, 15.xi.1982, J.T. Doyen; in ANIC. PARATYPES: NEW SOUTH Wales. ANIC: ‘Barren grounds’. SE uf Robertson, 12, 29x11, 1974, K.ALL, Key; Armidale, at light, 1 ¥, 27.13.1963, C.W. Frazier; Durras North. near Batemans Bay, 1 2 (with ootheca), 16,1.1985, H.M. Cumeron, ADDITIONAL MATERIAL QUEENSLAND. QMBA; Mt Moffat N.P., C. Qld, Mahogany Forest, 1000m, 1 2, 11-12.xii,1987, Mon- leith, Thompson, Yeates, EryMoLocy The specific name refers to the male's tongue- shaped supraanal plate. DESCRIPTION Male. Eyes extending well below antennal sockets (Fig. 7F), interocular distance about same as interocellar space, less than space he- tween antennal sockets. Pronotum with posterior half of lateral margins straight, weakly oblique, margin of anterior half convexly rounded, disk with pair of shallow oblique grooves on posterior half (Fig. 7A). First and seventh abdominal terga with large, dense, groups of setal tufts. Supraanal plate with concave sides, produced, tonguc- shaped, margins incrassate with small dark spines apically on each side of midline; paraprocts dissimilar, right one larger, spinclike; intercercal ridge present as a pair of pale plates hidden under paraprocts (Fig. 7C). Subgenital plate trigonal, convex, styles small, right one slightly more robust, located at apex of plate (Fig. 7B,D). Genitalia as in Fig. 7B: hook on left side, distal portion slender with subapical in- cision, basal arm portion broad, neck region ab- sent; median phallomere rodlike, enlarged distally, tapering to weakly divided, acute apex; right phallomere with cleft. Colouration. Yellowish brown, Female. Supraanal plate less concave than in MEMOIRS OF THE QUEENSLAND MUSEUM male, apex rounded (Fig.7E). Cubitus vein of hind wing with 5 complete and 3-5 (1 bifurcate) incomplete branches, apical triangle subob- solete. Hind wing with anterior field infuscated. ‘The ootheca is blackish, surface microscopically roughened, keel undulate with 26 shallow rounded teeth; 4mm high, 7.8mm long. Measurements (@ in parentheses). Length, 21,0 (21.5); pronotum length x width, 5.3 x 6.4 (5.9 x 7.1); tegmen length, 24.7 (25,0). REMARKS The shape of the pronotum is similar in glossa and australis and the differences in the male genitalia are small. The males are casily separated by the shapes of the supraanal plate, and styles, The supraanal plate of female australis is more triangular than that of glossa. The female record of glossa from Mt Moffat, Qld, should be confirmed by examining males, Neéotemnopteryx bifurcata n. sp. (Figs 8A-G,34) MATERIAL EXAMINED HoLoryee: ¢, Emu Ck, 27km of SW of Dimbulah, Queensland, 17,208 144.57E, 25-26.xi.1981, J. Balderson; in ANIC. PARATYPES: QUEENSLAND. QMBA: Bellenden Ker Range, Cableway Base Stn, 100m, Malaise trap in rainforest, 1d (terminalia slide 52), 25-31.x.1981, Earthwatch/QM, ANIC; Crystal Ck, 23 miles SSE of Ingham, 18.588 146.16E, 1d (terminalia slide 63), ¥.x11.1968, Britton and Miska; 6km SE of Chillagoe, 17.125 144.33B, 1d, 26.xi.1981, J. Balderson; Ewan Roud, 3-14 miles W of Paluma, 1d, 4-6.1.1966, J.G. Brooks; Mt Spec, Paluma Ra., 1d, 5- 71.1965, LG. Brooks; Cunningham’s Gap, McPherson Ra., Old, 1 (terminalia slide 177), 3-4.xii.1982, G. Theischinger. AUSTRALIAN CAPITAL TERRITORY. ANIC: Black MI, light trap, 12, (terminalia slide 58), 25.1,1966, 1d, 24.x1),1962. |. FB, Common. ETYMOLOGY The specific name refers to the bifurcated apex of the male's median genital phallomere. DESCRIPTION Male, Eyes extending below antennal sockets (Fig. 8B), interncular width greater than distance between ocellar spots. Pronotum suboval (Fig, 8A), lateral and posterior regions transparent. Cubitus vein of hind wing with 4-5 complete and 2-3 incomplete branches, apical lriangle small or AUSTRALIAN PARCOBLATTINI 549 Fic, 8. Neotemnopteryx bifurcata,n.sp. ¢o ¢.A,B, Holotype, pronotum, and eye (lateral view); C-G, Paratypes: C, Supraanal plate and paraprocts (ventral view); D-F, Variation in right paraprocts (ventral views); G, Subgenital plate and genitalia (dorsal view). Localities: A, B, Emu Ck, Qld; C. D, Crystal Ck, Qld: E. Bellenden Ker Range, Qld; F, Black Mt., A.C.T.; G, Crystal Ck. Qld. subobsolete. Front femur Type A3; pulvilli present on 4 proximal tarsomeres, tarsal claws simple, symmetrical, arolia very small], First ab- dominal tergum with a pair of dense setal tufts anteromedially, Seventh abdominal tergum un- specialised. Supraanal plate transverse, trapezoidal, midposterior region weakly depressed so margin appears undulate or con- cave in pinned specimen; intercercal ridge darkly sclerolised and spined on free border; paraprocts dissimilar, right one variable, with or without a straight, or curved spinclike process (Fig. 8C-F). Subgenital plate with dissimilar styles, right one more robust, at apex of plate, both with small dark spines, interstylar region membranous, less sclerotised (Fig. 8G). Genitalia as in Fig. 8G: hook on left side, with a subapical incision; median phallomere rodlike, apex divided (the bifurcation is visible in pinned specimens if the supraanal and subgenital plates are separated). €olouration. Light brown. Hind wing with distal region of anterior field darker. Female. See variant. Measurements (ACT specimens in paren- theses). Length, 15.0- 18.1 (20,.0-23.0); pronotum length x width, 4.1-5.0 x 5.1-6.0 (5.1- 5.4 x 6.3-6.7); tegmen length, 17.3-19.0 (23.3- 24.0). unt wa oS MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 9. Neotemnoapteryx bifurcata, n. sp., d variant from Broken River, Qld, 50 miles west of Mackay: A, Pronotum; B, Subgenital plate and genitalia (dorsal view), C, Supraanal plate and paraprocts (ventral view); d, Eye (lateral view). VARIANT (Figs 9A-D,34) MATERIAL EXAMINED QUEENSLAND. ANIC; Broken River, 50 miles west of Mackay, | (terminalia slide 67), 1%, at light, rain forest, 29.xi.1968,. Britton and Misko; Coominglah St. For., 22km NW of Monto, 1d (terminalia slide 73). 23.xti, 1982. J.T, Doyen, DESCRIPTION Male. Eyes extending below level of antennal sockets (Fig. 9D); interocular and interocellar distances similar, Pronotum (Fig. 9A) with lateral portions opaque or hyaline. Cubitus vem ofhind wing with 4-5 complete (1 bifurcate), and 4 incomplete branches, apical triangle subob- solete, Front femur Type Aj; pulvilli on 4 proximal tarsomeres, tarsal claws symmetrical, simple, arolia small. First abdominal tergum with a pair of dense setal tufts anteromedially. Seventh abdominal tergum unspecialised. Supraanal plate with hind margin convexly rounded, intercercal ridge on ventral surface deeply concave, minute spines on free border; right paraproct large, broad, robust (Fig. 9C). Subgenital plate with robust right style at apex, left style more slender, cylindrical, both covered with small spines (Fig, 9B). Genitalia as in Fig. 9B Colouration, Light brown. Anterior field of hind wing light brown, darker distally. Female. Cubitus vein of hind wing with 6 complete and 5 (1 bifurcate) incomplete AUSTRALIAN PARCOBLATTINI 551 Fic. 10. Nevtemnopteryx concava,n. sp. & paratype from same locality as holotype: A, Supraanal plate and paraprocts (ventral view); B, Pronotum; C, Setal gland on seventh abdominal tergum; D, Eye (lateral view); E, Subgenital plate and genitalia (dorsal view). branches, Supraanal plate trigonal, apex extend- ing slightly beyond hind margin of subgenital plate. Measurements (2 in parentheses). Length, 18.7-19,2 (23.0); pronotum length x width, 5.0- 5.2 x 6.0-6.5 (5.6 x 6.9); tegmen length, 21.4- 22.8 (24.7). REMARKS There are some differences between the variant and the typical specimens in shape of pronotum, hind margin of supraanal plate, and intercercal ridge. The most striking difference is in the shape of the male's right paraproct. However, the right paruproct of the typical morph is variable (Fig. 8D-F). Slendcrization of the more robust right MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 11, Nevtemmepteryx elliptica, n. sp.g 3; A, Pronotum,; B, Supraanal plate and paraprocts (ventral view); C, Eye (lateral view); D, Subgenital plate and genitalia (dorsal view). C, from holotype, A, B, D, paratype from same locality as holotype. paraproct of the variant (Fig. 9C) could result in a Structure similar to that shown in Fig. 8D or F. If additional collecting and study shows no inter- mediate forms between the variant and typical morphs then the former may prove to be a distinct taxon. Neotemnopteryx concayva n. sp. (Figs 1OA-E,34) MATERIAL EXAMINED Hovoryee; 3. Plot Forest Road, 3km NNE of Minyon Falls, Whian Whian State Forest, NNE of Lismore, New South Wales, 28.358 153.23E, Stop 93, 17.xi.1982, D,C.F. Rentz and C.D, MacNeill; in ANIC. PARATYPES: New SoutTH WALES. ANIC: same data as holotype. 1d (terminalia slide 64); Federal, near Dorrigo, 1d, 26.xi.-8.xi1.1985, A.M. Cameron, DARA: Baulkham Hills, 1¢, 5.xi.1977, B.E. Wallbank, BPBM: Harwood Island, 1d), 28.1.1967, C. Ryan, MCZH: Wentworth Falls, 2800ft, 10, 23.xi1.1931, PJ. Darlington, Harvard Australian Exp, (QUEENSLAND. BPBM: Brisbane, 1¢, 30.ix.1967, B. Bonnie, WESTERN AUSTRALIA, WAMP; Buningonia Spring (well), 31.268, 123.33B, 1d, light trap, 18-25.x1.1978, T F. Houston etal. AUSTRALIAN PARCOBLATTINI 35 ADDITIONAL MATERIAL. New SouTu WALES. ANIC: Willoughby, 1%, 8.11950, K.G, Campbell. QUEENSLAND, BPBM; Mi Glorious, 1 2, 13.11.1961, L. and M, Gressitt, ETYMaLOGY The specific name refers to the concave ex- cavation of the hind margin of the male’s. supraanal plate. DESCRIPTION Male. Eyes extending below level of antennal sockets (Fig. 10D), interocular and interocellar distances about same, less than space between antennal sockets, Pronotum suboval (Fig. 10B). posterolateral and hind border regions transparent. Cubitus vein of hind wing with 5 complete and 3-4 incomplete branches. Front femur Type A3; pulvilli on 4 proximal tar- someres, tarsal claws simple, symmetrical, arolia small. First and seventh abdominal terga specialised, setae on T1 numerous, dense, dark, those on T7 (which is hardly depressed medially) pale, fewer. less dense and difficult to see (Fig. LOC). Supraanal plate with hind margin broadly coneavely excavated, comers rounded; right and left paraprocts dissimilar, former with a spinelike process and setal plate; intercercal ridge present, free margin convex, with small spines (Fig. 1A). Subgenital plate broadly trigonal, right style at its apex, more robust than the left one (Fig. 10E). Genitalia as in Fig. 10E: hook on left side with a subapical incision, rodlike median phallomere apically acute, right phallomere with visible cleft. Colouration. Yellowish brown. Hind wings with veins and distal region of anterior field, yellowish, Measuremenis. Length, 16,3-22.0; pronotum length x width, 4.2-5.2 x 5.0-6.5; tegmen length, 18.5-23.3. Female. | am provisionally placing 2 females listed above, here, pending finding males in the same localities (Willoughby, NSW, and Mt Glorious, Qld). The supraanal plate is trigonal and its apex is shallowly, but distinctly excavated. Measurements. Length, 16.7-18.0; pronotum length x width, 4.8-5,2 x §.9-6.3: tegmen length, 17.7-18.5. REMARKS The shape of the hind margin of the male’s supraanal plate is distinctive, aS is the apically uw indented apex of the female‘s plate. The smallest male came from Harwood Island. Neotemnopteryx elliptica n. sp. (Figs 11A-D,34) MATERIAL EXAMINED Hotoryre: d, at light, 3 miles N of Eromanga, Queensland, 3.xi.1967, R.C. Lewis; in ANIC. PARATYPES. QUEENSLAND. ANIC: same data as holotype, 1d (terminalia slide 60); 62km W by S of Boulja, 23,028 139.18E, 1d, 16.x.1978, M.S, Upton. NORTHERN TERRITORY. ANIC: ‘Newcastle Waters’, 1¢, bevi.1929, T.G_ Campbell. ETYMOLOGY The specific name refers to the shape of the pronotum. DESCRIPTION Male. Eyes very well developed extending well below level of antennal sockets (Fig. 11C), interocular space slightly greater than interocel- lar width, about same as distance between anten- nal sockets. Pronotum essentially elliptical, preatest width at about middle (Fig. 11A). Hind wing with median vein bifurcate distad; cubitus vein with 3-4 complete and 3-4 incomplete branches, apical triangle absent. Front femur Type Aa: pulvilli present on 4 proximal tar- someres, tarsal claws symmetrical, simple, arolia present. First abdominal tergum with a dense, medial, setal tuft. Seventh abdominal tergum unspecialised. Supraanal plate trapezoidal (Fig 11B), hind margin deflexed medially, not ex- tending beyond hind margin of subgenital plate: intercetcal ridge large, pale ot darkly sclerotised, with or without a medial indentation, and with dense group of small spines on rounded free margin; paraprocts dissimilar, right one with a robust, curved, apically acute process (Fig. 11B). Subgenital plate trigonal, apex bearing a small knoblike right style, left style slightly longer, both with some small dark spines (Fig. [1D). Genitalia as in Fig. 1]D: hook on left side with a subapical incision, basal arm slightly wider than hook portion; median phallomere with an acute, darkly sclerotised apex; right phallomete with a reduced cleft sclerite. Colouration. Light brawn. Female, Unknown. Measurements. Length, 15.6-16.4; pronotum length x width, 3.8-4.0 x 5.1-6.0; tegmen length, 16.5-18.3. 554 MEMOIRS OF THE QUEENSLAND MUSEUM Fic, 12. Neotemnopteryx nana, n. sp. 3 holotype: A, Pronotum; B, Supraanal plate and paraprocts (ventral view); C, Supraanal and subgenital plates (dorsal view); D, Eye (lateral view); E, Subgenital plate and genitalia (dorsal view). Neotemnopteryx nana n. sp. DESCRIPTION (Figs 12A-E,34) Male, Eyes somewhat reduced extending slightly below level of antennal sockets (Fig. MATERIAL EXAMINED 12D), interocular space distinctly greater than Houortype: d (terminalia slide 51), BraemarS.F., via interocellar space, Pronotum subparabolic (Fig. Kogan, SE Queensland, 15-19.x.1979, G.B. Monteith 12A), sides deflexed, posterolateral and hind and QM; in QMBA, border regions transparent. Cubitus vein of hind wing with 2 (1 bifurcate) or 3 complete, and 3 incomplete branches. Front femur Type A3; pul- ETYMOLOGY villi on 4 proximal tarsomeres, tarsal claws sym- The specific name refers to the small size of — metrical, simple, arolia present. First abdominal the species. tergum with large setal specialisation. Seventh AUSTRALIAN PARCOBLATTINI 5 mis mi) Fic. 13. Neotemnopteryx stylipareda, n. sp. dd: A, Pronotum; B, Supraanal plate and paraprocts (ventral view); C, Eye (lateral view); D, Setal gland on seventh abdominal tergum; E, Subgenital plate and genitalia. Localities: A, C, Holotype, Wongabel State Forest, Old; B, D, E, Paratype, 21 km south of Atherton, Qld. abdominal tergum unspecialised. Supraanal plate transverse, trapezoidal, with horizontal groove on anterior half, hind margin not reach- ing hind margin of subgenital plate (Fig. 12C); intercercal ridge practically obsolete. repre- sented by pair of thin, lightly sclerotised plates with only a few minute spicules; right and left paraprocts are dissimilar (Fig. 12B). Subgenital plate broadly asymmetrically rounded, right style stouter than left one (Fig, 12C,E). Genitalia as. in Fig. 12E: hook on Icft side; median phal- Jomere with curved, darkly sclerotised acute apex; right phallomere with cleft sclerite. Colouraiion. Head brownish between eyes and medially on face, cheeks pale. Pronotum yel- lowish brown. Tegmina and wings dark brown, Female. Unknown. Measurements. Length, 12.5; pronotum tength x width, 3.4 x 3.8: tegmen length, 14.7, REMARKS This is the smallest known species of Neolem- nopleryx. Neotemnopteryx styliparedra pn. sp. (Figs 13A-E,34) MATERIAL EXAMINED HoLoryrr. d, Wongabel State Forest, near Atherton, Queensland, 17.20S 145,318, 18.xi,1981, J, Balder- son; in ANIC. PARATYPES: QUEENSLAND. QMBA: 21km § of Ather- ton, 1040-1100m, 1d (terminalia slide 50), DK. Yeates and G.J. Thompson; Bellenden Ker Range. Cable Tower 5, 500m, NE Queensland, 1°, x.- xii, 1982, 5, Montague; Kroombil Tops (Upper TA 47 Ck), 45km SSW Calliope, rain forest, 12, 9- 19.xii, 1983, G. Monteith and G. Thompson. ETYMOLOGY The specific name refers to the closeness of the styles. DESCRIPTION Male, Eyes extending below antennal sockets (Fig. 13C), interocular space less than distance between occlli and antennal sockets. Pronotum subpuarabolic (Fig. 13A). Hind wing with 6 com- plete and 4 (1 bifurcate) incomplete branches. Front femur Type Aa; pulvilli on 4 proximal tarsomeres, tarsal claws simple, symmetrical, arolia present. Abdominal terga 1 and 7 with medial setal specialisations, the latter (Fig. 13D) smaller and less dense than former, Supraanal plate transverse, hind margim subtruncate with small medial projection, long slendersetac along dorsal surface, and medioventrally; ventral sur- face with pair of large, dark, triangular, intetcer- cal ridges, cach densely covered with small spines or spicules; right paraproct with long, slender, spinelike process, left one with wider, shorter, apically upturned acute process (Fig. 13B). Subgenital plate with pair of closely spaced styles al apex, left one slightly more robust than right (Fig. 13E). Genitalia as in Fig. MEMOIRS OF THE QUEENSLAND MUSEUM 13E: hook on left side with subapical incision; median phallomere rodlike, apex unmodificd; right phallomere with reduced cleft sclerite. Colouration. Chestnut brown. Subgenital plate and part of supraanal plate may be darkly infus- cated. Hind wing with distal region of anterior field and most of costal vein area infuscated. Female. Eyes wide apart, interacular space greater than distance between ocellar spots. Supraanal plate trigonal, apex rounded. Hind wing infuscated as in male. Measurements (? in parentheses). Length, 20.8-21.0) (20.0-21.0); pronotum length x width, 5.4-5.7 x 6.6-7.0 (5.6-6.1 x 7.3-7.4); tegmen length, 23.5-27.0 (24.5-26.0). REMARKS The hind margin of the supraanal plate of Styliparedra (Fig. 13B) is similar to those of gloriousa (Fig. 3B) and some specimens of fulva (Fig. 2D). The right paraproct (Fig. 13B) is like that of bifurcata (Fig. 6F), and the intercercal ridge (Fig. 13B) resembles that of gloriousa (Fig. 3B), The position and closeness of the styles at the apex of the subgenital plate distinguishes styliparedra from other species in the genus. Neotemnopteryx dougtiasi (Princis) n, comb. (Figs 14A-J,34) Shawella douglasi Princis, 1963. p. 11, fig. 1 (3); Richards, 1967, pp. 37, 38, 41. MATERIAL EXAMINED Hovotyre ¢, Jurien Bay, Western Australia, Lime- stone Caves, 30.175 1 15.008, ix. 1958, associated with droppings of small cave dwelling bats, Epicecus pumilus, A.M. Douglas; Type no. 63-354, in WAMP. WESTERN AUSTRALIA. WAMP: same data as holotype, 2 nymphs (det. as Shawelle douglast by Princis in 1961). ANIC: Smithies Cave, Jurien Bay, on guany, 1d (terminalia slide 40), 12 nymph, 14.xi,1964, B. Muir (det. as Shawella douglasi Princis by Mackerras). DESCRIPTION Male. Eyes variably reduced not reaching below level of antennal sockets (Fig. 14B,D), interocular space greater than distance between antennal sockets (Fig. 14C), Tegmina reduced in length, meeting along midline of body, tapering posteriorly to rounded apex which reaches to TS or T7 (Fig. 14A). Hind wings vestigial, lateral, narrow, reaching T2 or hind margin of T3. Front femur Type Aa; pulvilli absent or with subob- AUSTRALIAN PARCOBLATTINI 557 solete pulvillus on fourth tarsomere; tarsal claws medially (Fig. 14J), Seventh abdominal tergum simple, symmetrical, arolia small. First ab- unspecialised. Supraanal plate transverse, dominal tergum with large, dense group of setae corners rounded, hind margin shallowly in- i " it awen? t 4 Deets tl i He Pe Hy Sean al t Fic, 14. Neotemnopteryx douglasi (Princis). dd: A, Habitus; B-D, Eyes (lateral and frontal views); E, Supraanal and subgenital plates (dorsal view); F, The same (ventral view); G, Supraanal plate and paraprocts (ventral view); H, Distral region of subgenital plate, and styles (dorsal view); I, Genitalia (dorsal view); J, First abdominal tergum and setal gland. Localities: A, D, Holotype, Jurien Bay, W.A.; B, C, E-J, Smithies Cave, Jurien Bay, W.A. 358 dented, fringed with setae along edge, 4 dense group of small dark spines.on distal ventral sur- face (Fig. 14E,F.G); right paraproct with pair of spinelike processes, intercercal ridge absent (Fig. 14G). Subgenital plate weakly asymmetri- cal, styles widely separated, right one slightly, larger, both covered with small dark spines dor- sally, interstylar margin straight (Fig. L4E,F,H). Genitalia as in Fig. 141: hook on left side, with subapical incision; median phallomere apically acute: right phallomere with large setose sclerite. Colouration. Light brown. Female. Unknown. Measurements (holotype in parentheses). Length, (17.5): pronotum length x width, (4.0 x 5.1) 6.2. 6.4; tegmen length, (9.2) 13.0. REMARKS The eyes of the holotype are smaller than those of the large male from Smithics Cave (Fig. 14B,D). According to Princis, douglasi lacks pulvilli. This seems true for the holotype, but in the other specimen the fourth tarsamere appears lo have a subobsolete pulvillus. As pointed out earlier, iL is sometimes difficult to decide if pul- villi are completely absent. In douglasi the pul- villi regions, especially onthe fourth tarsomeres, are lined on either side by a row of heavy spines, and in lateral view pulvilli appear to be absent. When viewed on their ventral surfaces, the apices of these segments are clear and colourless, and somewhat depressed (due to drying?), | have secn only 2 males, butitis possible that both eye reduction and loss of pulvilli could vary between populations of the same species, established in different caves, Individuals established longer in a particular cave might show greater loss of these structures (See races under Paratemnopteryx stonet), The well developed specialisation on T1 and the shapes of the right paraproct and median and night phullomeres (cp. Figs 1D,J and 14G_1) in- dicate that douglast evolved from a Neolemnop- teryx- like ancestor, Trogloblattella Macketras Trogloblariella Mackerrits, 1967, p.39. Type species: Trogloblatiella nullarberensis Mackerras, by monatypy. DIAGNOSIS Eyes greatly reduced or absent, Antennae very long. Tegmina and wings reduced. 4: first ab- dominal tergum weakly specialised, Front femur MEMOIRS OF THE QUEENSLAND MUSEUM with several large proximal spines followed by row of small piliform spinules, terminating in 3 large distal spines (Type Bs); pulvilli and arolia absent, tarsal claws simple, symmetrical. Male penitalia, subgenital plate and styles, paraprocts, and male and female supraanal plates similar to these structures in Neotemnopteryx. REMARKS Mackerras (1967, p.39) was correct in stating that Trogloblattella is related to surface dwelling Neotemnopteryx (Gislenia) and Paratemnap- teryx (Shawella), However, its male genitalia, subgenital plate, styles, and paraprocts are closet to Neotemnopteryx than to Paratemnopteryx. In the only other known species, Trogloblattella chapmani Roth from Sarawak (Roth, 1980, p.97), the male subgenital plate and styles differ distinctly from those of 7. nullarborensis and clearly did not evolve from a Neotemnopteryx- like ancestor. Distribution of 7. nullarborensis is shown in Fig, 35. Trogloblattella nullarborensis Mackerras (Figs 1SA-1,35) Trogloblattella nullarborensis Mackertas, 1967, p. 34, pl. JA-D, figs 1-6(¢ and ?); Norris, 1970, p. 110, fig. 5.6, HoLory rr (mot examined) do, Western Australis, Abrakurrie Cave, abour 24 miles WNW of Eucta, 27.ix.1966, D.C. and J. Lowry. MATERIAL. EXAMINED WESTERN AUSTRALIA. ANIC: The following were determined by Mackerras, and most of them were reported in her paper under distribution but were not indicated as paralypes: Mullamullang Cave, Nullar- bor, 1° (found dead about | mile in), 6.11966, J. Lowry, 12, Jun.1966, B. Robinson, 1d, W Crowle, 12, 8.1,1965. E.G. Anderson: Abrakurrie Cave, about 24 miles WNW of Eucla, | nymph, 27.ix.1966, D.C. Lowry and J. Lowry; Thylacine Hole, 19, 12.¥11.1966, 1 nymph, 23.x.1966, J. Lowry; Easter Extension, Mullamullang Cave, 1d (terminalia slide 80), 1.ix, 1966, J. Lowry; Roaches Rest Cave, about 27 miles NE of Madura, 13, 29.yiii.1966, D.C. and J. Lowry. and G. Hunt; Weebulbie Cave, 1 nymph, 3.x. 1966.5. Lowry. Sout AusTRALIA. ANIC: Koonalda Cave, 31.248 129,50E, 19, 11.4.1970, M.S. Upton and JE. Feehan, AUSTRALIAN PARCOBLATTINI Simm 559 Fo Fic. 15. Trogloblatiella nullarborensis Mackerras, A-E, d o : A, Habitus; B, Supraanal plate and paraprocts (ventral view); C, Subgenital plate and genitalia (dorsal view); D, Hind wing region showing reduced tegmina (in part) and hind wings, and glandular region on first abdominal tergum; E, Gland on seventh abdominal tergum; F-I, 9 9: F-H, Supraanal plates (dorsal views); 1, Tarsus, Localities: A, Roaches Rest Cave, W.A.; B-E, H, 1, Mullamulang Cave, W.A.; F, Koonalda Cave, S.A.; G, Thylacine Hole, W.A. DESCRIPTION Male. Eyes and ocelliform spots absent. An- tennae considerably longer than abdomen. Teg- mina narrow, leaf-like, almost reaching hind margin of metanotum, venation obsolete; hind wings resemble tegmina in colour and texture, smaller, reaching to about middle of T1 or T2 (Fig. 15A). Front femur Type B3; pulvilli and arolia absent (Fig. 151). First abdominal tergum weakly specialised with a medial elliptical shal- low depression divided medially by low, rounded, tapering, longitudinal ridge, which may or may not have a few setae anteriorly (Fig. 15D). Seventh abdominal tergum with a huge, round, medial depression filled with setae, and occupy- ing most of segment (Fig. 15SA,E). Supraanal plate subtrapezoidal, hind margin slightly deflexed (Fig. 15B); paraprocts as in Fig. 15B. 560 Subgenital plate weakly asymmetrical, styles dorsally spined, small, right one slightly larger and located in middle of hind margin (Fig. 15C). Genitalia as in Fig. 15C: hook on left side; median phallomere apically acute; right phal- lomere with a small setose plate and subobsolete cleft sclerite. Colouration. Yellowish brown to brown. Measurements (Mackerras’s measurements in parentheses), Length, 22.0 (24.0-27.5); pronotum length x width, 7.0-8.0 x 7.3- 8.4 (9.0 x 10.0), tegmen length, 5.1-5.8 x 2.4-2.9 (7.0 x 4.0). Female. Much larger than male. Supraanal plate broad basally, sides tapering to a subacute or rounded apex (Fig. 15F- H). Measurements (Mackerras’s measurements in parentheses). Length, 32.5-38.5 (34.0-38.5); pronotum length x width, 10.0-12.0 x 10.1-12.5 (13.0 x 13.0); tegmen length x width, 8.2-9.5 x 3.7- 5.0 (11.0 x 5.5). Paratemnopteryx Saussure Paratemnopteryx Saussure, 1869, p. 273; Kirby, 1904, p. 106; Shelford, 1908b, p. 9; Princis, 1954, p, 35. Type species: Paratemnopteryx australis Saussure, by monotypy. Shawella Princis, 1951, p. 61; 1954, p. 35. Type species: Blatta couloniana Saussure, Princis, 1951, p. 61). N. Syn. Franwalkeria Princis, 1954, p. 34. Type species: Franwalkeria glauerti Princis, by monotypy. N. Syn. REMARKS The armament on the front femur, and hind wing venation in fully winged forms of Paratem- nopteryx, Shawella, and Franwalkeria are similar. Male characters such as style morphol- ogy and their placement on the subgenital plate, morphology and position of the setal gland on T1, and the genital phallomeres also show basic similarities. The principal characters used by Princis to distinguish these 3 nominal genera are the degree of reduction of the tegmina and wings, and the presence or absence of pulvilli and arolia. I have discussed, under diagnostic characters, why these characteristics cannot be used as dis- tinguishing characters in these taxa, and there- fore I am synonymising them, Princis (1969, p.732) listed 5 species of Paratemnopteryx, one with a query. Two of these species are synonyms. I recognise 10 species of which 4 are new and 3 are new combinations. MEMOIRS OF THE QUEENSLAND MUSEUM Three of the taxa are cavernicolous. Some of the epigean species have reduced eyes and lack pul- villi and or arolia, and apparently are preadapted for cave dwelling. Based on front femur type, hind wing venation, male genital phallomeres and style morphology, Paratemnopteryx is closely related to Neotemnopteryx. Distribution of Paratemnopteryx species is shown in Fig 35; Paratemnopteryx centralensis also occurs in New Guinea. DIAGNOSIS Eyes usually somewhat reduced, not extending below level of antennal sockets (Figs 16D,18A,24B), sometimes well developed ex- tending below level of antennal sockets (Figs 29A,30A,31A). Tegmina reduced with hind wings smaller or vestigial (Fig. 16E,G), or both completely developed reaching beyond end of abdomen (Fig. 31B); in fully developed tegmina and wings, discoidal sectors of former oblique (Fig. 31E); hind wing with discoidal, median and cubitus veins straight, latter with 2-4 complete and 0 (rarely) to 4 incomplete branches, apical triangle absent (Figs 16A,20F,30F). Anteroventral margin of front femur usually Type A3, rarely Type Az; pulvilli present on 4 proximal tarsomeres (Fig. 18H), or absent from some or all segments (Fig. 18G), arolia present (Fig. 161), or absent. 3d: First abdominal tergum usually modified, rarely unspecialised; when present, gland area consisting of setae arranged along anterior border of tergum, sometimes part- ly hidden by overlapping hind margin of metanotum (Figs 17B,27C), or grouped anteromedially (Fig. 30D). Seventh abdominal tergum usually unspecialised, rarely with large dense group of setae, medially (Figs 18B,27A). Supraanal plate symmetrical without intercercal tidge on ventral surface; paraprocts dissimilar (e.g. Fig. 17C). Right and left styles usually similar, or almost so, widely separated, sym- metrically placed on hind margin of subgenital plate (e.g. Fig. 17D,E). Hooklike genital phal- lomere on left side. Ootheca rotated prior to deposition. Based on male characters, species of Paratem- nopteryx can be arranged in the following species-groups: 1. couloniana species-group. Right and left styles similar in size, covered in part with short dark spines. A) Supraanal plate with hind margin truncate, AUSTRALIAN PARCOBLATTINI rounded, or with a shallow indentation. Right paraproct similar to thal shown in Fig, 17A. a) Seventh abdominal tergum specialised, Pulvilli present or absent, arolia present. Species: stonei. The following lack a specialisation on T7 b) Pulvilli and arolia present. Species: couloniana. c) Pulvilli present, arolia absent. Species: glauerti. d) Pulvilli and arolia absent. Species: atra, australis, rufa. B) Supraanal plate with hind margin deeply excavated, right paraproct as. in Fig, 28A. Right genital phal- lomere as in Fig. 28B. Seventh abdominal tergum specialised, Pulvilli and arolia present. Species: broomehillensis. 2. centralensis species-group. Styles very small, similar, cylindrical (Fig. 31D), or dissimilar, right ane bearing long, curved setue, Jeft one minute, cylindri- cal, without long setae (Fig. 30E). Right genital phal- lomere as in Figs 2958,30G,31D- First abdominal tergum with or without specialisation. Species: centralensis, howarthi, suffuscula, This group, originally containing only centralensis, was in the genus Symploce (Roth, 1985b, p.300). KEY TO MALES.OF PARATEMNOPTERYX | Styles large, similar in shape, partially covered with short, dark spines (e.g. Figs 1 7E,23D)............ 2 Styles small, cylindrical, similar (Fig. 31D), or dissimilar (Fig, 3OE) ........ccsessseseessaereeneeeets 8 2. Seventh abdominal tergum with large, dense, setal specialisation (Figs [8B,27A) .......seeeeeeeees 3 Seventh abdominal tergum unspecialised ....,. 4 as) . Hind margin of supraanal plate shallowly concave (Fig. 18C). (cavernicolous) ........cc.eees stonet Hind margin of supraanal plate with U-shaped excavation me (Pig. Se pein $3 sbraomehillensia 4. Arolia present. Supraanal plate with apex of hind margin narrow, concavely excavated (Fig. 17B). (EPiQeaM)....eerecsccserreeereereeees couloniana Arolia absent. Supraanal plate not as above .. 5 la . Pulvilli present. Tegmina and wings fully developed. (epigean) ..........0.-006. glauerti Pulvilli absent or subobsolete ......-.:..-cceceeee 6, Supraanal plate broadly rounded (Fig. 22D). Teg- mina not quite reaching tip of abdomen, taper- ing towards. rounded apex (Fig, 22A). (CAVETTICOIOUS) .cscssesserseaseseererseesesswesseeeoes alra Hind margin of supraanal plate not as above, Tegmina shorter, reaching hind margin of T2 or TES (FIG. DEA) foccysescorscratdettggsstlepaeecctccceeptoodete 7 7, Hind margin of supraanal plate almost truncate, comers rounded (Fig. 24E). (epigean)....... rufa Hind margin of supraanal plate shallowly con- cave (Fig, 23C), (epigean) .......ee australis 8. Styles similar, cylindrical (Fig. 31D). First ab- dominal tergum unspecialised. (epigean) peldoclldoplaptleepUeosesetopebsatducteepedenneat soquccp de suffuscula Styles dissimilar, right one with curved setae, lefi one smaller, sometimes subobsolete, without curved setae (Fig, 30E). First ab- dominal tergum specialised (Fig. 30D) .....-....9 9. Supraanal plate broadly rounded (Roth, 1984: figs 18A,19F). Pulvilli present on 4 proximal tar- someres, (@piZean) ,........s01+00.-.. Centralensis Supraanal plate trigonal (Fig. 30H). Pulvilli absent. (cavernicolous) .......;.sss0. AOWAarthi Paratemnopteryx couloniana (Saussure) n, comb. (Figs 16A-H,17A-H,35) Blatta couloniana Saussure, 1863, p, 150, pl. 1 fig, 14 (¢ and 2); Walker, 1868, p. 87. Periplaneta concinna (nec Haan, 1842): Brunner, 1865, p, 229 (4); Princis, 1969, p. 731. Temnopteryx couloniana (Saussure), 1869, p. 240; Walker, 1871, p. 18; Tepper, 1894, p. 170; Kirby, 1904. p. 104, Blatia brunneri Kirby, 1903, p. 375; 1904, p. 139; Princis, 1954, p, 35. Ceratinoptera couloniana (Saussure); Shelford, 1908b, p. 19. Shawella couloniana (Saussure): Princis, 1951, p, 62 (¢); 1954, p. 35; McKittrick, 1964; Roth, 1968, p. 110, fig. 106 (ootheca); Johns, 1966, pp. 95, 96, 134, Paratemnopteryx blattoides Tepper, 1895b, p. 150, N. Syn. Paratemnopleryx australis (nec Saussure): Tepper, 1895b, p. 149 (4) (misidentification). 562 0.5 mm MEMOIRS OF THE QUEENSLAND MUSEUM Simm 2mm FiG. 16. Paratemnopteryx couloniana (Saussure). A-F, 3 3: A, Fully developed hind wing; B, Eye (lateral view) from ¢ with fully developed hind wings; C, D, Habitus (left tegmen removed) and eye (lateral view); E, F, The same; G, H, 2, habitus (right tegmen removed), and eye (lateral view); 1, Tarsal claws and arolium (frontal view); J, Tarsus (lateral view). Localities: A, B, 31 km west northwest of Tenterfield, N.S.W.; C, D, Canberra, A.C.T.; E. F. Black Mt., A-C.T,; G, H, Eildon, Victoria; 1, J, Same as A, B. Methana sp., May, 1963, p. 44, fig. 3 (misidentitica- tion); Johns, 1966, p. 134. REMARKS Shelford examined Tepper’s syntypes of Paratemnoptleryx blattoides and correctly con- cluded that one of the adults was a male of ‘Ceratinoptera’ couloniana (Saussure). How- ever, he incorrectly believed that the other male, which Tepper described as a female, was a valid specimen of blattoides. Actually this specimen alsa is couloniana, Shelford apparently was misled by the fact that the lectotype has fully developed tegmina and wings. whereas the paralectotype is brachypterous. Also the charac- teristic concavely excavated apex of the supraanal plate, typical of couloniana, and dis- tinct in the paralectotype of blattoides, is. not clear in the Jectotype because the edges of the plate are curled upwards and the weakly concave apex looks slightly different. The third syntype of blattoides is a nymph with the same locality as the adults; it is badly damaged and labelled Paratemnopteryx ?blattoides. It is not a Paratemnopteryx; there are no spines on the posteroventral margins of front and mid femurs (hind legs missing), and the anteroventral margin of the front femur has piliform spinules only and one large distal spine (Type C1). MATERIAL EXAMINED All specimens are brachypterous unless indicated as Mmacropterous (macr,): AUSTRALIA, MHGN; ¢& lec- lolype (here designated) and 1 ? paralectotype of Blat- ta couloniana Saussure. Victoria, NMVM: Narre Warren, ¢ (macr.) lec- lotype (here designated), and 1d (incorrectly labelled 2) paralectotype of Paratemnopteryx blattoides Tep- per (the lectotype has a handwritten label ‘copy from above’, j.e., copy from the label on the paralectotype which states ‘Ceraltinoptera couloniana’ Sauss.; Tep- per describes 2d ¢ as opposite sexes of blattoides, His 2 = ‘Cerat. couloniana Sauss., det. by Shelford, sent xii,1907". The paralectotype also has a label ‘sent to Shelford, xii.1907; see other specimen of Tepper with note by Shelford."), ANIC: 2 MacAlistair Crt, Surrey Hills, 1¢, inside house, 2.xi,1981, M.R. Harvey; Snobs Creek Fish Hatchery near Eilden, 19, inside AUSTRALIAN PARQOBLATTINI building, 13.11.1979, M.R. Harvey; Swift's Crk, 12, 10,x,1949, T.G, Campbell; Glen Wills, 20km E of Falls Creek, 1d, 13.4x,1972, M. Schwartz. The fol- lowing were collected by F.M. Cane: 28 miles NNE of Cavendish, 1c, 19, 11.iv,1951; Mangalore, 19. 9.iv, 1951, SAMA: Victoria, 19 ,C. French (incorrect- ly reported as Paratemnopteryx australis Sass. by Tepper, 1895, p.150). MCZH: Melbourne, 10 (macr.), H, Edwards, New Sout WALES. ANIC: Dorrigo, 13 nymph, 21.17.1954, E.F. Riek; Armidale, 1¢ (macr.). 11.1960, C.W, Frazier; Avalon, 1d, 12, 7.xU.1951, M.F. Day; Bankstown, 1d, 1%, xi.1953, Holmes; Coolamon, 15 (macr,), ta light, 81.1954, A.L. Dyce, 65km NW of Nyngan, 1d (macr,), 27.4.1949, ELF, Riek; 3tkm WNW of Tenterfield, 1¢ (macr-), 23.1, 1983, D.C\F. Rentzand M.S, Harvey; Nullo Mt., 20 miles NE of Rylstone, 1d, 11.x7.1950, T.G, Campbell, 16 miles E of Bungendore, 19, 18.47), 1951, H.M, Cane; Hearne Bay, Georges River, 12, &i 141. Mrs Day; Sydney, 222 nymphs, 20.v.L925. W.W. Froggatt; Sutherland, 1d nymph, xi.1927, M, Fuller; Braidwood, 19 nymph, 3.viii 1949, FJ, Gay; Nelligen, 19 nymph, (3,viii, 1949, 22 9 nymphs, lov.1949, KELL. Key, 12 nymph, 3.xi.1949, Cane and Gemmell. The fol- lowing were collected by A.M, Cane; 4 miles SE of Braidwood, | 2 nymph, 4.(i1, 1953; Gundy, 1d and1 2 nymphs, S.iv.1949; Towamba, 292 nymphs, 27.iv.1949; 3 miles SSW of Bega, 3¢d and 399 nymphs, 27.iv.1949, DARA; Bass Hill, 1d (macr.), 1%, 2.xii.1953; Roseville, 1d (macr.), infesting house, 30,xii,1952; Dapto, 29 2, 2.111956, Sydney, 1%, 11.1961; Gladesville, 1% (with ootheca), infesting house, 14.i1.1953, R. Wilson; Herne Bay, 12, 7.ix.1952; Waratah, 19, 1 nymph, infesting house, 5.xi,1952, R, Golledge; Belfield, 19 (with ootheca), in house, 29.71.1954; Seven Hills, 1°, 3.xii.1953, Mrs E. Grudnott; Bankstown, 16°, 26.xi.1953; Eastwood, Id, 2.xii,1953; East Hills, 4 nymphs, iii, 1953, 1 nymph in house, 11.11.1954; “Gilbula’, 16km S of Campbelltown, | nymph, J.vii.1984, 5.G. Hunter; Berridale, 1d, 20,xi.1961; Bankstown, 2¢6, 16.x1.1954; Wollongong; 1d, eating papers in cardboard, 9.xii.1960; Nundle, 19, 13.%i1958; Punchbowl, 1d, indoors, infesting linen cupboards, also naticed outside underneath wood, etc., 9.xii, 1954, Caller; Seven Hills, 1, in house, 7,7,1961; Wahroon- ga, 1, 7¥0.1952, HIM. Bungham, Cammeray, 19. 21.%.1957, Clovelly, 1d, 25.xi.1959, W.E. Wright; Northbridge, 1, x.1950, 1d, 14.v.1951; Groman, 1d, 6.xi,1958, TV. Bourke, Springwood, 19, 19.x1,1954; Beverly Hills, 1d, 12 nymph, 24.1.1955; Blacktown, | d nymph, 22.iii, 1964; Forestville, 3 J and |? nymphs, indoors, 16,iv,1962, Mrs Holder: 563 Rydalmere, 19 fyoiph, allacking dried insects in col- lection, 5.ix,1962, M.J. Nikitin; Blacktown, 14 and 12 nymphs, under bricks, 8.vi.1975, J.A. Longley; Lalor Park, 20.3 mymphs, 2.v.1961; Narwee, 1d nymph, infesting house and believed to be damaging articles of clothing, 8 jv.1954, A.R. Wood. The fol- lowing were collected by C.E. Chadwick; Dapio, 27 9, m house al night, 22.xi,1957; Wollongong, 13, under burk of Eucaiyptus, 15,x.1950; Beverley Hills, 203,12 nymph, in house, iv.1951; Lismore, 1¢, in garage, 10).x.1965; North Ryde, | 2, 28,ii1,1975, 19, 20.41.1975. SAMA: Sydney, | nymph, labelled Paratemnopteryx australis Sauss., by Tepper, 79.4x.1894, Froggatt, 12, Lea. SOUTH AUSTRALIA. ANIC; Ardrossan (Browns Serub), 1d, 19 (with ootheca), 19 (genitalia slide 45), 10, 24.vii, 1979, H.M. Cameron: Ardrossan, 19 nymph, 31, viii 1949, 19, 1.1950, 12, 5.4.1951, HM. Cane; no exact locality, 2d, 2-12i.1952, H.M, Cane. SAMA: Adelaide, public library, 1d (macr.) (incorrectly reported as Paratemnupteryx australis Sauss, by Teppet, 1895b, p.149. DARA: Gawler, 13, under bark of Eucalyptus sp. 24,xii.1967, CE. Chad- wick, QUEENSLAND, ANIC: [Skm S by W of Charleville, 26,328 146,126, 12 (macr.), 21.%.L975, M.S. Upton, AUSTRALIAN CAPITAL TERRITORY. ANIC: Canberra, 1d (macr.), 21.xii.1951, 1d (terminalia stide 41), 422 (1 with genilaha slide 42), E.F. Riek, 10, xii. 1949, 19,29 9 nymphs, 14.vii.1949, H.M. Cane, 229, iii,1944, DPF, Waterhouse, 19°, 15.xi.1954, H.M Cameron, 19, 8.11950, 19, 19.41.1950, K.HLL. Key, 1d, xii.1931, 12 nymph, in house, ji,1964; Canberra (Downer), 1d, 2292, 14,7,1968, K.ALL, Key; Canberra (Farrer), 35.225 149.05E, 19, 2,i1.1984, 19, 21,)1.1984, D.C.F. Rentz; Canberra (Turner), 12, 8.iv.1984, A.M. Hastings; Tinbinbilla, 12, 17,ix,1950,H.M, Cane; Black Mt., in glass house, CSIRO, 1,1 viii. 1948, R.W. Kerr, 1d, 21.%.1949, 19,%.1949, ‘culture parent adults from Black Mt. and CSIRO Lahs, ACT, killed 20,xi.1950', 1146, 229%, 53 2 and 32° nymphs, 24.vij, 1949, 29d and 19 nvmphs, 7.%.1949, 12 nymph, 21.x.1949, HM. Cane, 1d nymph, 13.14.1949, LF.B. Common; Cotler R., 229 nymphs, 16.xi.1949, H.M. Cane; Tinbinbilla Ck, 229 nymphs, 17-vii.1949, E.F. Riek; Canberra (Deakin), 180 3, 4 nymphs, 2.«i,1983, BB, Britton. DESCRIPTION Male, Eyes extend to about level of antennal sockets in macropteraus and brachyplerous morphs (Fig. 16B,D,F). Tegmina nonnal width, usually reduced in length reaching hind margin TS, wings more reduced, narrow, reaching Th (Fig. 16E); rarely, reduced tegmina are longer 564 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 2 . Measurements of Paratemnopterex couloniana. reaching T7 or supraanal plate, in which case hind wings longer and wider, reaching T2, or between T3 and T4 respectively; some in- dividuals have fully developed tegmina and the wings reach about middle of abdomen (Fig. 16C) or end of abdomen, if latter, median vein is simple or bifurcate apically, cubitus vein with 3-4 complete and 0-1 incomplete branches, api- cal triangle absent (Fig. 16A). Front femur Type A3; pulvilli distinct on 4 proximal tarsomeres (Fig. 16J), arolia present (Fig. 161,J). First ab- dominal tergum with setae grouped transversely along thickened anterior margin of segment, partly hidden by overlapping edge of metanotum (Fig. 17B). Seventh abdominal tergum un- specialised. Supraanal plate shallowly convex, raised portion delineated from flattened border by shallow groove paralleling tapering margins of plate, hind margin apex shallowly concave (Fig. 17A), sometimes almost rounded. Dis- similar paraprocts as in Fig. 17C. Subgenital plate symmetrical, transverse, styles similar, symmetrically placed, widely separated on either side of midline, suberect, or usually curved, apices lateral, interstylar margin straight (Fig. 17D,E). Genital phallomeres as in Fig. 17F: hook on left side with subapical incision; median phal- lomere rodlike, apex blunt, unmodified; right phallomere with cleft and spined sclerite. Colouration. Dark brown or dark reddish brown. Female. Eyes reduced reaching to about level of antennal sockets (Fig. 16H). Tegmina reach- ing to middle of T2 or T3; hind wings narrow, vestigial (Fig. 16G). I have seen no macropterous females. Supraanal plate trigonal, apex rounded (Fig. 17G). Genitalia as in Fig. 17G: right paratergite with an enlarged sclerotised plate (arrow). Ootheca rotated prior to deposition; i Pronotum length x width Tegmen length macropterous (¢) 13.2-15.5 3.7-4.7 x 5.0-5.8 12.0-15.5 brachypterous (¢) 12.8-17.8 3.6-4.8 x 5.3-6.8: 6.7-11.0 brachypterous (2 ) 15.8-20.9 4.5-5.7 x 6.8-7.6 5.4-7.5 dark reddish brown, 18 small, rounded teeth in keel, 17 egg cells, 3.5mm high, 5.6mm long (Fig. 17H Nymph. Resembles adult but lacks pulvilli, arolia and tegmina and wings. Measurements. See Table 2. REMARKS Brachypterous morphs of P. couloniana are more common than macropterous forms. All macropterous morphs are males, and the hind wings of brachypterous individuals in both sexes are smaller, usually rudimentary, than the reduced tegmina. Collection data suggest that this species may be domiciliary, and is the most common and widespread member of the genus. Outside of Australia, the species was introduced and is established in New Zealand (Johns, 1966, p.134). Paratemnopteryx stonei n. sp. (Figs 18A-H,19A,B,35) MATERIAL EXAMINED HOLOTYPE: d, Chillagoe Caves National Park, Royal Arch Cave, RA-117, NE Queensland, Australia, 6.vi.1985, F.D. Stone; in QMBA. PARATYPES: QUEENSLAND. QMBA: same data as holotype, 33d, 52% (1d, 19, retained at MCZH); Royal Arch Cave, 12, 7 nymphs, 2.viii.1984, F.D. Stone, T. Connery, and N. Sullivan, 22 2, 1 nymph, 2.vii. 1984, F.D. Stone, 1d (terminalia slide 46), 42 @ (1 with genitalia slide 47), 2d d and 49 2 nymphs, 19.vii.1982, Explorers Club, 32%, 1d and 229 nymphs, 29.v.1985, F.D. Stone, F.G. Howarth, and K. Eipel. ETYMOLOGY The species is dedicated to Dr F.D. Stone of Fic. 17. Paratemnopteryx couloniana (Saussure). A-F, 6d: A, Supraanal plate (dorsal view); B, First abdominal tergum and setal specialization; C, Supraanal plate and paraprocts (ventral view); D, Distal region of subgenital plate, and styles (dorsal view); E, Styles (rear view); F, Genitalia (dorsal view); G, °, supraanal plate and genitalia (ventral view; arrow indicates enlarged region of right paratergite; H, Ootheca. Localities: A, B, E, Bankstown, N.S.W.; C, D, F, Canberra, A.C.T.; G, Ardrossan (Browns Scrub), S.A.; H, Belfield, N.S.W. AUSTRALIAN PARCOBLATTINI 565 Hoty t 566 MEMOIRS OF THE QUEENSLAND MUSEUM AUSTRALIAN PARCOBLATTINI the University of Hawaii, who collected some of the specimens. DESCRIPTION Male. Eyes developed, not reaching below level of antennal sockets (Fig. 18A), interocular space about same as distance between antennal sockets. Tegmina reduced reaching to about hind margin of T6 (see variants), hind wings vestigial extending to about T1 or T2, lateral, narrow, membranous, with a few visible veins. Anteroventral margin of front femur Type A3; pulvilli essentially absent from all tarsomeres (Fig. 18G) (see variants), tarsal claws long, simple, symmetrical, arolia small. First ab- dominal tergum weakly specialised with a few setae along anteromedial border, often complete- ly covered by hind margin of metanotum. Seventh abdominal tergum with a large dense group of setae in an anteromedial depression (Fig. 18B), partly hidden by T6. Supraanal plate transverse, hind margin broadly rounded, convex or weakly concave; paraprocts dissimilar, right one with a large setose lobe and sclerotised spinelike process, left one with or without a short, stout, fingerlike process (Figs 18C,19A). Subgenital plate essentially symmetrical, sides upturned, styles erect (Fig. 18D) or deflexed ventrad, symmetrically placed, widely separated, spines on dorsal surface, interstylar margin weakly convex or straight, with a dense group of short spines along margin of plate at base of styles (Figs 18D,19B). Genitalia as in Fig. 19B: genital hook on left side, robust, median phallomere rodlike, broader on basal half, right phallomere with a cleft and setose plate. Colouration. Reddish brown. Female. Supraanal plate with hind margin rounded or with a shallow apical indentation; right paratergite with an enlarged sclerotization (Fig. 18F). Tegmina slightly longer than in male. Pulvilli essentially absent, but in some specimens they are weakly indicated on tar- someres 3 and 4 of the mid legs. Measurements (2 in parentheses): Length, 16.0-18.7 (16.8-20.0); pronotum length x width, 4,2-5.0 x 6.5-6.8 (5.1-6.1 x 6.9-7.7); tegmen length, 10.3-11.7 (10.5-12.0). 567 Nymph. Resembles adult except for absence of tegmina and wings. Eyes present, pulvilli and arolia absent. REMARKS Mackerras (in Richards 1967, p.40) may have recorded this species as ?)Gislenia sp. from 21 nymphs which could not be placed with certainty to genus. Some of her specimens came from Royal Arch Cave, Chillagoe Caves, and probab- ly these are the same as stonei collected in the same cave. Richards stated that ‘They have been collected from nine caves ranging in distribution from Northern Queensland to Central New South Wales, and also just across the South Australian border. Many are guanobites. Specimens from Ashford Cave in northern New South Wales, and Riverton and Viator Caves just across the Queensland border, all show loss of pigmenta- tion. The nymphs from Alexandra Cave, Naracoorte, are troglophiles. No bats occur in this cave. Cockroaches are present in large num- bers breeding in the cave, but show no sign of cave adaptation.’ Specimens of stonei, including its variants, show no loss of pigmentation. VARIANTS Specimens taken in different caves may vary from the typical material from Royal Arch Cave, in size, length of tegmina (hind wings are very small in all morphs) (Table 3), and presence or absence of pulvilli (all have small arolia). The important male characters such as genital phal- lomertes, subgenital plate, styles, supraanal plate and paraprocts, and tergal glands are so similar in all the forms that I consider them to be races of the same taxon (not paratypes). RACE A MATERIAL EXAMINED QUEENSLAND. QMBA: Chillagoe Caves: Clam Cave, 264,32 2,21.vi.1984, F.G. Howarth and F.D. Stone (1% retained at MCZH), 1¢ (terminalia slide 58), 322, 6 nymphs, 21.vi.1984, 1d, 2292 (1 with ootheca), 27.vi.1984, F.G. Howarth and F.D. Stone; Rhino Cave, 23 d, 1 nymph, 22.vi.1984, F.D. Stone; Spatial Cave, 1d, 22.vi.1984, F.D. Stone. Fic. 18. Paratemnopteryx stonei, n. sp., paratypes and races. A-D, dd: A, Eye (lateral view); B, Setal gland on seventh abdominal tergum; C, Supraanal plate and paraprocts (ventral view); D, Supraanal and subgenital plates and styles (rear view); E, Ootheca; F-H, 2 2: F, Supraanal plate and enlarged right paraproct (ventral); G, H, Front tarsi (lateral views). Races and Qld localities: A, E, H, Race A, Clam Cave; B, D, Race C, Barkers Cave, Yarramulla Station; C, F, G, Typical morph, Royal Arch Cave. 568 MEMOIRS OF THE QUEENSLAND MUSEUM Fic, 19. Paratemnopteryx stonei, n. sp. d, Race C, Undarra, Qld, Barkers Cave: A, Supraanal plate and paraprocts (ventral view); B, Subgenital plate and genitalia (dorsal view). AUSTRALIAN PARCOBLATTINI 569 TABLE 3. Variation in pronotum length and width and tegmen length in races of Paratemnopteryx stonei. Pronotum (mm) Length (do) Mean + §.D. Length (2) Mean + §.D. Width (2) Mean = §&.D. N es Tegmen Length (mm) Mean + §.D. (3) Mean S.D.(?) 1}.2+0.4 DESCRIPTION In the male, pulvilli occur on the 4 proximal tarsomeres of front and mid legs, and on tar- someres 3 and 4, or only on 4, of hind tarsus. In the female, pulvilli are present on tarsomeres 1 to 4 of front tarsi (Fig, 18H), 1-4 (rarely 3 and 4) of mid tarsi, and 1-4, 2-4, or 3 and 4 of hind tarsi. Two oothecae are dark reddish brown, surface smooth, keel undulating, with 14 and 17 egg chambers and 16 and 18 keel serrations (Fig. 18E); they measure (height x length), 3.7 x 5.2, and 3.7 x 5.8. Measurements (2 in parentheses). Length, 15.1-17.6 (15,6-18.5); pronotum length x width, 4.7-5.3 x 6,5-7.2 (5.1-5.9 x 6.6-7.6); tegmen length, 10.0-12.2 (9.6-11.0). RACE B MATERIAL EXAMINED QUEENSLAND. QMBA: Undarra, Bayliss Cave, 12 (on floor, halfway in), 22.v.1985, D, Irwin, 1d, 1 ootheca, 22.v,1985, F.D. Stone, | ootheca, 15.vi,1985, Howarth, Stone, and J, Bresnan. The following were Typical | Race A | Race B Race C 5.0% 0.2 | 4.62 0.2 6.60.1 /6,740.3 | 6.420.1 5 5 8 10,7+0,6 | 10.9+0.8 820. 24 7420.7 | 7.5+0.5 4 11 Significance between morphs (*significant at PS0.05) T&A=0.63; T & B=0.57; T&C=0.00*; A & B=0.83; A&C=0,01*:B& C=0.05* T&A=0.60; T& B=0,15 T&C=0.00*; A : A&C= 0.00*; B T& A=0,58; T & B=0,00* T&C=0.00*; A & B=0.00*; A & C=0.00*; B& C=0.73 T&A=0.00*; T & B=0.00"; T&C=0,00*; A & B=0.00*; A&C=0.00";B & C=0,.90 collected by Howarth, Stane, and Irwin in Bayliss Cave: 19, 21.v.1985, 2d d, 19 (on floor near low ceiling), 23.v.1985, 12, 1 nymph (roots, just before D.U.), 21.v.1985, 5d d (1 retained at MCZH), 19 nymph, 14.vi1985. DESCRIPTION Tegmina further reduced reaching only to mid- dle or hind margin of T3, significantly shorter than the typical morph and Race A (Table 3). Pulvilli present on tarsomeres 1-4 of front and mid tarsi; on hind tarsi pulvilli may be present on 4 proximal tarsomeres, or absent from the first and subobsolete on tarsomeres 2-4, Measurements (@ in parentheses); Length, 15.5-17.4 (19.5); pronotum length x width, 4.4- 4.8 x 6.2-6.5 (5.4-5.7 x 7.4-7.8); tegmen length, 6,4-7.0 (7.0-8,0), RACE C MATERIAL EXAMINED QUEENSLAND. OMBA: Undarra, Pinwill Cave, 17d d (1 with terminalia slide 56), 42 9, 40d and 729 an nymphs, 1 oatheea (1d, LY retained at MCZH), 20.v,1985, F.G. Howurth, D, Irwin, and F.D. Stone; Ikm SE ol Yarramulla, Bof M1 Surprise, Pinwill Cave, 14,19, 11.vii. 1984, D. Irwin, MU Garnet, Yarramulla Sia., Pinwill Cave, on Moor, 1d, 1? nymph, 18.5.1985, F.G,. Howarth; WUndarra, Barkers Cave, 444 (1 with terminalia slide 54), 23.v.1985, 69 2, 21.v.1985, Howarth, Irwin and Stone, 1d, 1¥, | ootheca, 15.v1. 1985, FGH, FDS, JB, 1d, 12.vii 1984, D. Irwin. DESCRIPTION Tegmina reduced as in Race 8 reaching only to TS or rarely as far as middle of TS. Pulvilli present on tarsomeres !-4 of all tarsi. Pinwill Cave specimens are the smallest of all the variants. Race C females from Barkers Cave are somewhat larger than those in Pinwill Cave, and their measurements are listed scparately’ (male size js similar in both caves). The ootheca has 14 egg chambers and measures 3.5mm high and 4.8mm long. Measurements (2 in parentheses): Pinwill Cave: Length, 12.2-16.6 (15.0-18.0); pronotum length x width, 3.6-4.7 x 5.0-6.1 (4.7-5.0 x 6.0- 6.6); tegmen length, 5,8-7,9 (7,0-8.3), Barkers Cave: Length, 14,5-16.6 (18.2-19.7); pranotum length x Width, 4,2-4.7 » 5.3-6,1 (5.2-5.6 x 6.7- 7,3); tegmen length, 6.2-7.6 (7.0-8.0). REMARKS This species is interesting from an evolution- ary view of cavernicolous species because of its various taces, Whatever the cause for the loss of pulvilli (see earlier discussion), their presence or absence varics in different cave populations. It may well be that the type of cave surface on which the insects walk influences the cventual loss af pulvilli, ie., certain surfaces may be detrimental fora species with pulvilli and arolia, even though the loss of pulvilli is not necessarily correlated with a cave habitat, There are epigean species, like Paratemnopteryx rufa (Tepper) which essentially lack pulvilli. Loss of arolia also isn't dependent on cave living; the epigean Paratemnopteryx glauerti (Princis) (=Fran- walkeria glavert/) lacks arolia but has pulvalli- The variation in pulvilli and arolia negates using them in generic diagnoses in this tribe (Parcoblattini) of cockroaches. In the case of stanei, they cannot even be used to separate Species. However, Paratemnopteryx howarthi, which is cavernicolous, lacks pulvilli and clearly evolved from epigean Pararemnopteryx centrulensts (Roth) which has them (see below). MEMOIRS OF THE QUEENSLAND MUSEUM The parent species of P. stonet probably had pulvilli on all 4 proximal tarsomeres of all legs. and Race C with pulvilli on all its tarsomeres may be closest to the original stock. The differences in the degree of loss of the pulvilli in the various races may be due to the lengths of time the morphs have been isolated in their respective caves. The typical morphs, which lack pulvilli on all tarsomeres, may have been established in their caves the longest. There js a marked difference in the size of typical morphs of stonei fram Royal Arch Cave, and those of Race C from Pinwill Cave. Perhaps differences in the kind of available food in the 2 caves account for this variation. Paratemnopteryx glauerti (Princis) n. comb. (Figs 20A-F,21A,B,35) Franwalkeria glauerti Princis, 1954, p. 34, fig. 32(¢). Paratemnopteryx australis (nec Saussure): Tepper, 1896, p, 358 (misidentification). MATERIAL EXAMINED HoLortype: & (terminalia slide 8), Merredin, Western Australia; in WAMP. WESTERN AUSTRALIA. 2ILS: Norsemen, | d (reported by Princis. 1954, p.34), NoRTHERN TERRITORY, ANIC: 9km N of Kulgera, 25.468 1A3.18E, 1d (terminalia slide 82), 2.x.1972, KHL. Key et al. SAMA: Alice Springs, 1°, Horn Exped., 1894 (incorrectly reported as Paratemnop- ieryx australis Sauss., by Tepper, 1896, p.358 (label reads Pararemnopteryx australis Sauss. vat,?). QUEENSLAND. SAMA: Cunnamulla, 1¢, 19, H, Hardcastle, QMBA: *Morney’,, 120km W of Win- dorah, 1d, 24.ix.1983, G.B. Monteith. DESCRIPTION Male. Eyes extending slightly or not at all below level of antennal sockets, ocellar spots well developed (Fig. 20D). Tegmina and wings fully developed, extending beyond end of ab- domen; cubitus vein of hind wing with 3-4 com- plete and 1-2 incomplete branches, apical mia absent (Fig. 20F), Front femur Type Aa; pulvilli present on 4 proximal tarsomeres, tarsal claws symmetrical. simple, arolia absent. First abdominal tergum with setae along a ridge near anteromedial margin. those in middle partly covered by hind margin of metanotum (Fig. 20C). Seventh abdominal tergum unspecialised. Supraanal plate transverse, hind margin broadly rounded (Fig. 20A); right paraproct with a setose lobe (Fig. 20B), Subgenital plate symmetrical, 571 AUSTRALIAN PARCOBLATTINI ISLA “D S(MaIA [EHUGA) SOOJdesed pur ayeyd yeueeidng ‘g . “BULM puify Sy S(MaIA TEAI) SaTAIS (Mata jesiop) ayeyd jeueeidng ‘y tadAjojoy ‘, a S(mata [esare]) a4q “Gq . wn319) }eurMoOpge ‘(stoulg) n4aannps xtadoumaping “(Zz ‘Old MEMOIRS OF THE QUEENSLAND MUSEUM 572 “(Mora [eSIOP) eIye}TUEs pue ajeId TeENUSsqns “WM “JOATY DOSIyoINW JuRlBA, ‘P “gq ‘(AMAIA [RSIOp) eI[RIUWAs pur ayeyd jeywadgns ‘adAjojoy p “y “(stI9UUg) Yuannj/s xCajdoumajwing [Zz -D14 AUSTRALIAN PARCOBLATTINI 373 pilose, styles similar, erect, robust, wide apart (Fig. 20E). Genitalia as in Fig. 21 A: hook on left side, with a subapical incision; median phal- lomere rodlike, unmodified; right phallomere with cleft and spined sclerites. Colouration. Brown, Head with face darker brown than vertex. Terminal abdominal terga darker, supraanal plate with flattened hind border region yellowish brown (Fig. 20A), Coxae with basal regions dark brown, remaining parts of Jegs lighter. The male from Northern Territory is darker than the holotype, its head is uniformly dark brown, pronotal disk with light brown mark, posterior border zones darker than lateral regions, Female, Habitus essentially similar to females of couloniana, but somewhat larger. Tegmina reduced reaching only to T2; hind wings much smaller. Genitalia similar to that of couloniana, with right paratergite enlarged (as in Fig. 17G). Measurements (2 in parentheses). Length, 14.0-19.0 (19.5-20.5); pronotum length x width, 3.9-5.1 x 5.4-7.3 (S.5-5.7 x 7.7-8.5); tegmen length, 13.5-18.0 (8.2). ?VARIANT MATERIAL EXAMINED WESTERN AUSTRALIA. ANIC: Murchison River, 27.498 114.41E, 1d (terminalia slide $1), 28,411,197 1, E.F, Riek. DESCRIPTION Male. This specimen appears to be a very small glauertt, [tis dark reddish brown, head uniformly dark except for pale clypeus, supraanal plate unicolourous its hind border not yellowish. Styles and genital phallomeres similar to those of the type, the right phallomere showing minor differences (cp. Fig. 214,B). Measurements. Length, 13.2; pronotum length x width 3.6.x 5.1; tegmen length, 12.2. REMARKS Paratemnopteryx glauerti is very close to P. couloniana and can be distinguished from it by the shape of the suprasnal plate. However, it is essentially a coulortiana that has lost its arolia. The absence of arolia also separates the females of the two taxa. It is possible that the 2 females with reduced tegmina which I have determined as glaverti are female coufoniana that apparently lack arolia. Paratemnopteryx atra Princis (Figs 22A-H,35) Paratemnupleryx atra Princis, 1963, p. 11, fig. 2 (3 and ?); Richards, 1967, pp. 37, 38, 41. MATERIAL EXAMINED HoLotyee, ¢ (terminalia slide 6), Marble Bar, Wesiern Australia, 21.078 119.41E, 10.x.1957, col- lected deep in mines on piles of dung of the bat Macraderma gigas, A.M. Douglas; in WAMP. PARATYPES: WESTERN AUSTRALIA. WAMP: same data as holotype.4d ¢ (1 incorrectly labelled %), 12 (genitalia slide 7). ADDITIONAL MATERIAL: WESTERN AUSTRALIA. WAMP: same data as holotype, 12,19 nymph, DESCRIPTION Male. Eyes reduced, not extending below level of antennal sockets (Fig. 22B), ocellar spots present. Pronotum flat, parabolic (Fig. 22). Tegmina slightly reduced, width normal, taper- ing towards rounded apex, not reaching end of abdomen (Fig. 22A); hind wings vestigial, lateral, membranous, reaching between T1 and T2. Front femur Type A3, pulvilli and arolia absent, tarsal claws symmetrical, simple. First abdominal tergum with a medial group of sctae along anterior margin, completely or incom- pletely covered by metanotum (Princis incorrect- ly stated that all terga are unspecialised). Seventh abdominal tergum unspecialised. Supraanal plate trigonal, hind margin broadly rounded (Fig. 22D); paraprocts dissimilar, tight one with large setose lobe, left one with spinelike process (Fig. 22E). Subgenital plate symmetrical, convex, styles similar, symmetrically placed, wide apart (Fig, 22C). Genitalia as in Fig, 22F: hook on left side with subapical incision; median phallomere todlike, apex unmodified; right phallomere with spined sclerite. Colouration. Black or black with reddish tinge. Female. Supraanal plate with hind margin con- cavely excavated (Fig. 22G). Genitalia as in Fig. 22H: right paratergite with an enlarged swelling (arrow). Measurements (9 in parentheses). Length, 19.0-25.5 (23.5-26.5); pronotum length x width, 7.3-7.7 x LO.0- 10.4 (&.1-8.8 x 11.7-12.5); teg- men length, 16.4-18.0 (17.2-17.3). REMARKS This, a true guanobe, is the largest species of cavernicolous Paralemnopleryx, 1Lis closely re- lated to P. couloniana. 574 MEMOIRS OF THE QUEENSLAND MUSEUM =e SS ——— 1mm FiG. 22. Paratemnopteryx atra Princis {rom Marble Bar, W.A.; A-F, dd: A, habitus; B, Eye (lateral view); C, subgenital plate and styles (rear-oblique view); D, Suptraanal plale (dorsal view); E, Paraprocts (ventral view); F, Genitalia (dorsal view); G, H, Females: G, Supraanal plale (dorsal view); H, Genitalia (ventral view; arrow indicales expanded region of right paratergite), E, F, holotype, olhers paralypes. AUSTRALIAN PARCOBLATTINI Ss? Paratemnopteryn australis Saussure (Figs 23A-D,35) Peratemnopteryx australis Saussure, 1869, p. 273, pl, 3. fig. 22 (2), Walker, 1871, p. 18; Tepper, 1894, p. 170; Kirby, 1904, p. 106; Shelford, 1908b, p. 9; Alexander, 1917, p. 98; Princis, 1954, p. 35; Roth, 1968, pp. 84, 110, fig. 141 (ootheca), HOLOTYPE (nol examined) 2, Melbourne (Victoria), Australia; the type ap- parently is not in the MHNG; I received only 12 specimen trom the Geneva Museum, supposedly of this species, but il was from Peak Downs, Queensland. MATERIAL EXAMINED WESTERN AUSTRALIA. WAMP: 11km SSE of Ban- jiwarn HS., 27.428 121.37E, 12 (carrying rotated ootheca), ex pitfall trap, 22-287/,.1980, TF. Houston et al,, Biol. Survey Sile BWRI, 6.8km NNW of Mt Linden, 29.195 |22.25E (with detached ootheca), 1°, 17-23.10.1979, T.F, Houston et a/., Biol, Sutvey Site YMR2; Yandil, 12, 1928 (reported by Princis, 1954, p.38); Kathleen Valley, 12, 1963, T. Moriarty; Bare row Island, ex Nusutitermes triodiae nest, 1d (tet+ rinalia slide 11), 12.v,1982, C.N. Smithers and W.H, Butler, 19 nymph, 16.11.1982, C.N, Smithers, DESCRIPTION Male. Eyes reduced, not extending below an- tennal sockets, interocular space less than dis- lance betWeen antennal sockets. Tegmina reduced in length reaching to about hind margin of T2, width normal, touching at midline, distal margin truncate, corners rounded; hind wings vestigial, narrow, lateral, reaching to about hind margin of Tl. Front femur Type Az; tarsal claws simple, symmetrical, pulvilll and arolia absent. First abdominal tergum with a low setae medial- ly on anterior margin, mostly hidden by overlap- ping hind margin of metanotum, only their tips visible. Seventh abdominal tetgum un- specialised. Supraanal plate with sides concave, tapering, hind margin shallowly concave, corners rounded, a distinct transverse groove near basal margin (Fig. 23A); right paraproct much larger than left, composed of 3 sclerites. one of them roundly bulbous and partially spined (Fig. 23C). Subgenital plate symmetrical, sides upturned, styles clongate, similar, upright (Fig. 23B), each located in posterolateral corners of plate, interstylar margin straight (Fig. 23D). Genitalia as in Fig. 23D: hook on left side with subapical incision; median phallomere slender, apex slightly enlarged, rounded; right phal- uw lomere With a cleft sclerite and small number of spines or setae. Colouration. Dark reddish brown. Female. Supraanal plate trigonal, apex rounded. Tegmina, hind wings, legs, and colouration as in male, Measurements ( in parentheses). Length, ? (16.9-20.5); pronotum length x width, ? (4.7-5.8 x 6.3-8.4); tegmen length, 6.5 (4.5-6.0). The following is Saussure's description of the female of P. australis. Head smooth. Pronotum semicircular, slightly convex, with traces of grooves, lateral borders deflexed, posterior angles slightly rounded, hind border weakly ar- ched, slightly projecting medially. Tegmina ter- minating on second abdominal} tergum and touch or cross each other along their inner borders, apex attenuated, rounded, nearly semicircular at apex; veins visible, anal field elongated reaching to four fifths of sutural border showing 6-7 axil- lary veins; anal groove strongly sinuate. Wings tudimentary. Legs greatly compressed, tarsi clongated and spined over their lengths, Arolia absent. Hind borders of abdominal terga straight, lateral borders weakly serrated, Supraanal plate triangular, apex slightly prolonged and rounded. Cerci large, flattened, fusiform. Colouration. Brownish black. Antennae hrownish black provided with gray hairs. Head with mouth, occllar spots, antennal sockets. brownish yellow. Tegmina chestnut brown. Legs and abdomen basally brown to brownish yellow, tibiae darker brown, Measurements. Length, 16.0; pronotum length ~ width, 5.0 x 7.0; tegmen length, 6.5. The female from the Geneva Museum which wus sent to me as Paratemnepleryx australis, from Peak Downs, Queensland, is smaller than Saussure’s specimen from Melbourne, or the material from Western Australia and is as fol- lows: Eyes reduced, reaching level of antennal sock- ets, interocular space slightly greater than dis- tance between acellar spots. Front femur Type Aa; pulvilli and arolia absent. Tegmina reduced, width normal; hind wings vestigial, reaching hind margin of metanotum, Supraanal plate trigonal, apex rounded, Head dark brown, clypeus and labrum lighter brown, Pronotum reddish brown. Tegmina lighter reddish brown. Abdominal terga and cerei blackish, with metal- lic sheen. Legs reddish brown, cerci darker. Measurements, Length, | 1.5; pronotum length x width, 3.5 x 3.0; tegmen length 4.3. 576 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 23. Paratemnopteryx australis Saussure, d from Barrow Island, W.A.: A, Supraanal plate (dorsal view); B, Subgenital plate and styles (rear-oblique view); C, Supraanal plate and paraprocts (ventral view); D, Styles and genitalia (dorsal view). AUSTRALIAN PARCOBLATTINI REMARKS The male from Western Australia which Prin- cis determined as Paratemnopteryx australis, and which I have described here as that species, differs from the male of Paratemnopteryx rufa, principally in the shape of the supraanal plate, paraprocts, and in the apex of the median genital phallomere. The females are similar and I doubt if they can be distinguished, unless collected with males. Colour is of little significance because it can be so variable. In general the specimens which I have described as rufa are larger than those of australis, but here too size in this group of cockroaches can be extremely varible. It is possible that australis and rufa are con- specific. They clearly are very close taxonomi- cally. Establishing cultures of specimens from different localities, especially topotypic material, and studying their biology, would help in deciding whether rufa is a valid species. Paratemnopteryx rufa (Tepper) (Figs 24A-G,25A-F,35) Periplaneta rufa Tepper, 1893, p. 101 (¢); in Horn, 1896, p. 363. Blatta rufa (Tepper): Kirby, 1904, p. 139; Shelford, 1910, p. 16. Blatta tepperana Gurney, 1942, p. 24 (footnote); Prin- cis, 1969, p. 733. Paratemnopteryx rufa (Tepper): Princis, 1954, p. 35 (2). Paratemnopteryx zeitzi Tepper, 1895a, p. 20 (¢d). N. Syn. Paratemnopteryx zeitzi Tepper: Princis, 1969, p. 733. MATERIAL EXAMINED HOLOTYPE: ¢ (terminalia slide 49), Oodnadatta, Central Australia (=South Australia), F. Juncken; in SAMA. SOUTH AUSTRALIA. SAMA: ¢ (terminalia slide 50) lectotype (here designated) of Paratemnopteryx zeitzi Tepper, L.(ake) Callabonna, far north South Australia, A. Zeitz, 1893. 22,0? 21.0 36d 23.0-24.0 ee 29.-30.0 New South Wales @ 26.0 3 holotype rufa 3 lectotype zeitzi Queensland 577 NEw SouTH WALES. ANIC: near Milparinka, 12, 5.i.1967, collected as nymph, xi.1966, under rock in 7" rainfall country, M.F. Day. QUEENSLAND. QMBA: 30km east of Betoota, SW Qld, 3d 5,32 9, 25.ix. 1983, G.B. Monteith. DESCRIPTION Male. Eyes reduced not extending below an- tennal sockets, interocular distance greater than space between small ocelliform spots and anten- nal sockets (Figs 24B,25B). Pronotum smooth, parabolic, weakly convex (Figs 24A,25A). Teg- mina normal in width, reduced in length reaching to about hind margin of T2, distal margin obli- quely truncate or subtruncate, veins visible; hind wings vestigial, lateral, membranous, not reach- ing hind margin of T1 (Fig. 25A). Front femur Type A3; pulvilli absent or subobsolete (in ventral view pulvilli may be represented by light- ly sclerotised areas bordered by several large spines, at the apices of 4 proximal tarsomeres), tarsal claws simple, arolia absent. First ab- dominal tergum with few short setae arranged in longitudinal row near anterior margin of seg- ment, hidden by hind margin of metanotum. Seventh abdominal tergum unspecialised. Supraanal plate transverse, hind margin subtrun- cate (Figs 24E,25C), or weakly convexly rounded: paraprocts dissimilar as in Fig. 24E. Subgenital plate almost symmetrical, styles, elongate, upright, partially covered by dense, dark setae, interstylar margin almost straight (Figs 24C,D,F,25D,E). Genitalia as in Fig. 24G,25F: hook on left side with a subapical incision; median phallomere slender, apically unmodified; right phallomere with a cleft and setose sclerite. Colouration. Shiny, reddish brown or dark brown to blackish brown. Female. Supraanal plate trigonal, apex broadly rounded. The specimen from near Milparinka, NSW is light chestnut brown. Measurements. See Table 4. 10.3-10.8 10.2-11.0 10.2 TABLE 4. Comparative measurements of Paratemnopteryx rufa. 578 MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS Tepper’s request he sent him a single male on According to Tepper (1893, p.102), ‘Mr. F. which the description of rufa was based. Juncken, ... mentioned in one of his letters that Presumably if these cockroaches were annoying, cockroaches proved rather annoying.’ At they are found in houses. But it is doubtful that Fic. 24. Paratemnopteryx rufa (Tepper), d holotype: A, Pronotum and tegmina; B, Eyes (lateral and frontal views); C, Subgenital plate and styles (dorsal view); D, Supraanal and subgenital plates (dorsal view); E, Supraanal plate and paraprocts (ventral view); F, Styles (ventral view); G, Genitalia (dorsal view). AUSTRALIAN PARCOBLATTINI 579 the ‘annoying’ cockroaches were Paraltemnop- leryx. A second specimen (in SAMA), from New Hebrides, which Tepper identified as Periplaneta rufa, actually is 4 male nymph of Periplaneta americana which is an annoying domiciliary pest. Superficially nymphs of P. americana resemble Paratemnopteryx rufa, but they have well developed pulvilli and arolia, and the male subgenital plate and styles are typical of a Periplaneta. Fic. 25, Paratemnopteryx rufa (Tepper). d lecltotype of Paratemnopteryx zeitzi Tepper: A, Thorax, tegmina (left one removed to show vestigial hind wing), and first3 abdominal terga; B, Eye (lateral view); C, Supraanal plate and paraprocts (ventral view); D, Distal part of subgenital plate and Styles (dorsal view); E, Styles (rear view): F, Genitalia (dorsal view). 580 Paratemnopteryx sp, 1 (Fig. 26A-C) MATERIAL EXAMINED QUEENSLAND. ANIC: The following were collected by S.R. Morton: 10km north of Sandringham Stn Homestead, at seed bait, Sandridge, 24d (1 with terminalia slide 49), 20.i.1980, 1d, 1%, 21,i1,1980; 10km west of Sandringham, 1 9, 12.i,1979; Sandrin- gham Sin, 19 nymph, 24.1.1979. OMBA: Birdsville, SW Old, ex pitfall trap 2d ¢, | nymph, 11-17.xi.1976, D. McGreedy; Canterbury, 90km W of Windorah, southwest Qld, 12, 27,ix.1983, G.B. Monteith. i mm MEMOIRS OF THE QUEENSLAND MUSEUM NORTHERN TERRITORY. ANIC: 15km south of Alice Springs, bat caves, pitfall trap 60m from entrance, 12, 2 nymphs, 15,vi.1976, G. Griffin, 26km WSW of Mulga Park H.S., 26.008 131.25E, 1 © (genitalia slide 83), 18.1.1982; D.C.F. Rentz, B.G.F. Rentz, and R. Honeycutt, Stop 10. DESCRIPTION Male. Eyes reduced not reaching below level of antennal sockets, interoculart space slightly greater than that between sockets, Pronotum sub- parabolic. Tegmina reaching to T2, width nor- mal, veins distinct. Hind wings vestigial Fic. 26, Paratemnopteryx sp, 1, 3d from 26 km WSW of Mulga Park H.S., Northern Territory: A, Supraanal plate and paraprocts (ventral view); B, Distal parl of subgenilal plate, and styles (dorsal view); C, Genitalia (dorsal view). AUSTRALIAN PARCOBLATTINI reaching Tl, Front femur Type A3 (rarely Aq); arolia absent, pulvilli variable, absent or subob- solele on tarsomeres 2 to 4. First abdominal tergum with row of short setae along anterior margin medially, covered by hind margin of metanotum. Seventh abdominal tergum un- specialised, Supraanal plate broadly rounded, tight paraproct with sctose plate (Fig. 26A). Sub- genital plate symmetrical, styles short, widely spaced, erect or curved ventrad around hind mar- gin of plate, interstylar margin weakly concave or straight (Fig. 26C). Genitalia as in Fig. 26C, Colouration, Light reddish brown. Female. Supraanal plate trigonal, broadly rounded. Females may be much darker than males. The female from Canterbury is the darkest with head ‘black except for yellowish brown clypeus, brown labrum; pronotum black; teg- mina dark reddish brown; abdominal terga and cerci black, abdominal sterna and legs reddish brown. Measurements (2 in patentheses), Length, 18.0-19.7 (18.0-22,0); pronotum length x width, 4.8-5.8 x 6.5-8.0 (5.2-6.1 x 7.2-8.5); tegmen length x width, 5,7-6,3 x 4,3-5,2 (5,1-8.3 x 4,4-5,5), REMARKS This specivs appears to combine characters of australis and rufa. lts size, and shape of male supraanal plate are more like those of australis (Fig. 23C) although it is not indented medially. The right paraproct and median and right genital phallomeres are similar to rufa (Fig. 24E,G). This may prove to be a distinct taxon, However, I have not seen the type of australis and, at least, topotypic specimens (Melbourne) should be ex- amined before deciding this question, Paratemnopteryx broomehillensis n, sp. (Figs 27A-G,28A,B,35) MATERIAL EXaMINep HoLotyre; &. Browmehill, Western Australia, 33.515 117.386, MV light trap, 91,7985, RP) Me- Millan; in WAMP. PARATYPE: WESTERN AUSTRALIA. ANIC: Wanneroo, (3, (lerminaliaslide 136), 1.1.1964,G.5. McCutcheon, DESCRIPTION Male. Eyes fully developed extending below level of antennal sockets; interocular space less than distance between antennal sockets. Pronotum with literal margins posterior to mid- dle weakly ablique (Fig. 27G). Tegmina and wings fully developed extending well beyond 381 end of abdomen; cubitus vein of hind wing with 3-4 complete and 4 incomplete branches, apical triangle absent. Front femur Type Ag; pulvilli present on 4 proximal tarsomeres, tarsal claws simple, symmetrical, aroha present. First ab- dominal tergum with large anteromedial transverse depression bordered by rounded clevation bearing very small spaced setae, these also present on either side of depression (sctac not in dense group) (Fig. 27C,D). Seventh ab- dominal tergum with large group of dense, black setae in shallow depression medially on anterior half of segment, partially covered by T6 (Fig. 27A.B). Supraanal plate produced, sides con- cave, hind margin with deep U-shaped excava- lion forming pair of apically rounded lobes bearing long slender setae dorsally, densely covered with small dark spines ventrally (Figs 27B,28A); ventral intercercal ridge absent, or at most represented by transverse, unpigmented, shallow elevation; right paraproct with very large spinclike process. (Fig. 28A). Subgenital plate symmetrical, styles similar, elongated, tapering, upright, bearing small, densely packed dark spines on one surface, each style located in posterior corners of plate, below cerci, interstylar margin convex (Figs 27B,E,F,28B). Genitalia as in Fig. 28B: hooklike phallomere on Jeft side, with subapical incision; median phallomere rod- like, apex acute; right phallomere with cleft-like sclerite, Colouration, Light brown, Pronotum hyaline on posterior half lateral to disk, and along hind border, Female. Unknown, Measurements. Length, 21.0-23.9, pronotum length x width, 5.7-6,0 x 7,2-7.6; tegzmen length, 22.1-23.5, REMARKS The general appearance of broomehillensis is that of a Nedlemnopteryx, but the shape and position of the styles-are typical of most species of Paratemnopteryx, The apex of the median genital phallomere resembles that found in P. centralensis (Fig. 29B). The shape of the supraanal plate of broamehillensis readily distin- guishes it from all other species of Paratemnop- teryx, or of Neotemmoapteryx. Paratemnopteryx sp. 2 MATERIAL EXAMINED NEW SOUTH WALES. Lord Mowe Island, Ned's Beach, LY (genitalia slide 84), xii. 1972. 