VOL. 117, PARTS 1 & 2 4 JUNE, 1993 Transactions of the Royal Society of South Australia Incorporated Contents Hutchings, P. A., Ward, T. J., Waterhouse, J. H. & Walker, L. Infauna of marine sediments and seagrass beds of Cpr: Spencer Gulf near Port Pirie, South Australia - - - - z : £ : 4 Barker, S. Seventeen new species of Australian Buprestidae (Insecta: Coleoptera) and a host plant of Castiarina uptoni (Barker)- - - - - - Beveridge, I. & Durette-Desset, M.-C., Adult and larval stages of Paraustrostrongylus ratti (Nematoda: Trichostrongyloidea) from Rattus fuscipes Campbell, R. A. & Beveridge, I. New species of Grillotia and Pseudogrillotia (Cestoda: Trypanorhyncha) from Australian sharks, and definition of the ey Grillotiidae Dollfus, 1969 - - - - E : 2 Baker, G. H., Barrett, V. J., Grey-Gardner, R. & Buckerfield, J. C. Abundance of life history of native and introduced earthworms (Annelida: Megascolecidae and Lumbricidae) in pasture soils in the Mount baie Ranges, South Australia - - - - - - - Gatehouse, C. G., Jago, J. B., Clough, B. J. & McCulloch, A. J. The Early Cambrian volcanics from Red Creek, eastern Mount Lofty Ranges, South Australia Austin, A. D., White, T, C. R., Maelzer, D. A. & Taylor, D. G. Biology of Etiella behrii Zeller (Lepidoptera: Pyralidae): a pest of seed lucerne in South Australia Lee, D. C. Incabates Hammer and Setincabates gen. nov. (Acarida: Cryptostigmata: Haplozetidae) from South Australian soils - - = i : Sprigg, M. & Bone, Y. Bryozoa in Coorong - type lagoons, Southern Australia Wells, A. & Cartwright, D. Females and immatures of the Australian Caddisfly Hyalopsyche disjuncta Neboiss (Trichoptera), and a new family placement- - Brief Communications: Johnston, G. R. & Richards, S. J. Observations on the breeding biology of a taierenyh frog (Genus Oreophryne) from New Guinea- - - Crossland, M. R. & Richards, S. J. The tadpole of the Australopapuan frog L Litoria nigrofrenata (Gunther, 1867) (Anura: Hylidae) - - PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 27 37 47 57 97 105 109 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 117, PART 1 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INC. CONTENTS, VOL. 117, 1993 PARTS 1 & 2, 4 JUNE, 1993 Hutchings, P. A., Ward, T. J., Waterhouse, J. H. & Walker, L. Infauna of marine sediments and seagrass beds of Upper Spencer Gulf near Port Pirie, South Australia Barker, S. Seventeen new species of Australian Buprestidae (Insecta: Coleoptera) and a host plant of Castiarina uptoni (Barker) - - - - - - - Beveridge, I. & Durette-Desset, M.-C. Adult and larval stages of Paraustrostrongylus ratti (Nematoda: Trichostrongyloidea) from Rattus fuscipes - - - Campbell, R. A. & Beveridge, I. New species of Grillotia and Pseudogrillotia (Cestoda: Trypanorhyncha) from Australian sharks, and definition of the family | Grillotiidae Dollfus, 1969 - - - - - - - - Baker, G. H., Barrett, V. J., Grey-Gardner, R. & Buckerfield, J. C. Abundance and life history of native and introduced earthworms (Annelida: Megascolecidae and Lumbricidae) in pasture soils in the Mount Lofty Ranges, South Australia - - - - - - - - - - - Gatehouse, C. G., Jago, J. B., Clough, B. J. & McCulloch, A. J. The Early Cambrian volcanics from Red Creek, eastern Mount Lofty Ranges, South Australia Austin, A. D., White, T. C. R., Maelzer, D. A. & Taylor, D. G. Biology of Etiella behrii Zeller (Lepidoptera: Pyralidae): a pest of seed lucerne in South Australia Lee, D. C. Incabates Hammer and Setincabates gen. nov. (Acarida: Cryptostigmata: Haplozetidae) from South Australian soils - - - - - Sprigg, M. & Bone, Y. Bryozoa in Coorong - type lagoons, Southern Australia - - Wells, A. & Cartwright, D. Females and immatures of the Australian caddisfly Ayalopsyche disjuncta Neboiss (Trichoptera), and a new family placement - - Brief Communications: Johnston, G. R. & Richards, S. J. Observations on the breeding biology of a a ti frog (Genus Oreophryne) from New Guinea - - Crossland, M. R. & Richards, S. J. The tadpole of the Australopapuan frog L Litoria nigrofrenata (Gunther, 1867) (Anura: Hylidae) - - - 27 37 47 57 67 77 87 97 105 109 PARTS 3 & 4, 30 NOVEMBER. 1993 Shiel, R. J. & Koste, W. Rotifera from Australian inland waters. IX. Gastropodidae, Synchaetidae, Asplanchnidae (Rotifera: Monogononta) - - - Tian-Rui, Lin & Jago, J. B. Xystridura and other early Middle Cambrian trilobites from Yaxian, Hainan Province, China — - - - - - - - Pell, S. D., McKirdy, D. M., Jansyn, J. & Jenkins, R. J. F. Ediacaran carbon isotope stratigraphy of South Australia — an initial study — - - Nicholas, W. L. Two new species of nematode (Nematoda: Enoplida: Thoracostomopsidae) from Lake Alexandrina, South Australia - - - - - - Hoste, H. & Beveridge, I. Interspecific and intergeneric relations between nematodes parasitic in the stomachs of kangaroos and wallabies —- - - - _ Ryder, M. H. & Bird, A. F. Effect of Acrobeloides nanus (Nematoda: Cephalobidae) upon the survival of Pseudomonas corrugata (Rubattetia) in ey pasteuy ae soil from Kapunda, South Australia < > : + Zbik, M. The Big Rock Donga Metcorite: A new H5 Chondrite from South Australia Tyler, M. J. & Godthelp, H. A new species of Lechriodus Boulenger (Anuras Leeptodicty lide) from the Early Eocene of Queensland — - - Barker, S. A new Australian species of Calodema (Coleoptera: Buprestidae) - - - Brief Communication: Wallman, J. F. First South Australian record of the carrion-breeding blowfly Calliphora nigrithorax Malloch (Diptera: Calliphoridae) — - - - - - Errata, Corrigenda, Addenda - - - - . ¢ + 2 3 * - insert to Transuctions of the Royal Society of South Australia, Vol. H?, parts 3 & 4, 30 November, 1993 171 179 183 187 191 193 194 INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF NEAR PORT PIRIE, SOUTH AUSTRALIA BY P. A. HUTCHINGS*, T. J. WARD**, J. H. WATERHOUSE** & L. WALKER*** Summary The infauna of six intertidal and shallow subtidal habitats were sampled in summer and winter near Port Pirie, Upper Spencer Gulf. The habitats included seagrasses and non-vegetated sediments. Over 12,000 individuals belonging to 372 taxa were collected in the survey. The species composition of this fauna had very little overlap (only 24 species) with that of a more recent survey of Northern Spencer Gulf. The density of the fauna in our study area was least (280 m‘~) in the unvegetated intertidal habitat, and highest (2612 m’°) in the intertidal seagrass habitat (Zostera), and was dominated by polychaetes, molluscs and crustaceans. The fauna of the Zostera and Posidonia habitats is compared with other similar habitats sampled in southern Australia, in terms of the number of species, density of individuals and feeding guilds of the fauna. Differences in species diversity and densities between this study and other studies are likely to reflect the intensity of sampling, as well as differences between locations in the density and species of seagrass and in the prevailing hydrological regimes. KEY WORDS: Posidonia, Zostera, seagrass beds, unvegetated sediments, infauna. Transactions of the Royal Sociery of S Ausz., (1993), LIT(), 1-45 INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF NEAR PORT PIRIE, SOUTH AUSTRALIA by BR A. HUTCHINGS* T. J. WARD**, J. H. WATERHOUSE** & L. WALKER*** Sumumury Hurcutnas, P_ A., WARD, T, J,, WATERHOUSE, J. H., & WALKER, L. 1993) Infauna of marine sediments and seagrass beds of Upper Spencer Gulf neur Port Pirie, South Australin. Trans. R. Soc. $. Ausr. Wy, 1-15, 4 uns, 1993. The infauna of six intertidal and shallow subtidal habitats were sampled in sammer and winter near Port Pirie, Upper Spencer Gulf, The habitats included seagrasses and non-vegetated sediments, Over 12,000 individuals belonging to 372 taxa were collected in the survey. The species composition of this fuuna had very litile overlap (only 24 species) with that of w more recent survey of Northern Spencer Gulf. The density of the tauna in our study arca was least (280 m) in the unvégetated intertidal habitat, and highest (2612 ny) in the intertidal seagrass habitat (Zastera),.and was dominated by polychaetes, molluscs and crustaceans. The fauna of the Zostera and Posidonia habitats is compared with other similar habitats sampled in southern Australia, in terms of the number of species, density of individuals and feeding guilds of the fauna. Differences in species diversity and densities between this stady and other studies are likely to reflect the incensity of sampling, as well as differences between facunons in ihe density and specios of seagrass and in the prevailing hydrological regimes. Key Worps: Fosidonia, Zostera, seagrass beds, unvegetated sediments, infauna. Introduction Upper Spencer Gulf near Port Pirie (33°19'S, 138°E), South Australia, is shallow (penenilly less than 20 m deep) and fringed with large expanses of intertidal mudflats, intertidal seagrasses (Zostera spp.), and subtidal beds of the scagrasses Pasudona australis and P sinwosa, These shallow expanses of seabed are fished commercially and recrestionally for a number of species of fish, including garfish and whiting, and for crustaceans including prawns and crabs (Jones 1979'; King 19777). Although these. and many other species, use the seagrass beds and intertidal flares ar various stages of their life cycle Jones 1979'), there have been few ecological studies designed to document other species of importance occurring in these habitats of the Gulf. Upper Spencer Gulf refers to the arca of the Gulf between Port Davis Creek (just south of Port Pirie) *« tnvertebrate Zoology, The Australian Museum, P.O, Bon A285, Sydney South, NSW 2000 ** CSIRO Division of Fisheries, P.O. Box 20,. North Beach, WA 6020 ‘e* jormerly of The Australian Museum now of 306, Moleswurth St,, Lismore, NSW 2480 WONES, G. I. (1979) Biological Investigations of the Marine Scale Fishery in Spencer Gulf. (Dept Agriculture and Fisheries, South Australia). 72pp. “KING, M. G. (1977) The biology of the Western King Prawit Fengeus latisulcaius Kishinouye and aspects of the fishery in South Australia. MSc. thesis, University of Adelaide, unpubl and Ward Spit, and Northern Spencer Gul! is defined as the portion of Spencer Gulf north of Ward Spit, 1.c. north of 33°S (Stefanson 1977). Spencer Gull, is hypersaline, with salinities gradually increasing from 35 “/ nto more than 60 °/ oo near Port Augusta at the head of the Gulf (Bullock 1975), Upper Spencet Gulf has an intermediate salinity regime, varying annually From about 41-47.2 °°... and water temperatures ringing from 10-26.2°C (Dow 1980°). Studies of fish and crustaceans in this area by Ward & Young (1982) have shown that they appear to be unaffected by the constant high salinines. In this study we examine the mainly invertebrate fauna Lying in the sediments in shallow-water habitats found in Upper Spencer Gull near Port Pirie, and its species composition. These results are compared with those obtained from sitmilur habitats elsewhere in southem Australia Similar hatitats which have been studied in detail elsewhere in southern Australia are those of seagrass beds and intertidal habitats. The infauna of seagrass beds has been examined in several studies in eastern Australia (Collett et af. 1984; Hutchings & Recher 1974), und in Western Australia (Hutchings et al. 1991, Wells et al. 1985) and reviewed by Howard es al, (1989), The infauna of intertidal habitats in Northern Spencer Gulf has recently been studied by Ainslie ev al, (1989), DOW (1980) Redcliff Project — Deaf Environmental Effects Statement. 178 pp. (Dow Chemical (Australia) Limited, Adelaide SA,}. te Materials and Methods Samples of sediment were collected at high tide from 18 sites representing six habitats, by divers using PVC corers (220 mm diameter and 160 mm depth) in winter (25Nii.1979 — 17.xiii.1979) and in summer (1.ii,1980 — 1.ii1.1980). Each corer had a sharpened leading edge and was capped at the other end by a PVC plate welded in place, A 50 mm hole in the cap covered with 1 mm fibreglass. mesh allowed water to escape as the corer was driven into the sediment, and, after coring, was sealed with a rubber bung. No attempt was made to avoid seagrass leaves when they occurred at a sampling =—— ¢ “= — P. A. HUTCHINGS, T, J. WARD, J. H, WATERHOUSE & L. WALKER site, and the sampling technique resulted in substantial amounts of live and dead seagrass leaf material, rhizomes and roots in the samples. The habitats sampled (Fig. 1) were from about mid-tidal level (on mudflats adjacent to the mangroves) to sediments in 10 m water depth, and are the same habitats in which detailed studies of sediments and epibenthic fauna have been carried out previously (Ward & Young 1981, 1982). The habitats included bare intertidal muds (Sites 1A, 1B, 1C), intertidal seagrass beds of Zostera mucronata (Sites 2A, 2B, 2C), subtidal beds of Posidonia australis (Sites 3A, 3B, 3C) and. Posidonia sinuosa (Sites. 4A, 4B, 4C) and bare unvegetated Ward Spit _.- SPENCER GULF Fig. 1. The Port Pirie study area in Upper Spencer Gulf. mangroves: Port = “Yy~pPirie smelter af River a PORT PIRIE INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF 3 sediments (Sites 5A, 5B, 5C and Sites 6A, 6B, 6C) in deeper water (Fig. 1). At each site three samples were collected haphazardly from each of three plots of about 4 m? separated by at least 5 m. This resulted in nine samples from each of the 18 sites, collected in winter and summer, except for summer, when one sample from the Posidonia australis habitat was lost. The samples were sieved in the field through 2 mm mesh and all residues were preserved in 10% formalin. In the laboratory the samples were sorted under water in large flat trays with the aid of low-power magnification. Hermit crabs were discarded because of the difficulty of extracting them from their shells, but all other non-colonial fauna was collected. Colonial organisms such as bryozoans and hydroids attached to the blades of the seagrasses were not sampled in this study, Because of taxonomic difficulties and a lack of funds the collection was stored for six years until resources permitted a detailed evaluation. The collection has been deposited at the Australian Museum, Sydney. Results and Discussion Species Composition of the Fauna The samples contained 12,396 individuals representing 372 taxa. These taxa consisted of 39% polychaetes, 26% molluscs, 22% crustaceans, and 6% echinoderms, with the remaining 7% being distributed among the minor phyla (Table 1). The distribution and abundance. of all taxa within the six habitats sampled is shown in Appendix 1. Of the 26 dominant taxa. defined as those represented by 100 or more individuals in the collection, 18 were polychaetes, four were molluscs, three were crustaceans and one was a holothurian (Table 2). The taxa identified as nemerteans and sipunculans may not be monospecific because of difficulties in identifying individuals to the species level. The most abundant species was Tanais dulongi, a tanaid crustacean. The six habitats sampled varied considerably in the number of species present. The four subtidal habitats all had a high numbers of species (173 — 193) compared to the intertidal Zostera habitat (116), and TaBie 1. Infauna of Upper Spencer Gulf! Numbers of species (5), density (d, number of individuals m°*), abundance (n, % of toral individuals in habitat) and total abundance (N) of the major taxa in each of the six habitat types sampled. All habitats were sampled with 54 cores (total surface area of 2,05 m*), except Posidonia australis which was sampled with 53 (total surface area of 2.01 nx’). taxa intertidal Zostera P. australis polychaetes Rn 57 94 d 243 1262 884 n(%) 87 48 76 crustaceans Ss 5 20 34 a 16 907 120 t1(%) 6 35 10 molluscs s 6 23 34 d- 13 330 §4 n(%) 5 B 7 ascidians S 0 4 3 d - ul 6 n(%) = 05 0.5 echinoderms 8 1 3 » d 0.5 39 37 n( 9%) 0.2 2 3 minor taxa* 3 6 d 8 62 27 ni %) 3 2 2 all taxa 26 15 Lil N S74 5362 2329 d 280 2612 1157 habitat P. sinuosa 5m bare Win bare total 9o7 1 88 148 630 501 242 626 73 76 49 62 44 23 18 BI 76 28 19 nH 9 4 4 i] 32 38 50 95 95 92 183 133 iT] 14 37 b 5 7 2 10 i 16 it 9 L 2 2 1 12 Q 10 20 38 14 32 za 4 2 7 3 4 3 4 18 ul 3 0 2 1 1 2 2 194 171 172 372 1764 Bs) 1017 {2396 860 660 497 1010 * Minor taxa includes fish, insects, sipunculans, poriferans, nemerteans, pycnuyonids, enteropneusts and hirudineans, 4 P A. HUTCHINGS, T. J. WARD, J. H. WATERHOUSE & L. WALKER TABLE 2. Abundances (%) of the dominant species — total abundances of 100 or greater — in each of the the six habitat types. ee ee a ee SE intertidal Zostera P. australis Tanais 26 (3) 819 (97) - dulongii Capitella 51 (8) 6l1 (92) - ‘‘capitata”’ Nereis - 31 (5) 577 (89) bifida Apseudes - 513 (87) 63 (H) dustralis Leitoscoloplos 31 (5) 414 (71) 121 (2)) Sp. Nephtys 109 (20) 99 (18) 13 (2) gravieri Ceratonereis 248 (65) 8 (2) 34 (9) sp. | Pista - 340 (91) 6 (2) violacea Platynympha 2 (0.6) 291 (90) 22. (7) longicaudaia Barunwlla sp, = 88 (31) 57 (20) Austromytilus - 45 (2) 2 (0.8) penectus Nematonereis - - 103 (41) unicornis Augeneria - 131 (55) 18 (8) verdis Tellina - 226 (95) - deltoidalis Zeacumantis 1 (0.5) 206 (99) - diemenensis Eunice sp. 3 - - Al (21) Notomastus 2 (lh 96 (52) sp. J Tellina 6 (3) TL (41) 20 (12) mariae Leptosynapta = 74 (49) 42 (28) dolabrifera Agyychis sp. - - 5 (4) Cirriformia - 43 (34) 43 (34) sp. 1 Caulleriella - 20 (18) 31 (27) sp. 4 Eunice sp. 1 2 - 5 (5) Diplocirrus sp. - 92 (90) 3 (3) Scoluplos 1 (i) 74 (73) 17 (17) simplex cas Pere ~ 7 (7) 35 (35) Sp. totals 475 (6) 4165 (52) 1354 (17) total abundance (all species) 574 5362 2329 % abundance dominant/abundance total ‘ 5 P sinuosa 3m bare 10m bare total - - 1 01) 846 - - = 662 29 (5) 70) 1 (0,2) 645 2 (0,3) 7) 3 (0.5) 588 13 (2) 1 (0.2) - 580 U5 (21) 149 (27) 67 (12) 552 14 (4) 69 (18) 10 (3) 383 12 (3) 3 (0.8) 12 (3) 373 8 (2) - - 323 88 (31) 46 (16) 6 (2) 285 2 (0.8) 57 (21) 211 (76) 277 18 (47) 25 (10) 7 (3) 253 49 (20) 26 (11) 16 (7) 240 8 (3) 5 (2) - 239 = 7 - 207 an 2548) B®) 45 (26) [2 (7) 18 (10) 172 33 (22) - I (0.7) 150 wey 4 38 (34) 21 (19) 3 (3) 13 81 (74) 19 (17) 4 (4) 109 4 43 (43) 14 (14) 1 (1) 100 859 (H) 661 (8) 439 (6) 7953 1764 1350 1017 12396 49 49 43 64 aE ee eee ee the intertidal mud, which was by far the most species- poor habitat with only 27 species recorded (Table 1). The pattern of densities amongst the six habitats was generally similar to that of species richness except for the Zostera habitat. This habitat had the highest density of fauna (2612 individuals ny?) with the second lowest species richness (115 spp.). Overall, polychaetes and molluscs were the two dominant groups in terms of both numbers of species and individuals, The crustaceans were generally similar to the molluscs in species richness and abundance in ithe two Posidonia habitats and the intertidal mud, but were considerably reduced in the two subtidal unvegetated habitats. Echinoderms, ascidians and the INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF LIPPER SPENCER GULF 4 minor taxa were represented by only Jew species and typically few individuals in all habitats. Of the 32,396 individuals collected 7,953 (64%) belonged to only 26 (7%) of the 375 species. The remaining 346 species were represented in most cases by only a few individuals (Appendix 1). Ninety-four species Were Tepresented by only a single individual in the collection and 155 species were present in only One babital. Similar patterns have been found in other studies of the infauna of Austratian seagrass beds {Collett er al, 1984; Hutchings et al. 1991). The 26 dominant species were distributed unevenly between the six habitats (Table 2). The two most dontinant species (Timwis dulongi and Capitella \‘capitata”’) were almost exclusively restricted to the two ineertidal habits, and predominantly in the Zastera. Several other species also occurred predominantly in the Zostera habitat — Apseudes gustralis, Leitoscoloplos wormalis, Pista violacea, Playympka fongicwidata, Tellina delroidatis, Zeacumantus diemdénensis, Diplocirrus sp. and Scolaplos siniplex. Nereis bifida and Eunice sp. | octurred oaainly in the PB australis and P sinuese habitats respectively, while Asychis sp. was mainly found in the Sm subtidal bare habitat. Only four of the deminan! species (Nepthys graveri, Ceratonerets sp, J, Tellina mariag, and. Scotoplos simplex) were present in fll six habitats. Tweaty of the 26 dominant were foutd in all the subtidal habitats. Thus the dominant species could be separated into two groups: those dominant in the intertidal and those dominant in the sabtidal, with few species common to both groups, A detailed analysis af the variability of species composition amongst individual habitats, sites, seasons and sediment characteristics will be presented elsewhere (Ward & Hutchings in prep.) Comparisons with other areas Intertidal unvegeiated sediments — A recent study by Ainslie et al, (1989) of the mfauna of intertidal unvegetated inudflats of Norther Spencer Gulf (about 60 km north of our suidy area) recorded a total af 99 species, These 99 species. included 31 species of polychaetes, 31 molluscs, and 22 crustaceans. This is considerably greater than the total of 27 species which were recorded from intertidal unvegetated habitats in this study. The densities of individuals recorded in the Northern Spencer Gulf intertidal habitats (about 550 in*} was also considerably greater than those recorded in this study (280 m*). These differences may result from the different scales of sampling in the two studies or to real differences between the. locations related to differences in the hydrological conditions between the two Spencer Gulf jocations; Similar sampling techniques were used in both studies although Ainslie etal, used a} sms sized mesh, and 20 replicates were collected af each site in summer and winter from Jane 1982 to June (986 inclusive, which is a far greater intensity of sampling than carried out in this study. The Ainslie et al. (1989) study saanpled a total area of 33 m? in Northern Spencer Gulf whereas we sampled about 2 m2. This difference in total atea sampled and the extended temporal scale of the Northern Spencer Gulf study may have contributed to the grealér number of species of infauna recorded. Alternatively, the Gulf near Port Pirie may have a depauperate incertidal infauna because of different salinity and temperature regimes, The species composition of the infauna found by Ainslie et al. (1989) in Northern Spencer Gulf ts not similar to that fund in this study, with very few species being common to both areas. Ainslie et al, identified a total of 100 taxa to species level, and in this study we identified 178 taxa to species. Of these only 24 species are common to both Upper and Northern Spencer Gulf. Obviously, some of the taxa ientified only to genus may, after further study, be found to occur in both regions. We suggest that this apparent difference in species composition may be related to the different hydrological regimes in these two regions of Spencer Gulf. Northern Spencer Gulf ts characterised by high summer salinities, up to 48 Soo (Nunes 4 Lennon 1986), and high summer teniperatures with wide seasonal Nuctuatlons LL — 25°C Yohnsen 1981*; unpublished data quoted by Ainslie et al. 1989). Northern Spencer Gulf is seperated from the Upper Gulf by a narrow channel just porth of our study area, and this could be responsible for maintaining a different set of hydrological conditions between these two study areas, ever though they are close to cach other Posidovia australis seagrass beds The low-intertidal seagrass habitat of Posidonia australis and P. sinuosa, in Northern Spencer Gulf, contained 147 species of infauma with a mean density af bout 170 individuals mt* (Ainslie er al. 1989), The fost similar habitat in our study (P aussreis) contained! 18) species with an average density of 57 individuals m*. Ainslie ef afl. do not give any indication of the density of seagrasses ii this habitat, but, because Aozidania is basically a suftidal seagrass species, we presume that thelt sample location comamed only scattered seagrass on the trrargins of more subsiantial subtidal beds of P australis, P. australis seagrass beds elsewhere In southern Australia have also been studied, In Table 3 the distritutions of the major phyla present in P. australis TJOHNSON, LE, (981) Hydrological dats from U Spencer Gulf 1975-1978 Fisheries Res. Paper No, 3; 0 (Dept Fistieries, Adelaide, SA, unpubl.) 6 P. A. HUTCHINGS, 'T, J. WARD, J. H. WATERHOUSE & L. WALKER TABLE 3. Posidonia australis: distribution of the major phyla (§ = number of species, d = overall mean density). site Polychaetes Molluscs Crustaceans Echinoderms Ascidians Minor Total Albany (Hutchings ef al. 1991) s 47 26 bt a iL 4 113 a** 73) 691 656 B32 124 ~ 2327 Spencer Gulf (this study) 8 94 34 a4 10 3 6 J81 u** 883 84 9 37 6 27 56 NSW (Collen et al, 1984) $ 134 1 106 # ? 32 323 a‘ 1656 285 lf) 13 2763 * amphipods not identified to species # echinoderms and ascidiany included in minor ™ densities (number of individuals Gulf = 2.014 m*, NSW = 5.74 m°*) in Albany, SW West Australia (35°03‘27"S 117°03° 27"W) (Hutchings er al. 1991) and at nine sites along the NSW coast from 32°13'S to 36°54'S (Collett et al. 1984) are compared with those occurring in Upper Spencer Gulf. Polychaetes dominated the infauna in these three geographical localities, both in terms of number of individuals and species, The average densily of infauna in NSW Posidonia beds was twice that of Upper Spencer Gulf, although the NSW data were based on samples totalling 5:74 m? in area, collected at nine locations along the NSW coast. Similarly, the P. australis sampled in Albany (total area sampled 1.2 m*) had a mean density of infauna about twice that found in Upper Spencer Gulf in this study. The species composition of the infauina of P. australis in (hese three geographical areas is different, indicating that although P australis has a wide distribution, occurring from Shark Bay in Western Australia around southern Australia to Wallis Lake on the central const Taste 4, Posidonia australis: distribution of species (%)). location herbivore suspension Albany (from Hutchings ef al., 1991) polychaetes 4 (9) 3.(7) molluscs 7 (33) 7 (33) cmistaceans - - uscidians . ut (100) echinoderms - 2 (iS) totals fi (hy 23 (22) Spencer Gulf (this study) polychaetes u (12) 12 (13) molluscs 15 (44) 13 (38) crustaceans I (3) 1 (3) as¢idians - 3 (100) echinoderms 1 (10) 1 (0) totals 27 (16) HW) (17) NSW (from Collett ef al., 1984) polychaetes - 28 (21) molluscs 10. (19) 24 (47) crustaceans 3@) (4) tonal B (5) 56 (19) phy it mm) calculated from total abundances and total area sampled (Albany = 1.2 m2, Spencer of NSW (Larkum & den Hantog 1989), the composition and density of its infauna vary according to location, The infauna may also be related to the density of the seagrasses themselves, but at the Upper Spencer Gulf and Albany sites the density and productivity of the P australis beds are similar (Watd 1987; Walker et al. 1991). Collett er al. (1984) report only a qualitative assessment of the density of seagrass blades which cannot be related to the numeric values determined for the Upper Spencer Gulf and Albany sites.. Therefore, hecause of both different sampling. intensities and seagrass densities, it is not possible to directly compare the density or diversity of the infauna found by each of these studies. In this study we also examined whether the feeding stmitegies employed by the infauna were similar regardless of geographical location. The fauna was assigned to one of five feeding categories (Table 4) for Upper Spencer Gulf, Albany and the nine locations of the fauna amongst feeding vategories at Albany, Spencer Gulf and NSW (number deposit carnivore omnivore 20 (44) 18 (40) - 1 (5) 6 (29) - 5 (36) 9 (64) - 4 G1) 5 (38) 2 (15) 30 (29) 40 (38) 2 (2) 40 (43) 30 (32) 1 (1) 2 (6) 4 (12) - 19 (56) 13. (38) 3 30) 5 (50) : 64 (37) 52 (30) 1 (1) 53 (39) 37 (28) ¥ (12) 7 (Id) 4 (8) 1 (2) 28 (26) 20 (19) 44 (41) ‘88 (30) 61 (21) 61 (21) INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF 7 along the NSW coast, The designation of feeding categories was determined from the literature (mostly for related species) and from personal observations, Camivores, including scavengers, comprised 21 — 38% of the total infauna in a geographical region. The proportion of deposit feeders tanged between 29% (Albany) to 37% (Upper Spencer Gulf). Suspension feeders, which may feed on suspended particulate ™matier as well as plankton, were between 17 and 22% of the fauna (Table 4). There were major differences in the herbivore and omnivore fauna between locations, with ranges of 5% (NSW) to 16% (Upper Spencer Gulf), and 1% (Upper Spencer’ Gulf) to 21% (NSW) respectively. These data suggest that the distribution of feeding guilds varies according to geographical location, Perhaps future studies should consider the feeding category of fauna as well as their abundance, productivity and biomass in order to estimate the ecological importance of a species in shallow marine systems, Edgar (1990) used size structure of benthic macrofaunal communities to estimate faunal biomass and secondary productivity but he did not assign the fauna to feeding guilds which may be an additional useful qualitative descriptor for benthic communities. Posidonia’ sinuosa seagrass beds P. sinuosa occurs from Shark Bay to the Great Australian Bight (Larkum & den Hartog 1989). However, the infauna of Posidonia sinuosa has only been sampled at one other location in southern Australia -— Albany in SW Western Australia (Hutchings et al. 1991). The density of the Albany infauna was twice that recorded from Upper Spencer Gulf in this study. This difference between the two faunas was largely the result of higher densities of molluscs, crustaceans and ascidians at Albany (Table 5). Conversely, Upper Spencer Gulf has a considerably increased species richness, the result of an increased number of polychaete species in Upper Spencer Gulf (Table 5). By comparison of the species lists in Hutchings e2 al. (1991) and Appendix 1 of this paper it is clear that there is. little similarity in the species composition of the infauna between these two localities. Therefore, as for P. australis, the intauna of P sinuosa appears to vary substantially between locations. Despite the different species composition there was a considerable similarity in feeding types between Upper Spencer Gulf and Albany (Table 6). Deposit feeders, carnivores including scavengers, and TaBLe 5. Posidonia sinuosu: distribution of the major phyla (S = number of species, d = overall mean density), site Polychnetes Molluscs Crustaceans Echinoderms Astidians Minor Toral Albany (Hutchin ge et al_ 1991} $ 24 18* 7 16 1 106 a 7 259 6a 54 40 - 1797 Spencer Gulf (this study) $ 37 3 4 2 5 4 192 a7 630 95 38 u u 300 * amphipods not identified to species = densities {number Gulf = 2,052 m’), of individuals m~) calculated from total abundances and total area sampled (Albany = 1.2 m?, Spencer Tame 6 Posidonia sinuosa: distribution of the fauna amongst feeding categories at Albany and Spencer Gulf (number of species (%))- location herbivore Suspension deposit camivore. omnivore Albarty (from Hutchings et a/,, 1991) polychaetes 3 (7) (3) 18 (39) 19 (41) - molluscs (2 (50) 4 (17) 4 (17) 417) - crustaceans - 1 vo m(s)) 6 (35) 9 (53) ascidiang i4 (100) . - - echinoderms - 1 (14) 3 (26) 2 (26) 2 (26) totals 15. (14) 26 (24) 25 (23) 31 (29) b () Spencer Gulf (unis study) polychaetes I4 (Ja) 16 (16) 38 (39) 28 (29) 1a) molluscs 9 (28) 12 (38) 319) § (25) - crustaccans (5) 22 (50) 20 (45) ascidians = 5 (00) - : = echinederms 2 17) 1 (8) 3 (25) 6 (50) - tolals 25 (13) 36 (19) 66 (35) 62 (33) Luh 4 FP A. HUTCHINGS, T J. WARD, J. H. WATERHOUSE & 1. WALKER Suspension feeders were jhe dominant feeding types in P. sinwosa beds in both locations. Concinsions While species diversity in the subtidal seagrass beds is higher in Upper Spencer Gulf thas in Albsny, it as lower than in the seagrass beds of NSW. The density of individuals is far lower in Upper Spencer Gulf than elsewhere. and this may reflect the different temperature and salinity regimes in Upper Spencer Gulf compared to Albany and the aine locations along the NSW coast, Collett et ai. (1984) concluded that hydrological conditions were Important determinants in the compostiion of the infauns of seagrass beds, and more important than latitudinal effects. This study and the study of Albany (Hutchings ef al, 1991) indicate thal the conclusions of Collett er al. (1984) may also hold for the broader geographical region of southern Australia. Hydnmlogical conditions may also be responsible for the difference between the Northerp Spencer Gulf intertidal infuuna (Ainslic et af. 1989) and the intertidal infauna of Upper Spencer Golf observed in this study. Acknowledgments Field collectlons were funded by the International Lead Zinc Research Organization, New York. Dr K. Tiller at the CSIRO Division of Soils, Adelaide kindly provided laborajory space. Vic Neverauskas, Rod Grove-Jones, Nigel Holmes, Kim Horne, and Annette Green assisted with field and laboratory work. Jurgen Sieg of the University of Osnabriick identified the tanaids. We appreciate assistance from the following people from The Australian Museum with the identifications: Helen Stoddart, Roger Springthorpe (crustaceans), Frank Rowe (echinoderms), Geol? Holloway (insects), Dug Hoese (fish), Jan Loch and Winston Ponder (molluscs). We also appreciate the assistance of Peter Davie (The Queensland Museum) who idemified some of the decapods and Gary Poore (The Museum of Victoria) whe identiled the isopods. References AINsuie, R.-C., JouNston, De A. & Orrcen, BE. W. (1989) idal communities af Negthemn Spenter Gulf, South Australia. Trans. R. Soc. 3. Anst. 13, 69-83, BuLLock, D. A. (1975) The general water circulation of Spencer Gulf, South Australia, in the period February to May. bid. 99, 43-53. CoLtett, L. C., HuTewinas, P. A., Ginss, P 3. & Cotums, A.J. (1984) A comparative study of the macry-benthic fauna of Posidonie australis seagrass meadows in New South Wales, Aquatic Bot. 3B, 11-134. Exar, G. £. (1990) The use of the size styucvare uf benthic macrofaynal comfiunities to elucidate faunal biomass and secondary production. J, exp. Mar. Biol, Ecol, 137, 5-214. Howarp, R K., Bpoar. G, 1, & Hutcrmas, P. A. (1989) Faunsl sssembluges of seagrass beds. pp 536-564, In A. W. D. Larkum, A.l McComb, & S.A, Shephend, (Ede) “Seagrasses ; A Treatise on the Bivlogy of Seagrasses with apecial reference io the Australasian Region” (Elsevier, North Holland}. Hutcomes, PA. & Recver H. F (1974) The faume of Careel Bay, with commeats on the ecology of mangrove and seaerass communities Aust. Zool. IB, 99-123. . FOR, Wetts, D, I. Watker, & G, A. Kenpeice, (1991) Sengrass, sediment and inf4una — 2 comparison of Fosidonia australis, Posidonia sinuove and hibolis eatarctica in Princess Royal Harbour, south-western Australia. 