RECORDS OF THE SOUTH AUSTRALIAN MUSEUM VOLUME 21 PART1. MAY 1987 CONTENTS: EDMONDS, §. J. Obituary of I. M. Thomas EDMONDS, 8. J. Echiurans from Australia (Echiura) HERCUS, L. A. Looking for Ditji-mingka JAGO, J. B. & PLEDGE, N. S. Obituary of B. Daily KERZHNER, I. M. Nabidae (Heteroptera) of Vanuatu LEE, D. C. Introductory study of advanced oribate mites (Acarida: Cryptostigmata: Planofissurae) and a redescription of the only valid species of Constrictobates (Oripodoidea) PATTERSON, C. & RICH, P. V. The fossil history of the emus, Dromaius (Aves: Dromaiinae) RILEY, J. & SPRATT, D. M. Further observations on pentastomids (Arthropoda) parasitic in Australian reptiles and mammals SUTTON, P. From horizontal to perpendicular: two recent books on central Australian Aboriginal painting TINDALE, N. B. Kariara views on some rock engravings at Port Hedland, Western Australia TRIGGER, D. S. Inland, coast and islands; traditional Aboriginal society and material culture in a region of the southern Gulf of Carpentaria WATTS, C. H. S. Revision of Australian Berosus Leach (Coleoptera: Hydrophilidae) ZEIDLER, W. The scaled-squid, Lepidoteuthis grimaldii Joubin, from southern Australian waters Volume 21(1) was published on 22 July 1987. Volume 21(2) was published on 24 December 1987. ISSN 0081-2676 PAGES 61-63 119-138 149-156 65-68 29-33 35-42 85-117 139-147 161-165 43-59 69-84 1-28 157-159 REVISION OF AUSTRALIAN BEROSUS LEACH (COLEOPTERA : HYDROPHILIDAE) BY H. S. WATTS Summary The Australian members of the hydrophilid genus Berosus are revised and redescribed. A key to species is give. Thirty-two species are recognised of which 18 are described as new: B. amoenus, B. arcus, B. aquilo, B. dallasae, B. gibbae, B. josephenae, B. juxta discolor, B. reardoni, B. macropunctatus, B. sadieae, B. nicholasi, B. niger, B. quadrapunctatus, B. trishae, B. timmsi, B. veronicae, B. ralphi and B. vijae. REVISION OF AUSTRALIAN BEROSUS LEACH (COLEOPTERA: HYDROPHILIDAE) ©. 1.5, WATTS WATTS, ©. HS. 1987, Revision of Australian Berosus Leach (Coledptera: Hydrophilidae). Ree: Se lua Mis. 211) 1-28. The Australian members of (he hydrophilid genus Berosyvs are revised and redeseribed. A key [Oo splecies is 2iven. Thirly-fWe species are recognised of which 18 are descrihed as new: 2 umoonus, Burens, B aquily, B dallasae, B. gibhae, B. josephenae, Bo justadiscolar, B. reardoni, B wnecropuncratus, B sacieae B. nicholas) B niger, B. quadrupuncralns, B trishae, B tiniest, B, vervnicae, B, ralphi and B. yijee, The following synonymies are proposed: B /lindersi Blackburn B. sinnlans Blackburn — B&. stigitatico/lis Fatrmaine: = B. auriveps B. approximans Pairmaire, Bo quartinus d'Orchmenr - B. queenslandicus Blackbur; B wrayis Blackburn B ovipentis kainmaire Wlackburn = & blackhurnt Zain B. discolor Blackburn: B wustratige Mutsarit. Co bLOS. Waus, Sourh Australian Museum, North Terrace, Adelaide. Soul Australia S000 Manuseripr received 24 July J98s. The hydrophilid fauna of Ausiralia js relatively rich in species and numbers of individuals. They form uw conspicuous part of tle aquatic fauna. In general they are well-colleeted and well-represented in vollections, Despite this they have received litle recent taxonomic attention. A major difficully in dealing with Berasus Leach, IS? in Australia was the inacequacies af existing keys which reflected a weak taxonomic base exens- plified by the facet that some [8 of the 32 known species were undesenbed, including the commonest southern species. At the same time some widespread species were deseribed under as many as five different names. The last biajor atlemipt jo revise Australian Berosus was by Blackburn in 1898, Some characters used in the key, such as leg colour and details of Lhe punctation, are nat totally reliable, but the male genitalia have provided good characters i all cases. The genus Berasus is a member of |he subfamily Berosinae which is characterized by a markedly deflexed head, seven- or cighteseemented antennae and meso- and metasternae without a continuous common keel, Berosus can be separated trom the other Australian genera in the subfamily (Regéinbartia, Gloharia and Amphiopsy by possessian af protruding eyes, seven-segmented antennae, and striae on elytra, although in some species these are partially masked by strong elytral punctations. Adults of all bur a few al the smaller species are yellow-brown in colour They are found mainly among vegetaiion at the boron and sides of ponds and pools. Adulbs are often taken at light sometimes some distance from water. The larvae are distinetive witha series of lateral filamentous appendages ard live in pools in similar situations to the adules. | know of no detailed description of the larvae of any Australian species, Berosus is a large cosmopolitan genus and many attempts have been made to subdivide it, none of which appear particularly satisfactory to me. In particular, the subgenus /¥ygrorrophus W. Macleay which includes only the species & (4.) nutans and B (H.) devisi, both Australian species, seems unwarranted and to be based on an undue emphiasts on the punetation of the upper surface. Within Australian species there are five more or less distinctive groupins. Group | species are small, have the midline of the mesosternum produced downwards in a distinet ridge, head black, metacoxal process weakly to moderately lobed laterally, last abdominal seement notched and the second elytra! stria 1/4-the length of the elytron (& discolor, B. approximans, B, reardoni, B. juxtadiscolor, B. limmsi). Group 2 species are small, with the midline of Ihe mesosternum produced downwards in a distinct pillac, head black, metacoxal process strongly lobed laterally, the last abdominal segment notched, the second elytral stria about 1/2 the length of elytron, and with strongly developed medial ridge on first abdominal segment (& macropunctatus, 8 quadrapunctatus, 8. trishae), Group 3 species are similar to Group 2 specics except for a less developed medial ridge on first abdominal! segment, strongly humpbacked which is associated with the presence of a space between elytral striae 7 and 8 and/or 8 and 9 in the middle of the elytron (A involutus. AL niger, & areus). Group 4 species are similar to Ciroup 2 but are te larger and have the second clytral sta reaching about two-thirds the lenpih of the eélytran (B duplopunctatus and B. queenslandicus). Group 5 species are generally larger with brown heads, midline of mesosiernum with a weak central keel, und metacoxal process withaul lateral lobes (BO nutans, B, pulchellus, B dallasi, 8B. macumbensis, B. imwittipennis, B. anoenus, 8, josephenae, & gibbae, B. majusculis, B. veronicae, B. aquila, B. australiae, B, decipiens, B sadieae, B nicholasi, B. vijae, B. subovatus, B. ralphi and B. debilipennis), Groups 1-4 have similarities with cach other and roughly correspond to the subgenus Berosus Hope. Group 5 species correspond to the subgenera Lnoplurus Hope and AHyzrotrophus W. MacLeay. Since Berosus is a world-wide genus, Surther consideration of subgeneric groupings should in- clude extra-Australian species and is beyond the scope ol this revision. The collections from whieh specimens were examined are listed under the following abbreviations: AM Australian Museum, Sydney, ANIC Australian National Inseet Collection, Canberra, BM(NH) &ritish Museuni (Natural History). London, OW Private collection of author. NMV National Museutn of Vivtoria, Melbourne, NSW DA New South Wales Department of Agriculture, Sydney. NTM Northern Territory Museum, Darwin, MNHN Museuni National d'Histoire Nalurelle, Paris. SAM South Australian Museum, Adelaide, WAM Wesiern Australian Museum, Perth. OP! Queensland Department of Primary Industry, Mareeba. KEY TO AUSTRALIAN SPECIES OF BEROSUS 1, Midhine of mesosternum produced downwards in a distinct pillar (Fig. 10), Aor projecting backwards heryeen metavoxue. Mead black imetallic), Metacoxal process lobed laterally (rudimentary in &. discolor). Last abdominal segment notched, Small (< 7mm). --.. ..2 | Midhine of mesosternum with weak central keel, projecting weakly forward between mesoconac Head completely or partially brown except B pulchellus whieh lacks elytal stfiae and B amoenus whieh has a predominantly black Pronolum and elytra also. Melacaxal process irvangular (Pig. 1) 22... + + ay, 2,(1) Second elytral soja about 4 feneth ‘of elvinontd 2, Second elylral stra 1/2 length of ebyiron ©. WS. WATTS 3, 42) 3. 4. (3) 4, . (4) §, b. (2) 6, 7, (5) 7. B. 16) R. 9. (4) 9. 10. (9) 10, 1) (Ry i. Rugose portion at metalentur <1/2 teneth of fomutr (Fig. 2), Mesosternul pillar flat.on ventral COG evens | tppceee & discolor Blackburn Rugose portion ol metalenmuy S12 lengil of fomtur. Mesosiernal pillar weakly concave of veritral edge rete:: | Riuizose portion of inesofernuir 12 2-2/3 length of fa) Ue ee er, awe | Rugose portion of mesofemur <1 /2 length ot femur . ot B approximans bairmane Rugose portion of (feta: and meso-lemora pale, same colour as rest of leg. B reardoni spnov. Rugose portions of meta- and meso-femora much darker |Man vest of leg 2.0, 2.2... 7 Second elyiral sija about 1/2 length of vlytiron, fugose portion of metafemur S1/2 length ot Femina ge sire ft iigaees. 9 cake, a) Second elytral seria aul least 2/3 lenerh af elytron, rigose poruion of metafemur <1 /2 lengih of PRAMS s SoS ae Oo =~ -e TF Punclures on sides af pronorum never in size, Interstvial pupclures small, subobsolete, hind lemurof male with triungilar bulge an hind edge we Banta’ sp.nev Panatiiver on 1 sittes of prenewum evenly sized with ouly a few smaller anes, inferstrial punctures moderately marked, hind femur of male normal ctr teers e ees B ytvtadiseoler sp.nov Space be} ween ely(ral striae 7 and 8 and/or 8 and 9 prealer in middle than at ends of elytron (Fig. 8), Medial ridge on firsr abdominal seyment 1/4 as deep as long, wilhoul backward projection on ventral edge Elytral striae 7, 8 and ¥ subparallel with each other, Medial ridge on fest abclontinal segment deep | 3-1/2 as deep-as long, wilh a backward projection on ventral edge -—,-.220,2--,,, 0 Elyrral siria 8 raised and cacinatein middle and towards apex. Distance between elyiral striae 8 and Y greatly enlarged i iniddle (Fig, 8), Interstrial punctures weak, subobsolete. Northern SEPRCIES Scare oicjciem O07 tan ae Dever aoe tll Elvtral stria 8 ot raised, ‘Interstria \ae 7 Ras Wide or wider than 8-9 (Fig, 9). Laterstrial punctures moderately strong, Southern species -- 46 B. invalutus Macleay Rowose por tion of metafemur approximately 1/2 length of femur. Panetures on dise of pranotum ol Ovo sizes, larger predominating. Posrerior partion of carinate eight) imtersta at right angles co edge of elytron. Black, except for interstria in apical 1/4 of elytron........... litt ty eeeet corer e B Riger spTDy, Ruwrense purhon of metafentur ‘tenehes 2/3 length of femur on rear edge, Punetures on diye ol! pronolum of abe size, posterior portion of catinate eighth imtersiria of approximately 45° to edge of elyiron (Fig. 8). Barcus sp.nov. Pronotum smooth, shiny, moderately covered With large puncitires often with spaces preater than 1/2 diameter of puncture or erealer between punctures. Pronoltum vellow with three longilidinal black Stripes... peace AD Head and proratiim rugase, densely. covered with large punctures, those an provotuny separated 15, (14) 18, 16, (15) 16, Vy, (14) 7, 8. (17) REVISION OF BEROSLIS by narrow nilges, Promotumt Black except far extreme lateral margins BR, IMACFOPUNCLALUS sp.Nov. } Rugose portions of mesofemur a little under 1/2 length of femur. . 8, quadropunctalus sp.m0y, Ruwose portions af mesofemur Ly 2-2/3 lengthy CU MANOR ese pe oe B. trishae sp.nov. Ponctures On prononiny af Iwo distinct sizes. Areas between elytral striae with shallow punctures... .8 daplopunctatus Blackburn Punctures on pronotum vary little in size. Areas belween elytral striae virtually impunctale .., B. queensiandieus Blackburn Punctures on elytron densely und evenly distributed, each with a short seta, Punctures in clyERAl striae ditfieull to distinguish froin (hose WM WMICPRIe wee Rete ne 15 Punctures on elytra mostly without setae. Phnetures if clytral ilersttiac seatrered, distines from those in striae Punetures on pronatuint expanded faterally forming a-dense network of closely spaced short striae orientated across the pronotum (Fig, 5) ; BL niteans W, Macleay Punciures ¢ on n prongtum MOTMA eee eee 16 Small (< 3.0 nod), elytvon without striae Head metallic black... . B. pulehellus W. Macleay Larger | >3.6%m), elytra wilh deeply impressed striae, Rear of head with dark markings ———. ; B. dallasi sp.nov. Interstrial purnctu ures towards sides of elyira not as mimcrous as as those on disc, olfen arranged TB SIMRO TOW ie ee ee ee eee 18 lorerstrial purtetures towards sides-al elytra mare numerous and larger (han (hose on disc, never arranged in single lines... . ~B macumbensis Blackburn Outer anical spine on elytron tong and (hin (Fig. 3) Interstrial punetures laterally subobsolete, arranged in one row aver most of elyiron. Rugese portion of meiafemur approni- mately 1/3-2/3 length of femur Pale ----- a Re bean Bo munitipennis Blackburn Nor with above combination of characters (Xi8; SV veep aecorresurat etogeo rere teres LS 19. (18) 19, of pronotum of one size (eg. Fig. 6)... 20, (19) Head black. 20, Head pale brown, 21, (20) 2), 22. (19) Rugose portions of mesc 22. 23. (22) stiria >22. Rupose portion of abdominal segment notched .,. 23 VEFONIEE vor yyy ss t fast 24, ” Punotures in jntersiria 3 of two sizes, with small size predominwing (Fig. 11), Punetures on dise of pronotum of two sizes mostly large with some smaller ones (ug. Fig. 7) 2.) 0-22.) 02... , 22 Punctures ininterstria 3 of one size, orwith only an occasionally smaller one. Punctures on dise 20 Interstrial punctures on elytron approximately the same size as those in striae Pi eb eee clavate mcm B. amoenus sp.nov. Inierstrial punetures on elytron smaller than (hose in striae... ,, 2) Rugose portions of femore dark-brown —.... 5 -B. Josephenue sp wov. Rupose Portion of femare pale-brown...,... . B. gibhde sp.nov Rugose portions of meso- and meta-lemore noticeably darker than rest of femursee Table |. Rugase portions of meso- and metafemore of same colour as rest of femur... oc... 23 larger (>3.9 mm). Number of punctures iw second ely(ral interstria lo level of end of second mesotemur approx. 1/3 length of lenitir. Male with apical iB. nemageulds Blackburn Sonaller (<6.5 mm). Number of punetures in second elyiral iniersiria back to level of end of sccond sifia usually <26. Rugase portion of inesofemur >1/2 length of femur except in B, NOC 24 Rugase portian of nievolerwur 2 1/2. Apical spines on elytra moderately developed (hig. 4) B veronicae sp.nov. Rugose purticrn of mesofemur >1/2 lengiti of femur ate Table 2. SYSTEMATICS Berosus discolor Blackburn (Figs 2, 13, 18) Berosus discolor Blackburn, !888(1889), p. 829 Berosus flindersi Blackburn, 1888(1889), p. 831, syn. Noy, TABLE L. Distinguishing characters for four species of Berasus, B. aquila sp. nov. B. australiae Mulsant B sadiae sp. nov. ; ilo sp. . I s & decipiens Blackburn j . 5 3.3-5,0 mm long Tips of elytron usually without spines. Third segment of male prolars) approximately gare length a5 segment, Elytral punctures weak, Strial punctures arranged in one line, over most of elviron. Coastal Norihern Territory, 6.5-9.0 mm long Tips of elytron with well developed spines. Third segment of male protarsi shorter than second segment. Elytral punctures usually strong, sirial punctures not artanged in one line inwards from. siria 7. Australia-wide. 6,1-8.0 mm long Tips of elytron with well- developed spines. Third segment of male length of second segment. Elytral pufierures usually weak, Strial punerures usually arranged in one line over most of elytron. Wyndham to Cairns. §-5.0 mm long Tips of elytron usually with well-developed spines. Third sewment of male length of second segment. Elytral punetures sually weak, strial punctiires usualy arranged th one line aver most of elytran. Coastal Northern Territory. ©. HS, WATTS TABLE 2. Distinguishing characters for five species oF Heresies, B, debilipennis Blaekburn 4.0-6.0 mm long Apex of elytron rounded or with weak spines. Punctures in Ist elytral interstria, scatiered, not arranged in a row. Lateral punctation on elytran moderale to STON. Tip of posiconul process Ii same plane as rest of underside, 10-18 punctures in second clyrral Strie. First segment of male protarsi much B. subovatus Knisch 3.0-3.0 iM lone Apex of elytron rounded of weakly truneated, Punctures in 1st elytral interstria, arranged ina Single row, Lateral punctation on elytron weak. Tip of posicoxal process in same pling as rest of underside. 6-7 punciures in second elyiral suria, First segment of male protarsi B. yiyae sp. nov. 3.0-4.5 mm long Apex of elytron with weak spines, Pufietures in Ist elytral interstria, arranged ina Single row, Lateral punctation on elytron moderate to strong. Tip of postvoxul process in same Plane as rest of underside. 8-13 punetures in second elytral stra, First segment of male prolarsi equal B. nicholasi sp. nov. 5.0-6.5 mm long Apex of élytron with weak to moddenale spines. Punetures in tor elyinal interstria, sealtered, not arranged in a row. Lateral puneciarion on elytron small and Weak. Tip of postcasal provess i Same plunge as rest of woiderside, LO<1R punchires in second elyiral stria. First sepment of male protarsi equal B, ralphi sp. noy. 3.5-4,.5 mm long Apex of elyiron truncated or with weak spines, Punetures in Ist elytral interseria, seattered, fol arranged ina single row, Lateral punclation on elytron small and weak, Tip of posicoxal process bent downwaruds av 45) to rest of Underside. Tel3 punctures in second ¢lytral Atria. First segment of male provars) longer than second. longer than second. second, in lenpth to longer than second. in length to second. 55 Thpes Berosus discolor: Holotype, male, Pr Lincoln, §.A., in BM(NH), seen. Berosus_/lindersi; Holotype, male, Pt Lincoln, S.A., in BM(NH), seen. Paralype, female, in SAM, seen. Descriplion (number examined 131) Length 2.8.8 mm. Oval, elytra moderately humped, highest behind middle where it is approximately 1/3 » height alshoulder: Apex of” ely(ron rounded, in some subavure and suggestive of a thick broad spine, Head relatively narrow, black, shiny, metallic, Pronotum black with sides broadly brown, often with small central brown spot or band behind front edge, shiny, metallic, Elytron shiny brown, with striae, punctures and a few spots black, Ventral surface black, appendages brown, except for extreme lips Of lapial palps and rugose porlion of meso- and metafemur which ean range from light-brown to black. Punctures small, dense. Punctures on head strong, regular, well-impressed, most less than 1/2 puncture width apart. Punctures on pronotum similar but slightly less dense. Elytral striae well-impressed. Second stria on elytron 1/4 length of elytron, Punctures in and between striae similar, about same size as on pronotum, strongest laterally, those on extreme humeral angles and belween first stria and suture smaller. Mesosternal keel weakly to moderately well-developed, pro- jecting backwards. Metacoxal process sharply produced backwards in niidline, lateral lobes raised, sides diverging towards lront, small narrowly oval shiny depression in midline. Keelin midline of first abdominal segment weak, reaching only |/4 to 1/3 of width. Rugose portion of metafemur approxr mately 1/3 length of feriiur. Meso- and meta-libia strongly spined. Apical abdominal seament with small weak noteh. Male: Protarsi foursegmented, first seament moderately expanded, Remarks Less. common in collections than B approximans. Readily separated from those other Berosus with a well-developed mesosternal keel and short second elyiral stria by the small area ol rugosity on the femora. Distribution (Fig. 79) Coastal southern Australia from Perth, W.A. ta Sydney, N.SW., and Tas, Berosus approximans Fairmaire (Pigs 10, 12, 17) Berosus approximans Fairmaire, 1879, p. $2. Berosus ovipennis Fairmaire, 1879, p82, syn. nev, wn REVISION OF BEROSUS She. Testa > BG) ost at %.5 $e lo 8 Gre Fe ess oy, FIGURES I-11. 1, Midventral region of B. nicholasi; 2, midventral region of B. discolor, 3, apex of elytra of B. munitipennis; 4, ditto, B. munitipennis; 5, pronotum of B. nutans; 6, pronotum of B gibbae showing punctation on disc; 7, pronotum of B arcus showing punctation on disc; 8, lateral yiew of elytron of B. arcus; 9, lateral view of elytron of A involutus; 10, lateral view of B. upproximans showing pronotal pillar (shaded); 11, elytron ot B. majusculus showing punctation of disc, Cc. H. S. WATTS | KY 18 16 ART See al — ie) 17 ie 22 3 : SHE? Ee 27 28 29 30 31 FIGURES 12-31. 12, dorsal view of male genitalia of B. approximans; 13, ditto, B. discolor; 14, ditto, B. juxtadiscolor; 15, ditto, B. reardoni; 16, ditto, B. timmsi; 17, lateral view of male genitalia of B. approximans; 18, ditto,B. discolor: 19, ditto, B. juxtadiscolor; 20, ditto, B. reardoni; 21, ditto, B. timmsi; 22, dorsal view of aedeagus and right paramere of B. duplopunctatus;, 23, ditto, B. queenslandicus; 24, dorsal view of aedeagus of B. arcus; 25, ditto, B. niger; 26, ditto, B. involutans;, 27, lateral view of aedeagus of B. duplopunctatus; 28, ditto, B. queenslandicus; 29, ditto B. arcus; 30, ditto, B. niger; 31, ditto, B involutans. 25 26 REVISION OF BEROSUS 7 yea a 38 39 40 = sega 0 ede - ot 46 = 52 Gao \S sae Ca 49 5 51 53 FIGURES 32-53. 32, dorsal view of mate genitaha of B dallasi; 33, ditto, B. gibbae; 34, ditto, B. aquilo; 35, ditto, B. vijue; 36, ditto, B. subovatus; 37, B. trishae; 38, lateral view of male genitalia of B. dallasae; 39, ditto, B. gibbae; 40, ditto, B. aquilo; 41, ditto, B. vijae; 42, ditto, B. subovatus; 43, ditto, B trishae; 44, dorsal view of male genitalia of B. debilipennis (thin form); 45, ditto, B. debilipennis (thick form); 46, ditto, B sadieae; 47, ditto, B. decipiens; 48, ditto, B. pulchellus; 49, ventral view of male genitalia of B. debilipennis (thin form); 50, ditto, B. sadieae; 51, B. debilipennis (thick form); 52, ditto, B. decipiens; 53, ditto, B, pulchellus, Q 0 C,H. S. 55 \ | 56 54 hy | | ) 60 61 Re 68 69 WATTS 58 57 59 ity 70 FIGURES 54-70. 54, dorsal view of male genitalia of B, majusculus; 55, ditto, B. josephenae; 56, ditto, B. amoenus: 57, ditto, B. nutans; 58, ditto, B. munitipennis; 59, ditto, B. macumbensis; 60, lateral view of male genitalia of B majusculus; 61, ditto, B. josephenae; 62, ditto, B. amoenus; 63, ditto, B. nutans; 64, ditto, B. munitipennis; 65, ditto, B, macumbensis; 66, dorsal view of male genitalia of B. nicholasi; 67, lateral view of male genitalia of B. veronicae; 68, dorsal, B. veronicae; 69, lateral, B. nicholasi; 70, lateral, B. macropunctatus. 72 FIGURES 71-74. 71, lateral view of male genitalia of B. australiae; 72, ditto, B. ralphi; 73, dorsal view of male genitalia 71 of B. ralphi; 74, ditto, B, australiae. 74 REVISION OF BEROSUS 9 Berosus stigmaticollis Fairmaire, 1879, p, 82, syn. nov. Berosus simulans Blackburn, 1888(1889), p. 832, syn. nov, Berosus auriceps Blackburn, 1889(1890), p. 447, syn. nov. Berosus blackburni Zaitz, 1908 p. 358, nom. nov. for B. auriceps. Types B. approximans. Holotype, male, Peak Downs, N-T. in MNHN, seen. B. stigmaticollis, Holotype, female, Peak Downs, N-T. in MNHN, seen. B. ovipennis, Holotype, female, Pt Mackay, in MNHN, seen. B. auriceps. Holotype, male, Northern Territory, in BM(NH), seen. B. simulans. Holotype, male, Rivoli Bay, S.A, in SAM, seen, Description (number examined 134) Length 3,0-5.2 mm. Narrowly oval, elytra moderately humped, higher behind middle where it is 1/3 higher than at humeral angle. Apex of elytron rounded or produced into a small, widely triangular spine. Head black, shiny, metallic. Pronotum brown with broad central panel darker, dark portion usually not reaching front or rear edges and often with lighter area in midline, shiny. Elytron brown, with striae, punctures and a few spots, black, shiny. Ventral surface black, append- ages lighter. Rugose portions of femora variable from light to dark-brown. Punctures on head strong, regular, well-impressed, most less than 1/2 puncture width apart. Punctures on pronotum simi- larly sized but less dense. Weaker on disc. Elytral striae well-impressed. Second stria on elytron 1/4 length of elytron. Punctures in striae and between them approximately the same size, as strong laterally as on disc. Reticulation of elytron variable, virtually absent in some, in others very strong and almost masking punctures, especially laterally. Ventral surface densely rugose punctate. Mesosternal keel strongly produced downwards forming a pillar. Front edge almost perpendicular to body, ventral edge straight, or concave in some specimens, with the front portion projecting downwards as much or further than rear portion. Lateral lobes of metacoxa raised, bluntly pointed, midline produced backwards in a point, small oval depression in midline devoid of sculpture. Keel in midline of first abdominal segment well-marked, reaching 1/4-3/4 segment. Rugose portion of metafemur 1/2-2/3 length of femur. Meso- and meta-tibiae strongly spined. Apical abdominal segment weakly and widely notched. Male: Protarsi four-segmented, first segment moderately expanded. Head narrower than in female, Remarks A southern species slightly larger than B. /lindersi. Readily separated from that species by the larger area of rugose sculpture on the femora. The notch on the last abdominal segment is also broader and weaker in B&B. approximans. Differs from B. juxtadiscolor and B. reardoni from northern Australia by having a smaller amount of rugose sculpture on the femora than in these species, in the generally strongly elytral punctures and in characters of the aedeagus. Its wide distribution, variability in size and in the strength of the dorsal punctation has led to it being described a number of times by earlier authors. Distribution (Fig. 76) Southern coastal Australia from Kimberley, W.A., to Rockhampton, Qld. Berosus reardoni sp. nov. (Figs 15, 20) Description (number examined 7) Length 3,2-4.5 mm. Narrowly oval, elytra moder- ately humpbacked, highest behind middle where it is 1.4 x height at shoulders. Apex of elytron bluntly pointed. Head shiny, black. Pronotum yellow-brown with two longitudinal black strips adjacent to midline, dark portions not reaching front or hind margin. Elytron yellow-brown with striae and many punctures on head strong, regular, well-impressed, most less than 1/2 a puncture-width apart. Punc- tures on pronotum moderately dense and strong laterally, weaker and sparser on disc. Dorsal surface weakly to moderately reticulate. Elytral striae well- impressed, less so on disc, second stria on elytron a little less than 1/4 length of elytron. Punctures in striae somewhat larger than those in interstriae, shallower and slightly larger laterally. Mesosternal pillar long, well-developed, ventral edge concave. Ventral surface densely rugose-punctate. Metacoxal process broad, lateral lobes lowered slightly, diverging towards rear, deeply and broadly pitted in centre, weakly triangularly produced backward in midline. Apical abdominal segment notched. Rugose portion of metafemur 2/3 length of femur, that of mesofemur between 1/2-2/3, that of profemur about 1/3 length of femur on hind edge. Male: Protarsi four segmented. First segment moderately expanded. Remarks This northern member of the approximans species group closely resembles B, juxtadiscolor, but differs in being lighter in colour, particularly the rugose portions of the femora, and in the shape of the aedeagus and parameres. Distribution (Fig, 75) Northern N.T. My tC HOS WATTS pes Halatype male ‘Katherine, NT, at light 9.11.68 -L oA. L. Warson’, in ANIC. Berosus jovtrdiscolar sp. noy. (Pigs I4, 19) Description (umber exited 18) Length 3,5-5,0 mm. Narrowly oval. Elytra weakly humpbacked, bighesr behind middle where itis 1.2 » fleizht at shoulder, Apex of elytron rounded Head shiny black. Pronotuin dark yellow-brown will two central longitudinal stripes darker. Elyiron darkish yellow-brown with punctures and strine darker, in thany individuals elytron almost com- pletely dark. Ventral Surface dark-brown, append ages other than pugose portion of femurs somewhat lighter. Punctures on head strong, revue and well- impressed, most less than 142 au puneture-width apatt. Punecures on pronoeruot moderately dense and strony laterally, weaker and sparser on disc, Dorsal surtace Weakly to moderately reticulare, Elytral strive well-impressed, less so on dise. Second siria on elyfron a little less (han a 1/4 length of clyiron. Punerures in strive sliglitly larger thar chose in interstriaé Punetures stronger towards sides. Interstriae not arranged ina single row, Mesosternal pillar long, well-developed, veritral cdee slopes downwards towards rear, sharply pointed on ventral rear ungle. ventral surface densely but shallowly rugose-punctate. Metacoxal process broad, square ish), lateral lobes weakly lowered, deeply and broadly pitted jn cenrre, triangularly produced backwards in midline. Midtine keel in first abdominal seument niodenuely developed in frant hall. Apical abdominal segment ootched. Rugose partion of metafemtur 2/3 length of fenur, that of mesofermur a little over |) 2, thar of profemur about 1/3 on hind ede, Male: Protarst foar-seemented, First sexnmient noderacely expanded. second seamient small chine segment long aid thin abour lengrh of firse and second semen, Remurks Chis species closely resembles B reardoni but has a more vomiplexly shaped paramere and dillerently Shaped aedeagus. It seems to be a darker species than B reardant which has the rugase portions of the femora the same colour as rest of leg, Only a lew speeiniens of & reardoriare known, A clearer idea of the relationships between these two species must awail (he colleciion of further dyalecial. The preater extent of rugosily an the legs separates i from the otherwise quite similar 2 disealon Types Holotype, mile, “LL 09'S 132-00, Black Point, Cobourg, N.T., 15-23 beh, 1977. 4. A. Weir’, in ANIC, Paratynes; six, same data as holotype, in ANIC: ong, same data as trolorype, in CW two.'Howard Sprites, WT, light 27.68, JO8. 1, Watson’, in ANIC, Distribution (Fig, 77), At present Known only from the coastal areas of the NVD, Berasns timunsi sp. nov. (Figs 16, 21) Description (dumber exanined seven) Length 3.4-4.0 nim. Oval, elytron weakly humped, highest behind middle where it is approximately 1.2 » heht of shoulder, Apex af elytvon founded er weakly trunedred. Head black, shiny, metallic Pronotun yellowish with central panel dark-brown, shiny, metllic. Elytron shiny yellow-brown, with sina, punetures and a few spots black, Ventral surface dark-brown to blak, appendages brown except for rugose portions of meso and metafemurs which are darker, Purciures on ventral surface mod vrately large, dense. Punctures an head strong. regular, well-iinpressed, mos at rear af head less than 1/2 puncturewidth apart, Punetures on pre- noruni strong, variable in size particularly lateralls less dense (han on head, well-impressed. Elytral striae well-impressed, with strong tendeney tor the inner edve of striae to he sharper than outer, Second strig an elytron 1/4 lenath of elytron, Punciuresin étriae strong and little larger (han largest on pro- nouuim, nlerstrial punctures small, subobsolete over most of elytron, Mesosternal keel moderacely developed, pillar well-developed, concave and dent: volate ou veutral cdee front and rear projections largest. Metacoxal process projecting backwards in midline, lateral lobes weakly raised, pointed behinu, weukly converging Lowards front, a wide shiny dig- mond shaped depression in midline, Keel in nividline of first abdominal segment moderately strone reaching weross mos! of seament. Rugose portian of metalemut approximately 2/3 length of femur, that an mesofemur approximately 1/2, that on pro- femur 1/2-173 length of femur Meso- and metatibiae strongly spined. Apical abdominal see- nent With song deep wide noteh lo about 1/2 distance of tiormally visible partion of segment. Mule; Protwarsi foursegmenred, first seunent stromgly expanded. Merafemur with distinet trie angular bulge in lower Wind margin midway alone rugoase poriian. female. Metafemur of sme speciniens show slight thickening in same arca as ip nyale. Remarks This species resembles the nrere widespread B Juntadisculor with whieh i t& sympatric, but differs from it (and all other Berasus) by the shape REVISION OF BEROSUS MN of the mule metafemut which is reminiseent of the femora of some male Dyriscids of the genus Antfiporus. The variably-sized pronotal punctures and very small interstrial punctures also separate it from B fuxtadiscolor and B. reardani, Disrribution (Pig. 82) Known only from the wpe locality at the tip of Cape York, Old, Tepes Holotype, male, 'Bamayza, Old, 29/6/83, Timms’, i SAM. Seven pararypes, same data as holotype; one male. one female, in ANIC; one male, three females in CW. Berusus niger sp. nov. (Figs 25, 30) Description (number examined seven) Length 3.3-4.0 mm. Oval. Elyira parallel-sided, humpbacked, hivhest just behind tbe middle where iris 18 ~ the height at che shoulders. Apex of elytron nol spined. Black, portions of elytral inter- siviae in apical hall and appendages of head, dark yellow-brown. Head and pronotum with large, dense, moderately-impressed punctures. Promoturm with some scattered much smaller punctures in spaces between larger punctures. Strial punctures large, shallow. luerstilal punetures weak, ill defined, lavking lalerally and apically, Second striae approxinuuely 1/2 length of clyiron. Spaces be- tween elytral striae 8 and Y, 9 and 10, and 10 and 11 greater in the middle of elytron than elsewhere. Elytral striae 1, 3, 4, 5, 6, 7, 8 and Il ridged, pronouncedly so an stra &. The stronely-ridged stria 8 bends sharply towards apex where if is abOUL 90° to the edve of elyrron when viewed lalerally, Ventral surface rugose-punctate. Mesosteroal pillar long, ventral edge weakly concave, anrerior ventral wogle wilh very small spine, Melacoxal provess broad, squarish, with large pit in nidling; projecting baekwatds sharply in midline, continued torwards in midline by well-marked ridge, Midline keel on first abdominal sezment sirongly raised for whole length of seyment. Apical abdominal segment moderately natvhed, lateral Manges moderately developed. Rugose portion of metafemur reaching 1/2 way on posterior edge and a little less on anterior edge, rugose portion of mesofemur about 13 length of lemut, that on profernur very small. Male: Protarsi four-seemented. Basal segments not expanded, first seement somewhat longer than seeoud and third which are subequal in size. Reatarks A north-eastern highlands species. Readily sep- araled [rom similar species by the almost completely black colour and the fori of the lateral striae on the elytron. Distribution (Fig. 7S) Nountalnous arcas on east coast north of Towns- ville, Old, Types Halotype, male, ‘Star Valley loaokoul & 3 km W of Paluma Qld, 3.ii.67, at light, LG. Brooks’, in ANIC. Paralypes: one, ‘13.6M up Whitfield Ra, Rd, Cairns, 3/11/70, J. G, Brooks, in ANIC, one, ‘Tinaroo Uk., NO. 1/73 AW.H.' in ANIC; one ‘MI, Spey, N.Q, 11/73 G.B.. in ANIC, one, “7.8M. NW Paluma, NO 5.11.73 J, G, Brooks, in ANLC; two, ‘Ausiralia N.Qid. Windsor Tableland via Mt, Carbine 11-12 Jon 1980 R. |. Store’, in QOPI. Berosus favolatus (W. Macleay) (Figs 9, 26, 31) Hyzrorrophus involutus W. MacLeay, 1873 p. 132. Types There are four specimens [rom Gayndah, Old in ANIC (on loan from the Macleay Museum), label- led ‘Syntypes’, Two of these are B palchellus and (wo B approximans. The original description does not indicate how many specimens MacLeay had be- fore him. Apart Irom the loealily there is little to connect these insects with a type series of RB favo/itus. Two specimens labelled *Hygrotrophus involutus M, L. W, Gayndah’, mounted on the same card, are in Lhe AM labelled ‘Holotype’ and num- bered K1953). They agree well with the bret des~ cription and belong to the species that subsequent workers such as Blackburn and Lea have identified as B involutus | therefore herein designate the lett- hand specimen as the leclotype of Hygratrophus invalulus W. Macleay, 1873. Description (number examined 475) Lengih 3.5-4.5 mim. Oval, elytra humpbacked, highes! just behind middle where about 1.5 + height! al shoulder. Apes of elytron pounced. Head and most of pronotum black, with metallic sheen. Front and side margins of pronotum yellow-brown, ovcasionally lacking, elytron brown, punctures and striae and variable number of markings black. Ventral surface black. Appendages light-brown, tip of labial palpi and rugose portions of meso- and metafemora black, Head and pronorum with derise lane evenly-sized nearly confluent pilpetures, Interstrial punctures on elytton Jaree and well- marked, almost touching, those lowards sides and humeral angles larzer and squarer. Interstrial puncture sinall, shallow, strongest on disc, obsolete laterally. Second sina approximately half length of I2 CPL S. WATTS elyiron. Distance between striae seven and eight and eight and nine often slightly ereater in middle, other singe approximately equidistant! fram each other. Ventral surface rugose-punctate. Mesosternal keel in form Of a long pillar, with a small downward point at tront edge, Metacoxal process broad, sides raised, Subparallel, deeply and broadly pilted in middle, pit devoid af sculpture, projecting backward sharply for short distance in midline, lobed. Midline keel of first. abdominal segment stronyly raised for whole width of segment, Apical abdontirial sexment broadly notched with well-developed lateral Nanges, a pair of very Small tubercules on edge of middle of notch. Rugose portion of metafemur dark, reaching 1/2-2/3 along femur on anterior edge, not as far on posterjor edge Male; Prolarsi four-se¢emented. Seemenis nol expanded, basal ttirge subequal in size. Remarks A common species in clear sireams and pools in the Great Dividing Range from Vie, to north Qld. Many specimens from the higher-areas of Vic, and N.S.W. are darker in colour and lack the distinctive yellow margios on the pronotum. Separated from the two related Species, B niger and Berens, by (he lack of strongly raised and bowed seventh elytral strid, tis also noticeably larger than B ares and is different in colour trom & niger. [tis only in the northern partion of ils range that B invelufus is sympatric with these two similar species. Distribution (Fiz, 76) Coastal eastern Australia (fom southern Gane York to south-eastern Vie. Rervsus areus sp, nov, (Figs 8, 24, 29) Descriptian (number examined 149) Length L-§-3,4 mm. Oval, elyira humpbacked, highest just behind middle where about 1.2 » heighr at shoulder. Apex of elytron without spines, Black, elytral interstriae, edges af pronotum apart from central portion of rear margin, apperidages apart from tip of labial palpi ajd rugose portions of meso and metalemora yellow-brown. hlead and pronotum with dense strong evenly-sized und spaeed punctures. Strial punetures on elyiron large, alnrose touching. Literstrial punetures large bur very shallow, stronger on dise than laterally. Second stria on elytron approximately 1/2 length of elytron. Distance between striae eight and nine greatly enlareed in middle to about twice distance between slmae nine and ten, which is also slightly enlarged in middle. Stria eight moderately carinate in middle section where distance from stria nine greatest. Ventral surface rugose-punclata Mesosternal keel in wide pillar, anterior ventral angle of pillar sharply produced downward inte small sharp point. Mera- coxal process broad, squarish, sides raised, sub- parallel, deeply and widely pitted in midline, projecling backwards sharply in midline, Midline keel on first abdominal s¢ement strongly raised for whole length ol segment, Apical abdominal seg- ment broadly notched wirh well-developed lareral flanges. Rugeose portion of metalemur reaching nearly 2/3 along posterior edge of femur and 1/3 On anterior edge, thal on mesofemur reaching a little over 1/2 length of femur an pasterior edge, 1/3 on anterior edge, that ou profemur 1/3 on posterior edge and less than. 1/4 on anterior edge, Mole: Protars| fourseamented, basal Uiree segments subequal afi size. Reniarks A northern species confined to the sireanis and niverside pools of the ranges around wand to the north of the Atherton Tableland, Smaller than B involutus and & niger and readily separated trom them by colour arid the form oF the eigtith elytral stra. Distribution (Fig, 79) Ranges of soucher Cape York and Atherton Table- land, Qld, Tipes Holotype, male, ‘S mi. N. Bloamfield Rn., N,O, 7-9. May 1970S, K, Curtis’, in ANIC. Paratypes, 11 same data as Holotype, in ANIC; one same data. ii CW. Berosus trishae sp. nov, (Figs 37, 43) Hescription (umber examined 39) Length 2.4-4.5 mm, Oval, Elytra parallel-sided, humpbacked, highest just behind middle where it is 1.20.2 higher than at shoulders. Apex of elytron rounded. Yellow-brown, head, three pough broad longitudinal patches on pronotum, about a dozen small patehes on elytron and vague areas of ventral surface darker. Head and pronotim moderately covered with large strong punctures, A single even row of slightly smaller punccures along lront and rear edges of pronotum. Strial punctures on elytron very large, almost confluent, stranpest laterally where they are ulmost square, Interstrial areas shiny lacking punctures except apically in interstria 3. Second elytral stria a litthe more than 1/2 length of clytron, Interstrial areas raised somewhat lowards apex of elV¥tron. Ventral surface moderately punctate, Mesosternal keel produced in robust pillar, ventral edge of pillar convex. Metacoxal process very broad, parallel sided, central portion with wide pit, lateral lobes project downwards at about 35°, narrowly and triangulacly produced backwards in REVISION OF BEROSUS i inidliie. Midline keel on lirst abdominal segorernt strongly raised for whole length of segment. Apical abdominal segment widely and deeply notched with well-developed lateral flanges, Rugase porion of metafemur abou! 2/3 length of femur thal on mesofemur |/2-2/3, and that on profemur aboul 1/2 length of femur, Male: Protarsi lour-segmented, basa] segments Hot expanded Remarks A northern Species, Separated (rom the quite simular & quadrapuncratys by ie ereater extent of rugose sculpture on the temura and characters of the aedeagus. The sculpture on the head is a Tit(le rougher than in & quadrupunciatus and the punctures on pronotum and élytra are smaller. Al- thouwh this size difference is clearly evident in direct comparisons of most specimens, | have been Unable 16 quantil’y it to enable the character to be used to separate the species. Most specimens are < 3,3 mm long bul a female from Charters Towers, Qld, in NSWD, is considerably larger at 4.5 mm, f ean derect no other difference from N-T. specimens. Distribution (Fig, 80) Darwin area, ST, Atherton Charlers Towers Qld. Tableland and, Tppes Holotype, fenvile, ‘NT. Lake Bennett area c. 25 km SE. of Manton Dam. 29-30 Dee, 1979, M, B. Malipatil’, in NTM, Paratypes; 20, ‘N.T., Ck. in Flolmes Jungle 131.1980 M, Malapatil’, in NTM, iwo same data in CW Berosus macropunctaius sp. Wav. (Fig. 70) Description (wmumber examined twa) Length 3.0-3.6 mm. Oval. Sides of elytra subparallel, humpbacked, highest about 1/3 dis- lance front apex of elytra where it is abaut 1.2 height at shoulders. Apex of clytron squarish. Yellow-brown, head and propotum black except for narrow yellowish band om lateral and hind edges, elytron wilh scattered darker patches, Head and pronotum strongly rugose-punckale, Stelal pune- (iires on clytron large, larger and squartsh towards sides, Interstrial areas shiny, impunetate exeept for a row of very small setae-beariny, punctures towards apex of third interstria. Lateral three interstrial areas raised into weak ridges, fnterstrial areas at extreme apex weakly raised, Ventral surface stromaly rugose- punctace. Mesosternal keel produced into a narrow pillir, Ventral edge of pillar slopes downwards rowards rear and has a small downward pointing spine an ventral anterior angle, Metadoxal pravess very broad, lateral lobes parallel sided, rather narrow, separated by wide pit in centre ol process, open towards rear, lateral lobes slope downwards at about 20%, broadly triangularly produced backwards in midline. Midline keel of first abdomi- val segmeni strongly developed over Whole length of segment, veotal edge serrale, apical abdominal segment deeply and widely norehed with well: developed lateral flanges. Rugose portion of metufemur 3/4 length of femur, (hat of mesolemut a little less than 3/4, thal on profemur 1/2 length of femur. Male: Protarsi four-segmented, basal segments not expanded, Remarks A distinetive species readily separated from & rishae and B quadrapuncrarus by the strongly rugose-punctate black head and pranolum. Distribution (Fig. 76) (N-T. (ype localities only,) Types Holotype, inale, ‘Burrell’s Ck Stuart H’way NT, 24 Nov, 1972 D.H. Colless'!, in AIDC. Pararype, one, 13°LS'S, 131°06'E, Adelaide River, NT, b6,x-72, M. 5. Upton’, in ANIC. Berosus quadrapunctatus sp. nov. Deseriplion (number examined 4) Lenerh 3.2-3.9 mm. Elytra humpbacked, highest just behind niddle where it is 1.3 height at shoulders, Apex of elytron rounded, weakly flanged. Yellow-brown With head black and three broad vague longitudinal patches on pronetum, scailered patches on ¢elytron and patches in the midline ventrally, darker, shiny. Head and pronotum moderately covered with large well-impressed punctures. Elytron with strial punctures very strong, particularly laterally Where they are square Punerures in adjacent laleral striae alnyost (Ouch. Interstrial areas shiny, lacking punctures excepl for some miuute setae-bearing ones Lowards apex if intersiria three, Second clytral stria reaches a little over 1/2 length of elytron, luterstrial areas towards extreme apex become strongly raised, Ventral surtace covered With numerous large bul separate rugose punctures. Mesosternal keel produced inti a narrow’ pillar, concave on ventral edge: Metacoxal process very broad, parallel-sided with large pit in midline, narrowly triangular, projecting backwards in midline, lateral lobes slope downwards at about 45" Keel on first abdominal segment strongly raised for Whole leneth of segment, serrate on ventral edpe Apical abdominal segment broadly notched, with 14 Cry Ss; well-developed lateral flanges. Rugose portion of metafemur a little over 1/2 length of femur, that on mesofemur a little under 1/2, that on profemur 1/3 length of femur. Male: Protarsi four-segmented, basal segments not expanded. Remarks A little known species differing from B. trishae by the less well-developed rugose sculpture on the femora and in the more uniform sculpture on head. A female specimen from Charters Towers, Qld.,(in NSWDA) may belong to this species. It is longer (4.5 mm), the rugose portion on the femora is a little greater, the sculpture on the head is rougher with some scattered very small punctures as in B&, frishae. In these characters it is intermediate between B trishae and B. quadrapunctatus. Distribution (Fig. 80) Known only from the type series from McArthur River, N.T, Types Holotype, male, ‘McArthur River, 16°47'S 135°45'E, 14 km S by W of Cape Crawford, NT., 25 Oct. 1975 M. S. Upton, in ANIC. Paratypes, three, same data as holotype, in ANIC. Berosus queenslandicus Blackburn (Figs 23, 28) Berosus queenslandicus Blackburn, 1898, p. 223, Berosus quartinus d’Orchymont, 1943, p. 6, syn, nov. Types B. queenslandicus, holotype female, Qld., in BM(NH), seen. Paratype, female damaged, in SAM, seen. B. quartinus, holotype, male, and paratype, in MNHN. Description (number examined 42) Length 4.2-6.3 mm. Oval, elytra humpbacked, approximately twice as high just behind middle as at shoulder. Apex of elytron rounded. Head black with metallic sheen. Pronotum shiny, widely black or dark-brown in midline, brown at sides. Elytron shiny, brown with dark-brown to black mottlings. Ventral surface mottled dark-brown and black, appendages brown, apical segment of labial palpi darker, rugose portions of meso- and meta femora dark-brown to black. Punctures on head large, subequal except for clypeal margin where they are smaller, those on disc separated by about 1/2 their width, a few scattered very small punctures in some specimens. Head with small midline ridge on back, sometime lacking. Punctures on pronotum as on head, closer together laterally. Punctures in elytral WATTS striae well-marked almost confluent in any one stria, stronger towards front. Interstrial areas vir- tually impunctate except for small area of large punctures on humeral angles. In some specimens some very shallow small punctures can be seen in some lights in interstriae on disc. Striae weakly impressed on disc, stronger laterally and apically. Second stria on elytron reaching 2/3 length of elytron. Striae approximately equidistant from each other. Ventral surface rugose-punctate. Mesosternal keel in form of high narrow pillar, ventral edge con- cave, keel on midline of mesosternum forward of pillar weak. Midline keel on first abdominal seg- ment well-marked, reaching almost completely across segment but weakening rapidly towards rear. Metacoxal process broad with well-marked lateral lobes, sides subparallel, with depressed shiny im- punctate area in midline. Rugose portion of meta- femur about 1/3 length of femur, ending in relatively straight edge across most of femur, that on metafemur 1/3, that on profemur 1/3 length of femur on hind edge. Apical abdominal segment deeply notched in midline with lateral flanges to notch. Male: Protarsi four-segmented, basal segments subequal not expanded. Remarks Separated from the superficially similar B. duplopunctatus by the slightly larger and differently shaped rugose portions on femora, the virtual lack of smaller punctures on head and pronotum and the virtual lack of interstrial punctures. Some specimens have some minute punctures in between the large ones on the head and to a lesser extent on the pronotum, but these are much smaller and fewer than those in B. duplopunctatus. Synonomy based on d’Orchmont’s description and illustration of male genitalia. Distribution (Fig. 78) Coastal south-eastern Australia from Caloundra, Qld., to Adelaide, S.A. Berosus duplopunctatus Blackburn (Figs 22, 27) Berosus duplopunctatus Blackburn, 1888 (1889), p. 828. Types Holotype, female, Pt Lincoln, S.A. in BM(NH), seen, Paratype, female, Pt Lincoln, S.A. in SAM, seen, Description (number examined 109) Length 4.5-6,4 mm. Oval, elytra humpbacked, at highest point just behind middle about 1.5 x height REVISION OF BEROSUS 1s at shoulder, Apex of elyiron rounded. bead relatively wide, black, shiny with metalhe sheen. Midline of pronotim black, sides brown. Elytron predominantly brown with punctures outlined in black plus some black spots, extent of black varies between specimens and is the daminani colour in many. Ventral surface black, appendages light- brown, lip of apical segment of labial palpi black, pupose portions of meso- and metafemora black, Head densely punetare with both large and small purictures. Pronotum with large and small punctures, less dense than on head. Punetures in elytral striae well-impressed, almost confluent, stronger towards front, about as sirong on cise as laterally, Interstrial punctures shallow, small, numerous, scattered, very weak taterally. Stnae moderately impressed, slightly weaker on dise Striac approximately equidistant apart. Second stria onelyvon reaches to 3/4 length of elytron. Hurneral angle of elytron serrate due to large punctures close to cdee. Ventral surfaee strongly ruose-punetate, Mesosternal keel in form of high narrow pillar, weakly concave on ventral cdae, keel in front of pillar weak. Midline keel on first) abdominal segment well-marked in front half, weaker im hind half of segment. Apical abdominal segment deeply notched, notch with lateral Tlanges. Metacoxal process with well marked Jateral lobes, converging towards rear and wilh small shiny inipurectate depressed urea in midline, Rugose portion of mesolemur about 1/3 length of femur on hind edge, ubout L/4 Jenerh on front edge of lemur, that on hind edge of metalemur 1/4, thaton profenmtue b/4 lenerh af femur. Male: Protarsi foursepemented, basal three subequal not expanded. Remarks Reudily separated from the superticially similar B queeusiaidious by the puncrace elytral intersinae ald the presence of Small punerures on the pronoun, Distribucian (Fix, BO) Coustal easrern Australis from Sydney, N.S.W. (0 Kangaroo Island, SA Tas. Berosus nutans (W. Macleay) (bigs 5, 37, 63) Hyerotrophus mitans W. Macl vay, 1873, p. 132. Berosus pallidulus Faatoaire, (379, pos, syn, aller Blackburn, 1898, p, 222, Types Two synivpes fabelled “Gayadah’ and mounted on the sane card are in the ANIC, The leti-hand side specimen, a female, is here designated as leerory pe. Descriplion (qumber examined 440) Length 4.2-8.8 mm, Elongate oval, hot humpbacked, Apex of elytron rounded or obliquely truncated, Dorsal surface brown with vague darker markings particularly on head aud pronotum. Elytron with steiae darker, Ventral surface dark brown bo black, appendages brown to dark-brown except for darker tip of Tabial palpi. Apex of abdomen occasionally brown, Punelures on head well-marked, weaker tawards front, transversely elongate. Sculpture on pronotum consists of a close nelwork of Vine lransyerse wrooves which cover most of the surtave, all but obscuring, punctures which themselves tend to be