7 Liepa 582 MEMOIRS OF THE QUEENSLAND MUSEUM 1mm 1mm Fic. 27. Paratemnopteryx broomehillensis, n. sp. ,d 6: A, Abdominal terga 6 to 10 (supraanal plate), and subgenital plate (part of T6 removed to show setal gland on T7); B, Abdominal terga 7 to supraanal plate; C, First abdominal tergum; D, Glandular region on T1; E, Supraanal and subgenital plates (rear view); F, Subgenital plate and styles (ventral view); G, Pronotum. Localities: A, C, E, G, paratype from Wanneru, W.A.; B, D, F, holotype from Broomehill, W.A. DESCRIPTION Female. Eyes reduced, interocular space greater than distance between small, distinct ocel- lar spots, and antennal sockets. Tegmina reduced in length, width normal, reaching to T7, tapering to a rounded apex. Hind wings slightly smaller, reaching to T6, cubitus vein with 4 complete and 2 incomplete branches, apical triangle absent. Front femur Type A3; tarsal claws simple, sym- metrical, pulvilli absent, arolia very small. AUSTRALIAN PARCOBLATTINI \ 25 me Fic, 28. Paratemnapteryx broamehillensis, n- sp. 3 paratype from Wanneroo, W.A.; A, Supraanal plate and paraprocts (ventral view); B, Styles and genilalia (dorsal view), Colouration. Head reddish brown, lighter on vertex and clypeal region. Pronotum and tegmina light reddish brown. Wing with yellowish tinge. Abdominal terga and sterna dark brown, Legs light brown. Measurements. Length, 13.2; pronotum length x width, 4.4 x 6.8; tegmen length, 9.0. Paratemnopteryx sp. 3 MATERIAL EXAMINED QUFENSLAND. ANIC: Noccundra, | 2, 8.xi.1949, ELF. Riek. 583 DESCRIPTION Female. Interocular space less than distance between ocellar spots, about same as space be- tween antennal sockets. Pronotum. transverse, anterior margin straight, hind margin weakly convex, lateral margins obliquely convex. Teg- mina reduced in length reaching T2, width nor- mal, apex rounded, veins distinct. Hind wings vestigial, lateral, reaching hind margin of T1. Front femur Type Ag; pulvilli present on fourth tarsomere only, tarsal claws symmetrical, simple, arolia absent. Supraanal plate trigonal, broadly rounded. Colouration, Head dark reddish brown, ocellar spots distinct, yellowish white, clypeus yel- lowish, labrum light brown. Pronotum reddjsh brown, disk lighter. Tegmina light reddish brown. Abdominal terga with basal 2 segments light reddish brown, remainder blackish. Ab- dominal sterna reddish brown becoming darker on posterior segments. Cerci blackish. Legs light brown. Measurements. Length, 16.0; pronotum length x width, 4.5 x 6.8; tegmen length, 6.5. Paratemnopteryx sp. 4 MATERIAL EXAMINED QUEENSLAND, QMBA; Undarra, Bayliss Cave, on floor beyond Duckunder, 19, 19 nymph, 23.v.1985, F.G. Howarth, D, Irwin, and F.D. Stone. DFSCRIPTION Female, Ocelli and eyes absent. Tegmina reduced in length, width normal, reaching to about hind margin of T3, anterior margin strong- ly convex, apex rounded, Hind wings vestigial, narrow, lateral, reaching to hind margin of T2, some venation visible. Femur Type Ag; pulvilli and arolia absent. Supraanal plate trigonal, lateral margins shallowly concave, apex rounded, Colouration. Pale yellowish brown. Measurements. Length, 21.5; pronotum length x width, 6,7 x 7.5; tegmen length, 9.3. Nymph, Resembles female except for size (length, 16.5mm), and absence of tegmina and wings. REMARKS The habitus of this female resembles that of Neatemnopleryx douglasi, but that species (known only from the male) has eyes (variably reduced), and small arolia, pulvilli essentially absent. The absence of cyes indicates it is 5k restricted to a cave habitat. Paratemnapteryx stonei also has been found in Bayliss Cave, but this specimen clearly is not that species, Parutemnopteryx centralensis (Roth) n. comb. (Figs 294,B,35) Symploce centralensts Roth, 1985b, p, 322, figs ISA- I8G, 19-190 (¢ and ?). MATERIAL EXAMINED Holotype and paratypes from Irian Jaya and Papua New Guinea Jisted in Roth (1985b). ADDITIONAL MATERIAL FROM — AUSTRALIA, QUEENSLAND. ANTC: Duanbulla Forestty Reserve, 13km NE by N of Yungaburra, 17.10S 145.39E, 1d (terminalia slide 106), 16.xi. (981, J. Balderson; 3km NEof M1 Webb, near Cooktown, 24 3, 3.x,1980, T.A. Weir and R.A. Barrett. The following were collected by LF.B. Common and M.S. Upon: 1 mile N of Kuranda, 1200ft, 1, 23.iv.1969; 3 miles W of Mossman, 1@. 14.111,1964, 1d, 13.i71,1964; 9 miles N of Kuranda, 1d, 12.11.1964; 9 miles E of El Arish, 1d, 711, 1964. BPBM: Cairns, 39.4, J.F. Mlingworth. OMBA; Flying Fish Pt, 2d &, 21,1.1965, E.C, Dahms. The following were collected by G.B, Monteith; 3km F of Lockerbie, C, York, 14, at MV Light, 30.i.- 441.1975; Lockerbie Area, Cape York, 1d, 13- 27.iv.1973. NoRTHERN TERRITORY. BPBM: Darwin, 1d (ter- minalia slide 459), M.V. Light Trap, 17.i,1964, J. Sedlacek. REMARKS Previously known only frum the island of New Guinea, this species was fully described by Roth (1985b, p.322). The cpigean species varics great- ly in size and colour. Specimens from Australia are generally smiuller, and some are darker brown than New Guinca specimens and very dark brown individuals may have a metallic sheen. The vyes are very large (Fig. 29A) and close logether. Pronotum and tegmina usually are covered with spaced upright hairs. Male supraanal plate, paraprocts, subgenital plate and minute styles, and genital phallomercs (Fig. 29B) are similar in Australian and New Guinea Specimens (Roth, 1985b, figs. 18,19), As indi- cated below, centralensis probably was the species trom which the cavernicolous howartht arose. Measurementx (do only, New Guinea specimens in parentheses from Roth 1485b); length, 9.5-14.4 (12.3-16.0); pronotum length x MEMOIRS OF THE QUEENSLAND MUSEUM width, 2.6-3,2 x 3,4-4.2 (3,0-3,7 x 3,9-4,9); teg- men length, 10.8-16.0 (14.2-20.0). Paratemnopteryx howarthi n, sp. (Figs 30A-J,35) MATERIAL EXAMINED Hocorvee: 4, Chillagoe, Queensland, Christmas Pot Cave, 29.vi.1984, F.D. Stone and D, Irvine, Explorers Club Expedition; in QMBA. PARATYPES! QUEENSLAND, QMBA: All specimens were collected on the Explorers Club Expedition in 1984 to the Chillagoe Caves, Queensland. The follow- ing are all brachypterous morphs unless indicated as macropierous (macr,); Tea Tree Cave, 1d (macr,), 1d (enlire Inseet mounted on slides 48a and 48b), 12 (genitalia slide 49), 2.vii., 1d, 24.vi., F.G, Howarth, 1%] ootheca, 2nymphs, 24.vi., 1d, 3 nymphs, 29.vi., F.G. Howarth, and F.D. Stone . 1d, 24.vi., F.D.Stone et al; Christmas Pot, 1¢, 22 2, 2 nymphs (swiftlet guano), 5 nymphs, 29.vi., F.D. Stone and D. Irvine, (1d (terminalia slide 240) and 1%, brachypterous paratypes retained in MCZH). ETYMOLOGY The species is dedicated to Dr Francis G. Howarth of the Bernice P. Bishop Museum, DESCRIPTION Male, Eyes well developed in macropterous morphs extending well below level of antennal sockets (Fig. 30A), but somewhat reduced in brachypterous forms (Fig. 30B), ocellar spots absent; Interocular space Jess (macr.) or greater (brachypterous) than distance between antennal sockets. Tegmina and wings fully developed ex- tending well beyond end of abdomen, or tegmina reduced in length (normal width), reaching to about middle of T5 or its hind margin. Hind wings smaller, membranous, reaching to about hind margin of T2 or T3; in fully developed wings, discoidal, median and cubilus veins straight, the former with 2-3 complete and 2-3 incomplete branches, apical triangle absent (Fig. 30F). Anteroventral margin of front femur Type As; pulvilli absent, tarsal claws simple, sym- metrical, arolia very small. First abdominal ter- gum with large, dense group of setae near anteromedial half of segment (Fig. 30D), Seventh abdominal tergum unmodified. Supraanal plate transverse, trigonal, right and left paraprocts dissimilar (Fig, 30C), Subgenital plate almost symmetrical, styles minute, the right somewhat larger with long curved setae, the left one cylindrical, cach located near posterior AUSTRALIAN PARCOBLATTINI 0.5 mm tn oo Ww Fic. 29. Paratemnopteryx centralensis (Roth), 3 from Danbulla Forestry Reserve, Qld; A, Eye (lateral view); B, Genitalia (dorsal view). corners of platc, interstylar margin straight (Fig. 30E). Genitalia as in Fig. 30G; hook on left side, median phallomere a slender curved rod with acute apex, right phallomere with an associated setal brush. Colouration, Light to dark brown. Lateral regions of pronotum hyaline. In dark morphs, the head may be much darker than abdomen, al- though the pronotum also may be relatively dark. Tegmina hyaline even in darkest forms. Measurements. Macropterous morphs in brackets, all others including holotype (in paren- theses), are brachypterous: Length, 9.5-11.0 (10.5) {10.0-10.5]; pronotum length x width, 2.6- 2.8 x 3.0-3.5 (2.9 x 3.1) [2.8-2.9 x 3.1-3.2]; tegmen length, 4.5-5.7 (5.6) [11,0-11.2]. Female. Eyes. narrower and slightly smaller than in brachypterous males, usually not extend- ing below level of antennal sockets. Tegmina and wings reduced reaching to T3 or hind margin of T4, wings smaller extending to about middle of T1 or T2. No macropterous females were seen. Supraana! plate distinctly triangular (Fig. 30H). Right genital paratergite with a small basal swelling (Fig, 301). An empty ootheca (Fig. 30J) is brown with 12 egg cells, 12 shallow keel serrations, and measures 2.5mm high and 3,5mm long, Measurements, Length, 8.5-12.3; pronotum length x width, 2.7-3,1 x 3,1-3.5; tegmen length, 3.9-6.1. Nymph. The eyes of small individuals are greatly reduced and are represented by a narrow strip of dark ommatidia. These increase in num- ber in large nymphs where the eyes almost reach adult size. Arolia and pulvyilli absent. REMARKS Paratemnopteryx howarthi is very close to, and probably arose from the epigean species Paratemnopteryx centralensis (Roth) (=Symploce centralensis Roth) from Irian Jaya, Papua New Guinea (Roth, 1985b, p.322) and Australia. The shapes of the styles and their position on the subgenital plate are similar in both species. Some differences are seen in the shapes of the supraanal plate (in both sexes), and in the apex of the hooklike genital phallomere in the males. The male paraprocts and right genital phallomere appear to have some small differen- ces, but this may be due in part to their orientation on the slides. Paratemnopleryx centralensis 586 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 30. Paratemnopteryx howarthi, n. sp., paratypes. A- G, d d: A, Eyes (lateral views) of macropterous and brachypterousu specimens respectively; C, Supraanal plate and paraprocts (ventral view); D, Setal specializa- tion on first abdominal tergum; E, Distal region of subgenital plate, and styles (dorsal view); F, Hind wing; G, Genitalia (dorsal view); H, 1, Females: H, Supraanal plate (dorsal view); I, Rignt paratergite; J, Ootheca, Localities: F, Christmas Pot Cave, all others, Tea Tree Cave, Chillagoe, Qld, FiG. 31. Paratemnopteryx suffuscula n. sp., dd. A, Paratype from Mt Boppy, N.S.W., head; B, Holotype, habitus; C, D, Paratype from Mt Arthur, N.S.W., supraanal plate and paraprocts (ventral), and subgenital plate and genitalia (dorsal); E, F, Paratype from same localily as holotype, right tegmen and hind wing. AUSTRALIAN PARCOBLATTINI 587 varies considerably in size, but generally it is larger than howarthi. The presence of both brachypterous and mac- Topterous morphs of P, Aowarthi in the same eave is of interest. Macropterous individuals have larger eyes. than brachypterous morphs, but both forms lack ocelli and pulvilli, The correla- tion of normal sized eyes with macropterous morphs, and their reduction in brachypterous forms has been shown in another cavernicolous cockroach, A//uaudellina cavernicola (Shelford) (Chopard, 1932). This species is found both in- side and outside of caves and Chopard arranged individuals in the following categurtes: 1) muac- ropterous and macropthalmic; 2) macropterous and micropthalmic; 3) micropterous and microp- thalmic. Chopard concluded that in Alfuaudel- lina, ocular reduction precedes wing reduction, The eye and wing reduction in P. Aowarthi does not reach the extreme losses found in Al- luaudellina, at least in the specimens | have seen. | have seen one male specimen which | am provisionally referring to P. howarthi, and which was not taken in a cave [24km N by W of Mareeba, Queensland, 16.47S 145.22E (ter- minaha shde 141, 24-25.%i,1981, J. Balderson (ANIC)]. ft has fully developed legmina and wings but its cyes are reduced and widely separated and has larger ocellar spots. There are small pulvilli on the 4 proximal larsomeres and the arolia are small, Its pronotur measures 2,4 x 3.0mm, and the tegmen length ts L0.2mm. IF this specimen is howarthcil may be an epigean form of the taxon. Paratemnopteryx suffuscula n. sp. (Bigs 31A-F35) MATERIAL EXAMINED Ho.otyre. 6, 1! miles S by W of Cocklebiddy, Western Australia, 32.128 126.036, 72.21.1969, Key and Upton, in ANIC, PARATYPES, WESTERN AUSTRALIA, ANIC; same data as holalype, 1d (legmen and wing slide 214) (Key's field nutes, trip 163, Stop 18683,6); Bullsbrook, Id, atlight, 4-xti.1978, K. and E, Carnaby; Python Pool, ne Mi Herbert, S of Roebourne, 24 d, 28.vili.1964, P,B. and L.C. Carne. The following were collected by NLS, Upton and JE. Feehan: 1km SSW of Millstream AS., 21,358 117.04E, 1d, 30.x.1970; 15km E of Millsiream H.S,, 21.355.117,12E, 1d (terminalia slide 190), 20.%,1970, WAMP: 70-75km ENE of Nor- seman, 1d, 10-16..1.1978, T.P, Houston ef al, SourH Australia. ANIC: Buckaringa Gorge, MEMOIRS OF THE QUEENSLAND MUSEUM Flinders Range, c, 20km NNW of Quorn, Ld, 19.17, 1985, €, Reid, P.W. Gullan, M. Lewis, New SouTn Wates, ANIC; Mount Boppy, 304, 25.xi.1949, E.F. Riek, Mt Arthur, nr Wellington, bd (terminalia slide 142), E.F, Riek. DESCRIPTION Male. Head exposed, cyes. large, extending well below antennal sockets, interocular distance about same as space between antennal sackets (Fig. 31.A); third and filth palpal segments longer than fourth. Pronotum suboval with a pair of depressions on distal half (Fig. 31B). Tegmina and wings fully developed extending well beyond end of abdomen, discoidal sectors of former, oblique (Fig. 31B,E), veins with short, upright hairs. Hind wing with subcosta reaching beyond middle, discoidal and median veins straight, cubitus vein straight with 2-3 complete and 2-3 incomplete branches, apical triangle ab- sem (Fig. 31F). Front femur Type Ag; pulvilli subobsolete or absent on proximal tarsomeres 1-3, present on fourth tarsomere, tarsal claws symmetrical, simple, arolia small. Supraanal plate with hind margin convexly rounded, not teaching hind margin of subgenital plate; right and left paraprocts dissimilar, right one with a plate bearing spines, and a long spinelike process (Fig, 31C), Subgenital plate with pair of widely separated similar, cylindrical styles, interstylar margin weakly convex (Fig. 31D). Genitalia as in Fig. 31D: hook on left side with preapical incision; median phallomere uniformly slender, apex unmodified; right phallomere very large, one of the plates bearing a row of large, stout, setae, Calouration. Brown, Head dark reddish brown, large ocellar spots white, clypeus yel- lowish white, labrum reddish brown, distal part lighter (Fig. 31A); last palpal segment light brown, other segments mostly pale. Pronotum with disk yellowish brown or dark brown, sur- rounding area hyaline or subhyalinc. Tegmina reddish brown, hyaline (Fig. 31E). Hind wing with subcosta and costal vein regions and distal half of anterior field infuscated (Fig. 31F). Ab- dominal terga brown, latcral regions and ter- minal segments darker, Abdominal sterna brown, scgment on distal half, darker, Cerci blackish brown dorsally and ventrally, Coxae with basal halves or more dark brown, remainder pale: femurs mostly pale sometimes infuscated, tibiae and tarsi darker, * Female, Unknown: Measurements. Length, 9.0-11-5; pronotum AUSTRALIAN PARCOBLATTINI length x width, 1.9-2.3 x 2.5-2.8; tegmen length, 9,9-11.8. REMARKS The habitus of P, suffuscula resembles that of some species of Dyakinodes. Keyella n. gen. TYPE SPECIES Keyella gayi n. sp. ETYMOLOGY The genus is dedicated to Dr K.H.L. Key of CSIRO. DIAGNOSIS Eyes well developed or somewhat reduced. Tegmina and wings fully developed in both sexes, sometimes reduced in the female; cubitus vein of hind wing with complete and incomplete branches, apical triangle absent. Anteroventral margin of front femur with some large proximal spines followed by a row of piliform spinules and ierminating in 3 large spines (Type B3); pulvilli on 4 proximal tarsomeres of front and mid tarsi only. and only on fourth tarsomere of hind tarsus, or pulvilli on all tarsal segments; tarsal claws simple, symmetrical, arolia present. d: first ab- dominal tergum specialised. Seventh abdominal tergum with or without a specialisation. Intercer- cal ridge on ventral surface of supraanal plate absent. Subgenital plate symmetrical or nearly so. Two styles, weakly dissimilar. Hooklike genital phallomere on left side. REMARKS This genus is closely related to Neotemnap- leryy and Paralemnopteryx and belongs in the Blattellidae: Blattellinae, tribe Parcoblattini. It is readily separated from the above 2 genera by Type B3 front femur (vs. Type A3). The morphol- ogy of the male setal gland on T1 also differs distinctly from those found in the related 2 genera. Keyella appears to be a link between Neotemnopteryx and Paratemnopteryx; one of the two species is closet to the former and the other is closer to the latter. Distribution of the 2 species of Keyella is shown in Fig. 35. KEY TO MALES OF KEYELLA SPP. !, Supraanal plate trigonal (Fig. 32B), Seventh ab- 589 dominal tergum unspecialised. Styles short, stocky, rounded (Fig, 32E) 0... BOVE Supraanal plate subtrapezoidal (Fig. 33B,D). Seventh abdominal tergum with a pair of large medial depressions containing a large dense group of selae (Fig. 33B). Styles elongate, lapering (Fig. 33E) «0.2... armidalensis Keyella gayi n. sp. (Figs 32A-E,35) MATERIAL EXAMINED Hotoryee: J, ‘dried out of spirit’, near Cairns, Queensland, Australia, viii.1966, H.M. Cameron; in ANIC. PARATYPES: QUEENSLAND. ANIC: Lake Barrine, 1d (terminalia slide 134), 15.vii.1933, H.A, Gay; Davies Ck, 20km east by south of Mareeba, 17.028 145.37E, 19, 19,xi,1981, J. Balderson. QMBA: Shiptons Flat, 35km S of Cooktown, 250m, 14, 22.iv.1982, Mon- teith, Yeates, and Caok, ETYMOLOGY The species is dedicated io H.A. Gay who collected onc of the paratypes. DESCRIPTION Male. Eyes not extending below antennal sockets, inferocular space greater than distance between ocelli or antennal sockets. Tegmina and wings fully developed the former with un- branched discoidal vein, branches of median and cubitus rami longitudinal. Hind wing with dis- eoidal vein branched or unbranched (may differ on the right and left wings), median. and cubitus veins straight, the former simple, the latter with 2 or 3 complete (some may be bifurcate) and 1 or 2 incomplete branches, apical triangle absent. Anteroventral margin of front femur with 6 large spines on proximal half followed by tow of piliform spinules and 3 stout terminal spines (the first of these is only slightly longer, but stouter, than the spinules preceding it): pulvilli large on 4 proximal tarsomeres of front and mid tarsi, present only on segment 4 of hind tarsi; arolia very small, Modification on first abdominal ter- gum consisting of a pair of exposed, small, dense yellowish setal tufts, with a distinct space be- tween them, and on either side an elevation bear- ing long slender setae (Fig. 32A). Seventh abdominal tergum unspecialised. Supraanal plate trigonal (Fig. 32B), reaching hind margin of subgenital plate; right and left paraprocts weakly dissimilar (Fig, 32D). Subgenital plate symmetrical, convexly rounded, hind margin pale, membranous, upturned (Fig, 32C). Styles, short, stocky, right one larger, pale ventrally (a continuation of pale margin of subgenital plate), dorsal surface densely covered with small ap- pressed dark spincs (Fig. 32E). Genitalia as in Fig. 32E: hook on left side; median phallomere uniformly slender, rodlike, right phallomere well developed with cleft. Colouration. Dark reddish brown, Portion of right tegmen covered by left, hyaline, colourless. Hind wing with aniecrior margin darkly infus- cated, veins brown. Femaie. Tegmina and wings fully developed. Lind margin of supraanal plate less trigonal than male, convexly rounded, Pulvilli present on 4 proximal tarsomeres of all tarsi but small on first and second segments on hind tarsus, arolia very small. Measuremenis (2 in parentheses): Length, 17.3-17.7 (25.0 flattened); pronotum length x width, 5.0-5.1 x 3.7 (sides strongly detlexed) - 6,8 (5.1 x 6.9); tegmen length, 15.8-17.4 (17.0). REMARKS The Widely spaced styles tend to resemble those of same Paratemnopteryx spp. Keyella armidalensis n. sp (Figs 33A-E,35) MATERIAL EXAMINED Hovoryee: 3, alight, Armidale, New South Wales, Australia, 14.1.1963, C.W. Frazier, donated by Univer- sity of New England; in ANIC. PARATYPES: NEW SOUTH WALES, ANIC: sume data as holotype except for collection dates: 1 (terminalia slide 135), (3.811, 1959, 26 d, 2.1,1960, MCZH: Salis- bury Cho 1, xi, Wheeler, DeSCRIPTION Male. Eyes well developed extending below level of antennal sockets; interocular space same as distance between antennal sockets. Pronotum with sides rounded, hind margin weakly convex. Tegmina and wings fully developed. Tegmina discoidal vein with an apically forked branch neat middie, branches of median vein lon- gitudinal, those of the cubitus vein longitudinal and weakly slanted. Hind wing discoidal vein straight, with an apically forked branch beyond middle, median and cubitus veins straight, the former simple, the latter with 3-4 complete and 2-4 incomplete branches, apical triangle absent. Anteroventral margin of front femur Type B3 MEMOIRS OF THE QUEENSLAND MUSEUM with 4 large proximal spines; pulvilli present on 4 proximal tarsomeres of all legs; arolia present. Firs! abdominal tergum with a pair of small separated tufts of setae and longer setae along a ridge anterior to them (Fig. 33A). Seventh ab- dominal tergum with a pair of depressions medially, in which is a large dense group of setae (Fig. 33B). Supraanal plate with hind margin truncate, right and left paraprocts dissimilar (Fig. 33B,D). Subgenital plate almost symmetrical, convex, hind margin upturned, Styles very close together near midline of plate, elongated, some- whal triangular, dorsal surface densely covered with small spines; right style slightly shorter and more robust than the left one (particularly in the pinned specimen, (Fig. 33C), but appears less so in the slide preparation (Fig. 33E). Genitalia as in Fig. 33E: genital hook of left side, with a subapical incision, median phallomere slightly enlarged and acute apically, right phallomere reduced, ; Colouration. Light to dark brown, Female. Unknown, Measurements. Length, 9.5-12.2; pronotum length x width, 2.5-2,8 x 3,.7-3,8; tegmen length, 10,5-13.2. REMARKS The closeness of the male styles resembles those found in Neoetenmopteryx stvliparedra (Fig. 13E). Keyella sp. MATERIAL EXAMINED NEW SouTH WALES. ANIC: Durras North, near Batemans Bay, 12, 23.xi1, 1984, HM, Cameron, DESCRIPTION Female. Eyes teaching slightly below level of antennal sockets: fifth palpal segment swollen, distinetly larger than the fourth. Tegmina reduced in length, width normal, tapering apical- fy rounded reaching to hind margin of T3. Hind wings smaller reaching slightly beyond hind margin of T2, folded longitudinally down mid- dic, veins present, Supraanal plate trigonal, apex rounded. Front femur Type Bs, pulvilli present on 4 proximal tarsomeres of all tarsi, arolia present, tarsal claws simple, symmetrical. Colouration. Pronotum and abdominal terga dark reddish brown. Head, hyaline tegmina, cerci and legs light brown. Hind wings infuscated. Abdominal sterna light brown, its sides and sub- genital plate darker_ AUSTRALIAN PARCOBLATTINI 591 0.5 mm FiG. 32. Keyella gayi, n.sp., 6 paratype from Lake Barrine, Qld: First abdominal tergum and setal specializa- tion; B, Supraanal plate (dorsal view); C, Subgenital plate and styles (rear view); D, Supraanal plate and paraprocts (ventral view); E, Subgential plate, genitalia, and styles (dorsal view). 592 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 33. Keyella armidalensis, n.sp., 6 6. A-C, Holotype: A, First abdominal tergum and setal specialization; B, Abdominal tergum 7 (with setal gland) to supraanal plate; C, Subgenital plae and styles (rear view); D-E, Paratype from Armidale, N.S.W.: D, Supraanal plate and paraprocts (ventral view); E, Subgenital plate, genitalia, and styles (dorsal view). Measurements. Length, 10.0; pronotum length x width, 2.5 x 3.6; tegmen length, 4.2. REMARKS The supraanal plate of this specimen resembles that of male K. gayi and it may be a brachyp- terous female of that species. Males from Batemans Bay are needed for specific determina- tion. The unnamed female looks like a very small specimen of Paratemnopteryx couloniana (Prin- cis) but its Type B3 front femur distinguishes it from that species (and genus) which has Type A3. ACKNOWLEDGEMENTS I thank the museums and curators mentioned in the introduction for sending me the material. I am grateful to the Bureau of Flora and Fauna, Australian Biological Resources Study (ABRS) 593 AUSTRALIAN PARCOBLATTINI o> "elpergsny ut ‘dds x(zajdoumajoan jo uoNNGIsIg “pe ‘D1 811 DI78ND & pypodnt tq vsso175 psno14016 V7 pupu © papouoo @ vapeipvd27 As (] asp. bnop & po14d111728 © (zupi1apa) vaynf (quprapa) vyapoanfie & paynf 4 _ whiazdouwa ,0aN ae . m x MEMOIRS OF THE QUEENSLAND MUSEUM 594 “popnyjour jou ase eAeL URI] JOJ aso) SA]UO endeg 10} UMOYS aE sisuajoajuas xXsajdoumajv1vg JOJ Sp10991 BIUINH MAN ‘elRISNY Ul yjaday pue ‘vyjayH1qQoO/ 80.4], ‘xKajdouwawang JO satdads JO UOTINGIISIG “SE “D1 szeuezvprmsD oynosnf{fns * 821 D478ND @ 2AvB © rygavmoy [| szsuazvazuao 01 7.@Aey you0is @ 14aanv16 @ $18U2112YaMW00Cq FY p{nig S2L8UaTOqdD11NU & paiqv Ww pupiuejnoa @ 01194301q071bouag whrezdouwazpapg ° = % ® A * * @ ® ° g ® \ ® a) ® % 7 * iv ® y f is OT AUSTRALIAN PARCOBLATTINI for pate support, and Dr D.C, Rentz for his aid with the maps, LITERATURE CITED ALEXANDER W.B. 1917. List of orthopterous insects recorded from Western Australia, (1915-1916) JR. Soe. W. Aust. 2: 97-103. ARNOLD,.J.W. 1974. Adaptive features on the tarsi of cockroaches (Insecta: Dictyoptera), Jnt. J. Insect Morph. Embryol. 3: 317-34. Bry-Brenko, G. YA. 1950. Fauna of the U.S.S.R. Insects, Blallodéa. - Zool. Inst, Akad. Nauk, 5.S.8.R., Moskva, 0.5, 40, 342 pp. (Russian). BRUNNING, C.F.A, 1948, Studies in Malayan Blat- tidae. Zodl, Meded. Leiden 29: 1-174, BRUNNER VON WATTENWYL, C, 1862. Ueber die von der kk. Fregatre Navara mitgebrachten Orthop- leren. Verh. Zoal. - Bot. Ges. Wien 12: 87-96. 1865, “Nouveau Systéme des Blaltaires'. (G. Braumiiller: Vienna). CHoparD, L. 1932. Un cas de micropthalmie lige @ l'atrophie des ailes chez une blatte cavernicole, Soc. Ent, Fr. Livre du Centenaire, June 1932, pp. 485-96, CHRISTIANSEN, K. 1965. Behaviour and form in the evolution of cave collembota. Evolution 19: 529- 37. Gurney. A. 1942. Studies in Cuban Blattidae (Or- thoptera), Bull. Mus. Comp, Zool: Harvard 89; 11-60, HEBARD, M. 1917. The Blattidae of North America north of the Mexican boundary, Mem. Amer, Ent. Sae. 2: 1-284, 1920, The Blattidae of Panama. Mem. Amer. Ent. Soc. (1919) 4: 1-148. 1929. Studies in Malayan Blattidae (Orthoplera). Proc. Acad. Nat. Sci. Philadelphia 81: 1-109, 1943. Australian Blaltidae of the subfamilies Chorisoneurinae and Eclobiinae (Orthoptera), Acad, Nat. Sci. Philadelphia, Monag, 4: 1-129. Jouns, P.M. 1966. The cockroaches of New Zealand, Rec, Canterbury (N.Z.) Mus. 8: 93-136, Key, K.H.L. 1970. Principles of classification and nomenclature, pp. 141-67, Jn CSIRO, ‘Insects of Australia.” (Melbourne University Press: Carlton). Kirnpy, W.F. 1903. Notes on Blaltidae etc., with descriptions of new genera and species in the collection of the British Museum, South Ken- sington. - No. IIL. Ann. Mag. Nat. Hist. (7)42: 373-81. 1904. ‘A synonymic catalogue of Orthoptera. Vol. 1’. (British Museum (Natural History): London), MACKERRAS. MJ. 1967. A blind cockroach from 595 caves in the Nullarbor Plain (Blattodea: Blattel- lidae). J. Aust. Ent. Soc. 6: 39-44. May, B.M. 1963, New entomological records, N.Z. Ent. 3: 44-53. MCKITTRICK. F.A. 1964. Evolutionary studies of cock- roaches. Mem. Cornell Univ. Agr. Exp. Stn 389: 1-197. Norris, K.R, 1970, ‘General biology’. pp. 107-140. fn CSIRQ, ‘The Insecis of Australia’. (Mel- bourne University Press: Carlton). PRINCIS, K. 1951. Neue und wenig bekannte Blattarien aus dem Zonlogische Museum, Kopenhagen. Spolia Zool. Mus. Hauniensis 12: 5-72, 1954. Australian Blattariae, Report from Professor T. Gislén’s expedition to Australia in 1951- 1952, Lunds Univ. Arssk. Avd. 2. Bd 50, Nr 13. Kungl. Fysiog. Sdllsk. Hand. N.F. 65, Nr 13: 3-49. 1959, Revision der Walkerschen und Kirbyschen Blaltarientypen im British Museum of Natural History, London, I}. Qpuse. Ent. 24: 125-50. 1963, Two new Western Australian cockroaches. J, R. Soc, West. Aust. 46; 17-2. 1969, Blaltariae, Suborder Epilamproidea. Jn Beier, M. (ed.), ‘Orihopterorum Calalogus,” Pt 13. (W, Junk: s’ Gravenhage). 1974, Ergebnisse der Ostetreichischen Neukaledonien-Expedition 1965. Blattariae ~ Schaben. Ann. Naturhist. Mus. Wien 78: 513-21. REHN, J.A.G, 1922, Contributions to our knowledge of the Dermapiera and Orthoptera of the Transvaal and Natal. Part, 1. Dermaptera and Blattidae. Ann. Trans. Mus. 9: 1-99. REnTz, D.C.F. AND CAMERON, H.M, 1983, Catalogue of Australian cockroaches. C.S./.R.0., Div. Ent. Tech. Paper 2431-37. Ricnaros, A.M. 1967. Cockroaches (Blattodea) from Australian caves, Helictite 5; 35-44. RotH, L.M. 1968. Oothecae of the Blattaria. Ana. Ent. Soc, Amer, 61; 83-111. 1969, The evolution of male (ergal glands in the Blattaria. Ann. Ent. Soc. Amer. 62: 176-208. 1980. Cave dwelling cockroaches from Sarawak, with one new species. Syst. Ent, 5: 97-104, 1985a. A laxonamic revision of the yenus Blattella Caudell (Dictyoptera, Blattaria: Blattellidae). Ent. Seand. Suppl. 22: 1-221. 1985b, The genus Symploce Hebard. I. Species from New Guinea. (Diclyoptera: Blattatia, Blat- tellidae). Ent. Scand, (1984) 1S: 299-331, 1986. The genus Symploce Hebard. V. Species from maintand Asia (China, India, lran, Laos, Thailand, South Vietnam, West Malaysia). (Dic- tyoptera; Blattaria, Blattellidae). Ent. Scand. (1985) 16: 375-97. 1987a. The genus Tryonicus Shaw from Australia and New Caledonia (Dictyoptera: Blattaria: Blattidae: Tryonicinae). Mem, Od Mus. 25; 151- 67. 1987b. The genus Episymploce Bey-Bienko. IV. Species from India. (Dictyoptera: Blattaria, Blat- tellidae). Ent. Scand. 18: 111-23. 1988. Some cavernicolous and epigean cock- roaches with six new species, and a discussion of the Nocticolidae (Dictyoptera: Blattaria). Rev. Suisse Zool, 95: 297-321. RoTH, L.M. AND WILLIS, E.R. 1952. Tarsal structure and climbing ability of cockroaches. J. Exp. Zool. 119: 483-51, SAUSSURE, H. DE 1863. Mélanges orthoptérologiques. 1° Fascicule, Blattides. Mém. Soc. Phys. Hist. Nat. Genéve 17: 129-70. 1869, Mélanges orthoptérologiques. IT ~ Fascicule. Mém. Soc. Phys. Hist. Nat. Genéve 20: 227-326. 1872. Mélanges orthoptérologiques. IV" Fas- cicule, Mantides et Blattides. Mém. Soc. Phys. Hist. Nat. Genéve 23: 1-164. SHELFORD, R. 1908a. On a small collection of Blat- tidae in the Naturhistorischen Museum zu Wies- baden. Jahrb. Nassauisch. Ver. Naturk. Wiesbaden 61: 27-38. 1908b. Orthoptera. Fam. Blattidae, Subfam. Phyl- lodromiinae. Genera Insectorum Fasc. 73: 1-29. 1910. Orthoptera. Fam. Blattidae, Subfamily Blat- yr MEMOIRS OF THE QUEENSLAND MUSEUM tinae (=Periplanetinae). Genera Insectorum Fasc. 109: 1-27. TepPeR, J.G.O. 1893. The Blattariae of Australia and Polynesia. Trans. Proc. R. Soc. §. Aust, 17: 25- 130. 1894. The Blattariae of Australia and Polynesia. Supplementary and additional descriptions and notes. Trans. Proc. R. Soc. S. Aust. 18: 169-89. 1895a. Descriptions of new or little-known species of Blattariae, Gryllacridae, and Stenopelmatidae collected at Lake Callabonna, S.A. Part I. Trans, Proc. R. Soc. §. Aust. 19: 19-24. 189Sb. Notes on Victorian and other Blattariae and descriptions of new species. Trans. Proc. R. Sac. S. Aust. 19: 146-66, 1896. Orthoptera. /n Horn, Scientific Expedition to Central Australia, IT. Zoology, Melbourne and London, pp. 357-79. WALKER, F. 1868. ‘Catalogue of the specimens of Blattariae in the Collection of the British Museum’. (British Museum (Natural History): London). 1869. ‘Catalogue of the specimens of Dermaptera Saltatoria and supplement to the Blattariae in the collection of the British Museum’. (British Museum (Natural History): London). 1971. ‘Supplement to the catalogue of the Blattariae’. (British Museum (Natural History): London). REVISIONARY STUDIES ON BLATTELLIDAE (BLATTARIA) FROM THE INDO- AUSTRALIAN REGION LOUIS M. ROTH Roth, L.M. 1990 08 31: Revisionary studies on Blattellidae (Blattaria) from the Indo- Australian Region. Memoirs of the Queensland Museum 28(2): 597-663. Brisbane. ISSN 0079-8835. The genus Shelfordina Hebard is reviewed and the following 10 species are redescribed: digitata (Hanitsch); fuscocastanea (Hanitsch); jarakae Hebard; latimarginata (Hanitsch), n. comb.; minor (Hanitsch); orchidae (Asahina), n. comb.; panamae (Hebard), n. comb.; spinistylifera (Roth), n. comb.; terminalis (Brunner); uniformis (Hanitsch). The following 7 new species of Shelfordina are described: cooki; philippinensis; robertsi; sumatrensis; tozerensis; wailimensis; yeatesi. Five species are found in Australia, one is from Panama and the others are mostly Indonesian. The type species of the related genus /mblattella Bruijning, /mblattella impar (Hebard), is redescribed. The Australian genus Dyakinodes Princis is reviewed. One species, centralis (Walker) is redescribed and the following 6 new taxa are described: bispinulifera; fraserensis; kurandensis; penibifida; waterhousei; uptoni. Keys for identifying males of Shelfordina and Dyakinodes are given. Three species of Anaplecta Burmeister are recognised in Australia, namely, A. calosoma Shelford from Queensland (previously known from Papua New Guinea), A. australiensis n.sp. from Queensland and Northern Territory, andA. brachyptera n.sp. from Queensland. The banded tegmina of calosoma are similar to those of A. vittata Hanitsch (Singapore, Sumatra, West Malaysia, and Indonesia), but based on male genitalia both taxa are distinct. Parectoneura bivittata n.gen., n.sp. from Northern Territory and Western Australia is described. Pseudectoneura Princis (New Caledonia) and Pseudectobia Saussure (India) are redescribed. Pseudectobia insularis (Saussure) (Mauritius) is transferred to Chorisoblatta Saussure and Zehntner. The subfamily placement of the last four genera is discussed. Parectoneura and Pseudectoneura are placed in the Blattellinae and Pseudectobia and Chorisoblatta are assigned to the Pseudophyllodromiinae. (1/nsecta, Blattaria, Blattellidae, taxonomic revisions, Indo-Australian. Louis M. Roth, Museum of Comparative Zoology, Harvard University, Cambridge, MA, 02138, U.S.A.; Correspondence: PO Box 540, Sherborn, MA, 01770, U.S.A.; 25 March, 1989. In this paper I review several blattellid genera, one of which, Dyakinodes with 7 species, occurs only in Australia. The Second, Shelfordina, has 17 species, 5 of which occur in Australia and the others are mainly Indonesian. In 1986 Dr David Rentz of CSIRO sent me some cockroaches that were found feeding on orchids in a greenhouse in Canberra. I decided the species was /mblattel- la panamae Hebard (in Rentz, 1987a) basing my determination on Asahina’s (1973) description of what he believed to be that species found on orchids in a Tokyo greenhouse. I was unaware that Asahina (1985) decided that the Tokyo species was not panamae and described it as Imblattella orchidae. After examining a paratype of /. panamae | concluded that it belonged in the genus Shelfordina Hebard, as did orchidae. One of the diagnostic characters of Shelfordina is the presence of a stylelike structure near the basc of cach style so that the male seems to have 4 styles rather than 2. I have examined the type species of /mblattella [I. impar (Hebard)] and find that it is not congeneric with Shelfordina. Other species now in Jmblattella should be reexamined, as should those species that appear to have 4 styles. Some species of Margattea Shelford [e.g., contingens (Walker), and lon- gealata (Brunner)] have accessory styles similar to those in Shelfordina, but they have Type B front femurs (Roth, 1989), rather than Type A In this paper I also describe 2 new species of Anaplecta, and redescribe Anaplecta calosoma, all from Australia, the latter also occurring in Papua New Guinea. | also describe Parectoneura bivittata n.gen., n.sp., from Northern Territory and Western Australia. Superficially this species resembles some species of Ectoneura, an Australian genus which Hebard (1943) placed in the Ectobiinae, and Princis (1969) included in the Chorisoneuridae. I redescribe Pseudectoneura 598 Princis, and Pseudectobia Saussure because the diagnosis of the former was similar to that of Parectoneura, and the latter name suggested a relationship to Ectobtus. The specimens used in this study were loaned to me by the following museums and their curators or collection managers: AMSA - Australian Museum, Sydney, New South Wales; Dr M.R. Gray and Mr B.J. Day; ANIC - Australian National Insect Collection, CSIRO, Canberra, Australian Capital Territory, Australia, Dr D.C.F, Rentz and Mr John Balder- son; ANSP - Academy of Natural Sciences of Philadelphia, PA, U.S.A., Mr Donald Azuma; BMNH - British Museum (Natural History), London, England, Mrs Judith Marshall; BPBM - Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A., Mr Gordon Azuma and Mr J. Strazanac; DARA - Biological and Chemical Research In- stitute, Rydalmere, New South Wales, Australia, Dr G.R. Brown; HDEO - Hope Entomololgical Collections, University Museum, Oxford University, Oxford, England, Dr G.C. McGavin; ISNB - Institut Royal des Sciences Naturelle de Belgique, Brussels, Belgium, Dr P. Grootaert; MCZH - Museum of Comparative Zoology, Har- vard University, Cambridge, MA, U.S.A.; MHNB - Naturhistorisches Museum, Basel, Switzerland, Dr M. Brancucci; MNHG - Museum d'Histoire Naturelle, Geneva, Switzer- land, Dr Bernd Hauser; NMVM - National Museum of Victoria, Melbourne, Victoria, Australia, Dr A. Neboiss; NMWA - Naturhis- torisches Museum Wien, Vienna, Austria, Dr U. Aspéck; QMBA - Queensland Museum, Bris- bane, Queensland, Australia, Dr G.B. Monteith; SAMA - South Australian Museum, Adelaide, South Australia, Dr G.F. Gross; UZMC - Univer- sitets Zoologiske Museum, Copenhagen, Den- mark, Dr H. Enghoff. [The United States National Museum, Washington, D.C. (USNM) has a type specimen which was not borrowed. Dr Syoziro Asahina sent me a paratype of ‘Imblattella’ orchidae Asahina which I have deposited in ANIC. ] Measurements are given in millimetres. Imblattella impar (Hebard) (Figs 1A-D) Neoblattella impar Hebard, 1920, p.58, pl.3, fig. 13 (do and 2): 1921a, p.163; 1933, p.115. Imblattella impar (Hebard): Bruijning, 1959, p.65; Princis, 1969, p.797. MEMOIRS OF THE QUEENSLAND MUSEUM HOLOTYPE (not examined), Tabernilla, Canal Zone, Panama, A.H. Jennings; in USNM. MATERIAL EXAMINED PARATYPES: PANAMA. ANSP: Porto Bello, 1d (ter- minalia slide 410), 15.11.1911, E.A. Schwarz; Al- hajuelo, 1d, 17.iv.1911, 2d 3, 9.iv.1911, A. Busck; Bejuco Riy., 1d, Wm. Schaus. DESCRIPTION Male. Interocular space less than distance be- tween antennal sockets. Tegmina and wings fully developed, former with discoidal sectors lon- gitudinal. Hind wings with proximal costal veins distinctly clubbed; discoidal, median, and cubitus veins straight, latter with 4-5 complete and () incomplete branches, apical triangle small (Fig. 1B). Front femur Type Ag; pulvilli present on 4 proximal tarsomeres, tarsal claws symmetri- cal, minutely serrated, arolia present. Abdominal terga unspecialised. Supraanal plate with medial portion broadly produced (Fig. 1A). Subgenital plate strongly asymmetrical, styles small, cylindrical, located on and near left posterior corner of plate; left style slightly larger, bearing some small, dark, robust spines, these absent from right one, interstylar margin an elongated oblique process bearing couple of rows of dark robust spines on proximal half (Figs 1C,D). Genitalia as in Fig. 1C; hook on right side. Colouration. Brown. Heads of the specimens are discoloured. According to Hebard, there are *...4 vertical lines on occiput, a moderately broad band between eyes, a moderately broad band, convex ventrad, between ocelli and face macu- late with blackish brown’. Pronotal disk light brown with picturing and dots of dark brown. Tegmina transparent, light brown. Wings with non-uniform brownish tinge. Abdominal terga light brown with dark suffusions on distal seg- ments. Abdominal sterna with shiny black maculation mesad on proximal segments. Measurements. Length, 9.0-11.1; pronotum length x width, 2.6-2.9 x 3.3-3.6; tegmen length, 10,8-11.8. REMARKS Neoblattella Shelford is a large South and Central American genus of Blattellidae, and the males show a great diversity of subgenital plate and style morphology. Hebard described many specics and separated them into species-groups, but did not provide diagnostic characters for these groups. Bruijning (1959) raised some of Hebard’s species-groups to genera, and selected INDO-AUSTRALIAN COCKROACHES Neoblattella impar as the type species of Imblat- tella and included Neoblattella litosoma Hebard as a second species with a general statement that the species included Hebard’s impar-species group. Princis (1969, pp.796-798) listed 14 species in /mblattella, most of them transferred from Neoblattella, and included Neoblattella panamae as a new combination. Hebard (1920, p.58) pointed out that the male subgenital plate of impar is ‘... of a type in no 599 way comparable to that found in any of the species ...’ of Neoblattella, or *... in any of the forms of Blattellites’. 1 consider the subgenital plate and style morphology important generic characters in this group of cockroaches. /mblat- tella panamae agrees with all the diagnostic characters of Shelfordina and | am transferring the species to that genus. The subgenital plate and styles of /mblattella impar are so unique that it may represent a monotypic genus. 0.15 mm Fic. 1. (mblattella impar (Hebard), d paratype from Alhajuelo, Panama: A, Supraanal plate (dorsal); B, Hind wing; C, Subgeniial plate and genitalia (dorsal); D, Left corner of subgenital plate (dorsal). win Shelfordina Hebard Shelfordina Hebard, 1929, p.46; Princis. 1969, p.871. Shelfordella (nec Adelung): Hanitsch, 1933, p.126; Princis, 1951, p.57. Payseina Strand, | 934. p.275; Bruijning, 1947, p.218; Princis, 1951, p,57. Ebnerella Hanisch, 1935, p.14; Princis, 1951, p.57, TYPE SPECIES Shelfordina fuscocastanea (Wanitsch), here selected [= Phyllodromia terminalis (nec Brun- ner), Hebard, 1929, p.46]. Hebard had 3 specimens when he selected terminalis as the type species of Shelfordina. One is a male of fuscocastanea and the other 2 are females of Shelfordina terminalis minor (Hanitsch) which L am raising to specilic rank, Since Hebard misidentified the type spectes | intend to apply to the commission to designate fuscocasfanea us the type species because there 1s no question as to the identification of Hebard’s male specimen. A male specimen of terminalis trom Borneo, the type locality, is unknown; [ have seen a male from Sumatra which I have provisionally deter- mined as that species. One of Hebard’s diagnostic characters of Shel- fordina was *.. sexes dissimilar, female with reduced organs of flight reaching apex of ab- domen only.” Hanitsch (1933, p.126) found that Hebard had misidentified the specimens he used for his description of the genus and the above ‘diagnostic’ character was invalid, Hanitsch renamed the genus Shelfordella, However, Skel- fordella was preoccupied and Strand (1934, p.275) replaced it with Passeina. In the interim, learning that Shelfordella was preoccupied, Hanitsch (1935, p.l4), renamed it Ebnerella. Hanisch had no valid reason for disearding Hebard’s original name and Prineis (1951, p57, footnote 2) synonymised the 3 invalid names. DIAGNOSIS Tegmina and wings usually fully developed, sometimes reaching only to end of abdomen; hind wings may be shorter than legmina, some- times almost vestigial (Fig. 10J). Discoidal sec- tors of tegmina longitudinal or sublongitudinal (Figs 6E.7G,4F). Hind wihg with some costal veins (usually the unbranched ones) clubbed (Vig, 11F), or thickened (Pig. 19D); discoidal vem straight, simple; median vein straight, usually simple, rarely with small apical branch; cubjitus vein usually straight, rarely weakly con- cave, With U (rarely) to 5 complete branches, MEMOIRS OF THE QUEENSLAND MUSEUM incomplete rami almost invariably absent, apical triangle present (except in greatly reduced wings). Front femur Type Ag, or A3 (if latter, first terminal spine only slightly longer than preced- ing one); pulvilli present on 4 proximal tar- Sometes, tarsal claws symmetrical, specialised (ventral margins serrated) (Fig. 11E; teeth may vary [rom distinct to sub-obsolete), arolia present. d: Abdominal terga unspecialised (in 2 species eighth segment enlarged, bearing spaced setae: Fig. 19E). Exposed portion of subgenital plate symmetrical. Styles widely separated, usually with a stylelike structure arising near their bases (hereafter called accessory styles) giving appearance of 4 styles rather than 2 (cp. Fig. 11C,c, Fig. 2G). Supraanal plate traasverse, symmetrical, right and left paraprocts similar plates, without spinelike processes (Fig. 11A). Genjtal hook on right side (Fig. 13B,e). REMARKS Princis (1969, p.871) listed 7 species of Shel- fordina from Malacca, Sumatra, Mentawai Is- lands, Celebes, and Japan. Of these S. japonica (Shelford) is a Symploce (Asahina, 1974) and 8. erythrecephala (Hanitsch) is an Episymploce (Roth, 1985a, p.211; 1986, p.180). In this paper | describe, redescribe, or discuss 17 species of Shelfordina of which 7 are new and 4 are new combinations. The male genital hook on the right side places Shelfordina in the Pseudophyllodromiinae (= Plectopterinae of McKittrick, 1964). The females in this subfamily usually do not rotate the ootheca prior to depositing it. A female of Shelfordina digitata (Fig. 2F), and one of S. orchidae were carrying oothecae in the vertical position, and they probably deposit the egg case from this position. SPECIES-GROUPS OF SHELFORDINA Based on the male subgenital plate and acces- sory styles I place species of Shelfordina in 3 species-groups as follows: 1. digitata species-group: Accessory styles ab- sent; interstylar margin unmodified, Species: divitata, 2. /atimareginara species-group: Accessory styles present, right and left ones similar in shape; interstylar margin strongly modified, bearing a puir of large, lobelike structures. Species: jarakae, latimarginata, sumatrensis. 3, fuscocastanea species-group; Accessory INDO-AUSTRALIAN COCKROACHES styles present, right and left ones similar or dis- tinctly different in shape; interstylar region not strongly modified, without setose lobes. Species: cookt, fuscocastanea; = minor; —archidue; panamae; philippinensis; robertst; — spinis- tylifera; terminalis; tozerensis; uniformis; yeatest;, wailimensts. Before receiving the type of Shelfordina uniformis, | considered erecting a fourth group in which the right and left accessory styles are strongly dissimilar; 2 species, spinistylifera and wailimensis would have been assigned to this group. However, §. uniformis has similar right and left accessory styles (placing it in the /us- cocastanea group), but its median genital phal- lomere and cighth abdominal tergum are so similar to those of spinistylifera that | consider them to be more closely related than spinis- tylifera is to wailimensis. KEY TO MALES OF SHELFORDINA The male of S. minor is not known, but if its tegmina are similar to the female’s (clear apical region), it will key to ferminalis. 1. Accessory styles absent (Fig. 2G) (digitata species- group), (Mentawai Islands) ............... digitata Accessory styles present, i iabattll appearance of 4 styles (e.g., Fig. 11C,e) ... Rerrerrerirr rad 2. Right and left accessory styles similar, or almost similar in shape (e.g., Figs 6G,14C) od Right and left aecessory styles strongly dis- similar in shape, left one larger than right one 3, Interstylar region strongly modified with 2 large lobes densely coyered in par| with setae (Figs 3H,4E,5F). (latimarginata species-group) .,...4 Inlerstylar region not as above (fuscocastanea SPECIES- BTOUP) -0... sc cecsseeseesereeenretretrers cae sstessaee 4. Pronotal disk light brown without distinet markings (Fig. SC). Region between interstylar lobes roundly raised, setose (Fig. SF). (Sumatra) seseacuasts . SUMAIFENSIS Pronotal disk with large light or dark reddish brown macula (Figs 3A.B,4A). Region be- oO! tween interstylar lobes not as above (Figs 5, Apex of median genital phallomere hardly enlarged (Fig. 3H), (Mentawai Islands) ...Ja/imarginata Apex of median genital phallomere greatly en- larged, rounded = Wig hi ain Jarak) i = om .. Jarakae 6. Tegmina reddish brown except for transparent colourless apical zone (Fig. 6E). (Borneo, Sumatra) 0... pec peee sec teerees eoycee reereese terminalis TegMina MOUS ADOVE voces eseseneeeeeneeens 7 7. Head with broad, brownish, transverse band on vertex, face very pale, immaculate (Fig. 12 A). (apan, Australia)... ccs erences orchidae Head not. as db0VE oo. eer eee & 8, Accessory styles with 3 or 4 dark spines on inner lateral margins (Fig, 9C). (Panama) ... panamae Accessory styles not aS above ..... ce eeeeeeete oo] 9. Apical margins of accessory styles with small setae; ventral surface of subgenital plate with small mound bearing group of dark setae anterior to each accessory style (Fig. 8D), (Sumatra) unsetesetipeassanpeastactban « fuscocastanea Accessory styles not as above. Mound-bearing selae anterior lo accessory Styles absent....... 10 10, Accessory styles short, bulbous, apically bifid (Fig. 16F). Tegmina and wings reaching to about end of abdomen. (Australia), ....0....- cooki Accessory styles not bulbous, Tegmina and wings extending heyond end of abdomen ..... 11 11. Head with a pair of transverse bands between ocellar spots and antennal sockets (Fig. LOC). Pronotum as in Fig. 10A. Interstylar margin convexly rounded (Fig. 10D), (Philippines) GiidaHscloatses-tatthartodateetiad talastauenectt philippinensis Head, pronotum, and interstylar margin not as BEY EW... 05 jos nap seossep eoreapoepacep doe vep sopaces desseproracepe 12 12. Cubitus vein of hind wing with 3-5 complete branches (Figs 14D, 19D) .cccsccesssesseeeseenes 13 Cubiius vein of hind wing with | or 2 complete branches (Figs 1 7B, 18D) oes eseecerneere 14 nj? 13. Accessory styles with 2 or 3small apical spines (Figs 14C, 15A). Fighth sbdominal tergum nor enlarged, withoul scattered setae. (Australia) -.- tozerensis Accessory styles spinelike, without apical spines or selae (Fig, 19G). Eighth abdominal lergum enlarged bearing numerous spaced setae (Fig, }9E). (Celebes) .............. aniformis i4. Interstylar margin undulate with pair of widely separated Setal brushes (Fig. eH aa tvnees dédideidericsclert eetrdanieie ws JEOLESL Interstylar margin trigonal, withoul setal brushes (Fig. 1SE). (Australia)... robertsi 15, Eighth abdominal tergum enlarged, bearing scal- lured spaced setae (as in Fig. 198; also see Rath, }O8S5b: Fig. (2G). (Sabah) ......... spinistylifera Eighth abdominal! tergum not enlarged, without scatlered selae (Sumatra)... watlimensts Shelfordina digitata (Hanitsch) (Figs 2A-G) Neoblatrella digitata Wanitsch, 1928, p18, pli, fig.7 (o and @). Shelfardella digitata (Hanitsch): Hanitsch, 1933, p. 126. Ebnerella digitata (Hunitsch): Hanitsch, 1935, p,14. Shelfordina digitata (Hanitsch): Princis, 1969, p.872. HOLOTYPE (nol examined) Mentawai Islands; in HDEO. MATERIAL EXAMINED Paratyres: Meniuwai Islands. HDEQO: Siberut, 1d (terminalia slide 232), 26.ix.1924, ELA. Karny, Type Orth, 329°/6; Sipura, 1 (with altached cotheca), 31x. 1924, H.H. Kurny, Type Orth, 3207/6. DESCRIPTION Male. Interocular space less than distance be- tween antennal sockets (Fig. 2C); palpal seg- ments 3 and 4 each longer than segment 5. Pronotum subelliptical (Fig. 2B). Tegmina and Wings fully developed extending beyond end of abdomen, former with discoidal sectors lon- gitudinal or almost so. Hind wing with subcosta and unbranched costal veins thickened on distal halves, discoidal, median and cubitus veins straight, latter with 5 complete and 0 incomplete branches, apical tnangle small (Fig. 2E). Front MEMOIRS OF THE QUEENSLAND MUSEUM femur Type A3; pulvilli on 4 proximal tar- someres, tarsal claws symmetrical, ventral mar- gins minutely toothed, arolia present. Abdominal ferga unspecialised. Supraanal plate transverse, trigonal, apex weakly indented; right and left paraprocts similar, weakly defined plates (Fig. 2D). Subgenital plate symmetrical, styles similar, cylindrical, widely separated, accessory styles absent, interstylar margin straight, un- modified (Fig. 2G). Genitalia as in Fig, 2G; hook on right side, apex acute without apical spine or preapical incision, median phallomere slender, terminating in threadlike filament, scleratised plate and 2 groups of setal brushes. Colouration, Head reddish brown, clypeus and labrum pale (Fig. 2C). Pronotum with reddish brown maculae on anterior half, blackish brown on posterior region, background colour at centre of disk yellowish, broad Jateral regions hyaline (Fig. 2B); the dark pattern apparently varies (cp. Hanitsch, 1928, pl.l, fig.7). Tegmina with mediastinal area subhyaline, pale yellowish, Temainder dark amber to light yellowish brown. Hind wing clouded on distal half of costal vein region and anterior field, and behind apical tri- angle (Fig. 2B). Abdominal terga light brown, darkly infuscated laterally, raund spots sub- laterally. Abdominal sterna light brown with lateral stripes and sublateral dots. Cerci yel- Jowish brown, basal 2 segments dark brown. Female. Cubitus vein hind wing with 3 com- plete branches (1 bifurcate). Supraanal plate strongly transverse, hind margin shallowly con- vex. Pronotal pattern as in Fig. ZA. Abdominal terga extensively infuscated. Abdominal sterna with large medial zone and most of subgenital plate dark brown. Ootheca. The female is carrying an ootheca in vertical position, keel dorsal (Fig. 2F). The egg cage is black with very narrow keel, respiratory serrations very small: surface covered with transverse striae, longitudinal cell divisions ab- sent: height, 2.6mm. Jength, 5.4mm, width, 2.0mm. Measurements (2 in parentheses). Length, 12.5 (12.8, contracted); pronotum length x width, 4,0) x 5.2 (4.5 x 5.7); tegmen length, 15.0 (14.5). REMARKS Were it not for having Type A front femur, this species could be mistaken for a member of the ceylanica species-group of Margaitea (front femur Type B). INDO-AUSTRALIAN COCKROACHES 603 2mm wai Islands: A, @ from Sipora, pronotum; B-E, ate and paraprocts (ventral); E, Hind wing; F, enital plate and genitalia (dorsal). Fic. 2. Shelfordina digitata (Hanitsch), paratypes form Menta G, 6 from Siberut: B, Pronotum; C, Head; D, Supraanal pl Ootheca attached to abdomen (lateral; from 2° shown in Fig. A); G, Subg 604 Shelfordina latimarginata (Hanitsch) n.comb. (Figs 3A-H) Neoblatella latimarginata Hanitsch,, 1928, p.19 (d and 9). ‘Shelfordina latimarginata (Hanitsch): Hebard, 1929, p.9. Shelfordella latimarginata (Wanitsch): Hanitsch, 1933, p,126. Ebnerella latimarginata (Hanitsch): Hanitseh, 1935, p.l4. Possoina latimarginata (Hanilsch): Bruijning, 1948, p.8s, Shelfordella jarakae (Hebard): Hanitsch, 1933, p.124 (incorrectly synonymised with latimareginata; Prin- cis, 1969, p.872 (listed as a synonym of latimar- ginata). HOLOTYPF (nol examined) 3, Mentawat Islands; in HDEQ, MATERIAL EXAMINED PARATYPES, MenTAWAt ISLANDS. HDEO: Sipora, 12 (terminalia slide 231), 22.x,1924, H.A. Kamy, Type Orth. 3307/5, WEST SUMATRA. HDEO: Siberut Island, Id, ix.1924, C.B.K. and N.S., Type Orth. 330°/5. DESCRIPTION Male, Interocular space less than distance be- tween antennal sockets (Fig. 3C); palpal seg- ments. 3 and 4 cach longer than segment 5. Pronotum subelliptical (Figs 34,B). Tegmina and wings extending beyond end of abdomen, discoidal sectors of former longitudinal (Fig. 3F). Hind wing with unbranched costal veins thickened on distal halves, discoidal, median and cubilus veins Straight, median with a small apical bifurcation, cubitus vein with 5 complete (1 with 2 small branches) and 0 incomplete branches, apical triangle distinct (Fig. 3G). Front femur Type Aa; pulvilli on 4 proximal tarsomeres, tar- sal claws symmetrical, minutely toothed, arolia present. Ahdominal terga unspecialised. Supraanal plate transverse, trigonal, apex of hind margin narrowly truncate, right and left paraprocts similar plates (Fig. 3D). Subgenital plate symmetrical, distal region directed dorsad (Fig. 3E); styles slender, cylindrical, similar, widely separated with accessory styles beneath them; interstylar lobe with pair of large structures their distodorsal surfaces covered with small dark spines (Figs 3E,H). Genitalia as in Fig. 3H; hook on right side with small apical spine, entire structure covered by sclerotised plale; median MEMOIRS OF THE QUEENSLAND MUSEUM phallomere, slender, curved, rodlike, apically modificd as rounded membrane; below phal- lomeres a large membrane, distal margin bearing huge brush composed of densely packed dark curved setae (in pinned male from Siberut Island this brush, which is normally hidden in the geni- tal chamber, was extruded and exposed). Colouration. Head with reddish brown occiput and vertex, transverse whitish band above anten- nal sockets, then a large reddish brown macula with pale areas within it, region below dark pat- tern, yellowish brown (Fig. 3C), Pronotum with broad reddish brown macula, pale areas within it, oblique lateral margins outlined opaque whitish, regions surrounding coloured area hyaline (Fig. 3B); pattern varies, disk may be light with dark oblique sides, whitish margins absent (Fig, 3A), Tegmina with mediastinal and costal vein area pale brownish yellow, remainder reddish brown, darker in anal area (Fig. 3F). Wings pale fuscous, darker in costal vein region and distal part of anterior field, and behind apical triangle which is pale (Fig. 3G). Abdominal terga yellowish brown, mottled with dark brown and lateral dots. Abdominal sterna yellowish brown, segments on basal half darker; subgenital plate dark brown on distal upturned portion below paired stylelike structures. Legs yellowish brown. Cerci dorsally light yellowish brown, basal segments darker, Female. Not seen. Measurements. Length, 13.0; pronotum length x width, 3.8-4.1 x 4,8-5.3; legmen length, 15.0- 16.8. REMARKS Differences in male genitalia and interstylar margin of subgenital plate clearly show that latimarginata and jarakae are valid taxa and not conspecific as claimed by Hanitsch. The brush- like membrane underlying the genital phal- Jomeres is much larger in latimarginata than in jarakae or sumatrensis, the other two members in the species-group. Shelfordina jarakae Hebard (Figs 44-E) Shelfordina jarakae Hebard, 1929, p.48, pl.3, figs 3,4 (9): Princis, 1969, p.872 [listed as a synonym of S. fatimarginata (Hanitsch)]. Shelfordella jarakae (Hebard): Hanitsch, 1933, p.126 [incorrectly synonymised with S. latimarginata (Hanitsch)]. INDO-AUSTRALIAN COCKROACHES 605 Fic. 3. Shelfordina latimarginata (Hanitsch), d paratypes: A, from Siberut Island, West Sumatra, pronotum, B-H, from Sipora, Mentawai Islands; B, Pronotum: C, Head; D, Supraanal plate and paraprocts (ventral); E, Subgenital plate (rear-oblique view); F, Left tegmen; G, Left hind wing; H, Subgenital plate and genitalia (dorsal). 606 MEMOIRS OF THE QUEENSLAND MUSEUM mm Fic. 4. Shelfordina jarakae Hebard, 3 holotype: A, Pronotum; B, Hind wing; C, Head; D, Supraanal plate and paraprocts (ventral); E, Subgenital plate and genitalia (dorsal), INDO-AUSTRALIAN COCKROACHES MATERIAL EXAMINED Houorvre; HDEG: 2 (terminalia slide 230), Pulo Jarak, I4.iv. 1921, V. Knight (with a Hanitsch label, Neoblanella latimarginata Han,). DESCRIPTION Male. Interocular space less than distance be- tween antennal sockets (Fig. 4C), Pronotum sub- elhptical (Fig. 44). Tegmina and wings fully developed reaching beyond end of abdomen. Hind wing with several costal veins clubbed, discoidal, median, and cubitus veins straight, latter with 4 complete (2 of them bifurcate) and 0 incomplete veins, apical triangle distinet (Fig. 4B). Front femur Type Aa; pulyilli present on 4 proximal tarsomeres, tarsal claws symmetrical their inner margins minutely toothed, arolia present. Abdominal terga unspecialised. Supraanal plate transverse, hind margin weakly trigonal, apex rounded, pgraprocts similar plates (Fig. 4D), Subgenital plate symmetrical; styles slender, cylindrical, similar, each located near posterolateral corner of plate with stouter acces- sory Slyle near us base; interstylar region with pair of large lobes densely covered with small spines on distal halves of dorsal surface; between these structures a large, round plate that seems to be the apex of the median genital phallomere (Fig. 4E). Genitalia as in Fig. 4E: hook on right side without apical or subapical spine or incision (in the drawing the hook is onented with us apex directed upwards); median phallomere apparent- ly expunded apically as a rounded plate: membrane beneath phallomeres with long row af dark setae. Colouration. Head reddish brown, occiput darker with 3 light transverse bands on face (Fig. 4C), Pronotum with large subtrigonal reddish brown macula, its lateral margins oullined brownish yellow, lateral regions hyaline (Fig. 4A). Tegmina brownish yellow. hyaline in mediastine area and greater part of marginal field, then reddish brown becoming paler distad (see Hebard, 1929: p].3. fig.3). Wings tinged with brown, upicul triangle clear (Fig. 4B). Ab- dominal terga brownish yellow, cach segment infuscated on distal half, with brown mesolateral dot. Abdominal sterna yellowish brown, darker brown lateral dots anu in large proximal area and bilobate portion of subgenital plate. Legs brownish yellow, small flecks of darker brown at bases of most spines, ventral surfaces of hind femurs washed with that colour. Female. Unknown, 607 Measurements. Length, 12.6; pronotum length x width, 3.5 x 4.5; tegmen length, 14.(). REMARKS Hanitsch considered jarakae a synonym of latimarginata undoubtedly because of the similarities in colour patterns and male terminal abdominal segments, However, differences in the median and left genital phallomeres and in- terstylar margins leave no doubt the 2 taxa are distinct. Shelfordina sumatrensis n.sp. (Figs SA-F) MATERIAL EXAMINED HOLOTYPE: ¢, Wai Lima, Z. Sumatra, Lampongs, no. 17K, xt-xii 1921, Karny and Siebers; in HDEO. PARATYPE: SUMATRA, HDEO: same data as holotype, 13, no, &6 (ierminalia slide 233). 2¢ d, 905.197 and 394,19, no. 190 (labelled Ebieredla sp. by Hanitsch); Peduda-B, Lampongs, 1d. i. 1922. DESCRIPTION Male. Interocular space less than distance be- tween antennal sockets (Fig. 5A); palpal seg- ments 3 and 4 each longer than segment S. Pronotum subparabolic (Fig. SC), Tegmina and wings fully developed extending beyond end of abdomen. Tegmina with discoidal sectors weak- ly oblique or longitudinal. Hind wing with sub- costa and unbranched costal veins thickened distad: discoidal, median, and cubitus veins straight, the median usually simple (rarely with asmall apical bifurcation), cubitus with 4-5 com- plete (one of the branches may be forked) and 0 incomplete branches, apical triangle small (Fig. SE). Front femur Type Aa or Ag (if latter, first ‘large’ terminal spine only slightly longer than one preceding it). Abdominal lterga un- specialised. Supraanal plate transverse, hind margin convex, right and lef paraprocts similar plates (Fig. SB). Subgenital plate symmetrical, distal half bent upwards at right angle (Fig. 5D); styles, slender, cylindrical widely spaced, at buse of each a stouter accessory style; interstylar region strongly modified forming large rounded plate bearing pair of denscly spined structures, floor between them longitudinally raised, setose (Fig. 5F), Genitalia as in Fig. SF: hook on right side, slender, without subapical incision or spine, with sclerotised plate at basal end: median phal- lomere slender rod, apex unmodified: membrane beneath phallomeres greatly reduced, roughly atchor-shaped, distal area covered by small 608 spines, the setae relatively light compared to those in jarakae and /atimarginata. Colouration. Head light brown, sometimes partially hyaline, without distinct markings (Fig. 5A); some specimens have faint indications of couple of transverse reddish brown bands. Pronotum with disk yellowish brown, without markings, surrounding region hyaline (Fig. SC), Tegmina hyaline, light brown without markings. Hind wing with thickened region of costal veins MEMOIRS OF THE QUEENSLAND MUSEUM dark, remainder with yellowish tinge, apical tri- angle clear (Fig. SE). Abdominal terga and sterna light brown, former laterally infuscated, latter with large dark macula medially on anterior seg- ments. Cerci light brown dorsally, apical seg- ment dark. Male from Pedada has more extensive infuscation on abdominal segments, and head has more distinct reddish bands. Female, Front femur Type A2. Hind wing with 1 of 4 cubitus vein branches, bifurcate. Supraanal plate transverse, narrow, hind margin convex. Fic. 5. Shelfordina sumatrensis n.sp., ¢ 3d. A,C,D, From holotype; B.E,F, from paratype, same locality as holotype: A, Head; B, Supraanal plate and paraprocts (ventral); C, Pronotum; D, Upturned region of subgenital plate (rear view): E, Hind wing; F, Subgenital plate and genitalia (dorsal). INDO-AUSTRALIAN COCKROACHES 609 FiG, 6. Shelfordina terminalis (Brunner), d from Wai Lima, Sumatra: A, Head; B, Pronotum; C, Supraanal and subgenital plates (dorsal); D, Subgenital plate (ventral); E, Tegmen; F, Hind wing; G, Subgenital plate and genitalia (dorsal). 610 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 7. Shelfurdina spp. A,B, S. terminalis (Brunner), head, and terminal abdominal segments (dorsal) of 2 holotype; C-G, 8. minor (Hanitsch), 2 from Fortde Kock, Sumatra: C, Head: D, Terminal abdominal segments (dorsal); E, Pronotum; F, Hind wing; G, Tegmen. Subgenital plate with large dark reddish brown macula, Measurements (@ in parentheses). Length, 11.0-12.7 (13.5); pronotum length x width, 3.5- 4.0 x 4.5-4.8 (3.8 x 5.1); tegmen length, 14.5- 15.5 (13.4). REMARKS The subgenital plate, styles and their as- sociated structures places sumatrensis close to latimarginata and jarakae. The median phal- lomere with its unmodified apex is more like that of latimarginata, The left genital phallomere is closer to that of jarakae. The greatly reduced INDO-AUSTRALIAN COCKROACHES membrane beneath the phallomeres, and the shape of the interstylar lobe and spination of the brushes, are distinctly different in sumuatrensis. Superficially sertatrensrs can be distinguished from its 2 close relatives by the absence of mark- ings on pronotum and tegmina, Shelfordina terminalis (Brunner) (Figs 6A-G, 7A-B) Phylladromia terminalis Brunner, 1898, p,206, pl. 16, fig. 11.09); Kirby, 1904, p.93; Shelford 1908, p. 13; Ianitsch, 1923, p.414, Shelfardina terminalis (nee Brunner): Heburd, 1929, pT [¥ = Shellordina minor (Hanitseh); J = Shel- Jordina Juseocastanea (Hanitsch), pl. 3, figs 1, 2]; Princis, 1969, p.S71. Shelfordella terminalis (Brunner): Hanitsch, 1933, p.126(¢). Passoina terminalis (Brunner): Strand, 1934, p.275; Broijning, 1948, p.88. Ebnerella terminalis (Brunner): Hanitseh, 1935, p.14. MATERIAL EXAMINED Hotoryee: 2, Barneo; in NMWA. ADDITIONAL MATERIAL. SUMATRA. HDEQ: Wai Lima, Z. Sumatra, Lampongs, |d (terminalia slide 228). xi xii. 1921, Karny (with following labels in Hanitsch’s handwriting’ Neoblatella terminalis Brun- ner, 3; She/fordelle). DESCRIPTION Male, Eyes very close together, space between them distinctly less than distance between anten- nal sockets (Fig. 6A). Pronotum subelliptical (Fig. 6B), Tegmina and wings extending well beyond end of abdomen, discoidal sectors of former longitudinal or sublongitudinal (Fig. 6E). Hind wing with costal veins thickened distad, discoidal, median, and cubitus veins straight, latter with 3 complete and 0) incomplete branches, upical triangle poorly defined (Fig. 6F). Front femurs und all tarsal claws missing (sce 9 description). Abdominal terga un- specialised. Supraunal plate strongly transverse, hind margin convex, not reaching hind margin of subgenital plate (Fig. 6C). Exposed portion of subgenital plate symmetrical, styles similar, cvlindrical Widely separated (Figs. 6C,D): near base of cach style a shorter, curved dark articu- lated accessory style, interstylar margin concaye- ly excavated (Fig. 6G). Genitalia as in Pig. 6G: genital hook on right side with preapical spine: median phallumere straight rod. apex un- 6} modified: Jeft phallomere with pair of sclerotised, cutved, spinelike processes. Colouration. Head reddish brown, darker on occiput and vertex (Fig. 6A). Pronotum with broad reddish brown area that extends from anterior margin almost to convex hind margin, most of lateral regions hyaline (Fig. 6B), Teg- mina reddish brown except for lransparent, colourless apical zone (Fig. 6E). Hind wing in- fuscaled_ thickened castal vein region somewhat darker, small area posterior to apical triangle also darker, apical region practically colourless (Fig. 6F). Abdominal terga light brown, laterally in- fuscated. Abdominal sterna brown, terminal sez- ments darker. Cerci dorsally with segments |-3, 9, 10, and 12 (apex) dark brown. others pale, ventrally segments darker but those in middle somewhat lighter than basal and apical ones. Female (holotype). Interocular space consider- ably less than distanee between ocellar spots and antennal sockets (Fig. 7A). Tegmina and wings extending well beyond end of abdomen, cubitus vein of latter with 4 complete branches. Supraanal plate sirongly transverse, convex, not reaching hind margin of subgenital plate (Fig. 7B). Front femur Type Ag; pulvillion 4 proximal tarsomeres, tarsal claws symmetrical, ventral margins with subobsolete teeth, arolia present, Colouration. Head blackish with reddish tinge, clypeus and labrum yellowish brown (Fig. 7A). Pronatal disk dark reddish brown. lateral borders lighter reddish brown. Tegmina dark reddish brown, large apical region transparent, colour- less. Hind wing with thickened costal vein region yellowish, distal borders of anterior and pasterior fields almost colourless, remainder weakly in- fuscated, Abdominal terga light brown, lateral borders slightly darker, terminal segments black- ish brown. Abdominal sterna reddish brown, subgenital plate blackish with reddish tinge. Cerei and leps reddish brown. Measurements (2 holotype in parentheses), Length, 8.3 (11.4); pronotum length x width, 2.6 x 3.7 (3.6 x 4.7); tesmen length, 11.5 (12.6). REMARKS Hebard (1929, p.46) had 2 females from Fort de Kock, Sumatra, which he determined as ¢er- minalis because their lepmina were *...suddenly transparent and whitish like ground glass, the other poytions being dark.’ He also identified a male from the same locality as the same species even though its tegmina Jacked the transparent, colourless apical region, Hanitseh (1933, p.126) pointed out thal this male was not /ermninalis and Ole Prineis (1969, p.872) correctly listed it under 5. fuscacastanea, whose type I have seen. Hanitsch had a male fram Mt Pai, Sarawak, and Wai Lima, Lampong, Sumatra, which had tegminal mark- ings like those of terminalis and he considered them that species. The male which [ am deserib- ing here as terminalis is Hanitsch's Wai Lima specimen; I have not seen his Mt Poi male, The females from Fort de Kock were named by Hanitsch terminalis minor, but I have raised this subspecies to specific rank (see below), Shelfordina minor (Haniisch) (Figs 7C-G) Neoblattella terminalis Brunner, minor Hanitsch, 1929b, p.273 (2). Shellordina terminalis (nec Brunner): Princis, 1969, 872|2 only; & = 8. fuscncastanca (Hanitsch)], Shellordina terminalis miner (Hanisch); Princis, 1964, p.872. MATERIAL EXAMINED Hororyer: 2, Fort de Kock, Sumatra, 920m, 1925, E, Jacobson; type Orth. 354'/3 in HDEO, PARATYPE: Pulo Toman, HDEO; 19, vwi.l915, ¥. Knight, Type Orth.354°/73. Additional material. Suntaira. ANSP: Fort de Kock, 12 (genitalia slide 415), [9.x 1913, 19, 920m, xii.1921, E. Jacobson (both reported as 8. erminalis by Hebard, 1929, p.47). DESCRIPTION Male, Unknown (see remarks). Female. Eyes wide apart, interocular space slightly less than distance between antennal sockets (Figs 7C). Pronotum subparabolic (Fig, 76). Tegmina and wings somewhat reduced reaching only slightly beyond end of abdomen, discoidal sectors of former, sublongitudinal (Pig. 7G). Hind wings with costal veins thickened distad, discaidal and median veins straight, simple: cubitus vein weakly concave with 4com- plete and O incomplete branches, apical triangle present (Fig. 7F). Front femur Type A3 (first of 3 terminal spines may be only slightly longer than preceding one); pulyilli present on 4 proximal tarsomeres, tarsal claws symmetrical with few subobsolete teeth, arolia present, Ter- minal abdaminal segments as in Fig. 7D, Colouration, Head reddish brown (Fig, 7C), Pronolum with Jateral borders yellowish, remainder dark reddish brown (Fig. 7E). Teg- mina reddish brown, apical region transparent, appearing whitish (Fig, 7G), Hind wing with MEMGIRS OF THE QUEENSLAND MUSEUM broad distal region practically colourless, remainder infuscated, thickened costal vein region somewhat darker, veins dark except in pale apical zone (Fig. 7F). Abdominal terga of terminal segments with broad lateral borders blackish, or dark brown, remainder light brown which extends medially to hind margin of supraanal plate. Abdominal sterna reddish or blackish brown. Cerci dark blackish brown with 2 pale preterminal segments. Coxae infuscated, remaining parts of legs reddish brown. Measurements. Length, 9.0-10.6; pronotum length x width, 3.1-3.3 x 4.0-4.6; tegmen length, 89-92, REMARKS Because the apical region of the tegmina are transparent in Sumatran specimens, as they are in Brunner’s Bomean terminalis, Hanitsch (1929b, p.273) described il as a subspecies, minor, stating that it differed from the nominate form in being smaller and in colour of the pronotum. There are other colour differences, hut more importantly, the interocular space is distinctly greater (Fig. 7C) in minor, than it is in the holotype of terminalis (Fig. 7A), and for this reason [ am raising Hanitsch’s subspecies to specific rank. Because the eyes of the male from Wai Lima, Sumutra, are even closer together than are those of the female terminalis, | am provisionally identifying it as that species, It is not Uncommon to find the interocular space somewhat different between the sexes of con- specific species, but the difference usually is less than that seen between Figs 6A and 7C, and more like that between Figs 6A and 7A. However, my interpretation of these 2 taxa should be con- firmed by examining males from ‘Borneo’ and Fort de Kock, and females from Wai Lima. Shelfordina fuscocastanea (Hanitsch) (Figs 8A-G) Neuvblattella fiisco-castanea Hanitsch, 1929b, p.274 (2). Ebernella fuscocastanea (Hanitsch), 1935, p.14, Possoina fuscocastanea (Hanitsch): Brutjning, 1948, p.8k. Shelfordina terminalis (nec Brunner, 1898); Hebard, 1929, p.47, ph 3, figs 1.2[4 only; 9 = Shelfordina minor (Hanisch), not S. teeminalis (Brunner) as stated hy Princis, 1969, p.872], Shellordina /uscecastanea (Manitsch): Prineis, 1969, p.872, INDO-AUSTRALIAN COCKROACHES 613 MATERIAL EXAMINED DESCRIPTION Ho.otyPe: ¢. Fort de Kock, Sumatra, 920m, 1926, Male. (see Hebard, 1929, pl. 3, fig. 1, for E. Jacobson; Type Orth. 355; in HDEO. habitus). Head slightly exposed beyond ADDITIONAL MATERIAL. SUMATRA. ANSP; same data — pronotum; interocular space less than distance as holotype, 1d (terminalia slide 416), i.1921 between antennal sockets (Fig. 8A), Pronotum as (misidentified as S. terminalis by Hebard, 1929). in Fig. 8B. Tegmina and wings extending well FIG. 8. Shelfordina fuscocastanea (Hanitsch), d d from Fort de Kock, Sumatra. A,B,F,G, From holotype: A, Head; B, Pronotum; C, Supraanal plate and paraprocts (ventral); D, Distal region of subgenital plate showing styles and accessory styles (ventral); E, Subgenital plate and genitalia (dorsal); F, Tegmen; G, Hind wing. 614 MEMOIRS OF THE QUEENSLAND MUSEUM LNW Sy Lage } Fic. 9. Shelfordina panamae (Heburd), d paratype from Rio Trinidad, Panama: A, Supraanal plate and paraprocts (ventral): B, Hind wing; C, Subgenital plate and genitalia (dorsal), beyond end af abdamen, discoidal sectors of former longitudinal (Fig. 8F), Hind wing with unbranched costal veins slightly thickened on distal halves, discoidal, median, and cubitus veins almost straight, latter with 3-4 complete and 0 incomplete rami, apical triangle distinct (Fig. 8G). Front femur Type A3; pulvilli present oa 4 proximal tarsomeres, tarsal claws with sub- obsolete teeth on ventral margins, arolia present. Abdominal terga unspecialised. Supraanal plate strongly transverse, hind margin convex, reach- ing hind margin of subgenital plate, right and left paraprocts similar (Fig. 8C). Subgenital plate with exposed portion symmetrical, styles widely separated arising from posterolateral corners of plate, each with slightly shorter accessory style near base, interstylar margin straight, upturned, unmodified (Fig, 8E); ventrally, anterior to base of each accessory style, a small mound bearing group of small dark setae (Fig. 8D). Genitalia as in Fig. SE: hook on right side with small apical spine; median phallomere, slender, rodlike, apex unmodified. Colouration, \lead reddish brown, labral and clypeal regions pale, face with light areas near cach antennal socket and on frons (Fig, 8A)- Pronotum with disk deep reddish brown, oblique margins of macula yellow, lateral regions semi- hyaline, posterior area very dark brown or black- ish (Fig. 8B). Tegmina reddish brown, yellow Spot near buse of axillary vein (Fig. 8F). Hind wing with subcostal and thickened costal vein areas pale, region posterior to thickened veins up to discoidal vein weakly infuscated, darkened region extending to apical margin, small area behind apical triangle weakly infuscated. Ab- dominal terga light brown, lateral borders infus- cated, supraanal plate with pale longitudinal macula medially. Abdominal sterna reddish brown, upturned interstylar region pale. Legs reddish brown. Cerci dark brown dorsally and ventrally, preapical segment pale. Female. Unknown. Measurements. Length, 10.0-11.5; pronotum length x width, 3.0 x 4.0-4.2; tegmen length, 12.5-13.6, Shelfordina panamae (Hebard) n.comb. (Figs 9A-C) Neoblattella panamae Hebard, 1920, p.56, pl. 3. fig. 12(0 and #). INDO-AUSTRALIAN COCKROACHES Imblattella panamae (Hebard): Princis, 1969, p. 798, HOLOTYPE (nol examined) oS. Roo Trinidad. Panga, (i112. A. Buseck; in United States National Museum. MATERIA, EXAMINED PARATYPE: ANSP: same locality as holotype, ¢ (ler minilia stide 409), Taw I9)1 A, Buseck, DisSCRIPTION Male. Interocular space slightly less than dis- lance between antennal sockets, Pronotum with anterior and posterior margins truncate, latter broader, lateral margins convex. Tegmina and Wings fully developed, discoidal ‘sectors of former. longitudinal. Hind wing with most costal veins distinctly clubbed, discoidal, median and cubilus veins straight, latter with 3-4 complete and Q incomplete branches, apical triangle small (Fig. 9B). Front femur Type Ag (first terminal spine may be only slightly longer than one preceding it); pulvilli present on 4 proximal tar- someres, tarsal claws with flange on inner mar- gins and with 2or 3 distinct bul microscopic tecth distad, Abdominal terga unspecialised, Supraanal plate transverse, hind margin convex, right and left paraprocts similar. simple plates (Wig. 9A). Subgenital plate almost symmetrical; styles similar, widely spaced, adjacent to each style basally, an aceessory style about as long as styles. with 3-oc 4 dark spines on inner margins: interslylar margin almost straight (Fig. 9C). Genitalia as in Fig. 9C: hook on right side without 4 terminal spine or subapical incision; median phullomere with apical and preapiecal spines, Colouration. General colour brawn, Mead with subobsolere broud brown intervcular band. Pronotal disk brawn with dark lines and blotches. Tegmina light brown, hyaline, Hind wing weak- ly infuseated, axillary Vein area lighter (Fig. 9B). Measurements. (Hebard’s measurements in parentheses). Length, 9.6 (10.0-11.5); pronotum length x width, 2.4 x 3.4 (2.5-2.7 x 3.6), tegmen length. 