VOL. 122, PARTS 1 & 2 29 MAY, 1998 Transactions of the Royal Society of South Australia Incorporated Contents. Prideaux, G. J. & Wells, R. T. Sthenurus baileyi sp. nov., a new fossil ea from the Pleistocene of southern Australia - - - Sheldon, F. & Puckridge, J. T. Macroinvertebrate assemblages of Goyder Lagoon, Diamantina River, South Australia - - - Anstis, M., Alford, R. A. & Gillespie, G. R. Breeding biology of Litoria booroolongensis (Moore, 1961) and Litoria lesueuri (Duméril & Bibron, 1841) (Anura: Hylidae) and comments on population declines of L. booroolongensis - - - - Kolesik, P. A new genus and two new species of gall midge (Diptera: Cecidomyiidae) damaging young branches of Eucalyptus spp. in South Australia - - - - - - - - = - Steen, Z. & Schwarz, M. P. Within-nest behavior in a eusocial Australian allodapine bee Exoneura SERENE tridentata Houston (Apidae: Xylocopinae) - = - Field, S. A., Keller, M. A. & Austin, A. D. Field ectibiey aaa BehaOnE of the egg parasitoid Trissolcus basalis (Wollaston) Hymenoptera: Scelionidae) - - - - - - -~ = = - Beveridge, I. Woodwardostrongylus petrogale sp. nov. (Nematoda: Cloacinidae), from the stomachs of rock wallabies fetrogale spp.) from Amhem Land - - - - - - - = Nicholas, W. L. | Mesorhabditis kinchegensis sp. nov. (Nematoda: Rhabditidae) from arid soil in Kinchega National Park - - - - = - Brief Communications: McDonald, K. R. First Queensland record of the burrowing frog Cyclorana cryptotis Tyler & Martin, 1977 (Anura: Hylidae) - -— - O’Callaghan, M. G., Ockleshaw, E. & Allen, J. The prevalence and distribution of nematodes in the large intestines of sheep in South Australia - - - - - - = = = = - PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 122, PART 1 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INC. CONTENTS, VOL. 122, 1998 PARTS | & 2, 29 MAY, 1998 Prideaux, G. J. & Wells, R. T. Sthenurus baileyi sp. nov., a new fossil Ranga from the Pleistocene of southern Australia- - - - - - Sheldon, F. & Puckridge, J. T. Macroinvertebrate assemblages of ‘Goyter Lagoon, Diamantina River, South Australia- - - Anstis, M., Alford, R. A. & Gillespie, G. R. Breeding biology of Litoria booroolongensis (Moore, 1961) and Litoria lesueuri (Duméril & Bibron, 1841) (Anura: Hylidae) and comments on population declines of L. booroolongensis- - - - - - = - Kolesik, P. A new genus and two new species of gall midge (Diptera: Cecidomyiidae) damaging young branches of sik bane spP. in South Australia - - - Steen, Z. & Schwarz, M. P. Within-nest bebavtelie in a eisontia Australian allodapine bee Exoneura chide tridentata Houston (Apidae: Xylocopinae)- - - Field, S. A., Keller, M. A. & Austin, A. D. Field enslony aod ohesditerr of the egg parasitoid Trissolcus basalis (Wollaston) Hymenoptera: Scsifonidas) - = -—§ =~ 5S & » = # = & = Beveridge, I. Woodwardostrongylus petrogale sp. nov. (Nematoda: Cloacinidae), from the stomachs of rock wallabies (Petrogale spp.) from AmhemLand- - - - - Nicholas, W.L. Mesorhabditis kinchegensis sp. nov. (Nematoda: Rhabditidae) from arid soil in Kinchega National Park - - - - - - Brief Communications: McDonald, K. R. First Queensland record of the burrowing frog Cyclonaien cryptotis Tyler & Martin, 1977 (Anura: Hylidae) - - - O’Callaghan, M. G., Ockleshaw, E. & Allen, J. The prevalence and distribution of nematodes in the large intestines of sheep in South Australia - - 33 45 55 65 73 79 85 87 PARTS 3 & 4, 30 NOVEMBER, 1998 Martin, H. A. Late Cretaceous-Cainozoic palynology of the Poonarunna No. | well, central Australia - - - - - - - - Kolesik, P. Rhopalomyia lawrenciae, a new gall midge species (Diptera: Cecidomyiidae) deforming leaves of Lawrencia squamata (Malvaceae) in South Australia- - - - - - - - Kolesik, P. Dasineura wahlenbergiae, a new species of gall midge (Diptera: Cecidomyiidae) damaging shoot tips of Wiphregrreipni stricta (Campanulaceae) in South Australia- - - Davies, M. & Watson, G. F. Developmental biology of Uperoleia talon Tyler, Davies & Martin, 1981 (Anura:Myobatrachidae)- - - Davies, M. & McDonald, K. R. A new species of frog (Anura: Mictany lice) fess Cape Melville, Queensland- - - - - - - - Davies, M. & McDonald, K. R. Developmental biology of Uperoleia altissima Davies, Watson, McDonald, ames & Werren, 1993 (Anura: Myobatrachidae) - - + oe - Coleman, P.S. J. Changes in a Mangrove/Samphire Ciiiaiitey, North ae Creek, South Australia - - - - - * + + = - Smales, L. R. New species of Seurechina (Nematoda : selsiiaion ‘sin in dasyurid marsupials from Australia- - - - Ferguson, M. A. & Smales, L. R. Spiroxys chelodinae Berry, 1985 (Nematoda: Spiruroidea) and Camallanus chelonius Baker, 1983 (Nematoda: Camallanoidea) from freshwater turtles ie Chelidae) in Queensland, Australia- - - - - ee Insert to Transactions of the Royal Society of South Australia, Vol, 122, parts 3 & 4,30 November, 1998 89 185 STHENURUS BAILEYI SP. NOV., A NEW FOSSIL KANGAROO FROM THE PLEISTOCENE OF SOUTHERN AUSTRALIA By GAVIN J. PRIDEAUX* & RODERICK T. WELLS* Summary Prideaux, G. J. & Wells, R. T. (1998) Sthenurus baileyi sp. nov., a new fossil kangaroo from the Pleistocene of southern Australia. Trans. R. Soc. S. Aust. 122(1). 1-15, 29 May, 1998. Sthenurus baileyi sp. nov., is described from Pleistocene deposits of Eyre Peninsula and the southeast of South Australia. The dentary is similar in size and morphology to S. occidentalis Glauert, 1910 but the cranium is much less inflated across the frontals and the rostrum less tapered anteriorly. Sthenurus baileyi is characterised by very low crowned molars, most similar to S. cegsai Pledge, 1992, S. brachyselenis Prideaux & Wells, 1997 and S. antiquus Bartholomai, 1963. Upper and lower premolars are similar to S. antiquus and §. brownei Merrilees, 1967. Overall, S. baileyi appears most closely related to S. antiquus and may represent the most plestomorphic member of the lineage containing the more brachycephalic sthenurine species. Key Words: Sthenurus baileyi sp. nov., Sthenurus antiquus, Sthenurus, Simosthenurus, sthenurine Kangaroo, Victoria Fossil Cave, Naracoorte, Brothers Islands, Eyre Peninsula, Pleistocene. Transactions ef the Royal Suciery of 8S. Aust, (1998), TI22( 1). 1-15, | STHENURUS BAILEY! SP. NOV., A NEW FOSSIL KANGAROO FROM THE PLEISTOCENE OF SOUTHERN AUSTRALIA by Gavin J, PRIDEAUX® & RODERICK T., WELLS* Summary PRipb AUN, G1 & Wrirs, RT. (1998) Srheniirus beilevi Sp. noy.. a new fossil kungaroo from the Pleistoeene of southern Australia. 7revey. R. Soe. 8. Auge, 122 (1), 1-15, 20 May. 1998. Sthenuras baileye sp. nov, is described from Pleistocene deposits of Eyre Peninsuli and the southeast of South Austrilia. The dentary is Similar insize and morphology to.8, eectenralis Glauert, 1910 but the cranium is much loss Inflated across the frontals and the rostrum less tapered anteriorly. Siemurus Dadlevé is characterised by very low crowned molars. most similar to8, ceesa/ Pledge, 1992. 8. brachyselenis Prideuux & Wells. 1997 and 4. untiquus Burtholomai, 1963, Upper sind Jower premolars are similar to $8. aarigqus and S. browne’ Merrilees. L067. Overall, §. badlevi appears most closely related to S. curignii and may represent the most plestomorphic iiember of the lineage vontuming the more brachycephilic sthenurine species. Kiy Worbds! Siediirus baileyi sp. nov., Sthenuras antiqiis, Sthenarus, Simeasthearus. sthenurine kangaroo, Victoria Fossil Cave, Naracoorte, Brotiers Islands, Lyre Peninsula, Pleistocene. Introduction Following its discovery in 1969, the extensive Pleistocene deposit within Victoria Fossil Cave iat Naracoorte. South Australia has yielded cremains from around one hundred vertebrate species, Included are slightly less than half of the known Pleistocene species of sthenurine kangaroos (subfannly Stheaurinuek Proceprodan rapha Owen, 187d, Sthenuruy anderson’ Marcus. 1962.5. browne’ Merrilees, 1967, S. gilli Merrilees, 1965, 8. maddocki Wells & Murray, 1979, 8. cecidentalis Glauert, L910. 8. pales DeVis, 1895 (Wells ef eal. 1984). and a oew sthenurine, & bailey sp.nov: The Species is also known from a single specimen collcered from an eroded cave on one of the Brothers Islands in Coffin Bay, Ryre Peninsula (Brown 1908: Fig. 1). Williams (1980) identified the cranium and ussociuted dentaries us Sifenurus cf moddock), bul tt is here designated us the holotype of 8. heilevi sp. nov, Deseription of the new species and a consideration of tts phylogenetic implications form the subject of this paper. Materials and Methods The material is housed in the South Australian Museum, Adelude (prefix SAMA) and Flinders University (prefix PU). Dental homology follows Flower (1867) and Luckett (1993). Dental nomenclature follows Tedford & Woodburne (1987), Ride (1993) or is stundard. Mensuration follows “Schou! of Biological! Scnnces, Phuders University of South Australia GMO Box 210) Adeluide S.Aust SOOT, Boil: wav ti prices @flinders cutiiut mADUha . = yeaycouRTe Pig. 7. Map of southeastern Australia showing location of deposits yielding Sthenurys bailey? sp. nev. Tedford (1966) and Wells & Murray (1979). Dental measurements (mm) are provided im Rible |. Systematics Order Diprotodontia Qwen, 1866 Suborder Phialungerida Aplin & Archer, 1987 Superfamily Macropoduidea (Gray. 1821) Fuimly Macropodidie Gray. 1821 Subfamily Sthenurinve (Glauvert, 1926) Genus Sthenurus Owen, 1874 Subgenus /’Simosthenurus Tedford, 1966 Sthenurus (?Simosthenuruy) baileyi sp. noy, (FIGS 1-8) Holotype > SAMA P13670, partial cranium (wath Li 3, dP2, dP3, M1-4. excivated P3; Fig, 2A,B, 3A, 4A, 2 G. J, PRIDEAUX & R, T. WELLS TABLE 1. Cheek tooth dimensions of Sthenurus baileyi. 8. brachyselenis, S. brownei (eastern form), S. cegsat and S-antiquus: mein, standard deviation (parentheses), range (brackets). Abbreviations: L= length, AW = width of anterior loph(id); PW = width of posterior loph(id); AH = crown height of anterior loph(id) on buccal side; PH = crown height of posterior loph(id) on buccal side; n = sample size. Note that crown heights are heavily dependent on degree of enamel wear, hence, frequently high standard deviations. Tooth — Species L AW PW All PH " UPPER DENTITION dP2 S. baileyi 10.5 75 10,0 62 6.1 | S. baileyi TYPE As above S. browne 10.9 (0.43) 8.7 (0.40) 10,8 (0,38) 7.0 (O51) 7.9 (0.00) 15 (eastern form) [104-1 1.9] [8.1-9,5] {10.4-11.4] 16.0-8,0] [6.6-8.7] No veusat = = = rf - = S. antiqaus . - - - - - IPS S. batleyt 10.6 99 108 - - | S. baileyi TYPE As abave S. brawnei 11.3 (0.29) 10.7 (0.32) 11.0 (0,37) 5,7 (0.54) 5.9:(0,46) 15 jeastern form) [10,6-1 1,7] [10,.2-11.3] [10,6-1 1.8] [18-64] [5.3-6.8| S. cegvat 93 8.6 92 63 63 | S. caentiyguns : 7 7 - PS S. baileyi 17.2 (0.14) 9.9 (0.28) 12.9 (0.21) 9.8 (0.28) 9.7 (1.06) 2 [E71 -17,3] (9.7-L0.1] [12.7-13,0] [9.6-10,0] {8.9-10,4] S. baileyi TYPE {7.1 10.1 13.0 10.0 14 S, browned 17.1 40,57) 10.9 (0.68) 13.7 (O.81) OLR (0.76) 9,7 (0.78) 21 (eastern form) {16.2-18,0] [9.0-11,8] {12.1-15.0] [S.3-LL.1] [8.6-1 1.3] S. cezsai = - - . - S. antiques 14.4) 6.3 11.3 bt 11 | Ml S. hailevi 12.3 (0.14) 12.3 (0.21) 12.3 (0.2L) 6.3 (0.07) 6.6 (0.42) 2 {12,.2-12.4| [121-124 {12.1-12.4] [6.2-6.3] [6.3-6.9] S. baileyi: TYPE 12.2 12 Pe | 6,2 63 S. brawnei 12.9 (0.43) 12.4 (0.40) 12.3 (0,34) 6.1 (0,76) 6,5 (0.62) 28 (eastern form) [12.2-13.6] {116-130 [ELB-13.1] [5.0-7.9| [S.3-8.1] S. cegyeail - : - - - S. antiques 12.4 12.0 12.) - - | M2 S. haileyi 13.8 (0.21) 13.2 (0.21) 13.1 (0.14) 6.9 (0.42) 7.2 (0.35) 2 [13.6-13.9] [13.0-13,3 }13.0-13.2] [6.6-7.2| 6.9-7.4) S. baileyi ‘TYPE 13,6 13.3 13.0 V2 7A S. brownet 14,1 (0.37) 13.6 (0.46) 13.1 (0.43) 6.6 (0.76) 6.9 (0.58) 23 (eastern form) [13.2-14.7[ {12.9-14.4 {12.5-14,3] [5.3-7.8| 5.8-8.0] S. cegsai - - - - 5.6 ! S. cautiquits 14,9 (0.07) 13.4 (0.14) 13.0 (0.21) 7.20113) 74 (1.34) 2 [14.8-14.9] [133-135 {12.8-13.1] [6.4-8.0] 6.4-8.3] M3 S. baileyi 14.5 (0.00) 13.7 (0.21) 12.6 (0.35) TA (0.14) 6.8 (0.28) 2 {14.5} [13.5-13.8 [12.8-13,3] [6.9-7,1] 6.6-7.0) S. baileyi TYPE 14.5 13.5 12.8 7A 6.6 SL browne 14,5 (0,39) 14.9 (0,50) 12.9 (0.54) 6.7 (0.62) 7.0 (0.53) 19 (eastern form) 13,7-15.5| [13.3-14,8] [12.3-14.5 [5.48.1] 5.9-8.1] NS cegseet 13.1 (0.07) 115 10.9 (O14) 5.4 (0.00) 5.5 (0.14) z 13,0-13.1] [10.8- 11.0 [5.4] 5.4-5.6| S. antiquus 16.0 (0.35) 13.6 (0.49) 12.9 (0.85) $.3 (0.35) 4.3 (0,00) 2 15,7-16.2] [13,2-13.9] [12.3-13.5 [8.0-8,5} [8.5] M+ S. hailevi 13.8 13,4 1L.3 6.7 6.1 | S_hailevé TYPE As above So hrawnet 14.0 (O43) 12.2 (1.48) 11.2 (0.40) &.1 (0.62) 7.4 (0.78) 16 (eastern form) 13,.2-14.5] [11,3-13,0] [10.6-12.0 [7A-9.6} [6.5-8.8] S. ceesat 12.2 10 o5 55 a4 | S. cantiyuun TABLE 1. — Continued NEW PLEISTOCENE KANGAROO LOWER DENTITION up2 ml m2 ms m4 S. baileyi S. buileyi TYPE S. brachyselenis S. browne (eastern form) S. cegsai S. aintiquus S, baileyi S. baileyi TYPE S. brachyselenis S, brawnet {custern form) SL cewyat S. antiqnus S. bailevi S, butileyi TYPE S. brachivselenis S. browne’ (eastern form) S. cegvat NS. cuvetigules S. baileyi S. baileyi TYPE S. brachyselenis S. brownet {eastern fort) S. cegsat S. antiquus S, baileyi S. baileyi TYPE S. brachyselenis SL brownet (eastern form) 8. eersal SL cgentreptiues: S. baileyi So baileyi TYPE S. betchyselenis S, brawnel {eustern form) S. cevsal S. autiquus S. bailevi S. baile TYPE S. brachyselenis S, browned’ (eastern form) S.ceusal Si antiguas 26 uA 8.1 9.8 (0.45) [9.2-10.9] 9.8 (0.14) [9.7-9.9] 99 10.2 10.4 (0.36) [100-111] 16.2 (0.78) [15.3-17.8 15.3 13,8 16,2 (0.53) [15.2-17.0 14.8 17.6 12.0 (41) [11.5-12,5 12] 13.9 13.1 (00.64) |12.1-14,7] 10,5 13.8 13.3 (0.46) {12.8-13.9| 12.8 14.7 (0.53) {13.8-16.1] 12.7 15.3 (0.85) [14.7-15.9] 14.1 (0.64) [13.6-15.0] 13.6 14.9 (0.51) (13.7-15.7] 12.8 16.0 (0.99) 115.3-16.7] 13.9 (0.11) [13.7-14.0] 13.8 14.0 (0.43) [13,2-14.5] 11.9 15.4 (0.49) [15.0-15.7] a 6.5 (0.34) [5.9-7,2] 8.90.71) [8.4.9.4] Sab 7.8 9,2 (0,46) [8.4-10.0] 8.0. (0.32) |7.7-8.4] 7.9 6.3 8.5 (1.34) [R095] 6.5 8.2 10.1 W.22) 19.9-10.4] 10.0 909 10.4 (0.49) [9.4-12.0] 11.4 (0,27) [110-116] hs 11.4 (0.39) {10.4-12.0] 10.8 11.9 (0.71) [A124] 12.2 (0.37) [11.7-12.6] 12.6 12.0 (0.43) [112-129] Ho 124 (042) [}2.1-12.7] 12.1 (0.26) [11.8-12,3] 2.2 12.2 (0.48) | 11.3-13.0] 10,3 8.9 (0.99) [8.2-9.6] 8.2 6,1 91 (46) [8.6-9.7] 9.1 (0.28) [8.9-9,3| 8.9 8.2 O93 (031) [8.7-9.8] 9.7 (O44) 9.1-10,3] 95 8.0 10.3 (0.58) 9.4-11.5] 7.2 10.1 0.1 (0.26) 9.8-10.4| 10,2 10.3 10.6 (0.43) [96-115] 1.0 H.2(041) [10.7-11.7] 7 11,7 (0.33) [11.1-12.2 10.3 12,1 (0.78) [11.5-12.6] 11,9 (0.42) [114-124] 12.4 12,2 (0,35) [11.4-12.9| 10,2 12.2 (0.49) [1 1.8-12.5] 11.1 (0.42) [10.5-1 1.5] 11.2 11.2 (0,40) {10.6-12,0] 8.6 11.0 73 73 5.7 7.5 (082) 9.5 (1.04) 17.9-1 1,0} 89 73 9.9 (0.85) [8.9-11.4] 8.3 7.1 (1.01) [6.3-8.5| 64 9.3 8.6 (1.04) 16.4- 10.0} 5.5 83 (0.49) [7.8-8.9] 8.0 9.5 (0.94) [8.2108] 59 lhe 8.2 (4.93) [7.0-8.9] 78 9.3 (0,61) (8.4-10.3] TA) 10,5 74 (043) [7.1-8,0) 71 8.1 (0,621 \7.4-9.6) 6.2.71) (5.7-6.7| 5.7 5.5 7TALU7 [6.40-8.2] 6.0 70 7.2 (0.63) [6.0-8.2] 9. (1.03 [7.3-10,2] 91) 6.7 9.9 (0.83) [8.9-1 1.3] 71 4 7.0 (0.76) [6.3-8.0] 6S 94 8.7 (1.07) [6,5-10.2] 85 8.4 (0.63) [7,8-9.2] 8.0 9,5 (0.80) [8.0-11.2] 59 10.4 TITY [6.9-8.4| TS 9.1 (0.73) 17.5104] 64 10,3 7.2: (0.35) [6.9-7.8] 7A TA(OTS) |6.5-8.8] 45.3 7.7 (0.35) [7.4-7.9] te i 17 ‘ G.I. PRIDEAUX & RT. WELLS SA) left and ngnt dentanes (with il, dp2. dps. nil - 4. excivated pa: Fig. 3B, 4B, 5C,D), apparently: collected from a bone breccia in-an eroded cave on the western end of west Brothers. Island (34 35' 8, 135° 20' EE), Coltin Bay. Eyre Peninsala South Austulin (Brown 1908: Williams L980: Fie. 1). Other mammals frona the deposit include Macrapus rufagrisens, Potoraus plaivops, Pyeudocteirus sp. Rarus fuscipes and Neopheco cinerea. A huge bird femur previously atiibuted to Gereverinty preween (Rich 1979) belongs toy Drwnailis rovaehollandice (J, MeNatnara pers, con, 1996), Age of type locality ty considered Pleistocene because all taxa identified to species are only known from the Quuteruiry. Sumhirly, the genus Sihenieuy appeurs nol lo have existed anywhere beyond the lite Picistovene, Details of collection are uncertain bul probably retrieved by DLR. George around 1902 (1 MeNcuirrunt pers, eaimm. 1996), Divaeiveasts Cranium similar iy srze to Stenuray ecenlentalin bul fronts less expanded. rostrum shorner and brouder with wider misals and larger minal aperture, PA similar to So deevwied but wath relatively marrow shallow longitudinal basin and mwa accessory cuspales unieror jo promiment poslerchuceut uceessary cusp. Upper molurs very low crowned. With shart preemmuluen weak pustprotoerista and very Well developed postparaerista. Dentary sivrlur ine sivve and morphology to S. vecidesitalis and 4S, cniieitys — Barthologiai, (463 but with mare posteriorly inflated pleryeoid fossu thiunm in any Siieniias species, Posteroventral batder uf neassclerie Fossa expanded laterally imu wide shelf, similar to SL cess, So aif and S. widddowki, iL intermediate belweens, occidentalix anil S. browees in genera! shape and degree of prociumbeney. pF must similar in morphology to 8. entiguds but lower erowned, with straighter lingual crest. Lower molars very low crawned. with anteropesterionly short trivonid, well-developed premetactistid. aud very reduced onstidd obhgua and parucnstid producure i morphialugy closest lo Neves, § uniques and S, hrachvselenis Prideaus & Wells, 199) hut wader relative to leagth, Description af holawpe Vertical portion of prenaxila thived dorsally prowidibg -clonwate contac)! with uasals. Diusterma shorn, unterinr V/s comprising premasxilla and posterior i minxitla. Tneisive foramina long. narrow, untecior border level with posterior extreme of 13 alveolus (Pig. 3A). Rostrum short, lapered anteriorly (Fig. 2A). Buceinitor fossa on maxilla rather shallow ameriolly, deeper postetiorly gthterion to zygomatic wich. Massetenic process well-formed, railier narrow, eroded off ventrally on leit and nent sides. Nasals very broad posteriorly dnd, although broken antenorly, clearly short. Nasotrontal suture gently sinusoidal (Fig. 2A). Prontals moderately inflated anteriorly: supriorbital crests only slightly developed (Fig, 2A). Temporal crests: moderately developed, not tully conversent upon sagittal suture, Large infraorbitul foramen positioned anteroventral und tnestal to Lichrymial foramen, just below orbital centre. Palatal vacuities extend anteriorly to anterior extreme of MI (Pig, 4A). Right lateral extremly of broken postpalatine bar devel wilh M4 jnterloph yullev. 1] crown ruther long. moderutely wide. wath vertical occlnsal surface faving posteriorly, 12 very small splint-like, Vasize of LL, 13 high crowned, but qyuile short iunteropostertarly (Pig. 2B). Small anterolingual lobe evidenton 3. dP2 reminiscent of P3, rognded in general outline. especially lingually, bur much shorter relative be width (Fig, 4A). Buceul and lingual crests siraisht, excepl for buccal curvature of lingual crest a posterior extremity, Anterior bisio sinh quite deer und separated from longuddimal basin by tow transverse ridgelet. Posterior basin appears to juve bee relabvely Targe, approx, hall size ol longiludinal basin. JP3 completely molarifurnm und similar in general outline t ML but differs by tiiving Lophs orientated obliqgely (not perpendiculird ta buceal and Trial sides of tooth (Fig. TA). In addition, precigulum very sha@bt terminating before reaching lingual extreme of tooth. Premetaerista appears well- developed. Profoloph appeurs to have been vary curved. convex anteriarly. No enamel crentlarans present i interloph valleys yery low. barely delectible postprotocristy positioned just Wagual of dP3 midline (Piz. 44). Postmetacwnulecrisia curves dorsobuecally frome metuconule to mect verticul postinetlcrishi. Saud) accessory crest positioned mesial ta postnetacristia, slight poslink centrally positioned on pasterior meraloph tuce, PS rounded jn oulline and tipered antenorly (Mig SA). Longitudinal basin shallow, Buceal erest barely excerding lingoul crest m height. Anteriarly, lingwil erest begins. tO cin purillel fa buceal crest then posterior 4/4 of crest curves oul Tingually. Small umertor busin present and separated from longitudinal basin by drunsverse ridge descendiny: from anterior buceal cusp terminating adjacent to anterior Tinguad casp. Posterior basin short. well- formed and separated [rom longitudinal basin by low tringverse ridge originating From posterior lingual cusp and onentated obliquely (slightly unterobueaily) to meet low down on buccal crest, Main posterobuccal aevessury cusp well-formed, nul quite as high as posterior parcot buceal erest. Three NEW PLEISTOCENE KANGAROO 5 = aw / —) , 2a * mM mp Vig. 2. Sitenaras ballew! sp. oy, ccaniuin, A. Holotype 1 P1AG70) dorsal view. BL Holotype lateral view, Seale bars = 10min Abbrevs; d = ditistema, [= front, i= niaxilla. mp = tasscterte process. W= basalonls = qesolrontal sature, p= parietal pire Prem. s = supragrbital erest. “ G, J, PRIDEAUX & R.'T, WELLS Hig. 4. Schein baileyi sp. nov. cranium and dentaries. Stereopai of holotype dentary cectusal view. Scale bars A. Stereopair of holotype cranium (PI3670) palatal view. B = 10mm. Abbrevs: a = anterior rool oF uscending tinus. b = buccinlor fossa an Alisa © = tandibular condyle, i = incisive foramina, pb = posteroyenteal border Of masseteric fossa, pv = palatal vacuities, small poorly separated accessory cuspules positioned anterior to mui accessory cusp (Pig. SA). Upper molars very low crowned, with protoloph equal in width lo metaloph in MI-2, but wider in M3-4 (Fig. 4A). Precingulum short. bueeal extreme terminating at distinct cuspule. representing cither stylir cusp A or B. Slight crest (probable puracrista) connects cuspule posteriorly ta puracone. Two to four slight vertical crenulations centrally located on precinguliim, with most togual probably temant preprotocrista (forelink). Postprotocrista weak, low, ascending buccally across fice of protoloph into interloph valley, uniting with vertical crenulation directed posteriorly from mid-pomr on protoloph, Postparacrista strongly developed, forming buccal border Of interloph valley, meeting slight premetierista on anterior face of metiloph (Pig. 4A) Interloph valley with few very fine to no enamel erenulations, Postmetaconulecrisit sweeps ucross posterior luce of metuloph terminating just posterior toend of postmetacrista, Two to three small distinet crenulations enclosed by postmetuconulecristy an metaloph posterior Mee, Pentary moderately proportioned, except for tt NEW PLEISTOCENE KANGAROO Fig. 4. Sthenurus baileyi sp, noy, cheek tooth rows, A, Stereopair of holotype (P13670) left upper cheek tooth row occlusal view, B, Stereopair of holotype (P13670) right lower cheek looth row occlusal yiew. Scale bars = 5 mm. Abbreys: co = crisid obliqua, pe = preciagulum, pd = paracrisud, ppe = postparacrista, ppre = postprotocrista, f = trigonid, Fig 5. Sthenarus baileyi sp. noy. premolars, A, Stereopair of holotype (P13670) left P3 close-up occlusal view. B, Stereapair of paratype (FUOQTG67) left P3 ceclusal view. C. Stereopuir of holotype left p3 close-up occlusal view, D. Stereopair of holotype Jell p3 occlusal view. Seale hars = 5 mm. Abbrevs: ac = uccessory cuspules. be = buccal crest, lb = longitudinal basin. le = lingual crest, me = main crest, my = median valley, phue = posterobuccal accessory cusp. x fF 1 PRIDEAUN ART WHEELS posteriorly intlited pterygoid fossa and lateral Axpinston of posteroventral border of mussereric fossee into wide shelf. Ramus moderately deep for wilh, partivularly in region of symphysis, Sympbysis gently tapered anteriorly and posteniorhy only extended short way beneath genial pit. below anterior root of dps. Digastrie eminence present hut Hol patticularly prominenr Digastric sulcus eatending from below anterior extreme ob pterygoid fossa to helow m2 hypolophid. Diustenta short, with fedian dorsal groave deep. relaively wide. Very shullow buceindter sulcus arises near posterior extreme of diastema, dorsal to large anterior mental forsmen. Bueeinalor sulcus deepens slightly posteriorly, terminates below ml hypolophic Posteriar mentil foramen positioned below m2 hypolophid. hallway between dorsal and ventral borders of mans, Anlenor ront of ascending ramus begins adyucent to m3 hypolophid (Fig. 3B), extending posteriorly to form huceal border of postuveolur fussa, Preryyor fossa inflated postertarly, projechine well beyond border Of Mmasseteric fossa when viewed Jaterally Masseterie toss deep. due largely ty laterally expanded posteroyentral border (Pig. 38), Ventral border of nasseterte fossa ut sume honeatal level as posterior region Of buceinator sulets. Musseteric Jofamen moderately latee. vertical in opeptation, Inferior mandibular foramen rither small, At untenior exueme of pterveoid fossa. anteromedial to iitenior mundibular foramen, dorseventally wade mylohyoid grouve present. This appears to hive heen partially overhung by shagp adterodorsally- directed process al anteremedial border of pterygoul fossa, and similurly-shaped posteroventrally - direcied process positioned helow postenoe extreme ot postaiveotur fossa. Mandibular condyle moderately laree (hig. $B) Angular process well- developed. rising dorsally to acute point. il rather shor. slender upturned. with oeclusal surface aba horicantal level just above Base of check well crowns. dp2 an both sides ef holotype toa wart or fragmentary to interpret. Likewise, dpa very worn, allhough clearly molinform, possessing, low bur well-defined parm premeta-. prehypo- unit preentu-cristids. p3 considerably longer than any molur, with main (Lingual) erest cxtending trom posterofingual vormer ly failing oi toeth anteriorly (Pig. SC.D). ‘Three cUspules form anterior part of main crest. with each bearing pair of literal ridgelets, one descending buccally. one fingtally. Buceal ridgelets lernimpate at fiw shelf formed by three conlluent cuspules, loved immediately anterior to buccal crest, Buccal crest straight. shor. equal in length to and mirroring shape af pestetior part of main crest, Median valley rather narrow. modenitely deep, Toward ts posterity (A, median valley waversed by coarse ridgelet (Fig. SCD). Lower molars very low erowned, with protolophid und hypolophid ecclusal suttaces finear and close to parallel. Trigonid very short, with paracristid low and composed of two moieties. Degree of separation of anterior and posterior muiglies increases from m1 to m4. Posterior part of paracristid sweeps smoothly dnterolingually across protolephid face, terniinates on bueeal side of anterior part Tn more posterior Molars, aMenor conjponeat of paraeristid shifted more lingually bul posterior extreme remains within buceal V/s of anterior protolophid face. originaliny well below lophid apex. A few fine enamel crenulations umse low down on antenor fuce ol prololophid and descend (ito tigontd basi, Lingual side of Ingonid bordered by welledeyeloped premetacristid. which termigates at puraconid Previngulid small and positioned anterobuceal ws paracrisiid, extending lingually as very thin pebinsija at antefior extreme of moku. Cristid obliqust (prehypnenistid)y low, sumihirly developed and aligned ia similar position on hypolophid: as pursertsod on prutolephid. Preentoctisuid’ very low and barely detectable. Asie from these weakly developed crests, shallow interlophid valley bears no enumel crenulations, Pastener face ol bypotoplre with low. shallow inflation: Paranpes: Prom Victoria Fossil Cave, Naracoorte South Australia (37 00'S. 140° 48) Ba, PUOOOA Jeit dnd nghe adult dentures: PUOLG7, let) PA, M1 (PS om Fis, SB), FUOL6S, right pa. FUOI94, partivl right denlary; SAMA PI6S31/P 16558. lett and right autuls denturies (left dentary in Fig. GA): P28287_ right juvenile derntary: P28659. right M2. lett M4, PUOON4, PUOH67 and SAMA P28659 may belang to sane I)dividual based of proximity ty deposit degree of enamel wear und occlusal hb Specimens collected’ by Prideauy. Wells and others. Age of deposit $s medial to lite Pleistocene (Wells er ul. 1O84: Ayliffe er al 10 yrress). Features out preserved adequately nm holotype cue deseribed from paratype SAMA P2822. Up? -cquil in length to dp3, yery similar iy morphology to p3 bul wider relative fo length. As in po. tinee cuspules dominute untenior half of main crest. cuch with tunsverse ndgelet on bieeal sule Ridgelets likely to have terpiinated 16 Gity Cuspules like p3, but due to considerable weur Sustained have become conflucnt with biiceal crest, vonveying an iHpression of More clopgale crest. Completely molaritorm. dps bears prololophn! tapered more tlaward Jophicl apex than hypolophid, As with dp2. wear has rernoved several leutures However, casdd oblique uppears more stmongly developed than in niolars and curved directly frou hypoconid apex into interlophid valley. terminate NEW PLEISTOCENE KANGAROO 9 Es Fig. 6. Sthenurus baileyi sp. nov, left dentary, A. Paratype (P16558) lateral view. B. Paratype (P16558) mesial yiew. Scale bars = 70 mm. Abbrevs: amf = anterior mental foramen, bs = buccinator fossa, d = diastema, de = digastric eminence. ds = digastric sulcus, imf = inferior mandibular foramen, mf = masseteric fossa, pmf = posterior mental foramen, ptt = pterygoid fossa, s = symphysis, Ww fi J PRINBAUX & &.T WELLS cen(tally on posterior protolophid faee. Very weak precarocristid also present, curving from entocenicd jolo interlophid yalley. lenmmating Tingual to eristiel obliqua. Enamel crenulations, similar to those on molars, gppear to have been present on anterior lophid faves. Slight, rounded posteingulid on posterior face oF hypolophid appears confluent with slight postentourstiut. Etyvinedtoey Named in honour of Mr Edwin Ed” Bailey whose efforts aver Une last 25 years have contibuted so much to fhe success oF pulaeontological work in the Naracoorte Caves, Veridian Unfortunately, puly one cranium is khown af, bakeyi sp now and variation within the upper denution ean unly be assessed by comparison of PS ind MI-3, which are each represented by two specimens, P3 is very similar in the holotype and PUOI67, with the slight occlusal wear in PUOTG7 responsible for most of the superficial differences between the specimens. In the holotype. PA is slightly wider anteriorly, both across the whole tooth and tie longitudinal basin. The lingual surlace of the holotype P3 is slightly more convex and rounded than FUOL67, The three cuspules anterior tothe main posterobuccul weessory cusp ure More sepuruted in FUOIGT. Only ote slight difference is deteerable on comparison of MI-3 of PI3670, FUOQT67 (MI) and P28059 (M2-3). The postparacrista 1s larger in the holotype. While greater weur sustiined hy FU A7 und P2S86S9 could decount for these differences, consideration of the mater mm which teeth oeelude suggests that they are more likely la reflect morphological variarion. Conplete or purtiu] denuiries are known for five maiwviduals, with three charucters clearly variable Depth and exten of the digastric sulcus is the most varidble character, Although deep and extending frony (he anterior extreme of the pterygoid fossa uy below the m2 hypolephicd in PIS670 and PLOS3 1/PL6S358. the suleus i much shallower und only extends to below the m4 protalophid: in FLOOO4. In P28282. the divastric suleus is even shallower thus pepating the diagnostic unlity of this chamacter, The degree lo which the plerygond (oss is inflaied posteriorly alsa yaries belween specimens, Inflation is greatest in PLO53 WP TASS8, slightly less mo the bGlatype (P13670). PUOOO4 and P28282. and ledst in PUO204, However. ibis sulficient in the latter (Oo mink fl US @ distinctive feature of So hetilevi. Dentary deprh relative te width js greater in PLOS31/P1 6558 (depth to width ratio below (2-3 = 179) compared fo FUOQ04 (1.65) and P2822 (1.61). The ratio is lowest in the hololype (1.46). Intraspecifie variation in dentary depth rehitive ta width ms cormonty observed in sthenurine species known Irom even small sumple sizes, Variauion in pa size is common in all sthenurines, Mivluding S$. ballew. While most of the paratypes are very similar in size. PIGS30/PI6551 and the holotype are noticeably shorter and narrower Morphology ovaries only shehily between individaals, primarily in the form of the buccal erest and minor variation in width of the median valley- The anterior half of the buccal crest in P28282 os slightly higher than the postertor half and curves posterolingually, becoming conlluent with a transverse ridvelet which crosses the median yalley This buceul crest roorpholoagy is not observed in any of the other specimens, although a very similar wansverse Pidgelet traverses the median valley in PI3670, Apart from this. feature, only the relative jofladon of the anterior region of the p3- varies slightly. A p3 referruble ro $8. baileyi as also known from Lindsay Hall Cave, near Madura on the Nullarbor Plain, Western Austra bat this speemmel remuins in the private collection of L. atehern Perth, This specimen is inseparable in sive and morphology from the South Australian Specimens. There iy little varfation in both sive and morphology of the lower molars, although the premetucrisud. paracrnishd and cristid obhqua of the paratypes are slightly more weakly developed than the holotype and the anterior lophid fees bear more fine eoumel crenulaGons, In addition. the postemeuhd os more shelf-like im each of the paratypes than in the holotype. except i) FUO2Od where there is a larger inflation of the ventrobuccal region of the hypalaphid posterior face. Comparison with other terxe Cranium. Although P3 was unerupled ine the holotype of S. baileyi sp. nov. the presence ol M4 in seclusion indicates that P3 eruption was imminent, An examination of other species for which a wuod age series ts known, reveals that lithe change in morphology or size in OSL uspects OF the eruniunt and dentary occurs From (his ontogenetic stage toile stage where P3 is erupted. This means that direct comparisons with older represemtuitives of ather tavu ave lenable. Leis worth noting that the two saniples al §. browne! and S. cecidenntlis with whieh 8. Auileys Is compared come from Naracoorte and ure considered fo represent the castent tons of both species, Although very similar in overall mormbology. they can he distinguished from the Loporypic Western Australiuy samples by their larger overul sive and Shelly smaller dentition relative to Ji Ane. NEW PLEISTOCENE KANGAROO Vl ‘The cranium of 5. barfleviis very similiar in size and brachycephaly to 8. eceidenralis, The premaxillac are also similar in relative size and morphology. Although rostral length of the two species is similar, the bucginator fossa on the side of the maxilla is deeper in 8. eceidentalix. This is coupled with a mesially concave aspect to the edge of the diastema, in contrast to the less distinet edge and shallow buceinator fossa in §. hadley’, This condition is more reminiscent of S. gilli and 8. andervant. The rostrum ofS. baileyi docs not taper to the same degree anteriorly as 8. vecidenialis, both becuuse (he frontals are less expanded and its qarial aperture is proporGonaly kuger Among the Siientiris species for which the splinehnocrunitun is known, lateral inflation Of the frontal region (particularly anteriorly) nd formation of supraorbital crests is greatest in 8, madidocki, S. occidentalis, S. stirling? Wells & Tedford, 1995 and §. Arawret The frontal region is relatively narrow in §. gilli, S. détdersant and S, Hindale? Tedford, 1966, The proportions displayed in S, baileyi are intermediate between these two groups, particularly bemween §. browne? and S. erlli. However. the nusals of S. beflev are very wide and constitute a greater proportion of the dorsal aspect of the fosteum than any other Sthemris species. except S. maddocki. Overall, the short and broad nature of the rostrum ts characterisue of S. balleve. The anterior extent of the palatal vacuities in 9 haley is akin te a number at other species, feyminatings close wo the GPS metaloph, or whit would be close to the posterior extreme of the P3 if it were in Geclusion, The masseteyic process appears iu have been well-developed, allowing for the damave in the holotype, and is intermediate between S. mudedocks and §. browner in size. Upper Dentition. In S\ baileyi sp. noy., the crown of V1 is slightly Jonger and broader than §, browne? wad is most stimulir to §. occidentalis, Weis not us high crowned us that of S. gi// and not as broad as i 8. anderson, 8. atlas (Owen, 1838). S. trrdalei or S. pales. The small, cvlindrieal 12 js iitermediate 1 size bewween 4, browne? and 8. vecidentaliy, 13 ts Most SHNIRW H size all general morphology tos. frrovene? but the buceul surfave iysmoolh and That, not hearing any vertically-orientated undulations. [n this respeel, S. beilevé is similar to &. eccidernteliy ands, will. though slightly shorter aad less inflated huerally thin 104, browne, dP2 OF S. batlev? sp. ouy. is closest in overull morphology to that species. Onentalion of fhe buccal and lieu! crests is alse similir but the posterior basin appears to have been larger nS. Aeiewin PA of S. hatlevi is amtust reminiscent af S. hawnel and S. eartiqutis in morpholowy, partivilarly iy the shape and aricntiwon of the buccal and Jingual crests and the anterior basin (Fig. 7). However, 8 bailey? possesses a shallower and narrower longitudinal basin and a prominent posterobaceal accessory cusp with ‘two cuspules amlertor to it (Pig. 7). The posterior basin is smaller than in either S. aariquus or S$. browne. Height of the linghal crest in §, aitiquay is considerably lower relative io [he buccal crest than iy either S. browne! or §. bailey’. In addition, the §. ctiquus P3 is also smuller relative to the size of the molars. The amount of wear that dP3 has undergone has obliterated several characters useful for compurison. However, the tooth appears to have heen generally similar to that oF S. browner but with a smaller precingulum, larger premetucrista and many fewer and finer enamel erenulations on the loph faces and interloph valley: The very low crowned nature of (he S. heilevi upper molars ts only upproached among Sthemuerta, hy S. ceased, So antiguas and § prddocki. Similar to S. cegsad and §. aatiqats, there are few erenulations on the Joph faces and the interloph yalley but the postprotocrista is more Weakly developed in S. hailey, The postparactista 1s) more strongly i ry j es 4 f . = - . —_ ea é 5 = —_ - i : a] } 1. , a it 4 - — HT I a am, Pig, 7, Comparntive sketches oh len PS inpechisal view. A Tivcdronemas pucktider Bo Sikes juirtipins (posterolinuiiil corer reconsiticted — ineaniplete 1 awtal speenmen) CN. hoilevt sp. naw DOS Ayo Sele burs= 3 Wii, Abbrevs! d= unteriar t= lineal 12 G. J. PRIDEAUX & R. 1. WELLS developed in S$. baileyi than any other species. including 8. hrawnei, The slight nature of the precingulum is similar to that of S. antiguas. Dentary, Th general morphology, the dentary of 8. baileyi sp. nov. is most similar to 8. occidentalis, 8. anriguus and 8. gilli, and to the former two in size. The ramus dilfers from §. eveidenralis in the following features: it is slightly narrower lor its depth, the symphysis extends only just beneath the genial pit. the il and symphysis ure slightly more procumbenot, the diastema is slightly longer, the divastric sulcus ts shallower and less extensive. the posteroventral border of the massetcri¢ fossa is more fared laterally and the pterygoid fossa is more inflated posteriorly. Sthenuruy baileyi differs froin §, antiguas uw its longer cheek tooth row relative to ramus depth. Morphology of the §. baileyi symphysis Most resembles that of S$. maddevki, where the symphysis tapers gently anteriorly and only extends slightly below the genial pit, However, unlike 4. meaddocki, the orientation of i} closely approximates that of the anteroventral border of the symphysis and in this respeer is similar to S, vceidentelis and S. browne’. Morphology of the iL crown and its degree of procumbency are intermediate between 3, vecidentalis aus. browne. Relative to the length of the ramus, the diustema of 3S. baileyi is proportionally longer than that of S. vecidentalls, 5. browned and S, ail/i, ets mast similar in length tos. maddacki bul is nol convex. dorsally as in this species. Depth and extent of the digastric suleus are similir to, but slightly more pronounced than in 8. meddock?, The degree of intraspecifie variation in depth and extent of the digastric sulcus also seen similar between (he lwo species. Literal expansion of the pusteravential border of the masseturic Tossa into wide Shell as similar tod. ceeseh So aly and §, maddocki, The pterygoid (ossa is more inflated posteriorly than aay other Siremumy species anu att his respeer, Y feadeve resembles Proceptodon, Size dnd morphology Gh dp2 most resembles that ot S. fie ied DUT Ss net us narrow unlerionky rohitive fo the posterior part of the tooth, The median valley moatso narrower Superficially, the dp2 tceal ens uppears Similar te fengthe no thalalS. Avo tal this Hipresvint es created because The wou sustaiedd babs resiicd Mm the erest becoming eontuent wilh the Sn cuspbles teins aiferiog dp3 is alse seine om size tnd ierplolaey fo that ots, Arewited bir the Erstit) ODT ia Ts Con Giet Yell Oe biypoconiied upes ine there cin Teweronamed crenulatioms on the tophid fees. In these ehanniiers he 8. bain ps more Closely resembles dat at \y aeeviteraeaten In morphology, stae relative to the olor. ame onenwiiion oF the tain und buccal crests. the 8 hetles!d oF is Similan te Hat lS. geting Be TY Fig, Ko Comparative sketches of right pa in oeclusal view. A, Sthenurus brachyselenis, BOS. uanguus, CS. baileyi sp. noy. D. 8) browne/, Scale bars = 5 im. Abhreva: a = anterior, |= lingual, differs by being smaller, lower crowred, slightly nove inflated anteriorly and having a slaighter main crest orientated from the posterolingual to the anterobuiccal dormer of the tooth. bn S. adit, the posterior part or (he main erest wends. anterohuceally (hen straightens anteriorly along the tooth’s midline, The S. baflevé p32 shures with S. brachyselenix the major features OF the tum erest aod a tidgeled traversing the median valley bub is easily Jistinguished by abs Larger sive, shiglitly grearer widrh relative to length (ratio Q.55 conipaned with 0,52) ane longer. straighter buecal crest (Pig. &). da size wud general outline, the S. betlevi ps is also sinitiar to. browne! hut is lower erowned aid hears o cobsidenibly shorter buecul crest. li sive and crown height. the lower migtars ahs. baflev ewe somewhabs imino those ofS. sees aad S cviguis, Sut ave Most stint ta these at § Ivacliyveleinis They differ from the later ru ther wrealey WIL polative to length (roti O54 compara wilh (72) ah the aulerioy! cinguhin oer hens symmetrically tapered auleriorly wou the shell-ike inflioem ci jhe posterior fare of the hyputophued heli much fess pronounced, In general db. duilew 3. hrachvseloma, 8S. ceases and 3. iets huve relutively smooth lophidh Mees. with only a lew line cnamel enenulations NEW PLEISTOCENE KANGAROO A Diseussion Sthenuras baileyi sp. neve retains a sure of cramoadental characters that suggest a fiirly plesiomorphic position within the genus, Although the deposits trom whieh the species ts known are Pleistocene in ape S. Aeileve is most closely comparable with the Phioeene S$, airiqgilix from Chinchilla in southeastern Queensland and 4, bravhyselenisx, a species of uncertain age from Wellington Caves, eastern New South Wales. p3 is very similar fo So @atiyius, but considerably nvore denved than S$. hrachyselenis, given its greater robustness relative to tbe molars ang longer. straighter buccal erest. In &, brachyvelenis, pd is quite arrow and has a short, erescentic buceal crest resincted to its posterobyecal corner, features. which are considered plesiomorphic for the genus (Podeaux & Welly 1897). The lower molirs are Intenmediaie herween S. hrachyseleniy ands. auuiquus i gener) morphology, bat. unfortunately, no appee moles lor Wie Former are known, However, the upper molars af S. baileyi are very sumilur to those of S. aeriguis. Based on a comparison of single fipper Molurs. these two species would be diffieule to separate. However, P3 ts notably more derived in 3. hiulevi, the lingual cizulum having become raised into a crest subequal in height with the buccal crest. lu 3. cartigaus itis markedly lower. Although the only known cranium is incomplete, S. baileyi can be clearly distinguished from all species Of Sphemens for which the cranium is Known. While exhibiting a similar degree ot brachycephaly (o 8. occidentaliy, §. bailey? possesses a shorter. broader rostrum and a less inflated frontal reoron (hun any of the other brachyeephalic Pleistocene species, Ineredsed inflation of the Irontils appenrs to have co-evolycd with jneredsed check tooth coniplexity in the fineaye (ar passibly linvages) lowding to the nore briehyveephalie (shorter-faced) Species. eg. $. brane &. eecidentatis wl 8, nidddbekh The modest degree at fronrel inflation. relatively stmple Jow crowed inolars and short bucwal crest not joming (he main vhest aiteriorly on pd provide a conceryable auWeCdent merpholowy to phese other specres, Uuformunaely. only one ramus and one munilha fraginent of So civigers see Known bul given: Lhe Jentil somihiritics between this species and A boiler, Nhe Vikelthood amay be thut these reflect overall cranial srmihidies. Althoweh the dentary ol Soaiiques ds incomplete. ole inypartant differends ja the craniuon of this speeres and §, barlert taay be nuiened by the Jonger cheek tooth row relitiye to denny depth observed: in aetiquas (nd 242 compared with 18S lor S. fades). Ths sugpesrsy a relatively longer dewlany und: Uerelon oa foie clongate cranium than fea S. baeeyy. This feature, tn conjunction with the shghily higher crowned molars, und more distinel ctisiid obliqua and parucristid. may make S. antiquis a possible structural precursor to the Jineaye that led to the more dolichovephaliv (longer-faced) Pleistocene species. This contention is supported by the fact that the ingualscrest of the §. antiguus P3 is notably lower than the bueciul crest. 4 featute shared hy the more dolichucephalic species. In (he more brachycephalic species the erests. tend to be subequal in height. Since musing oF the lingual cinguluny into a erest is a synapomorphy for all sthenurines excluding the plesiamorphic late Miocene Hadnmomas puckridel Woodburne, 1967 (Fig, 9), a lower crest muy be Tegarded is a more plesiomorphic condition Despite the celidnce on relatively limited Pliueene maternal, the suyilarities. between 8. bailev and Ss. antiquus imply a close relationship. They are more derived than Ss. cegved and §&. brachyselegis but more plesinamorphic than ony deserthed Pleistocene species. Features not shared with each other are either those shured with the more dolichocephilic species in the case of §, antiques, or with the more bruchycephalic species in the case of §. baileyi. We Tedford’s (1966) subgeneric (generie sensi Flannery 1983) definitions hold (ie. Simayieniray = brachycephalic, low-erowped cheek teeth with low links and many coarse enantel erenulations, Sdicnurns sense stricto = dolichocephalic, high erowned cheek teeth with Strong links and few fine enamel! crenulations), then 9. andfguns may represent ihe least demyed species in the subgenus Sihenusts, while S bullevi may fulfil a similar position in Simosthenurny (Fig. 9). Beeause 8. nolabitis Bartholomai, 1963, au apparently derived Johchocephalic species co-Gccars. with So antégines in the Pliocene Chinchilla deposit, the diverzence of the shorter- and longer-faced sthenaurine groups must huve oeeurred much eurher im the Pliocene Similarly, very derived species co-seeur with Ss. fuileyi an the Pleistocene, but all that thie demonstrates is thal 8) aetiquits anc S. bailey? anc striiciural precursors to the doli¢chocephalic une brachycephalic lineages. rather than part ol Wret direct ancestry So given ther verisimiltude, are ihe dillerneces between §. udev? and So amtignas sufhetent 1 Warrant phivement ji dillerent subgenera’! While they dee net possess muny of ie extreme character states Todfoed (MOGs uscd to deline the subeeneri, the questiau ts plywogenenedihy irrelevant so lors us Sritesuuriens ail Sehenireiy SA ae monophyletic The validity @f these tusa ty curreniy under Trvestiwanou Dy on of ts (GSP) and requires sine revision, sive (he Humber at descr beat erhenurine 14 G. J, PRIDEAUX & R. T. WELLS Srenurus brachyseleniy Sthemuris eegsal Hadronomas puckridgi longer faced = Syhenuruy — Sthenuris — shorter faced Sthenurus spp. antiqnis baileyi Sthenurus spp Fig. 9, Possible phylogram of basal relations in the Sthenurinae, based on the following synapomorphies. |, Cranium relatively large, neurocranium flexed dorsally relative to rostrum: occiput close to vertical. broad and deep with well- developed lambdoid crest; large palatal yacuities, narrow post-palatine bars; deep jugal expansion torming ectoglenoid process; Jaterally expanded supraorbital crests; ectotympanic thick, wide, cancellous and yentrally-kveled: ascending ramus relatively vertical. with pterygoid fossa elevated and deep: digastric sulcus / eminence well-developed: 12 very small and splint-like; 13 dominated by buccal crest. lingual crest restricted to anterolingual corner: upper meisors form V- shape when viewed yentrally, C1] absent: p3 bears posterobuccal cingulum: molars fairly short relative width and squarish in occlysal view: molar lophs relatively straight and close to parallel; lower molars with posterior face of hypolophid inflated ventrally. 2. Rostrum broad and deep; zygomatic process of squamosal relatively deep; dentaries unkylosed at symphysis; mandibular ramus deep and wide, with depth at symphysis barely shallower than beneath molars: P3 with lingual cingulum raised into crest, separated from buccal or main crest by longitudinal basin traversed by ridgelets: p3 with buccal cingulurn raised into crest, 3. p3 with curved buccal crest separated from main crest by wide median valley: p3 widened posteriorly: molars with more fine enamel crenulations. 4. p3 wider overall relative to length, with longer buceal crest: lower molars with crisud obliqua and paracristid shifted more lingually, 5, Cheek tooth row long relative to ramus depth, higher crowned molars, more prominent cristid obliqua and paracristid. 6. Cheek tooth row short relative to rumus depth, brachycephalic: retained lower crowned molars, low cristid obligua and paracristid. species has roughly doubled since Tedford’s (1966) review. Almost certainly, §. cegsai and 8. brachyselenis have va place within the two subgenera because they lack muny of the delimiting character states and appear to be the earliest derivations from the sthenurine lineage, post- Hadronomas piuckridgi (Fig. 9). We await the discovery of further Pliocene species to confirm exactly where S. cegyeai and 8. Arachyselenis fit within the sthenurine radiation. As more taxa become available more light will inevitably be thrown on this paramount phase in sthenurine diversification, Acknowledgments We thank N. Pledge and J. McNamara (South Australian Museum). R. Molnar (Queensland Museum) and R. Jones (Australian Museum) for the loan of specimens. L. Hatcher loaned the Western Australian specimen for comparison. J. McNamura is sincerely thanked for proyiding data on the type locality, preparing the holotype and his characteristic perspicacity when commenting on an early draft of this paper. We are alsa grateful to J. Long and M. Atcher for their constructive criticism of the manuscript. NEW PLEISTOCENE KANGAROO | a References Avtibhn Lo K., MARIANELI. PLC. MeCuntocn, M, T. Mortimer. G. E., HELLSTROM, J. C., Moriarty, kK. C. & WrLts. R.T. (in press) S00 ka preeipitabon record from southeastern Australia: evidence for interglacial relative undily. Geclowy. Brows, H. Y,L. (1908) Bone breccia and rock phosphate at Brothers Islands p. 343 de “Records of the Mines of South Australia, 4th edn” (Government Printer, Adelaide), FLANNERY, To FE (1983) Revision in The macropodid sublamily Sthenurinae (Marsupiala: Macropodoidea) wad the relationships of the species of Tropasodon and Lagastrophus. Aust. Marimet. @ 15-28. (1984) Kangaroos: 15 million years of Australian hounders pp. 817-835 Ja Archer, M. & Clayton, G. (keds) “Vertebrate Zoogeography and Evolution in Austria” (Hesperian. Perth). Fiownr, W. H. (1867) On the development and succession af teeth in the Marsuptala, Piri. Trans, R. Sec. Lond. B 157, 631-641. Lucnhnti, W. PF (1993) An ontogenetic as ment of dental homelogies in therian mammals pp. 182-204 fa Szalay. KS, Novacek, M_ J. & MeKenna. M. C: (Eds) “Mammal Phylogeny® (Springer-Verlag. New York), PRIpb AUX, God. & Wares. R. T. (1997) New Sthenarus species (Macropodidae, Diprotodontia) from Wellington Caves and Bingara, New South Wales. Prac, Linn Sec. NSW 1E7, ISL-L96. Rich, PV. (1979) The Dromornithidae, an extinet family of large ground birds endemic to Australia, Bie Min. Res. Bull, 184, 1-196, Rink, W. D. L. (1993) Jackmehoneva pen, nov, and the genesis of the macropodiform molir, Mem. Ass, Australas, Palaeontely 15, 441-459, TEprokD, RM. (1966) A review of the niaecrapodid genus Sthenurus. Univ, Calif, Publ. Geel, Sei. 87, 1-72. — & Woopsurknrt, M, O, (1987) The Uartidae, a new family of vombatiform marsupials from Miocene strata of South Australian and ain evaluation of the homology of molar cusps in the Diprotodontia pp. 401- 418 da Archer, M. (Ed.) “Possums and Opossums: Studies in Evolution” (Surrey Beatty & Sons, Sydney). Wetts, R, T. & Murray, PF (L979) A new sthenurine kangiroo (Marsupiala, Macropodidae) fram south- eastern South Australia. Trey. R. Soo. 8. Ast. 103, 213- 219. . Mortarty, KL & Witiiams. D. L. G. (1984) The fossil yertebrate deposits of Victoria Fossil Cave Naracoorte: an introduction to the geology and fauna, Ausi, Zool, 21, 305-333. Wiitams, Do L. G. (1980) Catalogue of Pleistocene vertebrate fossils and sites in South Australia. Trans. R- Soc, S. Aust. 104, (01-115. Appendix Material used for comparison with S. baileyi. See “Introduction” for abbreviations, except AM = Australian Museum, QM = Queenslund Museum. Species Sthenurus antiqtys S. brachyselenis S. browned (eastern form) S. cegsat S. gilli So mnaddocki S. acetdentalis (custern form) S. orecn S. pales Registration Number QM F293]. F2973 AM F31026 SAMA P2043, FU 0202, FU 0271 SAMA P31800 (tiolatype) SAMA P16528, P16629, P2()797. FU 0246 SAMA P16627, P16643. P16G73 SAMA P20798, P27799 QM F2923 (holotype) SAMA P27797 Locality Chinchilla, Darling Downs, Qld Wellington Caves, NSW Victoria Fossil Cave, Naracoorte, SA Corra Lynn Cave, Curramulkia, SA Victoria Fossil Cave, Naracoorte, SA Victoria Fossil Cave, Nuracoorte. SA Victoria Fossil Cave. Naracoorte, SA Darling Downs, Qld Victoriit Possil Cave, Naracoorte, SA MACROINVERTEBRATE ASSEMBLAGES OF GOYDER LAGOON, DIAMANTINA RIVER, SOUTH AUSTRALIA By FRAN SHELDON* & JIM T. PUCKRIDGE* Summary Sheldon, F. & Puckridge, J. T. (1998) Macroinvertebrate assemblages of Goyder Lagoon, Diamantina River, South Australia. Trans. R. Soc. S. Aust. 122(1), 17-31, 29 May, 1998. The wetlands in the arid zone of Australia have considerable significance as drought refuges. Despite this their biology is poorly documented. Goyder Lagoon is an arid freshwater wetland in the driest region of Australia, the central Lake Eyre Basin, South Australia. The abundance and richness of macroinvertebrates was examined at 11 sites within Goyder Lagoon. Insects, comprising 76% of taxa and 63% of individuals, dominated the assemblage. The prosobranch gastropod Thiara balonnensis (Smith) was the most abundant taxon, the prawn Macrobrachium australiense (Ortmann) comprised the greatest biomass. Key Words: Macroinvertebrates, semi-arid river, functional feeding groups, Goyder Lagoon, variability. Treinsaetions af ile Reveal Savierny ofS. Aas (199%) $2207), 02-31. MACROINVERTEBRATE ASSEMBLAGES OF GOYDER LAGOON, DIAMANTINA RIVER, SOUTH AUSTRALIA - HF py af by FRAN SHELDON & JIM T. PUCKRIDGH Sunimary Stiecuoy, B& Prccaipoe, 21) ( 1994) Mavcoinyertebrate assemblages of Goyder Liaoon, Didukintiia River, South Australian Tas, RoSed, S. Anse 12201), 17-3129 May. 199%. The wetlunds in the wie zone of Austialia have considerdble significance a4 drought refuges. Desptic this (heir biology is poorly documented. Goyder Lagoon rs an arid freshwater wetlind in in the driest region Ol Ausialia the central Lake Eyre Basin, South Australia. The abundance and richness of macroinvertebrates was examined at TT sles within Goyder Lagoon, Insects. vomprismye 760% of taxa and 63¢¢ of madiyiduals. dominated the assemblage. The prosobranch gasteupod Titan batonnensis (Siwith) was the mostubundant een, the prawe Membrchinncaustralieive (Orimang) comprised the grenlest biomass, Collectors were (he dominant fecding group ueross all habitats with tbe ridos of different Functional Feeding Ciroups (FRCS) suggesting that Goyder Lagoon is heterotrophic, dependent an allochthonous organie matter ys iLcurhon source, Multivariate analyses separated temponiry fron) perinanent habitats, The prawn Af ausirafiense, ihe yabbie Cheray destructor (Clark), We ephemeropterin Tagmaneceeniy arecata AtThu-Tercedor S& Suter anc the Wichppreran Bones. sp, dominuled at frequently inundated sites whereas infrequently inuridated Sites. Were dominated hy the nomnectid Aviiharey sp,, and the curinids Micrmiecte spp- There were suiking diNerences in assemblages trom different sites, even those From (he samme mesohabiiat type channel, Witerhole or billabong), ‘This may reflect diferent successional (rajectories wilhin the assemblage at cul site afler hydrological isolution. This supports the hypothesis that the variable flows characteristic of Gover Lagoon ace inporkint for mamtaining macvolnvertebrate diversily, Key Worps Macrainvertebrates, semi-arid river. funetional feeding groups. Goyder Lagoon, variability Introduction The establishment sind maintenance of specific types of wetlands amd wetland processes are controlled, it least in part, by the prevailing hydrological revime (Mitsch & Gosselink 1986). Although it sounds contradictory, wethuids do occur in and landscapes. Dryland rivers and their assocrated wetlands have a number of unique hydrotoical features (Molles & Dahm 1990; Walker etal, 995; Puckridge er al 1998). The dominant hvdrolovieal rhythms of drylkimd myers are not seasonal or annual but vefleet. in part, large weather phenomena such as the El Nifio Southern Oscillation (NSO) (Walker.e/ af. 1997). In the Australian arid fone, significant freshwater Wetlands. are mainly associated with nver channels wre floodplains and intermittent river Flow ensures that these wetlands do not accumulate sult from year to year und beeurne silt lakes (MeComb & Luke 1988). Wethids are amongst the world's most productive ceysysiems. Arid zone wetlands have consiteruble rewional significance for migratory birds (Breen {9Ul> Kingsford 1995; ANCA 1996), Despite this, Cooperitive Rescurch Centre Tur Prestivuter Ecology Departed of Zandogve Pie Unteeyerey ob Adelayde Aust. SUG their biology is poorly documented, in generul (Breen J991) und almost nothing ts Known of Australia's acid freshwater wetlands (Puckridge in press). This study describes the macroinvertebrate assemblage composition ul eleyen sites. within Goyder Lagoon. an arid freshwater wetland on the lower reaches of the Diamantina River. Lake Eyre Basin, South Australia (Pig. |), The Diamantina River system is one of the major wetland systems mn Austrutia thar remains substantially unmodified by auerresouree development and. as such. is likely to harbour w relatively pristine brota (ANCA 1996), Little, however, is known of the pracroinvertebrate fauna of any of the rivers in the central Lake Eyre Basin, Puckridge & Drewien (J98S%) and Rei & Puckridge (1990) list same of the taxa found in the Coongie Lakes system but there are no published data on the macroivertebrates of Goyder Lagoon, The highly variable Mows in Goyder Lagoon are likely to produce a diversity oF habitat types with a rutize of inundation frequencies and hence a spectrum oe different habiluts for aquatic invertebrates over time (Boulton in press), Multivariate analyses were used to identity the environmental yvariibles structuring the assemblages ai euch site and for different habiuat types. ‘The Functional Feeding Group (FFG) composition (Cummins & Klug 1979) of aysembliwes at cach site 18 F. SHELDON & J.T. PUCKRIDGE Diamantina River, Birdsville one G6 Eyre Creek — Diamantina River pshiplor Clifton Hills Outstation Pelican Lagoormh, + /ADHS Gr EGS PAIS Koonchera Dune G4&G5 ’ p CHHS Pootha Pootha Yelpawaralinna Ss G10 & G11 G9 We 0 50 100 km Fig. 1. Position of sampling sites (cf. Table 1) in Goyder Lagoon and on the Diamantina River. The heavy line marks the boundary of Goyder Lagoon. ADHS = Alton Downs Homestead; CHHS = Clifton Hills Homestead. MACROIN VERTEBRATES OF GOYDER LAGOONS 14 and in different habitats is explored with ratios of different FPGs Used to mdieate ecosystem altrihdtes. Miributes such as the rekuive jmportance of fictevotrophy or aulolrophy (Suggested by 3 domimunee of cither dearivarous eollectors or herhivafous: scrapers) und the relative amounts of Course Particulate Organic Matter (CPOM) and Fine Particulate Organic Matter (FPOM) in transport or ia slonige (suggested by the presence of filtering collegtors: Gr gathering collectors) (see Merrit & Cummins (996) were examined. Goyder Lagoon may he similar to (he Chow iki wethind of the lawer River Murray, South Austatat where colleetors domimuted The assemblige suggesting ollovhihaneus ordavie Cirbon was Ldeniinint Hood source (Boulton A& Tayi 997}, Materials and Methaeds Sry aired Goyder Lagoon is a 3030 kt inernddenr ternvil Comin & Williniis 1994) floodplain wethind sit ihe rerminus OF the Diaiantini/Georg ii river systen whieh has uo cachmenr aren of 365,000 ki (Pies. 1), Phe Diamantina is ane ol the most hydrologiowlly variable sven systems ih the world | Pochrupe er af, }9U8) Govder Lagoan beains 80 km below thie kiwi of Birds wille where fhe Diuinidna River splits ite an easter and a western unabraneh. These barnches then radiate into a redieuhite system of fine channels (o form the castern und western) intechal deltas. The custern internal deli ts larger and more frequently Hooden, Water thal makes ils way (hrough the maze of Cioyder Lagoon draiims to Lake Eyre vin Warburton Creek. The lagoon is bounded hy the dunefelds of the Sampson Desert to the west und northwest ane by the gibber plains ol Stiets Stony Desert to the cast. The survey wus conducted during November 1993 when wuler levels in the laoon were low. the fast pion flodding having oevurred in nid L991. AU bot the deepest waterholes Were dry dnd sampling was (herefore Confined lo these waterbodies. In the period JURS-1993 [lows occured samewhere in the lagoon Y oul ol LO yours (bandsat dati; South Agstealian Department for Eovironment. Hertlaye and Aboriginal Affairs). Flouding Crequency ane Cloned posicdence (imes wppeien Tram the Landsat record. diminish in the ligeon downstroam, and ins the northern fagoow tron) east la west. The inundation frequency oF cach site within the lagoon was ranked using an estimate from the overall gradien( in inundation frequency evident in the Landsat record (1986-1993) und cheeked with loeal Jandowners’ knowledize. Sites were chosen represent abo range of hydrological conditions and a spatial distribution aluog the lagoons axis (Fig. 1), In order of inundation frequency. snes meluded: * the permanent channel of the Diamantina Rivet upprosimately | Km dowastreat from the town of Birdsville (GG), * the pernunent channel of the main (northeasiern) branch of the Dianmntina in Goyder Lagoou at Chitton Hills Outstation (G7) and w closely associated backwater (GS), ‘un isolated waterhole at Andrewilla Station on it semi-permunent segment of the northwestern branch (G2), closely ussocnited backwater (Ci). and a onearby isokued billabong, Pelicun Lagoon (G3). ‘4a deepened but impermanent portion of the reticulate channel system ol the eastern dele formed by the iitrusion of Koonchera Dune (G4. GS), eo segment of the less frequently tlowded Warburton chanel io ahe southern dagounm, Poolttia Pootha Waterhole iGl0). und ai associutead billabong (G11), + un isoliled waterhole on a tess Hequently Mlooded wabraneb of the Warburton downstraant of GT, Yelpawuralinna Waterhole (G9). Aquitic’ systems such as Goyder Laguon can be desided Tote maero-, mesa. tind miero-habihuts (Walker ef af }9US). Lisuie fis hiertirehical system. Goyder Lagoon is a distinet mycrohabitat. Within Unis maerohubitut exist mesohabitats such as channels, waterholes aid billaboogs and within hese ae Microlabitits such ws emergent and submerged vewelation, submerzcd wood aad other substrate, In this study four mesohabitit types, channel. witerhole, backwater and billabong were sampled (Table 1). Overall microhubitats tneluded emerent vegetation such as sedges (Cyerky spp.) and semi- aquatic grasses (eg. Cyredon spp.) Torming sometimes dense stuns along the ed@es ot tre waterbodies, aquatic macrophyles (ee, Colvecuias sp. Miriepivilon spp. submerged woody debris (snags') represented by the roots or Lullen lambs of trees Sue os river redguntm® (Avelyn cimeldileasis Delinh. var ebtuse Blakelys Coolihah (Eyvealpai colihah Blitkely & Jacobs ssp arid (Blukely) b, Johnson & K. Hill) and River voobah (dca ia verephyla Cun ex Benth), course particulate organie matter such as packs of lea! litter und (wigs From riparian vegenition, and unvegedited littor) arews free of veeetition, woody detritus oF other cover. The latter were further divided into silt and clay suBstaitum, Sandy substratum: or rok substratum, Ninely (wo sumiples were collected from microhabitals present within the TT sites (Pig t, ‘able 1). 