transversely clongate as an head, A fine reticulation ts also present on some specimens. Scutellum seulptyred as pronotum, Elytron with siriae lacking or only Weakly im- pressed, rheir position allen oullined by rows of small crapsversely expanded. darkly pigmented spors. Elytrot densely and evenly covered with short rather stout setac, each seta arising from a small shallow transverse groove. Ventral surface shallowly rugose-punclale with small setae arising trom each puncture. Mesosternum with weak keel in midline, extended quite strongly buckwards between imeso- voxae. Midline of first abdominal segment with moderale keel in front 1/4. Metadcoxal process raised, sharply triangular with elongate oval area in midline devoid of sculpture, Rugese portion of metafemur 3/74 length of femur, that of mesofemur 2/3, that of profemur 1/2 length of femur, Male: Provarsi toursegmented, basal segment expanded, aboul as long as second and third segment combined, scvond segment expanded, same length as simple third segment. Reniurks A common and widespread inland species, readily recognised by the unique sculpture of the pronotum. Distrihutian (Fig. 78) Coastal and southern inland Australia. Berosus pulehellis WS. Nacleay (Figs 48, 33) Beresus pulchellus W. 5. Macleay, 1525, p. 35. Fiyeratraphus devisi Blackburn, L898, p, 225, svi alter d'Orehymont 1943, p. 421. Types B. pulchellus, Java, lovation unknown, Bo deviat fiolotype, ? female, Old., in BM(NH), seen, Description (number examined 233) Length 2,8-4.8 mii. Elongate oval, elytra tot humpbacked. Apex of elyiron founded, Head 16 C. H.S. black, shiny. Pronotum light-brown with front edge narrowly black and usually with wide central panel black, black area not reaching rear border and often with portion of midline narrowly light-brown. Elytral striae often darker. Ventral surface black, appendages light-brown except extreme apex of labial palpi which is darker. Head evenly punctate with moderately dense small punctures, punctures somewhat weaker towards front. Pronotum moder- ately and evenly covered with small punctures. Elytron strongly and evenly covered with similarly sized punctures, each puncture with a well-marked seta. Elytral striae lacking or very weakly impressed. Punctures in elytral striae same size as interstrial punctures and difficult to distinguish from them, Ventral surface evenly, shallowly rugose-punctate. Midline of prosternum with weak but well-marked keel, weakly projecting backwards between meso- coxae. Midline of first abdominal segment weakly raised in front half, Metacoxal process bluntly triangular, raised, midline with narrow oval area devoid of punctures. Apical abdominal segment deeply notched. Rugose portions of meta and meso- femora 3/4 length of femura, that on profemur 2/3-3/4 length of femur. Male: Protarsi four-segmented, basal two segments moderately expanded, basal segment about twice length of second which is about same length as third. Remarks This widespread and common species is readily separated from all other Australian Berosus by the small size, black head and virtual lack of elytral striae. It is most commonly collected from the sandy pools formed during the dry season in the beds of the major northern and eastern river systems. It has recently been collected in numbers in sewerage settlement pits at Whyalla, S,A., an unusual habitat and a long way from its normal range. Distribution (Fig, 82) Tropical Australia; Whyalla, S.A.; Asia. Berosus dallasae sp. nov. (Figs 32, 38, 70) Description (number examined 80) Length 3.8-6.8 mm. Elongate oval. Elytra not humpbacked. Head and pronotum shiny, elytron rugosely reticulate. Apex of elytron produced into two spines, outer one usually strongly and widely developed, inner often small or even vestigal. Dorsal surface light yellow-brown, rough patches on head, pronotum and elytron darker, elytron stria darker. Ventral surface dark-brown, appendages lighter, tip of lapial palpi dark, Head evenly and strongly punctate, punctures much larger than eye facet, much weaker towards front, Pronotum evenly WATTS punctate with large strong regular punctures except for some smaller areas along front and rear margin. Punctures on disc have a tendency to become longi- tudinally elongate. Elytral striae strongly impressed. Stria 2 with punctures a little smaller than those on pronotum and about twice size of those in adjacent interstriae. Interstrial punctures evenly, densely but shallowly impressed over whole elytron, each puncture bearing a small seta. Ventral surface rugose-punctate. Midline of mesosternum weakly and broadly carinate, moderately projected back- wards. Midline of first abdominal segment weakly carinate in front 1/4. Metacoxal process broadly triangular, midline weakly carinate in front of a small diamond-shaped area in centre, devoid of sculpture and bounded on front sides by weak edges. Rugose portion of metafemur 3/4 length of femur, that on mesofemur 2/3-3/4 that on profemur 1/2-2/3 length of femur. Male: Protarsi four-segmented. Basal two seg- ments moderately expanded, first segment about 1.5 x length of second which is a little larger than the narrow third segment, Remarks This widespread species of northern Australia is readily separated from the two other, equally wide- spread, species with setose elytral punctures, by the lack of strongly transversely elongate pronotal punctures (from & nutans), and lack of a black head (from B. pulchellus). Distribution (Fig. 75) Tropical Australia. Types Holotype, male, ‘| km N. of Millstream, W.A,. (21°35'S 177°04'E), 9-10.iv.1975, M. S. Upton’, in ANIC. Paratypes, nine same data as holotype, in ANIC; one same data as holotype, in CW. Berosus macumbensis Blackburn (Figs 59, 65) Berosus macumbensis Blackburn, 1896, p. 259. Types Holotype, male, Macumba Creek, S.A. in BM(NH), seen. Description (number examined 150) Length 5.6-7.8 mm. Elongate oval, not humpbacked. Apex of elytron with two spines, the inner usually small and the outer as long as the distance between the two spines. Elytra shiny. Dorsal surface light brown to brown with dark mot- lings. Elytral striae and punctures on disc darker. Ventral surface dark-brown to black. Appendages brown except for extreme tip of labial palpi which REVISION OF BEROSUS 17 7 7 @ B. majusculus e B. involutus a B. vijae © B. macropunctatus A B. dallasae a B. josephenae oO B. niger Oo B. approximans 4 B. reardoni 2 2 77 W 78 W e 8B. australiae e B. nutans a B. juxtadiscolor a B. gibbae gs B. ralphi A B. decipiens B. queenslandicus FIGURES 75-78. Distribution maps for Australian Berosus spp. 18 C. H. 8. WATTS 3 80 W 7 @ B. arcus e B. debilipennis m B. discolor a B. duplopunctatus A B. sadiae mw 6B. quadropunctatus 0 B. aquilo © B. trishae a B. nicholasi y 82 Vv) 8 a 8B. munitipennis @ B. pulchellus @ B. macumbensis O B. aemoenus A B. veronicae A B. timmsi 4 B. subovatus FIGURES 79-82. Distribution maps for Australian Berosus spp. (cont.). REVISION OF BEROSUS )¥ is durker. Head strongly and evenly punctate except towards front where punctures are considerably smaller and weaker, Punctures on pronolulh uneven in-size and often with a rather patchy distribution, A rew of very Small punctures along frontand rear margins. Elytral striae weakly impressed, lor the mosi part reduced to a row of Well-impressed pune- tures about the size of the larger punetures on pronatum. Inerstral punctures on disc markedly symaller than (hose on sides of elytra; in no case is there a tendeney for interstrial punctures to farm a single row, Interstrial punctures on dise tend to be uneven in size. Underside densely rugose- punctate, Midline of mesosrernum with weak rather rounded keel, projecting alittle distance backwards. Midline of first abdominal seament with litle or no keel, Metacoxal process raised, triangular, a narrow diarnond or oval-shaped area devoid of sculpture io the midline. Rugase portion of meta- femur t/2 to 2/3 Jength of femun that on mesofemur.a little over 1/2.and on profermura 1/3 toa 2 length of femur. Male: Pratarsi four-segmented, basal two sepmenrs expanded, basal segment larger than second, sevond and third segment subequal in length, Remarks A widespread inland species readily recognized Irom the other large Berosus by the pattern of punctation on the elytra, Where the outer punctures are much stronger than the inner punetures, Distribution (Fig, 81) Inland oAustraha. Berostis minitipennis Blackburn (Figs 3, 58, 64) Berosus munitipennis Blackburn, 1895, p. 30. Livper Holotype, ?mate, RM(NH), seen, Luke Callabonna, S.A., in Deseripnon (umber examined 48) Length 4,8-4.6 nim. Elongate oval, not humpbacked. Apex of elytron with yery long narrow ouler spine aud shor joner spine, outer usually Jonger than distanee between spine bases.. Light-brown, pranotum and head slightly darker than elytra. Suture fines and some punctures usually black on elyrron disc, Ventral surface dark-brown to black, appendages brown, ip of labial palpi darker. Head moderately fo quite densely covered with punctures, large and well-marked in basal halt, smaller towards froor. Elytron shiny or with weak to moderate reliculation, Pronotum sparsely or moderately and father unevenly covered with variably sized, well-marked punctures, front and fear margins with anly a few shallow small. punc- tures. Elytral striae weakly to moderately impressed, otien reduced oyer much of disc toa row of punc- tures. Serial punctures about same size as larger ones on pronotum. Interstrial punctures sparse, shallow, lacking completely over atuch of elytra in some specimens, where present tend to be arranged in single row, Second elyirae stm less than 1/4 length of elytron. Ventral surface shallowly rugase punctate. Midline of mesosternum weakly carinate. weakly extended backward, Midline of first abdominal seement weakly carinate in basal 1/2 oF 1/4. Mctacoxal process [riangularly-raised, with patch in midline withour sculpture, pateh bounded at rear by a slight V-shaped ridge extended back- wards in midline to tip af process. Rugose portion of metalemur |/3-2/3 leneth of femur, that of mesoferur I/4-1/2, thal on profemur 1/3 lengtt of fernur along ventral edge. Mule: Basal two seanmients of protarsi moderately expanded, First segment about 2 « lengih of second, which is.about the same length as the third, Remarks An inland species readily reeognized by the long, thin, outer elytral spines. and virtual lack of an inner ong and the weak development of sculpture on the elytra, This latter charaeter tends to be more marked in central and noerthert specimens compared with those from Vie. The eontrast of the pale upper side and black ventral surface in most specimens is quite striking. The extent of ruvosity an the femora ts particularly yariable in this species. Distriburian (Fig. 81) Inland Australia south of Barrow Creek, NLT. Berosus amoenns sp. mov. (Figs 56, 62) Distribution (number examined nine) Length 3.3-4.5 mm, Elongate oval. Weakly reticulate in some specimens. Elytra nol hump- backed. Apex of elytron truncated, spines vestivial, Dorsal surface dark-brown to black, edges of pro- notum arid elytron yellow-brown, sides of elyiren with three pale patches extending inwards from sides, (he apical ones more distinct, Dise of elytron variegated yellow and brown-black.. Ventral surface dark-brown, appendages a little lighter, apex of labial palpi dark-brown. Head sparsely covered wilh strong punctures, about. 1.5 x size of eye facet, those towards frant much smaller. Pronotum moderately but unevenly covered with strang punctures, a little larger than those on head, punctures tending to be 20 ©. 1 5. WATTS elongate longitudinally, a few isolated smaller punctures along front and rear margins, elytral striae moderately lo slrongly impressed. Second sina on elytron with 11 to 16 punctures. Punetures in striae, between siriae and on pronotum subequal in sizvé, Interstrial punctures lateral ro siria 9 apyoximately arranged in one line. Veriral surtace rugose-punctate, Midline of mesosternum weakly raised, moderately projecting backwards. Midline of first abdominal seement weakly carinate in front 1/4, Metacoxal process raised, triangularly pro- duced backwards with a shor! narrow keel in midline at extreme rear apd small narrowly oval area in widdle devoid af sculpture. Rugose portion of metalemur 2/3 « length of femur, that of mesoferur 1/2-2/3 that of profemur 1/4-1/2 length of femur, Male; Protarsi four-segmented, basal two segments broadly expanded, basal longer than second which is a litle longer that the small third seginent, Apical abdominal segment entire. Remarks A tare species from coastal NCT., readily recognised by the black head and predominantly black pronotum. The size and black head resemble the & approximans aroup of species bit the tinlobed mietacoxal process and lack of mesosternal pillar separate il from that group. Distribujian (Fig, 82) Coastal NT. Tipes Holotype, male. 12°52'S, 132°50'E Koonearra, \Skm E of Mt, Cahill, NT, 15.41.1972, M, §. Upton’, in ANIC, Paratypes; two, ‘16°28'S, 136°09'E 46 km SSW of Borroloola, NV. 28 Ot, 1975. M.S. Upton’, in ANIC; six, ‘King River NUT, W. McLennan Pres. H. L, White 17,1016", in NMV, Berosus josephenae sp, nav, (Figs 55, 41) Descripron (nuriber exaniined 19) Length §.5-6.0 mm. Elongate oval. Elyrra wor humpbacked. Apex of elytrou weakly to quite strongly produced backwards, ending in rwo spines, the outer well-develaped, the inner much less so, Dorsal surface light-brown, with patches on head, centre of pronotum, elywon, inner striae and pune- tures an elyiron darker. Ventral surface dark brown- black, appendages lighler, rugose portions of meso and metafemora and tip of lapial palpi dark brown- black. Head strongly punctate, punctures larger that eye facets, punctures weakening towards lront. Pronotum strongly densely and cvenly punctate, punctures ol one size execpt fora few stnaller ones on extreme front and rear edges; Elyival sitlae strongly impressed. Siria 2 with 15 to 23 punctures about size of thase an pronotum and 1.5 to 3 + size of {hose in adjacent interstrial area. Inferstrial punctures relatively large, uniform in size, well im- pressed, arranged in more than one row over most of elytron, larger but shallower towards sides. Ventral Surface strongly rugose-punciate, Midline of meso- sternum weakly to moderately carinate, moderately projecled backwards, Micline of first abdominal segment raised in front third, Metacoxal process raised, tritumgularly produved backwards, weakly ridyed in midline except for « broadly diamond- shaped area in centre which is devoid of sculpture. Rugose portion of mejafemur 2/3-3/4 length of femur that ol mesofemur 1/2-2/3, thal of profemmur about 1/2 length of femur. Male: Protarsi four-segmented, basal seaments moderately expanded, basal segment niuch longer than second seginent which is aboul same length as much narrower third segment, Apical abdominal segment entire, Remarks As yet known only by 4 few speciinens from coastal NT, this rather clongale species is distinguished by ils Strong even punctration on the pronatum ane cle black rugase portions of the femora. Distribution (Fig. 76) Coastal N.T. and Cape York, Qld. Tepes Holotype, male, 12°23'S 132°57'E § km NNW of Cahills Crossing, NT. East Alligator River, vi. 73, Upton & Foehan’, in ANIC, Pararypes; six, two same data as holotype, in ANIC: one, ‘Mareeba 22.1976 Ko & EF. Carnaby’ in CW. one, NUT, Junction of Arnhem Hwy & Genpelli Road MV. light 26-27 Sune 1980 M, B. Malipahi’, in NTM; one, NT, Lake Bennelt area © 25 km SE of Manton Dam 29-30 Dec, 1979. M. B. Malipatil’, in NT, Berosus gibhae sp. nov. (Figs 33, 39) Description (gumber examined '7) Length 4,0-5,0 mm, Elongate oval, Elytta jot humpbacked. Shiny but with elytvon of many specimens with a line-meshed reticulation. Apex af elytron wilh a stout moderately-sized outer spine and a virlually absemt inner one, Dorsal surface hzht-brown, elytral punctures and many striae and vague patches on head, pronotum and elytron darker. Ventral surface dark-brown, appendages lighter, except tip Of labial palpi which is dark- brown. Head moderately densely covered with REVISION OF BEROSUS 21 strong punctures, strong, 2 x size of eye facets except towards front where they become much smaller, Pronotum evenly and quite densely covered with strong punctures, the same size as those at rear of head, a few smaller ones along front and rear margins. Elytral striae moderately impressed, with punctures the same size or slightly smaller than those on pronotum. Second elytron stria with 12 to 18 punctures. Punctures in interstriae smaller than those in striae, larger but shallowly impressed towards side, over most of elytron arranged in one or two irregular lines, often bearing a single seta particularly those towards sides. Punctures in interstria 3 of uniform size, Ventral surface densely but finely rugose-punctate. Midline of mesosternum strongly raised, moderately projecting backwards. Midline of first abdominal segment weakly carinate in front 1/2-1/4. Metacoxal process raised, tri- angularly produced backwards, with a broad diamond-shaped area in middle devoid of sculpture. Rugose portion of metafemur 2/3-3/4 length of femur, that of mesofemur 2/3, that of profemur about 1/2 length of femur on rear face. Male: Protarsi four-segmented, basal segments weakly expanded. Basal segment longer than second, second and third segments subequal in all dimensions. Apical abdominal segment broadly but shallowly notched. Remarks This widespread northern species is rare in collections. The peculiar fine sculpture of the elytra, which is either irregular or with a fine grained reticulation, and the presence of a large number of setae- bearing punctures, are reminiscent of B, nutans and B. dallasae which show these characters to a greater extent. The more normal pattern of elytral punctation readily separates it from these species, in which the elytra are densely and evenly covered with punctures. The round rather than elongate pronotal punctures also separate it from these otherwise quite similar species. The light- coloured rugose portions of the femora separate it from B, josephenae. Distribution (Fig. 78) Coastal northern Australia. Types Holotype, male, ‘N.T. Katherine low level Native Park, 25 Ap. 1980 M. B. Malipatil’, in NTM. Paratypes; six, two same data as holotype in NTM; four ‘NT. N. P. Korlonjotlok Stream 18 Nov. 1979 M. B. Malipatil’, in NTM, Berosus majusculus Blackburn (Figs 54, 60) Berosus majusculus Blackburn, 1888 (1889), p. 824. Types Holotype and paratype in BM(NH), seen. Paratype, male, Adelaide, S.A. in SAM, seen. Description (number examined 336) Length 6.0-8.7 mm. Elongate oval, elytra not humpbacked. Apex of elytron with two well- developed spines, the outer longer than the inner. Dorsal surface brown with darker mottlings. Elytral punctures and striae black, strongly so on disc, weaker or absent on sides. Ventral surface black. Appendages brown, tip of labial palpi darker. Head relatively narrow, evenly strongly and quite closely punctate, punctures much weaker towards front. Pronotum unevenly covered with moderately dense, well-marked punctures, variable in size, a little stronger laterally with a row of small punctures along front and rear margins. Elytral striae well- impressed, second stria 1/4 ‘length of elytra. Punctures in striae well-marked, same size or larger than those on pronotum, interstrial punctures not in one line except in some lateral interstriae, well- marked, a little smaller than those in striae, particularly on humeral angles. Reticulation of elytron absent to moderately strong. Ventral surface densely and evenly rugose-punctate, Midline of mesosternum tending to form weak keel, weakly projecting backwards. Midline of first abdominal segment with weak keel in anterior 1/4. Metacoxal process raised, narrowly triangular. Midline with long narrow area devoid of sculpture. Rugose portion of metafemur 1/2 to 2/3 length of femur, that on mesofemur a little over 1/3 along ventral face, that on profemur about 1/3 length of femur along ventral face. Male: Protarsi four-segmented, basal two segments dilated. Basal segment about twice length of subequal second and third segments. Apical abdominal segment notched. Remarks This common southern species is readily separated from the rather similar B australiae by lack of black rugose, femore. It is more difficult to separate from B. veronicae, with which it is generally sympatric, but it is larger and has a greater number of punctures on stria 2. The notch on the last abdominal segment of the male also separates this species from others of its size. Distribution (Fig. 75) Southern coastal Australia from Perth to Sydney areas. Berosus veronicae sp. nov. (Figs 67, 68) Description (number examined 209) 22 CORLS. Lengitt 35-64 mm. Elongate oval, elytra not humpbacked. Apex of elytron with two Weak blunt spines. Dorsal surface light-brown with darker mottlings. Elytral punctures and striae black, strongly sa on dise in many specimens, Weakly or not ut all lacerally, Ventral suclace black, append- ages light-brown, tip of labial palpi a little darker. Head evenly, strongly and quite closely punctale Punctures toward front weaker. Pronolum evenly. strongly and quite closely punctate, svatered smaller punctures particularly on disc and front and rear margins, weakly reticulate in some. Elytron variably reticulate fram none to moderate, Elytral striae well impressed. Second stra about |/4 length of elytra. Strjal punctures about size of targer punctures on proootum. Interstriae | to S scattered, those between more lureral striue tending to be in one row except for extreme lateral edge. Ventral surface densely pugose-puretate, Midline ol’ pro- ‘sterniudy Weakly and evenly keeled, projecting backwards in weak prosrernal process. Midline of first abdominal segment weakly keeled in anterior 1/4. Metacoxal process raised, triangular, midline with diamond-shaped area devwid of sculpture in middle and shght keel to the rear of this. area. Rugose portion of merafemur berween 1/3 and 1/2 length of femur, that on mesofemur 1/3, that on profermur 1/4 length of femur. Male: Protarsi four-segmented, basal two segments moderately expanded. First sexment about twice length (and size) of second, which isa little longer than the narrow third segment, Apical abdominal segment entire, Remarks A cammon species in southern Australia, this species has frequently been mis-ideritfied in collections, usually as & majusculus, The un- notched apical seyment in the male separates it from this species. The smaller size and fewer punctures in the area between the lirst two elytral striae will separate all but the occasional specimens. Balbipes Fauvel is another name rhat has been altached to specimens of this species but this isa species from New Caledonia with differen aedeagus and other characters, Distribution (Fig, 8) South-eastern Australia, ‘Tas. Tipes Holotype, male, ‘Goulburn’, in SAM. Paratypes, same data as the holowpe, in SAM; two, ‘Albury NUSW. 1/61 CW", in CW. Berosus austratiae Mulsant {Figs 71, 74) Berosus ausrratiae Mulsant, 1859, 1p. 38, WATTS Berosus exlernespiaosus Pairmaire, i879, p. Rl sven. aller D’Orehymont, 1943_ Berosus gravis Blackburn, 1888 (89), p 826, syn, nov, Tipes Berosus australiae, not loeated (Hape, Dept of Ent. Oxford, revords i as having been sent ta BM(NH) in 1925). Type locality ‘Australia’, Berosus zravis; Holotype, male, S.A. in RM(NH), seen. Paratype, malein SAM, seen. Paratype, male, in SAM, seen. Type locality (Paratype), Murray Bridge, S.A. Berosus externespinosus, nol lovaled (?7MHNHI. lype locality, Rockhampton, Old. Description (number examined 455) Length 6.5-9.0 tam. Elongate oval, not tiump- backed, Apes of elyLron with two spines, the outer strong and usually abour two times length of inner, Dorsal surface brown to dark-brown with darker motlings. Elytral striae and punctures outlined in black except for punctures laterally, Ventral sur face black, Appendages brown, rugose portion of femora black or dark- brown. Punctures ov head nioderately strong and dense, much Weaker towards front, Pronotun) moderately covered wiih well- marked variable sized punctures, front and reat edges with rows of elasely spaccd small punctures. Elytron with sinae moderately impressed, strial punetures relatively small, aboul same size as those on pronotum. Interstrial punctures seatreredt, smaller, those in more lateral incerstriae tending to form a single line to varying degrees, Blyton surface shiny or Weakly to moderately reticulate with a fine reticulation. Ventral surface densely rugose- punctate, Midline of mesosternuin raised into weak keel, weakly projecting backwards. Midlinpe of first abdominal segment weakly keeled in front 1/4, Metacoxal process yaised, iiangular, with diamond Shaped area devoid of sculpture in midline. Rugose portion of metalémur 2/3-3/4 leneth of femur that on mesofemur a little more (han 1/2. that on pio- femur about 1/3 length of femur. Mule: Protarsi four-segmented, basal two scx ments strongly expanded, basal segment about 1.5 * length of second, whicl is tach longer than the small third segment, Apical abdominal segment entire. Remarks In southero and eastern Australia this large common spevies 18 readily recognized by the black rugose portions of the femora, In northern Australia some specimens have the nigose portions dark-brown and not greatly darker (han the rest of the les, and closely resemble B sadiewe and & decipiens, Separation of these three species is hest REVISION OF BEROSLS 44 done on aedeagi as. although & wivstraliae is nenerally larger, aud generally has stronger elytral purictupes, this is nor always so, Distrifuition (Fig, 77) Virtually Australia-wide, more common ii sauth- cast than elsewhere, rare in W.A., apparently absent from centre and north of thar state. Berosuy decipiens Blackburty (Fips 47, 52) Berasus decipiens Blackburn, |S88, p. 827. Types Holotype and pararype in BM(NH), seen, Paratype in SAM, seen. Descriplion (number examined (8) Length 6.1-8.0 mm, Elongale oval, nor hinmpbacked. Apex of elytrow with two weul spines placed close together. Reddish-brown with elytral siriae and punetures on elytron and pronotum black, particularly on dises, black areas on elywron coalesced in. some spevimens lo give extensive black areas on disc, Ventral surfaee black, appendages reddish-brown, rugosé portions of lemora darker Head moderately punctate with rather shallow punctures, those in front half smaller. Propotuin sparsely and rarher unevenly punclale wilh srvall ro Mmoderately-sized rather shallow punctures. Front and rear boarders with row of shallow small! punc- sures, Elytral striae weakly impressed especially on dise where for most part reduced toa line of punc- lures. Sina! punctures about same size as the large punctures on pronotum, Interstrial punctures sparse and shallow, those on disc smaller (usually much smaller) than those ii adjacent striae, weaker laterally, strong tendency for interstrial punetures to be arranged in one line over most of elytron, im intersiriae 3 and 5 there are some seatiered larger punctures. Venrral surface rugose-punctate. Midline of mesoslernum quite strongly and sharply carinute, weakly projecting backwards. Midline of first abdominul segment without keel. Metacoxal process raised, triangular wilh rather bowed sides, Midline weakly carinate hehind, with narrow area in middle unsculpcured. Rugose portion of metafemur 1/2-2/3 lenyth of femur, thal of mesofemur about 1/2, that of profemur a liltle over 1/2 length af femur Male: Protarsi four-segmented, basal twa segments moderarely expanded. First segment about 1.5 « length of second segment which is a little shorter than the narraw third segment, Remturks &. decipiens tsa northerm coasu species resembling B, australiae, \t differs (ram that spevies by generally being a hittle smaller and more clongate, and by the weaker and less numerous punctures on elytra and pronotum., The aedeagus and the much larger third segment of che male prorarsi separate it fram B ansiraliee. \ have fourid no characters that separate this species from B, sadieve other than the aedeagus. Distritnition (Fig, 78) Coastal northern Australia from Wyndham, W,A,, to Kuranda, Qld. Berosus sadleae sp, nov, (Figs 46, 50) Description (umber examined 2) Length 4.5-5.0mm Narrowly oval. Shiny. Elytrou nol humpbacked. Apex of elytron with two weak spines) the inner one smaller (han the outer, Dorsal surface light-brown, punctures on elylron and pronotym, elytral striae and patches on pronotum and ¢lytrou darker. Rear of head dark, centre and front lighter. Ventral surface dark-brown, append- ages lighter, rugose porlions of meso and meta- femora somewhat darker than rest of legs.. Tip of Jabial palpi dark, Head relatively narrow, relatively sparsely covered With well impressed punctures, about size of eye facet, smaller towards front. Pronotum sparsely covered with punetures, most about 1.4% puncture width apart, towards midline and in front punctures vary in size, Elytral strive weakly lo moderalely impressed, strial punctures about size of the larger ones on pronotuim, Intersirial punctures smaller. In interstriae 3 and 5 there are some scattered larger punctures. Punctures lateral to about stria 7 weak and arranged in one row, those inward of about stra 7 stronger and tend to be seartered. Stria 2 with I to 1S punctures which are about /wice as Jarge a5 these in adjacent inter strial areas, Ventral surface densely punctate. Midline of mesosternum raised, moderately pro jecied backwards, Midline of trst abdominal seg- ment weakly raised in front 1/3, Melacoxal process raised, broadly triangularly produced backwards, with srnall wide diamond shaped portion in middle devoid of sculpture. Rugose portion of metalemur 2/3 length of femur, that of mesofemur 1/2-2/3, that of profeniur 1/2 length of femur Male: Protarsi four-segmented, basal segment weakly expanded, first segment larger chan second which is sume leneth but-a little wider than third. Las| abdominal segment entire. Reimiurks A northern species Known for certain only from the iwo inale types. These are indistinguishable [rom B. deviniens excep for the distinctive aedeagus, Both species have been taken together at Howard Springs, N-T., and Lake Bennet, NC Further work a4 C1. S. WATTS is heeded to clarily the relationship of this northern group which also includes & wusfralive and B. aquilo, A series af specimens from Lake Bennet, NT. Dec. 1979, in NTM, includes both &..swcdieae and & decipiens but most are teneral which precludes. extraction of an identifiable aedeagus, hence J carol be sure to which species various individuals belong. Disiriburian (Big, 79) Known only from the wpe localiries, Types Holoiype, wale, “Howard Springs, NIL ar light, 27.1.68, E. Matthews’, in ANIC. Paratype, male ‘NT, Lake Bennett area & 25 km SE of Manton Dam, 29-30 Dec, 1979, M. B. Maliparil al MW, light’, in NTM Berosus aguila sp. nov, (Figs 34, 40) Description (number examined 11) Length 3.3-5.0 mm- Elongate oval. Elytra not humpbacked. Shiny. Apex of elytron with two spines, outer weakly lo moderately developed, inner one often lacking, Dorsal surface light-brown, punctures and striae on elytron darker, elycran with one or twa darker patches, rear of head darker Ventral surface dark-brown and black. Appendages lighler, rugase portions of femora dark-brown lo black. Head moderately covered with strong pune- jures about the saine size us facets of the eye or slightly larger, much smaller towards front. Pronotal puncttifes same size a8 those al rear OF head, rather unevenly distribuled, all approximately the same size except for a few smaller ones along extreme front inargin aid on disc. Elytral striae moderately impressed, stronger towards sides anc apex, Inter strial punctures smaller than those in striae, arranged in one line aver most of elylron, except between striae 1 to Zand 4to 5. Area bel ween striae 2 and 3 with a few larger punctures. Second elytral stria with 10 10 16 punctures which are about 3 size of those in adjacent interstrial areas, Ventral surface densely rugose-punctate. Midline of meso- sternum raised, moderately produced backwards, Midline of first abdominal segment carinate in {ront third, Metaconal process.raised, broadly triangularly produced backwards, midline weakly caripate, a sal] diamond-shaped area in centre devoid of sculpture, Rugose portion of metafermur 3/4 length of femur, that af mesolemur 2/3, (hat of prefemur 1/2-2/3 length of femur. Male: Protarsi four-seymenied, basal two segments strongly expanded, basal seament larger than second which is about same length as the unexpanded thin. Remarks Acsmall dark species from voastal NV The only smiall species in this grouping with the rupese portions of the femora darker (han the pest of the femur, and usally almost black. Distribution (Fig, 79) Known only from rhe type locality. and fram Daly River Missian, NE (ANIC). Types Holotype, orale, ‘Coastal Plains Rescarely Station, CS.LBR.O. Darwin, NT. at light, 30.66 E.C.B. Langfield’, in ANIC, Pararypes, ain saric data as holotype, seven in ANIC, two in OW, Berosus vijae sp. nov. (Figs 35, 41) Description (number examined 31) Length 3.0-4.5 mm. Narrowly oval, nat humpbaecked, apex of elytran with two weak broad spines, Yellow-brown, rear of head, areas adjacent to eyes, clytral striae, 2-3 vague patches behind imiddle of elytron, ventral surface except for appendages darker. Shiny. Head spursely punctured with variably sized punctures, sparser and weaker towards front and centre, Pronotum sparsely punc- tured with yatiably sized but quite well impressed punctures, sironger and denser laterally, front and rear margins with a single row of small punetures irregularly spaced, Elytral striae moderately to quite strongly impressed, particularly towards apex of elyiron, Punctures in striae well marked, about same size as the larger ones on pronotum. Intersrrial punctures a liule to much smaller than those in stride, arranged in single row oxcepr in interstrial 2to 3 Which has scattered large and small punctures. Second stria aboul 1/4 length of elytran with 9 to 14 punctures, Mesosternal keel weakly raised. Meta- coxal provess Iriangular with apex towards rear, a small approximately oval area in midline devoid of sculpture. Ventral surface, rugose-punctate with smallish punctures. First abdominal segment with weak midline keel in front 1/4, Rueose portion of metalemur 3/4 length of femur, that of mesofemur 2/3-3/4 that of profemur about 1/2 on frant edge and a little greater on hind edge, Male: Protarsi four-segnented. Mirst three segarents subequal in length, lirst moderately expanded, second rather less 50 and third net at all. Apical abdominal segment very weakly norehed with weak protruberence at each side of woteh. Remarks A small widespread porthern species closely resembling & ralph| and & veronicue, separated by range of Characters given in Table 2, Generally with REVISION OF BEROSUS larger elyival pluretures and more strongly intpressed striae allhough (hese are weakly impressed on (he dise in some specimens. Distribution (Fig, 75) Northern Australia, predoyninantly che western half, Types Holorype, male, “Tindal, NT 14=3PS, $32°22'R. 1-20 Dec, 1967, light trap. W. J. M- Vesijens*, in ANIC. Paratypes, four same data as holotype, in ANIC. Berosus ralphi sp. nav, (Figs 72, 73) Deseviptian (number examined 8) Length 3.54.35 mm, Oval, Not humpbacked, Apex of elytron iruncat or very weakly produced into Iwo broad short spines. Yellow-brown, purictures on elytron and a few small spats in apical 1/2 of elytron darker, Ventral abdominal segments orher (han upical are blotched darker. Shiny, Eltron alten moderately reticulate, Punetures of head moderately impressed, sparse, smaller towards [root and venter. Punctures of pronotum moderately impressed, sparse, variable ii size, With Yow of small punctures along extreme lront and rear margins, Elytron striae moderately to weakly impressed, Punctures on striae close, well fmopresscd, ubout size ol larger pronotal punctures, Unterstrial pupetures much Smalley than those in strlacand yeattered (in iler- sirtae One lo five, strong tendency to be arranged in a single row in interstriae lateral to striu 6. Interstria 3 with punctures of Uwe sizes. Stria 2 short, often poorly marked with 7 to 13 punctures. Mesosternal keel inoderately raised. Metavoxal provess, sharply trangularly produced backwards and downwards, a small slightly raised area iu midline in centre. Ventral surface rugose-punctate. First abdominal sezment with raised midline keel in tront L’4, Rugese portion of metafemur 3/4 length of femur, thal On mesalemur 1/2-2/3, thal on prolemmur t/4-1/2 length of femur. Male; Apical abdominal segment sinuate on hmd margin. First (wo segments of protarsi expanded, third segment not expanded, Firsi three seements progressively smaller in length, Remurks Similar in general facies to & vijae and B veronicae bur differing fron then) in the wrearer number and smaller size of the punctures in the first elytral juterstriae, Where they are arranged in more ihan one line, and in chargeters of the aedeagus. In all specimens so far examined the tip of rhe nietacoxal process is sharply deflexed downwards, 4 character not seen in other Australian Berosus. fh w Distribution (Fiz, 77) Coustal porth-westen) Australia and Queensland. Types Halotype, male, ‘Wyndham W.A, S. Stephens 20,2.01', in SAM. Paratyes 17; seven same data as hololype, i S.A.M, one ‘Derby NWA, W. DL Dodd’, in SAM; seven ‘Derby WIW.A,’, in SAM? two ‘Queenstand J.8261', in SAM. Berosus subavatus Kaisch (Pigs 36, 42) Berosus stictieus Fairmaire, 1879, yp. 82. Berosus sibovaTey Kuisch, 1924 (not Boheman, 1859) Type Holotype, female, Peak Downs NT, in MNHN, seen, Descriplian (number examined four) Length 4.6-3.4) mm. Elongate oval. Elytra not humpbucked. Shiny. Apex of clyiron rounded o1 weakly truncated, Dorsal surface light-brawn, punctures aud striae on dise of elyrron and vague patches on head, pronotum and elytron darker. Punetures in lateral striae on elytron with large squarish darkish subsurface markings. Ventral sur face dark-brown to black, appendages lighter, rugose porhions Of lemiura wilh a slightly darker hue thar rest of leg, Head with scaltered relatively small punctures, (hose at back of head a little larger than size of eye facet, those towards ventre and front of head smaller, Pronotum éparsely and unevenly covered wilh punelires, of twa sizes, larger aboul same size aS those ai rear of head, Elytral striac weakly impressed, over most of elytran little more than series of punetures. Second elytral stria with 6 to 7 punctures, about same size ora little smaller than the larger ones of pronoium 2 © size of those in adjacent interstrial areas, Punctures in interstriae 3 and 4 of two sizes. Inorerstrial puuctures lareral ro stria 6 arranged iu one line, Thase towants sides and apex small and weakly impressed. Ventral sure face finely rugose punctate. Midline of mesoster- num moderately and evenly raised, moderarely projected backwards. Midline of first abdominal segment weakly carinate in front 1/4, Metacoxal process raised, trlangularly produced backwards, midline carinate in rear 1/4 @ Targe roughly diamond-shaped area in middle, devoid of seulp- ture. Ruwose portion of metafemur 2/3 length of femur, that of metafemur 2/3, that of profemur about 1/2 length of femur Male; Protarsi four-segmented, basal two segments moderately expanded Basal segment larger than second which is larger than third. 26 tH & WANS Renwrks The relatyely Small pumiber of punctures in the sevond elyiral stria. strong lendency for punctures in literstriae to be arranged in one ling, and sparse and variably-sived pronotal and head punctures separate this species from related species, B vijae and B, raiphi are dit ficult to definitely separate trom this species apart from the form af the aedeagus which is vousiderably shorter than the parameres in B subovetus, Distribution (Fig, 814 Known from ouly two localities: Station Creek in north Old. (in ANIC), and Katherine, ST. fin ANIC). (The type is labelled s\ustral’) Berosus nicholasi sp. nav, (Figs 66, 69) Description (number examined $1) Lengih 5,0-6.5 mm. Glongate oval, bly(ra not humpbacked. Apex of elytron weakly extended wilh two small spines, inser one often small or lacking. Shiny, often moderately reticulate with a very fine even reticulation. Dorsal surface light-brown with punctures, elylral stride and patches on head, pro- notum and elyiron dark-brown, Ventral surface dark-brown to black with lighter movtlings. Appendages brown, tip of labial palpi darker. Houd relalively narrow, evenly and quite densely covered with slrang punctures, punctures weaker towards front. Pronotim moderately bul unevenly covered with strongly impressed punctures of 1wo general sizes with the larger ones predominating, rows of small punctures along front and rear margins. Elytral striae quite strongly impressed except towards sides and back. Srrial punctures relatively small, a little smaller than the larger ones on the pronotum. Interstrial punctures small, relatively sparse, those laleral (o stria 8 arranged in-one line, amaller than strial punctites aver most of elytron, Some of the more central interstriae with a few large punctures with shorl setae. Ventral surface densely rugose-punctare, Midline of mesosternum raised, moderately projecting backwards, Midline of first abdominal segment carinate only in extreme front, Metacoxal process raised, triangularly produced backwards, small elongate Oval area in centre devoid of sculpture. Rugose porrion of metafemur 2/3-1/4 length of metalemur, that of mesofemur 1/2-2/3, that of profemur 1/3-1/2 length of femur. Male: Protarsai four-segmented. Basal segments subequal, quite strongly dilated. Apical abdominal segment not notched. Remarks : One of a group of very similar northern specie which can only be confidently separated by characters of the acdeagys, Ip all but a few cases, jarger than other specles in the group ocher than B, dehilipennis, which usually has much sironger puncrures On the elytron, See Table 2, Distribution (Fie, 79) Coastal northern Australia from Derby, W.A. To Tawnsvifle, Qld, Types Holotype, male 12° 25'S 131'03'R Howard Springs. NC 24 ki S of Darwin 10.41.72. rainfovest, at light, E. Britton’, in ANIC. Paratypes, 12 same data ax holotype; L) in ANIC, one in CW. Berasies dehilipennis Blackburn (Figs 44, 45, 49, 51) Berosus dehilipernis Blackburn, 1898, pr. 223. Tivpe Holotype, temale, Cape York, Qld, in BM(NEL), seen. Description (number examined 74) Length 4.0-6.0mm. Klongate oval. Elyira nor humpbacked, Apex of elytron rounded or with (wo weak spines. Dorsal surface light-brown with pune- tures on pronotum and elytron, elytral striae and patches on pronotum and elytron darker, Rear of head dark-brown, centre and front lighter. Ventral surface dark-brown, appendages hghter, lip of labial palpi dark-brown, Mead wiilt sparse to moderately denise, strong purictures, a little larger than eye facets, grading 10 obsolete towards front. Pronotiun rather unevenly bur moderately covered with strong punctures the size of hose at rear of head, a few smaller punctures towards midline and front, Blytral striae quite strongly impressed parucularly laterally and towards rear. Second elytral striae with 10 to 18 punctures, 2x the size of those in adjacent inter- strial areas, Interstrial punctiires scattered ner arranged in a single row except in same places towards sides and rear, A few large punctures in interstfiae 3 and 5, Veritral surface densely rugose- punctale. Midline of mesasternum weakly raised, moderately projected backwards, Midline of first abdominal segment weakly carivate at extreme front. Metaucoxal process raised, narrowly triangue larly produced backwards, with a narrow diamonel. shaped area in. middle unseulptured. Rugose partion of imetafemur 2/3-3/4 length of femur, that of mesofenyur 1/2-2/3, thal of profeniur [/3-1/2 lenath of femur, Male; Protarsi four-segmented, First and second seymenrs moderately expanded, lirst much longer than sceand, which is about the same length as third, Apical abdominal segment broadly bur yeey weakly notched, REVISION OF BEROSUS 27 Remarks A widespread northern species which I initially con- sidered to be composed of at least two species. Further study may show that it inchides several closely related species. In particular, individuals can be placed in one of two groups according to the shape of the aedeagus. One group has the parameres thin and the aedeagus roundly hooded at the tip, the other has thick parameres and has a transverse ridge on the upper surface behind the roughened end portion of the aedeagus, These characters are to some extent variable and intermediates exist, eg. notably a specimen from Cunnamulla, Qld., in SAM. This species can be separated from the rather similar B micholasi by the usually much stronger elytral punctures and the relatively smaller second segment of the male protarsi (Table 2). Distribution (Fig. 80) Coastal northern Australia from Derby to Caims region. CHULCKLIST OF AUSTRALIAN BEROSUN (in alphabetical order) 8. australiaeg Mulsant = externespinosus Fairmaire = gravis Blackburn B. amoenus sp. nov. B. arcus sp. nov. B, approximuans Fairmaire = B. auriceps Blackburn B. blackburni Zatz = B. avipennis Fairmaire - B. simulans Blackburn ~ B. stigmaticollis Fairmaire aquilo sp. nov. dallasae sp. nov. debilipennis Blackburn decipiens Blackburn discolor Blackburn ~ B, flindersi Blackburn duplopunctatus Blackburn egibbae sp. nov. invalutus (W. MacLeay) Josephenae sp. nov. B. juxtadiscolor sp, nov, l Reah boB&aS ACKNOWLEDGMENTS The Curators of the collections listed earlier are thanked for the free and rapid access to their collections afforded to me. Dr E. Matthews kindly read portions of the manuseript and greatly improved it. Mrs P. Kidd’s expert typing of the various drafts is greatly appreciated as is Miss J, Thurmer’s drawings of elytrae and other structures, Mrs. Marianne Anthony, Librarian of the S.A. Museum, ferreted out obscure references without which progress would hitve been much slower . mlacropunciatus Sp, NOV, } macumbensis Blackburn . majusculus Blackburn munitipennis Blackburn nicholasi sp. nov. niger 8p. nov. nutans (W. MacLeay) = B. pallidulus Fairmaire B. pulchellus W. MacLeay — B, devisi Blackburn 8. queenslandicus Blackburn = B, quartinus d’Orchymont quadrapunclatus sp. nov. ralphi sp, nov. reardoni sp. nov. Sadieae sp. nov. suboyatus Knisch = 8B. sticticus Fairmaire (rishae sp. nov. timmsi Sp. Noy. } veronicae sp. nov. . vijde sp, nov, Dehra nan PaREtBe PRRDHA REFERENCES BLACKBURN, T, [888 (1889), Notes on Australian Coleoptera with descriptions of new species. Proc. Linn. Soc. N.S.MO (2/1011, 1S88(1889); 805-875. BLACKBURN, T. 1889 (1890). Notes on Australian Coleoptera, with descriptions of new species. Prove. Linn, Soc. N.S.W. (2)1V: 445-482. BLACKBURN, T. 1898. Further notes on Australian Coleoptera, with descriptions of new genera and species. Trans. R. Soc. 8. Aust, XXII, 1898: 221-223. BLACKBURN, T. 1896. Coleoptera (exclusive of the Carabidae) in Report of the Horn Scientific Expedition 28 in ALES: to Central Australia, Part Il — Zoology: 254-263. Melville, Mullen & Slade, Melbourne. BLACKBURN, T. 1895. Further notes on Australian Coleoptera, with descriptions of new genera and species. Trans. R. Soc. S. Aust, X1X: 27-60. WORCHYMONT, A. 1925. Contribution a l'étude des hydrophilides II. Annals Soc. Ent. Belg. LXV: 139-169. d’?ORCHYMONT, A. 1943, Neue oder interessante Sphaenidiien und Hydrophilinen der Malayischen region. Treubia Ill: 416-421. d@ORCHYMONT, A. 1943. Notes sur la tribus Berosini Bedel (Coleoptera Palpicornia Hydrophilidae). Bull. Mus. Hist. Nat. Belg. 19(42): 1-12. FAIRMAIRE, L. 1879. Déscriptions de Coléopteres WATTS nouveaux ou peu connus du Musée Godeffroy (Hydrophilidae). J. Mus. Godeffroy XIV: 80-83. KNISCH, A. 1924. ‘Col. Cat. Part 79: Hydrophilidae. 304 pp. Junk, Berlin. MAcLEAY, W. 1825. New Coleoptera from Java. Annul. Jay, 1825, 35. MacLEAY, W. 1873. Notes on a collection of insects from Gayndah. Trans. Ent. Soc. N.S.W. 11: 79-205. MULSANT, 1859. Australian Hydrophilidae. Opuse. Ent. IX: 58. ZAITZ, 1908. Catalogue des Coléoptéres aquatiques des familles des Dryopidae, Georyssidae, Cyathoceridae, Heteroceridae et Hydrophilidae. Horae Soc. Ent. Ross. XXXVIII, 1908; 282-359. NABIDAE (HETEROPTERA) OF VANUATU BY I. M. KERZHNER Summary Six species are listed, one of them (Stenonbis nitidicollis Kerzh.) is new from the Vanuatu fauna. Keys to adult Nabidae and to fifth instar larvae of Arbela occurring in Vanuatu are given. NABIDAE (HETEROPTERA) OF VANUATU 1, M, KERZHNER KER/ZHNER, |, M. 1987, Nabidue (Heteroptera) of Vanuutu, Rec, S. Ausr, Mus. 211) 29633, Six species are listed, one of them (Srenonabis nilidicollis Keren.) is new from the Vanuatu fauna. Keys to adult Nabidae and {o fifth instar larvae of Arbela occurring in Vanuatu are given. 1. M. Kerzhner, Zoolowical Institute, Academy of Sciences of the U.S.5.R., Leningrad 199034, U.S.S.R. Manuscript received (7 Pebruary 1986, Previous records of Nabidae from Vanuatu (formerly the New Hebrides) were based on the material in the Museum National d'Histoire Naturelle, Paris (Reuter 1908); the British Museum (Natural History), London (Harris 1938, 1939; Kerzhner 1970a); and the Universitetets Zoologiske Museum, Copenhagen (Kerzhner 1970b), In all, five species have been recorded. The material received from the South Australian Museum, the B. P. Bishop Museum and the U.S. National Museum of Natural History (thanks to the kindness of Dr G. F, Gross, (he late Dr J. L. Gressitt and Dr Th. J. Henry) contains all of the species already known from Vanuatu and one species new for the fauna. In the distribution lists below, only localities and collecting dates are mentioned, The data on expeditions, collectors and deposition of material for corresponding years are as follows: 1943 — collector Knight} deposited in the U.S. National Museum of Natural History, Smithsonian Institution, Washington, D.C,, USA; 1950 and 1970 — colleetor N. L. H. Krauss; deposited in the B. P. Bishop Museum, Honolulu, Hawaii, USA; 1957 — collector J, L. Gressitts the same museum, 1958 — collector Borys Malkin; the same museum; 1960 — collector W. W. Brandt; the same museum 1964 — collector R, Straatman; the same mMuUscUmm; 1965 — Biospeleological Expedition, collector G, P. Gross; deposited in the South Australian Museum, Adelaide, South Australia; 1967 — collectors J. & M. Sedlacek; deposited in the B, P, Bishop Museum; 1971 — Royal ‘Society and Perey Sladen Expedition; collectors — P. Cochereau (on Malekula J.), G. F. Gross (on all other islands), Masing Andrew (who assisted G, F, Gross on Tanna Jj); deposited in the South Australian Museum, but duplicates from large series will be sent 10 British Museum (Natural History), Museum National d'Histoire Naturelle, Paris, or will be retained in Zoological Institute, Leningrad; 1973 and 1976 — collector N, L, H. Krauss; deposited in the U.S, National Museum of Natural History. SYSTEMATICS Gorpis simillimus Harris (Figs 1-3) Previous records MALEKULA, Harris 1939: 150-I51. Material examined ESPIRITU SANTO: Apouna R. Camp 2, 146 m, 30.VIU,1971, at light, 1G) above Namatasopa, 400 m, 30. VILL.1957, 1 9; Hill EB, of Luganville, 100 m, Macaranga, 10.1X.1957, 1 o. General distribution Vanuatu (type locality) and Solomon Islands. Remarks Previously (Kerzhner |970a) | mentioned that there were some colour differences between specimens from Vanuatu.and the Solomon Islands. These differences are in the main confirmed by the new mutterial, In addition, shght differences in male and female genitalia (Figs 1-3) have been found. However the material at hand is insufficient for a study of individual variability and therefore | refrain from description of a new subspecies for the Solomon Islands population. Arbela immista Harris Previous records MALERULA. Harris 1938; 579; Kerzhner 1970a: 298; 1970b; 192, 0 L M. KERZHNER FIGURES 1-3, Gorpis simillimus Harris, 1, male from Vanuatu (Espiritu Santo), aedeagus; 2, female from Vanuatu (Espiritu Santo), vagina in ventral aspect; 3, female from Solomon Jslands (Guadalcanal), the same. Material examined ESPIRITU SANTO: Baldwin Bay, 17,VILL, 1958, } 2; 25 km NE Luganville, 12.1V.1964, 1 of, 1 9. PENTECOST (NE): 200-500 m, 27, 111.1964, 1 o. MALEKULA: Amok, 17.1X.1958, 1 0% South West Bay, 11 and 13.X,1971, by beating trees and sweeping grasses in the forest and along the river of the forest, 9 of, 10 9, 4 larvae, General distribution Vanuatu (type locality) and the Bistmarck Archipelago, Arbela costalis Stal Previous records BANKS, MALEKULA, ERROMANGA. Harris 1938; 581; Kerzhner 1970a: 298. Material examined API (‘Epi’): Vaemali, 10,VIII.1967, 1 oc. ERROMANGA: Ipota, 0-100 mi, March 1970, 2 @; Ipota, vicinity of Ipota, and River Camp between Ipota and Nuankao, 4, 5, 7, 12. VII1.1971, 4 o, 3 9. TANNA: Lenakel, 0-150 m, March 1970, 1 9; between Lenakel and Bethel, 28.VIL1971, 1 larva), ANEITYUM: vicinity of Analgahaut, 19.V11.1971, 1 o, 1 larva. General distribution Fiji (type locality), Samoa, Vanuatu and Solomon Islands, Arbela nitidula (Stal) Previous recards MALEKULA., Reuter 1908: 127; Harris (938: 568, Material examined ESPIRITU SANTO: Segond Channel, Santo, Aug. 1950, 2 9; Namatasopa, 300 m, 28.VII1.1957, 1 9; below Namatasopa, 250 m, 2 and 3.1N.1957, 1 o, 1 9; Tasmalum, 3 m, bush, 4.1X.1957, | or; Luganville, 23-28, V1I.1958, 1 or; Baldwin Bay, 28-30.1X.1958, 1 9; Narango, 90 m, May-June 1960, 2 9; Apouna R. Camp 2, 146.3 m, 2, 3.