10.6 (11.1-11.8). Shelfordina philipptnensts o. sp. (Figs IDA-J) MATERIAL EXAMINED Honoryre: 2 (lerminalia slide 470), Camarines Sur Proy,, M1 Triga, Luvon Island. Philippines, 500-600m, Hightirap, S40, 462 71M, Torrevillas: in BPBM, PARA RES Pri taist Ist ANDSs, BRBM: Negros On, a15 Mt Talinas. 900- | 200m, rain forest, | 29.01.1958, H. bE. Million, Luzon, Mt Prov, Wugao Mayoyao, L000: 1S00m, 1, 29.vi.1966, HOM. Torrevillas DESCRIPTION Male. Eyes close together, interocular distance distinctly less than space between ocellar spots and antennal sockets (Fig. 10C). Pronotum sub- elliptical (Fig. 10A). Tegmina and wings fully developed extending well beyond end of ab- domen. Tegmina with discoidal sectors lon- gitudinal (Fig. (OE). Hind wing with subcosta and unbranched costal veins clubbed, discoidal and median veins straight, simple; cubitus yein practically straight with 2 or 3 complete and 0 incomplete branches,. apical triangle well developed (Pig. (OF). Front femur Type Az; pul- villi present on 4 proximal tarsomeres, tarsal claws symmetrical, ventral margins distinctly loothed, arolia present, Abdominal terga un- speciilised, Supraanal plate transverse, hind margin convexly rounded: tight and left paraprocts, similar simple plates (Fig. 10B). Sub- genital plate symmetrical, styles widely separated, accessory styles present, similar, in- terstylar margin convexly rounded (Fig. 10D). Genitalia as in Pig. OD: hook on right side, hook portion short, apically rounded with a small preapical spine; distal end of median phallomere obliquely bent, with preapical sctose modifica- tion, Calouranion. Yellowish brown. Head yel- lowish with vertex brown becoming lighter on occiput, transverse brown bands between ocellur spots and antennal sockets (Fig. 10C). Pronotum with symmetrical pattern of dark maculae, lateral regions hyaline (Fig. LOA). Tegmina pale hyaline, mediastine vein weakly darkened for part of its length. Hind wing with clubbed region of costal veins dark, remainder with yellowish tinge (Fig. 10F). Abdominal terga yellowish brown, Abdominal sterna yellowish brown, basal half of subyenital plate with pair of large brownish black maculie on distal half of seg- ment. Legs yellowish brown, Female. Inicrocular space about same as tnter- ocellar distanee greater than space between an- tennal sockets. Tezmina reduced (Figs 10G,f), reaching to about T7; hind wings shorter than legmina, cubitus vein with | or no complete branches, apical triangle absent (Figs 10H,J). Supruanul plate with hind margin convexly rounded (as in ¢, Fig. 10B). Head and pronotal markings less intense than inmale. Tegmina with mediastine vein weakly darkened, anal vein pale 616 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 10. Shelfordina philippinensis n.sp. A-F, d holotype: A, Pronotum; B, Supraanal plate and paraprocts (ventral); C, Head; D, Subgenital plate and genitalia (dorsal); E,F, Tegmen and wing. G-J, @ paratypes, tegmina and wings: G,H, From Mt Talinas, Negros Or.;.I,J, From Ifugao, Mayoyao, Luzon. All tegmina and wings to same scale, INDO-AUSTRALIAN COCKROACHES 617 \ 3 mm Fic. 11. Shelfordina orchidae (Asahina), 3 from Black Mt., ACT., Australia: A, Supraanal plate and paraprocts (ventral); B, Terminal abdominal segments (ventral); C, Subgenital plate (dorsal); D, Pronotum; E, tarsal claws and arolium: F, Hind wing, Abbreviations: a, supraanal plate; b, right paraproct; c, subgenital plate; d, style; e, accessory style. or weakly darkened. Abdominal terga and sterna reddish brown, with interspersed light areas. Measurements. (? in parentheses). Length, 8.7 (7.7-9.6); pronotum length x width, 2.3 x 3.0 (2.3-2.6 x 3.1- 3.5); tegmen length, 9.5 (5.6-6.7). REMARKS This is the first record of the genus in the Philippines. Shelfordina orchidae (Asahina) n. comb. (Figs 11A-F,12A-D,13A- B,29) Imblattella panamae (nec Hebard), Asahina, 1973, p. 126, figs 9-12, 19-22; Rentz, 1987b, p.3, figs 1-8. Imblattella orchidae Asahina, 1985, p.6, figs 9-18, 20, 21, 23, 24(d and ?); Rentz, 1987a, p.44, figs 1-8; 1987c, p.14, figs.1-8. MATERIAL EXAMINED JAPAN. ANIC: 1d (terminalia slide 181), paratype of Imblattella orchidae Asahina, Totsuka, 5.11,1973, S. Asahina. AUSTRALIAN CAPITAL TERRITORY. ANIC: Canberra (Spence), 1¢, 1 nymph, 25.ix.1987, J. and S, Rickard. The following were collected by D.C.F. Rentz: Black Mt., 35.178 149.06E, Botanical Gardens, in orchids or orchid house, 1¢, 329 (1 carrying ootheca; 1 genitalia slide 176), reared from nymphs, 27.iii.1986, 1d (terminalia slide 175), 24.xi.1986, 1 9 22.vii.1985 (coll. as nymph, matured 10.x.1985), 12, 27.iii.1986 (coll, as nymph, matured 4.vi.1986), 19, 23.vii.1985 618 Cc 0.8 mm MEMOIRS OF THE QUEENSLAND MUSEUM 3 mm FIG. 12. Shelfardina orchidae (Asahina), 2° from Lake Barrine, Qld: A, Head; B, Tegmen; C, Pronotum; D, Hind wing. (coll. as nymph, matured, 17.ix.1985), 12, 10, it), 1987; Canberra (Farrar), 35.228 149.05E, 2¢ 3. 5.11, 1987, 15. 121.1987. New Soutt WALES. DARA: Hamilton. 23d, 52 9, ‘cating orchid leaves’, 23.11.1955, QUEENSLAND. Anic: Pialba (orchid nursery), x.1988, G. Cook (pers. comm, D. Rent); Dunk Island, 1°, 8.vili, 1950, R, Dobson, QMBA; Lake Barrine, Ather- ton Tableland, 17.158 145,38, rainforest, by torch, 19, 7.xti.1950, J, Balderson; Curtain Fig. 2km SSW of Yungaburra, Atherton Tableland, 17.175 145.34E, 19, 14.xi7. 1985, J. Balderson. DESCRIPTION Male: Interocular space distinctly less than dis- tance between antennal sockets (Fig. 12A). Pronotum suboval (Fig. 11D). Tegmina and wings fully developed, discoidal sectors of former sublongitudinal (Fig. 12B). Hind wing with proximal costal veins clubbed, discotdal, median, and cubitus veins straight, latter with 2-4 complete and 0 incomplete branches, apical tri- angle small (Figs 11F,12D). Front femur Type A3; pulvilli present on 4 proximal tarsomeres, tarsal claws symmetrical, ventral margins dis- linctly serrated, arolia present (Pig, 11E). Ab- dominal terga unspecialised. Supraanal plate transverse, hind margin convexly rounded, paraprocts similar, plates partially sclerotised (Fig. 11 A). Subgenital plate symmetrical, styles short, cylindrical, apically rounded, located in posterolateral corners of plate, euch with short contiguous, atticulated accessory style terminal- ing in pair of minute spines (Fig. 11B); between styles and accessory styles an upturned (in pinned specimens) plate (seen flattened in Fig. 11C). Genitalia as in Fig, 13B: hook small, on right side with small apical spine; apex of median phallomere terminating in pair of spinelike processes; accessory median phallomere with several large terminal setac; beneath median and accessory median phallomeres, a densely setose wrinkled membrane (this and dark brushlike structure visible through transparent subgenital plate); left phallomere much larger than right genital hook and is drawn further to the left of its INDO-AUSTRALIAN COCKROACHES 619 Fic. 13, Shelfardina orchidae (Asahina), from Black Mt., ACT., Australia: A, 2, genitalia (ventral); B, d, genitalia (dorsal). Abbreviations: a, first valvifer; b, left phallomere; c, accessory median phallomere; d, median phallomere: ¢, hooklike right phallomere. au normal position in order to show left side of aceessory median phallomere, Calouration. Head with occiput yellowish brown, broad brownish band on vertex, face very pale, immaculate (Mig, 12A), Pronotum hyaline, including disk which has symmetrical brown markings (Pig.11D), Tegmina transparent, very light brown, with some small dark brown spots basally along mediostinal and anal veins. Hind wings pale, translucent, veins dark. Abdominal (erga very pale, translucent, narrowly suffused with brown along lateral margins, supraanal plate with pair of dark maculae near hind margin. Abdominal sterna pale, brownish dot and sul- fusion near lateral borders. Cerci yellowish, basal segments, apex, und couple of preapical segments dark brown and separated from up by pale segment. Female. Supraanal plate transverse, hind mar- gin conves (Fig. 13A) not reaching hind margin of subgenital plate. Colouration as in male but a female from Lake Barrine, Queensland, has very dark markings on pronotum (Fig, 12C) and maculae on lezmina wre more extensive (Fig. 12B). Subgenital plate has pair of dark spots or single large macula near hind margin, Genitalia as in Fig. (3A; base of first valvifer greatly enlarged, femur-shaped, The ootheca is il- lustrated in Asahina (1985, Fig, 21). Nymph. The nymph is illustrated in Asahina (1973, fig.21; 1985, fig. 20) (these were also reproduced by Rentz (1987i-c). Measurements (9 in parentheses), Length, 8.6- 10.8 (9.6-11.0), pronotum length x width, 2.4-2.7 & 3,2-3.7 (2.5- 2.9 x 3.4-3.8), tegmen length, 10.4-11.8 (9.5-10,5) REMARKS This species apparently ts Well established in Australia, The carliest specimens were collected in 1955 in Hamilton, New South Wales., feeding on orchid leaves. Although most of the specimens were taken associated with orchids (including the type specimens from Japan), the species has been collected outside of green- houses, Dr Asuhina confirmed my determination of the Australian specimens According to Rentz (1987a-c), Shelfordina or- chidae 1s nocturnal, lives among roots of epiphytic orchids, and may fced on the growing root ips und damage flower buds and developing shoots of vegetative parts of the plants, They alsa live amongst pinebark and other planting material Used in pot grown orchids, Oothecae are secured lo tangles of roots, The insects hide MEMES OF THE QUEENSLAND MUSEUM among the roots or planting media and emerge and feed at night. Lepschi (1989) determined that orcludae is predominantly herbivorous, feeding on a variety of plant material including pollea, fungal hyphac and plant tissue. Tt has been col- lected on ferns and apparently is not restricted to feeding on orchids. Shelfordina tozerensis n.sp. (Figs 14A-E, 15A-C, 29) MATERIAL, EXAMINED Hororyern 3, 9km ENE of Mt Tozer, Iron Range Nat Park, 12.438 143, 17E, mixed low open forest, 10.40) 7986, DCP, Rentz; in ANIC. PARATYPES: OUFENSLAND. QMBA; Bellenden Ker Range, |/2km §. Cable Tower No. 7, 500m, 2d 4 (1 with missing abdomen), 29 9, 17-24.x.1981, Earthwatch/Qld Mus.: Iron Range, NQ, Id (pyrethrum knockdown/RF), 1d, 19, 50nm, in rain- forest, 3-10.aii, 1985, G. Monteith and D, Cook; Iron Range, Cape York Pen., NO, U2, 30.vi-4.vii.1977, GB. Monteith, ANIC, bkm ENE of Mi Tozer, 12.445 143, 16E, 12, 30.vi1986, T. Weir und A. Calder. The following were coflected by D.C,F, Rentz, in or near Tron Runge National Park! 2km NE by E of Mt Tozer, 12.448 143,198, 10,12, l.vi.1986; 3km BNE of Mt Tozer, 12.448 143.14E, rainforest margin 1.4 (ter- minalia slide 183), 322 (1 with ootheca; | with lemminalia slide 184), 28.vi.-4,vii 1986, [12, same data ex pan trap. JC. Cardate); km E by N of Mt Tover, [2.448 143.17E, 1d, 19, rainforest, 7.vit, 1Y86, same data as holotype, 2d d (1 with ter- minalia stide 180), 12, 7 nymph, -10.vit. 1986, L1km ENE of ML Tozer, 12.438 143.186, rainforest margin, ISP, L2 fal light). 11 vii 1986, DESCRIPTION Male, Eyes very close together, interocular Space less than length of first antennal segment (Pig, 14B), Pronolum suboyal (Fig. 14A). Teg- mina and wings fully developed extending beyond end of abdomen. Hind wing with subcos- ta and 5 or 6 costal veins clubbed, discoidal and median veins straight, simple, cubitus vein siraight with 3-4 complete and no incompicte branches, apical triangle small (Fig. 14D). Front femur Type Aa, or Ag (first terminal spine only slightly longer than preceding ones), pulvilli present on 4 proximal tarsomeres, tarsal claws minutely serrated, arolia present. Abdominal terga unspecialised. Supraanal plate transverse, hind margin convexly rounded, reaching to about hind margin af subgenital plate, right and left INDO-AUSTRALIAN COCKROACHES 621 Fic. 14. Shelfordina tozerensis n.sp., d paratypes from near Mt Tozer, Qld, Australia: A, Pronotum; B, Head; C, Subgenital plate (ventral); D, Hind wing; E, Supraanal plate and paraprocts (ventral). paraprocts similar, simple plates (Fig. 14). Sub- genital plate symmetrical, styles similar, cylindrical located in posterolateral corners, cach with an accessory style terminating with one or 2 small spines, arising ventrally on the plate (Fig. 14C, 15A); styles and accessory styles extend beyond supraanal plate and are visible in dorsal view; interstylar margin straight or weak- ly curved, generally reflexed dorsad. Genitalia as in Fig. ISA; hook on right side; median phal- lomere strongly curved distally. Colouration. Head with occiput and vertex dark brown, dark brown interocellar band, nar- row concave band between antennal sockets, more narrow maculae on frons (Fig. 14B). Pronotal disk with brown markings, lateral and posterior borders hyaline or semihyaline (Fig. 14A). Tegmina light brown, hyaline, basal part of humeral and anal veins darkly infuscated. Hind wing dark, not uniformly infuscated, anterior field lighter than posterior field, its api- cal region yellowish, region of axillary veins near apical triangle darker than rest of posterior field (amount of infuscation varies with some MEMOIRS OF THE QUEENSLAND MUSEUM ot “‘(yeaw9A) elpeiuad * a ‘Cd tRaayIOG “g ‘(JESIOP) elyeITUad pue azeyd pemuadgns ‘ p Sy :pld ‘1az0], IW skau woy sadAjrsed “ds'u s7suasazoy vulpsofpays “S| “O14 ww 1 INDO-AUSTRALIAN COCKROACHES 623 0.15mm 0.5 mm Fic. 16. Shelfordina cooki n.sp., 6 d. A,C,E- G, paratype; B,D, holotype: A, Head; B, Pronotum; C,D, Hind wings; E, Supraanal plate and paraprocts (ventral); F, Right corner of subgenital plate showing style and accessory style (ventral); F, subgenital plate and genitalia (dorsal). specimens much paler than others) (Fig. 14D). Cerci dorsally pale with blackish basal and Abdominal terga light brown, terminal segments — preapical segments, ventrally mostly brown. darker. Abdominal sterna light brown medially, Female, Interocular space about same as darker laterally, pale spots near each border. length of first antennal segment. Supraanal plate 64 strongly transverse, subrectangular, corners rounded, (Fig. 15C), hind margin not reaching hind margin of subgenital plate. Subgenital plate convex, produced, sides obliquely concave, apex rounded. Genitalia as in Fig, 15C. Abdominal sterna pale with narrow dark brown Jateral bar- ders and small pretateral round dots. Qetheca. Brownish black, kee! shallowly un- dulate with 2] small apically rounded teeth, with about 12 transverse, curved heavy ridges, finer ridges between them, longitudinal cell divisions absent. Length, 4.4 mm, height, 1,8 mim (Fig, 158). Measurements (2° in parentheses), Length, 8.7- 9.3 (8.7-10.3); pronotum length x width, 2.2-2.3 x 3.0-3.3 (2.2- 2.4 x 3.1-3.3); tegmen length, 9.9-10.3 (9,8-10.3). Shelfordina cooki n.sp, (Figs 16A-G, 29) MATERIAL EXAMINED HocLorype: ¢, Upper Boulder Ck, 1 1km NNW of Tully, NO, 850m, 16-19.xi.1984. Couk, Monteith, and Thompson; in QMBA (Reg. No. T11280). PARATYPES: QUEENSLAND. OQMBA: same data as holotype, 1d (lerminalia slide 64); Mt Bartle Frere, NO, NW/Centre Peak ridge, 1400-1500m, 16 (ab- domen missing), 7-8.xi-1981. ETYMOLOGY The species is dedicated ta Mr DJ. Cook who has participated in many Queensland Museum collecting trips (Monteith, pers, comm.), DESCRIPTION Male. Head slightly protruding beyond prothorax; interocular space about same as dis- tance between antennal sockets (Fig. 16A), Pronotum subparabolic (Fig. 16B). Tegmina slightly reduced reaching end of or slightly beyond end of abdomen. Hind wings smaller, costal veins weakly thickened, their number vari- able, fewer in more reduced wings; discoidal and median veins simple, straight, latter may arise from and look like branch of former, cubitus vein straight, simple or with 1 complete and no in- complete branches, apical triangle cssentially absent (Figs 16C,D). Front femur Type Az, first of 3 terminal spines may be only slightly longer than preceding spines which may include couple of piliform spinules; pulvilli on 4 proximal tar- someres, fursal claws distinetly serrated, arolia present. Abdominal terga unspecialised. MEMOIRS OF THE QUEENSLAND MUSEUM Supragnal plate transverse, hind margin broadly, shallowly convex; paraprocts similar dark plates (Fig. 16E), Subgenital plate convex, symmetri- cal, distal half reflexed dorsad; styles small, cylindrical, similar, each atising in posterolateral corners of plate: associated with each style, aris- ing {tom dorsal surface of plate, a bulbous, api- cally divided accessory style (Figs 16F,G). Genitalia as in Fig. 16G; hook on right side without subapical incision. Colouration. Head with occiput dark brown, broad pale transverse band on vertex, then a shiny blackish macula between eyes and anten- nal sockets, dull brownish macula on frons (Fig. 16A; dark markings in holotype are more intense and extensive than in paratype). Pronotal disk dark reddish brown with small yellowish spots, lateral borders yellowish on anterior half, remainder hyaline (Fig. 16B). Tegmina reddish brown, hyaline with suffused dark areas. Anterior abdominal terga brown with dark infus- cations, terminal segments dull black. Ab- dominal sierna mostly blackish brown, distal segments pale medially. subgenital plate dark except for narraw medial zone. Cerci dorsally with segments 1-3, 6, 7and 10 blackish (segment 7 may be pale), other segments yellowish, ventrally brown. Legs with coxae largely infus- cated, femurs with narrow dark brown bands distad on anterior surfaces. Female, Unknown. Measurements, Length, 7.2-8.0; pronotum length x width, 2.2-2,5 x 3.4-3.7; tegmen length, 5.6-6.5, Shelfordina yeatesi n.sp. (Figs 17A-F, 29) MATERIAL EXAMINED HoLotyPr & (terminalia slide 63), Thornton Peak via Daintree, NQ, 1100-1300m, 24-27.ix.1984, G.B. and S.R. Monteith; in OMBA (Reg. No. T11187). PARATYPES; QUEENSLAND OMBA: Thornton Peak, ilkm NE of Daintree, 1100-1200m, 19, 30.x.- 1.xi,1983, Monteith, Yeates, and Thompson; Thornton Peak via Daintree, 1000-1300m, 19 (with unthees: genitalia slide 66), 20-22.ix, 1981, G, Mon with and D. Cuok: Bellenden Ker Range, Cable Tower 3, 1054 m, 19 (genitalia slide 67), 25-31.x.1981, Earthwateh/Old Mus. ErYMOLoGY The species is dedicated to Dr D.K. Yeates who has done survey work in north Queensland (Monteith, pers, comm), Pr INDO-AUSTRALIAN COCKROACHES 62 FIG. 17. Shelfordina yeatesi n.sp., ¢ holotype: A, Tegmen, B, Hind wing; C, Apex of median genital phallomere (dorsal); D, Subgenital plate and genitalia (dorsal); E, Hind margin of subgenital plate (ventral); F, Supraanal plate and paraprocts (ventral). DESCRIPTION wings fully developed but reaching only slightly Male. Head hidden under pronotum; inter- beyond end of abdomen. Hind wing with costal ocular space less than distance between antennal veins distinctly clubbed, discoidal and median sockets. Pronotum subparabolic. Tegmina and veins simple, cubitus vein straight with 2 com- 626 plete and 0 incomplete branches, apical triangle small (Fig. 17B). Front femur Type A3 (first terminal spinc only slightly longer than preced- ing ones); pulvilli on 4 proximal tarsomeres, tarsal claws distinctly toothed, arolia present. Abdominal terga unspecialised. Supraanal plate strongly transverse, hind margin shallowly trigonal; right and left paraprocts similar dark plates with pale round central zones (Fig. 17F). Subgenital plate symmetrical with long cylindri- cal, similar styles in posterolateral corners, below them accessory styles with a small basal branch; interstylar border membranous, up- turned, with dense lateral setal brushes (Figs 17D,E). Genitalia as in Fig. 17D: hook on right side, distal region short, curved, without subapi- cal incision; apex of median phallomere modified as in Fig. 17C. Colouration. Reddish brown. Head with dark brown maculae similar to those of cooki (Fig. 16A). Pronotal disk reddish brown with indis- tinct pale spots. Tegmina reddish brown, hyaline, with large dark basal spot in anal vein region (Fig. 17A). Hind wing with costal vein region (except yellowish apex), and plical fold area darkly infuscated, area between discoidal and cubitus veins unpigmented, posterior field light- ly infuscated. Female. Supraanal plate transverse, hind mar- gin convexly rounded. Cubitus vein of hind wing with 1-2 complete and 0-1 incomplete branches (branching may differ on right and left wings of same female). Ootheca. Similar to that of S. tozerensis (Fig. 15B) except there are fewer heavy longitudinal tidges, most of them being very fine. Measurements (2 in parentheses). Length, 8.5 (9.0-9.6); pronotum length x width, 2.5 x 3.9 (2.7 x 3.8-4.1); tegmen length, 7.3 (6.7-7.6). Shelfordina robertsi n.sp. (Figs 18A-F, 29) MATERIAL EXAMINED HootyPe: ¢, Mt Finnigan [15.49S 145.17E], 37km S of Cooktown, NQ, 1100m, Pyrethrum knockdown, 20.xii.1982, L. Roberts; in QMBA (Reg. no. T.11178). PARATYPES: QUEENSLAND. QMBA: same data as holotype, 2d ¢ (1 with terminalia slide 61), 29 2 (1 with genitalia slide 62); same locality and collection method as above, 32 2, 850-1100m, 19-22.iv.1982, RF, Monteith, Yeates, and Cook; Mt Finnigan summit via Helenvale, NQ, 1100m, 1 (terminalia slide 65), 12, 28-30.xi.1985, Monteith, Cook, and Roberts, 1°, pitfall trap, rainforest, 28-30.xi.1985, G. Monteith and D. Cook. MEMOIRS OF THE QUEENSLAND MUSEUM ETYMOLOGY The species is dedicated to Mr Lewis Roberts who collected the holotype. He is a self-taught naturalist who lives at the foot of Mt Finnigan and who has guided many biologists to the sum- mit of this remote mountain (Monteith, pers. comm.). DESCRIPTION Male, Head almost completely hidden under pronotum; eyes wide apart, the space between them about same as distance between antennal sockets (Fig. 18A). Tegmina and wings reaching only slightly beyond end of abdomen. Hind wing with costal veins clubbed, discoidal and median veins straight, simple, cubitus vein straight with 1-2 complete and 0 incomplete branches, apical triangle small (Fig. 18D). Front femur Type A3 (first terminal spine only slightly longer than those preceding it); pulvilli present on 4 proximal tarsomeres, tarsal claws distinctly serrated, arolia present. Abdominal terga unspecialised. Supraanal plate transverse, broadly trigonal, apex rounded; paraprocts similar plates, darkly sclerotised around the borders forming a clear, circular, central zone (Fig. 18F). Subgenital plate symmetrical, sides upturned, distal half membranous, reflexed dorsad under supraanal plate; styles similar, cylindrical, non-articulated, located in posterolateral corners of plate, below them large dark accessory styles, the right and left ones weakly dissimilar, interstylar margin trigonal (Figs 18C,E; in pinned specimens inter- stylar membrane is collapsed and margin appears straight). Genitalia as in Fig. 18C: hook on right side, with small preapical spine; apex of median phallomere modified (Fig. 18B). Colouration. Brown. Head with pale vertex, occiput darker, large shiny, brown macula ex- tending from between eyes to level of antennal sockets, below this a symmetrical macula (Fig. 18A). Pronotal disk weakly mottled brown. Teg- mina hyaline, brown, slightly darker along part of anal vein. Hind wing weakly infuscated, sub- costal vein region slightly darker than rest of wing. Abdominal terga light brown, darkly in- fuscated laterally and on 3 terminal segments. Cerci pale dorsally, segment 6 and apical seg- ment dark. Legs pale, weak distal spots on anterior femoral surfaces. Female. Supraanal plate transverse, broadly rounded, hind margin entire, not reaching hind margin of subgenital plate. Colour markings may be more intense than in male. Abdominal sterna with large dark brown medial spots, and pair of INDO-AUSTRALIAN COCKROACHES 627 Fic. 18. Shelfordina robertsi n.sp., from Mt Finnigan, Qld. D, From holotype, all others from paratypes: A, ?, head; B-F, ¢ d: B, Apex of median genital phallomere (dorsal); C, Subgenital plate and genitalia (dorsal); D, Hind wing; E, Hind margin of subgenital plate showing styles and accessory styles (ventral); F, Supraanal plate and paraprocts (ventral). 628 MEMOIRS OF THE QUEENSLAND MUSEUM large, narrowly separated maculae in middle of Shelfordina uniformis (Hanitsch) subgenital plate. (Figs 19A-G) Measurements (@ in parentheses). Length, 7.9- 8.3 (7.9-8.5); pronotum length x width, 2.4- Shelfordella uniformis Hanitsch, 1933, p.127, fig.1 2.6 x 3.5-3.9 (2.4- 2.7 x 3.6-3.9); tegmen length, (d). 7.0-7.4 (6.5-7.5). Ebnerella uniformis (Hanitsch): 1935, p.14. FIG. 19. Shelfordina uniformis (Hanitsch), d holotype: A, Head; B, Pronotum; C, Supraanal plate and paraprocts (ventral); D, Hind wing; E, Seventh abdominal tergum; F, subgenital plate (ventral); G, Subgenital plate and genitalia (dorsal). INDO-AUSTRALIAN COCKROACHES Possoina uniformis (Hanitsch); Bruijning, 1947, p.218. Shelfordina uniformis (Hanitsch): Princis, 1969, p.872. MATERIAL /AAMINED HOLOTYPE: ¢ (terminalia slide 27), Lake Posso and neighbourhood, Central Celebes, ii.1895, Sarasin (labelled Shelfordella uniformis n.gen., n.sp.,d, R. Hanitsch, 7.vii.1933); in MHNB. DESCRIPTION Male. Interocular space distinctly less than dis- tance between antennal sockets (Fig. 19A); fifth palpal segment slightly smaller than fourth. Pronotum subparabolic (Fig. 19B). Tegmina and wings fully developed, former with longitudinal discoidal sectors. Hind wing with unbranched costal veins uniformly thick throughout, not clubbed, discoidal and median veins un- branched, straight, cubitus vein with 5 complete and 0 incomplete branches, apical triangle well developed (Fig. 19D). Front femur Type A3; 4 proximal tarsomeres with pulvilli, tarsal claws symmetrical, ventral margins minutely toothed, arolia present. Seventh abdominal tergum un- specialised. Eighth abdominal tergum enlarged, surface with spaced setae (Fig. 19E). Ninth ter- gum completely hidden under T8. Supraanal plate with hind margin thickened, convexly rounded not reaching hind margin of subgenital plate; right and left paraprocts similar darkly sclerotised (in part) setose plates (Fig. 19C). Subgenital plate elongate, sides broadly up- turned, distal region reflexed dorsad, concave; small styles flat, located in posterolateral corners, near base of each a large, curved, acces- sory style, interstylar margin weakly concave (Figs 19F,G). [Hanitsch, 1933, fig. 1, did not show the accessory styles in the illustration of the subgenital plate, nor did he mention them in his description]. Genitalia as in Fig. 19G: hook on tight side; distal half of median phallomere modified with a slender filament and a short dark sclerite connected to anterior half by a colourless membrane. Colouration. Head reddish brown, labrum yel- lowish brown, facial region darker (Fig. 19A). Pronotum dark reddish brown without markings, region outside disk somewhat hyaline. Tegmina light reddish brown, hyaline. Hind wing lightly infuscated, costal vein region darker (Fig. 19D). Abdominal terga light brown. Abdominal sterna reddish brown, subgenital plate darker. Cerci dark brown. Legs reddish brown. 629 Female. Unknown. Measurements. Length, 10.0; pronotum length x width, 3.3 x 4.3; tegmen length, 12.0. REMARKS The enlarged and setose eighth abdominal ter- gum, and the unusual median genital phallomere so similar to these structures in S. spinistylifera, clearly show that these two taxa are very closely related (see earlier discussion of species-groups). Shelfordina spinistylifera (Roth) n.comb. Symploce spinistylifera Roth, 1985b, p.154, figs 12A- G(é). MATERIAL EXAMINED SABAH. BPBM: Tawau Residency, Kalibakan R., 48km west of Tawau, primary forest,d holotype of Symploce spinistylifera Roth, 9-18.xi.1958, T.C. Maa (terminalia slide 381; tegmen and wing on 2 slides no. 382). REMARKS The subgenital plate and styles of spinistylifera are similar to those of wailimensis, but the acces- sory styles are much larger and differ in shape (Roth, 1985b; fig. 12C). The eighth abdominal tergum is greatly enlarged and has well spaced setae (Roth, 1985b; fig. 12G). I stated that this was a specialisation, and if it is, then spinis- tylifera and uniformis are the only two species of Shelfordina with a tergal gland. In my descrip- tion of spinistylifera I stated that the male’s geni- tal hook was on the left side. I re- examined my slide but I had separated the 3 main phallomeres and it was impossible to tell the original position of the hook. However, based on its position in uniformis and wailimensis, it probably is on the right side. Because of the similarity of T8 and median genital phallomere, I consider spinistylifera and uniformis more closely related than the former is to wailimensis. Shelfordina wailimensis n.sp. (Figs 20A-G) MATERIAL EXAMINED HOLotTyPeE: d (terminalia slide 234), Wai Lima, Z. Sum.[atra]., Lampongs, xi.xii.1921, Karny and Siebers (no. 221); in HDEO. 630 MEMOIRS OF THE QUEENSLAND MUSEUM DESCRIPTION ment 5 slightly smaller than segment 4. Tegmina Male. Interocular space less than distance be- and wings extending beyond end of abdomen, tween antennal sockets (Fig. 20A); palpal seg- former with longitudinal discoidal sectors. Hind Fic. 20. Shelfordina wailimensis n.sp., d holotype: A, Head; B, B, Abdominal terga 7 to 10 (supraanal plate), and subgenital plate (dorsal); C, Pronotum; D, Supraanal plate and paraprocts (ventral); E, Subgenital plate (ventral): F. Hind wing; G, Subgenital plate and genitalia (dorsal). INDO-AUSTRALIAN COCKROACHES wing with discoidal and median veins simple, straight; cubitus vein weakly concave with 5 complete and 0) incomplete branches, apical tri- angle well developed (Fig. 20F). Front femur Type Az; pulvilli on 4 proximal tarsomeres, tar- sal claws symmetrical, ventral margins minutely toothed, arolia present, Abdominal terga un- specialised. Supraanal plate transverse, trigonal, apex Weakly indented, paraprocts similar plates (Fig, 20D). Subgenital plate subsymmetrical, styles almost similar, cylindrical, small, each in posterolateral corners of plate; at base of each style a large articulated accessory style, rightone only slightly longer than style, left one much larger, apex curved, acute, slender, directed towards right style, interstylar margin concavely excavated (Figs 20B,E,G), Genitalia as. in Fig. 20G, hook on right side, apex acute, without subapical incision; median phallomere, slender. curved, terminating in small, slender, filament; between median and hooklike phallometes a curved, tapering sclerite. Colouration. Datk reddish brown. Head dark reddish brown from occiput to clypeus; basal half of elypeus light reddish brown, remainder yel- lowish brown, labrum light reddish brown (Fig. 20A). Pronotum with large dark reddish brown macula, areas on either side hyaline (Fig. Z0C). Tegmina light reddish brown, hyaline, darker along anterior barder, marginal field, and basal part of anal ficld. Abdominal terga 1-5 pale medially, broad lateral zones reddish brown, T6 mostly reddish brown, T7 with dark brown macula surrounded by whitish band anteromedially, laterally with poorly defined yellowish spats on light brownish background (Fig. 20B), Supraanal plate brownish with yel- lowish spots laterally (near cach cercus) and posteromedially (Fig. 20B). Abdominal sterna reddish brown, Cerci reddish brown dorsally, brown ventrally. Legs reddish brown, tarsi pale. Female, Unknown, Measurements. Length, 11.5; pronotum length x width, 3.6 x 4.6; tegmen length, 14.0. Dyakinodes Princis Dyakimodes Princis, 1951, p.66. Ty Ph SPECIES Ischnoptera centralis Walker = Dyakinodes \veeroriae Princis, by monatypy. DEAGNOSIS Small, slender, ..bitus usually similar to that 631 shown in Fig. 28A. Head exposed, cyes wide apart, sometimes bulging (Figs 22A,C); fifth pal- pal segment longer than fourth. Tegmina and wings fully developed usually extending beyond end of abdomen. Tegmina with discoidal sectors oblique. Hind wings with discoidal vein simple, straight; median vein straight, usually with basal part convexly curved anteriorly; cubitus vein straight except for concavely curved base, with 1, rarely 2, complete branches near distal part of wing, 3-6 small incomplete branches (some ap- pear like cross veins); a characteristic enlarged ‘cell’ is formed by the curved bases of the median and cubitus veins (Pig. 26A, arrow), apical tri- ingle absent (Figs 21D, 22F, 25B, 28E). Anteroventral margin of front femur with 1-4 small, stout spines succeeded by row of piliform spinules, terminating in 2 large distal spines (Type Bo: rarely with a third small distal spine), or rarely with piliform spinules only, terminating in 2 large spines (Type C2); venirul margins of mid and hind femurs sparsely spined (2-4 small spines);, pulvilli present on 4 proximal tar- somercs, tarsal claws symmetrical, simple, arolia present. o! First abdominal tergum un- specialised. Seventh abdominal tetgum with a glandular specialisation (Figs 21B, 22E, 23A,D,G), Supraunal plate symmetrical or al- most symmetrical: right and left paraprocts dis- similar (Fig. 26B,b); with or without a structure arising ventrally near base of cereus (subcercal process: Fig. 26B.a), Subgenital plate weakly asymmetrical, styles small, similar (Fig. 26) or weakly dissimilar (Fig. 22D), Hooklike genital phallomere on left side (Fig. 21B). REMARKS Princis (1969, p.994) lists only | species, Dyakinodes centralis, in the genus. I find 6 new species, described below. The genus is known only from Australia. The male genital hook on the left side places Dyakinedes in the Blattellinae (McKittrick, 1964). However, this should be confirmed by determining if the female rotates the ootheca prior to depositing 1 (Bolin, 1987) SPECIES GROUPS OF DYAKINODES Based on males. |, kurandensis species-group; Hind margin of supraanal plate convexly rounded, entire. Sub- cercercal processes absent. Species: kurander- Sus, uptont. 2. centralis species-group: Hind margin of supraanal plate excised, indented, or deeply divided medially. Subcercal processes present (sometimes greatly reduced). Species: bispinulifera; centralis; fraserensis; penibifida; walerhousei. KEY TO MALES OF DYAKINODES 1. Hind margin of supraanal plate roundly convex, not distinctly indented or excavated medially (Figs Hind margin of supraanal plate distinctly ex- cised medially (e.g., Figs 23A,24A) «0.0... 3 2. Styles almost equal, slender, cylindrical (Figs 21B,E). Setal tuft on T7 arising from distinct mound (Fig. 21B); Pronotum subelliptical (Fig. 22A). Front femur Type B .......... kurandensis Right style short, robust, left style smaller, nar- rower (Fig. 22D). Setal tuft on T7 in shallow depression (Fig. 22E). Pronotum suboval (Fig. 22C), Front femur Typé C.............0... Uphont 3. Supraanal plale transverse, subrectangulat, with pair of long, slender, medial processes (Fig. BERD slezseinisisciseisatesesrerbacosasineseadnsicets fraserensis Supraanal plate nol as above -...,-.)..00cc4F 4. Supraanal plate excised medially forming pair of large, broadly rounded lobes bearing several long spines ulong their margins (Fig. 26B) ssodtvaesacerascavsUasanevareettvawsabinlanittessastecsoaias centralis Supraunal plate nol as above «0... 5 5, Hind margin of supraanal plate with shallow medial excavation and small dark spines. along edge (Pig, 23C). Apex of median genital phallomere divided (Fig. 23B) ......s.ec eee pembifida Hind margin of supraanal plate much more deeply excavated medially forming pair of tri- angular lobes. Apex of median genital phal- lomere Not divided -..........ccccceeceseeeesereeesenee 6 6. Outer margins of triangular lobes of supraanal plate lined with small dark spines (Figs 24A,25A) Satetdgs stehbasorkiasinettpesctlaaheniaeseginorsens walerhousei Triangular lobes of supraanal plate each ter- MEMOIRS OF THE QUEENSLAND MUSEUM Mminaling in a spine, outer margins not spinose (Fig. 27B) ...c.ceesesseeeeeeseeseeeesesesseeae Déspinulifera Dyakinodes kurandensis n.sp. (Figs 21A-E,22A,29) Material EXAMINED HOLOTYPE: ¢ (lerminalia slide 458), Kuranda north Queensland, 200m, 13,iii.1956, J.L, Gressitt; in BPBM. PARATYPE: QUEENSLAND. QMBA: Windsor Tbld, 35km NNW of Carbine, NO, Bargoo Ck, RF, 850m, 19, 15-18.iv.1982, Monteith, Yeates, and Cook. DESCRIPTION Male. Interocular space about same as distance between antennal sockets, slightly greater than interocellar space; fifth palpal segment enlarged, longer than the fourth. Pronotum subelliptical (Fig. 22A). Cubitus vein of hind wing thicker than other veins, cell formed by bases of median and cubitus veins small, cubitus vein with 1 complete and 3 incomplete branches, apical tri- angle absent (Fig, 21D). Front femur Type B2 (with 4 small, stout proximal spines), pulvilli present on 4 proximal tarsomeres, tarsal claws simple, symmetrical, arolia present. Seventh ab- dominal tergum with large, semicircular, con- vexly taised, median elevation bearing dense setal tuft anteromedially (Fig. 21B). Supraanal plate with hind margin broadly rounded, entire, reaching hind margin of subgenital plate (Fig. 21B); paraprocts dissimilar, rectangular, right one larger without large, spinelike processes (Fig. 21A). Subgenital plate weakly asymmetri- cal, styles small, cylindrical, similar, widely separated, interstylar margin weakly oblique {Figs 21B.C,E), Genitalia as in Fig. 21E: hook on left side, with a subapical incision, median phallomere with distal region curved obliquely, Colouration. Head dark brown, lower part of face lighter, gradually becoming darker dorsad. Pronotum dark brown, lateral edges and disk lighter. Tegmina brown, transparent, anterior area darker then posterior region; hind wings weakly infuscated, veins very distinct, dark brown, Abdominal terga light brown, lateral bor- ders darker. Female. \nterocular distance slightly greater than space between antennal sockets. Hind wing with cubitus vein with 2 complete and 4 incom- plete branches. Front femur Type Ba, with 4 stout proximal spines, Supraanal plate trigonal, INDO-AUSTRALIAN COCKROACHES 0.5 mm 0.5 mm 633 Fic. 21, Dyakinodes kurandensis 0.sp., 3 holotype: A, Supraanal plate and paraprocts (ventral); B, Abdominal tergum 7 to supraanal and subgenital plates (dorsal); C, Subgenital plate (ventral); D, Hind wing; E, Subgenital plate and genitalia (dorsal). rounded apex almost reaching hind margin of subgenital plate. Top of head blackish shading to dark brown from middle of antennal sockets to yellowish clypeus and labrum. Pronotum with lateral edging yellowish brown, disk reddish brown, lateral and posterior regions dark brown, Tegmina reddish brown, humeral vein area dark brown, anterior edging and subcostal area yel- lowish, that part of right tegmen covered by left, colourless. Abdominal terga and sterna light brown. Cerci brown dorsally, ventral surface darker. Legs light brown. Measuremenis (@ in parentheses). Length, 7.5 (6.5); pronotum length x width, 1.9 x 2.7 (2.0 x 2.7); tegmen length, 7.7 (7.3). Dyakinodes uptoni n.sp. (Figs 22B-F, 29) MATERIAL EXAMINED HoLotyre: ¢ (terminalia slide 145), Bamaga, Cape York, Queensland, 26.ii1.1964, LF.B. Common and M.S. Upton; in ANIC, MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 22. Dyakinodes spp. A, D. kurandensis n.sp., d holotype, pronolum, B-F, Dyakinedes uptoni n.sp., 3 holotype: B, Supraanal plate and paraprocts (ventral); C, Pronotum; D, Subgenital plate and genitalia (dorsal); E, Tergal gland region on seventh segment; F, Hind wing. DESCRIPTION Male. Head with hardly any punctations be- tween ocellar spots; interocular space slightly less than interocellar distance; fifth palpal seg- ment larger than fourth, Pronotum suboval (Fig. 22C). Cubitus vein of hind wing with 1 complete (but apparently split in middle) and about 3 in- complete branches, cell formed by bases of median and cubitus veins, small, apical triangle absent (Fig, 22F). Front femur Type C2 (stout proximal spines apparently absent). Seventh ab- dominal tergum with setal tuft on small mound in shallow depression (Fig. 22E). Supraanal plate with hind margin convex, some minute dark spines along margin ventrally; right paraproct with large, curved, upically bifid process, left paraproct simpler, with several large setae distal- ly (Fig. 22B), Subgenital plate broadly rounded, tight style short, bulbous, located on thickened dorsal region, left style smaller, more slender (Fig. 22D). Genitalia as in Fig. 22D: hook on left side, median phallomere curved on distal half, apex acute; some sclerites of right phallomere were probably lost in slide preparation. INDO-AUSTRALIAN COCKROACHES Measurements, Length, 7.7; pronotum length x width, 1.6 x 2.0; tegmen length, 8.0. Female. | am provisionally placing the follow- ing female here (non-paratype), pending collec- tion of males from the same locality. QUEENSLAND. ANIC: 3km NE of Mt Webb, near Cooktown, 12, 3.x.1980, T.A. Weir and R.A, Barrett. The specimen Jacks the dark areas on the promnicen and base of the coxac, and the dark frown regions.on the subcostal and costal areas of the tegmina are greatly reduced. Measurements. Length, 8.0; pronatum length x width, 1,8 x 2.2; tegmen length, 8.2. Dyakinodes penibifida n.sp. (Figs 23A-F,29) MATERIAL EXAMINED HOLOTYPE; ¢@ (terminalia slide 148), Slaty Creek, 3ikm NE by E of Tenterfield, New South Wales, 16.xi.1982, D.C.F. Rentz and C.D. MacNeill, Stop 90; in ANIC. ETYMOLOGY The specific name refers (o the divided apex of the male’s median genital phallomere (virga). DESCRIPTION Male. Head with interocellar region and vertex minutely punctulaic; interocular distance about same as distance between ocellar spots; fifth palpal segment longer than fourth. Pronotum suboval (Pig. 23E). Cubitus vein of hind wing with I complete and 6 small incomplete branches, cell formed by curved bases of median and cubitus veins, small; apical triangle absent (Fig. 23F). Front femur Type Ba, with 2 stout proximal spines, pulvilli present on 4 proximal larsomeres, tarsal claws simple, symmetrical, arolia present. Seventh abdominal tergum with pair of deep, setose fossac, divided medially by mound bearing dense dark setal tuft (Figs 23A,D). Supraanal plate trigonal, apex shallowly excavated, deflexed, margins on either side of indentation with very small dark spines (Figs 23A,C); subcercal processes terminating in small spines: right paraproct with several robust, spinelike processes, left one simpler (Fig. 23C). Subgenilal plate with pair of widely spaced, al- most similar cylindrical styles (Figs 23A,B). Genitalia as in Fig. 23B: hook on Ieft side, 635 median phallomere apically bifid, right phal- lomere with subobsolete cleft sclerite. Colouration, Head dark brown, vertex and oc- ciput slightly lighter than rest of face. Pronotum dark brawn, medial region lighter (Fig. 23E), Tegmina dark brown, basolateral half darker than rest of wing cover. Hind wing with part of subcostal and costal area and apex, darkly infus- cated (Fig. 23F), Abdominal terga light brown, sides and terminal segments dark brown, Ab- dominal sterna with medial area up to subgenital plate, light brawn, remainder dark brown. Coxae with brown basal maculae, remaining parts of legs very pale. Measurements. Length, 10,7; pronotum length x width, 2.4 x 3.0; tezmen length, 12.0. Dyakinodes waterhousei n,sp- (Figs 24A.B, 25SA-E, 29) MATERIAL EXAMINED HoLotyrr: 3, Mt Moffat, N.P., CO, Top Sheller Shed, 1000m, 10-12.xii.1987, Monteith, Thompson, Yeates, in OMBA (Reg. No, T.11452). PARATYPES: QUEENSLAND, ANIC: Mogzgill, near Brisbane, | (terminalis slide 150). ix,1966, G. Mon- teith, QMBA: Lake Clarendon, SEQ, 2° 9, 11.xi.1971, E.C. Dahms. ETYMOLOGY The species is dedicated to Dr Douglas Watet- house, retired Chief of CSIRO, Division of En- tomology. DESCRIFTION Male. Head with few punciations on vertex; interacular space about same as distance be- tween ocellar spots; fifth palpal segment swol- len, longer than fourth, Pronotum suboval (Fig. 25C), Tegmina and wings fully developed. Hind wing with cell formed by curved bases of median and cubitus veins small but distinct; cubitus vein with 1 complete and some small incomplete branches and cross veins, apical triangle absent (Fig. 25B). Front femur Type B2, with 3 or 4 stout proximal spines: pulvilli present on 4 proximal tarsomeres, tarsal claws symmetrical, simple, arolia well developed. Seventh abdominal ter- gum with a pair of fossae divided by large dark selose mound (Fig. 25D). Supraanal plate hind margin deeply excavated forming pair of lobes with small dark spines near their acute apices; bottom margin of excavation truncate, membranous, colourless, with few dark spines 636 MEMOIRS-OF THE QUEENSLAND MUSEUM FIG. 23. Dyakinodes spp. A-F, D, penibifida n.sp., d holotype: A, Seventh abdominal tergum to supraanal and subgenital plates; B, Subgenital plate and genitalia (dorsal); C, Supraanal plate and paraprocts (ventral); D, Seventh abdominal tergum; E, Pronotum; F, Hind wing. G, Dyakinodes centralis (Walker), 3 from Upper Kangaroo River, NSW, Australia, glandular region of seventh abdominal tergum. INDO-AUSTRALIAN COCKROACHES 637 Fic. 24. Dyakinodes waterhousei n.sp., d paratype from Moggill, Qld: A, Supraanal plate and paraprocts (ventral); B, Subgenital plate and genitalia (dorsal). 638 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 25. Dyakinodes waterhousei n.sp., 3 variant from 4 miles southwest of Mudgee, NSW: A, Supraanal plate and paraprocts (ventral); B, Hind wing; C, Pronotum: D, Gland on seventh abdominal tergum; E, Subgenital plate and genitalia (dorsal). ventrally; subcercal processes reduced; left paraproct with single, apically rounded process, right paraproct with pair of large spinelike processes, one strongly curved, the other apically divided (Fig. 24A). Subgenital plate with pair of weakly dissimilar styles, each with a small lateral, apically spined tubercle (Fig. 24B). Genitalia as in Figs 24B (hook lost in slide preparation), 25E: hook on left side, median phallomere slender, apically acute, right phal- lomere includes a reduced cleft sclerite. Colouration. Head dark brown, occiput lighter reddish brown, clypeus pale, antennae with several black basal segments, rest pale. Pronotum with dark brown lateral areas which do not reach hind margin, remainder lighter (Fig. INDO-AUSTRALIAN COCKROACHES 639 Fic. 26. Dyakinodes centralis (Walker), 3 from Upper Kangaroo River, NSW: A, Hind wing (arrow indicates cell formed by curved bases of median and cubitus veins); B, Supraanal plate and paraprocts (ventral); C, Genitalia (dorsal); D, Abdominal tergum 7 to supraanal and subgenital plates; E, Distal region of subgenital plate showing styles (ventral), Abbreviations: a, left subcercal process; b, right paraproct; c, right phallomere; d, median phallomere; e, hooklike left phallomere. 25C). Tegmina with humeral area dark brown, remainder light brown, hyaline. Subcostal and costal vein areas of hind wing yellowish, remainder almost colourless. Abdominal terga yellowish brown with broad lateral zones dark brown. Abdominal sterna with broad lateral areas, and subgenital plate dark brown, remainder lighter reddish brown, Cerci dorsally pale with dark brown lateral edges, ventrally uniformly dark brown, Legs pale except for dark coxal bases. Female. Cubitus vein of hind wing with 1 complete and 5-6 incomplete branches and cross veins, Front femur Type Bz, with 2 small stout proximal spines. Supraanal plate strongly trigonal, rounded apex extending slightly beyond hind margin of subgenital plate. Head blackish, including occiput; antennae uniformly palc. Pronotum with more extensive dark brown areas than jn male, Measurements (2 in parentheses). Length, 10,8- 12,3 (8.3-10.4); pronotum length x width, 1,9-2.4 x 2.3-3.0 (1.9- 2.0 x 2.3-2.4); tegmen length, 10.0-1 1.1 (8.9-10.0). 40) VARIANT NEW SOUTH WALES. ANIC: 4 miles SW of Mudgee, 32,368 (49.356, 1o (terminalia slide 151), 18.x1.1968, Britton and Misko, This male shows some minor differences in the shape of the excavation of supraanal plate, and right puraproct (cp. Figs 24A, 25A), Dyakinodes centralis (Walker) (Figs 23G, 26A-E, 29) Ischnoptera centralis (Walker), (868, p.120 (2). Apolyta centralis (Walker): Tepper, 1893, p.48. Ischnoptera brunneonigra Tepper, 1895, p.155 (2); Shaw, 1916, p.86 (4) |[misidentification|; Princis, 1969, p 1002 (sp. meertae sedis); Roth, 1985a, p.212 (sp, dHeerfae sedis), o,5yn. Dyakinodes victoride Princis, 1951, p67, plo, figs 66,67 (cf), 1959, p.140, Dyakinodes centraliy (Walker): Princis, 1959, p.140; 1969, p.994, MATERIAL EXAMINED Horotyre:?, South Australia, presented by R. Bakewell: in BMNH. AppITiIONAL = MATERIAL: VICTORIA. NMVM: Fermshaw, 2 holotype (no, 4457) af Ischnoptera brun- neonigra Tepper. UZMC: ¢ holotype of Dyakinodes victortae Princis. Hauschild [no exact locality]. AUSTRALIAN CAPITAL TERRITORY. ANIC: Black Mt., 35.178 149.06E, Led, 54.1987, 2945, 19.41.1987, D.C.F. Rentz; Wombat Ck, 6km NE of Piccadilly Clreus, 750m, 35.198 148,51E, 19, flight intercept window (rough trap, i.1985, Weir, Lawrence, and Johnson: Mt Ainslie, 1g, TP. 10.1.1966, 1c, 11,1,1966, M.S. Upton: Black Mt,, light trap, 1c, 12.xi1.1967, 1.F.B. Cammon. New Soutit WALES. ANIC: Depol Beach, 10 miles NE of Balemmuns Bay, 1d. 27.x.1967, LF.B.Comman, 7 miles west of Rosebank, 1700ft, 1d, 8.x1.1961, LF.B. Common ind M.S. Upton; Chiswick Exp. Sin, | (terminal abdominal segments missing), 10.1.1970, RI. Roberts. MCZH: Upper Kangaroo River, 1d (terminalia slide 223), 281.1968, KHL. Key, DARA: Barrengr. F.R. [?Barrengarry],3d oat pressure lamp, 15.1.1966, C.E, Chadwick; Bundeen, 1d, 11- 12.x7, 1958, R.H. Mulder. SAMA: Porest Reefs, Id. Lea. QUEENSLAND. ANIC: ~Coonardou’, Fletcher, 14 (ler- minalia slide 147), 4.xi1,1966. T,.G, Campbell. DESCRIPTION Male. Head with interocellar tegion minutely punctulate; interocular distance about same as MEMOIRS OF THE QUEENSLAND MUSEUM space between antennal sockets; last palpal seg- ment enlarged. Pronotum suboval (Pig. 26F). Tegmina and wings fully developed. Hind wing with cell formed by curved bases of median and cubitus rami well developed, cubitus with 1 com-. plete and 2-S incomplete branches, apical tri- angle absent (Fig. 26A). Front femur with 1-4 large proximal spines, followed by row of piliform spinules, 2 (rarely 3) large terminal spines (Type B2 or Bs); pulvilli on 4 proximal tarsomeres, tarsal claws symmetrical, simple, arolia present. Seventh abdominal tergum with transverse fossa and setose medial mound (Figs 23G,26D). Supraanal plate with hind margin divided medially forming two broadly rounded lobes bearing large spines along the edges; sub- cetcal processes capped by several small spines; Tight paraproct large, bulbous, bearing stout, dark spines, left paraproct smaller, with fewer dark spines (Fig. 26B). Subgenital plate with pair of similar, cylindrical styles (Figs 26D,E), Genitalia us in Fig. 26C: hook on Jeft side with a subapical incision, median phallomere slender, tapering to acute apex. Colouration. Head dark brown, vertex yel- lowish brown; palps whitish or pale, terminal segments weakly infuscated. Pronotal disk yel- lowish brown, lateral regions dark brown (Fig. 26F). Tegmina with mediastine field and proximal half of scapular field dark brown. Ab- dominal terga brownish yellow, lateral portions and T9 and supraanal plate dark brown, Ab- dominal sterna medially brownish yellow, broad lateral regions dark brown, Cerci brown dorsally and ventrally. Coxae with basal dark brown maculae, those on mid and hind legs much larger iper spots on front coxae; remaining parts of legs ale. 4 The amount of dark brown on head, pronotum, tegmina, abdominal segments, and legs varies, A male from Fletcher, Queensland is very pale, lacks dark brown on pronotum and tegmina, although coxac and abdominal segments have dark markings, its supraanal plate, styles, and genitalia are similar to typical material, A male from Depot Beach, New South Wales., has all parts of the legs dark brown. Female. Supraanal plate trigonal, apex rounded extending slightly beyond hind margin of supraanal plate. In very dark forms, coxae are completely dark brown. Measurements. (2 in parentheses). Length, 11.4)-13.5 (11.7-12.8); pronotum Jength x width, 2.5-2.8 x 3.4-3.7 (2.6-2.8 x 3.4-3.6); legmen length, 11.8-13.8 (11.7-12.5). INDO-AUSTRALIAN COCKROACHES 641 E Fic. 27. Dyakinodes bispinulifera n.sp., ¢ holotype: A, Seventh abdominal tergum to supraanal and subgenital plates; B, Supraanal plate and paraprocts (ventral); C, Hind wing: D, Subgenital plate and genitalia (dorsal); E, Pronotum; F, Seventh abdominal tergum. in REMARKS Tepper (1895, p.155) described /schnoptera brunneontgra from a female from Fernshaw, Victoria, and indicated it is housed in the Nation- al Museum Melbourne. This specimen is very dark and greasy and it is difficult to determine its colour, However, it is clearly a Dyakinodes be- cause its wings, though badly damaged, show the characteristic cell formed by the curved bases of the median and cubitus veins. Two other male specimens from Fernshaw, and labelled /schnop- tera n.sp, by Tepper, have no type status, because he described only the female. One of these specimens is centralis. The other male lacks most of the abdomen, bul TI and T2 are present and the former segment is specialised and the median und cubitus veins do not form abasal cell. It is not a Dyakinades, although its front femur is Type B, Shaw (1916, p.86) described what he believed to be the male of /schnoplera brunneonigra from Healesville, Victoria, and called them cotypes. His description clearly shows that itis not brun- neanigra because TL and T7 are modified, and the front femur is Type A (4 large spines occupy- ing the middle third succeeded distally by a row of smaller spines). Dyakinodes bispinulifera n.sp, (Figs 27A-F. 29) Marerial EXAMINED HoLoryre, & (terminalia slide 149), Ruby Creek, 1 mile west of Amosfield, New Suuth Wales, 15.x01.1966, T.G, Campbell; in ANIC, EVYyMOLOGY The specific name refers to the 2 spines, 1 at the apex of each triangular lobe formed by the deep excavation of the supraunal plate hind margin. DESCRIPTION Male, Head minutely punctulate from inter- occllar region to vertex; interocular space vbout same as distance between antennal sockets. Pronolum suboval (Fig, 27E), Cell formed by curved bases of median and cubitus veins dis- tinet; cubitus vein with | complete and 5 incom- plete branches, apical triangle absent (Fig. 27C). Anteroventral margin of frant femur Type B2, with 2 Small, stout proximal spines; pulvilli present on 4 proximal tarsomeres, tarsal claws symmetrical, simple, arolia present. Seventh ab- dominal tergum with a fossa divided by a mound MEMOIRS OF THE QUEENSLAND MUSEUM bearing dense, golden, sctal tuft (Figs 27A,F). lind margin of supraanal plate with deep U- shaped excavation forming pair of deflexed lohes, apices curled, each lobe bearing an apical spine (Figs 27A,B); subcercal processes dis- similar, right one more robust without apical spine (this may be aberrant), left one trigonal with apical spine; paraprocts dissimilar, right one complex with dark sclerite bearing large spinclike processes (Fig. 27B). Subgenital plate trigonal, styles small, weakly dissimilar (Figs 27A,D). Genitalia as in Fig. 27D: hook on left side; median phallomere slender, rodlike, apex acute, Colouration. Head dark brown, occiput yel- lowish brown (Fig. 27E). Pronotum yellowish brown, lateral regions darkly infuscated for about two-thirds its Jength (Fig. 27E). Tegmina hyaline, pale, subcostal area weakly infuscated. Abdominal terga yellowish brown, broad lateral areas and terminal segments darker, Abdominal sterna with broad, light brown,medial stripe, lateral borders and subgenital plate dark brown. Cerci brawn. Basal regions of coxae dark brown, remaining parts of legs very pale. Female. Unknown. Measurements, Length, 11.0; pronotum length x width, 2.0 x 3,1, tegmen length, 12.5, Dyakinodes fraserensis n.sp, (Pigs 28A-F, 29) MATERIAL EXAMINED HoLotType: & (terminalia slide 144), Fraser Island, Queensland, 1¥km SSW of Indian Head, 25.108 IS3,07E. 11- 4.x.1978, D.C.F, Rentz and J. Balder- son, Stop 120; in ANIC. PARATYPE: Fraser Island. ANIC: same data as holatype, 1°. DESCRIPTION Male. (Habitus shown in Fig. 28A). Head with vertex minutely punctulate; interocular space about same us distance between ocellar spots; fifth palpal segment longer than fourth. Tegmina and wings fully developed extending beyond end of abdomen, Hind wing with a small cell formed by curved bases of median and cubitus veins; cubitus vein with | complicte and about 3 incom- plete branches and few cross veins, apical tri- angle absent (Fig. 28E). Anteroventral margin of front femur Type B2, with 2 stoul proximal spines; pulvilli present on 4 proximal tarsomeres, tarsal claws symmetrical, simple, arolia present. INDO-AUSTRALIAN COCKROACHES 643 Fic. 28. Dyakinodes fraserensis n.sp., 3 holotype: A, Habitus; B, Abdominal tergum 6 to supraanal and subgenital plates; C, Supraanal plate and paraprocts (ventral); D. Seventh abdominal tergum; E, Hind wing; F, Subgenital plate and genitalia (dorsal), 644 Duekinodes Vv centralia V waterhkouyzet @ freszrensia DO pentbifida @ biepinultfera O untond © kurandendis Shelfordina 4 orchidge W cooks tr yeacteat @ robertsi 4 toferensiia ret ee Fic, 29, Distribution of Shelfardina spp. and Dyakinodes spp. in Australia (the D. waterhousei record in New South Wales refers to the variant). Seventh abdominal tergum with large transverse fossa and medial setose mound (Fig. 28B); KOH cleared preparation shows a pair of large, rounded sacs bearing scattered setae, in addition to the dense medial tuft (Fig. 28D). Supraanal plate transverse with pair of long, dark, slender processes that curve ventrad, their apices reach- ing hind margin of subgenital plate (Fig. 28B); between these processes and cerci, supraanal plate is membranous, colourless, with small sub- cereal processes cach bearing an apical spine MEMOIRS OF THE QUEENSLAND MUSEUM (Fig. 28C). Subgenital plate broadly rounded, convex, styles cylindrical, right one slightly more robust, arising from small tubercle on dor- sal surface of plate (Fig. 28F). Genitalia as in Fig. 28F: hook on left side; median phallomere slender, slightly enlarged distally, apex acute, Colouration. Head black, clypeus pale, labrum brownish. Pronotum yellowish brown with broad lateral areas Very dark brown (Fig. 28A). Teg- mina hyaline, yellowish brown, subcostal and basal half of costal areas infuscated (Fig. 28A). Costal region of hind wing light yellowish brown. Abdominal terga pale, laterally infus- cated, Basal regions of coxae darkly infuscated, remaining parts of legs very pale. Female. Supraanal plate trigonal, apex extend- ing slightly beyond end of abdomen. Abdominal terga and sterna pale, lateral borders and terminal segments blackish. Measurements (9 in parentheses). Length, 8.5 (8.3); pronotum length x width, 1.9 x 2.3 (2.0 x 2.5); tegmen length, 9.1 (9.2). Dyakinodes sp. A MATERIAL EXAMINED NEw SoutH Wates. ANIC: Durras North, near Batemans Bay, 12, 10.i.1979, H.M. Cameron. DESCRIPTION Female. Head hidden under pronotum; punctulations on vertex indistinct; fifth palpal segment enlarged, longer than fourth. Pronotum subelliptical. Tegmina and wings fully developed. Hind wing with cell formed by curved bases of median and cubitus veins very large; cubitus vein with 2 complete and 3 in- complete branches, apical triangle absent. Front femur Type B3, with 6 stout proximal spines, pulvilli present on 4 proximal tarsomeres, tarsal claws simple, symmetrical, arolia present. Supraanal plate trigonal, apex rounded protrud- ing slightly beyond subgenital plate. Colouration. Yellowish brown. Head pale, light reddish brown area between antennal sock- ets and eyes, blending gradually with pale oc- ciput and lower part of face, interocular space almost smooth. Pronotum, tegmina, abdominal terga and slerna, and legs (including coxae), pale, dark infuscation absent. Dorsal surface of cerci pale, ventrally brown. Measurements. Length, 9.0; pronotum length x width, 2.5 x 3.3; tegmen length, 10.0. INDO-AUSTRALIAN COCKROACHES Anaplecta Burmeister Anuplecia Burmeister, 1838, p.494, Bruijning, 1948, p.43; Hanitsch, 1928, p.6; Hebard, 1929, p.27; Prin- cis, 1965, p.367. TYPE SPECIES Anaplecta lateralis Burmeister, by designation (Kirby, 1904, p.66) (type locality: Colombia, South America). REMARKS [have not seen Burmeister’s types of the type- specics Anaplecra lateralis from Colombia, However, | examined 5 males (1 with terminalia slide 426) from Panama (in ANSP) which Hebard (1920, p,15) determined as /ateralis. I have scén additional specimens in MCZH from Barro Colorado Island, Panama Canal Zone (1d, 19), and from Agua Caliente, Guatemala (143 4,1 with terminalia slide 248, and 109 ). The type-species agrees with the diagnostic char- acters used for the genus, including basic fea- tures of the male genital phallomeres. Wanitsch (1915, p.3J) in his diagnosis of Anaplecta stated that the males lacked styles. Later in describing various species in the genus he stated that styles were present in some taxa but not others. The styles are small and may be overlooked unless slides are prepared of the sub- Henital plate. This is true of the types of Anaplec- fa vittala which Hanitsch described as females bul are males (Fig. 31D). Females lack styles and have a valvular subgenital plate. Descriptions of Avaplecia spp. have stressed wing venation and colouration (c.g. Shelford, 1906,pp.240-7), Often only 1 or 2 specimens were available or used by describers and there is no indication of the extent of variation in ‘diagnostic’ characters. Bruijning (1948, p.44) pointed out that differences in the number of costal veins and in the number of cross veins between the discoidal and median rami are vari- able and have no specific value; this is probably true in many cases. Colouration also may vary, and sometimes colour is not distinctive, Many of the described species are based on females and even when males were available their genitalia were not studied. These structures should be used, if possible, for specific determinations and to show species relationships. McKittrick (1964, p.26) studied only 3 Anaplecta (2 South American and | African) identified to genus only and stated that *... more than | genus is repre- sented. to judge by differences in female 645 genitalia." The genus needs revision and at least the known species should be redescribed, using males when this sex is available. AFFINITIES Princis placed Anaplecta in the Anaplectidae: Anaplectinae. | follow McKittrick (1964, p.25) who believed the genus to be in the most primi- tive subfamily of the Blattellidae and one which has structures that ‘... bridge the gap between 2 superficially exceedingly different groups.’ (i.e., Polyphagidae and Blattellidae). DiAGnosis Size smal]. Tegmina and wings usually fully developed, rarely with reduced tegmina and ves- tigial Wings (1 species). In fully developed flight organs, tegmina narrow, usually with about 3 longitudinal or sublongitudinal sectors, Hind wings with cubitus vein simple, costal veins sometimes thickened distad, sometimes indis- tinct, with large subtriangular apical area (appen- dicular ficld) which when in repose is folded longitudinally and then reflexed over rest of wing: two weakly developed veins, one lon- gitudinal along fold, the other somewhat oblique, may occur in appendicular field, other veins lacking; hind wing often infuscated, costal vein region and part of appendicular field darker than remaining parts. Anteroventral margin of front femur with few large proximal spines followed by row of piliform spinules, terminating in 2 latge spines (Type Bz); tarsal claws simple, sym- metrical, unspecialised (not serrated); pulvilli absent, or apparent on fourth tarsomere only; arolia present. d: Supraanal plate with medial specialisation. Styles present, similar or differing in size and/or shape. Genital hook on left side. 2: subgenital plate valvular. KEY TO ADULT ANAPLECTA OF AUSTRALIA |. Tegmina reduced reaching to about T3 (Fig. 33A). Hind wings vestigial. Clypeus swollen (Fig- BSB) oeseceieere ee eel paar Drachyplert Tegmina and wings fully developed reaching to end of abdomen ar beyond. Clypeus not swol- TED (Piggy SY A)urractrinscnsnssosnsesescases usages texseeasesie 2. Tegmina with a pair of dark maculae separated by a transverse pale band (Fig. (VE), or maculae joined on one side (Fig. 30K) ........... calasoma 646 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 30, Anaplecta calosoma Shelford, A-C, 3 from 3km ENE of Mt Tozer, Queensland, head, pronotum, and terminal abdominal terga, D-K, Hind wings and tegmina: D,E, 3 from 11km ENE of Mt Tozer: F,G, 2 from Awar Bush, Papua New Guinea; H,1, ¢ from 6km ENE of Mt Tozer; J,K, 2 from Moses Ck, 4km north by east of Mt Finnigan (veins not shown in Figs E,G,I). Scales (mm): A-C, 0.5; D, 2.0; E-K, same as D. INDO-AUSTRALIAN COCKROACHES 647 Fic. 31. Anaplecta spp. A, Anaplecta calosoma Shelford, d from 11km ENE of Mt Tozer, Queensland, subgenital plate and genitalia (dorsal). B-D, Anaplecta vittata Hanitsch, 3 paratype from Singapore: B, Left tegmen; C, Left wing; D, Subgenital plate and genitalia (dorsal; genital phallomeres: a, L3; b, L1; c, L2vm; d, R3 [L = left, and R = right phallomeres - terminology after McKittrick, 1964]). Scales (mm): A, 0,25; B,C, 2.0; D, 0.25. 44 Tegmina brown, unbanded (Fig, 32C), some- limes witha pale spot behind the anal vein (Fig. F2DY cccccccseicscrisanpyityeeeeese ees OMStraliensis DISTRIBUTION Anaplecta is a large genus with more than 80 species in Central and South America, Asia and Africa (Princis, 1965, pp.367-80; 1971, p.1144; Bey-Bienko, 1969, p.8349; 1970, p.533; Asahina, 1977, p.277). Until now it has not been recorded from Australia. Rehn (1904, p,542) described Anaplecta platycephala from Cairns but Hebard (1929, p.27, footnote 14) pointed out that it did not belong in this genus and later (Hebard, 1943, p.10) placed it in Mediastinia Hebard. Distribution of Australian Anaplecta is shown in Fig. 34; not shown is the Northern Territory (near Darwin) record of A, australiensis, Anaplecta mmarated Shelford (Figs 30A-K, 31A, 34) Anaplecta calosoma Shelford, 1912, p.51 (2 ); Bruijn- ing, 1947, p.215; Princis, (1953) 1954, p.212, HOLOTYPE (not examined) Y, German New Guinea (= Papua New Guinea), Tamimundung (= mouth of the Tami River); Shelford did not indicate where the type is housed; it is not in HDEO. MATERIAL EXAMINED The following specimens have the legminal maculae completely separated by w transverse band (Figs 30E,G.D: ParuA New Guinea, ISNB: Awar Bush, 1d (ter- minalia slide 2), 19, 6.vi. L982, 1%, 21.vid982, P. Groutuert. QUEENSLAND, ANIC; 3km ENE of Mt Tozer, 12.445 I43.14E, 2k.v.- dvi 1986, 492 (Malaise trap/ethanol), J.C. Curdale, 4¢ d, 12 (Malaise trap), D.H. Colles», Quinn Park, Claudie River, 12.435 IS3.17E, 1d. 12- 20.1.1985, B.D. Edwards and B. Hacobian. The following were collected by T. Weir and A Calder) Skm ENE of Mt Tozer, 12.445 143.146, 30 3,32 9, 28.vi-4.vii. 1986) Ohi BNE of Mt Tozer, 12.445 143.16E, 1d, 30. vi,1986; 7km ENE of Mt Tover, 12.4235 143.166, 1%, L.vii. 1986; 9km ENE of MI Tozer, 12.435, [43.17E, 204d, 19%, 5- 10.viil. 1986; 1 Ikm ENE of ML Tozer, 12.435 143. 16E, 222. 1)-t6.vi 1986 The following were collected by D.C.F. Reate: Li km ENE of Mt Tozer, near Iron Range National Park, {2.438 143.18E, rainforest mar- gin, 19. 11.vii. P98G, at light, 12. (lerminalia slide MEMOIRS OF THE QUEENSLAND MUSEUM 185), 6342, 1022, 11.vit.1986; 9km ENE of Mt Tozer, 12.435 143, 18E, Iron Range National Park, at light in rainforest, 26 3, 10.vii.1986; 3km ENE of Mt Tozer, near Iron Range National Park, 12.445 143, 14E, at light, 54,69 2, 28.vi.-4.vii.1986; 2km NNE of MI Tozer, Iron Range National Park, 12.448 143,13E, at light, 19, 3.vii. 1986. OMBA: West Claudie R., Iron Range, flight trap, rainforest, 2d d, 39 2, 3-10.xi1.1985, G. Monteith and D, Cook. DESCRIPTION Male. Interocular space greater than distance between antennal sockets (Fig. 30A). Pronotum subelliptical (Fig. 30B). Tegmina and wings reaching to end of abdomen or slightly beyond. Hind wing venation as in Figs 30D,F,H- Anteroventral margin of front femur Type B2; pulvilli apparently absent, arolia present. Supraanal plate with medial specialisation, hind margin convexly rounded (Fig. 30C). Subgenital plate weakly asymmetrical, interstylar margin practically straight (Fig. 31A). Genitalia as in Fig. 31A; L.1 with long, slender, curved filament. Colouration. Head with labrum and clypeal region pale yellow, remainder dark brown (sometimes nat uniformly sa) (Fig. 304A). Prono- tal disk yellowish without markings, surrounding region hyaline (Fig. 30B). Tegmina with pair of dark brown to blackish maculae that are com- pletely separated by transverse pale band posterior to anal field, marginal field also pale or hyaline; size of maculae, especially distal one may vary (Pigs 30E,G,I; see variant noted below). Hind wing with appendicular field and costal vein area infuscated, remaining areas less so (Figs 30D,F,H). Abdominal terga brownish yellow, lightly infuscated, glandular setae on T10 golden, surrounding area very pale (Fig. 30C). Abdominal sterna, legs, and cerci, pale brownish yellow, Female. Supraanal plate with hind margin con- vexly rounded. Colour essentially as in male excep! base and lateral parts of valves near apex of subgenital plate reddish brown or dark brown. Measurements (? in parentheses). Length, 3,9- 4.7 (3,8-5,0); pronotum length x width, 1,1- 1,2x 1.5-1,7(1.0-1,4.x 16-18); tegmen length, 3.1-4.5 (3.5-4.5). VARIANT This variant generally is larger than typical material and the tegminal maculae are joined on their anterior margins along the marginal field (Fig. 30K). Female subgenital plate dark brown INDO-AUSTRALIAN COCKROACHES laterally on cach side of valves. Male genitalia similar to typical material. Most records near Kuranda. Measurements (2 in parentheses), Length, 4.5- 5,1 (4.5-5,6); pronotum length x width, 1.3- 15x 1.7-1,8 (1.3- LS x 1.8-.1.9); tegmen length, 4.2-5.1 (4.6-5.2). MATERIAL EXAMINED QUEENSLAND. ANIC: Kuranda State Forest, 3km north of Kuranda, 360m, flight intercept trap, rain- forest, 29 2, 25, vi.~ 3.viii, 1982, S. and J. Peck; Black Mt, Road, 30km north of Kuranda, leafmould, rain- forest, 19, 4,x1.1969,).G. Brooks; Moses Creek, 4km north by east of MU Finnigan, 12, 14-16.x.1960. T. Weir. The following were collected by Norris and Common: 3 miles north of Kuranda, 3¢d, 12, 24.1v.1955; 1 mile east of Kuranda, 1c (terminalia slide 186), 20 ¢,22 2, 4.v.1955. The following were collected by |.F.B, Common and M.S. Upton: 1 mile east of Kuranda, 12, 22.ii1.1964, Id, 429, 19.vi,1964; | mile north of Kuranda, 1200ft, 32 9, 23. iv, 1969, REMARKS Anaplecta calosoma has banded tegmina which are not characteristic of most other species of the group. Amaplecta vittata Hanitsch also has banded tegmina and because of this similarity, Bruijning (1947, p.216) suggested that vittata may be a synonym of calosoma. Although vitlata superficially tesembles calosoma, their male genitalia are distinctly different. Although Ihave not seen the type of calosoma, the specimens from both Papua New Guinea and Australia are similar and agree well with Shelford's descrip- lion, Many specimens of calosoma in the series from the vicinity of Mt Tozer were collected at mercury vapour light at night. A galerucine chrysomelid beetle Monolepta sp. (nr australis) was collected with the cockroaches. The colour and pattern of cockroach and beetle are so remarkably similar that a mimicry association is indicated (Rentz, personal communication). Anaplecta vittata Hanitsch (Figs 31B-D) Anaplecta vittata Hanitsch, 1923, p.396, fig, | (do not 2 as indicated); 1929a, p.S: Hebard, 1929, p.8; Bruijning, 1947, p.217, fig. 2; 1948, p.51. MATERIAL EXAMINED Honoryrr: J (not? as indicaled), Singapore, Thom- , 49 som Rd, Impounding Reservoir, xil.1922, C.- Saunders; Type Orth. 256 '/2 in HDEO. PARATYPE, SINGAPORE. HDEGO: same dala as holotype. (not? as indicated; terminalia slide 223), Type Orth, 25¢ 7/2. DESCRIPTION Male. Interocular space slightly greater than distance between antennal sockets. Pronotum subparabolic, hind margin weakly produced. Tegmina and wings reach end of abdomen; hind wing venation as in Fig. 31C. Supraanal plate with medial setal specialisation, hind margin convexly rounded. Subgenital plate practically symmetrical, styles small, widely separated, in- lerstylar margin symmetrically convex (Fig. 31D). Genitalia as in Fig. 31D; phallomere L1 wilh a Jong, curved filament. Colouration. Head orange. Pronotal disk oran- gish, surrounding region hyaline, Tegmina black with broad transverse whitish band across about middle, marginal field pale or hyaline (Fig. 31B). Hind wing infuscated as in Fig. 31C. Abdominal sterna straw coloured. Measurements, Length, 5.0; pronotum length x width, 1,3 x 1,7-1.8; tegmen length, 4.1-4.2 REMARKS Bruijning (1947, p.216) pointed out the dis- erepancies between Hanilsch's drawing (1923, fig. 1) of the hind wing of vitrata and that of the wing of another specimen (cp. Bruijning, 1947, fig. 2b) which Hanitsch had identified as vittata, My drawing (Fig. 31C) of the hind wing of the paratype of wittata (which Hanitsch used for his drawing) agrees with Fig. 2b of Bruijning and shows that Hanitsch erred in his original draw- ing. The male genitalia of vitta(a and calosoma are clearly different but both have a long curved filament-like sclerite (L1) indicating that both taxa are closely related (cp. Figs 31A and 31D), Anaplecta vittata was originally described from Singapore but Hanitsch (1929a, p.5) also recorded it from Sumatra (Medan, Tjinta Radja, and Arnhemia), West Malaysia (Kuala Lumpur, and Batang Padang, Perak). Bruijning (1948, p.51) had specimens from Bintang (Wes| Malaysia), Rhio Archipelago (Indonesia), and Medan, Anaplecta australiensis fi.sp. (Figs 32A-F, 34) MATERIAL EXAMINED 650 MEMOIRS OF THE QUEENSLAND MUSEUM FiG. 32. Anaplecta australiensis n.sp. A,B, ¢ holotype, head and pronotum; C, 2 paratype from 11km ENE of Mt Tozer, left tegmen (veins not shown); D-F, ¢ paratype from same locality as holotype, left tegmen (veins not shown), hind wing, and subgenital plate and genitalia (dorsal). Scales (mm): A,B, 0.5; C-E, 2.0; F, 0.25. INDO-AUSTRALIAN COCKROACHES HoLotyre; 3. %km ENE of Mi Tozer, QUEENSLAND, 12.438 [43.17E, Malaise trap/ethanol, 5-10,vii, 1986, J.C, Cardale; in ANIC (Reg. No. 8251). PARATYPES! OULENSLAND. ANIC: same locality and data as holotype, | 2,29 2, (Malaise trap). DH. Col- less, Ld (lerminalia slide 189), 39 2, T. Weir and A. Calder; 1 }km ENE of MI Tozer, 12.438 143.18E, 1° (Malaise trap/ethanol), ¢ (abdomen missing: ex pantrap), 11-16.vn,1986, LC. Cardale; 11km ENE of Mt Tozer near Iron Range National Park, 12.438 143.18E, rainforest margin, 12, 11 vii.1986, D.C.F. Rentz; Cape Tribulation area. 16,03-16.058 145.28E, littoral rainforest, 1d. 12, 21-28.ii11- 1984, A. Calder and T. Weir. NorRTHERN TERRITORY. ANIC: Holmes Jungle (nr Darwin), 12.2385 130.57E, rainforest. 