0, Distribution, composinon x abundance of amacrofauna, pp, 611-633. fn F. E. Wells, D, 1. Walker, H. Kirkman, & R. Letherbridge (Eds) “Proceedings of the Third International Marine Biological Workshop: The Marine Flora and Pauna of Albany, Westem Australia”. Vol. 2. (Western Australian Museum, Perth.) Larkum. A. W. D. & ovw Harroc, C_ (1989) Evolution and Biogeography of seagrass beds. pp.: 12-156, fe A, W, D. Larkum, A.J. McComb, & S. A. Shepherd, (Eds) “Seagrasses 4 Treatise on the Biology of Seagrasses with spectal reference to Ihe Australasian Region”. (Elsevier, Nort Holiand), Nunes, R, A. & Lanwon, G. W, (1986) Physical propeny Uistributions and seasonal trends in Spencer Gulf, South Austmulia: an inverse estuary, Aust, J) Mar, Freshw, Res. 37, 34-53, STEFANSON, R. (IS77) Spencer Gulf 9 review of the oceanography, marine biology aud the implications for development. pp. 13-16. Third Australian Conference oo Caastal and Ocean Engineering. (Institution of Enganeers Australia, Melbourne). Watken, D. 1., Hurcaas, P A & Weis FE. (1991) Seagrass, sxhunent and infauna- a rison of Fosidonis australis, Posidania sinuose and Amphibolis antarctica in Princess Royal Harbour, south-western Austrulia. [. Seagrass biomass, productivity and contribution 1 sediments. pp. 597-610. Jn F. E, Wells, D- T. Walker, H. Kirkman, & R. Letherbridge (Eds) “Proceedings of the Third Enietnational Marine Biological Workshop: The Marine Flora and Fauna of Albany, Western Australia’ Vol. 2. (Western Austrafian Museum, Perth.) Warp, T, 3, (1987) Temporal variation of metals in the seagrass Posidovic australis aid its potential as a sentinel accumulator near a lead smelter. Mar, Biol. 95, 315-321. & Youns P. C (981) Trace metal contammation of shallow marie scdiments near’ lead smelter, Spencer Gulf, South Australia. Aust. J. Mar: Fresh, Res, 32, 45-56. &___ (1982) Effects of sediment trace metals and Particle aiz¢ on lhe community structure of epiberthic seagrass fauna near a lead smelter, South Australia, Mar, Ecol. Prog. Ser. 9, 137-146, WELLS, F. E,, Rose, R, A, & LANG, S. (1985) An analysis of benthic marine invertebrate communities in subtidal Seagrass aml yand habitats in Shark Bay, Western Australia Rec. West. Aust. Mus. 12, 47-56, INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF 9 Appendix 1 Summary of all data Species Family Habitats intertidal Zostera Posidonia Posidonia Sm bare 10m bare sum australis sinuosa Harmothoe sp. 1 Polynoidae 5 7 3 1 16 Harmothoe sp. 2 Polynoidae 1 1 Harmothoe sp. 3 Polynoidae 2 1 3 Paralepidonotus ampulliferus Polynoidae 1 1 Sigalion sp. Sigalionidae i 2 6 3 2 ? Sthenelais sp. Sigalionidae 1 1 Chrysopetalum sp. Chrysopetalidae 1 1 Eumida sp. Phyllodocidae 21 18 15 5 59 Paranaitis sp. Phyllodocidae 1 4 5 Preryosyllis sp. | Phyllodocidae 2 2 Phyllodoce sp. A Phyllodocidae 49 y 4 6 6 Eteone n.sp. Phyllodocidae 9 5 1 L 16 Genetosyllis sp. Phyllodocidae | 1 2 Phyllodacid sp. 5 Phylladocidae I 4 3 8 Podarke tmicroantennata Hesionidae l 2 3 Padarke angustifrons Hesionidae ul 9 20 Typosyllis sp. 1 Syllidae 1 1 17 3 22 Exogone sp, Syllidae 2 2 4 Odantosyllis sp. 1 Syllidae. 2 2 Odontosyliis sp. 2 Syllidae 2 1 3 Pionosyllis ehlersiaeformis — Syllidae 10 14 2 2 28 Syilinae sp. 2 Syllidae 2 7 9 Typosyllis sp, 2 Syllidae 52 20 1 1 74 Ehlersia sp. Syllidae 15 22 1 39 Syllinae sp. 8 Syllidae 1 1 Ceratonereis mirabilis Nereididae 1 6 14 3 4 4 32 Neanthes sp. | Nereididae 1 1 7 9 Neanthes bassi Nereididae 10 15 3 2 13 43 Nereis bifida Nereididae 31 577 29 7 1 645 Neanthes kerguelensis Nereididae 6 9 36 3 74 Platynereis sp. Nereididae 65 5 2 1 73 Neanthes sp, 3 Nereididae 1 7 8 Neanthes vaalii Nereididae 45 i 1 1 49 Ceratocephela n.sp. \ Nereididae 1 L Neanthes sp, 2 Nercididae 1 1 Ceratonereis sp. 1 Nereididae 248 8 34 I4 69 Il 384 Ceratonereis aequisetis Nereididae 4 4 Nereid sp. 20 Nereididae 2 1 3 Gymnonereis sp. 1 Nereididae 2 2 Nephtys gravieri Nephytidae 109 99 13 15 19 67 542 Glycera americana Glyceridae 3 2 5 5 3 18 Glycinde sp. | Glyceridae 31 12 9 4 3 59 Goniada maculata Goniadidae 1 | Onuphid sp. | Onuphidae 1 B 21 1 36 Nematonereis unicornis Eunicidae 103 8 25 7 253 Eunicid juy sp. 1 Eunicidae I 1 Marphysa sp. 1 Eunicidae 1 1 2 3 34 41 Marphysa sp. 2 Eunicidae 2 oe) 1 2 2 22 Lysidice sp. 1 Eunicidae ! 5 6 Eunice sp. 1 Eunicidae 5 81 i9 4 109 Eunice sp. 3 Eunicidae 41 72 51 35 199 Eunice sp. 4 Eunicidae 7 6 2 IS Eunice sp. 6 Eunicidae I 14 39 8 62 Eunice sp. 9 Eunicidae 3 3 Eunicid sp, U Eunicidae 1 1 Eunice sp. 5 Funicidae 5 2 6 B Augeneria verdis Lumbrineridac 131 18 48 26 16 239 Lumbrineris sp. 1 Lumbrineridae 8 a 4 4 2 19 Arabella sp. | Arabellidae 9 8 ll 3 31 Schistomeringos sp. 1 Dorvilleidae 7 I 8 Schistomeringos loveni Dorvilleidae 19 16 6 16 7 64 Leitoscoloplos sp, 1 Orbiniidae 5 1 1 7 Phylo sp. 1 Orbiniidae 1 5 6 Naineris grubei australis Orbiniidae 4 6 10 10 PF. A. HUTCHINGS, T. J. WARD, J. H. WATERHOUSE & L. WALKER Appendix 1 Summary of all data (continued) Species Family Habitats intertidal Zostera Posidonia Posidonia Sm bare 10m bare sum australis —_ sinuosa Scoloplos simplex Orbiniidae 1 74 17 4 4 l 101 Leitosceloplos normalis Orbiniidae 31 414 21 B 1 380 Family Orbiniidae Orbiniidac 17 1 1 9 Aonides oxycephela Spionidae 1 ! Spio sp. B Spionidae 2 1 3 Prionospio sp. 2 Spionidae 1 2 1 4 Polydora sp. 1 Spionidae 2 2 Pseudopolydora sp. 1 Spionidae 5 ] 4 10 Boccardia sp. 3 Spionidae | 1 Boccardia sp, 2 Spionidae 1 1 Aquilaspio multipinnulata Spionidae 7 1 9 ! 2 20 Aquilaspio aucklandica Spionidae 3 2 3 8 Spio pacifica Spionidae 1 1 Scolelepis sp. 2 Spionidae. 5 17 5 27 Pseudopalydora sp. 4 Spionidae 1 5 6 Malacoceros sp. | Spionidae 10 28 38 4 90 Scolelepis sp. 1 Spionidae 2 4 6 Spionidae sp. 2 Spionidae 1 ] 2 Spionidae sp. 3 Spionidae L ! 2 Boccardia sp. | Spionidae 1 1 Aricidea sp. 2 Pilargiidae 1 1 3 1 6 Magelona sp..1 Magelonidae 2 5 1 1 9 sp. 2 Cirratulidae 1 1 Caulleriella sp, 1 Cirranulidae 4 1 5 Caullertella sp. 2 Cirratulidae 1 20 4 31 6 72 Caulleriella sp. 3 Cirratulidae 7 35 43 14 1 100 Caulleriella sp. 4 Cirratulidae 20 3] 38 21 3 3 Cirriformia sp. 1 Cirratulidae 43: 43 28 4 9 127 Cirriformia sp. 2 Cirratulidae 29 43 3 5 2 92 Cirratulid sp. 1 Cirratulidae 3 1 4 1 9 Dipilacirrus sp. Flabelligeridac 92 3 4 1 2 102 Scalibregma inflatum Scalibregmatidae 5 2 ll 2 20 Hyboscolex dichranochaetus Scalibregmatidae 1 1 Armandia intermedia Opheliidae 10 51 5 8 74 Heteromastus filiformis Capitellidae 26 5 31 Capitella. ‘‘capitata*’ Capitellidae 53 608 661 Barantolla. sp, Capitellidae 88 57 88 46 6 285 Mediomastus n. sp. Capitetlidae 19 19 36 7 7 RB ?Scyphaproctus sp. 1 Capitellidae 1 L 5 1 8 Scyphoproctus sp. 2 Capitellidae 7 2 7 I 17 Notomastus sp. 1 Capitellidae 2. 96 51 25 2 186 Notomastus torquatus Capitellidae 4 4 29 16 63 Leiochrides sp. 1 Capitellidae. 3 1 4 Poechilochaetous serpens Poecilochaetidae Ll 2 8 ll Asychis sp. Maldanidac 5 2 iS 19 1 Petaloproctus sp, Maldanidae 1 10 2 10 I 24 Praxillella sp. Maldanidae 1 2 2 5 Maldanid sp. 3 Maldanidae 3 3 Owenta fusiformis Oweniidae 5 3 8 Pectinaria sp. 1 Pectinuridae t 3 4 Isolda pulchella Ampharetidae 1 1 2 Ampharetid sp. 1 Ampharetidae ! 3 38 7 49 Ampharetid sp, 3 Ampharetidae 1 1 Lysilla pacifica Terebellidae i 4 7 26 Polycirrus sp. 1 Terebellidae 1 1 2 Polycirrus tesselatus Terebellidae 4 5 26 1 3 39 Polycirrus cf. nephrosus Terebellidae 1 1 ?Decathelepus Terebellidae 1 1 Thelepus plagiostoma Terebellidae 1 2 3 Thelepus extensus Terebellidae 1 1 Streblosoma sp. Terebellidae 33 24 57 Lanussa exelysis Terebellidac 6 18 1 25 Lysilla laciniata Terebellidae 2 1 1 4 Pista australis Terebellidae 18 16 1l 6 122 63 Pista violacea Terebellidac 440 6 2 3 2 473 INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF Il Appendix { Summary of all data (continued) Species Family Habitats intertidal Zastera Posidonia Posidonia Sm bare 10m bare sum australis sinuosa Neoleprea sp. Terebellidae 4 8 3 ah Loimia ingens Terebellidae 2 2 Lanassa acellata Terebellidae 7 2 9 Eypolymnia koorangt Terebellidae 24 26 3 1 2 56 Nicolea amnis Terebellidac 2 7 1 10 Lanicides fascia Terebellidae 1 i Amphitritinae sp. 3 Terebellidae 1 1 2 Amphitritinae sp. 4 Terebellidae 2 2 Terebellides stroemii Trichobranchidae 7 6 5 1 19 Trichobranchus sp. Trichobranchidae 1 4 4 1 10 Sabellid sp. 1 Sabellidae 37 1 3 91 Exchone variabilis Sabellidae 2 5 2 4 B Laonome sp. 1 Sabellidace 2 2. Sabellid sp. 3 Sabellidae 2 6 10 18 Sabellid sp. 4 Sabellidae ! I Galeolaria sp. Serpulidae 3 3 Serpula sp. 2 Serpulidae 1 1 Serpula sp. | Serpulidae 1 1 Hydroides sp. 1 Serpulidae 10 4 7 4 l 26 total polychaetes 498 2590 1779 1292 1029 497 7685 Haliotis sp. Haliotidae 1 1 Amblychilepas nigrita Fissurellidae 16 1 17 Amblychilepas cf. oblonga _—Fissurellidae 1 1 Amblychilepas sp, Fissurellidae 1 1 Emarginula sp. Fissurellidae 1 1 Collisella sp. Lotiidae l 1 Naccula parva Lotiidae 5 10 15 Isoclanculus dunkeri Trochidae 2 3 4 9 Phasianotrachus cf. apicinus Trochidae 3 1 4 Thaliotia sp. Trochidae 1 1 Gena sp. Trochidae 1 1 Micrastrea rutidoloma Turbinidae 1 4 4 9 Phasianella australis Turbinidae 1 1 2 Bembicium vittatum Littorinidae 1 1 Triphorid sp, 1 Triphoridae 1 1 2 Zeacumantus diemenensis Potamididae 1 206 207 Bittium sp. 1 Cerithiidae 2 1 3 Bittium sp, 2 Cerithiidae 4 5 Diala. sp. Dialidae 1 3 L 5 Gazameda iredalei Turritellidae 2 1 3 Sabia conica Hipponicidae 1 1 2 Zeacrypia immensa Calyptraeidae T 7 Sigapatella calyptraeformis Calyptraeidae 3 3 Eunaticina umbilicata Naticidae 2 2 4 Ectosinum zonale Naticidae 1 1 Cominella eburnea Buccinidae 2 I 2 2 7 Cominella sp. Buccinidae 1 1 Nassarius pauperus Nassariidae 1 1 Fiusinus australis Fasciolariidae | 1 2 Deniimitrella cf. lincomensis Columbellidae 2 1 3 Splendrilla hurpularia Turridae 2 Conus anemone Conidae 1 1 2 Scaeoleda verconis Nuculidae 3 3 Solemya cf. australis Solemyidae i 16 17 Barbatia pistachia Arcidae. 2 2 Glycymeris flammea Glycymeridae it L Glycymeris radians Glycymeridae & 8 Limopsis cf. tenisont Limopsidae 1 5 6 Limopsis sp. Limopsidae 5 5 Myrilus sp. Mytilidae 3 3 Austromytilus penetectus Mytilidae 3 Z 2 57 211 277 Trichomya hirsuta Mytilidae 3 3 Musculus cf. ulmus Mytilidae 2 2 1 4 6 15 Musculus cf. paulucciae Mytilidae 3 2 2 10 17 Modiolus albicostarus Mytilidae 1 1 2 P, A. HUTCHINGS, T. J. WARD, J. H. WATERHOUSE & L. WALKER Appendix 1 Summary of all data (continued) Species Family Habitats intertidal Zostera Posidonia Posidonia 5m bare 10m bare sum australis — sinuosa Brachydontes erosus Mytilidae 8 3 1 12 Atrina tasmanica Pinnidae 1 I Atrina sp. Pinnidae 2 2 Pinna bicolor Pinnidae 5 1 6 Electroma georgiana Pteriidae ul 5 2 3 3 24 Electroma sp. Pteriidae 2 2 Malleus meridianus Mallcidae 3 1 4 Valsella vulsella Malleidae 7 7 Equichlamys bifrons Pectinidae ] 2 3 Lima nimbifer Limidae | 1 Monia ione Anomiidae I 18 i9 Monia zelandica Anomiidae 4 4 Anomia descripta Anomiidae 2 2 4 Saccostrea *‘australis'' Ostreidae 1 3 3 19 26 Wallucina assimilis Lucinidae 4 60 2 5 4 is) Cavitidens perplexa Lucinidae 4 29 I4 1 1 49 Myrtea bractea Lucinidae 1 1 Venericardia sp, Carditidae 1 1 2 Fulvia tenuicostata Cardiidae l 1 Chama ruderalis Chamiidae i 1 Dasinia histrio Veneridae 1 i Circe rivularis Veneridae 1 1 3 5 Katelysia rhytiphora Veneridae 2 i 15 28 Katelysia sp. Veneridae | 3 1 7 Placamen flindersi Veneridae 13 3 Timoclea sp. Veneridac 1 1 3 l 6 Mactra sp. Mactridae 1 2 3 Téllina mariae Tellinidae 6 7 20 45 12 1g 172 Tellina deltoidalis Tellinidae 226 8 5 239 Tellina sp. 1 Tellinidae 1 1 Tellina sp, 2 Tellinidae 7 7 Semele monilis Semelidae 3 2 Laternula creccina Laternulidae il 48 2 6 67 Myadora complexa Myochamidae 1 2 4 7 Frenamya patula Pandoridac 1 1 Teredo Teredinidae 2 2 Bivalve 2 I 1 Bivalve 4 2 2 Chiton sp. 1 Ischnochitonidae 1 2 3 Stenochiton longicymba Ischnochitonidae 38 46 84 Stenochiton cf: pilsbryanus —_ \schnochitonidae 10 17 2 1 30 Chiton sp. 2 Ischnochitonidae 1 1 Chiton sp. 3 Ischnochitonidae 2 4 6 Dorid sp. 1 Dorididae 2 2 Dorid sp. 2 Dorididae 1 1 Dorid sp. 3 Dorididae 1 1 Philinopsis lineolata Aglajidae 1 I Philine sp: Philinidae 1 1 2 Atys sp. Haminoeidae 1 1 Salinator sp. Amphibolidae 2 2 total molluscs 25 677 169 194 185 374 1624 Nebaliidae sp. 1 Leptostracan 19 6 25 Paguroidea sp, 1 3 1 l 4 Grapsidae sp. 2 Grapsidac 1 1 Hemigrapsus spinosus Grapsidae 1 31 2 2 1 37 Halicarcinus rostratus Hy menosomatidae 1 3 4 Halicarcinus ovatus Hy menosomatidae 2 2 Portunus pelagicus Portunidae 1 i Ebalia intermedia Leucosiidae 3 3 6 Ebalia sp. A Leucosiidae 2 2 Macrobrachium intermedium Palaemonidae 9 2 2 13 Alpheus bidens Alpheidae 4 4 Alpheus euphrosyne richardson Alpheidae 2 2 Alpheus novaezealandiae Alpheidae 2 2 INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF B Appendix 1 Summary of all data (continued) Species Family Habitats intertidal Zostera Posidonia Posidonia Sm bare 10m bare = sum australis sinuosa Alpheus sp. Alpheidae 1 1 Metapenaeopsis novaeguineae Penaeidae 1 1 Apseudes australis Apseudidae 513 63 2 7 3 588 Apseudes sp. | Apseudidae 5 3 4 2 14 Apseudes sp. 2 Apseudidae 72 8 3 83 Kalliapseudes sp. Apseudidae 1 1 Tanais dulongi Tanaidae 26 819 1 846 Leptochelia ignota Tanaidac 1 1 2 4 Cymadusa sp, Amphithoidae 2 5 24 3 44 Ampithoidae sp. 1 Ampithoidae 2 2 Ampelisca toora Ampeliscidae 2 1 i 4 Byblis cf. bega Ampeliscidae 1 1 Pratolembos drummondae Aoridae 17 22 3 42 Xenacheira fasciata Aoridae 2 2 Bemlos strigilis Aoridae 1 i] Ceradocus ramsayi Melitidae 1 1 2 Maera mastersi Melitidae 6 1 2 9 Maera sp. Melitidae 1 2 3 Ceradocus dooliba Melitidae 6 6 Ceradocus rubromaculatus —_ Melitidae 2 2 Ceradocus serratus Melitidae l 2 3 Cottesloe berringar Melitidae 6 1 7 ?Parelasmopus s}). Melitidae 12 l 3 Melitidae sp. 1 Melitidae 1 1 Melitidae sp. 2 Melitidae I 1 Melitidae sp. 4 Melitidae 3 3 Leucothoe ?commensalis Leucothidae } L Leucothoe assimilis Leucothoidae 1 l Leucothoe sp. 2 Leucothoidae 1 1 Leucothoe sp. 1 Leucothoidae 2 2 Waldeckia sp. 1 Lysianassidae u 1 Waldeckia sp. 2 Lysianassidae 1 1 Waldeckia sp. 3 Lysianassidae 1 ! Parawaldeckia. sp. Lysianassidae 1 ! Tryphosella sp. Lysianassidae 1 1 Ischyroceridae sp. 1 Ischyroceridac 3 5 a Phoxocephalidae spp. Phoxocephalidae 1 23 5 4 33 Atylus sp. Dexaminidae 4 4 Paradexamine lanacoura Dexaminidae l 1 2 Paradexamine ?linga Dexaminidae L 2 3 Paradexamine moorehousei Dexaminidae l L Oedicerotid sp. 1 Oedicerotidac 1 ) 2 Eusirid sp. 1 Eusiridae 1 1 Eusirid sp. 2 Eurisidae 2 2 Eusirid sp. 3 Eusiridae l 1 Eusirid sp, 4 Eusiridae L 3 4 Hyale sp. Hyalidae 43 43 Tamituka doowi Platyischnopidae it 1 Natatalana wowine Cirolanidae 6 12 3 21 Platynympha longicaudata Sphaeromatidae 2 291 22 & 323 Cymodoce sp. 1 Sphaeromatidae 1 1 Cymodoce sp. 2 Sphaeromatidae 2 49 18 9 78 Cymodoce bidentata Sphaeromatidae 1 L Limnoria sp. nov. Limnoridae 5 6 Euidotea peronii Idoteidae 4 4 Crabyzos longicaudatus Tdoteidae | 1 Euidotea bakert Idoteidae 7 7 Leptanthura diemenensis Paranthuridae ; 1 ! Acculathura bassi Paranthuridae 2 L 3 Mesanthura stypandra Anthuridae 1 ] Amakusanthura olearia Anthuridae 3 l 4 Apanthura cf. isotoma Anthuridae 1 1 Haliophasma sp. 1 Anthuridac 3 3 3 1 10 Callianassa sp. Callianassidae 1 6 T Gomeza bicornis Corystidae 2 4 6 4 P, A. HUTCHINGS, T. J. WARD, J. H. WATERHOUSE & L. WALKER Appendix 1 Summary of all data (continued) Species Family Habitats intertidal Zostera Posidonia Posidonia 5m bare 10m bare sum australis sinuosa Ceratoplax punctata Goneplacidae 4 4 Litocheira bispinosa Goneplacidae 3 1 4 Actumnus setifer Pilumnidae 1 1 total crustaceans 33 1862 240 156 58 38 2387 Ascidiidae sp. 1 Ascidiidae 18 2 20 Ascidiidae sp. 2 Ascidiidac 3 1 4 Ascidiidae sp, 3 Ascidiidae 1 1 16 18 Ascidiidae sp. 4 Ascidiidae 5 1 6 Ascidiidae sp. 5 Ascidiidae 1 1 Ascidiidae sp. 6 Ascidiidae 2 2 Microcosmus sp. 1 Pyuridac 3 10 il 2 26 Didemnid sp. 1 Didemnidae 1 1 Pyura stolonifera Pyuridae 1 4 20 6 31 Polycarpa ?pedunculata Ascidiacea 4 4 total ascidians 0 25 2 22 32 22 113 Pentacta ignava Cucumariidae 1 I Cucumella mutans Phyllophoridae 1 I Thyone sp. Phyllophoridae 1 1 Scoliorhapis sp. ?nov. Chiridotidae 1 5 5 1 2 Trochodota shepherdi Chiridotidae 1 3 4 Leptosynapta dolabrifera Synaptidae 74 42 33 l 150 Order Synaptid Synaptidae 3 3 Thymio sycia Holothuriidae | L Goniocidaris tubaria Cidaridae t 1 Temnopleurus michaelseni — Temmopleuridae i 3 4 18 Tosia australis Goniasteridae 1 3 4 Patiriella exigua Asterinidae 1 1 Amphipholis squamata Amphiuridae 5 2 3 1 i Amphiura elandiformis Amphiuridae 1 1 23 25 Amphiura parviscutata Amphiuridae 10 4 2 20 46 Amphiura (Fellaria) sp. Amphiuridae 2 1 3 Amphiura trisacantha Amphiuridae 3 9 1 3 Amphiura sp. Amphiuridae I 10 15 1 27 Ophiocentrus pilosus Amphiuridae 3 I 4 Ophioconis opacum hiodermatidae 1 1 total echinoderms 1 80 74 78 28 66 327 Syngnathus phillipi Syngnathidae 3 3 Favonigobius lateralis Gobiidae 4 4 Pseudogobius