[X.1971, 10%, 7 9, 3 larvae; Malao Village, Big Bay Area, 23.VIII.1971, | larva. MALEKULA: Amok, 17,1X.1958, 3 o, 4 9,1 larva; South West Bay, 13-14.X.1971, beating small trees and sweeping grasses along the river to the forest and on swamp, 4 oc, 2 9, EFATE: Maai (Mat, Ambryn Vill.), 3 m, 1S.VIIL.1957, 1 9; Limestone, Plateau N of Maat, 100 m, 18, 20. VIIL.1957, 2 0; Vila, 0-100 m, Jan. 1976, 1 @, ERROMANGA: Ipota, 5.VIII.1971, 3 9, TANNA: between Lenakel and Balhel, 28.VIL1971, 1 o, 2 9. ANEITYUM: vicinity of Analgahaut, 18-20.VI1,1971, 3 ©, General distribution From India and south China to Vanuatu, Stenonabis nitidicollis Kerzhner (Figs 4-6) Material examined ESPIRITU SANTO: Nokowoula, 1100 m, 14.1X.1971, by sweeping, 1 brachypterous female, General distribution Australia, Vanuatu (new record). Remarks S. nitidicollis was described (Kerzhner 1970a) from a macropterous female, collected in New South Wales, Mrs N. Strommer (formerly N, Winkler), now in Heathmount, Melbourne, kindly informed me that she has examined macropterous and brachypierous males and females of S. nitidicollis trom Queensland. | compared the NABIDAE OF VANUATU 3 female (Fig, 4) from Vanuata with the holotype of S. nitidicollis. The yagina in the Vanuatu female (Figs 5, 6) is slightly larger (width 1.00 mm, in the holotype 0.93 mm), the dark coloration of the body is more pronounced and the hind lobe of pronotum has an intermediate longitudinal brownish stripe on each side berween the medial and lateral stripes, the antennae are longer (length of the first segment 0,93 mm, of the second 1.33 mm), and also the legs and rostrum are slightly longer. All these differences do not surpass the level of individual and geographic variability in related species. In the Vanuatu specimen, head width 0.76 mm, yertex width 0.34 mm, hind lobe of pronotum length 0.43 mm, width 1.29.mm, scutellum of equal length and width, body length 6.7 mm, width of abdomen 2.0 mm. Nabis (Tropiconabis) kinbergii Reuter, 1872 Nabis nigrolineatus (Distant, 1920). Nabis tasmanicus Remane, 1964. Nabis capsiformis auctt. (non Germar, 1838), part, Previous records ‘NEW HEBRIDES’ (as Reduviolus capsiformis). Reuter 1908; 114, TANNA (as Nabis nigrolineatus). Kerzhner 1970a; 355. FIGURES 4-6. Stenonabis nitidicollis Kerzhner, brachyplerous female from Vanuatu. 4, dorsal aspect; 5, vagina in dorsal aspect; 6, vagina in ventral aspect. 32 1M, KERZHNER Material examined ESPIRITU SANTO: tho exact locality, Oct, 1943, 1 @; Segond Channel, Santo, Aug. 1950, 1 ©; Luganville, 20, 23-28.VIL1958, 17 o,2 9, 4 larva; 10 km W. Luganville, JOJ1, 1964, 1 9) Santo, 13, 14.X 11.1965, 1 o, 19; Malao Village in Big Bay Area, 28VIILI97I, 1 9, MALEKULA: Amok, 17.1X.1958, 1 9, | larva; Lakatoro, 16-17,%, 1971, 1 o, EPATE: Vila, Aug. 1950, 1 ©; Vila, 0-100 m, Feb. and March 1970, | o, | ©; Vila, 0-200 m, Feb, 1973, 1 ©; Vila, 0-100 m, Jan. 1976, 2c; SE corner, JONVILA971, 1 oy 1 3, Plantation Gaillarde nr. Tagobe, 11. VIL.1971, 1 9, ERROMANGA: IL km W. of lpota, 100-200 m, Feb. 1970, L ony tpota, 0-100 m, Mareh 1970, 1 o; Ipota and vicinity of Ipota, 5, 6, 10, 12. VII.1971, 5 o&, 9G, TANNA: East Coast, 450 m, 8.011.1964, 1 o, | ©, Some specimens are taken at light. General distribution Australia and Pacific islands. The most remote records are Ryukyu and Bonin Islands, New Guinea, Samoa, Society Islands and New Zealand. Remarks The species was known formerly as N. tasmanicus and then as N. nigrdlinealus. However some doubts existed on {he last name because W, L, Distant described the species in Reduviidae (genus Sestrapuda) and mentioned a number of characters not appropriate to Nabidae. | examined the type series of N. kinbervii (Naturhistoriska Riksmuseet, Stockholm), which included | female from Sydney, belonging to N. nigrolineatus, and 2 females from Buenos-Aires, belonging to N, cupsiformis. | designated the specimen from Sydney as the lectotype of N, Ainbergii and this name should be used for the species occurring in Australia and surrounding islands (Kerzhier 1981; Woodward & Strammer 1982). KEY TO ADULT NABIDAE OCCURRING IN VANUATL Note, The key is intended only for determinaiun of Vanuatu! material, therefore same ol the characters mentioned are not applicable to all species of included genera nor (o all populations of the included species, 1, Fore coxal cavities closed behind. Fore coxae slender, greatly clongated, Fore tibiae curved, shorter than fore femora... 2... -. 0. Grorpis Stal One. species _ GC. simiflimus Harris — Fore conal cavities open behind, Fore caxae nearly canical, less than twice as Jong as (hick, Fore libiae straight, subequal in lengtl) to the fore femora... it 2 2. Nore and middle femora and tibiae with long slender spines, Ocelli vonliguous lrhela Sifl (see 3) —~ Femora and tibiae withouw ae spines. Ocelli well separated . . ee a oe a, Oe Ae, 3. Pronotum dirty yellow wilh (wo longitudinal brown ar black stripes, seldom completely light coloured. Hind lobe of pronotum not dull, bur Horse strongly shining as the fore lobe, Hind tibiae of lhe male wilh a densely pilose sub-basal thickening «0... 4. mit/dula (Stal) — Promotum (except in distinctly toneral specimens) complercly black or wilh only corners or sides of the hind lobe yellow. Hind lobe of pronotum either velvety dull, or as strongly shining as the fore lobe. Hind tibiae of the male without thickening, ...... 4... 000.04 4. Hind lobe of pronetum cull. vipepo ced. Immista y Harris - Hind lobe of peonotum strongly sev afl Os nfotefeten..ecfry-ommatet-tetam " jas LAL castalis sia! 5. Hind lobe of pranotum punctured. Connerxivum below not separated from the venter by a suture. Short- or long-winded. -----..,))......Slenonabis Reuter One species — S$. nitidienllis Kershner - Hind tobe of pronotum without punetures, Connexivun) below separated trom ihe yenter by a suture lying in a deep impression. Long- winged . One: species ~ N. Kinbergii Peder Kiy TO FLETH INSTAR LARVAE OF ARRBEI 4 OCCURRING IN VANL ATL |. Wing pads black wuh the cai third white - esas t0 por. ' --A immista Harris — Wing pads wnfealorous , REREAD SEER RIOCE Per- 2. Wing pads black . 4 hires 4 Costalis Stal — Wing pads light votoured. - A. nitiduta (Sia ACKNOWLEDGMENTS Iam very thankful to Dr G, PF, Gross, to the late Dr J. Le. Gressitt and to De Th. J. Henry for the interesting material and to Dr N. Strommer for the information on Stenonahis nitidicollis.. REFERENCES HARRIS, H. M, 1938.. The genus_4rhe/a Stal (Hemiptera, Nabidae). Man, Mag. rat, Hist, (ser, 11), te 561-584. HARRIS, H. M. 1939, 4 contribution ta our knowledge of Gorpis Stal (Hemiptera: Nabidae). Philipp. J. Sev. 69(2): 147-155. KERZHNER, |. M. 1970a, Neve und wenig bekannte Nubidae (Heteroptera) aus den tropisehen Gebieten der Alten Well. Acta ent. Mus. nain. Pragae 38 (1969); 279-359. KERZHNER, I. M. 1970b. Some Heieroptera Nabidae NABIDAE OF VANUATU 33 (Hemiptera) from the Southern Philippines and the © WOODWARD, T. E. & STROMMER, N. 1982. Nabis Bismarck Islands. Ent. Medd. 38: 117-194. kinbergii Reuter, the current name for Tropiconabis KERZHNER, I. M. 1981. Fauna SSSR. Nasekomye nigrolineatus (Distant), and its Australian distribution khobotnye, t. 13, nr. 2. Poluzhestkokrylye semeystva (Hemiptera: Nabidae). J. Aust. ent. Soc. 21: 306. Nabidae. Leningrad. 327 pp. (In Russian.) REUTER, O. M. 1908. Bemerkungen iiber Nabiden nebst Beschreibung neuer Arten. Mém. Soc. ent. Belg. 15: 87-130. INTRODUCTORY STUDY OF ADVANCED ORIBATE MITES (ACARIDA) : CRYPTOSTIGMATA : PLANOFISSURAE) AND A REDECSCRIPTION OF THE ONLY VALID SPECIES OF CONSTRICTOBATES (ORIPODOIDEA) BY D. C. LEE Summary The study of advanced oribate mites (Planofissurae, new name) is introduced as a further part of an ongoing study of sarcoptiform mites from South Australian surface soils. Morphology is considered with reference to a unified notation for hysteronotal chaetotaxy, notal pores, the form of leg trochanters, acetabula and apodemes. Constrictobates lineolatus Balogh and Mahunka is redescribed from South Australian material, the generic diagnosis 1s modified, Constrictobatinae (Fenicheliidae) is newly synonymised with Pseudoppiinae (Oribatulidae), and the superfamily Oripodoidea is considered. INTRODUCTORY STUDY OF ADVANCED ORIBATE MITES (ACARIDA; CRYPTOSTIGMATA: PLANOFISSURAK) AND A KEDESCRIPTION OF THE ONLY YALID SPECIES OF CONSTRICTOBATES (ORIPODOIDEA) D, C. LEE LEE, D. ©, 1987, Introdictory study of advanged oribate mites (Acarida> Cryptostigmata: Planotissurde) and a redeseription of the only valid species of Cunsirictabates (Oripodoidea). Ree, & Aust Mus. 211): 35-42, The study of advanced oribate miles (Planotissurae, new name) is intmodticed asa further part of ant ongoing study of sareoptiform mites from South Australian surface sails, Morphology is considered with reference to a unified notalion for hysteranotal chaetotaxy, notal pores, the form OM ee trochanter’, adetabula and apodemes, Constricrabates lineolarus Balogh and Mahunka isvedesenbed from South Australian material, the generic divgnosis is modified, Constrictobatinae (Penicheliidaed is newly synonymised with Pseudoppiinae (Oribalulidae), and (he superfamily Onpodajded is considered. Bb, C. Lee, South Australian Museum, North Terrace, Adelaide, South Australie 5000, Manuseript received 2] October (986, This is a further part of an ongoing study of sarcoptifarm miles from surface soil sampled from nine florally diverse South Australian sites. The primitive oribate mites have been considered elsewhere (Lee 1981, 1982, 1985), and here the study of advanced oribate mites {is introduced, The majority of the oribate mites sampled belong to this group which, because of morphological changes, requires a consideration of homology and notavon. Furthermore, because a new diagnostic character stale for these miles is recognised, and (he opinion that they should be unnamed (Lee 1985) is revoked, they are rediagnosed and dealt with under a new name (Planotissurae), The description of the primitive oribate mites in this sludy has been thorough but time- consuming. Balogh & Mahunka (1983) sugges! that ‘painstaking scrutiny, using some recently discovered features’ is not worth doing for only some members of a genus, Whilst appreciating this point, the paucity of a cammon denominator description is so limiting for many oribate mite groups thal a more subsranual level had to be undertaken, but gol to such an extent as in my previous work. The dorsal and ventral aspects af the sonta and the shape of the leg segments have been desvribed, but not the gnathosternum or the chaelotaxy and lorm of the hairs on the legs. The first superfamily to be considered is the Oripodoidea, partly because \t is a diverse and dominant group within the well-established Poro- hotae, atid partly because i is not only important in the study of soil zoology, but some of its members are intermediate hosts of anoplocephalid tapeworms, being infected by the cystercerooid (bladder worm) stage, The most recent work on the Oripodoidea is by Balogh & Balogh (1984), referring to it.as the “Oribatuloidea’ as well as excluding the Mochlozeidae and Parakalummidae, It includes 20 families in the superfamily, of which hall are listed ag new, The work gives great importance to whether or nor the hysteronoral foramina are nulliporose, sacculate or a mixture of both those character states. The keys and diagnoses use few character’ states, which for Constrietobatinae (the only family group so far considered) are in part inaccurate. Despite this, the work is valuable on the basis for studying oripodoids, Constrictobates was selected as an example, because it is unique within the Oribatuloidea in having |5 pairs of hysteronotal setae on the adult, This is only one pair less than in the holotrichous stale amongst ‘primitive oribate mites and is therefore valuable in homologizing the hystcronotal chactotaxy of the two groups. The South Australian mites examined are deposited in the South Australian Museum; the types have been returned to the Hungarian National Museum. MORPHOLOGY flystéronotal chaetotaxy There are three regularly used notational systems for ihe hysteronotal chaetotaxy, depending on whether they have a full complement of either 16, 1§ or 10 pairs of setae, The multiplicity of systems is based on uncertainties of homology, | introduced another system (Lee 1981) for the primitive oribate mites (16-pairs chaetotaxy) with the intention of applying it 1o all sarcoptiform mites, The chaetotaxy Ab DC. LEL of oripodoid mites, for which both the [5-pairs and 10-pairs chaetotaxies huve been used, is homologized here with the l6-pairs system. as illustrated (Figs |, 2). Because most advanced oribate mites have 15 pairs of hysteronotal setae on the trtanymph, both systems have sometimes been used for one species, Certainly, with the substantial vhange in form between. the nymphal and adalr stages, Any proposals of homology are uncertain, On the other hand, | consider that the heuristl advantage of a uniform notation oulweighs the disadvantage of using an uncertain homology, Previously (Grandjean 1954, Lee 1984) ir has heen stated (hat all Planoligsurae (= Circumdehiscentiae) lack seta J4 ( ‘islands’ <----=-------------- =~ == -------- ‘Saltwater country’ ------------------------------------ > ‘mainland’ ‘land’ wambalda INLAND, COAST AND ISLANDS 73 FIGURE 2. The physical environment in the coastal mainland area (Ganggalida country). The long sand ridges parallel to the coast-line are evident from the lines of vegetation. The area of saltpan in the background separates the coastal strip containing ‘islands’ (‘saltwater country’) from the ‘mainland’, in the Aboriginal perspec- tive. The saltpan and mangroves in the foreground occur only occasionally along this stretch of coastline. (Courtesy of Connah/Jones Aerial Archaeology, University of New England,) conceived (in general terms at least) as containing areas equivalent to the close off-shore islands.® However, this narrow coastal strip should not be considered as socially isolated in any way. Figure 3 indicates quite precise boundaries for the coastal estates, to the east and west; the boundaries are hence the sea to the north, and creeks and salt-arms to the east and west. But the boundedness to the south (i.e. in an inland direction) is much less precisely defined. All estates except one in the area shown on Fig. 3 extend across the saltpan to the ‘mainland’ to include one or more fresh waterholes. In the case of the exceptional estate (F), it includes site Fl (Gunamula), a large fresh water lagoon near the mouth of Cliffdale Creek, which is said to contain surface water for a substantial part of the dry season. Seasonal movements across the saltpan to waterholes, occurred particularly during the middle and late stages of the dry season, not so much because of a lack of fresh water in ‘saltwater country’ (for this is said to have been always available by digging ‘soaks’), but in order to obtain certain material resources apparently not at that time readily available on the ‘islands’ (e.g. water lilies, Nymphoea sp. and Nymphoides sp.). During parts of the dry season people are also said to have come to the coastal strip from a considerable distance inland; however, they apparently rarely remained there during the wet season because of their lack of tolerance of the increased number of mosquitoes then. The significant point here is that apart from the recognition of the environmental distinctiveness of ‘saltwater country’, people are said Example of named island -—--—>|sland" - vicinities “Wambilbayi Island” FIGURE 3, Ganggalida and Gananggalinda coastal estates showing the Aboriginal designation of the inland limit of the saltpan areas as the beginning of the ‘mainland’ (wambalda). Certain fresh water holes on the ‘mainland’ are shown for each estate. 4 DS TRIGGER 10 have moved in and out of it constantly, Nevertheless, it was viewed as the distinctive domain appropriate to 3 Conceptually separable group of ‘sullwaler people’. KaysHip AND SOCIAL ORGANISATION An Aranda (ype kinship system appears lo fave operated traditionally throughout the study regien. See Trigger (J985} for some discussion of its natitre, and also comments by Warner (1933; 68), Sharp (1935: 160-161) and Hale (1982). Evans (1985: 17-18, 48)-492) elaborales the Gayardild system of Bentinck Island and it is clearly of the same type as both others throughout the Wellesley Islands and also as Ganggalida, Garawa and Waanyi on the mainland; Tindale’s failure to consider the \ssue of this similarity (1977; 258-260) again renders his general suggestions about the lengthy isolation of the Bentinck Islanders jnadequate. Evans (L985: 21-22) in fact suggests “a relatively short period of isolation’ of the Gayardild (500 to L000 years), based partly on this matter of kinship system similarity, However, Tindale (1977: 258-259) does refer correctly to 1he marked lack of any system of tamed sovial categories (or ‘class’ system as he puts it) on Bentinck [sland in comparison with elsewhere throughout the region, though his expression of this fact Is at best imprecise in implying that the surrounding groups had ‘sections’. Evans’ expression of the comparison also lacks precision in suggesting that the Mornington Islanders and those on {he adjacent mainland had moieties and sections (Evans 1985: 17). What has operated throughout the region, except on Bentinck Island, is a system of named subsections (Trigger L985; 69-71, 350-340), though this system can be said to be ‘organized as unnamed patrilineal semi-moieties and moieties’ (Sharp 1935: 159; 1939; 455), Putting aside the distinctiveness of the Bentinck Islanders in this respect, and without discussing social organisation in detail here, we may simply note the situation of the coastal mainlanders (Ganggalida), They were included within the mainland blov stretebing to the Southwest in havine both male and female subsection terms, and were only partly similar to the inhabitants of the North Wellesley Islands im thar the latter had male Subsection terms only (Sharp 1935: 192, fn. 4). They lacked the named senii-morety terms which operated within at least che western parts of Garawa country (Sharp £935, 1939; Reay (962; Trigger 1987a), The coastal Gangyatida alse apparently lacked the |nstitutionalisad role distinction of “ownership and ‘managership’ in relation to land and ritual property, which operated to the sourhwese (Thigeer (985: Table 6). Although: it is difficult co reconstruct thelr pre-contact ceremonial life because of more recent influesices from the west, it is most likely that the coastal Garpgalida also lacked the major cult teremonies in whith the ownership/managership distinction ts so important. In these respects, they were identical to the inhabitants of all the Wellesley Islands, GENETIC EVIDENCE Several publications have described similarities and differences of a genetic nature between North Wellesley Islanders, South Wellesley Istanders and mainlanders, Among other findings, Simmons ef a/, (1962) discuss the unusually high B blood group gene frequencies among the Bentinek Islanders, as compared with groups in the North Wellesley Islands, However, in later work, Simmons ef aj, (1964) did flirther comparisons with nearby malnlanders and concluded thal the ‘Karawa’ also had a high frequency of this gene, and in fact have been ‘a main source in the spread of the B gene in the mainland tribes of this remote area’ (p, GR). This discussion about certain genetic features of the Garawa people is nor without methodological problems.’ However, the relevant point w be considered here concerns claims based on this genetic evidence by Tindale (1977: 254-255), He asseris (hat the ‘Karawa’ constituted a ‘small separate and {solated tribal envlave', and lived ir a ‘perhaps refuge area’. These assertions fit with Tindale’s suggestion that the ‘Karawa’, like the Bentinck Islanders (and one other very distant north Queensland group) witty whom they apparently share certain blood type characteristics, are ‘relicts’ of 4 previously more widespread population with this blood-type, Elsewhere, Tindale (1974; 122) himself notes that his speculation that the name ‘Karawa' can be (ranslated as ‘uplanders” or ‘hills people’, is ‘hazardous’, This is also the case tor his other speculations about this group. Neither the sociocultural evidence oor the mapped distribution of Garawa estates im ny data, support the Suggestion that Gatawa people have been in any way isolated, although oral tradition must be reearded as nal necessarily providing precise information for historical periods greater than perhaps four generations, Reay’s (1962: 95) comments about there ‘always’ having been ‘a great deal of contact, invluding inter-marriage, between Garawa and Anyula |[Vanyula]’, are similarly based on ethnographic data. However, ipter-marriage between these two language groups has also been indicated jn genetic studles carrled oat by White (1978: 42: 1979; 439, 445). To summarise the discussion so far, the coastal mainianders have been shown to be part of a linguistic sub-group extending throughout the Wellesley Islands, and imland for a substantial distarice (at least 1D) ke southwards to the INLAND, COAST AND ISLANDS 75 Nicholson River). To the west and south-west, neither Garawa nor Waanyi (nor Yanyula on the coast) are within this linguistic sub-group. A narrow coastal strip adjacent to the Wellesley Islands is classified in the Aboriginal view as containing areas which are environmentally similar to close off-shore islands, and is conceived as the separate domain appropriate to ‘saltwater people’. Nevertheless, substantial movement occurred between coast and adjacent inland, as well as between the coast and the North Wellesley Islands. The fact that the Gayardild in the South Wellesley Islands shared language and kinship system with others on the mainland and in the North Wellesley Islands, means that it is problematic just how isolated these particular islands were. The same type of kinship system extends throughout the study region, however there are differences in the presence of systems of named social categories. Finally, published genetic data assert various differences and similarities between populations in the study region. While there has been a suggestion that the Gayardild (Bentinck Islanders) and the Garawa may be ‘elict’ groups of a previously more widespread population, ethnographic and certain genetic data indicate a lack of isolation of the Garawa. Thus, my evidence certainly does not indicate any simple pattern of relationships between the various groups and cultural forms treated so far. We are confronted with a complex pattern of overlapping cultural forms extending throughout inland, coastal and island societies. The paper now turns to a detailed consideration of some aspects of material culture. The evidence here indicates major differences between coastal mainland and the inland Garawa/Waanyi traditions. In a number of respects, the coastal mainland traditions resemble those of the off-shore islands (particularly the North Wellesley Islands), in their differences from Garawa and Waanyi traditions. DIVERSITY IN INLAND, COASTAL AND ISLANDS MATERIAL CULTURE In this section I discuss a form of decorative marking, the use of (and attribution of meaning to) shell material, spearthrowers, spears, fishing artefacts and technology, and watercraft. Decorative marking The marking known as jinanggliyari in Garawa/ Waanyi is said to be absent from items traditionally produced in ‘saltwater country’, including both the mainland coastal area and the off-shore Wellesley Islands. The marking is a longitudinal fluted pattern made by an end of the adze-type tool known as biynmala. This tool consists of a gently curved stick of varying length with stone blades attached at both ends. One end is typically said to be a broad blade (known as gubija), while the more pointed blade is known as jinangyi, hence the name given to the mark it makes. Roth (1904: 20) refers to this tool as a ‘native gouge’. He presents figures showing it and the manner of its use (1904: Plate XIV, Figs 101, 105), and notes the Lawn Hill [Station] name’ as ‘gobija’. He does not, however, distinguish between the two ends of the implement; although the figure he gives elsewhere (1897, Plate XII, Fig. 235) for areas to the south of the study region, possibly indicates the blade at one end as broader than the other. In dealing with this implement for areas to the west of the study region, Spencer & Gillen (1904: 636-640) describe considerable variation in the size and shape of the attached stone flakes, and discuss the consequent different sizes of fluting made by the implement. Contemporary Garawa and Waanyi older people explain that the jinanggliyari mark was commonly applied to the convex side of two kinds of boomerangs: juguli, near-symmetrical shaped for hunting (QM QE6218), and man.guburana, hook- shaped for fighting (QM QE4274). The marking was also applied to wooden coolamons, jugiva (QM QES51), and sometimes to other items such as spears, clubs and shields (Fig. 4). It is said to have helped ensure that a weapon would ‘fly straight’, parti- cularly when combined with the right song (cf. Roth’s [1897: 146] comment concerning spears). From the inland perspective, the mark described above distinguished the item as belonging to the network of language groups stretching to the west from, and including, Garawa and Waanyi country; although it is clear from Roth’s account, that it was FIGURE 4. Three items showing the longitudinal fluting mark known as jinanggliyari, which is absent from items both on the coastal mainland and the Wellesley Islands, (Courtesy Queensland Museum.) 16 DS. TRIGGER also used in areas well to the south of the study regioli (see also Mulvaney’s (1976: 81] summary map showing ils distribution to the south and west of the study region). [tems with this mark may have been traded to the coastal ‘saltwater country’ (and to the North Wellesley Islands) but the mark was apparently not used by coastal artisans. Nor js iL likely that coastal people commonly used the tool which made the mark. Lt is absent From Meniniott’s (19794) list of the Mornington [sland artefact repertaire. Merimmott (1979a: 111) does state the contemporary claim by some “Lardil’ people thal the practice of imseribing fluting on hooked boomeranes ‘was obtained from the mainland in recent history’, and coastal mainlanders may also have partially incorporated the marking during post-contact times. Memmott (1979a; 113) alse notes the wooden coolamon as of possible recent mainland ongin, and like the ‘Lardil’, coastal Ganggalida people appear to have only made coolamons from Mefalewor bark (Roth’s ‘pleat-lype’ [1904: 30]), in pre-contact times.? The adze tool which was absent from the islands and the coastal mainland repertoires, was the implement typically sed in inland areas fp gouge out wooden covlamons, as well as to inseribe the fluted mark on them, Shell material Shel! material is of course plentiful jn the coastal environment, and the fourteen Ganggalida classifications that | have recorded represent only a portion of such knowledge. By contrast, the intand perspective appears tc regard ‘saltwater shell” as belonging to two broad size categories only: raluinve — ‘big one’ and malduwa — small one’. An example of the former is the large ‘bailer shell’ (Melo Amphora) used for carrying water leg QM QE593, obtained from Allen Island), Roth (1904: 29) notes that on the Wellesley Islands ‘the ventral curface of the last whorl of the. . ..shell is pierced for insertion of the thumb during transport’. Garawa and Waany! people are said to have not customarily used this item. Inlanders know that coastal people used both malduwa and rabunye saltwater shells lo construct small and larger (respectively) cutting and scraping implements, and this is seen as having been necessary and appropriate due to their lack of sujtable stone material. Freshwater mussel shell wag apparently used similarly, but only to a limited extent, in the Inland areas. Shellfish also formed a more sub- ‘stantial part of the coastal diel, as compared with the inland diet, However, the difference in use of shell ntajenal that inland people today remark upon, is the coastal people's lack of shell pendants known by Inianders as jaramara (OM QEI809, obtained from the Lawn Hill-Nicholson River area in 1900), This item is pearl-shell (Pinctada siaxitna), as are the others in several museum collections identified by an inland man as jaramara. These pearl-shell pendants were received by Garawa and Weanyi people from areas to the west and were considered items of great significance. Thelr general Iniportance appears to have been asa marker of certain formal social rales. For example, two individuals ‘promised’ to each other as marriage partners wore Jaraniara (for 9 short time at least) on the occasion of the promise being formally and ritually recognised, A further example isa woman wearing the item following her child’s death, particularly in any assoctated rituml settings. The /aramara thus marks its wearer as the focus of ritual related to an |mportant social situation, Occasionally, there may be a scratching, or engraving on it which may indicate the tie to # particular person, For example, such a scratching on the QM specimen noted above was Interpreted as representing the hand and extended fingers of its custodian. These items are inherited from parents, grandparents and other kin depending on circumstances, While not always considered highly secret, access to them Is restricted. It is thus the exclusion of coastal people froms the customary teceipt of these items from west of the study region, Which distinguishes them from the inlanders. Senior Garawa and Waanyi people now say that Lhe items were not passed on to Ganggalida people ‘because they didn't know’. In these respects, the situation of the coastal mainianders was identical with that of the Wellesley Islanders." Spearthrowers While inland artisans made three types of spearthrower, the coastal people along with all the occupants of the Wellesley Islands, made only one of these. It is known in Garawa, Waanyi and Ganggalida as miurnigu, and Menmott (19793; ILD) reports the same lerm (‘nrurraku’) for “Lardil’, Wis cut from one straight piece of cylindrical wood so that the distal end has a raised notch which slips into the hole typically at the back of spears (QM QEL6/113, see Fig, 5), Roth (1909: 200) describes it in some detail and refers to it as ‘a very primitive form of implement met With in the Wellesley Islands, and on the adjoining mainiand in the neighbourhood of Burketown , , .’, and illustrates itin his Plate LVI, Fig. 14 (and see also Roth 1902: 15). As Roth mentions, this is a very light implement; contemporary people say it is commonly made from the farrgwida tree (Thespesia pozgminecides), or one of several other light woods. Presumably Roth designates it as ‘primitive’ because of the notion that its construction is simple relative INLAND, COAST AND ISLANDS 77 QE Als} mere FIGURE 5. Three types of spearthrower made in the study region. The bottom one (QE 16/1113) was made throughout mainland coast, island and inland areas, while the other two were made only in the inland areas. (Courtesy Queensland Museum.) to other types, or at least somehow less developed. This is further made clear when he notes (1902: 15) that it is identical (except in its size) with a ‘toy wommera’ used commonly throughout north Queensland by boys emulating the fighting behaviour of men. However, both coastal and inland people state its advantage over other types of spearthrower to be that there is no problem with a separate attached peg coming loose. It can not be seen as under-developed in terms of functional performance. The two further types of spearthrowers made and used by Garawa and Waanyi people, but not in the coastal mainland area or on the islands, are termed ngaliga and wujula. The former is again a cylindrical straight piece of wood, but with a separate wooden peg (known in Garawa/ Waanyi as ngurru) attached to the distal end, usually with gum (QM QE11/53, obtained from Turn Off Lagoon on the Nicholson River). The latter type is flat and linear with a large notch forming a spatulate proximal end. The peg is attached to the distal end at an acute angle to the face (Robins 1980: 58), (UQ 1813; see Fig, 5). Roth (1909: Plate LXI, Figs 1-4) describes these two types and notes (pp. 200-201) them as ‘occasionally to be found in the area around Burketown, but certainly not of local manufacture, being brought in from eastward’. However, I am sure the editor’s footnote replacing ‘eastward’ with ‘westward and southward’ represents what Roth intended; though it can be noted that the editor’s correction was overlooked in McCarthy (1939: 419). My information is certainly that these two latter types were made and used in Garawa and Waanyi territory to the west and southwest of Burketown. Spencer & Gillen (1904: 668-670) describe and illustrate the ngaliga type as often found to the west of the study region, and state that it is traded ‘eastwards towards the Gulf’. The one they illustrate is in fact from the ‘Anula [Yanyula] tribe’ so the type apparently reached the saltwater domain to the west of the coastal Ganggalida area. It is shown with a human hair-string tassel at the handle end, and, although Roth (1909: 201 and Plate LXI) also shows such a tassel, this is now said not to have normally been added by Garawa and Waanyi people. In another publication, Spencer & Gillen (1969 [1899]: 578-582) give the name for this type (said to be made by ‘the Waimbia [Wambaya] tribe’) as ‘Nulliga’, clearly the same term as ngaliga given above. In both publications, Spencer & Gillen also show the wujula type in areas to the west of Garawa and Waanyi territory. While neither of these latter two types are now said to have been customarily manufactured by coastal people or Wellesley Islanders, some coastal incorporation of these items and the skills to make them did occur during post-contact times. One museum specimen (UQ 1814) of the ngaliga type obtained in the 1940s, is recorded as having been made by a man at Mornington Island who learned about the style from mainland sources. 2 Spears Four types of spear can be distinguished. In general, spear shafts are now commonly known as mugura, but are distinguished according to the kind of tree they are cut from. However, spear types are distinguished mainly according to the nature of their heads. The first type was used throughout inland, coast and island settings; it is known by the word for prong in all languages, and usually two or three prongs are attached to a shaft (QM QEII/53, see Fig. 6). The manner of construction described and illustrated by Roth (1909: 190-191) for the Wellesley Islands, appears applicable to both the coastal Ganggalida and the inland Garawa and Waanyi. (However, with the incorporation of European wire, the prongs have been commonly attached by wedging them into a hole made in the end of the shaft.) To summarise the earlier technique, the wooden prongs are fitted neatly against grooves cut along the shaft and then tied with string. These spears were used mainly for fish, but also for animals such as goanna in some circumstances, in both coastal and inland contexts. The second type of spear head (babagana [Garawa] and jimindi [Waanyi], but not known by a term in Ganggalida) is stone, and the spear was used to hunt large animals like kangaroo and emu (QM QE4278, obtained in the Burketown area), The stone items are said not to have been manufactured in the coastal area, but were apparently procured by coastal people from various inland sources. They were thus used by inland and coastal people both as knives and as spear heads fitted to shafts (cf. 78 D. 8. TRIGGER serrerses FIGURE 6. Four types of spear made in the region. The QE 4766/1 type was not used in inland areas, while the stone flakes for the QE4687 type had to be imported into the coastal area. (Courtesy Queensland Museum.) Spencer & Gillen’s [1969 (1899): 592-593; 1904: 640-656] accounts of the manufacture of these stone blades and their multiple uses, upon which McCarthy [1976: 35] appears to rely). Roth (1904: 18, 22) notes that the stone blades were made ‘along the ranges up the western border north from Lawn Hill, etc.’, and that small bundles of them rolled up in tea-tree bark were bartered. He specifically mentions (1909; 190) ‘the stone-spears of Burke- town, Point Parker [ie. coastal Ganggalida country], and the ranges along the Queensland Northern Territory Border [i.e. inland Garawa/ Waanyi country]’; and, it may be noted, he also comments (1904: 22) on a large import’ of such items occurring ‘from the Northern Territory, across the border from Wollogorang [Station] southwards’ to his North-West-Central districts. Tindale (1974: 122, 228) states that ‘Karawa’ people traded stone knife blades to the east generally, and that Mornington Islanders sent young marriageable girls in return for these items. He also mentions McCourt’s claim (published soon after [McCourt 1975: 80, 108]) to have discovered the origin of the long stone spear points found by Tindale ‘on Bentinck Island . . . and on the adjacent [mainland?] coastline’; McCourt says these were quarried and made in the Wollogorang-Calvert Hills Stations area, on the Calvert River, near Redbank Mine, etc, — that is, in Garawa territory. Contemporary old Garawa people certainly state that a number of places, including this area, were sources of stone used for both long knives and spear points. Indeed, they take pride in the Garawa knowledge of the manufacture of various kinds of stone tools. However, how far the stone blades made in Garawa territory reached, remains an open question. Although a comment from Roth (190la: 4) states that strong shell tools take the place of knives and scrapers on Bentinck Island, ‘as on the mainland’, his statement (quoted above) about the presence of stone spear-heads in coastal Ganggalida country is quite unambiguous. Yet none of Roth’s data describe these items for the Wellesley Islands, nor does Memmott’s (1979a) recent work with contemporary ‘Lardil’ people mention them. Thus, Tindale’s assertion that the stone spear-blades reached Mornington Island may be incorrect. Tindale (1977: 267) himself notes that such stone items were never used on Bentinck Island, though he was shown one on Sweers Island which had been brought by visiting mainland Aborigines accom- panying European men, The third type of spear is known in Garawa and Waanyi as ngarrgidigidi (ngarrgadaba =to spear), (eg QM QES56/1, obtained from Turn Off Lagoon). The long wooden head is barbed on one or both sides, making it difficult to remove. This is a fighting spear, said to have been used in a variety of conflict and dispute contexts. Coastal Ganggalida people are said to have similarly used this spear. Memmott (1979a: 110) also describes it for the ‘Lardil’. The final type!3 was apparently not made or customarily used by the inland people. It consists of a light wood shaft attached to a long smooth hardwood point of approximately the same length as the shaft (QM QE4766, Nos 1-7). The method of attachment is apparently unusual (R. Robins, Queensland Museum, pers. comm.), being the hafting together of the two sticks whose ends have been sliced at much the same angles. The join is bound in a distinctive fashion and no gum is used. This method of attachment is quite different from that customarily used by inland people. The spear was used for large marine animals such as dugong and sea turtle and is known in Ganggalida as miyalda. Memmott (1979a: 110) describes the same spear for the ‘Lardil’ on Mornington, where it is known as miya. Fishing and other methods of obtaining aquatic foods Both inland and coastal people used pronged spears and nets to obtain a wide range of aquatic foods. A large amount of knowledge is particular to certain environments, and this is especially so in relation to ‘saltwater country’. Yet coastal people clearly also used the same techniques as inlanders when ranging into the vicinity of fresh waterholes, for example, ‘poisoning’ techniques involving the placing of particular plants in the water, which cause fish to float to the surface where they can be easily obtained. As well, not all practices are the result of environmental determination. For example the ‘squeezing’ of stingray flesh in the coastal area or swordfish flesh in both saltwater and freshwater INLAND, COAST AND ISLANDS 79 areas, appears to be done solely because the taste of the meat is preferred without the ‘sticky stuff? which comes out when it is squeezed. Nets were used by both inlanders and coastal people. Despite size variation they were all made from string which was produced by rolling together either strands of certain grasses (QM QE1912), or strands of bark from certain trees — the bark of Hibiscus tiliaceus is particularly useful for this purpose in coastal areas, but other trees such as kurrajong (Brachychiton sp.) or black wattle (Acacia hemsleyi) are used throughout the study region. There are two methods of net usage in both coastal and inland areas. The first involves four people, one at each corner of a big net, dragging it so that two corners are held near the bottom and two at the surface, and the two ends are moved so that the net forms part of a circle, eventually dragging its contents on to the shore. In the second method, two sticks are attached to two sides of the net which is moved vertically through the water while other people frighten fish into it; when a weight is felt against the net, it is taken to the shore. !4 It remains uncertain whether line and hook fishing was done in this region. Roth (1903: 5) states that ‘before the advent of the Europeans, such articles in any material (shell, bone, etc.) were unknown’. Some Aboriginal people today agree with this view. Some remain unsure, and others claim that hooks and lines were in use prior to colonisation. Indeed, a shell type of hook certainly exists, obtained in 1882 from the Settlement Creek area (QM QE5924, see Fig. 7): a grass string line is attached to a groove cut around the top of the shank of a shell hook, and what are designated in the associated documentation as several shell sinkers, are tied to the line. Roth’s (1901b: 20) general comment for other areas where he found usage of fish hooks, that weights or sinkers are never used, is thus also problematic. It seems most likely that the hook shown in Fig. 7 was an isolated recent import into the region, possibly originating from the Torres Strait Islands. !5 The term wardugu has been given by several contemporary old people to refer to fishing lines complete with hook (cf. Keen’s [1983: 294] translation of the term as ‘fish, hook, wire, line’), and while the term bala is given for the hook itself, it is more generally applied to any kind of forked shape, whether as part of an implement or simply the fork formed by tree branches. Some people today, both inlanders and coastal people among them, state that hooks were also made from emu, kangaroo and pelican bone. One specific account from two older Garawa men is that ‘goanna collar bone’ and ‘catfish jaw bone’ are suitably hook shaped and were used as such by Garawa people. ‘molt 2 S & S06 Gee Boyt) fe es FIGURE 7, Fishing line with shell hook and sinkers, collected in 1882 in the Settlement Creek area. (Courtesy Queensland Museum.) Various kinds of grub, frog, insect and shellfish are now used as fish bait, as well as the fruit from the fig tree (Ficus racemosa). If line fishing has indeed developed since the early phases of European impact, it represents an innovation which has occurred throughout inland, coastal and island settings. Of particular interest in this region is a large complex of stone fish traps on the Wellesley Islands and the adjacent mainland coast. Stone walls as traps are said to have been constructed across inland watercourses at narrow places, and these were mainly used during ‘flood time’ (the wet season) when the stream ran strongly due to the extensive wet season rains. More temporary sapling fences were also constructed across both inland and coastal watercourses; in the coastal area stakes were stuck in the bed at an angle so that the receding tide would push deliberately placed bushes and other debris against them. However, the inland traps appear to have been much less important in the food production process than their coastal equivalents. The extensiveness of the stone traps throughout the islands and adjacent mainland coast, suggest their importance in the food production process. Over approximately 470 km of coastline, 334 traps have been recorded, one trap per 1.4 km of coast- line (although there is considerable variation in the density of traps throughout the islands and adjacent mainland); (see Robins et al. [n.d.] for details of the physical features of the traps, their distribution and relevant ethnographic data). Figure 8 shows one of the mainland trap sites (at Bayley Point), This series of traps extends along the shore-line for approximately 400 m, and from the shore-line up to approximately 150 m out on to mudflats, to the edge of an extending rock platform. The receding tide would leave contained (if not RO DLS. TRIGGER FIGURE &. Stone fish trap complex ar Bayley Point on the mainland coast opposite the Wellesley Islands. (Courtesy Connah/Jones Aerial Archaeology, University of New England.) exposed in some instances) fishes (including shark and stingray), and at times sea turtle and dugong, On a visit to these traps in 1983, Ganggalida people also used a pronged spear to obtaln crabs from Within crevices under the trap wall, The stone wall traps represent a highly developed (and apparently durable) form of food production technology, which is distinctive of the coastal economy of the mainland and the Wellesley Islands, Watercraft The distinctive triangular shaped raft of the Wellesley Islands area (known in ‘Lardil’ as walpa) has been described in various sources (eg. Memmiott 19792; 111), [1 is clear from contemporary accounts that these were also used by enastal mainlanders opposue the islands who refer to it ax walbywe. Roth (1908: 161) confirms this fact, and describes the call as: formed of atimerous logs of while Mangrove’ (ied toxelher al the baits as well as ar the extremities, with the result that it is much narrower foreward than al the stern, On top is placed some sea-weed, a sort of cushion tor (he voyager to sit upon. With such frail craft the [Aborigines] will not only visit island and istand, but even cross over (o the mainland, usually on the one course, making fora spol somewhere in the vicinily of Poine Parker. Elsewhere, Roth (1903: 3) notes again that: ‘There is no doubt whatever that Communication goes on here [irom Forsyth Island] between the mainland via Raines [later to be called Paines] and Bayley Islands and Bayley Point’. Davidgott (1935: 40), 10 his survey of Aboripinal watercraft, states that apart from in this relatively linnted area of the southern Gulf of Carpentaria, this type of raft existed elsewhere only on a section of the north-western Australian coast, and the question of its advantages and disadvantages for those who adopted it remains an interesting research issue. Davidson (1935; 39-45) reconsteucts an liistori¢al developmental sequence for watercraft where the triangular raft was invented earliest, but had then ‘given way’ along most of the northern Australian coast to the supposedly more sophistl- cated and preferred sewn bark and dug-out canoes, He makes no direct comment on why the raft was retained tm two regions: However, a cautionary comment is surely required here against the notion that the rafts would haye necessarily been inferior to the canoes, The pointis similar to the one made above concerning Roth’s attribution of primitive- ness to the coastal (and islands) spearthrower. While the raft may appear to involve less sophisticated construction techniques than the canoes, an assumption that the canoes were better suited than rafis for travel in all circumstances may well overlook certain advantages possessed by the latter; For example, .a feature of the rafts, particularly the larger ones, may have been greater stability which may have enabled safer transportation avross shallow sandbanks and channels, of larger numbers of children and valued items (including burning cinders to be used to re-kindle fires}. Within the study region of this paper, Ganggalida people on the coast just io the west of the Wellesley [slands are said to have used the raft, but to a much more limited extent than the Gananggalinda people directly opposite the islands, [t was not maniufac- tured or used by the inlanders. Garawa people are said to have been familiar with the manutacture of the sewn bark canoe (termed by them wulganyi) from the bark of the messmate tree; however, they only used it occasionally, most likely when moving over wide sections of the lower reaches of such watercourses as the Robinson and Calvert Rivers and Settlement Creek, While Gangpalida has a similar term for this type of canoe (wu/gunda), the coastal people are said to have used it even less than Garawa people. No suitable messmate bark |s apparently available in the coastal country. The sewn bark canoe was thug used only te a limited extent east of Yanyula territory, and the above data fill the gap mentioned by Davidson (1935: 35) concerning |ts distribution west of the Wellesley Islands, It was the Yanyula people from the coastal areas far to the northwest of the Wellesley Iskands, wha are now regarded throughout the region as having been the experts with bark canoes. It was they also who came from the northwest in dug-out canoes, which had apparently diffused as far eastwards in the Gulf as the vicinity of the Sic Edward Pellew Islands in Yanyula territory, having been initially INLAND, COAST AND ISLANDS 81 introduced into northern Arnhem Land by Macassan visitors well prior to the European presence in the Gulf (Davidson 1935: 20-24). Dug- outs are now known generally by the same term, muwarda, as that given to all European-style boats regardless of their size. Whether Keen (1983: 278) is correct in asserting that this term (her ‘muwata’) derives from the English word ‘motor’, it is most likely that dug-outs were known in the study area before motor-driven boats were brought in by Europeans. Contemporary coastal Ganggalida old people know how dug-outs were made from large- girthed trees of certain kinds (for example, Canarium australianum). However, while such trees may be found in their country, only a small number of dug-outs were apparently made here following the introduction of steel axes and the increased eastwards migration of Yanyula people with dug- outs after colonisation. Material culture — a Summary In summary, a longitudinal fluting mark was used in decoration by Garawa and Waanyi artisans, but not in the Ganggalida mainland coastal area nor in the Wellesley Islands. This mark is now recognised as distinguishing artefacts as belonging to the cultural bloc extending from Garawa and Waanyi territories to the west and southwest. The adze-type implement used to make the fluting was not manufactured in the coastal or island areas, and hence the wooden coolamon could also not be produced, There was no established stone tool working tradition on the coast or in the islands, whereas Garawa and Waanyi people speak with pride of the stone tool production techniques used in earlier times. Shell cutting tools appear to have taken the place of stone tools along the mainland coastal strip and in the islands. Pearl shell pendants were received by Garawa and Waanyi people through trade from far to the west and southwest, but these were not passed on to coastal Ganggalida people or to Wellesley Islanders. Coastal mainlanders and Wellesley Islanders made only one type of spearthrower, whereas the inlanders produced two further types as well. Of the four spear types in the study region, two were used across inland, mainland coast and island areas, one was produced only on the coast and islands, and one (with a stone blade as the head) was produced only among inlanders, although mainland coastal people (and possibly North Wellesley Islanders) received the stone blades through trade to a certain extent. Aquatic resources were obtained by use of spears and nets throughout all parts of the study region (and line fishing appears to have been adopted across the region, with the probable exception of the South Wellesley Islands, from early phases of European colonisation). Large stone wall fish traps were used only in mainland coastal areas and throughout the islands. Unlike both inlanders and Yanyula coastal people to the northwest, coastal Ganggalida people and Wellesley Islanders used only rafts as watercraft. CONCLUSION This paper has compared aspects of inland, coastal and island societies. The most detailed data have been given for traditional material culture, and this has indicated a substantial degree of similarity between coastal mainland and islands societies, compared with nearby inland society to the west and southwest. As ‘saltwater people’ occupying ‘saltwater country’, the coastal mainlanders can be regarded as part of the island cultural bloc in significant respects. Yet the material culture of coastal mainland society has simultaneously emerged as similar to that of inland society in certain respects, and if further aspects of material culture were to be studied closely, a number of differences between mainland coast and the islands could well become evident.!