1d, 6.xi.1972, R.W. Taylor. DESCRIPTION Male. Interocularspace about same as distance between anicnnal sockets (Fig. 32A). Pronotum subelliptical (Fig. 32B). Tegmina and wings reaching slightly beyond end of abdomen, Hind wing as in Fig, 32K. Anteroventral margin of front femur with 3 large proximal spines, fol- lowed by row of piliform spinules terminating in 2 large spines (Type Bz); pulvilli absent, arolia small. Supraanal plate with medial setal specialisation, hind margin convexly rounded. Subgenital plate weakly asymmetrical, styles small, interstylar margin convexly skewed ta left (Fig. 32F). Genitalia as in Fig. 32F; L1 without long, curved filament. Colouration. Head brown, clypeal region brownish yellow (Fig. 32A). Pronotal disk brown, the colour extending to the anterior and posterior borders, lateral regions whitish or hyaline (Fig. 32B), Tegmina dark reddish brown, marginal field hyaline (Fig, 32C), costal veins whittsh, sometimes with pale macula behind anal vein (Fig. 32D). Hind wing infuscated, darkest regions in costal vein area and anterior part of the uppendicular field (Fig. 32E). Abdominal terga light brown with some darker infuscation, tergal gland region of supraanal plate pale, area behind it dark brown, Abdominal sterna brown, legs and ceres (dorsally) lighter, straw coloured. Female. \nterocular space Jess than distance between antennal sockets. Supraanal plate trigonal, apex rounded. Abdominal terga dark brown, anterior segments lighter. Abdominal sterna brown, subgenital plate with valves and lateral regions darker, Measurements (9 in parentheses). Length, 4.0- 5.3.(4.1-5.5); pronotum length x width, 1.2- G51 1.3.x 1.4-1.8 (1.2- L.4% 1.5.1.9); tegmen length, 3.7-4.5 (35-45) REMARKS The genital phallomere L1 of australiensis lacks a curved filamentous sclerite suggesting that it does. not belong to the same species group as calosoma and vittata. Its close relatives may be determined by a study of the male genitalia of other species in the genus. Anaplecta brachyptera p.sp. (Figs 33A-C,34) MATERIAL EXAMINED HoLotyre: ¢&, Hinchinbrook Is., NEQ, Gayundah Ck, 10m, pitfalls. 8-18.%i.1984, Monteith, Thompson and Cook; in QMBA (Reg. No. Ti 1190). PARATYPES. QUEENSLAND, OMBA; same logglity as holotype, 18.228 146.13E, rainforest, sieved litter, 10m, 52.5, 12,10.xi,1984, 1 (terminalia slide 69), 12. 941.1984, 1a. 11.01.1984, Thompson and Gal- lon. DESCRIPTION Male (habitus in Fig. 33A). Eyes somewhat reduced, interocular space distinctly greater than distance between antennal sockets (Fig. 33B). Clypeus swollen (Fig. 33B). Pronotum sub- parabolic, Tegmina reduced in length reaching only to about T3, apically rounded. Hind wings vestigial, lateral, reaching only to T1, Anteroventral margin of front femur with 1 or 2 proximal spines followed by row of piliform spinules with 2 large terminal spines (Type Bz); pulvilli absent, arolia small, tarsal claws sym- metrical, simple. Supraanal plate (T10) with large setal specialisation located on mound aris- ing from depressed region of transverse segment, hind margin weakly convex (Fig. 33A). Subgeni- tal plate weakly asymmetrical, styles small, similar, interstylar margin weakly uneven, es- sentially not produced (Fig. 33C). Genitalia as in Fig. 33C; genital phallomere L1 with long, curved filament, Colouration. Clypeus, labrum and mandibles yellowish, rest of head (Fig. 33B), pronotum, tegmina, and abdominal (erga brown; glandular area on TIO, cerci dorsally, abdominal sterna, and legs, lighter brown. Costal vein of tegmina and lateral edge of pronotum pale. Female, Supraanal plate weakly trigonal, apex rounded. Subgenital plate valvular, Abdomen somewhat darker than male, Measuremenis (9 in parentheses), Length, MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 33, Anaplecta brachyplera n.sp. A,B, ¢ holotype, habitus and head; C, d paratype from same locality as holotype, subgenital plate and genitalia (dorsal). Scales (mm): A, 1.0; B, 0.5; C, 0.25. 3.2- 3.7 (3.6-3.7); pronotum length x width, 1,0- 1.1 x 1.3-1.5 (1.1- 1.2 x 1.5); tegmen length, 1.3-1.4 (1.3-1.5). REMARKS Anaplecta brachyptera apparently is the only member of the genus with reduced tegmina and vestigial wings. All other characters are typical of the genus. The male’s right genital phallomere resembles that of A, australiensts, whereas the left one (L1) has a long curved filament reminis- cent of those found in calosoma and vittata. Parectoneura new genus TyPe SPECIES Parectoneura bivitiata n.sp. DIAGNosis Sexes similar. Tegmina and wings fully developed. Tegmina with distinct venation, branches of median and cubitus veins lon- gitudinal, Hind wing when at rest folded along plical area, with apical triangle rolled up and curled in tight spiral (Fig. 35A); discoidal vein straight, unbranched, median and cubitus veins curved, former simple, latter with a complete branch, incomplete rami absent, apical triangle large (Fig, 35B). Anteroventral margin of front femur with some large proximal spines followed by row of piliform spinules, terminating in 2 large spines (Type B2); tarsal claws symmetrical, minutely toothed on ventral margins; ventral margins of mid and hind femurs with some large spines. d: Seventh abdominal tergum weakly INDO. AUSTRALIAN COCKROACHES Prine af Wang ASPEN r arty An cal es a \2 3 Pic Hee ~ Asn, Hint Munweve| _Shauurte aly ote farmer aa anaplacte ge a a er? — hos “at @ aclovoma ' ate, tow rad % \ D brachyptera i vpr aru Clavetraitenaie \ 7, 14a @ éatozona (usntan+) wi her By eet | 4, pee weer PENNA - Pe ee + ~~ \ o } ries ( a FE ie a) i er et, a Aibertnviltey fer > en a “the f " Fenner Hite Ve ee a owt cghinn tad = a a a ere enon ray Aap Gat By Le a 6 ar na Jitan Gay” Map Ld mis & > faric® £ Girowrains an reead ri \ Popvlrny Gyeer Bay Fo That Fay fo * ~— ae Repair Bay oe | ° = == ¥ betshed >See te be cr Hira | _ ra s a ~ it 7 ona jie isin aoa ce fmm at Me Steels Se La ie a bere FiG. 34, Known distribution of Australian Anaplecta species in Queensland (Northern Territary record of A. australiensis not shown), specialised (Fig. 35C). Subgenital plate subsym- metrical with a pair of minute, similar styles (Fig. 35D). Genital hook on left side, right phallomere reduced (Figs 35D,E). REMARKS Hchard (1943, p.12) placed Ectoneura in the Ectobiinae and the Australian species which pre- viously had been referred to Ectobius in a group Ectoneurae. ‘They agree with Ectobius in having the tegminal discoidal sectors oblique, the tarsal claws simple but asymmetrical, and the female subgenital plate simple. The general appearance of some of the species also suggests certain species of Ectobius. They differ widely from that genus and its allics, however, in having the ventral margins of the median and caudal femora unarmed, the ventrocephalic margins of the cephalic femora practically unarmed [there are fine hairs which are more slender than piliform spinules] with one or two large terminal spincs, and the intercalated triangle of the wings (in all individuals having these organs fully developed). cutled when at rest, instead of folded, with anal vein there diverging very strongly from the median fold’. as3 The new genus. Parectoneura superficially resembles species of Ectoneura, especially those taxa that have a pair of pronotal bands which continue along the full length of the tegmina, e.g. Ectoneura minima (Tepper), E. verticalis Hebard, and F. suffissa Hebard. ft agrees with Ectoneura in having the apical (= intercalated) triangle of the hind wing curled when at rest, and a gland on the male’s seventh abdominal tergum. It differs in having Type Bz front femur, sym- metrical, minutely toothed tarsal claws (vs. asymmetrical and untoothed) and some large spines on the ventral margins of the mid and hind femurs, which in Fctoneura are essentially un- armed. The Parectoneura male his a subsym- metrical subgenital plate with a pair of minute styles, whereas in Ectoneura the subgenital plate is distinctly asymmetrical and there is only one minute style. Hind wings in which the apical triangle is curled into a tight spiral when at rest are found in some other genera, e.g., Oulopteryx Hebard and Dasyblatta Hebard, both from South America (Hebard, 1921b; Rehn, 1951), Areolaria Brunner from the Malay Archipelago (Hebard, 1929, p.102), and Pseudectoneura Princis (New Caledonia), and Theganopteryx Brunner (Africa). Parectoneura bivittata n.sp. (Figs 35A-F) MATERIAL EXAMINED Houotype, 3, Tindal, Northern Territory, Australia, 14.318 132,22E, 1-20.x3.1967, light trap, W.J.M. Vestjens; in ANIC, PARATYPES: NORTHERN TERRITORY. ANIC: Tindal, § miles ESE of Katherine, 14.318 132.226, at light, 29 9, 30.xi.1967, 2d d, 6.xii.1967, 1d (terminalia slide 192), 7.xii.1967, 12, 8.xti.1967, 1 2, 9.x1i.1967, 12, 13.xi1.1967, 20 od, 14.xi3.1967, 12, 16.xii.1967, 12, 18.xi/.1967, 12,1 (ahdomen missing); 19.x1i1, 1967, 146 (terminalia slide 191), 20.x11.1967, W.J.M. Vestjens; Goose Lagoon, 11km SW by § of Borroloola, 1%, [8.vi1976, Key, Balderson, et al, [Key's field trip noles; trip 191, Stop 3614.8]; Daly River Miss., at light. 39 2, 8...1974, LF. Hutchinson, WESTERN AUSTRALIA. ANIC: 10km NW by N of mining Camp. Mitchell Plateau, Kimberley Distr., 1d, l1.v.1983, D.C.F. Rentz & J, Balderson, Stop 15. DESCRIPTION Male. Head hidden under pronotum. Inter- ocular space less than distance between ocellar spots and antennal sockets. Pronotum suboval (Fig. 35A). Tegmins and wings extending well f54 beyond end of abdomen. Tegmina with median and cubitus veins with longitudinal branches (Fig. 35A). Hind wing folded along plical area, intercalated triangle large, rolled up, tubuliform when at rest (Fig. 35A); median and cubitus veins curved, the latter with | complete and 0 incomplete branches (Fig. 356). Front femur Type B2 (with 3 large proximal spines); pulvilli appear to be present on the fourth tarsomeres only, tarsal claws simple, symmetrical, arolia well developed. Seventh abdominal tergum with distinct medial depression containing, some groups of short setac (Fig. 35C). Supraanal plate with hind margin convexly rounded, entire, not reaching hind margin of subgenital plate. Sub- genital plate almost symmetrical with 2 minute, similar, widely spaced styles on rounded apex (Fig. 35D). Genitalia as in Figs 35D-F; genital hook on the left side, its apical portion expanded, with preapical incision, basal arm uniformly slender, median phallomere relatively stout, dis~ tal region tapering, apex acute or subacute, tight phalloamere reduced to small cleft selerite bear- ing small proup of setae. Colouration, Head brown, Pronotum with pair of dark brown longitudinal bands, these continu- ing length of tegmina (Fig. 35A). Abdominal erga and sterna pale with lateral infuscation, Legs and cerci pale. Intensity of brown on head, pronotum, und tegmina varies and some specimens approach blackish brown. Female. Supraanal plate trigonal, apex of hind margin medially indented, reaching upturned margin of subgenital plate. Measurements (9 in parentheses). Length, §.2- 6.0 (4.9-6.2): pronotum length x width, 1.5 x L.9-2.0(1.5-1.7 x 1.9-2,3); tegmen length, 4.9- 5.6 (4.6-6.0). Pseudectoneura Princis Psendectoneura Prineis, 1974, p.517, TYPE SPECIES Pseudeetancura kalterbachi Prineis, by monotypy. DIAGNaSIS (based on 3) Tegmina und wings fully developed, former with unbranched discoidal vein, median and cubitus veins and their branches longitudinal. Hind wing with discoidal vein branched distally, median and cubitus vein practically straight, former simple, latter with complete and no in- complete branches, apical triangle developed, rolled spirally when al rest (Fig. 36D). Front MEMOIRS OF THE QUEENSLAND MUSEUM femur Type B3; pulvilli present on 4 proximal tarsomeres of all tarsi, tarsal claws symmetrical with subabsolete serrations on ventral margins, arolia present; ventral margins of mid and hind femurs with large spines. Seventh abdominal tergum strongly modified with deep fossa and large dense tuft of setae (Fig. 36). Subgenital plate strongly asymmetrical with single style on left side, right style absent (Fig. 36G). Hooklike genital phallomere on left side (Fig. 36G), REMARKS Pseudectoneura differs trom Parectoneura as follows: hind wing with discoidal vein branched, median and cubitus veins practically straight, gland on T7 much more camplex, subgenital plate strongly asymmetrical, and right style ab- sent. Pseudectoneura kaltenbachi Princis (Figs 36A-G) Pseudecioneura kaltenbachi Princis, 1974, p.514, fig. 4(d). MATERIAL EXAMINED Hovoryre: 6, New, Caledonia, Riv, Tiari, Niaouli- Wald, 17.xi.1965, Osterreichische Neukaledonien- Expedition, 1965 (PNK 80/2); in NMWA. PaRatTyre: NEW CALEDONIA. NMWA, same data as holotype, 1d (terminalia slide 55), DESCRINTION Male. Interocular space distinctly less than dis- tance between occllar spots (Fig. 36A); palpal segments 3 and 5 each longer than fourth, Pronotum subelliptical, widest point behind the middle (Fig. 368). Tegmina and wings fully developed, extending beyond end of abdomen. Teymina with median and cubitus veins and their branches longitudinal. Hind wing with costal veins weakly thickened, discaidal vein branched beyond middle; median and cubitus veins prac- tically straight, former simple, latter with 1 com- plete and ( incomplete branches, apical triangle developed, spirally rolled when al rest (Fig. 36D). Front femur Type B3 {with 4 large proximal spines, followed by a short row of only 5 piliform spinules); large spines on mid and hind femurs; pulvilli present on 4 proximal tar- someres of all tarsi (Princis incorrectly claimed that the hind tarsus has a pulvillus only on the fourth segment); tarsal claws symmetrical their ventral margins with subobsolete serrations, arolia present, First abdominal tergum un- INDO-AUSTRALIAN COCKROACHES 655 FiG. 35. Parectoneura bivittata n.sp., paratypes from Northern Territory. A, 2 from Daly River Miss., habitus (right tegmen removed to show the underlying folded hind wing; a = rolled up apical triangle of left wing); B, 2 from Goose Lagoon, hind wing; C-F, d from Tindal, 8 miles ESE of Katherine: C, Tergal gland area on T7; D, Subgenital plate and genitalia (dorsal; b = left phallomere; c = median phallomere; d = right phallomere); E, right genital phallomere; F, left hooklike phallomere. Scales (mm): A, 2.0; B, 1.0; C,D, 0.25; EF, 0.1. 636 specialised. Seventh abdominal! tergum with a large gland anteromedially consisting of a deep fossa and a dense group of setae (Fig, 36E). Supraanal plate with hind margin concavely ex- cavated, not reaching hind margin of subgenital plate; right and left paraprocts large dissimilar plates (Figs 36F). Subgenital plate convex, strongly asymmetrical, the left side excavated in which is located a cylindrical left style, right style absent; right half of subgenital plate produced as large lobe whose rounded hind mar- gin is divided and forms a spinclike process which Princis claimed is the right style. However this structure is not articulated and apparently is a modification of the hind margin but not the right style, which is absent (Figs 36C,E,G), Genitalia as in Fig. 36G; hooklike phallomere on left side; apex of median phallomere with a brushlike group of dark setae, right phallomere reduced to cleftlike remnant. Colouration. Head with vertex dark reddish brown, occiput lighter, interocellar area whitish, face yellowish brown with whitish areas lateral- ly, labrum brownish (Fig. 36A), Pronotum with disk yellowish brown and brown parentheses- like markings on anterior half, lateral borders hyaline (Fig. 36B). Tegmina pale, hyaline, humeral vein may be darkened. Hind wing with yellowish tinge along anterior border and behind apical triangle, grayish on posterior region of apical triangle and along edges of posterior field (Fig. 36D). Proximal abdominal terga light yel- Jowish brown with some infuscation, distal seg- ments dark brown, glandular region of T7 pale ycllowish, glandular setac dark, supraanal plate pale, laterally dark. Abdominal sterna light brown, laterally dark brown, Female, Unknown, Measurements: Length, £2.3-|2.5; pronotum length x width, 2.6 x 3.7-3.8; tegmen length, biT-1 18. Pseudectobia Saussure Pyeudectohia Saussure, 1869, p,234: Princas, 1951, p.65. Ty?e SPECIES Pseudeciobia luneli (Saussure) = Blatia luneli Saussure, by selection, Kirby, 1904, p.65. DiAGnosis (based on dD and modified from Prncis, 1951), Tegmina and wings fully developed. Tegmen MEMOIRS OF THE QUEENSLAND) MUSEUM with discoidal sectors longitudinal. Hind wing with cubitus vein straight and with complete and no incomplete branches, apical triangle poorly developed and not rolled spirally. Front femur Type Bo; ventral margins of mid and hind femurs sparsely armed. Seventh abdominal tergum with deep fossa that occupies most of the segment. Subgenital plate symmetrical with pair of similar styles. Hooklike genital phallomere on right side (Fig. 37E). REMARKS This genus differs strongly from Parectaneura and Pseudectoneura in hind wing structure and in having the male hooklike genital phallomere on the right rather than the left side, Pseudectobia luneli (Saussure) (Figs 374-E) Blatta luneli Saussure, 1868, p.355 (4); Walker, 1871, p.20, Ectobia (Pyeudectobia) luneli (Saussure), 1869, 235, Ectobia luneli (Saussure): Walker, 1871, p.15. Phyllodromia luneli (Saussure): Shelford, 1908, p.12. Pseudectobia lunvli (Saussure): Kirby, 1904, p.65; Princis, 1951, p.65,, pl.6, fig, 65 (0); 1969, p,996. MATERIAL EXAMINED Hotorvee; 3 (terminalia slide no, 1), Neelgheries, Indes [= Nilpiri, tndia], coll. Guérin; in MNHG. [The specimen is in very poor condilion; it is greasy, the tegmina and wings are badly damaged as are most of the legs - all tarsal claws are missing; Shelford (1911:156) examined the type and described it as ‘very shattered’, It obviously hasn't improved with age], DESCRIPTION Male. Eyes very close together, almost con- tiguous (Fig. 37B). Pronotum suboval (Fig. 37A). According to Princis (1951, p.65) the teg- mina and wings are fully developed reaching beyond end of abdomen. Median vein of tegmen separated from discoidal vein and supplied with 1 branch, discoidal sectors longitudinal; costal veins of hind wing very weakly thickened distad. The remains of the wing shows that the cubitus is straight and has at least 3 complete and 0 incomplete branches; the apical region is missing but Shelford (1911, p.156) stated that there is 'a small and ill-defined apical triangle’. According to Saussure (1869), the reflexed field of the hind wing is crumpled in repose (therefore not jubuliform as in Pseudectoneura), Front femur INDO-AUSTRALIAN COCKROACHES 657 FiG. 36. Pseudectoneura kaltenbachi Princis, d d. A,B,D, holotype; C,E-G, paratype: A, Head; B, Pronotum; C, Subgenital plate (ventral); D, Hind wing; E, Abdominal terga 7 to supraanal and subgenital plates (dorsal); F, Supraanal plate and paraprocts (ventral); G, Subgenital plate and genitalia (dorsal). Scales (mm): A-C, 1.0: D, 3.0; E, 1.0; F,G, 0.5. Type Bz (4 large proximal spines two of which are broken off, us is one of the terminal spines); tarsi damaged, none with claws, Seventh ab- dominal tergum (dried pinned specimen) with huge deep elliptical pit that occupies practically entire segment, posterior wall with minute setae (Fig. 37D); in KOH cleared specimen pit is seen io be pair of large depressions separated by lon- gitudinal wall. Supraanal plate transverse, hind margin practically truncate, not reaching hind margin of subgenital plate (Fig. 37D), right and left paraprocts similar plates. Subgenital plate symmetrical, lateral margins of distal region in- dented, bearing pair of similar cylindrical styles, interstylar margin produced and in pinned specimen appears to be a single projection (Fig. 37C), but when cleared in KOH consists of 2 small lobes (Fig. 37E). Genitalia as in Fig. 37E: hook on right side, apex truncate, without sub- apical incision or spine; median phallomere rod- like, tapering, apically acute; a pair of curved sclerites between median and right phallomeres, Colouration. The colour is difficult to deter- mine because the specimen is greasy. Head with occiput black, face yellowish with black mark- ings (Fig. 37B). Pronotum with broad, triangular black macula, lateral borders pale (Fig. 37A). Abdomen mostly ight brown, Female, Unknown Measurements, Length, 7.6; pronolum length x width, 2.0 x 2.9; tegmen length, ‘7. REMARKS Princis (1969, p.996) listed 2 species of Pseudectobia, namely the type species luneli, and insularts (Saussure) Which f find belongs in Chorisoblaita, and describe below. Chorisoblatta insularis (Saussure) n.comb. (Figs 38A4-G) Ectebta (Pseudectobia) insularis Saussure, 1869, p.236 (d). Ectobia iisularis Saussure: Walker, 1871, p15, Theganopteryx wisularis (Saussure and Zehntner), 1895, p.14. Lupparia insulariy (Saussure); Shelford, 1908, p.16. Pseudectobia insularis (Saussure): Kirby, 1904, p.65, Orian, 1957, p.517; Princis, 1969, p.996, MATERIAL EXAMINED Hocorvyer: 2 (terminalia slide no, 2), We de France [= Mauritius], Afrique, M.H, de Sausstre; in MNHG, MEMOIRS OF THE QUEENSLAND MUSEUM DESCRIPTION Male. Interocular space less than distance be- tween antennal sockets (Fig. 38C). Pronotum subclliptical (Fig. 38B). Tegmina and wings fully developed extending beyond end of ab- domen. Hind wings with costal veins weakly thickened distad, discoidal and median veins un- branched, cubitus vein straight with 3 branches that reach only to the vein demarcating the large apical (nangle (Fig. 38A), Front femur Type Bz (with 3 large proximal spines); pulvilli present on4 proximal tarsomeres, tarsal claws symmietri- cal, ventral margins bearing a few large teeth, arolia well developed (Fig. 38F). Abdominal terga unspecialised. Supraanal plate sub- trapezoidal, corners rounded, hind margin not teaching hind margin of subgenital plate (Fig. 38D). Subgenital plate extending well beyond supraanal plate, styles similar, widely separated, each located in posterolateral corners, interstylar margin convex, deflexed (Figs 38D,E); in dorsal view inlerstylar margin appears truncate because deflexed portion is not visible from above (Fig. 38D). Genitalia as in Fig: 38G: genital hook on right side, apex with minute spine; to right of hook an additional divided sclerite, its most posterior portion beating dark setae; median sclerite long, slender, apically acute. Colouration. Head with vertex and o¢cipul brown; lwo narrow, black transverse bands, one belween eyes, olher between upper part of anten- nal sackets (Pig, 38C), remainder of face largely hyaline. Pronotum yellowish, disk with brown dots and lines, outer border regions. partly hyaline (Fig. 38B). Tegmina with some yel- lowish and mostly reddish rectangular and dot- like spots between veins, Hind wing infuseated, anlérier margin at distal ends of costal veins, while; anterior field and region posterior to api- cal triangle darker than rest of posterior ficld and apical triangle (Fig. 38A); one wing that has been moumted on a card was completely bleached and was not infuscated; the wing that was still at- tached lo the specimen (and which I mounted on a card) had the infuseation deseribed by Saus- sure, Legs light brown, femur with dark brown stripe on dorsal margin, foreleg tarsomeres with narrow brown apical rings. Abdominal terga brown, darker infuscation laterally and on ter- minal segments. Abdominal sterna brown with broad, somewhat darker mediolongitudinal, Stripe. Female, Unknown, Measurements. Length, 9,8; pronotum length x Width, 2.6 x 3.8; tegmen length, 11.4, INDO-AUSTRALIAN COCKROACHES 659 FiG. 37. Pseudectobia luneli (Saussure), 3 holotype: A, Pronotum and head; B, Head; C, Subgenital plate (ventral); D, terminal abdominal segments (dorsal); E, Subgenital plate and genitalia (dorsal). Scales (mm): A,B, 1.0; C.D, 0.5; E, 0.25, 660 MEMOIRS OF THE QUEENSLAND MUSEUM FiG. 38. Chorisoblatta insularis (Saussure), d holotype of Ectobia (Pseudectobia) insularis Saussure: A, Hind wing; B, Pronotum; C, Upper portion of head; D, Terminal abdominal segments (dorsal); E, Subgenital plate (ventral); F, Tarsal claws and arolium of foreleg; G, Subgenital plate and genitalia (dorsal). Scales (mm): A, 3.0; B-D, 1.0; E, 0.5; F, 0.25; G, 0.5. INDG-AUSTRALIAN COCKROACHES REMARKS Princis (1969, pp.925-927) listed 11 species of Chorisoblatta of which 7 (one with a query) are from Africa, 2 fromm Madagascar, and 2 are from Mauritius. Of the Mauritian species one is C, liturifera (Stal), the type species of the genus, and the other is Chortsoblatta chopardi Princis (Princis, 1969, p.926). Choriseblatia insularis is the third species of the genus known from Mauritius. SUBFAMILY PLACEMENT OF PARECTONEURA, PSEUDECTONEURA, PSEUDECTOBIA AND CHORISOBLATTA According to McKittrick (1964), the Blattel- linae and Ectobiinae (based on 2 species of Ec- tohius) males have the genital hook on the left side and their females rotate the ootheca 90° prior to depositing them. In MecKittrick’s Plectop- terinae (= Pseudophyllodromiinae of Vickery and Kevan, 1983), the male's genital hook is on the right side and their females do not rotate the ootheca prior to deposition. Hawever, Bohn (1987) found that the males of some Ectobiinae have the venital hook on the right side (rather than the left), yet their females rotate the ootheca, Bohn concluded thal the position of the male's genital hook cannot always be used to place blattellids in their respeetive subfamilies and suggested that whether or not the female rotates her ootheca should be the criterion for placing a species in one or the other of these two sub- families. | have recently found that the males of Margattea nimbata Shelford have the genital hook on the right side but the female rotates its ootheca and I place this species in the Blattellinae (Roth 1989). Unfortunately, whether or not the ootheca is rotated is not known for most genora and species, including the genera under discus- sion in this paper, However, unti) oviposition behaviour is determined one can place a species in the Blattellinae or Pseudophyllodromiinae provisionally, if the position of ihe male's genital hook is known; subfamily placement can be con- firmed when the female's oviposition behaviour is determined, With this in mind | am placing Parecroneura aad Pseudectoneura in the Blattellinae and Pseudectobia and Chorisoblatta in the Pseudo- phyllodromiinae. Shelford (1911, p.l56) believed that Pseudectobia luneli ‘... is more suitably placed in the Ectobiinae’, but Princis placed jt in the Blattellidac rather than the Ee- tobiidae. | agree with Princis because the male 641 subgenital plate and styles, and genitalia of Juneli are more typical of the Blattellidae than they are of the Ectobiidae (nae). ACKNOWLEDGEMENTS I thank the museums, their curators and collec- {ion managers, and Dr Syoziro Asahina, men- tioned in the introduction, for lending me specimens, | am grateful to the Bureau of Flora and Fauna, Australian Biological Resources Study (ABRS), for partial support, and Dr D.C.F, Rentz and Mr G, Thompson for help in prepar- ing of the distribution maps. LITERATURE CITED ASAHINA,S. 1973, Taxonomic notes on Japanese Blat- taria. V, On three recently introduced blattellid species. Jap. J. San, Zool. 24: 123-8, (Japanese; English summary), 1974, Taxonomic notes on Japanese Blattaria, VI. What is “Phylledromica japonica”? Jap..J. San. Zool. 25: 161-66. (Japanese: English summary), 1977, Taxonomic notes on Japanese Blattaria, VIII. The Anaplectidae of Japan and Taiwan. Jap, Jour. San, Zool, 28; 272-80. 1985, Taxonomic notes on Japanese Blattaria, XV. A revision of three blatiellid species. Gen-Gen Sha, Miyamaru 8: 2-10, (Japanese: English sum- mary). BEY BIENKO, G. YA. 1969, New genera and species of cockroaches (Blattoptera) from tropical and subliopical Asia. Ent. Qhozr, 48: 831-62. (Rus- slam). 197), (translation of previous reference]. Ent. Rev. 84; 528-48, Bown, HW, 1987, Reversal of the right-left asymmetry in male genitalia of some Ectobiinae (Blattaria : Blattellidae) and its implications on sclerite homologization and classification, Ent, Seand, 18; 293-303, BRUNNING, C.F.A, 1947, An account of (he Blattidae (Orthoptera) from Celebes, the Moluccas, and New Guinea, Zool, Meded, R. Mus, Naw Aisi Leiden 27; 205-52. 1948. Studies on Malayan Blatlidae. Zool. Meded. R. Mus. Nat, list. Leiden 29; 1-174, 1959. Studies on the fauna of Suriname. and other Guyanus, (eds D.C, Geijskes and P.W, Wagenaar Hummetinck), No, 4, The Blattidae of Surinam. Vol. Il, pp.1-103. BRUNNER VON WATTENWYL, C. 1898. Orthopteren des Malayischen Archipels, gesammelt von Prof, Dr, W. Kiikenthal in den Jahren 1893 und 1894, ADA, senckenh, Naturf, Ges, 24: 193-288, BURMEISTER, H, 1838. Handbuch de Enlomologie, 2: 397-756, Berlin. HANITSCH, R. 1915. Malayan Blattidae. Jour, Straits Branch R. Asiat, Soc. 69 17-178. 1923. Malayan Blatlidae. Part Il. Jour. Malayan Branch R. Asiat. Soe, 1; 393-474, 192%. Spolia Mentawiensia. Blattidae. Bull. Raffles Mus, 1-44. 1929x. Dr E. Mjdberg’s zoological collections from Sumatra. Ack, Zool. 20A* 1-20, 1929b, Fauna Sumatrensis. Blatlhidae, Tijds, Ent. 72: 263- 302, 1933. The blauid fauna of Celebes, based chielly on the collections made by P. and F. Sarasin. Verh, naturf. Ges, Basel, 44: 119-50. 1935. On further blattids (Orth.) from Celebes. Stylops 4: 14-9, Heaarp, M. 1920, The Blattidae of Panama, Mem. Amer. Ent. Sac, (1919) 4; 1-148, 19214. A note on Panamanian Blultidae, with the description of a new genus und two new species (Orth.), Ent. News 32: 161- 69. 1921b, South American Blattidae from the Museum National D'Histoire Naturelle, Pans, France. Prac, Acad. Nat, Sci. Phil, 73 (Pt. 1): 193-304, 1929. Studies in Malayan Blattidae (Orthoptera). Prac. Acad. Nat. Sci, Phil, 81: 1-109. 1933. Notes on Panamanian Dermaptera and Or- thoptera. Trans. Amer. Ent. Soc, 59: 103-44, 1943, Australian Blattidae of the subfamilies Chorisoneurinae and Ectobiinae (Orthoptera). Acad. Nat. Sci. Phil., Monog. 4: 1-129, Kireny, W.P. 1904, "A synonymic catalogue of Or- thoplera, Vol, 1.” (British Museum (Natural His- tury)) London). LEPscut, B.J. 1989. A preliminary note on the food of Imblattella orchidae Asahina (Blattodea : Blat- tellidae), Aust. ent. Mug. 16: 41-42. MckKriigrick, FA. 1964. Evolutionary studies of cockroaches, Cornell Univ. Agric. Exp. St, New York State Coll. Agric., Mem, 389: 1-197. ORIAN, AJB, (957, Saltatoria, Phasmida and Dic- wwaptera of Mauritius. Ann. Mag. nat. Mist. (1210; 513-20. Princis, K. 1951, Neve und wenig bekannte Blattanen aus dem Zoologische Museum, Kopenhagen. Spolia Zool. Mus. Hauniensis 12; 5-72. 1954, Kleine Beilriige zur Kenntnis der Blattarien und ihrer Verbreitung, VI, Eat, Tidskr. (1953) 74: 203-13, 1959. Revision der Walkersehen und Kirhyschen Blatturientypen tm British Museum of Natural History, London WT Opuse, Ent. 24: 125-50. MEMOIRS OF THE QUEENSLAN]) MUSEUM 1965, Blattariae, subordo Blaberoidea. In Beier, M. ({ed.), ‘Orthopterogrum Catalogus. Pr. 7." (W. Junk: s’-Gravenhage), 1969. Blawtariae, suborda Epilamproidea, Fam. Blaltellidae. In Beier, M. (ed.), ‘Orthopterorum Calalogus, Pt. 13," (W. Junk: s'-Grayenhage). 1971, Blatlariae, suborder Epilamproidea, Fam. Ec- tobiidae. In Beier, M. (ed.), ‘Orthopterorum Catalogus. Pi. 14." (W. Junk: s’- Gravenhage). 1974. Ergebnisse der Osterreichischen Neuksledonien-Expedition 1965. Blattariae- Schaben. Annalen naturh, Mus. Wien 78: 513- 31. REuN, J,A,G. 1904, Studies in. ald world forficulids or eurwigs, and blattids or cockroaches, Prac, US. Natl Mus. 27: 539- 60, REUN, J, W.H, 1951, Classification of the Blattaria as indicated by their wings (Orthoptera). Mer. Amer. Ent. Soc. t4: 1-134, ReNTzZ, D.C.F. 1987, Jmblattella archidae Asahina,. an introduced cockroach associated with orchids in Australia (Blattodea : Blattellidae), News Bull, Aust, Ent, Sac. 23(2): 44-5. 1987b. Imblattella panamae (Heburd) a Panamanian cockroach associated with orchids in Australia. (Blattodea : Blattellidae). Bull. Or- chid Soc, Canberra 2; 3-4, 1987c. Danger in your orchid house. /miblattella archidae Asahina an introduced cockroach as- sociated with orchids in Australia (Blattodea ; Blattellidae). Aust. Orchid Rev., Spring 1987, pp.14-5, Roti, L.M. 198Sa. A taxonomic revision of the genus Blattella Caudell (Dictyoptera, Blattaria : Blat- tellidae). Ent, Scand., Supp. 22: 1-221. 1Y85b. The genus Symploce Hebard. TY. Species from Borneo (Kalimantan, Sabah, Sarawak), Sumatra and West Malaysia. (Dictyoptera: Blat- jaria, Blatlellidae). Ent. Seand., 16; 139-59. 1986. The genus Episynploce Bey-Bienko, II. Species from Sulawesi (Celebes). (Dictyoptera : Blattaria, Blattellidae). Ent. Scand. 17: 173-88. 1989. The cockroach genus Margattea Shelford, with a new species from the Krakatau Islands, and redescriptions of several species from the Indo-Pacific region. (Dictyoptera : Blaltaria : Blattellidae), Proc. Ent. Sac, Wash, 91; 206-29, SAUSSURE, H. DE 1868. Orthopterorum species novae aliquot. Rev. Mag. Zool. 20: 354-57. 1869. Mélanges orthoptérologiques. II" Fascicule, Blatlides et Phasmides. Mem. Soc, Phys, Hist. Nat, Genéve 20); 227-326. SAusSURE, H. DE AND ZFHNTNER, L. 1895, Révision de la tribu des Perisphaeriens (insectes INDO-AUSTRALIAN COCKROACHES Orthoptéres de la famille des Blattides). Rev. Suisse Zool. 3: 1-59. SHAW, E. 1916. Australian Blattidae. Part II. On the type of Ischnoptera brunneonigra, Tepper, with a description of the male insect. Vict. Nat. 33: 86-8. SHELFORD, R. 1906. XIV. Studies of the Blattidae. Trans, Ent. Soc. London 1906 (Part IT): 231-79. 1908. Orthoptera. Fam. Blattidae. Subfam. Phyl- lodromiinae. Genera Insectorum 73: 1-29. 1911. Preliminary diagnoses of some new genera of Blattidae. Ent. Monthly Mag. (2)22: 154-56. 1912. New Blattidae from New Guinea. Ent. Rundschau 29: 51-2. STRAND, E. 1934. Miscellanea nomenclatorica zoologica et palaeontologica. Folia Zool. Hydrobiol., Riga 6: 271-77. 663 TEPPER, J.G.O. 1893. The Blattariae of Australia and Polynesia. Trans. R. Soc. S. Aust. 17: 25-130. 1895. Notes on Victorian and other Blattariae and descriptions of new species. Trans. R. Soc. S. Aust. 19: 146-66. VICKERY, V.R. AND KEVAN, D.K. MCE. 1983. A monograph of the orthopteroid insects of Canada and adjacent regions. Lyman Ent. Res. Lab. Mem. 13: 680 pp. WALKER, F. 1868. ‘Catalogue of the specimens of Blattariae in the collection of the British Museum’. (British Museum (Natural History): London). 1871. Supplement to the catalogue of the Blattariae. (British Museum (Natural History): London). 664 MEMOIRS OF THE QUEENSLAND MUSEUM CORRIGENDUM RICHARD A.I. DREW, The tropical fruit flies (Diptera: Tephritidae: Dacinae) of the Australasian and Oceanian regions. Memoirs of the Queensland Museum 26: 1-521. Page 9. Line 9, paragraph 2, for ‘(9)’ insert ‘(10)’. Page 20. Line 8, paragraph 3, for ‘cross-vein’ insert ‘crossvein’. Page 31. for ‘Bactrocera) trivialis’ insert ‘Bactrocera (Bactrocera) trivialis’. Page 43. Line 7, paragraph 4, for ‘extremitiies’ insert ‘extremities’. Page 88. Under ‘Bactrocera (Bactrocera) pallilda’, ‘LOCATION OF TYPES’, should read: Lectotype ¢ (T. 5254) and paralectotype @ (T.5253) In QM; Page 184. Last line of paragraph 1 should read: (1 male, 1 female and the sex of 1 cannot be determined). Page 294. The reference Smith er al. should read: SMITH, E.S.C., CHIN, D., ALLWoOD, A.J. AND COLLINS, S., 1988. A revised host list of fruit flies (Diptera: Tephritidae) from the Northern Territory of Australia. Qd J. agric. amin. Sci. 45: 19-28. Page 295. Fic. 1. Caption, for ‘A1’ insert ‘Ay’. Page 521. Line 14, for ‘trivalis’ insert ‘trivialis’. TYPE. FIGURED AND MENTIONED FOSSIL INVERTEBRATES IN THE QUEENSLAND MUSEUM A.C. ROZEFELDS, E,D, MCKENZIE AND C, MOBBS RozefeldsA.C., McKenzie, E.D. and Mobbs, C. 1990 08 31, Type, figured and mentioned fossil inverlebrates in the Queensland Museum. Memairs of the Queensland Museum 242): 665-713, Brisbane. ISSN 0079-8835, A taxonomically arranged list of type, figured and mentioned fossil invertebrates held by the Queensland Museum is presented. Information for each specimen includes reference list, locality of collection, formation and age, Information on early collectors including de Vis, Hain, Daintree and Aplin is also included.[l7ype list, figured, mentioned, fossil, invertebrate, Queensland Museum, Hann, Daintree, de Vis, Aplin. A.C, Rozefelds, E.D, McKenzie and C. Mobbs, Queensland Museum, PQ Box 300, South Brishane, Queensland 4/01, Australia; 11 May, 1989. This list is the third in a series thal details the Queensland Muscum's holdings of type, figured and mentioned fossils; lwo previous lists recorded the plants (Rozefelds, 1986) and fish, reptiles and amphibians (Lees, 1986), The for- mat and coverage of our list follows the two earlicr lists in that synonymies and references to specimens are listed exactly as cited by authors. The present list differs from the earlier catalogues in that specimens are listed alphabeti- cally by their most recent generic allocations, For ease of reference, there is an index at the end of this paper that records all synonyms used for the specimens, For each specimen, the Queensland Muscum registration number (prefixed by ‘F’) and references are recorded, and, where known, formation and age, The list, in general, reflects the most recent published opinions as to status of the material in ihe Museum's collection. The dating of formations follows Day et al, (1983). Some material mentioned by carlicr researchers has not been located (e.g. Etheridge fil., 1892, 1909; Whitehouse, 1924). These specimens were neither figured nor referred to by registration numbers and therefore cannot be identified con- fidently. Some specimens were never in the col- lection. For example, Whitehouse (1926, p,214) erroneously attributed the neotype of Tropaeum australe to the Queensland Museum collection. It is in the collection of the Geological Survey of Queensland (Day, 1974, p.5). The carlicst fossil type material deposited in the muscum was studied by Etheridge (1872). This included the Richard Daintree. Collection and material obtained from the Gympie gold field by Christopher D'Oyly Hale Aplin. Daintree was appointed Government geologist for Northern District of Queensland on the 14 April, 1868 and Aplin was appointed aboui the same time for Southern District (1. Sanker, pers. comm., 1989). The history of the Daintree Collection 1s che- quered, Etheridge (1872, p.326) recorded that ‘the ship (‘Queen of the Thames’) which con- veyed both Mr Daintree and his large collection of minerals and fossils from Melbourne, was wrecked off {he east coast of Africa, near (he village of Bredarsdorp; and, as may be supposed, neither minerals nor fossils were much improved by their /ong submergence prior to recovery. The labour and anxicty of many years’ research ta establish the age and nature of the stratified rocks of Queensland was then nearly lost; and but for the fortunate position of the ship near the shore, fresh collections must have been made to elucidate the physical character and structure of the Colony.” Etheridge fil. (1892) refigured and discussed a large number of Daintree’s specimens studied by Etheridge (1872), Subsequently, Etheridge fil. and Dun (1906, p.3) thought that the Daintree Collection was lost, This is not the case, although the whercabouts of some material described by Etheridge (1872) remains unresolved. These specimens may have been incorporated into the Qucensland Muscum collection. This is unlikely because the bulk of Etheridge’s (1872) material was registered in 1917 and there is no mention of the missing specimens, Some of the material described by Etheridge (1872) was obtained by W.B. Clarke. It is possible that Clarke may have retained them and, subsequently, they were destroyed during the 1882 Garden Palace fire, Sydney (Etheridge fil, and Dun, 1906, p.3). There is no record of these specimens in either the Queensland Geological Survey (Susan ona Parfrey, pers. cormm., 1985). British Museum Natural History (R. Cocks, pers. comm. 1988), University of Queensland (Andrew Simpson, pers. comm., L989) or the Australian Museum (Fletcher, 1971). Aplin was involved in assessing the gold prospects in southem Queensland, including the Gympie area, The resulls of this field work ap- peared in six reports of the ‘Legislative Council Journal’ (Woods, 1964). The fossil collections made by Aplin from Gympie ate particularly significant because many of the sites on the min- ing field are no longer accessible. As well, the Specimens obtained were part of the initial col- lections of the Queensland Museum (Turner and Wade, 1986). Aplin, in June-September, 1871, produced the first catalogue of fossils in the Museum, At that time, he was responsible for the Gavernment’s mineralogical collections in the Parliamentary Binlding, Queen Street, Brisbane (Mather and Belcher, 1986; Turner and Wade, 1986). Responsibility for the collections passed trom the Philasophical Society to the government in that year, Aplin's catalogue included material obtained by Huckett, Daintree. Aplin and Gregory (Woods, 1964), This handwritten catalogue of 203 entries survives in the Queenslind Museum, Charles Walter de Vis became curator of the Museum im February, 1882 (Mather, 1984), He had collected extensively tram the Rockhampton District. De Vis did not record separate localities for his material but in an extract from a letter quoted by Etheridge fil, (1892, p,199) dated 25th July, 1888 de Vis says: ‘The fossils are trom the Agricultural Reserve; from the Fitzroy at Laurel Bank, about ten miles from Rockhampton, westward ta the Nine-mile Lagoon, thence to the Corporation Quarry, Athelstane Range, and to the northern outerop (al foot of Bersekers [sre}) of the synclinal bencath the township and hed of river’. Some of the de Vis collection is also from the Clermont District. Important collections were also obtained during the Hann Expedition, 1872, which ex- plored areas of Cape York Peninsula, Hann and Taylor ({he geologist of the expedition) collected specimens from 2 sites’ the first and major site wason the Walsh River, probably near ‘Boomers Hole’ (Fig.1), Hann (1873, p.1034) reports that: ‘Having completed and satistied myself of the nature of the country up the Walsh, [resolved to move on, which | didon the 15th July, by follow- MEMOIRS OF THE G“UEENSLAND MUSEUM ing down the tight bank of the river for three miles, through a poor country; here | came upon a limestone formation, which I found to be sinuilar in character to that on the Flinders and the Barcoo, In passing over the ridges.] noticed some fossils, which at once induced me (o draw up and prosecute a further search, The camp was fixed on the steep bank of the river, which here presented banks sixty feet in depth, composed, near the surface, of a light calcareous soil a few feet in depth, resting on a decp bed of shale, in which limestone boulders of all sizes were im- bedded and suspended, and in which the fossils were mostly found. A more interesting spot for a scientific man can scarcely be conceived; here he ts surrounded by the objects of his interest, they are under his feet like pebbles on the seashore, they are hanging above his head ready to crush him if not careful, he cannot move without sceing them around him onall sides; they were of all sizes, and numbers of them beautifully perfect; what, and how many to Save Was the puzzle, each new find exceeded the last one in beauty, until all the beautitul ones. were sufficient to load a dray, could we have saved them, and, as | had noteven one packhorse lo carry these and (he rack specimens, [was put tomy wits’ end how many to transport. However, Mr Taylor and myself collected the best of the various species, which we were content to secure and carry along with us. | found two or three bones of the vertebrae of a large animal, which were attached to each other by limestone.” The Hann Expedition matenal was registered into the Museum collections in 1882, [ was allo- cated the Donor registration number D212. Remarks in that register refer to it as part of the “Old Collection’. Presumably most of the material ob- tained was collected from the Walsh River, al- though Hann in his diary also recorded fossils on the Mitchell River. He says (p.1048) that ‘two or three varioties or forms were procured - not seen at the fossil camp’ (i.c. Walsh River locality), The specimens collected from the Mitchell River have not been identitied in the collection. Etheridge fil. (1892, 1894) described many of the taxa collected by de Vis from the Rock- hampton District and went on to research the Cretaceous ammonites of Queensland (Etheridge fil., 1909), Significantly, a large num- ber of ammonites were obtained by the Hann Expedition from the Walsh River areg, Whilchouse (Honorary Palacontologist, Queensland Museum 1927-1937) revised the Cretaceous ammonites in a series of papers, 667 INVERTEBRATE FOSSIL TYPES “IDALY YSTEAA UO ayts Surjdayjoo sofew soqeoIpur MOLY “(ZL8T ‘AleIq §,uueH WO 49RI]x9) pur[susaNGO Usajsee YOU ‘kar JOATY YS]eM/|]9YONIA oy) Wl UoITpedxg S,uuey jo 9jno1 Burmoys dep “[ “Dig al LON NS pases HPEy ssvub giyg ure] v907 ~umegr iy yemrsbuzege fagunog Buspeznpug poserué 2724 Aepunod wusdg meg (pees Ruagunma Pret) UGLZ © \ “uzume2 poo \ \<= FapYUEL TAILS Jeez ! hi ht, eng / : here POOF OO" _ r Fi Pp2ss RL ae ay, ft perne? KOR mews FOL we ! N prow yy xg Nn PEA POE EMO Fee “PONY SHINE YM surely uaa twang PS PEON PCY GF ey SnIB Ty UASg BOLL RA UPR: PPT he 668 Woods (1953, 1957) described the macrurous decapods and Brachyura from the Queensland Cretaccous. Evans (1961,1971) described Trias- sic Hemiptera from the Mount Crosby Formation in South Eastern Queensland. Recent foci of research in the museum have been the Cambrian trilobite faunas of North Western Queensland (Jell, 1970, 1975a, 1977); Ordovician nautiloids (Wade, 1977a,b); and Devonian crinoids Jell et. al, (1988). Abbreviations used in the text are the follow- ing: Fm, Formation; GSQ, Geological Survey of Queensland; GSWA, Geological Survey of Western Australia; QML, Queensland Museum Locality; UQL, University of Queensland, Geol- ogy Department Locality. SYSTEMATIC LIST Aconeceras walshense (Etheridge fil., 1892) F1272 HOLOTYPE Ammonites walshensis Etheridge fil., 1892, p.493, pl. 42, figs 10,11. Aconeceras walshense (Etheridge fil.) Whitehouse, 1926, pp.203- 4, Aconeceras walshense (Tenison-Woods); Whitehouse, 1927, p.114, pl.16, figs 24,b, text fig.6, Aconeceras walshense (Etheridge, 1892); Hill et al., 1968, pl. K8, fig.2. LOCALITY: Walsh River, NQ. FORMATION: Blackdown Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Collected Hann’s Expedition, 1872. Whitchouse (1927) incorrectly indicated Tenison-Woods as the author of this species. It was described by Etheridge fil. (1892). F1594 Aconeceras walshense (Etheridge fil.,) Whitehouse, 1926, pp.203- 4, pl.34, figs La,b, pl.37, fig.3. Aconeceras walshense (Tenison-Woods); Whitehouse, 1927, p.114. LOCALITY, FORMATION, AGE; As for F1272. REMARKS: Sce F1272. F1704 Aconcceray walyhense (Etheridge fil..) Whilehouse, 1926, pp.203-4. Aceneceras walshense (Tenison-Woods); Whitehouse, 1927, p.114, pl.16, fig.3, text figs 1, 7. LOCALITY: Walsh River, NQ. FORMATION: Blackdown Fm, AGE: Lower Cretaceous, (Aptian). MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS: See F1272. Whitehouse records the locality for this specimen as Walsh River al- though the Queensland Museum Palaeontology Tegister gives the locality as Western Queensland. The lithology is consistent with material from the Walsh River area. F1871 Aconeceras walshense (Etheridge fil.,) Whitehouse, 1926, pp,203- 4, Aconeceras walshense (Tenison-Woods); Whitehouse, 1927, p,J 14. LOCALITY: Walsh River, NQ. FORMATION: Blackdown Fm. AGE; Lower Cretaceous, (Aptian). REMARKS: See F1272. Actinocrinus sp. F17784 Actinocrinus sp. ind,; Etheridge fil., 1892, pp.207-8, LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. Alathyria pertexta Iredale, 1934 F6546 Alathyria pertexta Iredale, 1934; Hill et al., 1970, pl.Cz5, figsBa,b. LOCALITY: Maryvale Creek, 0.5 miles NW of homestead, NEQ. FORMATION: Unnamed fluviatile deposit. AGE: Pleistocene. Amerianna carinata (Adams, 1861) F655! Amerianna carinata (Adams, 1861); Hill et al., 1970, pl.Cz4, fig.11. LOCALITY: Gowrie Creck, Darling Downs, SEQ. FORMATION: Unnamed fluviatile deposit. AGE: Pleistocene. Anadara trapezta (Deshayes, 1840) F15649 Anadara trapezia (Deshayes, 1840); Hill et al., 1970, pl.Cz5, figs2a,b. LOCALITY: Bullock Pt, Wide Bay, SEQ. FORMATION: Unnamed marine deposit. AGE: Pleistocene. REMARKS: Previously registered as Mo2921 in .he Queensland Museum Mollusca register. INVERTEBRATE FOSSIL TYPES Anidanthus springsurensis (Booker, 1932) F2542 PARATYPE Linoproductus springsurensis Booker, 1932, pp.67-8, pl.3, figs 3,4. Anidanthus springsurensis (Booker); Hill, 1950, pp.10-2. Anidanthus springsurensis (Booker, 1932); Water- house and Briggs, 1986, pp.61-2, LOCALITY: Cattle Creek, Springsure Dome, CQ, FORMATION: Lower Bowen Basin. AGE: Permian. F2543 PARATYPE Linwproductus springsurensis Booker, 1932, pp.67-8, pl.4, fig.3. Anidanthus springsurensis (Booker); Hill, 1950, pp. 10-2, Anidanthus springsurensis (Booker, 1932); Water- house and Briggs, 1986, p.61-2. LOCALITY, FORMATION, AGE: As for F2542. F2544 PARATYPE Linoproductus springsurensis Booker, 1932, pp.67-8, pL4, figs 1,2. Anidanthus springsurensis (Booker); Hill, 1950, p.10- 2, Anidanthus springsurensis Booker, 1932; Waterhouse and Briggs. 1986, pp.61-62. LOCALITY, FORMATION, AGE; As for F2542, F2545 HOLOTYPE Linoproductus springsurensis Booker, 1932, pp.67-8, pl.3, figs 1,2. Anidanthus springsurensis (Booker); Hill, 1950, pp. 10-2. Anidanthus springsurensis Booker, 1932: Waterhouse and Briggs. 1986, p.61-2. Anidanthus springsurensis (Booker, 1932); Parfrey, 1986, pp.59-60. LOCALITY, FORMATION, AGE: As for F2542. F2546 PARATYPE Linoproductus springsurensis Booker, 1932, pp.67-4, pl.3, figs 5,6. Anidanthus springsurensis (Booker); Hill, 1950, pp. 10-2. Antdanthus springsurensis Booker, 1932; Waterhouse and Briggs, 1986, pp-61-2. LOCALITY, FORMATION, AGE: As for F2542, F2541 PARATYPE Linoproductus springsurensis Booker, 1932, pp.67-8, pl.4, fig.4. 669 Anidanthus springsurensis (Booker); Hill, 1950, pp. 10-2, Anidanthus springsurensis Booker, 1932; Waterhouse and Briggs. 1986, pp.61-2. LOCALITY, FORMATION, AGE: As for F2542. Aprathia rockhamptonensis (Mitchell, 1918) F1233 cf. Phillipsia woodwardi Etheridge fil., 1892 p.215, pl.44, fig.6. SYNTYPE Phillipsia roackhamptonensis Mitchell, 1918, pp.440, 459-61, pl. 50, lig.8 PARALECTOTYPE Aprathia rockhamptonensis (Mitchell, 1918) Engel and Morris, 1989 p.327- 332, fig.12h. LOcALITY: Rockhampton District, MEQ, FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. Engel and Morris (1989) incorrectly record the type locality as Corner Creek, Great Star River, north Queensland, Star Beds. The Palacantology register gives the locality as Rockhampton, MEQ. F1234a/b Phillipsia dubia Etheridge; Etheridge, 1892, pp.214-5, p44, fig.4. SYNTYPE Phillipsia rackhamptonensis Mitchell, 1918, pp.440, 459- G1, pl. 49, figs 9, 10, LectotTyre Aprathia rackhamptonensis Mitchell Engel and Morris (1989), p.327-332, figs 12a-c. LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE; Lower Carboniferous. REMARKS: Mitchell (1918) referred to this specimen as 716 which is a de Vis collection number, Engel and Morris (1989) erroncously refer to counterparts F1234a and F1234b as a paralectolype and lectotype respectively. The only paralectotype is F1233 and the lectotype is F1234a/b. (Sce also F1233), Archaeocidaris sp. F17785 Archacocidaris sp.; Etheridge fil., 1892, p.213. LOCALITY: Rockhampton District, MEQ. FORMATION: ?7Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. 670 MEMOIRS OF THE QUEENSLAND MUSEUM Fic, 2. A. HoLotyre Luciella (?) grayae, F1227, x3.2; B. Athyris roysii Leveillé, F1207, x3.7; C.Baylea konincki (Etheridge [il., 1890), F1222, x3.8; D. Collabrina cliftoni (Etheridge), F15648, x1.5; E. Collabrina cliftoni (Etheridge), F15648. x 2.0; F. Collabrina cliftoni (Etheridge), F15647, x2.0; G. Loxonema sp, F1225, x4.4; H. SYNTYPE Fdmondia (7) smithi Etheridge fil, 1894, F2837, x5.2: 1. HoLotyPE Cypricardella rectangularis Etheridge. F 15647, x5.4. INVERTEBRATE FOSSIL TYPES Armenoceras sp. F7229 Armenoceras sp. 4: Wade, 1977b, pp.308-9, fig.4. LOCALITY: W side of small hill, 23°19°S, 138°06°E, CWQ. FORMATION: Lower Nora Fm, AGE: Middle Ordovician, Astacodes sp. F2907 Astacades sp,; Woods, 1957, p.159, pl.4, fig.1, text fig.2, Astacades sp; Hill et al., 1968, pl.K11, fig.5 LOCALITY: ‘Currane’, 10 miles N of Dartmouth, ca. FORMATION: Allaru Mudstone AGE: Lower Cretaceous, (Albian). Athyris roysit Leveillé, 1835 F1207 (Fig.2B) Athyris roysii Leveillé, Etheridge fil., 1894, pp.529- 30, pl. 39, fig.4. LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Upper Carboniferous. Aucellina hughendensis (Etheridge, 1872) F3845 HOLOTYPE Avicula hughendenensis Etheridge, 1872, p.346, pl.25, fig.3. Aucella hughendenensis (Etheridge) Etheridge fil, 1892, pp.460-2, pl.25, fig.1. Aucella hughendenensis Etheridge; Etheridge fil., 1902a, p.67. Aucella hughendenensis Etheridge; Etheridge fil., 1902b, p14. Aucella hughendenensis Etheridge: Etheridge fil., 1907, pp, 321-2, Aucellina hughendensts (Etheridge Snr, 1872); Hill e¢ al., 1908, pl.K4, fig. 1. LOcALIty: Hughenden Stn, c. 0.5 miles from Hughenden Stn, c. 3 miles from Mt Walker, near Hughenden, CQ. FORMATION: Wallumbilla Fm, Ranmoor Member. AGE: Lower Cretaccous. Australiceras irregulare (Tenison-Woods, 1883) F1265 671 Crioceras jackii, Eth. fil.; Etheridge fil., 1909, pp.145- 8, p37, fig.2. Australiceras gracile (Sinzow); Whitehouse, 1926, pp.21 1-2. Australiceras irregulare (Tenison-Woods, 1883); Day, 1974, pp. 10-12, Table 1. LOCALITY: Walsh River, NO. FORMATION: Blackdown Fm. AGE: Lower Cretaceous, (Aptian). F1268 Crioceras jackit, Eth. fil.; Etheridge fil., 1909, pp.145- 8, pl. 36, fig. 1. Australiccras gracile (Sinzaw); Whitehouse, 1926, pp.211-2. Australiceras. irregulare (Tenison-Woods, 1883) in Day, 1974, pp, 1, 10-11, Table 1, pl.6, figs 2a-c, LOCALITY: Victoria Downs, Morven (see remarks). FORMATION: ? AGE; Lower Cretaceous. REMARKS: F1268. was previously registered in the Donor Register as D7710.12. D7710 is recorded in the Donor register as a ‘miscel- lancous collection of fossils sce Catalogue of Fossils’. Unfortunately no catalogue has been found. D7710 includes material from various localities, e.y., Gilbert River, Walsh River, NO, and the Simbirskites specimens (q.v.), Which are probably from North Germany. While Whitchouse (1928) records the locality for F 1268 (and the Simbirskites specimens) as Vic- toria Downs, Morven, the provenance of this specimen and other D7710 material is uncertain, F1269 Criovceras jackti, Eth. fil; Etheridge fil, 1909, pp.145- &, pl.35, fig. 1. Australiceray all, irregulare (Tenison- Woods): Whilehouse, 1926, p.210. Australiceras irregulare (Tenison-Woods, 1883): Day, 1974, pp, 10-12, Table 1. LOCALITY; Walsh River, NO. FORMATION: Blackdown Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Collected Hann’s Expedition, 1872. FI391 Australiceras irregulare (Tenison-Woods); Whitehouse, 1926, p. 210, pl.37, figs 1a,b. LOCALITY: Wrotham Park Stn, Walsh River, NQ. FORMATION: Blackdown Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Collected Hann’s Expedition, 1872. F1392 Australiceras gracile (Sinzow); Whitehouse, 1926, pp.211-2, pl. 34, fig.4. LOCALITY: not recorded. FORMATION: ? AGE: Lower Cretaceous. REMARKS: This specimen was not mentioned in Day's (1974) paper, but is probably referable to A, irregulare, Australiceras jacki (Etheridge fil., 1880) F1267 Crioceras fackil Eth. fil.; Etheridge fil., 1909, pp.145- 8, plas, fig.3. Australiceras jacki (Etheridge [il.,) Whitehouse, 1926, pp.209-10. Australiceras jacki (Etheridge Ir, 1880); Day, 1974, pp.9-10, Table 1. LocAuity: Hughenden Station, sheep wash water-hole, 800m from Stn, and 5km E of Mt Walker, near Hughenden, CQ. FORMATION: ?Wallumbilla Fm. AGE: Lower Cretaceous. F1390 HoOLoTYPE Australiceras transiente Whitehouse, 1926, p.212, pl. 34, ligs 3a,b. Australicerus jacki (Etheridge Jr. 1880); Day, 1974, pp.8-10, Table 1. LOCALITY: Walsh River, NQ. FORMATION: Blackdown Fm, AGE: Lower Crelaccous, (Aptian). F1396 Australiceras rebustum Whitehouse, 1926, p.211. LOcALity: Walsh River, NQ. FORMATION: Blackdown Fm. AGE; Lower Cretaceous, (Aptian). REMARKS: Day (1974) synonymised A. robus- unt with A. jacki. The matrix of this specimen originally contained F1271 and F1797, which were referred to ?Tunohamites taylori by Day (1974). P1597 Australiceras jacki (Etheridge fil.,.) Whitehouse, 1926, pp.209- 210, pl.34, fig.2. Australiceras jacki (Etheridge Jr., 1880); Arkell et al., 1957, pp.L211, L213, fig.240-6. Australiceras jacki (Etheridge, 1880); Hilleral., 1968, pl. K7, fig.1. Australiceras jacki (Etheridge Jnr., 1880); Day, 1974, pp.9-10, Table 1. MEMOIRS OF THE QUEENSLAND MUSEUM LOCALITY: Walsh River, NQ. FORMATION: Blackdown Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Collected Hann’s Expedition, 1872. Whitehouse (1926, pp.209-10) erroneously referred to F1597 as holotype of this species. The type material however had been originally figured by Etheridge (1880) and the lectotype was selected by Day (1974) and is held in the Geological Survey of Queensland (GSQ F1385). Aviculopecten laurienti Etheridge fil., 1892 F1209 SYNTYPE Aviculopecten laurienti Etheridge fil,, 1892, pp.268-9, pl.43, fig.4. FEuchondria laurienti (Etheridge, 1892); Maxwell, 1964. p.49. Aviculopecten laurienti Etheridge, 1892b; Water- house, 1982, p. &. LOCALITY: Rockhampton, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. Baylea konincki (Etheridge fil., 1890) F1222 (Fig.2C) Yvania konincki Eth, fil.; Etheridge fil., 1892, p.288, pl. 41, fig.7. LOCALITY: Rockhampton District, MEQ. FORMATION; ? AGE: Carboniferous. REMARKS; De Vis Collection. Generic assign- ment changed by Knight er al. (1960, p. L202). Bellerophon sp. F1223 Bellerophon sp. Etheridge fil., 1894, p.537, pl.39, fig.8. LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS; De Vis Collection. Bembicium melanostoma (Gmelin, 1791) F6552 Bembicium melanostama (Gmelin, 1791); Hill et al., 1970, pl. C24, fig. 15. LOCALITY: Nudgee, SEQ. FORMATION; Unnamed estuarine deposit. AGE: Pleistocene/Holocene? INVERTEBRATE FOSSIL TYPES Beudanticeras daintreei (Etheridge, 1872) F385] SYNTYPE Ammonites daintreei Etheridge, 1872, p.346, pl.24, fig. 1 (apertural view only). Ammonites daintreei Etheridge; Etheridge fil., 1892, p.495, pl.29, fig. 1. Beudanticeras (?) daintreei (Etheridge) Whitehouse, 1926, p,221, text fig.2. LECTOTYPE Bolileceray duinireei (Etheridge) Whitehouse, 1928, pp.203-4, pl.26, fig.2. Beudanticeras dainireeit (Etheridge); Arkell et al., 1957, p. L368. Locauity: Hughenden - about 800m from Hughenden Stn, and about Skm E of Mt Walker, CQ. FORMATION: ?Allaru Mudstone. AGE: Lower Cretaccous. REMARKS: Sce Beudanticeras perlatum (F1276). Daintree Collection, F3852 SYNTYPEAmmonites daintreei Etheridge, 1872, p.346, pl.24, fig.2. Ammonites daintreei (Etheridge) Etheridge fil., 1892, p.495, pl.29, fig.3. Beudanticeras (?) daintreei (Etheridge) Whitehouse, 1926, p.221. Boliteceras perlatum Whitehouse, 1928, p.204, Beudanticeras daintreei (Etheridge); Arkell ef al., 1957, p. L368. Boleticeras [sic] daintreei (Etheridge Snr, 1872); Hill eral., 1968, pl.K8, fig.4. LOCALITY: Hughenden, CQ. FORMATION: Wallumbilla Fm., Ranmoor Mem- ber. AGE: Lower Cretaceous. Beudanticeras flindersi (McCoy, 1865) F1382 Beudanticeras flindersi (McCoy); Whilehouse, 1926, pp.219-20, text fig. 1. Beudanticeras flinderst (MeCoy); Whitehouse, 1928, pp.201-3. LOCALITY: Coolibah, Hughenden, CQ, FORMATION: ?Wallumbilla Fm, AGRE: Lower Cretaccous. Beudanticeras mitchelli (Etheridge, 1872) F1274 Ammonites beudantiBrongn., vat. mitchelli Etheridge, 673 1872, p.345, pl. 23, fig.1 (laleral and apertural views). Ammonites flinderst McCoy; Etheridge fil., 1892, pp.494-5, pl 30, figs 1,2. Beudanticeras flindersi (McCoy) Whitehouse, 1926, pp.219-20, Lectotyre Beudanticeras mitchelli (Etheridge) Whitehouse, 1928, p,201, pl.25, fig.2. LOCALITY: Hughenden, CQ. FORMATION: ?Allaru Mudstone. AGE: Lower Cretaceous. REMARKS: Whitehouse (1928, p.206) etroneous- ly listed this Hughenden specimen as coming from Marathon (Daintree, 1872). F1275 Ammonites beudantiBrongn., var. mitchelli Etheridge, 1872, p.345, pl. 23, fig.2. Ammonites flindersi McCoy; Etheridge fil,, 1892, pp.494-5, pl.30, fig.3. Beudanticeras flindersi (McCoy); Whitehouse, 1926, pp.219-20. Beudanticeras mitchelli (Etheridge); Whitehouse, 1928, pp.201-2. Loca.ity: Hughenden, CQ. FORMATION: ?Allaru Mudstone AGE: Lower Cretaceous. REMARKS: Daintree Collection. Beudanticeras perlatum (Whitehouse, 1928) F1276 Ammonies daintreet Etheridge, 1872, p.346, pl.24, fig.1 (see remarks). Ammonites daintreei Etheridge; Etheridge fil., 1892, p.495, pl.29, fig.2. Houotyre Boliteceras perlatum Whitehouse, 1928, p.204, pl.26, fig.3. Beudanticeras perlatum (Whitehouse, 1928) Arkell elal., 1957, p. L368. Locatity: Hughenden, CQ, FORMATION: ?Allaru Mudstone. AGE: Lower Cretaceous. REMARKS: Whitehouse (1928, p.204) considered that Etheridge (p|.24, fig.1) was a composite figure based upon this specimen and F3851,. Etheridge (1872) figured most of his. material faithfully including imperfections and cracks. He also had complete specimens of this taxon to figure. It is therefore surprising and we think unlikely that he chose to produce a composite figure. However, no specimen in our collection matches Etheridge’s original figure and this presumably led Whitchouse to his interpretation. 674 Beudanticeras sutherlandi (Etheridge, 1872) F1273 HOLOTYPE Ammonites sutherlandi Etheridge, 1872, p.345, pl.21, tig4, Ammonites sutherland’ (Etheridge); Etheridge fil, 1892, p. 496, pl.29, fig.4. Beudanticeras (7) sutherlandi (Etheridge); Whitehouse. 1926, p, 222. Beudanticeras sutherlandi (Etheridge), Whitehouse, 1928, p.202, pl.25, fig.4. Beudanticeras sutherlandi (Etheridge Snr,, 1872); Hill et al., 1968, pl.K9, fig.2. LOcALITY: ?McKinlay’s Range or Marathon Stn, CQ, FORMATION: ?Allaru Mudstone, AGE: Lower Cretaceous, REMARKS: All previous authors record The locality of this specimen as Marathon or Marathon Stn, This specimen was previously registered as D222 in the Queensland Museum Donor register which records the locality as McKinlay’s Range. Beudantiella ogilviei (Whitehouse, 1928) F1897 Cophinoceras ogilviei Whitehouse, 1928, p.205. Beudantiella ogilviei (Whitehouse) Breistroffer, 1947, p.83. LOCALITY: Near mouth of Bynoe River, on Nor- manton-Burketown Rd, NQ. FORMATION: Normanton Fm. AGE: Lower Cretaceous, (Albian). REMARKS: BSeudantiella Breistroffer (1947) replaces Whitchouse's Cophinoceras, which had been previously occupied by Kophinoceras Hyatt. (Arkell et a/., 1957, p.L368). F2043 HooryPe Caphinaceras ogilviei Whitehouse, 1928, p.205, pl.26, figs 4a,b. Caphinoceras ogilvier Whitehouse, 1928; Hill et al., 1968, pi, KS, fig.5. LOCALITY, FORMATION, AGE: As for F1897. REMARKS: See F1897. Brachymetopus maccoyi var, spinimarginatus Hahn and Hahn, 1969 F2839 Syntyre Brachymiectopus dunstani Mitchell, 1918, p.486-8. pLSt. fig.4; pl52, fig... MEMOIRS OF THE QUEENSLAND MUSEUM LOCALITY: Rockhampton District, MEQ. FORMATION: 7Malchi Fin. AGE: Lower Carboniferous, REMARKS: De Vis collection No. 712. Other poorly preserved Brachymetopus pyzidial frag- ments are associated with this specimen (Mitchell, 1918). This specimen is referable to B. maccoyl Var. spinimarginatus Hahn and Hahn, 1968 (B. Engel, pers. comm., 1988). Bucania textilis de Koninck, 1883 F1224 Bucania textilis de Koninck ?; Etheridge fil., 1892, p.290, pl.41, fig.8. Locatrry: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. Burrows F1198 Burrow; Etheridge fil. 1892, pp.297-9, pl.44, fig.15. LocaALiry: Rockhampton District, MEQ. FORMATION: ? AGE: Carboniferous? REMARKS: The figure is laterally reversed in Etheridge fil., (1892, pl.44, fig.15). F1199 Burrow; Etheridge fil., 1892, pp.297-9, pl.39, fig.1. LOCALITY, FORMATION, AGE; As for F1198. REMARKS: specimen missing. F1200 Burrow; Etheridge fil., 1892, pp.297-9, pl.44, fig. 16, LOCALITY, FORMATION, AGE: As for F1198, F1201 Burrow; Etheridge fil., 1892 ,pp.297-9, pl.44, fig,17. LOCALITY, FORMATION, AGE: As for F1198, F1202 Burrow; Etheridge fil., 1892, pp.297-9, pl.44, fig. 18. LOCALITY, FORMATION, AGE: As for F1198. ?Cancrinelloides sp, F14414 Productus cora d°'Orb,; Etheridge, 1872, pp.328-9, pl.15, fig. 1. Productus cara d'Orb.; Etheridge fil., 1892, p.248, pl.l2, fig.14, LOcALitry: Caledonian Reef, Gympie, SEQ. (INVERTEBRATE FOSSIL TYPES FORMATION; Rammutt Fm. AGE: Permian. REMARKS: Waterhouse and Balfe (1987) con- sidered that P. cora was allied to Cancrinel- loides. P14415 Praductus cora d‘Orb.; Etheridge, 1872, pp.328-9, pls, fig, 2. LOCALITY: Gympie, SEQ, FORMATION: Rammutt Fm. AGE; Permian, REMARKS: See F14414 above. Carpocrinid indet. F14881 Carpocrinid indeL.; Jell ev al., 1988, p.364-6, figs7g-). LOcALITy: QMLS47 = (UQL5209) Burges~ topographic sheet [648459] prominent limestone knoll on left bank 100m from mouth of 2nd left bank side creck upstream from Jack Hills Gorge on the Broken River, Wando Vale Stn, 150km NW of Charters Towers, NO. FORMATION: Burges Fm. AGE: Early Middle Devonian, (Late Emsian- early Eifclian). Cercopoidea incertae sedis F6498a/b Cercopvidea incertae sedis; Evans, 1971. p.146, fig.4a. LOCALITY: Mt Crosby, SEQ, Upper Bed, 910805-911805, Ipswich 1 mile military map. FORMATION: Mt Crosby Fm. AGE: Early Late Triassic (Carnian). F6542a Cercopuidea tcertae sedis, Evans, 1971, p.146, fig.4b, LOCALITY, FORMATION, AGE: As for F6498. F6507a/b Cercopoidea incertae sedis; Eyans, 1971, p.146, LOCALITY, FORMATION, AGE: As for F6498. Charybdis callianassa (Herbst, 1789) F6559 Charybdis calliariassa (Herbst, 1789); Hill er al, 1970, pl.Cz 6, fig. 10- LOCALITY: Ayr, NQ, FORMATION: Unnamed estuarine deposit. AGE: Pleistocene/Holocene. 675 Chelodes whitehousei Runnegar, Pojeta, Taylor and Collins, 1979 F10087, FI0091, F10095, F10098-F10102, FIO104 PARATYPES Chelodes whitehousei Runnegar, Pojeta, Taylor and Collins, 1979, pp,1374-94, F10087, pl.2, figs 4-6; F10091, pl, 2, figs 12-14; F10095, pl.2, figs 24-25, 30; FI0098, pl.2, figs 36-38; F10099, pl.2, figs 42-44; F10100, pl.2, fig.45; FIOIOL, pl.2, figs 46-47; F10102, pl.2, figs 48-50: F10104, pl.2, figs 57-59. LOCALITY: QML278, G25, Glenormiston Stn road about 5kms E of No. 21 Bore, Glenormiston Stn, WCO. FORMATION: Ninmaroo Fm. AGE: Lower Tremadocian, (Datsonian). F10088, F10090, F10092-F10094, F10096- F10097, F 10103 PARATYPES Chelodes whilehousei Runnegar, Pojeta, Taylor and Collins, 1979, pp.1374-94; F10088, pl.2, fig.7; F10089, pl.2, figs 8-10; F10090, pl.2, figs 11-15; F10092, pl.2, figs 16-18; F10093, pl.2, figs 19-21; F10094, pl.2, figs 22-23; F10096, pl.2, fips 26-28; F10097, pl. 2, figs 33-35; F10103, pl.2, figs 51-53, Locality: QML278, G24, Glenormiston Stn toad ec. 8kms Eof No. 21 bore, Glenormiston Stn, WwCco. FORMATION: Ninmaroo Fm. AGE: Lower Tremadocian,(Datsonian). Chthamalis sp. F2026a Chthamalus sp.; Withers, 1932, pp.122-3, Chthamalus sp.; Hill et al., 1970, pl.Cz6, fig.3. LocaLity: Magnetic Island, NQ. FORMATION: Raised beach rock. AGE: Pleistocene/Holocene? REMARKS: Withers (1932) used F2026 for four different taxa. This specimen has been Tercgistered as F2()26a. Collabrina cliftoni (Etheridge, 1872) F15647-48, F16566 (Figs2D-F) Pleurotomarid clifioni Etheridge, 1872, p.347, pl.25, fig.4. Pleuratomaria | cliftoni Etheridge; Etheridge fil. 1892, pp. 484-5, pl.29, fig. 10 (refigured Etheridge’s, 1872 illustration), Pleurotomaria (?) cliftont Etheridge; Etheridge fil., 1902a, p. 79. oH LOCALITY! Head of Roper Creck, Gordon Downs Stn, CQ, FORMATION: ? Blenheim Sub-group. AGE: Upper Permian. REMARKS: Three specimens (F15647-48, F16566) from Gordon Downs are in the Queensland Museum collections and were col- lected by W.B. Clarke (Daintree, 1872, p.283). Stock Mortgages and bills of sale and other State Archive records indicate thal the only Gordon Downs Stn existing in Queensland in the 1860's (al the time of collection) was in the Clermont District, north of Emerald. Maps showing station boundaries in ihe [S80’s indicate that Roper Creck commenced near the northern boundaries of the station. Earlier maps that might show the extent of the station in the 186(’s do not exist. The Gordon Downs property and adjoining Mal- vern Downs Sta, which includes the headwaters of the Roper Creek, were owned in the 1860's by Samuel and Roderick Travers, respectively. Pleuratomaria cliftoni has been missed or ig- nored by subsequent workers possibly because of confusion regarding the locality. Etheridge (1872) considered the material equivalent to the English Oolite, presumably inferring a Jurassic age for this material. Etheridge fil. (1892) con- fused the whereabouts of the locality by suggest- ing it was near Roma, because of the existence of a Gordon Downs Stn in that urea. He referred the material to the Rolling Downs Group (Cretaceous) but noted that ‘the resemblance of this species to a typical Pleurotomaria is not a marked one, in fact itis very much more like the Palacozoic shells called Murchisonia with rounded whorls’. Daintree (1872, p.283) com- mented (hat the Gordon Downs material ‘on Mr Etheridge’s authority may be referred to the same horizon as the Pelican Creek Beds’. These beds. are now recognised as Permian in age (Olgers, 1969). Furthermore Daintree (1872, p. 286) cal- lected atypically Permian fauna of Spirifera spp. and Productus clarkei from presumably the same area. This collection was separate to the one made by the Rev. W.B. Clarke which included Pleurotomaria and bivalves. All the Queensland Museum material from Gordon Downs including a specimen figured as Myacites sp. by Etheridge (1872) were given the No. 010. The number does not pertain to any known Museum register. The Queensland Museum specimens of Pleurolomaria clifioni differ in size ftom Etheridge's original figure, however there is lit- tle doubt that they are part of the original material collected and are topotypes. Pleurotomaria clif- MEMOIRS OF THE QUEENSLAND MUSEUM toni is referable to the Permian genus Collabrina based on the high-spired shell with convex Whorls.and prominent selenizone and is probably conspecific with Collabrina parva (Wass), Topotypes from this locality, F15647 and F15648, are figured (Figs 3A,B). Conophillipsia grandis (Etheridge fil. 1892) F1232 SYNTYPE Phillipsia woodwardi Etheridge fil., (892, p.215, pl. 44, fig.sS. Phillipsia woodwardi Eth. fil.; Mitchell, 1918, pp.465- 71, pl.4Y, fig.9, pLSO, fig. 11, pl.S1, fig. 12. Conophillipsia grandis (Etheridge Ir, 1892) Engel and Morris, 1984, pp.53-8, figs18c, 19a. LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous, (late Tournaisian). REMARKS: De Vis Collection, Conophillipsia subquadrata Engel and Mortis, 1984 F2839 Phillipsia woodwardi Eth. fil.; Mitchell, 1918, pp.465- 71, pl51, fig, 14; pl.52, fig.1. Conophillipsia subquadrata Engel and Morris, 1984, pp.50-3, figs 16b,e; 17h. LOCALITY: Trilobite Ridge, Mt Morgan, MEQ. FORMATION: ? AGE: Lower Carboniferous, (mid-Tournaisian). REMARKS: The number 712 referred to in Mitchell (1918) is a de Vis collection number. Corbiculina australis (Deshayes, 1830) F6547 Cerbiculina australis (Deshayes, 1830); Hill et al. 1970, pl. Cz 5, fig.10. Locauity: Maryvale Creek, 0.5kms NW of homestead, NEQ, FORMATION: Unnamed fluviatile deposit. AGE: Pleistocene. Crassostrea commercialis (Iredale and Rough- ley, 1933) F6548 Crassostrea commercialis (Iredale and Roughley, 1933); Hill et al, 1970, pl-Cz5, fig.1. LOCALITY: Nudgee, SEQ. FORMATION: Unnamed estuarine deposit. AGE; Pleistocene/Holocene. INVERTEBRATE FOSSIL TYPES 677 Crinoid attachment bases F14858 Crinoid attachment bases; Jell ef al., 1988, p.397, fig.27p. LocALiry: UQLS318, Wando Vale [565395] 65-1 16m aboye base of formation in gully 2.2km NE of Storm Dam, Wando Vale Stn, 150km NW of Charters Towers, NQ. FORMATION: Papilio Mudstone. AGE: Givetian, Middle Devonian. REMARKS: Jell ef al, (1988) refer to this specimen as F14860 on p.397, The correct num- ber is F14858 as on figure cpation p.396. F14859 Crinoid attachment bases; Jell ef al, 1988, p.397 fig.27q, LOCALITY, FORMATION, AGE: As for F14858. Crinoid calyx F1194 ‘Crinoid calyx’; Etheridge fil., 1892, p.210, pl.44, fig.8. LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Early Carboniferous. Crinoid indeterminate F14951 Crinvid indet 1.; Jell ef al., 1988, p.397, ligs7a-c. LocaAtity:; QML547 (= UQLS209) Burges [648459] prominent limestone knoll on left bank 100m from mouth of 2nd left bank side creck upstream from Jacks Hill’s Gorge on the Broken River, Wando Vale Station, 150km NW of Charters Towers, NQ. FORMATION: Burges Fm. AGE: Late Emsian - Givetian. Early Middle Devonian, REMARKS: Jell et al. (1988) refer to this specimen as F14591 on p.397, The correct num- ber is F14951 as on figure caption 365. F14952 Crinoid indet 1.; Jell et al. (1988, p.397, fig.7e). LOCALITY, FORMATION, AGE; As for F14951. REMARKS: Jell ef al. (1988) refer to this specimen as F14592 on p.397. The correct num- ber is P14952 as on figure caption p.365. F14953 Crinoid indet 1.; Jell et a/., 1988, p.397, fig.7f. LOCALITY, FORMATION, AGE: As for F14591. REMARKS: Jell e¢ al. (1988) refer to this specimen as F14593 on p.397. The correct num- ber is F14953 as on figure caption p.365. F14954 Crinoid indet 1.; Jell er a/., 1988, p.397, fig.7d. LOCALITY, FORMATION, AGE: As for F14591. REMARKS: Jell et al. (1988) refer to this specimen as F14594 on p.397. The correct num- ber is F14954 as on figure caption p.365. Crinoid cf. Stemmatocrinus F1196 Basal cup of Crinoid cf. Stemmatocrinus’; Etheridge fil,, 1892, p.208, pl.44, fig.7. LOCALITY: Stony Ck, Stanwell, or Rockhampton MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous, REMARKS: Preserved on the same specimen are arms of crinoids referred to by Etheridge fil.(1892, p.210). Crosaphis anomala Evans, 1971 Fe508a/b HOLOTYPE Crosaphis anomala Evans, 1971, p.147. fig.3a, Loca.Lity: Mt Crosby, SEQ, Upper Bed, 910805-911805, Ipswich 1 mile military map. FORMATION: Mt Crosby Fm. AGE: Early Late Triassic (Carnian), Cucullaea semistriata Moore. 1870 F5595, F5598-F5600, F5602 Cucullaea semistriata Moore, 1870; Skwarko, 1974, pp.74-6,text fig.3. LOCALITY: ?Newmarracarra Limestone, Western Australia FORMATION: ?Newmarracarra Limestone. AGE: Middle Jurassic, (Bajocian). Cupressocrinites abbreviatus Goldfuss, 1839 F14582 Cupressocrinites abbreviatus Goldfuss, 1839: Jell et al,, 1988, pp.393-5. LocaALity: UQLS293, Wando Vale [524369] gully 400m E of the divide between Dosey and Page Crecks, Wando Vale Stn, 1S0km NW of Charters Towers, NQ. 7K FORMATION: Papilio Mudstone, AGE: Middle Devonian, (?Givetian). REMARKS: This formation was formally named by Lang etal. (1989) as Papilio Mudstone, F14585-F 14586 Cupressocrinites abbreviatus Goldfuss, 1&39; Jell er al., 1988, pp.393-5, LOcALITY: UQL5229. Wando Vale topographic Sheet (860371) right bank of Storm Dam Creek about 50m upstream from the confluence with the gully into which Storm Dam overflows when filled. Wando Vale Stn, 150km NW af Charters Towers, NQ. FORMATION: Papilio Mudstone. AGE: Middle Devonian, (Givetian). REMARKS: Jell ef al., (1988) erroneously record the locality for this specimen as UQLS348. Otherwise as for F14582. F14587-F 14589 Cupressocrinites abbreviatus Goldfuss, 1839; Jell e¢ al,, 1988, pp.393-S, LocALity: UQL5277, Wando Vale topographic Sheet [570409], 51-56m above base of section 300m ESE of The Valeano, Wande Vale Stn, 150km NW of Charters Towers, NO. FORMATION: Papilio Mudstone, AGE: Middle Devonian, probably early Givetian. REMARKS: As for F14582. F14598-F 14599, F14601, F14603 Cupressocrinites abbreviatus Goldluss, 1839; Jell er al,, (988, pp.343-5. LOCALITY: UQL5356, Wando Vale topographic Sheet [554367] east flank of Spongophyllum Hill, in head of eroding gully, Wando Vale Stn, 150km NW of Charters Towers, NQ. FORMATION: Papilio Mudstone. AGE: Middle Devonian, Givetian, REMARKS: As for F14582. FI4788 Cupressocrinites abbreviatus Goldtuss, 1839; Jell ef al.. 1988, pp.393-5. LOCALITY: From float in Burdekin River near Big Bend, north of Charters Towers, NO. FORMATION, ?Burdekin Fm. AGE: Middle Devonian. F14856 Cupressocrinites abbreviats Goldfuss, 1839; Jell er al., 1988. pp.393-5, fig.26g. LOCALITY: UQLS318, Wando Vale topographic Sheet (565395) 65-1]6m above base of forma- MEMOIRS OF THE QUEENSILAND MUSEUM tion in gully 2.2km NE of Storm Dam, Wando Vale Stn, 150km NW of Charters Towers, NQ. FORMATION; Papilio Mudstone. AGE: Middle Devonian, (Givetian). REMARKS: This specimen was omitted from the list of material on p.393 but is figured on p.394. Otherwise as for F14582. F 14868, F14873, F14875 Cupressocrinites abbreviatus Goldfuss, 1839; Jell et al, 1988, pp.393-5. LOCALITY; UQL5321. Wando Vale topographic Sheet [562390] creck section 3km NE of Storm Dum, Wando Vale Stn, 150km NW of Charters Towers, NO. FORMATION: Papilio Mudstone. AGE: Middle Devonian, (late Eifelian-Givetian), REMARKS: As for F14582. Cupressocrinites sp. ef. C. gracilis Goldfuss, 1831 F14841 Cupressocrinites sp. cf. gracilis Goldfuss, 1831; Jell etal., 1988, p.395, figs27ij- LocALity: UQL3579, Burges [687683] one of § localities collected from E to W along the fence linc, 600m E of Martins Well windmill, from upper purt of limestone which is slightly folded, Martins Well Area, 8km E of Pandanus Creek Homestead, 200km NW of Charters Towers, NQ. FORMATION: Upper Martins Well Limestone Member, Shield Ck Fm, AGF, early Devonian, (early Pragian). Cypricardella rectangularis Etheridge fiJ., 1894 F1216 (Fig.2]) HoLoryre Cypricardella rectangularis Etheridge fil., 1894, pp. $31-2, pl.39, fig.S. LOcaLiTy: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. Decorotergum warrenae jell, 1983 F12294 HOLOTYPE Becorotergum warrenae Jell, 1983, pp.195-9, figs la,b,e,d, text fig.2. LOCALITY: Kolane Station, S8km ENE of Taroom, SEQ, INVERTEBRATE FOSSIL TYPES FORMATION: Westgrove Ironstone Member, Evergreen I'm. AGE: Jurassic (Late Liassic). F12295 PARATYPE Decorvtergum warrenae Jell, 1983, pp. 195-9, figs le.tg,h, LOCALITY, FORMATION, AGE: As for F12294. F12296 Decorotergum warrenae Jell, 1983, pp.195-9. LOCALITY, FORMATION, AGE: As for F12294. Deltopecten limaeformis (Morris, 1845) F1208a,b HOLOTYPE Aviculopecten multiradiatus Etheridge, 1872, p.327, pl.13, fig... Aviculopecten mulriradiatus Etheridge; Etheridge fil., 1892, p. 2608. Aviculopecten squamuliferus (Morris); Fletcher and Dun, 1929. p,5, Deltopecten limacformis (Morris) 1845; Runnegar and Ferguson, 1969, p.251, ?Corrugopecten squamuliferus (Morris, 1845); Waterhouse. 