20 PF. SHELDON & J, T. PUCKRIDGE Pabur |. Lise of sites (ane their abbreviations) wrauped by niesohabitar. rhe number of samples collected from each microhabitat at each site ts alsa given. Mesohabitat Site Site Name Microhabitats No. Number Samples Backwater Gl Backwater of Andrewilla Waterhole Leaf litter 4 Silt 4 GH Backwater of Clifton Hills Waterhole Polygonum 3 Lignum 4 Silt 3 Waterhole (2 Andrewilly Wilerhole Rock 4 Sill 8 G4 Koonchera Dune Walerhole Sand 4+ (caster portion) Sill 4 GS Koonehery Dune Waterhole Lignum 4 (westem portion) Sand 4 Sill + G7 Clifton Hills Waterhole Polygonum 4 Lignum 4 Silt 5 Shag 4 ci9 Yelpawaralinnit Watterhole Emerg. vey 4 O10 Pootha Pootha Waterhole Emerg. veg 3 Silt 3 Channel Go Main channel of Diamantina River Polygonum 3 (Birdsville) Grass 3 Rock ‘f Silt 4 Billabong G3 Pelican Lagoon Billabong Silt 2 Gil Small Billabong Silt I Sample collection und processing Each site Was considered a distinct mesohabitat und was stratified into the microhabitats defined above (Table 1). The most prevalent of these were then sampled randomly (Boulton & Lloyd 1991), Macroinvertebrates were collected in replicate samples from each microhabitat by sweeping a SOO um mesh pond net ever 5 m-* for 20 sec, Samples were preserved in 70% ethanol and later washed through nested sieves (4000, 2000, 1000, 250 pm) and the organisms hand-soerted, enumerated, and identified as far as practicable, Unidentified specimens Were recorded as separate taxa (e.g, “tiny Zygoptera’”), Each taxon was assigned to one of four broad functional feeding groups (FFGs): collectors, predators und serapers (Cummins & Klug 1979), with collectors incorporating, filterers and gatherers. The assigned FPPGs, particularly “collectors”, ure tentative as the diets of most taxa are unknown; assumptions uboul diet were therefore based on MBollros, AW. (1988) Composition and Dynamics ot Macromyerebote Communities in “Pwo Intermittent Streatys. PhD thesis, Monash Uniwersity CGunpub.y 2 Seen A (}994) Lattetal Eeology ofa Repuluted Dryland River (River Murray, South Austratia) with Reference to the Gastyapoua PHD thesis, Unversity of Adelaide (unipub ). taxonomic affinities. Classification of taxa int functional feeding groups followed Cummins & Kluy (1979), Boulton (1988)', Boulton & Lloyd (1991) und Sheldon (1994)") At each sampling site. spot measurements were recorded of depth, Séevhi lranspurency, temperature, dissolved oxygen CYS] oxygen probe) and conductivity (Hanna HE 8733 conductivily meter): sulinity was computed trom temperature corrected conductivity values using the equations in Williams (1960). Data analysis Hierarchical analysis of variance (Underwood IO81) was used fo explore patterns in fichness and abundance between mesohabital und microhabitat scules. As the sampling design was unbalanced, subsets of the total dataset were used, Subsets were chosen so there were equal replicutes within each level making the design balanced. The lollowing scpurate unalyses were performed: i. Microhabitats (emergent vegetation, sill) bested within sites (G6, G7, G8, GIO), i. Microhabitats (ignum, silt) nested within sites (G5, G7, G&S). All analyses were performed in SYSTAT vy, 5.03 (Wilkinson 1990), Data were rendered normal by (ranslorming using log ),(X + 1). MACROINVERTEBRATES OF GOYDER LAGOON Zz Hierarchical patlerns in the dita were also explored using inultivariate statistics from the PATN software puckuge (Belbin 1993). Multivariate analysts allows patleriis. between samples to be explored. ‘lwo multivariate approaches were used: classifieation (oy derivation of discrete proupings of samples) and ordination (arringement of samples ina space ot a few dimensions). Both techniques urrange samples on jhe basis of their species conjposition, with those samples that cluster or group, together more likely to be similar in species composition (ler Braak 1947). Data were log,,(% + 1) ininsformed before analysis and then range standardised as suggested by Belbin (1993); the Bray-Curtiy coefficient Was Used as. the measure of dissimilarity between samples. AIL taxa occurring in one OC Mure Samples were retained in the analyses. Mlexible-UPGMA was used to cluster the samples, with the ANOSEIM procedure (Clurke 1993) used to test for significunt clusters of samples ata meso- and microhabilal scale, The SIMPER procedure Trom the PRIMER software package (Clarke & Warwick 1994) was used to identify the percentage contributions of different taxa to the simple groups. Semi-Strong Hybrid multidimensional scaling (SSH) ordinations were also computed trom the Brey-Curtis similarity matrix generated [rom the transformed and standardised data: setuugs of 30 itenutions and J ratio-ordinal cut value of 0.8 over 100) random starts was used, Solutions were calculated in two, three and four dimensions. The solution presented here had a stress of 0.18, The Stress ineasure gives an indication of the “goodness- of-lit” for the ordination (Kruskal & Wish 1978); an ordination with a good fil fur sumples within the ordination space basa stress value of less than 0,2, Sample groups from omesohubitats. sites and Mmicrohabitais were mapped on to the ordination plots. Rehidionships und between environmental community data are usually many, conrplex snd nonlinear (Gauch 1982). One aun of condueting multivarigte analyses is to detect major differences im Species composition for sample groups which are potentially related to environmental differences (Boullon 1948)', Spearman Rank cornlavons (Zar 1984) between the physicochemical variibles measured in the Held and the ordination scores on the SSH axes were calculated 10 examine relationships between environmental factors and macroinvertebrate ussemblige composition, Results Environmental conditions The environmental conditions for all sites are given in Table 2. The Ditumuntina River earries a high load of yery fine suspended sediment with purticle diameters of less than | pm. This contributed to Secchi depth measurements well below 20 em at all sites, Salinites were highest at those sites in the centre of the Lagoon which were the most chsconnected from any recent flow of water, sites on Andrewilla Station (G1 and G2),at Koonehera Dune (G4 und G4) and Pelican Lagoon Billabong (G3). Pootha Pootha Waterhole (GIO) at the southern end of the Lagoon had a lower salinity, probably the result of recent rain_ Sites on the main channel of the Diamantina (G6) and Clifton Hills Outstation (G7 and G8) also had relatively low salinities due to a recent small flow through the system. Al sites were well oxygenated (ncur saturation) with imstantineous Avaler temperatures manging from 20.34° C. Parterns of richness dnd abundance A total ef 7.363 individuals from 54 taxa was collected in. 92 sumples (Appendix), Insects were the dorminant group comprising 76% of (axa andl 63% of individuals (Fig, 20). OF the Insecta, the Diptera TabLE 2. Environmental conditions measwad in the livaral zene of each site in Gowler Lagoon, Diamantina River South Atuyinlia ta Noventber 199), Datawere not uvailable tof sites Go and G11 - n/a = nol avnilables Site Sample Depth Salinity (em) (moh't Gl Fd 309 (2 Oi 26) Ga aM) 8265 Ga 57 (299 Gn 4a 2133 Gin Wa wa G7 62 172 Gs 74 }2> Gy 63 769 G10) as 142 ml al no Seecht Temp Oxyyven (em) (°C) (Se Saluralion) 7 Wa ou 0) 6.0 230 ony 20.0 34.0 V0 40 227 ORD 40 24,7 WAY Wil ws ta a0 26,0 KA ay 25,4 THs 40 Td) Sho My 716 nl nia oil mi 22 PF. SHELDON & J.T. PUCKRIDGE comprised 39% of taxa and 41% of individuals (Fig. 2b), The prosobranch gastropod Thiara balonnensis (Smith) was the most abundant taxon, with the freshwater prawn Macrobrachinm australiense (Ortmann), the corixids Micronecta spp.. the predatory chironomid larvae Coelopynia sp. and the predatory caddisfly larvae Oeverty sp. also common. The prawo M, ausrraliense comprised the greatest biomass. Five taxa occurred only once. Observed at Koonchera Dune Waterhole, but not collected, was the freshwater crab Holthuisiena (Austroielphiuse) transverse (Martens), Both the number of taxa (richness) and the number ol individuals (abundance) differed between the inesohabitars (Fig. 3). Backwaters and waterholes had more individuals than the channel or billabongs. However, richness was similar for all mesohabitats, When the mesohabitats were split into siles there were differences between sites from the same mesohabitat. Of the backwaters, GI had many more individuals and a greater richness than G&, Similar differences occurred between the billabongs G3 and G11 and there was also variation in both richness and abundance for different waterhole sites (Fig. 3), Numbers of taxa und individuals also differed between microhabitats depending upon the site, Large numbers oF individuals were found in leat litter packs in GI and silt microhabitat trom beth C4 and G5, fewest individuals were found in snag and silt microhabitats from G7 and the rock microhabitat from G6. Emergent vegetation and submerged aress of hgnum contained the greatest number of taxa while the fewest number of taxa were from snug microhabitats. Bivalvia (CD Gastiopeda O Oligochaeta WB Hirudinea Ej Crustacea [J Insecta (_]Ephemeroptera | Odonata 3) Hemiptera ff] Coleoptera Diptera (]Trichoptera Fig. 2 fi Percent representation of the major invertebrate groups in samples from all sites within Goyder Lagoon, na? mber 1993, in terms of the number of (i) taxa in each group and (if) abundance of individuals. (b). Percent representation of the major insect Orders in samples from all sites within Goyder Lagoon. November 1993. in terms ol (iil) the number of taxa in cach Order and (iv) the abundance of individuals, MACROINVERTEBRATES QF GOYDER LAGOON Mean numbers of individuals (FP, , 6.246, p>0.05) did not differ between sites G6, G7, G8 and GIO bot mean numbers of taxa (Fy, = 12.178, p0,05) and individuals (Fy), = 1.607, p>0.05) did not differ for emergent vegetation und Sut microhabitals nested within these sites. There were also differences in mean number of taxa (FP, = 148.08. p<0.001) and individuals (FP), = 103.47, p<0.00T) between the sites G5, G7 and GS, with GS having more taxa and mdividuals than the other sites (Fig. 3). However, again there were no differences for either taxa (Vy), = 0.192, p20.05) or individuals (Fy, = 0.603, 750.05) between lignum and. silt microhubiliaty nested within the sites. Multivatiite dadlysis of samples UPGOMA classtlieation of the sample data indicated Wo main groups, The majority of samples foie the more temporary habitats (Cel, G2, G3, G4, G5, GY. G10) lonmed one group (A-D) while those from more permanent habitats (G6, G7 G8) formed 8 Abundance 8 250 g (b) Wa rs) Sit Lignum Silt Silt Leal Liter Polygonum (c) Nig. 3 23 the other (E-H) (Table 3, Fig. 4). The first group (A- D) further split into Group A-B, containing samples from the western anabranch of the lagoon (G1, G2) which is the more frequently inundated of the temporary habitats, one sample from G6 and one from G7, and Group C-D containing samples from the extreme temporary habitats on the lower section of the eastern anabrinch and the southern end of the lagoon (G4, G5, G9, GIO). The second group separated into Group E-G containing a majority ot sumples from Clifton Hills Outstation on the upper section of the eastern anabranch (GI and G&) and Group H containing most of the samples from the permanent channel of the Diamantina River al Birdsville. The single sample [rom the infrequently inundated billabong associated with Pootha Pootha Waterhole, GIL, was an outlier clustering with the nore permanent sites in groups A-D, SIMPER analysis showed thal the prawn Macrobrachium australiense, the yabbie Cheray destructor (Clack), the ephemeropteran nymph Tasmanoeoenis. arcuata Alba-Tercedor & Suter and the trichbopreran larvae Leveitiis sp, dominated the asseinbluge in the more frequently inundated sites, Ute | eee peas Channel Billabong eee = Mile = Y | Whoa! G3 oe G5 GIO Gif Silt Emerg. Veg. 5 80 Bo mK unui a | EgE= oO 2 = uz = §88a 3 5? & § i ge% 8 § 5 a & . Mean (486) of richness ind abundance for each of the three sampling scales. (a). Mesohahitats. Uh}, Sites. within Ealulontuat (c). Microhabitats within sites, collected from Goyder Lagoon, Novernber }993, Points indicure richness und bars abundance, 24 F, SHELDON & J, T. PUCKRIDGE TABLE 3, Flexible-UPGMA groups by site, mesohabitat and microhahitat. Those taxa contributing more than 5% to the similarity of the sample groups are also indicated. UPGMA No. Group Samples A 21 B I Cc 23 D 1 E 8 F 22 G 2 H 5 z S a > 06 5 00@°e@,. @r-eee @): ® @ o 7m 00 > No, Samples from Sites 7Gl 12 G2 | G6 1 G8 1 G7 2G3 1G7 12.G5 8 G4 4G9 6 G10 Snag 0.8940 22 22 No. Samples from Mesohabitats 8 Backwater 12 Waterhole 1 Channel 1 Waterhole 2 Billabong 21 Waterhole 10 Waterhole 3 Backwater 4 Channel | Waterhole 4 Channnel 11 Waterhole 6 Backwater 1 Billabong 2 Waterhole 5 Channel 1.0132 No, Samples from Microhabitats 4 Organic Matter 13 Silt 4 Rock | Polygonum 1 Silt 8 Sand 4 Lignum 4 Gruss 3 Sedge 3 Silt 3 Silt | Grass | Snag 3 Polygonum 1 Rock 2 Snag 7 Lignum 4 Palygonium 6 Silt 2 Grass 2 Silt 3 Rock 1 Silt | Polygonum 1.1324 1.2516 ' Contribution of taxa to sample group (SIMPER) Thiara balonnensis Coelopynia sp. Micronecta spp. Macrobrachium australiense no taxa Oevcetis sp. Coelopynia sp. Bezzia sp. Enithares sp. M. australiense Micronecta spp. Coelapynia sp. Anisops spp. M. australiense Austrogomphus australis Cherax destructor M. australiense Tasmanocoenis urcuata no laxa Ecnamus sp. M. australiense 1.3708 1.4900 57.6 40.6 Fig. 4. Flexible-UPGMA dendogram, using Bray-Curtis dissimilarity, of the 92 samples collected from Goyder Lagoon in November 1993. Entities in sample groups A-H are listed in Table 3. A pictorial summary of the microhabitat representation of samples is also given. Large circles depict greater than 70% of the samples from the microhabitat occur in the sample group, medium cireles 30-70% and small circles 1-30%, MACROINVERTEBRATES OF GOYDER LAGOON oF G6, G7, G8 (Table 3, Fig. 5). The prosobranch vastropod Thiara balommensis, predatory tanypod lurvae Coelopynia sp. and corixids Micronecia spp. dominated samples from sites at Andrewilla (G1, G2) whereas the predatory tuichopteran larvae Oecetiy sp. and Coelopynaia sp. dominated samples from sites at Koonchera Dune (G4, GS) as well as the temporary billabong G3, The infrequently inundated sites GO and G10 had an assemblage dominated by the highly mobile predatory hemipteran Enitheres sp.. the prawn M. australiense and corixids Micronecta spp. (Table 3, Fig, 5). Ordination highlizhted the differences between Thesuhabitats and between sites. When grouped by mesohabitat, those samples from backwaters. were dispersed across Axes | and 3 and grouped low on Axis 2 (Pig. 6). Samples trom waterholes were dispersed across all three axes whereas samples from channels grouped centrally on Axes | and 2 and low on Axis 3. Billabong samples were central on Axes | und 3 aind grouped high on Axis 2. When samples were grouped according to sampling site, their disthbution along Axis 2 refleeted the inundation frequency of the site (Fig. 7). Samples from the more permanent sites Go, G7 and G8 grouped low on Axis 2, those from {he less frequently inundated G1 and G2 prouped centrally on Axis 2 while those from the more extreme temporary sites G4, G5, G9 and G10 grouped high On Axis 2, Samples showed no distiner groupings according to microhabitat on any of the three axes (Pig. 8). The ANOSIM procedure suggested diflerences belween groups of simples at cach level: significant differences were located between sumple groups ata mesohabitat (R=L 118, p<0,001), site (R=1.362, pe0.0O01) and microhabitat (R= 1.079, p<0,001) level, Inman Frequent Infrequent Frequency —— ‘Siles Ge ial cx] fev) G7 G2 Ga a1 GB G&S UPGMA EFGH A c D Groupe Dorrwnant race Mecrowrachiun Thilcwrne Oecelissp Evntyes sp (SIMPER) dust abonioe padorinensle Crsuicean Gasimpod = Preciuiory —— ikphdy muatiiien Trenopteran predatary Biv 5S. Sites, UIRGMA itoups. and dormant tix Mren SIMPER in relation ty inuadatiua Frequeney, LIPCIMA eroup B has been omitted as i contains ont ore sacnple ind domindnt (Xu chniet be vendnited for one sample wroups using SIMPLER, Also the single sample from site Git (in LIPOMA feoup P78 Ongitied ae ie an edbier. being uniifrequently flooded Site that grouped with the Frequently Mouslevt sles. With regard to the jnstantaneous environmental variables measured, the sample distribution along the first axis of the SSH ordination was significantly correlated wilh Seechi depth, temperature and oxygen saturation (Table 4). Axis 2 showed significant correlations with salinity, temperature and oxygen saturation and all variables were correlated with sample chstbution along Axis 3. Functional feeding groups Collectors dominated the invertebrule assemblage of Goyder Lagoon, However, there was considerable variation in the FFG composition of the mesohabitats and of sites within mesohabitals (Fig. 9), Billabongs and waterholes contained similar numbers of collectors and predators. Plowever, collectors dominated the assemblage composition of the backwaters in both richness and abundanee, When the FFG composition of individual sites was further a Billabong @ Backwalor @ Walarhole © Billabong O Channel Aats 3 ao hig G, SSH ploronases fab dy, Z(t by a le) 2, Sal auimples callected front Goyder Ligwom November (U9 Samples ure hibelled aecontine to the fiesphabitats from whieh (hey wereecoteerce 26 F, SHELDON & J.T. PUCKRIDGE DNHAODOOKAGDH Qa S=OONTOBON + 5 oO e + 4 a a im) ° e * g Cliftan Hills i) Axis 2 Fiy. 7. SSH plot on axes (a) 1 y. 2, (b) 1 y. 3, (¢) 2-y. 3 of samples collected from Goyder Lagoon, November 1993, Samples are labelled according to the site from which they were collected, @ Leaf Litter Sand © Sedge &é Silt © Snag + Rock @ Lignum Emergent Vegetation é Grass \) Axis 2 Fig. 8. SSH plot on axes (a) | y. 2, (b) 1 y. 3, (¢) 2 y. 3 of samples collected from Goyder Lagoon, November 1993. Samples are labelled according to the microhabitat from which they were collected. Emergent vegetation = Polysonum sp, and Hooded Cyperus spp. TABLE 4+. Spearman Rank correlation coefficients between environmental Variables and the sample scores on the first three axes of the SSH of faunal data from all samples callected Jrom habitats in Gavder Lagoon, November 1993. Significance levels are indicated as follows: ns = not significant, * = p<0.05 pe0.01: = p<0,001, Axis | Axis 2 Axis 3 Salinity Secchi Temp Axis 2 0,032 ns Axis 3 -.143 (),324 ns ae Salinity O.071 0,698 O.416 ns ae a Secchi -0.658 0.029 0.375 O40) ns Hat eae femp 0.394 0). 302 -0,367 -0,237 -0,355 sees yes # # * Dissolved -0,365 O14 0.409 0,835 0.688 0.261 Oxyyen MACROINVERTEBRATES OF GOYDER LAGOON 120 Mesohabitats 80 Abundance 40 (a) water Channel Waterhole 9 tb : Billabong Back 75 250 Sites \ 160 Mesohabitats Richness a fi 0 : : Billabong Backwater Channel Waterhole Sites 2 Vig. 9, Mean (+SE) of (a) abundance and (b) richness for collectors (grey bars), predators (white bars) and scrapers (black bars) for mesohabitats and sites within mesohabitats, collected from Goyder Lagoon, November 1993. Git G3 Gi 2M Bf SHELDON & f 1. PUCKRIDGE TANLE a. Pudertonal Feeding Group rats as indicators af damindnt ecosystem Processes. { Auto = autotrophy: Hetero = Heterotrophy; CPOM = Coarse Parniculate Organic Mauer; PPOM = Fine Particulate Organic Matter; TFPOM = FPOM in transport; BFPOM = FPOM in substrate] (see Merritt & Cummins 1996). Eeosystem Functional Feeding Calculated Generul Criteria rao Evaluation Parameter Group Ratios Ratio levels Auto/Hetero Scrapers to Shredders + 0.21 Autotrophic Heterotrophic system, Votal Collectors >(),75 dependent on allochthonous organic matter inputs CPOM/ Shredders (o Total a Normal Shredder No shredders, riparian FPOM Collectors assoenition linked to zane functioning functioning riparian differently from thitt syslem predicted by Merritt and >t).25 Cummins, TEPOM/ Filtering Collectors to 12 FPOM transport (in low PROM in transport. BFPOM Gathering Collectors suspension) enriched = high particulate lou 20.50 Stable Serapers + Filtering (36 Stable substrates Limited stable substrates, Channel Collectors to Shredders plentiful habitats dominated hy + Gathering Collectors >0.50 sand wn silt ‘Top-down Predators to Total of all (ag Normal predator to Large mumber of control other groups explored (here were considerable differences among sites even within the same mesohubitat, The backwaters GJ and GS showed similar patlerns with both richness and ubundance dominated hy collectors. The two billubongs, however, differed with both the pichness and abundanee of G3 being dominated by predators compared with collectors in Gil (Fig, 4). Variation in PPG) composition of assemblages also differed between the different witerliole sites. The more temporary weterholes G4, G5, GY and G10 were dominated by invertebrate predarors whereas the more permanent walerholes G2 und G7 had aw larger number of collectors, Overall, there were few serupers. OF the total of S54 tisa collected from Goyder Lapoon sik were designated as ‘Scrapers’, three as “Filtering Collectors’, 25 45 “Guthering Collectors’ and 20 were ‘Preditors) no ‘Shredders’ were colleered from Goyder Ligoon (see Appendix), PPC ratios (Merritt & Cunmmins 1996), calculated using the PRG data indicate that Goyder Lagoon ts heterotrophic, has dow levels of invertebrute mediated leat litter breakdown, and the majority of PPOM in the system ts m storage (Pable 5), Discussion Samples taken ty Navember 1993 from habitats in Goyder Lagoon contiined a total of S4 macro- invertebrate bisa, Nearly all of these taxa have also been reeorded from the Coongie Lakes system prey balance <().15 invertebrate predators, reflecting temporary habitats late in succession (Sheldon unpub.) and from the lower River Murray und Darling River (Sheldon & Walker [998 ). However, a striking feature of the Goyder Lagoon assemblage was the presence of a diverse vroup of Motlusea, including two gastropod taxa. Neropeli wwhlincata (Conrad) und Thieres beloomensix, that have become extremely rare, if not extinct, in the River Murray and Darling River (Sheldon & Walker 1995a.b), Collectors were the dominant fecding group aeross all habitats du Goyder Lagoon, Funetional feeding group rahios (Merri{t & Cummins. 1996) suzgest that Goyder Lagoon is heterotrophic and dependent on dlochtionous organie matter asa carbon source (Table 5), This is not surprising sinee aquane plants were fare within the Lagoon. siall stiids of the mucrophyte Polygonum sp. being the nist conspicuous, The absenee of shredders iv the ussemmblave sugeests (hal for the invertebrates. the major allochittynous inputs do not come drrectly from ciparier lirter fall but rather from particulate organic mater presedbits PPOM withthe substrate. Decomposing plant material of inostly terrestrial floodplain origin would therefore form the dormant Food source for He agate (iagralhvertebrate. lowed webs. Organic mutter of Moodplain origin his been found to influence the strueture of invertebrate assemibhiwes mother lirge floodplain river systenns (Post & De La Crue 1977, Caley 188: Perry & Perry 99), Thorp & Delonge 1994) Meyer er at IQV7y MACROINVERTEBRATES OF GOYDER LAGOUS 19 All samples were collecied when water levels within Goyder Lagoon were low. Low water levels lend jo maximise (he “heliveen mesohablea’ differences (ice. differences between channel, waterhole and billabong). At bigher flows. between habitat differences would become inereasingly blurred as billubangs and witterholes beedme parr oF the channel environment. Alihough we expected the main differences in assemblage vomposition to occur henveen different meso- and mitrohabitats (regardless of site). there were striking differences in the Tichness aod wbundance of macromyertebrates collected from the different sites, even belween siles from the same mesohabitat type (Pig. 3), These differenves may reflect different suocessional Inyeclones oeeurring withi the assemblage at cach site after hydrological isolation (Boulton & Lake }992). Thus. the recent Looding/dryiig history of each site may be a Significant factor in determining the structure of the assemblive at any time, Sites, regardless of mesohabitut type. at Andrewi lla and Koonchera Dune contained the largest number of individuals and the greatest number of nixa (Pig. 3). These sites ure Intermediate m the range of Mooding Frequency (Fig. 5), As Tlooding is uw form of ecological “disturbance” this supports the notion of “infermediate disturbances’ being a driving Loree tn mainwining animal diversity within ecosystems (Ward & Stanford 1963). The sites ob Andrewilli Watethiole and Koonchert Dune. falling wathin thus band of intermediate flooding trequeney. were characterised by taxa such as the prosebranch wastropad Chiara fatlonnensis and the predatory cuddistly Gecens sp, (Pig. 5). Both these taxa tended to be fare in the more permanent sections of the Lugoon. Goyder Lagoon isan ared of high conservation significance (Morton el ad T99Sa), It provides habiiat for a ovariely of aquatic and terrestrial Organisms idan otherwise arid chvironment (Morton etal 199Sb: ANCA 1996). The muacroinvertebrate assetiblage, although not unique to Goyder Lagoon, did contain a number of molluses that are becoming inereasmyly encingered in other river systems. [bis the Geomorphology of the Lagoon that grves rise to the different mesohabitat types. Overlying Uns morphological template is the hydrology of the system. The variable (ows characteristic of Goyder Lavoon are intrinsie in Maintaining macro- invertebrate diversity, The waters of wethinds in aril regions are Inereasingly in demand by water resouiree developers (Walker e7 ul 1997). 1 Goyder Lazoon ts to remain an area of high conservation value then it is imperative that the hydrological diversity ehiracteristic of the system i maintained. Acknowledgments This study was condueted as part of the Biolugreat! Survey of the Lower Diamantina Floodplalii. a joint project of the Conservation Council of South Australia and the South Australian Department of Environment amd Plunning. The project was funded by the Australian Heritage Commission. We thank R. Molsher, J. 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J Ho & Drtowe, Mo Do (1994) The riverine productivity model: an heuristic view of carbon sources and orgiinie processing in large river ecosystems. Oiker 70, 305-308, UNprRWOOD, AL J. (1981) ‘Techniques ol analysis of yariince in experimental murine biology and ecology, Annw Reve Ovennogr Mar Biol WY. 513-605. WALKER, K.P, SHELDON, BE & Pockkipoe, J. T.(1995) An ecological perspective on large dryland rivers. Regul, Riv. UE, 85-104. > Peekripob, JT & Branch. S. (1997) Trrigation development on Cooper Creek, central Australia prospects for a regulated economy i a haom-and-Mist ecology. dynatic Convervations Mavine and bresiwvater ncosyatemy 7, 04-73. Warh, 1. Vv. & Srasvorny, J. AL (1983) The intermedia disturbance hypothests: an exphination for bio. diversity patterns in lotic ecosystems pp. 347-356 My Fontaine, T, DB. & Bartell, SoM. (Lids) “Dynamies of Lotig Beosystems” (Ann Arbor Science, Michigan), Winkinson, L. (1990) “SYSTAT: The System for Statistics: (SYSTAT Ine, Evanston, Ulinots). WiLAMs. W. D. (1966) Conductivity and the conventration of total dissolved solids in Australian lakes, Auss J) Man Prestavat. Res. V7. 169-176. ZAR, JW (1084) “Biostatistical Anulysin” New Jersey) (Prentice Mall. Appendix Species, functional feeding group [PC = filtering collectay, GiC = wathering collector: & = Scraper P= Predator] und total abundance for simples collected trom habits i Goyder Lagoon, Diamantina River, November 1994 Spee FFG Total MOLLUSCOA BIVALMIA Sphacriielac Sphueriun sp. KC 5 Corbieuhdie Cerio austniliy (Deshavesy FC an Lbyrifdae VWlesiuiin wilson? thew iC | EPASTRORPODA Aneylidiue frepisvic spp S | Manorbidke Glypraphysa sp. 5 4 Vivipuiridae Veropale sublireant (Oonrad) ic II Phiri Chinn halaineisis Canis Cit! 1a OLIGOCHAETA HIRUDINEA Glossiphoniidae CRUSTACEA CONCHOSTRACA Cyzicidae DECAPODA Palaemonidae Parastacidae INSECTA EPHEMEROPTERA Caenidae Baetidae ODONATA Gomphidae Cordulidae HEMIPTERA Corixidae Ochteridae Notonectidae COLEOPTERA Dytiscidae Hydrophilidae Hydraenidae DIPTERA Tipulidae Chironomidae: Tanypodinac Chironomidae: Chironominae Chironomidae: Orthocladinae Ceratopogonidace Muscidae TRICHOPTERA Eenomidae Leptoceridae MACROINVERTEBRATES OF GOYDER LAGOON Indeterminate spp. Indeterminate sp. Cyzicus sp. Macrobrachium australiense (Ortmann) Cherax destructor (Clark) Tasmanocoenis arcuata Alba-Tercedor & Suter Cloeon sp. Austrogomphus australis Selys Hemicordulia tau Selys Micronecta spp Cymatia sp. Indeterminate sp. Anisops spp. Enithares sp. Antiporus femoralis (Boheman) Allodessus sp. Eretes australis Erichson Cybister sp. Rhantus sp. Hyderodes sp. Sfernopriscus sp. Enochrus sp. Berosus sp. Limnoxenus sp. Octhebius sp. Hydraena sp. Indeterminate sp. Ablabesmyia sp. Coelopynid sp. Procladius sp. Cladotanytarsus sp- Tanylarsus spp. Chironomus sp. Chironomus cleacalis Atchley & Martin Cryptochironamus sp. Stenochironomus sp. Parachironomus sp. Dicrotendipes sp Paratendipes sy. Cricotopus spp. Bescia sp. Indeterminate sp. Ecnomus sp. Triplectides australis Navas Triplectides clongatus Banks Ovcelis sp GC GC Qwun vised nn BREEDING BIOLOGY OF LITORIA BOOROOLONGENSIS (MOORE, 1961), AND LITORIA LESUEURI (DUMERIL & BIBRON, 1841) (ANURA: HYLIDAE) AND COMMENTS ON POPULATION DECLINES OF L. BOOROOLONGENSIS By MARION ANSTIS*, ROSS A. ALFORD}? & GRAEME R. GILLESPIE£ Summary Anstis, M., Alford, R. A. & Gillespie, G. R. (1998) Breeding biology of Litoria booroolongensis (Moore, 1961) and Litoria lesueuri (Duméril & Bibron, 1841) (Anura: Hylidae) and comments on population declines of L. booroolongensis. Trans. R. Soc. S. Aust. 122(1), 33-43, 29 May, 1998. The embryonic and larval development of Litoria booroolongensis are described and compared to those of the closely related Litoria lesueuri. The habitat, behaviour and distribution of each species are compared and indications of marked population declines of L. booroolongensis are discussed. Key Words: Litoria booroolongensis, Litoria lesueuri, embryology, larval development, habitat, aggregation, population decline. Transactions of the Raval Societe of SMa (9OR), P2200) A4-d 44 BREEDING BIOLOGY OF LITORIA BOOROOLONGENSIS (MOORE, 1961), AND LITORIA LESUEURI (DUMERIL & BIBRON, 1841) (ANURA: HYLIDAE) AND COMMENTS ON POPULATION DECLINES OF L. BOOROOLONGENSIS hy Marion ANSTIS*. Ross A. ALFORDT & GRAEME R. GILLOSPIE: Summary Assis MM, Acrarn, R.A. & Girrrsme GR, 199%) Breeding biology of Lirarta aorodtongensiy (Moore. L06)) and Litera lesuen? (Dumeéril & Bibron. IS4)) (Anuew: Plytidae) and comments on populabon declines of LE, hoarowloreenves Trax 4 Sow S. Anst (2240), 33a, 29 May. 199k The erabryonic and fsrvail dewelopent af Litera: Becrnlongensns are deseribed and compared (0 those of the dlosely related Line feet’, The hubital, bebaviode dad: distribiition of each spews are compared and indicuGurms OF marked populition declines OF Ly owropfongensés ave discussed, Nhe two species have similar Jntic 1de-distories bin some ctifferences vecur ty body proportions, colyur in life, Dehavion! and habitat. iy both species the ewe nus isu compaer gelatinous clump, lypica) ot frogs breeding an vote environment. The tadpoles of both species are adapted to the lorie environment and hive streamlined bodies and suctoial moutb-parls. Adult Lo borroebuigenste vgerepate throughout the year and are wetve curnally (7 summer, Key Worns: Literla bourmelonmensiy, bite leidied, eoobryolopy, larval development habliit, dtgregatiott. popHlationd decline Introduction Literia, baorimlongensts was first desenbed by Copland (1957), us Ayia XY Moore. pending the type descdiption (Moore 1961). Moore deserbed tas an “uplind species” extending from the Armidale region to the Blue Mountains. in NSW. bur made no relerence to larval developotent, Anstis (/97+4) briehly described the tidpole as lotic and suctorial. with a footh cow formula of -/s und numerous oral papillae. The present paper provides a deseription ul embryonic und larval development or L- houwolongensis, With Comparisons to the sriibu’ tadpole of Lo leyueuri. Litarta hoeranlanensts is associated with flowing stretinis on the slopes and tablelands of the Great Dividing Range from the Queensland border to (he Victorian borer (Barker ef al, 1995). with the type locality (Guy Bawkes Creck near Bhor, NSW) and most records of this species indicaling that it commonly oecurs above S00 m in the region of the New England ‘Tablelands, NSW (Heatwole er ef. 1995), The most southern record of this species is from the Tumut River, Kosciusko National Pork and ithas revently bee found pr the adjacent Gooburragandea River, east of Tumult NSW (Hunter & Gillespie unpub.) Observations on the current shite of popularions (particularly in the New Lnghind region) ure discussed. Literia lesuened (Dumeril & Bibron) occurs in sunterh Australia tron) northern Queensland to Vittoria und almost vertiinly involves a comple of 26 Wideyiew Ral Bepieca | Lehr NSW 2082 » Department of Zoulogy aid Teapiwal Feelawy. Ties Cook Lnrversity Tawesville Ql dst Pepariniont of Addepy, Uivgedey of Metbornie Parkwitle Vie WU52, Anhur Rylud tostiowe HO os 147 Pecinelbere Wig. tO S44 sibling species (Moore 1967: Barker ef af, 1995) Vhe type Joeality is) Port Jackson, NSW. Observations on current population trends at same southern sites are reported. ‘The eggs of the northeastern. Queensland form have been deseribed by Rivhurds & Alford (1992) and Barker ef af, (19954 brielly describe the site of epe deposition by the populations found Fram southern Queensland to Victoria. The tadpole as found in the Melbourne ares has been brielly deseribed by Martin eral. (1966). As the distributions ef both species overlap Th places, 4 more detailed deseription of Lo lesmerit tadpoles from NSW is provided here for the purposes of comparison with L, boaraolengensis and lo assist in distinguishing tbe species. Distribution maps for both species together with localities studied m the present paper, are presented as Fiyure (- Numbered localities on Fig. DA relate to 1. heatadivierests und those on Piss [Bota L. lesen Materials and Methods Material esanined Literty baeratoigensis larvae Atstahan Museum (AM) R119062-64. 119087. 119071, (19073 (Serpentine River, Point Lookout NSW), RIIY0S5, | LOUBO. 1 P9083. 1 L985, 119N87. LTRS (arvae drom Back Creek. Porm Lookout NSW). Embryo/larval descriptions ure based en one cee mass fronian amplectant pair collected at Serpentine River (near Point Lookout NSW) on. 3.91. 1973, The pair was placed im an inflated plastic bag containing stream water, teeds and a rock, Until after oviposition, Larvae were maintained for up to three months i containers of 40 crm diameter ab water temperatures of 4 -25° C. Only larvae tram strearns 34 M.ANSTIS, RA, ALFORD & G, R. GILLESPIE i a a ee wir “i Vig 1A. Mup of eastern half of NSW showing distribution of Literia boorolvmgensts (shaded area = AM records: * = localities referred tin present paper). |, Back Creek, Point Lookout 30° 29' 20" 8. 152° 20° 48" E. 2. Serpentine Creek, Point Lookout 30° 28! 26" 8, 152° 19 26" B39, Near Tamworth 317 139° dO" 8. 150° (0 30" G4, tsaies Creek. Timor Caves 31°41 40'S, 151-07! 20" E44 Goobarragandra River 35° 24°02" §. 148" 26° 01" B. Populations no longer exist al loculities | a 2 (see tex), Bo Map of eastern hall of NSW showing distribution of Liria desuenet (shaded area = AM revords: #= localities referred to in present paper). 1, Quriimbalt Creek. 33° 19°45" 8, 151° 21' 38" B. 2, Near Palmadule AS (WATS, (St 20) 42" 3, Darkes Forest 34° 14" 02" 8, 190° 54 00" EL 4. Durkey Creek near Singleton 32" 44) 36" 5. 150" 50° 32" B.S. Coco Creck BE. of Cupertee 33° 08° 15" S$, 150° Ue 40" E. 6. Mongarlowe River, Braidwood 48° 29) 42"'S, 149° 57° 11" FO 7, Dingo Creek, Winghaty 31° 49° S50" 8. 152" E800" E, 8. Berrico 32° 04) 00" S, 151" 19 79" 6.9. Tir Creek. Bulga State Forest 31° 3149" S$. 152708) 21" B. 10. The Busin, Watazan State Forest 33° (6! 20" S, IST TP 4" B, LL. Bowong Creek 36° 21) 30" 5, 148° 2215" b, 12 Cotter River 39° 35'27"S, 148° 49! 0" BE. 13, Goodradighy River 35° 25' 0S" S, 48° 44" 40" E. The MacDonald River is shown on both maps. where L. leyvewr does hot ocear (in the region of the type Incality) were inchided in the study. Lérerie lesueurt laryacc- NSW localities: AM RLIOTOL - Glen Davis, 119099 - Ourimbah: 119104 - Allens Creek Picton: 119106, |19107 - Dingo Creek near Winghun; (IOS = Berrico Creek, 2 km LE, of Bertico; AYAS - Witiuean State Forest; A724 - Tirnill Creek (collection of M.A.) Tadpoles were reared Lo metamorphosis to confirm ideouty, Embryos and hirvae were measured with vermier callipers und an ocular micrometer attached fod Wild M5 stereoscopic Microscope, Larvae were observed if the Natural environment, then examined while anuesthetized with chlorbutanol, The staging system of Gosner (1960) was used, For L, hooroolmivensis, embryos studied included stapes 2 (n= 11), 8 (n= 15), 17- be (n=9), 19-21 (n=9), 22 & 25 (= 8). hirval stages included 27 (n= 1), 28 (4 = 4). 31 (n= 2), 32 (n = 1), 33 (m= 3). 34 (n = 4), 35 (n = 3), 36( n =4), 37 (n=2), 38 (n= 1), 40 (= 1), 41 () = 6) and inclamorphs al stipe 40 (n= 10). For L. fesueurt, larval stiges included 25 (n= 3), 27 (n= 1), 41 (n= 2), 32 (n = 1 34 (n= 1). 35 (n = 3), 30 (= 2). 37 (n =4). 38 in = 1), 39 (9 = 2), 4h (n= 2) and metamorpliy at stage 46 (n= 7). Ulustrations were made with the aid of drawing tube attached tu the microscope, The following abbreviations lor all morphometric variables are taken fram Anstis (1994), Lureral view: TL = total length, BL = body lenge BD = body depth: BTM = depth of tail musculature atbase of tails TD = maximum tail depth; DP =depth of dorsal fin (in tine with TD): VF = depth of ventritl ii Gu bine with TD); TM = depth of tail musculature (in line with TD); SS = snout to spiracte; S-E = np of snout lo anterior rim of eye: S-N = tp of snout to anterior ri of nutis; ED = diameter of eye. Dorsal view: BW = body width, EBW = body width at level of eyes; BTMW = width of tail musculature dt base of tail; 1O = interorbital spun, EN = edge ol eye to edge of haris; IN = internurial span. Ventral view: MW = maximum width of oral disc, The ubove morphometric variables. were logy, ronstormed prior to statistical aualysis, A principal components analysis (PCA) on the coviiriince matrix of these values was used to reduce the BREEDING BIOLOGY L. BOORQOLONGENSIS & (. LESUEURI 35 dimensionality of the data set and remove the effects of overall size (which is extracted ds the lirst component (Marcus 1990). The scores of each unimal on the second and third component (representing Shape variables) were plotied in a hiplot (Digby & Kempton 1987) in which the sealed couffivients of each variable on the second and third PC axes. were overlaid on the data. Examining this plot enabled us to determine whether the species differed in shape, which size-independent shape variable was most important in this discrimination and io postulate which of the original variables contributed most to the score on. that yartuble, We then calculated the slopes and intercepts of the relationshipr between log,,-transformed beady length and wl other Jog) )-transformed morphometric variables for each species. We used separate analyses of covariance (ANCOVAs) for each morphometne varible 10 determine whether the slopes and inlercepts of the regressions differed significantly belween species, Results Distribunon/hahirat Lilorin boorovlongensis A map of the eastern half of NSW. shows the general distribution of this species. as determined from specimens cegistered in the Australian Museu and the localities referred to in the present study (1- 5. Vis, PA), Hubrtats studied Were permanent Mowing seams running through wet or dry selerophyl| forest. or through semi-cleared grazing land in basalt Or granite range country (altitude range 450-1340 in), Field observations On this Species were cated out annually in the summer breeding time of December/anuary (1965-1974) at Back Creek, Point Tookout NSW (loculuy 1 Fig. PAL 30° 29° 29" 8, 152" 20) 38" EB), Serpentine Creek Clacality 2 Pie LA, 40" 28) 2608, (52° 19° 26" Ey and other neurby sirens al Potnl Lookout NSW (1250-134() om). {uring this period of nine years, gumerous tdpoles mehuniorphs and adults were readily lounct in the severil Streams inthe Pott Lookout and Arpaidialy rewion. A survey ol these same streams in December 190d yeveuled No adults or larvae. The only southern loeality. surveyed was Goubarragandeat River (Ingality 5 Bis, TA, 38° 2 O2" 5, 148° 2a’ O01" Ey where 13 adults were {once on Oxi 1996 along a BOO Mm stretch of the river. Litavia lesireiurt A nmap of the waster hall of NSW shows the eeneral distribution of this species as determined from speelmens registered in the Australian Museum, and a sample of localities studied trom 1972-1996 (Pius. TB, 1-13), Habitits. varied: from flowing streams to large dams in sanestone, melunorphic or granite country in rain forest, dry yeleeophyll forest or heath land (40-| 1000 m), ‘The frog was encountered 1h fairly low numbers in recent surveys in most streams in NE Victoria apd was absent from several sireams Where it would be expected 1a occur, on the hasis of habitat and focal distribution. Surveys in the Kosciusko National Park in 1996 (Hunter & Gillespic unpub.) found the species to be present inoply 15 of 40 likely streams. Behavionr Literig hooroneangensis During summer months in the 1960s-70s, adults were observed bisking if the su on exposed basalt rocks in mid-stream and frequently (ree or more individoals were found onder the same rock Within/beside the stream, ut the sorther Back Creek and wt Serpentine Creek. When disturbed. they immediately leapt under the flowing water Males were observed calling at night while suung on exposed rocks in shallow, flowing seclions of the stream. Six females (four gravid) and seven males (six with nuptial pads) were found by day under stones on pebble banks at the southern Goobarragundra River on 3) November 1996, Numerous Jarvac and metamorphs from stages 25~ 46 were observed at the two northern loculilies throughout December/January (1965-74). Tadpoles were commonly [ound on the substrite anonyse rocks. in shallow flowing sechons of stream including runs and riffles, and in shallow. slowly flowing inlets at the sides of the seam. The tadpoles pussessed at yiuimber of features typical of species inbubiting the lotic enyironment, including a suctorial oral dise fully surrounded by fhree or more rows Of papillite, a more streamlined hody form (especially the snout), a dicker tail musculature ung relatively shallow fajs, They were observed adhering to rocks with their suglorml mouths, tails bending tp the direction of water flow, Literia desueurt During winter, adults were found on ridve-lops in forest country away from streams. Adulis were ustully assecrited with [lowing streams. but alse bred ia completely isolated streamside pools ISPs) stots in bedrock shelwes. where the water was sul Males were observed calling beside dums in open forest near Ourimbah NSW (locality | Pig. 1B, 34" 19°45" 8, 191° 23' 38" B). Eggs were found laid in the stream on the edge oF rons in Slow water in conneeicd still pools, arin ISPs. At Bogong Creek (locality 11 Pry. 1B. 36° 21 30" S) 148° 1215" haa series of perched ISPs was consistently selected as breeding sites by several Fermales froin 1994 ~ 1996. In the first vear all but wne ladpole from two mir] M. ANSTIS. ROA ALPORD & G.R. CALLLSPIE cluiches survived, in the second year the pools dria up aud in the third year, newly-hatehed tadpoles could not be found after a tigh spring fload, No tadpoles have been found in the stream iiself in any OF (hese years, Ip Streams or fivers. Gidpoles were most commonly fuund on the substrate in shallow, slowly Mlowime sections of streams, segregated back waters wit reduced flow and perched rock pools or isolated pools Git times stagnant) beside the stream. They were yery agile when disturbed, capable at’ fast movement darting under rocks or leat fitter. As ind. hoomelongensiy, hey possessed a suctorial mouth, shallow fins, thick (ail musculature and steeambined hocly Oviposition and embryonic development Litaria hoaraoloneensis ‘The mean ege complement of four gravid females examined at Goobarragandra River near Tumut NSW was }S19 (range 1292-1784). A pair of frogs. foond in amplexas at Serpentine Creek on 3.x), 1973 was first observed at O740 he sitting in sunlight on an expased rock in a shallow inlet pool near the edge of the stream. After (O min they bad moved to a rock closer lo mid-streany ina shallow, Plowing seetion. The frogs were then collected ane phiced in i plastic bag, and at (300 ha single. compact exe muss was found partly adhering Co suspended veyenbon within the bag To avoid disturbance during development. eggs Were not counted. Embryos were al stage 2 when a Sample was first preserved at 1300 h. three or four hours after ees were lanl. There were two layers of jelly surrounding The embryo. In live emnbryas the animal pole was dark yrey aind the vevetal pole grey, The same preserved embryos examined in 1996, had a brown animal pole mening lo cream over the vegelal pole, A sertes of 1] preserved embryos at stage 2 ull had a diimeter of 146 mm. Approximately nine hours fier depostion they were at stige & and 15 embryos had a meun diameter of 15 mm_ The blastomeres were nore evenly divided tn the antinal pole than in the vegetal pole. Afier 36 h embryos were a stages |} and 12 and alter 56 h most were at stage | Stages 17-18 (lail-bud),. were reavhed after 67 by, Nine specimens at these stages ranged from 7.43- 3.36 Tan (mewn 2.96). A typival stage 17 embryo (Hig. 2A), had a rounded snout, prom ipent V-shaped ddhesive organs stomedaeal pity small gill-plate bulge; indistinct! pronephrie swelling: slight optic bulge: indistinct nurial pit slightly delineated with pigment, und rudimentary fins along the tail bud, Live embryos at stage 19 were light grey above with i pale grey yolk sac. Some embryos burst through the capsules duping le Mosculiir response in stage (8S but most did not begin hatching vad! stage 20, An embryo al stage 20 Measuring 5,76 mm (Pig. 2B) is deseribed us follows:- snout rounded; two pairs af externul gills - two branches on upper pair and 4-5 branches on lower; Optic and narial regions partly outlined with small crescent of pigment: adhesive organs well divided — remnant of V-shape below each: numerous fine museular tidges alone tail musculature: stontodaeal pit and fins both deepening, Dorsuni grey, yolk sac pale grey in live embryos, und pale brown/ycllow brown in preserved specimens, Live embryos at stages 20 und 2) were grey above with a lighter grey yolk suc and translucent grey fins. Nine specimens at these stitges ranged from 4.82 - 5.07 yim (ean 5.43), Mitchlings adhered strongly ty the jelly mass. The external gdls were visible macroscopically and when fully developed by stage 22 extended about 2/3 the length of the yolk sac, Ereht embryos ut early stage 25 ranged from 9.23 957 im Cea 846 ) dorsal pigmentation of larval Stages developing; dorsal edge of tail museularure finely edged with melanophores: fins. external body wall (ind venter clear (in preservative); spiracle and vent lube both functional: jaw sheaths pigmented, hut as yet without shape of older larvae; keraunisation incomplete, Tooth rows almosi complete except for 3nd lower row. which was beginning fo develop m two more advanced Iprvac at suige 25, measuring 11.5 and 12.15 nin. respectively. Litorta lesnenrt Most clumps of eggs observed were loosely udbering 10 rock substitte but some alse adtiered bo bottom sediments or Sedwes. There was no indication ——— B Fin. 2 Embreos of Liforne boerrploieensis A, Retnacee from capsule. stave (7, 3,24 nam, GB, Just hatched -stigee 20, 5.76 mm, Soule bur = finn. Stuwes ace (rom Gosner ( (960) BREEDING BIOLOGY L. BOOROOLONGENSIS & L. LESUEURI MY of nest-site excavation, as in the north-Queensland form, and 20% of clutches observed were deposued under, or partly under a smal! rock on the bedrock of the stream. Eygs were laid in a water depth of {0-20 cm, Clutch sizes of 14 egg masses examined in 1995- 1996 ranged from 610-3564 (mean 1878). The mean diameter of 10 eggs at stiges 43 (Gosner 1960) from Guoodradighy River (locality 13 Pig, 1B, 35° 25°05" 5S, 148° 44" 40" E, was 17 mm. No embryos beyond this stage were ayailuble for study. hurvee Analysis of covariance showed that the two species follow very similar lrajectories through larval body sives und stiges (tests lor differences i slope and intercept both p >> 0.05). The relationships between developmental stage and body size for both species are Wustraled in Fig. 3, which makes it clear chi jhey cannot be distinguished on this basis. The results of the principal components analyses (PCA) ure presented in Big. 4Al with the contributions of cach variable to the second and third vivenvectors Shown Ww Table f, The first (general size) principal component accounted for 87% of the variauon ih the data set. 'Uhe seeond and third (shape) components accounted for 4.4 and 4.7%. respectively. Despite the relatively small proportions of the overall variance accounted for by these components, ib ts clear from Vig, 4A, that in combination they diseriminate very effectively between the two species, indicating that the species 41 ss ! be 39 37 35 33 Mu Gosner Stage 29 27 # A L. booroolongensis 35 > Bai. lesueuri 5 10 15 20 Body Length (mm) Fig. 3. Gosner (1960) stives plotted agwinst body length for Litovia hooradloigensis and Lirarki lesneurt. Line represents pooled regression ol stage on length. y = 1.32 6+ 15.77. = 0.84, p< 0,000, do differ strongly in’ shape. Most of the diseriminatary power resides in Component 2, This means that the niorphometric variables responsible for the eigenvector loadings which are most closely aligned with axis 2 of the PCA in the biplot, should be most important in discriminating bebween the shapes of the two species. Tank 1 Reyresston of logy of morphomeric variables on leg), eo] hedy length: foreach species, with pevalies for vignificance af diflerences in slope and tntercept berween species fron an ANCOVA, and eigenvectors far the second and third principal components of lov jy ansfarned morphometric veriibles, Lo bowen gets Lh, leaueuri ANCOVA pevalues Principal Component Eigenveelors Dependent Variable Intercept Slope Intereept Slope Hileree pt Slope pC? PCS TL (2588 11077 (280 L077 H.0012 HOO UWnss 19 HPISRG BL - - - (03202 “04520 BW OTR ads 1623 DY19S (3654 WARIO Ud1304 NAME BD 0,123 (8400 (2574 (), 0625 0,000) (0327 27523 (LOSSSA BW “16 {hava4 “1512 (ASRGS UW (8t45 AY OS 2 ahs? WW) 04061 (8095 ASAT bleed? L000) PhO (HES (1245000 IN -U) a8UM (5303 9237 L004 OAS 0001 AROLO72 0.24724 LN A) hls4 (7780 9479 | Wo77 (1942 (0002 0.07379 UOTSHAT WM Ww 15500 1.795] 10958 1381 (5247 (00M ALUGITS 1), 4u54u BRIM 9476 1.2425 O.6718 |QQ07 O85) UA275 (i rset (423404 Th ANIISI 77a0 04308 (MMe OSE (0001 () 22844 Qtaj2o Db 04379 WATT -O,4984 ).2369 0.0014 1.0002 U260154 1-454 T™ (7799 04276 89%) hos7a O.U569 3348 (119224 OO 16840) ve (2827 ()4277 U9264 1udos (),7292 (0003 (20800) (34620 SS OLA O 449 “U1 894 9647 0.0016 1), 7200 -W0S245 (UALS Eb -],22355 [2728 (727) OSTA OAWIOT HOU W2AIle AY AT394 Vw 5 Lot “O19 8719 0.000) 0,006) “47415 0, OG78 SE 0.4276 TLOTSo -OA735 O.YTOS ONO NYIas “UO I6108 0.02676 SN O.N832 (RATS 7280 ORO72 O.0D01 1.5980, O,554d0 O11531 aK M. ANSTIS. R.A. ALBORD & ©. R. GILLESPIE 0.3 ray Litoria hoaraclongensis yr BC aLitoria /esueuri 0.2 2 oA ° a w o° oO a 0.0 -0.1 -0.2 0.3 -0,2 -0.1 0.0 04 0.2 0.3 PC2 Score N Pip + A, Results of Principal Component: Analyses. Vectors aire proportional to foadings of cach original morphometric variable in caleakiting scores on edieh PC axis. Bo Plots of the six (host Tiportant morplenmetric variables suggested by the PCA agaist body length Figure 4B presents plots of the six most inportant vuriubles suggested by the PCA against body length. These plots show that the species differ in these charactenstics, in the direction indicated by the PCA; L. heoravlongensis has relatively greater snout-naris length. mouth width, and snout-cye distance and relatively lower body depth, dorsal fin depth und eye diameter than £, fesneqad does at each body length, Literia bearvalongensis A composite description of wdpoles at stages 25- 39 is given. Using some anaesthetized specimens still showing colour in dite. A tidpole at stage 37 (36.2 mm) fron’ the egy mass laid at Serpentine Creek (locality 2 Pig. }A) is shown in Fig. 5 AVC. Dorsal; body ovoid, widest ucross branchial refion: snout broadly rounded: eyes corse-laterdl. positioned alinost '/3 along body length from cp ot snout, nares opening antero-literally: body wall uniforn rusty-brown with some darker mottling: darker browi band aeross urastylar region, brawt body colour continues along tal musculature as two longitudinal stripes, becoming lighter towards tail tip; young tadpoles ul stages 25-26 dark brown wilh irregular light grey band just anterior to darker urostylar tegion: Fibs iereasingly pigmented alter stave 34. Lateral: body streamlined, snout rounded, elongate; spiricle sinistral, moderately long and broad, aa ta an & 3 & Snout-Naris Length (mm) a0 E E20 Eu E = S 2 = 26 © = ie 53 2 Eel a op Sy 4 4 a ow of} E ' 3 = 3 5 = 6 E s a ¢ A FA a 5 4 o 3 te =] . . = “ “ Gedy Lange (rmeny Body Lengin (awn) B A Liroria nooreorongensie bitoni tomumurt tapering slightly from origin to postero-dorsal opening; oral disc directed ventrally: vent tube dextrak: body brown with part of golden ventral sheen visible, particularly over branchial region; itis golden: (ail musculature thick anteriorly, uniform brown with some darker patches, main antertor blood vessel and crevices between muscular ridges outlined with pigment: fins relatively shallow, dorsal lin rises eradually (or more acutely) to greatest depth at mid-peint of tail, or just posterior/anterior to i. yenural fin inereases slightly in depth in posterior half, but generally shallower than dorsal fin; fine hetwork of melanophore clusters (raced over Vaseuhie system on dorsal fi) and posterior’ hall of ventral tin (denser beyond stage 30); tal Up rounded, Venlral; venter with almost uniform copper/gokl sheen of iridophores, or pateliy sheen with darker areas showing through in between patches. Branchial region densely covered with — copper/golil Indopliores, Oral dise (Pig. 6B) oral dise wide, directed ventrally, band of papillae surrounding entice convoluted margin; 2 - 3 rows wound anterior, 4 - 6 uround dateral and 3 - 4 around postertor margins inner papitie on posterior murgin larger and) more widely spaced: papillae diminish iy size and inercase in OuMber through to outeemost rows two complete unterior und three complete posterior rows of labial teeth, all equal in length; juw sheaths moderitely massive, quite heavily keratinised, upper sheath with BREEDING BIOLOGY L. BOOROOLONGENSIS & L. LESUEURI 39 im, ot tier ~ , | al Dp Pies S. Laryue of Liferie booreoloneensiy und Literia leswenri, each al stage 47. AL Lo boc madongetyis. lateral view. Bob. leaneru7, lateral view, C. Le beorevlamernsts, dorsal view, D, L. fesnearr dorsal view, Seule burs = | nim. Stiging systeni oF Gosner (1960), Av M ANSTIS, RAL ALFORD & GR. GILLESPHB a tt B byt Oral discs of Litera byerectoneensis and hirortay leaueni A. Le lestetet Coeulilyy 10, Tig 1B, Stige 37), BLL, hooroolengertily Coeality 2. Pig, LAL Shige 34) Seale bars =) min. cental potch and small Waderlying keratinised ledge: inner mining OF jaw sheaths serrated, (edge partly warn in some individuals and dows not appear before Stage 28 in spechineus examined. Colour tm preserved specimens (composite deseription, stives 33-39): pigment slightly darker over abdomen, cramil, vetlebral and post-narial regions, Lut museulature evenly pigmented (dorsal view); abdomen opuqne wirh fine melihophores over snout and branchial region: dermnil body wall clear-with uw few seattered clusters of melonophores; iris black (lileral view); fine dense layer ol melunoplores over heart. abdomen and just anterior tO caeh gill; intestine visible; rail muscularice ahd liipbs unpigmented (ventral view). Liraria lesueuri Composite deseriptiom, live specimens. saytes 41 - 34 A radpote ar sie 37 (33-9 mm) from Watayain State Forest (loculuy LO Fig. 1B) is shown ia Pig, 3 B,D. Dersal; body ova, Widest aross abdominal region: snout rounded; eyes dorso-luteral ynd pusitaned less than '/4 wong body length from tip oF snout, nares ppen uncerolateraiiy: abdooinal, cranial. post narial regions und eyes appear darker below dermnl layer of golden Wophores covering most OF dorsum. except for urostylar region. where ubsence of widophorcs creates hroud, Uniform or broken darker bund; body wall sparsely flecked with simall chisters Ot metinophores in seme Specimens; soe brows palches over hal musculature; golden indophores denser aver Gills, snout, posterior rim of iris and just interior to urostylar tegion; young Wdpales. at stages 75-26 with darker dorsum (ess dridephares). and layer of cOnLUSHNg bund of pale grey pigment Just unlorior 16 darker upoptylar tegtons tail musculature sandy gal, or with Slight salmon rage, dark brown: patules seutiered ulang length of taik especiaily vlone wutertar dah limbs iicreasingly premented trom stage 34. Lider: body oveid: stout rouided: spiracle sinistral. broad at origin tapering lowards postero- Jorsal opemiug; oral dise direeted vertrally: vent tube dextrl; snout appears golden, layer oF copper/golil iridophores oVer majority of iris and lower part ot abdomen, chanps of tridophores outline lower cdee ofeach oil, cantinuing over venter: Gail musculature fairly thick. anteriorly, some seattered melaiuphare closiers overall but anterior lower edge; maim blood veasels, creviees between nmudeulir ridgys pigmented? fins relatively shallow to moderuely so. clear. with Joint wolden hue. line Seatleresd melunophore clusters over dorsal lin ane some dusky pigment over ventral fine-venation unpigmented brit Visible: dorsal fin rises gradually (or more deulely) te preatest depth anterior to, or dt, mid-pome ol tail, nui Lip narrowly rounded, Ventral opaque copper/gold sheen over abdomen, heart and some clusters of iridophores over stdes vl each gill, anterior half of venter otherwise clear hinbs-and tal oseulature unpiemented. Oni dise (Pig, GA. oral dise ventral in direenon fuirly wide: bund of pupiliae surrounding entue muraint 239 rews fine papillae around anterior, up ty six around lateral and 3-4 around posterior margin: inpermost row oF papilhie around the posterior margin very slightly larger; bwo complete anterior and three complete posterior labial touth rows, equil in Jength (PL row aw hte sherter in somrety jaw BREEDING BIOLOGY L. BOOROULUNGENSIS & Lo LESUEUWRI “y sheaths moderately missive, wath heratinisation greater on upper sheath; central notch On Upper Sheath, ti some individuals there is a small Underling keratinised ledge visible, similiur to tit shown in Fis. 6B for Le boervelongersis but less prominent, and absent or worn in ofhers; diner munis of jaw sheaths serrated. Coloar af preserved larvae (composite deseription stiges 3)-41)° body colour untlorn) brown uve abdomen, cranial, vertebral and: post-nacial regions. lighter elsewhere; dermal body wall clear, spirsely Nevked with small clusters of melanophores in some specimens. some brown patches aver ntl museulite (dorsal view); abdomen opaque, fine layer OF mekinophores over gill region, patehy in some specimens. becoming spurser over snouly pris hlaeke fins mostly elewe venauun uopigmented bu Visible (lateral view): fine layer of melanophores urpund sides Of abdoinen, (ntestine Visible, rest of venter clear: limbs and tal musculature unpigmented (ventral view } Metumerpheasts Lierice bern eeayin Dyes laid on 3.411973 were first metunorphosing from I8A1974, alter a larval life span of 21/4 months. Some were sill metimorphosmy by 14... 