© While the material culture repertoires of the mainland coast and islands are strikingly similar in significant ways, it would be inaccurate to define them collectively as a homogeneous material culture tradition completely separate from that of the inland. The same point was made when summarising the comparative data for the region on language, kinship and social organisation, and genetics. The paper has described a complex pattern of over- lapping cultural and other forms, with no area emerging as completely distinctive. This general- isation includes the situation of the Bentinck Islanders, despite the literature in support of their alleged lengthy isolation, At least wN% ffdemon- strated otherwise, we must recognise that the differ- ences between Gayardild society and that on the North Wellesley Islands and the coastal mainland, are apparently no more substantial than the differences between the mainland coast and inland. The main concern of the paper has been to consider the situation of coastal mainland dwellers adjacent to major off-shore islands society. The generalisation of this paper is that the relationship with the North Wellesley Islands societies in particular, has reinforced the maintenance of cultural differences between coast and inland societies on the mainland. This is not to argue for uni-directional influence from the islands to the coastal mainland. Indeed, some of the data indicate major areal influence from the mainland to both the North and South Wellesley Islands (e.g. the existence of an Aranda type kinship system RZ B.S, TRIGGER throughout the study region).. However, the North Wellesley Island societies should pot be viewed as without influence on mainland coastal socrety, In this sense, 1 am arguing thal those mainfand-island relations which entail regular contact should be viewed as similar to relations among peaples on the mainland, It may be that the envirorimental simi- larities in this region between the mainland coast and parts of the off-shore islands, particularly engender the cultural similarities dealt with im this paper (especially in the aspects of material culture traditions Which have been considered), However, the implication for studies in other regions is that when comparing island and mainland societies in Aboriginal Australia, particular attention should be paid to the environment and society of the immediately adjacent mainland coast, as well as to societies located further inland.. ENDNOTES |. The name of this language has been given as “Yukulta’ in the literature (Keen 1983), However, the term ‘Ganggalida’ was predominantly tised for the language and a group of people at the lime of my research, The larter term can be translated as meaning language’ or ‘talk’ (see Trigger (L985: 340-49} for further discussion of ihe relationship between these twa verms), 2. The language of the Mornington Islanders is otore conventionally spelt ‘Lardil’ (see eg, Memmott 1979p). However, except where quoling from works using this spelling, | will use the mainland Aboriginal expression ‘Layardiida’ 4. Elsewhere (Trigger 1987b), | have dealt if detail with the problematic issues entailed in using Janpuage names. in designate tr(bal” Units if this region, In this paper such cxpressions represeat a predominant View among Aboriginal residents which glosscs separable areas of land and associated bhadiles of tradnion by the use of language wares. 4 Indeed, Evans (1985; 7-8) describes the “basic ‘fsolation™” (\.e. distinctiveness) of the broader ‘Tangkie subgroup' of languages, meluding those throughout the islands and Ganggatida and Neuburinli oo (he maintand, when vompared with other Aboriginal languages. 5. Evans (pers. comm.) has speculated that the clymolagy of this term may be derived from kanatva — to buen’ and kafitva = Yo junp” (sce Keen 19R3: 272) fence, these people may have been known by a term for ‘ares jumping up’ or bushfire at times visible fram (he vantage point of other coastal peaple, 6, This is not to imply thal coastal mainlanders vagarded even the small close off-shore islands (Bayley and Pains) as withoul environmental features differentiating them in more specific respects frown (he ‘islands’ in maintand Saliwater country’ Certainly, the larger offshore istinds are much more diverse environments than both the smaller off-shore islands and the sand ridge ‘island’ areas within the mainland ‘saltwater country’; (see Memmott [1979b: 45-65) for a discussion of the land forms and ecology for Mornington, and Tindale |)962a; 280-288] for relevant information concerning Bentiack). [It would be interesting fo investigate whether the larger off-shore islands are designated as ‘slands’ (murndarmurra) in the coastal mainland perspective. Minimal data indicate that they are in fact regarded as wambalda (or and), hough coastal mamlanders would presumably identify as siurndamurra, areas within the larger off-shore islands similar to the sand ridges In mainland ‘saltwarer country’, 7, Simmons e/ @/, (1964: 75) themselves point out for the whole population tesied in this region, that ‘about one- half... were of mixed tribes and, of course, (heir earlier history ts unknown! Elsewhere (Trigger L987b), | have discussed the necessity of examining the linguistic (or tribal’) aljiations of individuals’ four grandparents if this matter is to be adequately researched. The basis on which individuals state primary affiliation with, say, father’s language rather than mother’s, involves a variety of social factors. Hence the possibility of defining clearly a relatively enduring endogamous population in ‘Wibal’ terms, from the slalements of individuals, is fraught with diffieulries. 8. Throughout the paper, items located in museum collections are identified by placing their registiation identifications iF parentheses, The letters ‘QM! indicate that the item isin the Queensland Museum collection, and UQ' indicates the University of Queensland Anthropology Museum cotlection, 9 Roi obtained wooden coolamons at Burketown in 1901 aod Forsyth and Mornington Islands in (903 (items 13359, 13360, 1336) respectively, in the Australian Museum register af the Roth Collection), however it is mast likely thal (hese would have been manufactured in inland areas. LO. Mr Tommy George, of Waanyi and Qarawa descent, inspected museum collections in Brisbane and Canberra 9 1980-ayid [) Brisbane during March 1984; some of the information he provided js confained in a transcription of an audio tape made alter his 1984 work (George 1984), 11, Several authors (see Mutlvaney’s (1969: 96; 1976: 83) summary maps) have documented peart-shell as an rem traded over very long distances, and by (hese accounts iL is possible that jt alsa came into the study region from af caglerly direction via Cape York Peninsula, However, itis rather the accounts of It coming originally (hur the very distant Kimberley coast (eg, McCarchy 1939: 96-98) which aré consistent with contemporary Aboriginal opinion in the study region, 1, Menmott (1979b; Table 7) notes ‘wuijala'as the name given by ‘Lardil' people to a spearthrower of the typical wert Cane York kind with attached shells atthe proximal end, and he says they reacted Morpington Island only in recent times. While it appears the “Lardil’ have thus borrowed his term. both mainland coastal and inland groups remain unfamillar with rie Cape York type and INLAND, COAST AND ISLANDS RS would mast likely simply referto it as being fram ‘another commry’, 13, Rowh (1902; 15) refers to a kind of toy spear used in games fy young boys: ‘so far observed . , . only amang the coastal blacks (o {he west of Burketown, and at Wollogoning (Northern Territory border)’, However while the few peaple of whom f have enquired stale thal it was used, | have no dala to compare with Roth’s description or ii, 14. The large ‘dugong net’ described for the “Lardil! by Memmoitt (1979: 112, 118) does fot appear to be known by contemporary Ganggalida people, though whether it was Once used in the coastal mainland area remains unvertain, 15, The hook collected from the region (Pig. 7) appears to be of the ‘bent pin’ type which is discussed by Massola (95: LL, 15) as having been possibly intraduced from the Torres Stralt, Anell (/955: 14-115) says that all fish hooks in Australla Were ‘undoubtedly’ intraduced from che Torres Sinait, and suggests that this accurred ‘relatively dale’, that 16 during the recent past. (6. Memmott's (1985) ongoing work comparing ‘Lardil’ and ‘Kaiadili' material culture repertoires indicates some significant differences between aspects of the material culture of the North and South Wellesley Islanders, the most striking being (hat (he Gayardild had a much smaller number of artefacts. Lt is most likely that further research would indicate that the Gayardild also diftered tran the mainland coastal Ganggalida in this respect, ACKNOWLEDGMENTS Research In the study region from early 1978 to 1983 was supported by the Department of Anthropology and Sovuidlogy, University of Queensland, and the Australian Institurce of Aborizinal Seudies, The staff of the Queensland Museum, particularly Mr Richatd Robins and Ms Julia Findlay, and of the University of Quoensland Anthropolagy Museum, parneulacly Ms Linely Allen, generously assisted in the location of items in the calleclions. Mr Richard Robins, and Dr Paul Memmott (Department of Architecture, University of Queensland) have provided helpful comments. Mr Tommy George, of Waanyi and Garawa descent, took particular interest in this research, and provided much Information during visits to museum collections in Canberra and Brisbane, REFERENCES ANELL, B. 1955, ‘Contribution to the History of Pishing if the Southern Seas’ Studia Ethnographica Upsahiensia No, 9, Uppsala. CHASE, A, & SUTTON, P. 1981. Hiinter-gatherers in a rich environment) Aboriginal coastal exploitation in Cape York Peninsula. Jn A. Keast (Ed.} ‘Ecological Biogeography of Australia’, Pp. 1818-1852. W. Junk, The Hague. CURR, E. M_ 1886, “The Australian Race’, Vol. 2. Government Printer, Melbourne. DAVIDSON, DS. 1935, The chronology of Australian watercraft, . Pol, Suc, 44: 1-16; 69-84; 193-207 EVANS, N.R.D. 1985. ‘Kayardild: The Language of the Bentinck Islanders of North West Queensland’ Ph.D jhesis, Australian National University, Canberra, GEORGE, T. 1984. Statement made aller examining the Gulf Country Collections ai the Queenstand Museum and University of Queensland Anthropalogy Museum Transeribed and edited by 1:5. Tripper from a tape recorded 26-3-1984, HALE, K. 1982. The logic of Damin kinship terminology, fa} Heavh, FP, Merlan & A. Rumsey (Eds.), The Languages of Kigship in Aboriginal Australia’, Pp- 1-97. Oceania Publications, Sydney. KEEN, S, 1983, Yukulla, Jn RJM. Oixen and B. Blake (Eds.), "Handbook of Australian Languages’, Vol 3 Australian National Universicy Press, Canberra. MCCARTHY, FD, 1939. ‘Trade’ in Aboriginal Australia, and ‘Trade’ relationships with Torres Strait, New Guinea and Malaya. Ovegnia 9: 405-438; 10; 80-104, 171-195, MCCARTHY, FD, 1976. “Australian Aboriginal Stane Iniplemerms! The Australian Museum Trust, Sydney, MCCOURT, T. 1975, “Abosigival Artefacts’, Rigby, Adelaide. MASSOLA, 4. 1956, Australian Fish Hooks and their Distribution, Mew, Mat, Mus, hic 22: 1th MEMMOTT, P, (979a. Lardil arifaets and shelters, Occasional Papers in Anthropology 9 07-142, MEMMOT, P. 1979b. ‘Lardil Properties of Place, PhD, thesis, University of Queensland, Berishane MEMMOTT, P- 1985. Aboriginal Fish Traps in the Gulf of Carpentaria und Wellesley Islands Research Project, Report on Data Collection and Compilation, Ms Aboriginal Data Archive, University of Queensland, Brisbane. MULVANBY, DJ, 1969. ‘The Prehistory of Auscratia’, Thames and Hudson, London, MULVANEY, D.J. 1976. ‘The chain of connection": the material evidence. Jn NM. Peterson (Ed.). ‘Tribes and Boundaries in Australia’, Pp, 72-94. Australian Institute of Abariginal Studies, Canberra. REAY, M. (962. Subsections at Borroloola. Oceania Ia: 9N-115, ROBINS, R. 1980. Wood identification of spearthrowers jn the Queensland Mueum ethnographic collection: an evaluation, Occasional Papers in Agthropalogy tO: 115-123. RORINS, R., MEMMOTT, P. & TRIGGER, BS. nal, Fish Traps of the Wellesley Islands and southern Gulf of Carpentaria, Ms. ROTH, W.E. 1897. “Eihnological Studies Among te North-West-Central Queensland Aborigines’. Government Printer, Brisbane. ROTH, W.E. 1900, Vocabulary list of Obarindi tribe, Ms, Queenstand Stale Archives Col/Ai9899. ROTH, WE. [90la. “The Carpentaria Blacks: the Wellesler Islands visit by Dr, Roth’, Ms, reprodueed by Dr P- Memmott, Aboriginal Data Archive, University of Queensland, Brsbane. ROTH, WF, (9016. Pood: its Search, caplire and preparation. North Queensland Ethnography: Bulletin No, 3. @ld. Parl. Pap. vy, 4 1391-1423, 84 D. S. TRIGGER ROTH, W.E. 1902. Games, sports and amusements. North Queensland Ethnography: Bulletin No. 4. Qld. Parl. Pap. 1: 1151-1174. ROTH, W.E. 1903. A Report from Dr Roth, Northern Protector of Aboriginals, Home Secretary’s Department, to the Under Secretary, Department of Public Lands, Queensland, 27-6-1903. Ms. Aboriginal Data Archive, University of Queensland. ROTH, W.E. 1904. Domestic implements, arts and manufacture. North Queensland Ethnography: Bulletin No. 7. Qld. Parl. Pap. 2: 2: 453-486. ROTH, W.E. 1908. Australian canoes and rafts. Man 8: 161-162. ROTH, W.E. 1909. North Queensland Ethnography Bulletin No. 13. Fighting weapons. Rec. Aust. Mus. 7: 189-211. SHARP, R.L. 1935. Semi-moieties in north-western Queensland. Oceania 6: 158-174. SHARP, R.L. 1939. Tribes and totemism in north-east Australia. Oceania 9: 245-275, 439-461. SIMMONS, R-T., TINDALE, N.B. & BIRDSELL, J.B. 1962. A blood group genetical survey in Australian Aborigines of Bentinck, Mornington: and Forsyth Islands, Gulf of Carpentaria. Amer. Phys. Anth. 20: 303-320. SIMMONS, R‘T., GRAYDON, J.J. & TINDALE, N.B. 1964, Further blood group genetical studies on Australian Aborigines of Bentinck, Mornington and Forsyth Islands and the mainland, Gulf of Carpentaria, together with frequencies for natives of the western desert, Western Australia. Oceania 35: 66-80. SPENCER, B. & GILLEN, FG. 1904. The Northern Tribes of Central Australia’, MacMillan and Co., London. SPENCER, B. & GILLEN, FG. 1969 [1899]. ‘The Native Tribes of Central Australia’, Oosterhout N. B., The Netherlands. TINDALE, N.B. 1962a. Geographical knowledge of the Kaiadilt people of Bentinck Island, Queensland. Rec. S. Aust. Mus. 14: 259-296. TINDALE, N.B. 1962b. Some population changes among the Kaiadilt people of Bentinck Island, Queensland. Rec. S. Aust. Mus. 14: 297-336. TINDALE, N.B. 1974. ‘Aboriginal Tribes of Australia’. Australian National University Press, Canberra. TINDALE, N.B. 1977. Further Report on the Kaiadilt people of Bentinck Island, Gulf of Carpentaria, Queensland. /n J. Allen, J. Golson & R. Jones (Eds.). ‘Sunda and Sahul: Prehistoric Studies in Southeast Asia, Melanesia and Australia’, Pp. 247-273. Academic Press, London. TRIGGER, D.S. 1982. ‘Nicholson River (Waanyi/Garawa) Land Claim’. Northern Land Council, Darwin. TRIGGER, D.S. 1985. ‘Doomadgee: A Study of Power Relations and Social Action at a North Australian Aboriginal Settlement’. Ph.D. thesis, University of Queensland, Brisbane. TRIGGER, D.S. 1987a. ‘Mugularrangu (Robinson River) Land Claim’. Northern Land Council, Darwin. TRIGGER, D.S. 1987b. Languages, linguistic groups and status relations at Doomadgee, an Aboriginal Settlement in North-West Queensland, Australia. Oceania 57: 217-238. WARNER, W.L. 1933. Kinship morphology of forty-one north Australian tribes. Amer. Anth. 35: 63-86. WHITE, N.G. 1978. A Human Ecology research project in the Arnhem Land region. Aust. Inst. Abor. Stud. Newsletter NS No. 9: 39-52. WHITE, N.G. 1979. The use of digital dermatoglyphics in assessing population relationships in Aboriginal Australia. ‘Birth Defects Original Article Series’, 15: 437-454, THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES : DROMATINAE) BY C. PATTERSON & P.V. RICH Summary The oldest known emu is Dromaius gidju n. sp. from the medial Miocene Kutjamarpu fauna at Lake Ngapakaldi in northern South Australia. This form, based on a partial hind limb, is smaller and has relatively shorter and less mediolaterally compressed hind limb bones, and less reduction of the medial and lateral digits than in the living form. D. gidju thus appears to be less specialized for a cursorial lifestyle, being somewhat intermediate between the forest dwelling cassowaries and the highly cursorial living emu, D. novachollandiae. Fossils from the Late Miocene and Early Pliocene may be allied to D. gidju, but more material is needed to allow confident assignment. D. ocypus from the medial Pliocene Palankarinna fauna at Lake Palankarinna, northern South Australia, is intermediate in size between D. gidju and D. novaehollandiae. In addition, its tarsometatarsus is decidely shorter relative to width than that in D. novachollandiae, thus indicating it is not as highly adapted for cursorial life as the living emu. Essentially all other emu fossils, Late Pliocene-Recent, appear to belong in D. novaehollandiae including : D. ‘patricius’, D. “gracilipes’, and ‘Metapteryx bifrons’, all defined originally by C.W. De Vis. The only exceptions are the King Island emu (D. ater) and the Kangaroo Island emu (D. baudinianus). Whether there was greater size variability in Pleistocene emu populations and whether a separate species of emu once inhabited Tasmania are problems yet to be resolved once larger collections of both living and fossil emus can be measured and analyzed. ‘THE FOSSIL HISTORY OF THE EMUS, DROMATUS (AVES: DROMAIINAE) C. PATTERSON & P, V, RICH PATTERSON, ©, & RICH, PV. 1987. The fossil history of the emus, Bronraius (Aves: Dromaiinae). Rec. S. Aust. Mus. 21(2); 85-117. The oldest known emu is Dramaius gidju n. sp. from the medial Miocene Kutjamarpu fauna at Lake Ngapakaldi in northert: South Australia, This form, based on a partial hind limb, is smaller and has relatively shorter and less mediolaterally compressed hind limb bones, and less reduction of the medial and lateral digits than in the living form. D. gidju thus appears to be less specialized for 4 cursorial lifestyle, being somewhat intermediate between the forest dwelling cassowaries and the highly cursorial living emu, D. novaehollandiae, Fossils from the Late Miocene and Early Plidcene may be allied to D. gidju, but more material is needed to allow confident assignment. 2. ocypus from the medial Pliocene Palankarinna fauna at Lake Palankarinna, northern South: Australia, is intermediate in size between D. gidju and D. nevachallandiae, In addition, its tarsometatarsus is decidedly shorter relative to width than that in 2, nevaehollandiac, thus indicating that it is not as highly adapted for a cursorial life as the living emu, Essentially all other emu tossils, Late Pllocene-Recent, appear to belong in D. nevaehollandiae including: D. putricius! D. ‘eracilipes; and Metapteryx bifrons; all defined originally by CW. De Vis. The only exceptions are the King Island emu (D. afer) and the Kangaroo Island emu (.D. bavdiniunus), Whether there was greater size variability in Pleistocene emu populations and whether a separate species of emu once inhabited Tasmania are problems yet to be resolved once larger collections of both living and fossil emus can be measured and analyzed. C. Patterson, Department of Zoology and P. V. Rich, Departments of Zoology and Earth Sciences, Monash University, Clayton, Victoria 3168. Manuscript received 19 August 1986. The living emu (Dromaius novaehallandiae) is B.P, Years belore present the second largest living ground bird, exceeded only c. Cranium, crania by the ostrich in size. Today and in the past, emus Cor. Coracoid i). have been restricted to Australia, and their origins CSIRO =~ Commonwealth Scientific and Industrial Research Organization, Division of Wildlife and. Rangelands Research, Canberra are nol understood. The fossil record of emus begins in the Miocene, with two now extinct species occurring one each in d Distal the Pliocene and the Miocene of northern South diapop. Diapophbyses Australia, The Quaternary King Island and est, Estimated Kangaroo Island emus seem to belong in two EB Femur separate species. All other fossil emus, mainly Fib, Fibula Pleistocene, however, are very similar fo and most HM Hunterian Museum, Glasgow probably conspecific with the living D. novae- Aurn, aon ; hollandiae. It is very likely, however; that the history hd ~_ f of emus on the Australian continent is much older M Mandible than curtently understood because of the general MM Geological and Mining Museum, lack of a pre-Miocene terrestrial record. Sydney Although the Pleistocene emus are currently NMY Museum of Victoria, Melbourne indistinguishable from the living emu, the Tertiary p Proximal species are distinct, The hlnd limb of the single Ph. Phalanx, phalanges Miocene form is not as cursofially adapted. This postzyg. Postzygapophyses 7 species hag a tarsometatarsus that is shorter and QM Queensland Museurn, ageing more robust, avd the lateral and medial digits of QVM ace Victoria Museum and Art : = ery, Launceston the foot are not as reduced as in the living emu, Rir} Right This paper reviews fossil emu material and SAM South Australian Museum, Adelaide outlines the major evolutionary trends demon- SIAM — Smithsonian Institution - American strated by the dromaiines during the last 20 million Museum of Natural History years, Expedition Field Numbers, The following abbreviations are used: Washington, D.C. and New York AM Australian Muscum, Sydney Sk. Skeleton(s), many skeletal elements AMNH = American’ Muscum of Natanil History, St. Sternum New York Syn, Synsacrum 86 C, PATTERSON & PV. RICH T2, 73, T4 Trochleae (1, [t, LV Tih, Tiblotarsus Tr. Tarsometatarsus Ku Vertebrate) UCMP = University of California, Museum of Paleontology, Berkeley WAM Western Australian Muscumt, Peeth PRewous Work There is surprisingly little in the literature concerning the fossil emus of Australia {see Table 1). The first reference to a specimen supposedly related ta emus was, in fact, a moa, ‘Dinornis queenslondiae’, described by De Vis (1884) from the Darling Downs, Queensland, Some later workers considered [his specimen (e.g, Hutton 1293, Miller 1963) to be related to the emus and cassowaries- Scarlett (1969), however, Found the fossil could be assigned ro Pachyornis elephantopus, probably collected from a midden on South tsland, New Zealand, and thus it is not a valid Australian record, and certainly not an emu. In 1888 De Vis described a new species af emu, Dromeius petricius, from a proximal end of a right tibiotarsus (QM 5547) and the distal end of another tibiotarsus (QM F5548), In the same paper he provisionally referred a left coracoid (QM FiL20) to the same species, These three fossils were from King Creek, Darling Downs, in south-eastern Queensland, De Vis (1892) considered the whale of the Darling Downs sediments to be much the same age, but it is now known that these fossil-bearing deposits represent a range of ages. The Chinchilla fauna is likely to be of Late Pliocene age. On the other hand, the Darling Downs fauna of the eastern part of the Downs, including King Creek, is of Late Pleistocene age (Woods 1960, Stirton ef af 1968, Rich 1979), Later De Vis (1892, 1905) also referred a femur fragment. three tarsometatarsi, and a partial synsacrum (QM F5549) to D. potricins. The referral of the synsacrum |s especially poteworthy, De Vis considered that because of its size, the fragment must have been from a cassowary er an emu, but: ‘as no extinct cassowary Is known yet in Australia, it seems almost necessary to attribute the present fossilio the emu DB. parriciis’ {De Vis 1905: 5). In. 1892 De Vis set up another species of emu, Dromaius. gracilipes, based on a distal left tarsometatarsus (QM F1142)- In the description De Vis omitted to note the location from which the Specimen was collected, but the museum Label associated With the specirneri Indicates that [t was from the Darling Downs, In this article De Vis (1892) also described a supposed kiwi, ‘Metapferyx bifrons', again without giving a location Spencer (1906) described minor of King Island, Bass Sirait, Tasmania. The previous exist- ence OF a Separate species of emu on this island was almost simullancously made by Legge (1907), but he withdrew the name. D. minor was tedefined by Spencer & Kershaw (1910) as more specimens became available, and recently the taxonomic status of this species has been discussed by Parker (1984), The status of the extiner Tasmanian emu is an as yet unresolved problem, Emus were introduced from the mainland in the 1800s, and interbreeding may have occurred (Howchin 1926). Le Souef (1903) gave the Tasmanian emu the specific name 2 diemenensis. Ridpath & Moreau (1966) considered it a subspecies of D newaehollandiae. The onty fossils and recent specimens of D diemenensis collected allve which are known to exist include 4 fernut, @ synsaccum, three tibintarsi, two tarso- metatarsi, a ces vical vertebra, and a leg lacking the femur and part of digit II (all at OVM) (Scott 1924, 1932), and three eggs (in private collections) (Campbell (900, Le Souef 1903, Spencer & Kershaw 1910, Dave 1926). Anderson (1997) deseribed an emu sternum, which is much thickee than those of the living DB. navachallandiae, from the Wellington Cayes, New South Wales. He suggested it might possibly belong 16 B patricius. Miller (1962) restudied Caswarites lpdekkeri, discussed earlier by Rothschild (1911). The type of the species is a distal right tibiotarsus (AM MEF!268). The type locality has been variously piven as Queensland, Cooma und Wellington Caves (Miller 1962), but its provenance is uncertain, lus preservation, however, is very unlike that of fossils from Wellington Caves, It ts clearly a-cassowary, however, and not an emu, Miller (1963) described a new species of emu Dromivetus (=Dromaius) ocypus based on an essentially complete tight tarsometatarsus (SAM P13444) from the Pliocene Mampuwordu Sands, Palankarinna fauna, Lake Palankarinna, South Australia, It is smaller than D, neveehollandiae, The tarsomietatarsug is eVidently the one referred to by Miller In Stirton e¢ af, (1961) as a new species of em with ‘proportions of the bone... intermediate between those of the emu and the cassowary’. Miller also assigned four phalanges (UCMP 56849, 60563, 94679, 94680) from Lake Kanunka (UCMP V-5772, Katipiri Sands or possibly Tirarl Formation) to the Dromornithidae, possibly Genyornis newioni (Stirton et af. 1961). As noted by Rich (1979), however, they actually belong in the genus Drormalas; Thus, no dromornithids are known from the Pliocene-Pleistocene Lake Kanunka fauna, and this adds another record for emus, Rich (1979) refers a left femur (SAM PL7104), from Brother's sland, South Australla to Genvornis newiori, but it conformis in all respects to Dromoafus and should be transferred to that taxon, THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) TABLE 1. Australian localities producing fossil emus (Dromaius). Locality Fossil Elements Rock Fauna Age 87 References Leaf locality, Lake Ngapakaldi, South Australia Bullock Creek, Northern Territory Alcoota (including Rochow locality), Northern Territory Lawson-Daily Quarry, Lake Palan- karinna, South Australia Lake Kanunka, South Australia Chinchilla, Queensland Darling Downs, Queensland King Creek, Queensland Thorlindah, Paroo River, Queensland Bingara, New South Wales Lake Menindee, New South Wales Lake Tandou, New South Wales Wellington Caves, New South Wales Wombeyan Quarry Cave, New South Wales ?Baldina Creek, near Burra, South Australia Brothers Island, South Australia Tmt., d Tib., Pes Dromaius gidju Tmt., Tib. Dromaius sp. Tmt. frags., Phs. Dromaius sp. Tmt. Dromaius ocypus; Tib., F. Dromaius cf. ocypus F., Ph., R., Tib., V. Dromaius novae- hollandiae, Dromaius sp. Syn., F., Tmt. Dromaius novae- hollandiae Cor., F., Tib., Tmt. Dromaius novae- hollandiae Dromaius novae- hollandiae Tib. Dromasius novae- hollandiae V., Syn., Tib. Dromaius novae- hollandiae, Dromaius sp. Ph., Tib. Dromaius novae- hollandiae Dromaius sp. Tib. Casuarius lydekkeri; St., Tib., Tmt. Dromaius novae- hollandiae, Dromaius sp. Tib., Tmt. Dromaius novae- hollandiae F. Dromaius sp. F, Dromaius novae- hollandiae Wipajiri Fm. Kutjamarpu Miocene Camfield beds Bullock Creek Late Miocene Waite Fm. Mampuwordu Sands Katipiri Sands Chinchilla Sands Unnamed Unnamed Unnamed Unnamed Unnamed sand lunette Unnamed Unnamed cave sediments Unnamed cave sediments Unnamed Unnamed aeolianite Alcoota Palankarinna Kanunka Chinchilla Darling Downs King Creek Unnamed Bingara Unnamed Unnamed Unnamed Unnamed Unnamed Unnamed Late Miocene Pliocene Late Pliocene or Early Pleistocene? Early to Middle Pliocene Pleistocene Late Pleistocene Pleistocene Pleistocene Late Pleistocene Pleistocene Quaternary Late Pleistocene Quaternary Quaternary Stirton et al. 1967, 1968; Rich 1979. Rich 1979 Woodburne 1967, Stirton et al. 1968, Rich 1979, Rich et al. 1982 Miller 1963, Rich 1979 Rich 1975, 1979 Woods 1960, Stirton et al. 1968, Rich et al. 1982 Woods 1960, Rich 1975, 1979 Baird 1986 Etheridge, 1889, Rich 1975, 1979 Anderson 1889, Rich 1975, Marcus 1976 Tedford 1967 Rich 1975 David 1950, Rich 1979, Dawson pers. comm. Hope 1971, Rich 1975 S.A.M. Museum label Rich 1975, 1979 88 C. PATTERSON & P. V. RICH —_—————————————————————————————————————————————————— Locality Cooper Creek, (includes Katipiri Waterhole and Wurdulumankula), South Australia Kangaroo Island, South Australia (Several localities) Lake Callabonna (lower stratigraphic level), South Australia Lake Kittakittaooloo, South Australia Naracoorte (Henschkes Bone Dig and Victoria Fossil Cave) South Australia Salt Creek, South Australia Warburton River, South Australia (includes Green Bluff locality and Kalamurina). Bone Cave, Western Australia A cave north of Moore River, Western Australia Irishtown, Tasmania King Island, Bass Strait, Tasmania Mole Creek, Tasmania Moybray Swamp, Smithton, Tasmania Scotchtown Cave, Tasmania Fossil Elements Fauna Age References F,M Tmt., Syn., V. Katipiri Sands Malkuni Pliocene- Stirton et al. Dromaius sp. Quaternary 1961, Rich 1975 Dromaiinae Sk.. Unnamed Quaternary Morgan & Sutton Dromaius baudinianus 1928, Rich 1975, Parker 1984 Ci, Sym, Vi5F., Tib, Lake Pleistocene Stirling & Zeitz Dromaius novae- Callabonna 1900, Rich 1975, hollandiae 1979 Tmt, Katipiri Sands Malkuni Quaternary S.I.A.M. Museum Dromaius novae- label hollandiae M., V., R., Hum., Unnamed Pleistocene van Tets & Smith Syn., F., Tib., Tmt., cave sediments 1974 phs. Dromaius novae- hollandiae, Dromaius sp. F. frag. Unnamed Quaternary Rich 1975 Dromaius cf. novae- hollandiae Syn., Tib., Tmt. Katipiri Sands Malkuni Quaternary — Rich 1975 Dromaius sp. Tib., Tmt. Unnamed Quaternary Dromaius novae- cave sediments hollandiae Tmt. Unnamed Quaternary —_ Rich (unpublished, Dromaius novae- 1971, field notes) hollandiae Tib. Unnamed Quaternary Scott 1924 Dromaius diemenen- sis. Needs review Sk. Unnamed Quaternary Spencer 1906, Dromaius minor sand rock and Spencer & Kershaw 1910, Jennings 1959, Parker 1984, Rich 1975 Tib. Unnamed Quaternary Scott 1932 Dromaius diemenensis. Needs review Syn., V., F., Tib., Unnamed Quaternary Scott 1932 tmts. Dromaius diemenensis. Needs review Le Unnamed Quaternary Gill & Banks, Dromaius diemenensis. 1956 Needs review THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 89 a Locality Fossil Elements Rock Lancefield, Victoria Tib., Tmt., Ph. Unnamed Dromaius novae- hollandiae McEachern’s Cave, Many skeletal Unnamed Victoria elements Dromaius novae- hollandiae Buchan Caves, Tmt. Dromaius Unnamed Trogdip Cave area, novae-hollandiae Victoria Fauna Age References Unnamed Pleistocene Gillespie et al. (26 000 B.P.) 1978 Unnamed Quaternary McNamara pers. comm. Unnamed Quaternary = Rich 1975. The bird remains in the Riversleigh fauna (Carl Creek Limestone) identified only as close to ‘Dromiceius’ in Tedford (1967) have been determined by Rich (1979) to belong to a dromornithid, Barawertornis tedfordi, and thus are not a record of emu. STRATIGRAPHY (see Table 1) Only a few fossil sites producing emus have been found thus far, and most are of Pleistocene age. Fossils of a new species, Dromaius gidju, proposed in this paper, have been found at the Leaf Locality (UCMP V-6313) on the eastern shore of Lake Negapakaldi, eastern Lake Eyre sub-basin, South Australia (Stirton et al. 1967). The sediments that outcrop here, known as the Wipajiri Formation, contain the Kutjamarpu fauna. Diprotodontid marsupials in this fauna are considered more primitive than those in the Beaumaris, Palankarinna, and Alcoota faunas, and have closest affinities with forms in the older Ngapakaldi fauna, known from localities listed in Stirton et al. (1968), in the Lake Eyre sub-basin and thought to be of medial Miocene age (Rich et al. 1982). The Camfield beds at Bullock Creek (Bullock Creek fauna), Northern Territory, of probable Late Miocene age, have produced Dromaius sp. currently under study by P. V. Rich. The Rochow locality (UCMP V-6349) at Alcoota, Northern Territory, near Alice Springs, has produced Dromaius remains that may be Miocene in age. But, as discussed by Rich (1979), the Waite Formation, which contains the Alcoota fauna, is not well dated at present. The diprotodont marsupials from this locale suggest a date younger than that represented by the Kutjamarpu fauna but older or contemporaneous with the Hamilton fauna. An unnamed rock unit containing the Hamilton fauna is capped by a basalt, which has been dated as 4.35 + 0.01 my. B.P. (or Early Pliocene) by Turnbull ef a/. (1965) and Turnbull & Lundelius (1970). The sequence is underlain by marine sediments of the Grange Burn Formation assigned to the Kalimnan stage. At present Alcoota is viewed as Late Miocene in age. Dromaius ocypus was recovered from the Lawson-Daily Quarry (or Lawson Quarry; UCMP V-5769) at Lake Palankarinna, eastern Lake Eyre Basin, South Australia (Miller 1963). The enclosing rocks, the Mampuwordu Sands, contain the Palankarinna fauna, and are overlain by the Tirari Formation and the Late Pliocene or Early Pleistocene Katipiri Sands containing the Malkuni fauna at Lake Palankarinna. An age of Middle to Late Pliocene is established by marsupial fossils, in particular Zygomaturus, which are more advanced than zygomaturines from Awe, Beaumaris, and Alcoota and yet more primitive than Pleistocene forms (Stirton ef al. 1968). Lake Kanunka (UCMP V-5772) in the eastern Lake Eyre sub-basin, South Australia, has also yielded Dromaius fossils. The Katipiri Sands or possibly Tirari Formation (see Rich 1979: 61) contains the Kanunka fauna dated as Pliocene or Early Pleistocene (Stirton et ai. 1961, Rich et al. 1982, Tedford pers. comm. 1985). The Chinchilla locality, south-eastern Queensland (Chinchilla Sands, Chinchilla fauna) also contains Dromaius. Several elements of the marsupial fauna appear more primitive than those in the Pleistocene eastern Darling Downs, and Woods (1960) assigned it a Pliocene age. Rich ef al. (1982) consider Chinchilla to be Early to Middle Pliocene in age. The distal part of a tarsometatarsus (AM F 58087) of an emu was found in the Australian Museum’s ‘old collection’ and labelled ‘mixed plus some from Lord Howe Island’. The fossil is very incomplete and appears to be from a juvenile individual. There are no reliable stratigraphic or locality data available for this form. All of the other known fossiliferous sites producing emus are Pleistocene in age. For these o) © PATTERSON & Py, RICH Pleistocene sites, as might be expected, some dates are better established than others, Depasition, tor instance, of the Darling Downs sedirnerts in Queensland may have occurred al several diflerent times (Rich 1979) during the Pleistocene, and definite ages for specific sites are difficult to determine, Thorlindah, on the Paroo River, Queensland, is thought by Rich (1979) to be Probably Pleistocene .-. the bird remains were collected along with fragments of ‘kangaroos’ and Diprofaden (Stirling & Zeitz 1900; 44) ina well 20 feet deep’, Diprolodon appears to be restricted ta the Pleistocene in all precisely dated situations. Emu material indistin- guishable from the living forms is known from Thorlindah, Vertebrate fossil-bearing localities at Lake Menindee adjacent to the Darling River and its major anabranch, western New South Wales, have been radiocarbon dated at 26 300 + 1500 B.P, and 1§ 800 B.P, (Tedford 1967), Fossils of Diprotedan, Thylaceieo, Phkaseolanus, Protemnedon and mracropodids have been recovered. UCMP Jocalities V-5371, Y-7185, V-67186 and Y-67187 have produced Dromaius fossils, Hope (1978) discusses the Slratigraphy of the Menindee area in some detail, with reference to the problem of dating the Pleistocene megafauna extinctions, At present the emu fossils from Menindee appear to be Late Pleistocene in age. At Lake Tandou, New South Wales, several Dromaius fossils were found in archaeological excavalions. Hope (pers. comm.) states that: ‘there is NOW 4 reasonable stratigraphy for the lumette [at Lake Tandoul], and a lot more cates; the oldest are in the order of 22 000-25 000) and lie ac the base of the uppermost scratigraphie unit. The problem ..- i8in Working out where Harry's [Harry Allen, who collected the specimens while doing research toward a Ph.D. thesis) matefial came from’, Tenta- Lively, a Pleistocene age seems appropriate for these fossils, Bingara in New South Wales has produced verlebrae and a tibiotarsus of Dromaius. The bone bed occurs in a fluviatile clay deposit about 39-N) em thick on the western side of Myall Creek, Remains of Diprotodon indicate a Pleistocene age (Anderson (889), Also in. New South Wales,. the Wombeyan Quarry Cave has yielded Dromaius Fossils. This is not the same cave as Broom Cave or Guineacor Cave, also in the vicinity of Wombeyan. The Wombeyan Quarry Cave has not been radiocarbon dated, but Hope (1982) beheves that it is of Late Pleistacerie age, It seems likely that the quarry deposit is older than the ‘Broom breccia’, but bath appear te be of Late. Pleistocene age. Other fossils recovered from Wombeyan Quarry Cave include Protemnadon, Sthenurus, Zygomaturus, Palorchestes, Thylacolea carrifex, Sarcophitus faniarius, and Progura gallinavea, There are several bone producing caves in the Wellington Valley area of New South Wales. Different levels and different caves may have trapped animals at various times in the Pleistocene to Recent, perbiaps even prior to this (L. Dawson pers, comm.) depending on when they were opened and rescaled (David 1950, Tedford 1967), Emu fossils have been recavered from caves in this area. Rich (1979: 58) states that Dromiaius remains were recovered from Cuddie Springs (Mara Creek, SSE of Brewarrina, 16 km ESE of Gilgoin), New South Wales, Anderson & Fletcher (£934) do not mention Dramaius in their, admittedly incomplete, list of fossils tecuyered from this, site. Wilkinson (1884) stated that; ‘bones of Diprotodon, Stherurts, Macrapus titan, latge wormbats, large birds probably emus, cracodiles and a gigantic camivorous lizard, Nofiosaurus .—. are found only within a few yards of the centre of the spring* Unfortunately, he does not describe or figure these bones, and the large birds may be Genpornis, specimens af which were later recovered by Anderson and Fletcher. MM F 19420, onlabelled when found in an old collection, has ‘the style of preservation [suggesting] that it comes from Cuddie Springs* {Pickett pers. comm.) but is too large to be Brorraius. lt appears, instead, to be the internal condyle of # titiotarsus of a dromornithid, perhaps Geryarnis. We have been unable to relocate (he specimens Rich (1979) assigned to Drorpeivs. Two Pleistocere cave deposits producing Droinaius fosals are Known in Westeen Australia, A cave norit of East Moore, Western Australia, has produced a tarsometatarsus of a juvenile emu (unregistered WAM). Bane Cave, near Jewel Cave, has produced an emu tibiotarsus and tarsorneta- tarsus. Four Pleistocene Dromaius tocalities are known from Tasmaria, Scott (1924, 1932) reported a tibiotarsus from Irishtown, 2 tibiotarsus from Mole Creek, and several elements (a synsacrum, Jemur, tibiatarsus, two farsemetatarsi, and a cervical vertebra) fram Mowbray Swaynp, near Smithton, in western Tasmania. The Mowbray Swatnp fossil site has been rachocarbon dated ait greater than 37 780 BLP. (Gall & Banks 1956), Another Mr Scott found bones of the Tasmanian emu at Scotchtown Cave in association with ‘Notorherium tasmanicum', Thylacoleo carnifex, and Palorchestey (Gill & Banks L9SA). DPromaius minor is known from the Bass Strait island, King Island, Tasmania. Anderson (1914) was of the opinion that the original Fossil matrix was a fairly hard, coarse, red-brown sand rock of shallow manne origin. Jennings (1959) stated thar THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 91 the fossils occurred in windblown sand dunes of Pleistocene to Recent age and that finds from different geological horizons had likely been brought together by winnowing. Separate from the King Island form, the now extinct Dromaius baudinianus, is known from Kangaroo Island (Parker 1984). Several localities along the south coast of Kangaroo Island (Cape du Couedic, Kelly Hill, Eleanor River, and The Brecknells) have produced fossil material (Morgan & Sutton 1928). Rich (1975) states that the age is Pleistocene. Three Victorian sites, all Pleistocene in age, have produced fossil emus. A partial tarsometatarsus is known from Trogdip Cave, part of the Buchan Caves said by Rich (1975) to be Pleistocene because of the nature of the marsupial fauna also preserved in it. Many fossils of the Australian megafauna have been recovered from a swamp near Lancefield, Victoria. As well as emus (less than 1% of the bones), Macropus giganteus, Protemnodon, Sthenurus, Diprotodon, and a dromornithid, probably Genyornis, were found. A sample of the bones themselves was radiocarbon dated at 19 800 + 450 B.P., while charcoal in the channel fill in and upon which the fossil deposit rests provides a maximum age for the bones of 26 000 + 500 B.P. (Gillespie ef al. 1978). A third Victorian site which has produced emu fossils is McEachern’s Cave in western Victoria. According to Wakefield (1967, 1969), due to the funnel shape of the entrance the cave has acted as a death trap for terrestrial animals. Gravitational movement, movement of trapped animals and water action were responsible for considerable mixing of cave sediments. The fossils are Late Pleistocene to Recent in age. A sample of mammal bones from the top layer of the Pleistocene sediments gave a radiocarbon date of 15 200 + 320 B.P. Extinct Pleistocene species found in the cave include Sarcophilus laniarius, Zygomaturus trilobus, Thylacoleo carnifex, Sthenurus spp. and Protemnodon cf. brehus. The remaining sites from which fossil emus have been recovered are all South Australian. From Brothers Island, Coffin Bay, about 50 km WNW of Port Lincoln, a femur fragment SAM P17104, referred to Genyornis newtoni by Rich (1979) but actually Dromaius, was found in an unnamed aeolianite of sand and shells. As similar deposits on the island have produced Sthenurus cf. brownei (Tedford in Rich 1979), a Pleistocene age is indicated. A number of Pleistocene localities collected by J. W. Gregory (1906) and later by joint expeditions from the University of California and the South Australian Museum, occur in the eastern Lake Eyre basin. The fossils were found as ‘float’ or in place in the Katipiri Sands, which contain the Malkuni fauna. Also collected by Gregory and later expeditions of the University of California and the South Australian Museum, are several localities on the Warburton River, including Green Bluff Locality (UCMP V-5771), Lookout Locality (UCMP Y-5776) and Kalamurina. The Warburton River is in the eastern Lake Eyre sub-basin, and has produced fossils from the Katipiri Sands. A Smithsonian Institution-American Museum (SIAM) expedition in 1970 recovered a tarsometatarsal fragment (SIAM 75) of an emu from the Katipiri Sands (Malkuni fauna) at Lake Kittakittaooloo. The Smithsonian Institution-American Museum Expedition and later a Museum of Victoria- Australian Army Expedition also recovered a number of Dromaius fossils from the lower level of Lake Callabonna in South Australia. This stratigraphic unit producing the emus also contained Genyornis newtoni, Diprotodon optatum, Phascolonus gigas, Sthenurus, Protem- nodon, and Macropus (Stirling & Zeitz 1900, Rich 1979) and has been dated at greater than 40 000 B.P. (Tedford 1967), but sometime during the Pleisto- cene. An incomplete femur (SAM P17103) bearing the museum label: ‘ ?Genyornis. Pleistocene locality unknown, possibly Baldina Creek near Burra, South Australia’ is actually Dromaius. If the location is in fact Baldina Creek, a Pleistocene age is suggested by the occurrence of known Genyornis newtoni (Rich 1979) and Diprotodon at this site (Stirling & Zeitz 1900). Several fossils of Dromaius were collected near Burra, South Australia by Mr R. E. Ireland and forwarded by the police department on 12 March 1935 to the South Australian Museum, They were found in a sandhill in association with Aboriginal (Homo sapiens) bones, SAM A25805 (information from museum label). The Aboriginal remains suggest a Pleistocene to Recent age. Two caves near Naracoorte (about 320 km SE of Adelaide near the Victorian border), South Australia, have produced Dromaius: Victoria Fossil Cave (van Tets & Smith 1974) and Henschke’s Bone Dig. Sediments producing the fossils in Henschke’s Bone Dig have been radiocarbon dated at about 33 800 B.P. (van Tets 1974). Smith (1971) stated that the bones in Victoria Fossil Cave are most abundant in the top 15 cm of the damp, friable, light brown earth forming the floor of the cave. She also states that the abundance of sthenurines, diprotodontids, and Thylacoleo suggests that the deposit was formed sometime during the Pleistocene and sealed before the Recent. Wells ef al. (1984) provide a complete discussion of current dates from this site. 92 C. PATTERSON & P. V, RICH SYSTEMATICS Only those features exhibited by the fossil specimens are discussed. For measurements see Table 2. Family CASUARIIDAE Brisson Members of the Casuariidae have a pterygoid that is inflated where it contacts the palatine; a palatine with a short shaft and an expanded medial plate; a long vomer; a palate that lies ventral to the parasphenoid rostrum and makes contact with the braincase only at the basipterygoid processes; maxillopalatines that are cone-shaped and open posteriorly; the cervical vertebrae are antero- posteriorly compressed; the atlas possesses lateral spines or occasionally complete vertebrarterial canals; the sternum is longer than wide with lateral margins concave laterally and has short, dorsally- directed sternocoracoidal processes and no sternal notches; the costal margin forms about 50% of the lateral margin; the antitrochanter of the synsacrum is located at the anteroposterior mid-point of the synsacrum; the ilium, ischium and pubis are subequal in posterior extent; neither the pubes nor the ischia are fused posteriorly along the mid-line; the ischium is deeper than the pubis; the ilium dorsal to the acetabulum is deep; the trochanter and head of the femur are subequal in proximal extent; the external condyle extends only moderately distal to the internal condyle; the popliteal fossa is elliptical and of moderate width; the distal depth and width of the femur are subequal; the posterior margin of the proximal articular surface is highly concave anteriorly; the external condyle and fibular condyle are subequal in breadth or the fibular condyle is broader; the cnemial crests of the tibiotarsus are little compressed mediolaterally; the inner cnemial crest extends far proximally to the proximal articular surface; the external articular surface extends far laterally; the margin of the external condyle is semicircular in lateral view; the tibiotarsus lacks a supratendinal bridge and also lacks an intercondylar eminence; the hypotarsus of the tarsometatarsus is narrow and centrally located; the hypotarsus extends decidedly further proximally than the intercotylar prominence; the internal cotyla is deeper than the external; the posterior shaft surface is deeply grooved; the anterior metatarsal groove is deep and extends the length of the shaft; trochlea IV extends distal to trochlea Il; trochlea III extends distal to trochleae II and IV; the phalangeal count for digits I, III, IV is 3-4-5; of the proximal phalanges that of digit III is longest; that of digit IV is shortest; the unguals are generally claw-like, except for the elongated ungual of digit II in Casuarius. Subfamily DROMAIINAE Vieillot Within the Casuariidae there are a number of characters which reliably distinguish Dromaius from Casuarius, the only other member of the family. In Dromaius the mandible is broad and rounded distally, not narrow and pointed distally; the mandibular articulation of the quadrate is step- shaped, with the external facet decidedly more excavated (in Casuarius the facets of the mandibular articulation of the quadrate are subequal); the pterygoid is not excavated dorsally; and the palatine and vomer are decidedly shorter than in Casuarius; the semicircular notch in the prearticular surface of the atlas is shallow and narrow, not deep and broad; the axis is longer, and the hypapophysis not as deep as in Casuarius; the cervical vertebrae possess long, not short, styloid ribs, which come to a point distally, and are not rounded; the neural canals and vertebrarterial canals are small; the excavation of the neural arch posterior to the prezygapophysis is shallow (from the eighth cervical posteriorly in Casuarius the excavation of the neural arch is deep); the thoracic vertebrae are similar to those of Casuarius, but the neural canals are smaller; the neural canals of the caudal vertebrae are small, with an elliptical cross-section, whereas in Casuarius the neural canals are large and triangular in cross-section; the sternum is only slightly longer than wide, not much longer than wide as in Casuarius; the costal processes lie in an almost horizontal plane, whereas in Casuarius they lie on a downward curve (antero-posteriorly); the sterno-coracoidal processes are moderately long, not very short as in Casuarius; the coracoidal sulci are short and overlap medially, whereas they are long and do not overlap in Casuarius; the body of the sternum is weakly concave dorsally, but in Casuarius it is strongly concave dorsally; the depth of the sternum anteriorly is shallow, not deep; the costal margin is long, whereas in Casuarius it is shorter; the supratrochanteric ridge is broader; the pre-acetabular synsacrum tends to be shorter than the post-acetabular synsacrum in Dromaius while the opposite condition exists in Casuarius; proximally and posteriorly the femur bears a large pneumatic foramen, lacking in Casuarius; in anterior view, the external condyle extends decidedly further proximally than the internal condyle, while the two condyles are subequal in anterior proximal extent in Casuarius; in medial view, the internal condyle is semicircular in outline, while in Casuarius it is triangular; the diameter of the head and the minimum diameter of the shaft at its proximo-distal mid-point are equal, whereas in Casuarius the head diameter is less than the shaft diameter; the shaft is almost straight, being more curved in Casuarius; the proximal extent of the cnemial crest is not as great as in Casuarius; anteriorly the external VHE FOSSIL HISTORY OF THE EMUS, OROMAIUS (AVES! IROMAIINAR) " condyle is rounded proximally, and it extends further proximally and is more pointed in Cusuarlus; aboye the anterior intercundylar fossa is. a. small ridge trending dorsally and laterally from the mid-line and ending in a small foramen, while in Casuarius this cldge is absent, but the foramen still exists; the tarsometatarsus and tibiotarsus are subsqual in length, unlike in Casuarius in which the tarsometatarsus is decidedly shorter; the secand trocidea is much more reduced than in Cagiutrllss; the intercotylar prominence is low and tends to be flat, while in Casvarius the intercotylar prominence is higher and convex dorsally; a distal foramen, which completely pencirales (he tatsometatarsus (antere-posteriorly), and a groove (occasionally a completely roofed-over forainen) running proximo- distally, are present, both absent in Cuswarius, the condyles af the phalanges of the foot tend to be greatly divergent plantarly; In Caswarius the condyles tend to be only moderately divergent plantarly; in distal view, the intercondylar fossa tends to be only slightly notched in a step-shaped fashion dorsally, while in Casvarius this notels rencds 10 be deeper and more V-shaped; the ungual of digit I11 is longest, and that of digit TV shortest, while in Casvarius the ungual of digit IL is longes!, and that OF digit IV shortest, Dromaiirs novuelollandise (Latham) Type Casuarius noveehdllandiae (Latham) Type Locality New Holland (Sydney, New South Wiles, Australia) (Table 1), Measurements Tables 2-13. Referred Fassil Marerial Bingura, New South Wales — , MM F16786, dorsal vertebra (¥.24-26°), neural spine, pre- and postzygapophyses, diapophyses and prearticular surface damaged; MM FI6797, dorsal vertebra (H24-267), neural apine and diapophyses net preserved, Tie, MM F16775, distal end and distal half of shaft. Pletstocene. Bone Cave (wear Jewel Cavel, Western Australia — Tih, WAM6S,5,34 (in part), shaft only. Tint, WAM 68,5.94 (in part), shaft only, Quaternary. Brothers Islan, South Ausirelia — &, SAM PI7I0d, proxiinal ead and proximal