1982b, pp.8,37. Squamultiferipecten. squamuliferus (Morris, 1845); Waterhouse, 1987a, p.157. Deltapecten limacformis (Morris) ?; Waterhouse and Balfe, 1987, pp.24, 30, pl.J, fig.10. LOCALITY: Lady Mary Reef, Gympie, SEQ. FORMATION: Rammutt Fm. AGE: Lower Permian, REMARKS: Waterhouse (1982, p.37) erroneously recorded this Daintree specimen as housed in the GSQ. F14391 Deltopecten limaefermis (Morris) '?; Walerhouse and Balfe, 1987, pp.24, 30, pl.l, fig.1 1. LocaLity: Gympie, SEQ. FORMATION: Rummutt Fm. AGE: Permian. Dielasma sp. F1206 Divlasma sp.; Etheridge fil, 1892, p.227, pl.4d, figs. 1,2. LOCALITY: Rockhampton District, MEQ. FORMATION: ? AGE: Carboniferous. 679 Dimitobelus sp. Fo089 Dimitabelus sp.; Hill et al., 1968, pl.K2, figs 1Sa,b,c- LOCALITY: Marion Downs, near Boulia, CWQ. FORMATION: ?Toolebuc Fm. AGE: Albian. Dolatocrinus peregrinus Jcll et al., 1988 FI4818 HOLOTYPE Dolatacrinus peregrinus Jell et al., 1988, pp.387-5, figs22a-d, LOCALITY: UQL5277 Wando Vale topographic Sheet [570409] 51-S6m above base of section 300m ESE of The Volcano, Wando Vale Stn, 150km NW of Charters Towers, NQ. FORMATION: Papilio Mudstone. AGE: Middle Devonian, (probably early Givetian). REMARKS: This formation was formally named by Lang et al. (1989) as Papilio Mudstone. F14866-F 14867 PARATYPES Dolatocrinus peregrinus Jell et al., 1988, pp.387-8; F14866, fig.22e; F14867, figs 22f,g. LOCALITY: UQL5320 Wando Vale topographic Sheet [563392] section on ridge 2.9km NE of Storm Dam, Wando Vale Station, 1S0km NW of Charters Towers, NQ. FORMATION: Papilio Mudstone. AGE;Middle Devonian, (late Eifelian- Givetian). REMARKS: As for F14818. F14872 PARATYPE Dolatocrinus peregrinus Jell et al., 1988, pp.387-8, fig.22h. Locauity: UQL5321, Wando Vale [562390] creck section 3km NE of Storm Dam. Wando Vale Stn, 150km NW of Chariers Towers, NQ. FORMATION: Papilio Mudstone, AGE: Middle Devonian, (late Eifelian- Givetian). REMARKS: As for F14818, Dysmorphoptiloides elongata Evans, 1956 Fo493 Dysmorphoptiloides elongata Evans, 1956; Evans, 1971, p.146, fig.2. Locanity: Mt Crosby, SEQ, Upper Bed, 910805-911805, Ipswich 1 mile military map. FORMATION; Mt Crosby Fm. AGF: Early Late Triassic (Carnian), 680 Echinalosia ovalis (Maxwell, 1954) F1205 Strophalosia gerardi King; Etheridge fil., 1892, p.260-2, pl. 40, fig.7. Straphalosia gerardi King; Prendergast, 1942, pp.45- 47. Echinalosia ovalis (Maxwell, 1954); Waterhouse, 1980, pp.37-8. LOCALITY: Banana Creek, CQ. FORMATION: Flat Top Formation. AGE: Permian, Edmondia (?) smithi Etheridge fil., 1894 F1219 undescribed bivalve; Etheridge fil., 1892, pl, 39, fig.8. SYNTYPE Edmondia (?) smithi Etheridge fil., 1894, pp.533-4. LOCALITY: Rockhampton district, MEQ, FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. The figure of F1219 is laterally reversed in Etheridge fil. (1892, pl.39, fig.8). F2837 (Fig.2H) SYNTYPE Edmondia (7) smitht Etheridge fil., 1894, pp-533-4, pl. 39, fig.6. LOCALITY, FORMATION, AGE: As for F1219, REMARKS: De Vis Collection. Enoploclytia sp. F1280 Callianassa (7) spi; Etheridge fil,, 1917. p.10, pl.2, fig. 4 Enoploclytia sp.; Woods, 1957, p.166. LOCALITY: Walsh or Mitchell River (?), NQ. FORMATION: Blackdown Fm. AGE: Aptian, Lower Cretaceous. Enoploclytia terraereginae Etheridge fil., 1914 F3234 Enopluclytia terrae-reginae Etheridge Jr; Woods, 1957, pp.166-8, pl.4, figs 5,6, text fig.6- Enoploclytia terrae-rceginae Etheridge, 1914; Hill er al., 1968, plL.K11, fig.6. LOCALITY: ‘Currane’, 10 miles N of Dartmouth, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). F3235 MEMOIRS OF THE QUEENSLAND MUSEUM Enoploclytia terrae-reginae Etheridge Jr.; Woods, 1957, pp.166-8%, pl.4, figs8,9. LOCALITY, FORMATION, AGE: As for F3234. F3236 Enaploclytia terrae-reginae Etheridge Jr.; Woods, 1957, pp. 166-68, pl.4, fig.7. Enoploclytia terrae-reginae Etheridge, 1914; Hill et al., 1968, pl.K11, fig.7. LOCALITY, FORMATION, AGE: As for F3234. Eoscartoides bryani Evans, 1956 F3693a/b Eoscartoides bryani Evans, 1956; Evans, 1961, p.20, fig.4a, Eascartoides bryani Evans; Hill er al., 1965, pl.T14, fig.2. LocALity: Mt Crosby Insect Beds, SEQ. FORMATION: Mt Crosby Fm. AGE: Early Late Triassic, (Carnian). REMARKS: The counterpart of this. specimen originally referred to as F3694 has been re- registered as F3693b. F3695a/b Eoscartoides bryani Evans, 1956; Evans, 1961, p.20, fig.4b, LOCALITY, FORMATION, AGE: As for F3693. REMARKS: The counterpart of this specimen originally referred to as F3696 has been re- registered as F3695b. F3697a/b Eoscartoides bryani Evans, 1956; Evans, 1961. p.20, fig.4e. LOCALITY, FORMATION, AGE: As for F3693. REMARKS: The counterpart of this specimen originally referred to as F3698 is re-registered as F3697b. Euastacus ? sp. F5740 Parastacid; Hill er a/., 1970, pl.Cz6, fig.4. Euastacus ? sp,; Sokol, 1987, pp.81-2, fig. 1. LocaLity: Brittains Quarry, Darra, SEQ. FORMATION: Datta Fm. AGE: Palacocene, Eucalyptocrinites rosaceus Goldfuss, 1831 F14533, Fl4541, F14544, F14546, F14548 Eucalyptocrinites rosaceus Goldfuss, 1831, in Jell et al., 9&8, pp.377-8; F14533, fig.15q; F14546, figs ISe- g: F14548, fig. 15m. INVERTEBRATE FOSSIL TYPES Locatity: QMLS12 Wellington topographic Sheet [799867] richly fossiliferous limestone (Unit 18 of Johnson, 1975) 870m NNW of Mountain View Homestead on Wellington Caves Rd, 9km SSW of Wellington, NSW. FORMATION: Garra Fm. AGE: Early Devonian, (Pragian, sulcatus biozone), Euporismites balli Tillyard, 1916 F14376 Euporismitey balli Tillyard; Lambkin, 1987, pp.295- 300, fig.3. LOCALITY: Clay Pit, New Chum (near Robert Strect, Ebbw Vale), SEQ. FORMATION: Redbank Plains Fm. AGF: Paleocene or Eocene. Euspira reflecta (Moore, 1870) F1259 Houotyre Natica lineata Etheridge 1872, p.342, pl.21, fig. I. Natica vartabilis Moore, Etheridge fil., 1892 pp.485- 6, 573, pl3l, fig.2. [Refigured Etheridge’s 1872 illustration]. Euspira reflecta (Moore), 1870; Day, 1967, pp.8-9. LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). Fissilunula clarkei (Moore, 1870) F1250 HOLOTYPE Cyprina expansa Etheridge, 1872, p.338, pl.19, fig. I. Cyprina clarkei Moore; Etheridge [il., 1892, pp.474-S, 568-9, Cyprina? (vel Cytherea 7) clarkei, Moore; in Etheridge fil., 1902b, pp.32-3. Fissilunula clarke: (Moore) Etheridge fil., 1902a, pp.36-7. Fissilunula clarkei (Moore) 1870; Day, 1967, pp.14-6. Fiysilunula clarket (Moore, 1870); Fleming, 1970, pp.8-9, LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Daintree Collection, Galene bispinosa (Herbst, 1783) FII8S 681 Galene bispinasa Herbst; Etheridge fil. and Me- Culloch, 1916, pp. 10-1, pl.3, figs3,4. LocaALiry: New Channel, Mouth of Brisbane River, Morcton Bay, SEQ. FORMATION: Unnamed estuarine deposit. AGE: Pleistocene/Holocene. REMARKS: See Scylla serrata (F1187), F6558 Galene hispinosa (Herbst, 1783); Hill er al, 1970, pl.Cz6, fig 8, LOCALITY: ?Moreton Bay, SEQ. FORMATION: Unnamed estuarine deposit. AGE: Pleistocene/Holocene. Gasterocomid indet. F14s40 Gasterocomid indet.; Jell e¢ a/., 1988, p.391, figs27a-h, Locauity: UQL3579, Burges topographic Sheet [687683] one of five localities collected from E to W along fence line 600m E of Martins Well windmill. Martins Well, 8km E of Pandanus Creek Homestead, 200km NW of Charters Towers, NQ. FORMATION: Upper Martins Well Limestone Member Shield Ck Fm. AGE: Early Devonian, (early Pragian). REMARKS: F14840c is erroneously referred to as G14840c in Jell et al. (1988, p,396). Gastropoda (not specified) F1764()-F17641 Gastropods: Wood, 1972, p.331. LOCALITY: Maroochydore, cnr Kuran Rd and Broadwater St, 1.5-6,.5m below surface, SEQ. FORMATION: Estuarine deposits. AGE: Holocene. REMARKS: These gastropods included Polinices conicus (Lamarck, 1822) and P. sordidus Swain- son, 1821. Georgina andersonorum Wade, 1977a F7159 HoLotyPe Georgina andersenorum Wade, 1977a, pp.4. 8. pl.2, figs 1-4, text fig.5, fig.6d. LOCALITY: Halfway Dam Area, Tobermorey Stn, 25°53°S, 137°50°E, NT. FORMATION: Top of Coolibah Fm. AGE: Lower Middle Ordovician. F7160-7165, 7167-7174 PARATYPES Georgina andersonorum Wade, 1977a, pp.4.8, text fig. 5, F7164, pl.2, figs 5,6; F7165, pl. 1, figs 1-4, LOCALITY, FORMATION, AGE: As for holotype. F7166 PARATYPE Georgina andersonorum Wade, 1977a. LocALitTy: Pulatera Hills, Glenormiston Stn, 22°57°S, 138°21°E, CWO. FORMATION, AGE: As for holotype. REMARKS: Wade, 1977a (pl.1, figs 1-4) is referred tao as F7135, the correct number ts F7165. Georgina beuteli Wade, 1977a F7219 HOLOTYPE Georgina beuteli Wade, 1977a, pp.4,9-10, pls, fig. 6, text Nig.6e, LOCALITY: near Halfway Dam, Tobermorey Stn, 22°53'S, 137°50'E, NT. FORMATION: Upper third of Coolibah Fm. AGE: Lower Middle Ordovician. F7220-7221 PARATYPES Georgina beuteli, Wade, 1977a, pp.4,9- 10, F7220, pl. 5, lig.7. LOCALITY,FORMATION, AGE: As for holotype. Georgina dwyeri Wade, 1977a F7206-7212 PARATYPES Georgina dwyeri Wade, 1977a, pp.4,10- 11, F7206, pl. 6, fig.1. LOCALITY: Base of Oadatra Point, Toko Range, Glenormistan, 22°53’S, 138°15'°E, CWO. FORMATION: Lower Upper Nora Fm. AGE: Upper Middle Ordovician. REMARKS: Half of F7200 is missing. Georgina linda Wade, 1977a F7175, F7178-F7185 PARATYPES Georgina linda Wade, 1977a, p.10; F7178, pls fig. 8; F7185, pI.S, fig.s. LOCALITY: W side of small hill in Lower Nora Formation, 23°19'S, 138°06°E, CWQ. FORMATION: Lower Nora Fm. AGE: Middle Ordovician. F7176 HoLotyre Gvorgina linda Wade, 1977a, p.10, pl.5, figs 1-3; ext- fig.dg, LOCALITY: 4.5 miles NW of 26 Bore, Glenormis- ton Stn, 22°47°S, 138°10°E, CWQ, FORMATION: Lower Nora Fm- MEMOIRS OF THE QUEENSLAND MUSEUM AGE: Middle Ordovician, F7177 PARATYPE Georgina linda Wade, 1977a, p.10, pl.5, fig.4. LOCALITY, FORMATION, AGE: As for holotype. F7186 PARATYPE Georgina linda Wade, 1977a, p.10. LOCALITY: Nora Fm, Adjacent to road at western end of Toko Range 22°43'S, 137°43’E, NT. FORMATION, AGE: As for holotype. REMARKS: specimen missing. Georgina taylori Wade, 1977a F7148 HOLOTYPE Georgina taylori Wade, 1977a, pp.4,8-9, pl.3, figs 1-4, text-fig.60. LOcALITy: Near Halfway Dam, Tobermorey Stn, 22°53'S, 137°50’E, NT. FORMATION: Coolibah Fm. AGE; Lower Middle Ordovician. F7090-F7152 PARATYPES Georgina taylori Wade 1977a, pp.4,8-9; F7091, pl.4, fig.5; F7101, pl.3, fig.5; F7106, pl.4, fig.6; F7111, pl4, figs 1,2; F7121, pl.4, fig.3; F7130, pl.4, fig.4; F7134, pl. 2, fig.7; F7139, pl.3, fig.6; ©7149, pl.4, fig.7. LOCALITY, FORMATION, AGE: As for holotype. F7153 PARATYPE Georgina taylori, Wade, 1977a, pp.4,8-9. LOCALITY: Pulatera Hills, Glenormiston Stn, 22°57'S, 138°21°E, CWO. FORMATION, AGE: As for holotype. F7154-F7155 PARATYPES Georgina taylori Wade, 1977a, pp.4,8-9. Locauity: 3kms S of Eurithethera Soak and 4-Skms WSW of first gap in scarp. Toomba Range, CWQ. FORMATION, AGE: As for holotype. REMARKS: F7114, F7122, and P7132 and por- tions of F7096, F7117, F7130 and F7111 are missing. P7091 and F7135 were figured in Wade (1977b). F7096 and F7109 are part of the same individual, and referred to one number, F7096. Glauconome sp. P1203 Glauconome sp.: Etheridge fil., 1892, p.223, pl.44, fig. 11. INVERTEBRATE FOSSIL TYPES LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Carboniferous. Glyphea oculata Woods, 1957 F3233 HoLotyPe Glyphea oculata Woods, 1957, pp.162-3, pl.4, fig.4, text figs4a,b. LOCALITY: ‘Currane’, 10 miles N of Dartmouth, cQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaccous, (Albian). Goniastrea aspera Verrtill, 1905 F12401 Goniastrea aspera; Pickett et al., 1985, pp.103-14. LOCALITY: S of Amity Point, N Stradbroke Is- land (Brisbane 1;100,000 Sheet, grid reference 441658). FORMATION: Unnamed marine sediments. AGE: Pleistocene. REMARKS: See Porites sp. Grammatodon (Indogrammatodon) robusta (Etheridge, 1872). F1242 HOLOTYPE Cucullaea robusta Etheridge, 1872, p.340, pl.20, fig. 1. Cucullaea robusta Etheridge; Etheridge fil., 1892, p.565. Idonearca robusta Etheridge; Etheridge fil., 1902a, p.68, Grammatodon (Indogrammatodon) robusta (Etheridge, 1872) Fleming 1966b, pp.13-6, pl.5, figs3a,b. Grammatodon (Indogrammatodon) robusta (Etheridge Snr., 1872); Hill et al., 1968, pl.K6, fig, 12. LocALitry; Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Daintree Collection. F1243 HOLOTYPE Cxcullaea costata Etheridge, 1872, p.340, pl.20, fig. 2. Cucullaea robusta Etheridge; Etheridge fil., 1892, p.565, Idonearca rabusta Etheridge; Etheridge fil., 1902a, p.68, Grammatodon (Indogrammatodon) robusta 683 (Etheridge. 1872) Fleming, 1966b, pp.13-6, pl.5, figs4a,b,c. LOCALITY; Maryborough, SEQ, FORMATION; Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Daintree Collection F5472 Grammatodon (Indogrammatodon) robuste (Etheridge, 1872) Fleming 1966b, pp.13-6, pl.6, figs3a,b,c,d,e,f. LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). Gyaloceras smithi Whitchouse, 1927 F304 HoLotyPe Gyaloceras smithi Whitehouse, 1927, pp.114-5, pl.17, fig.1, text fig.8, Gyaloceras smitht Whitehouse, 1927; Arkell et al., 1957, p. L286, fig.338-5. LOCALITY: Walsh River, NQ. FORMATION: Blackdown Fm. AGE: Lower Cretaceous, (Aptian). Gyroceras dubius Etheridge fil., 1892 F1231 HOLOTYPE Gyroceras dubius Etheridge fil., 1892, p.294, pl.4l, fig.12. LOCALITY: Rockhampton District, MEQ, FORMATION: ?Malchi Fm. AGE: Lower Carboniferous Hamites aff. H. maximus J. Sowerby F1601 Hamites alt, maximus J. Sowerby; Whitehouse, 1926, p.226, pl. 39, figs 2a,b. Locatiry: Ward River, head of Warrego River, sca. FORMATION: ? AGE: Lower Cretaceous. REMARKS: McNamara (1980) considered that F1601 was not referable to Hamites based upon the presence of a trifid lateral lobe. Hemiptera incertae sedis F6540a/b Hemiptera incertae sedis; Evans, 1971, p.149, fig.4d. LOCALITY: Mount Crosby, SEQ, Upper Bed, 910805-911805, Ipswich 1 mile military map. FORMATION: MI Crosby Fm. 684 AGE: Early Late Triassic, (Carnian). F6483a/b Hemiptera incertae sedis; Evans, 1971, p.149, fig.4e. LOCALITY, FORMATION, AGE: As for F6540. Hercophyllum shearsbyi Siissmilch, 1914 F2479 Hercophyllum shearsbyi (Sussmilch) Jones, 1936, pp.53-S, pl.S, figs la-g. Hercophyllum shearsbyi Siissmilch, 1914: Hill, 1981, p-F252, figs 160a-e. LOCALITY: Hatton’s Corner, Yass, NSW. FORMATION: ? AGE: Upper Silurian. REMARKS: This species was introduced as Cyathophyllum shearsbyi by Etheridge (1904, p.288), mentioned by Harper (1909, pp.39-43) and figured by Siissmilch (1914, fig.143); but it was not fully described by Jones until 1936. F2479 consists of a small remaining fragment, and the thin sections, which are figs la-g in Jones (1936). Heterochterus timmsii Evans, 1971 F6473 HouotyPe Heterochterus timmsii Evans, 1971, p.149- 50, fig.5. Loca.ity: Mt Crosby, SEQ. FORMATION: Mt Crosby Fm. AGE: Early Late Triassic, (Carnian). Heterojassus membranaceus Evans, 1961 F3701 HOLOTYPE Heterojassus membranaceus Evans, 1961, p.23, fig.Sb. LOCALITY: Mt Crosby Insect Beds, SEQ. FORMATION: Mt Crosby Fm. AGE: Early Late Triassic, (Carnian). Heteronella marksei Evans, 1961 F3699a/b HootyPe /feteronella marksei Evans, 1961, p.22, fig.Sa. LOCALITY: Mt Crosby Insect Bed, SEQ. FORMATION: Mt Crosby Fm. AGE: Early Late Triassic, (Carnian). REMARKS: The counterpart of this specimen originally referred to as F3700b has been re- registered as F3699b. MEMOIRS OF THE QUEENSLAND MUSEUM Hexacrinites interscapularis Phillips, 1841 F14580 Hexacrinites interscapularis Phillips, 1841; Jell et al., 1988, pp.368-70, fig.11a. LOcALity: UQLS305S Wando Vale topographic sheet [569404] south-westerly oriented gully 400m south of the volcano (not older than varcus biozone) Wando Vale Stn, NQ. FORMATION: Papilio Mudstone Fm. AGE: Middle Devonian, (late Eifelian-Givetian). REMARKS: The formation was formally named by Lang et al., (1989) as Papilio Mudstone. F14594, F14595 Hexacrinites interscapularis Phillips, 1841; Jell et al., 1988, pp.368-70, F14594 figs] le-h. LOCALITY: UQLS252 Wando Vale topographic sheet [551366] south flank of low hill, 1km south of Storm Dam Creek. (?varcus biozone) Wando Vale Station, 150km NW of Charters Towers, NQ. FORMATION, AGE, REMARKS: As for F14580. F14597 Hexacrinites interscapularis Phillips, 1841; Jell et al., 1988, pp.368-70. LOCALITY: UQL5318, Wando Vale topographic Sheet [565395] 65-116m above base of forma- tion in gully 2.2km NE of Storm Dam, Wando Vale Station, 150km NW of Charters Towers NQ. FORMATION: Papilio Mudstone. AGE: Middle Devonian, ( Givetian). REMARKS: As for F14580. F14596, F14600, F14602 Hexacrinites interscapularis Phillips, 1841; in Jell et al., 1988, pp.368-70. LOCALITY: UQL5356, Wando Vale topographic Sheet (554367) east flank of Spongophyllum Hill, in head of eroding gully; (varcus biozone) Wando Vale Station, 1S0km NW of Charters Towers, NQ. FORMATION: Papilio Mudstone. AGE: Middle Devonian, (Givetian). REMARKS: As for F14580. F14604 Hexacrinites interscapularis Phillips, 1841; Jell et al., 1988, pp.368-70. LocALiTy: UQL5335, Wando Vale topographic Sheet [620418] along northern tributary of Lomandra Creek, (ensensis zone), Wando Vale Station, 150km NW of Charters Towers, NQ. INVERTEBRATE FOSSII. TYPES FORMATION: Papilio Mudstone. AGE: Middle Devonian, (Givetian), REMARKS: As for F14580. Locality information for this site was omitted by Jell et a/. (1988). F14743, F14745, F14834, F14871, F14874 Hexacrinites interscupularis Phillips, 1841; Jell etal. 1988, pp.368-70. LOCALITY: UQLS5321, Wando Vale topographic Sheet [$62390] creck section 3km NE of Storm Dam, Wando Vale Stn, 150km NW of Charters Towers, NO. FORMATION, AGE, REMARKS: As for F14580. F14755, F14756, F14763 Hexacrinites interseapularts Phillips, 1841; Jetl eral., 1988, pp.368-70, Locality: UQL5320 Wando Vale topographic Sheet [563392] section on ridge 2.9km NE of Storm Dam, Wando Vale Stn, 150km NW of Charters Towers, NQ, FORMATION, AGE, REMARKS: As for F14580. F14843, F14845 Hexacrinites interscapularis Phillips, 1841; Jell etal., 1988, pp.368-70. LOcALity: UQL5218, Wando Vale topographic Sheet [559389] ¢. 68m above base of formation in gully on cast slope of Storm Hill, 1.2km N of Storm Dam, Wando Vale Stn, 150km NW of Charters Towers, NQ. FORMATION; Papilio Mudstone. AGE: Middle Devonian, ( ?late Eifelian). REMARKS: As for F14580. F14849 Hexacrinites interscapularis Phillips, 1841; Jell et al., 1988, pp.368-70. LOCALITY: UQL5267, Wando Vale topographic Sheet [543363] in gully, 1.S5km SW of Storm Dam, Wando Vale Stn, 150km NW of Charters Towers, NQ. FORMATION; Papilio Mudstone. AGE: Middle Devonian, (Givetian). REMARKS: As for F 14580. Hexacrinites spinosus Muller, 1856 F14746 Hexacrinites spinosus Miller 1856; Jell e¢ al, 1988, pp.370-2, figs 12d-f. LOcALITy: UQL5277 Wando Vale topographic Sheet [570409] 51-S6m above base of section 300m, ESE of The Volcano. Wand Vale Stn, 150km NW of Charters Towers, Nv. FORMATION: Papilio Mudstone. AGE: Middle Devonian, (probably early Givetian). REMARKS: This formation was formally named by Lang er al. (1989) as Papilio Mudstone. Homolopsis etheridgei (Woodward, 1892) F2796 Homolopsis etheridgei(H. Woodward); Woods, 1953, pp.50-2, pl. 2, fig.2. Homolopsis etheridgei (Woodward, 1892); Hill et al., 1968, pl. K11, fig.8, Homolapsis etheridgei(Woodward, 1892); Glaessner, 1980, pp, 173-4. LOCALITY: 10 miles N of Dartmouth, CQ. FORMATION: Allaru Mudstone, AGE: Lower Cretaceous, (Albian). F2843 Hamolapsis etheridge (H, Woodward); Woads, 1953, pp.50-2, fig. 1a. Homolopsis etheridgei (Woodward) 1892; Glaessner, 1980, pp. 173-4. LOCALITY, FORMATION, AGE; As for F2796, F2845 Hamolopsis etheridge (AH. Woodward); Woods, 1953, pp.50-2. Hamolupsis etheridge (Woodward, 1892); Glaessner, 1980, pp. 173-4. LOCALITY, FORMATION, AGE: As for F2796. F2846 Homalapsis etheridgei (H. Woodward); Woods, 1953, pp.s0-2. Homolopsis etheridgei (Woodward, 1892); Glaessner, 1980, pp. 173-4. LOCALITY, FORMATION, AGE: As for F2796. F2847 Homolapsis etheridget (HA. Woodward); Woods, 1953, pp.50-2, pl. 2, fig.3. Hamoloupsis etheridge? (Woodward, 1892); Glaessner, 1980, pp, 173-4, LOCALITY, FORMATION, AGE: As for F2796. F2848 Homolopsis etheridgei (H. Woodward); Woods, 1953, pp.s0-2. Homolopsiy etheridgei (Woodward, 1892); Glaessner, 1980, pp. 173-4. 686 LaOcaLiTy. FORMATION, AGE: As for F2796. F2875 HOLOTYPE Prosopon etheridgei Woodward, 1892, p.301, pl.4. Prosopon etheridgei H. Woodward; Etheridge fil., 1917, pp.5-7, pl.1, figs 1-4. Homolopsis etheridgei (H, Woodward); Woods, 1953. pp.50-2, pl.2, fig.1, Hoamolopsis etheridgei (Woodward, 1892); Glaessner 1980, pp. 173-4, pl.1, fig.2. LOCALITY: CQ. FORMATION: Probably Allaru Mudstone. AGE: Lower Cretaceous, (Albian). Homoptera incertae sedis F649 1a/b Homoptera incertae sedis; Evans, 1971, p.149, fig.4c. LocaLity: Mt Crosby, SEQ, Upper Bed, 910805- 911805, Ipswich 1 mile military map. FORMATION: Mt Crosby Fm. AGE; Early Late Triassic, (Carnian). Hoploparia mesembria Etheridge fil., 1917 F2908 Haploparia mesembria Etheridge Jr., Woods, 1957, pp. 169-71, pl.6, fig.4. LOCALITY: ‘Currane’, 10 miles N of Dartmouth, ca. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). F3239 Hoploparia mesemhria Etheridge Ir.; Woods, 1957, pp. 169-71. LOCALITY, FORMATION, AGE: As for F2908, F3240 Hoploparia mesembria Etheridge Jr., 1917, in Woods, 1957, pp. 169-71. LOCALITY,FORMATION, AGE: As for F2908. F3241 Hoplaparia mesembria Etheridge Jr; Woods, 1957, pp. 169-71. LOCALITY,FORMATION, AGE: As for F2908. F3243 Hoploparia mesembria Etheridge Jr.; Woods, 1957, pp. 169-71, pl.6, fig.2. Hoploparia mesembria Etheridge, 1917; Hill et al., 1968, pl. Ki 1, fig.3. MEMOIRS OF THE QUEENSLAND MUSEUM LOCALITY, FORMATION, AGE: As for F2908. F3244 Hoploparia mesembria Etheridge Jr. Woods, 1957, pp. 169-71, pl.6, fig.3. LOCALITY, FORMATION, AGE: As for F2908. Hylicella colorata Evans, 1956 F3686 Hylicella colorata Evans, 1956; Evans, 1961, p.15, fig. 1d. LOCALITY: Mt Crosby Insect Beds, SEQ. FORMATION: ML Crosby Fm. AGE: Early Late Triassic, (Carnian). Indeterminate heteropteron forewing F3702 Heteropteron wing; Evans, 1961, p.23, fig.Sc. LOCALITY; Mt Crosby Insects Beds, SEQ. FORMATION: Mt Crosby Fm. AGE: Early Late Triassic, (Carnian). Ingelarella strzeleckii (de Koninck, 1877) F946 Spirifer undifera var. undulata. F. Rom.; Etheridge, 1872, p. 330, pl.16, fig.3. Spirifer strzeleckii De Koninck; Etheridge fil., 1892, p.234, pl.10, fig.6. Ambikella strzeleckii (de Koninck) 1876; Ruonegar and Ferguson,, 1969, p.251. Lectotyre /ngelarella strzeleckii (de Koninck); Mc- Clung, 1978, p.43, pl.3, fig.25. LOCALITY: Lady Mary Reef, Gympie, SEQ. FORMATION: Rammutt Fm. AGE: Permian. REMARKS: Previously registered as F66 in Aplin’s 1871 Fossil Register. F3272 Spirifer undifera vat, undulata P. Rém.; Etheridge, 1872, p. 331, pl.16, fig.4. Spirifer strzeleckii De Koninck; Etheridge fil., 1892, p- 234, Ambikella strzeleckii (de Koninck) 1876; Runnegar and Ferguson, 1969, p,251. Ingelarella strzeleckii (de Koninck, 1877); McClung, 1978, p. 43. Locatity: Gympie, SEQ. FORMATION: Rammutt Fm. AGE: Permian INVERTEBRATE FOSSIL TYPES F3304 Spirifera undifera var. undulata F. Rém.; Etheridge, 1872, p. 330-3, pl.16, fig.5. Spirifera strzeleckii de Koninck; Etheridge fil., 1892, p.234. Ambikella_ strzeleckii (de Koninck) 1876; Runnegar and Ferguson, 1969, p.251. Ingelarella strzeleckii (de Koninck, 1877); McClung, 1978, p. 43. LOcALiTy: Lady Mary Reef, Gympie, SEQ. FORMATION: Rammutt Fm. AGE: Permian. REMARKS: Old No. F13A Aplin’s Register (1871). F5774 Ingelarella strzeleckii (de Koninck); McClung, 1978, p.43, pl. 3, fig.23. LOCALITY: Gympie, SEQ. FORMATION: Rammutt Fm. AGE: Permian. REMARKS: Previously registered as F37 in Aplins’ (1871) Register. Inoceramus carsoni McCoy, 1865 F1238 HOLOTYPE /noceramus pernoides Etheridge, 1872, pp.343-4, pl. 22, fig.3. Inoceramus pernoides Etheridge; Etheridge fil., 1892, p.464, pl.25, fig.12. Inoceramus etheridgei Etheridge fil., 1901, p.22-3. Inoceramus carsoni M’Coy, 1865, in Crame, 1985, pp.498-501. LOCALITY: Marathon Stn, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaccous, (Albian). REMARKS: Daintree Collection. F15642 Inoceramus sp. cf. I. pernoides Etheridge; Etheridge fil., 1892, p.464, pl.42, fig.7. Inoceramus carsoni McCoy, 1865; Crame, 1985, pp.498-S01. LOCALITY: Bowen Downs, cast of Muttaburra, CQ. FORMATION: ?Allaru Mudstone. AGE: Lower Cretaceous, (Albian). REMARKS: Daintree Collection Inoceramus elongatus Etheridge, 1872 F1240 Inoceramus multiplicatus Stol. var. elongatus, Etheridge, 1872, p.343, pl.22, fig.2 687 HOLOTYPE /noceramus elongatus (Etheridge) Etheridge fil., 1892, p.464. Inoceramus elongatus Etheridge; Etheridge fil., 1902a, p.70. LOCALITY: Marathon Stn, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). REMARKS: The concentric ribs and elongate form of this shell suggest affinities with J. car- soni group (sensu Crame, 1985). Daintree Col- lection. Inoceramus marathonensis Etheridge, 1872 F1239 HOLOTYPE /noceramus marathonensis Etheridge, 1872, p.343, pl. 22, fig.1. Inoceramus marathonensis Etheridge; Etheridge fil., 1892, p. 464, Inoceramus marathonensis Etheridge; Etheridge fil., 1902a, p. 70. LOCALITY: Marathon Stn, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). REMARKS: Daintree Collection. Inoceramus sp. cf. 1. sutherlandi McCoy, 1865 F1241 Inoceramus allied to I. problematicus, D’Orb.; Etheridge, 1872, p.344, pl.22, fig.4. Inoceramus carsoni McCoy in Etheridge fil., 1892, p.463. Inoceramus etheridgei Etheridge fil., 1901, p.22-3. Inoceramus cf. sutherlandi, McCoy, 1865; Crame, 1985, pp.S01- 2. LOCALITY: Marathon Stn, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaccous, (Albian). REMARKS: Daintree Collection. Labeceras (Labeceras) Whitehouse, 1926 compressum F1600 HOLoTYPE Labeceras compressum Whitehouse, 1926, p.228, pl.36, fig.5, pl.39, figs Sa,b. Labeceras (Labeceras) compressum Whitehouse; Reyment, 1964, p. 24. LOCALITY: Tower Hill, Muttaburra, CQ. FORMATION: ?Mackunda Fm AGE: Lower Cretaceous. 688 Labeceras sp. cf. L. compressum Whitehouse, 1926 F6096 Labeceras cf. compressum Whitehouse, 1926; Hill, et al., 1968, pl.K7, fig.5. LOcALITY: Currane Station, 16km N of Dartmouth, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). Labeceras (Labeceras) laqueum (Etheridge fil., 1892) F2148 Crioceras taylori, Eth. fil.; Etheridge fil., 1909, pp.162-3. LOcALityY: ?Head of Walsh River, NQ. FORMATION: ?Wallumbilla Fm. AGE: Lower Cretaceous. REMARKS: F2148 bears a label saying “Head of the Walsh River’, which suggests that it may be the Queensland Museum specimen referred to by Etheridge fil. (1909, p.162). However the lithol- ogy is not typical of the Walsh River material and this specimen, bears donor number D222 which suggests a McKinlay’s Range origin. See also Beudanticeras sutherlandi (F1273). F17743 HOLOTYPE Hamites? laqueus Etheridge fil., 1892, pp.496-7, pl. 42, figs 14, 15. Labeceras laqueus (Etheridge fil.) Whitehouse, 1926, pp.227-8. Labeceras (Labeceras) laqueum (Etheridge); Rey- ment, 1964, p. 22. LacaLity: Tower Hill, Landsborough Creek, a head tributary of the Thomson River, N of Mut- taburra, CQ. FORMATION: Mackunda Fm. AGE: Lower Cretaceous. REMARKS; The phragmocone and part of the living chamber of this specimen is missing but a plastoholotype (L598) is held in the Australian Museum (Fletcher, 1971), The figure in Etheridge fil. (1892, pl.42, fig.14) is laterally reversed. The other Queensland Museum specimen mentioned by Etheridge fil. (1892) has not been identified. Lasiocladia? hindei Etheridge fil., 1892 F5706 Hocoryre Lasivcladia? hindei Etheridge fil., 1892, p.199, pl.4i. figs 1.2. MEMOIRS OF THE QUEENSLAND MUSEUM Lasiocladia? hindei Eth. f.; Pickett, 1969, pp.9-10, pl.1, fig.3. Lasiocladia? hindei in Pickett, 1983, p.112. LOCALITY: Rockhampton, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. Pickett (1969) discussed possible relationships of this sponge. Leionucula quadrata (Etheridge, 1872) F1244 SYNTYPE Nucula gigantea Etheridge, 1872, p.341, pl.20, fig. 4, Nucula gigantea Etheridge; Etheridge fil., 1892, p.566. Nucula gigantea Etheridge; Etheridge fil., 1902a, p.74. Leionucula quadrata (Etheridge, 1872) Fleming 1966a, pp.5-8, pl. 2, figsla,b. LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Two specimens were illustrated by Etheridge (1872, pl.20, fig.4). Fleming (1966a) considered that the species description of N. gigantea would only fit the lower of the two specimens figured. He synonymised both specimens with L. quadrata, Daintree Collection. F1245 SYNTYPE Nucula quadrata Etheridge, 1872, p.341, pl.20, fig.3. Nucula quadrata Etheridge: Etheridge fil., 1892, pp.565-6. Nucula quadrata Etheridge; Etheridge fil., 1902a, p.74, Nucula quadrata Etheridge; Etheridge fil., 1902b, pp.23-4. LecTOTYPE Leienucula quadrata (Etheridge, 1872) Fleming 1966a, pp.6-4, pl. 2, figs2a,b. Locatity: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Daintree Collection, F1246 SYNTYPE Nucula quadrata Ethendge 1872, p.341, pl.19, fig.5. Nucula quadrata Etheridge, Etheridge fil., 1892, pp.565-6. Nucula quadrata Etheridge, Etheridge fil., 1902a, p.74. Nucula quadrata Etheridge, Etheridge fil,, 1902b, pp.23-4. INVERTEBRATE FOSSIL TYPES PARALECTOTYPE Leionucula quadrata (Etheridge, 1872) Fleming 1966a, pp.6-8, pl. 2, fig.3. LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE; Lower Cretaceous, (Aptian). REMARKS: Etheridge’s (1872) figure of F1246 is slightly smaller than natural size. Daintree Col- lection. Lepidoptera leaf mines F15346 Lepidoptera leaf mines; Rozefelds, 1988, pp.77-81, fig.2. LOCALITY: Clack Island, Princess Charlotte Bay, NQ. FORMATION; Battle Camp Fm. AGE: Upper Jurassic-Lower Cretaceous. Leucosia pubescens Miers, 1877. F2184 Leucosia pubescens Miers, 1877; Hill et al,, 1970, pl.Cz6, fig.6, LocaLity: Cleveland Bay, near Townsville, NQ. FORMATION: Unnamed estuarine deposit. AGE: Pleistocene/Holocene. Lithosmylidia baronne Lambkin, 1988 F14358 HoLotyre Lithosmylidia baronne Lambkin, 1988, p.447, fig.3. LOCALITY: road cutting on Baroone Rd, c. 3km ENE Gayndah, SEQ. FORMATION: Gayndah Beds. AGE: Middle Triassic. Lithosmylidia parvula Riek, 1955 F14359 Lithosmylidia parvula Riek; Lambkin, 1988, pp.447- 8, fig.4. LOCALITY: Mount Crosby Insect locality B, Mt Crosby, SEQ, FORMATION: Mt Crosby Fm. AGE: Early Late Triassic, (Carnian). REMARKS: Lambkin (1988, fig.4) incorrectly refers to this specimen as F1459, Lopha marshii australiensis Skwarko, 1974 F5594 689 Lopha marshii (J. Sowerby, 1812) australiensis Skwarko, 1974, pp. 89-91, pl.30, fig.6. LOCALITY: ?Newmarracarra Limestone, WA, FORMATION: ?Newmarracarra Limestone. AGE: Middle Jurassic, (Bajocian). REMARKS: This specimen was erroneously recorded in Skwarko (1974, p.90) as part of the GSWA Collection. Loxonema sp. F1225 (Fig.2G) Loxonema sp.; Etheridge fil., 1894, pp.536-7, pl.39, fig.7. LOCALITY: Rockhampton District, MEQ. FORMATION; ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. The specimen was poorly illustrated. It is refigured herein (Fig. 2D). F1226 Loxonema sp.; Etheridge fil., 1894, pp.536-7, pl.40, fig.6. LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. Luciella (?) grayae Etheridge fil., 1892 F1227 (Fig.2A) HOLotyPe Luciella (7) grayae Etheridge fil., 1892, pp.288-9, pl.41, fig.6. LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection, Etheridge’s figure of L. grayae bears little resemblance to the type specimen. Maccoyella alata (Etheridge, 1872) F1235 HOLOTYPE Avicula alata Etheridge, 1872, p.342, pl.20, fig.8. Pseudavicula ? alata (Etheridge); Etheridge fil., 1892, p.563, pl.24, fig.14. Maccoyella barklyt Moore; Etheridge fil., 1902a, pp.17-8. Maccoyella barklyi Moore; Etheridge fil., 1902b, pp. 11-2, Maccoyella alata (Etheridge, 1872); Fleming, 1970, pp.4-S. LOCALITY: Maryborough, SEQ, FORMATION: Maryborough Fm, 690 AGE: Lower Cretaceous, (Aptian). REMARKS: Daintree Collection. Maccoyella barklyi (Moore, 1870) F1236 HoLotyee Streptorhynchus davidsoni Etheridge, 1872, p.333, pl. 17, fig.1. Maccoyella barklyi Moore; Etheridge fil, 1892, p.455. Maccoyella barklyi (Moore), 1870: Day 1967, pp.21- 2 LOCALITY: ?Walsh River, NQ. FORMATION: Blackdown Fm. AGE: Lower Cretaccous, (Aptian). REMARKS; Etheridge (1872) considered this Daintree specimen to be derived from the Car- boniferous of the Peak Downs and Bowen River area. Etheridge fil., (1892) considered this locality was in crror and he referred it to the Cretaceous species Maccoyella reflecta or M. barklyi trom either the Walsh River, NQ, or Wallumbilla, SEQ. The lithology of the specimen is similarto other Walsh River material in the collection. Maccoyella corbiensis (Moore, 1870) F1237 HOLOTYPE Crenutula(?) gibbasa Etheridge, 1872, p.339, pl.lY. fig.3. Maccoyella corbicnsis Moore; Etheridge fil., 1892, pp.563-4. Maccoyella corbicnsis Moore; Etheridge fil., 1902a, p.2l. Maccoyella corbiensiy Moore; Etheridge fil., 1902b, p.l3, LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE; Lower Cretaceous, (Aptian), REMARKS; Duintree Collection, Macrocallista (?) plana Moore, 1870 F3850 Myacites sp. Etheridge, 1872, p.348, pl.25_ fig.7. Maerocallista (?) plana, Moore; Etheridge fil, 19024, pp.37-8. Locauity: Gordon Downs at the head of Roper Creek, CQ. FORMATION: ? Blenheim Sub Group. AGE: Permian REMARKS: See Collabrina cliftoni. Daintree Collection, Collected by Rev. W.B. Clarke. As MEMOIRS OF THE QUEENSLAND MUSEUM this material is Permian in age, assignment to a Cretaceous species 1s almost certainly incorrect. Macrophthalmus latreillei (Desmarest, 1822) F6557 Macrophthalmus latreillei (Desmarest, 1822); Hill ef al., 1970, pl.Cz6, fig.7. LocaLity: Moffat Head, Caloundra, SEQ. FORMATION: Unnamed estuarine deposit. AGE: Pleistocene/Holocene- Maltchiblastus australis (Etheridge fil., 1892) F1193 HOLOTYPE Mesoblastus ” australis Etheridge fil., 1892, pp.210-1, pl.44, fig.2. Malchiblastus australis (Etheridge fil.,) McKellar 1966, pp.191- 8, pl.24, fig.4. LOCALITY: Rockhampton District, MEQ. FORMATION: Neerkol Fm. AGE: Upper Carboniferous. REMARKS: Specimen is missing, FII95 HOLOTYPE Trievelocrinus ? carpenteri Etheridge fil., 1892, pp. 212-3, pl.44, fig.3. Matchiblastus australis (Etheridge fil..) McKellar, 1966, pp. 191-8, pl.24, figs 2-3,6. LOcALITY: Rockhampton District, MEQ. FORMATION: Neerkol Fm, AGE: Upper Carboniferous. Martinia (vel. (Sowerby, 1844) Martiniopsis) subradiata F948 Martinia (vel Martiniopsis) subradiata G.B, Sby., sp.; Etheridge fil., 1892, pl.43, fig.1. LOCALITY: Banana, CQ. FORMATION: ?Flat Top Formation. AGE: Permian. Meleagrinella sp. F5782a,b Meleagrinella sp.; Hill et al., 1968, pl.K6, figsl7a,b. Meleagrinella sp.. Fleming 1970, pp.5-6, pl.1, figs9- 10. LOCALITY: Ridge in the Gundiah |-mile sheet area, grid reference 675895. Bauple Mtns, SEQ. FORMATION: Maryborough Fm. AGE: Lawer Cretaceous, (Neocomian). INVERTEBRATE FOSSIL TYPES F5783 Meleagrinella sp., Fleming, 1970, pp.5-6, pl.1, fig.11. LOCALITY, FORMATION, AGE: As for F5782, F5784 Meleagrinella sp., Fleming, 1970, pp.5-6, pl. 1, fig.12. LOCALITY, FORMATION, AGE: As for F5782. F5785 Meleagrinella sp.; Hill et al., 1968, pl.K6, fig.16, Meleagrinella sp.; Fleming, 1970, pp.5-6, pl.1, fig.8. LOCALITY, FORMATION, AGE: As for P5782. Melocrinites tempestus Jeli et al., 1988. F14884 HOLOTYPE Melocrinites tempestus Jell et al., 1988, pp.372-4, figs 13d-g. LOCALITY: UQLS218 Wando Vale topographic Sheet [559389] c. 68m above base of formation in gully on Eslope of Storm Hill, 1.2km north of Storm Dam, Wando Vale Stn, 150km NW of Charters Towers, NQ. FORMATION: Papilio Mudstone. AGE: Middle Devonian, (?late Eifclian). REMARKS; The formation was formally named by Lang ef al. (1989) as Papilio Mudstone. F14853-F 14854 PARATYPE Melocrinites tempestus Jell et al., 1988, pp.372-4; F14854, fig. 13h. Locality: UQL5318/69 Wando Vale topographic Sheet [565395] 65-116m above base of formation in gully 2.2km NE of Storm Dam, Wando Vale Stn, 150km NW of Charters Towers, NQ. FORMATION: Papilio Mudstone. AGE: Middle Devonian, (Givetian). REMARKS: As for F14580, Mesaktoceras arachne Wadc, 1977a F7187 HoOLotyPe Mesaktoceras arachne Wade, 1977a, pp.4,11-14, pl.7, tigs 1-3, text-fig.7g. LOcALITy: 3km E of Halfway Dam, Tobermorey Stn, 22°53'S, 137°50°E, NT. FORMATION: Middle Nora Fm. AGE: Middle Ordovician. F7188, F7190-F7193 PARATYPES Mesaktoceras arachne Wade, 1977a, pp.4.11-14; F7188, pl.7, fig.4; F7190, pl.7, fig.S. LOCALITY, FORMATION, AGE: As for holotype. 69) F7189 PARATYPE Mesaktoceras arachne Wade, 1977a, pp.4, 11-14, pl.7, fig.7. LOCALITY: km E of Halfway Dam, Tobermorey Stn, 22°53’S, 137°S0’E, NT. FORMATION, AGE: As for holotype. F7194 PARATYPE Mesaktoceras arachne Wade, 1977a, pp.4, 11-14. LOCALITY: W side of small hill in Lower Nora Fm, 23°19’S, 138°06’E, CWQ. FORMATION, AGE: As for holotype. F7195 PARATYPE Mesaktoceras arachne Wade, 1977a, pp.4, 1I-14, LOCALITY: Valley between Toko Range scarp and QML319, 23°19°S, 138°06’E, CWQ, FORMATION, AGE: As for holotype. REMARKS; F7193 and F7195 are missing. Wade's (1977a) pl.7, fig.5 is referred to F7193, the correct number is F7190. Wade (1977a, pl.7) incorrectly records the locality for the holotype and paratypes F7188, F7190-F7193 as. 1lkm rather than 3km E of Halfway Dam. Mesocicadella punctata Evans, 1961 F368 a/b HOLOTYPE Mesocicadella punciata Evans, 1961, p,14, fig.la. LOCALITY: Mt Crosby Insect Beds, SEQ. FORMATION: Mt Crosby Fm. . AGE: Early Late Triassic, (Carnian). Mesothymbrts perkinsi Evans, 1956 F3682a/b Mesothymbris perkinst Evans, 1956; Evans, 1961, p.I5, fig. 1c. LOCALITY: Mt Crosby Insect Beds, SEQ. FORMATION: Mt Crosby Fm. AGE: Early Late Triassic, (Carnian). REMARKS: The counterpart of this specimen originally referred to as F3683 in Evans (1961) has been re-registered as F3682b. F3684a/b Mesothymbris perkinsi Evans, 1956; Evans, 1961, p.15, fig. 1b. LOCALITY, FORMATION, AGE: As for F3682a/b. REMARKS: The counterpart of this specimen originally referred to as F3685 in Evans (1961) has been re-registered as F3684b. Myloceras ammonoides (Etheridge fil., 1909) F1388 HoLotyPe Myloceras davidi Whitehouse, 1926, p.235, pl.37, figs 2a,b,c. Myloceras davidi Whitehouse; Reyment, 1964, p.30. Myloceras ammonoides (Etheridge, 1909); Mc- Namara, 1978, pp.231-7. LOCALITY: Bowen Downs, Thomson River, CQ. FORMATION, AGE: Lower Cretaceous, (Albian). F2230 Myloceras ammonoaides (Etheridge, 1909); Mc- Namara, 1978, pp. 231-7. LOCALITY: Rodney Downs, Aramac, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). F9262 Myloceras ammonoides (Etheridge, 1909); Mc- Namara, 1978, pp. 231-7, fig.4b, LOcALITY:Currane Stn,SkmN of Dartmouth, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). F9352 Myloceras ammonoides (Etheridge, 1909); Mc- Namara, 1978, pp. 231-7, Fig.4a. LOCALITY, FORMATION, AGE: As for F9262. F9396 Myloceras ammonoides (Etheridge, 1909); Mc- Namara, 1978, pp. 231-7, fig.2. LOCALITY, FORMATION, AGE: As for F9262. Myloceras auritulum McNamara, 1978 F9474 HOLOTYPE Mylaceras auritulum McNamara, 1978, pp.231,234-5, 237- 40, figs 4d,5a,7,9a. LOCALITY: [soray Station, 15.Skm SW of Tambo, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). F9475 PARATYPE Myloceras auritulum McNamara, 1978, pp.231,234-5, 237- 40), figs. 6a,b,9b. LOCALITY,FORMATION, AGE: As for F9474. Myloceras baccatum (Whitehouse, 1926) F1262 Crioceras Jlindersi McCoy, sp.; Etheridge fil., 1909, pp.153- 7, pl.40, figs. 3,4. MEMOIRS OF THE QUEENSLAND MUSEUM Flindersites aff. baccatus Whilehouse, 1926, p.237. LOCALITY: Saltern Creek, 30km W of Barcal- dine, CQ. FORMATION,AGE: Lower Cretaceous. REMARKS: Spath (1938) and Arkell er al. (1957) have referred Flindersites to Myloceras. This specimen was previously registered as D7710. See also F1268 (Australiceras irregulare). Myloceras flindersi (McCoy, 1867) F1261 Crioceras flinderst McCoy, sp.; Etheridge fil., 1909, pp.153- 7, pl.39, figs 2,3. Flindersites aft. flindersi(McCoy) Whitehouse, 1926, p.237. LOCALITY: Flinders River, NQ. FORMATION: ?Wallumbilla Fm. AGE: Lower Cretaceous, REMARKS: See F1262. F1263 Criaceras flindersi McCoy, sp.; Etheridge fil., 1909, pp. 153-7, pl.41, fig.2. LOCALITY: Queensland. FORMATION, AGE; Lower Cretaceous. REMARKS: See F1262. Myloceras intermedium Whitehouse, 1926 F1260 Crioceras flinderst McCoy, sp.; Etheridge fil., 1909, pp.153- 7, pL40, figs 1,2. HoLotyre Flindersttes intermedius Whitehouse, 1926, p.237. Myloceras intermedium (Whitehouse, 1926); Hill et al,, 1968, pl. K8, fig.3. Myloceras intermedium (Whitehouse, 1926); Mc- Namara, 1978, p. 236. LOCALITY: Mt Cornish, Aramac (neat Muttabur- ra), CQ. FORMATION: ?Allaru Mudstone. AGE: Lower Cretaceous, (Albian). F1364 Flindersites intermedius Whitehouse, 1926, p,237. Myloceras intermedium (Whitehouse, 1926); Mc- Namara, 1978, p, 236-7. LOCALITY: Longreach, CQ. FORMATION: ? AGE: Lower Cretaceous. Myloceras plectoides (Etheridge fil., 1909) F1389 INVERTEBRATE FOSSIL TYPES SYNTYPE Crioceras plectoides Etheridge fil., 1909, pp. 152-3. Aleteceras plecioides (Etheridge fil.) Whitehouse, 1926, p.232, pl.40, figs 2a,b,c, Myloceras plectoides (Etheridge, 1909); Hill et al., 1968, pl. K7, fig.6. Myloceras plectoides (Etheridge, 1909); McNamara, 1978, p.240. LOCALITY: Walsh River, NQ. FORMATION: Blackdown Fim. AGE: Lower Cretaccous, (Albian). Mytilops corrugata Etheridge fil., 1892 F1214 HOLOTYPE Mytilops corrugata Etheridge fil., 1892, p.272, pl. 40, fig.11- LOCALITY: Rockhampton, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. Nereites berneyi de Vis, 1911 F16373a HOLOTYPE Nerevites berneyi de Vis, 1911, pp.12-4, pl.3, fig.2, Locality: Wyangaria Stn, near Hughenden, CO. FORMATION: Allaru Mudstone. AGE: Lower Cretaccous, (Albian). REMARKS: Holotype is the longer trail. Nucula sp. F1210 Nucula sp.; Etheridge fil., 1892, p.274, pl.40, fig.10. LOCALITY: Rockhampton, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. Oblicarina carinata (Etheridge fil,, 1892) F1218 HOLOTYPE Chaenomya? carinata Etheridge fil., 1892, p.279, pl. 43, figs 5,6, Vacunella curvata (Morris) 1845; Runnegar, 1967, pp.63-7. Oblicarina carinata (Etheridge Jr.) Waterhouse, 1967, pp.53-7, pl.7, figs 2,3. Vacunella curvata (Morris); Runnegar, 1969, p,287, Oblicarina carinata (Etheridge, 1892); Waterhouse, 1969, p.40, pl.6, figs 4,7. Oblicarina carinata (Etheridge Jnr, 1892); Water- house, 1987a, p.175. 695 LOCALITY: Banana Creek, CQ. FORMATION: ?Flat Top Formation. AGE: Permian. REMARKS: Runnegar (1967, 1969) considered F1218.a crushed Vacunella curvata (Morris). De Vis Collection. Octomeris crassa Withers, 1932 F2026c HOLOTYPE Octomeris crassa Withers, 1932, pp.122-4, figs 1,2. Octomeris crassa Withers, 1932; Hill et al., 1970, pl.Cz6, fig.2. LOCALITY! Magnetic Island, NQ. FORMATION: Raised beach rock. AGE: Pleistocene/Holocene? REMARKS: Withers (1932) used F2026 for four different taxa. This specimem has been reregistered as. F2026c. Onestia etheridgei (Etheridge fil., 1892) F1258 Genus?: Etheridge, 1872, pp.339-40, pl.19, fig.4. HOLOTYPE Unicardium? etheridgei Etheridge fil., 1892, pp.569-70, pl.27, fig.1. Onestia etheridgei (Etheridge, 1892) Hill et al., 1968. pl. K6, figs Ta.b. Onestia etheridgei (Eth. fil. 1892); Fleming, 1970, p.7, pl.2, figs 2, 3. Onestia etheridgei (Etheridge Jnr.); Day, 1978, pp.37- 44, pl. 2, figs 3, 4. LOCALITY: Corporation Quarry (Baddow Quar- ty Area) Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Daintree Collection. Qonoton woodsi Giaessner, 1980 F2876 HOLOTYPE Qonoton woodsi Glaessner, 1980, pp.171, 173-4, pl.1, fig. 1, text figs 1a,b. LOCALITY: Currane Station, 16kms N of Dartmouth, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). Opisthotrigonia nasuta (Etheridge, 1872) F1248 SYNTYPE Trigonia nasuta Etheridge, 1872, p.339, pl.19, fig, 2a. Trigonia nasuia Etheridge; Etheridge fil., 1892, p.567. B+ Trigonta nasuta Etheridge; Etheridge fil., 1902a, pp.25-7. Opisthotrigonia nasuta (Etheridge Snr), 1872; Skwarko, 1963, pp.23-4. LOCALITY: Maryborough SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS; Daintree Collection. F1249 SYNTYPE Trigonia nasuta Etheridge, 1872, p.339, pl.19, fig.2. Trigonia nasula Etheridge: Etheridge fil., 1892, p.567. Trigonia nasuia Etheridge; Etheridge fil., 1902a, pp.25-7. Opisthotrigonia nasuta (Etheridge Snr), 1872; Skwarko, 1963, pp.23-4. LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Daintree Collection. Opsidiscus microspinus Jell, 1975a F6770-F6772 PARATYPES Opsidiscus microspinus Jell, 1975a, pp.80-1: F6770, pl.26, fig.7: F6772, pl.24, fig.1. LOCALITY: QML154, 150m NW of gate on 1st crest of Brumby Creek Rd, 19°33'S, 138°54°E, NWOQ. FORMATION; Currant Bush Limestone, Euagnos- tus opimus Zone. AGE: M. Cambrian. Pagetia fluitata Jell, 1975a F6811 Hototyre Pagetia fluitata Jell, 1975a, pp.42-3, 93, pl.8, fig, 12, Locality: QML122, Limestone ridges, left bank of West Thornton River, S of junction of Brumby Creek, 13°32’S, 138°54°E, NWQ. FORMATION: Currant Bush Limestone, Euagnos- fus opimus Zone. AGE: M. Cambrian, Fas09-10 PARATYPES Pagetia fluitata Jell, 1975a, pp.42-3; F6810, pl.8, fig.11- LOCALITY, FORMATION, AGE: As for holotype. Pagetia howard Sell, 1975a F6738-6758 PARATYPES Pagetia howardi Jell, 1975a, pp.43-5, MEMOIRS OF THE QUEENSLAND MUSEUM F6738, pl.4, fig.7; F6739, pl.4, fig.7; F6740, pl.4, fig.7; F741, pl4, fig.7; F6742, pl.4, fig.7; F6743, pl.4, fig.S; Fa744, pl4, fig.8; F6745, pl.4, fig.2; 6751, pl.4, fig.10. LOCALITY: OMLI117, Low hill, 1.6km N of D Tree Bore, E of road 19°41°S, 138°54°E, NWQ. FORMATION: Beetle Creek Pm, Xystridura templetonensis zone. AGE: M. Cambrian. Pagetia ocellata Je\l, 1970 F323 HoLotyPr Pagetia ocellata Jell, 1970, pp-303-13, pl.24, fig. 22; Jell, 1975a, pp.50-1. LocaLity: QML113, 2.5 to 3km N of Mt Mur- ray, 80km SW of Duchess, 21°S0’S, 139°58’E, NWO, FORMATION: Beetle Creek Fm, AGE: early M. Cambrian. F6170-F6322, F6324-F6326 Pagetia ocellata Jell, 1970, pp.303-13; F171, pl, 23. fig.31; F6175, pl.23, fig.30; F6186, pl.23, fig.29; F6187, pl.23, fig.28; P6191, pl.23, fig.27; F192, pl.23, fig.6; F6193, text fig.3; F6194, pl.23, fig.8; F6195, pl.23, fig.7; F6197, text fig.3; F6200, pl.23, fig.10; F6201, pl.23, fig.9; F6207, pl.23, fig.15; F6215, text fig.3; F6216, pl.24, fig. 15; F6217, pl.24, fig.14; F6218, pl.23, fig.18; F6219, pl. 23, fig.17; F6220, pl.23, fig.16; F6221, pl.23, fig.23; F224, pl.23, fig.25; F6227, text fig.3; P6229, pl.23, fig. 26; F6230, pl.23, fig.24; F6234, text fig.3; F6236-8, text fig.3; F6239, pl.23, fig.1, pl.24, fig.16; F624), pl.23, fig.2, pl.24, fig.17; F6244, pl.23, fig.3, text fig.3; F6245, pl.23, fig.4; F246, pl.23, fig.5; F6248, pl.23, fig. 11; F6253, text fig.3; P6255. pl.23, fig.12; F6264, pl.23, fig. 13, text fig.3; Fh266, pl.23, fig.19; F6268, pl.23, fig. 14; F6269, pl.23, fig.20; F6270, pl.23, fig.21, text fig.3; F6272, pl.23, fig.22, text fig.3; P6274, pl.24, fig.18, text fig.3; F6275, pl.24, fig.19, text lig.3; F6276, pl.24, figs. 6,7; F6278, pl.24, lig.1; F6283, pl.24, fig.4; P6285, pl.24, fig.2; F289, pl.24, fig.3; F6293, pl.24, fig.S; F6299, pl. 24, fig.13; F6306, pl.24, Tig.8; Fo308, pl.24, fig.9; F6309, pl.24, fig.12; P6314, pl.24, fig. 10; F6322, pl.24, fig.11; F6324, pl.24, fig.20; F6325, pl.24, fig.21; F6326, pl.24, tig.23, LOCALITY, FORMATION, AGE: As for holotype. REMARKS: Material consists of several hundred dissociated cephala, thoracic segments, pygidia, hypostomata and free checks. Specimens F6223 and 6245 are missing. Some minor damage to a few of the specimens isolated from matrix - INVERTEBRATE FOSSIL TYPES especially Fol86, 6270, 6275, 6289 - has oc- curred since description. The holotype and QMF6275 were pl.9 figs 2 and 3 respectively in Hill er al. (1971). F6938-6945 Pagetia ocellata Jel. 1970; Jell, 197Sa, pp.S0-1; F6938. pl. 28, fig.1; F6939, pl.27, fig.7; F6940, pl.27, fig.8; F941, pl27, fig.10; Fe942, pl.27, fig.¥: FoU43, pl2k, fig.2; F944, pl.28, fig.3; F6945, p.50. LOCALITY, FORMATION, AGE: As for holotype. REMARKS: F6938 was also listed erroneously as P. prolata in Sell (1975a, p52). Jell (1975b), figured F6942 (p.35, fig.Ja; p.39, fig.4c) and F6945 (p.39, fig.4d). Pagetia pollosta Jel, 1975a F6765 Hovotyre Pagetia pollesta Jell, 1975a, pp.53-4, pl.24, figs 7a,b. LOCALITY: QMLI54. 150m NW of gate. Ist crest Brumby Creek Rd, 19°33°S, 138°54’E, NWO. FORMATION: Currant Bush Limestone, Euagnos- tus Opimus Zone, AGE: M. Cambrian. F6759-F6764, FO766-FO769. F6773-F678 | PARATYPES Pagetia pollosta Jell, 197Sa, pp.53-4; Fo759_ pl24, fig.S; Fa762, pl.24. fig.9; P6763, pl.24, fig. 13: Fo266, pl.24, fig.12; F6768, pl.24, fig. 112 F676, pl, 24, fig.3; F6778, pl.24, fig. 14; Fo779, pl24, fig.8; FO780, pl24, lig.6; F6787, pl.24, fig. 10, LOCALITY, FORMATION, AGE: As for holotype. REMARKS: F677) and F6772. which refer to Opsidiscus microspinus, were erroneously in- cluded here in Jell (197Sa). Pagetia prolata Jel, \97Sa Fo872 HoLotyre Pageria prolate Jell, 197Sa, pp.39, 56-00, pLll, figs. LOCALITY: QML145. N bank of Playfard River, 400m W of Barkley Stock Route to Brunetic Downs, 19°4’S, 136°40°E, NWQ. FORMATION: Burton Beds, fempletonensis Zone. AGE: early M. Cambrian. NXystridura F6869-F687 |. F6S73-F6936 PARATYPES Pagetia proflata jell, 19754, ppS6- 695 60; Fos869 pli, fig.3; Fos870, pli, fig.4; F6871, pl.U1, fig.7: F6873, pl. 12, fig.2; F6874, pl.12. fig.3: F6875, pl.12, fig.4; Fo876, pl.12, fig.6: F6877, pl.12, fig.10, F6878, pl.12, fig.11, F6879, pl. 12, fig.12; P6880, pl.12, fig.14; F6881, pl.12, fig.16: FOS82, pl.12, fig.21; P6883, pl.12, fig.22. LOCALITY, FORMATION, AGE: As for holotype. REMARKS: Jell (1975a) erroneously included F6937 and F6938 here. These numbers refer to P. acellata. Jell (1975b) figured F6873 and F6874 (p.36, figs 2e-h). However, registration numbers are not cited for cach figure. Pagetia salebra Sell, 1975a F6786 HoOLoryre Pagetia salebra Jell, 1975a, pp.48,60-1; pl.13, fig. 9. LOCALITY: QML146, .8km N of Burketown- Camooweal Road, 19°317S, 138°52°E, NWQ. FORMATION; Inca Creek Fm, Euagnostus opimus Zone. AGE: M. Cambrian. REMARKS: The holotype was erroneously num- bered P6782 in the caption to pl.13, fig.9. (F6782 is an unpublished specimen referred to this species). F6788-F6793, F6796-6798, FO800-6803 PARATYPES Pagetia salebra Jell, 1975a, pp.60-1; Fo788, pl.3, fig-5; F679, pl.13, fig.l; F6791, pl.13. fig. 7, F6792, pl. 13, fig.3; P6793, pl.13, fig.4. LOcALITY: OQML136, Thorntonia-Burketown Road at Chummy Bore, 19°31'S, 138°52’E, NWO. FORMATION. AGE: As for holotype. F6783-F6785, FO787, F6799_ FO804 PARATYPES Pagetia salebra Jell, 19754, pp.60-1; F67S3. pli3, figs 2ajb; P6784, pl.l3, figs 2a,e; F6787. pll3, fig.6. LOCALITY, FORMATION, AGE: As for holotype. Pagetia thorntonensis Jell, 1975a F6822 HoLoty rk Pagetia thorntonensis ell, 1975a, pp.65-7, pL.22. lig.2. LOCALITY; QML128, 1,.6km W of road at gate N of Gum Lagoon Bare, W bank of W Thornton Creek. |5m trom base of cliff, 19°34'S, 138°55'E, NWQ. FORMATION: Currant Bush Limestone (Bottom 20m - Prychaunestus atavus zone). 696 AGE: M. Cambrian. F6812, F6813, F6815-F6824, F6841, F6843 Pagetia thorntonensis Jell, 1975a, pp.65-7; F6812, pl.22, fig. 9; F6813, pl.22, fig.1; F6815, pl.22, fig.3; F6818, pl.22, fig.5; F6823, pl.22, fig.10; F6824, pl.22, fig.6; F6841, pl. 22, fig.7; F6843, pl.22, fig.4 LOCALITY, FORMATION, AGE: As for holotype. F6814, F6825-F6840, F6842, F6844, F6845- F6866 PARATYPES Pagetia thorntonensis Jell, 1975a, pp.65- 7, F6814, pl.22, fig.8; F6846, pl.27, fig.2; F6847, pl.27, fig.1; F6848, pl.27, fig.3; F6849, pl.27, fig.4; F6850, pl.27, fig. 5; F6851, pl.27, fig.6. LOCALITY: QML132, hill south of QML128 19°34°S, 138°55’E, NWQ. FORMATION, AGE: As for holotype. REMARKS: F6860-F6866 are missing from col- lection. Jell (1975b), figured F6846 (p.37, figs 3a,b) and F6853 and F6852 (p.37, figs. 3c-3f. However, registration numbers are not cited for each figure. Pandanocrinus wellingtonensis Jell et al., 1988 F14532, F14537-F14539, F14542 PARATYPES Pandanocrinus wellingtonensis Jell et al., 1988, pp. 385-6. LOCALITY: QML512, Wellington topographic Sheet [799867] richly fossiliferous limestone (Unit 18 of Johnson, 1975) 870m NNW of Mountain View Homestead, Wellington Caves Rd, 9km SSW of Wellington, NSW. FORMATION: Garra Fm. AGE: Early Devonian, (Pragian, sulcatus biozone), Panopea acuta (Etheridge, 1872) F1251 SYNTYPE Panopaea (Mya) plicata, Sow. var. acuta Etheridge, 1872, pp.342-3, pl.21, fig. 3a. LECTOTYPE Panopea acuta Etheridge, 1872; Water- house, 1965, pp. 851-2. Panopea plicata acuta Etheridge; Waterhouse, 1969, p.72, pl. 6, figs 1,3. LOCALITY: Maryborough, SEQ (see remarks). FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: In an carly label and in Etheridge (1872) the locality for this Daintree specimen is given as Pelican Creek, Mitchell District. How- ever the specimen’s lithology is consistent with that of the Maryborough Fm and it also bears the MEMOIRS OF THE QUEENSLAND MUSEUM number 627 which is only on the Maryborough specimens and includes material studied by Etheridge (1872). The other specimen figured by Etheridge (1872, pl.21, fig.3) was referred to the Permian genus Vacunella by Waterhouse (1965). Panopea maccoyi (Moore, 1870) F1253 HOLOTYPE Panopaea sulcata Etheridge, 1872, p.342, pl.21, figs 2, 2a. Glycimeris sulcata Etheridge, Etheridge fil., 1892, p.571. Glycimeris maccoyi Moore; Etheridge fil., 1901, p.30. Panopea maccoyi (Moore, 1870) Fleming, 1970, pp.9- 10, pl.3, figs 3,4. LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Daintree Collection. Parallelodon costellata McCoy, 1844 F1211 Parallelodon costellata McCoy; Etheridge fil., 1892, p.274, pl.40, fig.12. LOCALITY: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. F1212 Parallelodon costellata McCoy; Etheridge fil., 1892, p.274, pl.40, fig.13. LOCALITY: Rockhampton district, MEQ. FORMATION: ?Malchi Fm. AGE: Lower Carboniferous. REMARKS: De Vis Collection. Parapisocrinus sp. F14842 Parapisocrinus sp.; Jell et al., 1988, p.391, figs27k-m. LOCALITY: UQL3579. Burges topographic Sheet [687683] one of five localities collected E to W along fence line 600m E of Martins Well windmill: all from upper part of limestone (lime- stone slightly folded), Martins Well, 8km E of Pandanus Creek Homestead, 200km NW of Charters Towers, NQ. FORMATION: Upper Martins Well Limestone Member, Shield Ck Fm. AGE: Early Devonian, (early Pragian, sulcatus biozone). INVERTEBRATE FOSSIL TYPES Parastacid gastrolith F7829 Crustacean gastrolith; Archer and Wade | 976, pp.383- 4, pl.54, fig.tf. Locatity: Terrigenous sediments underlying ‘Allensleight flow’ of the Nulla Basalt, overlying laterite and outcropping on Bluff Downs Station, along banks of Allingham Creek, 19°43’S, 145°36°E, NQ. FORMATION: Allingham Fm. AGE; Pliocene. Pedinogyra sp. F1358 Pedinogyra sp.; Hill et al., 1970, pl.Cz4, Tigs6a,b. LOCALITY: Limestone Quarry, Marmor, MEQ. FORMATION: Unnamed cave deposits. AGE: Pleistocene. Penarosa netenta Jell, 1977 F7059 HOLOTYPE Penarosa netenta Jell, 1977, pp.119-23, pl.21, figs 1a,b. LOCALITY: QML152, |.7km S of Chummy Bore, which is 6km W of Thorntonia Homestead (19°31.5°S, 138°52°E) and at Chummy Bore, NWO. FORMATION: Chummy Bore Fm, probably Euag- nostus opimus Zone. AGE: M. Cambrian. F7060-F7064 Penarosa netenta Jell, 1977, p. 119-23; F7000, pl.21, fig.5; F7061, pl.2l. fig.8: F7062, pl.2l, lig.6; F7064, pl.21, fiz 4 LOCALITY, FORMATION, AGE; As for holotype. F7065-7070 PARATYPES Penarosa netenta Sell, 1977, pp, 119-23; F7065. pl. 21, fig.3; F7068, pl.21, fig.2; F7069, pl.21, fig.7. Locaury: QML136, Chummy Bore on Thorntonia-Camooweal Rd, 19°31'S, 138°52’E, Nwo. FORMATION, AGE: As for holotype. Permasyrinx acuta (Etheridge, 1872) F5639 Spirifer bisulcata Sow., var. acuta in Etheridge, 1872, p.329, pl.16, fig. 1. 697 Spirifer trigonalis Martin var. acuta Etheridge; Etheridge fil,, 1892, p.230, pl.10, fig.12. Permasyrinx acuta (Etheridge snr) Walerhouse and Balfe, 1987, pp.24, 30, pl. 1, fig.3. LOcALiITy: Gympie, SEQ. FORMATION: Rammutt Fm. AGE: Permian. Phaenodesmia elongata (Etheridge, 1872) F1247 HoLotyre Leda elongata Etheridge, 1872, p.341, pl.20, fig.5. P Adrana clongata (Etheridge) Etheridge fil., 1892, pp.566-7, pl. 33, fig.6. Malletia elongata (Etheridge) Etheridge fil., 1902a, p.25. Malletia elongata (Etheridge); Etheridge fil.. 1902b, p.26. Phaenodesmia elongata (Etheridge, 1872) Fleming 1966a, pp.8-9, pl.4, fig.1. LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Daintree Collection, ‘Planorhis’ sp. F2819 ‘Planorbis’ sp.; Hill et al., 1970, pl.Cz4, fig.13. Locatity: Limestone Hill, Ipswich, SEQ. FORMATION: Silkstone Fm. AGE: Oligocene. Platyteichum coniforme (Etheridge fil., 1892) F122 HoLotTyPe Mourlonia (2) coniformis Etheridge fil,, 1892, pp.287-8, pl.41, fig.S. Platyicichum coniforme (Etheridge jun.) Dickins, 1961, pp.131-4, pl.17, figs 13-14. Platyteichum coniforme (Etheridge Jn. 1892), Water- house, [987a, pp.179-80, Platyteichum conifarme (Etheridge Jr, 1892); Parfrey, 1988, pp.20-1, LOCALITY: Banana Ck, near Banana, CQ. FORMATION: Flat Top Fm. AGE: Middle Permian. REMARKS; De Vis Collection. Plotiopsis balonnensis (Conrad, 1850) F6550 Plotiopsis balonnensis (Conrad, 1850); Hill et al., 1970, pl. C75, fig. 10. 698 LOCALITY: Darling Downs, SEQ. FORMATION: Unnamed fluviatile deposits. AGE: Pleistocene. Podophthalmus vigil Fabricius, 1798 F1184 Podophthalmus vigil Fabricius; Etheridge fil. and Mc- Culloch, 1916, p.9, pl.4, fig.3, pI.5, fig.3. Podophthalmus vigil Fabricius, 1798; Hill et al., 1970, pl. Cz6, fig.9. LOCALITY: New Channel, mouth of Brisbane River, Moreton Bay, SEQ. FORMATION: Unnamed estuarine deposit. AGE: Pleistocene/Holocene. Polinices sordidus (Swainson, 1821) F17639 Polinices sordidus (Swainson, 1821); Hill et a/., 1970, pl. Cz4, fig. 14. LOCALITY: Bullock Pt, Wide Bay, SEQ. FORMATION: Unnamed marine deposit. AGE: Pleistocene. Previously registered as Mo2930 in the Queensland Museum Mollusca register. Polychaete trail F16373b Annelid trail; de Vis, 1911, pp.12-4, pl.3, fig.2. LOCALITY: Wyangaria Stn, near Hughenden, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous. Porites sp. F12385 Porites sp.; Pickett et al., 1985, pp. 103-14. LocALity: S of Amity Point, N Stradbroke Is- land (Brisbane 1:100,000 Sheet, grid reference 441658). FORMATION: Unnamed marine deposits. AGE: Pleistocene. REMARKS: Pickett et al., (1989) redated this specimen, Goniastrea aspera (F12401) and Symphyllia sp. cf. S. recta (F12400). They referred these corals to isotope substage 5c, not Se as previously cited (Pickett ef a/., 1985). Productus sp. indet. F1204 MEMOIRS OF THE QUEENSLAND MUSEUM Productus sp. ind.; Etheridge fil., 1892, p.256, pl.40, fig. 4. LocALtty: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. AGE: Carboniferous. Prohysteroceras richardsi Whitehouse, 1926 F1783 Prohysteroceras richardsi pp.222-3. LOCALITY: Toliness Station, near Augathella, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). Whitehouse, 1926, F1784 HoLotyPE Prohysteroceras richardsi Whitehouse, 1926, pp.222-3, pl.38, figs 1a,b. Prohysteroceras richardsi Whitehouse, 1926; Hill et al., 1968, pI.K9, fig.3. LOCALITY, FORMATION, AGE: As for F1783. Promytilus mytiliformis (Etheridge fil., 1892) F1217. SYNTYPE. Modiomorpha mytiliformis Etheridge fil., 1892, p.273, pl.41, fig.4. Promytilus mytiliformis (Etheridge, 1892); Water- house, 1980, pp.106-7. LOCALITY: Banana Creek, CQ. FORMATION: ?Flat Top Fm or Barfield Fm. AGE: Permian REMARKS: Etheridge fil. (1892) referred three specimens to this taxon pl.14, fig.5, pl.38, figs 12, 13 and pl.41, fig.4 (which is laterally reversed), not pl.40, fig.4 as cited in Etheridge fil. Waterhouse (1980, p. 106) cited the specimen figured by Etheridge fil. (pl.41, fig.4) as the lectotype erroneously referring this Queensland Museum specimen (F1217) to the Geological Survey Collection. Waterhouse (figs 2; 6, 8) illustrated two specimens one of which (fig.2; 8) is a latex cast from an external mould, captioned ‘lectotype.’ Neither figure corresponds with Etheridge fil. (pl.41, fig.4), i.e., F1217. The num- ber (Waterhouse, 1980) associated with the lec- totype caption is F2188 which does not correspond with a Queensland Museum number. The specimen is apparently of Etheridge fil.(pl.14, fig.5) (GSQ F1488), (Susan Parfrey, pers. comm., 1988). The other specimen of P. mytiliformis figured by Etheridge fil. (pl. 38, figs.12, 13 (GSQ FI505)) is also in the Geologi- cal Survey Collections. Information for INVERTEBRATE FOSSIL TYPES Waterhouse’s fig.2; 6 is lacking and no matching specimen can be located in the Queensland Museum Collections, Waterhouse (fig.3; 1) also figured a latex cast of the anterior hinge of the ‘lectotype’ but (his specimen is not P1217 and is not a muscum specimen, Protacanites planorbiformis (Etheridge fil., 1892) F1230 HOLOTYPE Goniatiies planarbiformis Etheridge fil. 1892, p.295, pl.41. fig.9. Protocanites planorbifarmiy (Etheridge); Hill and Woads, 1964, pC 14, fig.6. Evcanites planorbiformis (Etheridge); Weyer, 1972, p.322. Protocanites planorbiformis (Etheridge); Campbell etal, }983. pl ll-2, fig.36. LOCALITY: Stanwell area, MEQ. FORMATION: Malchi Fm. AGE; Upper Carboniferous. REMARKS: As Campbell ef a/., (1983) pointed oul, the type locality (Lakes Creek) as given in Etheridge (1892) is Permian in age. De Vis col- lections from the Stanwell Area in the Museum had been included symongst material from the Lakes Creck locality in the past. [tis likely that the G. planorbiformis and the other ammonoids Etheridge described were collected from the Stanwell area. The lithology of topotypes of ‘Pseudarietites armmonitiformis’ is consistent with the Malchi Formation. The holotype of P. planorbiformis and the other Carboniferous am- monoids (F1228 and F1209) were loaned over- seas, but have been lost. ‘Pseudarietites’ ammonitiformis (Etheridge fil... 1892) F1228 Syntyee Naurilus ? ammontriformis Etheridge fil. 1892 pp.292-3, pl39, fig.9- Pseudarietites sp.; Bryan, 1929, p76. *Pseudarictites’ ammonitiformis (Etheridge); Hilland Woods, 1964, pl.Cl4, fig.&. ‘Pseudarietites” ammonitiformis (Etheridge); Campbell ef a/., 1983, pp.1 20-1, fig 49b LOCALITY: Stanwell area, MEQ, FORMATION; Malchi Fm. AGE: Upper Carboniferous. REMARKS: See Protecanites planorbiformis. This specimen is lost, although topotypes exist. F1229 699 SYNTYPE Nautilus ? ammonitiformis Etheridge, 1892, pp.292-3, pl.4l, fig.9. Pseudurietites sp.; Bryan, 1929, p.76, *Pseudarietites’ ammanitiformis (Etheridge); Campbell et al. 1983, pp. 120-1, fig.49a. LOCALITY: Stanwell Area, MEO. FORMATION: Malchi Fm. AGE: Upper Carboniferous. REMARKS: See Protocanifes planorbiformis. This specimen is lost, Pterinopecten devisii Etheridge fil., 1892 F1213a,b HOLOTYPE Prerinopecten devisii Etheridge, 1892, pp.270-1, pl. 40, fig.9. Prerinopecten devisti Etheridge, |892b; Waterhouse, 1982, p.d. LOcALity: Rockhampton District, MEQ. FORMATION: ?Malchi Fm. Ace: Lower Carboniferous. REMARKS: De Vis Collection. Pustulospiriferina etheridget 1970) (Armstrong, F6329 PARATYPE Functospirifer etheridgei Armstrong, 1970. pp.3 17-9, Pustulospiriferina etheridget (sic) Armstrong; Water- house, 1987b, p.45- Locality. UOL3127, Series of E-W Ridges, 0.5 miles E of ‘Homevale' Homestead, CQ. FORMATION: Tiverton Fm, AGE; Permian. F6330 PARATYPE Punctospirifer etheridget Armstrong, 1970, pp.317-9. Pustulospiriferina etheridgei (sic) Armstrong; Water- house, 1987b, p45. LOCALITY, FORMATION, AGE: As for F6329 F633 1 PARATYPE Punetespirifer etheridge Armstrong, 1970, pp.317-9 pl.25, fig.9, Pustulospiriferina ctheridget (sic) Armstrong; Water- house, 1987b, p.45. LOCALITY, FORMATION, AGE! As for F6329 Fo332 PARATYPE Punetospirifer etheridgei Armstrong. 1970, p.317-9, pl.25, fig. 17. Pustulospiriferina etheridgei (sic) Armstrong; Waler- house, 1987b, p.4s. 700 LOCALITY, FORMATION, AGE: As for P6329 Puzosia longmani Whitchouse, 1926 FI595 Hototyee Puzesia longmuni Whitehouse, 1926, pp.218-9, pl.37, fig.5, pl.39, figs la,b. LocaLtity: Burcoo River, CQ. FORMATION: ? AGE: Lower Cretaceous. Pyramus concentrica (Etheridge, 1872) F14398 Pyramus concentrica (Etheridge) Waterhouse and Balfe, 1YX7, pp.24, 30, pl.l, fig.9. LOCALITY: Gympie. SEQ. FORMATION: Rammutt Fm, AGE: Permian. Rhipidocrinus crenatus (Goldfuss, 1831) F14771 Rhipidocrinus crenatus (Goldfuss, 1831); Jell et al. 1988, pp.360, 362-3, fig.4k. LOcALity: UQLS320 Wando Vale [563392] section on ridge 2.9%km NE of Storm Dam. Broken River Wando Vale Station, 15Okm NW of Charters Towers, NQ. FORMATION: Papilio Mudstone. AGE : Middle Devonian, (late Eifelian- Givetian). REMARKS: The formation was formally named by Lung ef al. (1989) as Papilio Mudstone, F14869 Rhipidocrinus crenatus (Goldfuss, 1831) in Jell ef al., 1988, pp.360.362-3, fig.4l, Locauity: UQLS5321 Wando Vale [562390] creek section 3km NE of Storm Dam, Broken River Wando Vale Station, 150km NW of Charters Towers, NQ, FORMATION, AGE, REMARKS: As for F14771. Samarura sp. F12996a/b Samarura sp., Rozelelds, 1985b, pp.25-32, figs2a,b, LOcALity: Brassall Quarry, near Ipswich, SEQ. FORMATION: Aberdare Conglomerate. AGE: Late Triassic. MEMOIRS OF THE QUEENSLAND MUSEUM Sanmartinoceras fontinale Hudleston, 1890 F722 Sanmartinaceras alene (Tenison-Woods); Whitehouse, 1926, p. 205, pl.41, Figs3a,b. Sanmartinoceras fontinale (Hudleston); Whitehouse, 1927, pp. 116-7, text fig.4. LOCALITY: ?Walsh River, NO. FORMATION: Blackdown Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Whitehouse (1926) records the locality for this specimen as Walsh River. The Queensland Museum Palacontology register gives no locality. FI&69 Sanmartinoceras olene Whitehouse, 1926, p. 205. Sanmartinoceras fontinale (Hudleston); Whitehouse, 1927, pp. 116-7, pl.17, fig.4. LOCALITY: Walsh River, NQ. FORMATION: Blackdown Fim. AGE: Lower Cretaceous, (Aptian), REMARKS: Collected Hann’s Expedition, 1872. (Tenison-Woads); F1870 Sanmartinaceras alene (Tenison-Waods); Whitehouse, 1926, p. 205, Sanmartinoceras fontinale (Hudleston); Whitehouse, 1927, pp. 116-7, pl.17, fig.5, text fig.9. LOCALITY: Walsh River, NQ. FORMATION; Blackdown Fm, AGE: Lower Cretaceous, (Aptian). REMARKS: Collected Hann’s Expedition, 1872. Seylla serrata (Forskal, 1755) FI187 Seyila serrata Forskal; Etheridge fil, and McCulloch, 1916, pp. 9-10, pl.2, fig.) LOCALITY: New Channel, mouth of Brisbane River, Moreton Bay, SEQ. FORMATION: Unnamed estuarine deposit. AGE: Pleistocene/Holocene. REMARKS: Originally registered as F474, a ccl- lection of fossil crabs from Moreton Bay. F3237 Seylla serrata (Forskal, 1755); Hill et al., 1970, pl.Cz6, fig. LI. LocaAtity; Lime Pocket, near Donnybrook, Bribie Passage, SEQ. FORMATION: Unnamed estuarine deposit. AGE: Pleistacene/Holocene. INVERTEBRATE FOSSIL TYPES Serpula sp. F20)26c Serpula sp. Withers, 1932, p.122. LOCALITY: Raised beach rock, Magnetic Island, NO. FORMATION: Unnamed deposit. AGE: Pleistocene/Holacene? REMARKS: Withers used F2026 for 4 different tuxa on the same block. This specimen has been re-registered as F2026c. Shimantocrinus distinctodorsus Jell etal., 1988 ’ F14536, F14540 PARATYPE Shimantocrinus distinctodorsus Jell et al, L988, pp.390-1; FL4536, figs24m,n- Locauity: QML512, Wellington topographic Sheet [799867| nchly fossiliferous limestone (Unit 18 of Johnson, 1975) 870m NNW of Mountain View Homestead on Wellington Caves Road, 9km SSW of Wellington, NSW. FORMATION: Garra Fm. AGE: Early Devonian, (late Lochkovian- Pragian). Simbirskites morvenae Whitchouse, 1927 F1270 Perisphinctes Kayseri Neumahr and Uhlig, 1881; Etheridge fil., 1909, pp.238-9, pl.68. Simbirskites spp. noy. Whitehouse, 1926, pp.200-1, Simbirskites morvenae Whilehouse, 1927, p.111. LOCALITY, FORMATION, AGE: Uncertain, probably North Germany. ReMARKS: This is the larger of the two am- monites figured by Etheridge fil. (1909, pl.68). See F16438. F16438 ?Perisphinetes. kaysert Neumahr and Uhlig, 1881, Etheridge fil,, 1909, pp.238-9, pl,68 (doubtfully assigned specimen), Simbirskites spp. nov. Whitehouse, 1926, pp.200-1. HOLotyee Simbirskites morvenae Whitehouse, 1927, pl, pl.l6. fig. 1, text fig.S. LOCALITY, FORMATION, probably N Germany, REMARKS: This is the smaller of the two am- monites figured by Etheridge fil, (1909, pl.68). There are considerable doubts about the provenance of these specimens. Whitchouse (1927) records their locality as Victoria Downs, Morven. The specimens were originally registered in 1893 us D7710.12 and described in AGE: Uncertain, 701 the donor register as a “miscellaneous collection of fossils’, This number also applics to material from elsewhere in Queensland, including the Walsh River. Doubts about a Queensland provenance for the specimens were evident in Whitehouse (1927) and were firmer in Jater papers (1946, 1955). It now seems likely that they were not found in Queensland, but came instead from North Germany (Day, 1967), Whitchouse (1927) also says that a Mr Hurst was the donor, A manuscript note in a bound copy of the Etheridge fil. (1909) paper, originally belonging to R. Hamlyn-Harris, Director, Queensland Muscum (1910-1917), records. Mr Hunter.as donor. Spirifer sp. cf. §. bisuleata Sowerby F5641 Spirifera allied to Sp. bisulcata Sow. in Etheridge, 1872, p. 335, pl.t7, fig.4. Spirifera trigonalis Martin sp,var. bisulcara G.B. Sby.; Etheridge fil., 1892, p.230, pl.9, fig. 15. LOCALITY: Bawen River. CQ. FORMATION: ? AGE: Permian. Spirifera convoluta Phillips, 1836 P5640 Spirifer convaluta ? Phill, Etheridge, 1872, p.335. pl.17, fig.3. Spiriferconveluta Phillips; Etheridge fil., 1892, p.229. pl. 10, fig.1 1. LOcALiry: Bowen Basin, CQ. FORMATION: ? AGRE: Permian. Spirifera sp. cf. 8. aviformis M’Coy F952 Spirifera (allied to.S. aviformis McCoy); Etheridge fil., 1892, pl40, fig.3. LoOcALITy: Banana Creek, CQ. FORMATION: ?Flat Top Formation. AGE: Permian. Spirtfera striata (Martin) F5642 Spirifera striata (Marlin); Ethendge, 1872, pp.334-5, pl.i7, fig.5- Spirifera striata Martin sp? Etheridge fil.. 1892, pp.227-8. pl. fig. 16. Localtry: Bowen Basin, Peak Downs, CQ. 702 FORMATION: ? AGE: Permian. REMARKS: R. L. Jack footnote in Etheridge fil. (1892, p.228) noted that the Bowen River does not extend into the Peak Downs District. Strangesta sp. F6549 Strangesta sp.; Hill et al., 1970, pl.Cz4, fig.7. LOCALITY: Olsens Cave, near Rockhampton, MEQ. FORMATION: Unnamed cave deposits. AGE: Pleistocene. Strophomena analoga (Phillips, 1836) F945 Strophomena rhomboidalis var. analoga Phillips; Etheridge fil., 1892, p.245-6, pl.40, fig.6. LOCALITY: Rockhampton District, MEQ. FORMATION, AGE: Permian. REMARKS: Referred to as Strophomena analoga Phillips? in Etheridge fil. (1892, pl.40, fig.6). Struszocrinus dulciculus Jell et al., 1988 F14534 PARATYPE Struszocrinus dulciculus Jell et al., 1988, pp.367-8, fig.9n. LOCALITY: QML512, Wellington topographic sheet [799867] 870m NNW of Mountain View Homestead on Wellington Caves Road, 9km SSW of Wellington, NSW. FORMATION: Garra Fm. AGE: Early Devonian, (Pragian, assumed sul- catus biozone. REMARKS: Jell ef al. (1988) incorrectly refer to this specimen as F14543 in fig.9. F14535 PARATYPE Srruszocrinus dulciculus Sell et al., 1988, pp.367-8. LOCALITY, FORMATION, AGE: As for F14534. Symphyllia sp. cf. S. recta (Dana, 1846) F12400 Symphyllia cf. recta; Pickett et al., 1985, pp.103-14. LOCALITY: S of Amity Point, N Stradbroke Is- land (Brisbane 1:100,000 Sheet, grid reference 441658). FORMATION: Unnamed marine sediments. AGE: Pleistocene. REMARKS: See Porites sp. MEMOIRS OF THE QUEENSLAND MUSEUM Tellina mariaeburiensis Etheridge, 1872 F1254a/b SYNTYPE Tellina mariaeburiensis Etheridge, 1872, p.341, pl.20, fig.6a. Palaeomoera mariaeburiensis (Etheridge) Etheridge fil., 1892, p. 570. Tellina mariaeburiensis (Etheridge, 1872) Hill et al., 1968, pl. K6, fig.10. Tellina mariaeburiensis Eth., 1872; Fleming 1970, pp.7-8, pl. 1, fig.4. LOCALITY: Maryborough, SEQ. FORMATION: Maryborough Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: The counterpart of this specimen referred to as F1255 by Hill e¢ al. (1968) and Fleming (1970) was reregistered as F1254b. Daintree Collection. F1256 SYNTYPE Tellina mariaeburiensis Etheridge, 1872, p.341, pl.20, fig.6 (large valve). Palaeomoera mariaeburiensis (Etheridge) Etheridge fil., 1892, p. 570. Tellina mariaeburiensis Eth., 1872; Fleming, 1970, pp.7-8, pl. 1, fig.5. LOCALITY, FORMATION, AGE: As for F1254. REMARKS: Daintree Collection. F1257 SYNTYPE Tellina mariaeburiensis Etheridge, 1872, p.341, pl.20, fig.6 (small valve). Palaeomoera mariaeburiensis (Etheridge) Etheridge fil., 1892, p. 570. Tellina mariaeburiensis Eth., 1872; Fleming, 1970 pp.7-8. LOCALITY, FORMATION, AGE: As for F1254. REMARKS: Daintree Collection. Tetraclita sp. F2026b Tetraclita sp.; Withers, 1932, p.122. Tetraclita sp.; Hill et al., 1970, pl.Cz6, fig.1. LOCALITY: Magnetic Island, NQ. FORMATION: Raised beach rock. AGE: Pleistocene/Holocene? REMARKS: Withers (1932) used F2026 for four different taxa on the same block. This specimen has been reregistered as F2026b. Thalassina squamifera (Herbst, 1804) F679 Thalassina anomala, Herbst; Etheridge fil. and Mc- Culloch, 1916, p.7. INVERTEBRATE FOSSIL TYPES LOCALITY: Daly River, Anson Bay, NT. FORMATION: Unnamed estuarine deposits. AGE. Pleistocene. REMARKS: Campbell and Woods (1967) evaluated modern and fossil material previously referred to 7. anomala from Australia and placed it in 7. squamifera. F6556 Thalassina anomala (Herbst, 1804); Hill et al., 1970, pl.Cz6, fig.5. LOCALITY: Styx River, MEQ. FORMATION: Unnamed estuarine deposits. AGE: Pleistocene, REMARKS: See F679. F13694 Thalassina anomala Herbst; Etheridge fil. and Mc- Culloch, 1916, p.7, pl.1, fig.1. LOCALITY: Darwin, NT. FORMATION: Unnamed estuarine deposits. AGE: Pleistocene. REMARKS: This specimen is part of a collection of T. squamifera that was given the registration number F318. F13694 and the other figured material have been allocated new registration numbers. See F679. F13695 Thalassina anomala Herbst; Etheridge fil. and Mc- Culloch, 1916, p.7, pl.1, fig.2. LOCALITY: Darwin, NT. FORMATION: Unnamed estuarine deposits. AGE: Pleistocene. REMARKS: Sec F 13694, F13696 Thalassina anomala, Herbst; Etheridge fil. and Mc- Culloch, 1916, p.7, pl.2, fig.3. LOCALITY: Darwin, NT. FORMATION: Unnamed estuarine deposits. AGE: Pleistocene. REMARKS: Refer F13694. Tillocheles shannonae Woods, 1957 F3248 Tillocheles shannonae Woods, 1957, pp.171-3. LOCALITY: ‘Currane’, 10 miles N of Dartmouth, CQ. FORMATION: Allaru Mudstone. AGE: Lower Cretaccous, (Albian). F3250 Tillocheles shannonae Woods, 1957, pp.171-3. LOCALITY, FORMATION, AGE: As for F3248. F3251 Tillocheles shannonae Woods, 1957, pp.171-3, pl.5, fig.6. Tillocheles shannonae Woods, 1957; Hill et al., 1968, pl.K11, fig.4, LOCALITY, FORMATION, AGE: As for F3248. F3252 Ho.otyPe Tillocheles shannonae Woods, 1957, pp.171-3, pl.5, fig.5, text fig.8. LOCALITY, FORMATION, AGE: As for F3248. F3253 Tillocheles shannonae Woods, 1957, pp.171-3. LOCALITY, FORMATION, AGE: As for F3248. ? Tonohamites taylori (Etheridge fil., 1892) F1271 HOLOTYPE: Ancyloceras taylori Etheridge fil., 1892, pp.498-9, pl.42, fig.13. Crioceras taylori, Eth. fil., Etheridge fil., 1909, pp. 162-3. Toxoceratoides taylori (Etheridge fil.) Whitehouse, 1926, pp. 216-7. ?Tonohamites taylori (Etheridge Jnr., 1892) Day, 1974, p.14. LOCALITY: ‘Wrotham Park’, Walsh River, NQ. FORMATION: Blackdown Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Day (1974) noted that Arkell et al., (1957) regarded Toxoceratoides and Tonohamites as nomina dubia and possibly synonymous with Hamiteceras Anderson, 1938. Casey (1961, pp.76-7) resurrected Hyatt’s fami- ly Helicancylidae as a subfamily of An- cyloceratidae, and concluded that Toxoceratoides and Tonohamites ate recog- nisable taxa within this subfamily. Day (1974) was unsure of the generic placement of F1271 and F1797 but referred them to Tonohamites. Collected Hann’s Expedition, 1872. See also F1396 (Australiceras jacki). F1797 Toxoceratoides taylori (Etheridge fil.,) Whitehouse, 1926, pp. 216-7, pl.34, fig.S. ?Tonohamites taylori (Etheridge Jr, 1892) Day, 1974, p.14. LOCALITY: ‘Wrotham Park’, Walsh River, NQ. FORMATION: Blackdown Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Collected Hann’s Expedition, 1872. T4 This specimen was part of the same hand specimen which contains F1271. The shaft of the specimen is missing, but the external mould of the shaft remains. (See also F1271). Torynomma quadrata Woods, 1953 F2877 HOLOTYPE Torynomma quadrata Woods, 1953, pp.54-6, figs3a,e, pl. 2, fig.6. Tarynomma quadrata Woods, 1953; Hill ef al. 1968, pLAI1, fig. 9. Torynomma quadrata Woods; Glaessner, 1980, p.181, pl.2. fig. 8. LocALttty: 10 miles N of Dartmouth, CQ, FORMATION: Allaru Mudstone. AGE: Lower Cretaceous, (Albian). F2878 Torynomma quadrata Woods, 1953, pp.54-6. Torynomma quadrata Woods; Glaessner, 1980, pla. LOCALITY, FORMATION, AGE: As for F2887. F2879 Torynomma quadrata Woods, 1953, pp.54-6. Torynomma quadrata Woods; Glaessner, 1980, p.181. LOCALITY, FORMATION, AGE: As for F2877. F2880 Toarynomma quadrata Woods, 1953, pp.54-6, fig.3d. Tarynomma quadrata Woods; Glaessner, 1980, p.181. LOCALITY, FORMATION, AGE: As for F2877. F288] Torynemma quadrata Woods, 1953, pp.54-6. Torynomma quadrata Woods; Glaessner, 1980, p.181. LOCALITY, FORMATION, AGE: As for F2877. Triassocotis amplicata Evans, 1961 F3689a/b HOLOTYPE Triassecotis amplicata Evans, 1961, p.16, fig.ig. LOCALITY: MI Crosby Insect Beds, SEQ. FORMATION: Mt Crosby Fm, AGE: Early Late Triassic, (Carnian). Triassocotis australis Evans, 1956 F3687 MEMOIRS OF THE QUEENSLAND MUSEUM Triassocolis australis Evans, 1956; Evans, 1961, p.16, fig. le. LaOcaLity: Mt Crosby Insect Beds, SEQ. FORMATION: Mt Crosby Fm. AGE: Early Late Triassic, (Carnian), Triassacotis stricta Evans, 1961 F3688a/b HoLoryee Triassocotis stricta Evans, 196], p.16, fig. Hf , LOCALITY: Mt Crosby Insect Beds, SEQ. FORMATION: Mt Crosby Fm. AGr: Early Late Triassic, (Carnian). Triassohyponomus dinmorensis Rozefelds and Sobbe, 1987 F13436 Insect leaf mines, Rozefelds, 1985a, pp.80-1, pl.1, fig.a. HOLOTYPE Triassohyponomus dinmorensis, Roze- felds and Sobbe, 1987, pl. 51-57, figs 2,3. LOCALITY: Dinmore, 27°31°S, 152°51’E, SEQ. FORMATION: Tivoli Fm. AGE: Late Triassic. F13435 Triassohyponomus dinmorensis Rozefelds and Sobbe, 1987, p.51-57, fig.4d. LOCALITY, FORMATION, AGE: As for F13436. Triassothea analis Evans, 1956 F6515a/b Triassothea analis Evans, 1956; Evans, 1971, p.148, fig.ab. Locality: Mt Crosby, SEQ, Upper Bed, 910805-911805, Ipswich 1 mile military map. FORMATION: Mt Crosby Fm. AGE; Early Late Triassic, (Carnian). F6523a/b Triassothea analis Evans, 1956; Evans, 1971. p.148, fig.3c. LOCALITY, FORMATION, AGE: As for F6515. P6524 Triassothea analis Evans, 1956; Evans, 1971, p.148, fig.3d, LOCALITY, FORMATION, AGE: As for F6515. INVERTEBRATE FOSSIL TYPES Tricrosbia minuta Evans, 1971 F6520) HOLOTYPE Tricresbia minuta Evans, 1971, pp.145-6, fig. 1. Lacatity: Mt Crosby, SEQ. Upper bed, 910805-91 1805, Ipswich | mile military map. FORMATION: Mt Crosby Fm. AGF: Early Late Triassic, (Carnian). Trifidella perfecta Evans, )956 F3690a/b Trifidella perfecta Evans, 1956; Evans, 1961, p.18, fig.3a. Locatity: Mt Crosby Insect Beds, SEQ. FORMATION: MI Crosby Fm. AGF: Early Late Triassic, (Carnian). REMARKS: The counterpart of this specimen originally referred to as F3691 has been re- registered as F3690b. F3692 Trifidella perfecta Evans, 1956; Evans, 1961, p.18, fig.3b. LOCALITY, FORMATION, AGE: As for F3690. Fo504a/b Trifidella perfecta Evans, 1956; Evans, 1971, p.146. LOCALITY: Mt Crosby Insect Beds, Upper bed, 910805-911805, Ipswich, 1 Mile Military Sheet. FORMATION; Mt Crosby Fm. AGE: Early Late Triassic, (Carnian), Trigonta moorei Lycett, 1870 F5603 Trigonia moorei Lycett, 1870; Skwarko, 1974, pp.92-3. LOCALITY: ?Newmarracarra Limestone, WA. FORMATION: ?Newmarracarra Limestone. AGE: Middle Jurassic, (Bajocian). F5606 Trigonia moorei Lycett; Whitehouse, 1924, pp.4-6. Trigenia moore’ Lycett; 1870; Skwarko. 