alorum Gobiidae 2 2 Callogobius mucosus Gobiidae 5 5 Heteroclinus sp, 1 ] Ophiclinops varius l i Gymmapistes marmoratus Scorpaenidae 4 4 Platycephalidae 1 1 Vanacampus sp. i 1 Nemertean Nemertean 6 5 1 3 2 7 Enteropneust 5 5 Pycnogonid 2 2 Hirudinea 1 1 Sipunculan 1 10 4] 19 4 aN 96 insects 16 16 Porifera sp. 1 2 Porifera sp. 2 98 98 Porifera sp. 3 1 1 total minor taxa 17 128 55 22 18 20 260 Total 574 5362 2329 1764 1350 1017 12396 SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE (INSECTA: COLEOPTERA) AND A HOST PLANT OF CASTIARINA UPTONI (BARKER). BY S. BARKER* Summary Sixteen new species of Castiarina are described: C. aeruginosa sp. nov., C. antia sp. nov., C. anthrene sp. nov. C. chrysothoracica sp. nov., C. crucianella sp. nov., C. hasenpuschi sp. nov., C. indigesta sp. nov., C. luteofusca sp. nov., C. markhanloni sp. nov., C. melasma sp. nov., C. mimesis sp. nov., C. nebula sp. nov., C. rayclarkei sp. nov., C. tenebrosa sp. nov., C. woodi sp. nov. and C. xystra sp. nov.; and one new species of Themognatha, T. gordonburnsi sp. nov. Dicrastylis georgei Munir is identified as a host plant of C. uptoni (Barker). KEY WORDS: Coleoptera, Buprestidae, new species, Castiarina, Themognatha. Transactions of the Royal Society of S. Aust. (1993), 1I7(1), 15-26. SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE (INSECTA: COLEOPTERA) AND A HOST PLANT OF CASTIARINA UPTONI (Barker). by S. BARKER* Summary BARKER, S. (1993) Seventeen new species of Australian Buprestidae (Insecta: Coleoptera) and a host plant of Castiarina uptoni (Barker). Trans. R. Soc. S. Aust. W7(1), 15-26 4 June, 1993. Sixteen new species of Castiarina are described: C. aeruginosa sp. nov., C. antia sp. nov., C. anthrene sp. nov. C. chrysothoracica sp. noy., C. crucianella sp. nov., C. hasenpuschi sp. nov., C. indigesta sp. noy., C. luteofisca sp. nov., C. markhanloni sp. nov., C. melasma sp. nov., C. mimesis sp. nov., C. nebula sp. nov., C. rayclarkei sp. nov., C. tenebrosa sp. nov., C. woodi sp. nov. and C, xystra sp. nov.; and one new species of Themognatha, T. gordonburnsi sp. nov. Dicrastylis georgei Munir is identified as a host plant of C. uptoni (Barker). Key Woros: Coleoptera, Buprestidae, new species, Castiarina, Themognatha., Introduction The search for new species of Buprestidae occurring irregularly in isolated areas has been continued, with some success, by a group of dedicated amateur entomologists. Included in this new material are specimens of species rare in collections and some new species which are described herein as are some new species from older series which have been made available to me. The major gap in knowledge of this family in Australia is information on their larval biology and food plants. Castiarina uptoni (Barker) occurs in very isolated arid areas of the country. The larval food plant of the population which occurs in the Ashburton River district in W.A. has been identified as Dicrastylis georgei Munir. Gardner (1989a) in a generic revision of the tribe Stigmoderini pointed out that Polychroma Dejean held date priority over Castiarina LaPorte & Gory, but had not been used for over 130 years. She subsequently applied to the ICZN to have the name Castiarina conserved (Gardner 1989b) and that action has now been taken (Opinion 1628, Bull. Zool. Nomencl. 48 (1) March 1991, pp. 74-75). Materials and Methods Collection date and locality information listed for each specimen is a copy of all data written on each individual label. Male genitalia were prepared and displayed by the method described by Barker (1987) referred to in the text and if previously published, reference is given to the publication date. The specimens illustrated in all species but two are the holotype. Two allotypes are illustrated and the fact noted in the appropriate remarks section. Measure- ments given are mean total body length and width with standard error, except where there are insufficient * Department of Zoology, University of Adelaide, GPO Box 498, Adelaide, S. Aust. 5001. specimens available to make the last calculation, when only mean is given. Acronyms used in the text for museum and private collections following the four letter system of Watt (1979) are: ANIC Australian National Insect Collection, Canberra; NMVA National Museum of Victoria, Melbourne; SAMA South Australian Museum, Adelaide; WAMA Western Australian Museum, Perth; RCBA Mr R. Clarke, Byron Bay; JHIQ Mr J. Hasenpusch, Innisfail; MHSA Mr T. M. S. Hanlon, Sydney; MPWA Mr M. Powell, Melville; RMNA Mr R, Mayo, Wallsend; ASSA, Mr A. Sundholm, Sydney; GWQA, Mr G. Wood, Atherton. Castiarina crucianella sp. nov. FIGS 1C, 2B Holotype. &, 136 km NE Paynes Find, W.A., 31.ix,1984, Jones & Powell, WAMA. Allotype. 9, same data as holotype, WAMA. Paratypes. W.A.: 200, 399, same data as holotype, SAMA, MPWA. Colour. Head, antennae and pronotum blue-grey. Scutellum blue. Elytra orange with following blue markings: narrow basal margin; premedial fascia not reaching margin, reduced to elongate oblique mark in some specimens; post-medial fascia reaching margin; apical mark, all marks connected along suture. Ventral surface and legs blue. Hairs silver. Shape and sculpture. Head closely punctured, median sulcus broad, muzzle short. Antennae, antennomeres: 1-3 obconic; 4-11 toothed. Pronotum closely punctured, narrow basal fovea extending forwards to middle as glabrous line, basal notches represented by glabrous area on each side more marginal than medial; apical margin straight, basal margin bisinuate; laterally parallel-sided at base, rounded to widest pre-medially, rounded and narrowed to apex. Scutellum scutiform, glabrous, excavate, Elytra punctate-striate, intervals convex, smooth, laterally angled out from base, rounded at humeral callus, medially concave, rounded post-medially and narrowed to bispinose apex; sharp Fig. 1. Photomicrographs of male genitalia of the following Castiarina spp.: A. C. militaris (Carter); B. C. xystra sp.nov.: C. C. crucianella sp.nov.; D. C. aeruginosa sp.nov., E. C. sanguinolenta (C & G); F.C. mitnesis sp.nov.; G. C. inconspicua (Saunders); H, C. markhanloni sp.nov,; 1. C. antia sp.noy.; J. C. guttifera (Obenberger); K. C. tenebrosa sp.nov.; L. C, hasenpuschi sp.noy., M, ©. Inieofusca sp.noy.; N.C. wood sp.noy.; O. C. nebula sp.noy.; P, C. anelis (Saunders): Q. C. indigesta sp.noy.; R. C. rollei (Kerremans); 8. C. rayclarkei sp.nov.; T. C. chrysothoracica sp.nay. and Themognatha: VU. T gordonburnsi sp-nov, marginal spine, smaller sutural spine, margin rounded and indented between spines, apices diverging slightly. Ventral surface with shallow punctures, hairy, hairs medium length, edges of abdominal segments glabrous. S7; tuncate, medially indented in both sexes. Size. Males, 4.7 * 13.1 mm (3). Females 5.2 * 13.4 mm (4). Male genitalia. (Fig. \C) Parameres diverging from basal piece, more so post-medially. rounded to apex. Median lobe pointed, sides obtusely angled away. Apophysis of basal piece medium width, rounded apically. Remarks. Resembles C. crux (Saunders) and C. nota Barker. Differs from those species by being larger, with bright blue markings and different male genitalia (Barker 1990: Figs IK, IL). Name derived from crux L., cross. SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE Castiarina aeruginosa sp. nov. FIGS 1D, 2A Holotype. ot, 17 km E Mt Carbine, Qld, 23.1.1991, J, Hasenpusch & §, Barker, SAMA [ 21,241. Allotype, 9 Brumby Gulley, Mareeba-Kuranda Rd, Qld, 4.11981, 8. Barker, SAMA Lt 21,242. Puratypes. Qld: lo LO. Mt Molloy, 26.x1).1985, A Walford-Huggins, SAMA, W.A.. lor, 1 Q, 12 km E Broome, 5.iv.1986, D. Knowles, MPWA, Calour, Head coppery-yreen. Antennae purple. Pronotum and sculletum coppery-green. Elytra pale yellow with following green markings: narrow basal margin, Variable post-medial spot at margin on each elyiron; apical mark. Ventral surface; sternum coppery-green, ubdomen pale yellow: Legs purple. Hairs silver, Shape and seulpture, Head closely punctured, median Sulcus broad, muzzle short. Antennae, antennomeres; 1-3 obconic; 4-1 toothed, Pronotum closely punctured laterally. less so medially, basal fovea, basal notches more marginal than medial; apical margin projecting medially, basal margin barely bisinuate; laterally angled inwards trom base, rounded to Widest pre- medially, rounded and narrowed to apex. Scutellum scutiform, glabrous, flat. Elytura punctate-striate, intervals convex, shallowly punctured; laterally angled out frou base, rounded at humeral callus, medially concave, rounded post-medially and narrowed to bigpinese upex; small marginal spine. very small sutural Spine, margin rounded and indented between spines, apices diverging slightly, apical margin subserrate. Ventral surface with shallow punctures moderately hairy, hairs short, edges of abdominal segments glabrous. S7: males truncate; females rounded, slightly indented medially. i’ ea a, 5mm Fig. 2, Habitus illustrations of the following Castiarina specics A. €' -aerupinesa sp.nov. holotype; B.C crucianella 6p,nov; holotype. Size. Males, 4.3 % 11.2 min (4). Females, 5.) 13.5 mm (3). Male genitalia. (Pig. 1D). Short and broad. Parameres angled away from basal piece, more su post-mecdtally, rounded to apex, Median lobe sharp, sides obtusely angled away, Apophysis of basal piece mediune width, rounded at apex, Remarks. This species resembles. C. straminea (Saunders) but distinguished by greenish reflections of coppery markings and is smaller species, male genitalia differ (Barker 1986: Fig. 2N) and has northern distribution in Qld and W.A.. C. strained occurs in central and southern Qld and torthern N.S.W, Name derived from aeravo L., yerdigris. Castiarina chrysotheracica sp. nov. FIGS IT, 4B Hoelorype. a, Mt Carbine, Old, 15.7,)991, R. Clarke, SAMA £ 21,243. Allorype, 9 . same data as holotype, SAMA I 2),244. Paratypes. Qld: 299. Mt Carbine, #1.1992, R. Clarke, RCBA: Let, Mt Garnet, 94.1992, R. Clarke, MHSA; 1c", same data ax holotype, RCBA; 20 07, Mt Carbine, 16.1.1991, R. Clarke, ROBA; Ic, Mt Carbine, 8.1.1992, R. Clarke. RMNA, Colour. Head and antennae green with yellow reflections, Pronotum green with volden reflections. Scutellum yreen with yellow reflections. Elytra yellow with the following markings: blue-green narrow. basil margin and medial V-shaped mark surrounding scutellum; blue post-medial fascia reaching margin: blue apical mark, last two marks narrowly connected along suture. Ventral surface: sterum green with yellow reflections; male abdomen testaceous; female abdomen green except apex S7 testaceous. Legs green. Huirs silver. Shape.and sculpture. Head clasely punctured, median sulcus broad, muzzle short. Antennae, antennomeres: 1-3 obeonic; 4-11 toothed. Pronotum closely punctured. narrow basal fovea extending forwards to middle as glabrous line, basal notches more marginal than medial; apical margin slightly projecting medially, basal margin almost straight; laterally angled outwards from base, rounded pre-medially. tapered to apex, Scutellum scutiform, glabrous, excavate. Elytra Punctate-striate, intervals convex, punctured; laterally angled out from base, rounded at humeral callus, medially concave, rounded post-medially and narrowed in bispinose apex: sharp marginal spine, smaller sutural spine, margin rounded and indented between spines. dpices. diverging, apical margin subserrute. Ventral surface with shallow punctures, maderately hairy, hairs short, edges of abdominal segments glabrous, $7. rounded in both sexes. Size. Males, 14.3 + 0.24 » 3.5 4 O12 mom (A), Femiles, 13.5 * 5.1 mmm (3). 18 S. BARKER Fig. 3. Habitus illustrations of the following Castiarina species: A. C. rayclarkei sp.nov. holotype; B. C. indigesta sp.nov. holotype. Male genitalia. (Fig. 1T) Parameres slightly angled outwards from basal piece, slightly rounded medially, rounded at apex. Median lobe sharp, sides obtusely angled away. Apophysis of basal piece short and broad, rounded apically. Remarks. Colour and pattern of male resemble those of C. garnettensis (Barker) but that species has spineless apex and male genitalia are different (Barker 1989: Fig. 1G). Name derived from chrysos Gr., gold and thorax Gr., chest. Castiarina melasma sp. nov. FIG. 4D Holotype. 9 , Milmerran, Qld, 1.xii.1990, R. Clarke, SAMA I 21,245. Paratypes. Qld: 29 9Q, Milmerran, R. Clarke, RCBA, Colour. Head, antennae, pronotum and scutellum green with yellow reflections. Elytra yellow with black apical mark. Ventral surface and legs green. Hairs silver. Shape and sculpture. Head closely punctured, median sulcus narrow, muzzle short. Antennae, antennomeres: 1-3 obconic; 4-11 toothed. Pronotum closely punctured, basal fovea extending to middle as impressed line, basal notches on each side more marginal than medial; apical margin projecting medially, basal margin barely bisinuate; laterally parallel-sided at base, angled inwards then rounded to widest pre-medially, rounded and narrowed to apex. Scutellum narrow, scutiform, glabrous, excavate. Elytra punctate-striate, intervals convex, flat, punctured; laterally angled outwards from base, rounded at humeral callus, medially concave, rounded post-medially and narrowed to bispinose apex; small, sharp marginal spine, very small sutural spine, margin rounded and indented between spines, apices hardly diverging, apical margin subserrate. Ventral surface with shallow punctures, moderately hairy, sternal hairs medium length, abdominal hairs very short, edges of abdominal segments glabrous. S7: male unknown; female truncate and indented medially. Size. Females, 13.7 x 5.3 mm (3). 7.xii.1990, Remarks. Structure, colour and pattern distinct. Name derived from melasma Gr., a black spot. Castiarina indigesta sp. nov. FIGS 1Q, 3B Holotype. & , Kuranda, Qld, 2.ii.1992, J. Hasenpusch, SAMA I 21,246. Allotype. 9, Davies Creek, Mareeba, Qld, 18.i.1991, S. Barker, SAMA I 21,247. Paratypes. Qld: 200, 19, Kuranda, i.1948, G. Brooks, ANIC; 50° 0", 1 9, Kuranda, F. P. Dodd, SAMA; 1c’, Kuranda, 10.i.1980, G. Wood, RMNA; 19, Mt Molloy, ii.1987, G. Wood, RMNA; lo". Mt SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE 19 Fig. 4. Habitus illustrations of the following Castiarina species: A. C. xystra sp.nov. holotype; B.C. chrysothoracica sp.noy. holotype; C. C. antia sp.nov, allotype; D. C. melasma sp.nov. holotype. Garnet, 19.1.1989, S. Lamond, MHSA; Lo", Herberton, 1.1989, S. Lamond, MHSA; 20° 0", 19, Mt Garnet, 13.1.1991, R, Clarke, RCBA; 10°, 19, Mt Garnet, 7.1.1992, J. Hasenpusch, JHIA; to, 19, Mt Garnet, 9.1.1992, R. Clarke, RMNA; Lo, 19, Mt Garnet, 10,i.1992, R. Clarke, RCBA; Io", 19, Mt Garnet, 10.1.1992, R. Clarke, MHSA; I[o', Davies Creek, 18.1.1992, J. Hasenpusch, JHIA; 10, Kuranda, 22.1.1992, J. Hasenpusch, JHIA; 19, Windsor Tableland, 2.ii1.1992, J. Hasenpusch, JHIA. Colour. Head black with blue reflections. Antennae blue-green. Pronotum black medially (narrowly at apex, broadly at base) laterally red-brown, Scutellum black. Elytra reddish-brown with following markings: narrow dark blue basal margin; black sutural mark commencing medially, reaching apex expanded as black apical mark. Ventral surface and legs dark blue. Abdominal segments in some male specimens all brown. Hairs silver. Shape and sculpture. Head closely punctured, median sulcus, muzzle short. Antennae, antennomeres: 1-3 obconic; 4-11 toothed. Pronotum shallowly punctured, deep basal fovea extending forwards to apical margin as glabrous impressed line, basal notches more marginal than medial; apical margin projecting medially, basal margin bisinuate; laterally angled outwards from base, rounded to widest part pre- medially, tapered to apex. Scutellum cordiform, few punctures, glabrous, flat. Elytra punctate-striate, intervals convex, wrinkled and punctured; laterally angled out from base, rounded at humeral callus, medially concave, rounded post-medially and narrowed to truncate spineless apex; apices diverging, apical margin subserrate. Ventral surface with shallow punctures, sparse short hair, edges of abdominal segments glabrous. S7 rounded in both sexes. Size. Males, 16.8 + 0.32 x 6.4 + 0.10 mm (19). Females, 19.1 + 0.26 X 7.6 + 0.17 mm (10). Male genitalia. (Fig, 1Q) Parameres slightly angled outwards from basal piece, rounded post-medially and narrowed to apex. Median lobe sharp, sides obtusely angled away. Apophysis of basal piece short, medium width, rounded apically. Remarks. This species was confused with C. analis (Saunders) a smaller species occurring further south. C. analis has a prominent red margin around the elytra and pronotum, also their male genitalia are distinct (Fig, IP) Name derived from indigestus L., confused. Castiarina xystra sp. nov. FIGS. 1B, 4A Holotype. & , Black Mt, A.C.T., iii.1931, T. G., ANIC. Colour. Head light blue apically, dark blue basally. Antennae blue-green, Pronotum dark blue medially, red laterally. Scutellum blue. Elytra red with following blue markings: narrow basal margin, medially expanded into large rounded mark; sinuate fascia covering humeral callus, thin red strip on each side separating last two marks; post-medial fascia expanded anteriorly and posteriorly along suture; apical mark. Ventral surface: sternum predominantly blue medially, presternum red laterally with blue spot each side close to margin, presternal process