Ewo-lhinds of shaft, head and trochanter damaged, Quaternary. Chinchily, Queenstand — Yar. OM F143 din part), third trochlea only. Cooper Creck, South Australia — Spa, UCMP 56233, acetabular complex, (site 2, LICMP V5378). Ff, HM B775/869, entire, (Lower Cooper, locality 4). Tint,, UCMP 56313, distal end, fourth trochlea not preserved, (site 3, UCMP V5379). Late Pliocene or Early Pletstocene. Darting Downs, Queensland — &, QM FI43 (in part), distal, popliteal fossa region only (eastern Darling Downs). Tif, AM A97I3, proximal, see Figure 1; QM FSS47, proximal, figured (De Vis 1589); OM FSS48, distal, figured (De Vis L889), OM F1652, proximal end, most of cnempal crests noc preserved, (Condamine River, fear Dalby), Tors, QM F112), proximal frag.; QM F135, figured (De Vis, 1892), distal, juvenile; QM F142, distal Irae. QM F1143 {in part), distal end (eastern Darling Downs), Pleistocene, Lake Callabonna, South Australia — AMNH 9478, Vi, second cervical, posterior left side; third cervival, left side: Fourth cervical (articulates with third cervical), neural spine and ribs not preserved; sivth cervical’, ribs and right side postarticuliar surface not preserved; seventh cervical’, (articulates with sixth cervical), ribs and right stde prezygapophysis not preserved; ninth cervigal’, postarticular surface, ribs, right side postzygapophysis not preserved. SIAM 61, Si, fragments. AMNE 9677. F, distal end, internal vondyle damaged, AMNH 9676. Tif, entire, see Figure 2, Pleistocene, Lake Kanunka, South Australia — K, UCMP $6854, dorsal vertebra (#22 of 23), (UCMP 5772). F, UCMP RHTLO64, trochanter, condyles, and head partly eroded and crushed, (site 1, UCMI* V5772). Tib, UCMP 56845, distal end, most of internal condyle not preserved and remainder highly eroded (UCMP V5772), Ph, UCMP 56849, first phalanx, second digit, (UCMP V5772, UCMP 94679, first phalanx, third digit; UCMP 94680, second phalanx, third digit (UCMP V5772). Late Pliocene or Early Pleistocene Lake Menindee, New South Wales — Eggshell, UCMP 55948. Tih, UCMP 53825, two distal tibiorarsi with the same number, Tin, UCMP 53835, distal end most of third trochlea and second and fourth trochleaée not preserved. PR, UCMP $3832, dark colour (presumably burne), first phalanx, second digit, distal end; Ungual phalanx, second or fourth digit} flest phalanx, second digit, proximal end; second phalanx, Fourth digit (UCMP V5371), LICMP 53833, first phalanx, fourth digit and ungual (site I. VA7ISS UCMP 55983, first and second phalanges, third digit (UCMP Y67186), Late Pleistocene Lancefield,. Vietorta — Tid, NMV P43037, distal shaft; NMV P43041, distal shaft, juvenile: NMV Pd40l1, entire but articular surface worm; NMV PISO0(3, distal, Tw, NMV LS, distal; NMV P44012, proximal artitular surface eroded; NMV P44013, hypularsus eroded; NM¥ P4404; NMV P44015, proximal articular surface worn, sevond and fourth trochleae nol preserved, NMV P44016, 04 C. PATTERSON & PV. RICH proxtnial articular surface wort; NMV P4sD17, distal shaft; NMY P44018, distal; NMY P44019, distal; NMY P48392, sécond and part of third trochleace not preserved; NMV P150014, distal, fourth trochlea not preserved. Ph: NMV P4319, first phalanx, third digit, articular surfaces worn; NMY P43200, second phalany, third digit, proximal end. Late Pleistocene, 26 000 BP, MecWachern'’s Cave, Vieloria — C, NMV P157345, posterior fragment; NMY P157350, lower jaw, distal; NMY 9157353, posterior tragment. h, NMV PI57346, 2Jst or 22nd vertebra, neural spine, diapopbysis, might side prezygapophysis, part of centrun aod prearticular surface not preserved; NMV P175349, 23rd or 24th vertebra, neural spine, pre- and postzygapophyses not preserved, postarlicular surface worn: NMY PI57351, seventh cervical?, juvenile, ribs not ankylosed) NMY P157352, lith, !2th or 13th cervical, juvenile; NMV P15735%, third cervical, right side rib not preserved; NMV P1IS7364, 25th or 26th vertebra, leit side of centrum with prezygapophysis and diapophysis, but postarticular surface not preserved; NMV P157367, 20th or 23st verjebra, diagpophysis, part of neural arch, postzygzapophyses, and right side prezyzapophysis only, juvenile; NMV PI57368, about lth vervioal, prearticular surtace worn, left side prezygapophysis and ribs (not ankylosed) nat preserved, juvenile; NMV P157369, 22nd to 26th vertebra, right side of centrum only, juvenile St, NMV P157347, incomplete; NMV P15735§, entire. Syn, NMV P157361, fragment. Tih, NMV P157356, proximal; NMV P157357, distal; NMV P5736, distal; NMV_ P157365, proximal, Fih, NMYV P157363, proximal end. Tit, NMV P15S7344, distal, trochleae nol preserved, Quaternary. Cave sorth of Moore River, Western Australia — Trt, WAM-190 unregistered, (not seen, data from Pat Rich’s 197! field notes), Quaternary. Naracoorte, (Henschke's Bone Dig and Victoria Fossil Cave), South Australia — C, SAM PL7234, lower jaw, distal end only, (Henschke’s A3, 40"), KK, SAM PI7589, Léth cervical, ribs nol preserved; SAM P18246, fourth cervical; SAME P1I8247, 22nd to 24th vertebra, part of centrum and left side diapoplrysis, (Henschke'sk SAM PIS673, 15th to 1th cervical? vertebra, prezygapophyses and centrum damaged, (Hensehke's area X4, depth 17"); SAM Pi8830, 26th vertebra, prezygapophyscs, prearticular surface and diapophyses damaged, (Henschke's area X6, 15-30 em). Syn, unregistered SAM; SAM PI6501, acetabular complex only, (Victoria Fossil Cave, 0-10+RI0-0-12"), see Figure 3; SAM P17747, parts of alium, ischia and pubes not preserved, (Henschke’s A3, 39-42"); SAM PL&s00, distal tight ischium only, (Henschke's Af, 32-36"), J, SAM P2281? (in past), condyles badly erodod: unregistered SAM, intemal condyle darmaged. Tih, SAM PI7L49, distal end, internal condyle worn; SAM PI8829, distal, part of condyles not preserved, (Henschke's area X64, D-15 cm). Tint, SAM PL7B16, distal, (Henschike’s Al, 30-33" SAM P18693, 2 pieces, proximal, with articular surface badly eroded, and distal (Henschke's area A4, 150 om, western wall), Ph, SAM Pfs059, first phalanx, second digit (Henschke’s area A3, 33-36"); SAM P18248, second phalanx, third digit, (Henschke’s); SAM P18249, first phalanx, Fourth digit, (Henschke’s}; SAM Pi8252, first phalanx, fourth digit, (Hensclike's). Pleistocene, Salt Creek, South Australia — &, SAM P1710), proximal shaft-only, but head and part of trochanter not preserved. Quaternary, Thorlindah, (Paroo River), Queensland — 77b., MMF 12074, figured (Etheridge 1889), a cast (AM £516) has been made, distal end, condyles worn, Pleistocene. Trogdip Cave, Buchan Caves, Victoria — Trt., NMY Pi57343, shaft only. Pleistocene. Warturton River, South Australis — i, UCMP 56642, 2ist or 22nd vertebra, centrum only (Green Blut lovality, UCMP V-5775). Syn, UCMP 5647, fragment, fused sacral vertebrae only. F, HM 801/934, distal, moat of internal condyle nol preserved, {Kalamurina), see Figure 4. Ter, SAM P1318, distal end, second trochlea not preserved. (Stony crossing of Warburton, Six road miles west of new Kalamurina Station.) Quaternary. Wellingion Caves, New South Wales — 0b, AM ‘Ri’, distal end, external condyle missing; AM F10949, distal half, (J. Mahoney, in a note on the back of the museum label, disputes this locality), Tint, unregistered AM. proximal] articular surface badly worn; AMC’, proximal; AM F18935, discal end and part of shaft, second trochlea not preserved, other trochlea pitted, no distal foramen, juvenile, AM MBF7T7I, distal. Quaternary. Wombeyan Quarry, New South Wales — Tih, AM P58025, distal end and distal one-third of shalt. Tint, AM PS8026, proximal, Late Pleistocene, Comments and Descriplion Dromeius nevaehollandiae is the only extart species. A number of subspecies have been suggested (Condon 3975), but little is known of their ranges or morphological distinctness. The osteological characteristics of the species have been described above. There is a suggestion that a slightly smaller, as well as a larger form of Drowiaius novaehollandiag existed during the Pleistocene, The flame gracilipes proposed by De Vis for the smaller form was applied to a juvenile D. noveehollandiae {see below). Hence the smaller form, if real, is yer unnamed. As the evidence is limited, we have chosen nol ( create @ separate specific or subspecific names. De Vis alsa described a larger THE BOSSIL HISTORY OF THE EMUS, BROALAIUS (AVES: DROMAIINAE) 95 species of emu, B patricius, which we have also synionymized with B. navaehollandiae {see below}, Dromaius patricias (De Vis) Lecrolype (here designated) QM F847, proximal right tibiotarsus, King Creek, Darling Downs, south-eastern Queensland, Pleisiocene, Measurerients Tables 9 and WJ, Referred Material De Vis assigned a coracoid, and a proximal and 4 distal tibiotarsus to this species without naming a type specimen. The left coracoid, OM FI120, was only provisianally referred to parriciis (De Vis L888; 1291). In actual fact itis not even a bird bore. The bone is not hollow, and it projects too far lateral to the point taken by De Vis for the glenold facet to conform with an emu coracoid, Additionally, it lacks a pneumatic foramen, [t 15 too large, heavy, and robust to match any bird, ft is probably part of a mammalian pelvis. The distal ead of the left (not right as De Vis states) tibiolarsus, QM £5548, which De Vis assigned to D. pa(ricius is not distinguishable from D, novaehollandiae. De Vis also stated (p. 1290) that: ‘the rotular surface is relatively longer fore and all to a considerable extent and less concave transversely’, but he admitted (p. 1291) that this: ‘is perhaps in some measure due to abrasion’. The difference in the ‘eminences and ridges for niuscle ingertions’ anteriorly are also as De Vis states {p. 1291): ‘scarcely of specific value’, This specimen is within the size range of 2 noveehollandiae. Hence we designate the proximal right tiblo- tursus, QM F5547, as the lectotype of Dramans patricius, It is in most respects trivially different, if al all from BD novaehollandiae. The proximal width (of the articular surface) is greater than any 0 sovaekollandiae in our sample (57.6 mrn vs a maximum of 55.6. mm for xevaehollandiae, sample size, 2 =9), The inner cnemial crest (> 90.2 mm) is unfortunately not entirely preserved. lL may have exceeded (he maximum of our sample a! D. novae- hollandiae (103.1 mm), Concerning other points raised by De Vis, the fibular crest does oot attach more proximally, but the bone is thicker at the most proximal point of this crest, the external enemtal eresr does descend more distally and a groove between the external and internal cnemial crests does exist that is larger than is present in DB, novae- Hollandiae, We feel that the variability exhibired by D, patricius would not fall outside that of a large samtple af che lying emu, De Vis (1905) also referred a synsacral fragment (consisting of (He neiral canal of several syasacral vertebrae), OM FS549, to D, pareicins, though itis so incomplete as lo render diagnosis difficult. It is within the size range of D. nevaehollandiae and probably could be referred to that species, We have, however, chosen to assign it only to Casuariidae indeterminate, [t was collected from Wurdulu- mankula, a Cooper Creek tocality, De Vis (1892) referred a part of a distal end of a femur (likely to be QM F143, in part), the proximal third of a tarsometatarsus (likely QM F121), the ‘valcaneal region of another metatarse' (apparently lost subsequently) and a distal tarsometatarsus (likely QM FIl43 in pari) to D. patricius. As De Vis did not figure or describe these specimens in any detail, we are assuming chat the QM specimens listed are those referred to In his {892 paper, They agree with the (limited) description, were collected in the Darling Downs according to the museum labels accompanying the specimens, and appear to bear (on the fossils themselves) De Vis' handwriting. OF these, only the distal tarsometatarsus is. described, t is stated to be larver in almost all of its dimensions than the living Emu. It is, indeed, wider than any in our small sample both in the shaft and im ita lrochlear eXpansion, and trochlea 3 is deeper than those in our sample of B noyvaehollundiae. The proximal tarsometatarsus is very worn; it lacks both the internal and external cotyla, the intercotylar region, and the hypotarsus. It is within the size range of the living D, nevaehollandiae, The femur is. also very frazmentary; only the popliteal region is preserved, and it, too, lies Within the range of D. novaechollandiae. Etheridge (1889) referred a distal right tibiolarsus, MM F 12074 (and cast AM L516) to D. parricins, but contrary to his assertion, the specimen is net lareer than nor of a different shape to that element in the modern emu. I should be reterred to BD. novaehollandiae. Thus, some of the specimens that have been considered to be D. patricius are indistinguishable from DB. novaehallandiae. Others may he slightly larger in some measurements, but we doubt that D potricius deserves, on that account, specific status, because our sample of the living emu is still small. We, therefore, synonymize D, patricins will D, navachallandiae. Dromaius gracilipes (De Vis) Holotype Dnt. QM Fil4d2, Darling Downs, Queensland, Pleistocene, Measurements Tables 10 and 11. Deswvription De Vis based this species on a distal left tarsometatarsus, QM P1142, with the second and 6 © PATTERSON & PV RICH fourth trochieae not preserved and che margins of the Uvird rrochlea very eroded, The characteristics which De Vis used to distinguish thas fram 2 novachollonidiee are the lack of w distal foramen and associated miuscle canal, infecior size of the distal end, antero-posterior compression of the Shalt, and disproportionate size of the mesial trachlea, These are all juvenile characteristics, De Vis also states that the width of the third trochlea taken from centre of the lateral depressions (i.e, the ligamental pits) is greater in D, gracilipes than in D, novachollandiae. This is not the case. We therefore synonymize 2 gracilipes with D. novae- Aullendize, Metapterys bifrons (De Vis) The Tint. QM FII3S, locality not given, but presumably Darhng Downs, Queenstand, Pleistacene, Measturemnents Table 1. Deseriprian De Vis erected this genus and speries on the basis of a lef distal tarsometatarsus, QM F1I35, and allied it with the kiwis because: ‘the trochlea appear 10 be borne on the ends of moderately long stalks’ (De Vis 1892: 449), the lateral trochlear processes (i.e. the second and fourth trochleae) are almost equal in length, the medial trochlea extends beyond the orhers, the posterior surface of the shaft shows the lines of junction between the coalesced segments, and it lacks a distal foramen which perforales the shaft. De Vis considered the possibility that these might be juvenile characteristics, but unfortunately dismissed this idea. De Vis was of the opiiiion that the fourth trochlea Was shorter (ihal it was a righr tarsomelatarsus), but actually the second trochlea is shorter. The comparatively large intertrochlear notches, the rough pitted vaps on the trochleae, and the points raised by De Vis (presence of epiphyseal lines, lack of distal foramen) are indications thal the specimen is of a juvenile bird. As De Vis himself noted, the fossil does not have any articulation for the hallux possessed by kiwis, and the middle trochlea is too large for that of a kiwi. Metapteryx: bifrons in all respects conforms to a juvenile individual of 2 navaehollandioe, and we synonymize it with that species Dromaius ocypus (Miller) Holotype Tint, SAM Pi3444, Lawson-Daily Quarry, Lake Palankarinna, easfem Lake Eyre Basin, South Australia, Mampuwordo Sands, Palankarinna fauna, Phooene. Measurements Tables 8, 9, 10 and 11, Referred Material F UCMP RASS5176, condyles, trochanter, and mast or head not preserved. Th, UCMP RASSI82, distal, Same locality as type, Description Miller (1963) established this species from an essentially complete, but somewhat distorted and cracked right tarsometatarsus, SAM P13444, The overall length, width across the distal end, depth across the internal cotyla and proximal width are all less than similar measurements in living and fossil D, novaehollandiue. Additionally, the curvature of the intercotylar region is. moré pronounced (convex dorsally) than in D, novae- hollandiae, As noted by Miller (1963), the trachleae have already allained the size and proportions of 2 novaehollandiae. Vhe width across the distal end is smaller, in part because the intertrochlear notches are narrower than in D, noveehollundiae, The femur and Ubiotarsus are provisionally assigned (0 D, acypus, although they lie within the range of D. novaehallandiae in those parts which are preserved, because they were found |i the locality of the type specimen. The tarsometatarsus of D. acypus differs most noticeably from D. novae- hollandiae in its shortness. Unfortunately, the referred specimens are incomplete, and their length cannot be ascertained, Dramatus gidjur i. sp. Holotype SAM P26779. Associated Jett leg elements, Type locality Leaf locality (UCMP V6213), Lake Ngapakaldi, eastern Lake Eyre Basin, South Australia, Wipajirl Fm. Kuljamarpu fauna, Medial Miocene. Descripiion An incomplete left lez, consisting of the distal fragment of the ubiotersus (originally UCMP 71397), the proximal part of the tarsometatarsus (originally UCMP 71398), and a complete pes (originally UCMP RASS234}, The tibiotarsal fragment articulates with the tarsometatarsal fragment, The pes is complete but does not arti- culate with the tarsometatarsal fragment as the trochleae of the tarsometatarsus are not preserved. All fragments were found in close proximity and the assumption is made here thal they are from one individual, Dr R.A. Stirton in his field notes of 19 July 1962 assumed thal the leg and foot elements were all from one individual and assigned them a single field number RAS 5234. THE FOSSIL HISTORY OF THE EMUS, DROMA/US (AVES: DROMAIINAB) 97 Etymalogy From an Aboriginal word meaning ‘small’ (Anonymous 1965, language not specified). Meashrements ‘Tables 9, (0 and 12. Referred material None, Diagnosis A small emu with a slender antero-posteriorly compressed tibiotarsus and tarsometatarsus, The anterior lip of the Intercotylar region is convex dorsally in D. gidju (as in D. ocypus compared tu nearly Hatin D. novaehollandiae). The intercotylar region does not extend far proximal to the articular surface as it does in Casuarius, however, The lateral lip of the external cotyla is noticeably convex laterally in D gidju (weakly so in D, novae- hallandiae), The width and depth of the proximal articular surface are much less than in D. oeypus and D. novaehollandiae. The D, gidju tibiotarsus is much smaller than those In our D. novee- hollandiae sample, but is similar in general appearance and proportions of the distal end, Anteriorly the tarsometatarsus proximal to che condyles is somewhat crushed, The external ligzamental prominence above the external liga- mental pit is not as well defined in D, gidju as it is in D, eavoehollanciae. The anterior ligamental fossa appears proportionally larger and deeper than in D, novaehollandiae. The phalanges (except the second phalanx of Lhe second digit) are smaller than those ia & novaehallandiae, Digit 11 1s compara- tively longer relative to the other digits than in D, novaeliallandine (64% of the length of digit ILI versus 57,5-460.5% for our sample of De nevege- hollandiae (see Tables 2 and 12); the first phalanx of digic It is 88% of the proximal depth of the first phalanx of digit [11 versus 76.5-78% for our sample of 2D novaehollandiae, and 66% of the proximal width of digit LL yersus 52-53% for our sample of D. novaehollandiae), Digit 1V is also Compara- tively longer but less change has occurred (68% of the length of digit IL versus.62-64% for our sample of D, novaehollandiae, the first phalanx of digit 1V is 78% of the depth of the first phalanx digit IIE versus 73-75% in our sample of 2 novae- hollandiae). Excepting the proximal phalanges, the ratio of maximum proximal depth to width ts greater in 2 gidju; thus, the phalanges of A novue- hollondiae are more dorso-ventrally compressed than in DO gid/u. The ungual phalanx of digit IT in D gidjn is longer than the ungual of digit tH] (which ts poorly preserved). This is not duc to an clongation of the ungual of digit Hin gidje,as in Casuarlus, but the weak development of the digit JIL unpual. Comment From what 1s known of its hind limb structure, i) would appear that D. gidju was less cursorially adapted than D. novaehollandiae. This is based on the greater length of digits Wand LY relative to dapat Wiin D. gidjv as compared to D. novaehollandiae. This. foot structure is presumably adapted for Breater maneuverability in forested or less open conditions and greater ability Lo qove over regions of a somewhat unpredictable nature. Dromaius sp. indet. Several specimens because of their fragmentary nature and/or Wnusual proportions could pot be assigned (o species level, Other specimens were referrable only to Casuarlidac indet. (sce following section). Referred Material Tables 3-12 No Locality Data — Jini, AM F38087_ distal end, trochleae not preserved, Alcoota, Northern Territory — Tint, QM QA205, third trochlea only, QM QASOS, distal, fourth trochlea not preserved, UCMP RASS397 (in part), a third trovhica only. PA, OM QASO4, first phalanx, third digi; UCMP RAS $397 (in part), second phalanx, second digit. The Afvoota specimens are close fo & gidyu. Late Miocene. Baldina Creek? (near Burra). South Australia — Ff, SAM P17103, shalt only, partially reconstituted in plaster, Quaternary, Cooper Creek, South Australia — Tit, QM FIN21, proximal, badly eroded articular surface: Could be Pliocene-Recent in ave, Lancetield, Victoria — Tht. NM'V P35578, distal shaft, Pleistocene in age, dated at about 26 000 BP. (Qillesple ez af, 1978). Warburton River, Suuth Austratia — Db, OM F6668,, distal, condyles not preserved, (‘Kalie- murina?’ ts pencilled on the bane). Tit, QM F667), proximal, incomplete fusion of metatarsals dorsally, (atsal cajy not preserved, juvenile, Pleistoccne- Recent in age Wellington Caves, New South Wales — Si, AM F25218, figured (Anderson, 1934), incomplete, Thur, AM !A' distal, trachleae nor preserved; AM FLO8S0 (cf. DL nevaehollandiae), proximal end plus proximal part of shaft, articular surface eroded. Pletslocene. Dramaiinac inset. Referred Material Tables 4-12. No Location Data — Syn,, OM £6673, taecnent, fused vertebrae only. 98 ©, PATTENSON & PV. RICEL Bingara, New South Wales — Syn, MM FI6795, fused vertebrae only, Pleistocene, Couper Creek, South Australia — , HM BT76, dorsal vertebra, 24th-26th?, centrum and left postzygapophysis only, (Lower Cooper, locality 2). Ff, HM 8777, part of a lelt internal condyle and internal popliteal fossa region anlyy UCMP 60532, popliteal fossa region only, (Karipiri Waterhaole, UCMP site 9, V-586J), The locality suggests all Speciniens are Oromiais sp. Pleistocene. Lake Kanunka, South Ausiralia — r, UCMP 60560, third or fourth vertebrae?, proximal (dorsal) only, (Site |, UCMP V-S772). Probably Dramatus: other Dramaius dements known here Probably Late Pliocene, possibly Pleistocene. Lake Kittakittanolow, South Australia — Thr, SLAM 75, fourth trochlea only, pitted, juvenile. Late Pliocene, possibly Pleistocene, Mebachern's Cave, Victoria — KF or NMYV P157354, dorsal lrazmenls NMY PIS7358, dorsal fragment; NMV P157362, dorsal fragment; NMV P)57366, dorsal fragment, one facet not preserved, Probably Dromains. Quaternary, Naracoorte, South Australia — Kr, SAM P1807, third vertebrae?, dorsal Fragment, (Heuschke's Bone Dig); SAM PL8251, dorsal fragineut only but facers not preserved, (Henschke’s Bone Dig); SAM P8784, third vertebrae?, dorsal fragment (Hensthke’s Bone Dig}i SAM P22812 {in pari), third or fourth vertebrae?, dorsal fragment, The lovaliry sugpests these speciniens are Dvevrtuins. Quaternary, probably Late Pleistocene. Warburton River, South Adstralia — Syn., UCMP 56647, acetabulum and pectineal process only, Quaternary. Wombeyan Quarry, New South Wales — Tih, AM. P58027, ef, Bromaius, distal end. Late Pleistocerie, Aves indet, (previously assigned to Dromraius) Referred Material Tables 10 and 11 Kalamurina, South Australia — Tier, SAM P11552, distal end, second trachlea not preserved, large. Perhaps Dromornithidae (Table Li), Quaternary, Nu Locality Data — 7/b,, SAM P1748, proximal shall anly, articular surface and cnemial crests not preserved. Possibly net avian — the bone js rather dense. Dromaius ater and D. bauddinianus are pot reviewed here as Parker (1984) has. recently revised their taxonomy, Discussian Al least One species of emu, 2B gidju, was presen in central Australia in the mid-Miocene. Itis known from the Lake Ngapakaldi jn northern Sout Australia and associated with the Kutjamvarpu fauna. It does not differ suffictently from other emus to require erection of a new genus. While the intercotylar region of the fossil tarsometatarsus resembles the condition found in Casuariys in that is if not markedly flattened as in the Ilving Promaius, this character stale is also Inve of B ocypus, an undoubted emu from the Pliovene. The shape of the margin of the internal cotyla of the fossil ps similar to that of Dromaius and dissimilar to that of Casuarius, which is more excavated posteriorly. The posterior surface of the tarsometatarsus of D. gidju is unfortunately chipped and cracked, but the shape of the remaining fragment of the hypotarsus and Larsal cap appears more emu-like Lhan cassowary-tike. Comparing the pes of the Miocene fossil with recent emus it is obvious that lhe foor structure has undergone change through the last few million years, The second.alid, (oa lesser extent, the fourth toes haye undergone a reduction in size, This (rend is parallel with several other ratites (the ostrich and some of ihe Dromornithidae; Rich 1979), and would appear to be a cursorial adaptation similar to the reduction and eventual loss of Jateraland medial digits within horses (Equidae), The pes of D giejy is more cassowary-like than that of any known living or other fossil emus, implying that in the Miocene, emus were not as highly adapted to an open plains, cursorial lifestyle as they are now. The pes does not, however, contain the specialized ungual spike on digit U, which so characterizes the cassowaries, D, gigju appears to bea species that may be close to forms which gave rise to both emus and cassowarles. Based on material now ayailable, D gidju has a few specialized characteristics that seem to ally it with Dromaius, But, as our records of this form increase, there may be sufficient reason Lo separate it from both genera within this family as an early, quite Unspecialized form. A small emu of Miocene age, is also known from Riversicigh, Queensland. This material is currently under study by Waller Boles (Australian Museum), By Late Miocene or Barly Pliocene times a species near in size to D gid/u is kwown in the Alcoota fauna. It is represented by tarsometatarsal and pedal fragments. This form may be referable to DL gidju, The two phatanges dilfér slightly from those of the Lake Ngapakaldi form, which we have referred to D) gidju, bin are within the range expecred for intraspecific variation. Until more complete miaienal is available from Alcaota, however, Assignment (o Dremelus sp, indet, is preferable. It TIE FOSSIL HISTORY OF THE BMUS, BROMATLS (AVES: DROMAIINAE) 99 is certainly not referrable to Cisuarits, as the second trochlea ig much more reduced relative to the other trochleae, similar to the condition in Dramatis. By mid-Plioceng, a second species of emt, D. ecypus Miller, intermediate in size between D, gidjue and D. novaehollandiae. existed. It 1s known from a right larsometatarsus atid part of a fenjur and Gbiotarsus. The femur and tibiotarsus are, in those parts which are preserved, not unlike PD. novaehallandiae, but if complete would probably be shorter than the corresponding elements in Db novyuehollandiae, In any event, 2 novaehollandiae existed by the Late Pliocene or Early Pleistocene. Since then only D, novaehollandiae has been present on the mainland, [thas probably fluctuated slightly in size, presumably as a result of a host of selection Pressures such as climate (both temperature and rainfall), diet, predation pressure and competition. The species restricted in Recent times to King Island and now extinct may possibly extend Into the Pleistocene. Localities on King Island are as yet nol carefully dated, The emus of King Island (2 runor) and Kangaroo Island (D bavdinianus) appear to be separale species (Parker 1984), The populations oo both of these are most likely relics isolated by rising sea Jevels atthe end of the Jast glaciation of &@ population that perhaps Wag once more widespread, Other than their smaller size the King and Kangaroo Island emus differ but little from the mainland emu. The main osteolagical difference is in the shape of the skull (Spencer & Kershaw 1910, Morgan & Sutton 1928). Possibly also the distal foramen of the tarsometatarsus differs (Shane Parker pers, comm.), but there is considerable variability in this character in the mainland emu, The degree to which whe groove for che musculus adductor digiti 1V is roofed over by bone would appear to be age related. In juvenile crus the arch is almosi completely lacking, whereas in some adult specimens it is completely formed ln bone. Thus, this character is aot taxonomically significant for emus, Some Australian mainland fossil emus are within or jusr larger thaw the size range for D, miner tabulated in Rich (1979). At Lake Menindee, two distal tibiotarsi (UCMP 53825, includes RHT6 and RHT25) lic within the range of DB eninor for the width across the condyles and depth of external condyle but exceed DB minor in the depth of the internal candyle — both specimens measure 37.0 mm in depth of the internal condyle; the range of asample of 50 2. minor was 26.2-36,1 mm, the mean was 30.4 (Rich 1979, Table 33). What the relationship of these fossils is to 2 minor \s unresolved and will remain so until a much larger fossil sainple of mainland birds is at hand. De Vis (892) did deserjbe a smaller mainland Pleistocene species, D, gracilipes, but hls type specimen is undoubtedly an immature PD, novae hallandiae, Nevertheless, smaller emus did exist in Australia In the Pleistocene, OF specimens listed in the systematics section (above) the following lic below the range for modern emus in one ar mote measurements: AM — unregistered tarsomete- tarsus, “A', Fi0949, MF773; HM B775/869; QM Fit2l; SAM — P1318, PI8099,; UCMP — 53825, 53832, 35983, 60532, 79510, RHTIO64. The presently known fossils of mainiand emits smaller than the living 2 sovaehellandine are unfortunately few. We do not believe that they are represeniatives of D minor or D. haudinianus, because of the age of the fossils on the islands ly Late Quaternary. We favour the idea that speciation on King and Kangaroo Island could have taken place in very iittle (ime geologically speaking, Strong selection for dwarfism quite likely occurred alter chese emus became isolated al the beginning of the last interglacial (Le. the Holocene). The mainland emu, 2 nevarkallandiae, may have been at any One time in the Pletstocene bouk larger or smaller than at present. However, there is a possibility that the differences seen in fossil samples are more apparent than real, since the sample size of modern emus is still fairly small and some of the emus in osteological collections were recovered from zoological gardens, Whether or not extant wild emu populations differed significantly in size is largely unknown, As our sample did not contain representatives from the Northern Terntory or Western Australia, it is also unavoidably biased geouraphically. Periodic dwarfing of the mainland form may have been caused by the same selective agents which produced dwarfing in the island forms, We were unable to test che hypothesis that size changes were related 10 palaeoclimate or other environmental variables because there are coo few reliably dated emu specimens in the Quaternary collections. De Vis recognized 2 pairicius as a separate, larger species of emu, but we can see no significant size difference from B raveettollandiae, Perhaps D. patricius differed in its proportions from D. noveehollandiae, As so few complete bones are known, however, this is difficult to assess, Por example, a tibiatarsus SIAM 6) was found to have a smaller length to distal width ratio than most modern emus, bur this difference did not prove statistically significant (9>O0.405, t-lest)- There was a mass extinction of the Australian megalauna fihe larger macropodids, diprotodontids, dromornithids, ete just before the Holocene (Tedford 1967, Gillespie er al 1978)) suggesting a widely acting selective agent against large size, There is a suggestion thar the Tasmanian emu, UX) (~ PATTERSON & BP. RICH D. diemenensis, averaged slightly smaller than the mainland form, This idéa stenimed from the known eggs of the Tasmanian emu measuring slightly smoaller than those of the mainland emu (Dove 1926), and from the recollections of Legge (1907), Who saw the Tasmanian emu as a boy. On the other hand, Spencer & Kershaw (1910) report that the Rev, Knopwood captured an ‘Emew 60 lbs. weight’ on 9 October 1804 in Tasmania, Scoll (1924) gives the dimensions of a leg of the Lismanian emu collected by Gunn in the 19th century; it ts as large as those of the mainland, The fossils of the Tasmanian emu are large (Scott, 1924, 1932) indicating thal the larger Pleistocene form of the mainland also reached Tasmania, presumably ata tinie when Bass Strait did not exist — during a glacial period of lowered sealevel, Concerning the extinct Recent Tasmanian emu, the best evidence supports the view thal it was about the same size as (he mainland emu, Condon (1975), following Ridpath & Moreau (1966), treated the Tasmanian emu as a subspecies of D. novaehollandiae, Kathryn Medlock (Tasmanian Museum) is currently reviewing the status of this form, ACKNOWLEDGMENTS Our thanks go to many individuals and Histitutions that provided specimens utilized in this study ; W, Boles (Australian Museum), G, George (Sir Colin McKenzie Wildlife Sancinary, Hewles- ville), 1, Hope (formerly of the Australian National University), A. MceBvey (Museum of Victoria), M- McKenna (American Museum of Natural History), K, McNamara (Western Australian) Museum), R. Molnar (Queensland Museum), 8. Parker (South Australian Museum),.J. Pickett (Mining Museum), N_ Pledge (South Australian Museum), T. Rich (Museuny of Victoria), W, D. 1, Rolfe (Aunterian Museum), G, F van Tels (CSIRO, Division of Wildlife and Rangclands Research, Canberra), and D, Vernon (Queensland Museum), R. F. Baird, S, Parker, T. Flannery, F. Szalay, N, S, Pledge and G. F, van Tets critically read (he manuscript, and we are graletul for their time, We are sincerely grateful to all those field parties Which collected the original material, in particular those led by the fate R, A, Stirton (University of California, Berkeley) and R_ H. Tedford (Americar Museum of Natural History) Photography of specimens was provided by R. Smart, &, Coffa and E, M. Thompson (Museum of Victoria) and &. Morton and T. Zylstra (Monash University). Drafting was provided by D, Gelt, P, Hermansen, J, Muir and B, Shea typed varlous versions of the manuscript. Funding allowing the complelion of this project was provided by the National Geographic Sociery, the Australian Researeh Grants Scheme, the Commonwealth Education Department, and the Ingram Trust. We owe them our deep gratitude, Both the Garth Sciences and the Zoology Departments, Monash University, provided the venue for our work. REPERENCES ANDERSON, C, 1937, Sterne of a ratile bird (Dreurneier spt tron che Wellington Caves, New Seath Wales. Fae, Aust. Mus. 0: 76-77, ANDERSON, ©. & FLETCHER, H©. (934. The Cnddie Springs hone bed, Aust, Mie Mae, 5 152-158, ANDERSON, W. 1889, Gn the postTertiaey osaiferous clays, near Myall Creek, Aingara, Ree, Geol, Sri N.S, 2 116-126, ANDERSON, W. 1914, Nole an the occurrence of Lhe sandrock containing ones of extinet species of Marsupials (Emu (sic), Kangaroo, Wombat, vic.) of King Island, Bass Strait, Tasmania. Aust, Mus, Mer. 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OOLO, SJ 1977. ‘Ontogeny and Phylogeny’ The Belknap Press of Harvard Univ. Press, Cambridge, Mass. SO) pp. GREGORY, JW. 1906. ‘The Dead Heart of Australia’ John Murray, London xvi + 384 pp, HOWE, LH. 1978. Pleistocene mammal extinctions; the problem of Mungo and Menindee, New South, Walas. Aleheringa 2: 65-82. HOPE, J.H- (982. Fossil vertebrates [ropa Wombeyan eaves, Jn ‘Wombeyan Caves”. Sydney Spel Soc. Occ. pupor 6, 155-164 HOWCHIN, W, 1926, Same teterences (o (he literature concerding the extinet emus of Kangaroo Island and elsewhere. 8. Aust. Orn. 8(7); 244-293, HUTTON, FW, 1893, On Dinornis (2) queensiendige, Proc, Linm Soc. NSW 82): 7-10, JENNINGS, J.N. 1959. The coasial geomorphology of King (sland, Bass Strait, in relation to changes in the relative level of land and sea. Rec, Queen Vic, Mus, 1. KURTEN, 8. 1965. The Carnivora of the Palestine caves. Acta Zool. Fenn, WT 1-74. LEGGE, WN, 1907. The emus of Tasmania and King Island Ent fic 116-119. Le SOUEF, BD. 1903. A unique ecological specimen, Emu 3; V14-t15, MCNAB, & 1971 On the ecological significance of Bergman's tule. Bool 52; B45-854. MARCUS, L.F, 1976. The Bingara fauna. A Pleistocene vertebrate fauna fram Murehison County, New Sourh Wales, Australia. Univ. Calif. Publ. Geol. Sci. 114. MILLER, AH. 1962, The history and significance of ihe fossil Cusuerius lydekkeri. Rec, Ausi, Mas. 250) 235-237. MILLER, AH. 1963. Fossil ratire birds of the lace Tertiary of South Australia. Rec §. Aust, Mus, Waa 413-420, MORGAN, A.M. & SUTTON, J. (928. A critical deseription of some recently discovered bones of ihe euinct Kangare Island Emu (Qromaius diementanus), Emu 28 1-19. PARKER, S.A, 1984. The extinct Kangaroo [sland emu, a hitherta unrecognized species. Aull, Bri/. Orn. Cl, 1044], 19-22, RICH, PV, 197s. Antarctic dispersal routes, wandering vominents, and the origin of Austratia’s pon- passeriform avitauna. Mem. Nat. Mus. Wie. 36: 63-126. RICH, PY. 1979, The Dromornithidae, an extinci family of large ground birds endemic to Australia. Department of National Development. Bur. Min. Res. Geol. Geophys. Bull, 184, RICH, T.H., ARCHER, M., PLANE, M.. FLANNERY, T, PLEDGE, WN, HAND, 5S, & RICH, PF 1982. Australian Tertiary mamimal localities. Ja PV, Rich & E.M. Thompson (Eds.). “The Fossil Vertebrate Record of Australasia’. Monash University Offset Printing Unit, Melbourne. Pp. 525-572, RIDPATH, M4: & MOREAU, RE, 1966. The birds of Tasmania: ecology and evolution. fhis 108: 348-393. ROVHSCHILE, W, 19). Gn. the former and present distribution of the so-called Ratitae or ostrich-lke birds. bork. W Inter. Ornith. Cone. Berlin, 1910; 144-169, SCARLETT, R, 1969, On the alleged Queensland moa, Dinornis queenslandiae De Vis. Mem. Ola. Mus, 15(3): 207-212. SCHOLANDER, BA, 1955, Evolution of climatic adaptalion uf Romeolherms, Byol. % 18-26. SCOTT, H.H. 1924. .A note an the King Island Emu, Proe, R. Soc, Tas. 19242 103-107. 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Linn. deposits. Jn D. Hill & A.K. Denmead (Eds.), ‘The Soc. N.S.W. 10; 1207-1241. Geology of Queensland’. J. Geol. Soc, Aust. 7: 393-403. TABLE 2. Statistical summaries of the extant emu Dromaius novaehollandiae (X, mean; s, standard deviation; n, sample size). Range (mm) x s n Skull Length 140 -165 154 6.3 16 Width 58.7- 76.5 68.4 4.1 21 Depth 44.7- 50.7 48.0 1.6 19 Diameter of Foramen Magnum 9.3- 13.1 11.0 1.0 25 Length of Lower Jaw 131 -155 145 6.5 21 Symphysial Length of Lower Jaw 16.4- 23.0 21.0 2.4 23 Sternum Maximum Length 114 -164 143 11.4 25 Maximum Width 104 -141 125 8.2 25 Number of Costal Processes a eS 4 0.49 28 Width of First Costal Process 98.7-134 116 8.2 24 Width of Last Costal Process 76,4-112 § 96.8 8.2 24 Length of Costal Margin 42.4- 66.9 56.0 7.2 24 Length of Sternocoracoidal Process 16.3- 44.7 33.3 8.2 24 Width of Coracoidal Sulci 40.2- 62.2 51.9 5.9 24 Anterior Depth 14.4- 20.4 16.7 2.0 23 Scapulacoracoid Proximal Width 38.9- 55.6 45.9 4.9 20 Maximum Length 151 -187 168 11.3 18 Scapular Length 98.4-127 114 8.7 18 Minimum Width of Coracoid 13.1- 22.0 16.1 2,2 20 Minimum Width of Scapula 5.9- 8.9 73 1,0 19 Clavicles Length 35.2- 53,3 44.0 3.1 12 Maximum Width 3.6- 8.1 5.5 1.2 12 Depth 2.0- 4.9 3.4 0.8 12 Humerus Length 83.1- 98.7 90.3 4.2 20 Proximal Width 5.4—- 8.3 6.4 0.8 20 Proximal Depth 6.2- 7.9 6.9 0.5 20 Ulna Length 57.5- 73.0 64.9 4.5 20 Proximal Width 3.6- 4.8 4.2 0.3 20 Proximal Depth 3.3- 4.8 4.0 0.4 20 Radius Length 55.2- 68.9 63.1 4.1 20 Carpometacarpus Length 36.6- 50.6 43.6 3.6 20 Proximal Width 7.7- 12.3 10,0 1.2 20 Proximal Depth 4.6- 6.5 5.5 0.5 20 Distal Depth 2.5- 4.4 3.4 0.5 19 Manus Pl, Length 10.0- 26.6 13.6 4.1 13 Pl, Proximal Diameter 4.2- 6.9 5.0 0.8 Il P2, Length 4.8- 8.0 6.3 1.4 5 P2, Proximal Diameter 1.9- 3.7 3.0 0.7 5 P3, Length 6.1- 14.9 10.4 2.7 6 P3, Proximal Diameter 14.0- §.2 2.8 1.3 6 THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 103 Range (mm) x s n Synsacrum Length 337 -412 378 24.7 17 Diameter of Acetabular Foramen 12.3- 21.0 17.3 2.5 23 Width across Antitrochanter 90.9-109 101 5.3 22 Maximum Depth 88.7-109.2 99.3 4.2 23 Femur Length 175 -218 203 10.1 22 Proximal Width 62.0- 68.0 64.9 2.0 10 Proximal Depth (Trochanter) 55.8- 63.4 59.7 Zid 10 Diameter of Head 23.5- 29.7 26.4 ss 26 Distal Width 65.4- 79.2 yf Wes 3:2 26 Depth of External Condyle 64.0- 75.4 69.3 2.9 26 Tibiotarsus Length 340 -432 401 21.8 18 Diameter of Shaft, Minimum 19.5- 27.0 23.1 1.8 23 Diameter of Shaft, Maximum 24.6- 34.2 28.1 2.2 23 Proximal Depth 86.2-103 96.2 5.6 9 Proximal Width 47.4- 55.6 52.4 2.8 9 Length of Fibular Crest 74.6-110 90.4 120 9 Depth, Internal Condyle 38.7- 47.4 42.7 2.8 9 Depth, External Condyle 36.5- 45.9 41.8 2.1 26 Width, Distal End 38.9- 49.1 45.9 2;2 25 Fibula Length 231 -305 272 20.9 12 Proximal Width 13.9- 19.0 17.1 1.5 22 Proximal Depth 35.2- 48.7 38.7 pt 22 Tarsometatarsus Length 332 -422 383 18.0 22 Minimum Diameter of Shaft 11.6- 17.3 14.7 15 23 Maximum Diameter of Shaft 16.8- 23.1 19.9 1.5 23 Proximal Width 47.2- 54.0 50.0 2.1 25 Depth of Internal Cotyla 25.4- 27.6 26.6 0.8 9 Depth of External Cotyla 19.9- 23.7 22.0 12 9 Depth of Hypotarsus 36.0- 41.3 38.5 LSS) 8 Distal Width 47.4- 54.6 51.1 2.0 8 Tarsometatarsus, trochleae Width T2 9.0- 11.1 10.0 0.8 9 Width T3 21.9- 28.9 24.9 2.0 24 Width T4 12.2- 14.9 13.6 1.0 9 Depth T2 13.0- 17.6 15.4 1.4 9 Depth T3 19.0- 24.3 22.2 1.8 9 Depth T4 14.3- 17.2 15.6 IQ 9 Pes DII P1 Length 40.7- 52.8 47.4 2.8 14 Proximal Depth 16.3- 21.4 18.5 1,7 15 Proximal Width 13.2- 16.4 14.8 0.8 15 DII P2 Length 17.1- 22.4 19.5 2.1 13 Proximal Depth 11.6- 13.3 12.3 0.6 13 Proximal Width 12.4- 15.6 14.0 1.0 13 DII P3 Length 21.4- 28.2 25.9 22 9 Proximal Depth 9.4- 12.2 11.0 0.8 10 Proximal Width 10.4- 12.9 11.5 0.9 10 DIII P1 Length 58.1- 65.7 60.5 1.9 14 Proximal Depth 21.3- 27.3 23.6 155 15 Proximal Width 25.3- 31.1 275 2.0 15 DIII P2 Length 33.4- 42.8 38.8 2.3 13 Proximal Depth 15.0- 18.9 16.9 0.9 15 Proximal Width 20.3- 25.4 22.5 1.6 14 104 C. PATTERSON & P. V. RICH Range (mm) x s n DIT P3 Length 19.5 — 29.3 23.6 2.8 12 Proximal Depth 12,1- 14.1 13.1 0.7 13 Proximal Width 17.2- 20.8 19.0 0.9 13 DIII P4 Length 26.7- 34.4 30.4 2.1 9 Proximal Depth 11.7- 14.9 12.9 1,0 ll Proximal Width 14.8- 17.7 15.6 0.9 11 DIV P1 Length 33.7- 41.2 38.5 2.0 14 Proximal Depth 16.0- 19.8 17.3 0.8 15 Proximal Width 16.0- 19.2 18.0 0.8 15 DIV P2 Length 14.9- 18.2 17.0 1.0 13 Proximal Depth 11.1- 13.2 12.5 0.6 13 Proximal Width 14.2- 16.0 15.1 0.5 13 DIV P3 Length 10.5- 14.4 12.0 1.1 11 Proximal Depth 9.9- 12.1 11.0 0.6 12 Proximal Width 12.5- 14.9 13.2 0.6 12 DIV P4 Length 6.4— 12,3 9.7 1.6 ll Proximal Depth 8.7- 10.7 10.1 0.6 ll Proximal Width 10.4- 13.4 12.0 0.8 ll DIV P5 Length 19.2- 24,3 22.2 1.6 8 Proximal Depth 9.3- 11.8 10.5 0.7 10 Proximal Width 10.1- 11,5 10.9 0.5 10 Vertebrae Cl Length, Ventral 5.0- 7.9 6.1 0.8 12 Depth of Hypopophysis $.3- 6.1 5.8 0.3 11 Maximum Width across Arch 12,.8- 16.5 14.5 1.4 9 Depth 14.9- 17.7 16,1 0.8 12 Prearticular Surface 6.2- 8.5 Vl 0.7 12 Postarticular Surface 9.0- 11.3 10.4 0.8 10 Dorsal Length §.8- 7,3 6.4 0.5 10 C2 Depth 25,7- 30.9 28.1 1.4 16 Width across Postzygapophyses 20.4- 24.9 22.3 1,3 16 Width across Diapophyses 12.2- 15.5 13.8 1.0 16 Width across Postarticular surface 6.3- 8.7 7.3 0.7 16 Width across Prearticular surface 9.7- 11.4 10.5 0.5 14 Centrum Length 15.9- 20.5 18.3 1.5 15 Length from Pre- to Post Zygapophyses 14.1- 17.9 16.6 Ll 12 C3 Depth 21.0- 26.1 23.7 1.3 15 Postzygapophyses 22.7- 28:2 25.4 1.4 16 Diapophyses 17.7- 23.5 21.0 1.4 15 Postarticular Surface 6.9- 9.6 8.4 0.7 16 Prearticular Surface 8.1- 10.9 9.1 0.8 16 Centrum length 16.5— 21.2 18.9 1.4 16 Pre-postzygapophyses 22.3- 27.8 29.8 1.7 16 C4 Depth 18.1- 22.6 20.2 1.3 16 Postzygapophyses 22.1- 26.4 24.5 1.3 16 Diapophyses 21.6- 26.8 23.6 1,3 16 Postarticular Surface 9.5-— 13.0 11.3 1.2 16 Prearticular Surface 7,9- 12.0 10.3 0.9 16 Centrum Length 20,.5- 25.3 22.7 1.6 16 Pre-postzygapophyses 26.4- 32.3 29.4 19 16 C5 Depth 14.9- 19,3 17,1 1.2 17 Postzygapophyses 12.8- 19.5 17.0 1.6 17 Diapophyses 25,0- 28.6 26.2 1.1 l7 Postarticular Surface 13.6- 19.7 16.0 1.6 17 Prearticular Surface 10.5- 15.7 13.3 1.6 17 Centrum Length 22.4- 26.7 25.2 13 17 Pre-postzygapophyses 30.8- 40.6 36.0 2.5 17 C6 Depth 13.9- 19.7 16.5 1.3 17 Postzygapophyses 11.6- 16.4 13.6 1.2 17 THE FOSSIL HISTORY OF THE EMUS, DROMAJUS (AVES: DROMAIINAE) Range (mm) ¥ s n Diapophyses 17.4- 31.1 27.8 3.1 17 Postarticular Surface 16.8- 20.6 18,7 1,3 17 Prearticular Surface 14.5- 19.9 17.4 1.7 17 Centrum Length 25.3- 29.4 27.8 1.3 17 Pre-posizygapophyses 35,.7- 42.2 38.8 1.8 17 C7 Depth 15.0- 19.0 16.9 1.2 17 Postzygapophyses 11.5- 17.9 14.0 1.8 17 Diapophyses 27.8- 32.9 30.4 1.7 17 Postarticular Surface 1§.2- 20.1 18.0 1.6 17 Prearticular Surface 18.9- 24.1 21.0 1.4 17 Centrum Length 28.5- 32.3 30.7 1,3 17 Pre-postzygzapophyses 35.7- 41.4 38.4 1.5 17 C8 Depth 16.2- 19.9 18.0 1.1 17 Postzygapophyses 12.1- 19.7 16.8 2.2 17 Diapophyses 18.4- 34.2 30.4 3.6 17 Postarticular Surface 14.4- 19,1 16.9 1,4 17 Prearticular Surface 16.8- 32,2 20.8 3.5 17 Centrum Length 23.6- 35.6 32.9 2.8 17 Pre-postzygapophyses 36.5- 40.5 38.6 1.3 17 C9 Depth 18,2- 22,7 20,2 1,2 17 Postzygapophyses 16.3- 25.0 20.8 2.0 17 Diapophyses 28.4- 34.5 31.2 1.8 17 Postarticular Surface 13.4- 17.9 15.4 1.3 17 Prearticular Surface 16,1- 21.7 19.0 1.4 17 Centrum Length 34.2- 38.7 36.2 1.4 17 Pre-postzygapophyses 38.2- 45.3 40.4 1.9 17 C10 Depth 19.7- 23.6 21.7 1.2 17 Postzygapophyses 19.8- 27.2 22.9 1.6 17 Diapophyses 28.8- 34.7 31.0 1.8 17 Postarticular Surface 12.6- 16,7 14.8 1.2 17 Prearticular Surface 15.2- 21.1 17.5 1.7 17 Centrum Length 36.0- 40.8 38.4 1.6 17 Pre-postzygapophyses 40.5- 50.6 44,2 2.3 17 Cll Depth 21.0- 24.9 23.2 1.2 17 Postzygapophyses 21.2- 27.5 23.4 1.5 17 Diapophyses 29.4- 34,7 31.5 1.6 17 Postarticular Surface 13.8- 18.4 15.4 1.4 17 Centrum Length 37.7- 43,2 40.7 1.7 17 Pre-postzygapophyses 43.6- 53,2 47.3 2.2 17 Prearticular Surface 15.1- 19.9 16.9 1.6 17 C12 Depth 23.0- 27.1 24.6 1.3 17 Postzygapophyses 20.2- 27.5 23.8 1.7 17 Diapophyses 30.1- 35.0 25.0 1.5 17 Postarticular Surface 14.6- 19.4 16.5 1.5 17 Prearticular Surface 15.4- 23,1 17.9 2.3 17 Centrum Length 39.5- 44.4 42.4 1.5 17 Pre-postzygapophyses 44.9- 53.7 49.7 2.3 17 C13 Depth 23,.3- 28.2 25.7 i 17 Postzygapophyses 20.3- 27.0 23.8 1.9 17 Diapophyses 30.7- 37.3 33.7 1.7 17 Postarticular Surface 16.2- 20.6 17.8 1.4 17 Prearticular Surface 15.5- 22.6 18.9 1.9 17 Centrum Length 40.4- 45.9 43.6 1.6 17 Pre-postzygapophyses 47.7- 54.4 51.9 1,9 17 C14 Depth 24.3- 29.7 26.9 1.5 17 Postzygapophyses 20.7- 26.4 23.9 1.7 17 Diapophyses 32.0- 38.5 35.2 1.6 17 Postarticular Surface 17,3- 22.2 19.2 1.4 17 Prearticular Surface 17.1- 23.4 20.4 1.9 17 105 106 C. PATTERSON & P. V. RICH Range (mm) x s n Centrum length 41.0- 46.5 44,3 1.7 17 Pre-postzygapophyses 48.6- 56.6 §2.8 2.3 17 C15 Depth 25,6- 32.2 28.6 2.0 17 Postzygapophyses 21.4- 27.4 24.1 1.6 17 Diapophyses 34.5- 40.9 37,1 2.0 17 Postarticular Surface 19,0- 23.5 21.2 1.4 17 Prearticular Surface 17.6- 25.9 22.0 2.1 17 Centrum Length 41.2- 47.4 44,9 1.8 17 Pre-postzygapophyses 49.8- 58.4 53.6 2.5 17 C16 Depth 27.5- 33.5 30.5 2.1 16 Postzygapophyses 22.4- 28.1 25.4 1.7 16 Diapophyses 36,9- 45.7 40.7 2.6 16 Postarticular Surface 20.5- 25.7 23.3 1.6 16 Prearticular Surface 20,4- 29.2 24,1 2,2 16 Centrum Length 41.9- 48.5 45.3 1.9 16 Pre-postzygapophyses SL.1- 60.8 55.0 2.8 16 C17 Depth 29,3- 41.1 33.8 3.2 17 Postzygapophyses 25.1- 30.4 27.2 1.9 17 Diapophyses 41.8- 51.4 45.7 3.2 17 Postarticular Surface 20.9- 27.6 24.5 1.9 \7 Prearticular Surface 23.5- 32.4 26.5 2.2 17 Centrum Length 42.5- 48.7 45.5 2.0 17 Pre-postzygapophyses 50.0- 61.2 55.6 2.8 v7 C or V18 Depth 34.0- 56.3 41.2 5.5 16 Postzygapophyses 25.3- 32.7 28.4 2.0 16 Diapophyses 46.5- 68.4 53.6 5.7 16 Postarticular Surface 21.9- 29.0 25.3 1.8 16 Prearticular Surface 22.4- 31.1 27.6 2.6 16 Centrum Length 42.3- 48.2 45.1 1.8 16 Pre-postzygapophyses 48.4- 60.6 55.4 3.0 16 V19 Depth 43.0- 66.4 50,7 6,3 17 Postzygapophyses 25.2- 31.3 28.3 1.7 17 Diapophyses 56.5- 70.6 61.7 4.0 17 Postarticular Surface 21.6- 27.8 24.9 1.8 17 Prearticular Surface 23.4- 33.1 27.4 2.5 17 Centrum Length 40.9- 47.5 44.4 1.9 17 Pre-postzygapophyses 51.4- 58.4 54.2 3.1 17 V20 Depth 50.7- 77.1 62.6 5.7 7 Postzygapophyses 23.7- 30.2 26.5 1.9 17 Diapophyses 62.1- 81.8 67.5 4.6 17 Postarticular Surface 20.5- 25.6 23.3 1.5 17 Prearticular Surface 23.7- 34.0 27.6 2.7 17 Centrum Length 40.4- 47.6 42.8 1.8 17 Pre-postzygapophyses 49.7- 58.3 54.2 2.4 17 V21 Depth 56.6- 78.1 68.0 5.5 17 Postzygapophyses 23.4- 29,1 26.2 1.6 17 Diapophyses 61.3— 82.0 68.2 4.8 17 Postarticular Surface 19.6- 26.5 22.7 1.5 17 Prearticular Surface 21.4- 29.4 25.3 2.0 17 Centrum Length 37.2- 46.3 42.1 2.4 17 Pre-postzygzapophyses 47.9- 57.0 51.9 2.7 17 V22 Depth 57.1- 74.9 64.7 6.1 17 Postzygapophyses 22.4- 30.5 26.7 2.0 17 Diapophyses 61.0- 76.8 67.0 3,7 17 Postarticular Surface 19,8- 29,1 23.3 2.3 17 Prearticular Surface 21.9- 29.1 24.5 1.9 17 Centrum Length 38.3- 45.5 41.5 2.2 17 Pre-postzygapophyses 47.2- 57.8 50.7 2.9 7 THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) a Range (mm) V23 Depth 55.7- 72.9 Postzygapophyses 25.8- 33.1 Diapophyses 60.9- 74,4 Postarticular Surface 21.0- 26.7 Prearticular Surface 20.3- 29.0 Centrum Length 37.8- 43.7 Pre-postzygapophyses 43.8- 51.6 V24 Depth 60.2- 79.9 Postzygapophyses 26.1- 33.8 Diapophyses 62.1- 75.5 Postarticular Surface 19.8- 28.4 Prearticular Surface 21.3- 28.8 Centrum Length 37.5- 46.1 Pre-postzygapoplyses 43.2- 52.1 V25 Depth 63.7- 85.9 Postzygapophyses 27.9- 40.2 Diapophyses 43.4- 76.2 Postarticular Surface 20.3- 27.3 Prearticular Surface 20.4- 30.3 Centrum Length 36.8- 44.4 Pre-postzygapophyses 42.3- 52.9 V26 Depth 68.5- 87.8 Postzygapophyses 26.2- 40.9 Diapophyses 59.7- 75.6 Postarticular Surface 23.0- 30.4 Prearticular Surface 23.1- 38.7 Centrum Length 33.3- 49.4 Pre-postzygapophyses 41.2- 50.7 x nw n VA NENOwNY ND RFPANROR Ww “ ~ NN es ee * a New pw NNN ANY NN? CAwWWORUONMN NOUCIBON $C wi ~~ TABLE 3. Measurements (in mm) of skull material of fossil emus (Dromaiinae) from Australia. SPECIMEN NMV P157345 NMV P157350 NMV P157353 SAM P17834 Diameter, Foramen Width Depth Magnum 75.0 (est.) 47.4 10.8 >65.2 48.5 10.7 Lower Jaw, Symphysial Length 21.5 24.5 107 108 C. PATTERSON & P. V. RICH TABLE 4. Measurements (in mm) of vertebrae of fossil emus (Dromaiinae) from Australia. SPECIMEN MM F16786 MM F16797 NMV P157346 NMV P157349 NMV P15735] NMYV P157352 NMV P157359 P157367 NMV P157368 NMV P157369 SAM P17589 SAM P18246 SAM P18673 SAM P18830 SAM P18247 AMNH 9678 AMNH 9678 AMNH 9678 AMNH 9678 AMNH 9678 AMNH 9678 UCMP 56642 UCMP 56855 Length of Centrum 37,7 38.8 39.3 36.0 24.5 31.2 Length across Zygopop. 46.3 42.1 46.3 Vertebral Number V24? V26 or 25? V21 or 22 V23 or 24 c7? (juv) C11-13? Guy) Measurements See ene SS NN SS Oe Width Posterior Anterior Depth Postzyg. Diapop. Articulation —_ Articulation _ 42.2 (est.) = 25.5+ 30.0 (est.) -- 31.8 — 27.0 27.2 _ 23.6 _ 20.2 23,2 — — 62.8 (est.) 21.4 22.2 (est.) 14.3 9.7 24.6 16.9 12.8 18.2 17,7 — 11.9 12.4 (est.) 22,5 26.1 21.7 8.7 9.2 C3 — = 68.0 (est.) — — V25 or 26 _ 19.8 46.8 (est.) _ _ V20-21 (juv) 16.6* 16.6* 11.0 16.2 (est.) ?Cl1_— (juv) - _— _ 21.4 (est.) 26.0 (est.) V22-26 (juv) 29.7 23.1 39.7 21.8 23.1 C16 (15-17?) 19.9 23.9 22.5 10.7 9.8 C4 30.5. 27.5 37.9* 20.6 18.8 + C15-C17 _ 33.4 _ 27.4 > 25.6 V26? — _ 68.0 (est.) 24.8 V22 or 23 _ 26.0 (est.) -- _ — C2 24.2* 30.0 (est.) 22.0 (est.) = 11.2 (est.) C3 >20.2 29,1 25.9 12.6 11.7 C4 18.0 16.8 32.9 22.8 19.5 C6? 19.1 16.3 34.9 23.1 23.3 C7? 21.7 oo 36.4 = 22.5 C9? _ _ _— _ — V21 or 22 _ _ — 23.3 25.7 V22 or 23 —eeeee_ TABLE 5. Measurements (in mm) of vertebral ribs of fossil emus (Dromaiinae) from Australia. Seen eee Width of Facets >36.8 ~34.3 > 28.0 SPECIMEN NMV P157354 NMV P157358 NMV P157362 SAM P18107 SAM P18784 SAM P22812 (in part) UCMP 60560 SE 29.3 34.1 38.6 34.8 THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 109 TABLE 6. Measurements (in mm) of sterna of fossil emus (Dromaiinae) from Australia. Width Width Length Length Width Number of of first of last of of Sterno- of Cora- Maximum Maximum _ Costal Costal Costal Costal coracoidal coidal Anterior SPECIMEN Length Width Processes Process Process Margin Process Sulci Depth D. novaehollandiae NMV P157347 _ 124 3 118 — — — ~52,6 14.6 NMV P157355 — 112 4 109 90.9 49,7 16.4 — — D. sp. AMP 25218 147 118 4 127 109 64.2 —_ 70.5 17.9 Ow TABLE 7. Measurements (in mm) of synsacra of fossil emus (Dromaiinae) from Australia. ee ee eee ee ee Diameter of Length of Depth of Acetabular Width across Acetabular Acetabular SPECIMEN Foramen Antitrochanter Complex Complex D. novaehollandiae NMV P157361 — >83.6 56.3 _ SAM Unreg. 