1974, pp.92- 3. LOCALITY. FORMATION. AGE: As for F5603 F5607 Trigonia mooret Lycett; Whitehouse, 1924, pp.4-6, LOCALITY, FORMATION, AGF: As for F5603. Tropacum undatum Whitchouse, 1926 Fl 264 705 Crioceras jackii Eth. fil.; Etheridge fil., 1909, pp.145- 8. pL3l, figs 1,2. HOLOTYPE Tropaeumn undaium Whitehouse, 1926, pp.215-6. Tropacum undatum Whitehouse, 1926; Day, 1974, pp.6-7, Table 1. LOCALITY: probably Walsh River, NQ. FORMATION; ? AGE: Lower Cretaceous. REMARKS: Collected Hann’s Expedition, 1872, probably from the Walsh River. F1266 Crioceras jackti Eth, fil.; Btheridge fil., 1909, pp.145- 8, pl.32. fig.2, pl.34. fig.1. Tropaeum arcticum (Stolley); Whitehouse, 1926, p.215. Tropaeum arcticum (Stolley); Day, 1964, p.18. Tropaeum undatum Whitehouse, 1926; Hill er al, 1968, K7, fig. 3. Tropaeum undatum Whitehouse, 1926; Day, 1974, pp.1, 7-8, pl. 3, figs 2a,b, Table 1. LOCALITY: Roma, SEQ. FORMATION: Blackdown Fm, AGE: Lower Cretaceous, (Aptian). F1598 HoLotyPr Tropaeum rarum Whitehouse, 1926, p.216, pl.36, figs 1a,b, Tropaeum undatum Whilehouse, 1926; Day, 1974, pp.6-&, Table |, LOCALITY: Walsh River, NQ. FORMATION: Blackdown Fm. AGE; Lower Cretaceous, (Aptian). REMARKS: Collected Hann’s Expedition, 1872. F1605 Criaceras jackii Eth, fil. Etheridge fil, 1909, pp.145- 8, pl.38, figs 4-5. Tropaeum undatum Whitehouse, 1926; Day, 1974, pp.6-8, Table 1, LOCALITY: ?Walsh River, NQ. FORMATION: ? Blackdown Fm. AGE: Lower Cretaceous, (Aptian). REMARKS: Collected Hann’s Expedition, 1872. Vacunella curvata (Morris, 1845) F1252 SYNTYPE Panepaea (Mya) plicata Sow.. var. acute Etheridge, 1872, pp.342-3, pl.21, fig.3. Chaenomya (?) acuta Etheridge; Etheridge fil., 1892, p.280. Vacunella curvata Morris (1845); Walerhouse, 1965, p 852. 706 Vacunella curvatu (Morris), 1845; Runnegar, 1967, pp.63-7. Vacunella curvata (Morris) 1845; Waterhouse, 1969, pp.38, 71-2. pl.3, figs 5,6. LOCALITY: Pelican Creck, Bowen River, CQ (sce remarks). FORMATION: ? AGE: Upper Permian. REMARKS: Etheridge (1872) considered this Daintree Collection material to be Cretaceous, although the locality for this material (if ac- curate) would rule out this possibility, Etheridge (1892) doubted whether fig.3 and 3a were referable to the same taxon bul indicated a Per- mian age for F1252. Waterhouse (1965) con- curred in this assessment and referred F1252 to the Permian genus Vacunella and F1251 to the Cretaccous genus Panapea. Waterhouse (1969) erroneously refers to P1252 as F1952. Vacunella dawsonensis Runnegar, 1967 F18i4 Chaenomya? n.sp.; Runnegar, 1966, pp.374-S, lig.1d. Vacunella? dawsenensis Runnegar, 1967, pp.73-4, pl, figs 1,2. ?Australamya dawsenensis Runnegar, 1967; Run- negar, 1969, pp. 287-8, figs 3a,c. Vacunella dawsonensis Runnegar. 1967; Waterhouse, 19874, p.174. LOCALITY: Castle Creek, Dawson Valley, CQ. FORMATION: ?Barficld or Flat Top Fms. AGE: Lower Upper Permian Velesunio ambiguus (Philippi, 1847) Fl621 Velesunio ambiguus (Philippi, 1847); Hilletad., 1970, pl. C25, fig.11. LOCALITY; Darling Downs, SEQ. FORMATION: Unnamed fluviatile deposit. AGE: Pleistocene, Wyndhamia clarkei (Etheridge, 1872) F2887 SYNTYPE Productus clarkei Etheridge, 1872, p.334, pL.t7, fig, 2. Productuy clarkei E\heridge; de Koninck, 1877. p.203. Praductus clarket Etheridge; Etheridge fil, 1878, p.5l. Struphalosia clarker Evheridge; Etheridge fil, 1892, pp.25h- 60) MEMOIRS OF THE QUEENSLAND MUSEUM Strophalosia clarkei Etheridge: Prendergast, 1942, pp.42-3 LECTOTYPE Strophalosia clarkei Etheridge; Maxwell, 1954, pp.546-7, pl.56, figs 5, 6. Wyndhamia clarkei (Etheridge Sr); Dear, 1971, pp.11- 2, Wyndhamia clarke: Etheridge; Hill et al., 1972, pLP4, fig, 11. Locaity: Collinsville District, Bowen River, NO. FORMATION: Big Strophalosia Zone, Blenheim Fm. AGE: Middle Permian. REMARKS: Daintree Collection, Maxwell (1954) referred lo this specimen as holotype although lectotypic status is correct. (See the /nternational Code of Zoological Nomenclature). F2888 SYNTYPE Productus clarkei Etheridge, 1872, p.334, pl.17, fig. 2a. Productus clarkei Etheridge: de Koninck, 1877. p-203, Productus clarkei Etheridge; Etheridge fil., 1878, p.Sl. Strophalosia clarkei Etheridge; Etheridge fil, 1892, pp. 258-60, Strophalosia clarkei Etheridge; Prendergast, 1942, pp.42-3. PARALECTOTYPE Strophalosia clarkei Etheridge; Maxwell, 1954, pp.546-7, pl. 56, fig.8. Wyndhamia clarkei Etheridge; Hill et al. 1972, pl.P4, fig. 12. LOCALITY: Collinsville District, Bowen River, NQ. FORMATION: Big Strephalosia Zone, Blenheim Fm. AGE: Middle Permian. REMARKS: Maxwell (1954) erroneously referred la this specimen as F2887 in pl.56, fig.8. Daintree Collection. Zaphrentis profunda Etheridge fil., 1892 F119} Hooyer Zaphrentis profunda Etheridge fil,, 1892, p.200, pl 44, fig 1. Zaphrentis profunda Etheridge Jnr, 1892; Hill, L978, pp:29, 34. LOcALITY: Rockhampton District, MEQ. FORMATION: ? AGE: Permian. REMARKS: The figure in Etheridge fil. (1892) INVERTEBRATE FOSSIL TYPES appears to have been drawn from a plaster cast of F191. This holotype has not been sectioned so its generic affinities are uncertain but it may be referable to Eurvphvllum (Susan Partrey, pers. comm,, 1989). De Vis Collection. ACKNOWLEDGEMENTS The following people provided advice on taxonomic problems and/or collections in their control: R. Cacks, R.W. Day, B.A. Engel, G.I. Ingram, P.A. Jell, S. Parfrey, 1. Sanker, A. Simpson, S. Turner, M, Wade, and J.B. Water- house, Their assistance has greatly extended the coverage and depth of this catalogue. The library staff of the Queensland Museum obtained litera- ture relevant to the catalogue. The staff of the State Archives also helped by locating informa- tion on the Gordon Downs. Property. Peta Woodgate kindly typed the inordinate versions of this catalogue. LITERATURE CITED ARCHER, M. AND WADE, M. 1976. Results of the Ray E, Lemley expeditions Part t. The Allingham Formation and a new Pliocene verlebrite fauna from northern Queensland. Mem. Od Mus. 17(3): 379-U7. ARKELL, W.J., KUMMEL, B. AND WRIGHT. CW. 1957, Mesovoie Ammonoidea, pp.L&O-L4A71. dn Moore, R-C. (ed,), “Treatise on Invertebrate Paleontology, Part L, Mollusca, Cephalopoda, Ammunoidea’. (Geol. Sac. Amer. and Univ, Kansas Press, Lawrence, Kansas). ARMSTRONG, J. 1970, Queenslind Permian species of the spiriterid brachiopods, Punetospirifer and Cleiothyriding. Mem, Gd Mus, 15(4); 315-21. Booker, F.W. 1932. A new species of Productus trom the lower Bowen series, Queensland, Prac, R. Soe. Od 43) 66-72. Bresrroreer, M. 1947. Sur les zones D' ammonites dans L’Albien de France ef U Angleterre. Trav, Lab. Geol. Univ. Grenoble 26; 1-38, Bry An, W.H, 1929. Report of Carboniferous and Permo-Carboniferous correlation commiltee. Revent advanees in our knowledge of [he Car- boniferous and Permo-Carboniferous forma- tions of Queensland. Rep. Ansiralas. Ass, Adve Sei, 19: 74-77. CAMPBELL, B.M. AND Woons, 1 T. 1967, Quaternary crustaceans from northern Australia in the col- leclions of the Bureau of Mineral Resources, TUT Cunberea. Bull, Bur. Miner. Resour, Geal. Geapliys. Aust, 108: 44-2, CAMPBELL, K.S,W, 1959. The Martiniopsis- like spiriferids of the Queensland Permian. Palaeon- tology U4); 333-50. CAMPBFLL, K-S.W., BROWN, D.A. AND COLEMAN, AR. 1983. Ammonoids and the correlation of the Lower Carboniferous rocks of eastern Australia. Aleheringa 7, 75-123. CASEY. R. 1961. A monograph of the Ammonoidea af the Lower Greensand. Palacontogr. Soe (Moanagr). Part 1, 45-118. CRAMP, ILA. 1985, Lower Crefaceous |noceramid bivalves from the Antarclic Peninsula region, Palaeontology 28(3): 475-525. DAINTREF. R. 1872. 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Part 1 (Simbirskitidae, Aconeceralidae and Parahoplitidac), Mem. Qd Mus. 9: 109-120. 1928. Additions to the Cretaceous ammonite fauna of eastern Australia. Part 2. (Desmoceratidae). Mem. Od Mus, 97 200-6. 1946. A marine Early Cretaceous fauna from Stan- INVERTEBRATE FOSSIL TYPES 711 well (Rockhampton District). Proc. R. Sac. Qd Maroochydore, Queensland. Gd Govt Min. J. 73: $7: 7-20. 321, 1955. The geology of the Queensland portion of the = Woons, J.T. 1953. Brachyura from the Cretaceous of Great Australian Artesian Basin, Appendix G./n Queensland. Mem. Od Mus. 13: 50-7, Artesian water supplies in Queensland. Dep. Co- 1957. Macrurous decapods from the Cretaceous of ord, Gen. Public warks Parl. Pap. A, 56-1955, Queensland. Mem. Od Mus. 13(3): 155-75, Brisbane. 1964. C. D’Oyly H. Aplin, first government WITHERS, T.H. 1932. Barnacles from Magnetic Island, geologist for the southern distnct of Queensland. north Queensland. Mem. Qd Mus. 10(2): 122-4. Mem, Qd Mus, 14; 107-14. Woop, P.A. 1972. A possible Holocene shoreline at Aconeceras walshense, 668 Actinacrinus sp., 668 Adrana elongata, 697 Alathyria pertexta, 668 Aleteceras plectoides , 693 Ambikella strzeleckii, 686, 687 Amertanna carinata, 668 Ammanites walshensis, 668 Ammonites beudanti var. mitchellt, 673 Ammonites daintreet, 673 Ammonites flindersii, 673 Ammonites sutherlandi, 674 Anadara trapezia, 668 Ancyloceras taylori, 703 Anidanthus springsurensis, 669 unnelid trail, 698 Aprathia rockhamptonensts, 669 Archavocidaris sp. , 669 Armenoceras sp.. 67 | Astacades sp., 671 Athyris rovsii. 670, 671 Aucella hughendenensis, 671 Aucellina hughendensts, 671 Australiceras all. irregulare, 671, 672 Australiceras gracile, 671, 672 Australiceras trregulare, 671 Australiceras jacki, 672, 703 Australiceras robustum, 672 Australiceras transiente, 672 “Australomya dawsonensis, 7016 Avicula alata, 689 Avicula hughendenensis. 671 Aviculopecten laurienti, 672 Aviculopecten multiradiatus, 679 Aviculopecten squamuliferus, 679 Baylea konineki, 670, 672 Bellerophon sp., 672 Bembicium melanastoma , 672 Beudanticeras daintreei, 673 Beudanticeras (?)) daintreei, 673 Beudanticeras flindersi, 673 Beudanticeras mitchelli, 673 RBeudanticeras perlatum, 673 Beudanticeras sutherlandi, 674, 688 Beudantiella ogilviet, 674 Boleticeras daintreei, 673 INDEX Boliteceras daintreei, 673 Boliteceras perlaium, 673 Brachymetopus dunstani, 674 Brachymetopus maccoyi var. spinimarginatus, 674 Bucania textilis, 674 burrows, 674 Callianassa (2) sp., 680 2Cancrinelloides sp.. 674 carpocrinid indet., 675 cercopoidea incertae sedis, 675 Chacnomya (?) acuta, 705 Chaenomya ? carinata, 693 Chaenomya ? n.sp., 706 Charvhdis callianassa, 675 Chelodes whitehousei, 675 Chihamaluy sp., 675 Collabrina cliftoni, 670, 675 Convphillipsia grandis, 676 Conophillipsia subquadrata, 676 Cophinaceras ogilviei, 674 Corbiculina australis, 676 ?Corrugopecten squamuliferus, 679 Crassostrea commercialis, 676 Crenatula (2) gibbosa, 690 cninoid allachment buses. 677 erimoid calyx, 677 crinoid cf, Stenmmatocrinus, 677 crinoid indet. 1, 677 Criveeras flindersi, 692 Crivceras jackii, 671, 672, 705 Criaceras plectoides, 693 Crivceras taylori, 688 Crosaphis anomala, 677 crustacean gastrolith, 697 Cucullaea costata, 683 Cucullaea rabusta, 683 Cucullaca semistriata, 677 Cuppressocrinites abbreviatus £677, 678 Cuppressocrinites sp. ef, gracilis . 678 Cyathophyllum shearsbyt, 684 Cypricardella rectangularis, 670. 678 Cyprina clarkei, 681 Cyprina expansa, 631 Cyprina ? (vel Cytherea ?) clarke, 681 Decoreatergum warrenae, 678 712 Deltapecten limaeformis, 679 Dielasma sp., 679 Dimitobelus sp., 679 Dolatocrinus peregrinus, 679 Pysmorphoptiloides elongata, 679 Echinalosia ovalis, 680 Edmoniia (2) smithi, 670, 680 Enoploelytia sp., 680 Enoploclytia terraereginae, 680 Enoploelytia terrae-reginac, 680 Eocanites planorbiformis, 699 Eoscartoides bryant, 680 Euastacus ? sp., 680 Eucalyptocrinites rosaceus, 680, 681 Euchondria laurienti, 672 Euporismites balli, 681 Euspira reflecta, 681 Euryphyllum sp., 706 Fissilunula clarkei, 681 Flindersites alt. baceatus, 692 Flindersites all. flindersi, 692 Flindersites intermedius, 692 Galene bispinosa, 681 gasterocomid indet.. 681 gastropods, 681 Georgina andersonorum, 68 | Georgina beuteli, 682 Georgina dwyeri, 682 Georgina linda, 682 Georgina taylori, 682 Glauconome sp., 682 Glycimeris sulcata, 696 Glyphea aculata, 683 Goniastrea aspera, 683 Goniatites planorbiformis, 699 Grammatodon (Indogrammatodon) robusta, 683 Gyaloceras smithi, 683 Gyreceras dubius, 683 Hamites aff, maximus, 683 Hamites? laqueus, 688 hemiptera incertae sedis, 683, 684 Hercophyllum shearsbyi, 684 Heterachterus timmsti, 684 Heterujassus membranaceus, 684 Heteronella markset , 684 heteropteron forewing, 686 Hexacrinites interscapularis, 684, 685 Hexacrinites spinosus, 685 Tomelopsts etheridge, 685, 686 homoptera incertae sedis, 686 Hoploparu mesembria, 686 Hylicella colorata, 686 fdonearca robusta, 683 Ingelarella strzeleckti, 686 Inoceramus allied to 1. problematicus. 687 Jnoceramus carsoni, O87 MEMOIRS OF THE QUEENSLAND MUSEUM Inaceramus cf, sutherlandi, 687 Inoceramus elongatus , 687 Inoceramus etheridgei, 687 Tnoceramus marathonensis, 687 Inoceramus multiplicatus var. elongatus, 687 Inoceramus pernoides, 687 Inoceramus sp. cf. 1. pernoides, 687 insect leaf mines, 704 Laheceras compressum, 687 Labeceras cf. compressum, 688 Labeceras (Labeceras) compressum, 687 Labeceras (Labeceras) laqueum, 688 Labeceras laqueus, 688 Lasivcladia hindei,. 688 Leda elongata, 697 Leionucula quadrata, 688 lepidoptera leaf mines, 689 Leucosia pubescens, 689 Linoproductus springsurensis, 669 Lithosmylidia baronnae, 689 Lithesmylidia parvula, 689 Lopha marshit var. australiensis, 689 Laxonema sp,, 670, 689 Luciella (?) grayae, 670, 689 Macceyella alata, 689 Macceyella barklyi, 689, 690 Maccoyella corbiensis, 690 Maccoyella reflecta, 690 Macrocallista (?) plana, 690 Macrophthalmus latreillei, 690 Malchiblastus australis, 690 Malletia elongata, 697 Martinia (vel. Martiniopsis) subradiata, 690 Meleagrinella sp., 690 Melocrinites tempestus, 691 Mesaktoceras arachne, 691 Mesoblastus ? australis, 690 Mesocicadella punctata, 69) Mesothymbris perkinsi, 691 Modiomorpha mytiliformis, 698 Mourlonia (?) coniformis, 697 Myacites sp., 690 Myloceras aff. baccatus, 692 Myloceras ammonoides, 692 Mylaceras auritulum, 692 Myloceras baccatum, 692 Myloceras davidi, 692 Myluceras flindersi, 692 Myloceras intermedium, 692 Myloceras intermedius, 692 Myloceras plectuides, 692, 693 Mytilops corrugata, 693 Naticd lineata, 681 Natica variabilis, 681 Nautilus ? ammonitiformis, 699 Nereites berneyi, 693 Nucula gigantea, 688 Nucula quadrata, 688 INVERTEBRATE FOSSIL TYPES 713 Nucula sp., 693 Oblicarina carinata, 693 Octomeris crassa, 693 Onestia etheridgei, 693 Oonoton woodsi, 693 Opisthotrigonia nasuta, 693, 694 Opsidiscus microspinus, 694 Pagetia fluitata, 694 Pagetia hawardi, 694 Pagetia vcellata, 694 Pagetia pollasta, 695 Pagetia prolata, 695 Pagetia salebra, 695 Pagetia thorntonensis, 695, 696 Palaeomaera mariaeburiensis, 702 Pandanocrinus wellingtonensis, 696 Panopea ucuta, 096 Panopea maccoyi, 696 Panopaea (Mya) plicata, var. acuta, 696 Panopea plicata acuta, 696 Panapaca suleata, 696 Parallelodon castellata, 696 Parapisocrinus sp., 696 parastacid, 68U Pedinogyra sp., 697 Penarosa netenta, 697 Perisphincles kayseri, 701 Permasyrinx acuta, 697 Phaenodesmia elongata, 697 Phillipsia dubia, 669 Phillipsia rockhamptonensis, 669 Phillipsia woodwardi, 669, 676 ‘Planorbis’ sp., 697 Platyteichum coniforme, 697 Pleurotomaria ? cliftoni, 670, 675 Plotiopsis balonnensis, 697 Padophthalmus vigil, 698 Polinices sordidus, 698 polychaete trail, 498 Porites sp,, 698 Productus clarkei, 706 Productus cora, 674 Praductus sp. ind., 698 Prahysteraceras richardst, 698 Promyulus mytiliformis, 698 Prosopon etheridgei, 686 Protocanites planarbiformis, 699 ‘Pseudarictites’ ammonitiformis, 699 Pseudarietites spp., 699 Pseudavicula alata, 689 Pterinopecten devisii, 699 Punclospirifer etheridgei, 699 Pustulospiriferina etheridgei, 699 Puzosia longmani, 700 Pyramus concentrica, 700 Rhipidocrinus crenatus, 700 Samartra sp., 700 Sanmartinoceras Jontinale, 700 Sanmartinoceras olene, 700 Sevila serrata, 700 Serpula sp., 701 Shimantocrinus distinctodorsus, 701 Simbirskites morvenae, 701 Simbirskites spp. nov., 701 Spirifer allied to Sp. bisulcata, 701 Spirifer allied to S. evifarmis, 701 Spirifer bisulcata var. acuta, 697 Spirifer canvoluta, 701 Spirifer canvoluta ?, 701 Spirifer sp. cE. $. aviformis, 701 Spirtfer strzeleckii, 686, 687 Spirifer striata, 701 Spirifer trigonalis vat. acuta, 697 Spirifer trigonalis var. bisulcata, 701 Spirifer undifera var. undulata, 686, 687 Squamuliferipecten squamuliferus, 679 Stemmatocrinus (basal cup of crinoid), 677 Strangesta sp., 702 Streptorhynchus davidsoni, 690 Strophalosia clarket, 706 Strophalosia gerardi, 680 Straphamena analoga, 702 Straphomena rhomboidalis var. analoga, 702 Struszacrinus dulciculus, 702 Symphyllia recta, 702 Symphyllia sp. ef, 8. recta, 702 Tellina mariachuriensis, 702 Tetraclita sp., 702 Thalassina anomala, 702, 703 Thalassina squamifera, 702 Tillocheles shannonae, 703 ?Tonohamites taylori, 672, 703 Torynomma quadrata, 704 Toxoceratoides taylori, 703 Triassocotis amplicata, 704 Triassocotis australis, 704 Triassocotis stricta, 704 Triassohyponomus dinmorensis, 703 Triassothea analis, 704 Tricoelocrinus ) carpenteri, 690 Tricrasbia minuta, 705 Trifidella perfecta, 705 Trigonia moorei, 70S Trigonia nasuta, 693 Tropaeum arcticum, 705 Tropaeum australe, 665 Trapaeum rarum, 705 Tropaeun undatum, 705 Unicardium? etheridget , 693 Vacunella curvata, 693, 705, 706 Vacunella dawsonensis, 706 Vacunella (?) dawsonensis, 706 Velesunio ambiguus, 706 Wyndhamia clarket, 706 Yvania konincki, 672 Zaphrentis profunda, 706 MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA AND THE ORIGIN OF STARFISHES ANDREW B. SMITH AND PETER A. JELL Smith, A.B. and Jell, P.A. 1990 08 31: Cambrian edrioasteroids from Australia and the origin of starfishes. Memoirs of the Queenland Museum 28(2): 715-778. ISSN 0079-8835. The early Middle Cambrian edrioasteroids Stromatocystites reduncus sp. noy, and Edrio- discus primoticus (Henderson & Shergold), and the median Upper Cambrian edrioasteroids Chatsworthia spinosa gen. et sp. nov., Hadrodiscus parma gen. et sp. nov. and Cambroblas- tus enubilatus gen. et sp. nov. are described. C. enubilatus is a primitive edrioblastoid linking edrioblastoids to the cyathocystinid Edrioasteroidea. Chatsworthia and Hadrodis- cus are primitive isorophid edrioasteroids probably retaining biserial ambulacral flooring plates. The primitive starfish Archegonaster, from the Llanvirn of Czechoslovakia is redescribed. It is shown to be sufficiently generalised in its morphology to belong to the common stem lineage of both asteroids and ophiuroids. The most probable ancestry of Archegonaster is traced to Cambrian edrioasteroids with Edriodiscus as its closest relative. The origin and homologies of the ophiuroid jaw are clarified on the basis of Archegonaster mouth elements and it is argued that mouth angle plates are homologues of first ambulacral ossicles and the torus and its mouth spines are homologues of the first adambulacral ossicles and its spines. Edrioasteroid phylogeny is analysed using numerical cladistic methodology and an evolu- tionary tree constructed from stratigraphical occurrence data. Edrioasteroids diversified in paleoecological terms from being unattached low-level epibenthic suspension feeders to fill a number of different niches during the Cambrian and Early Ordovician. Different groups became semi-infaunal, vagile, semi-permanently attached or permanently cemented to a hard substratum, while one line even became convergent with crinoids by evolving a stem, The fauna described here suggests that edrioasteroids continued to diversify through the Upper Cambrian and that their rarity at this time interval is due to preservational factors. echinoderms, Cambrian, Ordovician, Australia, Czechoslovakia. Andrew B. Smith, British Museum (Natural History), Cromwell Road, London SW7 SBD, United Kingdom; Peter A. Jell, Queensland Museum, P.O. Box 300, South Brisbane, Queensland 4101, Australia; | August 1989. Echinoderms have a good fossil record that extends back to the Lower Cambrian and possib- ly even into the Precambrian (Gehling, 1988). Much attention has been focused recently on the early evolutionary history of this group (Termier & Termier, 1969; Ubaghs, 1975; Paul, 1977, 1979, 1988; Sprinkle, 1976, 1980a, 1981; Paul & Smith, 1984; Smith, 1984, 1986, 1988a; Campbell & Marshall, 1987). This has greatly improved our understanding of the morphology of early, primitive echinoderms but substantial gaps remain in our understanding of both phylogenetic relationships and evolutionary pat- terns as echinoderms diversified. In particular the origins of certain post-Cambrian lineages remains problematic because suitable inter- mediates, linking the derived forms to known Cambrian fauna, have not been discovered. Two factors greatly aggravate this problem; the ap- parent absence of a good Upper Cambrian record for echinoderms and the strong geographical bias in the known early fossil record. The dearth of identifiable Upper Cambrian fossil echinoderms was first pointed out by Sprinkle (1976) who noted that, although disar- ticulated elements of echinoderms were not un- common in Upper Cambrian limestones of western U.S.A., whole animals were extremely rare. Sprinkle (1980a, 1981) and Campbell & Marshall (1987) argued that this low diversity is genuine and that echinoderms underwent a two phase diversification, as postulated for marine invertebrates in general (Sepkoski, 1979). Smith (1988a), however, interpreted the Upper 716 Cambrian as a period of preservation failure be- cause although many Ordovician lines could be traced back to Middle Cambrian antecedents, few had described representatives from the Upper Cambrian. Of 80 species known from the Cambrian, only 12 (15%) are Upper Cambrian and of these, only 7 have formally been described. Similarly, of 40 named Cambrian genera only 7 (17.5%) come from the Upper Cambrian. Upper Cambrian echinoderms are thus considerably under represented. A second problem that is rarely considered is the strong geographical bias of the fossil record towards North America and western Europe. For Cambrian echinoderms 46 species occur in North America, 26 in Europe, 7 are Australian and one each is known from the U.S.S.R. and North Africa. Thus, excluding one species com- mon to both areas, 71 out of 80 (87.5%) species come from Europe and North America and only 9 (12.5%) from elsewhere in the world. This disparity must surely reflect the considerably greater amount of research effort that has been spent investigating Cambrian outcrops in western Europe and North America. Thus, al- though we may now have a fairly good idea of echinoderm diversity in the Cambrian of Europe and North America, there is great scope for new discoveries in suitable Cambrian sequences else- where in the world. Because Cambrian faunas outside Europe and North America are less well studied and because one of the few Upper Cambrian echinoderm- bearing beds is in Queensland, Australia, we felt that a search for echinoderms in the Cambrian of western Queensland might be highly rewarding. This paper is a direct result of a two-week field collecting trip and describes the edrioasteroid species that were discovered. A new carpoid from the Upper Cambrian of Queensland will described elsewhere (Smith & Jell in prep.). In addition we give a description of the primitive asteroid/ophiuroid antecedent Archegonaster pentagonus Spencer,1951 from the Lower Or- dovician of Czechoslovakia, as this has bearing on the phylogenetic diversification of Cambrian edrioasteroids. The first Australian Cambrian echinoderms described were Cymbionites craticula Whitehouse, 1941 and Peridionites navicula Whitehouse, 1941 from the early Middle Cambrian Thorntonia Limestone, north of Mount Isa, western Queensland. These were sub- sequently considered (Smith, 1982) to be the basal circlet of two different eocrinoids and our MEMOIRS OF THE QUEENSLAND MUSEUM collecting at the type locality in May 1989 con- firms this view. In 1971 Henderson and Shergold described Cyclocystoides primotica from the early Middle Cambrian Beetle Creek Formation, associated with Redlichia spp. This species is not a cyclocystoid (Jell et a/., 1985; Smith, 1986; Smith & Paul, 1982) and has been reassigned to the Edrioasteroidea and placed in its own genus Edriodiscus; it is elaborated upon below. Flem- ing (1977) showed the great abundance of dis- sociated echinoderm plates in the Lady Annie and Duchess phosphate deposits in the early Middle Cambrian Beetle Creek Formation, but made no taxonomic identifications. This material, collected from the vicinity of BMR locality D640 (Fig. 1), contains abundant, well- preserved disarticulated plates of edrioasteroids and is currently under investigation. Most of our knowledge of Australian Cambrian echinoderms derived from the work of Jell et al. (1985) who described Cambraster tastudorum and Ctenocystis jagoi from the medial Middle Cambrian part of the Cateena Group in northern Tasmania, isolated plates of Cambraster and probably Gogia from the early Middle Cambrian Coonigan Formation in western New South Wales, ?Stromatocystites sp. (=S. reduncus sp. nov. below) from the type locality of E. primotica, and the eocrinoid Ridersia watsonae and an unnamed isorophid (=Chatsworthia spinosa gen. et sp. nov. below) from the medial Late Cambrian Chatsworth Limestone near Chatsworth Homestead, 100 km north of Boulia, western Queensland. Thus the five taxa described below represent half of the known Cambrian echinoderm fauna of Australia. GEOLOGICAL SETTING OF LOCALITIES The Cambrian fauna described here comes from the eastern portion of the Georgina Basin in western Queensland (Fig. 1), Sedimentation in this epicontinental basin occurred from the late Precambrian to the Devonian, although not con- tinuously in any one area. Although the entire basin has been mapped geologically at a 1:125,000 scale, more detailed mapping has been confined to a few areas within the basin, mostly areas that were prospected for phosphate deposits in the 1960s and 1970s. The Burke River structural belt in the southeast has received considerable attention and the biostratigraphy of the echinoderm bearing Upper Cambrian Chatsworth Limestone in the area south of Lily Creek (Fig. 1) has been studied in great detail by CAMBRIAN EDRIOASTEROIDS FROM ALISTRALIA P Via TERRITORY 7 SSN S “i NS y 7) pe of l\\ o 7 NORTHERN "Pew nee 77 a——, - ~ -- --' 25°S Fig. 1. Locality map. A, showing extent of Cambrian outcrop in the Georgina Basin (diagonally lined), main roads, and centres of population in far western Queensland (redrawn from Smith. 1972), B, Comford Bore area on Yaringa Creek (from Henderson & Shergold, 1971). C, Chatsworth area (from Shergold, 1982, scale same as in B). D, Mount Murray area (from Jones & McKenzie. 1980). Triangles = water bores; stars = fossil localities mentioned in the text. Shergold (1982). He recorded pelmatozoan debris from a number of horizons and. at his locality K204, the eoerinoid aff, Macracystella sp. That fossil was later described as Ridersia warsonae by Jell et al., (1985), together with two specimens of an indeterminate tsorophid edrioasteroid. Further collecting by the authors at that locality has yielded many more specimens of Ridersia watsonae, the edrivasieroids Chatyworthia spinosa and Hadrodiscus parma and the edrioblastoid Cambrablastus enubilatus. Shergold (1982) described the Chatsworth Limestone in its type section, south of Lily Creek, as ‘dominantly sandy or silty pelletal skeletal yrainstone and packstone, with subor- dinate wackestone and clast grainstone; the en- vironment of deposition was high energy, shallow subtidal, intertidal and possibly aeolian; 718 MEMOIRS OF THE QURENSLAND MUSEUM CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA oral Gaver plate shesi floorina plates 719 ?hydropore ?qonopore Pic. 3. Stromatocystites reduncus sp. nov.; camera lucida drawing of plating in the oral region of the holotype f= flooring plate; ambulacra numbered [-V. cross-lamination and ripple marks are common: trilobites and echinoderms are predominantly dissociated and fractured skeletal parts are not uncommon; current winnowing and alignment of organisms widespread.’ Horizon K204 refers ta a4 m thick unit of thin-bedded limestone which is a pelletal grainstone containing’ rare ooids (Shergold, 1982, p. 97). Cross lamination occurs throughout this unit and shallow but large-scale channeling results in individual beds having limited lateral extent. Current alignment of fos- sils is often evident and we discovered a shallow channel infill, about 2 m by | m, with about 100 adult specimens of Ridersia watsenae lying in parallel alignment.Obvious hardground surfaces are absent. In addition to the echinoderms listed above, the fauna includes the trilobites /veria iverensis, Connagnostus sp., Peichiashania secunda, Prochuangia glabella, Pseudagnastus parvus and Wuhuia yilex, inarticulate and articu- late brachiopods. monoplacophorans, gastropods, sponges and un undescribed cothur- nocystid. Shergold (1982) placed this horizon within his Peichiashania secunda/Prochuangia glahella Assemblage Zone at the top of the [damean Stage, which he correlated internation- ally with the middle Franconian of North Amenca, The second locality from which we descnbe Cambrian echinoderms lies to the west of Mount Isa. At BMR locality QP243 (Fig. 1) on a low rise some 500 m south of Yaringa Creek at the bore some 3 km west of Cornford Bore situated on the north of Ardmore Station, 45 km west- southwest of Mount Isa (20° 49.57 S, 139° 03.5" BE) Henderson & Shergold (1971) discovered a single specimen with two individuals of Edriodiseus. primoticus. Here, laminated chert with silicified coquinite, algal chert and siltstone form the lower part of the Beetle Creek Forma- tion, testing unconformably on Precambrian quartzites. Accompanying these rare echinoderms are abundant exoskeletal fragments of the trilobite Redlichia. Opik (1970) described trom this horizon at the nearby M262 (Fig. 1) the species Redlichia versabunda, R. vertumnia and FiG. 2. Srraumatocystitey reduncus sp. noy., early Middle Cambrian from QP243 near Cornford Bore. A-C, (QMF17914, holotype. A, oral area, x7 (see Figs 3, 4A). B, oral surface, x17. C, aboral surface. x1.7.D. BMNH £63523, paratype, detail of oral surface with ambulacrum, margin of disc at bottom x3,Oral plating has been lost from the central band to reveul the internal surface of aboral plates. E, QMFI7920a, paratype, detail of oral surface showing interambulacral plating and, on the right hand margin, ambulacral flooring and cover plates, x3 (see Fig. 4B). All latex casts whitened with ammonium chloride sublimate. R, mayalis, and all three described morphologies are recognizable amongst our collection from QP243. Opik dated the horizon as late Ordian. immediately older than his Redlichia chinensis Zone, and early Middle Cambrian. A further specimen of Edriodiscus primeticus was collected and described by Jell et al. (1985) from this locality and on the same slab there was also a fragment of a Stromatocystites sp. We recollected from this locality and extended our search some 400-800 m further south along the ridge with QP243 at its northern end. This resulted in the discovery of an additional 35 specimens of the Stromatocystites and 5 specimens of Adriodiscus primoticus, described below. They occur in distinct layers associated with much disarticulated trilobite material. REPOSITORIES OF MATERIAL The material described below is housed in the following institutions: British Museum (Natural History), London [BMNH]; Commonwealth Palaeontological Collection, Bureau of Mineral Resources. Canberra [CPC]; Narodni Museum. Prague [NM]; Museum of Victoria, Melboume |NMVP]: Queensland Museum [QMF] SYSTEMATIC DESCRIPTIONS Class EDRIOASTEROIDEA Billings, L858 Stromatocystites Pompeckj., 1896 TYPE SPECIES Sirematocystites pentangularis Pompeckj, 1896; Middle Cambrian of Czechoslovakia and late Lower Cambrian of Newfoundland, DIAGNOSIS Edrivasteroids with fully plated aboral surface approximately as large as the oral surface. Un- differentiated oral mouth frame and cover plate series; biserial flooring plates and mulltiserial cover plate series. Interambulacra with well developed epispires. No differentiated peripheral rim plates. Interior of aboral surface with a series of divaricating ribs radiating from a central platform of larger plates. OTHER SPECIES S. walcom’ Schuchert, 1919 (late Lower Cambrian of Newfoundland), 8. reduncus sp, nov. (early Middle Cambrian of Australia). MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS Stromatocystites is the oldest and most primi- tive edrioasteroid known and is a possible an- tecedent to all later edriousteroid groups. The absence of differentiated peripheral rim plates separates Stramatocysntes from most other edrioasteroids and appears to be primitive for the class, as is the presence of epispires in interam- bulacra and the completely plated aborul surface, Although both previously known species have Straight ambulacra, the new species described below has curved ambulacra. This is also the condition seen in the even more primitive Lower Cambrian Camptostroma (Paul & Smith, 1984), although the direction of curvature is not the same. Stromatocystites reduncus sp. nov. Figs 2-9 Sromarocystites sp, Jell, Burre. & Banks, L985, p.192, figs 7e-f, 8. ETYMOLOGY From Latin reduncus curved back, in refer- ence to the shape of the ambulacra. MATERIAL Holotype QMF17914 part & counterpart; paratypes, QMF17915-25, 17938, 17944. BMNH E63519-22. E63525, Other material QMF17926-37,17939-43, 17944, BMNH E63523, 63524, NMVPLO7478. AGE AND O€CURRENCE From chert beds of the Yelvertoft Member. Beetle Creek Formation, early Middle Cambrian, exposed on the northern flank of alow rise 400-800 m south of the bore at Yaringa Creek. 50 km west-southwest of Mount Isa. Western Queensland (GR 980965). DIAGNOSIS A Stramatocystites with distally recurved am- bulacra; ambulacra | and VY curving clockwise, ambulacra LI, WI and LY anticlockwise. DESCRIPTION Specimens reach 53 mm in diameter and are rounded pentagonal in outline (Fig. 2B.C). The lower (aboral) surface is flat or slightly concave while the oral surface is moderately convex. Both surfaces are fully plated and there is no sharp demarcation separating oral from aboral surface or any distinct peripheral rim plating. CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA Ambulacra are long and slender, forming sharp, well-defined ridges over the oral surface and meeting centrally in a 2:1;2 arrangement (Figs 2A, 3). They are straight proximally but close to the edge of the oral surface they curve sharply so that the tips of the ambulacra come to face centrally (Figs 2B, 5C,D). Ambulacra J and V curve clockwise and ambulacra II, I -and IV curve anticlockwise. The recurved portion is about 1/3 of the radius in larger specimens. Flooring plates are biserially arranged (Figs 2A, 3). They are rather broad and short and there are large marginal sutural pores between adjacent flooring plates that almost reach the perradial suture. These pores lead to the interior of the theca, The outer edge of the flooring plates is exposed along the adradial margin of the am- bulucra (Pigs 2B, 3,4A.B) where they appear as a row of rectangular plates. Flooring plates are more fully exposed towards the distal ends of the ambulacra, Each flooring plate carries a large primary cover plate which is broader than tall with a blunt distal point. These cover plates sit directly on top of the flooring plates. Above the primary cover plates there is a sheet of smaller secondary cover plates without regular arrange- ment (Figs 2D, 4A,B). These are in two or three irregular rows and ate about 1/3 the size of the primary cover plates or less. Ambulacral plating decreases in size towards the arm tips. In the oral region the ambulacral series is undifferentiated (Figs 2A, 3), There are no en- larged flooring or cover plates and there appear to be four flooring plates along the postenor border of the oral area (Figs 2A, 3), The cover plate sheets continue across the oral area without differentiation. In the flooring plate closest to ambulacrum V there Is a crescentic opening which might represent the hydropore and a more ragged opening that might be the gonopore (Figs 2A. 3). Interarnbulacral areas are broad and composed of many plates. Towards the centre the plates are large (1-1.5 mm diameter) and stellate in outline with prominent epispires surrounded by a rim (Figs 2A, 3). The remainder of the plate surface has a reticulate omament but the rim and inner walls of the epispires are smooth (Figs 2D,E, 4C). The largest epispires (up to 0.5 mm diameter) are found centrally in each interam- bulacrum where they are oval in outline. Epispires decrease in size towards the margin of the oral surface and are more angular. either slit-like or, where developed at triple suture junc- tions, triangular in outline. Interambulacral | ta plates are smooth internally and haye nearly vertical sutures (Fig, 6A,B). The periproct is a cone of lath-shaped plates, some 20-30 in number, situated close to the margin on the oral surface mid-way between the two posterior ambulacra (Figs 2B, SC, 6B). Epispires reduce in size close to the periproct and are absent from the cone of lath shaped plates forming the periproct. There is no differentiated peripheral rim or marginal frame. At the edge of the theca, inter- ambulacra! plates become progressively smaller and rapidly merge into a narrow zone of small granular ossicles lying subambitally (Figs 5B, 7A,B. 8). This 2-3 mm wide band usually stands out as an elevated rim when seen from the aboral surface, suggesting that there might be some Fic. 4. Stromatocysrites reduncus sp. nov.; camera lucida drawings of plang. A. QMFI79]4a. holotype; lateral view of ambulacrum V at mid- length. B, QMF 17920a, lateral view of ambulacral planng. C, QMFI7927. two oral interambulacral plates in external view. cp=cover plate; fp= flooring plate; la p = interambulacral plate: 1° = primary; 2° = secondary, Scale bar = 1 mm, M J S OF THE QUEENSLAND MUSEL MEMOIR CAMBRIAN EDRLOASTEROIDS FROM AUSTRALIA 72 Pic, 6. Sromatocystites reduncus sp. noy, A, QMF17924. paratype, internal surface of aboral plates towards the margin of the dis¢e, x3.5, B, QMF17944, paratype, internal surface of oral plating in posterior interam- bulacrum with lath-like plates of the periproct evident, x4. Latex casts whitened with ammonium chloride sublimate. weak development of peripheral plates internally forming a more robust frame to the theca, On the aboral surface. inside the marginal zone of granular plates, the majority of the surface consists of a pavement of oval to sub-palygonal plates surrounded by rings of small granular os- sicles (Figs 5A,B, 7A,B, 8). The large plates are {5-2 mm in diameter and become larger towards the centre. They have a distinctive reticulate pattern of fine radiating ribbing (Fig. 5B). At the centre there is a small region of larger plates lacking interspersed granular plates around their borders. These are in no particular arrangement and there is no clear central plate (Figs 5A, 7A.B.E, 8). The interior of the aboral surface is very distinctive. There is an outer zone com- posed of rather thick stellate plates, resembling the epispire-bearing plates of the oral surface (Figs 2D, 6A). However. the spaces between the large plates are filled with small. thin platelets thal appear deep down in the depressions. This ouler zone is best developed interradially, Towards the centre, plates develop pronounced sharp keels that run radially and occasionally branch and merge (Figs 7C.D.F. 9). This zone of ribbed plates is extended radially to create a stellate pattern. Ribbing becomes stronger towards the centre and ends against an elevated platform of Some 12-15 large polygonal plates bounded by alow peripheral ridge (Figs 7C,D.F, 9). The plates around the edge of this platform have radiating ridges on their distal half, between 2 and 6 major ridges per plate (Fig. 9). These ndges ure continuous with the radial ribs on the more distant plates. The outer boundary of inter- nally ribbed aboral plates corresponds to the elevated pentagonal area that is seen from the exterior, REMARKS The incomplete specimen of Jel! ef al. (1985, figs 7C-F, 8B) from this locality could only tentatively be assigned to Stromarocystites, Jell etal, (1985) noted the internal ridging to aboral plates which had previously been unrecorded trom any stromatocystitid. However, we now know that a similar, but less well-developed, system of ridges occurs internally in the type species, §, pentangularis (Fig. 10), Re-collecting from this locality has produced a further 35 specimens and it is now the best known of all Siromatocystites species. It differs from the other two species of Stromatacystites in having distal- ly curved ambulacra. The oral area of the holotype is well-preserved and allows the plating of this region to be described in detail for the first time in Siramatoeystites. The crescentic opening here tentatively identified as the hydropore lies across a suture between two flooring plates and thus Fig. 5. Stromatoevstites reduncus sp. nov. A,B. QMF17916, paratype. A, aboral surfaces of two individuals, x L.4. B, detail of external ornament an aboral plates, x4. C/ BMNH E63519, paratype, oral surface showing ambulacra | and II, and the periproci, x2. BD, QMF19738a, paratype, oral surface in external aspect with two curved ambulacra and, in the upper half, the interior of the aboral surface. x2. Latex casts whitened with ammonium chloride sublimate. EMOIRS OF THE QUEENSLAND MUSEUM M 724 CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 725 ambulacrum marginal zone of granular plating plates of central disc FiG. 8. Stromatocystites reduncus sp. noy., QMF17925, paratype, camera lucida drawing of plating on the exterior aboral surface. Fic. 7. Stromatocystites reduncus sp. nov. A, BMNH E63522, paratype, aboral surface, exterior, x2. B, QMF17925, paratype, aboral surface, external, x1.8 (see also Fig.8). C, QMF17917a, paratype, aboral surface, interior, x2.3. D, BMNH E63521, paratype, aboral surface, interior, x1.7. E, QMF17919a, paratype, aboral surface, exterior, showing the central zone of larger plates towards the top, x3. F, QMF17915, paratype, aboral surface, interior, central part only, x4 (see also Fig. 9). Latex casts whitened with ammonium chloride sublimate. hr outer Stellate plates am MEMOIRS OF THE QUEENSLAND MUSEUM ribbed plates Fic, 9. Srromaroeysrires reduncus sp. nov, QMF17915, paratype, cameca lucida drawing of plating in internal aspect of the aboral surface (see also Fig. 7F). am=ambulacrum: ia = interambulacrun, resembles the arrangement in Toliglobus (see Bell & Sprinkle. 1978) and edrioasterids, The central part of the oral area was roofed over by what appears to have been a conunuous cover plate sheet and it seems probable that the mouth was permanently covered in §, reduncus. An unusual feature of the oral area ts the set of four small oval holes in the coyer plate sheet of the Posterior interray- The aboral system of internal ridges radially arranged around a central platform of plates is particularly intriguing from a functional point of view. A similar arrange- ment is seen in Toriglohus, although its en- tire aboral surface is reduced (o a small disc and the ridges occupy virtually the entire outer part. In Stromatocystites the exterior of the aboral surface usually shows # central stel- late region that is elevated above the more peripheral interradial 4ones and the outline of this elevated region corresponds to the outer border of the internally ribbed region, The invaginated appearance df interradial yones on the uboral surface and the radial arrangement of the surrounding ridges sug- gests that this structure might be developed for adhesion, The dise of Tor/g/obus undoub- tedly had to have had some form of aboral adhesive dise since its globose shape would have made it very unstable otherwise. Thus CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 727 FIG. 10. Stromatocystites pentangularis Pompeckj, New York State Museum E1673, from the Middle Cambrian of Ginetz, Czechoslovakia. A, aboral surface, exterior, showing the central zone of large polygonal plates, x3. B, aboral surface, interior, of a second specimen showing the central platform surrounded by a stellate zone of ribbed plates, x3, Latex casts whitened with ammonium chloride sublimate. it seems reasonable to interpret the aboral inter- nal structures in Stromatocystites as part of an adhesive disc. We suspect that the ridges mark the site of attachment for radial muscles and that these muscles were employed to elevate the central part of the disc to create a suction pad. The distal attachment point for the muscles is, however, weakly developed in Stromatocys- tites, although in Totiglobus there are stout peripheral rim plates to which muscles could attach. Possibly a marginal frame of peripheral rim plates arose to provide ngidity and to secure anchorage for the radial aboral muscles,andis connected with the evolu- tion of stronger adhesion in edrioasteroids. EENSLAND MUSEUM MEMOIRS OF THE QU 728 Pe a ay iy ee eae vee oye * s ee CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA marginal ring 729 ambulacrum Fic. 12. Edriodiscus primoticus (Henderson & Shergold), QMF17856, camera lucida drawing of plating in a juvenile. A, aboral surface. B, oral surface. (see also Fig. 11C,D). Edriodiscus Jell, Burrett & Banks, 1985 TYPE SPECIES Cyclocystoides primotica Henderson & Shergold, 1971, by original designation. AGE AND DISTRIBUTION Early Middle Cambrian of Queensland. DIAGNOSIS Disc-like edrioasteroids with a ring of more than 40 marginal ossicles which are an integral part of both oral and aboral plated surfaces. Am- bulacra extending over the marginal ossicles onto the peripheral rim. REMARKS This genus resembles Cambraster in having a well-developed marginal ring and ambulacra that extend onto and over this ring. Unlike Cambraster, however, it has a great many more marginal ossicles in the ring and these are tesse- lated into the aboral plated surface. In Cambraster marginal ossicles overlie the aboral plated surface and are not visible from beneath. Furthermore, the flooring plates of Edriodiscus are imperforate and the oral area resembles that of Stromatocystites more than Cambraster. In particular we have seen no evidence of there being a mouth frame of five large interradially positioned ambulacrals, which always appears prominently in Cambraster (Smith, 1986). Jell et al. (1985) had only a single incomplete oral surface when establishing this genus. Fur- ther collecting has produced an additional five specimens, two of which show oral surfaces. Edriodiscus primoticus (Henderson & Shergold, 1971) Figs 11-14 Cyclocystoides primotica Henderson & Shergold, 1971, p. 706, pl.138, figs 1-3. Fic. 11. Edriodiscus primoticus (Henderson & Shergold), early Middle Cambrian from QP243 near Cornford Bore. A,B, QMF17855a. A, general view of oral surface, x5. Note that much of the oral interambulacral plating has been lost to reveal the internal surface of aboral plates (see Fig. 13). B, enlargement of one ambulacrum to show the region of adradial plate alignment perpendicular to the ambulacral groove, x10. C,D, QMF17856, juvenile. C, aboral surface, x7. D, oral surface, x8. (see also Fig. 12). Latex casts whitened with ammonium chloride sublimate. 730 Sy ambulacral groove tM ya peripheral skirt RO Dp kone af ote OE marginals [Ao r f \ Oats oun \ — ant -—==s | - = : = MDM. WV BEATA ==> hy interior of aboral plate: ie iat ry —— 4 t. iw, interamb ae = z — plating“ == = =< S =: = = tlooring ve plates. = My aoa Cs Fic. 13. Edriediscus primoticus (Henderson & Sher- gold), QMF17855a, camera lucida drawing of oral surface. Much of the interambulacral plating has been lost to reveal the internal surface of aboral plates (horizontal lines). ‘Cyclocystoides’ primotica Henderson & Shergold: Smith & Paul, 1982, p. 677. Edriodiscus primotica (Henderson & Shergold); Jell etal., 1985, p. 190, figs 7a-c, 8. Edriodiscus primotica (Henderson & Shergold); Smith, 1986, p. 753, fig. 20. MATERIAL Holotype CPC11395; paratype CPC11396; other material NMVP107479, QMF17855-7, BMNH E63525,63526. AGE AND OCCURRENCE From chert beds of the Yelvertoft Member, Beetle Creek Formation, early Middle Cambrian, exposed at locality QP243 (=NMVPL1598) 400-800 m south of the bore at Yaringa Creek, 50 km west-southwest of Mount Isa, western Queensland (GR 980965). DIAGNOSIS As for genus. DESCRIPTION Individuals 9-35 mm in diameter. All are cir- cular in outline. The smallest has c. 55 marginal ossicles in the ring. These are differentiated from other plates and are longer (radially) than broad (Figs 11C,D, 12). Larger individuals have a similar number of marginal ossicles (50-60) but these are wider than long. At all sizes the mar- MEMOIRS OF THE QUEENSLAND MUSEUM ginal ossicles are an integral part of the aboral plated surface. Oral plating attaches to a weak ridge at the upper edge of the inner face of the marginals so that much of the inner face of the marginals must have been internal. The peripheral skirt is proportionally larger in small individuals but contains approximately the same number of plate rows at all sizes examined. At 9 mm diameter 4 irregular rows of plates make up the peripheral skirt, while at 15-35 mm diameter there are 4 or 5. Ambulacra are straight and meet centrally in a 2:1:2 pattern (no specimen shows all five am- bulacra but the best oral areas (Figs 1 1A, 13, 14) show two lateral ambulacra converging into a single ambulacral groove leading to ambulacrum III). Plating in this region shows no organization (Fig.14), suggesting that the cover plate sheet may have formed a permanent roof to the mouth. The ambulacra are straight and extend onto the marginal ring, where they overlie the marginal ossicles at a suture. In the smallest individual (9 mm diameter) the tip of the ambulacrum does not reach the outer margin of the marginal ring (Figs 11D, 12B) whereas in the largest the am- bulacrum completely covers the marginals ex- tending beyond to the edge of the peripheral skirt (Figs ILA, 13), No other oral plating extends beyond the marginal ring as far as we can tell. Ambulacral flooring plates are biserial and steeply inclined forming a deep, narrow groove. They are about 0.5 mm broad and a small part is exposed adradially. Sutural pores are not evident along the walls of the ambulacral groove (Fig. 13). Cover plates are small and multiple, except possibly towards the ambulacral tips in the smal- lest individual where there appear to be only primary cover plates (Figs 11D, 12B). A multi- plated skirt of cover plates roofs the ambulacra close to the oral area. The structure of the oral frame remains unknown but the oral area is similar to that of S. reduncus (cf. Figs 11A, 13 with Figs 2A, 3). Interambulacra are composed of large plates that are oval or subrounded in outline. Inter- spersed amongst these are smaller platelets of various sizes. There are no epispires or sutural gaps in the interambulacra (Fig. 11A,D). Platelets form an outer band close to the marginal ring and are also developed to the exclusion of larger plates close to the oral area (Figs 1 1A, 13). In the largest specimen the platelets close to the ambulacra are rod-like and preferentially aligned perpendicular to the ambulacral groove (Figs 11B, 13). Elsewhere platelets are randomly ar- CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA Fic, 14. Edriodiscus primoticus (Henderson & Sher- gold), NMVP107478, camera lucida drawing of plat- ing in the oral region. Note the apparently disorganised arrangement of cover plates. ranged. In juveniles (9 mm diameter) large plates dominate the interambulacral regions with little development of platelets (Figs 11D, 12B). This suggests that the large plates are laid down ini- tially and that, during growth, areas of platelets developed preferentially around their interam- bulacral margins. The aboral surface is a tesselate pavement of thick plates, In the smallest individual (Figs 11C, 12A) a large central plate is surrounded by a narrow ring of platelets. No circlets of plates can be recognized outside this region, The interior of the aboral surface is smooth and also forms a tesselate pavement (Figs 11A, 13). There is no sign of internal ribbing, as seen in Stromatocys- tites or Totiglobus, or of a central thickened platform. REMARKS This species was established for two specimens, both showing aboral surfaces. Henderson & Shergold (1971) interpreted this species as a cyclocystoid, but Smith & Paul (1982) later removed it from the cyclocystoids, interpreting it as a probable edrioasteroid. This was confirmed by Jell et al. (1985) when they discovered and described a third example showing part of the oral surface. Further collecting by the authors has yielded 5 more specimens, 3 of which are part and counterpart, show- ing details of both surfaces. The above description supplements those given previously. Alignment of rod-like interambulacral plates bor- dering the ambulacra is noteworthy because of its resemblance to the arrangement of virgals in Archegonaster. 731 Order ISOROPHIDA Bell, 1976 Previously edrioasteroids have been divided into three orders, Stromatocystitida, Isorophida and Edrioasterida (Bell, 1980). Stromatocystitids have a fully plated theca, biserial ambulacra, flooring plates with sutural pores, and interam- bulacral plates with epispires. Isorophids have an uncalcified aboral surface, a differentiated peripheral ring and uniserial flooring plates, while edrioasterids have biserial flooring plates with an exposed adradial portion, no differen- tiated peripheral ring and a partially or complete- ly calcified aboral surface. A fourth group, the Cyathocystida was recently erected to accom- modate turret-shaped edrioasteroids with single large ‘deltoid-like’ flooring plates (Bockelie & Paul, 1983). Smith (1986) modified this scheme somewhat by rejecting the Stromatocystitida as a paraphyletic ancestral group. The two Upper Cambrian edrioasteroid genera described below show mixed characteristics and appear to bridge the gap between Lower and Middle Cambrian stromatocystitids and the Ordovician and later isorophids. Synapomorphies shared with later isorophids include the disc-like body form, the imbricate plated peripheral rim, the form of the hydropore, the absence of aboral plating and the presence of simple cover plates rather than a cover plate sheet. Because of these features they are best regarded as primitive isorophids. How- ever both Hadrodiscus and Chatsworthia may have biserial flooring plates and the former has an oral frame of five interradially positioned deltoid plates of ambulacral origin. Neither have the typical uniserial flooring plates of Or- dovician isorophids as far as we can tell. The uniserial flooring plates of later Isorophida could have arisen in one of three ways. Firstly, each uniserial flooring plate could be the result of fusion of opposite pairs of originally biserial flooring plates. Secondly each uniserial flooring plate could have originated from a single biserial flooring plate by extending the amplitude of the V-shaped perradial suture until it reached the opposite edge, (the reverse of the suggested derivation of biserial from uniserial arm plating in crinoids - viz. Bather, 1900). Finally the uniserial flooring plates could represent a new structure not homologous with biserial flooring plates. The Upper Cambrian species described below show no evidence of extended V-shaped perradial sutures, having al- most straight perradial sutures instead. Further- more, the second hypothesis would require a ~ secondary series of primary cover plates and cover plate articulations to develop, No eviderice exists to suggest that this might have been the case, in contrast, for example, to the situation with uniserial crinoid urms, where sequential uniserial arm plates carry cover plates on alter- nate sides. Of the two genera described below Hadrodis- cus appears, tentatively, to be the more primitive, havinga poorly differentiated peripheral rim and small, closely fitting pentagonal cover plates. Flooring plate arrangement ts inadequately known but may have been biserial, In Chatswor- thia the peripheral nm plating is much better differentiated, as in most later isorophids, and there is a system of stout, spine-like cover plates that are non-contiguous when held erect. A similar cover plate arrangement ts found in early lebetodiscids such as Feerstediscus for example, However, in Chutswerthia the relasonship of cover plaies to flooring plates is still unclear. Either cover plittes ure firmly sutured on to the Mooring plates so that both generally act as a single ossicle. or the primary cover plates of later isorophids are in fact modified flooring plates that have taken on the role of protecting the radial water vessel. [In either case, it would appear that the uniserial flooring plates in more derived isorophids represent a new system of plates in- dependent of the biserial flooring plates of other edrioasteroids, Hadrodiseus gen. nov. ETYMOLOGY Greek hadros. elegant or delicate. TYPE SPECIES Fladrodixcus parma sp. noy. AGE AND DISTRIBUTION Median Upper Cambrian (Franconian) of western Queensland, Australia. DIAGNOSIS An isorophid with a poorly differentiated peripheral ring. straight ambulacra with small, lightly fitting cover plates and larger oral cover plates. Hydropore/gonopore a large, prominent opening, Oral frame composed largely ot stout, wing-shaped elements, REMARKS. The taxonomic position of MHadrodiscus remains problematic largely because itis incom- 3 MEMOIRS OF THE QUEENSLAND MUSEUM pletely known. Although the ambulacra are floored We clo not yet know what the plate arran- gement is. The relatively small size of the cover plates seen externally and the suggestion of furge, possibly biserially arranged plates forming the floor of the ambulacral grooves suggest to us that Hadrodiyeus had both cover plates and floor- ing plates, as in Srrameatocystites. However, the presence of a peripheral rim, the complete ab- sence of aboral plating, the position and form of the hydropore opening and the large oral frame plates that are interradial in position and are undoubted flooring plates, mark Hudrodiscus as more advanced than Stromeatocystites. Tt is less derived than Chatywerthia which has a system of large biserially arranged cover/flooring plates and a better ditferentiated peripheral rim. At present we consider it to be the most primitive known isorophid. Hadrodiscus parma sp. nov. Figs 15-18, 20C ETYMGLOGY Greek parme, a small shield, in allusion to its appearance. MATERIAL Holotype QMF1 7874; paratypes QMF17875, BMNH E63527, AGE AND OCCURRENCE All specimens come from locality K204 of Shergold (1982), from a grey, 4 m thick. lime- stone unit of the Chatsworth Limestone, 3.5.km south of Chatsworth homestead, 60 km south- east of Duchess, western Queensland, Peichiashania secunda-Prochuangia glabella Assemblage Zone, median Upper Cambrian (Franeonian). DIAGNOSIS: A Hadrodiscus with about 10-12. primary cover plates ina column at disc diameters of 6-7 mm, these primary cover plates being small, squarish and close-fitung. Peripheral rim corn- posed of 4 or 5 irregular rows of small imbricate plates not clearly differentiated from interam- bulacral areas. DESCRIPTION Specimens are circular in plan view and flat- tened hemisperical in profile. In diameter the holotype is c. 7 mm (Figs ISA. 16) and the two other specimens c. 6 mm (Figs [5B,C, 20C). The CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 733 Fic. 15, Hadrodiscus parma sp. noy., from the Chatsworth Limestone, median Upper Cambrian of Chatsworth, A, QMF17874, holotype, oral surface, x12 (see also Fig. 16). B,C, BMNH E63527. paratype. B. external oral surface, x10. C, internal view of same, x10 (see also Fig. 18). Silica replacements coated with ammonium chloride sublimate. aboral surface is uncalvified und the peripheral rim of the dise Mares outwards, There has been no post-mortem collapse of the disc relative to the peripheral rim so that the boundary between the two regions is not well demarcated, there being virtually a smooth transition fram rim to centre. The oral area forms the summit of the dise and is centrally positioned, Specimens appear to be unattached to any substratum. The peripheral rim ts composed of 4 or 5 irregular rows of small inbricate plates, 3 or 4 abreast in one interambulacral segment (Figs 15A, 16), These plates overlap towards the centre on their outer face. and have convex proximal flooring plates MEMOIRS OF THE QUEENSLAND MUSEUM edges. However, sutures are Weakly defined sug- gesting that the plate boundaries, though imbri- cate. were almost flush and thal the plates themselves did not override more proximal plates to any extent. The same is true of the inner face where no plate boundaries can be made out at all. None of the marginal plates is geniculate or has a thickened aboral edge. Instead the peripheral rim remains uniformly thin and only the base of the rm would have contacted the substratum. Plates in the rim are similar in size except towards the periphery where the outer- most plates are much smaller, The boundary between peripheral rim plating and the interam- peripheral rim cover plates hydropore oeriproct FIG, 16, Hadrediseus purma sp. noy., OMF17874, holotype. camera lucida drawing of the oral surfave (see alsa Fig. ISA), CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA Fic. 17. Hadrodiscus parma sp. noy., QMF17874, holotype, interpretation of cover plate arrangement. I-V = cover plates of ambulacra I - V; o = shared oral cover plates; ahp = anterior hydropore plate; php = posterior hydropore plate; scp = shared cover plate. bulacral plates of the disc is not marked, interam- bulacral plating appearing to merge gradually into that of the peripheral rim, except in the posterior interambulacrum. Again, plate boun- daries here are probably subtesselate as none stand out clearly (Figs 1SA,B, 20C). The ambulacra are relatively long and reach about 80% of the radius to the edge. They indent the peripheral rim plating to a small extent (Figs I5A, 16). In the holotype, ambulacra I, II] and V are straight, or virtually so, while ambulacrum II curves clockwise and ambulacrum IV curves anticlockwise (Figs ISA, 16). However, this is probably a post-mortem artefact since the disc has collapsed towards the anterior, and the 3 adjacent ambulacra (probably I, If and III) in one of the paratypes (Fig. 20C) are straight. The ambulacra are parallel-sided and have rounded, blunt tips (Figs 15A,B, 20C). In the holotype cover plates are present in all ambulacra except II, where they have been lost (Fig. 16). In am- bulacra I and V the cover plates are in the open position, in ambulacra [V they are in the closed position, while in ambulacrum III they are large- ly disrupted from life position. Cover plates were able to open widely along the length of the arm but probably formed a permanent cover to the oral area, as no clear perradial suture is apparent in this region (Fig. 15A,B). There are about 10-12 primary cover plates in a column at this diameter. The primary cover plates abut along most of their length and end in a V-shaped edge which may be hooked towards the oral region (Figs 15A, 16, 20C). Cover plates on either side of the ambulacrum alternate to interlock, al- though those seen in ambulacrum IV of the holotype are almost opposite due to distortion of the disc. Sutural gaps between adjacent cover plates are not apparent. Ambulacral grooves are moderately deep with a smooth rounded floor. Sutures are not visible suggesting that flooring plates were probably solid and robust although this appearance may be due to their coarse silicification. No part of the flooring plates can be seen along the adradial margin of the ambulacra and it would appear that the interambulacral plating abuts against the outer edge of the cover plates. In ambulacrum I] of the holotype cover plates have been lost revealing the ambulacral groove (Figs 15A, 16). Upper edges of flooring plates define the walls of this groove but again the detailed structure has been lost in silicification. The internal surface of one specimen (Fig. 15B,C) reveals the 5 ambulacral grooves. Around the mouth these are floored by large deltoid-shaped plates lying interradially and forming the mouth frame (Figs 15C, 18). Else- where ambulacral grooves are partially open (possibly due to incomplete silicification) al- though larger plates adjoin the ambulacral groove and may have floored the groove in life. If these plates are flooring plates then each plate must have carried 2-4 of the cover plates. In external view the oral area is best seen in the holotype (Figs ISA, 16) where the cover plates form an unbroken roof to the mouth. Plate sutures are nowhere clear and our interpretation of oral plating is highly tentative. The hydropore, however, is distinct as a deep depression in the posterior interambulacrum close to ambulacrum V (Figs 15A, 16). It has a narrow, bar-like am- bulacral cover plate on its adradial margin and 2 moderately large plates forming the posterior border. These posterior plates lie well below the ambulacra and would appear to be modified ambulacral flooring plates that have been en- larged into the posterior interambulacrum to incorporate the hydropore/gonopore. The remainder of the oral area consists of a number of small plates, to judge from the surface ir- regularities preserved. Left and right branching plates are easily identified but the plating in the 736 stone canal Fic. 18. Hadrodiseus parma sp. nov., BMNH E63527, paratype. camera lucida diagram of intenor of oral surface showing the mouth frame plates and arrange- ment of the water vascular system, Scale bar =1 mm, posterior part 1s less clear. A continuaus row of small, squarish cover plates may form the posterior edge of the oral area and connect the cover plate series of ambulacra | and V (Fig. 16). Conversely, this may be a single laterally elon- gale posterior oral plate thatis slightly fractured, Other plates lie anterior of this but no enlarged anterior oral plates are clearly differentiated. The interior of the oral surface (Figs 15C, 18) shows the 5 ambulacral channels around a central mouth, A sixth groove originates close ta the base at ambulacrum V and rapidly curves towards the outer surface: this is the hydropore groove and terminates externally at the hydropore. At the angle between ambulacra IV and VY is a large deltoidal plate. This has a median saddle and may be composed of 2 plates tightly sutured together, but the coarseness of the silicification prevents us from deciding whether this is indeed a single plate. In the posterior interray there appear 10 be 2 large polygonal plates, the plate closer to ambulacrum L having a prominent adoral projection. Other mouth frame plates are not evident and are presumably lost (possibly during etching). Interambulacral plating consists of a small number of apparently subtesselate plates (Figs I5A, 16). These have curved adoral plate boun- daries close to the peripheral rim but become more polygonal in outline towards the centre, MEMOIRS OF THE QUEENSLAND MUSEUM Only traces of sutures are seen in both specimens, A large periproct occupying most of the posterior interambulacrum (Figs 15A,B, 16) consists of a mound-like region of imbricate plates which become progressively smaller towards the centre. The middle of the periproct consists entirely of small platelets. REMARKS This species 4s readily distinguished from the contemporary Chatswerrhia spinosa by its smaller, more tightly fitting and more numerous primary cover plates, its more numerous and smaller peripheral im plates and more robust suluring between peripheral rim and interam- bulacra, In all specimens of C. spinosa the dise has collapsed well below the level of the peripheral rim and presumably marks a weak point. The marked difference in appearance of the primary cover plates between C. sprnesa and H. parma makes it most unlikely that the two forms simply represent different states of preser- vation of the same species (cf. Fig. 20A.B with Fig. 20C). The distinct groove leading from the cireum- oesophageal ring of the water vascular system and opening close to the base of ambulacrum V in the posterior interray proves that this opening is a hydropore. Smiley (1988) speculated that primitive echinoderms might have internal hydropore openings and that the opening generally referred to as the hydropore might rep- resent the gonopore. Chatsworthia gen. nov, ETYMOLOGY For Chatsworth Station, on which the specimens were discovered, TYPE SPECIES Chatsworthia spinosa sp nov. AGE AND DISTRIBUTION Median Upper Cambrian (Franconian) of Queensland, Australia. DIAGNOSIS An isorophid with short, virtually straight am- bulacra and undifferentiated oral cover plates. Coyer plates stout and erect,forming a single ulternalmg biseries; well separated when open. CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA Biserial arrangement of flooring plates meeting perradially to floor the ambulacral groove. REMARKS This genus 1s distinguished from Hadrodisens in the size and spacing of primary cover plates and plating of the peripheral rim and in the structure of the oral frame and flooring plates. In C_ spinosa there are only 6 or so primary cover plates ineach ambulacrum and the peripheral rim is composed of a small number (usually 10) of large plates with just 2 rows of small, outer platelets, In Hadrodiscus parma the primary cover plates are more slender and closely spaced, with about 12 in a row at an equivalent dise diameter, and the peripheral ring is composed of a-greater number of relatively smaller imbricate plates. C. spinesa is not sufficiently well preserved for the structure of the ambulacra to be determined unambiguously, but it appears to consist of a biseries of plates that meet perradial- ly to floor the ambulacral groove. Above and possibly continuous with these are stout, spine- like, cover plates. undoubtedly homologous to the primary cover plates of later isorophids. Chatsworthia spinosa sp, noy, (Figs 19. 20A,B.21-2%) Isorophid indet. Jell, Burrett & Banks, 1/985, p.195, fig. ¥. Fry MoLocy Latin spinasus, thomy, referring to ils cover plate series. MATERIAL Holotype QMFI7944; paratypes QMPF 1794541. NMVP107129, 108990, BMNH E63528-30. Other material OMF17952-9, BMNH E63531. AGE AND OCCURRENCE All specimens come from locality K204 of Shergold (1982), from a grey, 4 m thick, lime- stone unit of the Chatsworth Limestone, 3.5 km south of Chatsworth homestead, 60 km south- east of Duchess, western Queensland. Peichiashania secunda-Prochuangia glabella Assemblage Zone. median Upper Cambrian (Franconiin), DIAGNosis A Cheatsworthia with 5 or 6 stout, spine-like caver plates in each ambulacrum and a —J eA ~4 peripberal rim composed of c. LO large plates plus smaller accessory plates distally. DESCRIPTION Theca circular in outline with a diameter of 6=LO mm. The peripheral rim is well developed and differentiated trom the disc, which in all specimens has collapsed slightly beneath the level of the rim (Fig. 19), The theca has a low profile, though in life the disc may have been low domed, Only the oral surface is heavily plated. The aboral surface, where seen, shows no evidence of having hud large plates. How- ever, the surface always appears granular (Fig. 26) and there remains the possibility of a cal- cified lower surface of tiny granules. The petipheral rim is relatively broad, forming up to half the radius of the theca iftertadially, and indented radially forihe ambulacra (Figs (9B, 21). In profile the rim slopes steeply outwards at about 40°. The rim is usually composed of 5 large radial and 5 large interradial plates which alternate and overlap one another (Figs 21. 25), bul the arrangement is not strict and there may be an extra plate in one or other of the interambulacral regions, Peripheral rim plates have curved outer edges and are dif- ferentiated from interambulacral plates of the disc. On the distal part of the rim there are two irregular rows of smaller plates, the outermost series being the smallest (Fig, 27A). These also appear to imbricate. although plate sutures are rarely well marked. These plates may completely overlie the 10 larger plates and are commonly lost. When viewed from the interior the peripheral rim appears smooth and no plate boundaries can be identified. This would suggest that the plates are subtesselate and firmly bound together along sutures and explains why the peripheral rim is generally preserved in. an uncol- lapsed state, Peripheral rim plates are neither geniculate nor expand into a flattened base. Thus the rim has a uniform thickness and only the bottom of the rim would be in contact with the substratum. Ambulacra are straight. rather short, and broad, They extend to the peripheral nm which they indent but do not overlap (Figs 19.20A,B), The 4 to 6 large, spine-like cover plates in each ambulacral column (Figs 21-25) have # broad base and taper distally (Fig. 27B.C) and in some cases tips of lhe cover plates appear to be curved towards the centre of the disc, Cover plates may be preserved in the closed position with the am- bulacra appearing as tall, rather narrow ridges MEMOIRS OF THE QUEENSLAND MUSEUM 738 CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 739 (Fig. 19A,C). In such cases cover plates are channel. They alternate across the perradius. inclined towards the centre and overlap their More often specimens are preserved with the neighbours forming a canopy to the ambulacral ambulacra open and cover plates erect and form- Fic, 20. A,B.D, Chatsworthia spinosa sp. nov, A, QMF 17949, paratype, x7. B, NMVP108990, paratype, x6. C, Hadrodiscus parma sp. nov., QMF17875, paratype, x12. D, 7Hadrodiscus sp. QMF17943. internal surface, x7. Silica replacements from the Chatsworth Limestone. median Upper Cambrian, of Chatsworth, whitened with ammonium chloride sublimate. Fic. 19. Chatsworthia spinosa sp. nov., from the Chatsworth Limestone, median Upper Cambrian of Chatsworth. A, BMNH E63528, paratype, x8 (see also Fig. 22). B, QMFI7944, holotype, x7 (see also Fig. 21). C, NMVPI107129, paratype, x6.5 (see also Fig. 23). D, QMF17948, paratype, x8 (see also Fig. 25). E, QMF17945, paratype, x7, F, QMF17947, paratype, x9, Silica replacements coated with ammonium chloride sublimate. 