red, meso- and metasternum red medially; third coxae blue basally, red apically, abdomen predominantly red, bases of segents 5, 6, 7, 8 mainly blue. Legs blue. Hairs silver. Shape and sculpture. Head closely punctured, median sulcus deep, muzzle short. Antennae, antennomeres: 1-3 obconic; 4-1] toothed. Pronotum closely punctured, basal fovea extending forwards to middle as impressed line, basal notches on each side more marginal than medial; apical margin projecting medially, basal margin almost straight; laterally angled outwards from base, rounded post-medially to apex. Scutellum cordiform, punctate, excavate. Elytra punctate-striate, intervals convex and punctured; laterally angled outwards from base, rounded at humeral callus, 20 5, BARKER medially Concave, rounded post-medially and numowed to spineless apex; apices hardly diverging. Ventral surface With shallow punctures, moderately hairy, hairs long on stemun, medium length on abdomen, edges of abdominal segments glabrous, S7: male truncate: female unknown, Size, Male, 15,6 % 60mm () Male venittalia, (Pig. 1B) Parameres angled outwards trom basal picee, rounded ul apex. Median lobe sharp, sides woutely angled away, Apophysis of basal piece narrowed and rounded apically. Remarks. The elytral pattem is similar wo that found in C\ militariy (Carter) but ihe basal colours ditfer being red in C. vysiwa and yellow in ©. militaris. C.. tystva is a larger species and male genitalia ure different (Fiz YA), Name derived from xysrra 1... scraper, Castiarina rayclarket sp. nov. FIGS 1S, 3A Heloivpe. (t, Acaciw Plateau, NS.W,, 16.1,1992, R. Clarke, SAMA J 21.248. Allotype. @, Avdeia Plateau, N.S.W., R. Clarke, SAMA T 21,249. Paratypes. Old: 19, Warwick, 26.17.1992, R. Clarke, MHSA4, NSW: 19. Acacia Plateau, 711.1992. R. Clarke, RCBA, Colour Head and antennae bright green. Pronotum: bright gteen medially and along anterior margin and base; red laterally. Elytra yellow with following black markings wilh green reflections: narrow mark along suture from middJe to apex covering both spines. Ventral surface and legs bright green. Huirs silver. Shape and sculpture, Head shallowly punctured, glabrous, median sulcus, muzzle mediam. length, Antennae, antennomeres: +3 obconic: 4-f toothed. Pronotum shallowly punctured, glabrous, basal fovea; apical margin projecting medially, basal margin bisinuate, [aterally parallel-sided at base, rounded medially to apex, Povea-on each side at basal angles. Scutellum scutiform, glabrous, flat. Elytra puncrare- striate, inleevals corivex and smooth; laterally angled outwards from base, rounded at humeral callus, ticdiatly parallel-sided, rounded post-medially and narrowed W) bispingse apex, small marginal spine, lager sutural spine, apices diverging. Ventral surtace with shallow punctures, sparse short hair. edges of abdominal segments glabrous. 57: rounded in both SEXES. Male genitalia, (Fig, WS) Parameres angled outwards from basal piece. rounded near apex. Median lobe sharp, sides acutely angled away. Apophysis of basul piece medium width, rounded apically. Size. Male, 184 * 67 mm (i). Females, 20.6 * 3.0 mm (3), V1.1992, Remarks. Vhis species shows morphological similarities to C. rollei (Kerremmans). It diflers in colour, size and male genitalia (Pig. IR) from that species, Named after Mr Ro Clarke, Byron Bay. Castiarina antia sp. now, FIGS IL, 4c Holotype, Of. Auairuding, W.A,, 23. 4i1,1997, M. Powell & D, Knowles, SAMA I 21,250, Allorype. GY. 38 km N Binnu, W.A., 13. xi. 1988, M. Peterson, WAMA, Colow. Head, antennae, pronatumn, scutellum ventral surtace and legs purple-bronze, Elytra yellow with following dark blue markings: holotype with narrow basal margin, pre-medial fascia represented by medtal spot on each elytron (allotype has broad fascia reaching margin); broad post-medial fascia teaching maryan; mark covering apex, Ventral surface and legs purple- bronze, Hairs silver Shape and sculpture. Head closely punctured, medrin sulcus broad, muzzle very short, hairy. Antennae compressed, antennomeres: L3 obconic; 4'4-toothed; 5-10 touthed. Pronotum closely punctured, basal fovea extending forwards to middle as impressed line: apicul margin projecting medially, basal margin bisinuate; laterally angled inwards tront base, rounded to widest pre-medially, rounded to apex, taterally hairy, Scutellum seutiform, glabrous. flat. Elytra punctate- sinate, iitervals Convex, punctured: laterally angled out from base, rounded at humeral callus, medially concave, rounded post-medially, rounded abruptly to bispinose apex: small blunt marginal spine, minute sutural spine, margin rounded and indented between spines, apices diverging; Ventral surtace with shallow punctures, hairy. hairs long, edges of abdominal segments glabrous. $7: males truncate, medially indented; females rounded. Male genitalia. (Fig. (1) Short. Parameres parallel- sided at base, rounded outwards. pre-medially, rounded to apex. Median lobe blunt, sides acutely angled ayay. Apophysis of basal piece narrow, rounded apically Size. Male, 3.3 10.2 mm (1). Female. 3.3 © 10:7 mm (1). Remarks, Only Custiarina species Unat has reduced number of abtendomeres. The specimen illustrated is the allolype (Fig. 41). the first specimen collected. Name derived from «aria L.. forelock. Castiarina anthrene sp, nov. FIG. 6E Holarype. 9, 7 km W Nalbarra HS (28.398. 117,36). W.A,., 24-30 111.1981, T. FE Houston, WAMA, Paratype W.A.: | 9 same data as holotype, SAMA. Celouw, Head, antennae black with blue reflections. Pronotum black medially, blue laterally, Scntellum SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAL al Nig. 5, Habitus illustrations of the following Caytiarina species: A, OC) luteefiasca sp.nov, holotype; B. C, weed) sp.noy. holowpe. CC fenebrosa sp-noy, holotype. black, Elytra yellow with following black nuarkings: narrow basal margin; pre-media fascia not reaching margin anteriorly, extending to margin posteriorly; post-tuedial fascia reachiny margin; apical mark, last three marks connected along suture. Ventral surface and legs dark blue. Hairs silver. Shape und seulprire. Head closely punctured, median sulcus, very Short muzzle. Antennae compressed, antennomeres: 1-3 obeonic, 444 toothed; 5-LL toothed. Pronotum closely punctured, basal fovea extending forwards to middle as glabrous line, basal notches represented by glabrous area on each side more murginal than medial, connected by glabrous posterior margin; apical margin projecting medially, basal margin alinest straight; laterally parallel-sided at base, rounded fo widest medially, rounded to apex. Scutellum scutiform, glabrous. excayate, Elytra costate, intervals 3, 5, 9 raised at basal half wrinkled and punctured; laterally angled out from base, rounded at humeral callus, medially concave, faintly rounded post- medially, wpered, potinded to bispinose apex; small sharp marginal and sutural spines, margin rounded and indented between. apices slightly diverging. Ventral surface with Shallow punctures, moderately hairy. hairs long, edges of abdominal segments glabrous. $7: mule unknown; fetiales truncate, medially indented. Size. Females, 1.0 * 3.6 mm (2). Remarks. This is a very distinctive elongate species showing the typical modifications found in wasp/bee mimics. Name derived from anthrene, Gr., wasp. Castiarina markhanloni sp. nov, FIGS 1H, 6C Holotype, o , Round Hill Reserve, N.S.W., 10.1992, T. M, 8. Hanlon, SAMA 1 21,251, Allotype. 9 , same data as holotype. SAMA J 21.252, Paratypes. N.SW.; Boro. 109 9, same data as holotype. MHSA. Colour, Head, antennae, pronotum bronze-green with purple reflections. Scutellum purple: Elytra pale yellow with following dark blue markings with purple and blue reflechons: narrow basal margin, pre-medial fascia ends expanded anteriorly over humeral callas and posteriorly touching margin; post-medial fascia touching margin; mark covering apex, all marks connected along suture. Veniral surface and legs purpie-bronze with purple reflections. Hairs silver. Shape and sculpture. Head closely punctured, shallow median sulcus, short muzzle. Antennae, antennomeres; 13 obconic; 44 toothed; 5-11 toothed. Pronotum closely punctured, punctations small medially larger laterally, basal fovea extending forwards to middle as glabrous line then to apical margin as impressed line; apical margin projecting medially, basal margin bisinuates laterally rounded from base, widest pre- medially, rounded to apex, laterally hairy. Scutellum small, scutiform, glabrous, Nat. Elytra punctate-striate, intervals corivex, wrinkled, punctured; laterally angled outwards from base, rounded al humeral callus, medially concave. rounded post-medially and narrowed to bispinose apex; small blunt marginil spine, sutural spine minute, margin in most specimens almost truncate and indented between spines. apices diverging. Ventral surface with shallow punctures. long dense hair. edges of abdominal segments glabrous. $7: truncate both sexes. Size. Males, 9.7 + O18 * 3.5 + 0.05 mm (14)- Females, 10.2 + 0.22 * 3.9 + 0.09 mm (IN). Male genitalia. (Fig. |H) Short. Parameres parallei- sided from basal piece, angled outwards, pre-medially rounded then tapered, rounded at apex. Median lobe blunt, sides acutely angled away then widened. Apophysis of basal prece medium width, rounded apically. Remarks. This species could be confused with C. incvonspicua (Saunders). Main differences are the smaller scutellum and the strongly rounded pronotum. The male genitalia show some similarity in form 22 S. BARKER Fig. 6, Habitus illustrations of the following Castiarina species: A. C. nebula sp.nov. holotype; B. C. hasenpuschi sp.nov, holotype; C. C. markhanloni sp.nov. holotype; D. C. mimesis sp.nov. holotype; E. C. anthrene sp.nov. holotype. although those of C. markhanloni are narrow and the apophysis of the basal piece is more tapered (Fig. 1G). Named after Mr T. M. S. Hanlon of Sydney. Castiarina mimesis sp. nov, FIGS IF, 6D Holotype. o , Badjalling, W.A., 7.xi.1970, 8, Barker, SAMA 121,253. Allotype. 9 , same data as holotype, SAMA 121,254. Paratypes. W.A.: IS@o, 599, same data as holotype SAMA; 2.9 9, South Tammin Flora Reserve, 8.x1.1970, S. Barker, SAMA; 10, 70 km S Perth, Albany Highway, 19.xi.1970, S. Barker, SAMA; 20°", 59 9, Northam, C. G. Jessup, SAMA; 49 9, no data SAMA; 19, 34 km N Gin Gin, I1.xi.1990, M. Powell, MPWA, Colour. Head, antennae, pronotum bright green. Scutellum blue-green. Elytra orange with following blue markings: narrow basal margin; vitta on each elytron from humeral callus to pre-apex; line along suture meeting vittae at pre-apex. In some specimens vittae shortened at basal end leaving narrow mark over each humeral callus. Ventral surface and legs bright green. Hairs silver. Shape and sculpture. Head closely punctured, medium sulcus, muzzle short. Antennae, antennomeres: 1-3 obconic; 4/4 toothed; 5-11 toothed. Pronotum closely punctured, basal fovea extending forwards to apical margin as impressed line; apical margin projecting medially, basal margin almost straight; laterally parallel-sided from base, rounded post-medially to apex. Scutellum scutiform, glabrous, excavate, Elytra punctate-striate, intervals convex, heavily punctured; laterally angled outwards from base, rounded at humeral callus, medially concave, rounded post- medially and tapered to bispinose apex; very small marginal and sutural spines, margin rounded and indented between spines, apices hardly diverging. Ventral surface with shallow punctures, hairy, hairs long, edges of abdominal segments glabrous. $7: truncate both sexes. Size. Males, 8.9 + 0.19 x 2.8 + 0.07 mm (19). Females, 8.9 + 0.21 x 2.9 + 0.08 mm (18). Male genitalia. Parameres angled outwards from basal piece, rounded post-medially then increasing in width, notched apically. Median lobe blunt, sides acutely angled away. Apophysis of basal piece broad, rounded apically. Remarks. This species was confused with C. sang- uinolenta (C & G ). Their elytral colour and markings are very similar but male genitalia are very different (Fig. IE) and they are easily distinguished by differences in structure of their elytral spines. In C. sanguinolenta these are sharp and equal, in C. mimesis they are small and sutural spines are indented. They occur together and are part of a Muellerian mimicry complex, The name is derived from mimesis L., imitation. Castiarina tenebrosa sp. nov. FIGS 1K, 5C Holotype. o, 4 km W Paluma, Qld, 7.i.1986, E. E. Adams, SAMA I 21,255. Allotype. 2, 4 km W = Paluma, Qld, 4.i.1986, A, Sundholm, SAMA I 21,256. Paratypes. Qld: 29 9 , same data as holotype, ANIC; lor, 1 Q, same data as allotype, ASSA; 2, oo", 4km W Paluma, 6/7.i1.1986, A. Sundholm, ASSA. Colour. Head, antennae and pronotum black with green reflections. Scutellum green. Elytra yellow with black markings coalesced forming basal, pre-medial and sub- apical spots, first two coalesced to form vitta also spot on humeral callus covering margin. Ventral surface green with yellow reflections. Legs dark blue. Hairs silver. Shape and sculpture. Head shallowly punctured, broad median sulcus, short muzzle. Antennae, antennomeres: 1-3 obconic; 4'%-toothed; 5-11 toothed. Pronotum shallowly punctured, small basal fovea; apical margin straight, basal margin bisinuate; laterally parallel-sided SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE 23 at base, rounded to widest pre-medially, narrowed to apex, Scutellum cordiform, glabrous, flat. Elytra punctate-striate, intervals convex, puncvared; laterally angled outwards from base, rounded at husneral callus, medially concave, rounded post-medially and narrowed to bispinose apex; shatp marginal spine, small, sharp sutural spine, margin straight between spines, apices diverging. Ventral surface with shallow punctures, moderately hairy, hairs medium length, edges of atiominal segments glabrous. $7: rounded in both sexes, Size, Males, 11.9 + 012 x 4.2 + 006 mm (6), Females, [6 * 4.4 mm (2) Male genitalia. (Fig, 1K) Parameres angled outwards fram basal piece, rounded post-medially, parallel- sided, rounded tw apex. Median lobe sharp, sides oblusely angled away. Basal piece wide, rounded apically. Remarks, This species was confused with C. guitifera {Qhenberger) which has only been collected at Kuramia, Qld, The male genitalia differ (Fig. LF) Name derived from tenebrosus L., dark. Castiarina hasenpuschi sp. a0v. FIGS IL, 6B Holorype. o . Mt Lewis, Qid, 61.1991, 1. Hasenpusch, SAMA J 21,257, Allatype, 9. sare dataas holotype, SAMA I 21,258 Furatypes, Qld: Lor, 29 9, same data as holulype, JHIQ. Colour, Head black with blue reflections, Antennae black with blue-green reflections. Pronotum black with purple reflections.. Scutelluis blue, Elytra yellow wah following black markings: two fasciae and apical mark coalesced leaving a pre-nedial, post-medial and pre- apical yellow spot on each elytron and one on margin at humeral callus. Ventral surface preen, Legs blue. Hairs silver: Shape end sculpture. Head closely punctured, median sulcus, short muzzle. Antennae, antennomeres: 1-4 obaeanic, 5-11 toothed. Pronotum closely punctured, narrow basal fovea; apical margin projecting medially, basal margin bisinuate; laterally rounded from base, widest pre-medially, rounded and narrowed to apex. Scutellum cordiform, punctured, Mat. Elytra purictate- striate, intervals convex, punctured; laterally angled Gut front base, rounded al humeral callus, medially concave, Tounded post-medially and narrowed to bispinose apex; sharp marginal spine, small, sharp sulural spine, margin straight between spines, aptoes diverging, Ventral surface with shallow punctures, moderately hairy, hair short, edges of abdominal segments glabrous, Males: legs 2 and 3 with reduced pulvilll on tarsomeres 1-3, replaced with single median spine, $7: male trunctate, indented medially; female truncate, slightly indented medially. Size. Males, 9.8 * 3.8 mm (2). Females, 1.4 x 4.0 mm (3). Male genitalia (Fig. 1L). Paraineres short and wedge- shaped, Median lobe sharp, sides obtusely angled away, Apophysis of bass] piece medium width, rounded zpically, Remarks, Structurally this species closest to members of C. sexplagiata group except that elytra are smooth whereas they are roughened with punctures in most members of group, Named afier Mi J. Haseripusch, Tanisfail. Castlarina liteafusea sp. nov, FIGS 1M, SA Holotype &, Mt Lewis, Qld, 11.1992. G. A. Wood, SAMA I 21,259. Allorype, &, Mt Lewis, Qk, 6.11992, G. A. Wood, SAMA I 21,260. Paratype, 1o, Qhd: NQ., F P. Dodd, ANIC. Colowr. Head green. Antennae, aptennomeres: 1-4 green; 5-li blue-green. Pronotum: bronze medially; green apically. Scutellum green. Elytra yellow whh black matkings coalesced leaving yellow basal and medial spots on each elytron, these coalesced forming angled vitta, round yellow apical spot and yellaw mark on lateral surface of humeral callus. Ventral surface green. Legs dark blue. Hairs silver. Shape and sculpture. Head shallowly punctured, broad median sulcus, short muzzle. Antennae, aliienmnomeres: 1-3 obconic; 4-11 wothed. Pronotum shallowly punctured, small basal fovea; apical margin projecting medially, basa] margin bisinuate, latenilly parallel- sided ar base, angled inwards, rounded to apex, widest pre-medially. Scutellun: scutiform, glabrous, flat. Elytra