1974, Numerous metamorphs were observed aoninidly in Devember/ianuary at Back Creek and Serpentine Creck during 1965-74. No tadpoles or ielanorphs were observed a late auton at Back Creek on 1oy.1973. Ten newly mectumorphosed frogs trom Serpentine Creck (stage 46, (97+) ranged from $4.0 - 17.5 min (mean 15,23). Literia lesueurt Known observation dites lor metamorphosis ure 7 Apnl 1974 at Coco Creek (locality 5 Fig, 1B, 33° 08! 15S, (507 06'40" By god 1s Jamnary 1977 at Dingo Creek (locality 7 Pig. 1B, 31° 49°50", 152° [8 On" Ey. Tava lite spun is not known. Seven pewly Metunodphosed Frogs (staye 46) trom Dinga Creek ranwed fropy E17 = 15.0 anim (eat 14,94). Discussion Papnlation declines, hatitat While the present knowin distribution lor Litre Hoormlaiveiwes dong the mount range COU in NSWoos. tram the Queensland border to) the Viclooun border (he disinbutot and agius of Litarin lest aeeds clanfieation, with a likelilgod of omy of more species beth involved (Moore Oli Heatwole eo uf. (1995) report J, haoroalonieusis as “widespread” jo the New Englind region but the records they provide are only those of existing Museum specimens spain “a degree ol perjod from early 20th century ta 1990". Furthermore. extensive surveys for regional conservation planning in north-eastern NSW undertaken since 1991 through the casiern escarpment forests around Tenterfield, Armidale and Glen Innes by NSW National Parks & Wildlile Service (NP&WS), have failed ty locate this species (H. Hines, Queensland Departrnent of Environment & Ueritage pers. comm. 1997), Field observations at Guy Fuwkes River and nearby well-known loealities where the species was similarly abundant prior to 1980, have imdicuted few adults ever the past 17 years: The species could not be found in recent surveys at Ebor (type locality) god a hirge number of rivers i the area (M- Mahony. University of Newcastle pers. commit. 1997). Intense searches during 1995-96 in this area were who to no aya CR. Lars, University of New Boglind pers. comm. 1907). Surveys since (992 jn upland forests such as Glen Innes, Walcha, Mt Royal. Dorrige, Teatertietd. Coolah Tops and south to Tumul/Turbarurn ba (turgcting frogs in areas where £. hocuolimgenyts is likely to oceur), also resulted i hone al this Species being found (bh Lemekert, State Forests NSW pers. comin. 1997), Surveys in the southern region near Tumul NSW by Hunter & Gillespie (unpub. located only (3 frogs an 30 November 1996, along an BOO in stretch of the Goobarragandra River. Extensive searehies along the Tumut River. where L. beervaloneensiy had been recorded in the 1940s and in F987 failed to locate jany evidence of the Species (Hunter & Gillespie unpub). Examination of over L.Q00) specyinens of 1, honmotongensis in the Australian Museum revealed that uuly five specimens bave been vollected since HOO: One at Wombeyan Caves (34" 19S. 149° Su! EB) two at Canimbula, Blue Mountains (33° 41'S, 150° 12°F), one at Cos River (43° 28 8, 150704) FE} ant ane at the Abercrombre River, Governors Fat (a4) OF S$. 149° 41 EB), where Lo lesen are sympatric (IK, SmalloSydney University pers. com. 1997), Aceordingly, the spevies has heen Wominuted for inclusion in Part | of Sehedule | Cendtingered Species) OF the Threatened Species Conservation Act 1995 (TSC), More studies on current population numbers of £. badrohmeenscs over a mich broader range of Ts distribution are required to assess fucther the curred alaius Of this species, mm the Ligh) oF very significant frow deelipes oy the northern ranges between latiudes 31° 30° S and 24° 30S (north of the Macdonwld River ~ Pig. TAI, Fron present indications. the species uppears to have disappeared at ibe very lease fron its type locality and all known lowalites an the AcmidaleyGuvrwPornt Lookout 42 M. ANSTIS, ROA. ALFORD & G. ROGILLESPIL: regions where itonce was abundant. The only known northern area where adults have been observed ts locality 3 (Fig. 1A) near Tamworth (1994) in streams aha) altitude of 450-500 im (M, Mahony, Liniversity of Newcastle pers, comm. 1997), much lower than the Armidale/Point Lookout region. Further field surveys. will beed to be undertaken to verify the frog's continued existence here, but if so, this may relate to the data being gathered on frogs in perth Queensland whieh indicate that most declines are umongs| upland riparian species uboye 400 m (Richards ef al. 1993), Some observations in southernmost localities. near Turmul in 1996 also indicate a distinet drop in frag sightings (Hunter & Gillespie unpub.) sugvesting the species ts ikely to be endangered throughout is general distribution. Litorta lesueuri This species has a broader current distribution thin J. hooroatargensis. Barker etal. (1995) indicate thut UL desist LWO Species are currently included under L- lesnenri, “one confined (o northenstern Queensland une (he other extending down the couwst as fir is Vietoria.’ Comparative popakition studies on L. leven? are heeded, especially where the minges of the twa Speeres overlap, to determine whether this spectes 1s also undergoing a decline in certain localities. Surveys by Hunter & Gillespie in 1996 im Kosciusko National Park NSW showed that the species was present ip only 1S our of 40 likely streams, which may indicate a possible decline, While 4, boonmlongensts is restricted to Towing streants generally above about 400 in ZL. deswenry can breed in siréums. Strewmside pools und even shims. from 1100 4) (River Murtay and Snowy Kiver, ML Kosciusko NSW) to 40 m (Ourmbuh NSW), This ereuer versuoliny of altitude und breeding site selocnon may lave helped more populivions renin Thun for lL. boeroealorueryin Behaviour Neither Le feurvelengemsis mor L, lesueur® bas u yoru sae and each produces a sofl. low call obi serves OF short. repeated notes. Adulis are similar Nuiphalowiedily. beth breed in usseeiatian with Hlowrny sireains, ud have a similar tialjate winelr ps Adluple do fhe lotic environment Liteiia bevntaledtinensis agorevates Uiiler stones i hivee nuinbers dueine wire A aragp op Tu mutivadiits, was founel unger (he same stone beste Enis Creek NSW (locity + Fig, LA, Fle 440s. oF 2" Bion, bhai eet by fe Parke Populariuns here were Jarge, wath tom ah 15) frags ball sive clusses ohserved a 19 ei 1970 durin the day, under stones th tie eteek bed Males called during the day and night. The aggregation of L. hddredlongensis under rocks both during colder months and also in the breeding season during spriiig/summer, has. nol been reported for other Australian bylid species, Winter aggregation has been reported for Litaria svubolandulasa (Tyler & Anstis. 1983). (Tyler & Anstis 1975) and for /. péarseniand (Copland, 1960) (McDonald. & Davies 1990), but neither species has the same habit of commonly gathering in groups under rocks in the stream environment during its breeding season. Literia fesueuri adults, found on ridge-tops in forest uway from the stream during winter, are not known lo aggreaate. Lileria beorodlongensix is simular to the stream-dwelling 2. spencer’ (Spencer, 1901) (Watson ef af. 1991) in its diurnal behaviour. ofien basking in the sun on very warm rocks in mid- stream. Amplexus and oviposition also occurred diurnally in one obseryation for L. hoarrolongensts Lireria deyuteurt has been observed busking in sun. but not as frequently as bas been observed lor J. spencer) or L, houradlonze ysis. Ovipasition/embryes The egg inass.of L. booreolenuens’s found partly udhering 10 suspended vegetation in a plasie bag does not give a inte indication OF the mode af deposition in the natural environment, as the adults had separated and were swimming vigorously within the hag, constuntly disturbing the egy muss. Barker er af (1995) reported that the ege mass ot L, fooroclungensiy is deposited “among rocks. While the northeastern Queensland form of /. Jesuew lays caps ina single clump of two er more Tavers deposited in circular nests excavated in sund at the sides Of Mowing streanis. (Richards Ae Alton 1997) ees Masses ure so deposited among mocks ty streams where (he substrate is nob sand, The number af eyes mone clutch was estimated hy Richards & Alford to be 1200. Two other cliitches counted by Ss. Richards and M.A. ut Elphinstone Creek norith-Qld On Alvi.l996, numbered 1.738 and 1,674, respectively. Referring to southern populations, Barker ef al. (1895) state (hat several Huadred esp ire Jepositec in a solid) gelutious etunip whieh idheres te submerged recks or rhe hatam sediments We hive seen 2-3 inasses Juid beside evaeh athe, partly comneei oe. Whilst cee deposition marily Queenslind form apf. desterad nay he exe hdtest Nests ji sand bats, eggs af the southern lori have ony been observed i mchy areas. wher such excivalion is mgt possible As [rows have been observed urauud dais (og, ub deealigy 2 Fig. bt Where Ho nicks Were present. ane cudpales Found i sundy atrediows (locubitnes 1. 3. 5. 1 Pige FBS, further studs of the mode of deposition mi the Somuliont for silts ot the BREEDING BIOLOGY L, BOOROQLONGENSIS & L. LESUEURI Wa References is needed (9 determine whether or not nest excavation may occur al suilable sites. The mean egg complement of L. fesmear? was ereater than in ZL. hooroolongensix. Both species have a compact. gelatinous egg mass. similar in form to that of other streani-breeding liylid foes such as L. pecrsentiahd (MeDonald & Davies 1990), L. wibeglandulosa (Anstis & Litdejohn 1996), L. genimeculata (Horst, 1883) (as L, encnemis, Davies 1989), andl. spencer. The egg mass of L, fesuewrt is loosely attached to the substrate, while those of 4. ysubelandulosa, L. spenverd and Lo pearyoniana adhere more strongly. Larvde Although larvae of bouh species are the sane size atany given stage (Pig. 3) and appear superficially similar they differ considerably in shape. he tadpole of L. hoeroelongensiy has a relatively larger, wider ofal dise afd broader, more elongated, streamlined snout than 4. lesuear? (Table 1. Pigs 4.5). The distances from the tip of the snout to the anterior rim of the eye and to the anterior rim of the naris are relatively greater in L. booreclongensiy and the body depth, dorsal fia dept and eye diamerer are IMARANTILLL Ch. Gihonsmin Gok FesunG, S$. (106) Observilions Of oviposition sites of the spotted tree frog Litt spencert, “In vhe Spettieht 2 fd td less. The eyes are positioned more medially and directed a litte more dorsally. The presence of the small keratinised ledge underneath (he central noich of the upper jaw sheath was voted in populations of both species, but nor found as consistently qur us prominently in Z. desveut. This feature has not been recorded for larvae of any other Australian lrogs. The jaw sheaths ure commonly more heavily keratinised in L. bvorvoloneensis, but this feature may be variable umongst populations, us has been observed io L. desuenr?. add ip northern and southern populations of the tadpole of L. verreawsi (Dumeril, 1853) (Anstis 1976), Acknowledgments The Australian Museum is acknowledged for the loan of specimens. A grunt to M.A. from the Peter Rankin Trust Fund in 1996 has assisted this study, and is gratetully acknowledged, Valuable Observations on population declines of F. hoeredlongensix were provided by H. Hines. P. Webber, K. Harris, M. Mahony, F Lemekert, J. Reesei, K. Thumm and K. Small, Constructive comments on the munuscripl were given by $. Richards, FP. Parker, M. Mahony. G. Watson and M, Davies. References AwsTis, M. (J974) An introduetion ja the study of Austratian tadpoles. Herpetfaniad 7, 914, (1976) Breeding biology and birval development Ol Liferie verreauee (Anurd, Flylidae), Trans RO Sie §- Anish 100, 193-202, (1994) The lirval deyelopnient of Lirarie brevipalmete (Anura, Mylidae), Mem, Old Mis, 37, 1-4, — & Lerrecious. M_1.11996) ‘Phe breeding biolagy OY Litoriee Subghindutosa ind Le citrepe (Amur Hylidae). anda re-evaluation of thei!) geoeraphic tistabulion. Quen. ROA Ant 120, 83-99 BarKwie, 1, GRIGG. GC & TYLER. Me J. (1995) 7A Held guide fo Australian frogs (Surrey Beathy & Sones. Chipping Norton NSW), Copnann, SI (1957) Australian ree frogs of the genus Myla. Pree. Loe She. NSW 829-108, Davin. M, LIY89) Developmental bielogy af the Avsttulopupuiin fiylid hon datura ehevenius C(Anure: Hylidhed. frais, RSet. 8 Aas PV TPS 220 & Riiakos, Sod. (TYO) Developmental brolowy obibe Australian hy bd frog Ave cares lav? (Crlinther), Trans RK See 4 Aust, V4, 207-271, Hicry PG. NLR Kean RAL COST) Multivariate wnalysis of ceoloyicu) communities” (Chapman & Chill, Londons. Piercuer. JJ. (1889) Observations on the oviposion and habits of certain Australian hatrachians. Proc. Linn. See. NSW ser, 2. 357-387, Gossnk, BL (1960) A simplified table for shiwing aura embryos and durvae with nutes on tdenuticaion, Herpetelagica VW, V83-190. Hiavrworl, HO DL Ravay. 2. WeiHeR. Po & Wik, G, (1995) Fauna survey of New England. (Vo The Frogs, Mom, Qld Mus, 38, 229 249, Mancous. Lo L990) Traditional tmorphonetrics pp. 77 122 In Roni, FI & Bookstuin, [Musi “Proceedings at the Michigan Morphatogical Workshop, Spee. Pub. no. 27 (Li Mich Mus Zool, Ann Arbor Michi). Martin. A. A Lintneiniis, Mob & Rawirssen. PA (i966) A key to che dius ces oF the Melbourne arcs. and dn addition to the anurin (aun, Wer Nab 83. 312. FAS. MeBosarn, KR, & Poyins, MM. C1990) Morptiotoyy: and higlopy of the Australien tree Woe Liner pencils (Caphind Amur: bhyldiiel Cami. Ro Sues, Aust 04 145-156. Moorn, J A CM6t THe Hogs al caster New South Wales Ball Amen Mies. Net Hivn 21, 149-486, 44 M. ANSTIS, R. A. ALFORD & G. R. GILLESPIE RICHARDS, S. J. & ALFORD, R. A. (1992) Nest construction WATSON, G. F., LITTLEJOHN, M. J., HERO, J.- M. & by an Australian rainforest frog of the Litoria lesueuri ROBERTSON, P. (1991) Conservation status, ecology complex (Anura: Hylidae). Copeia 1992, 1120-1123. and management of the Spotted Tree Frog (Litoria , McDonaLp, K. R. & ALForD, R. A. (1993) spenceri). Tech. Rep. Series 116, Arthur Rylah Declines in populations of Australia’s endemic tropical Inst. Environ. Res., Dept. Cons. Environ., Vic., 40 rainforest frogs. Pacific Conservation Biology 1, 66-77. + vi pp. TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 122, PART 2 A NEW GENUS AND TWO NEW SPECIES OF GALL MIDGE (DIPTERA: CECIDOMYIIDAE) DAMAGING YOUNG BRANCHES OF EUCALYPTUS SPP. IN SOUTH AUSTRALIA By PETER KOLESIK* Summary Kolesik, P. (1998) A new genus and two new species of gall midge (Diptera: Cecidomyiidae) damaging young branches of Eucalyptus spp. in South Australia. Trans. R. Soc. S. Aust. 122(2), 45-53, 29 May, 1998. Two new gall midges are described from galls on young branches of two Eucalyptus species in South Australia and a new genus, Okriomyia, is described to contain them. The new genus belongs to the tribe Asphondyliini and the subtribe Schizomyiina. It differs from other Schizomyiina in the shape of the aedeagus, the solid tooth of the gonostylus and the cerci-like female tenth tergite. Okriomyia schwarzi gen. et. sp. nov. was found on Eucalyptus gracilis and O. flabellidentata sp. nov. on E. cosmophylla. Infested branches fracture at the site of the gall as the trees mature. Males, pupae, and larvae of both species and the female of O. schwarzi are described. The new species differ from each other in the morphology of the male genitalia, the pupal face, and the pupal prothoracic spiracle. A key to the Australian genera of the tribe Asphondyliini is given. Key Words: Gall midge, Cecidomyiidae, Okriomyia schwarzi, Okriomyia flabellidentata, Eucalyptus gracilis, Eucalyptus cosmophylla, South Australia. Transactions af the Reval Seviety of S. Aust (1Y98) 122(2), 45-53 A NEW GENUS AND TWO NEW SPECIES OF GALL MIDGE (DIPTERA: CECIDOMYIIDAE) DAMAGING YOUNG BRANCHES OF EUCALYPTUS SPP. IN SOUTH AUSTRALIA by PETER KOLESIK* Summary Kovesin. P.( 1998) A new genus und two new species of eall midge (Diptera: Cecidomyridae) damaging youny branches of Kucelypris spp. in South Australia, Trans. A, Soe. S. Aust, 122(2). 45-53. 29 May. 1998. ‘Two new gall midges ure deseribed from galls on young branches of two Licalvpius species in South Australia und anew genus. Okrmnwia, is described to contain Mem, The new genus belongs to the ibe Asphoniylinn und the subtribe Schizomyiina. lt differs from other Schizamytina in the shape of the aedeagus, the solid tooth of the vonostylus und the verci-like female tenth tergite, Okriomyta sehwers gen. cl sp. now was found on hawalvprs graciis an O. flabellidentata sp. noy. on E. cosmophylta. Infested branches fracture at the site of the wall as the trees mature, Males. pupae, and larvae of both species and the female of Q) sefnvarc are described. "The new species differ from each other in the morphology of the male genitalia, the pupal face, and the pupal prothoracie spiracle. A key to the Australian tenera of the tribe Asphondyliini is given, Rey Wokns: Gall midue, Cecidamyiidae, Okrionivia seliwarsi Okrionyia flebellidentata. Bucalyyitus grata, Eucuiypray cosmoplivila, South Australia Introduction Nacalypoes, the dominant genus of most Australian woodlinds and forests, hosts a whole suite of gall- lormings inseets, many of them undescribed. The present paper describes two gall midges, new to science. which were found damaging young branches of two eucalypts in South Australia, Galls of Okriomvia schwarei sp. nov. on Eucelyptus eracilis TF Muell, (Pig, 1) were found at two localities: Nadda. in the southern part of South Australia near the Victorian border and Porestville, a suburb south-west ol Adeluide. Galls of O. Mabellidentita sp. nov. on LL cosmophylla P. Muell. (Mig. 2) were found at Cleland Conservation Park, near Adelaide. The newly-described gall midges were found only in moderate abundance. However, heavy infestations could have the potential to impact seriously on the population dynamics of their hosts, since (he infested brinches fracture at the site of the galls as the trees mature Eucalyptus gracilis sa 3 - [2m high shrub or tree distributed through the mallee belt of continental southern Australia. [is an ari zone species usetul for firewood und crosion control and is) highly revarded for honey production (Cunningham e7 al. 1981, Chippendale 1988). Ibis offen used in urban planting. © Department oF Horticulture. Viticuliure ind Oenology Faculty of Nerieultucal diel Nateral Resource Serences, Che University. ut Adeline PME OT Glett Osmond S. Aust. S064. Fig. |, Gall of Okriomvia schwerce sp. ney, on brinch of Arecelypris gracilis, Seale bur = 20 min, young at 1 KOLDSIR nN = Pik, > Gallo Ohkrionvie flabellidentata sp. nov, on youne brinch of Rac yyas cosmaplivila, Seale bar = 20 niet Eivalvprs cosmophylla is a South Australian shrub or tree, usualy 5 LO high, that oceurs from the Mount Lolly Range to the Fleurieu Peninsula and Kangaroo [sland in open shrubland. low. open forest and heathland nearthe sea (Chippendale TO88) Tis Widely dsed in urban planting, The new gall midses do not resemble any Known eens so a new genus has been erected for then. Ohrfomyia becomes Australia’s fourth known gers of the ibe Asphondyliint und the venem endemic to Australia. A key fo the Australian eeperd of Asphondy lint ts given in the present paper, Material and Methods Gls on branches Of Evealvyptus viacilis were Porestville (19477999) abel Naddy (b2eyi T9960), Two, one, Hires. Tour and one galls Irom branches of A. cesmophyila were collected at Clehind and Morintta Conservation Parks 27,41 1992. TS 199F 5S. und 12 P99S. and 23.1 1997, respectively. Ja the labortory the pulls were cut open and the larviie processed i two ways. “Ao small number was preserved in 7O% ethanel AO darter collected at Wind of Sulnzomyiini. ad subtribe consisdnge exclusively of number Was Wansterred ilo tearing pots where Ure larvae dug themselves into wet sand. Pupation took plice i (he sand, Several qales and ternales emerged from the galls from Ay graeiliv, OF the galls collected from E. vesmoplydie adults emerged only from the suinple collected on 23.11.1997 12 mules und ong femules. Emerged adulis were preserved together with their pupal skins i 70% cthaeiel Microscope mounts of the type series were prepared weording to the feehnique outlined by Kolesik (19950). The type series and other material retained in 70% ethanol, together with dried galls, ae deposited inthe South Austithian Muscum, Adelnide {SAMA}, the Australian National Inseet Colleetion, Canberra [ANLC| and the State Herbarium of South Australia, Adehiude [SHUSAJ. Descriptions aid measurements reler to the holotypes and paratypes Terminology of adult tnorpholosy follows That of Gaené (DOS) ): larval terminology Totows that ot Cane (1989) Genus Okriomyia wen. nov. Type species: Gkrioivic schwirel sp. mov. Adtilty Heeul. Fye haces heaasonoid, eye bride H-8 fieets longs medially, Antenna with 12 Thigelloneres, distal ones nol shortened. Plagellomeres eylinadrical. sessile. first ane second not fised. with short sehuw ind hearing low, finely reticulate circumlila. Suape 4s long as wide, pedicel hulfias long as wide, Labella hemispherical, each wilh several setae, Palpus with 4 segments. Theres Wings: Rs joing Coat apex. sliehtly bowed anteriorly, Rs absent. Ry joining C neve mia length, Cu forked, birst Girsomere lacking: vente distal spine, tarsal claws Simple, as long as empodia \hdomen. Tergites | o- & with selie eventy distribuied, forming dense row posteriorly, Sterna | Hot sclerotized, aselose; stermites 2 - 8 willl seuie mM iWO separate areas: wide, anterior field and narrow, posterior band. Pemale abdominal sternite 7 15) % stermile @ Male lemminialis gonocoxite with apico ventral lobe: genostvlus short and wide. with tool in form ot serrate plate no more strongly pigmientea than remuinder: cereus hilobed, deeply divided medially, eniutyinuted posteriorly, with several posterior setoe; parameres. small, sefuses hypoproct WITH posterion Mari coneuve, each faberd lobe with J ~ 2 apteal setae, aedeagus comprising Iwo pits; dorsal part robust, conical, ventrally eovered with selerotived Vili on apical thicek ventral part sinmeerh Thin ie katerul view, shallowly enurgnded apieatly in dorso-yventral view, usetose. Pemule terminalia OVIPastlor short, Peshy, tergunmy 9 and stevia, 9 sclerotizeds terguin LO in form of two hinge, cory NEW Cb like lobes. more selerottzed uinteriorly, evenly selose; verer large, diserele, more selerotiaed: posteriorly, evenly selose: hypoprect small hilabed. cach lobe with apical sen. Pipi Aen hors strongly piemented, cephalic swellings, fugral protuberances. prothoracic spiracle, dorsal spines of abdomen stghily pigmented: abdominal Skin not pnented Adtennal horns blunt onantlerior suchice, produced anteto-ventally Hitoau aeule ridge, Cephalie selenite wilh pair of swellings shorter (hin antennal horns. Cephalic pair ef papillae With long serie. Moos with one or two sclerouzed protuberdnees on cach side, One Of two lower facial papllic With seni, one Of three lateral papillae with sel. Abdominal seginents | - 7 awilh pair of setose ventral papilie. 2 pairs of setose pleural pupilhw, 2 pairs OF usectose and pair of setose dorsal papillae, Abdominal seements with pair of ventral papillie, 2 pits Ol pleural pupiite, purr of dorsal papillae, all selose, Abdominal seunients 2 - & dorsally with field cf strong. one- or tweepetited spies on auiterior tall, Lave Inlewument covered wilh finy, Sparse Spiculac, Head: strongly selerotived, postern-luleral apodemes longer than head capsule, antennae 2 x Jomger than basal width. Neck segment with pair of dersul papillie, Thomeie segment, with pair of ventral pupils, 2 pairs oF pleural papillae. pair oF sternal papilhie. 3 pairs of lateral papillite. 2 pins of dorsal pupillae, Spatula bilobed, with shut. Abdominal segments. |= 7 with paitol veuteal papillae. 2 pais of pleural papitlae, 3 pairs of dorsal papilfie, Abdominal segment 8 with pair of ventral papithie, 2 pues Gb pleural papillae. puir of dorsal papilhue, Terniindl segment with pair of aml papillae on short lobes, pair of terminal papillite on prolonged lobes, ATT pupillie aselose, Livintelasy The prefix, “Okfio?” ts fram the Greek okries. Meanie roughness, referting to the pigeed ventral surfice OF the dedeavus dnd distinguishing the new genus trom other Sehimomyimi The suttix -myiae is Cireek for fly, Bematks Okrtonvie gen. hoy, belongs ta the trike Asphondyliini on the basis of the following shared dpomorphies: the presence Of a ventrosupied! lobe on he SondeoXite with wonestylis consequently sihdatcal dorso-ventrally; the short, quadrate pomosty lis, He presence of puramercs: and the lange female slempite 7 thal iy 1.5 4 is long as sterniie 6. TDOMYTDAE PROM EUCALYPTUS 47 The new genus belongs tothe subtiibe Sehivorny iit beewise il lacks a yentro-apical spine on the first lursomere. bas male parameres. hus a short Meshy ovipostor and the pupil miegumentis unpigmented, The new genus is unique among the Sehivomyiina because of the divided acdeapus. the solid loath on the gonostylus and (he cerci-lke female tergumt 1 The Australian gents. Leeinoricariio Fell the oat other genus assocniued with galls on /vealyptan spp. (Kolesik 1995a). most closely resembles Gkrionie wen. now OGdrtoumia shires wilh Keciedicarna the lone Tobes on the lerminal larval segment and the Neshy ovipositor with divided ceren. whieh represents the most plesiomoerphic ovipusitar in Asphondyliini. The tibe Asphondy lit is Known in Australia front 12) species distributed among four gener Ayphandviia Loew, a large. eosmiopolitin genus. belonging lo the subiribe Asphondyhina, contains sever species: A, aithocercidty Kolesik (Kolesik er a V99T), A. dodonevae Kolesiky (19950). A, a arimis Kolesik (1997), A, lal Fava (1916). \. inte Kolestk (L997. AL loews) Skuse (LASS) and A mibieunda Skuse (1888). The other three genera, all belonging (oO the subteibe Schizornyiina. are Knows only from Australi and contain five species: Bnvineticornitoustalasiae Felt (POU) Eo malarska Kolesik (1995u). Shuserurtee atlecusuarimee Kolesik (1995b) and the new species. Okrinnvid selec and Ch fubellidenrata Key to Australian genera of Asphondytiini 1. Pirst farsomere With spur male pumimiere absent) fenile with pair or dorsal Tobes at base ob needle tite eV positon, ptipal slain conmplerely piaiiented as = 1 Asphandylia First tursomere without spur: male paramere presen female witholt pur Of clarsal febes at base ol ovipostlot or owiposrlor not oeedle-like; pupal skig mat preme med ona leastubdomen., (al owepiefoes email estate re seebitven so 1 Pitee fteriniinal female Thigelotieres slickossively anu progressively shorten; ovipostor needle-like, mule purneres Tiree, us wide as postcrion lobes of coerce, pupal cephalie swellings longer tian antennal horns p cobkinens otto eres porrerma acto comeaphracerna vorcpon xo egypt toe REELS FAL Three tering female Hagellomeres subequal in length: ovipasitor shert, fleshy, wath untiised cerch mate purdnieres smth meh narkower than posterior lobes at cerci! pupal cephulie swellings shorter than antennal TIGHT PT ns coceccenenncasiiilers conse nsaitidesMensriestleescesibintecen gonbhiidenn al ‘aa Tooth on sonosty lis ConsEstiqg ol seven separate leah, fermale with pair at smal) dorsi lobes posterior (0 eighth terete: fareest dorsi sping vn pupa serrated Upiclly on ved EOC Orne ‘Tooth an AYNsIYLUS “sot im tai ut Wo pllwen female withbut pir of dorsal lobes pustercar Wo with wererbe Uiiesal spines: cn PPA ONE Ort wed-perntedl A krienive 1s P KOLRSIR Okriomyia schwarzi sp. ny, (PIGS 1, 3-6, 19-23, 27-30) Hoalowpe: &. Nadda. South Australia {34° 37" 40° 54° FE], Lali, (990, reared by PL Kolesik fram branch gallon Aucalyptas gracitiy b Muell, larva calleeted [2vii 1996 by J. Sehware, [21338 |JSAMA| Paratypes; 255, 3 9°. 4 pupal skins [SAMA, L21SA9-121347], 2 8A, 2 FP, 3 pupal skins [ANIC], same ditt but emerged 12.-20.Vin, Love: 4 larvae |SAMA, T21348421350). 2 larvae [ANIC], collected with holarype hie. 3, Gall OF Oknaniiin sofivercr sp. nov. on Bucalypins wis ~ longitudinal section. Arrows mirk larval exit hiles., Scale Py 10 na. Other niuterial (SAMA: 4 od. 24 23 pupal skins, 3 pupuc. sania datas 17 larvae, gall collected Will holotype: 2 9°22 pupal skins, Porestwille South Australia |44'56°$. 14836 Ef. 23.1, 199s. P. Kolesik, reared from branch galls on 2. ereesliy larvae collected POAT 993; 4 gulls collected with holotype [SHSA I, Deseriprion Male (bigs 4-6, 10-15) Colour antennae grey; head black: thotax brown: lees yellows ahdomen with selerotized parts and Wing length 2.2 mm (19 9 25). Genitals gonocoxite covered wilh short setae, with 2 short, thin, posterior lobes; apico-ventral lobe oon wonoeoesite long, deleulate: tooth oon gonostylus setae black, Hon-sclerotized parts orange dorso- medial narcow. finely serrated: aedeaetis narrow distally in lateral view; hypoproct with durge lobes, ats loug its Hedeagus. Fennile (Fuss 10-20) Wine length 3.0 mn (2.8 © 3.3). Circumlila oo Nagellomeres uboul ball density of mile ones. 15 x (63 -b.0) lonver than sy Abdumind stern 7 sternilc 6, Setac on cere 2 X shorter and much denser thin on lergite 10, Ovipositor as Jong as lergites 7 und § towether, Colour as in male. Pupii (Figs 21-23) Colour: Antennal horns brown, cephalic swellings. facial protuberances, prothoraeie spiracle. dorsal spines pale brown, abdominal skin grey. Total length 4.3 mm (3.8 - 4.0). Antennal hortis 86 pin (77 ~ 109) long. Cephalic sewe lol tin (138 - 18h) longs Cephalic swellings 46 Win (36 © 65) Jong, Upper five with 2 pairs of sclerotized protuberiunees, inner pail SI pin (48 ~ 54) long, outer pair 30 pam (29 — 38). Selae on lower facial papillae 122 pin (103-143) long. Prothoracic spiracle with slight, eridual curve, 244 Hin (206 267) long, trachea ending al uper, Learvai (Hiss 27.30) Colours pink to ofange. Total length 3.9 mo (44 7.8), Head capsule width at base 92 fim (90 ~ 94), lengih 70 pm (63 - 74). length of postero-laleral upodemes 116 tin (110-127). Antenna 26 pum (25 - 27) long, Sternal spatula 445 po (461 - 543) lone. With apical enlargement (OO pin (83 — 130) wide depth of incision 46 Lim (29 - 68), Termmal lobes 160 um (113-233) long Etyinology The speaies is named after the collector of the lurval stage of the Wpe specimens. Julie Schwarz, Department of Plant Serenee, University of Adelaide Gall and bielows Young branches of Anecelvplus wraciiy are swollen lo form galls 8 - 20mm in length and 7 - 9 mim in diameter. with outer walls 1 - 3 rim thick (Fig. 1). The gall outer surface ts scabrous, reddish brown i colour, Inside there dire 1 - 5 ovoid chambers, each occupied by | = 13 larvae, Larval colour may yviry from pink to orange between chambers of the site wall but is the same within a chamber, No association berween the colour and the age of Jarvae was apparent. Gall walls contain less woody Ussue than Wnaffected parts Of the branch, which results i the will being springy lo the touch and cruhehy when cut with a kaife., This characterisuc is shared with alls OO. flabellidentata. When the kirvae are Tally #rown, they Jeave the galls through one or (we wireular openings that develop in cach chomber (hig. 3). Pupation tikes place tn the stil, Okriomyia flabellidentata sp. nov (FIGS 2, 7-9, 24-26. 41-34) Holotype: &. Cleland Conservation Park. South Australia [34° 58° 8. Tak! 42" Ri, boa bee? P Kolesik. reared trem beaneh gall on Brealyprits NEW CECIDOMYIIDAE FROM EUCALYPTUS AY Fives 4-6. Male of Okriomyia sclwwersi sp. nay. Pig. 4. Acdeagus in frontal view, Fig, 5, Aedeagus in lateral view. Pig. 6, Genitalia in dorsal view (inner part of cerci diagrammatically cut out for beter clarity). Pigs 7-9. Male of Okrionn te flabellidentate sp. noy. Fig. 7. Acdeagus in frontal view. Pig. 8. Aedeagus in literal view. Fig. 8. Genitalia in dorsal view (imer part of ceret diagrammatically cut out), Seale bars = 100 tim. Abbrev.: a. aedeagus: e. cercus: ga, gonacaxal apodeme; ge. gonocoxite: gs, gonostylus; h. hypoproct: 9, paramere, pa. parameral apodeme. 50 P. KOLESIK Figs 10-20. Okbionrvia scliwarel sp. nov, 10-15 male, 16-20 female. Piz. 10. Head in froma view. Pig. Pl. Wine, Pig. 12. Last three Magellomeres. Fig. 13. Sixth flagellomere, Fig. 14, First tarsomere. Fig. 15. Last tarsomere with claw and empodium. Fig. 16, Sixth flugellomere. Pig. 17. Last three flagellomeres. Fig, 1S. End of abdomen in lateral view. big. 19, Oyipositor in lateral view. Pig. 20. Ovipositer in yentval view. Seale bars = LOO im 10. 12. 17. 19.20: S00 pm |: 50 pon 14-16: 200 tm Ls, Abbrev; c, cercus: h. hypaproet: 5. sternite: & tergite. NEW CECIDOMYIIDAE FROM EUCALYPTUS \ \ . y ) dN ) ° \ 7 \\ \ | \ . 8 \ \ (oe) os \ @ D ms * Sa . 9 Ne f \ " \ | fret { \ I, | i SS. \ , i | \ if | y i 32-——_ 33-— ce Vigs 21-23. Pupa of Okriomyia schwurci sp. noy. Fig, 21, Anterior part in dorsal view. Pig. 22. Anterior part in lateral view, Pig. 23. Prothoracie spiracle. Figs 24-26. Pupa of Okriomyia flabellidentata sp. noy. Fig. 24. Anterior part in dorsal view. Pig. 25. Anterior part in lateral view. Fig. 26. Prothoracic spiracle. Figs 27-30. Larva of Okrioniia schwarci sp. nov, Pig. 27. Wighth and terminal abdominal segments in ventral view. Pig, 28. Eighth and terminal abdominal segments in dorsal view. Fig. 29. Head in ventral view, Fig. 30. Spatula with adjacent papillae. Figs 31-34. Larva of Okriomyta flabellidemata sp. nov. Pig. 31, Bighth and terminal abdominal segments in ventral view. Fig. 32. Eighth and terminal abdominal segments in dorsal view, Fig, 33, Head in ventral view. Fig. 34. Spatula with adjacent papillae. Scale bars = 200 bun 21, 22, 24, 25, 27, 28, 31, 32: 100 Lm 23, 26, 30, 34; 50 um 29, 33. 52 P-KOLESTR casnuiplivilea b Muell.. larva collected 23.17.1997, 121351 |SAMA],. Puratypese 2.33, 3 pupal skins (SAMA, 121352- 121356). 2 22,3 pupal skins [ANIC], same dita bul emerged! (5-17.11, 19972 3 larvae [SAMA, J21357- 121359]. 2 larvae [ANIC], collected. with holotype. Oren material (SAMA): (all collected: fron, branch sally on A. casmophylla by Po Kolesikhe 7 oct. 4 pupal skins, sume data bul emerged |-f4iv. 997: 12 larvae, gull collected with holotypes 3° larvie. Morialta Conservation Park [34° 54° 8. 138° 44 0], 27K, 1992; 9 hurwac, Clehinal Conservation Park. 3, & FQN, 1995; wall, Cleland Conservation Park, 73.) 1943 [SHSA]. Deseviprion Male (bigs 7-9) Colour: as in QO. sehwears Wing length 2.90 in 27-40). Genitalia: gonueoxtte covered with long sche. WHT Awa Shork, posterg-dorsal lobes, one thin, One Wider apleo-wentral labe on gonocoxite stot, routed, tooth on gsonestytas wide. coursely serred: acdeags wide distilly ja lateral view: liypoproct with thin lobes. much shorter chin Wedeagus, Fennile Unknowea Pupe (Pigs 24-206) ‘Total length 3.9 fain (3.7 © 4.0), Antennal hors 62 um (St - (15) long. Cephalic setae 147 um (137 1605) long. Cephalic swellings 25 tm (20 - 29) Jong. Upper lave with parrof sclerotized protuberances, 31 jn (25 - 34) long. Setae an Jawer facial paplac 38 frm (32 - 45) lone, Prothorieie spiriele bowed: il distal third, MO um (174 204) Jong, trachea ending abapex. Otherwise as in Qn se/warst. hein Wigs 31-34) Colour pink (o orange. Total length 4.4 pam (3,7 - 5.0), Head eapstile width ait base 99 Lh (95 = 102), length 7O Lin (69 81). Teneth of posteroshiteral apodemes 125 pny (LOO - 140), Antenme 24 pn (24 - 25) Jong. Sternal spatula 425 pm (398 - 475) long, With upiedl enlargement 127 pun (LOS - 14) wide, depth of ineision S4 ta (50-59), Terinmal lohes [48 pun (2b = 160) long, Livros The name "tlabellidentata™ is i eompound Palin ddjecrive Tram “flabellun’’ meaning lao. and “dennis. meaning toothed. refering ta the shape ol The tooth on the gonostylits. Gall wid dielagy Youn branches of Auacelyday cosmoplvila are swollen ti form galls 10-70 tim in length and (0 = 1S mm in diameter, with outer walls 2-4 imo thick (hig. 2), The gall outer surface is smooth to scubrous. green to brown i colour, Inside there ure | - 4 inegulirly-shauped Chambers, each vecupied by 5 - IS larvae. Pupation takes place in the soil. The gulls ren tecornisable on the branches for several years afler (hey have been formed. Many branches hater fravtuwe at the site of the gall since the gall Hissue as less rigid than that of the tree. The same phenomenon was observed iW On yefwerci, The galls OF O fibellidentata on i. cosmoply tie are common in the nature conservation parks around Adelaide, Remarks The two new species difler frome cach other in several characters. The males of OAriemiyta selves Haye wnarrew loth on the 2onosiy lis, the hypoprect is as long as the aedeagus, the gonocoxite has two Wil, posterior lobes dorsa-medially. wid the apices ventral lobe om the pgonocoxile is aceite: Thre mules of OQ. flebelfedenane have w wide tooth oa the gonustylus, (he bypoproek is mueh shorter (at che vedas, the gondcoxite has no posterior lobes dorsa-mediilly bul tas one thin und one wide lobe dorsally, and the upice-verntral lobe oan the gonccoxie is short and rounded. The pupae ab c, vohwurs! have bwo pairs ol sclerotized protuberances on the upper faee, long setae on the lower facil pipilac, and an evenly-henl prothorucic spiracle The pupae of O. ffebellidentata have one paw of sclerotized protuberunees on the upper luce, short setae on the lower facial papillae, dnd a distally bowed prothoricie spiracle, Thal as many as 12 males and ne females were reared from the one gallon kieahyres cosmophaila eolleeted JAI 1997 suggests that females of G. flabelidentata produce unisextual progeny, a phenomenon found im Centarinne vorghicole (Coquilletty (Baxendale & Teetes TYh)) and Cystiphora sonchi (Brent) (MeClay 1996). In arder lo verily (he production of unisesual progeny iO, flabelhidentata, and perhaps ©. sehwerg. inure adults have fo be reared from: separate galls. This Indy require rearing larvae rom a larger number of Wills ds OL flabel/idertata seeiis Hot lo be an easily reurcd species. Prom some 1500 lirvae originating frome TO galls inehided in this work only the }2 mates emerged. Acknowledoments The Department of Environment and Natural Resourees, South Australia kindly permitted Collecting in the Cleland and Morialia Conservation Parks, M,C, O'Leury, Stale Herbariin of South NEW CECIDOMYIIDAE FROM EUCALYPTUS Australia Adelaide, courteously identified the host plant species. Special thanks go to J. D. Gray, Department of Horticulture, Viticulture and Oenology University of Adelaide and R. J. Gagné. 4 Systematic Entomology Laboratory USDA Washington DC USA, for commenting on an early draft of the mansucript. WwW References BAXENDALL, EP. & Trevis, G. L. (1981) Production of unisexual progenies by (he sorghum midge, Contarinie sorehicola, Auns. Entom. Soc, Am. 74, 412-413, CHIPPENDALE. Go M. (1988) Eucalyptus, Anguphora (Myrtaceae) pp. 1-447 In George, AS, (Ed.) “Flora of Australia” Vol. 19 (Australian Government Publishing Service, Canberra), CUNNINGHAM, G. ML, Mutinam, W. £., Manriorer, PL. & Lreicu, L. H. (198l) “Phints of Western New South Wales” (New South Wales Government Printing Office, Sydney). Epwarps, FW. (1916) Two new Australian Diptera, Av. May. Net. Hist. 103, 496-502. Ping, LP. (1915) New genera and species of gall-midges. Proc, US Nath Mus, 48, 195-211, GaGnh. Ro J (L981) Cecidomyiidae pp. 257-292 In MeAlpine, J.P, Peterson, B. V.. Shewell, G. O., Teskey. HI. Voekeroth, J. R. & Wood, P.M. (Rds) “Manual of Nearetic Diptera.” Vol. 1 (Canadian Government Publishing Centre, Quebec). _ —— (1989) “The Plant-Meeding Gall Midges of North America” (Cornell University Press. Ithaca, New York), KOLestk, P. (1995a) A new species of Lecineticornia Pelt (Diptera: Cecidomyiidae) on kucalyprus fasciculasa in South Australia. /. Aust ent. Sov. 34, 147-152. (1995b) Shusemvia alocusitarinde, a new genus and species of Cecidomyndae (Diptera) damaging lateral branch buds of drooping sheoak, Al/locesuariid verticilata in Australia. Trans. R. Soc. S, Aust, 119.4140, (1995¢) Asphondvlia dodonacue, anew species of Cecidomyiidae (Diptera) damaging leaves and branches of hop-bush, Dodonaea viscose (Sapindaceae) in Australia. Zoid. 119, 171-176. ~ + 6 il (1997) Two new species of Aspliondylta (Diptera: Cecidomyiidie) trom Iilosercia spp. (Chenopodiaceae) in South Australia. hid. 121, 59-66. _ Wiirtremore, R, & Stach, H, M. (1997) Asphonidylia athocercidis, a vew species of Cecidomyndac (Diptera) galling flowers of Anthoverciy litterce (Solanaceae) in Western Australia. /bzel, 121, 157-162. McCay. A. S. (1996) Unisexual broods im the gall midge Cystiphora senchi (Bremi) (Diptera, Cecidomyiidae), Canad. Entom, 128, 775-776. Skusy, FA. A. (ESS88) Diptera of Australta. Part 1. Prac Linn. Soo, NSW. (2nd Series) 3, 17-145. WITHIN-NEST BEHAVIOUR IN A EUSOCIAL AUSTRALIAN ALLODAPINE BEE EXONEURA (EXONEURELLA) TRIDENTATA HOUSTON (APIDAE: XYLOCOPINAE) By ZETA STEEN* & MICHAEL P. SCHWARZ* Summary Steen, Z. & Schwarz, M. P. (1998) Within-nest behaviour in a eusocial Australian allodapine bee Exoneura (Exoneurella) tridentata Houston (Apidae: Xylocopinae). Trans. R. Soc. S. Aust. 122(2), 55-63, 29 May, 1998. Understanding the processes involved in the evolution of social behaviour has become one of the most challenging areas of modern biology. Since bees and wasps exhibit a variety of social organisations they are particularly useful for addressing social evolutionary questions. Allodapine bees are especially useful for examining social evolution, since species display varying forms of social organisation from solitary to eusocial. This study examines within-nest behaviour of Exoneura (Exoneurella) tridentata, a native Australian allodapine bee. This species has the largest known colony sizes of any allodapine bee and exhibits striking size variation among female nestmates suggesting that sociality may be regarded as highly eusocial. Key Words: Exoneura tridentata, social behaviour, allodapine bees, aggression. Piawurrions of de Navel Nentety ab hive (1QO8L 122(2). 55-63 5S WITHIN-NEST BEHAVIOUR IN A EUSOCIAL AUSTRALIAN ALLODAPINE BEE EXONEURA (ENONEURELLA) TRIDENTATA HOUSTON (APIDAE: XYLOCOPINAE) by ZPTA STEE ’& Mictiart P Scuwarz Summary Sihtn. Z. & Scuwans MOT Cfg9Ss Withinenest behavioul ia etsicial Australian alledapmne bee fe tenenc (vareuredta) iidentatoa Houston tb Apidae: Xylocopiuie), Tris A Seo SN Advi 92202), 55-04, 29 May, 198, Understanding the processes invelved in the evolution of sociil behaviour has become ene of the most Chillensing areas of modern biglowy, Sinee bees and wasps exhibit a variety of sect) orgunrsations they are paicuherly Use for addressing soetl evalitioniny questions. Stlodapime bees are especially sel for WT Soci evolution, since species displiy varying forms of sachil ordination fron solitary 1 eusacial. This Study examines WHh Tn nest hebaiviodl ol Bievewia (Evonearete) tridentate, a niative Austertian allodapiic boo. This species his fhe lamest Known colony sizes of any allodapioe bee anc exhibits siriking size vuriaiion among female nestimtes susgesting thal sockliiy may be regarded as bighly cusoeil, Here we assemble a bobavivical cutulosie Lor this species atid show dat although many behaviors are similar la those recorded: (or other allidupines. this species differs by the marked presence oPovertaesression displayed in the form of biting. Overly dgonshc bebaviodi's have fet been recorded Om ather Australian allodapiaes and lave been recordca ony rarely ay offer allodapine (iin, Evenedaye fridentaie appedes Wo differ from ether bizhly eusocil speetes where (here s uscally HAle or no aggression bul iste “gentle despatisnn', Key Worbs, Jowioie: pide, social beliavinur, allodupie bees. aeeresston Introduction The atlodapine bees provide opportunities for LOIpUvE approwehes to the evolugan af social behaviour becouse of the wide range Of social Orunsaiion WHE the betwee spectes and genera One sinall and endenie Australi subgenus Prone. contitins fours species that ringe from the predominantly solitary fiveaenra densa Rayment (Michener 1965) to the eusecial & fedemate (louston (977: Hurst & Setware 1996), In most comparative studies of insect social evolution (bere is an impheit assumption that small coluny sive is assoeiated woth flexible und hehaviourally mediated reproductive skew. The ailehance of dominance hrerarehies vie phy steal awoniSnh is conmsmWercd wi opranmiiye trail (Wilson O71), Correspondingly, Jurge colony sizes with strom repraduaive skew und) pon-ugonmsrieally mintiined hierarehics are usually regarded as move derived traits. Wilson (1971) stiggested that less Sophisticated Forms of social ordbisation would iwolve physreal neelinisms OF control such as igeression Wwilhimew colony: bul (hal this ts rephiced by “gentle despotism" in more advanced lorms of soenilty. [lis alse generally assumed that a high level of behavioural specialisation wa more derived Ira and that this ean lead to higher levels of volony eHlhieney (leanne P98So) However. dhe idea that dillerent formas oF sacl orginisutiog cin be Seal al Biolomel Seieniss, Flinders University of South MSHI GHD Wax STD) Adlethoidle’ So Aviat SCH TD tibial: Aa Steen Ge Tider scabies dFunged ina sequence of ‘prmnidve to Tadwaneed? his been questioned (Kukuk 1995) bul few studies have explicitly investigated whether jpronitive’ or ‘udvaneed! Loris OF soctulity Within tixu correspond 10 husal or distal positians within phylogenetic trees, Eronenre tridentata is ao Austrabir allodapine hee thal lives in semi-arid environments. This species hits the hirgest Known colony stees of inv allodaprne bee and exhibits morphologicul differentiation. between putative castes (Houston 1977, Hurst & Selwirs 1996), Much of the information aboul social organisation bas been inferred frou disseetion af Hest ovcupants and brief observations of females ouside of their nests Clouston 1977, Hurst unpub), Tas suspected that this species exhibits caste dilferenliation, where large females (lermed ‘Miajors') ure queen-like and smatler Females ('Minors') act as workers within the colonies (Homston 1977: Horst 1996). Llowever: wilhih-fest behavioural studies have not heen curried Ou) Loouaséss Whether These Iwo trorpls realhy ane behaviourally distinet, Colony size and the wsseenition between morphology une reproductive shitty sugeest thal (his species more closely approaches the bightly cusecial fon oF organisation characterishe obapinie. micliponine and highly eusocial hahietme hees. than uny other allodaping hee. This study Mvestigates \witiichest behaviour rn observation colonies ob /. tridentata. A repertoire ol hehuviours ts presented fiere in the term of i bebaytoural catalogue and compared with other behavioural studies ob wlodipines. These data well Who be used for specitic anulysis of behavioural speculation, which wilhappear ing (uti series of a0 ¥ STERN & MP SCHWARZ poblications. Ty addition, the Wea that morphological caste differentiation and linge colony size are ussocniled with low levels ob agenisat im colony intevration is discussed in relulion lo the social ormimisuhion ol A. prredemtaret. Materials and Methods Stitely vite Exonenra tridentart tests were collected fram Lake Gilles Conservation Park (13048 bk, 32 54 8) loaded in the ori east of Eyre Peninsula. South Austrulig. In this aren, 22. aidentde Wests were prin oipally ty disused beetle burrows excavated in Aewe fr pupyrecarpo Benth, (Western Myall) and Halyerryou aleiafoliun (Dest). (Bulloek Bushy Deal branches ol both tree species were examined for nest enifinces ie. the exit holes made by the original beetle oecupalts Trtaet colonies were collected daring February 1995, Field collection of nests took place when lemperatures were cool (12 C-20) Cy. 0 ensure Thiet all Gecupinty were present. Onee un entrance hole was locwted, the brunch was removed, entrances were blocked With tHssuic paper, fhe brinch was placed ina watemrool bay and stared qh ai insuhured container with jee for Hransport to blinders University. At Flinders University the nests were stored ana constant Lempertture roon ab approximilely 1 ¢ for provessing. Nests were opened using a knife cil ull nest aceupants. meludiig brood and: test contents sdch us pollen. were transferred to a Perr dish. Adults were individually marked) using Uambrol!™ and Testors! enamel pauls apphed to The thorax and merisond, Bee colonies were then Prubsterred to arlifienl observation Wests, Arniticial nests were similar in design te thoxe described by Selwear? & Overholt (1993) hut were imade Ob pine wooed tstead ob balsas Bach dest consisted uf a rectingulie peewee of untreated: pine woo 7108 20% [Ss mint A gropye was gouged inte one longitudinal fee (3 rm dian x 200 mm tent), The groove wus smobtiicd out wilh a mend rod ta remove any Splintdrs ab woud. A piece of glass. 210 X Shon, Was placed Mushy suuiinst the groove aid secured! al both ends with iipulador gape. A black curdbourd cover was placed over (he hiss Wy exehidhe Hivht henween observation periods. Observalion nests were set up oo subhori4ontal trays ina shade house at Flinders Unrversity. One end of the shade hotise wis open so (hab bees could. fords freely outside, Nest entraiees faced fhe oper end ol the shade house, A inuxinian gl four nests wats plased on each dray with approstmately TS ern herween cael) nest Observalinn nests were list placed in the shade house at disk Sy days athe colleetion and opening. This ensured that the bees hud approximately b2 hours tinue aruhenl nese to allow their odours 10 permeate the nest before i wits possible for them to leave (the next morning) Sticks were haphazardly placed near nests ty wet as visual cues for returning bees, Behavioural observations Ofice Observation Nests Were ser tip Aves were alowed to adjust Lo their new enviroment lor one week before observations began. Data collection ivolved ‘sean’ and ‘focal’ sampling techniques (Altman 1974). Sean sampling invalved recording the position of cach individual a the observation nest. using a S prim seale whong the gliss and wos vonducted immediately before and alter focal simpling. This was done to determing whether certain bees were spending more Gine than oliers 1 verlain areas of the nest. for example, uew the entrance or near the hrood. Focal sumpliny iiwolyed 2 min observations of cach bee mat nest. Nests aod individuals were randomly Selected euch day for order of observations. A headband magnifier (4 5,2 magnification) was used to observe (he behuytour ol individuals. All behaviours pertormed ina 2 min period for each individual were recorded into a voice opened recorder Observations were trie sermbed on to data sheets ata later date. These behavioural dala were used to eonsiruel te bebay foul catalogue ind liter to caumine behavioural speciilisanion. Behavioural observations look place ia the after noon, (1300-1700 hp, when temperatures were = 20 Cound bees were active Tn total, 1 nests were observed with up to lotr bests here Gbserved i any one sessian. Table | provides information about whieh nests were observed. when Wey Were observed ind how miny minutes of observation each bee per vest received) In addition, the numbers of bees (hal Were present for the ital and Gaal observatin Periods are saved. Results Field-callected Wess The contents of nests collected TH February Ln are summarised in Table 2. Durtne these simpli peniods. colonies used for behavioural observations were ceubings brood, dT curly Februiry colanios contimed brood Gf all developmental siames. Le, owes, larvae, prepupae and pupae By bite February fondle bees i tire colonies: haa aliiost ceuacd ces hiyin ind brood mostly comprised: Taryae, prepupyie und pupa, There was a great deal of vartabion i tie number ol adult lernales present ima nesh rine Mun T-TR (Pie TS BEHAVIOUR IN AN ALLODAPINE BEE ST TABLE |. Details for nests of Exoneura tridentata obyerved in this study. Nest First Last Total number Total minutes — Initial no. of Final no, of observations observations of of Individuals Individuals observation observation periods per nest per bee per nest I 7 Mar: 14 Apr. 15 30 8 a 6 7 Mat. 14 Apr. 15 30 9 Sa y 7 Mat. 14 Apr. 15 30 ) 12h 12 7 Mar. I4 Apr. 15 30 13 15h 3 5 Apr. 4 May i) 38 5 5 4 5 Apr 4 May 19 a8 4 5b 20 5 Apr 4 May 19 38 4 5¢ 30 26 Apr. 16 May 2) 40 6 x 43 26 Apr. 16 May 1) 4) 3 4b 56 29 Apr. 16 May 20) 40 4 4 Decreases in the number of indivduals were probably due to death whilst foraging or dispersal to other nests“ Increases were due to the addition of newly eclosed bees?, or intruders which swapped nests°. Taser 2. Summary af nest contents for colonies of Exoneura tridentata collected in February 1993 from Lake Gilles, Soutlt Australie. Mean value (+ S.E.) for early February Nest contents Mean value (+S.E.) for late February (N=24) (N=13) Eggs 1.21 (0.57) 0.08 (0.08) Larvae 75 (0,03) 0.62 (0,27) Prepupue 0.67 (0.28) OAT (O13) Mupiae 242 (0,72) 2.23: (0,70) Majors L.L7 (O16) 1.23 (34) Minors 4.17 (0.83) 4.48 (1.40) Males 0.33 (O13) (LA8 (0.21) Behavieural repertoire In the following section behayiours observed during ihe study ure presented as a behavioural catalogue. Observed behaytours are classified into four functional groups (often interconnected or overlapping): (1) sell maintenance behaviours, (1) nest maintenance behaviours. (ii) inter-adull behaviours. and (iy) aduli-brood interactions, StL MAINTENANCE BEHAVIOURS INACTIVE Bees were recorded us being "inactive" when no Number of nests other behaviour was being performed, Inactivity often occurred within a behavioural sequence. For example. a bee could stop grooming, be inactive [or same time. and then travel forward in the nest. Bees could ether be stitnding ‘upright’ or they could be lying ‘upside down' on the floor of the nest. Maeta ef af. (1992) included slight movements in their | description of a similar behaviour, "Resting" However, in this study bees were only recorded as 20- 10- 16-18 I 2-3 4-6 7-1) A1-15 Females per nest ‘a. 1. bistegran of colony sizes (number of females per nest) of Aveneure tridentate collected from Lake Giles South Australia, Bebruary 1995 3K 4, STEEN & M,. P. SCIIWARZ inuehive When Wey were motionless, veers iWdentata spent a large amount OF Lime inactive, Since inaciyvily can oecur within and between behavioural sequences i) is dilficull ta) show Humerigally the wmuunt ob time spent juctive hecutise of the way the dite were colleeted, Generally, though, We bees were more active Wher lemperilures were 220°C and/or when a lorager returned, SELL GROOMING “Croomine” wits observedt frequently, and included wy uelivily where the body surface was cleaned Sequenves lop clean different areas of the body wore similar to those reported: for Braaiseps leiierr Cameron and Cerufina spp. (Mactieral 1992), The most Common sequences were: (a) head cleaned by imtialhy wipta a foreles with the probascis then lorcloy wised ta wipe the leneth of the antennae. beginning dt the base: foreleg (ain wiped with the probosers, followed by the wiprig of the head wilt the forelegs, (bh) the metasomi was cleanee! by using the Tibial spurs ae the hindlegs to serupe all dust/pollen, (&) the thors was cleaned with the mid fous (the metasoma and the thorax were often groomed al the sine lime wath the different legs) (a) ihe wing surfaces were groomed by drigging the wins under the metisoma with the bind lees, Wiping then between the metasoma and hind legs, une then Thiekite then baek inte position, Grooming did not oceur as one long uninterrupted sequenve as hus been observed Tor Bo hewirt (Macta ef af, 1992), Grooming could be brie! or last for the whole 2 mi observation period, SHICHI BODY SION EMEA S This was intermittent behaviours whieh was fie Gbserved dung lone bouts ol naetivity, and bebiy- OUP Comprised SHANE Movements OF head, body or lees which diel nob anvelye any other type ol behav- TOU, PAWEL LING “Teeelling’ tnvalved movin: forwards Gt back sands up er down the west for 1-20 en, Bees that were travelling Were usdally very detive but the wavelliny: speed varied, Travelling forward offen resulted ina bee coming inlo contact with others and was usually followed by "passing! (see below), VURENING "Turning was used to desermbe a change of direction in the nest. Turning involved cuming the body and semersaultina. resulting the bee facing the opposite direction. Both Majors and Minors uppeared fo turn with equal ease, This behaviour vwevirred anywhere in the nese, uolike that in Cerainad spp. whieh have ov luring burrow enlurgement nea the nest ertynee (Maki en al, 1992), Thiming ollen ocenrred as parlor a sequence of behaviours during interactions between idividiials, ie. ih Gould becur daring sequenees which involved “nudging”, “passing” Or “uvoidance™ (see below), (fa hee upproached but avoided another bee, it mish either "Wavel” up Lo the bee, and then back away or il might "tare" and “travel” ithe e@yposite deen, MECTAR DEHYDRATION Individuals were observed flexing and bending the proboseis and. although droplets of neeiar would hat be seen with at the magnifications used, i wes assumed that they were dehydrating eet! as fas been observed in other allodapines idler leecdime (Michener 1972; Macta ered, (992), Same bees slow ly fully extended and retieted the whole pru- hoseis without bending if. The proboseis was extended and held out lor about 20 see thes retracted before being extended again. Some individuals spent the whole two mio ubservation period perlorming: this behaviour NEST ABSENTEEISM When individuals were regularly absent (rer the Hest Ho owits assumed that they were forage. However, i they were absent for more dni 3 obser JUUON sesslobs in a cow. He Was assured: (hat they were elther dead or had dispersed, Absenieeisi (ur forging aclivity) was only observed when tenper- atures Were 2 25°C. Foriwers were dentitied when they were seen returning to (he pest, Upen return. Hg, forrers usiidly worked (herr way dlown (he mesh pissing: dnc interucting. wilh) ether job Vidiels. often having “bueeal contac!" with vtber muds. presumably providing thei with mee- tir (see inber-adult behovieurs) Often such wv bee Would then leave the nest again amd retien filer Poragers were not observed feeding bir vue, Nest MOMNTENANEL BEHAVIOURS GL ARDING A bee was recorded iis “suurding" when ifoceupicd The position closest lo the nest entrance wilh ils body oriented se that its head wats facing away trom the entrance, Suelo position atiows the metisomu ta block the nest entrance from intruders, as reearded for other allodapine bees Bo lewitn (Maeur er al, 1999), Al miata (Batra er at 1993) aind L. biealor (Melia & Schwary 1993), During guarding the hee wus tnactive either on ths back or standing upright, i a bee was closest fo and facing the vest enivance, it Wus mot tecorded os guarding, sinee bees i thes postion would offen be we the process ol leaving BEHAVIOUR IN AN ALLODAPINE BEB ) the Hest, Minors were often seen guarding und in done fests, Majors, particulirly ege-layers, were aot seen to atid at all, Guarding did) hot always oevur near the nest cuirance, In some nests the “wuard" was stutioned hel ol the wiry down the nest bul was the bee closest to the nest entrinee, These guards were somedimes seen tn patrol’ the nest fron thitt secuon Up to the catrance. This involved the bec rapidly “travelling” forward, whilst rapidly aptennating (“Inspeetine”) the nest linen before returning to the guard position, Tn some nesis Th also appeared thal (wo tadividuals would) gud alternately or one ii frontal the orien Although there were times when More Than cone Hel vidal Was Seen ith The g@etara postion, there were individuals who ever "ouurded". During the study. no other invertebrates Were Observed entering the Hests. Sinve there was my interference from other invertebrate predators in the CAPUVe SIQATION, guardiig te this study tay ot reflect natin! behaviour ob this speectes. INSPECTING This hebaviour invelved a bee alternately anter- nate Obyects. lor example the nest will or brood, Epes were frequently antennated fin this way, Sometimes bees travelled up and down the Hestrospeetine the lumen wall Dirring this