15.5 _ — SAM P16501 18.0 108 — —_ SAM P17767 12.2 104 60.7 33.8 UCMP 56333 17.4 — 62.3 36.8 TABLE 8. Measurements (in mm) of femora of fossil emus (Dromaiinae) from Australia. ee Proximal Depth of Proximal Depth Diameter Distal External SPECIMEN Length Width (Trochanter) of Head Width Condyle D. novaehollandiae HM B775/869 190 oe — — 59.6 61.2 R HM B801/934 — — — — — 76.2 d,R RHT 1064 190 65.1 — 27.6 — — R SAM Unreg. 7 _— _ 27.4 >76.3 76.7 SAM P17104 _ ~67.2 ~61.2 ~28.0 _ — pL SAM P22812 204 67.4 61.4 25.5 68.8 — L AMNH 9677 _— — — — 83.3 70.7 dR D. cf. ocypus UCMP RASS176 190 59.5 — — — — pb oe 110 C. PATTERSON & P. V, RICH TABLE 9. Measurements (in mm) of tibiotarsi of fossil emus (Dromaiinae) from Australia. Length, Depth, Depth, Width, __Diameter of Shaft. proximal Proximal Fibular Internal External Distal SPECIMEN Length Minimum Maximum Depth Width Crest Condyle Condyle End D. novaehollandiae AM 49713 — _— —_— 113 57.8 — a a — p,R AM ‘B’ — — _— — a —_— 45.1 — 46.6 d,L AM F10949 — — _— — = — >35.0 31,2 38.1 d,L AM P5802 — — _ _ = 41.0 42.1 47.3 d,R MM F16775 —_ — — _— — _ 44.9 44,3 49.5 d,R NMV P44011 = 22.1 26.6 >76.0 >44.1 93.1 _ — — R NMV P150013 — 24.9 32.7 — a 95 44.7 40.2 50.4 d,L NMV P157356 —_— _— _— >92.2 47.4 _ — — p,R NMV P157357 — — os — — _ 39.1* 39.1 43.3 d,L NMV P157360 — — = — ~ — >39.3 39.3 41.8 d,L NMV P157365 — 21.1 29.2 >91.5 49.2 96.4 _ _ — »p,R QM F1652 — — oa — 56.1 >80.5 — a — p,L SAM P17149 _— — _ — — — 42.7 — 43.6 d,R SAM P18829 _ _ — — — > 38.8 41.3* >37.5 d,R SIAM 61 384 23.3 32.3 102 56.9 88.6 45.8 43.9 51.8 L UCMP 53825 (RHT6) —_— —_ _ _— — os 37.0 35.1 37.4 d,L UCMP 53825 (RHT25) _ a a = — _— 37.0 34.9 35.6 d,L UCMP 56845 -- = = — = _ — 38.8 40.6+ d,L WAM 68.5.34 ve 21.6 26.3 — = 107 — _ _— L D. patricus’” MM F12074 o — — —_— a —_— 35.5 34,3* 41.3 d,R QM F5547 - — — — 57.6 >90.2 — a — p,R QM F5548 — ~ _— a — — 46.2 45.7 48.6 d,L D. cf. ocypus UCMP RASS5182 _ _ — _ — _ 43.0 40.8 46.0 d,R D. gidju SAM P26779 = = — — — _ 31.8 31.3 32.0 d,L THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 11 TABLE 10. Measurements (in mm) of tarsometatarsi of fossil emus (Dromaiinae) from Australia. Depth, Depth, _ Diameter of Shaft = Proximal Internal External Depth, Distal SPECIMEN Length Minimum Maximum Width Cotyla Cotyla Hypotarsus Width D. novaehollandiae AM Unreg. _— — a 56.7 25.9 24.5 42.5 — p,R AM “C’ _— — — 53.9 25.2 21.3 40.5* — pL AM F771 — 11.8* 20.5 — — — — 47.5 d,R AM F18935 — 14.0 19.4 — — — — >46.2 dL (juv) AM P58026 — _ 53.0 23.2* 19,1* 39.5* — p,R NMV L5 — — — —_— a = — 54,17 d,L NMV P44012 — 14.1 21.9 >42.3 = _— >34.0 52.4 R NMV P44013 — 16.1 22,2 48.3 26.3 22.5! >35.1 55.9 R NMV P44014 — 14.9 19.8 49.8 24.2 21.4 37.5? §2.2 R NMV P44015 _ 14.0 19.5 46.5 24.0' 19.8 >37.0 — R NMV P44016 — 15.5 20.4 — 23.0 >21.8 > 38.2 51.2 L NMV P44017 _ 13.3 21.7 — — - >50.5 dL NMV P44018 — 13.8 23.1 — — = -- 54.0° d,L NMV P48392 _— 12.2 21.0 — — — — — dR NMV P150015 _ 16.7 23.6 —_— — — — — d,R NMV P157343 oo 13.0 19.3 >39.0 = a >32,4 — L NMV P157344 _ 14.2 19.6 = — — — - d QM F1143 (in part) = — 15.4 25.7 _ oe = — 56.1 L SAM P13118 a 12.6 -- _ = — — — 4d,R » SAM P17816 — 16.0 23.4 — _ — — 53.5 d,L SAM P18693 _ 16.7 23.9 49.0 (est.) = a — 56,3 L UCMP 53835 13.9 18.7* — _ — — 39.0 d,R UCMP 56313 — 13,57 os — — — — 4d,R WAM Unreg. 190 — — a 33.8 — — — — (juv) WAM 68.5.34 >330 16.6 18.2 —- —_— _ os _ L D. gracilipes’ QM F1142 _ 10.9 21.5 = - — _ — 4d,L (juv) ‘Metapteryx bifrons’ QM F1135 _ — os — = _— 34.1 d,L (juy) D, ocypus SAM P13444 330 — — 47.1 21.2 20.9 35.5+ 53.0 R D. gidju SAM P26779 — — = 35.0 18.9 14,.8* 26.9+ — pL Dy sp. AM ‘A’ aa 10.3 17.1 os — _ == d (juv) AM F10850 — — _ 46.4 ~ — = — pL NMYV P35578 — 16,2 28.0 — — —_ _ — 4d,R Aves, indet. SAM P11552 at 21.2 28.0 _ _ — — 63.7 dL um 112 C. PATTERSON & P. V. RICH TABLE 11, Measurements (in mm) of the distal ends of tarsometatarsi of fossil emus (Dromaiinae) from Australia. —_----—————e—”:,—O0W0—_?ee—— — Width, Width, Width, Depth, Depth, re T3 T4 T2 T3 D. novaehollandiae AM F771 9.1 23.8 12.2 12.1 20.3 AM F18935 a 24.1 12.3 os 24.1 QM F1135 5.9 15.7 45. 8.8 13.8 QM F1143 (in part) 10.9 27.2 14.3 15.3 25.0 QM F1143 (in part) _— 23.0 a _ 22.6 NMV L5 > 10.5 24.8+ 13.2+ > 14.0 25.2+ NMV P44012 > 9.2 24.3 12.8+ 13.1+ 22.3* NMV P44013 9,8+ 24.3 14.5+ 15.1 23.8 NMV P44014 10.9 26.4 13.2 14.8 22.5 NMV P44015 _ — = _— 21.47 NMV P44016 10.3 25.9 13.3 13.8 21.4 NMV P44017 > 8.8 >23.3 >13.1 14.8 23.3 NMV P44018 > 9.0 25.0+ 14.8 13.4+ 20.4+ NMV P48392 — —- >10.5 >21.5 NMV P150014 1133 28.3 — 15.0' 24.0 NMV P150015 14.9% 25.1 _— 11.6 29.8 SAM P13118 _ _— 11.9 — 20.2 + SAM P17816 10.0 27.8 >13.3 15.0 23.1 SAM P18693 12.6 29.8 15.6 >15.3 25.6 UCMP 53835 — > 18.4 — _ UCMP 56313 10.7 27.9 — 15.3 23.1 D. §racilipes’ QM F1142 — 16.6 = — 16.5 D. ocypus SAM P 13444 10.6 27.8 13.5 14.7 21.9 D. sp. AM ‘A’ —_ 16.5 _ —_ -14.7 QM QA205 _ ~16.1 _— — >14.4 QM QA416 13.3* — _ 23.9* _ QM QAS505 8.5 17.5 _— 11.9 16.8 UCMP RAS5397 (in part) — 15.7+ — _— 14.0* Aves indet. SAM P11552 — >26.3 > 14.5 — >27.5 * perhaps T4 THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMAIINAE) 113 TABLE 12. Measurements (in mm) of phalanges of fossil emus (Dromiinae) from Australia. Proximal Proximal SPECIMEN Phalanx Length Depth Width Element D. novaehollandiae SAM P18059 P1,DII 45.0 20.0 14.8 L SAM P18248 P2,DII 37.6 17.8 23.8 R SAM P18249 P1,DIV 38.2 17.6 18.8 R UCMP 53832 P4,DII or DIV 20.6 10.6 10,5 UCMP 53832 P1,DII a 16.7 13.4 p UCMP 53832 P2,DIV 16.3 12.3 14.1 L UCMP 53833 P1,DIV 34.2 16.1 17.8 L UCMP 53833 P5,DIV? 20.0* 9.6 8.6+ UCMP 55983 P1,DIII 53.2 20.8 25.7 L UCMP 55983 P2,DIII 36.2 13.9 21.3 L UCMP 56849 P1,DIU 43.0 18.9 17.0 L UCMP 94679 P1,DI1 48.4 18.9 24.0 UCMP 94680 P2,DIII 37.0 18.8 21.4 L D. gidju SAM P26779 P1,DII 33.1 14.2 12.9 L SAM P26779 P2,DII 23.5 11.8 11.6 ie SAM P26779 P3,DII 19.3 10.1 8.9 L SAM P26779 P1,DII1 45.1 16.7 19.5 L SAM P26779 P2,DIII 31.7 12.9 15.2 L SAM P26779 P3,DII 22.5 10.5 13.0 L SAM P26779 P4,DIII 18.4 9.8 10.5 L SAM P26779 P1,DIV 29.2 13.0 14.0 L SAM P26779 P2,DIV 16.1 10.6 10.8 L SAM P26779 P3,DIV 10.8 9.2 9.4 L SAM P26779 P4,DIV 8.8 7.9 8.2 L SAM P26779 P5,DIV 15.6 8.8 7.7 L D. sp. QM QA504 P1,DIII 41.5 15.3 16.8 R UCMP RAS 5397 P2,DII 19.8 11.9 10.9 L TABLE 13. Measurements (in mm) of the fibula of fossil emus (Dromaiinae) from Australia. SPECIMEN Proximal Width Proximal Depth Element NMV P157363 17.0 36.4 p,L 114 C, PATTERSON & P. V. RICH “a tem FIGURE 1. Dromaius gidju, n. sp. Type from the Wipajiri Fm. Leaf Locality, Lake Ngapakaldi, Kutjamarpu fauna, Miocene. A,B, stereo pair of pes, dorsal view. C,D,E,F, tarsometatarsi in posterior (C), anterior (D) and proximal (E,F, stereo pair) views. G,H,I,J,K, distal left tibiotarsus in anterior (G), posterior (H), distal (I), internal (J), and external (K) views. Scale indicates 1 cm. THE FOSSIL HISTORY OF THE EMUS, DROMAIUS (AVES: DROMALINAE) 1s TT FIGURE 2, Mid-Cainozoic emu fossils from northern South Australia and living casuariids. A-D, posterior views of tarsometatarsi of (A,B) the extant Dromaius novaehollandiae, (C) D. ocypus (SAM P13444), and (D) the extant Caswarius unappendiculatus (from Miller, 1963). E,F, stereo pair in anterior view, tarsometatarsus, the type specimen of Dromaius ocypus, SAM P13444, Pliocene, Lawson-Daily Quarry, Mampuwordu Sands, Lake Palankarinna, Palankarinna fauna, G, distal tibiotarsus in anterior view of D. cf. ocypus, RAS 5182, Pliocene. H,!, femur of D. ef, ocypus, in posterior (H) and anterior (I) views, RAS 5176, Pliocene, 116 C. PATTERSON & P. V. RICH Tem FIGURE 3. A variety of late Cainozoic emu fossils. A, sternum of Dromaius sp. in dorsal view, AM F25218. B, pelvic fragment of D. novaehollandiae in lateral view, SAM P16501. C, distal left femur in posterior view of D. novaehollandiae, HM B801/B934. D,E, proximal right tibiotarsus in lateral (D) and proximal (E) views. F, left tibiotarsus in anterior view, SIAM 61. 7 tem tem FIGURE 4. A variety of late Cainozoic emu fossils. A, distal right tibiotarsus in anterior view, ane of the original type specimens of Dromuius putricus'De Vis, QM F5548. B, QM F120 considered by CW. De Vis to be a coracoid af D. patricus, but non-avian. C, distal left tarsometatarsus of ‘Metapteryx bifrons' De Vis in anterior view, QM F1135, originally designated a kiwi but actually a juvenile D. novaehollandiae. D, distal tarsometatarsus of D. gracilipes’ De Vis in anterior view, QM F1142, a juvenile D. novaehollandiae. E, distal left tarsometatarsus of Aves indet. ef. dromornithid in-anterior view, SAM P1552, assigned previously as emu. F,H, tarsometatarsus fragment of a small D. novaehollandiae in anterior (F) and posterior (H) views, AM ‘A’, no locality or age data available, G,I,J, second and third trochleae of a small Dromuwius in posterior (G), distal (1), and posterior (J) views, QW QA505. K-N, phalanges of D. novaehollandiae in dorsal views. (K) phalanx 1 digit Lf, UCMP 94679; (L) phalanx 2, digit Ll, UCMP 94680; (M) phalanx 1, digit Il, UCMP 56849; and (N) ungual phalanx 4, digit 111, UCMP 60563. ECHIURANS FROM AUSTRALIA (ECHIURA) BY S. J. EDMONDS Summary Seventeen species of euchiurans [i.e. echiurans] are recorded from Australia of which nine are redescribed, and none are new. One, Anelassorhychus porcellus adelaidensis, is given new status. Listriolobus bulbocaudatus Edmonds, 1963 is now considered a junior synonym of L. brevirostris Chen & Chen Chang, 1958, Ochetostoma myersae Edmonds, 1963 a junior synonym of O. baroni Greeff, 1879 and Austrobonellia mjoebergi (Fischer, 1921) a junior synonym of Pseudobonellia biuterina Johnston & Tiegs, 1919. A key to the Australian species is given. ECHIURANS FROM AUSTRALIA (ECHIURA) S, J. EDMONDS EDMONDS, S. J. 1987. Echiurans from Australia (Eehinra). Reo. S. Aust. Afus, 52(2) 196138. Seventeen species of euchiurans are recorded from Australia of which ning are redescribed, and none are new. One, Anelassorhynehus porcellus adelaidensis, is given new status. Lisiriolohus bulbocaudatus Edmonds, 1963 is now considered a junior synonym of L. brevirostris Chen & Chen Chang, 1958, Qchelostorne myersue Edmonds, 1963 a junior synonym of O. baron Greeff, 1879 and Austrobonellia mjoebergi (Fischer, 1921) a junior synonym af Psexdoboneliia hiuterina Johnston & Tiegs, 1919. A key to the Australian species is given. 5. J. Edmonds, Honorary Associate, South Australian Museum, North Terrace, Adelaide, South Australia 5000, Manuseript reeetved 2) October 1986, Echiurans are soft-bodied, protostomous, coclomate, worm- to sausage-shaped, marine Inver- lebrafes thal resenible annelids and sipunculans. Because they are largely subtidal and occur in burrows and protected places echiurans are not always easy to find. They have, however, been collected in tropical, temperate and polar seas and are known from the littoral to the ultra-abyssal resions of the oceans. More than 150 species have heen described. Records of Australian echiurans are few and scattered (Haswell 1885, Whitelegge 1899, Augener 1903, Hedley 1906, Dakin 1916 and 1952, Fischer 1919 and 1921, Johnston & Tiegs 1919 and 1920, Monro 1931, Edmonds 1960, 1963, 1966 and 1982, Nielsen 1963, Dartnall 1970 and 1976) and only 16 species have so far been reported, of which three are well known, In Australia echiurans have been found in burrows in mud and sand, in limestone rocks and in coral, in tangled roots of sea-grasses, under stones and in cracks and fissures in rocks, Some have been dredged. The best known Australian echiurans (and ones that can be readily collected) are Merabonellia haswelli, Anelassorhyn- chus parcellus adelaidensis and Ochetostoma australiense. Scuba divers report the presence of Jarge numbers of the first in shallow water at Encounter Bay, S,A,, between Wright J]. and the Bluff and from the islands comprising the Banks Group in Spencer Gulf, S.A, and of the second species from Coobowie and Bdithburg in St Vincent Gulf, S.A. Large numbers of O. austra/iense occur at Caloundra and Dunwich, Qu., the species feeding from the surface of intertidal mud flats at low water, In other parts of the world they have alsa been found in the empty shells of molluscs and in sand dollars, Most of the specimens examined in the present study were found intertidally by collectors or subtidally by divers and are now in the collections of Slave Museums, The classification used in this paper ts that outlined by Stephen & Edmonds (1972), which in tum is based on that of Fisher (1946). LIST OF AUSTRALIAN ECHIURANS The species marked ‘*' have not been examined by the author. Records of species marked “?" are considered doubtful. Family Bonelliidae Metabonellia haswelli (Johnston & Tiegs) Pseudobonellia biuterina Johnston & Tiegs 7* Archibonellia michaelseni Fischer + Zenkevitchiola brevirosiris Murina * Sluilerina alba Murina * Vitjazema ultraabyssalis Murina * Protohonellia papillasa Murina Family Echiuridae Anelassorhynchus porcellus porcellus Fisher Anelassorhynchus partellus adelaidensis Fdmonds 7, statis. ”* Anelassorhynchus vegrandis (Lampert) Arhynchite hiscocki Edinonds Listrialobus hrevirosiris Chen & Yeh Chen-Chang 7* Listriolabus sorbillans (Lampert) Ochetostoma baroni (Greeff) Ovheiostoma australiense Edmonds Thalassema sydniense Edmonds Family Ikedaidac Ikeda sp. The following abbreviations are used in tis paper} AMS (Australian Museum, Sydney), MV (Museum of Victoria, Melbourne), SAM (South Australian Museum, Adelaide), WAM (Western Australian Museum, Perth), TMH (Tasmanian Museum, Hobart), N.SW, (New South Wales), Qu. (Queensland), S.A. (South Australia), Tus, (Tasmania), Vie. (Victoria), WA. (Western Australia,), The anatomy of a ‘generalised’ echiuran is shown in Fig. 1. 120 8. J. EDMONDS Key To GENERA OF AUSTRALIAN ECHIURANS The doubtful genus Archibonellia has not been included in the key; it would key near Pseudobonellia, I Erpbeyscis fifty seth o ei ic eee pes | 2 Proboscis not bifid. .-.--,..............02.. 3 2. One nephridium, with distally placed, stalked nephrostome..,............4- Metabonellia Two nephridia with a small sac carrying a male between them, ,....,.....,. Pseudobonellia bands ..... J ovur: Eile nga e 5.9 MONTE edbs-t 4 Longitudinal musculature of body wall not grouped Hato Mandy): ppt ok. stay hte nate hh wens 6 4. Nephridia less than 10 and arranged in pairs; trunk Jong pr SHOP 5 ws: ASC L9b Dee. 300k ee 5 Nephridia very numerous and not in pairs; trunk very Jong, up to 400 mm... ..... 0.2. eee Ikeda 5. Intervals between longitudinal muscle bands traversed by small bundles (fascicles) of oblique muscles Bhd § Seay. 5 5 SRC hae. 38 Sb oo Ochetostoma Intervals between longitudinal muscle bands not traversed by small bundles of oblique muscles + Annet tiveeeerereesaeeess Listriolobug 6. Nephrostome distal ................. ‘Vitjazema Nephrostome basal ........0....0000e00s cease 7 7, Ventral setae none........ 2... cece cee eee 8 Ventral setae tWo os eid ee eee eu eesewaene 9 8. Anal glands long and slender; posterior region of proboscis surrounded by a collar Zenkevitchiola Anal glands bushy or feathery; posterior region of proboscis modified to form a cup... Sluiterina 9, Nephrostomal lips long and spirally coiled...,... ann! $0-seaee eevee eees es Anelassorhynchus Nephrostomal lips long or short but not spirally coiled Le ee oon oe ring t ae nibiitdeteebictis tenet 10 10. Anal glands tubular or sac-like.,.....,...... il Anal glands feathery or bushy .... Protobonellia 11. Proboscis with expanded or fan-like anterior extremity Pticleite ati p tinge eters gpg bt we teen 8 Arhynchile Proboscis without expanded or fan-like anterior CREPES sey ate Mey celI ae om Thalassema FIGURE 1. A generalized diagram to show some of the internal anatomy of an echiuran; dorsal view. Most of the much coiled intestine has been omitted. av, anal DESCRIPTION OF GENERA AND SPECIES vesicle; c, caecum; cf, ciliated funnel; cg, ciliated groove: cl, cloaca; dy, dorsal vessel; 2, gonad; i, intestine; in, Genus Metabonellia Stephen & Edmonds interbasal muscle; Ib, lateral vessel; m, median vessel; n, nephridia; ne, nerve cord; nl, nephrostomal lips; nv, _Afefahonellia Stephen & Edmonds, 1972, p. 394, neurointestinal vessel; 0, oesophagus; p, pharynx; rv, ring vessel; s, siphon; vy, ventral vessel. Type-species: Bonellia haswelli Johnston & Tiegs, 1920, (Bonellia gigas Nielsen, 1963, which was named as type by Stephen & Edmonds (1972) is now AUSTRALIAN ECHIURANS 11 considered to be 4 junior synonym of B. haswelli). The genus contains only one known species. Diagnasis Female of medium to large size, with proboseis bifid, grooved and ciliated on ventral surface. Pale to dark green. Two vevitral setae. Single nephridium with nephrostame on a short stalk placed about two chirds of way along vephridium towarils its distal extremity. Anal vesicles branching. Intestinal siphon present, Male worm-like, as long as 20 mim and without setae, Pound in nephridium of female, Motaboneiia haswelli (lohnston & Tiegs) (Figs 2-4, 18) Bonelfig fleswelli Johnston & Tiegs, 1930, pp. 73-76; Ednionds, 1940, pp. 95-96. Bonellia gigas Nielsen, 1963, pp. 41-67, Bonellia rasmanicu Dartnall, 1970, pp. 69-74. Metabonellia haswelli Stephen & Edmonds, L972, pp. 394-395; Darthall 1976, pp. 1041-1043; Edmonds 1982, pp, 314-316, Halowpe: AMS Gll22 and paratype O)261. Type locality, Sydney Harbour, NSW, ‘under stones just above the limit of law water’. Previous Australiun reeards. Jolnston & Tiegs (1926), Edmonds (1960), Niclsen (1963), Durtnall (1976). Description of femule trunk: Medium to liurge, shape vartable (sausage {a sub-ovoidal), light to dark green, length 20-80mm (Nielsen 1963; length 80-120mm, maaimum width 40mm), Skin smooth to rough, usually covered anteriorly und posteriorly with near circular rows of rather Flattened papillac. Thickness of body wall variable, longitudinal jusculature CoOmLMUOUS, Proboseis: Firmly attached, long, bitid and cupable of great extension, maximum length in fixed specimens 260 oum- Anns shore and usually of abour equal length, 20-30 mm. Lateral edges tend to roll inwards, veritral surface ciliate, dorsal surface smooth, Setoe Two, together with reserve setae, ventrally placed and posterior to mouth. Setal but no interbasal muscle present Alimentary Cunal: Mouth at base ef preboscis, canal much coded and fastened to body wall by numerous strands of muscle. Consists of (1) foregut (divided into pharynx, oesophagus, crop and gizzard), (2) midgut qwider and longer than foregut), associated for most of its length with a coll#leral litestine or siphon and (3) hindeut and gloaca, Siphon jubular, of smaller diameter than lutestine and arising near beginning of midgut. A ciliated groove hes in wall of much of posterior intestine. No precloacal caecum, Faecal matler forms pellets. Anus at posterior extremity of trunk, Nephridium: Single, attached to coelomic wall of (rink just posterior to level of setae, Size variable, depending on reproductive condition of animal. Wall of anterior and posterior regions usually thicker than thin, often transparent, middle region, which seems capable of much extension and even sacculation, Nephrostonie distal, situated about three-quarters to two-thirds length of nephridium away from sephridiopore. Nephrostomal lips frilled or cremated, situated at end of short stalk- Nephridium holds eggs and/or a male (two males in one specimen). Eggs develop in coelom along a mesentery assoviated with ventral blood vessel and nerve cord. Diameter of largest eggs 0.50-0,55 mm, Vascular System: Thin walled ventral blood yessel, Single neurointestinal vessel, arising posteriorly Tram two short arms on each side of anteriormos! region of intestinal siphon, joins ventral vessel in anterior half of trunk. Thin walled dorsal vessel] fuses with intestine near posterior exlremily of foregul. Anterior continuations of ventral and dorsal vessels extend into probasels. Neuroimestinal vessel often vesiculated or superficially roughened and coloured dark yellow to orange. Anal Vesicles; Two, much branched, tufted and each a complex of tubes and tubules, Johnston & Tiegs (1920, p. 75) described them thus: ‘Into the cloaca open fwo anal vesicles ... Into each open about 18 tubules, some quite short, others much longer. These tubes give aff smaller or larger numbers, al times very large numbers, of secondary nephridial [? excretory] rubes, cuch ending in a narrow heck Which bears a circular disc with the nephrostome opening in iis cenlre. The dise is composed of a ring of compressed elongated cells, with strongly staining nuclei and fringed with a ring of cilia’, In larger specimens the vesicles are larger and the branches more tufted. Descriplion of male (based on four stalned and mounted specimens) Located In wephridia of females but not permanently aitached like male of Pseudobanellia diuierina, Long, thin or flat but swollen or rounded anteriorly, lendifiy to taper posteriorly; largest about 20 mm long, maximum ‘width 1,2, lacking setae, Body ‘wall very thin, Outline of whal might be a rudimentary gut tuns through most of animal; body cavity contains developing sperm morulae. Cilia on some regions of body surface, No males were found by Johnston & Tiegs (1920). The male af Bonellia gigas Nielsen, 1963 is 19 mun long and 0,5-(,0mm wide, lacking setae bur 122 5, J. EDMONDS possessing a posterior sucker or clasper, The male of Bonellia tasmanica Dartnall, 1970 is 7 nim long, 0.8 mm wide, tapering posteriorly and lacking both setae and clasper. Systematics Bonellia gigas Nielsen, 1963, described from a very large bonelliid collected at Western Port, Vie., was considered to be different from & fuswelli chiefly because it lacked an intestinal siphon, Although I have not been able to dissect the type of B gigas, [ have dissected one specimen trom Port Phillip Bay and another from Flinders. Both have siphons. Further, Fig. 2 of Nielsen, 1963, clearly shows a siphon attached to part of the pur, especially the posterlor part. Nielsen apparently mistook the siphon for a vontinuation of the neurointestinal vessel and labelled it ‘intestinal blood vessel’ Dartnall (1970) described B tasmanica trom northern Tasmania, atguing that it differed from & haswelli because it lacked a siphon and from beth B& haswelliand & gigas because its nephridia were sacculated. It is probable that what Dartnall described as ‘an intestinal vessel, Which runs closely along the wall of the intestine for about the posterior two thirds of its length’ and then ‘leaves the gut and joins the ventral vessel’ is in part a siphon and in part a neurointestinal vessel. The sacculated condition of the nephridium of & tasmanica is'a doubtful character and was probably caused by temporary nvuseular contractions of the organ at the time of fixation, [n one specimen trom S.A, the thin walled part of the nephridium is constricted near its middle so-as to form twa sacs. From a study of male and female specimens of RB haswelli, B. gigas and & fasmanica it is concluded that the three species are synonymous, the first name having priority, Stephen & Edmonds (1972) transferred the species to a new genus Metetionellia on account of the distal position of its nephrostome. Johnston & Tiegs’ specimens from N.SW, were small but Nielsen’s from Victoria were very large. Specimens examined and localities New South Wales; Fairlight (near Manly) (1) AMS W56i2 and (3) AMS W4702 (two of these clearly show (the relationship between the neurointestinal vessel and the siphon); Camp Cove, Sydney Harbour (1) AMS W8703; locality unknown (1) SAM E1404. Victoria: Port Phillip Bay near Porr Arlington (2) MV coll. and at Mornington (1) MV coll; Flinders, Western Port Bay, SAM EL400 (1) and type male and female of B& gigas MV G2696 and 2697; Port Phillip Survey area 31 ‘inside buoy’ (1) SAM E1407. Tasmania; Jacobs Boat Harbour (north Tasmania), paratype of B /asmanica, TM K226, South Australia; Spencer Gulf — Boston f. (near Port Lincoln) (2) SAM EI401; at following islands in Banks. Group, Marum tl. (3) SAM E1475, (3) SAM E1492, Lusby 1, 0) SAM E1488, Langton 1, (1) SAM E1504, Hareby 1. (1) SAM E508; Se Vincent Gulf — Edithburg jetty (near base af outer piles) (2) SAM E1403, (1) SAM El4§7, (3) SAM 1458 (3) SAM Bi502; Marino Rocks (3) SAM E1406; Aldinga Reef (i) SAM E1402; Victor Harbour - near Rosetla Head (3) SAM B145], (3) SAM E1483, (3) SAM E1466, (5) SAM Et45], (3) SAM El4%) and near Whalers Wharf (3) SAM E1474; St Francis t. (Nuyrs Archipelago) (4) SAM 1469, Western Australia: Mistaken 1. (King George Sound) (1) WAM 37-85; Garden I, (1) WAM 10-73; off Carnac 1. (1) WAM 71-75; off Dansborough (LU WAM %73; Houtman Abrothos (Ac N. end of Morley |, Easter Group) (1) WAM 279-85, Distrifurian Known trem south-western, southern and south- eastern Austratia, from the Abrolhos J. in Western Australia to Sydney Harbour in New South Wales. Usually collected subtidally, occasionally intertidally. No records atfrer than from Australia. Habitat ‘In South Australia individuals of this species live in crevices between rocks and Under rocks in Sheltered, calmer water where there isa deposition of fine, muddy silrs, such as occurs on the lee side of the Blu/f at Victor Harbour and at Edithburg jetty on Yorke Peninsula. The gteatest concentration appears to he at the perimeter of the rubble-reef ‘area, especially where the latter abuts open flat where sea-prasses grow, The proboscis extends only at night and then over the bottom adjacent to the burrow for a radius of about one nietre' (NW. Holmes pers. comm). Genus Pseudobonellia Johnston & Tiege Pseudehonellia Johnston & Tiegs, 1919, pp. 213-230; Stephen & Edmonds, 1972, p. 401; Datta Gupta (976 p. 115, Type-species: Psevdobonellia buiterina Jobnsion & Tiegs Diugnasis Female with bifid proboscis Trunk with twa ventral setae, Two nephridia (gonoducts) with distally placed mephrostomes; nephrostomal lips crcnated. Anal vesicles branching, Male carried in small blind tube thal projects into coclam between nephridiopores, Type species: Pseudobonellia AUSTRALIAN ECHIURANS FIGURES 2-4, Metahonellia haswelli, 2, anterior vegion dissected. 3, seta (scale line = 0.3 mm), 4, portion of anal vesiele, hiuterina Johnston & Tiegs, 1919, The genus contains only one species. Pseudobonellia biuterina Johnston & Tiegs (Figs 5-8, 19) Pseudobonellia hiuterina Johnston & Tiegs, 1919, pp, 213-230, pls 9-11; Monro 193], p, 33; Fisher 1948a, p. 856; Edmonds 1960, pp. 96-97, fig. 5; Stephen & Edmonds 1972, p, 401. Archibonellia mjoebergt Fischer, 1921, pp, 6-8; Austrobonellia mjoebergi Fisher, 1948a, 856; Edmonds 1960, p. 97. Holotype: AMS G477; type locality, North West [slet (Capricorn Group), Qu. Description of female Trunk: Small, maximum length 23 mm (mostly 6-14), maximum Width 3-6, pyriform, sub-ovoidal to globular, pale io dark green. Body wall thin (sometimes transparent) except in anterior and posterior region, usually thinnest on dorsal side. Surface usually wrinkled by large numbers of closely set papillae. Nephridiopores and opening of male tube on anterio-ventral surface often very noticeable, Proboscis: Long, bifid, adherent and capable of much extension; in fixed specimens 1-10 times as long as trunk. Arms shorter (3-10 mim), normally about equal length. Ventral surface ciliated, lateral edges tending to curve inwards. Mouth at base of proboscis, Setae: Two (in addition smaller reserve setae usually present), golden brown, slightly iridescent, with free end forming a weak hook. Johnston and Tiegs state that larger hooks are 2-3 mm long and smaller 0,7-0.8 mm and that a strong muscle pad joins their internal ends ‘evidently serving to impart fo them a lateral pincer-like movement’, The description aptly fits the structure and function of an interbasal muscle. Although there are well developed setal muscles in the present specimens, none clearly show the presence of an interbasal muscle, Nephridia (gonoducts or uteri): One pair, size variable, usually prominent, tapering distally, each with a slightly frilled, distally placed nephrostome borne on a short stalk. Largest eggs in nephridium 0.25-0.35 mm (Johnston & Tiegs 1919 give 0.[1 mm). Nephridiopores on each side of nerve cord in anterior trunk region, Male tube (androecium): Small, 1,5-3.0 mm long, opening externally near nerve cord at about level of nephridiopores. Opening often very noticeable. Tube encloses small, wormlike male attached to distal end of tabe. Alimentary Canal: Foregut short; midgut long, thin walled and associated for much of its length with an intestinal siphon; hindgut short. No precloacal caecum. Faeces form sub-ellipsoidal pellets. 124 5 1. EDMONDS Vascular System; Thin walled ventral vessel runs alongside of nerve cord, Thin walled dorsal bioad vessel (fastened to foregut and body wall by mesenteries) fuses with fut near junction of fore- and mid-gut, Single nevrointestinal vessel, arising from wall of midgut just posterior to anteriamast extremity of siphon, joins ventral vessel in anterior half of trunk. lotestinal extremity of neurointesunal vessel usally arises from two short roots which lie on each side of the siphon and in close contact with it, Neurolntestinal yessel sometimes vesiculated, Anal Fesicles; Fwo branching, tuft-like masses of fine tubes, arising on each side of posieriormost section of hindgut, In this respect they differ from the type description ia which ‘each anal tree consists af masses af very delicate, simple, cylindrical tubes Opening separately into the rectum’. Edmonds (1960, p, 96) remarked that ‘the anal vesicles do not seem to communicate With the cloaca as simply as described by Faohnston and Tiegs’and (hat the tubes branch to some extent. The anal vesicles of dissected specimens in the present collection fram Queensland and Western Australia are branched. In one of the paratypes (AMS 6477), the vesicles arise fram about! 12 shar! tubes which branch and sometimes rebranch into simple cylindrieal rubules that open to the coelom through slightly dilated funnels fringed with cilia, Ovaries: Jotinston & Tiegs (1919) slate thal the ovaries arise from the peritoneum lining the muscular strands (hat hold the posterior portion of the rectum in position and that they |le transversally on the frenulae. In the present specimens developing egys lie more longitudinally in the posterior third or half of the body cavity in close association with the ventral vessel, Description af male Accarding ta Johnston & Tiegs (1919) the male lives permanently in the androecium of Ihe female, the two being fused distally. Without definite mouth or anus, although a rudimentary canal ls presen, No setae. Two seminal vesicles present. How the male performs its sexual function is not known. Johnston & Ticgs (1919) suggest that ‘the sperms may be liberated into the cavity of the androecium whence they reach the exterior through its canal and enter the adjacent openings. ft 18 possible, however, that the male may be protruded through the canal of the androecium and actually liberate eperms in the female apercure’. Systemulics Although the specimens examined 17 the present collection from North West |. and Heron 1, differ in a few respects from the lype description, they are considered to be ? biuierina. No satisfactory character has been found tat distinguishes any of the Western Australian bonelhids from Aiaterine. Those collected from the CSIRO laboratory a} Waterman Bay (from under pots standing on sand in an indoor aquarkum through which sea Water was conlmually passed) Were pale green while Lhose collected at Heron 1 were dark green. How important colour di flerences are is noc known, Agius & Jaccarini (1981) have shown that the unpigmented trochophores of Bonellia viridis when kept under constant Wlumination develop into unpigmenied adults, Whether the depth of colour of diverting depends on the amount of light received is Unknown. Twe other bonelllids closely related to P biuterina haye been described from W.A., Archiborellia michaelsen Fischer, 1919 and Austrehanellia myoebergi (Fischer, 1921), the former from Rottnest 1, the latter fram Broome, both localities being places where # bivéerina has been found. Bath Fischer's species were described from single speciinens. Anstrobonellia mineberg) Fischer, (921, is now being placed ip the synonymy of FB biuterina Johnsten & Tiegs, (919 Fischer described his specimen thus; trunk oval, 45 mm lung, light prey and |ransparent; proboscis short, 18 mim long, with two arms of unequal length, Setae two with recurved tip, no interbasal muscle. Nephricia tava, thin walled (containing eggs) with distally plaved néphrostames and a median, unpsired, smailler, thick-walled ‘Segmentalorgan’ or ‘Uterns’, with a basully placed nephrostome, Intestine lone and convoluted, Anal vesicle dise-like and expanded With J2e15 deniritic main stems the branches af which possess lateral funnels. Lf Fischor's Segmentalorgan’ is a male tube and if injury accounts for the inequality in the length of the probosels arms, then the description of A_ thioebergi fits very well that of P divrerine. Fischer's other specimen from W.A., Austrabonellia michaelseni, differs iy other characters and is Possibly a separate species, Specimens exvarined and locglities (these are additional to those recorded in Edmonds, 1960) Queensland; Heron 1, (1) AMS W3719; North West F (3) AMS WISO7, (l) AMS W691; Whilsunday Group (2) AMS W3029: One Tree 1. (1) AMS W9275 Westen Australia! Barrow f, (2) WAM 139-8); Riddell Pt, Broome (3) WAM 50-85; Roebuck Bay, Broome (2) WAM 47-835; Abrainos I, (1) WAM 102-79; Yanchep Reef (2) WAM 98-79; Waterman Bay (CSIRO Manne Laboratory) 110) SAM E1439-1441; Garden I, (1) AMS W3720; Albany (S. side of Princess Royal Harbour) (2) WAM 147-81; Lookour Pt, Cheyne Bay (1) SAM E1482. Distribuwlion Queensland: Phe Crear Rariice Reet, from AUSTRALIAN ECHIURANS 125 Capricorn Group in the south to Low I. in the north. Western Australia: From Barrow LI. in the north to Cheyne Bay in the south. Whether the species extends from Queensland to Western Australia through Torres St is not known. It is not known from S.A., Vic., N.S.W. or Tas. Other record: New Caledonia (Stephen 1976). Habitat Specimens have been collected intertidally in coral and limestone reefs; also from under stones and objects resting on sand. Genus Archibonellia Fischer Archibonellia Fischer, 1919, p. 283; Fisher 1984a, p. 856. Type-species: Archibonellia michaelseni Fischer, 1919. Diagnosis Female with bifid proboscis. Trunk with two ventral setae. Three nephridia, two being paired, lateral and very small and the third median, large and unpaired. Position of nephrostome not known. Intestine very short with globular caecum. Male not described. i FIGURES 5-8, Pseudobonellia biuterina, 5, anterior region, digestive and vascular systems; 6, nephridia (gonoducts) and male tube; 7, small portion of anal vesicle; 8, setae from different specimens (scale line = 0.5 mm). 126 8. J EDMONDS ?* Archibonellia michaelseni Fischer Archibonellia michaelseni Fischer, 1919, pp, 283-285; 1926, p, 207; Fisher 1948a, p, 856; Edmonds L960, p. 97. Type-specimen; Not Known, type locality: west coast of Rottnest J,, near Fremantle, W,A, Description of female (after Fischer 1919) (Male not kriown) Trunk: About 12 mm long, grey in life, Proboscis anteriorly forms two lappets. Setae two, with reserve pair and an interbasal muscle. Two small, paired nephridia (‘Segmentalorgane’) placed below a large unpaired ‘Uterus’, Intestine very short, consisting of a single loop in anterior half of trunk and a bew- shaped tubular part in posterior half. Globular caecum present, Ovary lies along posterior region of nerve cord, Anal vesicles arise as single stems an either side of rectum and branch at tip. Position of nephrostome not known. Systematics A, michaelseni resembles FP bititerina Johnston & Tiegs in many respects, especially in the possession of an unpaired, median ‘Uterus’ lying between paired lateral nephridia, Moreover, Fischer (1919) thought that he saw a miale in the median structure, In PR bivterina, however, (1) the lateral nephiridia are much larger than the medial tube; (2) the alimentary canal is very long and convoluted and not very short (incredibly short) as shown in Fischer 1919, fig. 6; and (3) neither caecum nor interbasal muscle is present, While the two species may be synonymous the described differences between them are considerable and until the type of more specimens became available it ig probably best to consider them as dilferent. Genus Protobonellia [keda Protobonellia Ikeda, 1908, p, 259; Fisher 1948a, p, 854; Datla Gupta 1976, p.1L5. Type-species; Profohonellia mitsukurli Ikeda, 1908, Didgnosis Proboscis of female long, tubular, non-bifid, Ventral setae two, One nephridium. Nephrostome stalked, fimbriated, basal, Anal vesicles tong, dentrine. Male not known. * Protobonellia papillosa Murina Protobonellia papillase Murina 1978, pp. 112-113, fig, 4. Description (atter Murina, 1978) Trunk 28 mm long, 16 mm wide, bearing rounded papillae 0,25-1,25 mm in diameter, densest anteriorly and posteriorly, Proboscis light grey, width 5.5 mm, distally blunted; basal part (near mouth) has form of oval collar with thick, wavy, pigmented borders and two long processes laterally. Setae two, golden-yellow, bent or curved, 0.5 mm long, 0,15 wide, Nephridium single, rounded, lying oo right side of nerve cord; nephrostome short, basal. Anal vesicles form dense bushes on each side of rectum. No clearly visible anal rosette. Specimen and locality Described from one female specimen collected during cruise of ‘Dmitrii Mendeleef', Stn 1245, 30° 24'S, 161° 57'E near Lord Howe 1, al 1200.m. No other record, Genus Sluiterina Monro Sluiterina Monro, 1927, p. 618; Murina 1976, p. 840. Type-species: Mamingia sibogae S\uiter, 1902, Diagnosis Proboscis of female non-bifid; lateral edges turin inwards piving structure a tubular appearance; edges fuse near mouth to form a cup, Nephridium single, nephrostome basal. Anal vesicles bushy or brush like. Male unknown, * Sluitering alba Murina Sluiterina alha Murina, 1978, p. 11, fig. 3. Description (after Murina 1978) Trunk 44 mm long, 8 mm wide, posterior region damaged, Body wall white, thick and not transparent. Proboscis 12 mm long, 6 mm. wide (distal part damaged), with lateral margins folded inwards making i tubular in form. Nephridium single, suc-like, 7 mm long, 3 mm wide, with centrally located nephrostome, About 50 white eggs with diameter 0.12-0,13 mom in cavity of body. Between posterlor coils of gut are bunches of bight yellow material, probably remains of anal vesicles, Specimen and lacality Described from a single female collected during eruise of ‘Dmitrit Mendeleef', Stn. 1373, Great Australian Bight, 33° 48S, [27°07°E at 1080- 1100 m. No other record, Genus Vitjazema Zenkevitch Viljazema Zenkevitch, 1958, p, 193; Datta Gupta 1976, p. 115. AUSTRALIAN ECHIURANS 12) ‘Type-species: Vitjuzema ul/raabyssalis Zenkevitch, 1958. Diagnosis Probosics of female non-bifid; anterior region, however, expanded into a shghtly widened ‘head’ with thickened festoons along anterior border; under festoons are triangular flaps directed inside a ventral gutter. Setae two. Nephridia two, nephrostome distal, Anal vesicles sac-like. No male known, * Vitjazema ultraabyssalis Zenkevitch Vitjazema ultraabyssalis Zenkevitch, 1958, pp. 195-197, fig, 3; Murina 1978, p. 115. Description Trunk green, length I4-15 mm. Proboscis 9-27mm Jong with deep funnel on ventral side; anterior region widened with festoon-like border consisting of 5-6 triangular lappets, Setae 2, large with bent blades and blunt tips. Nephridia one pair, wilh distal nephrostomes at end of long tube. Anal vesicles unbranched, covered with small funnels, Specimens and localities ‘Two female specimens collected during cruise of ‘Dmitrii Mendeleef’, Stn 1365, Great Australian Bight, 34° 25°S, 128° 12° 5B at 3880 m. Distribution Kurile - Kamchatka Trench (at 5560-9700 m); Marianne Trench; Great Australian Bight (at 3880 m). Genus Zenkevitchiola Murine Zenkeviichiola Murina, 1978, p, 108, ‘Type-species: Zenkevitchiola brevirostris Murina, 1978, Diagnosis Proboscis long, non-bifid. Trunk without setae. Single pephridium, nephrostome basal. Anal vesicles, two, long, slender, filamentous. Male not known, * Zenkevilchiola brevirostris Murina Zenkevitehiola hrevirostris Munna, 108-109, fig. 1. 1978, pp. Description (alter Murina, 1978) Trunk 68 mm long, 28 mm wide, anterior third and posterior quarter covered with low, rounded pupillae 1.5 x 0.8 mm. Coils of gut visible through body wall, Proboscis white, transparent with lateral margins turned up or folded, length 65 mm, width 5-7 mm, distal extremity curved and widened, proximal extremity forms cup-with a slit on ventral side. Nephridium single, }imm Jong, 5mm wide, located on right side of nerve cord; anterior region swollen and filled with eggs 0.25-0.3 mm in diameter, posterior half with thicker walls and na eggs. Nephrostome basal, stalked and with rosette at distal end, Anal vesicles two, dark brown, tapering distally. Gut coils 10-12. Anus forms weak rosette, surrounded with small papillae. Specimen and lacality Described from one female collected during voyage of ‘Dmitrii Mendeleef’, Stn 1345, near southern Tasmania, 43° 47'5"S, 147° 51°E at 755 m. No other record. Genus Anelassorhynehus Annandale Anelassorhynchus Annandale, 1922, p. 148; Fisher 1946, pp. 221-22; 1949, pp. 480-481; Stephen & Edmonds 1972, pp. 443-444, Type-species; Thalassema branchiorhynchus Annandale & Kemp, 1922. Diagnosis Proboscis well developed, usually long, never bifid. One pair of ventral setae, Longitudinal, circular and oblique musculature of body wall continuous. Nephridia, |-3 pairs, Nephrostomal lips long and spirally coiled (thus differing from genus Thalassema}, KEY TO AUSTRALIAN SPECLES OF ANELASSORH YNCHUS |. Nephridia, (wo pairs and post-setal........... 2 Nephridia, three pairs and post-setal.., A, vegrandis 2. Trunk globular to ovoidal, sandy-grey to light brown in colour ..... A. poreellus porcellus Trunk sausage-shaped to clongate, green in colour... 20. veeey Ay porcellus adelaidensis Anelassorhynchas vegrandis (Lampert) Thaslassema yegronde Lampert, 1883, p. 341; Monro 1932, p. 33. Anelassorhyachus vegrandis Visher, 1946, p, 222: 1949, p. 481. Dipe-localin Philippines, 128 5. J, EDMONDS Description (atler Lampert 1883) Proboscis lacking. Nephridia three pairs and post-selal. Nephrostomal lips spirally coiled, Anal vesicles long and without ciliated funnels. Remarks Monro’s specimen Irom the Barrier Reef was in poor condition and his identification wag made with some reservation, The species is not well known Anelassorhynchus porcellus porcellus Fishvei (Fig, 20) Anelassorhynchus porcellus Fisher, 1948b, pp. 274-277, figs la-d; Edmonds 1960, pp. 91-92, pl, le. Type-specinren: US. Nat. Mus., Washington D.C.. Type-locality, Honolulu, on reef south of hartrour. Description Yrunk: Globular to ovoidal, colour sandy grey to light brown, length 25-40 mm, maximum width 15-29 mim} skin rather thick but wrinkled with tumerous flat papillae; musculature continuous, Selae two, golden-brown, lying posterior to mouth; no interbasal muscle present, Proboscis; Fleshy, readily deciduate, usually tapering antetiorly, 8-20 mm long. Alimentary Canat: Very long and fragile, tilled wilh fragments of coral, small shells and coral sand (which usually rupture the thin gut wall as soon as one tries to free the intestinal coils); presiphonal secon of mid-gut very long. Vascular System: Consists of dorsal vessel, ring vessel, 1WO neuiro-intestinal vesgels and a ventral vessel. Serae: Two pairs, post-setal, with nephrostomal lips long, slender and coiled. Anal Vesicles: Two long, with small, utstalked funnels, Intestinal siphon present bul no precloacal caccum. Sysfematics The specimens from Heron I, correspond closely with two of Fisher’s specimens of A. porcellus from Kakaoha Reef, Hawaii (U.S. Nat, Mus, part 26423). Fisher was unable to recognise any ciliate funnels on the anal vesicles of his specimens. In the Australian specimens the funnels, though small and sparse, are definitely present. One of Fisher's Speciinens when dissected was found ro possess three pairs of nephridia, Specimens Exarrined Qu; Heron I, (Capricorn Group) (3) SAM E1425; North-West 1, (Capricorn Group ) (1) AMS W214; Ingram J, (1) SAM ElL43]; Brockhurst Reef off Townsville (1) SAM ELdY4, Distribution Western Pacitie Ocean at Hawaii and Great Barrier Reef, Qu, Anelassorhynchus porcellus adéelaidensis Edmonds fi. stat, (Figs 9-U, 21) Anelussarhynchys udelaidensis Edmonds, 1960, pp. 92-93, pl, 2a. Anelassortynchus porcellus (in part) Edmonds, 1982, p. 316. Type-specimen; AMS; type locality Aldinga Beach, S.A, Desvription A number of specimens Which have previously been called 4. adelaidensis and A, porcellus (in part) are now being referred to as a new subspecies, A. porcellus adelaidensis. The new subspecies differs from the nominate form in size, colour and distribution, Thank: More elongate than nominate subspecies, length 15-90 mm, maximum width 10-40 mm, always lighi to dark green, surface wrinkled and made verrucose by many, large, flat, white papillae (most Numerous anteriorly and posteriorly); secretes copious amounts of mucus making animal very slippery to held, Probescis, Fleshy, readily deciduate, up to 37 mm long, lateral edges. may be wavy but never with processes, See: Two golden brown, 2.8-5 mm long, no interbasal muscle (setae lost in some specimens). Nephridia: Two post-setal pairs (occaslonally an extra single nephridiufm or pair); nephrostomal lips filamentous, weakly lo sirongly coiled, Alimentary Canal; Very long and much convoluted, with very long presiphonal section of mid-gut; faecal matter not in form of pellets; intestinal siphon present but no caecum. Anal besicles: Two, long, thin walled, tubular bul usually expanded basally, and attached (especially basally) to body wall by numerous mesenteries or muscles; coelomic surface bears sparsely distributed ciliate funnels and numerous brown spots which appear to be aggregates of very small pigmented granules, Vascular system: Dorsal, ring, two weurointestinal- and ventral vessels, Hand cut sections of proboseis show two lateral and one median vessel, Well-developed ventral nerve cord extending Into proboscis. Specimens and lacalities Vicioria; Port Phillip Bay, S4M E429, Brighton, SAM B1432.. AUSTRALIAN ECHIURANS FIGURES 9-11. Anelassorhynchus porcellus adelaidensis. 9, dissected specimen; 10, setae (scale line = 1.0 mm); II, ciliated cup from anal vesicle. 130 §. |. EBMONDS South Australia; St Vincent Guil-Porl Willunga, SAM E1427 (1); Aldinga Reef, SAM E1428 (1); Cape Jervis, SAM E1430 (1); Coobowie, SAM E1454 (1), SAM E1455 (1), SAM E1456 (5), SAM El46l (3), SAM E1462 (1); Brighton, SAM E1463 (5). Spencer Gulf-at following islands al Banks Group, Winceby 1_, SAM E1476 (3), Marum £., SAM 61477 (2), SAM E1486 (1), SAM E1493 (3), Lushy lL, SAM E1489 (3). Eyre Peninsula-Port Lincoln, SAM E1479 (2); Port Turton Jetty, SAM B1S03 (1); Venus Bay (under jetty), Blanche Pt (Streaky Bay), Smokey Bay, (coll, K. Gowlett-Holmes).. Systematics The internal anatomy of the South Australian specimens corresponds very closely with that of 4. porcellus, Their shape, however, is more elongate and their colour green. Fisher does not mention the colour of his specimens bul specimens of A, porcellus collected by the author at Heron I., Qu,, were light brown to sandy grey, The green subspecies is commonly collected by divers in S.A. but no brown forms have yet been found in the State. The distribution of the two subspecies consequently seems different. A. gangae Bisewar, 1984, recently described from Natal, South Africa and A, porcellus adelaidensis are closely related species. Distribution Southern Australia from Port Phillip Bay ( Vie.) to Streaky Bay (Eyre Peninsula) in S.A, Genus Arhynchite Sato Arhynchite Sato, 1937, pp. 142-143; Fisher 1946, p. 485; Stepheri & Edmonds (972, p. 414, Type-species: Vhalassema arhynchile \keda. Diagnosis Proboseis long slender, often ribbon-ike, sometimes deciduate; anterior extremity expanded into @ fan-like structure, Two ventral setae with strong interbasal muscles. Nephridia two, with nephrostomal lips expanded into a leaf-like structure. Anal vesicles long, thin walled and unbranching, Vascular system with or without ring vessel Arhynchite hiscocki Edmonds Arhynchite hiscocki Edinonds, 1960, pp. 9U-91, Ng. 3, pl. 1, fig. 1b; 1966, p. 178; Stephen & Eximonds 1972, p. 417, Holotype: AMS W3714; type locality: Dunwich, Qu., ‘dug from sand pit, 18" below surface’, Description Trunk: Elongate, slender, pencil-like, length 100-120 mm, width 4-6 mm, fixed specimens yellow-brown to zrey green. Surface made verrucose by numerous near rows of elevated papillae, slightly larger at anterjor and posterior extremities. Masculature continuaus, Proboscis: Delicate, slender, about 30 mm long, {.5-2.5 mm wide, still attached to trunk in holotype; anterior extremity flattened and fan-like. In one specimen from Victoria proboscis is shorter, deciduale, with antenor extremity more spoon-like, Selae: Two, connected to bady wall internally by strong radiating muscles and to each other by strong interbasal muscle. Nephridia: Two, sub-cylindrical, slender, length variable; postsetal, Nephrostome basal was expanded, frilled or leaf-like lip. Alimentary Canal: Midgut with siphon; no precloacal caecum. kascular System: Dorsal blood vessel fuses with foregul at posterior extremrty of latier; neurointestinal vessel conneets with anterior section of midgut near anterior extremity of siphon, No Ting vessel observed. Anal Vesicles: Two, very slender, brown, about one third to a quarter as long as trunk, fastened (hroughout their length to posterior region of alimentary canal but to body wall over last quarter of their length, Numerous ciliated furmels scattered over their surface. Systematics Sato (1937) erecled the genus Arhynchile for a group of echiurans Jacking a proboscis, Fisher (1949), having found cwo species possessing a long deciduate proboscis, redescribed the genus. The proboscis of the helotype of 4. hiscocki is still attached but that of one of the Victorian specimens is detached. The genus contains six species collected from places bordering the Pacitic Ocean; A, californicus -Monterey {U.S.A,), A. inamoenus - Monterey (U.S.A), A, pugettensis Puget Sound (U.S.A.), -4- rugosus -Shantung (China), 4, arhivachite —Japan. Some of the spevies are closely related and difficult to distinguish, 4, Aiscacki is not a well known species und needs re-examination and revision when more species are found. Specimens and localities Queensland: Stradbroke |, (Edmonds 1960), AMS W714 (2) Victoria; Port Phillip Bay (Editiends 1966) VM coll, (2) AUSTRALIAN ECHILIRANS it South Australia; Spencer Gulf, north of Port Lowly, SAM E1524 (1B). Genus Listriolobus Spengel Listriolobus Spengel, $912, p. 316; Fischer 1926, p. L10; Fisher 1946, p, 233. Type-species: Listriolobus bahamensis (Fischer), (designated by Fisher 1944), Diagnosis Proboscis of variable length, truncate but never bifid, Two setae with interbasal muscle, Longitudinal musculature of trunk wall grouped into bands (not always well developed in young specimens), Oblique musculature not banded or fasciculated as in Ochetosiama. Nephridia two to three pairs, nephrostomal lips long and spirally coiled. Anal vesicles sac-like to tubular and without branches. Kry 10 AUSTRALIAN SPECIES OF LISTRIOLORBUS |. Longitudinal musculature in 7 bands L.. brevirostris Longitudinal musculature in 13 bands 1. sorbrllans ?”* Listriolobus sorhillans (Lampert) Thalassema serhillans Lampert, 1883, pp. 340-341; Augener 1903, p. 349. Listriolebus sorbillans Fisher, 1946, p. 234. TWpe-locality: Philippines. Australian record: Sydney (call. Dr Schutte, 1876) in Augener 1903, p, 349. Description (after Lampert 1883) According to the type descriprion (based on a single specimen) the trunk is 65 mm long and proboscis 24. Longitudinal musculature in 13 bands, Nephridia three pairs with spirally coiled nephrostomal |ips, first pair presetal in position, Setae small, Anal vesicles long, brown, bearing tmicroscoupic ciate funnels, Small rectal caecum. Augener’s description of his single specimen is brief. Trunk about 42 mm long, proboscis 18 mm, Whole body covered wirh papillae which are smallest in mid-trunk and largest posteriorly, Anal vesicles about two-thirds length of trunk, Remarks This Australian record needs conlirmation. If the oblique musculature of Augener’s specimen was fasciculated then it might have been Ocherostoma australiense, Listriolobus breviewstris Chen & Yeh Chen-Chang (Figs 12-13, 22) Listriolobus brevirostris Chen & Yeh Chen-Chang, 1958, pp. 273-278, fig. 7 A-D; Stephen and Edmonds 1972, p, 424, Listriolobus bulbocaudatus Edmonds, 1963, pp- 243-244, pl. 1, Fig. 1. Type-locelion, Kiao-chow Bay, Shantung, China. Description Trunk: Sub-cylindrical to cigar shaped, length 2-85 mm, maximum width 10-20 mim, fixed specimens light to dark pink, Surface cavered with while papillae, lying almost in rows. Posterior extremity way sometimes be modified and expanded Into a fleshy, bulbous, conical structure: bearing three or four rows of prominent, pointed or mamillate, white or pink papillae. Longitudinal rnusculature arranged in bundles, often difficult to discern externally. Dissected specimens show seven (eighi in one specimen) well spaced longitudinal bands, occasionally weakly developed where the body wall is thin. Oblique musculature between bands of longitudinal musculature continuous and not in fascicles. Proboscis; Non-deciduate in all specimens, In fixed condition short, stout, 10-16 mm long, 6-12 mm wide. Small papillae on dorsal surface. Lateral margins wrinkled, folded, indented or crenated. Ne lateral processes as in Anmelassorhynchus branchiorhynchus Ammandale & Kemp. Selae: Two (with smaller reserve setae), length (measured in straight line from base co tip) up to 7.2mm, strongly hooked and sickle-shaped. Strong interbasal muscle. Nephridia: Two post-setal pairs, in one specimen three nephridia on one side and two on other. Length variable, some extending almost to posterior extremity of trunk. Nephrostamal lips long and much coiled, Alimentary Canali Long, much coiled, Presiphonal section of midgut long. Precloacal caecum present. vascular System: Dorsal vessel expands into a thin walled saccular vessel or heart. Well-developed ring vessel at Junction of fore- and midgut gives off two neurointestinal vessels which join before they reach the interbasal muscle and then bifurcate to form a loop around the muscle Ventral vessel pressed close to nerve cord and terminates in the cloacal caecum. Anal Vesicles; Long, thin walled, brawn, swollen basally in most specimens, bearing numerous, small, ciliated cups some on very short stalks. Anteriorly placed cups more sparsely distributed. 