punctate-strate, intervals convex, punctured; laterally angled outwards from base, rounded at humeral callus, medially concave. rounded post- medially and tapered to bispinoase apex; sharp marginal spine, smaller sharp sytural spine, margin rounded between spines, Ventral surface with shallow ponciures, few short hairs, edges of abdominal segments glabrous. $7: trunctate in both sexes. Size. Male, 12.4 x 4.7 mm (1). Females, 12.7 * 48 mm (2), Male genitalia (Fig. IM), Parameres angled outwards from, basal piece, rounded to apex. Median lobe pointed, sides acutely angled away. Apophysis of hasal piece medium width, rounded apicully, Remarks. Largest of black and yellow C, producta (Saunders) group mimics and with C. tenebrosa and following new species, only ones that have yellow vitae oo elytra. Name derived from /uteus L., yellow and fuscus L,, black, Castiarina woodi sp. nov. FIGS IN, 5B Hoterype. oF, Mi Lewis, Qld, 1.11992, G. A. Wood, SAMA I 21,261 a4 S, BARKER Alloype, 9. same data as holotype. SAMA 21,262. Colowr, Head and antennae blue-green. Pronotum green with bronze reflections. Scutcilum blue-green. Elytra yellow with folkxwing dark green marks; matkings coalesced to form yellow basal and medial spols also coalesced forming long vitta covering humeral callus and margin; large round pre-apical spot, Ventral surface green. Legs blue-green, Hairs silver. Shape and sculpture: Head shallowly punctured, broad tmehan sulcus, short muzzle. Antennae, antennomeres: 1-3 obconic; 4-Ii toothed. Pronetum shallowly punctured, deep basal foves; apical margin projecting medially, basal margin bisinuatc, laterally parallel sided at base, rounded to apex, wides! pre-medially, Scutellum scutiform, glabrous, flat. Elytra punctare- striate, intervals cortver, punctured; Jaterally angled outwards from base, rounded at humeral callus, medially concave, rounded post-meiially to unispinose upex; sharp marginal spine, sutural spine represented by very small notch, margin straight betweert spine and fotch, apices diverging. Ventral surface with shallow punctures, moderately hairy, hairs long and denser in medial strip, edges of abdominal segments glabrous. S7: truncate in both sexes. Size. Male, 1.4 x 4.5 mm {l). Female, 11.9 «4.5 mm (1). Male genitalia, (Fig. IN) Pacameres more or less paraliel-sided from basal piece, rounded to apex. Median Jobe pointed, sides obtusely angled away. Apophrysis of basal piece short, narrowed and rounded to apex, Remarks. Elytral markings of this species arc similar tothese of C, luteofusea but distinctive male genitalia and clytral spines distiguish this species. Named after Mr & A. Wood, Atherton, Castiarina nebula sp. nvy. FIGS 10, 6A Solerype, of, Mt Lewis, Gid, 6.11991, 3, Hasenpusch, SAMA I 21,263 Atiaype. 9 , same data as holotype, SA MA 121,264, Perarypes. Qhd: 20-0, Kuranda, 1.19.1990, 5,71, 1990, G. A. Wood, GWAQ; 1¢, Kuranda, Qld, 2.1.1978, A, & M. Walford-Huggins, SAMA; 29 9, Kuranda, S.0.1990, J. Hasenpusch, JHIQ; 1¢, Kuranda, 141.1990, G, Wood, GWAQ, Colour Head and amennae green. Pronotum black with blue of green reflections, Scutellum green. Elyira yellow with following black markings: fasclae and marks coalesced leaving six yellow spots, Largest pre- medial, smallest post-medial and intermedial on margin at humeral callus. Veniral surface green. Legs blue. Halrs silver. Shape and sculprure, Head shallowly punctured, median sulcus, medium teagth muzzle Antennae, antenaomeres: t-3 oboome: 4/-loothed; 5-IL inethed., Pronotum shallowly punctured, very small basal fovea: apical margin straight, basal margin bisinuate; laterally angled inwands from base, then rounded from near base to apex, widest pre-mecdially. Scutelhim scutiform, few punerures, glabrous, flat. Elytra punctate-striate, intervals convex, smooth; laterally angled outwatds from base. rounded at humeral cullus, medially concave, rounded post-medially and tapered 10 bispinose apex; sharp marginal spine, smull sharp sutural spine, margin rounded and indented between spines, apices diverging. Ventral surface with shallow punctures, few short hairs, abdoniinal segments glaheous. S7: truncate both sexes, Males: legs 2 and 3 with cach pulvillus om tarsomeres L-3 replaced by median spine, Size. Males, 10.9 * 3.8 mum (3). Females, IH x 4,2 mm {4}, Male genitalia (Fig, JO). Parameres widened from base, rounded at apex. Median Jobe sharp, sides obtusely angled away. Apophysis of basal piece medium width, rounded at apex, Remarks. Elyiral markings of this species are similar to those of © actasignata (Carter, 1919). However C. octosignato difters stucturally as elytral spines are differem shape, Male C. octestenata Unknown. Name detived from nefufosus L., dark. Themognatha gordonburnsi sp. nov: FIGS WW, 7 Holetype, ct. 69 km N Galena Bridge, WA, 29.ix.1992_ M. Golding & M_ Powell, WAMA. Allotye. 2, 100 km N Murchison River, W.A,, 14.i%.1980, G. G. Burns, NM'VA, Faratypes, WA.; 20-9, same datd as allvtype, 14,ix.1980, 17.ix.1980, G. G. Burns, NMYA: Sor cr, same uatt as holotype, SAMA & MPWA, Colour, Head and antennae black. Pronotum black with yellow literal murgins, Scutellam black. Elyira predominantly brown with yellow fateral tmargins and following black markings: narrow basal margin; females with faint pre-apical vitta over each humeral callus-and faint markings on suture, post-medial tascin aot reaching margin; spade-shaped pre-apical mark extending over apex. Ventral surface: male predom- inantly black pre-sternuns with yellow mark on meta- slettal coxae; abdominal sclerites yellaw wih testaceous edges; fernale black with yellow mark oa meta-sternal caxae nd on lateral edges of all visible abdominal segments. Legs black. Haars silver. Shape ard sculpture. Head punctured, hairy, apex medium length, Antennae, antennomeres: 1-3 oboonic, 4 semi-toothed; 5-11) toothed. Pronotum punctured, apical margin projecting medially, basal margin faintly sinuous, laterally rounded from base to apex, bulbous pr:-medially. Scutellum almost circular, flat, without punenues Elytra punctate-striate, intervals convex and SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAG 2S 5mm Fig. 7 Habitus illustration of Themoynatha gordoaburnsi spunoy, allorype smooth; laterally angled outwards from base, rounded at humeral callus, medially more or less parallel-sided, rounded post-medially to spineless apex. Ventral surface punctured, hairy. S7: trunctate, medially indented in mule: rounded in female. Male genitelia. (Fig. 1U). Parameres angled outwards from basal piece, rounded at apex. Median Jobe blunt, sides acutely angled away. Apophysis of basal piece medium width, rounded apically. Size. Males, 20.6 + 0.35 x 8&4 + 013 mm (10). Females, 24.3 * 9:6 mm (3). Remarks. Although thé locality data on the holotype and allotype appear different they refer fo the same area. The allotype has a narrow median yellow band on the pronotum, not reaching base or apical margin. T. burnsi is a spring emerging species, adults have been collected on the Mowers of Grevillea sp, and Bursaria spinosa, On the basis of its structure and male gemitalia, it can be grouped with 7 oleata (Blackburn) u blue and red species which emerges in late summer and autumn and is usually associated with mallee Mowers. The specimen illustrated ts the ullotype (Fig. 7), the first specimen.collected, Named to commemorate my friend and colleague the late Mr Gordon Burns, Mornington, A host plant of C, uptoni (Barker) Although the adults of many Castiarina species are commonly collected, the pre-adult stages and host plants of even the commonest species are mostly unknown. Castiarina upiuni (Barker, 1979) was described from dead adults found inside tubular highway: marker posts, north af Barrow Creek, N.T. Presumably they had been dropped inte the cavities by predaccous insects that had been feeding on them, possibly asilid flies which commonly prey on buprestids (Barker & Inns 1976). Live specimens were subsequently collected in the same general area but their food plants were not recorded. Several years ago Herbert Demarz collected live specimens of C. uploni on the leaves of Dicrasrylis georgei Munir, a very low shrub growing in an ifterdune situation on Yanrey Station, 4-31 km south of Barradale roadhouse on the Great Northern Highway, W.A. On 10 August 1990, I visited the locality with Mr Demarz and we examined the same plants in that area, but found no adult beetles on the leaves, Many of the stems were dead and detached from the individual plants. Examination of the live part of cach cut stem showed frass and when dissected, every one was found to be galleried and to conlain a living beetle. Most were adult, but a proportion were newly pupated or in various stages of post-pupal metamorphosis, OF fourteen specimens dissected out, twelve were C. upteni and two were C. quadrifasciata (Saunders, 1869). The twa species belong to different species groups on basis. of structure of their male genitalia, but are similar in hody colour and pattern and thus form a Mullerian mimicry group. When I visited the same area on 20 August 1990, miuny C. uptoni adults were found on the leaves of the host plant as well as two specimens of C. quadrifasciata, As D. georgei does not occur in the N.T., beetles from the type locality must have another host plant which could be a second species of Dicrastylis, possibly D, gilesii F, Muell. which occurs in that area (Jessop 1981). Acknowledgments 1 thank the following for their assistance: Dr K. Walker, NMVA; Mr T. A. Weir. ANIC: Ms H. Vanderwoude, Department of Zoology, University of Adelaide. 1 thank the following collectors for the loan of specimens: Mr E, E. Adams, Edungalba; Mr 26 S. BARKER As D. georgei does not occur in the N.T., beetles from the type locality must have another host plant which could be a second species of Dicrastylis, possibly D. gilesii F. Muell. which occurs in that area (Jessop 1981). Acknowledgments I thank the following for their assistance: Dr K. Walker, NMVA; Mr T. A. Weir, ANIC; Ms H. Vanderwoude, Department of Zoology, University of Adelaide. I thank the following collectors for the loan of specimens: Mr E. E. Adams, Edungalba; Mr R. Clarke, Byron Bay; Mr H. Demarz, Wanneroo; Mr T. M. S. Hanlon, Ride; Mr J. Hasenpusch, Innisfail; Mr A. Walford-Huggins, Yeppoon; Mr R. Mayo, Wallend; Mr M. Peterson, Perth; Mr M. Powell, Attadale, Mr R. I. Storey, Mareeba; Mr A. Sundholm, Elizabeth Bay; Mr S. Watkins, Caparra; Mr G. A. Wood, Atherton. References BARKER, S. (1979) New species and .a catalogue of Stigmodera (Castiarina) (Coleoptera: Buprestidae). Trans. R. Soc. S. Aust. 103, 1-23. (1986) Stigmodera (Castiarina) (Coleoptera: Buprestidae): taxonomy, new species and a checklist. Ibid. 10, 1-36. (1987) Eighteen new species of Stigmodera (Castiarina) (Coleoptera: Buprestidae) bid. M1, 133-146. _______ (1989) Contributions to the taxonomy of Australian Buprestidae (Coleoptera): new species of Astraeus and Stigmodera (Castiarina) and a key to Astraeus (s.s.). Ibid. 113, 185-194. (1990) New species of Castiarina (Coleoptera; Buprestidae) and a redescription of C. diversa (Kerremans) and C. elderi (Blackburn). Jbid. 114, 169-178. & Inns, R. (1976) Predation on Stigmodera (Themognatha) tibialis by a fly. WA. Naturalist. 13, 147-148. Carter, H. J. (1919) Proc. Linn. Soc. N.S.W. 44, 137-173. GARDNER, J. A. (1989a) Revision of the Genera of the Tribe Stigmoderini (Coleoptera: Buprestidae) with a discussion of phylogenetic relations. Invertebr. Taxon. 3, 291-361. (1989b) Castiarina Gory & LaPorte, 1837 (Insecta, Coleoptera): proposed conservation. Bull. Zool. Nomencl. 46(3), 170-172. Jessop, J. (Ed.) (1981) ‘Flora of Central Australia? (A. H. & A. W. Reed Pty Ltd, Sydney). SAUNDERS, E. (1869) Descriptions of fifty new species of the genus Stigmodera. J. Linn. Soc. 9, 460-483. Wart, J. C. (1979) Abbreviations for entomological collections. N.Z. Zool. 6, 519-520. ADULT AND LARVAL STAGES OF PARAUSTROSTRONGYLUS RATTI (NEMATODA: TRICHOSTRONGYLOIDEA) FROM RATTUS FUSCIPES BY IAN BEVERIDGE* & MARIE-CLAUDE DURETTE-DESSETT Summary The morphology of the adult and the third and fourth larval stages of Paraustrostrongylus ratti from the native rodent rattus fuscipes is described, with particular reference to the synlophe, as well as the mechanism of attachment of the nematode to intestinal villi. The synlophe of the fourth larval stage has an oblique axis of orientation, in contrast to the frontal orientation in the adult stage, and most closely resembles the synlophe of species of Dessetostrongylus parasitic in dasyurid marsupials. Ontogenetic data therefore suggest that Paraustrongylus evolved from an ancestor resembling Dessetostrongylus. KEY WORDS: Nematodes, Trichostrongyloidea, Paraustrostrongylus, ontogenesis, larvae, morphology, rodents. Transaartions of the Royal Society af S. Aust (1993), 17(L), 2736 ADULT AND LARVAL STAGES OF PARAUSTROSTRONGYLUS RATTI (NEMATODA: TRICHOSTRONGYLOIDEA) FROM RATTUS FUSCIPES by LIAN BEVERIDGE* & MARIE-CLAUDE DURETTE-DESSET+ Summary Bevermeoe, 1, & Durerre-Desser, M,-C, (1993) Adult and larval stages af Paraustrostrong ylus raiti (Nematoda: Trichoatrongyloldea) from Rattus fuscipes, Trans, R, Soc. $8, Aust, U7), 27-36 4 June, 1993, The morphology of the adult and the third and fourth larval stages wf Paraustrostrongylus rari from the native rodent Rastus fiscipes is described, with particular reference to the syntophe, as well as the tnechanis¢n of artechiment of the nematode to intestinal villi. The synlophe of te fourth larval stage hasan ublique axis «vf orientation, in contrast to the frontal orientation in the adult stage, and most closely resembles the synlophe of species of Dessetostrangylus parasitic in dasyurnud mursepials. Ontogenctic data therefore suggest that Purausirongyus evolved from an-ancestor resembling Dessetostrengylis. Key Worps: Nematodes, Trichostrongyloidea, Paraustrostrongylus, ontogenesis, larvae, morphology, rodents. Introduction The trichostrongyloid nematode subfamily Herpetostrongylinae occurs in the small intestines of Australian marsupials and is one of the few tichostrongylold groups in which there is apparently a close evolutionyry parallel between hosts and parasites (Durette-Desset 1982, 1985; Humphery-Smith 1983; Beveridge 1986). Three distinct lineages have been recognised among the cight component genera in Marsupials, each sharing a probable comron ancestry with Woolleya, a contemporary genus which occurs in dasyurid marsupials. Woolleva shares fearures in common with the genus Viaraaia, belanging to the family Viannaiidae, which is present in South American marsupials and rodents (Humphery-Smith 1983). Two of the herpetostrongyline genera, Austrostroneylus and Parausrrostrongwus, occurring in diprotedon! marsupials, with one species in the marsupial mole, Norerveres typhlops, and one in a rodent, Ratius fixectpes, are of particwlar morphological interest due to the development of paired lateral cuticular inflations, unique within the Trichostrangy- toidea, termed “floats” by Durette-Desset (1979). The evolutionary development of these floats was investigated in Austreastronyylux and in a related penus, Sutarostroneylus, (Beveridge & Durette-Desset 1986) and species with intermediale or primitive morphological features, that is either with a single float or without floats, were identified. This study suggested that Sutarostrongylus, parasitic in Thylogale spp., exhibited a number of primitive characters, ancestral to those seen in Ausirosirongylus, but provided no * De Melbourme, Parkville, Vic, 3052. ' Laboratoire de biologie peresilaire, protstolovie et hélminthologie Muséum national d"Histiare naturelle, él, rue Buffon, Paris. France. mmem of Veterinary Science, University of additional insights into the possible relationships between a postulated Mbolleya-like ancestor and Sutarostrongylus. Desyetostrongylas is one possible intermediary between Rbollepa and Sutarostrongylus (ste Beveridge & Durette-Desset 1986) as it has a synlophe. or camplement of body ridges, identical with that of Siarestrongylus except for the fact that the axis of oriettatian of the synlophe is objique in Dessetostrongylus but frontal in Sutarastrangylus, Humphery-Smith (1983) by contrast, placed greater emphasis on the frontal onentation of the syntophe of Austrestroneylus and derived it direcdy from an ancestral stale resembling that found in Maolleye sprenii. Cassone er al, (1986), studying new species of Rbolleya and Parricialinag from dasyurid marsupials confirmed the dire Hbolleya - Austrostrongylus relationship identified by Humphery-Smith (1983), but considered that Dessetostrengylus was a sister group 10 Austrostrongylas. All studies on the evolution of the Herpetostrongy- linae to date have relied on the comparative morphology of the adult nematodes and particularly on differences in the anatomy and orentation of the syniophe. The ontogenesis of larval stages is an important source of phylogenetic information mn the Trichostrongyloidea (Durette-Desset 1985), but has not been exploited in the case of the Herpetostrongylinae ather than in the case of Beveridgiella pearseni (see Humphrey-Smith 1980), because no life cycles are known. We decided to investigate the morphology of the various life-cycle stages of Paransrrostrangylus rat, the anly member of the genus to occur in a eutherian mammal (Obendorf 1979) to anempt tm obrain additional information on the evolution of the synlophe of the peous. Due to its abundance in Rartus fiuscipes and the eage with which Infected rats could be obtained and kept in the laboratory. # rutti was considered to 2x I. BEVERIDGE & M.