beh jour bees moved their Heads shehuly aed apiclly Hoved antennae, MOVING HUBRIS Debris in (he vest was moved by passing it under the body with the forelegs to the bind legs ther pushing backwards with the hind legs or metasorn This behaviour was rare (approx, Q.39 oF the observation Time) sinve the nests avere in hard, Mie-vriammed wood wich required litthe main- ftenanes, Debris observed Tm) the nest eluded cauynie jie. oecusronally, dead individuals, "Moving debris" was not usually observed doless lumperutires were = 25°C. PAPER ADE TT AE TAMIGUIES WOTPARC “Avondinee', a combination OF other behaviours, iInvelved Gne dodividdal daawelling: towards another mdividual dnd “Untcueiing either the metusonn or lace of thatindividdal and thea suddenly bucking away oe Tinie an tavellinges Ti the opposite alirection, ANTE AAT CON TACT "Antennal contaet® accompanied most itereadul behaviours, When wt Trdividiael Game tn te contact With another mndividtial a criti “antermiited" the ofher's metasona or faee. HW tidiyidtids were hice losfaee the two individuals tapped each other's antennae, PASSING “Passing” is the eschanve of positions by nest mates. Passing oeeurred when individuals were either ficing euch other or the "passer" was lieing the tnetasonw of the individual she intended to piss. In cach cause, iudividttals ooented (hentselves ven ler-lo-venter, essentially walking over each other A pass was either simple or complex. “Simple pass- ing” Gnvelved the smouth exchange el) positians, with individnals usually flattening thei bodies ugainpt The nest wall, “Comples. passing” involved one individual biting al anether medividualls buaty parts. und/or strupeling and grasping cueh other with the legs. Either one or both individials would Hits, Sometiones one midiyidial would bile the othe on the ventral sig between the ietusonit ane the (horas. near the articuluion between the tochunte und the thors. Passing somehimes mvalved briel "buveal camtuce” between the two individials. Although it Was often difficult to determine clearly whether buecal contiet tad actually geeurred. Ue Wan not always cusy to Uistinuiish between the passer adind the "passed", except when onde was intbially stationary and doarher wes teevelli pe HEICCAL CONTAC EP lnelivaduiils were often observed to toueh cuch other's open mundibles with their ewrn épen aindibles: this was termed "buccal cantict. When Hhdividuadls were Involved dn such intenichons. one individual was standing upright und the other was positioned: upside down, Tnehvidtals dibse engaieed tn briel hoceal contact during passing. During Approximately 54 oF biiweal contact iileractions. neekiur flow between the mouth purty of individuals Was Observed ancl individuals were observed placing thew probosers bebween the mandibles oF another individual, Protflering of lobules of nectar (Meloa & Sehiwirs 1999). wits nolohperved in by drideniale. UDO Ne! "Sudging"” involved one individual using Us fice to nudge or hut) the metisoma or face of another individual, The hee that "nudged" was usually upright. Nudging usually resulted in one af the Following: a) The nudged individual (ined and (he nudser retreated, which sometimes involved (he nudged hee opening iis mandibles. bh) UP nudged frewr behind. the bee beige gudved would sometiines position its antennmie laterully (oul tothe side). then 7 nudecd again ib mnght open is mandibles, ‘This eventially resulted in the hee a Z. STEEN & M, P.SCIIWARA juming, jnvesigaling the "nudger’, and thea simple or complex passing and/or buceal contact ¢) The nudged or the nudger pissing and "bring" uel oer, MANDIBULATISG Mandibulating, Le. the opening und closing of the mandibles not associated with eating, appeared i occur before biting enconnters. I some cases 1 Uppeared that mandibulating was a signal thal one individual was reject an approuch from another individual, Por example “Ab approached ‘Bt, ‘A’ nudeed "BY "BR then opened mandibles, “Ab (hen retreated. “Biting” encounters someuimes followed, Similarly. an imdividual was nudged rom behind it sometimes opened its mandibles and/or turned aed faved the nudger often opening the mandibles wan In wddition, Matlening of the antennie laterally often ocenrred during mandibulating, This sometimes oevurred when midividiails came face to face or il one was nudged from behind, BEEING In this Siidy aggressive cheounters were Observed for oh. weldentata. These the thor, around the coxue ung metusoma. Olen When one individdal tried te escape from such in eheounter the other bee would pull it buek osing ts forelegs. "Bilin encounters were olten vomples, For exwihple. (A> used its fie fe nudge "B'S lave. Then one or hath bees opened the mandibles and a complicated pass followed. Whilst the bees were venien oO venter and strigeling (holding cach other with leas; one would hite the other ou the ventral side ol the thorwa, After a sttiiggle the bitten bee wis oflen observed on dis hack while the biter hele the other bee's antennae in is mandibles, ia “ug Whawar" Gheounter This tug-of-war could: last far 120 see, botlowiie a fie-olewyr encounter the Tdividiial whtel bad daiiaited (he pass (he bition) SOMetiMmes ullenipledl (er puss ius and Ofer a siiiple pass would follow, ADRUEIE BROOD IN TR ACTIONS PY ANINATION OF WROOH Excunation of brood was acceanplished with ve ailennae, aid, faa desser exten, ie ieuth putts (open and clostime mimdibles on che bread) Individuals Lippe pupae. larvae or ees. WIE cael HAW NEEDING HROOD Brood Were sainetimes nudged before (hey were moved, This belivioie dich dot fesull, Hewever, iy jhe bro ipprestibly chanutas: position invelved bituay ab mandibles, auitennae, peek, legs, the ventral side of MOVING: BROOD Older brood (ate instar larvae, prepupae and pupae) were usually moved in a way similar to the way debris was moved in the nest. In A. iridentate. simik to EL biedlor (PS. VMurst pers. comm, 1995). the bee initiiy held the brood with the fore (arsi then passed them under the body and pushed ther backwards using the hind legs. Repositioning of brood occurred allen within the nests of A irideniua. Sometines a bee would move edeh pupa until it reached the end of the nest, then it would move then all back again: seeonds later another individual sometimes did the sane thing. Some Minors which consistently stayed near the broad were oflen observed performing this behaviour, fh addition, hees sometimes simply handled the pupae with the fore lows bur did pot aetially reposition them. GROOMING BROOD Bees ovcasromuly extended the proboseis to the hrood or bit gently al the brood with thei mandibles: such bebaviour wes eateporised as “grooming broad’. This behaviour was rarely observed. Greating jay have occurred during movirns or with Tiindlings but i owas difficult to observe the finer movernents of such behaviour becuuse of the speed oF movement of the prohoseis and the lirited magnification, OVIPOSTTION When "Ovipositing”. the female oriented herself so that the head pormted towards the nest entrance During eve laying hees were observed fone of thice positions: vennal sartice lyeing Upwards, dorsal Surface facing upwards und lateral surface facie upwards, Prior ound during "Oviposition” the sting Was Gaternled. Onee an ene had been deposited qa the floae or the nest, Ure bee retracted the sai Approximately ba punt passed betore the fenyale iuirovel around and inspected (We eee willy (he aalennie. Oviposition @ecurred close La the mest end (5 and was vibserveel ford Majors and | Miner ¢§ sepacdte colonies) Tadividhials Tok approx inidtely SO Wii fo discharpe un cep Plowever oie Majer look F8 ni to Tay at eas. Discussion Beliviouwr has prev daily heen stodied i denial far RB. lrewinli (Maeta er iil. 1999), A ote, AL kallawe (Baumer af (99%), AMlodapie exelent Strand) (Mason lOX8)} dind 2 bleelor iMehine & Seliweure 1093), Bvonenme (rrdentiata wenerallyospencdd iy Lite HMOURL OT Ce Tachive, SHH Teather Rees CN aot y eral (902: Batra ela (998), Activity temled lor pre wicuter ua dlays wheo the temperahines were uhove BEHAVIOUR IN ANS ALL ODAPISE BEE ol 28 C. Sinihily, wher the temperdture was warmer bees tended to forge more und, especially ater return of a forager to the mest. gener) activity appeared lo increase. Exon iidednita was bat dbserved to exhibit (he types of fest maintenance behaviours found in other allodapines. probably due to the hard nature of he nest substrate. Most allodapines excivate (here OWE NOSES I pithy Sudstrite imiterial, whereas &. Iridentatd do pot Although observation nests provided no opportunity for nest walls to decay during the course of (he sludy, natural nests are WAG unbRely to require fepines to the nese wall or entre, Since Lhey also ocetir in line grammed wood. fhe contrasts With ak, biceley which pertoriis varius nest miintenanee aetivities suely as Clearing aod famping (removing loose material (rom the nest wall and shape gest lumen) extending the nest lumen (excuvaling rear of the burrow). collar Construction (ping Wood imto a collar near nest entrance) und remoying debris dwood strimds. Zienewne rridenta may exhibit HOSE IUTIeNUNee SeOVTes Lod prenter degree when Hew nests ane lotinded and tere is a need to remove brine Tet behind by been larva, This study is the first lo deserbe eee Tayroy ip at Aagnenre species. Bee laying was only observed duting, (he diy althotirh iarkiy idsi have oeeurred at feht (obserwtions Were only tnade dupe the iliy) Meo living wieyscoilier bo that deseribed far B mrte (Batra oral WOD) un Bde (Maer et 1907). However, bwo of the three 2, tridmire majors That were observed OVIPOST Te were runely or never seen euardun. Phe thin mijor was seen bo sajurd bol she was sully S em fron the base of Hie nestamd fot pear the entrance. This differs from &, Jeovien (Maen et al O07) aad bo dicaler (Howerdoory de Selivare P9988: Buller atin pless) where reproductive glominants wre guards, Exe faving in his species appears da be uw yery slow process compared wath other bees (G8 see. Bo tear tH) (Maeta ef al 1992). in terms Of both the Time Taken to deposd af ems and (he freqdeney ob eee laying, Que tenile, i particuhae spent 4s min depositing aces whieh tray have heen related to the [acl that the femperture was low that dia ¢< WC). and bees were generally less active al lower temperadiunes. However, these ohservariins did pot cover the perrod of imaasrtal eee prduction and stow be treated with edution. Aggressive belityiour bas net been reported far olber allodipine bees except rarely henveon fH wiylitanel its Social parasite B halfage (Batti ef af (W984) and jnlrequently for A. eveforuee wed B. freee (Mason L988). Phe ayonistic behaviour described for These speeies rnuinly coosisned of nudging, biting Ol legs amd bodies ind blocking passage. hut alse included singing (Baten er vf. (993. Mason TOSR) Agoniste behaviour between a hast and fs parasite is nol uncommon and Often results i sither host G1 parasile being removed from the nest (Batra el af. 19034), Ageressive ieteractions are alse found in social Species OF the bee tibes Halictini and Nylocopiny (Breed et ah 1978) Michener 1990). However, 2. frfdeniiir Was offen observed ti cngave in gegressive encnunlers which inyolycu a prea dewl of bidne ound strugeling, with seme eneouniers Becoming quite savupe, Sach cneounters were offen preceded by nudging ind folliwed by passing. The miandibilating that ecard same- limes. eiiber prior lo er in response to nudging iil Hite. might idso be aggressive oy mature Cane Xe Michener (1983) found that some 2eeavewre: spp. produce iratuis whieh cligit Vigeroos geoming responses in predatory als, Batra ed af, (b984) deserthed oo mundihulating durrog aeeression heiween Bo ate und its sucnil parasite BL keviiges wil sugeested (hat mandibilar seerenons were involved, Hinay therefore be suggested that when & Tridemate mandibulate al eweh other, (ley alse release chemical seervetions which rity be weorishic or relay information wheut donee: stctus. The agonistie behaviours observed mL. ident sugeest Wal dominance hierirehies nny he present wilbin colonies, Te appears (hat some ijdividddls engave im cern oypes af behayinur whieh could he IMerpreted as assertion oF duminunce. Bees that are offer nudged or bitten and those that exhibit avonlance behaviour muy have more subordiiate roles in the nest. Dilferences in the way ined;viduals respon be other individuals in terms of these behaviours pay he related to dominance (he. when Some individuals wre nudwed (hey cheaee Ha simple: pass. whercus when other individuals ure nudged and/or bitben they Copuge iw complicated puss), Brothers & Michener (1974 found that ‘qneens’ ob Lenvreadiassan sephyran) were the maximal nudgers in the colony They steeestecdh (hal nodeing belie indicates dominance sinilar to hal observed in other prin Hively cusuen wasps amd bees Brothers & Michener (197-4) experimentally showed toe 7. séphyriin. that mudenig by the quicen pris ae role in the division of Libour anion the workers hy imbibing evar development, During this study gourding behastorr was not We sete us thucobserved in field studies ol Ey frfefeyitetia, Pewith the abdomen curled and used ta block the entrance from predidors suchaits anes UP Hirse unpub.) This may be rekited ta the Piet that there savas no predation pressure in the shade house environment unlike studies of AL Pieefar conducted in shide houses where ants were at problem (Bulls Furst") However females that were guardine were always facing the betitom of the nest which sugeests du ot 4 STUEN & M, PSCTIWARA, they were dna posmon to block the nest a the need arose. Trophalladis as altruistic behaviour, foragers engage in energetically cosdy and risky behaviour to vhiiin food whieh they reloigiuish to others. Trophallaxis isamportant in (he soci! organisation OF iiny seeial insects (Wilson 1971), Thy allodapines (here nay be differences m the way in whieh Wophallaais is performed. Evaneura brealer Wave been Observed te engage i solicition behaviour before rophallasis oeeurs (Melia & Schwarg 1993), Solieurion dvelved: individuals rapidly: stroking cach other's antennae prior to buecal eontaer, Trophatlasis in 2. bieular can adso invalve one iidividiel proflering a slobuile oF liquid: ty another (Melia & Seliware 1993). Peotfering of globules was not obperved in“. tridentera und UP solicitation occurred, ib was Loo fast to he identiticed, However, 1 is Hikeby that midividtals which engaged i "huecul conuiel where heel flow was observed, were frequently engaging i trophylhasis, Trophiallaxis Allows tenes to feed without leaving the nest. The presence Of Wrophullaxis in AL trideviie therefore allows behavioncal specialisation where only some Ol the females huve to lorige and other females ean perform other duties in the nest. Axonenura tridenieio exhibits a similar repertoire of behaviours loother allodapines (Maeta er ef, 1992, Bawa oe wh 193; Melna & Schwarze 1993), Behaviours recorded in (his study, ineludiie adult Udult fiteractions and adull-brood inleractions, ure TBC. Nr 1994) Puseehility ia Neatidand: populition ofan Astrid Atlodiuping bee, Eveneure bicolor Smith (Apidie. Xylocopime), BSe Cons) thesia, Plindens University of South Australia (unpub). Hers, POS (1004) Repradtictive hiamarehies by on Austealian ATodapine bee. Evaneiiid Bical Sith (Anthaphenrtdae, Sylogapaned, BSe (tons thesis, La’Probe University. (ipl), HIP Simihie to those found for other species, sugeestiny thal such behaviours are likely lo be ancestral an thal development of novel behavioural elements is Hot Heeessury Tor soci) oruinisation to evolve tron small family groups to large groups wilh morphological ifferennation among colony members, However, unlike other allodapines, 4. piedenraore esbibirs frequent und overt ugonistie behaviours among nest mates. Such agonistic behaviour tas offen been associated wall more primitively soci speeres, According la Wilson's (1971) erileria, & wridemiata can be lussed as highly eusocial because there female morphalogiea! dimorphisin associated with reproductive division Of labour, Pherelore, &. frdeweta doesn't cortorn to Wilson's (197 |) sugeestion hith averession within a colony can be replied by "gentle despotism’ as socklity involves larger eroup sive und requires a greater degree of integration, Most other highly eisociit species display distinct morphs WAH are directly associted with diserele behavioural costes. invelving Minimal or noageression Considering the presenee OF aeyressive iilenielions with) 2. trident colonies, if would seeny (hut incrensedvolony size ane the developracnl af mor Polo ditfercotiaton among colony mernhers need not he acconmpanied by decreased levels ol over iitra-eolony dye ression, Acknowledgments We would like to thank friends. and laboritory Inembers who assisted with field work and N. Bull, S. Reyes. PB Tuest, J. Bird aid bwo anonyitous relerees foriuviee on the manuseripl. This researeh was puirtially finded by wats from the Ansteatian Research Couneil to M, POS. Field work was curricd oul WH permission from (he South) Australian Depurtimeal af Environment and Natural Resources. Per Ney, Q23256-03. issued to M.S, Reterenves Alona fe CIM 74 Observational stuely al lelavieue sapling methoels. Bedaeine 49, 227267. Baton, So Wo To Sakagiiin S22. & Moen Yo boos) Relwivinwn al the Tndicn alladapine bec. retina Adie. i SeTUE psi dn he festh uh AD iste (Hy menoptenn: Anthoptiaridieh 2 Neascas brrtenid Sen Of, 345-460. Broce M1. Silverman. I ML ode hell, Wo (17s) Aponistiv Behwiour soci infericuons, aml hehavioueal specialisation im a primitively eusocil bee frsecten Sactaia, Cavis, IS, 351-364, brothers, Do & Michenern ©. 1D. (1994) Ttentetions ia colonies al perontively soc bees TH Ethalowy al division owl hike ine Lentiiglossi se plivrane (ymehopteri Halide) 2 Comp. Physi a |e Jan. Bick Nd. Minti. ACC. NORTMAIST, Yoo ubvEsIVN. B Lo & Souwary, M. PO (in press) Caving your duliehters Wie esve he quenthing reproductive dominance in a oprimilively wackil bee, fre. Ney, Soe Set (Serres B), Combo Hook Micueare, cof) (1983) Chvanistey aod Hichon of nanidibular ehind praduets af bees of the Lenus Laenenra (HY ienopterd: Anthaphoridieh A. Chen, Evol, 8 (825-1531, Tlonbsbeurn, Kd SonWwane. MT (it press) Chardin Spee saion jn pre-repramuenve Colonies Pine etl bochap yi ie bee Kavtenre bicalvn dal boul eval WW Hrnsres oT, F197 7) Nesting: Woloy of fhtec alocappite: hoes WE THE subgenis Crema Michener Pins, RL Sey SN day TOE G1 ba BEHAVIOUR IN AN ALLODAPINE BEE 63 Hurst, P. S. & SCHWARZ, M. P. (1996) Morphological differences among females of the eusocial allodapine bee, Exoneura trideniata (Hymenoptera: Apidae). Proceedings of the XX International Congress of Entomology, Firenze Italy 25-31 August, p. 413. JEANNE, R. L. (1986) The evolution of the organisation of work in social insects. Monitore zool. Ital. 20, 119-133. Maeta, Y., SAKAGAMI, S. F. & MICHENER, C. D. (1992) Laboratory studies on the behaviour and colony structure of Braunsapis hewitti, a Xylocopine bee from Taiwan (Hymenoptera: Anthophoridae). Univ. Kansas Sci. Bull. 54, 289-333. Mason, C. A (1988) Division of labour and adult interactions in eusocial colonies of two allodapine bee species (Hymenoptera: Anthophoridae). J. Kansas Entomol. Soc. 61, 477-491. MELNA, P. A. & SCHWARZ, M. P. (1993) Behavioural specialisation in pre-reproductive colonies of the allodapine bee Exoneura bicolor (Hymenoptera: Anthophoridae). Insect. Soc. 38, 1-18. MICHENER, C. D. (1965) The life cycle and social organisation of bees of the genus Exoneura and their parasite, /nquilina. Univ. Kansas Sci. Bull. 46, 317-358. (1972) Activities within artificial nests of an allodapine bee. J. Kansas Entomol. Soc. 45, 263-268. ——— (1974) "The social behaviour of bees: a comparative study" (The Belknap Press of Harvard University Press, Cambridge, Massachusetts). (1975) A taxonomic study of African allodapine bees. Bull. Amer. Mus. Nat. Hist. 155, 67-240. SCHWARZ, M. P. (1994) Female biased sex ratios ina facultatively social bee and their implications for social evolution. Evolution 48, 1684-1697. & OVERHOLT, L. A. (1993) Methods for rearing allodapine bees in artificial nests (Hymenoptera: Anthophoridae). Aust. ent. Soc. 32, 357-363. WILSON, E. O. (1971) "The insect societies" (The Belknap Press of Harvard University Press, Cambridge, Massachusetts). FIELD ECOLOGY AND BEHAVIOUR OF THE EGG PARASITOID TRISSOLCUS BASALIS (WOLLASTON) (HYMENOPTERA: SCELIONIDAE) By S. A. FIELD*, M. A. KELLER*® & A. D. AUSTIN® Summary Field, S. A., Keller, M. A. & Austin, A. D. Field ecology and behaviour of the egg parasitoid Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae). Trans. R. Soc. S. Aust. (1998). 122(2), 65-71, 29 May, 1998. The ecology and behaviour of Trissolcus basalis (Wollaston), a parasitoid of the eggs of the horehound bug Agonoscelis rutila (F.) and numerous other species of pentatomid bug, were studied in the field over two years near Adelaide, South Australia. The adult bug population declined sharply early in summer due to the combined effects of senescence of host plants, egg predation and parasitism by T. basalis and a sympatric species, Trissolcus ogyges (Noble). Hyperparasitoids of T. basalis were recorded for the first time in South Australia. Key Words: Trissolcus basalis, Scelionidae, egg parasitoid, horehound bug, Agonoscelis rutila, egg masses, defensive behaviour. Treailvdetionts af.the Bowl socrety of S Mest C2998), 122¢2), OS-71 O35 FIELD ECOLOGY AND BEHAVIOUR OF THE EGG PARASITOID TRISSOLCUS BASALIS (WOLLASTON) (HYMENOPTERA; SCELIONIDAE) by S.A. Finep*®, M.A, KELLER® & A. D. AUSTIN ® Summary Rib boS AL KELL ML Aw ADs in, AOD. Meld ecology and behaviourof the euy parasitoid Disses Masalis (Wollaston) (Hymenoptera Scehontlaes 7s. A, See, S. Aust. (YOK). 12242). 65-71. 29 May. 1998, The veylovy and belay of Frissadens baseless (Wollaston, i pairisitaid af the eggs af (he horehound buw Agenoscelts ratily (VO aid numerous other species oF pentatamid bug, were studied ithe Held aver hwo years ned Adehtide. South Mustialia. The adult bus population declined sharply curly in summer due to (he Combined GHects of senescence of host phints cae prechition and parasitism by 7 bawafis and a sympatric species. Trisyolcny peyges (Noble). Ayperparisitolds of 2 basis were reconded for the tiest tine in South Auseratia, Compennon among femule purusitonds for ueeess io host age masses differed widely between the Gwe years (sume season). and lemiles displived uduptitions ( earpatition, They patrolled host cae lasses where alone and defended then ugeressively in divect coests with conspeciics. These ohsenwitions rejaforye previous liboialory Work ond suaoest hither avenics al peseueh on the behavipural sinilegies used by f. brasalis during Helerce al cue miisnins Ky Wovon: Cielo busulis. Scelionidie. eee parasitoid, horehound bug, Agonaseedis eunila, eo Wiisses. Uelensive behaviaur Introduction frissoleuy bavalix (Wollaston) is uo solitary parasitoid of fhe wees of the itrodueed preen yeectuble bus, Nesare viridila (Loy (Hemiptera: Pentitomidae), dnd aw number of other pentatornied species (Cimber 1964), ineludim Lhe qurive horehound bug. Avancayceliy come (EL, Sinee its first WMpertation tito Austria in 1933 (Noble 1937), 7 basaliy has heen released a number of times (Clarke ISM) Due lo is perceives! importince as a biocontrol avent worldwide. many aspects af its hinlogy trave been documented (e.g. Wilson L96t, Cumber 164: Powell & Shepard 1982: Bin ey af. 19s6; Volkol & Colazza 192: Matiaeci ef al 1904), Although the field eculogy of 7 haselis in Australi is best Known Hon its assocnition with WV, Write We Turner 1983, Clirke 1990), its biolagy when parusitising A. futile on the iitrodueed weed Horehound, Merrubriun vulgare (yh has also been Tivestimsited in view of tts potential for maintain purus numbers in cropping areas (Kelly |987) When feeding on horehound. reprosuetive maturity of AL varity is dependeat pow the availabiliy of Hlowers, and so both hust and parasitoid populavon dynamics ire closely linked to Seasonal eyeles of plant growl, Although activity and populion peaks Ol both host and purasitonl comeide with the major srowth phase of (he plant in spring and early sumer neither species aippears to unter dhapause Departinit al Crap Protection. Waite Coonyins Pho Lirtiversityot Adelie VO Cer Csengiud Si Nest SClie, over winter and a high rate ab parasitism (> 70% ) is maimained throughout the year (Kelly }987). Due to the smallsize and rapid movement of 7 hevelis. field observations are difficult and dati have only been collected foi) one study on host searching under semi-field conditions (Turner 1983) Oviposition behaviour, exploiiwion of host egy Intsses (= pulches) und competition have not been studied in the field. This paper reports fell data on the ceology and behaviour of 7) basalis parasitising Ac ruil, Us most common hosr un the Adeliide region OF South Australia (Pig. 1). In spring and summer OF 1994-5 und 1995.6, date were collected on the seasonal Tuetations of fost plant and host populations. sources OF host mortality and behaviour Of parasitoids as (hey exploited, and competed tur. masses of lost cees. The purpose of this work was ulso to provide the Fetundkition Tor more detaded laboratory-based stidies ol patel exploitation and defence (Micld ef af. 1907). Materials snd Methods Hast plants anel tiases Data on the ecolowy and behaviour al 7 basalis i (he held were collected trom late October to late March in 1994-5 ind 1995-6 in the Brownhill Creek Conservation Park, in the Adelaide foethitls, Sampling sites were selected by taking Lwo 500m Iradsects dloug candor direchions through a patch ol horchound, The transcets were divided inte 10) intervals of equal length and a random paint was taken along cach fferval, The nearest horchound pling, or discrete cluster of plants. to cach of these porns was marked ws a sampling site, [stems veased fy S.A, PIELD.M, A. KELLER & A.B, AUSTIN Competing agq parasitoid: Thssaleus ogyges yb ey *S : *! Host plant 14 Horehound 4 Marru brtum vulgare 4 ruiiis.eaa mess {hest patch) —— ‘Alternative host plant: Salvaton Jane. Ecthum plantagnaeurnt | Vis. 1. Surmoiary of the natura history of the A. radile to show any green foliage during sampling. the marker was removed and replaved on the nearest stem bearing foliage. When no phints with eight or inore stems With green folipge remuined, sampling was discontinued, On 21 days between 26 October and 2) Mareh 1994-5. dala on host plant and host population, and parasitoid behaviour were collected, The numbers of open towers on stems were recorded and ai ides of the A. Afi population was obtained by counting the total number of adults on all simple stems at LO.00 am. Parustioid egy lee The nuinbers of eges earried by female parasitoids (n= 31) were assessed by collecting wild 7) fetsetis females on seven days between 30 November and 20 December in 1994, and dissecting them in the laboratory, In addition, exe maturation under laboratory conditions was studied by determining parasitoid egg load when females were between one and 10 do old. Prior to disseetion, wasps were held individually in vials supplied with honey solution for L-10 do at 25° CL without ovipositing, before being frozen at -60°C, The metasoma of individual wasps was removed and dissected ina drop of water on a eavily slide and the number of eves in the ovaries wournted, Behavioral observations In both years, behavioural observations on parusiloid oviposition behaviour, patch exploitation und competion were made. Host patches were created hy glueing O-1 d-old ege mitsses of laboratory-reared AL vad, each containing between Eqgq parasitoid wasp: Trissoleus basalis — Hyperparasitoid wasy: Acres es sp. we Main host insect: Horehound bug, Agonoscelis rutila ‘Natural! host insect, Green vegelable bug, Nezara virdula \ Alternative hostinsccts | ~-~~—__ Cermatuius nasal Oeshalla solinellanterg: L leiscilis system used in this study (see text for deserption) 12 and 24 eggs, on to small squares of green cardboard and stapling them to leaves on randomly chosen sample stems, Patches were laid ont belween O.O0 um. and 2.30 pm. and the oumber of female wasps on cach ege mass was recorded every 30 min until O30 pam. tn 1994, between one and 12 exp masses were observed on each of five ditys between 26 November and 12 December. In 1995, 16 exe asses Were Observed on cach of eight days between & November and 26 December, An index of daily competition for eye masses was obtained by taking ihe maximum number of wasps observed in any one sumple during the diy for each exe mass and calculating (he mean aeross all egg masses, To facilitate Comparison between data sets for the dillerent years, hese means were taken it (he period 2.30 p.m. to 6.30 pam. as some data from 1994 were collected only during these times of day, To compare rates of disvovery of egg miusses, Kaphin-Merer vstintes Of survivor functions (Haccou & Meelis 1994) lor the time until discovery of egg masses were caleuhued for data pooled within seasons, ‘The survivor functions plat the cumulative proportion of weg masses discovered its a function of time, and this provide an estimate of the instantaneous rate ol discovery of cap masses. To observe pateh exploitation and) defence behaviour in detiil, pitch visits by single wasps (= 4). amd by pairs of wasps (n= 6) to randomly selectad artificial patches were videotaped und converted to behaviour sequenee records in the laboratory using a TRS-3O Model 100) portable computer programmed with event recording software (The Observer, Noldus Information BEEAVIOU RAL ECOLOGY OF TRISSOLCUS BASALIS Technology Wageningen The Netherlands). Behaviour was divided into categories representing host @aimination, oviposiuon. puleb-leaving and, for pales of fenmes.agonistie behaviour (Field!: Field in press) When patch contests between two femules one individual (he resident’) usually estiblishes dominance and aggressively exclides the other (ihe intruder) (Watson L961, Pield'). The Mttuder then waits nearby and periodically returps to (he cue mass fo allempl further oviposition, Where bout deneth simple sizes pernrilted (n> 20), intruder ‘retreat’ behaviour (detined ay the time between being driven away Hom the ege Mass and returning) Wils tested Tor abrupt chinges: (i bout lengh using a HON pPUVMNetne multiple change point lest (Haceou & Mechs 1994). Where changes were sinifeunt at the udjusted levels saweested by Haccou & Meelis (1994), they are usteated with cumulative bout fongth plots. Mec, Results Hast pitts, hosts, porasttords aned liyperparasitoiads I) 1994 ink 1995, host plants und host insect populations underwent marked HMuetuations. Natibers of lowers peaked in late November, and therwuller declined steadily Wich mid January (Pie. 2) when all plants showed very Jide or ne green yerehition und no flowers or leaves. Popukiion counts of A. rita deereased in’ parallel wath: the deeline in plant quality, stabilising (ic low devels in mid—Kinuary (Pig, 2) Addit A, cidi/a also appeared to be susceplible lo high femperuiures, as many died during dhol. windy spell carly in December. The lirst Hyiiphs appeared at this time, indicwting the vibergence OF The first weneration of the season, Numbers of adull ad, rafiiae over summer remained nich lower thin the peaks observed in sprog, Weeds other than horehound. in particulir Salyvaiion Jane, Mehiane planteaginiia (L.) were alse abundant at the fieldbsite in (995-6, and were utilised for feeding ane reproduction by A. rutile (Fig 1), Adults were observed leeding. mating and eying eyes On 7 plieeinesen, allhounl ip wits uitelear whether pymphs were able to complete development solely on this phint. 7rixsoleus basealiv were also vbserved foraging on A. plantergineuin plants, Apart fram 4 puffle. other pentatomisdts accusioially observed ou horchound at the study site were N.oniridite, Cermeatulis neasaley (Westwood) uml Gechalia Schieber (Cuérin-Méneville) (Fig 1) alhof whieh have previously been recorded vs hosts Tor 7 havelis. Rees Ob ON, viridider cid C. I Bae S.A CU) Patel eQphoruiid cmt dletemeas (ip the exe Pibiitonl Frases Hii Wolken Up iteepiene Sev Honthue) MAD Hiisis. Phe Criversity al Adehiishe Canputs Total bugs/80 stems o — —— Adult bugs = Flowerheads Mean no, flowerheads/stem 1) OS Fem A 7 Vig. 2. Numbers of a raila adults sind mean jurbers ef Nowerheads (47 SD) on horchound sfetos in 1994-95, 10 it) 2 6 4 Age (days) Fin. 4 Mean number of eaes (+1 SD) ii the ovaies at lomitle 7 baxalis trom [10d after enemenee Nunihers below error hues indieale sample sivas nasdliy were Nol seen bul those of CL wehaedenbury were colleetud and both To beaselis and Trisseleres avyues (Noble) were reared from them (Fig. 1) Trivsoleuy oavges did not complete vevelupment i A. ridley eves, so O. selnellenbergd appears to have been its min hostal the site. Triwoleuws ovy ees could be distinguished from 7 easaliy in the Held and laboratory by @ distinet difference in host marking behaviour, Ruther than dragging the ovipoesitor smoothly ever the hosteve ia igure 87 mation as in TD) basaliy (Wilson 1961, Weber etal L996), 7 egvges Woved the ovipostlor horizontally aeross the oge With a “bouncy, jagged” motion, Females defended eye misses similarly lo 2 bavelis and Hilerspeciie vontests were observed. bul nor recorded in detail As ne objective of This study wits fy gauge the overall levels oF competion far aveess lo hosts among 7. hawaliv, the data presented below inglude Observations in which J auvees was alse present Tp 1994, The proportion of loti observiligns in Which 7 veyves occurred was nol recorded but in JOYS TL was upproximilely |O%, In addition to the primary parisituieds. females of the hyperparasitoid devevdiseides sp. (Ciraule & Dodd) — (Hymenopteri: — Pleronmalidae) — were vecusionilly Observed silting an egy Masses fon) December 1904 to Linuary 1985 (ria. 1), OA SA PIBLD. M.A. KELLER & A.D. AUSTIN Parasitotd eae lente Disseetions of Liboratory-reared female 7 hasaliy revealed (hal they emerged with a substantial complement of eyes and then slowly mittured firther cogs over lie (Pig. 4). showing that 2 basis isa syHoVigere species, Most of the parasitoids collected From the field were carrying substantial numbers of midure egys (Fig. 4). indicating that ege-limitation was not common inthe field at this time, In curly November in 1994, three different fembles that discovered egy inasses inthe afternoon reniined On then overbight alibough no intruding eonspecilies were observed, Two of these females were disseeted and found te have exe loads of one and three eggs, respectively, Behavioural observations Yirastioids searched for hosts by flying between horchound stems, and then walking rapidly up and down the stems, palpating the surface with thei antennae until they had located an cee mass. While scurching a stem, they offen passed within a few centimetres OF an ese mass without detecting if and so-did nol appear to be detecting exe miusses using visual orchenical cues, Thus, location seemed to he by physival contuch Upon contiaeling un ese mits, Wusps examined unly one ob a few host ewes. then commenced oviposition (Bin ef al. 1994). 1 one or more conspeeiies were present, wasjys engaged in agonistic behaviour (Wilson 1961), Competnion for egg masses differed widely herween the (wo years (Pigs 5.6). Tn 1994, there was a peak of piwastlold activity Orn December 2 (ig. 5). On this dale several remarkable Observations af parasitoid competition were recorded, Th one instanee. maximo oF 14 parasitoids was observed simultaneously Competing for aceess loa single ege Hass. dn another observation, five purasituids bad discovered an ese mass as it was being laid by the female AL rutile. They were following the bug, parasitising the eaes immediately they were laid aod Eqg load Fig. 4. Mature exe loads of female To basaliy collected benveen 30 November aml 20 December pn (994, were fighting each other for possession of the incipient egg muss. Lhree other cases of immediate patch discoveries were recorded on the sume din. Avenoseelis ratila showed only rudimentary delensive behaviour, occasionally kicking at the parasitoid but this had no deterrent effect on the 7 dasaliy, Instead, the parasitoid sometimes responded by directing its aggressive behaviour toward A, rulilid. In contrast to the high peak of competition observed a (99d. gompetition remained Tow thronghout the entire sampling period m 1995 (Pig. 5), This difference in intensity of parasitoid aeivity between the Iwo years is alse reflected i the time until disvovery of patehes (Fig, ©) In 1904-95, almost allege masses were discovered within 7 hat being laid, whereas in 1995-96, the vast majority of eve masses remained undiscovered jy the same time period, resulting ina highly significant difference between the vu curves (Log-rank test, e* = 100.9. | dh. p< 0.001), Sample sizes for continuous time records of patch exploitation and defence were small and observations were sometimes incomplete. precluding un extensive anmulysis of the time and sequence strueture of behaviour Flowever, the observatrons did confirm that the patterns of behaviour seen in previous laboratory studies (Wilson 1461, Cuniber 1964) rellected those oceurring under Tield conditions. When alone, wasps successively examined, oviposited in and marked hosts hefore examining the surrounds of (he cee mass and finally leaving. dn cach of the four observitions. sell: superparasitisen (i.e. double oviposition in the same host cog by the same femule parasitoid) did not oecer before the egg mass Was fully depleted (ic. all host eves in the eve muss were parasitised), and only becurted after that in one observation, when the wisp sell-superparasinsed three times before leaving, Upon depletion of the pateh, in two of the Tour observations wasps embarked on penods ol Mean (Max. no of wasps) Hig. 5. Comparison at patel competition herween years mean (+) SD) af maximum noniber al 2 devalis onary Ce THUNS BEHAVIOURAL ECOLOGY OF TRISSOLCUS BASALIS oH —— 1994/95 —— 1995/96 % Discovered Time (hours) lig, @. Comparison of pate discovery limes between years Tor 1994 (26 November -12 December) and 195 (23 Novernber- 14 December). ‘defence’ af (he egg muss. These defence periods consisted oF alternating bouts of ‘stabonuary? behaviour (motionless, sitting on the ege mass) aod ‘patrolling’ behaviour Gripily darting from one side of the ega mass to the other). This apparently pre- cnipuve pateh defence behaviour continued for upproximately Tl omin and 2h 30 min in the two observations, respectively, despite the fact that no comipelilors Were present. When contests between two individuals occurred, sequences Of agonistic behaviour developed. These sequences exhibited the same major characterises as those observed by Wilson (1961) and Cumber (1964), including the establishment of resident intruder roles. Fights occurred cither on the first encounter, or after a brie! period of mutual tolerance, The tendency for individuals to light appeared to Ncrease alter successlul completion of one of more ovipositiens, although occasionally individuals becume aggressive jmmediately upon arriving alan eer miss, and before examining the host eggs or oviposifing, Following the onset of ageression tind role establishment, the itrider usually retreated tor the underside of the leaf when attacked by the resident, out of view of the resident (eae fapses were always phiced on the upper side of the teal), Cumulalive boul lenatty (fH) 18] n wu ao iu HU (io hb Flout number hig. Comiiative beat leverh plat fin crete belay vai OF omiculer im opuiewise vantest showed two ahnapt Chee Th Bout ena Tesi an Hieeedise Chere grraw yen a deereused Gripe arrow), Cumulative bout length (h) c [=> _ iv] § 10 15 20 25 Boul number Mig. 7 Cumulative bout length plot for retreat behaviour of the iitruder ti a paiiwise contest. showing a single ubrupt increase in bout lengahy Garrow )- searched the or fed on nectar in und cither sul motionless, groomed, surrounds OF the eee ass, Nowerheuds. In two cases Of pairwise contests, abrupt changes in bout length were observed. Inthe first observation (Fig. 7), the intruder switched from short to long retreat bout lengths, suggesting that it had ceased to compete for possession of the cue iass, and had begun wailing for the resident lo leave, In the second observation (Fig, 8). which lasted 36 min longer the intruder swilched twice: lirst to longer relteat bout lengths, and then back to shorter ones, which were SUT longer than those inthe initial period, The intensity of bouts of aggression varied widely: from no-contest encounters in which one individual retreated without retaliation, to infense escalated fights. in whieh individuals locked together and rolled across the leal for up to 300s. Although no obvious injuries were observed, sometimes both individuals tell from the teal, temporarily losing Opportunities to exploit the egy mass amd leaving i unvturded, Although some individdsls recovered their position on (he egg mass quite rapidly. others did not return, showing that in addition too the possible risk af injury, engaging in an escalated fight also entitled the risk OF permanently losing HWecess 1 the eee pass, Conspecilic superparusilisny (oe, oviposition tn the same host eee previnusly parasidised Once or mare by a different female parasitoid) was common, either Wiel Wasps Visited cae iasses sequently or simualiineously, In one extreme case, a mass of 20 set oul oon the day of peak competion (December 2) in 1994 recenved 126 ovipositions over p period of 9 hh, Discussion The host popuhition observed im this study underwent marked Puctuarions Populitions peaked i ospriig. bot eges hed ly the overwintering population ob adulis ab this time seasonal vw SN PIED MA KEELER A A DD ALS TIN tiered high levels of predition und parisitisn aid few pyniphs cmerged to form the new generation, Hhis mortdity, combined with) the deeline tn food avaihibihity resulting frond the senescence of host plants, Ted tou sharp decrease To host population levels by Tie nd Gf the December Therefore, bor the parasitoid Gere was a peak iy absolute plumbers ot host exes dyailable for approximately tWwer ionths front late Qetober to early Deeember Alita depletion ol wee reserves mi 7D hesaliy was rare during this period Th inay hive had significant efheets On forum Behoyiour when (lh oecupted, eaustie wasps fo shay leger and fund ese misses. rather Thun Contin Faring. The eteunetnce oF hyperparasitism ly Avrrcliveides sp. may also hive exerted G seloetve pressure Or wisps awit low cae load. causine twendeney rewired staying dnd giuinding egg passes. Aerlyiidey spi previously been reebvercd Tair eve jWasses ool Vo viride it custern Australia by Churke & Seyntone (1992), These quthars shawed (hil the species isa fy perparusttont of 7) bawalin, bul sugeested iLmay be only casually associated with 7, hasalis, Clarke & Seymour (1992) did not repert Which stage of Z drisetiy the hyperpanisitoid attacks bul in observations made i Wie present study: wasps only sal on (he exe masses withbul probing or OVIPOSMp, Suggesting that they may have heen wilting for the immuiine 7 beweliy da reach a hater larval stage, ar the pupal stage. Attempts to Observe behaviour oF the dereefiveides: sp. incl cour it in Whe lubaratory were vosuceesstil, as ik was only dbserved a few Uines in the Jield and: was oever recovered from A, itil Gee MUSKeS The Trequent ocearrenee of superparasitsny by 7 beiliv in Whis study provides lurther eymence that superparasitien) is vominTon iT nature Canssen [Yad van Alphen & Visser 1990). tis eeolowival importinee far Lo hitsaliy as underscored hy the Hichine That (he probability of The superparasttis ie Terie oblige offspring (oii a host is very high vt she Superparasiises soon aller the other fennale oviposits inthe host Wielledal 1997), As Wis leaves the oftspring of the first female ta oviposit al subsuintiil ask. the high frequeney of vecarrenee af stiperparasidisin and its high fitness pay-allrnay Have favoured the evolution af pateh defence in F baselis. Th adition lo the Rack thal Gee pousses are of a delendable size, us noted by Winige (1982), The drop in the pay-ofl fren) superparisilisny lowards vero ifter & ob (Kield er a 1997) may explain the observation that fenules aiborted Gviposiiign in an eae uss tit had Been parisitised as much as 9 h earlier Still this raises the question of the Mechanisin af host discrimination, which needs ta be pursticd ie Pitare: studies, Observahions of puleh exploit ancl delence revedled behaviour patterns simi kin te those seen i previous laboratory studies (Field!) and sugwest init belaviote is adapted to bial levels of competition, When alone on an ee miss, wasps engaged in extended periods of defensive behaviour boll before and alter the eve irtss wits Fully depleted. suggesting a high ie expectation” of conspecifics. arriving and superparusiising. Chis behaviour could result from oa combination of dinate expectation of competing set by natural selection. aid a flexible response based On experience from previous eez messes, Alted restdentypteuder tales were established tn pairwise contests, Tilruders showesl ubrupt vacredses in periads of rethaat behaviow CUFT COMENES wh Walled OUL OL Visto Of residents (ander he dear), suggesting Whey were waite for residents Wy leave the eee tnass. Alihiweh patuder hehavinne was pot followed jo deri. tn the second comeust analysed (rig. ®t ts probable tat the fest switel resulied from the intruder rikiligh an extensive search Of the sUPOU ine are Wy check Cer other Uinoceupied: cyan rasses, uneb the second: (roy its retuuming to the origimal patch, Exploring the SUPPOUNd ws Lor aHeniiive HOST Gee masses 1 he current one May be particularly important wher sihy (nubvidaals ure Competing Tor a) Ege rns ard giining access iS difficult: a Siteition thal was Hequently Observed in-times af peak eompetirioan The dynamics of patch delence im such sittiitions invelving more thap lwo individualssire contplex dn! difHeull lo fifer trom: the present study, Altiougl subslantial Muetuations in the levels of compelition (lo beet and conipetitian is someimes less iitense, periods of extreme Competition muy nevertheless have played a dmpertiadt tole ta the evalytion at foraging behaviour. The Observations made in ibis siidy reveal sere Hovel patterns TH Lie agonistie belmwiour of 2. Leselty dnd indicute the Appropriate ceolovicn) context a WHICH Lo tivestigane them Tureen ie the liberatapy The factors leading to the onset ofigeression did the Mechwisiis Ob Coplest fesoliton Che. eslablistiment Ol resident-intruder roles) hiwe rarely been studied ny parasitoids (bul see Petersen & Tardy T296) and are the subjeet oF current jvesdivation iin 7 heryeelia (Pield & Calbert unpub, Also, a qiuntibative Mi Vsis. OF The Hime-struetire OF interactions betwee Hehting purasitioids fas nol preyiwstly beer allemnpled. and the present results mmaicute (hat (hs muy he both possible and ol wreat interes! to 7 Pasatin. Acknowledements feo wish to thank G. Thoms fer the tse ol bis property, C. Culbert for programining the multiple chinge-point test and 1. Mound, M. Kakkinn and [ Bird for eriticisms of the manuscript BEHAVIOURAL ECOLOGY OF TRISSOLCUS BASALIS TI References VAN Atpibn, J. J, Mo & Vissrk, M. E. (1990) Superparasitism as an adaptive strategy for insect parasitioids, A, Rev Ent. 35, 59-79. Rin, F, Vinson, S$. B. STRAND, M. Ru. Conazaza, 8. & Jones. W. A.J. (1993) Source of an egy kairomone Sor Trissedcus bayalis, a parasitoid of Nesara viridule, Physiol, Ent, WB. 7-15, Charkh. A, R, (1990) The control of Nesura viridula 1 with introduced eggs parasitoids in Australia, A review of a ‘landmark’ example of classical biological control Aust. Jari, Res. 41. 1127-1 146. — & Srymour., J, &. Aerocliseides Girault and Dodd (Hymenoptera: Pleromalidac) parasitic on Trissolens bayalis (Wollaston) (Hymenoptera: Seelionidae). a parasitoid of Mesure viridila (lL) (Hemiptera: Pentatomidae). 4 Ausi Ene See, SL 299-300, Cimapk, Ro A, (1964) The egg pitrasite complex (Seelionidae: — Tymehoptera) of shield bugs (Pentatomidae. Acanthosomidae: Heteroptera) in| New Zealund. NZ J) Sei, 7. 536-554. Piteb, S.A. (in press) Pateh exploitation. patch-leaving and pre-emptive patch defence jn the parasitoid wasp Trisseleus basaliy (nisecta: Scelionidie), [liology: ———. Kenner, M.A. & CALBERY, G. (1997) The pay-off from superparasitism in the cee parasitoid Trryselcus basatis, invelation to patch defence. Aeol Kur, 22, 142 149. Goprray, A. C. J. (1994) “Parasitoids, Behavioral und Fyolutionary Leology” (Princeton University Press, Princeton, New Jersey). JANSsiN, AL (1989) Optimal host selection by Drosophila parusitoids inthe field. funet Beal, 3, 469-479, (1992) "Two species of Marriacet, 1... VINSON, S. Bo WILLIAMS, HJ, ALbrici, J, Ro & Bin, (1993) A long-range attructamt kairomone for egg parasitoid Trrsselous basalis, isolated trom defensive secretion of is host, Nesara virtdula. Chene. Evol, 19, (167-1181. Nosik, N.S. (1937) An egy parasite of the green vegetable bug. Agric. Gaz. N.S.W. 48. 337-341. Prrirsin, G. & Harpy, 1 C. W. (1996) The importance of being larger: parasitoid intruder-owner contests and theie implications tor clutch size. Ania Behe $1, 1363- 1373. PowenLl. JE. & Sumparp. M. (1982) Biology of Australian and United States strains of Trisyeleuy Paxalis, a parasitoid of the green vegetable bug, Nesara viridisse. Aust. J, bel 7. 81-186. Turner, J. Wo (1983) Influence of plant species on the movement OF Triwelcus basaliy Wollaston (Hyimen- optera: Scehionidae) - a parasite of Mesure viridula Let. Aust. ent. Soc. 22, 271-272. VotgKork, N, & Conazza, S, (1992) Growth patterns of teratocytes Mm the inimiature stages of Trissadouy bewelis (Woll.) (Hymenoptera: Scelionidue). an eye parasitoid of Nezara viridula (1) (Heteroptera: Pentatomidae}. fat. J. Insect Morphol. & Enbrval. 21, 323-336. Waact, £K, (1982) Sibmating and sex ratio strategies in scelionid wasps. Keol But. 7, 103-012. Weber, C. A. SMILANICK. J. ML, Ennnr, LL, EL & ZALoM. B. G. (1996) Ovipositional behavior and host dis- crimination in three scelionid egg parasitoids of stink bugs. Biol. Control 6, 245-252, Wir son, (1961) Adult reproductive behaviour in Aweleus hasalis (Hymenoptera: Scelionidae). Aust Zool. 9, 739-751. WOODWARDOSTRONGYLUS PETROGALE SP. NOV. (NEMATODA: CLOACINIDAE), FROM THE STOMACHS OF ROCK WALLABIES (PETROGALE SPP.) FROM ARNHEM LAND By I. BEVERIDGE* Summary Beveridge, I. (1998) Woodwardostrongylus petrogale sp. nov. (Nematoda: Cloacinidae), from the stomachs of rock wallabies (Petrogale spp.) from Arnhem Land. Trans. R. Soc. S. Aust. 122(2), 73-78, 29 May, 1998. Woodwardostrongylus petrogale sp. noy. is described from the stomachs of two species of rock wallaby (Marsupialia: Macropodidae), Petrogale concinna Gould, 1842 (type host) and Petrogale brachyotis (Gould, 1841) from Arnhem Land, Northern Territory. The new species is distinguished from congeners, W. woodwardi (Wood, 1931) and W. obendorfi Mawson, 1976 by the presence of four pairs of oral denticles compared with six pairs in W. obendorfi and 16 pairs in W. woodwardi, by the spicule length which is 0.90-1,07 mm in W. petrogale sp. nov. compared with 1.4 mm in W. woodwardi and 1.7-2.1 mm in W. obendorfi, by the length of the female tail which is 0.22-0.23 mm in W. woodwardi, 0.18-0.22 mm in W. obendorfi and 0.11-0.17 mm in W. petrogale sp. nov. In addition, the vagina is 0.7-1.0 mm in W. woodwardi, 0.8 mm in W. obendorfi but only 0.31-0.48 mm in W. petrogale sp. nov. The characteristics of the genus are considered as well as its relationships within the Cloacinidae. Key Words: Nematoda, marsupials, rock-wallabies, Petrogale, Woodwardostrongylus, new species. Tre ans ef the Revel Somaya So Auge, (L998), 9222) 73-78 7 WOODWARDOSTRONGYLUS CLOACINIDAE), FROM THE (PETROGALE PETROGALE SP. NOV. (NEMATODA: STOMACHS OF ROCK WALLABIES SPP.) FROM ARNHEM LAND by J, Bhyeringn® Sumnuiry BEVPRIDOD, TP (TOUR) Wanelivcdostndiy petoagale sp, now (Nematodes Cloacinnlie), fromthe stomachs 0 rock wallabies (Perrouele sppo bron Armbent Land, Tres We Soe. XS Meare, 122 (2) 75-78 29 May Ys. Woodiidosnuney los petal sp. nov ops desernthed: from the stemaehs ol iwo species a? roek walhiby (Marsupiatia : Maeropodidie), Perogale conenuid Gotld, 1842 (ype hosti and Pariwdle becliveniy (Gol, PO) trom Arnhem Lund, Northern Territory. Phe new species is dintingiished from Gongeners. Wo wrcedtvesdi (Wood, LOS 7) ae Wi endian de M vwaert L976 by Te presence of four pps al onl denticles Compared willy sty pens ie WE oPbeydeedi and 16 pars in Wo woedword:, by the spicule tone th whitch ts 0.90-L07 min in i perraueate ap ney, conipared with Frm Wo ypeodiweritt and 17-20 im in Wo obentorfi, by the lentil of the female Toth whieh is O. 22-0, 23 lo We teed. OTK “27 jan in WE ebcudoeti and QE, Pim WW, petreigale sponoy tnaddibonm the vagimiiis (7 DOs We hod Ue inn in Wo abendor? bitonty 131 a8 iin WW. petiagele sp. ney. The characteristics of the genus-are considered as wells ibs relationships within (he Cloweurilie Kiev Woks. tiiroduction One ol the most dnusudl eenerd of nematodes OoC Une Wy the stomaehs ot Kirigarous und willibies is the cloacinid genus Weeewurdn Mraneviiy Wahid, 1964 which ts found in tannels i the superficial squamous epithelinan of the stamach und Oesophagns rather than ja the lane ob the stomuch oor hare maitestine oor coiled around vesuphageal papilliwas is the case with most of phe other members of the Eimily (Woed 193.2 Miwsen 1971, 1976; Beveridoe Ac Spratt 1996). Two species are cirently Keawe willl this genus, the: type species, WM weerdwird? (Wood, 193.1) origtimally deseribed From Woodward's wallaroa. Moers mens Weeder? Thormias. LOOT, frond the north Wo Western Austrilia (Wood [98d. Wahi 964), slibsequcnlly redesenbed from Pearson Island: rock wilhibies, Petrogale lafeclis pearson’ (Thos. 1927) Troms Soul Australie (Mawson LU7 by ane We dhendori Mawson. 1976 froin tre whiptiiled williby, Maeropis pares) Beret, ES3S (ype host, theewalllaroo, AL robust rebuatis, Gould, (S41 and the red-neeked wallaby, AA rifowerseus (Desmurest, IS17), trom: north-eastern New South Wales ane souti-eastend Queenshind (Maiwsoo 1976). The litter species Wits subsequently reported as a comiion parasite Of brush-railed rock wallabies, Mmefibers of the Mucovule poeniciiaa complex (Po aystiniliy Kuinsay. TA77. 2 eodman Thomas. 1923, 2 herberti Thoms, 1926, 2 iernate Gould. 1842. 7 muireeba Eldridge & Close, 1992, BO penned (Gray. 1825) and Po shaman’ Eldridge & Close, J992) Tron eastern Queensland by Beveridge etal. Deparment of Verrinery Semmes, The Liveenty al Melbourne Purkville View A062 Neruda, (hirsupluls, ruck-willubtes. evade. Weed iiyreiiis. new species (O89) aind bus since heen found alser in the agile walhiby, Mecropus cudis (Gould, TS842), und (he swamp wallaby, Willabia bicolor (Desmurest, S04) (see Spratheral 1991). Spratt eral (1991) dsted ain additional andescribed species Ob Werle: stiome vis from the nabarlek, Petourle concn Gould, [S42 and the shori-eured tock walliuhy, P hrachyenis (Gould. E84), fram the Northern ‘Territory. Tnothe present paper the undeseribed species fisted by Spratt es a (1991) is deseribed, te didanostic feattites Of the eens are reassessed und iis poston within the existing classification is discussed since the venus bas in the past heen variously allocated to the Strongyloiden (Walid 1064) and the Trichostrongyloided (Mawson 197 1), Materials and Methods Nematodes were recovered fran’ dhe preserved cureases OF rovk wallabies provided by Dr J. bk, Nelson, Monash University Melbourne Vie, Immediately afer shooting, curcases were perlusca with TO% formol saline yin the Jel ventricle lullowed by immersion of the entire carease in LO% formalin, Nenitodes revovered trom the gastric Mucosa Were slored in 70 ethanol and were clawed in hiclophenol for cyanitiation. Drawings were made with the amd of a drawing tbe atlached to an Olyinpus BH microscope, Measurements are presented in mmous the range fer 5 specimens followed by the mean in parentheses. To examine the localisation OF nematodes within the SUstric MUCOSA small pieces of purasitised stomach wall were embedded i ip was and sections, cubata thickness of S in, were stuned with haematox sti and ¢osin, All nemalode specimens have been deposited im the South Australian Museum. Adelaide (SAMA). id | BEYERIDGE Woodwardostrongylus petrogale sp. ny. (FIGS 1.13) Melitype : Atehem Land vt SAMA AHC 30592. sume dita. SAMA ATIC 30593, Tarai ype BO ue dite. SAMA ATIC | 47490, Giher mitrerial exanrinert (ron stommieh ot Felravale brachyots, Artem Land NV) edt dk, Netson, Dandeye i ee, 20-99, SAMA ATIC 17, 5 from Stomach ol Pedragale cence, IxXtl977. coll do bo Nelson, Allan. <) Devseriphion Slender elongate nenmlodes, cuticle covered wath numerous fine transverse anniliiions. Mouth opening Uny. dorsoventrally elongate, apparently rigid Joteral murgins of mouth opening, each with row of four relraetile, dome-shiapeu denticles: adhhionil parrot hoy denticles al dorsal and ventral extremities of aich row. Two amphids and) four suibmediin papullie lateral to rows. of denticles; verve tissue extending posteriorly and literally fran scuisury pilpiiae. Sabcutieular region of anterior extremity heavily selerotised. Buccal capsule thick walled with Liint triisverse striaitlans: anlerilin part of buceal capsule dorsoventrully clongate in apical vielvs of Dead, become ciecolir posteciorly, buceal castle Supported externally by LO promiment bands of muscle running from external surface ob buccal capsule ta longitudinal somatic musculature. two bundles of inuseles present dorsally and two ventradiy, lwo thick lateral musele bundles anid: four slender submediin bundles. Oesophagus long und slender corpus eylindtieal oarrowing slightly to form short isthimuss isthmus merging tito elongate bulb. Nerve ring surrounding OesOphagenl Corpus ail SHIT: secretory excretory pore in mid+egion ol oesophageal bulb: deirid at level of seeretory-esereLory pore: Male Total length 131-155 (140): maximonm width 0.1 7-0,23 (0.20%; buceal capsule O.020-0,037 (0.032 } in length, wrth pn biter views O01 7.020 (0.019), in dorso-ventral views (L0TO-OQ.013 (0.012), oesophagus 0. 79-086 (O83)2 nerve ring lo anterior end O4A0-048 (42): sveretory-exuretory pore tu anterior end Q.46-0.57 (0.52). deirid to anterior end 0.60 10.60). Bursal lobes short, of approximately equal length; ventral lohes joined yentealiy; dorsal lohe small, slightly shorter than lateral lobes, nol clearly demarcated from lateral lobes. Ventral rays slender. upposed. reach margin of bursa. extemo-lateral tay shehtly stonter than other literal rays, divergent [rom Vineiien of them, shefiily recurved peur extremity nol reachine margin of bursay medio-kiteral and veatro lateral rays slender apposed. reaching margin of bursa extemo-literal riy arises from lateral trumk, serail, does nobrewch pabeny of bursa. Dorsal cay brtireites close to Origine branches long, Slender, arctate: seoundury Hfavision into branchlets Geeurs nedr cxremily abrayy external branchlets short, direeted postero-hitevally, da not teach matin of bupsy: internal brinchlets longer, dirceted posterorly, almost reach margin of bursa. Genial cone prominent. anterior lobe hatte. conical extenus almost to lini oF veutral lobes: posterior lip spall Will) parr oof prominent postermorly directes appendages; pubernaculanm absenk; central cordite and lateral paired elongate thigkenings ol spicule shewths present at then junctions, spicules wlungale. alie, O90 1,07 (0.97) long: anterior extremities ircceuhurly Knobbeds distal Gps blunts die diminishes Th width towards spice tip, Hemale Length T64-22.4 (200) masini width Obs O29 10,27), buccal capsule O.027 0.087 (030) in Tenth. width in lateral views 0.020. in dorse-ventral views (010-0015 (L013), desophigis 0.92 [28 (LOLS nerve ring te anterior end O,24-044 (O39) seercloryeexerclory pore to anterior end 0,470.65 (0,62); deirid (o anterior end O.S58-0,63 (0.62), Tat short. confeal (LT 1-OLE7 (05). vulva inomectiarely anterior to anus, 023-0. 42 (26) trom posterior ond, Vaging short, straight. direeted anteriorly, 031-048 (0.39); vestibule ta form of Ye wath thick pmiscutae wills, O.1S-0:20 (0.174) Tong: sphineler aril INT bolo GP approximately equal lengths. very variable in length, each O.08-0.25 longs utent parallel. ru anteriorly trom intundibula ege ellipsoidal (). 03-0, 1A (O13) 4 0,072.09 (0,08), Locetisation within sramach Nemiutodes lig in small sinuous tunnels tm the superticral ayer Ob the stratified squamous epithelium of the forestomach. Nematodes were not found penetrating as deeply as the lavine propre and the presence of the nerpatodes in the squires epithelium provoked no inflammatory response. The umerior ends of the nematodes were buried i tunnels while the posterior ends lity (ree in the gastric lumen, Discussion The nematode species described above belongs to the Cloacininae Stossich, 1899 beeaise it possesses a cylindrical bueewl expsule. a burs in whieh the dorsal lobe has four branches. the externe-dorsal ray arises from the kiteral truink and a cervical groove js NEW NEMATODE FROM ROCK WALLABIES TS LX .\ea 3 | | Pigs 1-12. Weodwardosrongvius petrogale sp. voy, 1, Anterior end. Tateral view. 2. Cephalic extremity, Jateral view. 3. Cephalic extremity, dorsal view. 4. Cephalic extremity, apical view, 5. Transverse optical section through anterior end al’ buccal cupsule. 