132 5. J. FDMONDS Svstematics These Australian specimens closely resemble Ochetostorna septemvotum Datta Gupta, Menon & Johnson, 1963 from Quillon, India, In none of the Australian specimens, however, has the oblique musculature of the body wall been found to form fascicles, like that shown for Ochetostoma acionyotum by Fisher 1946, pl. 23, fig. 2, and pl. 24 (in the spaces between muscles labelled MYL, ML and MDL) or like that shown in the transverse sections of Ochelostoma bombayense by Mathew 1976, fig, 5, Because fasciculation of the oblique musculature is a character of Ochetostoma, the Australian spetimens are considered different from O. septemyvoruin, Edmonds (1963) originally described the specimens from Queensland as Listriolobus bulbacaueatus. At the time he was unaware of 1, brevirostris Chen & Yeh Chen-Chang (1958) from Kiao-chow Bay, Shantung, China. At a laler date Stephen and Edimorids (1972, p. 424) considered the two species were distinguished by three or four rows of prominent papillae and a bulbous structure both present at the posterior region of L. bulbocaudatus. More recently the author has examined three specimens from Queensland, in which the rings of prominent papillae and the bulbous structure are much reduced, This information brings the specimens. within the range of Z, brevirastris. Consequently L. bulbocaudatus \s now considered to be a junior synonym of LZ, breviresiris Chen & Yeh Chen-Chang, (1958), Speciniens examined and localities Queensland; Yeppoon, SAM E434 (1); Mud t., Moreton Bay, SAM E1433 (1); Round Is, Hervey Bay, SAM E1436 (1); Bramble Bay, SAM EL460 (1); ? Bramble Bay, SAM E1435 (4). Dredged from mud. Distributian China at Shantung, Australia: Queensland. Genus Ochetostoma Leuckart & Rueppell Ochetosioma Leuckart & Rueppell, 1828, pp. 7-8; Fisher 1946, p. 240; Stephen & Edmonds 1972, p, 426, Type-species: Ochetostoma erythragrammon Leuckart & Rueppell, 1828. Diagnosis Proboscis long, Capable of much extension, non- bifid. Trunk medium to large with longitudinal musculature lying in well-defined bands. Intervals between bands crossed by numerous fascicles ar small bundles of inner oblique musculature (Fisher 12 7 13 FIGURES 12-13, Lisirialobus brevirnstris, 12, anterior region dissected; 13, seta 1946, pl, 23, fig, 2), Nephridia in one to seven pairs, with long spirally coiled nephrostomal lips. Setae two, with or without interbasal museles. Anal vesicles long, more or less tubular, unbranched Rectal caecum usually present. KEY TO AUSTRALIAN SPECIES OF OCHETOSTOMA 1, Longitudinal musculature in /1-\3 (\1-14) bands; three pairs of nephridia, One pair presetal, (wo pairs postsetal. Fasciculation of oblique musculature usually well developed. No intérbasal muscle ii todelgh hers eege tevy tera Ch yustentivose Longitudinal musculature in 17-19 (17-21) bands; two pairs of postsetal nephridia. Fusciculation of oblique muscualture well developed, Strone interbasal muscle.) 1.00.60. cove OQ baroné Ochetostoma australiense Edmonds (Figs 14-15, 23) Ochetostoma australiense Edmonds, 1960, pp. 93-94, tig. 4, pl. 2b; Datta Gupta & Menon 197], pp. 177-178, figs. 2c, 2e. Tipe-specimen: AMS; type loeality, mud flats at Dunwich (Stradbroke [.), Qu. Description Trunk: Usually large, sausage-like, cigar shaped or elongate, pale to dark red, length 40-130 mim, maximum width 15-30. Thickness of body wall variable, sometimes Very thin, Surface, especially in anterior and posterior fegions, covered by numerous small, fal, fleshy to wartlike papillae. Usually 12-13 (11-14) well developed longitudinal muscles best counted in dissected specimens. Jn 2¢) dissected specimens the maximum number of bands was 14 in 3, 13 in 10, 12 in 6 and J) in one, Two AUSTRALIAN ECHIVRANS 35 bands, one on each side of the nerve cord, lie very close together and may appear to be one. Oblique musculature between longitudinal bands usually grouped into numerous fascicles, which may be weakly developed or even absent in parts of some animals. Proboseis: Jn living animals is highly extensible (150-200 mm), pale and ribbon-like. Ln. fixed specimens shorter, fleshy, up to 60 mm long, with lateral margins rolled inwards on yeritral side. Usually adherent or non-deciduate, Anterior extremity may be flattened somewhat and lateral Margins may be slightly wrinkled. No lateral PIPOCeASES, Serae: Two, 2.5-3,1 mm long (measured in a Straight line from tip to midpoint of base), golden, encased in sheath connected to body wall by a nuniber of sera] muscles. No interbasal muscle. Nephridia: Three pairs, the first presetal, others postsetal, Length variable, sometimes over half length of trunk. Nephrostomal lips elongate and spirally coiled, although Sometimes only weakly, Jn one specimen only five nephridia present. Digestive System: Mouth at base of proboscis. Foregut short, midgut very long and much eoiled, Presiphonal section of midgut long and traversed for part of its length by cilrale groave, Siphonal section of gul also long, Well developed caecum present. Gut contents largely mud and sand; oo faecal pellets. Vasenlar System: Consists of dorsal blood vessel (sometimes well expanded), ring vessel or sinus (may also be expanded), two long neurointestinal vessels and a ventral blood vessel. Two neurountestinal vessels fuse to form one short vessel which joins the ventral vessel at about the level of the setae, Posterlorly ventral vessel gives off a branch to caecum, Anal vesicles: Two, long, slender, thin walled, light to dark brown, opening into cloacal region of intestine. Posses small, unstalked ciliate funnels. Systematics Edmonds (1960) considered! these specimens with 11-14 muscle bands to be different from O. ervrhrograminod Leuckart & Rueppell which possess 14-18 muscles. @ erytArogrammon and O, auslraliense. however, are closely related, The neurointestinal vessel of all the Australian specimens exaniined in the presen study is double for most of its length, Datta Gupta & Menon (1971) state that the corresponding vessel in their specimens Of & ervthragrammon is single, [f this difference always exists, ir further distinguishes the lwo closely allied spevies, Sato (1999, fig. 9), however, shows two ceuroiptestinal vessels for his specimens af C2. ecvifrogriniunert. Stephen & Edmonds (1972) list nine species of Ocherostoriu that possess three pairs of nephridia and in which the number of longitudinal muscles varies from 12 to 22. Wesenberg-Lund (1939) and Sato (1959) considered that they were conspecilic. If they are correct then O. australiense would become ©. ervthrogrammon. Specimens examined and localities Queensland: Dunwich (in mud flats-in front of cemetary) SAM E1410 (4) and B14] (1); Caloundra (dug at low tides from mud flat; opposite a small mangrove island) SAM El415 (7); Myora (in mud flats at law ude) SAM ES413 (20). New South Wales; Goodwood [. (near mouth of Clarence River) AMS W3186, W3375, W387; Brunswick Heads (1) AMS coll, Habitat At Goodwood 1., the worms are found ‘between high and low water marks in rather dark sand Situated close to some small mangrove clumps. The proboscis is white and fleshy and protrudes from a hole in the sand. [t lies along the surface of the sand and is about 6” long and 1/2” wide. In this condition it appears to be quile flat (like a ribbon) and does not appear to take ona tube-like shape as in preserved specimens. The body of the worm is Solt and bright red in colour. Twelve longitudinal muscles show up clearly’ (P, Durie pers, comm.) Distribution Eastern Australia from Caloundra (Qu.) to Goodwoad Et. (N,S.W.), Andaman 1. (Datta Gupta & Menon, 1971). An inhabitant of intertidal mud f}ats. Ochelostoma barani (Greeff) (Fig. 24) Thalassema barenit Greeff, 1879, pp. (41-192, pl. 6, figs 62-67- Ochetostoma baronij Mackie, 1951, p. 247; Stephen & Edmonds 1972, p, 429; Amor (974, p, 123-124. Ochetostoma nyersae Edmonds, 1963. pr. 245-246, pl. 1, fig. 2. Previous Ausiralian record: N.SANV, (Edmonds 1963), Description Trunk: Small to moderately large, sac~ sausage- or cigar-shaped, length 21-70 mm, maximum width 9-25 mm; anterior region rounded, posterior sometimes almost pointed. Green. Surface covered with soft, almost white, shghtly clevated, wari-like papillae, largest in posterior region of trunk. Longitudinal musculacure in 18-19 (1 in one specimen) bundles, Oblique musculature between 134 S. J. EDMONDS longitudinal muscles forms fascicles which sometimes are only weak. Proboscis: In preserved specimens about half to fifth length of trunk, either deciduate or non- deciduate. Lateral margins tend to roll inwards so as to form a tube. Plump and almost concial in largest specimen. Setae: Two, up to 3.1 mm long, golden, connected by strong interbasal muscle. Alimentary Canal: Mouth at base of proboscis. Gut much coiled and filled with coral and shell fragments; obviously animal is able to ingest larger particles than Ochetostoma erythrogrammon and Bonellia viridis (Chuang 1962, Jaccarini & Schembri 1977). Nephridia: Two post-setal pairs, nephrostomal lips long, weakly or strongle coiled. Largest ova 0.09-0.11 mm in diameter. Anal Vesicles: Two, very large, slender, tapering distally and bearing numerous, small, brown unbranched ciliate cups or funnels. Vascular System: Dorsal blood vessel, ring vessel, two neuro-intestinal vessels and ventral vessel. Neurointestinal vessels long but fusing to form one short vessel which joins ventral vessel at about level of setae. Systematics Ochetostoma myersae Edmonds, 1963, was described from N.SW. as possessing 18-21 longitudinal muscles, two pairs of post-setal nephridia and unbranching ciliate funnels, At the time the species was considered to be different from O. baronii (Greeff) in which the ciliate funnels were described as being branched, a fact confirmed by Fischer (1895: 19). Amor (1976, p. 123), however, after studying specimens collected at Canary Is (type locality), Brazil and Galapagos Is. found that ‘amongst the 38 specimens examined there did not exist any branched outgrowths in the anal vesicles’, The examination of three specimens of O. baroni from Arrecife, Canary Is, collected by A.K. Totten and identified by A.C. Stephen (B.M. 11.7.7.37). confirms Amor’s finding that the fuhnels are unbranched, In view of this evidence (especially as Amor examined 38 specimens) the statements of Greeff and Fischer about the branching of the ciliate funnels are questionable. Consequently the chief reason given by Edmonds (1963) for separating O. myersae and O. baronii is now invalid and O. myersae becomes a junior synonym of O. baronii. Amor (1976: 123) also considers O. edax Fisher, 1946 and O. kefersteini (ten Broeke, 1925) as junior synonyms of O. baronii. O. punicea (Dartnall, 1976) is very closely related to O, baronii if the ciliated funnels of the latter are unbranched. O. punicea has 18-19 longitudinal muscles, fasciculated oblique musculature, setae about 2.25 mm long, two pairs of post-setal nephridia with spirally coiled lips and anal vesicles with unbranched ciliate cups. No interbasal muscle, however, is present and the left anal vesicle is not symmetrically placed in relation to the right. Whether the last character is taxonomically significant is doubtful. The species was described from Great Tulear Reef, off south-west Madagascar. Specimens examined and localities New South Wales: Long Reef (near Sydney) (1) AMS 3357; Collaroy, AMS W3368 (1); Minnie Waters, ‘intertidal region of low tide’ AMS coll. (1). Queensland: Bird I. (Moreton Bay) (1) SAM E1417. Distribution Eastern Australian from Sydney (N.SW.) to Moreton Bay (Qu.). Extra Australian, wide: Atlantic Ocean (Canary I., Bermuda, West Indies, Senegal, Florida, Brazil); Indian Ocean (Zanzibar, Amboina); Pacific Ocean (Papua, Loyalty I., Galapagos). FIGURES 14-15. Ochetostoma australiense. 14, anterior region dissected; 15, setae (scale line = 1.0 mm). AUSTRALIAN ECHIURANS 135 Genus Thalassema Lamarck Thalassema Lamarck, 1801, p. 28; Fisher 1946, p. 230; Stephen & Edmonds 1972, p. 452. Type-species: Lumbricus thalassemus Pallas, 1776 = Thalassema thalassemum (Fisher, 1946). Diagnosis Echiuridae with well developed, long, non-bifid but usually truncated proboscis Two ventral setae; lacking anal setae. Longitudinal, circular and oblique musculature continuous, Nephridia in one or LWo pairs; nephrostome basal and nephrostomal lips neither elongated nor spirally coiled. Thalassema sydniense Edmonds (Figs. 16-17) Thalassema sydniense Edmonds, 1960, p. 89-90, figs. 1-2, pl. la. Holotype: AMS G11219; off Watson Bay, Sydney, N.S.W. Description (based on four specimens reported in Edmonds, 1960 and two additional ones) Trunk; Small, grey-brown, sausage to sub- ovoidal; length 2.5-8 mm (most about 5), width 1-2.7, Surface covered with papillae, lying almost in tows and largest posteriorly. Musculature continuous. Proboscis: Firmly attached, as long as trunk or less, becoming narrower anteriorly. Setae; Two, golden brown, 1.0-1.1 mm long, with strongly recurved tip; strong interbasal muscle and well developed system of setal muscles, Nephridia: Two pairs, post-setal. Nephrostome on short stalk near proximal extremity of nephridium; lips expanded but not elongate or spirally coiled. One specimen with only 3 nephridia. Alimentary Canal: Very long, intestinal siphon present but no caecum. Anal Vesicles: Two, expanded towards base; surface with ciliated cups. Systematics These specimens resemble Thalassema steinbecki Fisher, 1946, which occurs along the Pacific coast of North America from California to Ecuador and which has also been reported from the Indian Ocean (Datta Gupta 1975), T sydniense differs from T. steinbecki because its nephrostomes are on short stalks or peduncles. 7. sydniense is known only from six specimens, four of which are very small. it is not a well known species and needs redescription when more specimens become available. Whether the species is a small one or whether the specimens so far collected are simply small ones is not known. Specimens examined and localities New South Wales: Watson Bay, AMS G11219 (4). Victoria: 40° 39°0"S, 144° 56’E (Bass St Survey) MV G3386 (2). No other records. Q \ 17 FIGURES 16-17. Thalassema sydniense, 16, entire animal (scale line = 1,00 mm); 17, seta (scale line = 0.25 mm), Genus Ikeda Wharton Ikeda Wharton, 1913, pp. 260-261; Fisher 1946, pp. 220; Stephen & Edmonds 1972, pp. 471-472. Type-speécies: Thalassema taenivides |keda, 1904, Diagnosis Trunk very long with longitudinal musculature thickened to form bands, Proboscis very long, non bifid. Nephridia very numerous and unpaired. Ikeda sp, The yery long proboscis of an echiuran has been noticed and collected a number of times by divers 136 8. J, EDMONDS in St Vincent Gulf, S.A. [Edmonds 1982, pl. 23 (4)]. They report that the organ Is able to extend for more than 1.5 m, The echiuran itself, however, has proved very difficult to collect on account of the depth of iis burrow and the problem of digging in sand at depths of 6-10 m. So far only one specimen has been collected. Unfortunately, it was considerably damaged so that only a limited amount of information can be given about it, Several intact probosces of other specimens have been collected, The worm resembles in some respects /keda taenioides (Ikeda, 1904), known from six Japanese specimens. A specific identification of the specimen from S.A_18 nat possible on accotint of the damage to its nephridial and anal regions. 1 am, however, tentatively assigning it to the genus Ikeda, Description Trunk: Long, slender, worm-like, rather flat jn preserved condition; 290 mm long, 7-11 mm wide, pinkish-brown when collected but dark brown when fixed. Langitudinal musculature grouped in 5 bands prominent externally; numerous. small sub-globular papillae cover surface of much of trunk. Proboscis; Flat, about. 400m long, 5-10 wide, with margins in fixed specimens slightly frilled; one surface cream-brown in colour marked with almost transverse brown-black stripes; posterior region of proboscis (near mouth) modified to form cup-like Structure, Seftae: Two, about 10 mm long, with well developed setal muscles. Nephridial region much damaged, Anal Vesicles: Missing, Alimentary Canal: Very long and convoluted, Some eves with maximum diamrer 0.35-0,38 mm entangled im gut. Specimens examined end localities ‘Kemps Ground’, off Glenelg, St Vincent Gulf, S.A., at 9-10 m, 13 March, 1986, ane spec. coll, N. Holmes and S. Parker; SAM E1509; separate probosces of other specimens SAM E1587, Coffin Bay (near Black Spririgs), Eyre Peninsula, S.A- (probosels only). ACKNOWI EDGMENTS Thanks are due to the following far help with specimens: from W.A,, Mrs L. Marsh, Mra S$, Slack-Smith, Dr E. Hodgkin and Dr B. Wilson: fram 8.A4., I.M. Thomas, Mrs K, Gowletl-Holmes, N, Holmes, Miss H. Kald and W, Zeidler; fran) Vic, Dr B, Smith and Mry H, Black; from Tas., Dr A. Dartnell and Dr A. Green; from N.SW., Dr P. Hutchings, Miss P, Wearne, Miss E. Pope and Miss E. Bennett, trom Qu,, Prof W. Stephenson, W. Green, S. Cook, ©. Kelly and Mrs M. Specht. Mr P. Kenipster (University of Adelaide) took the photographs and Dr 1. Beveridge (Institate of Medical and Veterinary Sciience, Adelaide) made sections of the body wall of one apecies, Mr R. Sims and Mr E, Easton (British Musuem of Natural History, Londen) and Dr M. E. Rice (Smilhsonian Institute, Washington D.C.) kindly sent specimens on loan, Dr E, Matthews (SAM) helped with translations {rom Russian. REFERENCES AGIUS, L, & JACCARINI, V. 1981. Development of Plament if tie echiuiran worm Bonellia viridis, J, Zoal., Lond. 193; 25-31, AMOR, A. 1976. Genera and species of Echiura known from South America Proc Internat. Symp, Biol. Sipunecula and Echiira Kotor 1970 (publ. 1976). 1 119-125 ANNANDALE, N_ 1922. The marine clement in Me fauna of the Ganges. Bi/dr Dierk, 22; 143-154, AUGENER, H. 1903. Beitrage vir Kenntnis der Gephyteen nach Uutersuchung der im Gottinger zoologischen Museum befindlichen Sipwncoliden und Echiuriden. Arch. Natyreesch, 69: 297-371. BISEWER, R. 1984. A key to the species of Anelassoriivnchus (Echiura) wilh. a description of a new species from the east coast of South Africa S. “fn & Zool, 19 {L): 16-21 BROEKE, A. ten, 1925. Westindische Sipuneiliden und Echiuriden. Sijdr. Dierk, 24; 31-6, CHEN, Y. & YEH CHEN-CHANG, 1958, Noles on some Gephyrea of China, Sera Zoologica Sinica We 265-177 (With Brglish transl}. CHUANG, SH. 1962. Feeding mechanisnr of the eghiuroid Ocketastoma erythrogramiman Leuckart & Rueppell, 1828, Biol. Bull, Mar, Lab, Woods Hale 123: 80-85. DARIN, W.J, 1916, The marine biology of Western Australia. Proc. R. Soc. Wo Aust, 1 1-27. DAKLN, W.J. 1952, Gephyrea, Jn “Australian Seashores’. Angus & Robertson, Sydney, Pp. 157-158. 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Tras, R. Soe, 5, Aust, 83: 89-96. EDMONDS, 8.5. (963. Two tiew echiuroids (Echiuroidea) from Australia, Trans, R. Soe, 8, Aust. 87: 243-247. EGMONDS, §.J, 1966, Port Phillip Survey 1957-1963: Sipunculoidea and Echiuroides. Mem, Nata, Mus, Wier, 27; 175-178, EDMONDS, §,J. 1982. Echiurans. In S.A. Shepard & I, M. Thomas (Eds). “Marine [Invertebrates of Southern Australia’ Parr |. Handbook of the Flora and Fauna of South Australia, Govt Printer, Adelaide FISCHER, W. 1895. Die Gephyrcen des narurhistorischen Miseums 2u Hamburg, ADA. Geb. aaturw. Mamburg 13; 1-24. FISCHER, W, 1919 Gepliyreen der Sud-westkuste Australiens. Zoal, Anz. Sb: 177-285. FISCHER, W, 1921, Results of Br E, MyGbere’s Swedish scientific expedition to Australia 1910-1913, 27 Gephyreen. K. svenska Vetensk - Akad. Hund. 61 (8): 1-8. FISCHER, W, 1926, Sipuficuliden und Echiuriden der Hamburger Sudsee-Expedition 1908-1908. Mitr, Zool. Staatinst, und Zool, Mus, Hamburg 42 109-117. 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Tokyo 24; 1-64 JACCARINI, V. & SCHEMBRI, PJ. 1977, Feeding and particle selection jn the echiuran worm Bonetlia viridis Rolando (Echiura ‘ Bonelliidae). 1 aq. Afar Biol. Ecol. 28; 163-168, JONNSTON, T.H. & TIEGS, GW. 1919. Pseudobanellia, a new echiuroid from the Great Barrier Reef. Proc. Linn. Sac, N.SWE 44: 213-239. JOUNSTON, TWH, & THEGS, QW, 1920, A new species of Bonellia trom Port Jackson, Sydney, Ree. Aust, Mus [3% 73-76. LAMPERT, K, 1883, Uber einige neue Thalassema, 2. Wiss. Zool, 34: 334-342, LAMARCK, J.B.P de M. LROL. ‘Systeme des animaux sans vertebres’, 432 pp, LEUCKART, MS. & RUEPPELL, W.7S. 1828. Neue wirbellose Tiere des roten Meers, In W.P.S, Rueppell. ‘Atlas zur der Reise in Nordliche Atvica’, |, Zoologic: 6-9. MACKIE, 6.0. 1961. Echiurids from the Canary Islands, Ann. Mag. Nat, Hist. 13: 247-251. MATHEW, J. (976. Integuments in echiurans, In Proc Internal. Symp, Biol. Sipuncula and Febivra, Kotor, 1970 (publ, 1976) [hy 151-158. MONRO, ©.A, 1927. 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FIGURES 18-24. 18, Metabonellia haswelli; 19, Pseudobonellia biuterina; 20, Anelassorhynchus porcellus porcellus; 21, Anelassorhynchus porcellus adelaidensis; 22, Listriolobus brevirostris; 23, Ochetostoma australiense; 24, Ochetostoma baroni, FURTHER OBSERVATIONS ON PENTATOMIDS (ARTHROPODA) PARASITIC IN AUSTRALIAN REPTILES AND MAMMALS BY J. RILEY & D. M, SPRATT Summary A collection of adult and nymphal pentastomids representative of at least four genera is described. Two species of Raillietiella and one species of Parasambonia from snakes (Pseudechis australis, Pseudonaja textilis and Cryptophis nigrescens) are probably new, but more specimens are required before their status can be confirmed. Mature and immature Waddycephalus longicauda, W. superbus and W. punctulatus (all Riley & Self 1981) are identified from snake hosts but specific determination of five lots of specimens was not possible. Evidence endorses an earlier suggestion that there may be two species of Waddyccephalus in tiger snakes ; W. scutata from island populations and an unnamed species from mainland populations of the Notechis scutatus/ater complex. Nymphal specimens of Waddycephalus from marsupials (Parantechinus apicalis and Dasykaluta rosamondae), a snake (Cryptophis nigrescens), a gecko (Heteronotia binoci), a skink (Hemiergis decresiensis) and frogs (Ranidella remota and Palmatorappia solomonis) all bear characteristic double hooks. The accessory spine above the hook arises from a point midway between the hook and the fulcrum and appears to be an integral and functional part of the hook. Armillifer australis Riley & Self 1981 is described from infections in four pythons (Morelia amethistina and Morelia spilota) ; the latter is a new host record. A single nymph recorded from the body cavity of Rattus leucopus is identified as A. australis on the basis of abdominal annulus counts. FURTHER OBSERVATIONS ON PENTASTOMIDS (ARTHROPODA) PARASITIC IN AUSTRALIAN REPTILES AND MAMMALS J. RILEY & D, M, SPRATT RILEY, J. & SPRATT. D, M, 1987. Further observations on Penjastomids (Arthropoda) parasitic in Australian reptiles and mammals, Ree, 8, Aust, Mus. 21(2): 139-147. A collection of adult and tiymphal pentastomids representative of at least four genera is described. Two species of Raillietiella and one species OF Parasambonia from snakes (Pseudechis australis, Pseudonaja textilis and Cryptophis nigrescens) are probably new, but more specimens are required before their status can be confirmed. Mature and immature Waddyecephalus fongicauda, W; superbus and W. puncrulatus (all Riley & Self 1981) are identified trom snake hosts Dut specific determination of five lots of specimens was not possible, Evidence endorses an earlier suggestion that there may be two species of Waddycephalus in liger snakes; Wi scufata from island populations and an unnamed species from mainland populations of the Notechis sculalius/ater complex, Nympha) specimens of Waddycephalus from marsupials (Parantechinus apicalis and Dasykaluta rosamondae), a snake (Cryptaphis nigrescens), a gecko (fleteronotia binoei), a skink (Hemiergis decresiensis) and. frogs (Ranidella remota and Palmatorappia solomonis) all bear characteristic double hooks, The accessory spine above the hook arises from a point midway between the hook and the fulcrum and appears to be an integral and functional part of the hook. Armmillifer ausiralis Riley & Self, 1981 is described from infections in four pythons (Morelia amethistina and Morelia spitota); the latter is a new host record. A single nymph recorded from the body cavity of Rattus /eucopus is identified as A, gusiralis on the basis of abdominal annulus counts, J. Riley, Department of Biological Sciences, University of Dundee, Dundee DDL 4HN, Scoiland, UK. & D. M. Spratt, Division of Wildlife and Rangelands Research, CSIRO, P.O, Box 84, Lyneham, A.C.) 2602. Manuscript received 21 October 1986. In an historical review of Australian Pentastamida, Riley, Spratt & Presidente (L985) recorded seven genera comprising 17 species occur- ring in Australian reptiles and mammals, and identi- fied nymphal Weddycephalus spp. and Armillifer spp. from marsupials, This paper reports primarily on a pentastomid collection in the South Australian Museum (SAM) and describes further adult and nymphal material, attributed to the genera Waddycephalus and Armillifer, from reptiles, amphibians and mammals. The double nature of the hook of nymphal Waddycephalus (Riley et al. 1985) is. confirmed, as are earlier observations (Riley & Self 198ib) of significant anatomical differences beiween mainland and island forms of Waddy- cephalus infecting the same species of snake, Two Jarge raillietiellids and two parasambonids from snakes are described, however more material. is required before their specific status can be con- firmed as new. MATERIALS AND METHODS ‘The material exantined in this study was collected primarily in eastern mainland Australia, Tasmania and neighbouring offshore islands, It is supplemented by nymphal Waddycephalus (or Elenia?) spp, from ibe Solomon Islands and New Guinea. The methods are those outlined in Riley e¢ af- (1985) and the hooks of railhietiellids were measured accotding to the convention of Ali ef al, (1982) i. barb length AB (notation AD in error, p. 42, Riley et al, 1985), shank length BC, Overall hook length of the double hooks of nymphal Waddycephalus spp. was measured according to the convention illustrated in Figure 2. All measurements are in micrometers with the exception of body length, which is in millimetres. Most specimens are deposited in the South Australia Museum (SAM), Adelaide; two lots are deposited In the Queensland Museurn (QM), Brishane Reptile nomenclature follows Cogger, Cameron & Cogger 1983; dasyund marsupial nomenclature follows Archer 1982, Order CEPHALOBAENIDA Raillietiella spp. trom snakes Railiietielta sp. a Material Examined From lung of Pseudechis qustralis (Gray), locality unknown (died in Melbourne Zoo), in SAM No, NI9BO183, SPRATT J. RILEY & D. M. 140 AUSTRALIAN PENTASTOMIDS 14] Description Female (n = 4). Length 33-52 (K = 42), with 45 or 6, 41 or 2, 36 (2) and 39 (?) annuli respectively. Posterior hook of 52 mm specimen, AB 240, BC 370. Male (n = 1). Length approx. 9, annuli uncount- able and therefore slide-mounted. Posterior hook, AB 135; BC 220. Base of copulatory spicule massive, maximum diameter 520 and covered with a reticulum of tubular elevations (Fig. 1A). Discussion The heavily ornamented male spicule, 520 um across at the base (Fig. 1A), is virtually identical to those of R. orientalis and R. agcoi (Ali, Riley & Self 1982). R. orientalis infects colubrid, viperid and elapid snakes in south-east Asia and Taiwan whereas R. agcoi is found only in cobras in the Philippines. The females of these two species are distinguished in a number of ways: the former has bigger hooks (see Fig. 3, in Ali et a/. 1982), more annuli (33-47 contra 30-35) and is generally longer and stouter than R. agcoi. The overall shape of the present species, and its hooks, are reminiscent of R. agcoi but its annulus count, though variable, is within the range of R. orientalis. The host, the king brown snake, Pseudechis australis, is an endemic species and its raillietiellid parasite may be unique by virtue of geographic isolation. This is probably a new species, but, because it combines important characteristics of these two closely related species we have left it unnamed, pending more specimens and more refined diagnostic techniques. Raillietiella sp. b Material Examined From lung of Pseudonaja textilis (Dumeéril, Bibron & Duméril), Townsville, Queensland, in SAM No. N1985149. Description Female (n = 2). One headless abdomen; length other specimen 50, annuli uncountable, specimen therefore slide-mounted. At least 30-40% of eggs in uterus contain fully-developed primary larvae; the specimen is therefore mature. One posterior hook measured, AB 400; BC 510. FIGURE 2. Diagrammatic representation of hook of nymphal Waddycephalus sp. The distance between the larger arrows indicates our measurement of the overall length (a = apodeme; ah = accessory hook; f = fulcrum; hb = hook barb) (scale bar = 120 ym). Discussion Apart from the species recorded above, only two raillietiellids are known from Australia; R. amphiboluri from the bearded dragon, Amphibolurus barbatus (Cuvier), (Mahon 1954, Riley et a/. 1985) and R. scincoides from the eastern blue-tongued lizard, Tiliqgua scincoides (White), (Ali, Riley & Self 1984): the latter has blunt-tipped posterior hooks and R. amphiboluri is smaller than the present species with much smaller hooks (AB 200-220; BC 370) (Ali et al. 1985) (notation AD in error). The intact specimen clearly belongs to the Group VI taxon of raillietiellids (Ali et a/. 1985) which includes all of the species from snakes. The two species, R, orientalis from south-east Asia and Indonesia and R. agcoi from the Philippines are, zoogeographically, most proximate to the present specimen, but it has much longer hook barbs (dimension AB) than either of these species (compare with Fig. 3 in Ali, Riley & Self 1982). This is almost certainly a new species but the poor state jn ee FIGURE 1. A, Copulatory spicules of male Raillietiella sp. from Pseudechis australis showing massive base covered with tubular elevations (scale bar = 500 um). B. Cephalothorax of male nymph of Waddycephalus sp. from mesentry of Satanellus hallucatus [described by Riley ef al. (1985)]. The double nature of the outer hooks is obvious but the inner hooks are not in the plane of focus (m = mouth) (scale bar = 500 pm). C. Outer hook of nymph of Waddycephalus sp. from Hemiergis decresiensis showing spinous extension. The back of the extension forms a long apodeme (a) (f = fulcrum) (scale bar = 100 zm). D. Hook of adult Armillifer sp. from python (S.A.) illustrating typical unornamented porocephalid hook (a = apodeme; f = fulcrum) (scale bar = 200 pm). 142 J, RILEY & D. M. SPRATT of preservation and lack of males precludes specific identification, Order POROCEPHALIDA Parasambonia spp. from snakes Parasambonia bridgesi Riley & Self Material Examined From lung of Pseudechis porphyriacus (Sha\w), Healesville Sanctuary, Victoria, in SAM No. N1980173. Description Female (n = 2), Length 26, with 50 and 54 pre- vaginal and post-vaginal annuli. Male (n = 1). Length 7, with 53 annul Discussion There are no uncertainties regarding the status of these specimens, All of the characters fall well within the ranges described for P bridgesi by Riley & Self (1982). Parasambonia sp. a Material Examined From lung of Austrelaps superbus (Gimther), 5 kim south of Bowral, New South Wales, in SAM No, N1986192, Description Male (n = 1 plus 2 anterior ends), Length 8, with 51 annuli. Heads slide-mounted, outer hook with finger-like extension, AD 175, 190; BC 95, 100. Discussion The outer hooks possess the projecting spine characteristic of the genus Parasambonia, The low annulus count is more characteristic of P minor than P bridgesi, however the hook dimensions are much smaller than those of A minor, the dimensions of which do not overlap with the much larger hooks of P. bridgesi (Riley & Self 1982). The absence of fully gravid females precludes confident specific identification. AR minor, but not RB bridgesi, has been recorded from the copperhead (Riley & Self 1982). Parasambonia sp, b Material Examined From lung of Cryptephis nigrescens (Ginther), Mogo 5.K, New South Wales, in SAM Noa N1986191, Description Female (n 1). Length 29, annuli uncountable. Outer hook with projecting spine, AD 465; BC 245, Discussion Hook dimensions in this gravid specimen are larger than those reported in species of Parasambonia from Australian snakes (Riley & Self 1982), suggesting that it represents a new species. Additional specimens in good condition are required to resolve this matter. The stomach of this small-eyed snake contained a partly-digested eastern water skink, Sphenomorphus quoyii (Duméril & Bibron)- - Waddycephalus spp. from shakes Waddycephalus longicauda Riley & Self Material Examined From lung of Demansia psammophis (Schlegel), Queensland National Parks and Wildlife Service, Moggill, Queensland, in QM No. W12193. Description Female (n = 1). Length 27, with 49 pre-vaginal and 7 post-vaginal annuli, preadult, Male (n = 6). Length 8-11, with 56-59 annuli (X = 57). Discussion The features of this material, particularly the long and finely tapered post-vaginal tail, are characteristic of HW longicauda (Riley & Self 1981b), Waddycephalus sp, a Material Examined From lung of Pseudonaja lextilis (Duméril, Bibron & Duméril) taken at Halls Gap, Grampians, Victoria, in SAM No, N1980204. Description Female (n = 1). Length 37, with 63 pre-vaginal and 4 post-vaginal annuli, Hooks removed from one side, BC 530, 540; AD 860, 890. Discussion The only pentastomid described from PF textilis is a female which was tentatively identified as porphyriacus purely on the basis of similarities in annulus counts: hooks were not measured (Riley & Self 1981b). The present fully mature specimen has far fewer pre-vaginal annuli (63 contra 75) than the type series of H4 porphyriacus. Hook dimensions are very much smaller than those of H¢ porphyriacus and similar to W. superbus from a AUSTRALIAN PENTASTOMIDS i] copperhead, Austrelaps superbus (Giimher), and to an unnamed species from a mainland tiger snake, Notevhis scutatus (Peters) (Riley & Self 1981b), Two other characters, the number of annuli and the attenuated caudal extremity, also place it close to these species. However, the type series of HW. superbus was denved from copperheads taken in Tasmania and nearby islands (Riley & Self 1981b), as was a tnore recently described infection (Riley eral. 1985). To date, WH. superbus is recognised only as an island species, although copperheads are known on the mainland where their range overlaps that ot F fextilis. Different dietary preferences have been reported in these snakes (Cogger 1983) however, the food habits of different populations of a snake species over ils geographic range are not well known. Recent evidence suggests strongly that dietary preference is different in regions where food resources are skewed or Limiting (Schwaner 1985), Until more specimens become available the status of the Haddycephalus from Pseydonaja textilis remains uncertain, Waddycephalus sp, bh Material Examined Froes lung of Notechiy ater atger Kinghorn, Reevesby |, South Australis, in SAM No. NIS8SISI, Description Female {(n = 2). Both specimens in very poor condition, length about 36-38, na detail of annulation could be discerned. Hooks fram one Specimen, BC 400, 410; AD 705, 690. Discussion The hooks are much smaller than ihase of all recognised species of Waddycephalus except VW Scusata, algo taken from a tiger snake, on St Francis Island, South Australia, Riley & Self (1981b) recorded the host of the type as Notechis scutata (= scufaius), based on the collector’s label in the vial. Cogger (1983) recognised N. sculatus as a purely mainland species being replaced by N. afer on islands off the coast of South Australia. However, recent studies of morphological variation in tiger snakes on Kangaroo Island have revealed banded, unbanded, red-bellied and melanistic forms believed fo belong Lo the same species complex (Schwanec 1984), Thus W{ scufata js currently recopnised as a parasite only of island populations of the N. seu/atns/ater complex, its principal distinguishing characteristic being its particularly small hooks. Hook dimensions in the present specimens form a cluster distinat from. Wt scwtata (compare with Fig. 6 In Riley & Self 1981b and below) and sugyestive of a new species of Waddyeephalus. Wiaddycephalus sp. c Moterial Examined From lung of Nolechis scutatus (Peters), Grampians. Victoria, in SAM No, NISROI71. Description Female (n = 7), Length 32-43 (k = 38), with 60-63 pre-vaginal (x = 61.5) arid 2-3 (X — 3.2) past- vaginal annuli.. Hooks taken from one side of 32 mim and 40 mm females, BC 495, 535; AD 885, 910 respectively. Male (n = 4), Length 14-20 (R = 16), 62-68 annuli (X — 64), Discussion Jn their review of the genus Waddycephalus Riley & Self (J98)b) separated the species from tiger snakes (Wolechis spp.) into two groups distinguished principally by marked differences in hook size: small hooks are churacteristic of WH. seu/ata from island populations of the Notechis scutatus/ater complex whereas larget hooks are found in specimens from mainland tiger snakes (see Fig. 6 in Riley & Self 1981b), ‘They concluded that geographical isolation was responsible for the observed differences and that two hosts may be involved. The present findings substantiate these differences and combine to suggest indeed that there may be iwo species of Waddycephalus infecting tiger snakes. More sophisticated diagnostic techniques, preferably utilizing live matenal, are required 10 confirm this postulate. Waddycephalus superbus Riley & Self Material Examined From hing of Ausirelaps superbus (Giinther), (a) Launceston, (b) Longford, Tasmania, in SAM Nos. (a) N1980175, (by NI9BO205. Description Female fa) (n = 3), Length 37-41, with $9-63 pre- vaginal and 3-4 post-vagiral annuli, Hooks from one side of 40 mm specimen, BC $30, 530; AD 900, 970, Female (6) (n = 2). Most of abdomens missing, annuli uncountable; both apparently mature. Hooks dissected from one side of both females, BC 525, $25; AD 840, 840 respectively. Male (b) (n = 1). Length 15, possibly 62 annus. Discussion These specimens are Very similar to the type serles of HW! superbus (from the same host species also taken in Tasmania) except that one specimen From Launcesion has rwo more abdominal annuli and the hooks are shightly larger than those described by 144 J. RILEY & D. M, SPRATT? Riley & Self (J981b) and Riley er wl. (L985), Nevertheless, all of the honk dimensions measured to date combine to form a discrete cluster group and this species at least, is now well characterized. All specimens recovered thus far come [ram Tasmania, endorsing the suggestion that Waddycephalus teretiusculus Baird, 1862, the type species of the (axon and also occurring in the copperhead, |s prabably a maintand species (Riley & Self 1981b, Riley ef a/. 1985). Lungs of specimens of A. superbus held in the Australian National Wildlife Collection were examined for teretiusculus from the following mainland localities (numbers of Specimens in parentheses) but pentastomids were not recovered: Mi Gingera, ACT (1), Ginini Plats, ACT (1); Captain’s Flat, NSW (3); Pepper Creek on Big Badja Mountain via Numeralla, NSW (1); Kosciusko National Park near Kiandra entrance (1) and near Peak River, NSW (1); Tumbarumba, NSW (1) Portland, Vic. (2); Flinders 1. (1). Waddycephalus sp. d Material Examined From lung of Drysdalla coronoides (Giinther), Fenelon !., South Australia, in SAM No, N1985152, Description Female (a = 1). Immature, slide-mounted. Length 10, with 56 pre-vaginal and 5 post-vaginal annuli. Hook measurements BC 280, 300; AD 430, 420. Male (n = 1). Immature, slide-mounted, Length 9, with 64 annuli. Discussion The host snake was originally recorded as Denisania coronoides but species of Drysdalia were formerly Included in the genus Denisonia. The immature state of the present specimens precludes specific identification, Waddycephalus punctulatus Riley & Self Materiul Examined From lung of Dendrelaphis puncrlata (Gray), Northern Territory, in SAM Noa. N1984153. Deseription Female (n = 1). Length 33, with 52 prevaginal and 11 posl-vaginal annuli, Male (n 1), Length 14, possibly 4! annuli. Discussion There is no confusion concerning the status of these specimens from the common tree snake; their size and annulus number agree well with the original description of WY punctulatus (Riley & Self 1981b), Waddycephalus sp. e Material Examined Fram lung of Morelia spilota (Lacépéde}, Sa Francis §., South Australia, in SAM No, N1985154, Descriptian Female (n = 1). Immature, length 9.5, with 52 pre-vaginal and 4 or 5 post-vaginal annuli, Discussian The anterior part of the cephalothorax, inchiding the pair of inner hooks and the mouth are missing. The outer hooks lack the projecting spine characterisuic of Parusambonia spp. (Riley & Sell’ 1982). The position of the vagina places the specimen in the family Sambonidae and the annulus count indicates that it is a species of Waddycephalus hut, it has far fewer annuli than the immature female described previously from the same host species and tentatively identified as HW. porphyriacus (Riley & Sell 1981b), Nymphal Waddycephalus spp. Material Examined {i) from Parantechinus apicalis (Gray), locality unknown, in SAM Ne, NI9B0210. (li) Encysted in a skink, Hentierpis decresiensis (Cuvier), South Australia, in SAM Na, (985155, (ui) Eneysted in abdomen of Daspkalura resatnondae Ride (a) Woodstock Station, (b) Abydos Station, near Marble Bar, Western Australia, in SAM Nos, (a) 1985156, (b) NI980182. (iv) One nymph, from below posi-orbital skin of a frog, Ranidella remota Tyler & Parker, Papua New Guinea, in SAM No. N1985157. (Vv) 14 nymphs encysted in intestinal connective tissue af Cryplophis nigrescens (Giinther), Mogo S.F, New South Wales, in SAM No, N1986190. (vi) 3 nymphs eneysted in gecko, Heteronotia binvei (Gray), Girraween National Park, Wyberba, Queensland, in QM Na. W12194, Description (i) Three nymphs dissected from cysts and slide- mounted. Length about 6, | specimen (sex unknown) with 74 annuli, 1 male with 70 annuli. All hooks double, overlain by accessory spine (Fiz, 2) the base of which arises from a point between fulcrum and hook, Spine an integral part af hook and attached to fulcrum only by thin, flexible sheet of cuticle, Overall hook length (measurement as |llustrated in Fig, 2) 300-340, (ii) Three nymphs dissected from cysts and slide mounted, Length about 4-5, with 62-63 annuli, sex indeterminable. Hooks double, overall length 210-250 (Fig. 1C). AUSTRALIAN PENTASTOM(DS 145 (iia) All dissected from cysts and slide-miounted, Length about 6, with 56-62 annuli (% = 60), Hooks double, overall length 250-280, (iiib) Six small cysts opened and nymphs slide- mounted, large composite cyst containing many larvae left intact, Length 4-5, wilh 56-59 annuli (3 counted). Hooks double, overall length 230-260, (iv) Male nymph, length approximately 4, with 56 annuli, Hooks double, not dissected and measured. (v) Three nyoiphs.slide-mounted. Length about §, with 56-58 annuli (% = 57), Hooks dauble, overall length 170-180. (vi) Length about 5, with 56-61 annuli (% = 58). Hooks double, overall length 195-215, Discussion Riley ef a/. (1985) ascribed a double-hooked male nymph from the northern quoll, Satare/lus. hallucatus (Gould), to the genus Waddycephalus exclusively on the number of annuli, which vary from 5$-78 in adult males of the genus (Riley & Sell 1981b). This is considerably more than occurs in the two other genera which may have double- hooked larvae, Elenia and Parasambonia (Reymons (929, Riley & Self 1982, Riley ef a/. 1985). All of the present specimens are placed in the genus Haddveephalus for the same reason, although specific identification is nol possible. The specimens from D. rosamondae probably belong (0 the same species, those from Abydos Station being at a slightly earlier stage of development. The nymph trom &. remota may be punctulatus, This species was first described from the common tree snake Dendrelaphis punctiulata (Gray) in Australia (Riley & Self 1981) bul this host also occurs in New Guinea (Cogger 1983), The prey of tree snakes consist of frogs and birds, although reptiles and small mammals are occasionally eaten (Cogger 1983). Clearly frogs are probable intermediate hosts of Ho punciulatus, Either Waddycephalus or Elenia sp Material Examined One nymph, from submandibular lveophatic sac of a frog, Palmatoreppia solamenis (Sternfeld) Solomon. Islands, in SAM No, NL98SIS58.. Deseripiion Male. Length 5, with 48 annuli, Hooks double, overall length 220-230. Discussion ‘The generic status of this nymph 1s uncertain. Waddycephalus komodoensis and Wo radiata are known from Indonesia (Riley & Self 198th) and Elenia vitiensis is known from the tslands of Fiji (Heymons 1932). The low annulus count of the specimen may just preclude it being a species of haddyeephalus, the lowest annulus number known in mature males is 52, occurring in HW Kormodaensis (Riley & Self 1981b), Armillifer sp. fram snakes and rodents Armillifer australis Riley & Self Material Examined (i) From yiscera (the specimens probably in- habited the membranous lung which fs often mistaken for the abdominal cavity) of a python (species unknown), South Australia, in SAM Na. N1980207, (i) From lung of Morelia amethistina (Schneider), Melbourne Zoo, in SAM No N1980206, Gili) From lung of Morelia spilota (Lacépéde), Queensland, in SAM No, Ni980208. (ivy) From lung of Morelia spiloia (Lacépéde), Melbourne Zoo, in SAM No. N1I980172. (v) Encysted nymph from body cavity of Ratius lercopus cooktownensis Tate, Queensland, in SAM No, N1980209, Descriplion (i) Female (n = 2) Length 63 and 67, both with 3L annuli and 2 incomplete annul on terminal segment. Hooks removed from one side of one female, AC 410; AD 624 (Fig. 1D), Male (n = 2), Length 21 and 22, with 40 snnull; first 12 annuli with pair of projections pointing backward from posterior lateral angles. (ii) Female (n = 15), Length 34-53 (one punctured female not included) (X = 42.5), with 29-32 annuli (% = 30,6) and 2 (or in two cases, 3) incomplete armuli on terminal segment, Hooks from a 49 mm specimen, AC 440; AD 635. (iii) Female — mature (n = 1). Length 47, with 32 annuli (plus two incomplete segments terminally). Female — immature fn = 4). Lensth 16-27 (X = 19), with 29-32 annuli (plus 2 incom- plete). Male (n = 3). Length 16-17, with 36-37 annuli, anterior 1-12 bearing backward-pointing projections, (iv) Female (n = 1), Length 46, with 32 annuli (plus 3 incomplete). (v) Female?) (n » 1). Length 6, with 31 or 32 annuli. Hooks simple but could not be measured. Discussian Adult specimens from the four snakes are vomistakably 4, australis and all characters accord perfectly with those of the type series (Riley & Self 198la). Morelia spilota is a new host record, 146 J, RILEY & BM. SPRATT Hooks of the nymph from R, /evcoprs are simple (i.e, without an accessory spine) and (he annulus count is within the range (29-35) of mature female A, atistralis (Riley & Sell 19814), The present specimen almost certainly belongs to this taxon, as other species of Armillifer described from Australian hosts have more annuli (Riley & Self 198la, Riley ef wil. 1985), DiscUSSION In our earlier review of pentastomid parasites in Australian reptiles and mammals (Riley e/ a/, 1985) we noted that the state of our knowledge of taxonomy ts embryonic, and this is particularly true of the genus Waddycephalus, Our original finding of a double-hooked larva, which we attributed to the genus Haddycephalus rather than Blenia solely on the basis of the number of abdominal annuli, was the first implication of mammals as intermediate hosts in this genus. This single male larva, from the mesentery of Satanellus hallucatus was cleared and mounted (Fig. 1B) and we observed that the sharp spinous extension overlying the hook appeared to be an integral part of it and separate from the fulcrum. This is unlike the situation in the related genus Sambonia where the accessory spine is clearly an extension of the fulcrum (Fain & Mortelmans 1960), The relative abundance of nymphal Waddycephalus material in the present study has permitted more detailed observations of hook morphology and these have confirmed our earller interpretation. The spine is a functional part of the hook, 1b possesses an apodeme, onto which muscles attach and extend from it down into the fulcrum (contrast Figs IC and D), The relative positions of the hook, its spinous extension and the fulcrum sré presented diagram- matically (Fig, 2). The seven species of Maddycephalus currently recognised in Australia infect boid, colubrid ane elapld snakes (Riley & Self 19816) which prey upon a variely of vertebraies (mostly frogs, lizards or mammals — see Cogger 1983) and the present report of Waddycephalus nyniphs from these three classes of vertebrates is to be expected, particularly since vertebrate intertriediate hosts are usital in porocephalid life-cycles (Nicoli & Nicoli 1966), Also, there is growing evidence from experimental infections (Esslinger 1962, Vargas 1970, Winch & Riley 1986), and from recoveries of nymphs in intermediate hosts (Sachs, Rack & Woodtord 1973}, that the definitive number of annuli is present in porocephalids by the infective stage. Our tentative diagnoses are based on the assumption that this occurs in the genus Waddycephalus. From the viewpoint of host dietary regimen, itis equally likely thal the related genera Parasambonia and Elenia also utilize vertebrate intermediate hosts but in all cases, experimental evidence of these life-cycles is required, ACKNOWLERGMENTS We gratefully acknowledge che co-operation of Dr David Lee who arranged loan of specimens from the South Australian Museum, Mrs F, Walter who examined lungs of mainland copperheads for pentastomids, Br Terry Schwaner who read an earlier draft of the manuscript and offered! valuable comment on the biology of snake species, and Dr (an Beveridge who collected many of the speciinens and ¢riticized an earlier draft of this work. REFERENCES ALI, J, HL, RILEY, J, & SELF, J.T. 1982. A revision of the taxonomy of Ruillictiella boulengeri (Vaney & Sambon, 1910) Sambon, (410, 8. erientalis{Hett, 1915) Sambon, 1922 and R. weca/ Tubangul & Masilungan, 1956 (Pentastomida: Cephalobaenida). Sys/. Parasitol. 4: 285-301. ALL J. H., RILEY, J, & SELB, J.T) 4984. 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Some observations on the taxonomy and systematics of the pentastomid genus Armillifer (Sambon, 1922) in South East Asian and Australian snakes. Syst. Parasitol. 2: 171-179. RILEY, J. & SELF, J. T. 198ib. A redescription of Waddycephalus teretiusculus (Baird, 1862) Sambon, 1922 and a revision of the taxonomy of the genus Waddycephalus (Sambon, 1922), pentastomid parasites of Asian, Australian and Indonesian snakes with descriptions of eight new species. Syst. Parasitol. 