-C. DURETTE-DESSET z 7 ay Ba mel jatar Taw! Ss ae Figs 1-15. Paraustrostrongylys ratti Obendorf: Adult. 1, anterior end, lateral view, dorsal aspect on left hand side; 2, anterior end, dorsal view, 3, apical view of mouth opening and lips; 4 optical transverse section through hexagonal buccal capsule, with dorsal tooth, 5, transverse optical section through anterior end of oesophagus: 6, anterior region, left lateral view; arrows indicate origins of ridges; 7, anterior region, right lateral view: arrows indicate origins of tidges: 8, bursa, lateral view; numerals indicate ray numbers according to Durette-Desset (1983); 9, bursa, ventral view; 10, gubernaculum, lateral view; ll, spicule tips, ventral view; 12, spicule tip, lateral view; 13, genital cone, ventral view of papilla 0; 14, genital cone, lateral view showing papillae 0 and 7; 15, fernale tail, lateral view. Scale Hines 0.01 mm: figs 1,2, 10, 1-14 to sane scale; figs 3-5 to sarne scale; figs 6.7 to same scale; figs 8,9 to same scale. Legend: a, amphid; d, deirid; e, excretory pore; 1, lip; p, posterior atrophic uterus; s, submedian papilla; sp, sphincter; 1, dorsal tooth; ve, vestibule: LIFE CYCLE STAGES OF P RATT! 29 be mom suitsble fot investigation than species occurring in marsupials, The morphological data presemted here also provide (he basis for subsequent ultrastructral studies. Methods Naturally infectex! rats, Rattus fuscipes (Waterhouse), were trapped at Blackwood, Victoria (37°29’S, 144°19"E), killed im the laboratory and the small intestine was divided into segments and opened in warm 0.09% saline The itnestinal segments were Placed im an incubator for two hours to allow Neinstodes to migrare ins the saline. Nematodes were then washed in saline and fixed in hot 70% ethanol, Small numbers of nematodes were fixed in 2.5% glutaraldehyde in phosphate buffer at 4°C. Adult, fourth and parasitic third-stage nematedes were cleared in Jactophenol and examined, using Nomarski interference contrast microscopy. Transverse sections of jhe trody of male and female nematodes were cul using a cataract sealpel, mounted in tactophenol for examinauon and orented using the methods of Durette-Dessei (1971). Apical views of the anleriar extremity were made by similar means, Specimens fixed in glutaraldehyde were embedded in resin, Sections cul ata thickness of |j.m were stained with toluidine blue and were used to confirm mormphelogiwal featires seen in hand-cut section, Additional specimens were dehydrated in a praded ethanol series, dried in a critical point drier, coated with pold and examined with a Siemens Autoscan scanning electron microscope. Ridges of the synlophe were numbered in an anticlockwise fashion beginaing with the lefi-ventral ridge, in order to demonstrate homologies between stages. The numbering system for the ursal rays and papillae: follows that of Duretle-Desset (1985), Faeces from naturally infected rats were collected, mixed with an equal quantity of charcoal and cultured on moist filter paper in Petri dishes at laboratory temperature. Frve and eight days later, larvae emerging from the faccal-charcoal mixcure were collected in distilled water and concentrated by sedimentation. Third-stage larvae were examined live in water as well as after having been immobilised by heating. Some larvac were killed in hot 70% ethano) and cleared In glycerol by transferring to a mixture of 70% ethanol and glycerol and allowing the cthanol to evaporate. Measurements were made either with an ocular micrometer or from drawings made using 4 drawing tube and are presented in the text in millimetres as the range followed by the mean in parenthescs. Morphological terminology for the synlophe follows that of Durerie-Desset (1985), A drawings are oriented with the dorval aspect uppermost and the left hand side of the nematode body towards the left margin of the page, Parasitic third-stage larvae of Aoorani were distinguished from the synhospiralic nematodes Nippustrongylus magntes and Odilia bainae by feanures of the tail and cephalic extremity, based on material obtained from monospecific experimental infections with each of these two species. To obtain a monospecific infection of adult F ratri, one naturally infected R. fuscipes was killed, all P rare’ in the small intestine were sorted while being maintained in warm saline, and were transferred by enlerotomy to a laborgtory-reared R, fuscipes under general anaesthesia. Four weeks Later, the recipient rat was killed, the intestine removed and. 10% buffered formal saline injected rapidly into it to fix nematodes in: site, Fragments of intestine with worms attached subsequently were dissected and prepared for scanming electron microscopy us deseribed above, Additional segments were dehydrated; embedded in paraffin, serially sectioned at a thickness of 10 um, and the sections stained with haematoxylin and eosin fur histological examinaticus. Results Adult mile snd female nematodes, fourth-stage larvae and two parasitic third-stage larvae were examine! morphologically undes the light microscope, 33 well as third-stage larvae cultured from faeces, Light microscopic observations were compared with scanning electron micrographs of the adults. Poraustrastronegylus nati Obendarf, 1979 FIGS 1-38 Description: Adult: small nenxstodes, red when live, spirally coiled in 3-5 tight coils, ventral sucface innermost, capable of uncoiling and beconung straight when maintained in warm isotonic solutions. Cephalic extremity with prominent cephalic vesicle, 20-30 fine transverse ridges on vesicle (Fig. 1). Mouth opening, surrounded by four sub-median papillac and two amphids, no fabial paplae visible; six rounded lips project into mouth opening. Buccal capsule prominently developed, heavily scleratised, circular to sub-hexagonal in transverse section (Fig. 4), walls, arched inwards, increase in thickness towards base, lumen increases in diameter twards base; prominent tooth projects inwards from dorsal Jobe of oesophagus; sub-ventral teeth absent, esophagus elongate, slender, clavate, widened at anterior extremity; excretory pore yanable in position, usually anterior, occastonally posterivr, to vesophago-intestinal junction; deirid tiny, domed, al level of exuretory pore; nerve ring in mid- oesophageal region, visible in few specimens only. 30 1. BEVERIDGE & M.-C. DURETTE-DESSET “My Figs 16-23, Paraustrostrongylus ratti Obendort: adult. 16-22, transverse sections of body, 16-19, male, 2.3 mm long; 16, at posterior end of cephalic vesicle, 0.06 from anterior extremity; I7, in oesophageal region, 0.30 mm from anterior extremity; 18, in posterior half of body, 1.60 mm from anterior extremity; 19 in cloacal region, 0.10 mm from posterior extremity; 20-22, female, 2.6 mm long; 20, oesophageal region; 21, 0.50 from anterior extremity; 22, 1,90 mm from anterior extremity; 23, posterior end of male, ventral view, showing termination of ventral ridges (arrows) and termination of floats (f). Legend: d, dorsal; J, left; r, right; v, ventral; ridges are numbered (1-7) in an anticlockwise direction from the left ventral tidge. Scale line: 0,01 mm. LIFE CYCLE STAGES OF PRATT Hv Body covered with numerous fine transverse annularions (Fig. 36); two lateral, fluid-filled cavities (= floats) present on either side of body, extend from unmediately posterior to vesicle to posterior region of nematode. Synlophe composed of three ventral ridges (1-3) (Pig. 36), oriented From right wo left, diminishing in size from left to right; right Sloat with pwo dorsal (5,6) (Fig. 35) and single ventral (4) ridge directed towards left dorsal, ridges b-3 commente posterior to vesicle; left floal, with single ndge (7) (Fig. 35) directed perpendicular Lo body, commences posterior to mid-oesophagus; right float with two dorsal (5,6) and one yentcal (4) ridge; ridec 6 commences postenor lo vesicle, followed by 5 then 4 in mid-oesophageal region. Male (measurements of 10 specimens). Total lenge LYB-2.66 (2.30); maxitnutn widih (without floats) ,050-0.070 (0057); cephalic vesicle 0055-0065 (0054) long) oesuplragus 0,27-0.30 (0,28) lung, nerve ning -cire@ 0.15 from anterior extremity; excretory pore 0.21029 (0.25) from antenor extremity; deitid 0.22-0,30 (0.26) from anterior extremity: spicules 0.23-0.32 (0.27) long; gubernaculum 0,025-1045 (0.037) long, Synlophe; ventral ridges ard right float femiitiate near anterior extremity of spicules: teft Moat continues tn leve) of gubermaculum,. Bursa symmetri- cal, lobes indistinct, dorsal labe not separated fron lateral lobes; rays 2 to 6 of bursa grouped im patter of 3-2 (sensu Durette-Desset 1983) Ur 1-2-2 allowing for highly divergent ray 2; ray 2 slender, short, divergeit, nol reaching margin of bursa; rays 3 to 5 more robust, directed postero-faterally, of appreni- mately equivalent size, not quite reaching margin of bursa, my 6 shares common origin with ray 5, robust, blunt, directed posteriorly, reaches margin of bursa; dorsal trunk separate (rom lateral trunk; ray 8 uriges from dorsal trunk, slender. does not reach margin of bursa; rays 9 slivhily asymmetrical, shor, slender, arise Close to origin of ray 10; ray LO stout, divides hear extremity inte + branches; outer pair of branches more rubust; fmal branches do not reach margin of bursa. Spicules simple, elongate: alate; anterior exthemitles irregularly knobbed, distal ups jouted: each spicule with fine, spiniform ventro-lateral branch arising in distal 1/6-1/7 of spicule; spicule tips surrounded by expanded sclerutived [ange in \lorso- ventral view; each main branch of spicule terminates in two fine spiniform projections within flange; pubernaculum elongate, rectangular in dorso-ventral view, composed of two layers; genital cone heavily selerotised, complex, cumical in shape, c. 1020 long, base D020 wide; papilla 0 at tp of ventral lip of genital cone; paired papillae 7 on dorsal lip of cone. Female: (measurements cf (0 specimens), Total length 2.48-2.95 (2.70): maximum width (without floats) 0.060-0.070 (0.066), with floats circa (110; cephalic vesicle (060-0070 (0,063) long, oesophagus 026-032 (Q.29) long) nérve fling circa O18 from anterior extremity; excretory pore 0.21-0.27 (1123) fram anterior extremity; deind 0.22-0.26 (0.24) from ameriog extremity; bail OO8-O.U (0.10) lung; vulva to posterior extremity 014-021 (0.17); egg 0,065-0.080 (0,074) by 6.035-0.045 (0.038). Synlophe: ventral ridges extend to vulva; floats disappear in region of uterus, approx, 0.44 from tail, Tail extremely long, conical; vulva immediately anterior to uterus, opening to exterior on slight prominence; female genital system monodelphic though with postendr uterus patent and ovary persisting in vestigial form: vagina, vestibule and sphincter crea 0.06 long; infundibulum short, circa O04 long, prodelphic, leads to elongate uterus containing 1-4 eggs; eges thin-shelled, ellipsoidal. Fourth stage larva: Small nematodes, spirally coiled in 34 cous, ventral surface innermost, cephalic vesicle absent; mouth opening surrounded by four submedian papillae and two wmphids; lips absent, Hupcal capsule sub-cylindrical, heavily sclerotised. teeth absent. Oesophagus elongate, clavate, nerve ring in mid- vesophagesl region; excretory pore in region of oesophago-intestinal junction. Synlophe: body floats absent; five ridges; three ventral ridges (1-3), anenied from right to left. diminishing in size fron Jefi to right, ventral ridge (4) on right hand side, oriented from right to left; single dorsal ridge (5) on right hand side, onented dorsal from right fo lefi; orientation of synlophe oblique from right ventral to left dorsal at about 60° to saginal axis. Tatl elongarm, conical. Male (measurements of five specimens), Total length 1,09-1,76 (1.44), maximum width 0.033-0.046 (0.040), oesophagus 0.21-0.28 (0.26), tail 0.049-0,085 (0.066), Female (measurements of five specimens). Total length 1.47-2.00 (1.80), maximum width 0.030-0,052 (0.044), oesophagus 9,25-0.29 (0.27), tail 0,052-0,143 (0,082), Specimens with developed genitalia show distinct posterior uterus, recurving into short ovary (Fig, 32), Third stage larva» Parasitic: Two parasite third stage: larvae were recorded, but detailed morphological comparisons were possible from one only. Small nematode, 0.76 long, spirally coiled in three coals; buccal capsule cylindrical, very lightly sclerotised; oesophagus slender, clavate, 0.18 bong: excretory pore in posterior pesnphageal region, 0.12 from antenor extremity; tail elongate, conical, with dorsal and ventral projection. Syniophe composed of rwo pairs of alse beginning on lateral aspects of body, perpendicular to body surface; towarde mid-region of body, alae gradually shift in position to dorsal and ventral; m md hody region, the left ventral parr are larger with one iu almost a mid-ventral position, and second ridge to 32 I. BEVERIDGE & M.-C, DURETTE-DESSET Figs 24-32. Paraustrostrongylus rarti Obendorf: larval stages. 24-27 third. larval stage cultured in vitro; 24, entire larva, lateral view; 25, cephalic extremity, lateral view; 26, tail, lateral view; 27, transverse section in mid body region; 28, transverse section of parasitic third-stage larva from small intestine; 29-32, fourth-stage larva; 29, anterior end, lateral view; 30, buccal capsule, lateral view; 31, transverse section of larva in mid-body .region, arrow indicates oricntation of synlophe; 32, tail of female fourth stage larva with posterior branch of genital system. Scale lines: fig. 24, 0.1 mm; figs 25-32,.0.01 mm; figs 25-26 to same scale. Legend: p, phasmid; u, posterior uterus and 0, oyary; ridges are numbered in an anticlockwise direction from the Jefi-ventral ridge. LIFE CYCLE STAGES OF P RaTiyY as cme sick: of i; two dorsal alac smaller, one almost doisal Lo position, Other lo one side of it. freedivine: (measurcinents of five specimens). Mender, elongate larvae, 0.50-0.55 (0.53) long, maxima width 0.016002) (0.018); baceal capsule cylindrical, ¢. 0.005 long, 0.002 wide, continuous with selerotised internal lining of anterior oesophagus: oesophagus slender, 044-016 (0,15) long; nerve ring CLOTS-0.083 (0.082) from anterior end; excretory pore CHOR 1.099 (0,096) fromanterior end; intestinal cells filled with granules, number of cclly not ascertained: genilal primorhium avoid, 0.008-0.013 (0.010) by TODS-O008 C007), 128-0.33 (0,31) from anterior extreme, tail clomgale, conical, 0.065-0,089 (0,075) lon, with dorsal and ventral spike close to tip; larva with four longitudinal alae, two on cach side of body. in Jateral position, whois perpendicular to holy wall, ditachment ta the intestine! nurse Adult nematodes are coiled ventrally around pitestenal villi (Pigs 33, 37) usually with the tail near the distal Hip of the villus; when fixed. the coils of the body are mameained (Fig. 34). At the site of altachntent, nenalodes compress the willl (Figs 37, 38) and, although they generally retract from the site of attachment when lixed, the impressions of the ventral ridges remain in the intestinal epithelium (Fig, 38). Changes in the epithelium at the site of attachment include cuboidal to squamous cpithelial cells, clongahon (= flattening) of nucler associated with the change 0 ao siyuumous cell type, loss of cytoplasmic differentianon and the loss of the brush border of muicavillt, Although sormetimely squamous, no defects were detected in the epithelium, No marked iiflanimatory changes were detectel but there were significant numbers of mononuclear cells, macrophages und lymphocytes, present in the Bani propria together with a small number of cosinaphils Discussion Morphology of the adult The description of the adult provided here supplements the original description by Obendort (979), which was found to be accurate in all essentials. Obendort (1979) however, did not provide an apical view of the untenior extremity and provided only a single, anonenied drawing of the sy nlopbe of the adult. In the present examination, the synlophie 1s described in detail, including the oripins and terminations of the body ridges, Duretre-Desset (1979) and Beveridge & Durelte-Desset (1986) have shown that the number of bendy ridges changes in the posterior region of the body in several species of Axytresrrongyluy and Paraustrastrangylus, but comparable data were lacking for P. rat, This study demonstrated that in P rarti, the principal nelges arise tm the oesophageal region and persist to the bevel of the spicules in the male and to the level of the vulva in the femile. In the male, the bef, but nol the righe Moat extends almost to the level of the bursa, as iv does also in P rrichosarl and in A. safovialas, the latter being d spoctes which possesses only one float F retti also resembles A. safestatus in having three rather than the four ventral longitudinal ridges present momost other members ol tfese gener, The three principal ventral ridges arise Close to the cephalic yesiele, while the ridges on the floats arise somewhat more posteriorly, with the ridge on the left float arising mid-way between the vesicle and the exeretory pore and the two dorsal ridges on the right Nout arise in the anterior oesophageal region, The ventral ridge arises midway between the cephalic vesicle and the exeretory pore. The onginy and lerminations of ridges haye been repomed in few confamilial species, but have been shown to be of considerable taxonomig use at the species level in genera such as Nerriaredirus (see Lichrenfels & Pilitt 1983). Preliminary observations by Beveridge & Durette-Desset (1986) on species of Adstrostronyylus suggest that this may be the case in the Merpetostrongy fae, but features have been described in too few species to allow any firm conclusions to be diawa, An imeresting feature of the morphology of / farri, noted by Obendor! (1979) was the presence of 4 vestigial posteriog branch to the female genial system. His observations were confirmed inthis study, and the same structure was also seen in the Jourth larval stage. The genera Austrostrongylus and Paraustrostrongylay wre considered to be very closely related but can be separated on the basis of the presence of a sclerotised genital cone in: Paraustrostrongylis, the absence of ventral teeth in Pirastrostrongylus and the position of ray 2 (Beveridge & Durette-Desset 1986), Species of Paraustrostrong\las are invariably, monodelphic whale most species of Austrostrongylues are didelphic, The evolution of monodelphy has oceurred repentedly in the Trichostrongyloidea (see Duretie-Desser 1985) and the vestigial posterior uterus. in P rari provides an obvious connection between the mortxtelphic and didelphig dorms seen in these two elosely related genera, [n other trichostrongyloid genera such as Neohelignonedla, the posterior uterine branch imuy persist in the adult nemiutode, but does so only as a small collection of cells, posterior to the vulva (Durette- Desset & Cassone 1987), rather than the almost fully tormed bul diminutive posterior branch seen in P mare. Anachment ta villi Results presented here indicate that? rani attaches to intestinal villi by coiling spirally qroond them, as in certain other tachosirongylod neraiodes (Duretie Desset 1985), The ventral body ridges clearly press ime the intestinal epittelium, and may theref ire assist 34 I, BEVERIDGE & M.