6, Transverse optical section through posterior part of buccal capsule showing muscle bands running midrally from outer wall of capsule, 7. Bursa, lateral yiew, §, Bursa. dorsal view, 9. Bursa, ventral view. 10, Posterior end OF inale, dorsal view showing spicules and thickenings of spicule sheaths at their junction. U1. Female tail lateral view. 12. Pemiale genital system, lateral view. Seale bars = 0.1 mm 1, 7-12; 0.01 mm 2-6. Legend : a, amphid: d, denticle: de. deinids e, seeretory-exerelory pores i infundibulum: UW, lateral thickening of spicule sheath; m, mouth opening; a, nerve ring: s. submedian cephalic papilli: sp. sphineter: t, cential thickening at junction of spicule sheaths: v. vasina: ve. vestibule, Vy 1 BEYER = ’ " er oe Tuy ~~} se Pip LA Histilogigal keaton af othe: stimngel wall al Perrugade ciniiane —oshrowing — laeuliatient al Wialivedeaviaie is poravele spo nove Cy Tt ol super Hepat aquumels epithelial timely eros! formed by the nematode dad the lack of any leitlannniory reavnon ithe epitteliian, Seale bare dh barn hicking, Ho belongs to the genus Woodwarde- Mromgylay becuse 1 possesses a heavily selerotised nonil rezion. av Gransversely stated buccal capsule and a vow of sclerotised denticles on cither side at (he mouth opening. the latter characteristic being the INust obvieus featipe defining the genus. The specnmens deseribed above ure distinguishable fron Woowemlward) and Woobenederfi primarily by the Humber ol pairs of selerotised ural denucles, Woodiedrdostraneylis \woddwerdl possesses: 16 pairs Of denticles wile Wo ofencdori possesses six pars, Inailbol the specimens from Pelrageale spp. Tron Arnhem Land there are four large pairs of denticles: Avcaeh end of the cows oF dentteles. there ys a pair of tiny denticles which has hot been meludedl in the determmation of the punmber of pairs of denticles becuuse these denticles ure oot obvious in baler views and are only vlearly visible in an upical view ol the motth revion, The same terminal pairs oF tiny denticles ure evident mp ihe seunning electron Wicroeruph af the cephalic extremity al Wovhendarfi published hy Mawson (1976, fill) although the feature is Hat entioned ip (he deseription and was nob taken into considerbon when determining (he number of pairs of denticles on each side of (he mou opening, Whether these same terminal pairs ol tiny denticles ure present in Wo Wreeelwerned? 1s not known, In addition. the specimens described above differ fron eongeners in spicule length, being 14 THM HL WO seeeediverredé (Wood 1931), 172.7 mine i Wo ebendorfi (see Mawson 1976) compared with 090-107 nim in the species described above, There is.alsow corresponding difference in the length of the yagi which is O.7-L.0 mim tong in Wo weenie, O68 mm in Wo ahenderi and O31-0.48 mim in the species deseribed here. Therefore, the material from rovk wallabies trom the Northern ‘Lerriory ts considered Lo represent a distinet species amd tle ANG Weeedwerdastnaneyiay peirorale sp. roy is proposed for al, the name being derived fron) the generic narne of the hosts. Th compuniig the deserpuon of We new species wilh those of its Congeners, several morphological characterises of the genus warrant comment dae te ipparent inconsistencies or errors ip publishes Jeseriplions, The vesophagus isdeseribed in the other species ius heing slender und clavate yet in We perragele there is a distinel constnction ab tie level of te nerve ring und the oesophagus is clearly divisible jalo corps, isthmus and oan elongate bulb. Specimens af ie dhendorfi were examined und the same subalivisiats Of The oesophywus wre evident althotigh they have tet been dlisted or desertbed in the Titermtuce, The structure of dhe nesephagus is of considerable hiagnuine sizmficanee heeause i is ahayaie woneru ol the tribe Cloucininea (Stossich, (ROO) af the Cloawciiinae bat i subdivided tite corpus, mihnnus and bulb in the tribe Pharyneostroney lines Popova, 1952 into whieh Wheewerdes ay is bas been placed (Liehtenfels TORO) The revised Wlerpretaion Of these morpholowieal charaeters Therefore beeares eoncorduit with the: earrent laxononne position of the genius, Th the original deseription of the genus, Wahn (Mod, fig, Th) desetibed and Uhistrated a “wubermiculin’ fh the type species, We Waoeeiver till Th the orginal desemptton of the same species, however, Wood (199-1) had heen muh more caulions qd had deseribed ~* yin accessory piece... present is iin frresulur shaped siueiure’ which “uppeats: to finetion as a auide for the spicules’. Mawson (1976) slated That a SUDerNACUTUNY Wis present uy Pbendorfi ul did not Whustrabe the stricture, Durele Desset & Beveridve (M980) illustrated the “gubermuculan’ ob Wodhenderfi but iiselour (rei the tHusteation thi the structure 4s not gubernaculin but the cordate dhiekening at the junction af the spicile sheaths (Beveridge |982) A woberpaculit is absent in Wo perreauade, allhougly at Joes possess a cordate thickening and paired Tater thickenings of the spicile sheaths af their junction fo vommnon with all other cloaemids iad ehubertiids that have been examined for the presenee ot tlie structures (Beveridge |9O87) I therefore seems thost likely that a gubernaculum is absent in this genus and the cordiule central thiekening at the junction of the spicule sheaths hus been mistaken fora vubernaculum jn the past. This problem pertains to several genera of the Pharyngostrongylined and has becn discussed in detail by Beveridge (1982). The morphology of the feniale genital systenr hus not been described in detail for Wo wacedivereli, NEW SEMAPODE PROM ROCK WALLALIES i? allhough Mawson (M7 by rllusteated an essentially: Y- shaped ovejeeloe with parallel, wmphidelphie uleri in her redeseriphion of thts species from rock wallabies from South Australia. The ovejector was deserihed lor Wo ohendori by Durette-Desser & Bevericle (IYSO) who HfUstrated a short Yoshaped vestibule will) short sphincters und infundibula und sugvested that the morphology of the oyejector was Wtermedite between thar vound in the Trivbostrongyloidea and the Strongylotdea. Lachtenfels (PYs80) classified the oweyector of Woodwardosinnevties as lypieally stroneyloid and us her Yoshaped supertivially. but resembling relaed gener with J-shaped Gveyeciors im that the sphincters and tofindibyhe are short, The wvejeetors OF. pelrosale are sumilu to those of Wo ebendorss and confirm Pichtenfels’ interpreuiion. Beveridge (1Y87) ilustmted the uveyector ab We petragude (deseribed stmply as Weedlvuandosironelits sp.) and CONT Ted thal it foo ereed with the deseription and ullocalron suggested by Lichtentels (1980), The oveyeetor jn this genus «§ therefoce considered te he aw moditcd J-shaped ovejector uceording to. the delinitions of Lichtenfels (lOs80). wath the modification probably heme a alivect result of the slender clongaty body structure imposed upon this nematode genus by its cocalisation within epithelial tunnels OF the Bitstric Meesit, The owwsternatie position oof the genus Woodweardoyiongvii bas been the subject of some uneeruitinly. The type species was initially deseribed as Mioryitvostrae yas seared? indicating a close telanonship with the genus Phearwigostronyy- lay Yorke & Maplestome (926, Walnd (1904) Subilivided the genus Pharvigestrongyiias and erceted the new venus Woeeiinedistooerlis Toe We Wado) bul provided no explain of as possible pelathanships with Phaivigesreneytin. Mawson (197) ) redesceibed 2 woodward? front tock willities nd ereetod the mew venus Craviiecpay whieh she plaved in the trichostroneylon! family Aimidustomitidie (Travassos. 1929) based on the pipers al Inghs (i965. 1968) dealin with copvermence OF (he cephilie Icatures ot penitodes OkCUfide buted Hb the stom: linn OF herr hosts whe Tis view thal inany of the stroneylale nemidodes of Nistratian marsupials were members of the Tichostroney low hamily Armdostonmiutidie: Subsaqeatly. ab desenbinw Wi eben dari. Mawson ()926) recognised the synensiny ob Crateceps with Woodiostrornevlis Hub placed Wewnltvared- oviesiies we the An dostomaticiw, Dhupete- Dessel & Beveridge (F980) ry contrast relerred: tte suri Co (he Stronsylolein amt Prchlentely (148d) pluzed the genus m the strongylord inthe Phuryogostrongylinea characterising the tribe primarily on the basis of a huveal capsule wath transverse strimions und therehy re-ussociuing Woodididosmoane\ tis wilh Planiicosmunertys. (he genus with which to owas irst linked by Weod (1931). Beveridye (1982), in a revision of whe Phiryogostrongylinea. omitted Waedweredestien fy/as on Ure basis of nneertiinities ds to tts affinities. The addition of a new species confirms. the characlers upon whieh the cents wits erected while providing some medification to the definition af the genus. prinevipally in relationship tthe morphology OF the oesophagus and Oveyeedor and the absence of a tlie euberaculun, Te assocnuion with mernbers of the Pharyngostrongyloided ts Supported on the buss of aw transversely strated buccal capsule. although this character beeurs alsin certain Benen OF the relted (ribe Zoniolaiminea (Popewa, (952) (Beverndve 1984). This inerphologiew! character appears to be the only feutire upon which alfinilies evan be judped because other characteristies ol the genus are so highly moadihed ta acommuodite its tnusudl mode of existenee within the stomach wall What they are plylowenctcally unijformative. Therefore, in view of the luck of evidence to the contrary. and wilh the limited or even equivocal evidence OF assocnibons bused on the presence ol a siriited buecal cypsule, jt secs reasonable to consider Wordiwardestrong yas asa highly modified member of the tribe Pharyowostongylinea, The host and veographival distributions of menibers OF the gents dre not yet fully elucidated. On the basis ob current evidence, Woobendarff occurs ina variety of roek wallabies: scrub wallabies and hinguroos wong the eastern coast of Queenstand and New South Wales. Wo weedeard? is known from Kangaroos erie ning From the northwest of Wester Australie Galthourh based on ae ¢oo record: tron Brikua) ind from rock wallibies in South Austalia, while Wo permayale ts knewn tren nek walhibies IrontArmhem Land tn ihe Northern Territury. The feature common to all members of the genus ts thal they Pardsitise rock Wallabies But Hest cetatianshl ps Warrant ingore (herough tivestisatron bebop apy conelisivos cun he clawe Trond this Ghsenvalion, Acknowledgments ‘Thats are dite fo De Jb. Netson for prosgsianal The Parasiles Hpor which thes deseriplion is biosee ain ty Dr DM. Spratt for readline a drat ob (he HOLSET ip, 78 I. BEVERIDGE References BrveripGr, 1. (1982) A taxonomic revision of the Pharyngostrongylinea Popova (Nematoda : Strong- yloidea) from macropodid marsupials. Aust, J. Zool. Suppl. Ser, No, 83, 1-150, (1983) Taxonomic revision of the Zoni olaiminea (Popova) (Nematoda : Strongyloidea) from macropodid marsupials. /bid. No. 91, 1-88. (1987) The systematic status of Australian Strongyloidea (Nematoda). Bull, Mus. Nat. Hist. Nat., Paris, 4° sér. 9, 107-126, & Sprart, D, M. (1996) The helminth fauna of Australasian marsupials : origins and evolutionary biology. Adi, Paraitel. 37, 135-254. ’ , Chosp, R. L.. BARKUR, S.C. & SHARMAN, G. B. (1989) Helminth parasites of rock wallabies (Perrogale spp.) from Queensland, Aust. Wild/. Res. 16, 273-287. Durerrr-Desser, M.-C. & BrveripGcr, |. (1980) Sur ta position systématique du genre Woodwardostrongy/is Wahid, 1964 (Nematoda : Strongyloidea). Bull. Mus. Nat. Hist. Nat., Paris, 4° sér. 2, 77-80. INcLis. W. G, (1965) The nematode parasites in the gizzards of birds: a study in morphological convergence, Proc. Zool. Sac. Loni, 135, 125-136. (1968) The geographical and evolutionary relationships of Australian trichostrongyloid parasites and their hosts. J. Linn. Soe, (Zool.) 47, 327-347. LICHTENFELS, J. R. (1980) Commonwealth Institute of Helminthology Keys to the Nematode Parasites of Vertebrates. No. 7, Keys to genera of the superfamily Strongyloidea. (Commonwealth Agricultural Bureaux, Farnham Royal). Mawson, P. M, (1971) Pearson Island Expedition 1969, 8, Helminths. Trans. R. Soc. S. Aust. 95, 169-183. (1976) Woodwardostrongylus obendorfi: new species (Nematoda : Amidostomatidae) from kangaroos, Ibid, LOO, 121-124. Sprarr, D. M., Beveripor, I. & WALTER, E. 1. (1991) A catalogue of Australasian monotremes and imarsupials and their recorded helminth parasites, Ree. S, Aust, Mus. Monogr. Ser. No. 1, 1-105. Wadip, S. (1964) A preliminary revision of the genus Pharyngostrongylus Yorke & Maplestone, 1926, J. Helminthol. 38, 181-190. Woop, W. A. (1931) Some new parasitic nematodes from Western Australia, Rep. ast, Anim, Path. Unive Canils. 1, 209-219, MESORHABDITIS KINCHEGENSIS SP. NOV. (NEMATODA: RHABDITIDAE) FROM ARID SOIL IN KINCHEGA NATIONAL PARK WARWICK L. NICHOLAS* Summary Nicholas, W. L. (1998) Mesorhabditis kinchegensis sp. nov. (Nematoda: Rhabditidae) from arid soil in Kinchega National Park. Trans. R. Soc. S. Aust. 122(2), 79-84, 29 May, 1998. Mesorhabditis kinchegensis sp. nov. was collected in an anhydrobiotic state in dry red sand under a bluebush. Maireana pyrimidata (Benth.) Wilson, 1975. This is not the usual habitat for Mesorhabditis which is commonly associated with rich organic matter. The same species was also found in agricultural soil. Key Words: Anhydrobiosis, Australia, Mesorhabditis, nematode, soil, taxonomy. Transactions of the Boval Sect ofS. Aust (L998), L222). T9BA 19 MESORUABDITIS. KINCHEGENSIS SP. NOV. (NEMATODA; RHABDITIDAE) FROM ARID SOLL IN KINCHEGA NATIONAL PARK by Wakwiek L. NICHOLAS Summary SHOHOLAS, Web. (1998) Mesarhabeits kinchegensiy sp. noy, (Nematoda: Rhabdiidae) from aed soil in Kinchevsa National Park, Tras. WSS, Aust. 122 (2), 79-84, 29 Mity, 1998, Mesorhubdilis kinehevensiy sp. voy, was collected in an anhydrabiotie state in dry red sand under a hluebush Maireane pirinidare (Benth.) Wilson. 1975. This is not the usual habital lor Mesorkabdiiy which is dominantly fscice WHR rie organi rouner. The same species was also found in agricultural soil, Distinguishing features of this species ure that tn the made the fps ab the long. almost straight, distally lrsed spicules, are abrupily (irned ventrally, The formula for the aerapgement al the bursal papillae ts (2 br 3) with none olthe pupillae (used at (hei bases. tithe female the Gil is long and pointed so thatthe distunee thai the posreribn Vulva to the anus is aboutone anda bull tines the tal length. Key Warns: Anhycrobtosis, Australia. Weserhadiditis, nematode, soil. laxononny, Introduction Most species of Meyerhabeitiy ive been reported from tieh deeaying organic matter such as humus, rolling wood or dung, Several speeies are usually found in close association with insects, Pew species hve been found in au habitats, The species deseribed herein was collected in an anhydrobiote stile from dry sandy sor with fiithe organic matter. One other-species of Mexorhaldilis, Mo spiculigera (Steiner. 1946) Osche, 1952 has been reported lo survive anhydrobiosis (Sudhuus 1978), Osche (1952) subdivided the very large genus Rhabditis into seven subseneri, One of which was Mesorhabditis, with the type species Rilwalylitiy ypiculigera Steiner, (936. The taxonomy of Rhabditidie has been extensively reviewed by Sudhaus (1974, 1976, 1978) Whe has retained Mesorhabditiy al subgeoerne rank. Phis view was not supported by Andrassy in his autboriGiive mongemiph on the suborder Rhwbditina (Andrassy 1983) in whieh he considered Meserhahditiv to he a sepunile venus within the Mesorhabditinue, a rank accepted Hi this paper Sudhaus (LOOT) was not, however, persuaded by Andrussys acuments (Mut Mosorhobditis has aenerie canking, The difference i niniking rests an the Gasonomist’s inclination towards Tuinpimee’ or ‘split’, Within the suborder Rhabditina, the combination ol charieters that distinguish Mesorhabditis area Monodelphie femme with the villva: Well posterior to the Hd-poral ob Mie body wid a ponited conical til. The male has long, more or less straight spicules that are Jistally Tised, The male bursa is peloderian wath “Diwewton oak Motiv aid Zogloy Nawtnilign Nabeul Onieersary Content ACE C200, pared borsal papitlie arranged in three groups, lypivally bwo preecloucal, five peri-cloaeal and three closer lo the ip of the tail expressed by The bursal formula (2+54+3). Material and Methods Several samples of dry sandy soil were liken with aeylindrical metal corer, 12.5 cm lou. 5 cin miternal diameter, close to and below a bluebush on 4 November [984, The siunples were phiked in plistic bags and returned to the laboratory in Canberra, Ten days later subsamples of 5 2 were placed on tissue paper in fap water in Baermanmn funiels. Atier 18h the funnels were drained and the nematodes collected. From one subsample. Giken tron directly heneath the bluebush, fifteen specimens of a new species Of Mesorhabditiy were found Clopether with many other nematodes). "This species was nat found inainy of the other samples The specimens ol Mevorhabeditiy were fixed i Sti formalin and transfered ta 5% aqueous glycerol, whieh wis concentrated by evapetanon at 4a Cc, then mounted on slides in anhydrous glycerol with cover Slips Supported by ghiss heads and ringed with Cilyceel (Gurr). Drawings and ricusurements were made will a ediera ducida wdachient on a Zeis Ulfraphot micrpscope, Type material hus been deposited in the Nations Nematode Collectu|l (ANIC) atthe CSIRO Division of Entomolosy. Cunberr ACT. Mesorhahditis kinchegensis sp. nov. (MIGS |S) Healowpes 9 Kinehega National Park. NSW. 4oxy 1984, ANIC Nematode Collecuorn side QOOQ00S , RO speeimen QO000007, Paratypes: 6 34,5 29, Kinchesa National Park, Axi 194, ANIC Nematode Collection slides QOOO006-12, specimens QOQDQ008- 19. Measurements: Table 1, Measurements in im. Deseripiian of Halewpe male Body cylindrical, slightly tapered at head, rather bluntly truncated ut hind end (Fig. 1), tail short with peloderan bursa (Figs 1.3). Cuticle finely annulited, lateral field appears as Uiree parallel lines beginning in cervical region and extending as far as tail (Pig. 3), Six offset, rounded, clearly separated lips, each bearing a prominent labial papilla (Pig. 5). Buecal cuyily cylindrical, without pharyngeal collar. glottis possessing minute denticles, probably two (Fig, 5). Pharynx with strong muscular corpus. slightly expanded af metacorpus. narrow — isthmus, sutrounded by nerve ring, valved pharyngeal bulb lerminuling in very short trilohed cardia. surrounded hy intestinal tissue (Fig. 2). Secretory-excretory pore. ventral, level with base of isthmus (Fig. 2). Intestine, initially filling pseudocoel. becoming compressed about halfway along body by gonad, followed by rectum opening at cloaca (Fis. 1). Tail short. sharply pointed. Single testis reflexed dorsally, leading to short vesicula semipalis and long yas deferens, Clouca surrounded by a peloderan bursa wilh 10 pairs of papillae arranged (24+5+3) (Fig. 3). W, L, NICHOLAS Bursal papillae not fused it base, short posterior pair curled over, Two long narrow nearly straight spicules, capitulum distinet, distally fused, lips abruptly angled ventrally at about 25° to the main part, just beyond & slight constriction (noteh) (Fis, 4). Gubernaculut a short straight rod, Posterior deirids at level of middle of spicules (Fig. 3), Paratypes and other meles Measurements: Table |, All the inale paratypes closely resemble the holotype. The level at whieh the spicules fuse, about 50% of ther length, can only be clearly seen by squashing and rolling the specimen under a cover slip, which renders the specimen useless us a lype specimen Fomale paraiypes Measurements: Table (. Female paratypes vlosely resemble males (Pizs 6,7) except for reproductive organs and lait (Fig. 8). Homodromous ovary reflexed dorsally i nid revion of body. Uterus extending to just beyond short Vagina and vulva. One paratype female (Figs 6-8) possesses sperm i) a short transitional region between ovary und uterus and six developing eges. about (5 pin in diameter and varying from 15 - 24 fin in dength, Amphid fovea, a minnte oval slit at base of lateral lips, visible only in this. paratype Varin 1. Measerements of Mesorbabditis kinchevensis speci, ---—————————— CO _ocn— (“—_— Sex/Type Male/Holo Male/Para n=6 Femuale/Pura n=5 Mean SD Range Mean 5D Range ee" Length 524 AK? 49.30) 432-533 543 7196 407 -H62 Max, width 2k 40 3.83 25-34 AQ) 24)4 JAAS Bueeul cavity 15 15 2347 I-16 (5 151 [416 Corpus Ol 54 1,27 S137 ahi] (51 56-60 Phavyix Pa) 126 37 [1-140 [22 6.9] }1R-128 Heal to nerve rine 92 72 11,25 5A-KA 7 6,22 61-70 Head to secretory /eseretory pore 116 92 142 77-100 Kd 2010 55-13 Head to intestine |? 138 5.01 132141 135 K OY 1al-2 Heid to gonad flexure 224 mit) 19.85 [88-245 242 56,08 205-284 Heit to vulva - 439 OG ATS 455 Head to anus 4a) 454 IO AMW)-SO7 493 57.2! 455-518 Gonad length 2Nh 278 13,97 268-295 AI [34.14 THOS 45 Rectum length 34 3| 108 23-3 a2 6) BU ER i} Yul 22 23 34 IR27 ait 14.77 32.70 Vulva to anus - - - 05 7.89 37-10 Spieule Is 4h 4.48 415 - Guberneulinn Ih a Mala) [8-24 Do Man's a (8.7 7 1.6 12.7-184) au 242 S427 De Man's b i 4 37 3A | 4.4) 47 AAT Do Mais 238 aI I4 17-95 0 5 24) 47-145 De Mais Vu e - hI 732 77-82 _ CC rr C ——n— ANHYDROBLOTIC MESORHABDITIS rye Wuillad lad aL Wha dat IAIN 1 ets 50um 2 2a 50um 346. 10um 5 se 107 Figs 1-5. Holotype male. 1, Entire male. 2, Ceryical region. 3, Cloacal region with spicules, gubernaculum, bursa and bursal papillae and lateral line in lateral yiew, +. Spicules orientated to show fusion. 5. Head and buccal cavity. 82 W. L. NICHOLAS female (Fig. 7). Tail conical and sharply pomted (Fig. 8). Vulva posterior and distance from vulva to anus about 1.5 x tail length. Lateral line marked by three incisures extending from mid pharyngeal revion to caudal region (Fig. 6). Differential diagnosis. Mesorhahditis spiculigera (Steiner, 1936) Osche. 1952, 1s the only other species reported to survive periods in anhydrobiosis (Sudhaus 1978). [t has a world-wide distribution and has been reported from New Zealand but not from Australia. It differs from M, kinchegensis sp. noy. in possessing a longer narrower buccal cavity, fusion of the bases of bursal rays 4, 5 and 6 and the tips of the spicules, though notched, are not angled ventrally. The ratio of length to width of the buccal cavity in M. spiculigera is about 10: 1 (illustrated by Sudhaus 1974 Pig. 7). whereas in M. kinchegensis it is about 4 : 5. Two other species, M. ssunvoghi Andrassy, 1961 and M. WIHT Chic habdadanbuandd " 6 50um 7, ee FS um Qo Se es 50um Figs 6-8, Paratype female, 6. Cervical region also showing the three incisures of the lateral line. 7, Head. §, Posterior body showing reproductive orgiuns, ASTYOROBIOTIC MESORMARDYTIS u4 Inmeuspiemlosa (Sehuaurmans Stekhoven, 1905) Dougherty. 1955, abso have a notch close Lo the tip at the spicules, bul antike 7. kinchevensiy, heir spicule Ups are nol angled ventrally beyond the notel. Andrassy (1983) provides a useful key to the 17 species he recognises. a summary Of diagnostic Chiricters and references to taxonomic deseriptions, Several species, WM, oscher Osdtner in Osche, 1952) Dousherly, 1955, MW megectilis (Sudhaus. 1978) Andrussy: 1983. A frreveheds UxGener a Osehe, (952) Dougherty, 1955. A vennwagli, WL Jiglandicola (Puehs, (937) Dougherty, 1955, M7, saliasy anid MO irariiensis (Meyl, 1953) Douvherly, 1955 ci be clearly distinguished by havingoa shorter female wil so that the distinee (rom YUL Ta Uns iS TCH preater Hain the tail length. Up the new spevies (he distance ts only about 1.5 x the hal length, A vanely ol features distinguishes other species From Mo Afmohewensis, Ui WL aiethi (Sudhuas. 197%) Andirissy, 1983 the spicules tire uch Shorter (29 36 pon compared With 41-5) ain in Mo hinchevensiy) The bueewl cavily of At cnivonterplea (Sudtius, (978) Andrassy, 19X83 is asymmetric. the phanyin of dL erigenmearenniy (Khem. 965) Andiassy. 1083 is unusually long, one third ol body lenwth, Mesarabedins africans Andrassy, HY82 bos labial papitlie curved: inwaruds, Mo uffime (ROrner in Osehe, 1952) Dougherty, 1955 has ported tips. MM. renoipreulim OsGrner in Osche, 1952) Dourherty, 1955, MW. belari (Niven, 1949) Dougherty, [O54 qn A. Gtritensiy possess only Wine borsal niys, the middle group faving four Tasted OF fhe nmere tise) five. di uM yerteriee Dassonville & Heyis. P984. deseribed by Dussonville & Fleyis (hO84) afer the publicaion of Andrissy’s monograph, the laueral line has five jneisures rather (hin the more typienl three, as in A Kineheversiy, Sudhaus (97K) tas ohserved uberralons in fhe tadl length ane) besa Muvidil spectinens bab the characters used) to disthiviish MM, Rineheveusys ate consistent mall ihe ype specimens deserthed in (hts paper Htabilar Soil around plik too. The type specimens were onlleeted ie diy sind ie ai anhydrabiotie stare fren AVOUT The roots a blchtish, Medrecie pyciniidiite, rays at in Kinchega National Park, NSW, Three males of the Mune species Were collecled by M. Hodda from a field ol lupins on The Soil Conservation Experimental Farm at Cowra. NSW. These ure in the ANIC. collection, Nematode Colleetion slides H001290, 0001295 ail 0001286 butare bot ieluded vs pantlypes, as they come from uovery different habitat and ure mounted on slides with several other species of nematodes, Disteibunon AU present the species is known from only two lowawlities in New South Wales. Discussion The type-specimens ol Mesamiabditiy Machevensis sp. ney. come front an atypical habitat for Mesorhabditis, namely, arid soil with tithe orgame Maller TH Kinchega National Park. although this speeies bas also been collected front aeccultival Jund, Kinehega National Park lias a very irregulie annual rain, averiing 235 rim, and tin smnual evaporacion rite of 2000 din, Teniperitures reach 49" Co suniier aie fall to 0) Coin-winler, Most of the previously described species of Mesenhabedtay hive been found i cdlecomposiig organic nutter sueh as modldy or rotting wood, or Iiinis (Andrassy W983). Severdl others hiwe been found in close ussecnition wil imseety such as searabid) heeile lurve. Mevorhatilits meveehiliy wits ussucnied with hymenopteran nests CAndtassy 183) Mesorhabditiy sudhetusi has been reported tony soi CMndrassy 1983) and AY srriadtes tron fresh walter (Dassonville & Tleyns [Ondi Wesuelinbeitin spicaliverd, the olher species kaowe to survive in awhydrubioses, Has been Toundh i roti: wood: and horse due. Ths ducer larvae were associated with dune beetles (Sudhans (978), Acknowledements Jauneratclal lo Ms J. Sit for collecting samples Hom Kinchesa National Park ane) Dr ML boda. for Inmaking specimens fram CSIRO] Nematolopy collection wadhible lar stay. References Anmmasny lh (lO84) A Tesontimic Review of the Suborder Khaliliiina (Nemarike Sacerentid (Gditmens de VOlhee de ke Researches Selentiique ah lechmique Coulee Hers Peis). JJASSONMIEDD, AF ok HBV SL C1984) Proshwaiter nenitades treme South Adrien 7. New pind Kineawn species ooullewted ott Skirmmerepruiy Mietoria, Phyrophvlaenien Obs a, OSCE OG. FINS I Systematik Und Phy lerie den Cartan Kivibdity (Nem ALO) Mae Tb Avs SMD, 190 80 SUD MS Wo CPE) fur Systoriatik, Verbotene, Okeloure und Bralosie Heuer amd weg bekunoler hibuitlen (Nermiohad 1, Teal dba POE S. 174. 122 (19700 Sonim h hilonisele Rerinerkiiwen uber Arenund Gattungen der Uiiterkumtie Rhabditinae vertu five (Rhahditidae, Neroatodiy, Vewnrafagien 22, 4976, 84 W. L. NICHOLAS = (1978) Systematik, Phylogenie und Okologie (1991) Check list of species of Rhabditis der holzbewohnenden Nematoden-Gruppe Rhabditis sensu lato (Nematoda: Rhabditidae) discovered between (Mesorhabditis) und das Problem, “geschlechtsbezogener” 1976 and 1986. Nematologica 37, 229-236. Artdifferenzierung. Zool. Jb. Syst. 105, S 399-461. FIRST QUEENSLAND RECORD OF THE BURROWING FROG CYCLORANA CRYPTOTIS TYLER & MARTIN, 1977 (ANURA: HYLIDAE) BRIEF COMMUNICATION Summary During fauna surveys conducted in Cape Melville National Park (150 km north-west of Cooktown) and adjacent areas, new species and new records of vertebrates and earthworms were obtained’’. Following a thunderstorm on 21.xi. 1995, large numbers of frogs were found in a localised area (14°34°45’" S, 144°29°50°° E) approximately 7-9 km west by road of the Wakooka Outstation. A call unfamiliar to me was in the large chorus. Observation revealed a species of burrowing frog of Cyclorana not recorded in Queensland. Thirteen males and one female were collected. The call was recorded, tissues sampled and photographs taken. Trimesaetions of te Reval Sorte of SM (199%) 122(2) KS-RO BRIEF COMMUNICATION HIRST QUEENSLAND RECORD OF THE BURROWIN ~~ vi 7 FROG CYCLORANA CRYPTOTIS TYLER & MARTIN, 1977 (ANURA : HYLIDAE), Durie Faun surveys conduerd i Cupe Mehatle National Park C150) kan ooriewest af Cookdsva) aad AUJUCONL reds: Hew species an new cecords of vertebrates und Garth woes were obiined!: Following a thunderstorin on 2bsr 1995, latee pubes oF frows were found in a localised aren ChE ANAS S, 144" 20550" LY approximately 79 kny west by roud of ihe Wakooku Qutsntion. A vall Unhunilin tome Wee oy the hire chorus; Observation fevenled aospeeies of burrowing trae ob Cyclaraier net recorded in Queenshind. ‘Thirteen males aid ane fenuile Wer Collected. The cull was recarded, issues sampled ane pholowrophs taken Qn the basis of norphotopy and oll | tentatively denis the Hop us Cvefarenn eryptatiy, a seal burrowing Hag previously Known from northern Wester Aust and the Northern herritary Hadividiibs conform in colour and appearunve to the Veseription af) ceyproris Tyler aod Martin, wath) a bility mottled dorsi of slute, erey uml saloon with a ulistinet silo phsteorbital bur (Fie Li The appearence is Stokingly siovlin to the phitayeriph in Tyler ened, (O82. 1 preservadive, (he salmon colon ranol was lost The raptial pads Were salmon aad ladecl in proservative. The ventral sirtaee wars white ind intles had a shite thro, The LWP wits Gevered Will skin i all animals. Toes were hall webbed wilt no expanded terminal dieses Nip t, Male Cvilera eryprens Trem west of Wakoaka Outsuien, Queershinel, The general appeaninge is of idiimpy ain fobust frog (Pip. 1). Meastiremerts, following previous faemliods . fall Within the tange ol oreplaris! Sioul vent length ranges from 357-459 nim, with the only female measuriys 38, )mim, Legs wee short (PL/S-¥ 033-0039) yn the eve i naris distince is wremer Or Jess than the intenarial spar (LE N/IN O.955-1.141) (Table 1), Frogs Were Found Hi temporary, rere fifledl pools alone a draiuge Whe ta lows open woodkind on chu soils, The verehtion wis duminited by Aveldleuee steneisray lever anid Enealypras leprophieba (vip, 2 Alvitude was 40+ 1 in Call dination was 503 tasce witha pulse repetition rite at [13 pulsesssee ! anda dominiune frequeney of SOU He The pulse reperiiow Mite is lawer thin those rages al oe cryplotiy described im the Tilerature and Lake Argyle area (P83-193 pulses see 2o hin Nie Like Argyle Tourist Villugets 145-160 pulses see! [Go b> Watson pors. anit. 1997}. lake Argyle 1a8 pulses see for holorype, Daly Watters) CRible 2), Tenmpenaures at the callie site were 2H.2 CO (wae) and 27.2" C (ane) Cv lorie ceoypons wos calling while Coating in walter, When the voeul sae inflided the anterior halt ot the ives Wath Hfled, The inflotig and deletion of the yoeul sae cured the body ta rock ia Toner similar to Nerden delaneseaphuy vbserved elsewhere im Cape York, Arotter localities in its tange Co ervgetatis usually calls whilst Hoorn a water (GE Watson pers, comm, 1997) although the hololype ovis auling from dand OA. AL Martin pers. cam, 1997), The snite fenile was collueied Mowting in the water in ayilhiry aniplexus. Two other species ot Cyelarana. Co brevipes and CL pevertalteendion. entled froin ihe banks Ob pomls whereas Linaeedyre ster crctes called with Co erypfediy in the pools The presence of ©. erypreriy near Witkooku O)uistutigns represents worminge esrenston oF YOO ken east ror prey tous foculity records obtuined by Davies. Tyler und Watson at Borroloola, Northern Territory (SAMA R43702, 16°12 8 13637) EF). The intervening area accoss The gull pkainis fas hot heen exteasively sampled and additional populations may be vapeeted, The location of all records far € ervyprotis is tit band across nerthers Acstitia (ham Berhy, Western Australia to Wakooke Qutstigen mur Cape Melville between 1430'S gd [2 20'S, Other frogs collected with ©) cevyplons were 2 €. brevipes, Co unvertinftandin. literia albounitata, by coerulea, b. ribelli, Lined ynasies atuerity, Noveteler melanoscauplies and Clperaleie miunula. Voucher specimens of Cerys celleered ai the Wakooka site ie tn the Atherton office of Deparment al Lovironmenteatlection Sos SN 30000, SN 7I0b8 20, 72074- 25, 7245-36, 7204044 Taki) Morphofogical meannrenents of dd Cycloranu cry ptotis fem west af Wakooke Quisterion. Queensland Abbrevintions fallow Tyler & Martin (1975), SV] TL/SVL HW/SVL- HW/TL RD/HW L-N/IN Rare A745 9 (1339-0), 39 036-043 (1924-1, 169 (1.26-0.343 955-1141 Meun 49.5110 356 O.386 1 URS (). 2M 1.65 RO % J Morphologically these specimens conform closely wath Pda | th : i ft C. ervplotis. However in view of the size of its range ; j extension as well as the differences in cull structure, the % Ate identification should be regarded as provisional until iS Ae age! b= Substantiated by biochemical analysis. ‘ Me ae Dr M. Davies, University of Adelaide und Dr A. A. = giorceeand ES, raat Martin, Royal Melbourne Zoological Gardens reud un early version of the manuseript and provided helpful comments. Assouile Professor G. PF. Watson. University of Melbourne provided call analysis dati. Field assistance was provided by M. Blackman. Q. Hart and R. Worall. C, Prith supplied the C. cryptotis photograph. The referees, Assovrate Professor M. J, Tyler and Dr A, A. Martin. made significant fe contributions to the manuscript. Alf uf this assistance is eratelully ucknowledged., Fig 2. Habitat of Cyclorana eryptoriy west of Wakooka Outstation, Queensland. Froes were calling from the Lermnpordiry pool. Taunn 2. Call variation within Cyclorana eryplouis. Superscript numbers in source colurmm refer to references. W - Water lemperature, A - air temperature. at calling sites, Source & val Locality Dominent Call duration No.of pulses Pulse see! Calls min! ‘Temperature °C sumple frequency — (milliseconds) This paper n=] Wakooka, Qld S00 503 58 113.3 TA 28.2 (W) G.F Watson Lake Argyle area. WA 920 439-455 65-74 {45.8 83.9 (pers, comm. 1997) n=2 Holotype! Daly Waters. NT 1060 530 - 158 24,1 (A) Tyler erals n=l Lake Areyle area, WA &S0-1 100 330-348 61-70 183-195 36.9 (A) ‘Jamieson, B. G. M. (1997) Mem. Qld Mus, 42. 233-270. Tyler, M. J., Davies, M. & Martin, A. A. (1982) Copeiy MeDonald, K. R. (1997) /hid. 42, 307-309. 1982. 260-264. ‘Tyler, M. J. & Martin, A. A. (1975) Trans. Ro Soc S ‘Tyler, M. J. & Martin, A. A. (1977), Ree. S. Aus. Mus. Aust, 99, 93-99, 17, 261-276. K, R. McDONALD, Conservation Strategy Branch Queensland Department of Enyironment PO Box 834 Atherton Qld 4883. THE PREVALENCE AND DISTRIBUTION OF NEMATODES IN THE LARGE INTESTINES OF SHEEP IN SOUTH AUSTRALIA BRIEF COMMUNICATION Summary Nematodes from three genera (Trichuris Roederer, 1761, Oesophagostomum Molin, 1861 and Chabertia Railliet & Henry, 1909) have been identified from the large intestine of sheep in Australia’. However, there is relatively little information on their distribution and prevalence. Oesophagostomum venulosum (Rudolphi, 1809) and Chabertia ovina (Fabricius, 1788) are believed to be widely distributed, particularly in winter rainfall areas** and Trichuris ovis (Abildgaard, 1795) and T. skrjabini Baskakov, 1924 are common species in sheep and goats’. Transactions of the Royal Soci at S. Aust (V99R), 122(2 87-88 BRIEF COMMUNICATION THE PREVALE ‘EK AND DISTRIBUTION OF NEMATODES IN THE LARGE INTESTINES OF SHEEP IN SOUTH AUSTRALIA Nennitodes from three gener (| 7richwis Roederer, 1701. Qesaphagesionin Molin, (S61 und Chabertin Ruillier & Henry, P09) have been identified from the large mlestine of sheep in Austr’, However, there is rebitively: [ttle information oon ther distibuiton aml prevalenee. Oevaphaygestonman venitoxiun (Rudolphi, 1800) and Chabert ovine (Fabricius, L788) are believed to he widely distributed, puuticulaely ta winter cotlall qurcas>! and Tricity ovis (Abildpaard, 1795) and 7) wkrjabini Bushakoy, (924 dre comiion species in sluep gad goats’. Beveridge and ord surveyed the distribution and prevalenee in South Australia of the ceanonncally Mportint Uichostoiwylod nenkdades of sheep, which occur iy the stonmuch and small intestine. and reported that some were betrer adapted to hot dry cnavironments (han others?) denmtodes. in the Taree intestine were identified only ineidemtally. Ty the present study the prevalence und regional distr iion of nematodes of the lhirge intestine of sheep in South Australi was determined by examining freshly collected caecu and colons of 313 sheep collected from 116 Widely distributed localities. (Pig, 1) from [991-)993, Information On farm minagement, age. sex or breed of sheep Wats Unavailable and) provision Far the possibiliy of seasonal yarhilions in infection was not possible in the collvetion of material Intestines Were opened. emptied. washed in water andl the mitcenal suclice was inspected for nemilodes whieh occasionally adhere ty (he gir wall A iin subsample of the Tatestinal contents was examined for Tre HeTalodes UsiNg Un OLVTpus stereo-microseone, The remaining contents were washed through a 670 urn sieve Nematodes were collected. fixed) in LO bulfered formalin und cleared Th lacloephenol Tor tend feution, Male and female nematodes were identified hy comparison with published deseriptions of species! Four tematode species from Uiree gener were found) (heir prevalence in three rain kul zones is shown in Table b. Ritty-cieht percent of sheep were infected with nematodes. The mean numbens wi rie OF burdens of adult nematodes and the number of localities from which each species was shown i Tuble 2 Trivhiwis ovisand 1, skejahind were conn and witlely Wisttibuted with the hitter species more prevalent, Both ferodes aeenrred i 140 OF animals and in 7% al these cuses 7 oskryabind owas the predaumaitt speeies. Sihificantly fewer 20 avivowere found in the low: paint! cone (<250 nin than mares with higher raimtall yg = 41 ps O02), Onophagestomion venilosunr wos (he most commen nematode reeovercd and was most previlent lin sheep fran wrens with more than S00 mit of rainfall Nematodes of all three gener occurred concurrently in only four animals. all from the highest rainfull gone (S500 i. Pourth stage larvae of. veniesin were the galy Timature henatades foundk these wer prescot in 38 animals from 24 foculities, V7owith un annul rainfall greater than 300i, Weller areus of South Australia are upparently more favourahle for the Iransmission OF O. verufestnr (han drier areas, Banks! detected ©. evive in chotable” siimbers tn South Australia forty vears qe but did not vonsider it ta be Importint. The intlience of highly effective unthetniniies, sitce (heir itroduetion ta fhe P9608, njay Have contributed tothe current low prevalence of CL avid Prialiity spp, are generilly regarded as harmless! bur lave been assocned fecovered une Tabi l. Prevndlence (% ) according to rainfall of neniatodes inte large imestine ap steep in South Austratia. Rainfall zone *urusile Overall 200 ~ 349m 350 - 499mm =S00r0 Triclieis aves att 1 ais 28 Trichawis \krjabini 32 42 a4 28 Oevoplagestonuil 43 30) 29 SY venulosin Chubertia ivina 5 6 i 5 TABLE 2. The mean diner uid range of nennitode burdens and the number of localities (in 250-349 1m (et), 350-499nn Oh), >SO00m (ao) rainfall zones) from which cach spevies wus recovered, Nematode species Mean Trichucls avis 7 Hrichwvis skrjabini 1 Vesophagestamin venulosum 15 Chabertia ovina 4 Range No ol localities i by v 0-63 I 13 \5 (60 alt) a 7 (261 22 att 20 10 4 6 4 The tohal Tocalities tn cach rainfall gone - Gi) = 23, (hy = 4h ic) = 49 88 with deaths of sheep in Australia during drought conditions®. Oesophagostomum venulosum is non- pathogenic’. lmmature C. ovina cause intestinal damage during their development but adults are less pathogenic!. Natural nematode tmfections in sheep offen consist of a mixture of genera and species, some of which appear to have hittke effect on their own but may contribute to disease caused by more pathogenic species!, Although the parasites found in this survey are not considered to be economically important, the data supplement previous records of gastro-intestinal nematodes of sheep in this region of Australia a oO gy? Yoo ADELAIDE O Fig. |. Distribution of samples examined from sheep in South Australia. One to 4 sheep were examined at each locality Negative for nematodes (9, one nematode species LU, two species A, three species @, four specics MH, ‘Cole, V. G. (1986) “Animal Health in Australia. Vol. 8, Helminth parasites of sheep and cattle” (Austratian Government Publishing Service, Canberra). ?Forsythe, B. A. (1953) Aust, vet. J. 29, 349-356. ‘Banks, A. W. (1958) Ibid. 34, 20-26. iBeveridge, I. & Green, P. E. (1981) /bid. 37, 141-142 ‘Beveridge, 1. & Ford, G. E. (1982) /bid, 59, 177-179. Goldberg, A. (1951) Proc. Helminthol. Soc. Wash. 18, 36-47 7Herd, R. P. (1971) Int. J. Parasitol. 1, 189-199, ‘Farleigh, FE. A. (1966) Aust. vet. J, 42, 462-463, °Goldberg, A. (1952) J. Parasitol. 38, 35-47. M. G. OTCALLAGHAN, E, OCKLESHAW and J. ALLEN, South Australian Research and Development Institute, 33 Flemington Street Glenside S. Aust. 5065. VOL. 122, PARTS 3 & 4 30 NOVEMBER, 1998 Transactions of the Royal Society of South Australia Incorporated Contents Martin, H. A. Late Cretaceous-Cainozoic palynology of the Poonarunna No. 1 well, central Australia - - - - - - Kolesik, P. Rhopalomyia lawrenciae, a new gall midge species (Diptera: Cecidomyiidae) deforming leaves of Lawrencia squamata (Malvaceae) in South Australia - - - - - Kolesik, P. Dasineura wahlenbergiae, a new species of gall midge (Diptera: Cecidomyiidae) damaging shoot tips of Lense Stricta (Campanulaceae) in South Australia - - - Davies, M. & Watson, G. F. Developmental biology of Uperoleia a “Tyler, Davies & Martin, 1981 (Anura:Myobatrachidae) - - - Davies, M. & McDonald, K. R. A new species of frog (Anura: Seats from Cape Melville, Queensland - = - Davies, M. & McDonald, K. R. Developmental biology of Cnanise ei Davies, Watson, McDonald, Trenerry & Werren, 1993 (Anura:Myobatrachidae) - oe M > 7 Coleman, P. S. J. Changes in a Mangrove/Samphire Community, North Arm Creek, South Australia - — - - - = - - Smales, L. R. New species of Seurechina (Nematoda : Seuratidae) paras in dasyurid marsupials from Australia - - - Ferguson, M. A. & Smales, L. R. Spiroxys chelodinae Berry, 1985 (Nesumade: Spiruroidea) and Camallanus chelonius Baker, 1983 (Nematoda: Camallanoidea) from freshwater turtles (Pleurodira: Chelidae) in Queensland, Australia - - - - PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 89 139 185 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 122, PART 3 LATE CRETACEOUS-CAINOZOIC PALYNOLOGY OF THE POONARUNNA NO. 1 WELL, CENTRAL AUSTRALIA By HELENE A. MARTIN® Summary Martin, H. A. (1998) Late Cretaceous-Cainozoic palynology of the Poonarunna No. | well, central Australia. Trans. R. Soc. S. Aust. 122(3), 89-138, 30 November, 1998. Palynomorphs found in Late Cretaceous-Cainozoic sediments are described. The Winton Formation yielded the Cenomanian Plicatella distocarinata Zone, but the uppermost part contained an equivalent of the late Paleocene Lygistepollenites balmei Zone, showing it should be reassigned to the Eyre Formation. The Eyre Formation also includes sediments that are an equivalent of the mid Eocene Lower Nothofagidites asperus Zone. An abundance of Asteraceae and Chenopodiaceae/Amaranthaceae pollen in an assemblage at shallow depth is thought to be Pliocene-Pleistocene in age. Key Words: Central Australia, Palynology, Late Cretaceous, Tertiary, Palaeovegetation. Transactions of (he Royal Secrery of S Aust (1998), 12203), 89-138. LATE CRETACEOUS-CAINOZOIC PALYNOLOGY OF THE POONARUNNA NO. | WELL, CENTRAL AUSTRALIA by HELENE A, MarTIN® Summary Marvin, HAL (1998) Lite Cretaveous-Cainosoie palynolowy of the Poonarunna No. | well. central Australia. Trans. R. See, 8. Aust. 1223), 89-138, 30 November, 1998, Palynomorphs found in Lite Cretaceous-Cainozole sediments are described. The Winton Formation yieldeal the Cenomianian Plieaella distocurinate Zone, but the uppermost part contained an equivalent ol the late Paleovene Lygivtepallenites babe’ Zone. showing it should be reassigned to the Eyre Formation, The Eyre Formation also includes sediments that are an equivalent of the mid Eovene Lower Nerhefiigidites ayperus Zone, An abundance of Asteraceae and Chenopodiaceac/Amuaranthaceae pollen in an assemblage at shallow depth ts thought fo be Plioacene-Pleistocene in age. The vegetation of the late Paleocene and auddle Rocene was mainly forests with minor herbaccous SALT communities, Gymnosperm pollen dominated the lute Paleocene palynoforis and proteaceous taxa were very diverse. Pollen of Cononiaceae/Elaeocarpaceae is moderately common and there is a wealth of a@iospern pollen, Inthe mid Rocene, pollen of Arauvariaceae, Casuarinaceae anda lithe Norhofages were dominant and there Was a great diversity of angiosperms. The Plivcene-Pleistocene palynofloras haye a limited diversity with pollen of the herbaceous/shrubby Cyperaveae. Poaceae, Asteraceae and Chenopodiuceae/Amaninthacesae Jominunt, Cusuuriniceae and Myrlaceae are the only likely trees and there is relatively lite pollen of these families, henee the vegetation was open shrublands similar to that found in the region today. There are, however, some dispare Gixa inthis Pliocene/Pleistocene palynoflora that are unknown in the wrid region today. Key Worps: Central Australia, Palynology. Late Cretaceous, Tertiury, Palaeovegctation. Introduction 1dr an Taz . ey 4 @ Bore of this study This study of the Dethi-Santos-French Petroleum Boles hiportad BV POONARUNINA No. | Co, (Aust.) Poonarunna-| well was undertaken inthe Sluttet (1291) hope that ir would shed some light on the evolution # Macrotiossil site of the arid flora and vegetation. The location of the well, northeast of Lake Eyre. and within the first sand ridges of the Simpson Desert (Magnier'). is strategically placed for this purpose (Fig. 1). Finding preserved pollen in wrid/semi-arid regions isd major problem, Preservation requires anaerobic conditions in permanent likes. swamps, bogs ete, Once the climate becomes dry. these permanently- Soh \ Peachayarina wel sites disappear, Alternate wetting and drying in wes _. Branohe seasonal swamps and lakes destroys petlen. Rulon “3 Moreover, when pollen ts deposited in permanently- : MW MAREE - L Callabora wel sites, 1 must he hunied deeply enough to escape the effects of a later fluctuating water table of future drier climate, HW iLis to reniin preserved. Deep weathering has undoubtedly destroyed much of the Vig 1 Locality map. pollen record, but where pollen has been preserved, Hood pulynofloras were recovered. This paper Cretaceous sequences intersected ii (he Poona documents (he pollen species recavered and reports runna-l well on the palynostratigraphy of the Tertitry and upper Geology ‘ ) Wowie! Science. Universi of New S Wale , : “ae font WNcience, Mniversy OE Nery, Sault Wales Poonarunna-f was sunk to 1,696 im with the SVU ONY Moe. , * - . \ Manet 1 Heol) Well gumptetign report el! Poonmuner Ni | Intention of exploring the Palavozoic or Proterozoic SOU AT SLT Cupid basement strait beneath the Permian or Mesoaoie on) HA MARTIN cover beds. The well established a good reference for Mesoaore studies (Magnier), bul there was no jnterest in the Cuimozoie. The present study concentrates on the Tertiary palynostrutmsraphy tert includes Che uppermost of the Creticcous which vsuiblisbes the Tertiary/Cretaceous boundary, In ihe well completion report (Mugnier'), the following units ure defined from the lithology and correlation ol aaimmieriy lows with supplementary information fran sonie Jogs und resistiviry. The ‘Tertiary and Quaternary were defined as consisting of Quaternary sand and alliviun (025.9 m) and white dolomite limestone interbedded with grey murky. sandy clay (5,.5-95.4 mi). The Lite Creticeaus iConamanian) (95,4-667,5 m) wis dedined as the Winton Pormution, consisting of alternating grey lo dark grey silt, fissile Shale und siltstone wih disseminuted pyrite. The present study records the palynology of the upper 146 m only, Le. the Calnozoie and the upper part of the Late Cretaceatts Sequence, The upped part of the Winton Formation examined in (his study is remarkibly Upiforn in appearance. From the palynolowy, the Cretiecous- Tertiary bonndivy occurs at about 110m. und the interval from 95-110 ti Should be reassigned to the Eyre Bormiation, since the Winton Formation elsewhere mW the Luke Eyre Basin js cortelated with the upper Phimepollenites, pannesus and Plieutelhe dixte- corinatis Zones. of Albin te Cemomaniin age (Krieg epal 1991), The Eyre Formation (Callen et af (995) is a widespread and distinctive fluvial to fluviokicustrine sund unit, but Jithologieal variations oveur i some chunnel facies. Plant fossils are characteristic. cmd Jocally abondint, Caurbonaceous horizons wat the Eyre Formation contain spores, pollen and a few dinoflagellates of late Paleocene to middle Bocene age. The Byre Formation was deposited in large menndering and braided streams (srieg era! T8997), The unit between 5.5 and 95.4 mis correlated with the Etadurina Mormation of probable law Oligocene (6 Pllagene ave. hut it ms most Tikely largely carly Minveng, Phe Bladen Formation was deposited i a evaporative food plan-hicnsting environment, vader ad clinute ner (han thator the Lyre Parmation (Knee eral WO]: Callen er al, T9YS), Matertals aod Methads Only curlines Were available for the present study. The possibility of conmbiminatiol is greater with cuuiags. bul avilh proper drilling and sanmpling provedures. reliable samples may be oblained. This topic as fully discussed by Martin (P98) In the present Sty, rhe Late Cretaceous assemblages conluincd véey few ‘Tertiary grains und barren samples in the Camnozoic section suggest thal coptimination is niinimnal anc these samples are reliable. The samples were freated with hydrochloric and hydrofluore acids to remave mineral matter. Controlled oxidation with cold Schulze solution and polussiuin carbonate was used to eloar the residues. which were then mounted jt elycerine jelly (Brown 1960: Ciray 1965). The palynofloras were assigned to vores using (he ranges of diagnostic species. The Tertiary palyuo- floras were quantihed with counts.of about 150-200 erains and percentages were bused on the total pallen count. The pollen spectra derived from the counts provide a hasis for interpretations af the palsen- verotillon, An assessment of the abundiiee ab micposcapie curbouted particles in the Tertiary ussemblases was mide. The formation of these particles t) conlro- versial, When they are found in mid-late Tertiary and Qualernury sediments, it is generally accepted thal they were formed from burning aid we ehareod particles (Luly era/, L980, Martin 1987; Kershitw ef a. W901). On the other hand. Sehopf (1975) has suvacsted that black curbonised particles huve formed by oxidation at the surtace OF swans However. there have heen numerous studies which shaw hit chareoal may be distinguished from other black varbonised materiul (Scouw L989; Cohen & Spach 1977, Sander & Gee 1990) and chureoul muy be found in sediments of any age, Palynostraligraphy hale Crehiweods paulynosteytigraphy is based on tharol Helby etal, (1987), the systematic paly nology is presented in Appendix |. the species identifedt ae given in Table band the sires of diagnostic speeres in Preute 2, 1/0 146m, Pligatelly distocarinita Zane. Comonentan The assemblages in this zone lack the distinerin Phyllocludidies meaysani’ and Prafeuctlies spp. ol the Romans? Zone Plroytells distucertnade acl Trilobesporites irierereu/ans. whnse ranges gral with the 2 omewsenin Zone are presenr These assemiblawes thus fit the 2 disrecartuna Zune of Cenoommniun wwe (Fig, 2), Addsapallis enicifarns is usally present in the Cenomanint it the Luke lyre Basi (N, F Alley pers, comut 1995), bot it has for heen revarded tronr these assemblages, Burger (1993) reports (hit Foreriimiyprads daily, bas nel been recurded trom the Po diareeerinue Zone, bur i ispresent ip (he sample fram LLS-119 mob this study (Table |). Spores ol ferns, tycopods and bryophytes: are PALYNOLOGY OF THE POONARUNNA NO. | WELL ‘TABLE 1. Late Cretaceous species identified from Poonarunna-1. +, present. ++, common. Depth (m) Spores Aequitriradites spinulosus A. Verrucosus Baculatisporites comaumensis Balmeisporites glenelgensis B. holodictyus B. tridictyus Camarozonosporites australiensis Ceratosporites equalis Cicatricosisportes sp, of Burger Cicatricosisportes spp. Clavifera triplex Crybelosporites punctatus C. striatus Cyathidites australis C. minor Dictyophyllidites sp. Foraminisporis dailyi F. wonthaggiensis Foveogleicheniidites confossus Gleichentidites circinidites Laevigatosporites ovatus Microfoveolatosporites canaliculatus Ornamentifera sp. cf. O. Sentosa Perotrilites jubatus Plicatella distocarinata Polycingulatisporites sp. Reticuloidosporites arcus Retitriletes austroclavatidites Ruffordiaspora australiensis R. ludbrookiae Sestrosporites pseudoalveolatus Stereisporites antiquasporites S. pocockii Stoverisporites Microverrucats Trilobosporites tribotrys T. trioreticulosus Triporoletes sp. ct. 1. simplex Gymnosperms Alisporites sp. ch. A. grandis Araucariacites australis Corollina sp. cf, C. classoides Ginkgoeycadophytus nitidus Microcachryidites antarcticus Podocarpidites ellipticus P. exiguus Podosporites sp. Trichotomonosulcites subgranulatus Angiosperms Cupuliferoidaepollenites cf. C. parvilus Dicopopollis sp. Foveotetradites fistulosus 110- 115- 128- 134- 143- 113 119 131 137 146 + + + ~ + + + + + + + + + + + + oo + + + + + + + + + + ++ ++ +e ++ ++ + + a + + + + + ae + + ++ + ++ - ++ + + + + + + + + + 4 + + 7 + + + + + 4 + + + + + + + + + + oo + + + + + + + + + + + + +f + + + + + + + + + + + + + + + ++ ++ + + + + 92 HLA. MARTIN Manne t continued... Depth (in) atte 115- 28 1ad- \43- 13 119 al 137 lao Lihtucidites sp. et. (. Kattangutaensts + Lillavidites sp. + Phinopollenites augathellaensts ' + 1 PL panninsas t + + Seroeloretrdi(es Varirenicilags t Tricalporites sp. ck To uparyexiiuts + Alpae Roryocncous briaait + Hovologinella sp. + Lecuniella sp. + Saeplodininim gravattensix + + + Sehizosporis relicnlatus + t Acritarch sp. inde. + ACE ZONE DIAGNOSTIC SPECIES Phimopollenites augathellaensis, Po pennosas and Toei: [asameeaen - Foveotetradites fistilosus being the most common MEDCENE | tae ct (Table 1). The angiosperm palynoflora may he swasrrcarn | (Tne placed in Suite IH, of Lite Apuan-Early Cenomanian ‘ 4 age (Burger 1990), for it lacks the triporate forms of & Suite 1V which starts within the Cenomaniin (Burger Tavinrilinenites CAMPANIAN, Nevtrcohaypeditess SONIBLILE ur i = oO | 4 Theoiporites t FE | pawtouan ones CONIACIAN | py jyettnios Hine ve sis TURONIAN TEM a wes 3 nae 3 Ele 15 CENOMANIAN in| ra By a E z 2 ALBIAN Phimopoienies s a PAGS Fig. 2. Ranges of diagnostic Cretuceous species, from ie & Helby ef al, (1987), particularly diverse and abundant in this zone, Cyathidiies miner is very common in all of the sumples and Kuf/ferdiaspera/Cicatricosisporites spp., Foruminisporis wenthagelensis, Gleiehe- niidites cireinidites, Laevigutesporites evetus and Srereisporites untiqitasporites are sometimes abundant, Gymnosperm pollen is diverse and well represented with Podosporites spp. common in some sainples, Most of the trees in the palacovegetation would have been species of gymnosperms, There is a small angiosperm pollen content with 1993), The five sequential samples ure essentially the sume age. but there are quantitative differences. The three oldest assemblages are diverse with a good representation of gymmnosperni pollen, but the angiosperm content is restricted in diversity and abundanee. The abundance of gymnosperm pollen suggests that forest would hive have been a major part of the vegetation. The assemblage at 115-119 m has fewer gymnosperms than the other assemblages and an unusual abundance of Srereixporites antiquasporiies, with affinities to Splegitien, and Foraminsporis wonthaggiensis, similar to the living hepatic Nothylas bruetelii (Dettmann 1963). This assemblage suggests that there were extensive bogs or Wetlands and fewer forest trees in the immedite vicinity. The uppermost assemblage has a diverse gymnosperm palynoflora, indicating forest similar to that of the oldest assemblages, and an inereased angiosperm palynoflora. The megusporangiate water ferns, Balmeisporites spp., are also well represented at this level, The microphinkton content of these assembliges 1s low with only a few forms represented, Lecaniella sp. and Sehizesparis reticulaiuy have probable affinities with the Zygnemetaceae (Grenfell 1995), a family of filamentous green algae usually found 1 shallow, flowing fresh water. Berryececcus braunit may be found in fresh and brackish water (Pentecost 1984) and Saeptodinium vravatiensiy is a tresh water dinoflagellate (Harris 1973), PALYNOLOGY OF THE POONARUNNA NO. | WELL TABLE 2. Jertiary species identified in Poonarunna-1. Percentages of total pollen count are given. +, present but not counted in the counts of 150 or more grains. Depth (m) 6- 73- 9 76 Spores Agalla sp. + Camarozonasporites amplus C. bullatus Camarozonosporites sp. 1.2 Ceratosporites equalis 0.6 Cyathidites australis 0.6 C. paleospora 1.3 5.4 C. splendens + Dictyophyllidites concavus 8.4 Foveotriletes lacunosus Gleichenia circinidites 18 Grapnelispora evansii Laevigatosporites ovats 1.2 Polypodiacoisporites sp. ct. P. relirugatus + Poalypoeliidites spp. 0.6 Retitriletes austroclavatidites Stereiyporites sp, Todisporites sp. Triletes sp. ct. 1) tuberculifarmis 1.8 Triporoletes reticilanis 0.7 + Verrucosisporites Cristatus + Unidentified 0.7 Gymnosperms Araucariaciates australis 1.3 14.4 Cupressaceae/Taxodiaceac 0.6 Dacrycarpites dustraliensis Dilwynites eranidatus 1.2 D. tuberculanis 0.6 Lygistepollenites florinii 8.4 Microcachryidites antarcneus Phyllocladidites mawsonii 0.6 P. retienlosaccatus Podacarpidites spp. 0.7 4.2 Trichotomonosuleatus suiberanulatus Angiosperms Acaciapollenites inyriosporites 2.0 Aglaaridia qualamis Amosopolliy dilwynites Arecipiles sp. cl. A. minutiscabratus Austtalopollis obscurus Beaupreaidites elegansifornus + Chenapodipollis chenopodiaceoides 19.2 Cunoniaceae (tricolporate) 0.6 Cunoniaceae (bicolpate) Cupaneidites orthothechus 0.6 Cyperaceae 10.6 1.8 ef, Dodonaea 0.7 Rlaeocarpaccae Evicipites crassiexinus LS Gothanipallis bassensis Graminidites monoporites 73 05 94. 97- 1O1- 104- 107- 97 101 104 107 110 (16 1.3 0.6 0.6 1o 1.3 7.2 i) 99 24.3 1.9 5.9 34 19 1,9 0.6 0.6 i=) =e NS Ke 3 nt m= te be by S 2 2 3.0 0.6 0.6 “a rm 0.6 13 0.6 0.6 0.6 3.9 0.6 0.6 94 TABLE 2 continued... Depth (m) Haloragacidites haloragoides H. harrisii Hexpollenites austroclavatus Lewalanipollis cf. L. rectomarginis Lewalanipollis cf, Persoonia Liliacidites lanceolatus Malvacearumpollis sp. Malvacipollis diversus M. subtilis Milfordia homeopunctata M. hypolaeniodes Myrtaceidites eucalyptoides M. parvus M. verrucosus Myrtaceae unidentified Nothofagidites emarcidus N, deminutus N. faleatus N. vansteenisii Nuxipollenites kempii Polvyorificites oblatus Polyporina granulata Propylipollis ivanhoensis P. latrobensis P. ef, pseudomoides Pct. P. reticuloscabratus Propylipollis sp. Proteacidites adenanthoides P. adenanthoides/crassus P. angulatus P. annularis P. cooksoniae P. crassus P. fromensis P. grandis P. incurvatus P. sp. cl. P. ineurvatus P. ef. obseurus P. cf. slipplatis Proteacidites spp. Quintiniapollis psilatispora Rhopites alveolatus Rhopites sp. cf. R. alveolatus Santulumidites cainozoicus Sapotaceoidaepollenites rotundus Simplicepollis meridianus Simpsonipollis sp. 0.7 Sparganiaceaepollenites barungensis 0,7 Tricolpites sp. cf. T. asperamarginatus T. sp. cf. T. confessus T. sp. ef. T. discus LT. phillipsti T. thomasti Tricolporites angurium T. lewray H. A. MARTIN 7T3- 94- 716 9O7 12.0 3.9 0.6 + 0.6 0.6 + 0.6 1.8 0.6 13 + L.8 %Q6 + 0.6 0.6 1.8 5.3 + + 3.3 + 13 10 + 0.6 0.6 2.4 18 19 + + 0.6 0.6 1.3 + + + + + 4.8 0.6 3.0 0.6 101- 104- 107- 104 107 110 3:2 2.0 3.9 0.6 4 0.6 1.3 0.6 0.6 13 0.6 2.0 1.9 0.6 + 0.6 + 0.6 0.6 0.6 13 2.0 La 0.6 + + 0.6 0.6 13 1.3 1.9 1.3 + + + 0.6 70 3.3 19 0.6 0.6 + + 0.6 a 1.9 2.6 0.6 + OG 20 PALYNOLOGY OF THE POONARUNNA NO. | WELL 95 TABLE 2 continued... _ ---—--- On —————— —— SsSsSseSsSeSFSFSFSFFFFFssFFFSFFFsffsseF Depth (m) 6- 73- 9 76 Triculparites substriatus 41, paenestriatnus Tricolperopollenites endobaltreus 0.6 Trierites minisculits Triporepallenites ambignis + Lubulifloridites untipodica/simplis 27.8 Unidentified angiosperms 79 6.0) Alpae Borrvocwecus ++ Debarya + Morkallacysta pyramicdelis Pediastram + Summary of major pollen groups Spores 2.0 21.5 Gymnosperms 14 29.9 Casuirinaceae 5.6 12.0 Myrtaceae 3.3 2.4 Cunomaccae/Elaeoearpaceae Nothofagus 10.8 *Proteacede” 3.0 Asteraceau 27.8 Cyperaceae 10.6 1.8 Poaceae 7.3 Reshonaceae 2.4 Sparganiaceae 0.7 0.6 Chenopod type 19.2 94- Q7- LO1- HO 107- oT 101 1o4 107 110 2.4 + 2.5 9.1 13.6 15.0 143 + + + 7 + + 92 O7 Wd [1.8 3.7 Si3 51.5 Cama) WS 42.5 3:9 3.) 