3: 243-257, RILEY, J. & SELF, J. T. 1982. A revision of the pentastomid genus Parasambonia Stunkard & Gandal, 1968: a new generic character, a description of the male and a new species. Syst. Parasitol, 4: 125-133. RILEY, J., SPRATT, D. M. & PRESIDENTE, P. J. A. 1985. Pentastomids (Arthropoda) parasitic in Australian reptiles and mammals, Aust, J. Zool. 33: 39-53. SACHS, R., RACK G., & WOODFORD, M. H. 1973. Observations of pentastomid infestation of East African game animals. Bull. epizoot, Dis. Afr. 21: 401-409. SCHWANER, T. D. 1984. The identity of red-bellied black snakes on Kangaroo Island. Trans. R. Soc. S. Aust. 108: 137, SCHWANER, T. D. 1985. Population structure of black tiger snakes, Notechis ater niger, on offshore islands of South Australia. Jn G. Grigg, R. Shine & H. Ehmann (Eds). ‘The Biology of Australasian Frogs and Reptiles’. Surrey Beatty & Sons, Sydney. Pp. 35-46. WINCH, J. M. & RILEY, J. 1986. The development of Sebekia oxycephala (Pentastomida) from a South American crocodilian (Caiman sclerops) in experimentally infected fish. Z. Parasitenkd 72: 251-264. VARGAS, M. V. 1970. A contribution to the morphology of the egg and nymphal stages of Porocephalus stilesi Sambon, 1910 and Porocephalus clavatus (Wyman, 1847) Sambon, 1910 (Pentastomida). Revita Biol. trop. 17: 27-89, LOOKING FOR DITJI-MINGKA BY L. A. HERCUS Summary Ditji-mingka was an important Aboriginal site in the north-east of South Australia. The present paper contains a Wangkangurru text describing what has happened to this site. LOOKING FOR DITJI-MINGKA L. A. HERCUS L. A. HERCUS 1987. Looking for Ditji-mingka. Res. S. Aust. Mus. 21(2): 149-156, Ditji-mingka was an important Aboriginal site in the north-east of South Australia. The present paper contains a Wangkangurru text describing what has happened to this site. L. A. Hercus, Faculty of Asian Studies, Australian National University, P.O. Box 4, Canberra, Australian Capital Territory 2600. Manuscript received 13 May 1985. Ditji-mingka, ‘Sun-Cave’ (lit. ‘Sun-Hole’), south- west of Etadunna on the Birdsville Track in north- eastern South Australia, was one of the most important places in Diyari mythology. Evidence of this comes from J. G. Reuther who worked for eighteen years (1888-1906) as a missionary amongst Diyari and neighbouring people at Killalpaninna on the lower Cooper. There was nobody who could rival the achievements of the Reverend Reuther in documenting the languages and traditions of the Lake Eyre Basin. Without his great work of fourteen volumes, recently translated by P, Scherer (Reuther 1981), much information would be totally lost. He gives the following account of the Ditji- mingka site in his list of place-names. Ditji = ‘sun’; minka = ‘cave’. ‘There is a cave at this spot, where the female sun muramura is said to have first risen. But since she made conditions too hot for her followers (‘people’) she wandered inside the earth towards the east and has been rising there ever since’ (Reuther 1981; VII: 29). There are two expanded accounts of the Ditji- mingka story given by J. G. Reuther (1981, VIII: 20 and X: 20). In the latter he describes how Ditji, the Sun, was one of the two wives of the Wild Onion Ancestor ‘Jelkabalubaluna’ (i.e. Yalka parlu- parlunha, ‘a wild Onion peeled’); Ditji crawled into the earth at Ditji-mingka. He makes other references to the site; the most telling of these is in XI: 163 where he describes ‘tthe enchantment of the sun’, ie. a form of magic for creating hot weather. This magic was carried out with the use of ‘ditjipupara’ (earth from Ditji-mingka). Reuther adds: ‘Everyone who is a devotee of the female muramura, Ditji, has some of it in his wurley’ (1981, XI: 163). There can thus be no question about the importance of the site. The Reverend Reuther — as is clear from Wangkangurru evidence — usually gives a ‘cleaned up’ version of mythological traditions. We cannot be certain whether he himself censored the myths, or whether Aboriginal people, who respected him, were reluctant to tell him things they knew would offend him. His accounts are dull and humourless: all the spice has been taken out. There was certainly more to the story of Ditji- mingka than he implies, and there were associated rituals. The police trooper Samuel Gason was the first to publish details of Diyari traditions. He was not a missionary, but after all, he lived in the Victorian era. He was clearly shocked by the Sun rituals and gives the following version: ‘Their traditions suppose that man and all other beings were created by the moon, at the bidding of the Mooramoora. (This term simply means “Ancestor”). Finding the emu pleasant to the sight, and judging it to be eatable (but unable, owing to its swiftness to catch it during the cold that then prevailed), the Mooramoora was appealed to to cast some heat on the earth so as to enable them to run down the desired bird. The Mooramoora, complying with their request, bade them perform certain ceremonies (yet observed, but too obscene to be described), and then created the sun’ (Gason 1879: 260). Other versions of the sun-legend, all involving Ditji-mingka, the cave of the sun, are given by Siebert (1910: 44-45) and by Howitt (1904: 427), Aiston gives yet another, different account of the Sun History, in which the Sun Ancestor is regarded as male. The cave however still figures prominently: ‘The moora, however, escaped, but so annoyed was he, that he sank and sank, until at last, at the place now called in memory of the event Ditchaminka, he plunged into the ground. To this day is shown the hole in the stony plain where the sun disappeared’ (Horn & Aiston 1924:131). Diyari rituals gradually fell into oblivion because of the disintegration of the group and because of pressure against ‘paganism’ from the missionaries. Nevertheless the myth and the knowledge of the Ditji-mingka site did not immediately fade from the minds of Diyari people. H. K. Fry (1937: 189-194) published a brief Diyari text, with English translation, of a myth called ‘Ditji-mingka Mura and Pinja, Sun Cave Mura Revenge’. Apart from the name in the title, this text does not actually mention that the female Sun Ancestor is involved. There can however be no doubt that it is an 150 1. A. HERCUS uncensored version of the same myth as in Reuther (1981, X: 20): the Old Woman in Fry’s story, like Reuther’s ‘Ditji’, is one of the two wives of the Wild Onion Man ‘Jelkapalupaluna’. This text was given to Fry and Vogelsang by an old Diyari man ‘Sam’ Dintibana Kinjmilina, who owned the myth at that lime, in the mid-1930s, Sam also told Fry that he had a ‘sun-bag’ filled with light blue earth from Ditji-mingka which he claimed could be used to ‘sing the sun and make it very hot’ (Fry 1937: 193). [t is of the nature of cult-heroes that they may be larger and more extreme than the ordinary things of life. The female Sun Ancestor and her older sister in Sam Dintibana’s story certainly were: they are depicted as grotesque and fearsome old women who got their husband ‘Jelkapalupaluna’ killed, cut him into small pieces and mutilated the body, and then mutilated themselves. The Diyari text only gives us an outline, but we can begin to imagine why the full story does not appear in Reuther’s work and why Gason was shocked. The important matter is that despite the missionary disapproval of earlier years, the myth was still alive in the 1930s in Sam Dintibana’s mind. Even after Sam’s death the knowledge that Ditji-mingka was a special place, a ritual centre, lingered on and the older people with Diyari associations had a deep affection for it. This became clear to me over the years that I was travelling on the Birdsville track for language work at both Marree and Birdsville. Ditji-mingka was often mentioned by Mick McLean Jrinjili, by Jimmy Russell Wanga-mirri (‘Many mornings’), and also by Ben Murray Palku- nguyu (‘One mass of clouds’). Whenever we headed north on the Birdsville track we would pass Blaze’s Well, a well which is now totally silted up. It was in a small depression which they called Thidna- kurduni [‘Making a (deep) Footprint’) because it was there that Ditji, the Sun Ancestor, stepped into muddy ground while looking for wild onions. As we continued north towards Cannuwaukaninna Bore those older men would point nostalgically towards the west, saying; ‘Over there is Ditji- mingka’, Mick and Jimmy were Wangkangurru, Ben's mother was Arabana, but they had all been associated with Diyari people (Hercus 1980, 1986; Austin 1981), They had learnt from them to admire the cave of the sun and were anxious to see it again. They often described it: it was in one of the more inhospitable areas on Etadunna station, in rough country. [t was high up on the western slope of the ridge that faces the bed of the Ditji-mingka creek as it nears Lake Palankarinna. There was a soakage in the creek nearby and the old people used to camp there for their ceremonies (see Fig. 1). The walls and the ceiling of the cave were of glistening gypsum which reflected the last rays of the setting sun: it was the home of the sun. There were rock-carvings inside, circles which symbolised the sun; Jimmy Russell had seen them. The two PIGURE 1. The edge of the bed of the Ditji-mingka Creek, where the soakage used Lo be. (Photo B. Jeffery.) LOOKING FOR DITJI-MINGKA 151 cave-openings were slightly upward; this meant that occasionally animals, particularly feral goats, fell in and died, not being able to get out again. It was a haven for snakes and this irritated the owners of nearby Etadunna station. Mick McLean and Jimmy Russell spoke of the cave so often that I wrote letters to have it protected; Ditji-mingka was truly a site of significance. Moreover it was only a few miles from Lake Palankarinna which was a declared geological reserve because of important fossil finds (Rich, van Tets & Knight 1985: 46 ff). We were determined to see Ditji-mingka and tried a number of times in the early 1970s. Mick McLean was then well over eighty years old, he could not scramble up over the ridge, and on our own we failed to find it despite his instructions. Next time in August 1974, Jimmy Russell came to help, he was only in his seventies and his sense of direction was uncanny, like Mick’s. He was deeply ashamed, because he too could not find it. Later of course it became clear why! Finally in June 1976 we had quite a group of people, which included Jimmy Russell and Ben Murray, as well as members of the South Australian Government’s Aboriginal Heritage Unit, along with linguists Tamsin Donaldson and Peter Austin who was then writing his grammar of Diyari (Austin 1981). We again went looking for Ditji-mingka (see Fig. 2). Later in camp Jimmy Russell spoke in Wangkangurru about what happened:! Text 1. thika-rna arniri wadna-ya-rna Ttatinha yadla-ku Come-IMP we run -SP-IMP Etadunna close-DAT kari-rnda uta arniri yani-ngura mudlu-nga-thu arniri see-PRES now we say-CONT sandhill-LOC-EMPH we marrili tharni-thika-Ihuku Ditji-mingka-ruku mingka this side eat-return-PURP Ditji-mingka-ALL cave nhanhi-lhiku. see -PURP. FIGURE 2. Ben Murray (left) and Jimmy Russell (right) setting out to look for Ditji-mingka. (Photo B. Jeffery.) 152 2. wantali Separately yurrakati-nga west side-LOC intja Ditji-mingka? Where Ditji-mingka? L. A. HERCUS thika-lhuku arniri come-HIST we yuka-lhuku. go -HIST. 3. arniri yuka-ka partjarna wadlhu yurrakati-nga We go -PAST all place west side -LOC thika-lhuku arniri, wadlhu pidla Thita-pulumanha. return-HIST we, place name Thita-pulumanha, 4. uta arniri partjarna karla-nga pathara midla-nga, Now we all creek-LOC box tree nose-LOC, thidna-ra -ki yuka-rna mingka-thu wapa-rna thika-rna foot-CAUS-EMPH go-IMP cave-EMPH seek-IMP come-IMP thadna-rna, wadni-rnda watungunta nguthi thika-lhuku. leave-IMP, follow-PRES rest reverse come-HIST. 5. nhararda waru katha-liparna, ipali katha-rna, Here long ago walk-ANC, before walk-IMP, malka-thu mingka nhanhi, Ditji-mingka. not-EMP cave see, Ditji-mingka. kutha irtjirtja thuntiripa-rna kutha kathiwiRi-ri Water soakage cover over-IMP water big -ERG tjiRi-ri. flood-ERG, padni-li punga nhanhi-ra, thanpi-liparna thiRi-ri Not-ADV humpy see-PUNC, knock down-ANC flood-ERG wanpa-rna. carry -IMP. 6. kayi kadnha awukinta kadnha arniri wadna-yi-rnda. Here rock this-ALL tock we run-TR-PRES. yadla witji-yangu, ayi! tjarlpa tharka-tharka-rnda close become-PLUP, hey! tree stand-stand-PRES nhararda yada... here close.. 7. thanpi-thanpi-la-rda Destroy-destroy-ALT-PRES Ditji-mingka! Ditji-mingka! LOOKING FOR DITJI-MINGKA & L, What was it like before? J. ngurka arla, parluru, antha iparli katha-nangka-rda Good true, smooth. I before travel-CONT S-PRES marna parkulu nhanhi-ka neurku-nhaku! kanhangarda mouth two see-PAST good-EMPH! There mingka tharka-tharka-rnda mingka-rda. cave stand-stand-PRES gape-PRES. 9. antha nguyu katha-liparna nhantu-ra, marna-nga I alone travel-ANC horse-CAUS, mouth-LOC tharka-rnda. mintjt-mintji-rnda, muyu katinari yantakara stand-PRESS. Shine-shine-PRES, sun beyond west muyu mintjiyva-ra. sun shine-TR-PUNC. 10. malka antha wintaku-ra, antha nguyu-nguyu. Not I go in-PRES, I alone-alone. kudju-ru pirda-lira nguyu-nguyu mingka-nga. kurdaitcha-ERG kill-LEST alone-alone cave-LOC, 11. antha thadla-ra waya-rna, thadla nhinka-rna-li. 1 fear-CAUS want-IMP, frightened squint-IMP-EMPH. mintji-mintji yalkiri-ri. muy round one. shine-shine kopi-INST. Sun. 12. thanpi-thanpi-rda, partjarna thanpi-la-rda Knock down-PRES, all desiroy-ALT-PRES parluru-ku level-DAT. Translation 1. We came back (from looking at the lower Cooper) and drove in close to Etadunna. We had a look around and then started saying ‘we'll have our lunch this side of the sandhill and then go to Ditji-mingka to have a look at the cave.” 2. (We split up) and walked separately coming and going over the west side of the sandhill. Where is Ditji- mingka? 3. We went all over the country on the west side and back again, that area is called Thita-pulumanha. 4. Then we all walked around in the creek bed, where the box-trees run out to a point. We were going about on foot, looking for the cave, we went and turned back again, we left, we followed the others but they came back as if they had got to a dead end. 5. ] used to travel around here long ago, but I can’t see that cave, Ditji-mingka, The soakage had been washed out by big rains, by flood-waters. [ couldn't see the humpies (that had been there), they must have been knocked down and carried away by the floods a long time ago. 6. ‘Ah, here is that rock!!’ (I said). So we all ran straight over towards the rock. We got near: ‘Ah, that tree is standing over there, we’re really close!” 7, They have blown up Ditji-mingka! 8. L. What was it like before’? J. It was lovely, all smooth. When I was travelling about a long time ago I saw the two openings of the cave. It was beautiful. I stood there for a while just looking at the cave. 9. |] was travellng about alone on my horse, and I stood in the mouth of the cave. It was gleaming and glistening, the sun was shining in from the west. 10. I didn’t go in, | was on my own, (I thought) a kurdaitcha [revenge killer] might come and kill me if I was all alone in that cave. 154 L, A, HERCUS 11. | was terrified. I peered in, very frightened. It was glistening with kopi. There was (engravings of) the sun there, round ones. 12. It has been destroyed, completely blown apart (with dynamite), razed to the ground! (see Fig, 3). CONCLUSION In 1879 Gason, as was typical of the period, had a very restricted view of Aboriginal religion, when, as quoted above he states with obvious lack of enthusiasm: ‘Their traditions suppose that man and all other beings were created by the moon, at the bidding of the Mooramoora’ (Gason op, cil.), We tend to take an equally simplistic view nowadays when we identify sites on a one to one basis with traditions: we are often told for instance in popular reporting that the destruction of a particular tree might result in the Aboriginal myth associated with it being forgotten. There is more to traditional mythology than that. This has often been said, and it was expressed with special clarity in 1965 by W.E.H. Stanner in his well-known article ‘Religion, Totemism and Symbolism’, where he shows the subtlety and intricacy of Aboriginal religious thought. Speaking of the totemic symbol- function he states that it has: ‘four elements (i) living men (totemists) serving as interpreters of (ii) signs (totems and totem-places), by using (iii) vehicles that form and express affective conceptions of (iy) sign- objects, which are the significance of the Dream Time marvels’ (Stanner 1965: 228). If we look at these elements with regard to the Diyari Sun Myth we can see that the first to be impaired was (iii), in that the ritual, the dances, the songs and the detail of the stories fell into oblivion, Therefore (iv), the deeper symbolism, the marvel of the Dream Time Sun Myth has disappeared for ever, All we are left with is the bare outline of the story. Until recently we still had (ii) the totem-place. The situation was no worse than for much of the mythology of eastern Australia, where we usually just have minimal stories and the totem place. Although we lacked the vehicle of interpretation provided by the chants and dances, we might still have had a glimpse of one aspect of the marvels of the Dream Time through the engraving and through the appearance of the cave, but this has now gone through the destruction caused by European activity. This leaves us only with (i), the living men. For many years, since the death of Sam’ Dintibana, there has not been anyone who ‘belonged’ to the Sun Myth and identified with it. There are only two people living to whom the mythology of Ditji-mingka and the surrounding areas still means something: they are Jimmy Russell who has suffered a major stroke and Ben Murray who is now in his nineties. Therefore it is sadly true that soon there will be no one who can stop on the Birdsville Track at the right spot south of Etadunna, look across to the west, and visualise that not far away is Lake Palankarinna- an expanse of white fading away into the distance into an endless plain (see Fig. 4). No one will remember the humour of the story of the wicked Old Man Markanjangkurla who chased the Seven Sisters across the lake and across the plain beyond and recall what he did there, FIGURE 3. The heap of rubble that used to be Ditjimingka. (Photo B. Jeffery.) LOOKING FOR DITJI-MINGKA 155 FIGURE 4. Looking towards Lake Palankarinna from near Ditji-mingka. (Photo B. Jeffery.) (Lake Palankarinna, which Europeans know mainly as a fossil reserve, is really parla-ng-kari-nha, a crude name, since parla means ‘semen’ and kari means ‘to chase’). Very soon there will be no one that can even visualise the cave south of Lake Palankarinna and point and say: ‘Over there was Ditji-mingka’. We have lost the Sun Myth with all its symbolism. ENDNOTES 1. The text transcribed in this paper was recorded in June 1976 as Hercus field-tape 739, a copy of which has been deposited with the Australian Institute of Aboriginal Studies, Canberra. For ease of reference the text has been split into numbered sections. The divisions are on the whole in accordance with intervals in speech, In this paper a practical orthography has been used for Wangkangurru: Plosive consonants other than the retroflex plosive have been written as unvoiced (k, p, th, ¢), but prestopped consonants have been written with voiced plosives as this corresponds most closely to the pronunciation, hence bm, dn, dnh, dnij, dl, dth. Retroflexes have been written as r + consonant, i.e. rl is retroflex / rn is retroflex n rd is retroflex f Interdentals have been written as consonant + A, hence nh, th, lh, Palatals have been written as consonant +/, hence ¢/, nj, //. ng has been used for velar 7. The three r-sounds have been transcribed as follows: r = the alveolar flap rr = the trilled r R = retroflex r. The following abbreviations have been used for linguistic terms in the interlinear gloss: ABL Ablative case ACC accusative case ACT active stem-forming suffix ADV adverbial suffix ALL allative case ALT altruistic stem-forming suffix ANC ancient past CAUS causative case CONT continuous participle CONT $ continuous stem-forming suffix DIST aspect showing distance EMPH emphatic clitic ERG ergative case EXCL exclusive pronoun HAB habitual aspect HIST historical past IMP imperfective LOC locative case NAR narrative past PAST past tense PERF perfect PLUP pluperfect POS possessive suffix PRES present tense PUNC punctiliar present PURP purposive SP speed form, indicating action undertaken before departing TR transitory aspect 156 L, A. HERCUS REFERENCES AUSTIN, P. 1981. ‘A Grammar of Diyari, South Australia.’ Cambridge University Press, Cambridge. FRY, H. K. 1937 Dieri legends. Folklore 48: 187-206; 269-287. GASON, S. 1879. The Manners and Customs of the Dieyerie Tribe of Australian Aborigines, Jn J. D. Woods (Ed.). ‘The Native Tribes of South Australia’. Pp, 258-306. E. S. Wigg, Adelaide. HERCUS, L. A. 1980. How we danced the Mudlunga. Aboriginal History 4: 5-31. HERCUS, L. A. 1985, Leaving the Simpson Desert. Aboriginal History 9: 22-43. HORNE, G. A. & AISTON, G. 1924, Savage Life in Central Australia’. Macmillan, London, HOWITT, A. W. 1904. ‘The Native Tribes of Southeast Australia’. Macmillan, London. HOWITT, A. W. & SIEBERT, O. 1904. Legends of the Dieri and kindred tribes of Central Australia. J. R. Anth. Inst. 34: 100-129. REUTHER, J. G, 1981. The Diari’, translated by P. Scherer. Microfiche edition, Australian Institute of Aboriginal Studies, Canberra. RICH, P. V., VAN TETS, G. F. & KNIGHT, F. 1985. ‘Kadimakara’, Pioneer Design Studio, Melbourne. SIEBERT, O. 1910. Sagen and Sitten der Dieri und Nachbarstaemme in Zentral Australien. Globus 97: 44-50; 53-9. STANNER, W. E. H. 1965. Religion, totemism and symbolism. Jn R. M. & C. H. Berndt (Eds). ‘Aboriginal Man in Australia’. Pp. 207-237. Angus & Robertson, Sydney. THE SCALED-SQUID LEPIDOTEUTHIS GRIMALDII JOUBIN FROM SOUTH AUSTRALIAN WATERS BY W. ZEIDLER Summary The first substantiated record of Lepidoteuthis grimaldii from Australian waters (i.e. within the 200 nautical mile fishing zone) was by Lu & Phillips (1985) but they gave no details of specimens. The purpose of this paper is to provide more details of these specimens, in particular of one in the South Australian Museum (SAM) which was not seen by Lu & Phillips, and thus highlight this interesting record from Australian waters. THE SCALED-SQUID LEPIDOTEUTHIS GRIMALDIT JOURIN FROM SOUTHERN AUSTRALIAN WATERS The first substantiated record of Lepidereuris arimalaii from Australlaiy waters (1G within the 20) nautical mile fishing zone) was by Lu & Phillips (1985) bur they gave no details of specimens, The purpose of this paper is to provide more details of these specimens, in particular of one in the South Australian Museum (SAM) Which was not seen by Lu & Phillips, and thus highlight this interesting record from Australian waters, Lepidoteuthis grimaldil was first deseribed by Joubin (1895) from two mantles froin a sperm whale’s stomach and from a fragment of @ Risso’s dolphin, both caught off the Azores, However, up until 1960 there had been only four records of this species and only two included the head (Clarke 1960) and a complete description was not available until 1962 (Clarke & Maul 1962), Since then the species has been recorded from most of the world's oceans, North and South Atlantic (Clarke 1966), Indian Ocean (Clarke 1980), Pacifie Ocean between the New Hebrides and New Caledonia (Rancurel 1970), Pacific Ocean olf Japan (Okutam ef al. 1976), Tasman Sea (Clarke & Macleod 1982) and Southern QOvean (Lu & Phillips 1985)- Complete specimens are still rare and apart fram juveniles caught in nets (Clarke 1964 & 1980, Lu & Clarke 1975, Roper & Young 1975) and the reoord of Lu & Phillips (1985) all specimens have been obtained {rom the stomachs of predators, ovainly the sperm Whale, Plivsefer catoden, but also fran Rissa’s dolphin, Granyus griseus (Joubin 1895); the taneet fish, Alepisaurus ferox (Rancurel 1970}; the black-scabbard lish, 4phanopus curbo (Clarke & Maul 1962) and the tuna, Gera ahesus (Clarke & Maul 1962), The first published evidence that 1, grrmvaldu might oecur in Australian waters was provided by Clarke (1980) who recorded the buccal mass of two specimens from the stomachs of sperm whales caught by whaling, Ships Operating out of Albany, Wesiern Australia, Clarke & MacLeod (1982) also recorded the remains of specimens from the stomachs of sperm whales killed ii the Tasman Sea between 3305, 172°R and 40S, 155°R but this is at least 550 km south-east of Cane Huwe, eastern Australia, The Australian specimens referred to by Lu & Phillips (1985) and the one iy SAM are noteworthy in (hat they are the first records of adulls from other than predators’ stomachs und indicate that the species oceurs relatively close in- shore along the south coast well within teach of commervial crawlers. Derails ol speciinens are as follows: 1. Male, 122 mm dorsal nant leneth, 97 km cast olf Broken Bay, New South Wales (33° 28'S, [52° 43’ E), depth 0-1000 m, Engel mid-water trawl, FRY ‘Kapala’, J. Paxton, 14 December 1977 (Australian Museum, Sydney AM, C111782). 2. Sex undetermined (viscera missing), 753 mm mantle Jength, approx. 30 km south-west off Beachport, South Australia, trawled in 550 m by ‘Margaret Phillipa’, 6-10 September 1982 (Museum of Victoria MY, F53159), 3. Sex undetermined {viscera decayed), 790 mm dorsal mantle length, approx, 40 km south-west off Beachport, South Australia, trawled in 220 m, obtained fresh from fish processor in Portland, Victoria by W. Zeidler, 22 October 198t (SAM, DI7589), Specimen I, a juvenile, is in relatively good condition and only the tips of the arms are missing. Generally it agrees with the description of young stages given by Clarke (1964) and some body measurements are given if Table |. TABLE |. Lemdoteuthis erimaldii body measurements, Character Measurement (mm) Specimen | Specimen 3 Manele length (darsal) 122 ‘790 Mantle length (Ventral) 115 740) Mantle width (max) 26 210 Fie length 60 410 Fin widih tmax) sy 240 Gladius length - 790 Gladius width (max) - 3 Rachis length - ang Rachis width (max) - 28 Max. width of scales 14 Ww Specimen 2 measurement® Specimen 3 (Pig. 1) when collected was in good condition with only the tips al the arms missing. However, it was inadvertently left out of the freezer and deteriorated considerably before being measured and preserved, The head js too damaged for accurate measirement, olher bady measurements (Table 1) are according ro Roper & Voss (1983) and beak dimensions (Table 2) are according to Wolff (1984), Some measurements arc inaccurate due ta the damaged nature of the specinien a. the fins are contracted, dorsal mantle length probably is longer as Ihe up oF the tail fs damaped and abou! 70 mm is missing and the ntantle is probably not as wide when all internal is too damaged for accurate 158 W. ZEIDLER FIGURE 1. Ventral view of Lepidoreuthis grimaldi, SAM, D. 17589. organs are intact. The beaks (Fig. 2a-c), radula (Fig. 2d) and gladius have been adequately described for this species by Clarke & Maul (1962) and the SAM specimen does not differ from that description. TABLE 2, Lepidoteuthis erimaldii (SAM, D17589) beak dimensions, Character Measurement (mm) Upper Hood length 39.5 Rostral length 17.6 Wing width 9.5 Rostral tip to inner margin of wing 26.0 Wing to crest length 43.2 Crest length 59.6 Jaw angle width 10.0 Lower Rostral tip to inner posterior corner of lateral wall 41,2 Rostral length 17.3 Rostral tip to inner margin of wing 35.2 Wing length 18.3 Jaw angle width 93 ACKNOWLEDGMENTS | wish to thank Tan Loch, The Australian Museum, for the loan of Specimen 1, Dr C, C, Lu, Museum of Victoria, for information on the MV specimen and for his constructive criticism of the manuscript and John Glover, SAM, for providing common names of fish predators. The photographic expertise of Roman Ruchle (Fig. 1) and Jan Forrest (Figs 2 and 3), both of SAM, is also gratefully acknowledged. REFERENCES CLARKE, M. R. 1960. Lepidoteuthis grimaldii — a squid with scales. Nature, Land. 188: 955-956. CLARKE, M. R. 1964. Young stages of Lepidoteuthis grimaldii (Cephalopoda, Decapoda). Proc. Malac. Soc. Lond. 36: 69-78. CLARKE, M. R. 1966, A review of the systematics and ecology of oceanic squids. Jn FE. S, Russell (Ed,), ‘Advances in Marine Biology’, Vol. 4. Academic Press, London and New York. Pp. 91-300. CLARKE, M. R. 1980, Cephalopods in the diet of sperm whales of the southern hemisphere and their bearing on sperm whale biology, Discovery Rep, 37: 1-324, CLARKE, M, R, & MACLEOD, N. 1982. Cephalopod remains from the stomachs of sperm whales caught in the Tasman Sea. Mem. Nain. Mus. Viet. 43: 25-42. CLARKE, M. R. & MAUL, G, E. 1962. A description of the ‘scaled’ squid Lepidoteuthis grimaldii Joubin 1895, Proc. Zool. Soe. Lond. 139: 97-118. JOUBIN, L, 1895, Céphalopodes recucillis dans l’estomac d’un Cachelot capture aux iles Acores. CR. Acad. Sci. Paris 121: 1172. THE SCALED-SQUID LEPIDOTEUTHIS GRIMALDI JOUBIN 159 LU, C. C. & CLARKE, M. R. 1975. Vertical distribution of cephalopods at 11°N, 20°W in the North Atlantic. J. mar. biol. Ass. U.K. 55: 369-389. LU, C. C. & PHILLIPS, J. U. 1985. An annotated checklist of the Cephalopoda from Australian waters. Occ. Pap. Mus. Vict, 2: 21-36. OKUTANI, T., SATAKI, Y., OHSUMI, S. & KAWAKAMI, T. 1976. Squids eaten by sperm whales caught off Joban District, Japan, during January-February. Bull. Tokai, reg. Fish. Res. Lab. 87: 67-113. RANCUREL, P., 1970. Les contenus stomacaux d’Alepisaurus ferox dans le sud-ouest Pacifique (Cephalopodes). Cah. ORSTOM sér Oceanogr. 8: 4-87. ROPER, C. F. E. & YOUNG, R. E. 1975. Vertical distribution of pelagic cephalopods. Smithson. Contrib. Zool. No. 209: 1-51. ROPER, C. F. E. & VOSS, G. L. 1983. Guidelines for taxonomic descriptions of cephalopod species. Mem. Natn. Mus. Vict. 44: 49-63. WOLEFE, G. A. 1984. Identification and estimation of size from the beaks of 18 species of cephalopods from the Pacific Ocean. NOAA Tech. Rep. NMFS 17: 1-50. W. ZEIDLER, South Australian Museum, North Terrace, Adelaide, South Australia 5000. Rec. S. Aust. Mus. 21(2): 157-159. FIGURE 2. Lepidoteuthis grimaldii (SAM, D17589); a. Upper beak; b, c. Lower beak; d. Radula. REVIEW : FROM HORIZONTAL TO PERPENDICULAR : TWO RECENT BOOKS ON CENTRAL AUSTRALIAN ABORIGINAL PAINTING BY P. SUTTON Summary Albert Namatjira, the life and work of an Australian painter by Nadine Amadio, Anne Blackwell, Jonah Jones & Daniel Thomas. Macmillan, Melbourne, 1986. ix + 102 pp. illus. Cloth only. $A29.95. Dot and circle : a retrospective of the Aboriginal acrylic paintings of Central Australia edited by Janet Maughan & Jenny Zimmer. Royal Melbourne Institute of Technology Communications Services Unit, Melbourne, 1986. 207 pp. illus. Paper only. $A20.00 REVIEW FROM WORIZONTAL TO PERPENDICULAR: TWO RECENT BOOKS ON CENTRAL AUSTRALIAN ABORIGINAL PAINTING Albert Namatjira, the Life and Work of an Australian Painter by Nadine Amadio, Anne Blackwell, Jonah Jones & Damel Thomas, Macmillan, Melbourne, 1986, ix + 102 pp. Illus. Cloth only, $429.98. Dot and Circle: a Retrospective of (he Aboriginal Acrylic Paintings of Central Australia edited by Janet Maughan & Jenny Zimmer. Royal Melbourne Institute of Technology Communications Services Unit, Melbourne, (986. 207 pp- illus. Paper only. $A 20,00, “The interaction between the European and Aboriginal artistic tradulons can produce a renaissance potentially as significant (or Australian life as that which was laifiched Upon Europe by the spread of the “new knowledge” from Constan- tinople in the sixteenth century’. With these dramatic words Dr H.C. Coombs launches his introduction to the most recent book on the wark of Albert Namaliira, compiled and edited by Nadine Amadio, arts editor of Sydney's Sunduy Telegraph. De Coombs is certainly right ta point out that Aboriginal graphic.and sculptural imagery conslilules a distinctive body of sources for the arts in Australia generally, allhough their role thug far might more aptly be likened to that of African and Oceanie works in the rise of twentieth century primitivism in Western Eupopean art, than to that of the manuscripts of Constantinople cirea 1453. The Renaissance was, among olher things, a scholarly revolution. By contrast, Picasso was-able to remark, in effect: “Everything | need to know about Africa is in those objects’ (Rubin 1984: 74). However, Albert Namatjira, on the face of it, is a reverse case: an Aboriginal person who painted in the Buropean-Australian watercolour landscape style. The handsome new book of around 30 colour plates and nearly 40 pages of text which is part of the subject of this article begins with a ‘celebration’ of the painter’s life by Nadine Amadio, Here we are fhelplully told of the important retrospective exhi- bition of Namatjira's work held at the Araluen Arts Centre in Alice Springs in 1984, and curated by Mona Byrnes. This signalled a revival of interest ing painter who had become an Australian house- hold name by the 1950s bur whose reputation had never been very secure among curarors and scholars of art and whose reputation declined afier his death in Iragie circumstances in 1959. Amadio's chapter carries the obligatory post-l960s enlightenment regarding the falsity of applying the wrdinary sense of the term ‘primitive’ to Aboriginal art, While bending a knee briefly to Namatyira’s belatedly recognised (and poorly documented) concern with cvuniry and sites of mythological significance, this chapler succeeds nevertheless in rescuing and promoting Namatiira as a haive artist (my term, not hers) in another sense: instead of High Chocolate Box we now have Grandma Moses, Amadio is especially concerned to frame Namatjira’s work within the traditional key symbols of western European art mythology. He was a ‘true painter’ (p. 5) with his ‘own vision! (p. 6), ho ‘Knew the joy of a full creative life, He must have possessed an extraordinary drive and passion’ (p. 2). His ‘drive’ or ‘creative drive’ is referred to repeatedly, Strangely posed next to this supposed ‘drive'is Namatjira as Nature’s Gentleman, a man whose ‘dignity’ and ‘natural dignity’ (p. 10) are to be remarked upon, and whose pictures are noted for their pellucid, airy calm and balanced repose, They are also pictures of beautiful (and largely unpeopled) landscapes, excluding even a hint of the nasty side of tile available in and areund the Alice Springs of his times Jonah Jones’ short picce on the 1984 exhibition, where a remarkable 56 Namatjiras were assembled, is of particular historical value, althoagh one could have done without che rmystificatian of the comparlson with Piero della Francesea on page 18 (‘It is all there’, What Is? And what is it?), Daniel Thomas's chapter re-evaluales Namatjira’s relationship ro the art. world. tt is the best chapter by far and say$ same new and substantial things abou( Namatjira’s wark, He points out some snajor reasons Why Namatjira’s work was anly slowly recognised by art professionals; it came our of rhe tourist industry; it Was exhibited in unprestigious locales for much of his hte; yice-regal patronage inade art scholars suspicious, watercolours were strongly associated (as they still are) with amateurisin; and Namatjira painted in a style that was already considered conservative and backneyed by ‘professional, vocal, modernist’ arrists. Yet he painted with the best of those vharavteristically Aboriginal artistic qualities of “extrenie delicacy, refinement, and gentleness’ (p. 26), Finally, Thomas makes Lhe important observation that Namatjira’s ant. which was seen in his own lime a$ “Buropean’, Io2 is now ‘re-Ayboriginalised’ because of three maitt factors: iis reference to signifeant country; is symbolic repetitiveness; and 18 fineness of touch, ‘Our altered understanding pertnits us now (o admire it more fully, and lo be moved by i (p26), The other exeellent chapter in the book ts that by Anne Blackwell, @ young archaeologist and historian who sadly was killed ina war accident in Central Australia in 1986. While Thomas’ chapter excels in ideas, Blackwell's excels in historical fact. The history of Hermanusburg, che roles of Carl Strehlow and FE W. Albrecht there, the creation of the Aboriginal uraft industry, the biography of Namatjira, and the emergence of 'a Hermannsburg, watercolouris! school’, are presented with scholarly atrention to detail and handsomely dlustrated. Iv is here that we find, amazingly, the only remark on Namatjira’s work by an Aboriginal person: ‘That old man — he painted’ This is reminiscent of Amadio’s comment about the Papunya acrylics painters on p. 33: ‘Every painting they point to, they say simply, “this is my country” .. .', Typically, s lengthy explanatory discourse or even a brief gushing eulogy or eritival sideswipe is an unlikely Central Australian Aboriginal response to a question about a painting. This does not mean that exlensive records cannot be made of what people know, think and teel about a picture, Nh does mean Thac much patient and laborious Investigation has 16 be invested in the process by the outside enquirer, Forrning the last section of the book, the colour pistes are perhaps the best guide to the aesthetic upparatus of lovers of Namatiira’s work, because Iliey are accompanied by capuons which use phrases such as; Tichly-toned —, . warmth and sensuality .., simple beauty of form —., an earthy power’ {p. 46); delicacy and economy’ (p. 48): a gentle and lyrical painting, teslrained in volour' fp, 89), Throughout evalirations like these both here and earlier in the bawk the recurring thenves ate: 1. Innocence, naivete, gentleness, restraint 2. Fineness, delicacy, tonal subtlety, economy, accuracy 3. Lightness, enerey, vitality, drama, vibrancy, urgency 4_ Optimism, celebration, fanctfulness, warmth, sensual respose. 5. Vasiness, distance, openhess of Space 6 Repetinion, 1 think Chis lends some weight to my suggestion that Namatjira has emerged as 2 naive but technically proficient and respectable painter, in the assessment of fnany of his admirers. [in the past, il was his Falsely suppased lack of sophisheation Which played such a commanding rate in his lion- isttion and demotie fame among suburban Ans- Traliatis. His lotellectual sophistication is now seen 10 lie in the relanouship between his pictures and the geographical sites and geographical religion they represent, rather than in some happy fusion of his High Culture with his manner of representation, Had he done the same works in the 1980s, though, this may not Have been the case al all, Namatyira's is the gentle innocence and ‘warural dignity’ of someone ‘close to the earth’ and Unpolluted (or just about ta be polluted) by the industrial civic culture. This is not the gentleness (gentility, geoteelness, gentrification, gentleman- liness} of restrained passion so characteristic of ‘civilisation’, where some sign of the wildness that has been domesticated must remain i art is to avoid being too wishy-washy. This is the clarity of Eden. Ard | suggest, also, that this analysis might to a lesser degree apply to the generally positive response by art lovers to the Aboriginal acrylics which have so rapidly displaced Hermannsburg watercolourism from centre stage in the Centre, Technically variable but usually Far from technically incompetent, the acrylics are — especially when documented — genuifely appealing depictions of the concerns of a tradition which seems [ree of the tired baggage of self-comment. While Albert Namatjina painted (lor sale, at least) the typically horizontal view of landscape of the Enropean art tradition, the acrylics painters presenc us with images of place viewed [ram a perpen- dicular, aerial photographer's perspective. Papunya paintings, Papunya Tula Ari, Prntup! paintings, Western Desert paintings, Aboriginal acrylics, the dot paintings — the tera) has yet to settle down, perhaps reflecting two persistent uneertaindes. Que of Ufese wicertainiies resis on the taet chat most of us who talk about the works haye at best a partial grasp of their social and cultural location, or indeed that of any art or artefactual style in Aboriginal Australia, Amather ineertainry arises from the fact that, fifteen years after the birth of che Aboriginal acrylics ‘mave- went’, i is sill manly Known through exhihirion epheniera, TV programmes, bric! mayurine arpctes, and spall sections of survey works such as the excellent introductions to Aboriginal art by Berndt, Berndt & Stanton (1982) and tsaacs 41984). In other wards, there is no mayor, susiaimed, scholarly study of these paintings available, and which is on a scale comparable to Nancy Munn's Halbiri fcunography (1973) or Howard Morphy's Unpublished thesis on northeast Arohem Land bark paintings (1977), Such a study would form an authoritative basis for characterising the works. An historically ortented study of this kind ix now urgently needed, (ur the acrylics are not only a ‘srvle’, uw ‘genre’. & ‘Thovemen!” (spreading phena- oienon, at east) and an “indusiry’ but alse a rapidly developing ‘phase’ in a very old graphie tradition — in feet, the very one Munn was writing about iwerily Years ago The catalogue of book Bar arid Circle, edited by Janel Maughan & Jenny Zimmer, lies Somewhere between ihe handy but hghiweight ephemera, which usually Throws up unintegrated fragments of face (and fiction), and the fullecale scholarly study Which turns an array of facts into a body of knowledge and interpretation, In one-of its facets, Dot anc Circle isa listina and reproduction of the 102 pictures shown in the exhibition of [he same name lent by the Flinders Universtiy Acl Mrseum to the Royal Melbourne Insotute af Teclunology Gallery in April-May 1985_ They later returned to be shown at Flinders University in Adelaide, The catalogue is valuable nol only because it lists the works and their atten considerable documentation, but also because it reports the exhibviion itself in the form of photographs of the installation and 1s opening, lists of case exhibit¢ and exhibition photographs, and 4 reproduction of the exhibition text labels, Inventories of the firse year's consignments of aervlics from Papunya te the Stuarr Art Centre 1971-2, provided here by Pat Hogan, and the appendices detailing works painted by David Corby Kjapaltjarrt and Tiirkey Tolsen Tyupurrita while Aboriginal Artists in Residence at Flinders in 1979, may alse be mere lists but they will be of much greaicr historieal interest as Gme goes by. In its ather, more discursive role the book does provide a collection of brief essays on the ongins and sigoificance of the acrylics movement by a variety of authors, There is tile imtelleciual coherence berween the essayists here, however. Geoff Rardon’s account of the sudden adoption of aeryhe painting at Papunya durjig his time there it 1971, here reprinted from his 1979 work (Bardon 1979}, is uw reminiscence of importance to Australian art history and also a work of greac honesty, even if at times it shows an unnervingly slight grasp of the culuire of Aboriginal people. Dick Kimber's much mare savvy recollections of the same period follow those of Bardon and very usefully cite the names of the key European supporters of the ‘movement’ (As Kimber says, rather paradoxieally, iris ‘only a movement in Buropean eyes accustomed to crowds ralher than individuals’ (p. 19).) lis clear that the aerylics ure, and always have been, very much a dveet prodact of particular Aboriginal-European interactions, nol stmply ‘Aboriginal products’ sought and boughl by Europeans. Another reminiscence in this colleetion, Radney Morice’s ‘The Kdnekayunt experience’, is the exceptional essay in the lilerary sense if as beuurifally written, it is alive and conveys the quality of daily experence in a remote Aboriginal camp, wd ib rings true, ay Ure work of someone Who was Wis there long enough and had sufficient sensibility to get it nght, As a means of orienting the reader to an unfamiliar World it is excellent, even though its reiation to the business at hand ls rather gerieral (and like the Bardon piece itis a reprint (in Uns case, from Over/and |978)), The heavy dominance of the overview in the presentation of the Aboriginal arts is, 1] hope, soon coming to an end, Andrew Crovker’s essay, ‘Potential and pitfalls', promotes |he view Lhat Aboriginal works should be allowed to take ‘their place in the contemporary arnstic Jorum’, aiid supports the view that the evolution of Aboriginal art “enables those Aboriginal artists with an exceptional artistic flair to (loursh'(p. 47}, opposing the restrictive attitude of ‘traditional is better’. The possibility that the Papunya paintings emerged from an assertion of traditionalism by older men sits uneasily here — it does not invalidate Crocker'’s view, bul it does cry out for consideration, The sections of the book which will perhaps. be ol greatest iaterest fo scholars of art history and material culture are those by Janet Maughan and Vinvent Megaw. (Ry ‘material culture’ here | meat io include aesthetic anthropology, as well as the less theoretically-oriented branches of that field.) Megaw, an archacologist by training, has published a number of papers on the Aboriginal acrylics phenomenon. This one, ‘Dreamtime diseipline or alien adulteration?’, stresses a theme similar to that of Crocker’s piece which precedes it} Chal the “fine art” status achieved by Aborginal works is legitimate, and that commercial and insti- tutional recognition (or should it be vonstitition”?} of thal fact is inevitable, People who complain of capitalist penetration of Aboriginal society via art are idealising, romanticising and alrempring to fossilise its culture, Megaw suggests that change in Aboriginal art is not adulteration of a pristine Table culture ty a greedy, foreign one bul 4 sign of Aboriginal ealtural vitality tn the modern world, Like his other papers on similar themes, Megaw's essay in this case questions the rationality and the moraliry of distinctions such as fine art/tourist art, art artefact, art /crall and so on, meanwhile raising the problem of applying aesthetic judgements across cultures. Several pasitions on these subjects are adumbrated, none of them is particularly rigorously pursued, and few of them are plumped for ottier (han gingerly, The profession of arehac ology is Mentioned jnan act of dissociation of the author from the philosophy of art or art criticism, Jenny Zimmer's preface (oddly, one of two, rhe other being by Maughan & Megaw) provides much usetul information on jhe creation of the exhibition itself, As a performance it must go down as one of the most archestrally complex in Abonginal art history, She alse offers some revealing thoughts an 164 why such an exhibition was considered useful as part of the education of some Melbourne art students. There is not room here to argue all the points she raises, although most of them are arguable. Her most interesting point is summarised in her last paragraph: ‘Cultural Convergence can only serve to make our culture richer and more authentic. This exhibition and booklet are dedicated to the concept of Cultural Convergence’ (p. 13, bold print original). Perhaps this is an implicit stand against the recent avalanche of Left criticism in this field, which resounds with phrases like ‘cultural genocide’, ‘astheticisation’ and ‘neo-colonialist encapsulation’. It may also be a recognition of the fact that a new Australian sub-culture has been emerging for fifteen years or so among people of no Aboriginal ancestry, many of them resident in Central Australia. They are creating a partly new way of life, one affected by a perception of Aboriginal culture and by interaction with Aboriginal people, yet distinct. This theme is reflected in the final section of the exhibition and catalogue, which is called variously ‘White artists using Aboriginal designs’ (p. 18), ‘The Western Desert image and the European art context’ (p. 25), ‘Across the cultural divide’ (p. 191) and ‘Across the cross-cultural divide’ (p. 205). Tim Johnson is a non-Aboriginal artist who has used graphic devices drawn from the Aboriginal acrylics and who has painted jointly with an Aboriginal artist (see cat. nos. 97, 98). Whether this is best described as ‘convergence’ or ‘incorporation’ (etc.) has been a matter of debate. Whether it has yet resulted in ‘good works of fine art’ has also been a matter of debate. The latest wave of European artists influenced by Aboriginal graphs can probably claim a moral, if not an aesthetic, advance over predecessors like Preston, whose contact with Aboriginal people was generally much more marginal and whose borrowing was done from photographs and museum collections rather than ‘with the people’. Not only artists but collectors and curators interested in the Aboriginal acrylics are beginning to talk of them as the first truly Australian art, and as a case of cultural convergence between Aborigines and European Australians. This perception, problematic as it may be, is gathering force. I have left Janet Maughan’s contribution to the end, both because it is the longest and because it is comparatively rich, She outlines the history of the genre and the referential meaning system of its images, and attempts a style analysis which, as she herself points out, fails to get off the ground (p. 17). I do not agree that such an analysis is not possible, however. The effect of variety perceivable among the pictures is great, certainly, but it is one which rests on the recombination of a limited set of motifs and of ways of placing them in a symmetrical field. The list of motifs or recurrent symbols in Figure 2.(p. 16) seems to me seriously incomplete, and also conflates sets of symbols which ought to be kept distinct. No structural typology of the images is offered, although the works are eminently suitable for this kind of first-sort. It is difficult to see how a style analysis could procede without some kind of sorting into the various kinds of axial symmetry, for example. Maughan then goes on to discuss the major subgroups of works in the exhibition and the rationale for their subgrouping, beginning at the beginning (the early paintings), and moving on to look at two artists specially represented in the exhibition, Tim Leura Tjapaltjarri and Toby Brown Tjampitjinpa. This concentration on two indivi- duals was important to establishing an idea of the style and output of particular artists over time. Institutional roles in the acceptance and promotion of Aboriginal acrylics are highlighted here. The particular efforts of Visual Arts staff and students at Flinders University are given their due, though for better balance there could have been more emphasis, for example, on the historic role of Robert Edwards and the Aboriginal Arts Board in promoting both the production and the sale or public collecting of these works in the 1970s and 1980s. ‘Children’s paintings’ (for, not by) come next, followed by ‘Women’s paintings’ (by, not for), where it is made clear that women are under-recognised as painters in the genre and tend to produce for the ‘craft’ market. The question of how much this may be a reflection of gender roles in matters of religious authority is not opened up. Maughan ends by relating Papunya paintings to conceptual art, but, in the absence of a discussion of what that is, many readers may find the connec- tion obscure. It is a pity there was not room enough in her essay for a substantial discussion of the relation of the acrylic designs to those largely very similar ones of the same region which have been recorded since the earliest European times, but in other media: the rock intaglios, rock paintings, body decorations, shield designs, and the sand drawings of daily conversation and play. Among Maughan’s more interesting observations are the view that the complexity of figure-ground relations in the acrylics has increased over time, and her important statement that there is a gaping lack of published information on Aboriginal criteria for merit in paintings. Great credit is due also for her long and detailed work on the catalogue annota- tions. The production quality of the book leaves much to be desired, but most of my reservations there would have been pre-empted by more generous publication funding of typesetting and layout, printing and binding. The page size (A4) is unattractive; it needs to be more squat. The 13 colour plates are fine but the monochrome plates are rather muddy. In one case a signature has been chopped in half (p. 104) and in another the alignment marks have intruded (p. 111). Lines wobble. Perhaps more serious are editorial lapses such as uncaptioned plates (pp. 1, 4, 8, 12, 41, 48, 93), typographic errors, and the entirely wrong information under the plate on p. 20 which says: ‘Western Desert: Demonstration of circular designs which have in part replaced the more complex traditional ground paintings. Photo: Penny Tweedie.’ This is actually a photograph by C.P. 165 Mountford taken at Haasts Bluff in 1942. This mistake is perhaps partly compensated for by the humour value of the caption to the upper photo on p. 46: ‘Old Mick Tjakamarra with Daphne Williams recording the story of the painting, 1982. Photo: Vincent Megaw’. Ms Williams, apparently short of paper, is on close inspection writing down the details in her Bank of New South Wales cheque-book. . . ACKNOWLEDGMENTS I am grateful to Christopher Anderson, Philip Jones and Andrew Pekarik for their helpful comments on an earlier draft of this paper. REFERENCES BARDON, G. 1979. ‘Aboriginal Art of the Western Desert’. Rigby, Adelaide. BERNDT, R.M., BERNDT, C.H. & STANTON, J.E. 1982. ‘Aboriginal Australian Art: A Visual Perspective’. Methuen Australia, Sydney. ISAACS, J. 1984. ‘Arts of the Dreaming: Australia’s Living Heritage’. Landsdowne, Sydney. MORPHY, H. 1977. ‘ “Too many meanings”: an Analysis of the Artistic System of the Yolngu of Northeast Arnhem Land’. Ph.D. thesis, Australian National University, Canberra. MUNN, N.D. 1973. ‘Walbiri Iconography’. Cornell University Press, Ithaca. RUBIN, W. 1984. ‘Modernist Primitivism: An Introduction’. Jn W. Rubin (Ed.). ‘ “Primitivism” in 20th Century Art: Affinity of the Tribal and the Modern’. Museum of Modern Art, New York. P. SUTTON, Head, Division of Anthropology, South Australian Museum, North Terrace, Adelaide 5000. Rec. S. Aust. Mus. 21(2): 160-164. RECORDS NY THE SOUTH AUSTRALIAN MUSEUM VOLUME 21 PART 2 NOVEMBER 1987 ISSN 0081-2676 CONTENTS: ARTICLES 69 D.S. TRIGGER 85 119 139 149 157 161 Inland, coast and islands: traditional Aboriginal society and material culture in a region of the southern Gulf of Carpentaria C. PATTERSON & P. V. RICH The fossil history of the emus, Dromaius (Aves: Dromaiinae) S. J. EDMONDS Echiurans from Australia (Echiura) J. RILEY & D. M. SPRATT Further observations on pentastomids (Arthropoda) parasitic in Australian reptiles and mammals L. HERCUS Looking for Ditji-mingka NOTES W. ZEIDLER The scaled-squid, Lepidoteuthis grimaldii Joubin, from southern Australian waters P. SUTTON From horizontal to perpendicular: two recent books on central Australian Aboriginal painting