740 periproct MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 21, Chatsworthia spinosa sp. nov., QMF17944, holotype, camera lucida drawing and plate interpretation diagram. Vertical hatching = peripheral rim plating; horizontal close hatching = interambulacral plating; stippling = ambulacral zones undifferentiated; blank = cover plates; black = missing. Ambulacral cover plates are labelled as follows: I-V = cover plates of ambulacra I - V, bp = lateral branch plate, o = shared oral plate, hp = hydropore plate. ing a border to the food groove (Figs 19B,E.F, 20B). In such cases the cover plates are well separated leaving broad gaps between. Spine- like cover plates are less well-developed over the oral area. The ambulacral groove itself may be preserved in the open position, as a broad U- shaped channel, or in the closed position, as a narrow slit-like trench. This alone indicates that the ambulacrum is floored by a biseries and in cross-section, although a perradial suture cannot be identified due to the coarse nature of the silicification, a discontinuity of shape indicates where it probably lay (Fig. 27B). Furthermore, from the interior the perradial line appears as a sharp ridge and in one specimen (Fig.26) the interior of the ambulacra show possible biserial flooring plates. However, the arrangement of flooring plates is nowhere clearly seen, due to the coarse nature of the silicification, and the junc- tion between cover and flooring plates remains problematic. In the oral area, the spine-like plates become less evident and cover plates appear more plate- like. Coarse silicification again hinders inter- pretation of oral plating. The arrangement of plates is probably best seen in the holotype (Fig. 21) and two of the paratypes (Figs 22, 23). The ambulacra meet in a 2:1:2 pattern with small lateral branch plates at the inner angles of am- bulacra I and II, and IV and V. No noticably enlarged oral plates are developed. A row of 4 cover plates forms the posterior part of the oral area, the outer 2 abutting the lateral branch plates. In BMNH E63528 (Fig. 22) only a single large plate occupies this region. This could rep- resent an oral frame plate devoid of its two cover plates. The central 2 posterior plates lie opposite 2 anterior cover plates and roof the central part of the oral area. No secondary or accessory cover plates can be distinguished, but this could be due to poor preservation. The oral frame, if our inter- pretation is correct, is composed of 3 large floor- ing plates(Fig. 22), 2 anterior plates occupying the angles between ambulacra II and III and between III and IV, and a single broad posterior plate. It is not known whether there are also enlarged flooring plates between the lateral am- bulacra that also contribute to the oral frame. The hydropore/ gonopore opens in the posterior in- terradius close to the base of ambulacrum V. There is a large U- shaped plate that forms the posterior and interradial border to the opening and a narrower plate forms its adoral border. CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 74) FiG, 22. Chatsworthia spinosa sp. noy.. BMNH £63528, paratype, camera lucida draw ing and plate interpretation diagram. Symbols as in Pig. 21. Both have the position and appearance of being modified ambulacral floormg plates, When viewed from the interior (Fig. 26) the oral area stands out as a C-shaped ridge open towards the posterior. The ridge on the hydropore side lies more distal and creates a more open curve than the ridge on ambulacrum [. The posterior oral flooring plate must therefore have considerably Jess depth than the other oral frame plates. Interambulacral areas are relatively small and wedge-shaped. No specimen shows undisrupted plating but small ovoid plates can be discerned ina number of specimens. There is no evidence of epispires between interambulacral plates which were presumably either imbricate or sub- tesselate. The penproct is a conical structure occupying much of the posterior mteram- bulacrum (Figs 19, 21, 24, 25). About &-10 wedge-shaped plates form the periproctal cone and possibly smaller granular plates occur ul the summit. REMARKS This species was first Ulustrated and described by Jell er al. (1985) on the basis of 2 moderately well-preserved specimens from our type locality. They recognized it as an isorophid but, because certain key characteristics could not be identified from that material, it was Icft under open nomenclature. We have since collected a further 21 specimens and, although none is particularly well preserved, there is now sufficient material to establish the basic morphology of this species. A major uncertainty. remains as to whether there are separate covering and flooring plates, since once again no clear sutures can be made out, Jt 1s probable that the flooring plates form the U-shaped groove and the cover plates are the spine-like projections that arise from this. How- ever, we have not seen any specimen in which clearly separated cover plates and flooring plates can be unambiguously distinguished, and it is therefore possible that only a single set of plates is present, acting as both flooring and cover plate, articulating perradially and carrying a long spinous projection. The similarity of the spine- like plates in C. spinesa to cover plates of other isorophids, such as Foerstediscus, ts striking. und there cun be little doubt that the plates seen in Chatsworthia are indeed homologous to those of other 1isorophids, C. ypinose is distinguished from the co-oecur- ring isorophid, H. parma, by ts. simpler and stouler peripheral rim and more robust am- bulacral cover plates that are much fewer in number. MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 23. Chatsworthia spinosa sp. nov., NMVP 107129, paratype, camera lucida drawing and plate interpreta- tion. Symbols as in Fig. 21. Order EDRIOBLASTOIDA Fay, 1962 Family ASTROCYSTITIDAE Bassler, 1938 TYPE GENUS Astrocystites Whiteaves, 1897 [Type species: Astrocystites ottawaensis Whiteaves, 1897]; Tren- tonian (Caradocian, Ordovician) of Ottawa, Canada. OTHER SPECIES Astrocystites distans Webby from the Lower Caradocian of New South Wales [possibly a subjective junior synonym]: Cambroblastus enubilatus sp. nov. from the Franconian, Upper Cambrian of Queensland. RANGE Median Upper Cambrian to mid Ordovician. DIAGNOSIS Edrioasteroids with a polyplated stalk and a bud-shaped theca with 5 basals. Ambulacra prominent with primary cover plates only. Flooring plates exposed adradially, primitively biserial but may be reduced to a single mouth angle flooring plate situated at each interradial angle. REMARKS Edrioblastoids are probably the rarest of echinoderm groups that have been given class status. Although Astrocystites ottawaensis has been known since 1897 few specimens have ever i wy he =n IG) i) fo) Fic. 24. Chatsworthia spinosa sp. nov., QMF17951, paratype. camera lucida drawing and plate interpretation. Symbols as in Fig. 21, CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA periproct 743 FiG. 25. Chatsworthia spinosa sp. nov., QMF17948, paratype, camera lucida drawing and plate interpretation. Symbols as in Fig. 21. been collected. The species was erected on the basis of 3 specimens from the Trenton Limestone (Middle Caradocian) at Kirkfield Quarry, On- tario; 2 held in the Canadian Geological Survey collections of which one is the lectotype and the other is lost, and a third held in the British Museum (Natural History). Mintz (1970) described a fourth specimen from the type locality and horizon. A second species, A. dis- tans, was described from the basal Caradocian Cliefden Caves Limestone of New South Wales (Webby, 1968) based on two tolerably complete specimens and a number of fragments. Mintz (1970) believed that A. distans was probably a subjective junior synonym of A. oftawaensis. Sprinkle (1982) described a small number of isolated deltoid plates of an edrioblastoid from the mid Ordovician Bromide Formation of U.S.A. and deltoid plates also occur in the Upper Ordovician Boda Limestone of Sweden (C.R.C. Paul, pers. comm. November, 1989). Ordovician edrioblastoids have a highly dis- tinctive morphology and have been difficult to relate to other echinoderm groups. When they were originally described they were thought to be most closely related to the diploporite cystoid Asteroblastus (Whiteaves, 1897). Bather (1900, 1914) redescribed the material then available and placed the species in its own family within the edrioasteroids. Hudson (1925, 1927), however, argued for their close association with blastoids as did Fay (1962, 1967), while both Webby (1968) and Bell (1980) have related them to edrioasteroids and Mintz (1970) to eocrinoids. Paul (1988) commented that Astrocystifes was so unusual in its morphology that it could not be placed in his phylogenetic scheme of cystoids. 26. Fic. NMVP108990, paratype, internal surface, x6. Chatsworthia spinosa nov., sp. 744 MEMOIRS OF THE QUEENSLAND MUSEUM ambulacral tunnel ——_, Fig. 27, Chalsworthia spinesa sp. noy., camera lucida drawings, A, QMF17948, peripheral rim plating in side view. B, QMF17946, view of one ambulacrum looking from the mouth cavity, C. reconstruction of ene cover plate, based on NMVP108990 and QMF 17948. C)_ external aspect. C2 internal aspeci. Scale bars =| mm. The phylogenetic placement of Astrocystites has been particularly difficult because there were no clear intermediate forms linking it to other echinoderm groups. It was therefore difficult to establish homologies. The discovery of a primi- tive form of edrioblastoid from the median Upper Cambrian of Australia bridges the gap between the mid Ordovician Astrocystites and certain edrioasteroids, making the phylogenetic place- ment of edrioblastoids much clearer. PHYLOGENETIC POSITION OF EDRIOBLASTOIDS Even prior to discovery of Camibrablasrus the link between edrioblastoids and blastoids (Hud- son, 1925,1927: Fay, 1967) seemed untenable. The lack of brachioles and a homologue of the hydrospire system and lancet plate clearly distin- guished edrioblastoids. Furthermore, the an- cestry of blastoids is now generally agreed to lie with coronates such as Srephanocrinus (Sprinkle, 1980b, Brett ef a/., 1983, Smith, 1984, Donovan & Paul, 1985, Paul, 1988) which resemble edrioblastoids even less. Two Cambrian groups seem worthy of con- sideration as possible ancestors to edrioblas- toids. namely eocrinvids and edriovasteroids. Many eocrinoids possess a polyplated stalk like that of Cambroblasius that is differentiated trom a bud- like theca. They also have the rudiments of an organised thecal plating with differentiated circlets of plates including basals, just as in Cambroblastus. The ambulacral arrangement is much different. however. Bocrinoids have short ambulacra restricted to the oral pole and these give rise to slender brachioles. Brachioles attach to the flooring plates and are believed to be homologous with the cover plate series seen in primitive echinoderms (Paul & Smith, 1984). The absence of any indication of brachioles in the Upper Cambrian edrioblastoid is strong evidence that edrioblastoids are not specialised offshoots from some eocrinoid lineage. The derivation of edrioblastoids from edrioasteraids seems much more likely, The am- bulaecral structure in Cambroblastus is very like that in edrioasteroids. with large primary cover plates roofing a broad and deep food groove that covers much of the oral surface. The reduction of flooring plates to a single mouth angle plate. as occurs in Astrocystites has its precedents amongst edrioasteroids. since this is precisely what occurs in cyathocystids and probably also in rhenopyrgids. Furthermore, both of these edrigasteroid groups have elongate thecae, with the aboral surface developed into a stalk. In the Fic, 28, Chalsworiiia spinosa sp, nov. reconstruction: interambulacral plating stippled, CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA cyathocystids the stalk isa single (?fused) calcite plate in the form of a long-cylindrical tube (Bock- elie & Paul, 1983), while m rhenopyrgids the stalk 18 composed of imbricate plates. One of us has argued previously (Smith, 1986) that the stalk in rhenopyrgids is homologous with the aborul plated surface in stromatocystitids and that the circum-oral plates at the top of the stalk are homologous with the peripheral rim plates in other edriousteroids. [f this is correct then the plated holdfast in Canthroblasius would be homologous. with the large central plates in Stromareeystites or Tatiglohus and the plated holdfast of smaller platelets homologous with ihe peripheral rim of subroarginal platelets. Cambroblasius has a rigid basal part to its theca and a more flexible upper part, The lower part consists of 3 circlets of tesselate plates and from its position, between the extremities of the am- bulacra and the flexible subambital zone of platelets, would appear to be a direct homologue of the peripheral tim plates of other edlrioasteroids. ‘The upper semi-fexible part of the theca is composed of interambulacral and ambulacral areas sitnilar in arrangement to those found in other edrioasteroids, especially in the hiserial arrangement of Mooring plates and the large simple cover plates, Over the oral area the 3 primary cover plates meet centrally, exactly as they do in eyathoeystids and rhenepyrgids, Since the plating in Cambroblastus can be directly homologized with that of edrinasteroids (Fig, 34), whereas eocrinoids differ in the fun: damental structure of their ambulacra, we believe edrioblastoids are derived forms of edrioasteroids. Three principal groups of edrioasteroids stem from a Srromatocystites-ly pe ancestor (Smith,!986); 1, Isorophida, which are disc-shaped encrusters with an unplated aboral surface; 2, Edrioasterida, which are glabose but- tom dwellers that have retained biserial flooring plates. and have lost the central plating of the aboral surface; and 3, Cyathocystida which are lurret-shaped forms which have formed a peduncle from the aboral plated surface and have reduced ambulacral plating to 5 interradially positioned flooring plates that carry multiple cover plates, Both edriousterids (Wealcattidiscus) and isorophids (Chatsworthia) had become dif- ferentiated by the median Upper Cambrian, but the earliest cyathocystids are mid Ordovician in age, Edrioblastoids have expanded their aboral surface into a stalk and in later forms reduced their ambulacral flooring plates to 5 interradial elements (the deltoids of Asfracysritey), They are 745 therefore likely 10 represent a sister group to the Cyathocystida. Cumbrahlasrus seems to be tae specialised to represent 4 common ancestor to either cyathocystids or rhenopyrgids however, although the reduction of flooring plates.to the 5 primary plates is a putative synapomorphy of Astrocystites plus cyathocystids plus rhenopyr- gids, Until intermediates linking rhenopyrgids, cyathocystids and primitive edrioblastoids are found, relationships of these 3 groups will remain problematic. TRANSITION TO ASTROCYSTITES Although Cumbrohlastus closely resembles Astrocystites in general organisation (both have apolyplated stalk, a bud-shaped theca dominated by large ambulacral plates thal extend below the ambitus and are roofed by primary cover plates} it retains a number of primitive characteristics, Several morphological changes must have taken place in the evolution of Asrrocystites from Cambroblastus, Firstly the thecal plating be- came simpler with the reduction to 5 basals and 5S radials. [t also became completely rigid, flexibiliry along the adradial margins of the ani- bulacra apparently being lost. Ambulacral plat- ing was also simplified with the loss of all but the first (interradially positoned) ambulacral floor- ing plate. This flooring plate expanded into the deltoid plate and its distal tips grew over the adoral portion of the radial plates. The simplification of thecal plating through the loss of circlets and all bur the first formed ambulacral flooring plate is most likely to have been achieved through heterochrony, with the ap- pearance of juvenile features in adult forms being the product of neotony or pacdamorphosis, Elongation of the primary eral piases into de|- toids and the multiplication of coverplates would have occurred ata later stage. A second major difference between Cambroblastus and Astrocystites is inthe system of intrathecal pores developed along the adradial margin of the ambulacral groove on delloid plates. No such system of pores is preserved in Cambroblastus which would there- fore appear to he a specialization of derived edrioblastoids. However, preservationis such that we cannot entirely rule out the possibility that minule pores were originally present but have become obliterated with silicification, The func- tion of the pores in edrioblastoids has never been properly assessed, They could be the equivalent MEMOIRS OF THE QUEENSLAND MUSEUM 746 CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA basals q47 cover plate B periproct external portion of flooring ‘ plate intrathecal ledge on flooring Cc plate Fic. 30. Cambroblastus enubilatus sp. nov., QMF17872, holotype, camera lucida drawings. A, lateral view of postertor interray. B, lateral view of ambulacral plating in ammbulacrurn IL. C, detail of periproctal region. Scale bars = 1] mm, of the umpullar sutural pores in primitive edrioasteroids but these are absent from Cambroblastus as far as can be told and would have to represent an atavistic structure, Further- more. they are not sutural pores, as in Stromatocystites or Edrioaster, but are a series of pores within a single ambulucral flooring plate. Whatever their function, they are probably unique to Asrrocystites. The only other significant difference is in the organization of stalk plating. Whereas there ap- pears to be no clear lineation to stalk plating in Cambroblastus the stalk of Astrocystites.1s made up of 5 columns of plates (although only the proximal part of the stem is known). This again iS a specialization of Aytrocystites. Cambroblastus gen. nov. ETYMOLOGY From Cambrian and the Greek h/astos, a bud, in allusion to the shape of the theca. TYPE SPECIES Cambroblastus enubilatus sp. nov. AGE AND DISTRIBUTION Median Upper Cambrian (Franconian) of Queensland. Fig. 29, Cambroblastus enubilatus sp, nov. from the Chatsworth Limestone, median Upper Cambrian of Chatsworth, A, QMF17873. paratype in side view, x8 (see also Fig. 32). B-D, QMF17872. holotype. B, lateral view of posterior interradius, x7.5 (see also Fig. 30). C, oral view. x7.5 (see also Fig. 31). D, lateral view o! ambulacra If and TV, x7.5, Silica replacements whitened with ammonium chloride sublimate. 748 oral cover plates periproct Imm Fic. 31. Cambroblastus enubilatus sp. nov., QMF17872, holotype, camera lucida drawing of the oral region. DIAGNOSIS An edrioblastoid with an irregular polyplated stalk, 3 irregular cycles of thecal plating and ambulacra composed of biserially arranged flooring plates bearing simple cover plates. No intrathecal pores to the ambulacral grooves. REMARKS This genus is sister group to the Ordovician edrioblastoid Astrocystites on account of its very similar thecal shape and ambulacral organisation and its polyplated holdfast. It differs from that genus in having an additional circlet of thecal plates between the radials and basals, in having multiple biserial flooring plates to each am- bulacrum rather than a single oral plate (deltoid), and no system of pores along the margins of the ambulacral grooves. Furthermore, the plating of the stalk is less well organised in Cambroblastus. Cambroblastus enubilatus sp. nov. (Figs 29-33) ETYMOLOGY Latin enubilatus, freed from clouds, an al- lusion to its importance in revealing the origins of edrioblastoids. MEMOIRS OF THE QUEENSLAND MUSEUM TYPES Holotype QMF17872, paratype QMF17873; no other specimens are known. AGE AND OCCURRENCE Both specimens come from locality K204 of Shergold (1982), from a grey, 4 m thick unit of the Chatsworth Limestone, 3.5 km south of Chatsworth homestead, 60 km southeast of Duchess, western Queensland. Peichiashania secunda-Prochuangia glabella Assemblage Zone, median Upper Cambrian (Franconian), basal disc Fic. 32. Cambroblastus enubilatus sp. nov., QMF17873, paratype, camera lucida drawings. A, lateral view of complete specimen. Hatched lines = missing; black = plate surface badly etched. B, ob- verse of stalk. CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 33. reconstruction. Fic. Cambroblastus enubilatus sp. nov. DIAGNOSIS As for genus, DESCRIPTION The theca is 10-11 mm long and 6-7 mm in maximum diameter with the widest part being slightly above mid-height. The stem is 5.5 mm 749 long and 1.5-2 mm wide, with a flared base (Fig. 29A). The theca tapers gradually to the stalk and has a rounded apex. In oral view the theca is rounded pentagonal in outline, with the angles formed by the prominent ambulacra (Fig. 29C). Interradial zones are concave. The mouth lies at the apex of the theca and the periproct close to the apex in the posterior interambulacrum (Fig. 30A). There are 5 large and prominent ambulacra stet that meet in a 2:1:2 pattern at the apex (Figs 29C, 31). Ambulacra are c. 2 mm wide at the apex and taper slightly to a broad rounded point situated a littke below the ambitus. The am- bulacral groove is completely roofed over by a series of simple cover plates. There are 9 or 10 cover plates in each column. These are largest at the apex and become progressively smaller dis- tally (Fig. 30A,B). Cover plates are tall and form a spacious vaulted canopy to the ambulacral groove. The cover plates fit together tightly with no sutural gaps. They interlock along a zig-zag perradial suture. Individual plates are pentagonal in outline and have a single moderately well- developed sagittal strut on their inner face. The suture between cover plates and flooring plates is nowhere clearly seen. In ambulacrum II the cover plates appear to alternate with the flooring plates (Fig. 30B). Elswhere they may sit more directly above the flooring plates, although plate boundaries are nowhere clear. No secondary cover plates are present. Over the oral area there are 5 large cover plates that meet centrally, one from each interradius (Fig. 31). The posterior interradius may, however, have 2 such plates, to judge from the plate outline. These are the oral plates and the smaller adjacent cover plates may attach directly to them. The floor to the ambulacral groove is U-shaped and smooth, with slight undulations marking the plate boundaries. Flooring plates are biserial and in places the sutures have been preferentially etched. However, no sutural pores are consistent- ly developed and in less corroded ambulacra no pores whatsoever can be seen. We therefore believe that pores were either not present in the ambulacral groove or small and obliterated by silicification. Flooring plates are exposed along the adradial margin on the outer side of the ambulacra (Fig. 30B). Beneath the exposed por- tion of the flooring plates a distinct concave facet runs the length of the ambulacrum (Fig. 30A). This is the intrathecal section of the flooring plates and is an imbrication flange on which the interambulacral plates rest. In the posterior inter- ambulacrum of the holotype, interambulacral 750 oral Rhenopyrgus MEMOIRS OF THE QUEENSLAND MUSEUM Astrocystites Cambroblastus Fic. 34. Diagram indicating the homologous regions of plating in rhenopyrgids (Rhenopyrgus), Cambroblastus, and the Ordovician edrioblastoid Astrocystites. plates are preserved in position against am- bulacrum I, covering the facet, but have slid off the flooring plates of ambulacrum V on the other side to reveal the concave facet (Figs 29B, 30A). This suggests that the ambulacral/interam- bulacral boundary was imbricate not rigid and that the ambulacral flooring plates had some limited degree of flexibility. The sutures between flooring plates are well seen in the intrathecal section. Interambulacral plating, best seen in the posterior interambulacrum (Fig. 29B, 30A), con- sists of a pavement of small polygonal plates with slightly swollen centres. These are 3-5 abreast, reducing to a single plate adapically. There may be a single elongate plate in the centre of the posterior interambulacrum. The interam- bulacral plates tend to hold together even though they are displaced from the adradial suture, sug- gesting that they were firmly bound together. The periproct opens close to the oral area in the posterior interambulacrum (Fig. 30A,C). It faces towards the apex and is set back on a ledge. It is composed of a low cone of wedge-shaped plates, relatively few in number. Above the periproct and close to ambulacrum V is a small conical structure with a central depression (Fig. 30C) interpreted as the gonopore. No trace of the hydropore is seen, probably because of the coarse preservation. The lower part of the theca is cup-like and appears to have been rigid. Plate boundaries are obscured except in the posterior part of the theca (Figs 29B, 30A). Here traces of 3 circlets of plates are visible. In the paratype there are 3 plates forming the base of the cup on one side (Figs 29A, 32) and, by inference, there must therefore be a basal circlet of 5 plates, which we homologise with the basals of Astrocystites. Above this, in the holotype are large hexagonal plates. Because only traces of sutures can be distinguished we are not certain of a regular pentameral pattern to this plating but there are 2 further circlets of polygonal plates above the basals. Some of these appear to be slightly con- cave and all are covered in a fine retiform orna- ment, At the base of the ambulacra an indistinctly defined plate may be the homologue to the radials in Astrocystites. The stalk (Fig. 29A) is cylindrical, narrowest immediately beneath the theca, and expanding slightly towards the base. The surface appears CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA FiG, 35. Incertae sedis echinoderm fragment, QMF 17859. early Middle Cambrian from QP243 near Cornford Bore, x4. A, external surface. B, internal surface. Latex casts whitened with ammonium chloride sublimate. different in texture to the theca, being much more granular and with many small pits. As a number of these pits appear to be linearly arranged we interpret them as traces of plate boundaries and the stalk to be composed of a relatively large number of small granular platelets. Towards the base these platelets become better defined and are polygonal in outline. At the base the stalk A flares outwards into a cup-shaped holdfast of large polygonal plates (Fig. 32). The stalk is broken at the bottom and the full extent and shape of this holdfast remains unknown. REMARKS A comparison of this species to the only other known edrioblastoid, Astrocystites, has been FiG. 36, Incertae sedis echinoderm fragment, QMF 17859, camera lucida drawings of internal (A) and extemal (B) surfaces. given above and its phylogenetic position has also been dealt with fully. Cambroblastus was an erect, medium-level, suspension feeder using its ambulacra to capture suspended food particles. The stalk, which is relatively well-developed for Cambrian echinoderms, would have elevated the food capturing surfaces well above the sub- stratum. Cambroblastus must have been at- tached to a substratum by its basal disc but, because only part of this is known, we are not sure as to how fixation was achieved. However, the shape of the basal disc is reminiscent of basal attachment discs of Echinosphaerites for attach- ment to hard substrata, and is unlikely to have been a rootlet structure for attachment within unconsolidated sediment. The apparently tesse- late nature of the stalk platelets suggests that the stalk was relatively stiff and inflexible, unlike the imbricate stalk of rhenopyrgid edrioasteroids. Furthermore the stalk is relatively robust and has held together well, not having collapsed during fossilization. Both observations indicate that the stalk of Cambroblastus was more or less rigid and not extensible. The broad imbricate facet between the floor- ing plates and the interambulacral plates implies that this boundary was flexible and that a large degree of movement was permitted between the two plate systems. Interambulacral areas are often separated from the adradial suture and are also variable in width (Fig. 29B,C,D). Further- more, there is a distinct change between the lower cup- like portion of the theca, where plates are rigidly sutured, and the rather irregularly concave surfaces of the interambulacra between the arms. The flexibility between ambulacral flooring plates and interambulacral regions sug- gests to us that the ambulacra retained a certain amount of movement and that the food groove could be widened or closed by lowering or rais- ing the flooring plates. The absence of pores in the floor of the food groove requires that the water vascular system, presumably lying in the floor of the groove, lacked internal ampullae. Incertae sedis (Figs 35, 36) MATERIAL A single specimen, part and counterpart preserved as a mould in silica, QMF 17859a/b. MEMOIRS OF THE QUEENSLAND MUSEUM AGE AND OCCURRENCE From chert beds of the Yelvertoft Member, Beetle Creek Formation, early Middle Cambrian, exposed on the northern flank of a low rise some 500 m south of the bore at Yaringa Creek, 50 km west-southwest of Mount Isa, western Queensland (GR 980965). DESCRIPTION The specimen is 10 mm x 12 mm, although the posterior(?) edge is fractured and missing. It has a narrow anterior border and 2 long straight sides that diverge at c. 80°. The external surface (Figs 35A, 36B) shows a flat, unornamented platform surrounded by a | mm wide ledge. This ledge is concave, dipping steeply from the platform before levelling out. It is crossed by vertical sutures approximately 0.8-1.0 mm apart. There is a central plate on the anterior border and adjacent plates occupy each of the angles. No sutures are seen on the platform. The interior of the specimen shows a steeply dipping marginal rim surrounding a crescentic region of labyrinthine struts (Figs 35B, 36A). There is a central platform towards the posterior of the specimen. The rim is crossed by vertical sutures, at least anteriorly. Struts from the labyrinthine region run to the centre of each marginal plate. The anterior struts are rather short and dip into the labyrinthine region but there are 2 long struts that run from the side walls right to the posterior platform without branching. The floor of the labyrinthine region is pitted and the specimen must have been exceedingly thin at these points. The posterior platform is flat and smooth. Towards the posterior there are 6 small (0.2 mm diameter) pores. It is not known whether these perforate the plate as the corresponding region is not seen in the counterpart. No sutures are seen in either the labyrinthine region or the posterior platform. REMARKS This fragment is an enigma. We feel confident that it is echinoderm, even though calcite plating is not preserved. This is because of the presence of plate boundaries along the peripheral ledge. The absence of sutures in the main body of the plate suggests to us that the fragment is a com- posite element, constructed of several plates fused together. The peripheral flange and shape of this element are reminiscent of large inter- radial first ambulacral plates; the flange being an adradial ledge either formed of fused ambulacral flooring plates or directly flooring the am- CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA bulacra. Large deltoid plates. situated inter- radially and bearing ambulacral structures, are known in edrioasteroids, blastoids, parablastoids and edrioblastoids, The interradial plates in edrioasteroids form the oral frame and are suspected of being formed from fused am- bulacral flooring plates (Bell, 1976, Smith, 1986). However, they are much smaller and none 1s known to have such a distinctive labyrinthine interior. In edrioblastoids the interradial deltoids are also supposedly composed of fused am- bulacral flooring plates (Mintz, 1970, Bell. L980) and here the plate is large with an adradial ledge. However, such deltoid plates are known only in the Middle Ordevician Astracystires and were apparently absent in the Late Cambrian Cambroblastus, The deltoids in blastoids carry part of the hydrospire system and are not closely comparable. The deltoid in parablastoids is a large plate perforated by a system of canals termed cataspires (Sprinkle, 1973. Paul & Cope, 1982). It does not underlie the ambulacra but abuts the flooring plate system and no species is known to have a labyrinthine internal plate or- namentation. Because it may represent an inter- radial plate of fused ambulacra we suspect it night be some form of edrivasteroid derivative, but it represents a major departure in torm from any reported echinoderm group. The crescentic zone of labyrinthine struts creates an extremely thin-walled surface and we believe that this may be an adaptation for gaseous exchange, The struts provide strengthening sup- port to the theca, bracing the ambulacral zones, while the pits are sites of gaseous exchange. Extreme thinning of calcitic plates associated with gaseous exchange is a common feature in many echinoderm groups. Amongst cystaids. for example. thin-walled regions of the plate ure found in paracrinoids (deep pits), diploporites (the exterior of haplopores is a thin membrane of calcite) and dichopoarites. (where tube-like pas- sageways from the interior run immediately beneath ihe surface of the plate). The arrange- ment of pits and struts in such an irregular man- ner rules out the presence of any sophisucated circulatory system as in most cystoids, Gaseous exchange must have taken place through simple diffusion across the thin plate. “Subclass SOMASTEROIDEA’ Family ARCBEG ORL TERIDAG Spencer, 95) Archegonaster Jaekel, 1923 TYPE SPECIES Archegonuster pentagonus Spencer, 195], Archegonaster pentagonus Spencer, 1951 (Figs 37-51) Archegonasrer pentagons Spencer, 1/981, p. 107, fies Y, 70, 12-15, 34, 37-40, 54, 55. Archegonaster penagenus Spencer; Spencer & Wright, 1966, p. 041. fig. 39C. Archegonaster pentagonuys Spencer; Smith, |YR8b, p. 89, pl. 7.1, fig. 4, pl.7.2, fig, 6. MATERIAL The Narodni Museum, Prague houses 65 specimens, including the types. AGE AND OCCURRENCE All material comes from the Llanyirn Sarks Formation of the Osek district. Prague, Czechas> lovakia. DEseriPTlon Archegonaster is.a pentastellate echinoderm with a stout marginal frame of ossicles and nar- row projecting arms (Fig. 37). There is no separa- tion between disc and arms, the two merging imperceptibly into one another. Its size is dif- ficult to estimate since so many specimens are meomplete or distorted. The most complete specimen (Fig. 37) has a radial length of 35 mm and an interradial/radial diameter of 60 mm, but the tips of its arms are missing. Specimens cover a large size range, from about 15 mm radial length (Fig. 49B) to around 50 mm, The body ts flattened. Ambulacra. These are composed of biserially arranged ambulacral ossicles (=flooring plates) arranged alternately and forming an open L- shaped channel (Figs 38-40), In oral view they appear as T-shaped ossicles. measuring about 3 mm perradially and with a 2 mm transverse bar in larger specimens (Figs 40A, 41B-E). The per- radial suture 1s virtually straight and successive ambulacral plates overlap one another, the distal! part of one plate passing beneath the proximal part of the next distal plate. About one third of the plate imbricates beneath ihe adjacent am- bulacral plate. The perradial face, forming the ambulacral groove, is relatively smooth and slightly depressed centrally. The distal portion of the plate is more tapered and composed of denser stereom: this is the portion that passes under- neath the next distal plate. There is a well marked rim to the top of the ambulacral groove. This runs i fram the proximal end of the plate to the (ransverse bar, where it ends abruptly ata notch (Fig. 40A). Presumably, this is the notch for the lateral branch of the radial water vessel leading toa tube foot. In Jurger specimens the floor to this notch has asmull pit. When ambulacral plates are in life position (Figs 38E, 41B-E) the distal wall of the notch for the radial water vessel 1s formed by the proximal edge of the succeeding am- bulacral plate. The transverse bar is tall and narrow, wilh steeply sloping sides. The ridge to the ambulacral groove continues along the top of this bar, At its abradial end the transverse bar is tripartite, with a large flatuish central face that slopes outwards (the articulation face for the adymbulacral ossicle) and two narrow rims marking the edges of proximal and distal podial basins (Figs 39G, 40C). The articulation face expands aborally and in some may be slightly convex, though itis usually flat, The flanges on either side are steeply dipping and flare out towards their base, The Mange on the distal face is considerably better developed than that on the proximal face. which hardly flares out at all (Pig. 40A). Thus only the proximal part of each podial basin is floored and the narrow rims presumably mark the site of tube-foot attachment. In aboral aspect the ambulacral plates are also T-shaped, with the transverse bar generally ap- pearing much more clearly tripartite at its up (Figs 40B, 41A), The perradial zone is rather flat and smooth, there being no evidence for dorsal inter-ambulacral muscle attachment sites. The proximal end of each ossicle is more pointed than the distal end und is marked off from the rest of whe plate by a small rim. This rim becomes progressively more pronounced in plates towards the mouth and marks the limit of plate imbrication, The imbrication surface is inclined away trom the perradius. The outer part of the plate also slopes downwards away from the per- radius, The three regions at the tip of the transverse bar correspond to the central articula- tion face for the lateral arm plate and the two lateral Manges flooring the podial basins. As seen orally. the distal flange is much better developed than the proximal flange. The two flanges form a rim to the transverse bar. The way in which opposite plates articulate perrudially is not clear. There is no evidence for one series having over ridden the other or for overlapping of alternate plates. "The perradial MEMOIRS OF THE QUEENSLAND MUSEUM edge of ambulacral plates is thin and no tongue and groove structure could be identified. As the perradial line is virtually straight the ambulacral plates may simply have abutted along a narrow face. In juvenile specimens, or towards the arm tips, the ambulacral plates become less strongly im- bricale and more squarish in oujline with a less pronounced transverse bar (Figs 40D,49 C), Adambulacrals, Ossicles that articulate onto the ambulacral (flooring) plates are called adam- bulacrals in asteroids and Jateral arm plates in ophiurotds. As both structures are homologous we choose to refer to such plates as adam- bulacrals, In Archegonaster the adambulacrals have a complex torm, best understood by ex- amining the camera lucida drawings. that show the plate in different orientations (Fig. 42). The ossicle is L-shaped in side view and more ir- regular in adradial (j,e, the fice that articulates with the ambulacral plate) and abradial (ic. dis- tal to the articulation facet) aspects. The ossicle hus a laterally flattened shatl, articulating onto the ambulacral plate at its base, a swollen head and a laterally Matlened abradial projection which articulates with the first virgal distally (the virgal projection). Short (1 mm) spines attach to the head (Fig, 414). The proximal fave of the shafl has a median ridge running up the length of the shaft (Fig. 42A), There is a clearly defined pit on the adradial side of this ridge towards the head of the shaft. This is presumably a muscle pit. The abradial edge of the shaft may have a small projection or flange beneath the main abradial projection, but only in larger ossicles close to the mouth, It is laterally flattened but never achieves the size of the virgal projection, The virgal projection is square-ended and com- pressed in the proximal/distal plane. It has a smoothly rounded distal face but is flatter or slightly concave on its proximal face and hears shallow grooves. The upper edge of the virgal projection is a well-defined rim in Jarger ossicles and this is separated from the median ridge of the shaft by a clear notch, Theecistal face has a prominent flattish platform forming the head. The upper (adoral) face of this platform carries 6 or 7 spines. The spines are largest on the proximal end and decrease in size significantly towards the distal end (ie. towards the virgal projection). In oral view the spines sit in slight FIG. 37. Archegonaster pentagons Spencer, Llanvien of Osek, Crechoslovakia, AJB, NM EH30. A, oral. B, Aboral. Latex casts whitened with ammonium chloride sublimate, 2, CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA EMOIRS OF THE QUEENSLAND MUSEUM M 56 b = Pager nS ‘ed og. a at, . CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA depressions, the largest being proximal at the heel of the L-shaped plate. Towards the mouth, the adambulacral ossicles become more slender and elongate along both the shaft and the virgal projection, while towards the arm tips the adambulacrals are much more block-like, with hardly any shaft or virgal projec- tion. In the outer part of the ambulacra these adambulacrals attach directly to the marginal ossicles. Interradial plating. The interradial zone, lying between the adambulacrals of the ambulacra and the marginal ring, is a complex, plated surface and commonly disrupted by post-mortem decay. Towards the tips of the ambulacra interradial plating is absent and the adambulacrals attach directly to the marginal ossicles (Figs 43C, 49C). At the interradius plating is also absent, leaving wedge-shaped uncalcified zones (Figs 37, 38A- D,43A,B). In between these two regions the adambulacrals are connected to the marginal os- sicles via one or more slender rod-shaped plates termed virgals (Fig. 43). The number of virgals in a row increases towards the centre as the interradial space becomes broader. The first vir- gal attaches to the abradial projection on the lateral arm plate (Figs 43C, 45, 46) and is verti- cally orientated with slightly expanded ends, The distal face is smoothly rounded, with a central ridge that is continuous with a corresponding ridge on the lateral arm plate. The proximal face is flat or slightly concave and has spoon-shaped depressions at each end, possibly the site of muscle attachment (Fig. 44). The ends of the plate are flat suture faces. Up to 5 other virgals are present in the row (Fig. 46). These are generally similar in shape but some may fork abradially and the lower edge is often stepped (Fig. 44B). Those plates that fork show a double suture at one end and the ridge on the distal face also branches. Some plates have their plane of flattening rotated between adradial and abradial ends, The rows of vertically elongate virgals are separated by rows of intervirgal plates (Figs 38E, 39F, 41C,D, 43C, 45, 46). These are more ir- regular in shape, some cross-shaped with four sutural faces. Where virgals are well preserved there is always an aboral set of plates that attach to the inner edge of the virgal row (Fig. 45). The 757 first of these also attaches to the abradial edge of the lateral arm plate (Figs 41C, 45A). In the best preserved specimens (Fig. 39E,F) these plates form an almost complete covering to the inter- radial regions. There is a suggestion that the virgals form vertical ribs with the intervirgals lying flatter in the spaces in between. However, it is possible that the intervirgals are only con- nected to one side of the virgals and do not link adjacent virgal rows. Mouth frame plates. The 3 most adoral am- bulacral plates in each column are modified in Archegonaster into a jaw. Although a few specimens show large gaps down the perradial line of ambulacra (buccal slits of Spencer, 1951), this we interpret as post-mortem artefact. In well- preserved individuals only the first 2 ambulacral plates are completely separated from one another at the perradius (Figs 37, 38A). The first am- bulacral plate lies interradially and is elongate radially (Fig. 47D). It is always paired with the corresponding plate from the adjacent am- bulacrum and is the most highly modified of the ambulacral plates. The second plate is stouter, more cylindrical and is set obliquely to both the first and the third and subsequent ambulacral plates. The third ambulacral plate is only slightly modified from the remaining ambulacral plates in that it has a stout proximal end. Opposing third ambulacrals meet along the perradius in life. The first ambulacrals, or mouth angle plates, are flattish plates that abut interradially along their aboral edge (Fig. 47A). The aboral face is crossed by 2 grooves. The more proximal groove is clearly marked and Vs towards the interradius. This, by comparison with the jaws of Recent ophiuroids, is the site of the circum-oral nerve ring. There is a strong projection immediately distal to the nerve ring groove and a shallower groove distal to this projection, which marks the site of the circum-oesophageal water vessel ring. The distal portion of the aboral face is a ledge and the groove for the water vascular ring runs obliquely away from the interradius along the edge of this. In lateral view the interradial face (Fig. 47A2) has a broad, relatively flat platform distally and a zone of pits and transverse bars proximally. This pitted zone is presumably the site of inser- tion of muscle or ligament that binds the pairs of Fic. 38. Archegonaster pentagonus Spencer, Llanvirn of Osek, Czechoslovakia. A,B, NM 41/83, x2. A, oral. B, aboral. C.D, NM 42/83, x2. C, oral. D, aboral. E, NM L10446, one ambulacrum showing ambulacrals, adambulacrals, virgals, intervirgals and marginals, x7. Latex casts whitened with ammonium chloride sub- limate. MEMOIRS OF THE QUEENSLAND MUSEUM 758 CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 759 plates together interradially. The platform is bar is smooth, uniformly convex and set oblique smooth and ts an abutment surface. The proximal — to the aboral face. The perradial face in lateral A B ambulacral imbrication Surface : adambulacral podial ‘4 articulation basin : Bae Z. : face notch for water vascular branch adambulacral articulation face Fic. 40. Archegonaster pentagonus Spencer, Llanvirn of Osek, Czechoslovakia, Camera lucida drawings of ambulacral plates. A, oral face of three plates, arm lip towards bottom (slightly restored from NM L10149), B, aboral face of 3 plates, arm tip towards bottom (slightly restored from NM L10438), C. abradial face of one plate arm tip towards right (from NM L10464). D, aboral face of 5 plates towards the arm tip, arm tip towards the left (based on NM 1.35050). Scale bar = 1 mm. Fic. 39. Archegonaster pentagonus Spencer, Llanyvirn of Osek, Czechoslovakia. A-C, NM L10434. A, detail of ambulacral and adambulacral plates in oral view, 43.5. B, general View, x2, C. madreporite plate, x5. D, NM L10435, ambulacra with adambulacrals, x4. E.F, NM L10143, x2. E, oral. F, aboral (note interradial plating). G.NML25454, lateral view of ambulacrals, aboral surface uppermost, with adambulacra and interradial plating, x3. Latex casts whitened with ammonium chloride sublimate. MEMOIRS OF THE QUEENSLAND MUSEUM 760 ' ’ - =) he oe ‘ M os a wae? a - _ — ‘ 2. y - - A * _ Se : ’ . miya oF CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 761 view (Fig. 47A)) has part of a podial basin dis- to this proximal edge, but the plate could only be tally and a thickened semi-cylindrical bar studied where the proximal plates had been dis- proximally, This proximal bar is smooth and — rupted and presumably mouth spines may easily uniformly convex. Nospines wereseenattaching have been dislodged or lost. spines virgal projection pit ledge for spines primary spine pit ambulacral articulation face Fic. 42. Archegonasier penragonus Spencer, Llunvirn of Osek, Czechoslovakia, camera lucida drawings ol adambulacra, A, lateral view of proximal face (NM L10438), B. kiateral view of distal face (NM L10434) C outer (oral) face (NM 110435). D. adradial face (NM L10495). E. inner (tboral) face (NM L10150). Scale bar = ! mm and all to same scale. Fic. 41, Archegonasier peniagonus Spencer, Llanyirn of Osek, Czechoslovakia, A, NM L10438, aboral view of an ambulacrum with second ambulacral at the top. Note the adambulaerals seen i lateral view, x6. B.D, EF. NM 110537. B, oral face of one arm, 43. D. same enlarged to show virgals and intervirgals, x3. B. adoral portions of two ambulacra in oral view, x3. C, NM 43/83, interradial plating in one arm, x3- Latex casts whitened with ammonium chlorice sublimate. 762 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 43. Archegonaster pentagonus Spencer, Llanvirn of Osek, Czechoslovakia. A,B, NM 39/83, fully articulated specimen showing the interradial gap in plating, x2. A, oral surface. B, aboral surface. C, NM L10069, one arm showing ambulacrals, adjacent adambulacrals and complex interradial plating, x5.5. Latex casts whitened with ammonium chloride sublimate. The second ambulacral plate is slightly pinched about one third of the distance from the proximal end (Fig. 47B, C). It carries two half podial basins on its perradial side separated by an oblique ridge. The smaller proximal basin faces laterally while the larger distal basin faces more adorally. The proximal half podial basin fits against the half podial basin on the distal end of the first ambulacral to create a completely enclosed basin. There is a broad distal tongue that imbricates with the third ambulacral plate. The interradial face is smooth and cylindrical. There is a small lateral arm plate that attaches to the transverse bar. The third ambulacral plate resembles more distal ambulacral plates except that it has a much larger proximal end with a prominent raised rim abradially marking the limit of overlap of the second ambulacral plate (Fig. 47C, D). Madreporite and stone canal. The madreporite is seen in anumber of specimens (Figs 37A, 38B, 39B,C, 41D), It is a large button-shaped ossicle, circular in outline, situated in one of the interrays close to the mouth. Because the madreporite is not part of a tesselate surface and has often collapsed after death, Spencer (1951) was not sure whether it was aboral or oral in position. However, 11 specimens in which the madreporite can be seen show the central part of the disc with ambulacra undistorted and all five ambulacra facing the same direction. In all 11 specimens the madreporite lies with its outer surface facing aborally and so we conclude that the madreporite lay on the aboral surface. The outer face of the madreporite is ornamented with a series of branching grooves radiating from the centre (Fig. 48A). The inner face is more domed and is smooth (Fig. 48B). Towards the centre there is a scroll- shaped perforation which marks the site of attachment for the stone canal. The stone canal is best seen in NM L10150 and L10438, but is present in other specimens also. It is a tube built up of a single series of stacked cylindrical ossicles. These ossicles are oval in outline and show a U-shaped outer calcitic sheath surrounding an anchor-shaped core (Fig. CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 763 Fic. 44. Archegonaster pentagonus Spencer. Llanvirn of Osek, Czechoslovakia, camera lucida drawings of interradial elements; A, NM L10438, two first virgals. B, NM L10457, two first virgals (v) and an intervirgal (iV) in part and counterpart view. Scalebar = | minzall to same scale. Fig. 45. Archegonaster pentagonus Spencer, Llanvirn ot Osek, Czechoslovakia, camera lucida drawings of interradial plating. A, NM 43/83. B, NM L10446, C, NM L43183. ad = adambulacral plate. vj, v2, v3, = first second and third virgal plates. iv = intervirgal plate. m= marginal. Scale bar = | mm. 764 Fic. 46. Archegonaster pentagonus Spencer, NM L10450, Llanvirn of Osek, Czechoslovakia, camera lucida drawing of one half of an arm from the second ambulacral to the ninth showing relatively undisrupted interradial plate arrangement. am2-8 = ambulacrals 2 to 8. ad2-9 = adambulacrals 2 to 9. V2-9 = virgals forming row from adambulacrals 2 to 9 (successive virgals are labelled as i, ii, iii, iv, v). [V = intervirgal plates. MEMOIRS OF THE QUEENSLAND MUSEUM 48C). A scroll-shaped cavity runs through each ossicle. Marginals. The complete rim of stout mar- ginal ossicles (Figs 37, 38) is formed by cylindri- cal ossicles that abut along smooth, slightly concave lateral faces (Fig. 50 ). Three specimens of disarticulated Archegonaster plates from a slightly higher horizon (D y2 of Zbytky hvesdic, Sarka) have a crenulate rim to the lateral faces of marginal ossicles and may represent a separate species. The marginals are largest interradially and decrease in size towards the arm tips (Fig. 49A,D). They are slightly triangular in cross-sec- tion, with the two longer and slightly flatter faces being the external aboral and oral faces. These faces, in large ossicles, have a narrow smooth border and bear a distinct, but fine, pitted orna- ment (Figs 49A, 50). The inner face is more rounded and lacks the pitted ornamentation. There is a distinct ledge on the aboral face mark- ing the boundary between inner and outer faces. Some ossicles are distinctly saddle-shaped. The inner face usually has small projecting facets that are rounded or vertically elongate (Figs 38E, 43C, 50). There is usually a single facet to each marginal ossicle, but occasional ossicles may bear two. The most interradial 5 or so ossicles lack facets (Fig. 49A). These facets are the at- tachment points for the interambulacral skeleton. The 17-27 marginal ossicles from the inter- radius to the arm tip, indicate that there were some 170-270 marginal ossicles in the complete ring. The marginals diminish in size markedly towards the tips of the arms, where they become almost U-shaped in cross- section with a slightly concave inner face. In the outer part of the arm the marginals sheath the ambulacral groove (Figs 49B,C, SOD). Aboral surface. Aboral plating is composed of small (c. 0.05 mm diameter) granular plates, closely packed together. In one specimen this tegmen coats the entire surface and is wrinkled, showing that it was flexible (Fig. 49C). It ends at the small ledge on the aboral face of marginal Fic, 47. Archegonaster pentagonus Spencer, Llanvirn of Osek, Czechoslovakia, camera lucida drawings of proximal ambulacral plating. A, first ambulacral (= mouth angle plate). Aj, perradial face. Ag, interradial face. A3, aboral face (two adjacent plates abutting. slightly restored) drawn from NM L10149 and L10150). B, second ambulacral plate in perradial view (NM L10476). C, second and third ambulacrals in interradial view (NM 41/83). D, proximal am- bulacral plates in NM EH30, plating slightly dis- rupted. Scale bars = 1 mm. CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 765 notch for water A; vascular ring Ao ligament interradial pits muscle pad 3 bloat) distal it podial basin 766 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 48, Archegonaster pentagonus Spencer, Llanvirn of Osek, Czechoslovakia, camera lucida drawings. A, exterior of madreporite plate (NM L10434). B, interior of madreporite plate: note the scroll-shaped perforation (NM L10444). C, end view of a single barrel-shaped ossicle of the stone canal (NM L10438). Scale bar = | mm, ossicles marking the boundary between the pitted ornament of the external surface and the smooth internal surface. REMARKS Archegonaster was described by Spencer (1951) but, because all the material was preserved as moulds and high quality latex casts were unavailable to Spencer, the description is inadequate in many respects. Spencer, for ex- ample, believed that the virgals formed a single series and that they were developed only towards the arm tips. Arche gonaster is considerably derived in com- parison to any known Cambrian echinoderm and shows several characteristics that link it to ophiuroids and asteroids in particular. There is a madreporite and stone canal. The aboral position of this madreporite suggests that Archegonaster lived with its mouth facing the substratum, since the madreporite would need to be clear of sedi- ment to function efficiently. It also has adam- bulacrals that articulate onto transverse bars from the ambulacrals and which carry spines. The most proximal ambulacral plates are modified into a jaw and the ambulacral tips ex- tend outwards as arms, though still bound within the marginal frame. However, it remains relatively primitive in comparison to ophiuroids. The radial water ves- sel is not enclosed and the ambulacral plates, at least proximally, are strongly imbricate. The adambulacrals attach directly to the marginals or via a series of interradial plates and the oral surface is bounded by a well-developed marginal frame. Spencer (1951) believed that Ar- chegonaster was the most derived of his somasteroids (largely because it was slightly younger than Villebrunaster and Chinianaster), but both Villebrunaster and Chinianaster have enclosed radial water vessels and reduced mar- ginal ossicles and are thus morphologically closer to ophiuroids than Archegonaster. The plate arrangement towards the tips of the arms of Archegonaster resembles that of asteroids, with the adambulacrals attaching directly on to the marginals, both marginals and adambulacrals being stout, block-like ossicles. The imbricate ambulacrals are peculiar; no Cambrian edrioasteroid has such strongly imbri- cate plating, although limited imbrication is present in all. Nor do any of the Ordovician asteroids or ophiuroids have a comparable arran- gement. However, the ambulacral plates towards the arm tip in Archegonaster strongly resemble ambulacral plates in Cambrian edrioasteroids such as Cambraster walcotti (Smith, 1986, text- fig.7). Imbrication is best developed towards the mouth and is least developed towards the arm tips, thus allowing greatest flexibility of shape towards the centre of the disc. This may be associated with the adoption of a macrophagous diet. Mouth frame plating. The structure of the am- bulacra closest to the mouth is particularly revealing when considering the origins and homologies of the ophiuroid jaw. However, con- flicting nomenclature and concepts of homology are to be found in the literature and it seems CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 767 Fic. 49, Archegonaster pentagonus Spencer, Llanvirn of Osek, Czechoslovakia. A, NM L25172, one arm in aboral aspect. x3. B, NM L10462, two arms towards their tips in oral aspect, x8. C, NM L35050, one arm in aboral aspect, x5: note the aboral integument of granules which is wrinkled towards the base of the arm. D. unnumbered specimen in the Narodni Museum showing the tapering arms; note that the disc is folded over upon itself, x2.5. 768 MEMOIRS OF THE QUEENSLAND MUSEUM virgal facet virgal facet intervirgal Fic. 50. Archegonaster pentagonus Spencer, Llanvirn of Osek, Czechoslovakia, camera lucida drawings of marginal ossicles. Aj-4, Aboral, oral, internal and lateral aspects of a large marginal of the central part of the disc (drawn from NM L10150, L.13228 and L10438). B1-3, internal, external and lateral aspects of a marginal towards the arm tips (drawn from NM L10462). C, marginal with virgal attached (NM L13228). D, distal tip of an arm in oral aspect showing how the marginals protect the ambulacral groove (NM L10462). Scale bar = | mm. advisable to correct this in the interests of unifor- mity for future descriptions. Spencer (1914, p. 29; 1925, p. 263) believed that the most proximal set of 10 plates in the ophiuroid mouth frame were interradial in origin and therefore modified interambulacral plates rather than ambulacral plates. These most proximal elements he named ‘mouth angle plates’. In his scheme the second plates of the ophiuroid jaw, distal to each mouth angle plate were the first of the ambulacral plates and, ac- cordingly, he termed these plates ‘ambulacral 1’. This nomenclature has been followed by many subsequent authors and was reinforced in the Treatise by Spencer & Wright (1966), despite their acknowledgement (Spencer & Wright 1966, pp. 13,14,29) that the mouth angle plate is part of the ambulacral series. Fell (1963) af- firmed the ambulacral origin of the mouth angle plates and Bjork et al. (1968) went so far as to denote the plate behind the mouth angle plate (which Spencer (1925) and Spencer & Wright (1966) referred to as ambulacral 1) as ambulacral 2. However, Kesling & Vasseur (1971, fig. 8) reverted to Spencer’s terminology, and the Treatise terminology (Spencer & Wright, 1966) has become standard over the last 20 years. Such terminology contradicts the origins and homology of the mouth plates and ought to be abandoned. We therefore advocate, and hence- forth use, the following set of terms: the most proximal plate of each ambulacral column, situated interradially will be the ‘first ambulacral’ or ’mouth angle plate’; the succeed- ing plate is the “second ambulacral’ and sub- sequent ambulacral plates are denoted accordingly as ambulacrals 3, 4, etc. CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA The first ambulacral bears the anterior half of a podial basin in Archegonaster (Fig. 47) and a slight depression from the aboral water vascular ring canal marks the position of the lateral branch leading to this podial basin. We interpret this groove as the site of a branch of the water vas- cular system leading to the first tube foot in that podial basin. The position of this branch leaves it vulnerable to damage from larger particles that were ingested. Evolution led to enclosure of that branch of the water vascular system, giving greater protection, and this condition is found in most subsequent ophiuroids, Similar enclosure of a canal in the second ambulacral would lead to the arrangement that is found in Lapworthura miltoni (Salter), for example (Spencer, 1925, fig. 191), Likewise in each arm segment the passage connecting each podial basin with the radial water vessel, which in Archegonaster appears as a notch, is ultimately enclosed within the am- bulacral plate. In Lapworthura the opening of the canal leading from the water vascular canal to the first podial basin has shifted onto the second ambulacral and thus pierces through two plates. As the first and second ambulacrals are closely fixed together, passage of this canal from one plate into the other would pose little problem. The situation in post-Palaeozoic ophiuroids and in Palaeozoic euryalinids is for there to be a single aboral pore in ambulacral two which divides internally to feed both the first and second podial basins. This is a derived condition and cannot be used to argue that the second ambulacral plate is a compound plate, as has been done by Bjork eral. (1968). The general one to one correspondence of podial basins to am- bulacrals and adambulacrals to ambulacrals in early forms such as Archegonaster, Eophiura (pers, obs.; Spencer & Wright, 1966, fig. 70-1), and Praedesura (pers. obs.; Spencer & Wright, 1966, fig. 23-3) among others, gives us con- fidence that no ambulacrals are compound plates. Only one element has not been identified in these early forms and that is the adambulacral element of the first ambulacral. We believe that the torus that abuts the proximal end of am- bulacral 1 represents the modified adambulacral 1. On this interpretation the spines on the torus would be the modified lateral spines of the adam- bulacral. It is not difficult to envisage the most proximal adambulacrals in primitive members becoming modified for food manipulation in the same way that the proximal tube feet became modified into buccal tentacles. Although most ophiuroids have 5 torus plates, one for each pair 769 of interradially joined first ambulacrals, several do have 10 plates, one for each first ambulacral (e.g. Spencer, 1925, fig. 188C). Although the situation in Archegonaster remains unclear, we believe that the primitive condition is for there to be ten torus plates bearing mouth spines. Structural and functional specialization of the more proximal elements around the mouth were undoubtedly associated with the change from suspension to deposit feeding. In suspension feeding a fixed mouth frame gives strength and rigidity, but for deposit feeding flexibility of the mouth is essential. The proximal ambulacrals thus became very different in appearance to more distal arm plates. Significantly the homologies that we propose allow for the development of the ophiuroid jaw from the simple ambulacral and adambulacral series through only a few steps, each relating to specific functional needs. We do not need to invoke the loss or gain of any plate, nor any compounding of plates. Furthermore it gains support from the most recent embryologi- cal work of Hendler (1978) who has also argued for the ambulacral origin of the mouth angle plates in living ophiuroids. The most proximal elements (ambulacra 1) are highly modified in shape and carry a groove for the circum-oesophageal ring nerve and water vascular ring. Their general shape and soft tissue associations leave no doubt that they are homologous to the mouth angle plates of ophiuroids, In Archegonaster these plates are in direct continuity with other ambulacrals and carry a partial podial basin towards the posterior which is shared with the next ambulacral plate. Spencer (1925) thought that the first plates in the ophiuroid jaw were modified adambulacrals, since the first podial basin is found on the second of the jaw plates and the adambulacrals are in direct alignment with the first jaw plates (mouth angle plates). That the mouth angle plates cannot be modified adambulacrals of the so-called ‘first ambulacrals’ is further reinforced by the fact that these plates bear a small adambulacral in Ar- chegonaster discrete from the mouth angle plates. Mouth angle plates of Archegonaster are similar to those of primitive ophiuroids such as Eophiura in lying in the same plane as the rest of the ambulacrals. In later ophiuroids the second ambulacral plates are highly modified and project aborally over the third ambulacrals. However, the basic pattern of having the first ambulacral plates interradial in position and the second ambulacral plates set oblique to both the vH) proximal elements and the rest of the ambulacta, is common to both Archegonasrer and all ophiuroids. The connection between Archegonaster anu asteroids is less clear-cut. Unquestionable Or- dovician asteroids (viz. Platanaster) differ from Archegonaster in having block-like ambulacrals that firmly abut, but have a comparable arrange- ment of plates around the mouth, As noted above,, the ossicles become much more block-like dis- tally along the arms in Archeganuster and the radial water vessel is not enclosed, as in all asteroids. The marginal ossicles are probably directly homologous with the marginals or sub- marginals of Ordovician asteroids, and forms such as Petraster have prominent block-like marginals framing the oval face. Antecedents ol Archegonaster are still un- clear, Although Archegonaster is morphologi- cally primitive in comparison to asteroids and ophiuroids considerable differences remain be- tween itund any known edrioasteroid. However, despite the limited evidence with which to trace the Cambrian antecedents of Archegonaster, wetake the pragmatic view that, amongst known forms, every laxog has another thal is its closest known relative. The stout marginal ring fram- ing the body and the projection of ambulacral lips onto the marginal ring of Archegonasrer are suggestive of cither Cambrasrer or Edriodiscus, However, both of these genera have a peripheral skirt of platelets and an oral periproct, neither of which is found in Archegonaster, The mouth plating in Archevonaster was flexible and presumably under muscular control, whereas Canibraster has a fixed mouth frame of 5 large interradial elements that represent fused first am- bulacral plates from adjacent ambulacra. The mouth frame of Edriodiscus is unknown, but externally resembles thal of Stromatocystites which consists of the LO most proximal ame bulacral plates (Smith, 1986), Since we do not beheve that the flexible mouth plating of Ar- chegonaster, comprising the most proximal 10 ambulacral ossicles, 18 likely to be derived from a form in which proximal ambulacrals have become fused and modified into 5 large interradial elements, a Cambrian edrioasteroi With unfused proximal ambulacrals seems the most plausible antecedent to Archegonaster and its descendents. Thus current evidence points to Adriadiscus as the closest known rela- live of Archegonaster amongst Cambrian echinoderms. We predict that further inter- mediates berween edrinasteroids and = Ar- MEMOIRS OF THE QUEENSLAND MUSEUM Chegenaster Will be found im the Middle and Late Cambrian. GENERAL DISCUSSION PHYLOGENY There are five principal echinoderm groups in the Ordovician Whose ancestry lies in the Cambrian edrioasteroids, namely edrioasterids.. isorophids, rhenopyrgids plus cyathocystids, edrioblastoids and Arche eonaster and its deriva- tives (including both asteroids and ophiuroids), Smith (1986) presented a cladistic analysis of a number of these groups but specifically excluded Archegonaster und its relatives, which had not at that point been studied. In this paper we have accumulated more evidence about the morphol- ogy of primitive edrioasteroids and added a num- ber of new taxa. as well as documenting the morphology of Archegenaster. It is. therefore appropriate to reanalyse edrioasteroid relation- ships ithe light of this new information. A data matrix was assembled (Fig. 51; Table |) based on as much morphological information ass Currently available. This was then subjected to 4 computer parsimony analysis using Hen- nigS6 (Farris, L988). All characters were treated as unordered and Camprostrama was used to root the tree, A total of LO8 equally parsimonious trees were found of Jength 32 and consistency index 62 (Fig. 51). Stratigraphical occurrence data for relevant early Palaeozoic cleutherozoan taxa (Fig. 52) include all Canybrian edrioasteroids and (he eur- liest occurrences of all major post-Cambrian cleutherozoan lincages. By combining the cladistic analysis wilh stratgraphical data we arrive at ihe best corroborated evolutionary tree (Vig. 53), The origins of edrioasteroids may be traced back to the recently described late pre-Cambrian Arkarua (Gehling, 1988) from the Pound Sub- group (Rawnsley Quarizite) of South Australia. Arkarua is still poorly known, and we do not even know whether it had a blind gut or an anus. However, it shows more resemblance to Cambrian edrioasteroids than to any known ediacaran species and we tentatively accept it as an early echinoderm and the oldest known pen- taradiate species. The Cambrian fauna is better known and Paul & Smith (1984) suggested that Camptostrama represented an intermediate stage bridging (he morphological gap between pel- matozouns such as Kinzercysiy and CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA eleutherozoans such as Stromatocystites. Pel- matozoans and eleutherozoans were already dif- ferentiated by the late Lower Cambrian. Our cladistic analysis suggests that Stromatocystites represents a generalised eleutherozoan and that the origins of later eleutherozoan groups ¢an be traced back to It. There are several species currently included in the genus which is probably paraphyletic. We recognise two distinct lineages originating within Stromatocystites. From S. pentangularis one lineage gave rise to Totiglobus. Totiglobus has internal struts on aboral plates that Bell & Sprinkle (1978) interpreted as an attachment device. Such struts are also found in Stromatocystites pentangularis and S. reduncus, where they are better developed. Two important differences distinguish Totiglobus from Stromatocystites, the structure of the oral frame and the enlargement of the oral surface relative to the aboral surface. Whereas Stromatocystites has an oral frame formed from the 10 most proximal ambulacral ossicles Totig/obus and its relatives have an oral frame of 5 large interradial elements. This character is also seen in the Late Cambrian edrioblastoid Cambroblastus as well as in the Ordovician and younger Cyathocystida (i.e. Cyathocystidae and Rhenopyrgidae). The late Middle Cambrian ‘Totiglobus’ lloydi Sprinkle (1985) has a similarly shaped theca but the aboral portion of the theca appears in side view and, although broken off, could be inter- preted as forming a short peduncle of sub-tesse- late plates. In the slightly younger Cambroblastus the aboral surface has been greatly expanded into a polyplated stalk and this lineage continued into the Ordovician to give rise to the edrioblastoid Astrocystites. Cambroblas- tus is essentially the body of Totiglobus raised on a stalk. Because cyathocystids have an elongate, stalk-like aboral surface and because they have reduced their flooring plates to a single large deltoid lying interradially (Bockelie & Paul, 1983), we believe that they represent a sister group to Cambroblastus and later edrioblastoids. Rhenopyrgids probably have a similar reduction in ambulacral flooring plates, to judge from ex- ternal plating, but the precise arrangement has never been described. They differ from cyathocystids in having a polyplated, imbricate aboral surface rather than a single massive turret that is a single calcite crystal. We therefore agree 771 with Bell (1980) and Bockelie & Paul (1983) in suggesting that cyathocystids represent a distinct line of descent from isorophids and edrioasterids, and are more closely related to edrioblastoids. However, we also include rhenopyrgids within this group because of their aboral stalk pending confirmation of their ambulacral flooring plate arrangement. Another group with an aboral attachment disc and with an expanded oral surface relative to the aboral surface includes Walcottidiscus and its descendents the edrioasterids. These we believe represent a separate line of descent from a Totiglobus-like ancestor. In this lineage the aboral surface became uncalcified. The origin of isorophids remains problematic, but Hadrodis- cus may represent an intermediate morphology between edrioasterid-like forms and later isorophids. Hadrodiscus still retains large del- toid-like oral frame plates that lie interradially and may have had biserial flooring plates, al- though in external appearance it has a peripheral rim and the flattened streamlined appearance of an isorophid. The peripheral rim plates in isorophids appear not to be homologous with the peripheral ring in forms such as Edriodiscus, to judge from the cladistic analysis. They certainly appear rather different; in Edriodiscus the mar- ginals are stout, barrel-shaped ossicles that abut, whereas in isorophids the marginals are thin im- bricate plates that overlap. However, there remains the possibility that isorophids were derived from a more disc-like form such as Cambraster. Stromatocystites walcotti, from the Lower Cambrian, appears different from other Stromatocystites spp. in being flat and disc-like, with slightly differentiated marginal ossicles. This dise-like form and rather flat, simple am- bulacral ossicles are also seen in the early Middle Cambrian Edriodiscus. As argued above, we believe that Edriodiscus is the sister group to Arche gonaster and thus a possible precursor to the major post-Cambrian eleutherozoan groups of asteroids and ophiuroids. The position of Cambraster is problematic because it shows a mixture of characteristics. On the one hand it has an oral frame composed of 5 large interradially positioned flooring plates (Smith, 1986) which link it to the Totiglobus group. On the other hand it has a virtually iden- tical peripheral ring and body form to Edriodis- 772 & . ‘| I o rae 5 ow» 3 hd Ks o-— 5 Bs 02 5 3 € 6 8 & 46% 4 6 o & © ww 9 a -. 3S © o a gt G ees — E ~ &@ ge ®% 5 on 4 OW WU 16088 @® 0 7? 09 0 145 0 0 0 oo 8 @ 14 00080@¢6U%7?tUCUehlU8 13 0° 008 000 12 0008 0 8B @ 11 0908 5» @®@ o @ © 10 © @ 50 © co oc 9 09090 8 © 8 2 8 © oo 09 0 PF o T 9 09 89 80 Geo 6 © 0 &®© @ ®©@ 9 oOo 5 © 90006 ? 