42 2.0 3.9 La 0.6 20 54 24 6.4 13.2 5,8 12.4 144 12.8 9.2 8.5 1.9 24 13 0.6 1.2 13 2.0 13 [.3 1.9 3.3 0.6 TT These Cenomanian palynofloras are generally similar to those of Bathurst Island, northern Australia, described by Norvick & Burger (1975), except that the former contains freshwiater microplankton whereas the latter has a very diverse marine dinoflagellate flora. Norvick & Burger oecurrences of (1975) illustrate the known Cenomanian palyoofloras and, except for three non- marine localities in the Eromanga Basin, all the others are around the northern, western and southern periphery of Australia. The stratigraphically important species of the Cenomanian have been studied. but other than this study, the report by Narvick and Burger (1975) is the only Australian report to document all the palynomorphs in Ceno- mantin assemblages. Tertiary palynostratigraphy follows Stover und Partridge (1973), Macphail (1996) and studies in Central Australia (Krieg ef af, 1991, Sluiter 1991; Alley ef al. 1996; N, F. Alley pers. comm. 1994). The systematic palynology is given in Appendix 2, the species identified in Table 2 and the pollen diagram in Figure 3. The definitions of the pollen groups are given in Table 3. Taser 3. Definition of the major pollen groups used in the pollen diagram, Mig. 3. A tut list of taxa is presented Appendix 2. Name on diagram Taxa included in the group Spores All spore taxa Arauicariaceae Arancariacties australis Podoearps Padocarpidites spp. Lagvarestrobes Phylloclaclidites mawsanti Dacryelisn Other gymnosperms Lygistepollenites flurinii All other taxa under gyninosperms Cunoniaceae Cononiaceae and Elaeoearpacede Casuarinaceae Haloragavidites harrisii Myrtaceae All species of Myrtaeeielites Nothofagus All species of Nothofagicites *Proteicene” ALL species of Beaypreaidires, Lewalanipollis, Proteacidites and Propylipallis ‘Callitriche” Anstralapollis obscurus Cyperaceie Cyperaceae Poweoae Graminidites monoporites Asteracene Tuhuliflovidites spp. Chenopad type Chenopodipollis chenapodiaceatides ey HA, MARTIN Samples examined MAJOR POLLEN GROUPS 9? 4) & jotal pollen count ZONE AND AGE LAKE FROME SURFACE POLLEN SPECTRUM POONARUNNA No.1 WELL Acacia seen joses of s es 5 1 eeee Onher aCage a2 ahacese Gymnosperns Pagocspus Daenydiun Cunonmacsaey Elagsocar Lagarosrrobios Casuarnnace: Fi. 3. Pollen dis Pilocene-Pleislocane Ss SS SS IS SS S NSS SSVOV SS SSS SSS Ys Y oy % wy Barrer Barren Upper L.baimer equiv Lala Paleaoene BeaA P aistacarintata Cenomanan Cyperaceae Astirat Crenapod type ucam. For definitions of the pollen groups, see Table 3. The surface pollen spectrum from Lake Frome (Shutter & Kershaw 1982) represents the present vegetation ined is added for comparison, J 2 WTO om. Upper Lygistepollenites balmei Zee equivalent, later Paleocene Gymosperoy tax are diverse and their pollen may constitute mare thin half the count (Pig. 3. Appendix Ay. Podocarpus (Podocurpidites spp... ts the most common. and Lagerestrobos (Phyllaclaciidites mawseail) is well represented. Pollen) trom the unviosperm taxa Cunoniaceue/Elacocurpaceae. Casnurimuceae (Heloragacidites harrixit) and Myrlaceae (Myriacidites sp.) is present but that fron Norhafirei (Nothofigiditey) has not been detected in this study. although is is usually present bul extremely rare tn this zone (N, FL Alley. pers. eonm, N04) There is a profusion of the proteaccous lax Bedipreidiies, Lewelanipaltis, Prapilypelliy and Profeacidites spp, (see Appendix 2) with Protedidites cadenanthaides, Po cana PF crass, 2 frameusix, Po cookseniae. Po leduhtonit ad Poreflesus the most common, OF Cellfirfehe Wustalopollis obsciris), Restionaeaw GWilfontia spp.) and Cyperaceue (Cyperaceaepalliy sp.) are present alsa. There is i number al anidentuhed digiospermn pollen ivpes present. and while they mary not be miportane for swatieraphie Consideration, they Indivale wrieh and aibunckint aneiospeon element i the hue Paleocene, The content of curbortsed prirlicles ts bas These asembliges aee usstened to the Upper L baline! Zone equivalent. The oceurence wf Cihapirelspert evitsee ied Trinealpits eho- confessus, whose vanges end ab the top af the Maastrichtiun (Helby et al. 1Y87) ts anomalous. but they may be reworked. Other distinctive taxa usually found in the Muustrichtian are lacking and quantitatively, these assemblages are unlike those of the Maastrichtian. Comparisons When compared with the L. Meliner Zone of the Gippslind Basin, the stratigraphic ranges of some diagnostic und churucleristic speaies are not the same in the two regions. For example, Proteccidites erassus, Po lelehionii and Po reflevtis. present in Poonarunna hegin their ranges in the Hocene of the Gippstind Basin and PB framensis in Poonarinni his not been reported from the latter site (Stover & [yin 19732 Stover & Partridge 1973). In contrast, Cunonidceac, Cyperuceae and Milfardia lypel- cqenrodes OF this study are not found in the Gippstind Basin. The rich and abundit profeaceots tori are a feature Of The assemblages i central Avstraha bat they are “heither particularly diverse nop partiectatly common im those of the Gippsland Basin (Stover & tvans 1973. p. 59). Phere ure similarities. however, Hat “vinnosperm Lisa aire diverse aad common in both localities, although the Hontinite species of the gone. Lo belmer, as not been found to this study. Phyllocladiditey spp. und Australopelles obscutiis dre characteristic of both regions, Che late Paleoeene L. bale’ Zone is reported trom the Southein, Mono ow the highhinds of southeast Australi (fayloc ead M00) Gyomosperi poten PALYNOLOGY OF THE POONARTINN ANOLE WEL n Jominates some of the ussemblayes and Neviofayns may be abundant in others, Angiosperm taxi are well represented anid the proteaveous clement may be relatively abundant in sone: Austredopotliy-vbycuruy wid spewies of Mvriceeidifes have not been recurded from the Monaro, unlike the other tare Paleocene lovalitivs, Cunoniatede, Cyperaceue and Resti- onideac are lacking, in keeping With other localities in Southeast Australia. but in contrast to thal of central Australia In the Otway Basin, the Gembierina edwerdsii Zouule (Harris 1971) i lite Paleacene in age also (Stover & Partridge 1973), Gyimnosperm pollen is abundant and A. abscess usually present The protetceaus HAA are Well Pepresented in tie Otway Basin. but Cunonincene, Cyperaceae andl Resti- onaceae have not heen recorded, Sluiter (1991) records date Paleocene palynotioras from the BMR Mulvorina 2 bore. southeast of Lake Pyre. The assemblages are dominated by an excepuonall abundanee (up to 07% ) of Cunontacene. Gymnespenn pollen is combo aid Netiofages mire. similar lo those of Poonarunna-!. but the proteuceous taxa ure net particularly abundant, The late Puleoeene-carly Hovene suite is similar but with wil mcrease in Myrtaeeae. the proteaceous taxa and Ansralopillis obscurys, With an overall decrease in evanospern pollen (Stuiter 1991), The palynofloras ob these lwo localities near Lake Tiyre are thos very similar bur Pounarunnd-) has much less Cunonmicede PO-U4 ot Pollen is extremely sparse in dis interval and is insufficient for stiuly. Berrvececens, however, is sometimes abundant, indicus a fresh-beackish Water lake environment, Carbonised particles ure present throughout in small amounts bur are particukuly abundant at tbe 79-85 m level. 73-70 nt Lower Nothofacuires wsperus “evr eauevatent, und kocene The eymnosperm pollen cantenct ts less Una teat of the 2. botnet Zone equivalent but os soll comsidenible, Attuciriieene — LArdiderridcies aistrlty, Dilwwnites -arenatardin) and Detereliinn (Lariepallentios foraniy ire the ost abundant in (his group, The anwsiospeyin Hora is paruculathy eet) With Carstiarijaccae (Haleragectites hurisyit) wid Netholasus (Nutialagtdites spp.) moderately abundant The spore vontent (ferns. bryophytes. Iyeopods) is mocerate. The proteaceaus content 1s restricted in diversity and vihundanee. The sarhonised particle coment ts low, This assembluve is similar to that of the mid Bovene unitot Peachawarinna-2 of Lake Eyre Basin (Sliiter (294) As well es (he general chaructertstics expressed ubove. Cunonfuecac or Myrtaceae (Myrraceidites spp.) may be common and the proleaccous conten! may be more diverse in the usseinblages of Peachwwariana-2. [also contains species which Hirst appear ut the late Eovene Middle Nothofugidites asperus Zooe of the Gippsland Basin, View Auluoridia qualmix, Proleacnlites ch PB, wipplatus and Tricolpites themisii (Stover & Partridge 1973. 1982). However Slumer (1991) rewards all of these species fron the mid Bovene unit of Peachawarinna-2, Assemblages al Nelly Creck, southern Lake Eyre Basin (Alley eral. 1996) are dominated by pollen of Casuaripaceae (Aelerugacilites Aeisyiiy anil gyninosperm pollen is relatively common with Araucatiaceae (Araneurtacites australis, Dilyyiites granwaiuy) the most abundant Angiosperm pollen is well represented and there is a swwealth of protcaveous species. Myriuceae (Myrracesdites) and Nathafagius (Nothofiiyidies) are consistently present, Tieolpites thamasirand Proleccidiies stipplaiis, with ranges of Lower and Middle V. asperus Zone equivalents in central Nustralia. are present alsa (Alley ev ef, 1996), The Poonurunni-| assemblage thus compares well with those from Nelly Creek, with the exception Unt the proleacequs componenl is more diverse and abundant in the latter, The Poonirunna-! assemblage is therefore assigned to the Lower NO dsperny Zone equivalent of mid hocene age. Commperiveny Mid Eocene macrofossil assemblages at Nelly Creek. Poole Creck and in Some of the Silerete floras have a number of taxa with botanical affinities i common with the Pooparunna-b microfossil assemblage, and the weneral characteristics are similar Four proteaceous teal faxu. Ageriis (Ariucariicede). Podovarpaceae. Civinaastonia (Casuarinaceae) and Myrceipln thon (Myrtacese ). ire listed (Christophel af al 1992), More detailed comparisons OF macro- and microfossil assemblages ure pat possible beeause of the dillerence in provenance, transport and preservation of the plant parts Fouad br (he owe types of assemblages. leat physidgnomy yields invaluable evidence about the veseraition and (his topic (§ discussed baler, Tie Pooburunna-l assemblage bus less Nothofiguus und more Casuaringceat when compared qwith the Lower Norlofieidites ayperus Zone of the Gippshuid Basin (Stover and Partridge J973. 1982. Partidye 1976) Aswell. the minges of muny Species are not the same in the tyr localities. Phe nen-woody swamp taxa of Cyperacede. Restionacedeé and Sparcaimnicear have not heen repented fra the Lower Nothofagidites oyperus Zone of the Gippsland Basin. This Poonarunna-| palynoflora is probably time equivileal to the Prateacidtites pachypolas Zone af OM HLA. MARTIN the Ouwuy Basin in Victoria (Harris 1971) whteh os rich In Nothefavas and proteaceous taxu, Mid Eocene floras are also found ifthe St Vincent Basin, South Australia, where the palynofloras have wy abundance of fern spores but frequencies of symnosperm, Casuarinacene und Notiofagis pollen are Jow, There is a diversity of proteaceous and other angiosperm pollen. will no single wroup dominant (Alley 19872 Alley & Broadbridge 1992). When the mid Eocene palynoflora ab Poonarunna is compared With those of Anglesea (Christophel e7 a, 1987), the latter has more Nothofevus and fewer gymnosperms, Palynofloras at Bungonia (Trussell & Owen 1988) also lave few pymnosperms when compared with those in Poonurunma. 61-73 m Pollen was not recovered from this interval bul the alea, Momyveceecus, is present and very abundant from 67-73 m. The 70-73 ny level has Peers also. These algal species indicate a fresh-hrackish water lake cnvironment. The carbonised particle eontent is low through (his interval, except for te 67-70. m level where tt is high, YO] i Pollen was. not recavered from this interval. Curbonised particles are present ihroughoul, increasing Up the section, except for the 16-20 in level where (hey decrease. f-Goin. Plineene-Pleisiecene Asteraveae (1 7ithulifloridites spp). Poaceae (Cadmmuriites monoporites) and the chengpou type (Chenepeadipullis chenopodiacevidess make tp the bulk ol the pollen coun, Casuarinacene (Halaniuaciudites — harristiy antl Myrlacene (Mvrfaciddes spp) are present. but pyiibospern pollen ind spares gre miniial, Curbonised particles dre extremely common, This type of assemblage has mob heer feconted previously from central Australia and there is ne direer means of dating 7h The general quantitative uspeers of the ussemblage suggest oy Pleistaeene ae. bused upon experienee in southeastern Australia (Marti) 1987) ohut i licks the distinctive Pleistocene Hihuhflavidies plewstecenteus (Macphinl 1996). Palvporing aranidara of late Miodene Pleistocene (Mawphi) (86) os present. Preqhertcies similir to (hose shown in Fig, 3 may be Polit i shieibliids and regions lonay (Stuiter & Kershaw TY82), When the Pouonarugia-T assemblave ts compared with surface sainples lront shroblands vround Lake brome. ii is romnurkubly similar, except Cora greater CY periceae pollen content. suggestiiy hat there was yore swam vesetation in the Pliseene Pleistocene Hin there sat Lake brome today Tubaliflaridies spp. Yurse appear in the rad Miocene of Southeastero Australia (Stover & Partridge 1973), bat there are wuimerous reports: ol these species from the early Miocene aid atew fray the Oligocene (Muller l98}io My experience bis shown that most pollen types are not common inthe carly part OF their ranges. but become more abundant faite, AL Poodaruinna, Tidiiliflertdites is the mast abundant group, Suggesting a relatively your ape. ie. Miovene if no Pleistocene. 1-61 No pollen sas recovered, The charcoal) particle: content was very high. Palacoveyetation and palaeoenvironment The Cenomiuniin palynoloris are donmimiued by bryophytes, lycapods. pleridephytes und wymno sperms but the angiosperm content is relatively small and this Suggests that the vegetation was mainly Forest with gymposperms forming the (major part ol the canopy, if not the whole of it One palynoflory sigwests more extensive bogs or wellands, its discussed previously, These palynofloras predate the #ppearance of proteaceous and Netlofaguy pollen types Which become a distinctive clement in-yourper palynofloras, The hte Paleocene palynofloras of Poonsrinn-| indicate what the region was iainly forest. The eymnosperms Padecarpuy (Padacdnpidites spp. Lawaraytrobos, the Huon pine Ce aaledidites maser) and Daervdiuin (Lypistepollenites Hlermnin) were common. ind there was a diversity of aiher Yyminosperm tind, 2. Decrvearpis (Dace rarpites), Microcachyrs (Microcachiryidites). Cusuia- rinucewe (Heloregacntites herrisii), Nethafasus (Nathofasidties spp), Myrtaceae (Myrracerdifes spp.), Caunomaceae/Elaeoctrpicede were pact of the forest Canopy, though probably a relatively (ior part. There is a wealth of profeadeous pollen ty pus bul mest of them cannot be identified with living tina. AL leust some of ther are Jikely to have been lprest trees, Similar to the protwacenus (rees foand iy rainforests toray. Proteuceous species have iW very luw pollen representation (Kershaw 97, O71 1976, Murti 197%) hence. Wese eXciner praleneeniis haku Were probably fur mere ubiidont i the Maleocene vevetation than is suggested hy the pallens frequencice, Swamp communities were Revie i extent aad conlayned me on-wwoorly fina ef. ‘Cullivicvhe® (Avatralepoltis, ebscuruy) (Macphal 990). Cyperaveae ane Restionuceie, Other wigiospenms were diverse (Table 2) and if they were law pollen producers. ds is tie case with most insect pollinated species. they were probably mote abundant than is Sugwestéd by the pollen canits PALYNOLOGY OF THE POONARUNNA NO. | WELL 99 A lresh water Jake environment with copious Botryococcus and some Pediastrum succeeded the late Paleocene floras, through most of the presumed Eocene, There is. however, insufficient pollen for study through most of the sequence. Within this lake sequence, there is a layer of sediment with a goud pollen content, suggesting that the lake had receded from this area and that it had become vegetated. The id Bocene palynoflora in this layer suggests mainly forests with little non-woody swamp vegetation. Arwucariiceac, Dacrvdiam, Casuarinaceae and Nothofagus were prominent, the proteaeeous content Was low at this particular location and there was a greal diversity of other angiosperms, The Pliocene-Pieistovene vegetation was probably open woodland or shrubland, generally similar to that of today in arid: abd) semiarid regions, ‘Trees, if present, were probably Cusuarinaceae and Myrtaceae. bul both of these families contain shrubby Tarr +. Register vf strated specimens. taxa as well as trees. Acacia (Acaciapollenites miyriosporites) is present, and as Acacia has a very low pollen representation (Sluiter & Kershaw 1982), it may have been common in the vegetation. similar to that of today. The very Jow frequencies of Araucariaceae and Podecarpus may have resulted from long distance dispersal. or there may have been small, rare stands in the landscape. IL is remarkable that a Plio-Pleistocene palynoflora has been preserved at all, A wetter climate and some condition(s) which allowed a more rapid burial of sediment must have prevailed for a brief interval, IP itis accepted that most of the carbonised particles result from burning, then the Tertiary recerd (Fig. 3) suygests that burning increased during the presumed Neogene, reaching a maximum in the Pliocene/ Pleistocene. There was, however, episodic burning in the Eocene. Very lew carbonised particles are found in the late Paleocene and Cenomunian palynofloras. CRETACEOUS SPECIES Acriuirch sp. indet. In Nequiniradites splinieasius 4A,B Aeainriradites verracisus 4b F Afisporires spect. As aranedts ON Arainaridedtes duatralts OB Bacilalisporttes con ancusts AG Baliieisporites wlenelwensis SA Balmeispovites hialodicivus 4H Balmeiwporiles ftaludictus Al Bilmeisporites tridictyus 4B. C Balineisporites tridietyuy 40 CONTE TOS IMOSPONTIOS USI OHNTS Ay CONN ONPOVUON UUSIELTOHNTS 4K Ceratosporiios equatis SLL FP Creatricasinportes sp, of Burer 6C Chewiricasixportes sp, of Burwer 6D, F Claviferi triples 6] Clavifera triples 6k Corellia sp. ch Oo clesseidos eo Cryhelosporues puanerdtin OL. M Crybelosporttes [ACHAHIN tC Cryhelaspurites sirhaus TALK Crpulifereidecpoullenties el, Co prarvuliey oI Cyaridites ustealis ol Cyathulites minor 6H Picopapalliy sp. OM. N Dictvophyttidites sp. GN Forwninisports daily ra8) Peel ispeeis won ae edenyis it) hoveauhliheniidites confossus Th Poveatetradies fistilasus OA, B CM HOLES Cire dadidites oP Horalegiedht sp. HON, Slide No. England Pinder coordinates 3060-3 Wa3-0 BOGS A O35) 3064-1 163-0 A060-3 K4l-2 A064-| p55.4 3064-| 135-| 3060-1 035-1) 3UO0-3 J51-1 A0O4-| 560-0) 3060-| M34-1 3062-| 056-3 3063-1 P47-2 3060-3 V53-\ 3062-3 Cid4-4 3004-3 S40-4 3004.3 R44) 3065-2 MS5O-( 3003-1 R29.) 3002-3 Vaud SOOO 4 O40) AMY Var AOO2-] R440) OH4- | W322 $64 3 S5N-] WAL 3 Md5-| 3000-1 Jas-| 3060-3 S454 3060-4 Pls.) S004. | W53-1 A -3 33-0) A003 P31-2 WH 3 S40-2 Mi 42 L603 LOO H. A. MARTIN TABLE 4 continued... Species Fig. Slide No. England Finder coordinates Laevigatosporites ovatus 7G 3060-3 Y42-4 Levaniella sp. 10C 3063-2 M54-0) Lecaniella sp. 10D, E 3063-3 O#0)-2 Levaniella sp. 1OF,G 3063-3 F50-0 Lecaniella sp. 101 3061-1 H36-3 Lifiacidites sp. cl. L. kattangatdensis oP 3060-1 $33-0 Liliacidites sp. 9S 3060-1 N29-3 Microcachryidites antarcticus 9D 3060-3 V42-0 Microfoveolatosporites canaliculatus 8K, L 3063-1 QO43-0 Ornamentifera sp. cf. Q. sentosa 7TH, 1 3062-1 O53-0 Perotrilites jubatus 71..M 3064-2 Feo-4 Perotrilites jubatus 8C 3064-2 N49-2 Phimopollenites augathellaensts 90 3060-3 O36) Phimopollenites augathellaensis 9Q,R 3064-2 Q52-0 Phimopollenites pannosus 9U 3060-3 O36-3 Phimopollenites pannosus OV 3060-3 GS0-0 Plicatella distocarinata 4c. D 3060-3 N37-4 Podocarpidites ellipricus OF 3960-3 X43-2 Podocarpidites exiguus OF 3060- | M294 Podosporiies sp. OH. 1 3060-3 X39-4 Podosporties sp, OI IK 3060-3 Y44.2 Polycingulatisporites sp- SA. B 3060-3 F54-3 Reticuloidesporites arcus 8G 3063-3 039-0 Retiriletes austroclavatidites 8D 3060-3 Y45-0 Retitriletes austroclavatidites 8E, F 3060.2 P40-0 Ruffordiaspora australiensty 6A 3003-3 J54-3 Ruffordiaspora ludbrookiae 6B 3060-3 M36-0 Saeprodinium gravatiensis 1OH 3060-3 Y37-0 Schizosporis reticulatus 1OK 3064-3 O40-4 Senectotetradites varireticulatus oT 3060-1 R40-3 Sestrosporites preudvalveolatus 60 3062-3 K41-3 Stereisporites antiquayporites SN 3061-3 O50-0 Stereisporites antiquasporites 80 3061-3 H46-3 Slereisporites pocockit 6F, 3063-1 Q47-0 Stoverisporites microverrucatus 8M 3061-2 O59-0 Trichoromonosuleites subgranulatus 9G 3060-2 U65-0 Tricolporites sp. ch. 1) apoxyvexinus OXY 3060-1 M6l|-3 Trilohosperites tribotrys of 3064-3 H45-0 Trilobosporites trioreticulosus 8H 3060-3 R48-4 Triparaletes sp. ef. T. simplex &J 3060-3 X4A0-4 TERTIARY SPECIES Acaciapollenites iyriosporites 13C 2983-1 N34-4 Aglaoridia qualamis 7G 3000-3 P4]-2 Amosopollis difwynittes ls 3057-2 $40-3 Amasapoallis dihvynites 13F 3057-1 N36-0 Amosapollis dilwynites 13G 3006-3 D31-0 Araucariaciales australis 121 3000- | O40-() Arecipites sp. ct. A, miniuiseabratius 13D 3006-1 X42-1 Australopallis obseurts I3N.0 3059-2 TS3-0 Acolld sp. HW 3000-2 J45-0 Beaupreaidites clegansiformis I3I,K 3000-2 LAS-3 Bairyocoecus braunit I8B 3000-1 V42-3 Barryecoccus braunis 1sD S000- | V42-] Cumarozonosporires amplus WA 3006. | W38-4 PALYNOLOGY OF THE POONARUNNA NO, 1 WELL 101 TABLE 4 continued... Species Fig. Slide No, England Finder coordinates Camarazonesporites amplus 11B 3000-3 042-1 Camarozonosporites bullans 11D 3059-2 O40-3 Camarozonosporites 8p. H1E,F 3000-2 E47-0 Chenopodipollis chenopodiaceoides 13L 2983-1 Y53-0 Cunoniaceae (tricolporate) 13S, T 3058-3 O41-0 Cunoniaceae (bicolpate) I3L.V 3057-2 T37-1 Cupressaceae/Taxodiaceae 12D 3058-2 $55.3 Cyathidites paleospora UW 3059-1 O60-2 Cyathidites splendens HG 3059-1 V45-3 Cyathidites splendens UH 3059-1 V52-2 Cyperacenepollis 13M 3000-3 D39-0 Dacrycarpites austratiensis 12L 3059-1 $50-0 Dicopopollis sp. 18k 3000-1 29-0 Dilwynites eranulatus 12) 3000-1 R42-0 Dilwynites gramtlatus 12k 3006-2 W39-3 Dilwynites eranilatus IBA 3006-2 V58-0) Elaeocarpaceae 13Z, AA 3005-1 K39-0 ‘Ephedra’ notensiy 13B 3006-2 133-4 Ericipites crassiexinuy 13P.Q 3000-! R30-0 Gleichenia circmnidues 11c 3059-3 LS9-2 Gothanipollis basxensis 144A, B 3006-3 W57-1! Grapnelispora evansit 124 3059-2 Q43-3 Halavagacidites haleragoides 13R 2983-1 030-2 Huloragacidites harrisit 1G 3000-1 V4l-4 Hexpollenites austroclavatns 14D, E 3059-2 49-4 Lewalanipollis tf L. rectoniareinis L6G 3006-1 F43-4 Lewalanipollis of. Persoonie ISI. K 3000-3 C55-1 Liliacidites lancealatus L4P 3058-3 V50-0 Lygistepollenttes florinti 12G 3059-2 P4)-1 Malvacipollis subtilis 14h 3000-1 Q28-0 Micracachrvidites antarcticus 12N 3059-2 L34-1 Milfordia homeopunctate 14c 3000-1 228-0 Milfordia homeapunetata lay 3000-1 PAT-3 Milfordia hypolaeniodes 14k 3006-3 049-0) Myrtaceudites eucalyptoides 140 2983-1 Q33-1 Morkallacysta pyrantidalis 1kC 3059-2 Q28-1 Myrtaceidites eucalyptoides 14R 2983-1 V41-2 Myrtaceidites verriicosus 148 3000-1 S40-1 Nathofagtdites emarcidus 14L 3000-1 K58-2 fothofagidites demitutiis 14N 3000-3 T48-0) Nothofagidites faleatus 14M 3000-2 P36-4 Nuxipollenites kempti 140, P 2983-1 J46-1 Pancolpate sp. [SE 2983-1 139-0 Panporate sp, 18G 2983-1 Q40-1 Pediastrum sp. tSA 3000-1 R53-2 Phyllocladidites mawsonii 12E 3059-3 R43-0 Phyllocladidites reticulosaccams 12H 3059-2 Q42-4 Podocarpidites exigiuus 12M 3059-3 K39-1 Polyorificites oblatus I4BB.CC 3000-1 O59-1 Polypadiacoisporites sp. ct. P. retiregatus 11L,M 2983-1 J43-2 Polyporina granulata I7A,B 2983-1 L46-0 Propylipollis ivanhoensis 147 3006-3 Q47-3 Propylipollis ivanhoensis 14W 3058-2 Q58-2 Propylipollis latrobensis 14X 3005-1 U40-0 Propylipollis sp. cf. P. pseudamoides 16B 3006-2 V64-0) 102 H. A. MARTIN TABLE 4 continued... i Species Fig. Slide No. England Finder coordinates Propylipollis sp. ef. P. pseudomoides 16C 3058-1 $56-4 Propylipollis sp. cf. P. pseudomoides 16D 3006-3 Q47-3 Prapylipollis sp. cf. P. reticuloscabratus 14U 3006-3 $433 Propylipollis sp. cl. P. retieuloscabratus 14V 3058-1 $56-2 Prapylipollis sp- 14y, 7 3006-1 X39-1 Propylipollis sp. l4AA 3005-1 X43-4 Proteacidites adenanthoides ISA. B 3058-3 O49-2 Po angulatus 15G 3057-1 X57-0 Po angulatus 15H 3059-3 I38-0 P. cooksoniae 16A 3006-2 MS50-0 P crassus Isc, D 3059-3 159-2 P. fromensis 1365.7 3059-1 049-2 P. grandis IS] 3059-1 R60-4 P meurvatus ISN 3000-1 $40-0 Psp. ef, Po ineurvetis 16N, O 3057-1 K34-1 P. ef, obscurus 16d. K 3006-1 K46-1 Pel, stipplatus lol 3006-2 $58-0 Proteacidites sp. | 16E. F 3059-3 O41-0 Proteacidites sp. 2 16L,M 3006-| MS1-0 Proteacidites sp. 3 16H 3059-3 Q51-2 Quantiniapollis psilatispora I7L. 3000-3 C53-4 Retitriletes austroclavatidites 12B,C 3059-3 N59-2 Rhopites alveolatus I7E 3000-1 $40-2 Rhapites sp. cf. R, alveolatus 17P,Q 3059-2 U65-2 Santuldmidites cainozoteus 17F 3000-1 X44-2 Sapotaceoidaepollenites rotundus IIR 3059-2 T62-1 Simplicepollis meridianus 17C, D 3000-1 KS4-2 Simpsonipollix sp. 13H, 1 3059-2 $27-0 Tricolpites sp. cf. T. asperamarginatus 17H, 1 3000-1! P50-4 Tricolpites sp. cf. T. confessus 170 3059-2 R41-0 Tricolpites sp. cf. T. discus 17M, N 3000-1 Q40-0 Tricalpites phillipsii 177 3005-1 W434 Tricolpites thomasii 17U,V 3000-1 W39-1 Tricolpites sp. 180, P 3058-2 Q5}-2 Tricolporites angurium WW4 3000- | P29-2 Tricolporites leures I7TW, X 3005-1 Q46-0 Tricolporites sp. | 17€C 3058-2 N41-0 Tricolporites sp. 2 I7AA, BB 3057-1 T33-4 Tricalporites sp. 3 18] 3058-1 P55-2 Tricolporites sp. 4 [8s K 3058-1 W530) Tricalporites sp. 4 I8L 3058-2 N42-4 Trivolporites sp. 5 18M, N 3057-1 W33-3 Tricolporopollenites endobalteus I7R,S 3000-1 062-0 Triletes sp, ct. Ty utbereuliformis LIK 3000-1 N54-3 Trierttes minisculus I8Q 3058-2 $55-3 Triorites sp. 18H 3059-3 T33-3 Triporoletes reticulatus 12F 3059-2 S57-3 Triporopolenites ambiguus I7Y 3000-1 L43-4 Tubulifloridites antipodica/simplis I8R 2983-1 §39-2 PALYNOLOGY OF THE POONARLINNA NOL | WELI 104 The development of the arid Mora and vegetation The aim of this study was te find fossil evidence about the development of the arid flora and veweuition, The late Paleocenesmid Bocene palynotloras deseribed here. with a@ substantial tainforest element. are clearly not arid adapted. The vegetation, however. grew ol the floodplains und depositional basin, and such habitats would not be the first in the hindscape to register aridity. Middle Rocene macrofossil assemblages from other lowalities in the Lake Eyre Basin have some large- leuved Land. Consistent with inforest. and ip variety! oF smaller leaved taxq, suggesting selerophytlous yegetulian (Christophel e/ af, 1992; Christophel 1904). The veeetation is interpreted as being gallery nudtorest along the watercourses und selerophy Tous vegelutiin, adapted both 1 low fertility and scusonally dry conditions. in the intertives. These maveofossil assemblages are unique to cenrral Australi (Greenwood ef a! 19980: Greenwood 1904). The diterval betWeer the imid BKocene and the Plincene-Plerstocene cid not yield pollen, The Phiocene- Pleistocene assemblage is penerally similar to thitt produced by the extint arid Shrublanus ot Lake Vrome (Fig. 3) but contains some disparate clements, Dodomeiea Iriqguetra (Nuxipollentions kempii) ts tound in this assemblage, bul the modern species is restricted to wet seleraphyll forests of the southern half of the cast coust of Australia (Martin 1997). Dodenaed tiquetre 15 also present in the mid Bocene of the Lake Eyre Basin (Sluiter 1991) and probably became extinct in this region at some time fier the Pliocene-Pleistacene. There huve been miuny studies On the Mora of the arid zone that have generated Various hypotheses ubout it} orgins, As many of tbe tisit to the arid one show affinities with related taxa in adjacent regions. miost of the hypotheses involve reeruiiment from the floras surrounding the arid yone (Barlow 1981). Such studies, however. do not reveal taxa which once grew in the arid zone and have beeome extinet there. such as Dedonuea triquetra, ‘Vhe fossil record sueesis thatthe vegetatn developed by continneus adaptation of some of the taxa already in the fegion (by evolution of new species) to a drying environ: ment. Those taxa that could nor adapt te the arid environment disappeared from the region. Acknowledyments Tam indehted ta N-P Afley of the Department of Mines and Energy Resourees (formerly the South Australian Depariment of Mines and Energy) for Invaluable assistance with this project which seas supported by wn Austrilinn Research Grant, References ALLEY, Nb (1987) Middle Bovene jee of ihe megalissil florid at Golden Grove. South Australias Preliminary report and Comparison wth the Mastin Bay Bor, Zraves. Ro Sde SMa, Gd 2212, — & Beoabweibar, b. M- (1992) Middle Eacene palvaotlorin form the One ‘Tree Hill urea, St. Vineents Basin, South Austria. Adhering bb, 241-267. — Reog. GW Cathy. RAL (1996) Trarly ‘Tortiany Lvre Formation, lower Nelly Creek. southern Luke Evee Basin Austrtlia: palynolsgieal dibinge ul nuterafioras win Silerete. and palteaelimatic inter prenitians, Aust df Eetrnh Set. ABT | B84. Uacrionsh. J F978) Palynologicnl zonition of the Lave Jitessic apd barly Cretaceous sediments of the Yarrujadee Formation, central Perth Basig, Western Austalit, Rep. Geol, Sern WA, 1-33, Baniow BAC I9RT Phe Australiin Floris Its Origa and hevolurion pp. 25-75 Ja George, Ao. ded.) "Plova or Austiilia Voll (Australian Government Publishing Service. Canberra). Bist, ALN. (1981) An early Plideene assembhige front Lake Tay. sauth-western Australia, and its phyt geographic inplications. Aryio2 Bot 29, 277 201. BE ACKHURE, KB. (1936) Betryecorcys and the aleal caals, Try Roy, Soe, Edinburgh 38, 840-854, Buows.C. Av (1960) *Palynologicnl echniques” (houston Stare University, Bato Rouge, Burcbk, D970) Farly Cretaceaus angiosperm petlen erdins trom Queensland, BMR Geol. & Ceaphys Bull 6. 1-16, (1973) Palynological observations im the Carpentaria Basin, Queensland. 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Vier 77. 1-148. __ (YTS) Angiospermous pollen fram Albian fa Toroniin sedimeats of ester Austraba Gael Sues dad Spacey Prebl, A, 3-34. & Janzen, DM, (1496) Pollen of proteacenus type from flesh Cretaceous sediments. sotith-enstert Atistralia. Weheringa 20, 103-160. & Prayhokh. G LlY6K) Thkanemy ob sone Cretaccoys spores and pollen urpins romp eslern Austr, Pric. Rea, Soo. Vien 8h, G1-b3, 5 ~& (4969) Palynology of the Australian Creétuceous: A review pp. 174 200 Ay Campbell. KOS. W. thd i Suvitigraply jd Palaeaatolopy. Fskays i Honour of Derathy JAP CAustraliin National University Press. 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(1994) Palacobetanical evidence for Australian Tertiary climates pp. 4-59 J Hil R. S. Fal.) “Australian Vegetation History. Cretaceous ta Recent” (Cambridge University Press, Cambridge). Cappny, R. AJ & ALLEY, No F (1990) The correlation and depositional environment of Tertiary strata based on macrofloras in the sourhenm Lake yee Basin, South Australia. Wepre Mines and ken SA Rey Bk 90/15. 1-57. HALL. J. (1974) Cretaceous Salvininceae, avai Me Mot, Gen OL, 354.367 Thageis, WOK, (1965) Basal lertiary micnfloras from (he Princetowrarea. Vielorut. Australia. Pulieoniee BS, TA-106.. (1971) Tertiary stratigraphic palynalugy, Onway Basin pp. 67-87 Ja Woploer, He &e Doulas, 1 OL (baba) “The Otway Basin oF south-eastemt Australi’ Spee, Hall, Geal. Survy SA anid Viet (972) New form species Tram seater Australian early Tertiary: sediments. Trai Ae Sac 8, Mant Of, 53-65. {1973) Terlitry ponemarine dingthywellile eyst assemblages from Australia, Geol Sop Anan Spey. Publ. 4, 159-} 66, Hashkti. T. Ro 11968) Siteoate pollen grams beam ite Lower Cretaceous at the Great Arlesiin Busan Australie Univ Ott Papers, Dept Grol. @. 21-243 Newer Uh (1972) Pollen and spore assemblies. fran Oneenslind leetlary sediments, Geel, Suri of Gk Paty, ASS. Palavontol. Pap, 30. 1-31 Dhihy, Ro. Moan, RO & Bane. AL AL ORT) A palynolowical comation of the Australian Mesozoic pp. | 100 7y Jel, PA, (eel) "Studies im Austeatiin Mesovone Palynology (Assocnibon of Australes Palen tologists, Sydney). Hrusshe. Cod (1971) "Pollen and spores of Chile! (line ol Arizona Press. Tucson). hin, BO M1976) Farly Perkivy pollen frum Napperby, eentral Aushalia, BM Jonnie! af Creal & Geeplys 1 fod (14, & Hagens, WK, (1977) The palynaloey oF early ‘ferimery sediments Ninetyeast Ridge, Indian Oeean Filaveautol. Assoc. Spéc. Pap. in Petaweny. 19. 1-70. KER Ww A OP (1970) A pollen digerin front lake Ihuramoo, north-east Queenshind, Australia, Mew Phytel OY, TRAAROS, - (M971) A pallen diagram fram Quincan Cries, north east Queenshind. Australia, Ibid, 70, 669-68 |. A (1976) A lato Pleistocene and Molocene pellen diagram trom Lyochs Crater. narth-oastern Queenshund, Australia, /hre, 77. 464-408, (1985) An extended Miler Quaternary vegetation record tra northecastem Queensland and tis implicuhions Fur the seasonal tropics oF Ausinilia, Prow Evel See. Anat 1. 1792 ).89 —, Dievisra, DB, M.. Mek wen Mason, BO) & Warisivpe By Et 1991) Polynological evidente tue Quulerury eoviTorments of the maith south-eyster i Austria. Qua Sei Rew Th 391404, RandoG. Wo. Reis. PAL CALLAN. ROAL PREEMA®. 2 J. AULEY, NOOR & Fokehs, B.ooG. (99h) [250,000 Geological Series Exphiiatory Notes, Chedimurka, South Australia, SA Dept Mines &e Hinensy- Loy. J. Spcrrnk, ER. & Kershaw. A, PCL9RD) Pollen studies of Tertiary brown coals: Preliminary anabyses al lihwtypes within the Latrobe Valley, Vietorias Muniyli Publ. Geogr 23, \-78- Macreane M. BK. (1990) Anvrredepalliy obsereris (Maen) Krutzeh emend. Stover & Partridge. 2A News! 26-6. 2. PALYNOLOGY OF THE POONARUNNA NO, | WELL. (05 (1996) Palynostriligraphy of the Murray Basin. infand south-eastern Australia, AGSO Record 1996/57. & TRESWELL, EM. (1984) Palyniostratigraphy of ihe central west Murray Basing AMR J Aus), Ceol, & Ceophyys, TL 304-331, Marrin, A, R, ER (1973) Reappraisal of same palynomorphs OF supposed proteaceous affinity 1, The venus Beaupreaidiey Cookson et Couper and. the species Promacidites hakediles Couper. Geol Soc Aust. Spee. Pall A TRTS. Martin, HA, (1973) Palynology of some Tertiary and Pleistocene deposits, Lachlan River Valley, New South Wales, Aut, 2 Bat. Sapp. 8, 1-57. (1973b) Upper Tertiaty palysiology in southern New South Wales. Geol Soe. Ausi. Spec. Publ, 4, 35 S46 (NYT) The history of Hey (Aquiloliacene) with speciil reference to Australias Lvidenee trom pollen. Aust 4 Bor. 25. 655-673, _ (978) bvalution of the Austrian fora and Veyelitioh thrauuh the Ternary, evidence front pollen. Afeheringa 2, }&b 202, (198du) The use of quantitateve eclationships and Paleoewoloy in stealigeaphic palynology af the Murray Basin in New South Wales, dbl 8, 253-272, (IYK4b) The statieraphic palynalogy oF the Mitnay Rasin tn New South Wales, J) The Marrumbidgee area, J. Prec. Rey Soe NSW AULT, 35-44, (7987) ‘The Cainoaoie history of the vepetation aid climate aC the Lachlan River Region, New South Wiles. Prac, Lint, Soe SW 109, 24-257, il (1904) UNUstaliin “Tertiary phytowcoursphy. evidence [rom palynolagy’ pp, 104 (42 Jn Hill RS, (hd) ‘Australie Vegelation History, Creticeuns te Keven’ (Cambridge Unversity Press. Canibridpe ). (1997) Tho use of ccolugioal tolerances for the reconstruction of Tertiary palneoelimates, dawn Bear 45. 475402. MoM, A, (L003) Palynology of Sites 815 and S24; the Neogene yegerition Instory of coastal north eastern Austiilia, Pee, ODP Serenifie Res, 133, 115-125. Metwryph. D, 1. (1965) Some new pollen species trom New Zeuland Tertiary deposits, N74. Bor, 3. 204-214, (L968) Murther bew patlen species fant New fool Tertiary and uopermost Cretaceous depasils, hid, @, (77-204, Mitbpstan, B,C. & Pockyan. PT (1989) Migecne- Plemtovene spores und pollen Fram Central Obie, South Ishin. New Zealand MZ Geel, Suet, Palaenutol, Bill. SY 1-126 Mibsb, 1. F. ClO88) Palynology oF a late Kocene dignilie sequence [rom the western marin of the Cuela Basi, Weartern Australia. Meni Awe. Ane Pulawomtol & 285-310, Muitern, J. 11981) Fossil pallen records of extant ingiasperms. Bor Kev 47, 1-142, Nomick, MOS. & BukGek, D. (1975) ‘Palynology of the Cendoninian Ob Bathurst Island. Northern Territory, Austr’ (Alisiiiiiin Govt Publishing Service, Canbera), Pyne, ALD. (1976) The geological expression of custiiy inthe Barly Tertiary of the Gippstind Basin, AMEA drarenel Wi, 73-74 Pea thous! A. (198d) “Introduction to Freshwater Algae (Kingprint, Richmond. Surrey), Pres. M, D.& Crrmsropnne. D.C. (1978) Anytrasequoia Wwinfonensiy..a new taxodiaceous cone from Queenskind, Alisrratia. Cattad. 1. Bot. 36, 3119-3128. Prhavrorp. G. & DirverMaAnn, Me FL (1978) Pollen of Hacrydivn franktinit Hook, Po and eompurable early Tertiary microfossils. Pollen er Spares 2, 513-534 PockNAnL, DB. T & Croskike Yo M- (1982) Tayenomic revision of some Tertiary trivalporate und) uicolpate gris trom New Zealand, VZ0 Bor 20,715. & MOEDeMIALE. PC (1984) Miocene-Pleisiacene spores amd pollen from Central Otage, Seay Island, New Zealand, NA Geol Sur Malaceireal. Bull, SY, L128. Sasbek, POM, & Gk, CLT. (1990) Fossil charcoal fechniques and applications, Rey Peleeohat Palynel. 3. 269-279, Seroer J, MA (1975) Modes of fossil preservation. Jon. 20, 27-53. Seay. ALC. (1989) Observations on the nature and orivin of fusain, dateraut do Cot Geol. 12. 443-475. SpucrER, LOR, KK. (199)) barhy ‘Tertiary yeeetalion and climates, Lake Byre region, north-eastern South Ausralia pp, 9% 136 Ja Williais. Mo AJ. de Deckker P& Kershaw, AP (Bes) The Caimovoie in Australia: Reappraisal al the evidenve’ Geal Saco Aust Spec. abl. 1. ®& Kershaw AL P9822) The nature of the late ‘Perliary vevetation in Austell A ermge 6. 24) -222. Stover, by B.& Byans. BOR, (1973) Upper Crericeors spore pollen yvonution. offshore Gippsland Basin. Australia. Geol. Soo. Aust Spee, Publ, AS5-72. & Parigipat. A. DO. (1973) Verity and Lare Crelaecous spores aiid pellen from the Gippstand Basin suuthedstern Australia. Pree. Rea See Mier, 88. 237-280 —& (1982) Eoeene spore-patlen from rhe Wenllup Pornation. Western Qustalia Pulyielnuy @ HO-OS, ow CEOS) A new file Cretagepus meguspore with grapnel-like appendage dips flem Australia and New Zealand. (bid 8. 139-b44. Taylor, G PRESWELL, EM. McQubes, RG. d& Brown, M,C, (1990) Early Tertiary palieageayriphy, Gindform evolution, and palacoclimates of the Soulhern Monica. NSW, Australia, Palaeoueng. Pulaeoelim. Pulieoeent, 78. 109-)34. TRUSWEHI LL Ma Shbrimk, 1 RL & Harr. WK. (POSS) Palynology of the Oligocene-Miocene Sequence in the Qakvale-1 -corehole, western Murray Basin, South Austvalia. BMR J Aust. Geol Geaphys. 9 767-295 oo & Owen, J.-A. (1988) Boeede pallen trom Bungonit. New South Wiles. Mew deseo Anat Pethetcontal S. 259-289 TSENG CHENG, Tuan C1972) Pollen Plog of Taian’ (National Taiwan Univ, Press, Triped. Tunin, J. R Tavhok, G. & TRuswine, be Me (1442) Palynolowy of Term Luke Bunyen, Cooma, New South Wales. AMIR. Ads, Genel & Geophys. 7, 255-208, 106 L.A. MARTIN Appendix | Late Cretaceous Systematic Palynology, For the distribution of the species in the bore, see Table 1. Por the register of illustrated specimens. see Table 4, Spores Genus Aegultriradites Delcourt and Sprumont emend, Cookson & Dettmann 19Ot lype species: Aeyutriradires dubius Delcourt & Spromont emend. Delvourt, Dettman & Hughes 1963 Aequitritudites spiuilasus (Cookson & Dettmann) Cookson & Deltmunn 1961 FIG. 4A, B Comments. The spinulose clements over the distil sut- face ure about | tim in diameter and the exime | um thick, Compare with A. verrivesis. Spore body, 45 pm. overall, 55 um, Stratigvaphic Range, Marly and Middle Cretaceats (Detunann 1963), Aequitriradites avcus/spinilasas, Tron the Muroypara florida Zone. late Jummssie, into Phyllocludidites mansani Zone, Tironiin-Contacion (Helby et et, 1987), Aequitrirudites verrucasins (Cookson & Dermiunn) Cookson & Deltriunn 196) FIG. 4E. F Comments. The verrucate elements over the distil sut- face ure 2-3 pint in diameter and the exine 2-36 yun thick, Compare with A, spiudeasits. Spore body, 48-57 pm, over- all FO-RS pom. Stratigeaphic Runge. Widely distributed in south-eastern Australia in the Upper Mesovorw (Detain 1963). Cienus Bevalatisporites Thomson & Pflug 19534 IWpe species: Keculerisparites pennarins (Wolff) Thomson X& Pilg P54 Bacutadsporties cameiunensis (Cookson) Potonie L956 MiG, 4G Stratigraphic Range, Prom the late Jurassie-larky Cretaceous. it is common throughout the Upper Mesocote (Detimann 1963), Genus Balmeisporites Cookson & Dettmann L958 Type species: Balineisperites halodicyins Cookson & Derrmann [958 Balueisporiter gleneleensis Cookson & Dettmann 1958 FIG, 5A Comments. This species is similar to Bo falevtictviey bul the spore body is larger and the exine thicker: The inner homogenous layer is 5 um thick on these specimens. conte pared with 12 pin on A. fralpdiecivus, Spore body, 162 4 112 um, Stratigraphic Range, Within the Plicdsela distocertretis Zone. Cenomanian, to within Tricelpires puchvextrens Zone, Santoman, of south-eastern Australia (Detimann & Playford 1969), Cenormanian, possibly Turaniiun of nerth- west Australia (Norvick & Burger 1975), Balmeisporites Ralodtcwus Cookson & Dettmann 195k FIG. 4H, 1 Comments. Most of these lurge megaspores are broken, The spore exine Consists Of un toner, homogenous layer b2 po thick and an outer granular aver about 5 yigr thick, the fat ter supporting the muri of the reticulum. Spore body, #7- 110) pm equatorial diameter, overall, 137-166 pin xs 170-235 pny Stratigraphic Range, Cyhelasperites sirtaiusy Subsone, Late Aptian-Early Albian, to the lower part of the Appendivispories distocarmans Zone. Cenomaniao (Dettmann & Playford 1969), Balmetspovites tridiciyuy Cookson & Dettmann |YS% FIG, 5R-D Comments. The absence of a reticulum, a thick. ier homogenous layer (5 pin), an outer granular layer. 1-2 pm and the large membranous winglike oulgrewths distin- guish this species (Cookson & Detiinann [Y5K). These specimens show sinuous ridges 4 pot high (arrow), whieh MAY anastomose, Spore body, $2-85 8 42-110 pin. Stratigraphic Range, Aptiin-Albiun (Cookson & Detimunn 1958; Detonann 1963), Cenomunian. this study, Genus Cumarezenesporires Pant 1954 ex Potonie 1956 cinend, Klaus 1960 Type species: Refasperiies eretaceauy Weyland & Krieger 1953 Camarosonosporites austrutiensis Norvick & Burger 1975 FIG. ALK Comments, The distal surface has coarse rugulae about 3 um wide, Separated by grooves | yim wide, On the proxinnuil surface, the pattern ts finer und the contact surfaces ure almost smooth, This species is smatier than CL aanplies, 2%- S7 um compared with 57-109 pin respectively (Norvick & Burger 1975; Dettmann & Playford 1968). Equatariel diameter, 40-44 pum. Stratigraphic Range. Albian of the Great Artestan Basin und Albian into Turoniaia of Northern Australia (Norvick & Burger 1975). Genus Ceratosperites Cookson & Detumanin LOSS Type Species: Dettmann 1958 Ceratasporites equilis, Cookson & PALY NOLOGY OF THE POONARUNNA NO. | WELL 407 E Figs 4. Cretaceous species. A, B. Aeguitriradites spinutosus. C.D. Plicatella distocarinatis. Eb. Aeguitriradites verruce- sus. G. Buculatisporites comaumensis. UW, 1, Balineisporues holodieytus. J. K. Canimoconosporites australicnsis. Scale bars = LO pm. 108 HA, MARTIN Cerafosparites equalis Cookson & Detain 1958 FIG. SE, FE Stratigraphic Range, From the Revitrileses wetheroensis Zone, latest lurassie (Helby en uf 1987) trough the Paleocene (Hurris L965; this study), Clomiricusisperties sp. oF Norvick & Burger 1975 FIG, 6C-E Deseription. Spore tilete, ainb tiangulin, sides slightly convex lo deeply concave, Most spores hive deeply con- cuve sides, appear 3 lobed and present in equatorial view, Lasiree ex tench almost To equator, membranaus fps 4-5 put high, Three sets of parallel muri dinastomose in radial region OF distal surface. Edges OF muri irregular or wavy with small Fovule Within muri, especkily where sets of mur anustomose. Four muri and intervening grooves, &- 10 ym, Equatorial diamerer 43-53 ain, polar diameter 35-40 pin, Comments. This species is distinguished trom: Auifferd- laspord dustralaiis by the Tegular ature et the muri Norvick & Burger ()975) fuure an undeseribed species, pl, 20 Tig. 4. similar to this one. Strativraphie Range. Norvick & Burger (1975) note that their undescribed species in the Cenomanian iy ehuraenristic ofthe Upper Albian in Queensland. Cenomanin, this study. Genus Chavifera Bolkhovitena YOo ‘Type species: Clavierd frinfex (Bolkhovilinwy) Balkhoy- Hind YOO Clayifera iiples (Bolkhoyitinad) Bolkhovitina 1966 FIG. Ot, K Comments. The distal surface is strongly arched and the proximal pytaniidal, The nifterhadial crassitudes ure 4-5 200 thick. features Which distinguish it from Gleietenidites. Us alfinities are with the Gleicheniaceue, Stratigraphic Range. From within the Caproyporna par deve Zone, Albitn. ii northern Australia und rare in the Micatela distocainde Zone, Cenomaniin, southeastern Austiihia, through the Forcipites Jamu Zone. Maasti- chlian (Helby ef cal L987). Geous Crybelosporires Detinann 19634 Type species: Crybelesporites sirtatuy Cookson & Detimann L958 Crybelasporifes punctatis Detimann 1963 FIGS 61, M, 7 Overall diamerer, 36-55 4 28-48 pon. Stratigraphic Range. Lower Creticcous (Dettinann 1963), Cenomanian, this study, Crybeloyporites strats (Cookson & Dettman) Dettmann 964 PIG. 7A, B Comments, The sclerine is 4-5 pind thick with a henge. neous ioner layer | pin thick and a ruffled outer layer which is irrewularly Strigte on the proximal side and ceticulite on the distal surface. The mur are thin and sinuous, and the lumina 4-4 pm wide, All these feattires are a guad fit with Co sirfeeties. Stratigraphic Range, Crvbelesporites steletis Zone, inte Clavifera Triples Zone. Late Aptian into Turenian (Dettmann & Playford 1969), ©. sérietius Zane through Phimopollenites pannosus Zone. latest Aptian through Albian oF gorther Australia und continuing inte the Tubulifloridites litter Zone, early Maastrichtian of southern Australia (Helby ef ef 19ST). Cendimaniin of northern Austrailia (Norvick & Burner 1975) this study), Genus Cwathidites Couper |953 Type species: Cyathidites australis Couper 1954 Cvarhidites australis Couper [953 FIG. 61 Stratigraphic Range, Common throughout the upper Mesozoic in southeastern Australian (Det 1963), Prom the Permian (Poster $979) into the Tertiary (Hitrris 1965). Cyaidites miuar Couper 1953 FIC), 6H Comments. Very conten to this study — Strtigriphie Runge, Prom the Jurassic (Dettinane 1963) info the Tertiary (larris 1963), Genus Dicnophyflidites Couper emend. Detlimann 1965 Type species: Dictvophvlidites harris’ Couper (58 Dictyvaplryllicites sp. FIG. 6N Comments. There is considerable variation inthe popilie tion assigned to this genus, Genus Feruininisporis Keutgsch 1959 Porenmninisports wort egien sis (Cookson & Dettmann) Detimann (63 FIGS 60. 7E, F Stratigraphic Range, Prom the Rufferdiaspora austrulient viN Zone, cartiest Cretaceous. 10 the Phyllectudidites niin yet Zone, Turoniin-Coniieiin (Helby ef el. LOST) Forannisporis dear (Cookson & Detiminn) Detmiunn 1903 FIG 7D Stratigraphie Range. Widely dispersed in the Upper Mesozore of southeustert, Australia (Dettmann 1963). Gettus Poveduleichentidites Norvick & Burger |Y75 Type species: Fovewaleichentidites (al. Glejehentidiies 3 comassiy (Hedlund) Norvick & Burger 1975 Foveauleichentidites Comfossus (Hedlund) Norvick & Burger 1975 FIG. TILK Description, Amb toiangular with rounded upives, (rilete lusurae thin and straight. reaching to apices, Exine | yin thick with interradial crassitudes up to 6 pm wide, Fovulae. < | pm in diwmeter. spaced up te | pin apart accur on bork surfaces, Equatorial diameter, 30-32 yim. PALYNOLOGY OF THE POONARUNNA NO. | WELL LOY B Vig. 5. Cretaceous species continued. A. Balmetyparites glenelgensix. B-D_ Balmeisparites iridietvus, EE. Cerataspurites eglalis, Seale bars = 10 um, 110 L.A, MARTIN Stratigaphie Range. Cenoniunian of northern Australia and sporadically in the Albian of the Great Artesian Basity (Norvick & Burger 1975). Genus Gleicheniditey Ross ex Delcourt & Spruniont. emend. Dettmann 1963 Type species! Cleiehentidites senonivus Russ 1949 Gleicheniidires crecinidites (Cookson) Dettmann [963 PIG. OP Strutizvaphie Range, Gleieheniidites spp. first appear in the Ewly Jurassic CHelby ef ak PORT Ch cireduiedites ts vommon in Upper Mesozoic sediments af southeastern Austalia (Detimann 1963), tis comparable to Gleielienin ond ranges through the Tertiary tothe present diy, Genus Loevigatosperites Vruhan (Y33 hype Species: Leerivafesporites vileerds brim) brat 1933 Linvigaresporites ovalis Wilson & Webster 1946 hIG. 7G Comments. A conmmnion und widely distributed species in the Upper Mesovene (Dettinanh 1963. Norvick & Iurger 1975) and (hrough the Tertiary, This very common in some af the samples al this study, Genus Microforeolaiasporitey Kriteseh L959 Type species: Micrafoveolatosporites conalicilatis Dettmann 1963 Micrafoveatuiasporiies canulicniaus Detunann 1963 HIG, aK, L Stratigraphic Range, Albian ob the Great Artesian Basin (Detlniann 1963; Norvick & Burger 1975), Cenomanian of northern Australia (Norviek & Burger (975) und central Australie (this Study), Genus Orawnentiferd Bolkhovitina [966 Type species: Ornamentiferd echima (Bolkhoviting) Bolkhovitind [Go Ornamentferd sp cl QL ventavet Detionn & Phivtord 19o8 Pie, WL | Deseniplion, Ah wiinetlay with romnded angles, illete sear will elevaled pnrembranous lps. Inferridial crass itudes. Ho 20 tin long, 5-6 pio wide, bear simious ragulae <1 yr tinh aod tpn wide, Rugulae extend over distal surtace, Prosonel surlace palteried witht low verrucae, Diamerer, 32 a) Comments. The rugulite pattern ayer the eriussitudes di fers Hom OO) seitesa whieh has erassitides with serrate fein. Phe pater atthe distil sures Covers the entire supface On TES form whercis ibis restricted to a reiieular area. With the pices io the inlerradial region on GQ) sents (Detmano & Plinford 1968). This specimen is similir fo the ane Hanred by Norviek & Burger (1975, pl, 23, fig. 3), WiHhOUL deseripnien, Stutighiphie Range. Por OQ) ventas. within the Ticalporites apeyvetinus Zome ti within the Nothafavidties Microtlora, Conmeiin to Canpamin (Deumann & Playford 1969), From the Trreedperites apoxvextins Zone through Parcipites lorigus Zone. Santonian through Maastrichtian (Aelby ef al 18) Ornamenifera ch, OO. setosa, Conemanian (Norvick & Burger 1975: this study), Genus Perotrilites Ledtiman ex Couper 1953, emend. Evans L970 Type species: (designated by Couper 1953) Mororeifites vranulatus Couper 1953 emend, Evans 1970) Perotrilitey jubatus (Dettmann & Playlond) Evans 1970 FIGS 7L.M.8C Comments. This species is distinehve with two math Vidges bering spinose crests, running mere or less parallel to the trifere lisurae on the distal surlice (Cpseudomuri ol Norvick & Burser 1975). Spore body. 45-58 pin diameter, Zona, 25-30 pun wide, Stratigraphic Range. PAinepollenites penmoasis Zone, Lite Albian through Claviyere ripley Zone, Early Turoniu (Detimann & Playford 1969). Genus Plicatela Maljavkina Tt Type species: Pleaella trichacanthe Matjay kina 110. hy subsequent designation of Potonie [G0 Plicatella dixtocerinita (Dettmann & Plavtord) Dayies }O8S HG, 4c, D Comments, Parallel muri occur on both distal und proai- mal surlyee. “The three sets of mut run parallel to the equa tor ind on the distal surface, coalesee ta form a fin-like pre jection ii the radiil region, height 5S pind and projecting 5-8 yn beyond the equator. The muri are 2-4 pin Wide dnd the grooves, 13 pm, Equatorial diameter, 54-62 kum, Stratigraphic Range. Fron within the Captesycre pave doxa Zone do within the Plivllocladidites nidiwsenil Zone, Albiun through “Purontan, stirling cartier fi narthern Australia thar in the southeast (Helby ef g/, 1987), Geos Molvementarisporites Simonesics & Kedeves ; ! . enend, Playford & De ina 1965 lype species: Molveingulattsperites corculiy Simmonestes & Kecleves 1961 Palveingulatisparnes sp. FIG, BAL B Descoiphon, Amb sabelreulir, tilete lasunie have shiek ened nmureins, 2-3 pin wie, wah rewular stviations about 2 ben apant, Leuatarkal thickening 293 pin wide, distal supkice hears (Wo circular concentric ridges. Surface is psi Dinneter, 48 pn. Comments. A rare species ih this study. Species oF Palvciigilatispariies are more Cypieal Of the Jurassic, but some May be found i the Cretaceous (Phivtard & Detinain 1965. Hetby ef al CLOT). Genus Merenvidusperites Plug 1953 Type species! Reticulaldasporitey deutdtiis (PAlag) Llue ast PALYNOLOGY OF THE POONARUNNA NO. | WELL 111 Fig. 6. Cretaceous species continued. A. Ruffordiaspora australiensiy. B. Ruffordiaspora ludbrookiae. C-K. Cicatricosisporites sp. of Norvick & Burger, PL G, Sfercisporites pocockii. H. Cyathidites minor. 1. Cyathidites australis, J. K, Clavifera triplex. L. M. Crybelosporites punctanis. N. Dictyoplyllidites sp. O. Sestrosporites psuedoalveolatns. P. Gleicheniiditey cercinidites, Q. Foraminisporis wonthaggiensis. Scale bars = 10 pan. 12 H. A. MARTIN Reflenloidasporites wenus (Balme) Dettinann 1963 FIG. 8G Stratigraphic Range, Jurassic and Lower Cretaceous sed- dents (Delton 963) Cenomanian, this stidy, Genus Retiietliies van der Hammen ex Pierce emend. Doring. Krutzsch, Mat & Schultz in Krutzsch 1963 Type species: Revitrileres globasuy Pierce 1961 Remarks. Lyeopediumsporites has been restricted to forms with loveo-reticulie sculpture formed by pits closely spaced to form a reticulum. Reritileres aecommodiates a positive reliculite sculpture formed of raised muri (sec the discussion in Backhouse 1978). Reditriletes austreclavertidites (Cookson) Doring, Krutvsch. Mai & Schultz in Krutasch 1963 FIG, SD-F Stratigraphic Range, Widely distributed in Jurassie and Creluevaus sediments. Cienus Kufferdiaspera Dettmann & Cliltord 1992 Type species: Raffordiaxpora Gil Madirigisparites) ais- Haliensis (Cooksont Dettmann & Clifford 1992, by subse- quent desmnation af Dettmann & Chitford 192 Kuffordiaspora australiensts (Cookson) Dettmann & Clifford 1992 FIG, 6A Comments, The narrower muri distinguish this species trom &. fdbreakiae, The muri have straight edges, thus tt is distinctive from Cleatricosisparites sp. of Norvick & Burger (1975). Stitigeaphic Range. Prom the Nuffordiaspora australien- viy Zone, earliest Cretaceous (Helby er af TY87) te Clavifera triples Zone. early Coniacian (Detumann & Playford 1969). Ruffordeispora tidbrookiae (Dettmann) Dettnnann & Clifford 1992 PIG, 6B Comments. The wider muri distinguish this species from © pustraliensin. Swratigraphic Range. From the Crvbelosporites stylasis Zone. earliest Cretaceous, lo the base of the Coptespure juiradoxa Zone, latest Aption-earliest Albin (Dettmann & Phiylord 1969), This species is rare in Lis study and sone: whit corroded: henee Wo may be re-worked, Genus Sesraspertties Dettmann 1963 Type species: Cinewlatiyporites prendoeleulatas Couper 1958 Seatrapuriies predidodiwalainiy (Couper) Detar 1963 FIG. 60 Djaumeler, 35 pir, Stratigraphic Range, Upper Mesozoie of southeast Australie (Denman 1963). Canis Srereispariios PV (99 Type species: Srereisporiies srereaides (Pole & Venité) Pilug 1953 Sieveisporites aniiqueasporiies (Wilson & Webster) Detomann 1963 FIG AN, O Comments, Pullers of low verrucae, <1 pt henht, 2-5 um wide. creates negative reticulum on distal surliee, Pattera varies [rom barely perceptible (Mig, $O) on small specimens to conspicuous (Pig. SN). usually on lurger spee- imens. Equatorial diameter, 26-46 jm, Stratigraphic Range. Mesozoic and Tertiary strata, Sreisparties pacodckit Burger L980 FIG. OF, G Stratigraphic Range, Vneommon in the Barly Cretaceous, Similar forms bave been found in the Cenomanian (Burger JO80), Genus Staveriyperiies Norvick & Burger 1975 Type species: Sloveriyparttes aferaverrucaniy Norvick & Burger 1975 Stoverisparites micraverrucatuy Norvick & Burzer 1975 FIG, SM Comments. Creseentic shaped elevations which delinnt or partially enclose circular or elliptical shallow depres: sions are characteristic of the genus, This species dilfers from Staverisporites tintariy (Cookson & Dettmann) Novick & Burger 1975 in that So mierwerricatiy his Mmicroverrucate ornamentation. Diameter, 32-35 jim. Stratigraphic Range. Cenomanian of Bathurst Ishind (Norvick & Burger 1975), probubly the Albian of the Carpentaria Basin (Burger 1973). Cenomianian of Central Australia (thts study). Genus Trilehayporites Pant ex Potonié 1956 Type species: Trilehosperites hanwenious (Delcourt & Spumont) Patonie 1956 Trilebosporites tribotrys Dewmann 1903 FIG, Bt Striligeupice Runge. Lower Cretaceous ta the Olwiay and Great Artesian Basins (Detimunn L963), Cenaniiinan. this stuicly Hilohosparites Woreticulosis Cookson & Detimiann 1958 PIG. SH Statigeaphic Range. Fran the Coplasperit pandas Zone too within the Plicatella distocutinaia Zone, latest Aptian into eurly Cenomuntiia (Dettmann andl Play lord 1969), Cenomanian of northern Australie (Noryick & Burger (W753), Genus Friperaletes Michedlishvilt 1960 emend. Playlord 1971 Type species: Jriporeleies simentarty Miehedlishyilh ih Mtchedlishyili & Samotavieh [960 PALYNOLOGY OF THE POONARUNNA NO. | WELL 113 B Fig. 7, Cretaceous species continued. A, B. Crybelosporites striatus. C. Crybelosporites punctatus. D, Foraminisporis dailvi, B, F. Foraminisporis wonthaggiensis, G. Laevigatosporites ovatus. H, L, Ornamentifera sp. cf. O. sentosa. J, K. Foveogleicheniidites conjossus. L, M. Perotriletes jubatus. Scale bars = 10 pm. V4 HOA MARTIN Triporoletes spel, To smmplex (Cookson & Dettmann) Playford 1971 FIG 8] Comments. The specimens of this study. when compared with 7. simplex, have a thinner mner layer of the selerine, 279 Be ARR pupae, TD) bervie JSAM seine dit as paratypes: gall. coteeted with holaiype, ADIGA SLO? [STISA],. Mei (Pigs 27) Colowe Head and thomas brown, abdomen with splerotiseadl paris. browu anh fon-selerotised parts wry Head: Antennas seape broadest distally, as long as dicta! brewdth. [Sx length pedieel pecieed broader Hie lone Mopellomeres ba 14 re rumiben fest and sevehd fol Tased, neck about ey lenwih node: CHCUHHH COMprisMme (WO transverse and two Jonna! Bands. PaAdpus three-seaypented, Bye facets pounded, close together, spapser al verles, eye bridve 5-6 fitels lone. Labelle rowehly tere spherical. literally with 2-5 selae. Prony with [2-20 selie per side, Thorns: Wing length 2.2 mim (hO-24 n= 50. width) LO im (O,8-1be Rs same thickness: cntine length. slightly curved posteriorly. joining C anterior to apex; Ry joining C near wing mid-length: Se cell pigmented and together with Ry and adjacent part of R; bearing scales. Claws toothed, empodinn as long as claws. pulvilli hall length empodiun. Abdomen: All tergites with pair of sensory sete in anterior corners and row of setae posteriorly. lergites Tand 8 additionally with few selue scattered meso- laterally; stermites 2-8 with pair of sensory setae anteriorly, a row of sctue posteriorly and a band of sctue mesally. Genitalia: gonocoxites cylindrical, ventral articulation with wonostylus longer than dorsal articulation, setose and setulose: gonostylus uboul same width entire length, sctose and setulose throughout, with strong tooth, comblike distally; cerer separate, selose and setulose: hypaproet bilobed. with one seta apically on cach lobe, setulose: purdametes setulose, with G-8 setose apical papillae: wedengus come, Fumule (Vis 813) Colours lead and thorax brown, abdomen with sclerotised parts brown and non-seleroused parts ced, Head: Flagellomeres 12-13 i mamber. terminal ones sometimes fused. neck about Yj length node. Thorax: Wing length 2.0 mm (ha, = 5), wiih OS min (0,6-0.9). Tergite § Wath single pare of scosory setae anteriorly, sclerotisation undivided, am shape of Jeter “sa. Ovipositor: cert fused into single, termingh spheroid lamelli, setose and setulose; Aypoproct rounded apically i dorsn ventral view, bearing (wo selue posteriorly, setulose. Other characters as in male. Pupa (Figs (4-16) Colours antennal and frontal hoths pigmented, brown, remaininie parts anapimented, Leagil) 26 mm (2.5 2.8.0 = 5). Antennal hops strana, biti, JO pm 6172-206) long, Proms on eae sides are frontal ford, pare oot papillae on lower fice. one selose, one asetose: Fiplet ot lateral taeil papillae gue setose. wa uselose, Prothoricte spitiels with several irregular protuberanges apreally. (ches ending between hall and distal third of spracte. Intesurment of ubdeming segatents euvered with spice. very dense dorsally. ii daesul spities Present, Lest inyter teary Cees be, 08) Colours piokish ped. hength 2.5 iy t2a-2.6, 1 = 0). Integuiment covered with spieuhie, ead with postero-lareral aporlemes shorter (tan heal length, No sparahe present, ATL papillae with shark setae, Thoricie and first ablomioil segments will pain al ventral pupillae. (wo pairs of pleneal papilhte. tree pairs of dorsal pupillie, Abdomind segment & with pain of yeotal papiliie. Ave pairs of pleural papillae, ANEW GALL MIDGE FROM LAWRENCIA SQUAMATA I41 Figs 2-7. Male of Khopalomvia lawrenciae sp. nov. 2, Head in frontal view. 3. Last three flagellomeres. 4. Sixth Magellomere, 5. Last larsomere with claw, empodium, and pulvillus. 6. Genitalia in dorsal view. 7, Wing. Scale bars = 100 tim 2, 3; 50 pm 4-6; 200 um 7. 142 P. KOLESIK Rises fh. ; / oe 10 ‘o"o @ Wr ity te Hein rtater My Mee Pelee 13 Figs 8-13. Female af Rhopalomyia lawrenciae sp. noy, 8. Posterior end of abdoinen in dorsal view, 9, Posterior end of ES | ve I ovipositor in ventral view, 10, Last three flagellomeres (paratype 121397), 11. Sixth flagellomere. 12. Posterior end ot abdomen in ventral view, 12. Last three flagellomeres (paratype 121398). Scale bars = LOO pm &. 10, 12. 13) 50 pum 9: 25 fim TL. fay > ae a ee, A NEW GALL MIDGE FROM LAWRENCIA SQUAMATA H4i Figs 14-18. Rhopalomyia lawrenciae sp, nov.. 14-16, pupa, 17,18, larva. 14, Anterior part in ventral view. 15, Anterior part in lateral view, 16, Prothoravie spiracle. 17. Two terminal segments in dorsal view. 18. Head capsule. Scale bars = 100 pt 14, 15.17: SO um 16, 18. pair of dorsal papillae. Terminal segment with four pairs of terminal papillae. Anus ventral, Etvinalowy The specific name meuns “of Lawrencia’, the host plant, Gall and biology Leaves of Lawrencia squamata infested by this gall midge are several times larger than normal in volume, 4-6 mm long and 3-4 mm wide (Fig. 1). Each gull contains a chamber occupied by one larva. The chamber wall is lined with a thin, hard, pale-brown layer of tissue at the time the larva is fully-grown, Pupation takes place inside the gall. The pupa raises two thirds of its body outside the gall before the adult breaks through the antertor end of the pupa. On 8 Seplember, 1996. on the southeastern coast of Hindmarsh Island (Fig. 19), the galls contained larvae and pupae, with the first adults already emerging. On this occasion, the host plants were about 20 cm high and about 50 cm in diameter and approaching the end of flowering. Lawrencia squamata accounted tor some 10% of the ground covering of the dense, herbal, coastal vegetation at this locality. The population density of the new gall midge was high, comprising up to 10 galls per host plant. Many galls were found cut open, possibly by birds, a phenomenon described for other cecidomyiid galls previously (Struble & Osgood 1976; Tscharntke 1990), Id P KOLESTE Pie 18. Tindtidest Iskind Soul Australia — aerih voew at (he pypie loealiny nf Réepeedanein Javed spe iia | (iimdinarsly feline 2, Sty Riehid Poriisuta. 3 Younstiistend Pernaali White aerawe tates (he type hieality Cre toile disnledion Calls or We pew wall unide Were teumel on huicreneta aynrneta phils collected from the tollawiay lnoiities on South Austratias SW ol Marte jr SYS lav SU BY wack to Fisher [abe 204 Pa Sb 6S ke Wool’ Nullarbor [3b 27S. bane hb a0 hm Sor byre Highway [sl 47'S, tale 82 Pit km Soot Voomibre [AVP Ses, Pate ES by, Veet Sinelaie [32 7S. ate ST ey. Thevenand (S27 YS, [AV WU del sineky Busy Jar? 1 So 14s" Pe], Vale Pies Dice |evEe bat 1av° 4 El, Nit luce Stalin (22° 2358, (35° Heh Camere [42° 2655, PAM? 32 Fay, Fue Psheret |92" 28S, 138 TY by]. Pee Istund AD IY’ SS. 1S" AE] bs kin No iliscn (vw Ww 19S ay be. St brivets stand ae 3148 PAR TR EL). Mastin bela | $2" 40S. 144s 7 Eb] Teal Ian pat? sao Sas 17 RL Redein Suevey are pat 4S Lee al UY Strike Bay ja de Ss. (oe SE] 6 kin Et Cyneens [9° 43/ Hh. Ir UE Vellistens [33 GMS, Pade SA) bf, Cywell (S5° 45S, 136° 55 be), Buea Creek Plain elas kit NS of Mt Mary bh] 34° S95 A. 18? 26], AW side eal Lake Lhamntin [93° 97S. 138° 8 leh. Port Hoel JS42 05S. 1578 38 De), Mit Mlory Pade i Sade on i), Acetate (liter Llarbour |34° 48 KR ) AR 20) FY] Weel Cupe (iynes National Burk) [38° 1S. 1 3e° 50 1) Minkilon (rt! das, 147° Ae), Pom Linch jae 44S. LAS S211, Porrens Isham La4° 44° 5, ISK" 32 EY Part Adelaide (24° S05 8, Fee SE], Bort Noamlinga [Sa 098° S. Lage 28" A]. Pondaliwie Vay [AS ES. 146 SOUT. Pinditmeste Istana [3 Th S, Lane S07 |) Malinois [3° 96S. 139° 40 by. IX kin N of Meningiv [35° 37° S. 1aHe 2h ty Vennavhar Porat (Rangsran isto |a5° 33S. ban 4? E|, Coovong [40° Js S, LAV 43 TE] The highest qibuevlamee al calls wep Camigh ou plats wolleetwal Ta PY fy NooM Wace urn tbe exposed roeky Head hind of Dog blued whine plans Were subject to salt spray duping stariis Che phans From (his area whicll ve depasited if SHSA aye pall, dense shrubs. whoo! (OOo figh gid E50 nein diameres, eaelt bering seme LOU cally at tlie new sll inntee Oher loealions wilh fel uhundunee pl ahs were The Coorany ephints collected tos 1 8A! by ME. aR, Sti) yd Added iter Harb (plants collected Tah PT by A. Gh Spronenh Renrirks BNO TS WE GT Getty SSE SEE EES Species farming walls an plants af ihe Lanny Asteraceuc. J lene ane Pear alist ete Hypa icons ul sriips fin ie eens, one Conlinis species (list Maye hima with spatula tbseat ind pape will) alent hones presenp pie orher ennniins spee wes thar five eve WOT Spatula Brose Tih crid) puijoue: Watlh sate nital Horiis shscnt (Gane FOOD) Dre nw species belts te tips faeiier group The enly ather kien Australian atte Afapaferayig, gente rites Rolesik (1990), a species defurtinie stone at Cromena Tame J. Black (Quodenimeeney in the lathe Fayre region. beloms te tle latter sedup Boil spevies belongs 1 Syivews (1975) biulegical eroup V of promaey eall prdueers will lurvue eed A NEW GALL MIDGE FROM LAWRENCTA SOQUAMATA 145 solitarily und pupation taking place in the gall) The gall of Ro goodenide comprises & conglomerate of individual chambers whereas the gall of R. luwrenciae sp. nov. consists of a single chamber. Because only these two species of Rhopalomyia are known to be native to Australia, it is loo early for a general characterisation of the genus on this continent. Below. a key is given to adults of the two nalive species and R. californica Felt, an American species introduced into Australia to control Bacchari. 8 helimifolia L, (Asteraceae) (McFadyen er al.': Gagné & Boldt 1995), Key to adults of Australian species of Rhopalomyta 1. Tarsal claws toothed... R. lawrenciae Tarsal cliuws simple 2. Palpus with 3 or 4 segments: length of papillae on male parameres !/s - '/> paramere Width... wR, goodeniae Palpus with | or 2 seements; length of papillae on male parameres about '/2 paramere width.......... -R californica Acknowledgments W. R. Barker and M. C. O'Leary. both of the State Herbarium of South Australia Adelaide, courteously identified the host plant species and assisted in examination of dried host plant specimens, respectively. D. Eastburn, Murray-Darling Basin Commission, kindly gave permission to print the photagraph in Figure 19 1 thank J. D. Gray, Department of Horticulture, Viticulture and Oenvlogy University of Adelaide. and R. J. Gagne, Systematic Entomology Laboratory USDA Washington DC, for commenting on an early draft of the manuscript. References CivGnr. Kd. (1904) * The Gall Midges of the Neotropical Region” (Cornell University Press. Ithaca, New York). — & Bonn, PE. (1995) The gall midges (Diptera: ~ Cecidomyiidae) of Baccharis spp. (Asteraceae) in the United States, Pree, Aur, Sac. West, 97, 767-778. Jessor J.P. (1986) Family Malvaceae pp. 821-848 In Jessop. J Po & Toelken, Ho R. (Eds) “Flora of South Austratia,” Part 2 (South Australian Government Printing Diyision, Adelaide), KOBPAER IJ 1896) Newe Mitthedungen iiber Gallmiicken, Wiew. bat. Zi@ VS. 85-105, Komrsin. P(1995) A new species of Evctiectioarnia Pelt (Dipteras Cecidomyiidae) on Eyecafypiis faseiculasa in South Australia, /. Aus ent See. 34, 147-152, (1996) Rhopulomvie geudeniin, a new species of Ceeidomylidae (Diptera) damaging Ceonenin linata (Goodlentaeeae) in inland Australia. Trans. Ro Soe. 8. Vass. 120, (SS-160, PMePAb VES. Poh. Danses, Ch Bod “Tompnas AL J (P9RSy Bielujieal control ab grommet bush: pp. 2a 40 fn Harvey, GI (edo) “Australian Weeds KReseurch Newslenor' (The Ala Ploncher Research Sein. TSCHARNTEKE, TT. (1997) Two new species of Asphondylia (Diptera: ¢ ‘ecidomyiidae) from Halosarcia spp. (Chenopodiaceae } in South Australia. (id. 121. 59-66, RUBSAAMEN, BEL 41892) Die Gallmiicken des Kanighchen Museums fiir Naturkunde zu. Berlin, Berl, Ent 237. 319-411. pls VIEXVUEL, Stkiiser, BD. B. & Oscoon, F. A. (1976) Predation on larvae ot the balsam wall undee, Desinewra bulsanicala (Diptera: Cecidomyiidae), within galls in Maine. Ceneel, Ent. 18, (443-1444. SYEVEN, bk. (1975) Study on relabonships between hibits and external structures in Oligotrophidi larvae ( Dipterin: Cecidomyiidae). Zvel. Scripta 4, 55-92. (1990) Vogellruss heemtrachtigt dic Dichteregulanion einer Gallmiickenpopulation dureh Parasitoide: Weeliselwirkung ewischen vier trophischen hbenen. Min, Deutsch Ges. Alle. Anwew. Biri. 7 552- 544. DASINEURA WAHLENBERGIAE, A NEW SPECIES OF GALL MIDGE (DIPTERA: CECIDOMYIIDAE) DAMAGING SHOOT TIPS OF WAHLENBERGIA STRICTA (CAMPANULACEAE) IN SOUTH AUSTRALIA By PETER KOLESIK* Summary Kolesik, P. (1998) Dasineura wahlenbergiae, a new species of gall midge (Diptera: Cecidomylidae) damaging shoot tips of Wahlenbergia stricta (Campanulaceae) in South Australia. Trans. R. Soc. S. Aust. 122(4), 147-151, 30 November, 1998. A new South Australian gall midge, Dasineura wahlenbergiae, that damages shoot tips of Wahlenbergia stricta (R.Br.) Sweet, a common plant of grassy habitats in Australian and New Zealand, is described. Two leaves of the shoot tip of the host plant are malformed into a globular, hollow, hairy, partially discoloured gall, 2-5 mm in diameter. The male, female, pupa and larva of the new species are described. The new gall midge is the fourth Dasineura species known from Australia. Key Words: Gall midge, Cecidomyiidae, Dasineura wahlenbergiae sp. nov., Wahlenbergia stricta, South Australia. Transactions of tre Reval Society of §. Aust. (1998), 122(4), 147-184. DASINEURA WAHLENBERGIAE, A NEW SPECIES OF GALL MIDGE (DIPTERA; CECIDOMYHDAE) DAMAGING SHOOT TIPS OF WAHLENBERGIA STRICTA (CAMPANULACEAE) IN SOUTH AUSTRALIA by PETER KOLESIK* Summary KOLUSIE, P1998) Masitenia willenbergiae. w new species of gall midge (Diptera: Cecidomyiidac) danagine shoot lips of Waitenbergia s(ricta (Campanulaceae) in South Australia. Trans, Ry See 8. Auyt, 12204), 47 151. 30 November, 1998, Anew South Australian gall midge. Dayinenta wohlenbervice. that damages sboot fips ob Walilenbergia sirickt (R, Bry Sweet. common plant of grassy habitats in Australia und New Zealand, is described. Two leaves of the shoot tip of the host plant are malformed into a globular, hollow, hairy, partially discoloured gall, 2-S min in diameter The male, female. pupa and larva of the new species are desenbed. The new wall midge is the fourth Dasineura species known (rom Australia, Key Words: Gall midge, Ceewlamyiidue. Dasinenra wahlenbergiae sp. nov... Wehlenbergia siricni. South Australia, Introduction The new gall midge deseribed here was found it malformed shoot fips of the tall blue bell. Wahlenbereia stricta (Ro Br) Sweet (Campan- uluecae) at) Moriah Cooservation Park, fear Adelaide, Wahlenbergia stricta is a perennial herb, (00-9000 nim high with large, blhe flowers and is common al grassy sites in various yegetation types throughout Australia and New Zealand ¢Siith 1986). The plants grow on slopes at the Morialta Conservation Park and in the spring the shoot buds of many of them are moditied ime globular, tary gills. Some plants have all their shoot tips galled and consequently do nor reproduee, Materials and Methods Shoot tip galls on Wallenberaia strict were collected at Moriali Conservation Pairk on 1S September, 1996 ind brought to the laboratory where a few of the galls were pecled open and the developmental stages Of the gall inducer examined. Same of the galls contained young larvae, some malure larvae, some cocoons and others were emply The cocoons contaned either larvae or pupae. A small number of the mature larvae wits preserved in 70% ethanol A few eocoons were torn open and the larvae and pupae preserved as above. The majority of the galls was laid on wet sand within a pot to allow ' Deprrtmeat of Hortealiove, Vitieultne ain Oenology. Waite Cuyats, The University of Adelaide PMB | Glen Osmond S, Aust. 5064 them to develop into adults. Pupation took place Within the galls. Emerged adults were preserved in 70% ethanol Canada balsam mounts of the type series were prepured for microscopic examination according lo the technique outlined by Kolesik (1995), Measurements refer to the holotype and paratypes. The type specimens. and other material retained in ethanol, are deposited in the South Australian Museurn, Adelaide [SAMA], the Australian National Insect Collection. Canberra JANIC| and the Swedish Museum of Natural History [SMNH]. Dry samples af the galls are deposited inthe State Herbarium ef South Australia, Adelaide |SHSA], Genus Dasineura Rondant, US40 Dasineura Rondant, P40; 12 & 17 Proposed type species Vipile aisverbrii: Schrank, 1803) Gagné eral. (1997) Dasineura is a large, cosmopoliti Yenus UF some 2) species containing Oligetraphini with four scemented pilpi, toothed tarsal claws, an Ry wing vein that meets C oanteriar to the wing apex, and the female ciehth tergite divided (hte two longitudinal sclerites, Dasinenra wahlenbergiae sp. nov. (FIGS )-15) Holawpe: 3, Morialta Conservation Park, South Australia [34° 54° 8. 138° 44° BJ, 20.14.1996. P. Kolesik, reared from a shoottip gall of Wallenbergia aricia (R, Br.) Sweet collected 15,i,1996, 121384 |SAMA],. 148 P, KOLESIK Figs 1-6. Male of Dasineura wahlenbergiae sp. nov. |. Head in frontal view. 2. Last three flagellomeres. 3. Wing. 4. Last tarsal segment with claw and empodium, 5, Sixth flagellomere, 6, Genitalia in dorsal view. Scale bars = 100 um 1, 6; 50 hm 2.4.5: 500 um 3. ANEW GALL MIDGE PROM WAMLENRERGIA STRICTA du Parniypes: 39 2.3 pupae [SANA, 121385-12 1390), PP 2 7 2 pupae fall ANIC] same dina but emerved 17-25.i8, 19962 3 larvae |SAMAY, 3 larvae [ANIC], collected with holotype, Omer materials 34 5 [SMNUL, same dati as holotype bul enierged 20,-25ax,1996: 37 Taryae. 5 pupae within cocaons [SMNI|. eollected with Holotype: gall [SUISA, AL99747 199). collected with holotype. Deseripron Male (Bigs 1-04 Coluur eyes bhicks bead. thorax und) abdomen orlinesred: leas, auitennie, pilpi setae and settles ufey. Walleles oralige brown, Hew; Antero seape square in frontal view; pedicel sphernide 10 fiwetlomeres. Crstund second fused, nevks as long or slightly longer than podes: CHCHMTKG coliprisnig Uwo Uupsyerse yn bwe Jongitudioal bands, Palpus four-seamented, segrrents progressively longer, Bye fieers rounded. eluse together excep on veriex where small area ol ne Hicets separtites the eyes, Labelle tapered clistally, Kilenlly wall © setae, Proms wath) 23-26 serie per stile. Thorax: Wing length 2.) mim (2.022. n = 2), wWidih OO inn (8-91 Rs joining C anteriorly to upers Ri jouing C slightly anteriorly to mid-leneth; Ks not obvious. Claws (nothed, emporia as lone as chiws, Abdomen: Tergites |S with pair of sensary sete ih ablerlor copners, lereites 1-7 With stile setal poy posteriorly and seules scutlered evenly, fergile & ta form of mimrow. selenatised, anterior bund wathoud seme. Stermites 2-8 with pair of sensory sete anteriorly, setie i Wide band anterierly ated narrower band posteriorly, area between Lwo bands of setae mere weokly selerorised. Genitalia: gonueoxile evlindrival. setose and setulose; BOHOSEVIUS Lupered distally, sparsely setose, Setulose basally up te “/) oF ats Tengu ventrally and '/> dorsally, sparsely striate beyond, bearing distal comb, cere) large. cach with several selue uipieally, selulose: hypoproct deeply and widely divided, with one sel on each Jobe, selulose: parameres sheathing vcdeagus. with subylobulit disteusions dorso- basally. with --S selose papillae upiealhy; aedeagus long, stout, Female (vise 7-10) Colours ais in nile. Heid: 16 (lagellomeres, evlindrical, with necks /\- yy node's leowth, creumbilte comprising iwe transverse and lwo longitudinal bands. distal Irinsvere band with loop, cireumililin attachment points very dense. Labella with 7-10 setae laterally. troos with 22-28 sete hiterally. Thorax: Wing length 2.) mm (2.42.3, p = 5), Wilt O24 nin (O8-O.9), Abdomen; Tergiles 1-8 with pair of seasory setae inanterior corners, tergiles 1-7 with single setal row posteriorly and seales seatlered evenly, tergite 8 ciyicled THto two donaitudinal selertes, Stermites 2-7 wilh pair Of sensory selae anteriorly, setae im wide hand anteriorly and narrower band posteriorly, urea between Iwo bunds of selue more wenkly selerotisud. sternite Snot developed. Qvipasitas protractiles elongate, O.7 mm (0.0-0.7) long (anterior Linnil of genial chamber to werminal tip distance), 31% (29- 45) of wing length: cerer fused medially into single. prolonged, terminal lamella, setose und setulose: AV poproet WHA Wo setae, setalose. Prpe (Pig, 1) Colour antennal horns brawn at upex. remaining puts yellow, Length 2.00 mit CLS-200 nm = 5), Antenml horns small, pointed, Frons on each side: three frontal papillae two of thent setose. asetose one someumes ducking) three usctose lateral facial papilla, Cephalic papilla with seta 94 pin (LS8Y 201) Jong. Prothorucre spiracle 230) pint (220-244) ling, trachea ending at apex. Tnresument af abdomind segments covered with spiculac shelly lonver dorsally, second through seventh abdominal seenicutls with group of dorsal spines ad anterior hall) First through eighth abdominal segments with wwe pairs oF dorsal asetosxe papillae, one pair ol Setose pleural papillae, Gve pairs af asetose ventral papillae, Last destar larva (Figs 12-141 Colour: red. Lenwth 24 gin (2028 w= 6) integument covered with rounded phites about LO pin ti dhameter, veninilly wth several transverse rows of spiculae on anterior fall af thorieie ane abdominal semments, Plead swith postero-hiterul apodemes as Jon as hed lenguh. Spatula bilobed, with long shat, longth $47 jn (177-169). Papitlae charuererisue of Dasineura laksa (syWwen L975), Livinology The name wellenbersiae is derived trom the genene mome vl the host plait. Gall and hiolaen The new gall Widwe modifies bwo deaves ob the shoot bp at Wehlenbergia stricto into a globular, hollow, hairy, partially discoloured gall, 2-5 main diameter (Fig. 15). On (5 September, 1997, at Morita Conservation Park most galls contained mature lurve. but some gulls contined young lurvue, Some Cocouns With larvae or pupae within or 150 P. KOLESIK ‘ ; egy ~ a Figs 7-15. Dasineura wahlenbergiae sp. nov.: 7-10 female, | 1 pupa, 12-14 larva, 15 infestation symptoms. 7. Posterior end of abdomen in dorsal view. 8. Posterior end of ovipositor in ventral view. 9. Sixth flagellomere. 10. Last three flagellomeres. 11, Anterior part in ventral view. 12. Spatula with adjacent papillae. 13. Head. 14. Two terminal segments in dorsal view. 15. Gall on Wahlenbergia stricta (R. Br.) Sweet. Scale bars = 100 um 7, 11, 14; 50 um 8-10, 12, 13; 10 mm 15. A NEW GALL MIDGE FROM WAARLENKERGIA STRICTA IS] empty cocoons, und others Contained no remnants of the gall inducer, Up to 20 larvae were found within a gall. The adults reared in this study originated from larvae pupated within the galls. Discussion Dasinetiva, the largest genus of Cecidomyiidae, comprises species oecurring in all zoogeographical regions of the world, Four species are known from Australias DL aedeiaelongifoliae (Skuse, 1890) (Gagné & Marohasy 1993) and 2D. diedyé Riibsaamen (1916) which damage Mowers of Acacia longifolia (Andr.) Wild. (Mimosaceae) and A. evelopy Cun, ex Dou tespeetively, D, divbaath? Kolesik & Skuhravit (1997) which ts an inquiline in Mower galls on Hybanrhus floribundus (Lindley) Muell, (Violaceae) Wduved by an unknown gall midge. and the new species described here. Dasineura wallenbergiae sp. nov. belongs to Sylyen’s (1975) biological group EH of gall midges whose larvae are primary yall inducers, feed gregariously abd pupate in both the soib and the plant. ‘The adults of the new species reured if) the present study originated from larvae that pupated within galls, bul the fact that some galls were found empty with neither cocoon remnants nor parasitoids within. suggests that part of the turval population pupates in the soil. This conforms with the behaviour of Sylven’s (1975) biological group LI, Dasiniera diybanthi. the only other Australian species of this genus described in detail. belones to gruup LL of gall midges Whose Jarvae are inquilines. feed gregariously and pupate in the soil. The new species differs ftom D. liyhanthi in several morphological characters. In D, wahlenbergiae, the wing vein Rs is not obvious, the tooth on the tarsal claw is much smaller than the claw, the [emale Hlagellomeres are much longer than wide, in the male genitalia the gonostylus is tapered distally, Ue mitle cerci and parameres are nearly as long us lhe aedeagus, and the larva has no medial papillae between the terminal papillae. In D. Ayhanrhi, the Rs is evident, the tooth on the tarsal claw is nearly as larve as the claw, the female flagellomeres are as long as Wide. inthe male genitalia the gonostylus is abou! the same width through its entire length, the Male cerei and parameres are much shorter than the wedeagus. and the larva has several medial selose papillae between the terminal papillie, Acknowledgments ] am grateful to H.R. Toelken. South Australian Stite Herbarium for the identbication of Wahlenhersia sricta A. Stark, Halle Germany courteously provided a copy of Ribsadme's paper, Special thanks go lo J. D, Gray, Department of Horticulture. Viticulture and Oenology University of Adelaide, R. J. Gagné, Systemuue Entomology Laboratory USDA Washington DC, and E, Sylveén, Swedish Museum of Natural History Stoekholin for commenting onan early draft of the manuscript References GAAGNE, 1, J, HARRIS, KL ML SKEET A, ML. Satins. M amt Svives, Be (L987) Daamenra Rondam, [840 (lise. Dipreni: proposed designanon oat lipali yviibel Solmaink, 1802 as the ype species. (Case 2986, Ball. Zoal, Nein, 54, 92-44 & MAROUASY 1. (1993) The gall Widges (Diptera: Cecidomyiidie) of Acccig spp. (Miimasacene) a Kenya. lnseota Mundt 7, 77-| 24, Ko esik, BCP9oS) A new species ol Locinetieorme Fell (Diptera: Cecidomydidue) on Lucedypray fusetenfesa in South Australian. A Aus. enn Soe. 34, 147-152. & Skuneava, M. (1997) Dasmenra lybanthi spec, nov. wo Hew inquiline species of Ceeidemytidac (Diptera) from galls on Hybanthas flaribundas (Violieeae) in Australia Sradi Dept 4. 240-246, RISAAMEN, le Lh (M916) Bentrae zur Renniniis ausserecuropitiseher Gallmiicken, Sifsaiueberichte ir Gesellachaft Natieforsehender Ereninde oi Bertin W915. 43)-48 1. SKUSE, BAL AL (TRY0) Diptera of Australia Neratacens. Supplement 1, Pree, Linn, See, NSW (2nd series) F. 97 A- 4l3. Svrvi, Bot (M86) Pamily Campunukivaie pp. tA76-| 443 Mt Jessop. d. Po & Toelken. H.R. (Rds) Plor of South Australia, Part 1 (Polemoniiceae - Compositie)” (South Australian Government Priiting Division, Adelaide), Syivin. E. (1975) Study on relationships berween habits und externyl straciires in Olivotrophidi luryae (Diptera, Cecidomyiidae). Zoel. Seripta 4, 55-02, DEVELOPMENTAL BIOLOGY OF UPEROLEIA TALPA TYLER, DAVIES & MARTIN, 1981 (ANURA: MYOBATRACHIDAE) By MARGARET DAVIES* & GRAEME F. WATSONT Summary Davies, M. & Watson, G. F. (1998) Developmental biology of Uperoleia talpa Tyler, Davies & Martin, 1981 (Anura: Myobatrachidae). Trans. R. Soc. S. Aust. 122(4), 153-157, 30 November, 1998. Uperoleia talpa is a small fossorial frog restricted to the southwestern portion of the Kimberley Division of Western Australia. The frog breeds in the monsoonal wet season, and lays clumps of eggs in single capsules in ephemeral ponds. Larvae hatch at stage 19. Later-stage larvae have strongly arched tail fins, a sinistral spiracle, extremely large, cavernous external nares and a larval tooth row formula of two upper and three lower rows of labial teeth. Labial papillae are clearly interrupted both anteriorly and posteriorly. Larval life span is about 71 days. The large and conspicuous external nares have been found in a further five species of Uperoleia and are suggested as a possible diagnostic character for some larvae of the genus. Key Words: Uperoleia talpa, larvae, embryos, generic character, life history, tadpole, Myobatrachidae. Tramaetions a the Raval Secien of S Yast (1998), 122(4). 153-157. DEVELOPMENTAL BIOLOGY OF UPEROLEIA TALPA TYLER, DAVIES & MARTIN, 1981 (ANURA:MYOBATRACHIDAE) by MARGARET DAVIES & GRAEME F, WATSON’ Summary Davirs. M. & Warsus. GF (1998) Developmental thology of Uperuleia talna Tyler, Davies & Marrin, 1981 CAnira:Myobatrachidie). Jeans A. See. S. Aust $2204). 153-157. 30 November. [908. Uperoleia talpa iss small fossarial frog restficted to the southwestern portion of the Kimberley Division of Western Australia. The [row breeds in the monsoundl wel season, and lays clinips ofeges Ti sige capsules in ephemeral ponds, Larvae hath arstige 19, Literstage larvae have strongly arched tail fins. a sinistral spirale, extremely large. cavernous external nares umd a larval loath row formule of two upper und three Inwer rows of lubial teeth, Labial papillie are clearly interrupted both anteriorly and posteriorly Larval life span is about 71 days. The lacie and comspicious external nares have been found ina further five species of Uperalear sind are suggested asa possible diagnostic charter for some binvae of (he genus, Key Worns: Uperoleia telpa. larvae. embryos, generte chacaeter, life listory. tadpole, Myohatrachidae Introduction Uperalen Gray. L841 is a genus of small bur rowing, myobulrachine frows with a wide-ranging distribution deress Austealia in areas Of poor winter rainfall, Prior la the revision of Tyler efal (1st), the genus comprised three species. bul with the description ol Uperieia altivsine (Davies er ul. 1993), now ineludes 24 taxa. However, very Tile is know of the tarvad biology at the genus Moore (1961) deseribedt the larva ol U) merrvrereter (how eonsidercd ta bet! deevigar Keterstein, 1467 (Davies & Liulejohn 1986)) whilst Watson & Martin (1973) described the larva of what was thought to be UU. mermorala. bul whieh is now considered to be a representative Of LL tyler? (Davies & Lillleyohn }O86). Tyler er el. (1983) recorded the lHfe history of Hh jaundape Vyler, Davies & Martin, 1981) Davies e al, (1986) described the larva of 00 dittomoda Tyler, Davies & Martin, 198T and Richards & Alford (1993) provided a description of (he larva of U. mintila Davies, McDonald, Corben & Ingram, LO86, Pull life bistory data of Lhese species are scarce, Uperoleia tipa ‘Tyler. Davies & Martin, I98T is a large member of the venus (males 26-40 mim 5-V, females 35-38 qn) CPyler en al 1994), with a restricied distribution in the southwestern portion of ihe Kimberley Division of Western Australia. The species Was originally described from three [rogs colleeted ona very dry night south of Derby (Tyler ey dh 198)) and the description has been amplified by ) Peptol Agology. Liliversity ob Adelaide Astle 30Gb). ( Deplol Zamlony, University of Melbourie Parkville Vie. 052, Davies & Martin (L088), who provided addigonal morphological, ostevlowieal aid distributional in- formation and described the call, In early Pebrusey 1994 we collected ampleetant pairs of Lh talpe the spawn of which Wwe reared to metumnorphosis. thus allowing the description af the Tle history of the species (hat we report here, We also discuss sone features that may aid in generic recognition of larvae, Materials and Methods The sevies of Uperoleta talpa was reared. trom spawn deposited in phistc bags by ampleetant pairs collected in the field, Larvae were initially reared tn dwrated water at ambient temperature in the field hefore being transported to Adelaide where they were mulled i) a COnstail temperature room at 30 41° C in dechlorinuted tip water, Larvae were fed on boiled organie lettuce leaves supplemented with commercial goldlish Wakes (Bioser), Material was preserved in Tyler’s Hui (Tyler 1942) and Hlustrarions were mude with the aid of a Wild M8 sterea dissecting microscope with attached camer lucida, Mewsurements were mie using ai eyepiece micrometer Developmental stages are those of Gosner (1960), Material eanmined: Davies collection: Uperoleia talpa series: U. lithomiuda serws, U, allixsime series; U. dnundata series (basis of data used hy Tyler er al. 1983); larvae of Crinta (Rentdella) scanifera (Girard. 1853), 6. (Ro) riparia (aitleyohn & Martin, 1965). Peendophiyie VPitvinger, 1843. Tadpoles of &. luevivala were provided by Harold Phmann and | 54 M. DAVIES & CF WATSON (poles of (/ russedie (Loyeridge. 1933) were cx- amined i the collection of the University ol Michigan, Museum of Zoology, Ann Arbor, Michigan Results Pwo amipleenint pairs of Uperaleia talpua were enilecied al 2345 hoon 3.11, 1994 at a site (2.2 km south af the Gibtr River Rd tumoll on the roud south of Derby in the Kimberley Division of Western Australia (Figo 1). The night wits extremely hol, humid and stormy althouzh na rai fell in the Inmmednie area. Two other species of Uperalera (Ui mberad (Andersson, 1913) and 4 aspera Tyler Duyies & Martin, ISS) were calling at the same pom Upereleta dap was calling trom the Ury yeuetation tanhest trom the water, Uo sijeberus was calling from the edee af the water and Oo aaperie was calling From the intermediate areas, The choruses ol Oo defpecanel Uh apera were substantial whilst thar OF LL nyiherey was less viworous. Loarta rubella (rry. TAZ) und c velarene cistraliy (Gray. 842) were also culling around the pond. When we visited the same site the following evening. there was much less aeHivity With a single) feldme and cery tew L aspera calling, Avila time, tewly mecinorphiised in C. arvptotiy Tyler & Martin, 1977, CL loauipes Tyler & Martin, L977 and Cy australis Were located, The captive pars were retained in pond water in inflated plastic bags, supparted by feeereum containers. The U. Kepe spawned vurly on 6.111994, AL O700 on 71,1994, the eges hid reached fate scustrulit slige b2. A single capsule surrounded the oyun Mean capsule diameter of six egies wits LS mm (range 1.78-2.04) and the ova had a mean diameter of 138 mim (ring 1.30-1.60 mim), At 1300 on 71,1994 embryos were at stave 17 (tail bud) (Pig. 2) with the tail being better developed thar the Head. Hatehing (stage 19) was completed by TITS an Ai 1994 (Mig. 3). The newly hatehed birwae fad vo eaterial gills: (he eyes were very dillicull to defect ind (he mouth had not perforated: adhesive glands Were HOF pigmented ul this stage. By HOS on 11.1, 1994, same preserved lirvac webe already at Stage 25, The spiracte hud formed and the adhesive whinds were pigmented, There was no keraunmsation of the beak ar teeth (Wis. 4). Mulerial preserved on [241 1994 inelided some Iirvie stil at Stage 22/23 (hig. 3) in which the nostrils hil not perforated althouph the adhesive ghinds were pigmented, The spiracle hud oor toriiea aU ulthouwh the mouth was perlorated there was no kervinisation on (he beak or teeth Pin. bo Site abo wlttct ampleetan pairs OF Moerutera tafpa were collected. Se kit S Guth Bier Relient on Piehiway t eH al Pder liv, WA DEVELOPMENTAL BIOLOGY OF UPEROLEIA TALPA 155 _ Fig. 2. A. Lateral and B. Dorsal views of Stage 17 (tail bud) embryo of Uperoleia talpa at 2200, Scale bar = | mm. Fig. 4. A. Ventral and B. Lateral views of Stage 25 larva of Uperoleia talpa. Scale bar = | mm. Sal eal Fig. 5. Lateral view of Stage 25 larva of Uperoleia talpa. Scale bar = | mm. Fig, 3. A. Lateral view of newly hatched larva at Stage 19. B. Lateral view of larva at Stage 22 of Uperoleia talpa. Scale bar = 1 mm. Tarte l. Measurement (in min) ef body and total length of larvae of Uperoleia talpa as mean and range. N = number of individuals. Stage (Gosner 1960) Body length (mm) Total length (mm) N 11,-20.11,1994 25 4.19 9.66 5 (3.84-4.64) (7.84-8.16) 14.11.1994 26 3 8.0 4 (3.2 - 3.6) (7.84-8,16) 26.11.1996 27 5.44 12.8 - 13.8 2 2.-29. 11,1994 28 5.46 13.72 7 (4.8-5.92) (12.96-14.58) 6.-29.i11.1994 29 6.46 15.33 5 (5.44-8.48) (14.24-17.44) 29.i11,-13.iy, 1994 30 6,99 16,69 3 (6.08-7.52) (15.36-17.6) [3.iv. 1904 3] 6.08 16.00 | I3.iv, 1994 32 7.84-8.64 18.56-21.8 2 13.iv. 1994 33 9.28 21.92 i] 13.iv. 1994 34 8.72 21.8 4 (8,0-9,28) (20.32-22.72) [3.v.1994 35 9.28-10.4 23,2-26.4 1 13.iv.1994 36 10.72 26,08 3 (10,24-11.2 (24,8-27.2) I3.iv. L994 37 9.92-11,2 25.28-26.58 2 U3.iv, 1994 38. 10.8 25.12 i 156 M. DAVIES & G. EF WATSON Fig 6, A, Lateral and B. Dorsal views of Stage 36 Larva of diperetedt tulpu, Seale bar = 5 mm. Fig. 7, Oral disc of Stage 46 larva of Upeviteta talpe. Seale bar= | mn. Larvae al stave 25 preserved at 1115 an 14.11.1994 (Fig. 5) had perforated nostrils, which were round. but these were not as conspicuous us is seen at later Stages (see below), The horny beak was keratinised aod kerauinisation of one upper and 2-3 lower rows of labial teeth was beginning. Yolk sull remained in the ut. although the cloaca was open dextrally, Larvae remained at stage 25 until after 20.0).1994, period of up to 9 days since the onset of this stave. Measurements of larvae at stages 25-38 of Gosner are viven in Table 1 The following description is of a larvaat Stage 36 (Pig. 0), Body ovoid, widest at midpoint of body. Snout evenly rounded i dorsiul and lateral view. Nares dorsal and extremely large and cavernous. Eyes moderately conspicuous, Spirucle sinistral, short, opening dorsally and visible when viewed from above. Anal tube broad and dextral to ventral fn, Dorsal fin more strongly arched than ventral fin. Pins rounded terminally. Dorsal fin commencing on posterior part of body, deepest about half way along its length, Veotral fir commencing posteriorly to body. approximately same width along its length, Tail musculature moderately thick, tapering to fine point. Oral dise small and ventral. Labial papillae widely interrupted anteromedially. Also. interrupted DEVELOPMENTAL BIOLOGY OF LIPEROLEIA TALFA 1s7 posteromedially. Two rows of upper leeth and three rows Of lower. the second of which is divided (Pig, 7). Short P3 row supported on flexible flap, ‘Tail musculature and fins moderately heavily suffused with pigment. Small dark-brown islands of pigmentation un the body. Larvae reached metamorphosis at stage 46 on I5.iy, 1994, 71 days aller spawning, Discussion We have now examined tadpoles of six species of Upervleia, as wells several olber myobulrachine species, [eis clear that the external pares of muny Uperoleia larvac are unusuilly lire and cavernous (Vig. 6). OF the species examined, this feature was present mi all but (. imurdare, Richards & Alford (1993) provided measurements of €. minuda, but these do not allow ialiree: comparison of the data we provide here, since diuneter in relation to the width of the head could not be ascertained, These authors do not conmiment on the relative size of the nares. H, however, the nares ure nol particularly large, this feature would be useful in separating larvae of C) iimile (rom ff fithomoda - a species: pair in whieh the adults are difficult ta Sepirate beth mor- pholowieally, and, a high temperature. by call OK. R. McDonald pers. comm. 1986) There is no generic tooth row formula for Uperoleda, Uperaleta minuta, U. lithomoda and tnundata share a formula of 2¢2)/3 whilst U/ laevigata has a formula including no undivided rows of 3. Uperoleia talpa has 2(2)/3(2). whilst that of (!. altissima is 2(2)3(1.2) (Davies & McDonald 1998). The dark tail tip recorded by Richards & Alford (1993) in early stage larvae, whilst shared by UL mimula, UL lithonioda UL laevigata and UU. ariysima (Davies & McDonald 1998), is not present in. talpa, The heavy pigmentation of O. talpa larvae is Shared by O/ tvleri (Watson & Martin 1972), The flexible flap supporting P3 labial teeth is recorded inal Uperafeia lo date, but is not unique Lo Uperoleia, being found in Criniat (Reanidelle) sienifera, C. (Ro) riparia and as a larger structure in Pseudophryne species (Watson & Martin 1973; Davies unpub.). Thus the possibility of using thes leature for generic recognition foreshadowed by Richards & Alford (1993), cannot be sustaited, The large nares may he uselul in some species assemblages, being absent in only OU. (auadera iimony the species wwe have examined to date, The unusual tail bud stage in whieh the tail is better developed than the head was noted alsa by Moore (M61) in U4. feevigata. This developmental condition merits further investigation Acknowledgments Fieldwork was supported by the Anstralian Research Committee and we thank Mo. Tyler foe companionship in the field and the referees tor helpful comments Reterences Davins. ME Ae Li tortoise, Med (86) Phe brag sents Uperalda CXnutihepadienytidie: in southeastern Australia, dics A Soe So Mash EM TED ps. _ — oN Maris, “AOA, (PORK) Redetiniian ot Uperoele tala Tyler, Davies & Martio, 188) CAnurie Leptodiery tiie: Myobutraelinae), fe 112, 47-89, & MeDosarb. BK. R. (1908) Develupmental hiology Of Upertein vitissime Davies, Walsor, MeDonild, Lrenerry A Werren. 18a CAN Ura Mycbatinelieliwe. Malek 22, 167 172, & CORBIN, ©. (IYsO) The rents Hiperafaia — Ciray CAnFE Leploduery Hetiey i Queensland, Australia, Hae, R, Soe, Wek 98. 147 E88, Wastin, Ch B MIC DOs anne Ke Ro TRPNs ay. M ode WiRRE SS, CE 1 1L9UR) A new apecies of Myrreralola (Anura: Leptodjetylidae: Myobatrachiniey tren fortheastern Australia, “tem, Old Atay, 33. 167 U4. Goss, KROL. C00) A simplified table for staging anaes embryos and Larvae Wil totes on identiteatiarn Herpotalowion Ube (R20) Mouikh. AS UIYOD The Togs OF eastern New South Wales Kull, Ain, Mus. Nat. Hive, (20, 140-386, Rieiiykos, Sb AR ALO, RAL (1995) The tadpoles ul twe Queenshind frees GN: Pylithie. Myabut rachidne), Mea Ghi Wis. 33. 337 340. Ty MA 1962) On the preservation atanurun Gidpales, Nest, J Set 25, 222, eCROOK. CAL ADAVIES. ML CIOS a) Reprodietiye bidlvey of the frags of the Maeele Creek Systeni Northern Territory, Mec S. Ata Was B.S hi Davis, Mak MARTIN AL ACTOS TE Adstentiar frogs of the leptodaerylid venus Upenvena nay, Vive ot fool Stippl Ser 7 1 O4, Satie bo AL Ae Teminstosr, RO (POU bis ot Western Australia’ Revised edicin (Western Anatraliin Museum. Berth). Warsom, Gob & MARIN A, AV T973) Lite histary, leval morphology and rehitionships of Australian lopradaery ia Hows, Lraas A Saw, S. Aur. OT ON dA, A NEW SPECIES OF FROG (ANURA: MICROHYLIDAE) FROM CAPE MELVILLE, QUEENSLAND By MARGARET DAVIES* & KEITH R. MCDONALDT Summary Davies, M. & McDonald, K. R. (1998) A new species of frog (Anura: Microhylidae) from Cape Melville, Queensland. Trans. R. Soc. S. Aust. 122(4), 159-165, 30 November, 1998. Cophixalus zweifeli sp. nov. is a relatively large member of a genus of microhylid frogs restricted to New Guinea and the Cape York Peninsula of Queensland. The new species is found in boulder fields in the Cape Melville National Park. Females are characterised by having flame-scarlet axillae, groins and hidden parts of the hind limbs. Males have not been observed. The finger discs are expanded. Morphologically the species is allied to C. saxatilis, but unpublished mitochondrial DNA sequences link it with C. infacetus. The description of this taxon brings the number of species of the genus in Australia to 13. Key Words: Cophixalus zweifeli, new species, osteology, Microhylidae, morphology. Transactions of the Royal Sactery of S. Aust. (1998), 122(4), 159-165 A NEW SPECIES OF FROG (ANURA: MICROHYLIDAE) FROM CAPE MELVILLE, QUEENSLAND by MARGARET Davies’ & Kerrd R. MCDONALD! Summary Davies. Mad MeDonNanh, KR t}998) A new apecies of frog (Andra: Microhylidae) from Cupe Melville, Queensland. Tas, A. Soe S. Aut (2204). 159-165, 30 November 1998 Caphivalus caveifedi sp. toy. is wrelatively barge member of a genus of microlylid Trays restricted to New Guinean and the Cape York Peninsule of Queensland The new species is found. in boulder fields in the Cape Melville National Park. Females are characterised by having Clame-scarletaxillie. geoios un hidden parts of the hind fimbs, Males have not heen observed, The finger dises are expanded, Morphologically the species ts allied oC) sexes, bul unpublished mitochondrial DNA sequences lok it with Co dafdeenis. The description of this Jason brings the nuniber of species of Ihe genus in Australia to 13, Kiy Wonos: Cophiveles sveffedi few speaes, ostealogy. Microhytidae, morphology, Introduction Faun surveys have been conducted in Cape York Peninsula by the Queenskind Bavironiment (ind its predecessors) sinee 1975, Information on the vertebrate fauna of the area bas been reviewed by Winter & Lethbridge 1995! us part of Stave }ool the Cape York Peninsula Land Use Stody, Subsequently fauna and fora surveys in Cape Melville National Park have lovaled significant pew records for mammals, reptiles, frogs, earthworms anil verelition types (Stanton 1994+: Little & Hall 1996; Stanton & Fell 1996": Jamieson 197, McDonald 1997, 1998, unpub.) The area of Cape Melville National Park was increased front 36 000 ha lo 137 O00 hit in L995, thus incorporating @ greater diversity of habitats and an increase in the runge uF Hora and fauna in the park, The new area includes assemblages ol topography, geology and vexctation types unique to Cape York (Stanton 1944, so the Cape Melville National Park is can aued of proven and potential codenisnd (Covaeeviel & figean L978: Stanton & Pell }990% danieson 197: Me Donald 1997), © Dephel Zoology. Darversity ab Adetitley Ansisiiia S002 ) OM preecvalion Stiteey BAO. Depa hice ob Terrien iO Bos kad Alherin Olt ass 3, Wires J OW. Tragic POD TUS) Lerres tei) Gertolinte fot Cape York Penosebe Cape York Peatusuli tan tae Some Satgngh Resources Amatyae Prem, Bribie Cush, OF we at (ie Cee ordiitern Cigna SHhvstion. dh PECPOM @uypy Mebwilie Sutra) (rks Rasetirie Thar Cheer repiet Hor Hie Oticenshanel Departenn ot LWIPO nen ie Peri ite) Siwy 1 PMT DL EYSGO Kaniitiida) al Capa York (hikari rept faite Quive osha Deporte i) Deieaments Department of A large hylid tro (Liforia andiiremalin MeDonald, 1007) was discovered in boulder fields of the Melville Ringe. In addition, a second few frog species was Tocated amongst boulders, This species wats recognised as a mentber of Microhylidae. a family well represented in New Guinea bur with Aastraliin representatives restneted to the — subtamily Genyophryinae in Wwe genera Cophivulus and Spheneplirvne. Aastralian microhylils are contined to hortheast Queensland, with the exception of Sphenophryne adelphe Zwerlel, TUS. a species found in the north of the Northern Territory (Tyler & Davies 1986), Australian microhylids were reviewed by Zweilel (1985) who recognised 16 species. sever of whieh he deseribed at that time. Richards ef a, (1994) described Cophivaluy monticola (rom the Carbine Tablelands, northeast Queensland and here we deseribe a further Cophivalus from Cape Melville. Viaterials and Methods The omatertal stodticd is deposited) in the Queenstind Museum. Brisbune (QM) and the South Australian Museum. Adelaide (SAMA). Measure- ments were made with dial calipers reading wo 0.01 mm, Measurements tiken (ii mind) were > tympanum diameter (T), eye lo naris distinee (hb), eye diameter (TL). foot (F). hand (Hi), head with (EW), fiewd lengih CHL), internarial span (IN), snout ty vent length (SV). tibia length (PL). width oF third finger dise and of penultinite phahinx. width ot fourth toe disc qind of penuldimate phaliis. length ob hand and lenwth of foor and follow Zweifel (1985) tind Tyler (1068), Material was cleared apd stained using at modification of he method of Dingerkus & Uhler lal) M DAVIES A KR MCDONALD (O77), Description aid discussion ol Ostvolory Lollow ZAwenlel (1985), Results The new speeies ts assigned to Copliveluy on the basis of the following features: denuiries mot in conlict; vertebral Coluinm prococlous: longue !/y [ree hehind with ne furrow or pouched pocket: maxillie nat ome comtiet (relationship) with premuasilite indaiernmnable). ‘This combination af feattires assigns the species to the Genyophryninae (Zweitel 1971), In addition, the species jacks procoracoids and vlavicles. has snout that is not narrow er clongate ind lueks at hypertrophied serous sland an the snout, The other defining feature oF Caphivalis, the aliry Process being typically slender and not merging insensibly into the body of the bone, could not he determiiued, Cophixalus zweifeli sp. nov, (FIGS 14) Holoiye: \' QM Jodess (formerly QNPWS N29789) Cape Melville National Park, 14° 15! 3S, eh? 27 40 BE, altitide 60-80 mi. 17,1, 1995. Coll, Ke. R, McDonald and L. A, lackson, Paruiypess & SAMA RS1L080 16.41.1995, Same location and collectors as holotype: 2 QM J64889 y+ Hi LP yatidiee fet sp. aM VTE ES Woah Sain (formerly (QNPWS N73038) Cape Melville National Park, Permwinent Camp Qld (rear type locality), allitude 40 m, T4.411,1995, Coll J, O'Shea (cleared and stained) Definiiion A large species (YY 40,1-45.4 mm SV) wilh long legs, large finger dises with third finger dise larger than fourth toe die. ai elonyate snout: dorsal eolounmtion brown with Mamescarlet axilla, thigh Nushes tind ventral lon markiigs. Deserimion of Holonype Head slightly narrower than body) legs moderately long CPL/SV OST), snout truneate from pbove, straieht and slightly projecting in profile (rigs 1,2) canthus rostralis straight, loreal region steeply sloping: nares anterolateral on Up al snout eye to hares stance ereaer Chon iitornianial span CliEN/TN = 1.125), eyes moderately urge, commend! outline cleautly visible from beneath, interorbilal width greater than widlh of upper eyelid. Tympanim linge. obseure dorsally, diameter greater than hall eve diameter. Relative lenuths ab fingers 3=d=2=1, the first slender and approximately hall the length of the second (Fig, 3). Dises of lingers 2-4 greally enlarges and trumeate, That of first barely extending, beyond Width OF penultimate phalans (Fig. 3): sabarticuler tubercles founded. moderuely prominent. bow a 4 NEW SPECIES OF COPHIXALUS If] Fig. 2. Cophixalus oweijeli sp, nov, A. Lateral and B. Dorsal views of head of holotype (QM J64888), Scale bar = 5 mm. ovoid inner and outer palmar tubercles. Relative lengths of toes 4>3>5>2>1 (right foot). toe three abnormally short on left foot; length of first toe approximately half that of the second. All toes with enlarged truncate dises with terminal grooves. Discs on first and fifth toes smallest and approximately same size. Toe dises smaller than those of fingers 2-4 (Big, 3). Subarticular tubercles rounded, moderately prominent. Low elongate ner meta- tarsal tubercle. no Outer metatarsal tubercle. Dorsal and ventral surfaces smooth. Colour and pattern: dorsum tan with darker brown pigment spots above insertion of arm, alony flanks and superior to inguinal region and along midvertebral region; large faint mark between and posterior to eyes: dark canthal stripe [rom tp of snout, through nostril and eye and above and slightly posterior to tympanum. Pale crescent along anterior R) Fig. 3. Cophixaluy cweifeli sp. nov. A, Palmar view of left banc. B. Plantar view of right foot of holotype (QM 64888). Scale bar = 5 mim. 162 M, DAVIES & K. KR, MCDONALD rim of tympani and paler stripe along lower rinr al eye, Dark brown pigment patehes on dorsal surfices of hand and forelimb, Lesser pigrient spots on dorsal surkive of Took dark patches along anterior edee of tibia, Throat very lightly dusted with pigment, more concentrated around margin of jaw and speckled with white, Measurementy SV 40,L0TL 20.6; IW 13. 9 TLL 13.4: B45; 73.1; RN 3,6; IN 3.2; third finger dise 2.2 (penultimate phalanx 1.0); fourth toe dise 1.9 (penultimate phalanx 0.71); hand 11.7: foot 7.3: ThISV O31; HW/SV 0.35; HL/SV 0.33: TIN/IN 1.125; HIL/AW 0.96, BESY Ob IN/SY 0.08. third tager dise/SV O.054; fourth toe dise/SV¥ O,047; hand/sV 0.20: foot V O45: ENS V (0902 7/E ULAe Colour ia life Dorsal surhice beige when lirst observed at night darkening to lan during Ihe day with widely seatleredt, irrenulie brown speckles. Brown moti on aim wd thigh dorsal surface, Black canthal streak trom snout (rough eye and above tyimpianin, Lateral dark brown mottled marking between axilla aml groim-. Axithi gromn. hidden parts of thigh. ventral dbia aod miner halt of Foot flume searlet (Smithe 1975). Ventral surtice densely mottled Fight purple on throat and ches! becomme more ditfuse posteriorly. Ventral sutfiaee of femuciund arm mottled with brown, Dull yellowish waste on lower third oF abdomen and ander the femur, Brown ventral surface to hand and foot. Varnintan The iwe paritypes have the following meisure Mens: SAMA RSIO80: SY 41.5; TL 22.2: IW 128: TT Ide BOS05 LE 43s RN 40k IN 4.2: third Hinwer dise 25 (penultimate phalang 12)e fourth foe aise 17 (penullimare phakins O.6% hand lds fet TT, PLS 0.54: DIW/SV 0,40; HEYSV 0,45; AYES O95: TNS TW 113, BYSV 0.12) TIN/SY O18 (hint finger ise/SV 0.06; Fourth toe dise/SV O.04; laml/S'¥ (29: thovSV 0.46: EN/SV (1.006, T/E 0.66. OM [6g885. SV sh Th 20.3 IW 14 2: HE IAL be oP 34: PN SAIN Ay did finer dive 245 (penultimate phaking Lay: forth te clise 19 (penualtiniite plains Uoer hand bOo8: foat 1S: TES Gate: HAW SV O32: HLiSV 0.475 BN/IN 108; TILZTIW 0.92; BySV 0),U8: TN/SV O77. hid finger tite’ ¥ INS 3; fourth toe HiseYS¥ 0.0425 hand/SV 0.242 thovS¥ 143, PNVS\! WNT be ORS Dori) caleut os more hrewe than tin i SAMA ROVORD ciel He nurkitges are more clistiet The SCenN a! SUF. In puticnloralthe thine and pheno WHOM er ds flore eavaly ails irrenuda hl) prerie ted walla Laral witike stripe amehialhy. The undersurtice ul the (highs ts rere heavily spechted, Comparison with ather species Cuophixalus cweifeli sp, nov. is uw very large species of Australian mierohylid comparable only with ©. sevadiliy Zweifel & Parker, 1977. In addition, the (hird finger dise of the Hew species is hunger than (hat ofthe fourth tue, a feature shared by Cy sewerifiy aml ©. vrnares (Pry, 1912). This latter species is smatler than eiher CL aweifeli ar C. saxarilis, The eanthus rostralis is straight in C, Sweijfels compared with a pounded canithus in Cy seediliy, The snout of ¢, cwelfele is longer than thal of ©. seveailis. The distinctive Thame-searler colouration on the hidden surfaces af legs is not found in any other Caphivalies it Australia. Peniies of ©. yeaediliy ure canary yellow at night, darkening tow light tan during the day, Unpublished cata of ©. Hoskin from mitochondrinl DNA sequences show C. pieifels lr be i sister taxon to CL dafacenis Zweitel, 1985 and ia separate clade from ©. savediliv, Cophivatuan infeicetis is asmall species (females te 17-6 mm SV) with a rounded eanthus rostralis. features not shared by C\ cweifeli. In life, CL infaeetis is dark grey on (he uiderside, compured with [he purplish colour of the (hroat and chest ole. cweiletr. Osreology One paritype wis cleared und stained, but untortinitely becuse of poor preservation. the material did not remain iitwet throughout the indeeration process. However, charicteristie and diagnostic features were obtainable, Skully The skull is lootiless with well-deyelaped and well-ossified nasals and frontoparielals. The ybanrtojigal arrieuhwtes with the minithi ‘Che eleutheroenathine condition af the premaxillae, typical OF the Genyophryiie, could nou be confirmed, The oloceipilal region (prootic and exoceipial) is ossified and the bones are vlosely aansociited with each other The vemers have aw well developed transverse arin (probable fised vomer yc pulling) arising froth an expanded arew in the inidline of the palate. and an anterior ani that passes mesial to and then anterion to the jiternal mares. he lraunisverse arm teaches The manithiry shell althoueh remaining lied Uy the maxilla by cartilaee (Pig. 44, The pleryuoid is extremely robust Mhete ts u thickened median portion al the lye plate and. (he posterior cornu huve well-developed Nanges it ig 4). The pewtoral girdle lacks chreictes wid a Sery saul medial projection niw represen| a yestioal ormostembin (Pig. 4). Cale icanen is whsenr in the mesosterimal region Presnerml vertebrae ure nate-inbricnte, Relative Willlhs af Lrninsverse pricisses andy ISS DelVotl SVSVISVIISVITL Vestiziol taunsverse priesses lin APPUreAL on the Urosty le NEW SPECIES OF COMIINALUS lo Fie 4. Caphiivelis sweifeli sp. nev. AJ Right yvomerine bone in ventral view. B. Dorsal view of hyoid plate. C. Ventral elements af pectoral girdle, D. ‘Terminal phalanx of finger (Paratype QM Jo4889). Scale bars = 1 mm. ‘The tps of the terminal phalanges of the hands and feet are ‘T-shaped (Fig. 4), Comparison with other species Zweitel (1985) examined the osteology of 1) species of Australian Cophixvalus and the current comparison is with these data, The otoceipital region of C. zweifeli is similar to that of CL savatilis, C. concinnus Tyler. 1979, and C. exiguus Zweifel & Parker, 1969. Cophixelus infacetus and C. hasmeri Zweitel, 1985 have the ossification of the prootics restricted to buried nubbins, as seen from above. The other species examined by Zweitel have an intermediate condition between these two extremes, The fused vomers and palatines of C. cweifeli approach those of C. coneinuus in their relationship with the maxillary shelf. whilst the mesial extension upproaches that of C. orntatus, The anterior portion of the complex approaches that of C. saxurilix although itis more robust in C. zweifeli. Zweitel did not recognise characters in the hyoid us being useful in interspecitic comparisons. Some Cophixelus (including C. sexarilis) have a small cartilaginous protrusion on the anterior ventral midline of the pectoral girdle (7 vestigial omost- ernum). A smaller process is apparent in C. cweifeli. The terminal phalanges lack a median notch found in C. infacetus, C, saxatilis and C. ornatus. Distribution The species is known only from the type locality in Cupe Melville National Park. Hahirat The habitat of C. zwe/feli sp. nov. is restricted to boulder fields of Altanmoui granites (Fig. 5). The holotype and paratype (SAMA 51080) were located at the base of rocks at night near a ereek flowing through the rock formation, No calling was heard. Paratype (QM J64889) was found on a rock in a stream flowing out of the boulders. Kivmology This species is named for Richard G. Zweitel, former Curator of Herpetology at the American Museum of Natural History, New York, whose revision of the Australian microhylids is a standard reference. We honour his contribution to herpetology and his friendship. lod MM. DAVIES & Ik. Discussivn Morpholiwically Cupliccdiay swerfell appears lo he a SISTET Spec wetoc \waeilis Bathare Titer Vr iN; ihe lest of any oVisteahian microhylil and semnilar m only jUOpartions Che fame-searler colgutuen m the AN are amt on the legs is uniqoe WO) cwetfele bh auldition to morphologiont appemrenee. ciel und C. wawveeliy uclive Similar habitats al eranitic boulder fhelds with patches at veeehilion iemost pockets (Pig. 5: Parker 1977, Pics 7 for the habitat of ©. This Taem oF taba is restrivted to the Melwatle Range ind Black Mountam im Cape York Peninsula, Sith sonal aveas of just a few hectives are found in nimenws locelons it eastern Queenstind (Sianton $994) The direer distanee Tront Black Mountain ty the Melville Ringe is (75 ki, Ramfall araynd the Cape Melville Rane is estimated 1 be as Hiwh) as 2000 mim (seme 700 iim bisher than the surrounding country) (Stanton booed’). Nouvithsiinding the merphologienl taku abe owedeli with CL savartlis, Tloskin™s dane trom mitochondriil DNA sequences fink the specurs wilh C oinfveems in separate clade from ©, wavarslis Zweitel (LOSS) atempled ta denve a tree ot closed see Fawejlel & WE WALES). melitionships iimonest Cuphivalis usm extertol {Me DONALD Ms momphohwicul characters bul fem thas foo be “ursaislying (Aweilel PRS p. 470) net belidve that uny ane af his miest plrsinenions Zwerlel did lites Woe detemsible, Given We Porn-congiueroe mourphilesicnl anu biechermtenl lata here. ito ots vleue tr a robust beiw cen micheuted more marpholowical study ust other morpholoeres (ie exrenvial featares aS Needed eas a lostat the roObustess of the mitochondrial DNA diliie Th the allie ane copious, Indepemdem and evenly cisiributee aevess the branches of the tree, phylogenies Cream any dary (494) gine coneruenee between trees trom diferent cutie sets set fend to converge (Mishler stich provides strong ewilente far any hypothesis ol phylugenerie® bistary Hos clear bowevor thar whitewer the data set ised i derive felanonships, monophy fy iy Coplivdas Hest most be deniwnstrited 4iistralun Acknowledgments We thank L. Jickson and J. O'Shea for feld assistamwe and Co Lloskin for permission ti quete unpublished dita from his BSe tHoms) Chess and further Studies, M. J, Tyler critically read the manuseript We oalo thank the refenses fot can- SPH Wet SUSOUSTIOIA Mark where Caliitilies Wedfel sy HOY IN hold NEW SPECIES OF COPHIXALUS 165 References CovaAcevicH, J. & INGRAM, G. J. (1978) An undescribed species of rock dwelling Cryptoblepharus (Lacertilia : Scincidae). Mem. Qld Mus. 18, 151-154. Dincerkus, G. & UHLER, L. D. (1977) Enzyme clearing of Alcian Blue stained whole small vertebrates for demonstration of cartilage. Stain Technol. 52, 229-231. JAMIESON, B. G. M. (1997) Some new and_ previously known earthworm species from Cape York Peninsula (Annelida: Oligochaeta; Megascolecidae) Mem. Qld Mus. 42, 233-270. Litre, A. & HALL, L. S. (1996) Preliminary observations on the bats of Cape Melville National Park. N. Qld Nat. 34, 53-57. McDonatp, K. R. (1997) A new stream-dwelling Litoria from the Melville Range, Queensland, Australia. Mem. Qld Mus. 42, 307-309. (1998) First Queensland record of the burrowing frog Cyclorana cryptotis Tyler & Martin, 1977 (Anura:Hylidae). Trans. R. Soc. S. Aust. 122, 85-86. MISHLER, B. D. (1994) Cladistic analysis of molecular and morphological data. Am. J. Phys. Anthropol. 94, 143-156. RICHARDS, S. J., DENNIS, A. J.,. TRENERRY, M. P. & WERREN, G. L. (1994) A new. species of Cophixalus (Anura:Microhylidae) from northern Queensland. Mem. Qld Mus. 37, 307-310. SMITHE, F. B. (1975) “Naturalist’s Color Guide” (American Museum of Natural History, New York). TyLer, M. J. (1968) Papuan hylid frogs of the genus Hyla. Zool. Verhandl. (Leiden) 96, 1-203. & Davies, M. (1986) ‘Frogs of the Northern Territory’ (Conservation Commission of the Northern Territory, Alice Springs). ZWEIFEL, R. G. (1971) Results of the Archbold Expeditions No. 96. Relationships and distribution of Genyophryne thompsoni, a microhylid frog of New Guinea. Amer. Mus. Novit. 2469, 1-13 (1985) Australian frogs of the family Microhylidae. Bull. Am. Mus. Nat. Hist. 182, 205-388. & PARKER, F. (1977) A new species of frog from Australia (Microhylidae:Cophixalus). Am. Mus. Novit. 2614, 1-10. DEVELOPMENTAL BIOLOGY OF UPEROLEIA ALTISSIMA DAVIES, WATSON, McDONALD, TRENERRY & WERREN, 1993 (ANURA: MYOBATRACHIDAE) By MARGARET DAVIES* & KEITH R. MCDONALDF Summary Davies, M. & McDonald, K. R. (1998) Developmental biology of Uperoleia altissima Davies, Watson, McDonald, Trenerry & Werren, 1993 (Anura: Myobatrachidae). Trans. R. Soc. S. Aust. 122(4), 167-172, 30 November, 1998. Uperoleia altissima is a small fossorial frog restricted to upland areas in northeast Queensland. The frog breeds in the monsoonal wet season, and lays clumps of eggs that fall to the floor of ephemeral ponds. Larvae hatch at stage 19. Later-stage larvae have moderately strongly arched tail fins, a sinistral spiracle, large, narrow, cavernous, external nares, and a larval tooth-row formula of two upper (second divided) and three lower rows (first and second divided). Labial papillae are strongly interrupted both anteriorly and posteriorly. Later-stage larvae are strongly pigmented although the strongly-pigmented tail tip of earlier larvae is less so. Larval life span is about 39 days in captivity. Key Words: Uperoleia altissima, larvae, embryos, life history, tadpole, Myobatrachidae. Fransaettons of the Royal aectery ofS. Aust (1998), E224), 16772 DEVELOPMENTAL BIOLOGY OF UPEROLEIA ALTTSSIMA DAVIES, WATSON, McDONALD, TRENERRY & WERREN, 1993 (ANURA:MYOBATRACHIDAE) by Marcarer Davies’ & Kern R. McDoNaALb Summary Davies, M, & McDonanion KR. (1998) Developmental bialagy of Uperoferds aid Gin Davies. Watson. MeDonald, ‘Prenerry & Werren. 1993 (Anurw Myobatrachidie). Pras. Ae Soe. S. Ause 122(4). 167 172. 30 November, (998, Unperaleia alixsines is asmall fossorial frog resinictes 10 upkind areas in northeast Queenshtud, The frog breeds ithe monsoonal wer season. and keys elueips of eggs that fall te the Toor of ephemeral ponds. Larvae Taich al stage 19. Lalter-stage larvae ave moderately strongly arched tail Tins, a sinistral spiracle, large. narrow, cavernous. extemal nares and a larval tooth-row formula of lwo apper (second divided band three Tower rows (lirst and second divided), Labial papillae are strongly interrupted both anteriorly and posteriorly. Taiterstage larvae are strongly plemented although the svongly-piemented tail lip oF earlier Lirvae is Tess so, Larval tle span ts about 39 days in caprivity KY Wokds) Uperledd altivstine, hurvaed. crbryas, lle listory. tadpole, Myabatraehidie Introduction Uperolerd altissiita Davies, Watson, McDonald, Trenerry & Werren, 1993 1s a Woothed member of a speciose venus of small fossorjal frogs distributed across mainiind Australia except for the southwest of the continent. Uperaleia altissimea is confined to elevated sites on Ue western Wet Tropics Bingeoyraphic Reyion (Stanton & Morgan 1977) from Princess Hills, Lumbolz National Park. north bo the Windsor ‘Tablelind of northeastern Queensland, The species is found tn moist cucalypt forests and woodlinds above 600 metres. Although deseribed i 1995 from freshly collected material, the species had been collected but vot identified previously. Little was known of its breeding biology. In carly February 1997, We cneountered a breeding chorus af the froe following heavy miinkdl at a site onthe Atherton Tablelands. Amplectant pairs which luier Spawned were collected, and the resultant larvae were reared to metumorphasis. The deseriptian of this life history adds to the scarce dita available on dite histories of the 24 species of Uperolere (Moore 1961; Watson & Martin 1973; Tyler e/ al. 1983: Davies ef al, 1986; Richards & Alford 1993: Davies & Warsan 1998), Materials and Methods The series of Uperalera altissmia was obtained from spawn deposited by amplectui adults collected Depl al Aoolwy. Criiveriiy of Adelie Anata S005, 7 Conservalidn Strateey Branch. Peparnnientol Lnvironmedt, Po Box S47 Wherton Ole 4883_ in the field. Larvae were initially reared in aerted water at ambient temperature (water temperature approximately 24° Cy) in the field before being iransported (a Adelaide Where they were maintained in wv constint temperture room at 3h + 1 Can dechlorinated tap water. Larvae were led an boiled organic lettuce eaves supplemented with come mereial goldfish flakes (Biosera). Material wis preserved mi ‘Tyler's fluid (Tyler 1962) and illus trations were made with the wid ola Wik M& sterco dissecting microscope with attached camera lucida, Measurements were taken using an eyepiece miero- ineter, Developmental stages are those of Gosner (1Y60), Results Four amplectant pairs of Uperdlita alussiie were collected on 1.4.1997 at a quitery near Carriigton “alls (17° 19" SI" S, 145° 26! 42" &). The site tsa quarry with gravel pits some of which have regrowth vewetation, We had visited the site on the previotis night utter min, but although O. adtissena was calling, no breeding was observed, However, Litaria rubella (Gray, (842) was calling and breeding took place later (hat night. Other species calling when Li. alussima was breeding included Crinfe remota (Tyler & Parker, 1974). 2. rasure (Gray, 1842), 1 rothii (DeVis, 1884). 