4 o oo 8s @® @ » @ co ee ee ee ee ee) 2 0 #e @® @ ® @ o 1°s @ @ @ @ @ eeee80e88000080 0 e Totigiobus MEMOIRS OF THE QUEENSLAND MUSEUM e@oMBee008800084000 Rhenopyrgus eeMe00e880008007 Cambroblastus @eeoeeseonvo0onro 0n Walcottidiscus e@ee08880880080 0-~ Edriophus eeee0006886888 0 0 ~ Hadrodiscus @0ee08808y0 0 @~ 0 0 Chatsworthi eeee8e00 MP ee © Bo 0o~ Foerstediscus eee eo0evryv 3007000 ‘7’ Ioydi Fic. 5). Character matrix and cladogram for Cambrian edrioasteroids and related taxa. Characters 1-16 are listed in Table 1. symbols are as follows:- open circles = plesiomorphic state. solid symbols = derived state (circles, squares and triangles indicate different character states). ? = character state unknown. cus. For {he moment we have accepted its place- ment as shown in the cladogram (Fig. 51), pend- ing further information. ECOLOGICAL DIVERSIFICATION Having examined the phylogenetic relation- ships of the major edrioasteroid groups we can now consider the palaeobiological significance of the various evolutionary changes that oc- curred. We trace edrioasteroids back to the Lower Cambrian Cumprostroma, which lies close to the latest common ancestor of pel- matozoan and eleutherozoan echinoderms (Paul & Smith, 1984, Smith,1988b). Camptostroma has a broad, conical aboral surface that was con- tractile and it lived with its pointed base inserted into unconsolidated sediment. It was a sedentary suspension feeder using its curved ambulacra to collect food particles which were then passed to the mouth along the food grooves. Most later groups specialised for attachment to firm or hard substrata and were thus able to colonize higher energy environments and presumably gained access to a richer food source. The evolution of attachment in edrioasteroids is thus seen as a direct adapta- tion for resisting dislodgementin currents. The earliest forms (Sfromatocystites) retained a fully plated aboral surface which was modified into an attachment pad. They were certainly not per- manently attached to a substratum but probably used their aboral surface to adhere to firm sedimentary bottoms. No Strematocystites is known to be attached to any form of hard sub- stratum, such as a shell and they appear to have been low level suspension feeders living in CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA 773 Thecal plating multilayered Thecal plating Imbricate aboral plating Conical base Ambulacra curved Epispires present Aboral surface fully plated Oral surface weakly domed Cover plates a multiplated skirt ] layer only Aboral plating a tesselate pavement (@) flat (@) aboral stalk Ambulacra straight Epispires lost Aboral surface partially or completely uncalcified Oral surface strongly inflated (@) Cover plates of large primaries plus small secondaries (@) Cover plates large, with intrathecal portion Mouth frame of 10 ambulacral plates Anal opening a cone of lath-shaped plates No peripheral mm of plates Marginals absent Mouth frame of (@) five fused interradially positioned elements (®) five radial elements that are not flooring plates. (&)five uniserial flooring plates (@)Anal opening a periproct of small platelets (®)anal opening aboral or lost i Peripheral rim plates (@) marginal ring of large imbricate plates (®) marginal ring of stout block-like ossicles Flooring plates of S, pentangularis type Ambulacra restricted to oral surface Marginals No intemal ribbing on aboral surface Flooring plates of (@) S. walcotti type (®) isorophid type. Ambulacra extend through marginal rim Marginals intergral part of both oral and aboral surfaces. Internal ribbing developed TABLE 1. Characters used in the phylogenetic analysis of early Palaeozoic eleutherozoans (autapomorphies ommitted). moderately protected environments. Edrioasterids, from Walcottidiscus onwards, continued in this mode of life but became uncal- cified over the central part of their aboral surface, presumably to improve adhesion. From this basic attachment style there developed 4 specialised offshoots. Firstly, Totiglobus evolved towards becoming a higher level suspension feeder by reducing its aboral attachment pad and expanding its oral, food- gathering area into a globular surface. Attach- ment remained identical to that in Stromuatocys- tites and we do not believe that Totiglobus was capable of resisting strong currents, Secondly. isorophids became much more streamlined in profile, with a well-developed peripheral rim of plates to strengthen the theca. Isorophids have completely uncalcified aboral surfaces and are generally found attached to hardgrounds or shells. They thus lived limpet-like in areas of relatively strong current activity and were able to invade many shallow water habitats previously unavailable to edrioasteroids. Thirdly, cyathocystids evolved even more secure attuch- ment by cementing themselves to hardgrounds. Their entire aboral surface became a single cal- 774 n 4 ° _ c 3 & < ie) a > ie) ° so] = ie) — o ~~ ” 0 a & E Oy oO c S&S = a E oak o -— 5 5 20 9 sO o oa 6 ” 3 | ~ Qa wn E ~ © Archegonaster \Edriodiscus MEMOIRS OF THE QUEENSLAND MUSEUM “ ” 3 6 oO ° os a rr) 3.4 5% as) > Of S o =c oo > s a) Q Ro Y no) x inom, i id eo = if] x oy % af ° 3| of >|? o uw wg choi = 9 rr) f = 2 % 2.5 o ~~ O Ss &/ ~ ” 5 3 vo Wo bya as aI oD ° ra) So o oe a 5° 2.5 = 3. fk = of ¢ x ® e (75) ° camaial, Fic, 52. Stratigraphical range chart for early Palaeozoic edrioasteroids and related taxa. Heavy lines = known stratigraphical range. thin lines show implied evolutionary relationships. cite tube which was firmly attached to a hard substratum such as a shell. These forms must have been particularly well adapted for life in high energy regimes. Fourthly edrioblastoids also evolved an aboral attachment disc and fixed to hard substrata, though precisely how is still unknown. Their aboral surface became trans- formed into a stiff stalk which elevated the food gathering area well above the substratum and they represent the only edrioasteroids to become relatively high-level suspension feeders. Presumably they were adapted for habitats of intermediate current strengths. Two edrioasteroid lineages departed from the attached mode of life to gain access to different habitats. Rhenopyrgids became infaunal suspen- CAMBRIAN EDRIOASTEROIDS FROM AUSTRALIA high-level suspension feeder semi-infaunal in mobile sediments Pelmatozoans Rhenopyrgids semi- infaunal in unconsolidated sediments aes Camptostroma Stromatocystites sessile epibenthic Suspension feeder Arkarua low-level Cyathocystids Totiglobus 775 permanently fixed to hard substratum high-level Edrioblastoids low-moderate energy semi-permanently attached Edrioasterids moderate -high energy semi-permanently attached to hard substratum lsorophids unattached sessile benthic deposit /suspension feeder; mouth up vagile deposit feeder mouth down Edriodiscus Archegonaster Fic. 53. Ecological diversification of early Palaeozoic eleutherozoans. sion feeders in mobile, unconsolidated sedi- ments. Their aboral surface is modified into a long, flexible and highly extensible peduncle which terminates in a basal sac. This peduncle we believe was inserted into the sediment so that rhenopyrgids were firmly anchored. The peduncle could then be used to retract the animal into its burrow, in an analogous way to the brachiopod Lingula, and was presumably adapted for life in more mobile sedimentary substrata. The other lineage to depart from attachment includes the flattened disc-like forms Cambraster and Edriodiscus. Both either lost, or never evolved, an aboral attachment disc and have simple aboral plating instead. They presumably were low level suspension feeders in protected habitats where attachment was un- necessary. Both have a stout marginal frame to strengthen the body. This line we believe gave rise through inversion to the free-living forms such as Archegonaster which were deposit feeders living with their oral surface towards the substratum. From them evolved the asteroids and the opuroids. 7716 THE CAMBRIAN DIVERSIFICATION As noted in the introduction, there are two contrasting explanations as to why Upper Cambrian echinoderms are uncommon. One ex- planation is that evolutionary diversification oc- curred in two phases, the first in the Lower and Middle Cambrian and the second in the Or- dovician (Sprinkle, 1980a, 1981; Campbell & Marshall,1987). The alternative explanation is that diversification was continuous throughout this period but that Upper Cambrian faunas were rarely preserved (Smith, 1988). The two models make very different predictions about the kinds of Upper Cambrian echinoderms that should be discovered, Under the two phase diversification model, Upper Cambrian echinoderms are numerically depauperate and were remnants of the Middle Cambrian fauna which survive through to the Early Ordovician, whereas, under the continuous diversification model Upper Cambrian echinoderms are expected to comprise intermediate taxa linking Middle Cambrian and Early Ordovician faunas. Our collections from the Chatsworth Lime- stone, described here, represent the best Upper Cambrian echinoderm fauna so far reported. Al- though the number of Upper Cambrian species is small, they clearly bridge the gap between Mid- dle Cambrian and Ordovician taxa. Ridersia is intermediate between Gogia-like eocrinoids and glyptocystitid rhombiferans, Hadrodiscus is in- termediate between Stromatocystites-like edrioasteroids and Jebetodiscid isorophids and Cambroblastus is intermediate between edrioasteroids and edrioblastoids. From this it would appear that echinoderms continued to diversify through the Upper Cambrian but have simply not yet been discovered, due to preserva- tional failure. We predict that many more groups will be found to have an Upper Cambrian record when suitable fossiliferous horizons are dis- covered, ACKNOWLEDGEMENTS The research for this paper was made possible by a Royal Society Study Visit Grant to A.B. Smith; field work in western Queensland and laboratory preparation of specimens was funded by the Queensland Museum. Material was made available to us for study by R. Prokop (Narodni Museum, Prague) and D.J. Holloway (Museum of Victoria, Melbourne) for which we are grate- ful. Chris Paul (Liverpool University) reviewed MEMOIRS OF THE QUEENSLAND MUSEUM an earlier draft and made many useful sugges- tions. We should like to thank the photographic studio of the BM(NH) for some of the photographs of Archegonaster and Gary Cranitch for assistance with the other photog- raphy. LITERATURE CITED BASSLER, R.S. 1938. Pelmatozoa, Palaeozoica. Fos- silium catalogus 1: Animalia, 83.(W. Junk: Ber- lin). 194pp. BaTHER, F.A. 1900. The Edrioasteroidea. /” Lankester, E.R. (ed.), ‘A treatise on zoology, part 3. The Echinodermata.’ pp. 205-216. (Adam and Charles Black: London). 1914. Steganoblastus. Geol. Mag. n.s. dec. 6 1: 193-203. BELL, B.M. 1976. A study of North American Edrioasteroidea. Mem. N.Y. St. Mus. 21: 1-447. 1980. Edrioasteroidea and Edrioblastoidea. Univ, Tenn, Dept. geol. Sci., Stud, Geol, 3: 158-174. BELL, B.M. AND SPRINKLE, J. 1978. 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AND SHeERGOLD, J.H. 1971, Cyelocystoides from the early Middle Cambrian of northwestem Queensland. Palueontology 14: 704-710. HENDLER, G, 1978, Development of Amphioplus ab- ditus (Vernll) (Echinodermata; Ophiuroides), U, deseription and discussion of ophiuroid skeletal ontogeny and homologies. Bre, Bull, 154: 79-95. Hubsos. G.H. 1925, The need for improved technique in illustration../. Geel, 33: 642-657. (927. The surface characterises of Asrroeystites (Stegunoblastus) ovawaensiy, Rept Vermant St. Geal.. 15: 97-110. TabKeL, O. 1923. Zur Morphogenie der Asterozoa- Palaont, Z. 3: 1-28. JeLt, PLA,, BURRETT, C.F. AND BANKS, MLR. 1985. Cambrian and Ordoyician echinoderms from eastern Australia. Aleferinga 9: 183-208, Jones, P.J. AND McKenziz, K.G. 1980. Queensland Middle Cambrian Bradoriida (Crustacea): new taxa, palaeobiogeagraphy and biological af- finities. Alcherinua 4; 203-225, KRSLING, R,V. AND VASSEUR, D. LE, 197]. 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British Palaeozoic Asterozoa, pt. 6. Palaeon- togr. Soc. Monogr., 237-324. 1951. Early Palaeozoic starfish. Phil. Trans. R. Soc. B 235: 87-129. SPENCER, W.K. AND WRIGHT, C.W. 1966. Asterozoans. pp. S4-107. Jn Moore, R.C. (ed.), ‘Treatise on invertebrate paleontology. Part U, Echinodermata 3.’ (Geological Society of America and University of Kansas Press: Lawrence, Kansas). SPRINKLE, J. 1973. Morphology and evolution of blas- tozoan echinoderms. Spec. Publ. Mus. comp. Zool. Harvard, 284pp. 1976. Biostratigraphy and paleoecology of Cambrian echinoderms from the Rocky Moun- tains. Brigham Young Univ. Geol. Stud. 23: 61- 73% 1980a. Early diversification. Univ. Tenn. Dept. geol. Sci., Stud. Geol. 3: 86-93. 1980b. Origin of blastoids: new look at an old problem. Geol. Soc. Amer. Abstr. Prog. 12(7): 528. 1981. Diversity and evolutionary patterns of Cambrian echinoderms. U.S. geol. Surv. Open File 81- 743: 219-221. MEMOIRS OF THE QUEENSLAND MUSEUM 1982. Astrocystites. Paleont. Contr. Univ. Kansas, Monogr. 1: 307. 1985. New edrioasteroid from the Middle Cambrian of western Utah. Paleont. Contr. Univ. Kansas, Pap. 16: 1-4. TERMIER, H. AND TERMIER, G. 1969. Les Stromatocystitoides et leur descendance: Essai sur |’evolution des premiers Echinodermes. Géobios 2: 131-156. UBAGHS, G. 1975. Early Palaeozoic echinoderms. Earth Planet. Sci. Rev. 3: 79-98. WEBBY, B.D. 1968. Astrocystites distans sp. nov., an edrioblastoid from the Ordovician of eastern Australia. Palaeontology 11: 513-525. WHITEAVES, J.F. 1897. Description of anew genus and species of cystideans from the Trenton Lime- stone at Ottawa. Canadian Rec. Sci. 7: 287-292, 395-396. WHITEHOUSE, F.W. 1941. The Cambrian faunas of north- eastern Australia. Part 4: Early Cambrian echinoderms similar to the larval stages of Recent forms. Mem. Od Mus. 12: 1-28. LIMNODYNASTES FITZINGER (ANURA: LEPTODACTYLIDAE) FROM THE CAINOZOIC OF QUEENSLAND MICHAEL J. TYLER Tyler, M.J. 1990 08 31: Limnodynastes Fitzinger (Anura: Leptodactylidae) from the Cainozoic of Queensland. Memoirs of the Queensland Museum 28(2): 779-784. Brisbane. ISSN 0079-8835. Seventy-six frog ilia representing the leptodactylid genus Limnodynastes are reported from the Tertiary of Riversleigh Station, Queensland. Four of these are L. sp. cf. tasmaniensis and the remainder L. antecessor sp. nov. Two ilia of L. ornatus (Gray) are reported from a Quaternary cave deposit at Riversleigh Station. [JAnura, Leptodactylidae, Limnodynastes, Cainozoic, Queensland. Michael J. Tyler, Department of Zoology, University of Adelaide, Box 498 GPO, Adelaide, South Australia 5001, Australia; 9 September, 1989. The frog genus Limnodynastes Fitzinger is rep- resented by 12 extant species (Frost, 1985). The geographic range of the genus extends throughout Australia but for the Nullarbor Plain and western Eyre Peninsula in South Australia. One species, L. convexiusculus (Macleay), reaches southern New Guinea (Zweifel, 1972). The fossil record of Limnodynastes is prin- cipally from Quaternary sites, and includes four extant species (Tyler, 1989a). The Tertiary iecord is restricted to the holotype of L. archeri Tyler (1982) from the Etadunna Formation at Lake Palankarinna, north of Lake Eyre, South Australia. The phylogenetic affinities of L. ar- cheri clearly lic with L. ornatus (Gray) and L. spenceri Parker: a pair of species that is so dis- tinctive that recognition of a separate genus for them may be justified (Tyler, Martin and Davies, 1979; Roberts and Maxson, 1986). Given that the latest evidence suggests that L. archeri is from the Oligocene (Lindsay, 1987), it is clear that Limnodynastes (sensu lato) is an old lineage, and the fossil record assumes particular significance. A large number of frogs has been recovered from Tertiary sites at Riversleigh Station in northwest Queensland. The first species to be reported there was Lechriodus intergerivus Tyler (1989b) and, to date, 315 (50%) of the ilia located represent that species. Numerically the second most dominant genus in the samples is Lim- nodynastes. Here I report the Tertiary taxa, and describe one as a new species. In addition | document the finding of L. ornatus at a Quater- nary site there. MATERIAL AND METHODS The specimens reported here were obtained at the Vertebrate Palacontology Laboratory at the University of New South Wales. They are now deposited in the collections of the Queensland Museum (QM), and the South Australian Muscum (SAM). Letters preceding registration numbers are departmental identifications. Unas- signed specimens retain the Vertebrate Palaeon- tology Laboratory reference numbers, which are prefixed AR. The material consists of isolated and common- ly fragmentary ilia. Their safe manipulation has been greatly simplified by the use of a Cosy M200 vacuum pump with an M202 probe which has a terminal compatible with Luer hypodermic syringe needles. The size of the ilia examined was such that 21 to 26 gauge needles were used. To buffer the contact between needle tip and specimen, a short sleeve of silicon rubber tubing was attached to the needle tip. The descriptive format of the material follows Tyler (1976, 1989b). SYSTEMATICS Family Leptodactylidae Limnodynastes Fitzinger The generic characteristics of the ilium have been defined by Tyler (1976), and exhibit varia- tion according to the species group involved. Hence L. ornatus and L. spenceri exhibit a dis- 780 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 1. S.E.M. of QM F17517: distal end of paratype of Limnodynastes antecessor. tinct crest to the ilial shaft, the 2. dorsalis group a groove upon the medial surface of the shaft, and L, tasmaniensis an anteriorly protruding. ventral acetabular expansion. The extension of the dor- sal prominence and dorsal protuberance is equal- ly variable. All specics share an clongate and tapering dorsal acetabular expansion. Limnodynastes antecessor sp. nov.’ (Fig. 1) MATERIAL EXAMINED Hovotyee: QM Fi7510. An almost entire right ilium collected at Gag Site, Riversleigh Station, Queensland. PARATYPES: There are 18 paratypes: C.S, Site: SAM P29794, P30038, P30042, QM F17518: Gotham Site: SAM P29795; Upper Site: QM F17511-13, 17516-17; SAM P29797, P30050;, Gag Site: QM F17514-15; Wayne's Wok Site: QM F17616-18: 17517 subject of S.E.M.). DESCRIPTION OF HOLOTYPE Ilial shaft slender, slightly curved, cylindrical and lacks medial groove and dorsal crest. Acetabular fossa small, deep and with ex- tremely prominent rim anteriorly. Dorsal margin of acetabular fossa situated slightly superior to inferior margin of ilial shaft. Pre-acetabular zone largely vertical to ilial shaft and inferiorly merges insensibly into moderately developed ventral acetabular expansion. Dorsal acetabular expansion extremely elon- gate, tapering to acute point and extending supe- riorly above ilial shaft. Dorsal prominence extremely pronounced, rising high above the dorsal margin of ilial shaft. Dorsal prominence situated largely anterior to anterior rim of acetabular fossa, and exhibits CAINOZOIC LIMNODYNASTES 781 localised, circular depression on level with supe- rior margin of acetabular fossa. Dorsal protuberance poorly developed, Length of ilium 11.0mm; DAE-VAE 5,9mm; acetabular fossa maximum diameter 2.4mm, VARIATION Almost all specimens are incomplete to vary- ing degrees. The largest specimen is SAM P30038. Tt is complete and has a length of 21,5mm, The extent of the elevation of the dorsal prominence varies, such that in some paratypes L$ nol as pronounced as in the holotype. The localised cireular depression proximal to the dor- sa] prominence js variable, and is scarcely detec- table in several specimens. REFERRED SPECIMENS Portions of an additional 53 ifia are referred to ihis specics but ure so incamplete that they do not contribute to an understanding of specific char- acteristics. For that reason they have been ex- cluded from the type series. Unless indicated specimens have been lodged in the Queensland Museum: R.S.O. Siles F17591, (7527-8; Upper Site; FI7519, 17522-23, 17529, 17595, SAM P30005, P30050, AR14433, F17621-2; C.S. Site: F17520-1, 17592, 17825, 17599, 17587, 17589, 17623, SAM POO, P3039, PRU043, P30046-47, ARI1617; Gag Syte; F17524, 17000, 17596, 17526, 17594, 17530-2, 17534, 17619. 17625-5; R-TS.2 SAM P30044-45; Wayne's Wok: F17593, 17587, 17620, 17624; Out- asite: F17533; Henk's Hollow: F17588, SAM P30048; R.V, Site: 17590; Neville’s Garden Sile: SAM P30040-41, P30049, Four of these specimens (SAM P30005-4, AR11617, 14433) are substantially larger than other referred specimens and the type series, Reconstructions indicate that they are derived from individuals with a snoul-vent length range of approximately 60- 80mm. Either they repre- sent the maximum size attained by L, antecessor, or they represent a distinct species. In the event that the latter suggestion 1s correct, sub-adull specimens may be indistinguishable from L. an- tecessor. Given the few specimens available | prefer to accept at this stage that they are con- specific; resolution awaits the recovery of com- plete specimens. COMPARISON WITH OTHER SPECTES From L. orraius (Gray) and L. spencert Parker the new species is distinguished by the lack of a dorsal crest upon the ilial shaft. From L. tas- maniensts Giinther it differs principally in the nature of the pre-acetabular zone and ventral acetabular expansion which protrude con- spicuously in that species but not in the new species. Members of the L. dumerilii species group have the superior rim of the acetabular fossa above the level of the inferior margin of the ilial shaft, whereas in. attecessor it lies below iL. Limnodynasies salmint Steindachner has a very small, laterally situated dorsal prominence and protuberance, whereus these features in the new speeics extend superiorly and thus are more conspicuous. Limnodynastes convextusculus is distin guished by possession of a slightly developed dorsal crest to the lial shaft, absent ind. anteces- sor. The closest atfinity of the new species lies with i. peron (Duméril and Bibron). Both species have the acetabulum situated in a position in- (erior to the ilial shaft, a well developed acetabular rim, and a lack of longitudinal inden- lations upon the lateral and medtal surfaces of the ilial shaft. Limtnodynustes antecessor is distinguished from L. perani by its less pronounced dorsal prominence, a more acute angle between the dorsal acetabular expansion und the dorsal prominence, and by having a more concave ventral acetabular expansion. LOCAL DISTRIBUTION AND-AGES Limnodynastes antecessor occurs over a wide tange of the Riversleigh Station sites. Archer, Godthelp, Hund and Megirian (1989) indicate that more than 97 sites and corresponding local faunal assemblages have been found there. They have attempted to equate the major of those sites with Sites A-E of Tedford (1967). The 1, an- fecessor localities are thought to date from early to late Miocene. ETYMOLOGY The specific name is the Latin for ‘ancestor’, so alluding here to the ancestral nature of the taxon. Limnodynastes ef. tasmaniensis Gunther MATERIAL EXAMINED Four ilia, SAM P30007 R.S.O. Site, SAM P30008 Upper Site, OM F17627-28 CS. Site, Riversleigh Station. Queensland, 782 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Photograph of SAM P29798: Limnodynastes ornatus from Message Stick Cave, Riversleigh Station. DESCRIPTIVE NOTES The specimens form two distinct size groups for whereas SAM P30007 lacks most of the ilial shaft, the length of the proximal portion (4.7mm) equates roughly with a 15mm ilial length, and an approximate body size of 4Umm, the remainder are from much smaller individuals. One (SAM P30Q08) is complete and measures 4.6mm. A second (QM F17627) is 5.0mm and the remain- ing individuals incomplete but of comparable size. These measurements indicate a body length of 15-20mm. Thus the conspecificity of the material, let alone identity, remains suspect. This is likely to be resolved if more representatives become available. The association with L. tasmaniensis hinges upon the protuberant nature of the sub-acctabular zone so thal the margin anterior to the acetabular region is sigmoid (narrowing in the pre- acetabular zonc). Contrasting with this evident affinity is the finding that there ts a distinct ontogenetic change in the form of the ilial shaft in L. tasmaniensis. Metamorphlings and small sub-adults exhibit a dorsul crest characteristic of the L. ornatus species group. Such a crest is Jost in ontogeny and is lacked by the small fossils. The significance of this feature is being inves- tigaled as part of a broader study of ontogenctic trends of skeletal features in the genus in col- laboration with M. Davies. Limnodynastes ornatus (Gray) (Fig. 2) MATERIAL EXAMINED Two ilja, QM F17598, SAM P29798, Message Stick Cave Site, Riversleigh Station, Queensland. DESCRIPTIVE NOTES The identification is based upon a combination of two features: firstly the slightly developed dorsal crest of the ilial shalt (shared by L. ornatus and /.. spenceri), and the fact that the ilial shaft is curved; itis straight in L. spenceri. SAM P29798 is a complete ilium, with a total length of 12.7mm, a DAE-VAE distance of 4.2mm, and a fossa diameter of 1.8mm. QM F17598 lacks the distal portion of the ilial shaft and measures 11.0mm; the DAE-VAE distance is 4.1mm, and the fossa diameter is 1.6mm. COMMENTS The donor of the larger of the ilial specimens would have had a snout to vent length of ap- proximately 31mm (Fig. 3) which ts at the base of the size range of adult males, and less than the range of adult females (Tyler, Smith and Johnstone, 1984), Message Stick Cave Site is an unreported Quaternary site, The record of L. arnatus represents the first fossil record of that species, and the first Quaternary Trog to be reported from Queensland. DISCUSSION To date 623 anuran lia have been recovered from the Tertiary sites at Riversleigh Station, Of that total 76 represent Limnodynastes, indicating that the genus was numerically a signficant com- ponent of the fauna. It is of interest to examine whether the presence of Limnodynastes supports the concept of Riversieigh being cool, temperate rainforest in the Tertiary: a conclusion indicated by the predominance of Lechriodus there (Tyler, 1989b; Tyler, Hand and Ward, in press.). CAINOZOIC LIMNODYNASTES “40 35 . ey ee ee eee fe 30 /\ o/ y= -225 +2.62« if 35 r2~ 0.98 } f s i a | on | | | Snout to Vant Length (mm) fe oa 2 4 8 16 12 14 i6 6 (lum Length (mm) Fic, 3. Regression line of ilium length of Lim- nadynastes ornatus wilh snout-vent length. Es- timated length of largest fossil specimen indicaled by broken lines, t-value for slope 21,9735, p, .001. Por x= 12.7mm, y = 31.06mm (95% confidence limits = 30,79-3 1,33). Modern Limnodynastes occupy a diverse range of habitats, and two adaptive modes can be recognised: robust, fossorial forms with shovel- shaped, metatarsal tubercles, or more slender, sharper snouted frogs lacking enlarged metatar- sal tubercles (Tyler, Watson and Martin, 1981). There is no absolute association between these modes and the environment. This is because fossorial adaptations occur in species that inhabit extremely demarcated wet-dry climatic regimes (c.g. L. spenceri), and also in those that spend shorter periods below the ground in temperate zones (e.g. L. dumerilit). The phylogenetic affinities of the Riversleigh Tertiary Limnodynastes species lic with two taxa that are not fossorial, Both of these species ex- tend throughout the East and Southeast of the continent, breeding in static or slow-moving water, but more frequently encountered in open country rather than in rainforest. Thus their presence does not conflict with the assumption of a temperate rainforest but neither does it sup- 733 port that interpretation. Limnodynastes species may be too labile in terms of their ecological requirements to be good indicators of palacoen- vironments. A biological feature shared by Lechriodus and Limnadynastes 1s the habit of depositing eggs in foam nests that float upon the surface of water. The process by which the foam nest is produced in the two genera is identical (Tyler and Davies, 1979), Given that this shared biological feature originated in an ancestor to the genera, the origin of the habit is clearly ancient, and which was exhibited by 65% of the individuals at Riversleigh in the Tertiary. ACKNOWLEDGEMENTS lam extremely grateful to Dr Michael Archer, Dr Suzanne Hand and Mr Henk Godthelp of the University of New South Wales, for theit con- tinued aid and support throughout my study of the fossil frog fauna at Riversleigh Station. Laboratory studies have been funded by agrant from the Australian Research Council. Miss Veronica Ward played a vital role in the sorting and documentation of the material, and prepared the illustrations. Research facilities were provided by the University of Adelaide. The assistance of Parke Davis Pty Ltd is gratefully acknowledged. The materials upon which this siudy was based were obtained through the support of the follow- ing funding bodies and organisations lo M, Ar- cher, S. Hand and H, Godthelp; Australian Research Grants Scheme; Department of Arts, Sport, the Environment, Tourism and Ter- ritories,; National Estate Programme Grant Scheme; Wang Computers Pty Ltd; Australian Geographic Pty Ltd; Mount Isa Mines Pty Ltd; the Queensland Museum; the Australian Museum; the Royal Zoological Society of N.S.W.; the Linnean Society of N.S,W.; An- sett/Wridgways Pty Ltd; Mount Isa Shire Coun- cil; the Riversleigh Society and the Friends of Riversleigh. LITERATURE CITED ArcHer, M., GopoTHELP, H., HAND, S.J. AND MEecigian, D, 1989. Fossil mammals of Riversleigh, northwestern Queensland: prelimi- nary overview of biostratigraphy. correlation and environmental change. Australian Zoologist 25(2): 29-65. Frost. D.R. 1985. ‘Amphibian species of the world". 784 (Allen Press and Association of Systematics Col- lections: Lawrence, Kansas). Linpsay, J.M. 1987. Age and habitat of a foraminiferal fauna from near-type Etadunna Formation, Lake Palankarinna, Lake Eyre Basin. South Australian Department of Mines and Energy Report 87/93, unpublished. ROBERTS, J.D. AND MAXSON, L.R. 1986. Phylogenetic relationships in the genus Limnodynastes (Anura: Myobatrachidae): a molecular perspec- tive. Australian Journal of Zoology 34: 561-73. TEDFORD, R.H. 1967. Fossil mammals from the Carl Creek Limestone, northwestern Queensland. Bulletin of the Bureau of Mineral Resources, Geology and Geophysics, Australia 92: 217-36. TYLER, M.J. 1976. Comparative osteology of the pel- vic girdle of Australian frogs and description of a new fossil genus. Transactions of the Royal Society of South Australia 100: 3-14. 1982. Tertiary frogs from South Australia. Al- cheringa 6: 101-3. 1989a. ‘Australian frogs’. (Viking O’Neil: Mel- bourne). 220 pp. 1989b. A new species of Lechriodus (Anura: Lep- todactylidae) from the Tertiary of Queensland, with a redefinition of the ilial characteristics of the genus. Transactions of the Royal Society of South Australia 113: 15-21. MEMOIRS OF THE QUEENSLAND MUSEUM TYLER, M.J. AND DAviEs, M. 1979. Foam nest con- struction by Australian leptodactylid frogs (Am- phibia, Anura, Leptodactylidae), Journal of Herpetology 13(4): 509-10, TYLER, M.J., HAND, S.J. AND WARD, V.J. in press. Analysis of the frequency of Lechriodus inter- gerivus Tyler (Anura: Leptodactylidae) in Oligo-Miocene local faunas of Riversleigh Sta- tion, Queensland. Australian Zoologist . TYLER, M.J., MARTIN, A.A. AND DAVIES, M. 1979. Biology and systematics of a new limnodynas- tine genus (Anura: Leptodactylidae) from north- western Australia. Australian Journal of Zoology 27: 135-50. TYLER, M.J., SMITH, L.A. AND JOHNSTONE, R.E. 1984. “Frogs of Western Australia’. (Western Australian Museum: Perth). 108 pp. TYLER, M.J., WATSON, G.F, AND MARTIN, A.A. 1981. The Amphibia: diversity and distribution. pp. 1277-1301. In A. Keast (ed,) ‘Ecological biogeography of Australia’, (W. Junk: The Hague). ZWEIFEL, R.G. 1972. A review of the frog genus Lechriodus (Leptodactylidae) of New Guinea and Australia. American Museum Novitates 2507: 1-41, MARINE TURBELLARIA (ACOELA) FROM NORTH QUEENSLAND LEIGH WINSOR Winsor, L.. 1990 08 31: Marine Turbellaria (Acoela) from north Queensland. Memoirs of the Gueenslund Museum 28(2): 785-800, Brisbane. ISSN 0074-8835. Seven species of free-living acoels are now known from Australia. A new genus, new species and new combination of haploposthiid acoels ure reported. They are Waminoa lilus gen. el sp. nov., lwo non-sexual Waminoa sp., Convolulriloba hastifera sp. noy., and C. Japonica (Rata, 1951) vomb. nov... The diagnosis forthe genus Convelutriloba Hendelberg, and Akesson, 1988, is emended. The heterogencous convolutid genus Amphiscolops is reviewed, and the genus Feferochaerus reinstated and emended ta accommadute MM. australis Haswell, 1905, and //. sargassi (Hyman, 1939) comb. nov. New records of 7. australis and Convolutriluba ef. retrogemma are reported. A key to free-living acoel genera Jeseribed [rom Australian waters is provided. 7 Acwela, topical marine flarworms, iayonomy, key, Waminoa, Convolutriloba, Amphiseolups, Heterochaerds. Leigh Winsor, Depariment of Zoology, Jamey Cook University of North Queensland, Townsville, Queensland 4811, Australia; 23 Seprember, 1989, Little is known of the free-living turbellarian acvel fauna of Australia, Until now only two species, both intertidal convolutids, have been described. They are Amphiscolops australis from Port Jackson, New South Wales, (Haswell, 1905) and Wulguru cuspidata from Townsville, north Queensland (Winsor, 1988), This paper is mainly based upon collections of acocls from north Queensland waters. Itincludes descriptions of new taxa, a reappraisal of the genera Convolutrilaba (Haploposthiidac), Am- phiscolaps and Heterochaerus (Conyolutidac), and a key to genera of tree-living Acoela described from Australia. MATERIALS AND METHODS Squash preparations of specimens were ex- amined by differential interference contrast microscopy. Other material was narcotized with 7.5% magnesium chloride and fixed in marine Bouin's fluid. Specimens were processed to paraffin wax, serially sectioned at S-7}.m and stained with iron hacmatoxylin-eosin, trichrome stains and selected histochemical methods (Win- sor, 1984). Wholemounts were stained by car- mine, iron haematoxylin and histochemical methods. Specimens for clectron microscopy were fixed in 2.5% glutaraldehyde in sea-water. For scan- ning electron microscopy (SEM) specimens were critical point dricd, gold coated and ex- amined jn secondary mode usingan ETEC-SEM. Tissues for transmission electron microscopy (TEM) were post fixed in 1% osmium tetroxide. processed to Spurr’s resin, sectioned at 70- 9Onm, stained with uranyl acetate and lead citrate, and examined using a JOEL FX200 TEM. Terminology follows thal proposed by Cannon (1986). Type and voucher specimens prefixed G are lodged with the Queensland Museum (Townsville Branch). Material from the Australian Museum, Sydney is prefixed AM.W. The following abbreviations ure used in the Figures: a - anterior end, bursa; as - algal sym- biont; b - bursa; be - bursal canal; bg. - basiphil glands: ¢- cuboidal epithelium, ce - concrement cells em-cell mass (7 sensory function); d- duct; e - eye; em - epidermal musculature; f - lateral body Jold; fa -female antrum: fo- frontal organ; fv ~ false seminal vesicle: g - ganglion; m - mouth; me - matrix cells. ; nm - nucleus; 6 - oocyte; OV - ovary: pS - prostatic secretions; pt - pit, frontal organ; r - rhabdoids; s - sperm: sb - sagittocystoblast: sg - sagittocyst; st - statocyst- statolith: sv - seminal vesicle; t - testes; y - yolk-producing part, ovary; d- male genital pore; ¥- female genital pore Haploposthiidae Westblad, 1948 Waminoa gen. nov. DIAGNOSIS Haploposthiidae discoid to obcordate in shape; colour resulls from pigments and symbiotic algae; often with two coexisting specics of algal symbionts: brain insunk; mouth ventral and in 786 posterior third of body; rhabdoids present, ovaries and testes paired; with sub-terminal male genital pore; male atrium ciliated; seminal bursa with paired lateral bursal canals. Statocyst- statolith, eyes, frontal gland, female genital pore and penis absent. Epizoic on corals. TYPE OF GENUS Waminoa litus sp. nov. ETYMOLOGY The genus name Waminoa is an Aboriginal word meaning companion and alludes to the occurrence of this acoel on corals. The specific epithet is from the Greek /itos meaning plain or unadorned, refers to the drab appearance of the species. Waminoa litus sp. nov. (Fig. la-f) Haplodiscus sp. (in part) Trench and Winsor, 1987 MATERIAL EXAMINED Type Loca.ity: Geoffrey Bay, Magnetic Island, 19°08’S, 146°50’° E. Collected by G. Bull from soft coral, 9.9.1979, HOLOTYPE: G23003, four microslides, longitudinal sagittal 5j.m sections, iron haematoxylin and eosin. PARATYPE: G23004, wholemount, iron haematoxylin. OTHER SPECIMENS EXAMINED: Geoffrey Bay, Mag- netic Island, collected by P. Osmond from the soft coral Sarcophyton sp. March, 1983; 30.4.1985; 21,5.86; 9.12.87; 13.4.89; Nelly Bay, Magnetic Island. Collected by P. Osmond from Sarcophyton sp. 13.4.1989. DESCRIPTION At rest living specimens are obcordate in shape and cinnamon- brown in colour with a translucent margin. Pigmentation is due to algal symbionts. The body is slightly narrower anteriorly than posteriorly, with a pronounced mid-caudal notch. Sexual specimens attain about 2mm maximum width. The mid-dorsal region ‘is generally slightly elevated and less pigmented than the rest of the body. Dorsal cilia are 54m long and are less numerous than the ventral cilia, 7m long. The dorsal epidermis is 2.5m thick and the ventral epidermis 5m thick. Longer sensory cilia are sparsely distributed over the body. Underlying the epidermis are circular, oblique and lon- gitudinal muscle layers. The dorso-ventral mus- MEMOIRS OF THE QUEENSLAND MUSEUM culature is weakly developed. Circular muscles surround the mouth and male genital pore. The brain is insunk and consists of a bilobed ganglion. Immediately posterior to the ganglion is anon-glandular cellular mass 251m by 50m, possibly sensory in function. The cells are slight- ly ellipsoid, 6-8y.m diameter with large evenly staining nuclei, prominent nucleoli and pale evenly staining cytoplasm. A statocyst-statolith, eyes, frontal gland com- plex and concrement cells are absent. Two species of algal symbionts are present. An Amphidinium sp. measuring 16um-24um diameter and a smaller Symbiodinium sp. ap- proximately 8m in diameter. The algae lie im- mediately below the epidermal musculature, and in some specimens appear almost completely to occupy the central parenchyma. Fusiform packets, measuring 27j.m x 5m, of translucent acicular rhabdoids are present in the dorsal epithelium. There are two types of non- thabditiform glands: a granular type fairly even- ly distributed over the dorsal surface, slightly more abundant anteriorly and absent in the region of the caudal notch. The granular secre- tions are found to be composed of neutral mucopolysaccharides. The other type of gland is mainly concentrated along the margins and ventral surface: These glands have amorphous acid mucopolysaccharide secretions. In living specimens numerous spherical refrac- tile bodies were observed underlying the epider- mis. In stained sections they are acidophilic and measure 5-7.m diameter. They do not appear to be associated with any gland or particular struc- ture, but lic within the parenchyma. These bodies give negative histochemical reactions for mucins and lipids. The mouth lies ventrally in the posterior third of the body. The endocytium contains amor- phous sedimentary particulate matter, diatoms and remains of unidentified crustacea. Testes are paired and lie dorso-laterally to the ovarics. Sperm pass posteriorly and accumulate before passing to the male antrum. The male genital pore is situated ventrally and subter- minally on a muscular papilla. The ciliated male antrum, which appears to be a simple invagina- tion of the ventral epidermis, communicates with a duct. The duct is well defined distally but indistinct proximally. There is no definite penial structure. The ovaries are paired, ventrally situated and extend posteriorly behind the mouth. The semi- nal bursa lies ventro-medially and posterior to TROPICAL ACOELS 787 @ 7 \ 1 RS ee \ o ; Ns ea hr ———— = | mur NiRigasenyra woe Fic, 1, Waminoa litus, (a) Dorsal aspect, living specimen, (b) ventral aspect, whole mount (for clarity the ovary on right side is not shown), (c) longitudinal section (d) male copulatory apparatus, (¢) bursa and bursal canal, and ({) dorsal view of bursa showing bursal canals (unscaled sketch), 788 MEMOIRS OF THE QUEENSLAND MUSEUM Fic. 2. Waminoa sp. 1. (a) dorsal aspect, living specimen by transmitted light, and (b) by incident light (black dendritic clusters represent pattern of concrement cells),(c) ventral aspect, wholemount. the base of the muscular papilla and is partially surrounded antero-ventrally by a cellular mass, in part comprised of matrix cells. From either side of this structure two short weakly sclerotised bursal canals project antero-laterally into the parenchyma. A female genital pore is absent. No duct communicates with the bursa. Waminoa sp. 1 (Fig. 2a-c) MATERIAL EXAMINED LOcALITy: Marine aquarium Australian Institute of Marine Science, Townsville. Collected by Dr S. Col- lard from corals, 19.11. 1981, and fixed in mercuric chloride-acetic acid. VOUCHER SPECIMENS: G23011 Grenacher’s carmine stained wholemount and G23012 a single microslide, horizontal 61m trichrome stained sections. DESCRIPTION Living specimens are approximately 2mm long and ovate to obcordate in shape. Examined under incident light, the dorso-lateral surfaces are mot- tled with dendritic iridescent-white concrement which is retained in fixed specimens. In colour and pattern the acoels exactly resemble the retracted polyps on the coral on which they were found. Under transmitted light these acoels ap- pear translucent brown, less dense along the dor- sal mid- line. The colour is due to symbiotic algae. Three unpigmented zones are present anteriorly. Fixed specimens are approximately 1.2mm diameter. The mouth is ventral and situated in the posterior third of the body. Statocyst-statolith, eyes and frontal gland are absent. The bilobed cerebral ganglion is insunk. There are two species of algal symbionts: the smaller species 6-74m diameter and the larger less numerous species 9-12.54m diameter. Male and female gonads and copulatory organs are absent. Waminoa sp. 2 MATERIAL EXAMINED LOCALITY: Pandora Reef, 18°49°S, 146°26’E. Col- lected by P. Alino from Acropora longicyathus 22.11.1987, and fixed in the field in marine Bouin’s solution. VOUCHER SPECIMENS: G23013 Grenacher’s carmine stained wholemount and G23014 a single microslide of serial horizontal 64m trichrome stained sections. DESCRIPTION Specimens are Imm diameter, discoid with slight mid-caudal notch with mouth in the ventral posterior third of the body. The colour when live was not noted. There are no dendritic flecks on the dorso-lateral surface. Two species of algal symbionts are present: a numerous small species 7-8.54.m diameter, and a larger less numerous specics | 1-14j.m diameter. Statocyst, eyes, fron- tal gland, male and female gonads and copulatory organs absent. TROPICAL, ACOELS SYSTEMATIC DISCUSSION In external morphology Waminoa species closely resemble those of the planktamie con- volutid Haplediscus Weldon, 1888. However with ventral subterminal male gonopore, simple tubular male antrum, and penis absent or possib- ly present only as a small papilla, Waminoa is assigned to the Haploposthiidae as defined by Cannon (1986). Within the Haploposthiidae Waminoa is cleat- ly differentiated from the other eleven genera by a combination of characters: the presence of two coexisting species of algal symbionts from which Waminea species derive their colour, presence of rhabdords, paired ovary and testis. an isolated seminal bursa with two weakly scleratised bursal canals, and absence of statocyst, cyes, and frontal gland. Warminoa is closest to Pseudoltaplogonarta Dérjes, 1968, from which it is distinguished chiefly by the presence of a well formed seminal bursa with paired lateral bursal canals, and from Deuterogonaria Dérjes, 1968, by the presence of the paired bursal canals and absence of a vesicular granulorum, The presence of an Amphidinium sp. and Sym- biodinium sp. in the tissucs of Waminoa, described by Trench and Winsor (1987) repre- sents the first observation of two dinoflagellate Species co-existing in the same cell of the inver- (ebrate host, though Yamasu und Okazaki (1987) have reparted the presence of two different algal symbionts in two species of the convolutid Am- phiscolops. At present Waninoa litus is the only species found with copulatory organs. Although there is no obvious penial structure in this species, the atrium may evert to form a penis. In two mature specimens a small ventral sub-terminal papilla is present in the region carresponding to the posi- tion of the male pore. Unfortunately the absence of sperm in the immediate vicinity of the papilla makes it difficult to confirm the nature of this structure. Waminoa sp.1 is distinguished from the other two species by the presence of concrement. Waminea sp. 2 found on a seleractinian coral, is smaller in diameter than W. /itus and Waminoa sp.1. As host specificity of the species is un- known it is considered prudent at present to distinguish W. litus and Waminoa sp. 2 though they may prove to be conspecific. The Haplodiscus sp. of Yamasu and Okazaki (1987) from Sesoko Island (Ryukyu [slands, 789 Japan) found on coral is possibly a species of Wamunaa. In their preliminary report the authors remark thal “no specimens bore male and female germ cells or vential organs. Body is brown due to symbiotic algae, an Amphidinium type dinoflagellate.’ Conyolutriloba Hendelberg and Akesson (1988)(emend) DIAGNOSIS Haploposthiidae with colour and pattern resulting from pigment and symbiotic algac: frontal gland present; statocyst may be absent; brain insunk; mouth ventral; rhabdoids and sagit- tocysts present; ovaries and testes paired: ovaries divided into oocyte-producing and yolk-produe- ing parts; with two ventral genital openings; male antrum ciliated passing directly into semi- nal vesicle: without penis; female antrum and vagina ciliated; seminal bursa with single sclerotised bursal canal, TYPE OF GENUS Convalutriloba retrogemma Hendelberg and Akesson, WKS Conyolutriloba cf. retrogemma Hendelberg and Akesson, 1988. MATERIAL EXAMINED Locacivy: The marine aquarium, University of New South Wales, found on juvenile Tridacna gigas clams. Specimens fixed in Bouin’s fluid were forwarded by L. Goggin. VOUCHER SPECIMENS: G23007 wholemount stained Grenacher’s carmine, and G23008 four microslides of serial horizontal 6um sections, haematoxylin and eosin, DESCRIPTION Non-sexual specimens |-2mm long are similar in external morphology to C. retregemma and correspond to the original description. Sagit- tocysts are nol present in whole mounts or sec- tioned specimens. Algal symbionts measured 5.4- 9m diameter. Conyolutriloba hastifera sp. nov. (Figs 3a-f, 4a-f) MATERIAL EXAMINED Tver Locauimy: Australian Institute of Marine Science (ATMS) Cupe Cleveland, near Townsville, 74H north Queensland. Collected from the aquarium by Dr Clive Wilkinson 21.9.1983, HoLotyrr: G2Hi05 four microslides, longitudinal saggital sections Sum, trichrome stain. PARATYPE: G23006 wholemount, Gower's carmine. OTHAR SPACIMENS EXAMINED, Sexual specimens found on soft coral, Geoffrey Bay, Magnetic Island, collected by G. Bull 9.9.1974: specimens collected from the marine aquarium at AIMS hy Dr Clive Wilkinson: November 1983, 27.11.84, 30.1.85. ETYMOLOWY The specific epithet Aaséifera is derived from the Latin hasta - a spear, and ferens - bearing, and refers (0 prominent anterior batteries of sagitlocysts present in the species. DESCRIPTION Living specimens at rest measure up to 3mm long and 1.5mm wide. The body is dorsoventral- ly flattened, broad and rounded anteriorly, It narrows slightly in the mid third of the body, then broadens posteriorly with two lateral caudal lobes and smaller median lobe. Jn life the median lobe is generally slightly clevated and does not project posteriorly beyond the lateral lobes. The ground colour is a translucent greenish- yellow to greenish-brown and is primarily the result of symbiotic algae which are fairly uniformly distributed dorsally und ventrally. A fine orange-red pigmentis concentrated dorsally mainly uround the base of the median caudal lobe and along the lateral body margins. It is scattered sparsely mid dorsally and anteriorly but is absent ventrally, Iridescent-white dendritic flecks seen under incident lighting are present over the whole dorso-lateral surface apart from two unpig- mented, slightly depressed arcas (eye fields) about 364.m in diameter situated anteriorly, Ag- gregutions of flecks mid-anteriorly, mid- posteriorly and laterally at the ‘waist’ form the points of » cruciform pattern, Both dorsal and ventral epidermis are 24m in thickness, The entire body surface is covered by cilia 5-6..m Jong. Sensory cilia 14-15um long are spaced 75-S5y.m apart. Underlying. the epidermis are the usual cireular, oblique and longitudinal musele layers, Dorse-ventral muscles are abundant, particularly laterally. Two eyes are present within the unpigmented anterior areas, Bach eye is oval, measures 19m antero-posteriorly and 344m medio-laterally, and is composed of minute reddish brown MEMOIRS OF THE QUEENSLAND MUSEUM granules, The cerebral ganglian is imsunk and bilobed. There is no statocyst-statolith, Algal symbionts (Prasinophyceae, ? Platymonas) measure 7-12j.m diameter and un- derlic the cutaneous body musculature dorsally and ventrally. Three types of rhabdoids are present. The most conspicuous ure sagitlocysts, refractile acicular bodies 18-20um long and about 2.5-3.6um diameter with central stylet 1j.m diameter, lying in the parenchyma at right angles to the surface with only the lips protruding from the epithelium. They are particularly numerous in batteries anteriorly, and toa lesser extent dorso- ventrally and laterally. They are sparse posterior- ly. Ventrally they surround the male pore, Histochemically the sagittocysts are strongly basiphilic and periodic acid-Schiff positive. Sagitlocysloblasts are localed deeper within the body and ultrastructurally comprise a lateral nucleus and thin granular cytoplasm surrounding, the sugittocyst. The sugittocyst consists of a fibrillar, concentrically lumellate cortex about 3.5-44.m diameter surrounding a central stylet. The stylet is a lym diameter membrane bound tube. The wall is approximately 300nm thick and composed of short fibrils, with an inner thin electron dense band. In the central lumen is an cccentrically situated electron dense rod about 140nm diameter, The conspicuous reddish-orange epidermal pigment noted macroscopically is due to orange- coloured fusiform and bacilliform rhabdoids 4,5- 6.3m long and 0.94m diameter, contained within packets 7-364m diameter and 24-60j.m long, lying at a shallow angle to the surface of the epithelium, Fusiform envelopes measuring 8-1] 8j.m long and 27.7- 3.6,.m diameter filled with numerous aciculur, refractiie microrhabdoids 2.7-3.6m long and Q,25-1,8jm diameter, lie in the epithelium at a shallow angle or parallel to the surface, Within (he dersal and ventral epidermis are gland cells containing translucent cytoplasmic granules which give a positive alcian blue his- tochemical reaction for acid mucopolysac- charides. These glands are particularly numerous anteriorly. A distinct cluster of strongly basiphilic glands form a frontal gland complex und open into # antero-ventral pil, Parallel with the dorsal surface is a network of concrement cells which are the iridescent white flecks ob- served macroscopically. The mouth opens ventrally and is situated in TROPICAL ACOELS 791 FIG. 3. Convolutriloba hastifera. (a) dorsal aspect, living specimen by incident light, and (b) fixed specimen (SEM). Note depressed eye-fields (arrowed) (c-f) epidermal structures: (c) anterior sagittocyst (d) sagiltocys- toblast (e) pigmented rhabdoids and concrement cell (f) packet of acicular rhabdoids. 792 MEMOIRS OF THE QUEENSLAND MUSEUM WwAQOS eS eae, Gas fo—=—., UT ee ¥ © SD) Fic, 4. Convolutriloba hastifera. (a) ventral aspect, whole mount (for clarity the ovary is not shown on right side) (b) longitudinal section (c) frontal organ and pit (unscaled sketch) (d) male copulatory apparatus (e) female copulatory apparatus, plan view (unscaled sketch) (f) female copulatory apparatus. TROPICAL ACOELS the anterior half of the body. Ingested amphipods were observed in the endocytium of live specimens, The testes are paired and are distributed dorso- laterally. Sperm pass posteriorly, accumulate in false seminal vesicles then continue into the male copulatory organ. This organ consists of an un- ciliated chamber filled with sperm. Basiphilic granular secretions form a plug in the genital pore, and are derived from prostatic glands equatorially surrounding the male organ. The male gonopore is ventral and ringed by sagit- tocysts. The ovarics are paired and lateral. They are divided into oocyte producing and yolk produc- ing parts. The oocyte-producing portion is ventrad and extends posteriorly 0.5mm from the anterior tip to just behind the female genital pore. The yolk-producing part is dorsad and begins just behind the mouth and terminates posterior to the oocyte portion. The female genital pore is ventral ap- proximately midway between the male pore and mouth. A short, ciliated female antrum leads into the bursal complex. The bursa is elongate, thin walled and filled with basiphilic granules. It communicates with the proximal end of a single anteriorly-curved sclerotised bursal canal. SYSTEMATIC DISCUSSION Hendelberg and Akesson (1988) noted that the male copulatory organ in Convolutriloba is a different type from that of Convoluta, but con- sidered the possibility that in their asexually reproducing specimens the male organ was not fully mature. For this reason they expressed some uncertainty as to whether the genus should be assigned to the Convolutidae or Haplopos- thiidae. The male copulatory organs of C. retrogemma and mature C. hastifera are similar. This suggests that the male organ described from the former specics was mature. Both specics lack a penis. Convolutriloba is therefore confidently assigned to the Haploposthiidae, Specimens of Convolutriloba from Tridacna at the University of New South Wales were non- sexual and unable to be identified fully. The specimens arc not C. Aastifera as anterior sagit- tocysts are absent. Examples of C. hastifera the same size as the Sydney material are found to have antcrior sagittocysts. The specics is similar to C. retrogemma. Convolutriloba hastifera was first observed in 1976 by Dr John Collins of James Cook Univer- 793 sity (JCU) on the rocky part of the base of Goniastrea sp. and Platygyra sp. corals collected from the intertidal zone Geoffrey Bay, Magnetic Island, They were also present in 1976 in the aquarium at JCU, and in March 1981 collected by Dr Clive Wilkinson from the aquarium at AIMS The characters which clearly distinguish C. hastifera from C. retrogemma are the presence of batteries of sagittocystoblasts and sagittocysts anteriorly, presence of cyes and an ovary with oocyte-producing and yolk-producing parts. The dorsal pattern of concrement in C. hastifera dif- fers from that of C. retrogemma (Hendelberg, in litt.). Convoluta japonica Kat6, 1951, (and Kawakatsu, 1983) considered by Dérjes (1968) to be a species incertae sedis, is now assigned to Convolutriloba. Convolutriloba japonica (Katé, 1951) comb. nov. is characterised within the genus by the presence of a statocyst, a rounded posterior with disc-shaped terminal adhesive gland, and absence of lateral caudal lappets. Other features accord with the generic diagnosis. No separate diagnosis was provided by Hen- delberg and Akesson, 1988 for the genus Con- volutriloba, the generic characters being those of C. retrogemma. As other species have now been referred to Convolutriloba it is appropriate to revise and emend the generic diagnosis. Unfor- tunately the presence or absence of caudal lap- pets is not a reliable generic character in the acocla. Asexual reproduction with reversed polarity has so far only been observed in C. retrogemma, and is therefore not included in the generic diagnosis. In both C. hastifera and C. japonica the paired ovary is divided into oocyte-producing and yolk- producing parts. At present it is uncertain whether the ovary in C. retrogemma is divided into the two parts as in the other species, In C. retrogemma, Hendelberg and Akesson (1988) describe a germ layer in which some cells are considered to be developing oocytes. Other cells present may be yolk-producing. Confirmation of a bipartite ovary in this species must await ex- amination of sexually reproducing specimens. From the combination of characters which now define Convolutriloba, the presence of a paired ovary divided into oocyte-producing and yolk- producing parts distinguishes the genus from all others in the Haploposthiidac. Otherwise this genus is closest to Pseudohaplogonaria Darjes, 1968, which also has a single bursal canal. Although sagittocysts are present in several 704 species of acocls they have only received scant attention at the light microscopic level (see Graff, 1905; Ivanov, 1952; and Marcus, 1959). In structure and mode of secretion, but notin size ot chemical composition, sagittocysts share many features of a discharge-type rhabdite, defined by Smith, Tyler, Thomas and Rieger (1982). According to these authors the Acoela seem to lack true rhabdites. Further altrastruc- lural investigations are required to ascertain whether there is any homology between acoel sagittocysts and the true rhabdites of other tur- bellarian orders, Convolutidae Graff, 1904 Heterochaerus Haswell, 1905 (emend) Heterachaeruy Haswell, 1905 p.425 : Bresslau, 1933 p.264, Amphiseolops Luther, 1912 p52; beauchamp, 1961 p.186, DIAGNOSIS Convolutidac oblong in shape, with caudal lappets; colouration and pattern due to symbiotic algae and concrement granules; rhabdoids, statocyst and statolith may be present; with eyes: brain insunk; frontal gland absent; mouth ventral in mid-body; ovary und testis paired; male geni- tal pore ventral in posterior third of body; mus- cular penis surrounded by seminal vesicle: female genital pore anterior to male pore; female antrum ciliated; bursa T-shaped and bilobed, each lobe with two or more tubular bursa! canals; in mature specimens a copulatory canal (ct. Laurcr’s canal) may be present dorsally opening into female antrum. TYPE SPECIES Heterochaerus australix Haswell, 90S. Heterochaecus australis Haswell, 1905 (Fig. 5a-c) Heterochaerus australis Haswell, 1905 p.425. Amphiscolops australis (Haswell); Luther, 1912 p.52; Marcus, 1950 p.21; Dirjes and Young, 1973 p,350. non Amphisculups sp. (australis?) Trench and Win- sor, 1987, MATERIAL EXAMINED Syntyres; AM W387 Port Jackson , New South Wales, Australia, found in shallow intertidal rock pools, Approximately 33 Carmine stained serial sec- MEMOIRS OF THE QUEENSLAND MUSEUM tions of a group of 17 specimens on a single micros- lide. The fifteenth section is marked with ink. Voucner SPECIMENS: Collected at Pioneer Bay, Or- pheus Island 18° 37'S, 146°30°E, sexual, from sedi- menis, collected J. Gray, 29 May, L984. G23009 wholemount, picro-carmine. G23010 three micras- lides of sernjal horizontal 7um sections, iron haematos vlin-cosin, OTHER SPECIMENS EXAMINED: Queensland: Saltwater Creek 19°05"S. 146°25°E, nonsexual, {rom the surface of stones, subtidal zone in the creek, collected I. Kneipp, 28 May. 1986; Shelly Bay near Cape Pallarenda, Townsville 19° |S, 146°45'E, non-sexual, from subtidal sediments, collected L. Winsor, 9 August, 1987. DESCRIPTION Specimens ate spatulate in shape with two clangate Jatero-caudal appendages, translucent brownish-yellow in colour with a clear margin, |-2mm by 0.5-0.75mm. The dorsal surface ex- hibits iridescence under incident light. A Stalocyst and stalolith are present lying midway between two minute eyes. Brain bilobed and insunk. A frontal gland ts absent. Algal sym- bionts, possibly an Amphidinium sp.. measure 10-14y.m in diameter, Testes und ovary are paired. Female genital pore is ventrally Situated |80j.m posterior to the mouth and §0Qj.m anterior to the male genital pore. The bursa is bilobed and T-shaped with lobes present as bulbous extremities on the transverse arm. Each lobe contains 2-3 bursal canals. Sperm accumulate in a seminal vesicle which surrounds a conical muscular penis. SYSTEMATIC DISCUSSION The copulatory organs and other features of the Queensland specimens agree closcly with Haswell’s (1905) clear description, and com- parison with the syntypes. There are minor dif- ferences in length of the caudal appendages of the specimens examined. Those from Orpheus Island have elongate appendages whereas in specimens from Saltwater Creek and Shelly beach the caudal appendages are short and rounded. In some specimens from both sites the caudal appendages are unequal in length, possib- ly due to accidental amputation of the normally long delicate structures. Variation in the number of barsal canals in cach of the nwo lobes. of the bursae of present material is within the range reported for the species. The presence of Laurer’s canal could not be confirmed in preparations of the new material, TROPICAL ACOELS Heterachaerus australis is characterised by the presence of multiple bursal canals in each lobe of a T-shaped bilobed bursa, rhabdoids, statocyst-statolith, and in mature specimens what may be a copulatory duct between the female antrum and the dorsal surface, The species lacks a frontal organ, and this was the principal reason Haswell (1905) considered australis was precluded from the genus Amphiscalops, all members of which had this structure. He there- fore erected the genus Heterochaerus to accom- modate this species. However ina footnote Luther (1912 p53), con- sidering only external features and multiplicity of bursal canals, synonymised Helerochaerus within Amphiscolops. Luther's action was not disputed in the literature until Bresslau (1933) without comment listed Heterochaerus in the Convolutidac, separate from Amphiscolops. Reinstatement of the genus was noted by Marcus (1947) who nevertheless included australis in a key to the species of Amphiscolops (see Marcus 1950) thus tacitly agreeing with Luther (1912). In his comprehensive treatment of the Acocla Dorjes (1968) apparently overlooked A, australis 795 as if was not included in a listing of Amphis- calops species. However he obviously accepts the validity of the species as Dérjes and Young (1973) reported the occurrence of Amphiscolops australis in Kenya, To evaluate the systematic position of A. australis it 18 necessary to examine the taxonomic characters of all species in the genus Amphiscalops (Table 1), As it presently stands this genus is clearly heterogeneous. Westblad (1946) drew attention to the different types of bursal canals in Amphiscolaps and with respect lo these structures (Westhlad, 1946, 1948) remarked on the heterogeneity of the genus. Later Stcinbock (1955) commented that the ab- sence of a stalacyst in four (now seven) Amphis- colops species might possibly justify the erection of a new genus. such as “Alithicum’ to accom- modate them. The current diagnosis for Amphiscolops is as follows (after Dérjes, 1968): Coloured through pigment or symbiotic algae; statocyst may be absent; brain insunk; frontal gland weak; mouth ventral; rhabdoids may be absent: posterior as a rule with two pronounced wdoos Fic. 5. Heterachaerus australis. (a) dorsal aspect, living specimen and (b) with elongate caudal lappets (Orpheus Island specimens) (c) Ventral aspect wholemount (for clarily the ovary is not shown on right side). 796 SPECIES MEMOIRS OF THE QUEENSLAND MUSEUM AUTHORITY MORPHOLOGICAL FEATURES 7 8 9 10111213 1415 16 17 is 19 GROUP 1 cinereus, TYPE (Graff, 1874) ;1905 5000 R O + + + + + + 12 BLL 2 + s fulgineus, Peebles, 1915 3000 R O + + + + 2? + Ll 1 BLL 2 + s Reidl, 1956 ? 600 R O + 2? + 2? 2? 2? 11 Sn Cc 2 + ? 2 daponicus, Katd, 1947 mM 6000 F 0 0 + + + ? O 1 1 SPL 4 + oT s evel inacy Marcus, 1947 N 2000 F O # + 0 + QO ? A gemet Liperus Marcus, 1954 x 600 F O + + + 0 ? 0 2 2 v GROUP 2 bermudensis, Hyman, 1939 mM 2000 T + ? + # + + # Z2 1 SL LB 2 + 7? § sp. 2. Yamasu & Okazuki, 1987 M R500. PO + Seo Boece 2 + dy Be Le ? v Ss sp. 3. Yamasu & Okazuki, 1987 BOG. RE Cee ete Foe Td ar, Gy 2 Vv s is] GROUP 3 PS langerhanst (Graff, 1882); Hyman, 1937 4600 F + 0 + + + ? + 1 1 SM 38 + Ss fal earvalhot Marcus, 1952 2000 F + 7 + + + 0 + 1°21 =SM 2-6 + s sp. 4. Yamasu §& Okazuki, 1987 M 2500 B+. ? O + @ 97 1. 1 SMF 7, Vv s Oo GROUP 4 cf. langerhanst sp. L. sp. form castellonensis sp. form gerundensis Sp. GROUP 5 sp. 5.4 Yamasu & Okazuki, Yamasu & Okazuki, Steinbdck, 1955 Steinboéck, 1955 Ehlers & Dérjes, 1 Yamasu & Okazuki, 1987 1987 9793 1987 N 1200 N 1300 N 3000 N 4000 N 2000 M 5000 F + Fo4 Fo o+ ? “3 Oe 2 4 A Oo + + ? A Ook 27 A Oo + 7 2 A oO + 0 0 ? GROUP 6 tralts — (Haswell, 1905) Hyman, 1939; Marcu s, 1950 M 4000 J 3000 ia) AS TABLE 1. Morphological and other taxonomic characters of species of the genus Amphiscolops, together with a commentary on species groups within the genus. Group 6 has been transferred to Heterochaerus (this paper). TROPICAL, ACOELS 797 KEY TQ MORPHOLOGICAL FEATURES IN TABLE |. += present; O = absent; ? = not mentioned in original descripnion or uncertain. sexual maturily of specimens upon which description based. N = nonsexual, J = juvenile; M = malure- miximum length reported (in 2m). shape of posterior end. R = rounded: F = forked (bilobed); T = trilobed. algal symbiont. pigment, . frontal organ. . Statocyst-statolith. _cyes. 9, rhabdoids. 10, conerement cells. 11. number of female genital pores. 12, number of male genital pores. 13. type of seminal bursa. BL = bilobed with paired lateral bursal canals; SL = spheroidal with paired lateral bursal canals; SM = spheroidal with multiple central bursal canals; SP = spheroidal With paired ventral papillae each containing two bursal canals (japonicus); TM = T-shaped bursa with multiple lateral bursal canals. 14. type of bursa) canal, C = corkscrew shape (zeii); L = long tubular: P = papillate, large and small. 15. number of bursal canals; for taxa with bilaterally arranged bursal canals, half (his number are present on each side. 16, seminal vesicle ~ penis sac. 17, posterior position of mule genital pore. ‘T= terminal-sub terminal; V = ventral, 18. reproduction apparently by A = urchilomy or $ = sexually. 14, birth of young. O =oviparous; V = viviparous. Senay bun NOTES ON SPECIES a. cinereus, Paired vaginac open into female genital duct. b. fulgineus. Eyes with refractile granules, Speci¢s may be conspecific with cinereus. c. japonicus. Eyes with refractile granules, Adhesive sucker presen! between mouth and female genital pore, d. evelinue, Antero-dorsit! fossa present. e. bermudensis. Frontal organ pit present, Paired vaginae open ventrally, f_ sp. 5. Has Iwo species of algal symbionts (as does sp, 3). g. australis. Dorso-atrial duct (cf, Laurer’s canal) present in mature specimens, h. sargassi. Paired efferent female openings where ovarian stroma interrupts ventral epicytium. Seminal bursa in this species and in australis differ from the bilobed bursae in other species of Amphiscolaps. COMMENTS ON GROUPS GROUP 1 In all members of the group (which includes the type of the genus), algal symbionts are absent, the male genital pore is terminal in position, bursae have paired lateral tubular bursal canals, The taxonomic position of evelinae and gemilliporuy is uncertain (species incertae sedis) GROUP 2 Similar to Group |. Algal symbionts present, male genital pore ventral in position, bursae with paired lateral tubular bursal canals. The position of the penis in bermudensis is uncertain. Amphiscolaps would be fairly homogeneous if if was restricted to Groups I and 2, with appropriate emendment of the generic diagnosis. GROUP 3 A homogeneous group characterised by presence of algal symbionts, numerous papillate bursal canals in the bursa, and ventral position of the male genital pore. The structure of the bursal canals apparently differs (rom those structures in Groups | and 2. This group should be separated from Amphiscolups. GROUP 4 All the species In this group have been described from non-sexual specimens, They have been placed in Amphiscalops primarily because they have algal symbionts and caudal lappets (bilobed posterior) although the type of the genus lacks both these features. At present these species cannotbe reliably assigned even toa family level and are more appropriately regurded as species incertae sedis. GROUP 5 Although little data on this species is available, the large size, presence of (wo species of algal symbionts, multiple bursal canals and terminal male genital pore suggest it may belong to a new genus reported from the western Pacific (Winsor, pers. obs.). GROUP 6 This group is characterised by the presence of algal symbionts, ventral male genitalia pore, multiple lateral bursal canals and T-shaped bursae, A frontal organ is absent. This group has been transferred to the reinstated genus Heterochaerus (this paper). TR posterior lappets: ovarics and testes paired; two genital openings, or they may be manifold, semi- nal bursa with two or more curved, or corkscrew shaped sclherotised bursal canals; male copulatory organ complex conical, pyramidal or tubular, a male antrum and seminal vesicle may be absent. Thorough revision of the genus at present is hampered by insdequate data for many of the taxa. Also some of the characters used (o define the genus, in particular the frontal gland and the type, number and position of the bursal canals - key clements in the generic diagnosis - need to be re-evaluated. Ideally this should be under- taken at both light microscopic and ultrastruc- lural levels, However despile these problems Species groups within the genus can be recog- nised (Table | - comments on groups) thus in- dicating directions for further revision. OF the Amphiscolops species for which ade- quate data are available, those comprising Group 6~A. australis and A, sargassi Hyman, 1939—are the only members of the genus in which a frontal organ is absent. Both species have similar mor- phology, in particular the presence ola T-shaped bilobed bursa, cach lobe of which contains lwo or more bursal canals of the same type. As both species exhibit ua combination of characters clearly different from Amphiscalops Group 1, which includes the type of the genus A, cinereus, they should be removed from Amp/tiscolops, Luther's (1912) synonymy of Heterochaerus within Amphiscalops is rejecled, Heterachaerus Haswell, 1905, is now reinstated and diagnosis emended. Both A. australis and A. serpasse dre transferred to this genus, Known only from non-sexual (Hyman, 1939) and immature (Marcus, 1950) material Heterochaerus sargassi (yman, 1939) comb, nov. nevertheless exhibits the essential charac- ters of Helerochaerus, Hyman (1939) indicated that in this species there ‘are no evident frontal glands nor frontal pit’ (her use of the term frontal glands referred specifically to the group of glands forming a frontal organ, rather than cutaneous glands which open from the anterior margin). Marcus (1950) corroborated these find- ings noling *As glandulas frontais desembocam, por via de regra, separadamente no boride anterior, exceptionalmente coalescem alguns dutos efcrentes delas, simulando um orgao fron- tal. Fosscta frontal nav ocorre.” Deseribed are cutancous glands which normally open from the MEMMnIRS OF THE QUEENSI.AND MUSEIIM anterior margin; occasionally the ducts of some of these wlands coalesce, simulating a frontal organ, The statement by Ehlers and Dérjes (1979, p14) thal ‘Marcus... beschreibt zwar cin Frontalorgan fiir A. sargassi..” is therefore incor- rect. The type of bursa, and the type, number and position of bursal canals in H. sargassi are similar to those of H. australis. In A. sargasstthe ovarial Stroma interrupts the ventral epithelium (Marcus, 1950), These structures may be efferent openings for the discharge of eggs. Haswell (1905) concluded that in H. australis eges were discharged through the female genital pore. However he drew attention to a blind anterior diverticulum in the female antrum which he con- sidered might be a vestige of a temporary passage formed to discharge eggs. Multiplicity of female ducts (vaginae) is known in Amphtscolups cinereus, A. fulgineus, A. bermudensts, A. gemel- liporus (Table 1) and in Wulguru cuspidata, At present it is not Known Whether these ducts are copulatory of efferent in function, and whether they are homologous. The absence of 4 statocyst-statolith in H. sar- gassi may be & consequence of reproduction by architomy, noted for other acoels (Hendelberg und Akesson, [98&). The Amphiscalaps sp. (australis ?) reported from the marine lakes in the Republic of Belau (Trench and Winsor, 1987, p.2) is now con- sidered to belong to a new genus to be described elsewhere (Winsor, pers. obs,). The Amphis- colops sp. 5 of Yamasu and Okazaki (1987) may possibly be referable to this genus. KEY TO FREE-LIVING ACOEL GENERA IN AUSTRALIAN WATERS The following artificial key is provided to facilitate recognition of free-living acoel genera presently deseribed from Australian waters. It is based upon external features evident in living and fixed specimens, Known distributions and habitats are provided, The key is intended to supplement Cannon’s (1986) invaluable guide to turbellarian famihes and genera, Full identifica- tion of acoel taxa requires microscopical and histological cxamination of sexual living and fixed specimens. |. Body discbid to obovate in shape, posterior margin rounded Or notched medially: occ Waminou (13mm diameter, found on hard and soft corals; north Queensland and western Pacific) TROPICAL ACOELS 1°. Body elongate, posterior pointed or with lateral Caudal lappets oo... sccseesesepeerseiessseensenenenereen 2 2, Posterior terminaling in single pointed lip, lateral caudal lappets absent... ieee Wuleure (1-3mm long found intertidally in sand and sediments; north Queensland) 2’. Lateral caudal lappels present, posterior with or without median JOBE woe o 3. With posterior median lobe .......... Convalutriloba (3-4mm long found in marine aquaria Townsville, Sydney and Gothenburg, Sweden; on corals in north Queensland waters; one species without caudal lobes found m Japan) 3°. Posterior median lobe absent .......... Heterochaerus (1-4mm long, found in rock pools Port Jackson, New South Wales; in subtidal sand north Queensland; in shallow pools.on a sandy beach, Mombasa, Kenya) ACKNOWLEDGEMENTS The following people kindly obtained specimens on my behalf, provided access. to specimens in their care, or generously provided additional information on yarious species: Perry Alino, Gordon Bull, Prof. J. Gray, lan Kneipp and Phil Osmond of JCU: Dr Lester Cannon, Queensland Museum, Brisbane; Dr Sneed Col- lard, University of Western Florida, Pensacola; Louise Goggin, University of Queensland, Bris- bane; Dr Jan Hendelberg, University of Giteborg, Géteborg; Dr Pat Hutchings, Australian Muscum, Sydney; Prof. Bob Trench, University of California, Santa Barbara, and Dr Clive Wilkinson, AIMS. The electron microscopy was undertaken by my wife Heather, Prof. D. Griffiths helped with identification of algal symbionts, Sally Jeston tracked down obscure literature and Dr John Collins provided enthusiastic help and advice in many ways. To all [| am particularly grateful. CSIRO Science and Industry Endowment Fund generously pravided some of the equipment used. LITERATURE CITED BEAUCHAMP, P. DE. 1961. Classe des Turbellairés. /n Grassé, P-P., (ed.), Traite de Zoalogte. IV: 35- 212. (Masson e} Cie: Paris). BRESSLAU, E, [933. Turbellaria, pp. 52-320. Jn 799 Kukenthal, W. and Krumbach, T, (eds), ‘Hand- buch der Zoologie.” (Walter de Gruyter & Co: Berlin and Leipzig). Cannon, L.R.G, 1986, *Turbellaria of the World. A guide to families and genera,’ (Queensland Museum: South Brisbane). 136 pp. Doryes, J. 1968. Die Acoela (Turbellaria) der deutschen Nordseekliste Und ¢in neues System der Ordnung. Z. zoal. Syst. EvalFarsch. 6; 56-452. DORJES, J. AND YOUNG, J.O. 1973. 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Zool. 36: 321-327. HYMAN, L.H. 1937. Reproductive system and copula- tion in Amphiscolops langerhansi (Turbellaria Acoela). Biol. Bull. 72: 319-326, 1939, Acoel and polyclad Turbellaria from Ber- muda and the Sargassum. Bull. Bingham aceanogr. Coll. 7: 1-26. Ivanov, A.V, 1952. [Turbellaria Acoela from the Southern Coast of Sakhalin) Travdast. Zool. Acad. Set, URSS 12.; 40-132, 85 figs. [In Rus- sian] Karo, K. 1947, A new species of the Conyolutidae (Acoels, Turbellaria). Seibursu Suppl. No.1: 58- él, 1951. Convoluta, an acoelaus turbellarian, destroyed the edible clam. Mise. Rep. Inst. Nat. Resoure., Tokyo 19-21: 64-67. KAWAKATSU, M.R. 1983, Commentaries on two Japanese papers of the late Dr. K. Kalé describ- ing new Species of acoelous turbellarians, with their republication, Occ. Publ, Biol. Lab. Fuji Women's College. 10; 1-16, 800 LUTHER, A. 1912. Studien uber acoele Turbellarien aus dem Finnischen Meerbusen. Acta Soc. Fauna Flora fenn. 36: 1-59. Marcus, E. 1947. 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MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum ‘CONTENTS ALLSopP, P.G. F Anoplognathus hilleri sp. nov. (Coleoptera: Scarabaeidae: Rutelinae) from southeastern Queensland and notes:anmA. lindersensig Came «4.0. 25 bn eh ee i ke ee 377 BAEHR, M. Two new species of Trichotichnus Morawitz from north Queensland (Coleoptera: Carabidae: Harpolinacye a5 jesse eae seep te ease. Re oenieeed Sit b A ens ru re de eee eecte? as 383 Bruce, N.L. Hysterothylacium Ward and Magath, 1917, and Ichthyascaris Wu, 1949, ascaridoid nematodes from Auusttalinndemersaltrsbes 0504.4 49 ies Seis pegs ee th ee ea 4 eG 389 Bruce, N.L. ; Redescription of the ascaridoid nematode Hysterothylacium scomberomori (Yamaguti) from Australian spanish mackerel Scomberomorus commerson (Lacepéde) ............-. 427 CANNON, L.R.G. Apidioplana apluda n, sp., a turbellarian symbiote of gorgonian corals from the Great Barrier Reef, with a review of the family Apidioplanidae (Polycladida: Acotylea) .............-. 435 Davin B., DAVID, M., FLOOD, J. AND FROST, R. Rock paintings of the Yingalarri Region: preliminary results and implications for an archaeology of inter-regional relations in nothern Australia... 1... 2 epee ee ee ee es 443 DAVIE, P.J.F. New and rare crabs of the subfamily Dotillinae (Crustacea: Ocypodidae) from northern Australia and Papua Mew Guied 4 jp Se bes = Sect pe ee ee pe Ee be 8g 463 INGRAM, G. AND CORBEN, C. Litoria electrica: a new treefrog from western Queensland ... 2... 2 ec ee eee eee es 475 KLEINSCHMIDT, K. “Yes, I well remember that Mrs W.C.C. Wright....’ The Elsie Wright Needlework Collection . . . . 479 KOHOUT, R.J. A review of the Polyrhachis viehmeyeri species-group (Hymenoptera: Formicidae: Formicinae) . . 499 KOHOUT, R.J. AND TAYLOR, R.W. Notes on Australian ants of the genus Polyrhachis Fr. Smith, with a synonymic list of the species (Hymenoptera: Formicidae: Formicinae)... 2.2... 6 eee ee ee ee ee ee 509 RICHER DE FORGES, B. AND GUINOT, D. A new Cyrtomaia, C. griffini, from Australia (Crustacea: Decapoda: Brachyura) ..........- 523 RoTn, L.M. A revision of the Australian Parcoblattini (Blattaria: Blattellidae: Blattellinae) ........... 531 Roth, L.M. Revisionary studies on Blattellidae (Blattaria) from the Indo-Australian Region ..........-. 597 CORRIGENDUM RICHARD A.I. DREW. The tropical fruit flies (Diptera: Tephritidae: Dacinae) of the Australasian and Oceanian regions. Memoirs of the.Queensland Museum 26: 1-521 .......-...+5-+65 664 ROZEFELDS, A.C., MCKENZIE, E.D. AND MOBBS, C. Type, figured and mentioned fossil invertebrates in the Queensland Museum ............ 665 SMITH, A.B. AND JELL, P.A. Cambrian edrioasteroids from Australia and the origin of starfishes .........+25+200% 715 TYLER, M.J. Limnodynastes Fitzinger (Anura: Leptodactylidae) from the Cainozoic of Queensland ......-. 7719 WInsoR, L. Marine Turbellaria (Acoela) from sacred Chueetistaed oo 7 a8 8 py Red bal be de ehere ents 785