0. fallax (Peters. S81) and Limnodvaastes: terracresinae Very, VOUS. The night was humid following torrential rain in the nearby Herberton Range, although rain did not appear to have fallen at the site. Males were culling from the gravel areas surrounding the temporary pools (Pig. 1). often 1M M DAVIES All. RL MeEDONALID Pig, 1. Callin tale Uperaleke aliisinie ab Careington Balls Quuirey site (SY approximately 24 nim}. ae SS". - hee 2 Atnpleenint (aierdeda adnan a Coecipton Pah quarry st (SY oat nile approwonately 24 mn UPEROLEIA ALTISSIMA DEVELOPMENT loo facing away trom the water toward the surrounding vegetation. Pairs in inguinal amplexus were found moving toward the shallow water (Fig. 2) The U. altissimea spawned early on 2.11,1997. At 1310.0n 3.41,.1997, the eags had reached stage 12. late vastrula. A Single capsule surrounded the oyun. Mean capsule diameter of four eggs was 2.27 mm (range 2,22-2,32) and the ova had a meun diameter of 1.36 mm (range 1.32-1.40 mm). At 1300 on 4.41.1997 embryos were at stage 17 (tail bud) (Fig. 3). with the tail being better developed than the head. Adhesive elands and the stomodaeal pit were prominent, They Fie. 4. A. Dorsal. B. Lateral views of Stage 17 (tail bud) embryo of Uperoleia altisstna, Seale bar= | mm. had reached stage 18 by 5.11,1997 (Fig. 4). Hatching (stage 19) was completed on 6.111997 (Pig, 4). The newly hatched larvae had no external gills: the eyes were very difficult to detect and the mouth had not perforated: adhesive glands were pigmented at this stage. By 0900 on 7.i,1997, some larvae were at stage 20. Adhesive glands were well developed on stalks and both the mouth and the external nares were perforated. Larvae were still at stage 20 at 1140 on 8.111997, the cornea was not transparent. and heavy pigmentation was apparent on the tail fin. Larvae had reached stage 26 by L000) on 14.11,1997. The horny beak was keratinised as were upper und lower labial tooth rows, The adhesive glands were reduced to patches of pigmentation. The nares were large and cavernous and the tip of the tail was particularly heavily pigmented (Fig. 5). Stage 2% was reached by 1100 on 19.i7.1997_ Later-stage larvae lacked the heavily pigmented tail tip. Pig. 4. A. Ventral. B. Lateral views of Stage 18 (muscular response) embryo. C, Lateral view of newly hatched larva of Upervleta altissima at Stage 19, Seale bars = | mm, 170 M. DAVIES & K.R. MCDONALD hiv. 6 A. Lateral. BL Dorsal views of Stage 34 larva of Uperaler alrissinia, Seale bar= 5 mon. UPEROLEIA ALTISSIMA DEVELOPMENT 171 Tarn tl. Measurements (in mm) of body and total lengih ef larvae of Uperoleia altissima as mean and range. N = number of individuals, Stage (Gosner 1960) Body length (mm) 28 6.16 (6.08-6.24) 29 6.83 (0.36-7.36) 30 6.72 (6.72) 32 7.84 35 8.42 (8.00-8.64 36 9.36 (928-944) 37 936 (9.12-9.60) 38 9.09 (8.64-9.6) 39 10.08 (10.08) AQ 9.60 4] 974 (9,6-10.56) 42 10.00 (9.99-10.08) 43 10.27 (9.6-10.94) 44 10.24 (10,08-10.40) 4s 10.44 (10.4-10.56) 4h 10.24 (10.08- 10.400) Measurements of larvae at stages 28-36 are given in Table |. The following description is of a larva at Stage 39 (Pig. 6). Body ovoid. widest at midpoint. Snout evenly rounded in dorsal and lateral view, Nares dorsal, Jarge. narrow and cavernous. Eyes conspicuous, Spiracle sinistral, moderately long, opening postertorly and searcely visible when viewed trom above. Anal tube broad opening dextral to veniral fin. Dorsal fin more strongly arched than ventral fin, Fins rounded terminally. Dorsal tin commences on posterior part of body and is deepest about halfway along 1s Jength. Ventral fin commences posteriorly io body and is deepest about halfway along its length. Tail musculature moderately thick, tapers to point. Oral dise small and ventral. Labial papillae widely interrupted aateromedially; less widely interrupted posteromedially, Two rows of upper teeth, secand divided: three Jower rows, first anel second divided (Fig. 7). Short P3 row supported on Mexible flap. Tail musculature and fins heavily Total length (mm) N 13.68 2 (13.12-14.24) 16.53 3 (15,68-17.12) 16.56 2 (16.32-16.8) 19,20 I 19.31 3 (18.24-20.48) 23.68 7 (23.52-23.84) 23.68 4 (21.92-24.96) 24.08 6 (22.24-25,92) 26.08 2 (25.92-26.24) 25,28 | 26.26 y (25,76-27,36) 2 7 qd 4 i suffused with pigment. Dark-brown islands of pigmentation on body. Larvae reached metamorphosis at stage 46 on 13.11.1997, 39 days afler spawning. Discussion The complete larval biologies of Uperoleie altissima, UL inihdarea Tyler, Davies & Martin, L9st (Tyler ef af 1983) and Lo talpa Tyler, Davies & Martin, L98t (Davies & Watson 1998) are noy known as are tadpole morphologies of U, ryleri Davies & Littlejohn, 1986 (as UL maciearcuti. Watson & Martin 1973). Of) lithamoda Tyler, Davies & Martin, D98t (Davies ef al. 1986) and LL iminitla Davies, McDonald & Corben, L986 (Richards & Alford 1993). ‘These latter two species occur in or near geographic locations of OC. affissiima, hence a comparison of their salient features is of value for identification of tadpole assemblages. Uperaleia mime and U. lithamoda share a looth row formula of 2(2)/3 whilst that of LO eltissfra is 172 M. DAVIES & K. R, MCDONALD f OUTTIMM NT “ é Wie Fiz. 7, Oral disc of Stage 37 larva of Uperofetu elfivsinu, Seale bar = ] mm. 2231.2). The flexible Map supporting P3 labial teeth is recorded in all Upereleia to date, but is not unique to Uperolera (Davies & Watson 1998), The dark tail lip in early stages of U. mimulea recorded by Richards & Alford (1993), is shared by U. lithamoda and U. altisyima. The heavy pigmentation of later L’, altissima larvae may be greater than the diffuse pigment of U. lithomoda and U. minnula. and is a feature of other Uperoleia (Davies & Watson 1998). There are considerable differences in the length of the spiracle, that of U. alrissima being intermediate between those of U2 niimul/a and U. lithomodea. The unusual tail bud stage in which the tail is better developed than the head was noted also by Moore (1961) in LL feevieata Kelerstein. 1867 and Davies & Watson (1998) in UO) talpe. Acknowledgments This research was supported by the Department of Zoology, University of Adelaide and the Queensland Department of Environment and Heritage, We thank M. Tyler for critically reading the manuscript and the referees for their helpful comments. References Davies. M., McDonacp. K. R, & CorRen, C, (1986) The genus Uperoleia Gray (Anura: Leptodactyhdae) in Queensland, Australia. Proc, R. Soe, Vict 98, 147-188, & Watson. G. FE. (1998) Developmental biology of Uperoleia talpu Tyler, Davies & Martin. 1981 (Antira:Myobatrachidae), Trans. Ro Soe, Ausi. 122, 153- 157. . MCDONALD. K. R.. TRENERRY. M. P. & WERREN, G, (1993) A new species of Upernieia (Anura: Le ptodaetylidae: Myobatrachinae) from northetstern Austraha. Ment, Qld Mus. 33, 1607-174. Gosnek, KL. (1960) A simplified table for staging anuran embryos and Jarvae with nates on identification Herperalugien Ub, 182-190, Moori. J, A. (1961) The frogs of eastern New Seuth Wales, Bull. Am. Mus. Nat, Hist. 121, 149-386. Riewaros, S.J. & ALFORD, R.A. (1993) The tadpoles of iwo Queensland frogs (Anura: Hylidae. Myobat- rachidae). Mem. Qld Mus, 33, 337-340. SrANTUN, J.P. & Morcan, M. G. (1977) "Project RAKES - a rapid appraisal of key and endangered sites, Report No. 1: the rapid selection and appraisal of key and endangered sites: the Queensland case study” (University of New England School of Natural Resources, Armidale}. Tyrer. M, J.(1962) On the preservation of anuran tadpoles. Atist. J. Set, 25,222. Crook, G. AL & DaAvins. M- (1983) Reproductive ~ biplogy of the frogs of the Magela Creek Systeim, Northern Territory. Rec. S. Amst. Mus. 18, 415440, Warsos, G. F. & Martin A. A. (1973) Lile history, larval morphology and relationships ot Australiana leprocdacty lie frogs, Trans. Ro Sac. 8. Att, 97. 33-45, CHANGES IN A MANGROVE/SAMPHIRE COMMUNITY, NORTH ARM CREEK, SOUTH AUSTRALIA By PERI S. J. COLEMAN* Summary Coleman, P. S. J. (1998) Changes in a Mangrove/Samphire Community, North Arm Creek, South Australia, Trans. R. Soc. S. Aust. 122(4), 173-178, 30 November, 1998. Use of a computer GIS package to study aerial photographs of North Arm Creek (1979-1993) confirmed previous studies suggesting a landward migration of the grey mangrove, Avicennia marina, but seaward progradation was also apparent. Samphire communities were reduced in area by nearly two-thirds, with the majority of the lost area overgrown by mangroves. At the same time samphires colonised unvegetated areas and some areas previously occupied by mangroves. From 1979-85 the area colonised by samphire was similar to the area lost, but was less from 1985-93. It is suggested that several factors are responsible for the changes in distribution. Key Words: Avicennia marina, Halosarcia, Sarcocornia, mangrove, samphire, saltmarsh, temporo-spatial change, progradation, colonisation. Pranic tions af the Rove sacieny af S. Atist (1998), 122) (4), 173-078. CHANGES IN A MANGROVE/SAMPHIRE COMMUNITY, NORTH ARM CREEK, SOUTH AUSTRALIA by Peri S. J. COLEMAN’ Summary CoLeMAm®. PS. J. (1998) Changes in a Mangrove/Samphire Community, North Arm Creek. South Australia, Trans. KR. Soc, 8S. Aus, 122(4), 173-178. 30 November, 1998. Use of a computer GIS package to study gerial photographs of North Arm Creek (1979-1993) confirmed previous studies suggesting a landward migration of the grey mangrove. Ayieennia marina, bu seaward progradation wat also apparent. Samphire communities were reduced tn area by nearly two-thirds. with the majority of the lost area overgrown hy mangroves. At the same time samphires colonised unvegetated areas aud sume areas previously occupicd by mangroves. From 1479-85 the area colonised by samphire was similir to the area lost, but was Jess from }985-93, It is suggested that several factors are responsible for the changes in distribuion, Key Worps: Aticcnnia marina, Halosarcia. Sarcocurnid, mangrove. samphire, saltmarsh, temporo-spatial change, progradation, colonisation, Introduction North Arm Creek drains from the Wingfield/Dry Creck area of Adelaide northwards into the mangrove zone of Barker Inlet (Fig.1). The zone comprises a seaward tringe of the grey mangrove Avicennia marina (Porst) Vierh. var resinifera (Forst) Bakh., backed by a salt marsh comprising nixed samphires of the genera Hulosarcia P. G. Wilson and Sercocentia A, J, Scott. The mangroves und sumphires form bands of variable width on both banks of the creek. The ereek has been used for the reception of stormawaters, sewage effluent and trade wastes, The wel coustal ecosystem edging the creck has been considerably modified since European settlement, In the fite 1800s seawall embankments were builtalonge the mangrove/samphire interface, and the samphire Aone Way used as papturage, Salt production on the eastern side of the creek began in 1934 and progressively much of the low lying area inland ot the seawall embankment has been ponded, On the western side of the creek the low lying lam behind the seawall became a muanicipal refuse tip, In the JO70S 8 series OF groynes supporting power pylons Was butt thioueh te taingrove/samphire zone ubulling the creek, The more recent changes have resulted in changes to the water flows and tidal dynamics inthe area. tn ella Bavironimentil Cotati, b2 Beach Reak St kil Aus ST) Hinvoriey, POL CEM ROy Haseltine Stmdy and rely: bvaloetion OF TCT Solt bvapergden Ponds” Bitterns Discharge on the Mangrove Continuity. sith Supplement Repunt provided ta (C] Ncatoilias Ackelatde, CU api. ), Sttily Aten oe Nak Puinp Torrens Island Swat Ale Lefevre Af Mier: Peninsula € ( Norlt ray ry jf Port Adelaide North Ne wich Fig 1. Map ot the region. J986 Bradlev! recorded large seale dieback of bath mangroves und samphires im the yiemity ol the power pylon grovnes and recent field inspections have revealed that the area is only slowly berry recoloaised, Aerial photographs of North Arm Creck, taken between 1949 and 1993. show changes in the mangrove and samphire communities. Some changes are marked. such as areas of dieback. or the inland wdyance of mangroves. The use of Geographic Information Systems (GLS) technology has allowed a closer look af the changes in ane small region of the 174 PS.) COLEMAN North Arm Creek coustal wetlind, the drainage area uf the Dry Creek Salttiells' No. & Pump. The area is hounded On the east hy wt seawall and om the vest by North Arin- Creek, Running ecntally veross the area is a small greek (hat has been formed by the discharge of bitterns (brine that remains after salt erystalligation is complete) from the saltiields ‘The No. & Puinp and its supply drains are clearly visible on aerial photographs, Previous studies Burlon (1982) studied mangrove development north of Adelaide to the River Light using aerial photogriphs eovering the period from 1935-1982. Ve noted that the manyroave stinds showed dilfereat direvtions of growth at the (wo extremities af the study zone. Crenerally the northerly inungroves were prograding (extending seawards) while the southerly Inangeraves near Swan Alley were reureutine tlie aerass The samphire Mats. Burton's paper discusses Ihe possible causes of tis difference, i parlieukar discussing ferrigenbus supply and relative scu-level tise (eustatie rise and land subsidence). Durie 1985-6 Bradley! examined the mingroves in the vieinity of the Noo S Pump on North Arn Creek, He tsed Visual comparisons of aerial photographs of the area laken jn 1879 ard 1085, He peeved two trainscer fies weross the uresaued ntupped a Faget bi anus = Advpttine * SAUNA orth 143 vn free Pinan f 0 all uD Lin Tie 2 Tie neue ey ae PP ad Od the distribution, heallh and age oF the veweralion along these, Plastie 30 cm rulers were attached to the transect pees to deterniine passible sedimentation patterns. The alignment of the substrate on the rulers wis recorded, A further study of the North Aro Creek te Swan Alley area was undertaken by Blackbuen® in 109d, He used GIS (eebniques to asvertain distribution changes within the mangrove and ssimphire communities. Blackburn? did nol physically visit the North Arn Creek. but the photographs he examined indieated both landward and seaward progradation Gl mangroves, The present study re-cxamines the area reported by Bradley! (Pig. 2). The review of the area combines a GIS analysis of vertil photographs (1949-1993) with 4 ground survey using Bradley's! existing: transects. The study was constramed by growth ob Maneroves making access dilficull, lass of some sediment rulers and loss of transect pegs nearer the seaward fringe: Materials vod Methods Department of Environment and Natural Re- sources | 1480 seale cokugements of four aerial survey photographs dated 1 January 1949. 19 March 1979, 18 February LO8S and & December 1993 were manually dribsed tate a orn saitable fay use in the GIS mapping package TNTonips Lile. The three more recent photowraphs. duting, from 1Y79 cw 1993, were veorcetified Using man-made structures onthe neihbouring sallfielvl as control ports, alone with Woluwted nmingrove GAL meri) lees Liat were ideniltiuble through the series of photographs. ‘Lhe 1949 photogriph shows a lindscape so different from the present thal geurceuilication cold only be waccomplished by matching Ue angles of narrow ‘barrow-pits” along the sea-wall. so vate from ir were Guy used ina general manner im the present Study, The prmeipal components af the mapped ares were dete is mangroves, samphires.ar jeither (water or bane mud An amilysis of the Timuts of vegetavon hver each OF twee periods (U7 )O85 und 185- YOU wos underiaken to boy blo vdewromine whit the dyvnuimics af the verehiuve vhange wore, Iieldwirk wh 1296 ineluded Winding: the Leaner pegs plived by Bradley’ in 1985. Vhe vexetution alone he Twe trynsects Was recorded and its heigl Weaswred Wy meres MSIE Lape Keudines were alse rewonled where sediment mifeds were still in puree Het oa Brarnues, Bo ciel) Mopiige Mayepase yin Salrenarss) Commmoties Sear ee Bleeets Digelwwe Area kintall Poti Atldale Report te Peirce Seen Mroelitere CE a piber CHANGES IN A MANGROVE / SAMPHIRE COMMUNITY 7 Typur |. General cheanves in the vegetation. 1979-1093, Mar 79 Mangroves (17) 73902 Saniphire (ns 14173 Vevehtlive cover (m2) RSOTS a beb—-$S Dee-"93 Change, *79-"93 TROOA N8459 + 14557 12769 5340 ~ $833 91377 93799 +5774 Tash 2. Change rates ef the prineipal campenents af ihe vegetation. Area of etch chine ehiss of the yezetalion (a) Annnal rate of change 1979-1985 Mangrove to niangrove (6 change) 69175 Saimphive te iminennve 202 Mamnurove to samphire 2913 Neither (o iuingrove 35) Mingreve to neither” 124) Neither’ (9 sumphire ARTO Siimphire to neither! 1924 Saniphive jn samplire (ao change) S004 Veretative loss 3163 Vegetative dhercase 7396 Rale of change in loti vegetative cover 1945-1993 TO7OTU8S IUSS. 1993 THY74 - - GOUS a4 762 20 46 7s 3758 587 47th ahh 207 74 W163 646 145 AOR 32| 400 2051) - 4576 527 a72 442] i254 613 706 43 * “neither” indicines areas of bare mud or water Because the A, mwrine trees had grown considerably during the TL years since the transect pegs were pliced, locating the pegs past the 80 mH mark on Transeet A and the 110m mark on Transeet Bomight possibly fave resulled in damiage to the mangal and so no dala were collected beyond these ports, Results Vevetarton imeppinge A comparison for the period 1979-1093) shows extension Of the mangrove canopy, and a reduction in {he aren covered by samphires. with an overall inerease in vegetated area (Table 1). As the study zone is delimited on the landward side by the seawall embankment, the gain must either be the overzrowing of previously bare mud patches, or some seaward uccession, The results of the analysis of the limits of vegelation Tram 7979-1985 and 1985-1993) are summarised in Table 2. The latwest change over the period was a jnerease of mangroves at the expense of the samphire community. tlowever, the extension of both mangroves and samphires over bare mud and into water areas ts alse oecurring, along With samphire colonisation ol arcus previously supporting mangroves. Lirosion is oecurring in some wreas of samphire, Samphire has given svay lo mangrove at their interface as the mangrove has. extended inland. Almost the entire central samphire zone has been suceeeded by mangroves and the trees have also oecupied many of the creck lines as well as colonising the bare mud dreas along the seawall embankment. Site visits ia duly and September |996 revealed juvenile mangroves growing along the No. 8 Pump discharge channel and specimens more that 4in high growing along the seawall within 40m of the discharge pout Some mangrove areas have been replaced by sumphire or by bare mud. This has mainly occurred in the southern part of the study area hut alse along the bitterns discharge creck Along the seaward edge, progmiadation ol mangroves is apparent along the entire length of the study zone. The extension is most marked i the southern areas, with a maximum advance of approxinnitely 25-30 min the 14 years from 1979- 1993, In the northern purl of the study zone the seaward progression consisted mainly of infilling the many invaginations and embayments around isolated trees and the advance was between 10 and 15 m. The cause of the slower progradation ol mangroves in the northern urea is uncertain but the milling of semi- enclosed areas suggests that low water flow rates in the sheltered areas were conducive to sediment wecretion, whereas the actual seaward fringe may have been exposed lo stronger wave action, The samphire community has also been extending, and has beeome established on previously bere mud: there ure now samphires along the seawall within [Sm 176 Taunt. 3. Sedimenrarion recdings along the trdasects. P.S.d, COLEMAN Distance A Transect GB Transeet along transect 1985 1986 reading 1996 reading, 1985 1986 reading 1996 readin initial (change) (chanpe) initial (change) (chanpe) redding readin in Ts 63 nit 5.2 63 nit ( tdbem) (-1) em) Hin bf K 3 45 46 7 (-lL4em) (r5.0 em) (-U,1 cnt) (2.4 om) Aan 43 52 4 4 34 ww (-0.9 em) (44.2 em) (+0.2 em) 50 m 8 7S ta 7 62 ie (40.5 cm) (+0.8 em) oom 6.1 6.1 Nib 63 63 3.5 (no change) (no chanve) (+408 om) Note 1: 1985 & 1986 readings from Bradley (|986)!, Note 2: Readings are the alignment of the substrate agaist 30 ch plastic rulers attached to the transect pegs. Zero is to the lop af the ruler, of the discharge point As the creeks are being iitilicated by miatueroves, new areas for samphire colonisation have appeared. Much of the new growth is along the seawall and to the north of the study urea, Where the ETSA eroyne his altered the tidal circulation. The bitterns discharge does not appear to have affected growth Of samphires negatively. possibly because (he species are adapted to surviving in high salinity regimes, but samphire has been eroded away in some areas along the bitterns discharge creck. Treisects Figure 3 presents the 1985 ind 1996 heights of the vegetation along Bradley's! existing transects and shows the muturation of young stands of mangroves and the new colonisation (by mangroves and sumphire) of areas closer to the discharge point. The 1996 date were collected along the transects in September, The forests are now so dense that accessing the pegs is difficull and so the iranseets do pat continue fo the original (50m point, Sedimentiaion The bitterns discharge creek has formed since the 1949 geri! photograph was taken and Bradley's! report capressed same coneerh that erosion might be oecurring in this creek near the discharge point, He examined sedimentation patlerns away lrom the iminediate disebarge point by attaching plastic rulers to the transect pons and recording the celative hehe Of (he substrate at each location, In response to Bradley's! finding during the initial Observation period (1985-1986) that some erosion wis oveuriie lea’ the discharge point, saltfield personnel deposited concrete blocks in the drain to break up the flow. To determine the types of change that might have been occurring since 1985/86, the rulers were examined ip September |996 where they sull existed. The few remiining sediment rulers indicated that the hydrology of the area may have changed. These 1996 readings are presented (Table 3) together with Bradley's! L985 and 1986 readings. The southern transect (Transect A) shows deposition to have occurred within tO m of and possibly closer to, the discharge point. The lopography of the transect is smooth, with no creeks crossing It, so sedimentation may be relatively uniform across the urea, The northern transect has several small evecks crossing il, and the Crosion/sedimentition pattern is more complex. The lack of rulers makes iCdifficult to Interpret, However, the urea closest to the discharge point has eroded somewhat over the list LO) yours, forming a creek tine, Al low tide any clischarse follows the existing creeks (slightly to. the north before turning westerly), which have become more detined as mangroves have colonised the Mats around them. The creek at 60 Mm is not recorded as having it sediment ruler on Bradley's! original sediment lable, but a reading Of the topographic plan of the transect done ih 1985 shows the creek to be about 15 em deep; the current reading is 14 em, The ruler an the JOO m Wansect peg in the main forested area along the northern transect shows a small sediment sain. Discussion The detailed GIS study was possible beeause sufficient Markers were visible iN veri photog raplis CHANGES LN A MANGROVE / SAMPHIRE COMMUNITY Distribution along Transect A 1985 Height of vegetation (mm) 177 60 70) 80 90 100 110 0) 100 Ho 10 20 30 40 Ss) A Distance along transect (m) Distribution along Transect B 19&S 4 1996 0 --2ee->- u x 2 = | B Distance long tumseet (my Fig 3A. Heteht distribution of vegetation along Transect A” B. Height distribution of vegetation along Transect Bs, to allow precise yeorectification. The 1949 photograph lacked some markers, reducing confidence in the precision of its georectification, However, this earlier photograph provides some insight into changes in vegetation patterns. The main differences include: 1. a larger area of vegetation in 1949 benween the seawall and North Arm Creek: mangroyes extended further out into the creek, no creek in the location of the current bitterns discharge creek and the land inside the seawall was grazing land. a wide expanse of samphires, with mangroves penetrating trom North Arm Creek in) towards the seawall along depressions, and 4. areas of mangrove dicback just behind the seuward [ringe of mangroves. i] to The 1949 photograph showed that the mangroves in North Arm Creek were already retreating inland, so the seaward expansion visible in the post-1979 photographs must have started before 1979 but atter L949, It is postulated that there has been an advance and regression of the mangroves with relatively small changes in water flow patterns. According to Hodgson et al. (1966) North Arm Creek received the flow of effluent trom the Islington Sewage Farm from 1881 through to the opening of the Bolivar Sewage Treatment Works in the 1960s. During the operation of the sewage farm, nutrient rich water would have been released into North Arm Creek, The effluent may have supported algal blooms that could have caused the sporadic oxygen depletion in the waters of the creek recorded by Hodgson (1959)* . Induced anaerobic conditions are 178 P. S.J. COLEMAN reported to cause the asphyxiation death of areas of mangroves (Diop et al. 1997) and this may explain the areas of di¢back visible in the 1949 photograph, The changes in the mangrove/samphire com- munities vistble in the 1979-1993 photographs confirm previous studies that have sugvested that a landward migration of A. marina is occurring in the southern reaches of Barker Inlet resulting in a reduction of the area of samphires, However mangroves are also prograding seawards and covering a larger area, suggesting that the growth and distribution pattern is not a response Lo a single factor. While land subsidence/sea-level rise (Burton 1982) may be responsible for the landward progradation, it cannot account for simultaneous seaward progradation. Sedimentation readmgs from. the transect rulers indicate that sediment is accumulating over much of the area and that any lowering of the Jand surface is likely to be a widespread Jandform settlement (PPK 19925) of the sedimentary coastal deposits rather than a lack of sediment supply or erosion per se, except in specific areas such as creek lines and patches of mangrove dicback. Samphire communities over the period 1979-1993 were reduced in area by nearly */; despite the overall gain in vegetated area. Most of the lost area was overgrown by mangroves, However, the direction of change was not entirely one way, as samphires colonised areas previously occupied by mangroves plus areas of mud/water. During the carlicr period, between 1979 and 1985, the urea of new samphire growth cach year nearly matched the area lost. so that there was an apparent loss of only 200 m? of samphire annually. The later period (1985-1993) showed a slowing im newly colontsed areas of samphire. Although the area overgrown by mangrove or eroded each year remained about the same asin the earlier period, the rate of loss appeared higher (LO0O nv annually) because there was Itttle colonisation of new areas by samphire References Burpon. T. E. (1982) Mangrove Development North of Adelaide. 1935-1982. Trans. Ro Soo. S. Aust. 6, 183- TRY, Hopason, HL J. N., Lewis, RK. Wo. MILes, K. Ro, Jupp, Po& GILCHRIST, J, W. (1966) Report of the Committee of Hnguiry ite the Udlisation of Bffluent from Bolivar Sewage Treatment Works. Government Printer, Adelaide. (Unpub. ). ' Honcson, H, JN. (1959) ‘Treatment and Disposal of the Sewage of the Adelitde and Salisbury-Klizabeth-Gawler Drainage Areas. Engineering and Water Supply Department, Adelaide. (Unpub.), PPR Consultants (1992) MFP Australia Gillman/Dry Creek Urban Development Proposal-Drafi Environmental tmpaet Statement prepared for the Premier of South Australia, Adelaide. (Unpub.} Diop, E. S.. SoumARE, A., DIALLO, N. & Guise, A. (1997) Recent Changes of the Mangroves of the Saloum River Estuary. Senegal. Mangroves and Salt Marshes 1, 163- 172, NEW SPECIES OF SEURECHINA (NEMATODA: SEURATIDAE) PARASITIC IN DASYURID MARSUPIALS FROM AUSTRALIA By L. R. SMALES* Summary Smales, L. R. (1998) New species of Seurechina (Nematoda: Seuratidae) parasitic in dasyurid marsupials from Australia. Trans. R. Soc. S. Aust. 122(4), 179-184, 30 November, 1998. Seurechina hobbsi sp. nov. is described from the stomach of Phascogale tapoatafa from Western Australia. It differs from S. chaneeti, the type and only described species, in being a larger worm (6-8 mm compared with 3.1-3.8 mm) with longer spicules (500-630 wm) for S. hobbsi compared with 185 ym for S. chaneeti. Seurechina spratti sp. noy. 1s described from the stomach and small intestine of Sminthopsis leucopus and Antechinus agilis and is most closely related to S. hobbsi from which it differs in having three lateral papillae extending into the caudal alae rather than two, oval rather than spherical eggs and the absence of a large projecting lip anterior to the vulva. Key Words: Seurechina, nematodes, Seuratidae, Echinonematinae, Australia, Dasyuridae, marsupials. Tronsaetions afte Ravel Societhy af S Aust (1998), T2204). 179-184. NEW SPECIES OF SEURECIIINA (NEMATODA: SEURATIDAF) PARASITIC IN DASYURID MARSUPIALS FROM AUSTRALIA by L. RL SMALES Sunumary Sviapes. LR. 1998) New species of Searveting (Nemiuoda 2 Scuratidie) parasitic ieckisyuridl iarsupiils lean Australia, Teaiis Be Soc Ss Alist E2204), 179-18, 30 November, 1998, Semrechine hebbsi op. WoW is described from the stomach ot Phiscogale tapoanife tron Western Australi. I differs Trou. ehaneer. (he (pe wid oAly deserbed species, i berg a larger worm (6-8 in compared: with 3.1 3.8m) with longer sprees (500-630 pind for §. frefbat compared with TSS pm lor S cfemeed, Searectiia yyrrotté sp. nay, is described [rom the stomach and small intestine OF Saintiopaty dedeopis and Antechinesy agilis and is mast closely related (oS. frabAsd from whiek it differs in having three fateral papillae extending inte the caudal alae rather thon two, oval father (han spherival eves and the ubsence of a hinge projecting Hip anterion to the valyva Kry Wokbs: Seweniiaa. nematodes, Seuratidac. Eehinonematinge, Australi, Dasyuridae, marsupials, Introduction Nemmlodes of the furnily Seuratidie are parasites oF reptiles, birds, bats. redents and Australian marsuphils (Chabaud }978) The fumily inelides venerd in Which the mouth is dorsu-ventrally elongated and fimked by paired lips and genera in which (he mouth opening ts triingulir or hexaeonal (Inglis 1967) All three genera oecurring in Australian marsupiils. Sewreehina. Pradechine and Linstewitwned spp. are contained in the subfamily hehinonematinue Inelis, 1967, characterised by a lurve mouth opening with no lip lobes, the anterior end of the body being swollen as a cephalic bulb hearing hooks, no pre-cloucal sucker on the mile and cloacal region covered by cuticular granulitions Although originally placed in the Sehnerder- nematidae by lntlis (1967) the affinities of Linviowinema Smiues, 1997 (lormerly Behinonema Linstow. 1898 preaecupied) with the larvae ol a species of Senratiu Hall, (976 resulted in Quentin (1971) placing the Eehinonermiatinue in the Seuratidie, The genera Linstvwdnemea and lngleettineg Chabaud, Seupeau. Beveridge. Bain & Durette- Desset. 1980, vont species with a triangular mouth Opening on a swollen cephalic bulb beariny hooks. The monatypie genus Sevreching Chabuud, Seureal. Beveridge, Bain & Durette-Desset, [980 however, has neither a tritngelar mouth opening nor a swollen cephalic bulb bearing hooks, although il does have other characteristics of the subfamily. In this paper, Iwo new species of Sevreciine are ' School of Biolowieal ane havevnmenkd Sefenees, Ceara Quesmsbindl Criversily Rocs himpton Ohl 4702. described, The delinition of the sublinmily Hehi- Honematinae js re-evaluated and the yelationships belween the venera discussed, Materials and Methods Nematodes collected from Sriiitliapsin dencapuy ad Anlechinus auilis were fixed ta bot LO%e formalin and then stored in 70. ethanol The preservation history of the specimens from Phetsireete wepoctefit is unknown although they were stared i 7% ethanol. All hematades were examined after clearing in lactophenol. Measurements for more than tour specimens are given in micrometres, as the range followed by the mean in parentheses, and were made with the aid of an ocular micrometer or drawing tube ahd bap Measorer, Drawings were made with the aid Ob ad drawing tube. Type muterial has been deposited in the South Australian Museum. Adekude (SAMA) wid voucher specimens are held in the collection of CSIRO Wildlife und Ecology (CSIRO), Seurechina habbsi sp. nos. (FIGS 1-10) Types? Hulotype cd. allolype 2. paratypes 4 dd. 17 oS) from stomach of Plawecavale tapoutafa (Meyer 1793), Manjimup (34° 15'S. 16) 09" LE) WA, June 1992, coll, S, Rhind, SAMA AHC 31262, ATIC 31263 und ATIC 31264, respectively, Minerial examined: Prom Phiscogale tapectafea fypes. 180 L. Ro. SMALES Figs 1-10. Seureehine hobbsr sp. noy. 1, Anterior end optical section. arrow indicating laminae Uateral view). 2, Cephalic end, optical section (lateral view), 3. Cephalic end, optical section (dorsal view). 4. Cephalic ene, arrows indicating laminae (ateral view). 5. Cephalic end (en fuce view) 6. Male posterior end (ventral view). 7. Body spines. a. From oesophageal region. b. From mid body region. c¢. From posterior body region, 8 Male posterior end (lateral view), 9, Female posterior end (lateral view). LO. Vagiia (lateral view). Seale bars = 200 um ts 10d Ss 100 pov 6.8.9 FO: 50 ui 7:25 pum 2. 3, 4 NEW NEMATODES FROM DASYURID MARSUPIALS 181 Deseripnon Small worms, body with tine transverse cutie kur unnulations, Cephalic extremity without spines, remainder of body with up to 50 rows of spines (nid body of female) at cach annulation: extending over Ye ohody dorsally to caudal ake ventrally of male. over entire body of female: spines beeommnig progressively smaller towards posterior end, Anterior extremity wilh mouth opening and oral cumily, elonsted dorse-veoually. bearine 2) pairs double cephahe papillae, pair aniphids: without lips or lip-like structures, |Sntertar end af vesophagus cupped by 2 oval, dorso-ventrally aligned sclerotised tings, enlamwed dorsally and ventrally. Qesophaztis surrounded at untertar end by 4 pairs hunminue &O lon, Oesophigus simple chivilorm oh. body lenwth, Nerve fing aid seeretory-exeretory pore net sce, deirids linge. eonival. al level-al Sth row oF spines. Mite (i=5 unless olheryine shite) (bigs 1-8) Length 0-70 (0.5 mim), width 300-370 (440), Ocsuphugus 502-569 (536) Jong, Deirtidls 77-94 (85) from anterior end. Spieules equal sine. without whe, 5006630 (590) lone, about i) body Jength. Gubermaculum 50 (n=1) long. Tyo pies lateral pre wloavcul papillag. | pair extending inte lateral alae: | pay lateral ud-cloicul papillie extending inte lateral alice, 3 pais postelotcul papillae, | pair phasnuds well posterior to Glaaua near tail tip, Pettelouwal pupillae not seerr “Tail | 30-170 (150) long. Fenle (v=35 unless otherwise stated) (Pigs 9, 10) Length ZUR U8 vin), width SIO-3580 (550). Oesophiwus 536-670 (910) long, Deiids nov seen, Vaying 550 (n=l) long: vulva opening behind a large projecting Lip, 2600-3450 (4000) from anterior end. Monoudelplig, (gil 215-280 (255) long. Fags spher- ieal 1) 54 (47) diameter. Livniales! The species is named aller Mr R. Hobbs who his been Helpful iy providing material for.this work Kemiarks The method ot fixation used Tor this materith wats not ideal most specimens being contracted and distorted, [Was impossible to determine the number and arrangement of the peri-cloueal papillie on mule speciinets but lateral pre- ad- and) post-cloucal papillae could be seen, Their number and artange- ment are similar to those of the bype and only other species, SL avaneen. Semvehind Hobbst. 6-8 nan Jone. is at kirwer worm thin dS. efazeen. 3.13.8 pin. With longer Spicules (500.630 in 5. febhyt conipured with 185 in & eheneety. The posterior ventral body: spines cover the entire ventral body surliee of mate §. hobby whereas those of & chuneetd terminate in two dateral bands (Chubuud ef af 1980, Fig. Lp p 430), In S Nehbse the lemale tail (215-280) is longer than that ofS. efaneeté (120), the spherical cues are larger (47 diameter compared with 40x35), the vulva iS pre-equaturiitl compared with a post-equatorial vulva in So hancert. Sencechina habhsi 1% monodelphic, whereas 4. chareet/ is didelphic, Seurechina sprattt sp. voy (FIGS 11-19) Types: Voloype 2. atlolype So trom: stomach) of Sminthopsis leucopus (Gray. (842), Sidlings Swamp South, Vimbillica State Purest (47° TBS, 149° 45' Ty, NSW, 25.483, coll. P Hiyeork, SAMA AHC 31265 und AFIC 31266, respectively. Meterial evanied: Prom Snitnihepsis lencapuis: NSW, tpes: From stomach Aavtechinus cagilis Dickman, Parraby. Crowrher & Killa. (998: 20 4 7 Y 2. Sidhinus Swamp North, Timbillrea State Forest. NSW) Law 87. S188, coll PO Haycock and EB. L. Walter, CSIRO N2841, N2977, Dese riplion Simall worms, body with fine transverse cuticular annulitions. Cephahe extremity without spines. reminder ot body with up to 46 rows spines (mid body of fenale) an eaeh aipntilation, extending Over YY hody dorsally ta caudal akre ventnlly, of mle, over enti body of female: spines becoming progressively Soiuller towards posterior end, Ajiterioy extvemity with mouth opening and oral cavity cloned dorsa-ventrally, hearin 2) pains double sub-median cephalic papillae. pair lateral aniphids: without tips or lip-like structures. Anperiot cud Of oesophagus capped hy 2 oval derso-ventrally aligned selerotised rings. enlarsed dorsally vind ventrally, Ousophaus surrounded al anlerior end by 4 pairs laminae 110-165 long, Ovsophagus simple: chayv-ifort, Yee '/je body length. nerve rin aileron lo. detrids. deirids Linge, Comma aitlevel of about ath- 7th row of spines: Seerefory-exerelary pore nol seen, Mole Oneasiwenients of falotype falliwed by measurements of 2 cde fron A. wu7fis) (Plas Fh 12, 14.17. 18) Length 44. 45-6 pimsy width 220. 270-340, Oesuphagus 470. 355-470 long. Nerve ving #3: derrids VEO, RO-T10) fromt antetior end. Spicules equal, sitar without ale, SOQ. 350-000) long, about My body length. Gubernaculiunt fot seen i holotype, SO-38 in specimens trom A. again. Two pairs of hiteral pre-cloacal papillae: | pili extending mtg fateral alae, 2 paws lateral ac-cloacal papillae 182 LR, SMALES AS a) han he Me. Figs }1-19. Searechina sprati sp. noy. 11. Antertor ead, optical section (lateral view), 12. Cephalic end. optical section (lateral view). 13. Anterior end (en face view). 14. Body spines, a. From oesophageal region. b. From mid body region. 15, Cephalic end, optical section (dorsal view). 16, Cephalic end (dorsal view). 17, Male posterior end (lateral view). 18. Male posterior end (ventral view), 19, Vagina (lateral view). Seale bars = 100 pm Th. 19: 50 pm 12, 17, 18; 25 pin 14, 15.16: 10 pm 13. NEW NEMATODES FROM DASYURID MARSUMALS 183 extending lhto Literal ahie, 3) pairs peri-cloucal pupiliie; 3 pairs post-cloweal papillae, 1 pair phasmids well posterior to cloaca near tail tip. Anterior lip oF eloaea with swallem taterul edyes Simulating pair of supplementary papillae. Tail | Ab), {O5-200 long, hemale (neastiienients of allitype tollawed measurements Ob 7 Ly from A, audi) ies ba. 1 16, (9) Lenath 6.6, 6.07.0 nit (6.9), with 470, 300-630 00). Oesophugus G00. 380-570 (485 lomo. Nerve runt, Secretory-excretoey pore, demds not seen, Varia 340, 450 (n=1) long, Vulva 2950, 2975-3485 (3150) trom anterior ead, Monydelphie, Tail 240, 1-240 (2501) hone. Pugs oval 34-53 147) lone by 23-33-(27) wide. ny 3, Eivinelosy The species is named after Dr BoM, Spratt in revo Tion af ls Conte bathed Lo Gur understinding of the hehotinths of dasyurids. Renirks The scurclory-cxereiory pore. oay un coneedled between body spmws close fa the anterior end. is vilen dieult lo defeet in echionernatines. Ta lis species, The anterioremls of all worms from A, aerliy were Contucred, tod ereater or lesser extent during Futon, obscuring the seeretory-exerelory pore. This may have occurred beeduse the heads. of the worms were embedded ii ihe mucosa al post- mortem cxamimnaion (DO Mo Spratt pers. comm. }O98) Meustinanents of oesuphagesl length were Als uiffeeted by the state OF Fixation, those ol lermiles from A. aailis bere apparently slterter dati that of the female Prom S$. feneepius, Other measurements oF specimens Trontthe two hosts were consistent Wilh their belonging tow single species, Sevechind sprit most elosely resembles hohbsi a size, length oF spieates, distribution of body spines, posifion of yvulyi. bene monodelphie and the length of the tail in both the mate and female. AU of these characters distinguish both 8. habbsr and So osprane foin S. cheaneen. Searee hina spradé can be veadily distinguished from both hobhsi ind So chaiect® in having three rither than two fare dateral papillie extending (ito the caudal lace, Searechina sprend lias oval egas whereas those oF S. tabby: ate spherical. Senreefine spear lacks Hie lunge projecting lip anterior to the vulva Pound in &. hobbse, Semechinag chinteecn wis described front eaves allneatis. Gould, {842 from Koolan [shind, off the vost of fort Westerns Australia, 8. bobs? fron P fapodtate (Meyer 1793) front thre southert miintand of Western Anstralia and §. spre tram S- leucopus (Gray, L842) ahd AL eediy Dickman, Parnaby, Crowther & Ring. (998 fron southeastern New South Wales fear the Victorian border ‘The differences between S$. Chance and the other two species may be explained, at lease partly. by acngraphic separalion. The similarities between 4, hobhsi and So spreui would be the resale ot a common ancestor in coastal Vieloria and/or Nesw South Wales. where the ranges of the three host species overlap (Sumner & Dictkmun T4998; Soderquist 1993), Discussion Inglis (1967) created) the subfamily leh Hanematinae to wccommodate the genus Aeliauend (se). and pliced it within the Sehnercernenttiduc. niher than the Seurattdite because OF its long sinple spicules, short gubernaculuiri und a Lriradiate mouth opening without fips. The aftiity of the Beli nonemiulinae with the Seuratidie was diseavered by Quentin (1971) and confirmed by Chabaud. cn al (1980), They linked the presence of a simple, lip less. triraciate mouth opening. two pairs of doubled, subiediah eeplalic papillae. a very shallow vheilostome. and chanacteristie spines an the body culicle oF adults will the Seuralidae, sie ith) harwal development in Seapatua sp. When the genus Senpechiia wis erected by Chabaud ef a/. (1980), these authors diseussed tts lack of Gephalie hooks but placed it in the Eehi nonematinie wlones with the genera, Lonalondrena (formerly Achinonene prenee.) and dieleehine. Which also ocour ih dasyurid hosts. hey did not comment upon the fuct that Senreching has an oval mouth openiny wor upon the signitieatwe of the dorso-ventrally elongated culeahir structure behveen the oesophagus and the mouth opening AL present this chitinaus cup. the walls of which are made up of bwo Superimposed rites (Chabaud ef ef. JOSO). ds NOL Tterpreted as part ala cheiostonre ws defined by Inglis (1967) bul rather as assoeialed with un pesophastome, The four pairs of sublate rid laminae found in the cephalic hypodernis wene howe by Chabaud er af (1880) us appearing To be dilations of the hte! Telds. possibly playing a roke tk mechanism ty hold the eervien! Spies steady when they are embedded jn the gastric mucosa, These structures hive not been found in other echinonemiuatines (Chabaud er uf TY80; States LOOT), For the time being it is eanveniont To renin Seureehine within the Eehinoaematinae until the developmental relationships of the mouth, oese- phygus and assoviated structures have been detenmined, The senus could ether be moved ta 184 L. R. SMALES the Seuratinae on the basis of a bilaterally symmetrical mouth opening, necessitating emendation of the diagnosis of the Echino- nematinae to accommodate adult worms with an oval mouth opening, or, alternatively new groupings could be established. Acknowledgments ] am grateful to R. Hobbs and D. M. Spratt who made available the specimens for this study, and I. Beveridge who criticised an early draft of the manuscript. References CHABAUD, A. G. (1978) Keys to genera of the Superfamilies Cosmocercoidea, Seuratoidea, Heterakoidea and Subuluroidea No. 6 pp. 1-71 Jn Anderson, R. C., Chabaud, A. G. & Willmott, 5. (Eds) “Keys to the nematode parasites of vertebrates” (CAB International, Farnham Royal), . SEUREAU, C., BEVERIDGE, L., BAIN, O. & DURETTE- Desset, M.-C. (1980) Sur les Nematodes Echino- nematinae. Ann. Parasitol. hum. comp. 55, 427-443. INGLIS. W. G, (1967) The relationships of the nematode superfamily Seuratoidea. J. Helminthol. 41, 115-136. Quentin, J.C. (1971) Sur le cycle évolutif de Seuratian cadarachense Desportes, 1947 et ses affinités avec ceux des Nematodes Subulures (Ascaridia) et Rictulaires (Spirurida), Ann. Parasitol. hum. comp, 45, 605-628. SMALLS, L. R. (1997) A revision of the Echinonematinae (Nematoda : Seuratidae) from bandicoots (Marsupialia : Peramelidae). Trans. R. Soc, S. Aust. 121, 1-27. Soperouist, T. (1995) Brush-tailed Phascogale: Pliuss- cogale tapoatafa (Meyer, 1799) pp. 104-106 Jn Strahan, R. (Ed.) “The Mammals of Australia” (Reed Books. Chatswood). SUMNER, J. & DICKMAN, C. R. (1998) Distribution and identity of species in the Antechinus stuartii ~ A, flavipes group (Marsupialia: Dasyuridae) in southeastern Australia. Aust. J, Zool. 46, 27-41. SPIROXYS CHELODINAE BERRY, 1985 (NEMATODA: SPIRUROIDEA) AND CAMALLANUS CHELONIUS BAKER, 1983 (NEMATODA: CAMALLANOIDEA) FROM FRESHWATER TURTLES (PLEURODIRA: CHELIDAE) IN QUEENSLAND, AUSTRALIA By MERYL A. FERGUSON*® & LESLEY R. SMALES* Summary Ferguson, M. A. & Smales, L. R. (1998) Spiroxys chelodinae Berry, 1985 (Nematoda: Spiruroidea) and Camallanus chelonius Baker, 1983 (Nematoda: Camallanoidea) from freshwater turtles (Pleurodira: Chelidae) in Queensland, Australia. Trans. R. Soc. S. Aust. 122(4), 185-189, 30 November, 1998. Spiroxys chelodinae was found in 22 of 77 Emydura krefftii, in three of six areas sampled. This is a new host record. Camallanus chelonius, previously reported only as occurring in the pleurodiran turtle Pelusios sinuatus from South Africa, was found in seven of eight Elseya latisternum, 19 of 77 Emydura kreffti1 and one of one Chelodina expansa from five of six areas sampled, new host and locality records. This finding provides a link between pleurodiran turtles on three continents. Key Words. Spiroxys, Camallanus, nematode, freshwater turtles, Pleurodira, Australia. Treecions of te Royal Seciely af S Navi (1998). T2204), PKS E84, SPIROXYS CHELODINAE BERRY, 1985 (NEMATODA;: SPIRUROIDEA) AND CAMALLANUS CHELONIUS BAKER, 1983 (NEMATODA; CAMALLANOIDEA) FROM FRESHWATER TURTLES (PLEURODIRA: CHELIDAE) IN QUEENSLAND, AUSTRALIA, by Meryl A. PERGUSON' & Lestey R. SMALES' Summary. Percuson, MA & Smares. (Re (1998) Spica ehelodinve Berry. }Y8S (Nematoda: Spiruroideay and Camallanis chelonins Baker, (983 (Nematoda: Camullanoided) fron freshwater turtles (Pleurodira Chelidae) it Queensland, Australia Tas. A bee Ss. Ati. 122(4), 185-189. 30 November. 1998. Spiraxyy cheloedinne was found inn 22 of 77 Enveira Reefs, in three OF Sis areas sampled. This is a new host record, Camadlanus chelonius, previously reported only us oceurring mi the pleurodiran torte Pedisies sinus from South Africa, wits found in seven of ent Elsevier dadiscernmmn Lol 77 Leeyedire hrefffit and one of one Chelodina expanse trom five af six areas sampled. new hoseand tneality records. This finding provides a link benween pleuradinan firtles on three continents, Kie Wonis: Spires, Canrrlfanes, nematode, [reshiwater tirtles. Plearodico, Auatiaha Introduction Two miuyor routes af orgins for the nemtode parasites of reptiles aod amphibians have been suuuested (Baker lO84), Groups with sporadic representatives in ainiphibians and reptiles are considered to tave beer captured from invertebrates or other vertebrates. The majority of nematodes, however, Fave evolved wilh their amphibian and repuilian hosts, The spirurid family Camatlinidae ts an example of the Hest mode of origi This family is suggested to have evolved in fish in tropical Asia. the region with the wreatest diversity and riehest eamallanid: hiune (Strombery & Crites 1974), Boecal morphology suigwests Hal He subdamily Camallaninae, including he venus Cimallanuy Railliel & Henry, 1915S is the tnost reeently evalved ane this is supported by the fier that 43% of Camella species oceur in turtles, frogs ane smukes (Strombery & Crites 1974). In Australis there are flo Species known trom freshwater fish ar turtles, although Serpinenn aclorugatas (Baylis, 1933) Petter, 1979 has been reportest front a eryptodiran turtle. Heavens uranix, in Malaysia (Baylis 1933). The-spirurid family Gnathostomidae, including the monovenerie subfamily Spiros ine. is an example ar the second mode of origin, Le, evelving primarily in imphibians and reptiles (Baker 1984), The genus Spiros Schneider, 1866 probably originated in the ' Sehool ooh Bivlogieut vind Choveeenenil Senices, Cernig Oueenslind Ciniyersity Rock lmplon Olt 4702, holarctic or oriental regions us perssites of non Inaring cheloniins of the Suborder Cry ptodira (Berry LOSS), Spirovys chelodinde Berry, 1985 was first recovered from Cheledina sp. Prony Australia (South Australia. New South Wales. Queensland and Western Australin) and New Guinea | Berry 1985). Estat turtles are vrouped inta Lwe suborders on the basis of differences in ncek vertebrae Mexion during head retraction, AIP Australian turtles are members of the Suborder Pleurodira, a group characterised by sideways Texion al the neck vertebrae, whieh had a cosmopolitan distribution before the separation of Gondwanakine and Lanriesi 120-100 million years ago (mya) (Paugh eral 1990). Theit modern distribution is restricted to the Pelomedusidie of Soath Africa, the Podocnemidie of South America und the Chelidae of Sout America. New Guinea and Australia (Cogper 1996), All remaining freshwater and marine titles are members of the Suborder Cryptodira, a group characterised by vertical Texton ofthe neck, Th this study. §. ehedadinae wis found in Lanvedeere kroffii Gray in Eastern Australia, Camella chelenius Baker, 1983 Is reported for the first time from the pleurociriun turtles Bvivdure krefitit, Blveva latixiernum Gray, Chelodina expansa Gray and ©. loneicallis Shaw from Australia Muterials and Methods A total of 86 turtles, | Chelading expansa & Elseya lufisiermun and 77 Endura Krefiri, Was taken fron 6 Queensland catchments using wt batted crah pot or 86 M.A, FERGUSON & 1. R. SMALES COOKTOWN 15°5 fag CAIRNS @ . TOWNSVILLE @, 20°S ROCKHAMPTON . BUNDABERG #; OS'S Hig. 1 Distiburaan ol the Nemimdes Craellanis Chitty and spires ehelvdtiae (ran Baevdure Arelini Plsewa hvisternin and Claladime expanse tall locales sumpled HW Queenstind from May [9% to Dee WOT B= Camaflinay elretunins, dy = Spiracys echelon fiand line. The lollowing localities were sampled: Cooktown, 1S 28° 8, 145° (So RE (2 EF, letivtermunys Couns 16° 45°58, 145° 47° E C1 A. leatisternuin | Ben Aeeditiy: Townsville 19 16" S. 146° 49° RIS Ene Kveffit, Proserpine 20" 24'S. 148" 35' (Sb. lativieranin, WO Bin, krefftiiy;, Rockhampton 23° 22'S, (50° 32° BCL ©) expanse, 500 Eat Arefftity and Bunduberg 24° 52'S, 152" 2 ECL Ean hrefftii) (Fig, }). "The turtles were cothanased by cervical injection of at least 2.5 ml oof Pentobarbitone sodium (Nembund o) per ke of turtle, Che turtles were dissected then ulboruns were examined with the acl Ol a dissecting microscope, All nematodes found Were washed in saline, fixed in near boiling 10% formalin then stored jn 70% ethanol, Speeiiens from the Queensland Museuin. Brisbane (OM) fron ©. lungicallis, localily unknown (G207571). Tran &. favisternin, Mulgrave River, Glo Atish and Edinontou. Queeushiwad (G2132398, G213239, G213241). and Troi Min, Aresfii, Mulgrave River, Queensland (G213240), were alse examined for comparative pHeposes, Specimens for detailed Microscopic eauminulion were cleared in Juctophenol, All measuremenis are in im unless olherwise indicated vind were made using an eyepiece micrometer. Voueher species, dos Photomicrograph af antecior at Camathanrnes tetas Fron Eaadure krefiiiin Arrow chitnous bueeul nidees, Seale bur = 35 yun, b Nip 2 Tichicales G213990-G 14004. bave heen deposited i the Queenshind Museum, Brishane. Ecological terminology conforms to the definitions of Marsalis ebal, (WY82), Results Adult Apacs efeladie were tecovered Lom granulomas in the stomiuch of 22 of 77 hin, kreffiii examined (28.6% prevalence), but not from other species examined. Examination shawed that the granuiomas originated oi the mucosal layer, but in some cuses had extended into the submucosa une muscularis, and could be seen in the serosa on the extermil surface of the stomach. Eneysted hirvae were also found in the intestinal walls. mesenteries and liver, but numbers were not recorded. Adult Comins chelonius (Pig. 2) were recovered (hom the small intestine of 19 Of 77 Bin. Areffiti (24,7 prevalence), 7 OFS FE, lafisternon (87.3% prevalence) and the single ©. expease examined. OF 77 specimens examined. 4 bi Areffii were tnfectcul NEMATODES FROM AUSTRALIAN TURTLES 1K7 TABLE L. Comparative body measmements, ov um unless otherwise stated, of male and female Spiroxys chelodinae froma Emydura kreffit fro Rockhampron, Queensland und the paratypes of S. chelodinae, (paratype measurements from Berry POSS |. Means in parenthesis. Specmnens from Enryeura kref{tit Paralypes Male Female Male Female n=5 iL=4 n=10 nw=10 Total length Gm) [8-25 (21) [4-27 (22) 73-273 TY-28.6 Masini width 429-516 (457) 374-563 (491) 179-545 171-860 Lenuth pseudolabiam 36-495 (42) 40 (n= 1) 31-65 23-645 Width pseudolabium 83-112 (101) 76 (n= 1) 53-114 hd-136 Length muscular pharyax 435-415 (A80) 408 (n= 1) - - Leneth glandular pharynx 2500-3100 (2883) 2400 tn = 1) - - Length pharynx (mm) 3.26 (n= 1) 261 (n= 1) 1.534-3.61 1,72-3.59 Nerve ring from anterior 536-71 (O09) 4&9 (n= 1) 332-21 435-643 Secretory-excretory pore from anterior YOL-1173 (L037) 424-K 11) AIO-89 | Deirids from anterior (02-1224 (1099) 624-| 334 O44-[312 Leneth gubernaculum 115-168 (142) 108-295 - Length spicule 1230/1630 (1455) 770-2410 - Length tail (74-201 (189) 245-268 (254) 96-281 121-459 TABLE 2, Comparative bouv measurements, tr unt iwiless othenvise stated, of male cud fenuile Camatlanus cheloaius fron Elseya fatistermum fre Queensland ane C. chelonius fron South Africa (measurements from Baker PYs3). Means in parenthesis. Specimens from Specimens from Queensland South Africa Male Female Male Femiale n=5 n=5 n=4 n-4 Total length (mm) 9-14 (12) 13-24 (18) 10.6-16 17.2-23 Maximum width 181-409 (261) 340-516 (388) - - Length buccal valves 101-127 (119) 134-151 (140) - - Width buccal valves 107-147 (121) 134-168 (151) - - Length muscular pharynx 429-502 (474) 489-594 (550) 506.595 518575 Length glandular pharynx 608-1020 ($22) M84- 1054 (949) S3K-YS0 O31 1094 Nerve ring from anterior 231-241 (237) 235-288 (270) 214-251 275-281 Secretory - excretory pore fromanterior = 523 (n= 1) - Deirids from anterior 663-705 (714) Vulva from anterior (mim) < TUS) - h4-12.4 Length right spicule 450-460) (455) - 522-572 - Length left spicule 450-460 (455) - 325-384 - Length tail 100-175 (132) {81-261 (214) 135-175 222-428 with both C. chelonins and 8. chelodinae, 1S were infected with ©. eheloniuy only (19.5% prevalence) and 18 were infected with 8. ehelodindae only (23.4% prevalence), Splrayys chelodinae was tound at Bundaberg, Rockhampton and Townsville only. Camallanus chelonius was found at all localities except Bundaberg, but this is probably because only one host specimen was examined at this locality, Measurements of $8. chelodinae trom this study are in the range reported by Berry (1985) for 8. chelndinae occurring in Chelodina sp. from Western Australia, South Australia, New South Wales. Queenslind and Papua New Guinea (Table 1). The quality of the female specimens was such that few measurements could be taken, but no measurements were outside the range reported by Berry (1985). Specimens of C. cheloninus from this study contorm to the description by Baker (1983) especially regarding buccal yalye morphology and the number and arrangement of caudal papillae. The 18S M.A, FERGUSON & LR. SMALES | Fig, 3, Drawing of tip of right spicule OF ¢eonallares Chelonins trom Emyedtire kref/iii showing differenee trom South African speeunens of Baker (1983), Scule bar = 25 ju measurements OFC, chelonius (Table 2) contorm for the tost parr to these given by Baker (1983) although there are differences in’ spicule length (shorter in Queensland specimens) aod female tail length (shorter in Queensland specimens), Also the slender, sharply pointed clongate process on the tip of the right spieule as figured by Baker (1983, Fig, 1p. 163) appears to be shorter in the South Atrican specimens than the Queensland specimens (Fig 3). However, these differences do not appear to be significant enough to propose a mew species. Discussion Although specimens of Llseya and Laver were exuumined for nematodes by Berry (1985), the sourees ob these hosts Were not given. Spiros cleloddinae wis not recovered by Berry in either host genus: but has now been found in ban Areffir fron Bundabers, Rockhampton and Townsville, The finding ofS. thelodinae in kay kreffii in Central and Northern Queensland is a new host and docality record, This is the first record of ©) chelates from Australiin turtles. Camallanus chelonius has now known been reported from Australian, (this study) and South African (Baker 1984) pleurodiran turtles, The only other report of a camallanid from a pleurodiran turtle ts Serpinema amaconicus (Riberio, 1941) Petter, 1979 from South America (Riberio 1941). All other Serpinemai spp. are found in cryptodiran turtles, with each geographical region haying its own species (Petter 1979). Camallanus spp. have large numbers of unseparated buccal ridges (Fig. 2). while Serpine nie spp. have smaller numbers of distinctly separated buccal ridges. The similarity of spicule morphology and distribution of caudal papillae between C. chelonins and Serporenia spp.. however, indicates i close relitionship (Baker 1983). Also, the buccal morphology of S. aaacenicus und C. chelonius has been suggested to be intermediate between Case- Hanus and Serpinema. This suggests that Sespinentia may have evolved first in pleurodirans (Baker 1983), A camallanid ancestor of S$. amezenicus may have been captured by early eryptodiran turtles and radiated with the major cryptodiran radiation around 120-90 mya (Shafter et af. 1997). The geographical distribution of Serpineme in cryptodiran turtles shows that the greatest diversity of fauna ts in South and Central America and tropical Asia (Stromberg & Crites 1974). The finding of §. ammezenieus in South America. an intermediary form between Camallanus and Serpineme. suggests that South America may haye been a point of origin for this genus. which then moved into North America and Asia with its hosts. The oecurrence of C. chelonifay in both Australian and South African pleurodires suggests that this species originated before the separation of Australi and South Africa, approximately 120-lOUmya. The close link between South Africa and South Amerion us recently as 90 mya und the similaritics between chelonius, S amezenicous and other Serpinemea spp SuBBeSsL i COMMON Origin for these groups, Acknowledgments Thanks go to C.J. Parmenter for assistance in turtle collection and jdentifieation, and to L. Cannon and K. Sewell of the Queensland Museum for providing access ta Museum Specimens. NEMATODES FROM AUSTRALIAN TURTLES 189 References Baker, M. R. (1983) Nematode parasites of the turtle Pelusios. sinuatus (Pelomedusidae: Pleurodira) from southern Africa. Syst. Parasitol, 5, 161-167. (1984) Nematode parasitism in amphibians and reptiles. Can. J. Zool. 62, 747-757. Berry, G. N, (1985) A new species of the genus Spiroxys (Nematoda; Spiruroidea) from Australian chelonians of the genus Chelodina (Chelidae). Syst. Parasitol. 7, 59-68. Bays, H. A. (1933) On a collection of nematodes from Malayan reptiles. Ann. Mag. Nat. Hist., Ser. 10 11, 615- 633. CoaGcer, H. G. (1996) “Reptiles and amphibians of Australia” (Reed Books, Melbourne). Maraous, L. G., Escu, G. W., HOLMES, J. C., Kurs, A.M. & ScnaAb, G, A. (1982) The use of ecological terms in parasitology. J, Parasitol, 68, 131-133. Perrer, A. J. (1979) Essai de classification de la sous-famille des Camallaninae (Nematoda, Camallanidae). Bull. Mus. natn. Hist. nat., Paris, 4 sér, Sect. A 1, 991-1008. PouGH, F. H.. HkISER, J. B. & MCFARLAND, W. N. (1990) “Vertebrate Life’ (Macmillan Publishing Co.. New York). Riperto, D. J. (1941) Pesquisas helmintologicas realisadas no Estado do Para. VIL - Camallanus amazonicus n. sp. parasito de Podocnemis expansa (Schw.). Mem. Inst. Oswaldo Cruz 35, 723-727. SHAPFER, H. B., MEYLAN, P. & McKnicut, M. L. (1997) Tests of turtle phylogeny: molecular, morphological and paleontological approaches. Syst. Biol. 46, 235-268. STROMBERG, P. C. & Crites. J. L. (1974) Specialisation, body volume and geographical distribution of Camallanidae (Nematoda). Syst. Zool, 23, 189-201. ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED Patron: HIS EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA SIR ERIC NEAL, AC, CVO OFFICERS FOR 1998-99 President: M. A. J. WILLIAMS, BA(Hons), MA, PhD, ScD Vice-Presidents: T. C. R. WHITE, BSc, BSc(For), PhD N. F ALLEY, BA(Hons), MA, PhD Secretary: Treasurer: O. W. WIEBKIN, BSc, PhD J. H. BRADBURY, BSc, MSc Editor: Assistant Editor: J. BIRD, BSc N. F, ALLEY, BA(Hons), MA, PhD Librarian: Programme Secretary: S. BARKER, BSc(Hons), PhD Minutes Secretary: Membership Secretary: C. R. WILLIAMS, BSc(Hons) A. J. McARTHUR, BE Members of Council: P, KOLESIK, BSc, PhD A. F. BIRD, BSc, MSc, PhD, DSc J. E. PATTISON, MA, BSc, MSc, Grad Cert Ed M. J. 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