-C. DURETTE-DESSET the nematode in maintaining its attachment to the villus. The ventral surfaces of both body floats are also in close apposition to. the epithelium and effectively increase the surface area of the nematode in contact with the intestinal epithelium. The ventral ridge of the right float (4) and to some extent the latero-dorsal ridge (4) of the right float (5) also cause indentation of the epithelium, and may therefore also assist in attachment. Once in place on a villus, dorsal ridges would seem to have little function in attachment, and one of the features of P ratti is that it has few dorsal ridges. However, when observed in warm isotonic solutions, the nematode is capable of uncoiling completely, and evidence from the localisation of experimentally transplanted nematodes indicates that they are capable of migration within the intestine as is the case with CP" _.. aT — ~ a oe ye Figs 33-38. Paraustrostrongylus ratti Obendorf, 33-36, scanning electron micrographs. 33, entire nematode coiled spirally around villus in small intestine; 34, entire nematode, 9, showing body coils (4); anterior end to left; 35, dorsal surface of nematode showing ridges 5 and 6 of dorsal aspect of right body float and ridge 7 on lateral aspect of left float; 36, ventral surface of posterior region of body showing ridge 7 on left body float and ventral ridges | and 2; 37, longitudinal section through villus showing posterior part of nematode coiled around yillus, with anterior end extending to left of field; 38, histological section through villus at point of attachment of P. ratri, with nematode retracted, leaving sites of indentation (arrows) of ridges in epithelium. Scale bars: figs 33, 34, 37, 0.1 mm; figs 35, 36, 38 0.01 mm. LIFE CYCLE STAGES OF Fl RAITI a8 other trichostrongyloids (see Crofl & Ma 1977), The dorsal ridges may therefore be of use daring nematode migrations wijhin the small intestine, when they are uncoiled and are moving between villi, Morpitology of Larval surges The morphology of the buccal capsule of the fourth stage of P ratii resembles very closely that of the fourth stage of B. pearseni (sec Humphery-Smith 1980) In peramelid marsupials and Globocephaloides trifidospicularis: (see Beveridge (979), a species parasilic ln macropodid marsupials. The Globocepha- loidinae, to which the latter genus belongs, was placed Within the Herpetostrongylidae by Durette-Desset (1983) bated on features of the bursa and buccal capsule, Since members of ihe sub-family Jack a synlophe, their precise alfinities have not been established, However, the similaritles between. the fourth stage larvae of G, trifidospiculanis, & pearsuri and P rat provide additional evidence that the two subfamilies are related. Only two parasitic third larval stages of A rated were found, However, they were identified by the characteristics of the tail, which was identical to that of larvae cultured from faeces. Both parasitic larvae were spirally coiled, as are the fourth stage and adult The pairs of alae Which were clearly lateral in position at (he anterior and posterior extremities of the body were slightly shifted in position in the mid-body region 26 that the larger, left pair were almost ventral in position, while the smaller, right pair became almost dorsal in position. This gradual shift in ridge pasitlon and the hypertrophy and ventral shift of one pair of alae presumably aids jn attachment, as is the case in the adult, Two pairs of lateral alac were evident in the free-living third larval slage but they remained in the Jateral position throughout the length of the nematode body. Several other teichostrongyloids have paired lateral alae tn the third stage larva (Ecker & Schware 1965; Duretie-Desset & Cassone 1987), The apparent change tn Orientation of the ndges in the mid-body segion during the initial parasitic phase of the life cycle appears to assist the nemtapode in sttachiny to villi, bur (he mechanism by which this might occur is unclear. Usually there is no change in the synlophe unless a Moult Occurs, bul lf this ingiance, the change in position of the alae is vistble in the entire nematode as well as in sections, Additional observations are clearly needed co confirm the results reported here. Evolutionary relationships The description of the fourth larval stage of F rarsi feovides additional insight into the evolution of the Herpetustrongylidac. The type ef information which can be provided by the larval synlophe has been discussed by Duretie-Desset (1985), Generally, the larval synlophe demonstrites primitive features compared with thar of the corresponding adult and frequently resembles the adults of other, related genert thereby allowing phylogenetic reconstructions, In the ease of P revi, as in other tichostrongyloids, the larval synlophe differs markedly from that of the adult_ The larva lacks floats; in the larva, the axis of orientation is oblique rather than being frontal, as occurs in the adult and the synlophe has iwo fewer ridges, lacking those found og the left flout and on the dorsal right float of the adult, The farval syolophe of P neni most closely resembles that found in the adults of Swarostronzylus and Dessetostrongylus but differs from that of Sutarcstrongylus in baving one. rather than two dorsal ridges, ane sn onentation since the synlaphe of Sutarasrronevius has a frontal orientation, simular to thar af the adult Prati. The oblique orientation of the synlophe of the larval stage of Ft rare| resembles that of Desselastrongylus, from which it differs only in having fewer dorsal ridges. The homology of ridges between. adult and larva here considered wo be most likely is that ridges | to 3 of the larva correspond with ridges 1 to 3 of the adult, while the two additional ridges of the Jarva (4 and 5) correspond bo two of the ridges on the nght float of the adult (4 and 5). An alternative possibility would be that the first four ndges from left to right correspond to the four yentral ridges present in rnost species af Austrastrangylas and some species of Puruustrestrongvlus, wilh the filth ridge corresponding to one of the ridges on the right float. This interpretation invalves postulating the loss of # yentral ridge in adult P rani and the appearance of two rather than ane new ridge for the right float, The more parsimonious of the two hypotheses has been chosen here, 11 is also consistent with the hypothesis of Humphery-Smith (1983) that species of Wooileva with three left-ventral, obliquely atiented ridges, were the likely ancestors of Dessetosirongylus. The suggested intermediate farms in the proposed lransition series for the evolutionary lineage hetween Uholleya and Austrostnongylus/Paraustrastrong wis have been Heveridgiella (see Durette-Desset 1982; Humphery-Smith 1983) and Dessetrstranewus (see Beveridge & Durette-Desset 1986). Potricfaline, an additional possibility, has a frontally symmetrical synlophe. while both Beverideielfa and Dessete- Strong ylug have an oblique orientation to the synlophe. Patriciatina was considered to have been derived from Beveridgielia by Humphery-Smith (1983) and Cassone et al (1986). Beveridgiella has 4 greater number of dorsal ridges than Dessetestrongyius, and increases in the number of ridges oocur in # number of evolutlonary lineages within the Trichostrungyloiden (Durette- Desset 1985), suggesting that the synlophe in species af Reveridgicifa is probably derived from a Dessetostrangylus-like ancestor. This i fact is shown in the larval stage of 8 pearseil (see Humphery-Smith 36 I, BEVERIDGE & M.-C, DURETTE-DESSET 1980), which has a synlophe close to that of Dessetostrongylus. The synlophe of the fourth-stage larva of P rarti differs from that of adult Dessetostrangylus moorhousei only in lacking an extra dorsal ridge, Because of the close correspondence between. their synlophes, it appears likely that Paraustrostrongylus evolved from an ancestor vesembling contemporary species of Dessetostrongylus, thus supporting the hypothesis proposed by Beveridge & Durette-Desset (1986) and Cassone et af, (1986), Acknowledgments Rats were obtained under permits from the Victorian Department of Conservation and the Environment (90-053 and 91-095), We wish to thank Christine Andersen for expert technical assistance, Lilliana ‘Tatarczuch for ibe preparation of the thin sections and Dr D. M. Spratt for comments onthe manuscript. This work was supported financially by the Austrahan Research Council, References Bevenipce, 1. (1979) A review of the Globocvephaloidinas Inglis (Nematoda: Amidostomatidae) from macropadid marsupials. Aust J. Zool, 27, 151-175, (1986) Coevolutionary relationships of the helminth parasites of Australian marsupials. pp 93-117. In “Coevulutiun and Systematics” A, R. Stone & D. L, Hawksworth (Eds). The Systematics Association. (Clarendon Press, Oxford) & DURETTE-DESSET, M.-C, (1986) New species of Austrostrongylus Chandler, 1924 (Nematoda, Trichostrengylvides), from Australian marsupials, with a redescription of A, minutus Johnston & Mawson, 1938, and description of a new genus Sutarastrongylus. Bull. Mus. natn. Hist. nat., Paris 4éme sér. 8, 145-170. Cassone, J., DURETTE-DesseT, M.-C...& PRESIDENTE, P, J, A. (1986) Nouvelle hypothése sur I'évolution des Herpetostrongylinae (Nematada, Trichostrongyloidea) parasites de Marsupiaux ausiraliens. /bid, 267-283. Croit, N. A. & Ma, K. (1977) The location of parasites within their hosts: the influence of surgical manipulation of the intestine and blood supply on the dispersion of Nippostrongylus brasiliensis in the rat. Int. J. Parasitol. 7, 21-26, Durette-DesseT, M.-C. (1971) Essai de classification des Nématodes Heligmosomes. Correlations avec la paléobiogeographie des hates. Mem, Mus. natn, Hist. nat,, Paris, sér A, Zoologie 69, 1-126. (1979) Compléments morphologiques a letude des enres Austrostrongylus Chandler, 1924, et raustrostrongylus Mawson, 1973 (Nematoda, Trivhostrongylidae). Bull, Mus. natn, Hist. nat., Faris, d2me sér 1, 1023-1030, —____ (1982) Relations hdtes-parasites chez les Trichostrongyloides. Mém. Mus. nam. Hist, nat., Paris ser. A,, Zoalagie 123, 93-100. _—_—. (1983) CIA Keys to the Nematode Parasites of Vertebrates, No, 10 dn R, C, Anderson & A. G, Chabaud (Eds). Keys to genero of the superfamily Trichostrongyloidea. Commonwealth Agricultural Bureaux, Farnham Royal, pp.6s. (1985) Trichostrongyloid nematodes and their vertebrate hosts: reconstruction of the phylogeny of parasitic group. Adu Parasitol. 24, 239-306. & Cassone, J. (1987) Sur deux Nematodes Trichostrongyloides parasites d'un Muridé africain. TT - Chronologie des cycles, description des stades larvaires et des immatures, Ann. Parasitol. Hum, Comp. 62, 133-158, Ecxerr, J. & SCHWARZ, R. (1965) Zur Struktur der Cuticula invasionsfhaiger Larven einiger Nematnden, Zeit, f Parasitenk. 26, 6-142. HuMrPHERY-SMITH, J. (1980) Beveridgiella n. gen., Dessétostrongylus 0. gen. (Nematoda, Trichostrongyloidea} parasites de Marsupiaux australicns: Bull, Mus. natn. Hise. nat., Paris, déme sér. 2, 999-1012, (1983) An hypothesis om the ¢volution of Herpetostrongylinae (Trichostrongyloidea: Nematoda) in Australian marsupials, and their relationships with Viannaidae, parasites of South American margupials, Anse. J. Zool. M, 931-942. LicuTenre.s, J. R. & Piuirr, P. A. (1983) Curicular ridge patterns of Nematedirys (Nematoda: Trichostrongyloidea) parasitic in domestic ruminants of North America. with a key to species, Proc. Helminthol. Soc, Wash. 50, 261-274. ORFNDORF, D, L. (1979) The helminth parasites of Rattus fuscipes (Waterhouse) from Victoria, including description of two new parasite species. Aust. J. Zool: 27, 867-879. A NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTIA (CESTODA: TRYPANORHYNCHA) FROM AUSTRALIAN SHARKS, AND DEFINITION OF THE FAMILY GRILLOTITDAE DOLLFUS, 1969 BY R. A, CAMPBELL* & I, BEVERIDGEF Summary Two new species of trypanorhynch cestodes Grillotia amblyrhynchos sp. nov. and Pseudogrillotia spratti sp. nov., are described from the spiral valves of carcharhinid sharks in Australian waters, G. amblyrhynchos is distinguished from congeners by possessing seven hooks per principal row, four intercalary hooks that merge with a wide longitudinal band of small hooks on the external tentacular surface, and a basal armature with microhooks on the external surface. Pseudogrillotia spratti sp. nov. is the fourth species in the genus and differs from congeners in the absence of a basal swelling, lack of a pars post-bulbosa, 9-10 hooks per principal row, absence of a band of hooks from all but the basal region of the tentacle and a single row of intercalary hooks. The subfamily Grillotiinae Dollfus, 1942 and the family Pseudogrillotiidae Dollfus, 1969 are discussed. The subgenus Paragrillotia and family Pseudogrillotiidae are rejected. Grillotiidae Dollfus, 1969 is formally defined for the first time and the three genera, Grillotia Guiart, 1927, Progrillotia Dollfus, 1969 and Pseudogrillotia Dollfus, 1969 are admitted with revised diagnoses. KEY WORDS: Grillotia, Pseudogrillotia, cestode, new species, Australia. ‘Dransactions of the Royal Sactery of S Awss 993), WIE, 37-46 NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTIA (CESTODA: 'TRYPANORHYNCHA) FROM AUSTRALIAN SHARKS, AND DEFINITION OF THE FAMILY GRILLOTIIDAE DOLLFUS, 1969 by R. A. CAMPBELL* & |. BEVERIDGET Summary CampPaeLe, R, A, & Bevirmce, |, (1993) New species of Grilloria and Pseudoprillotia (Cestwda; Trypanorhyncha) from Australian sharks, and definition of the family Grillotiidae Dollfus, 1969. Trans. X. Soc, §. Aust. 117(1), 37-46, 4 June, 1993, Two new species of trypanorhynch cestodes Grillotia amblyrhynchos sp. novo and Pyeudogrillutia spratti sp. nov., ate described from the spiral valyes of carcharhinid sharky in Australian waters. G. amblyrhynchas ix distinguished from congeners by possessing seven hooks per principal row, four intercalary hooks that merge with a wide longitudinal band of sinall hooks on the external tentacular surface, and a basal armature with microhooks on the external surface. Pseudogrillona spratd sp. nov. is the fourth species in the genus and differs from congeners ip the absence of a basal swelling, lack of a pars post-bulbosa, 9-10 hooks per principal row, absence of a band af hooks from al! but the basal region of the tentacle and 2 single row of intercalary hooks. The subfamily Grillotiinae Dollfus, 1942 and the family Pseudogrillotiidae: Dollfus, 1969 are discussed, The subgenus Prragrilloria and family Pseudogrillotiidae are rejected. Grillotiidse Dollfius, 1969 is formally defined for the first tume and the three genera, Grilloda Guiart, (927, Pragrillotia Dollfus, 1969 and Psewdogrilloria Dollfus, 1969 are admitted with revised diagnoses. Key Worps, Grillotia, Pseudogrillotia, cestode, new species, Australia, Introduction Few records of Griflotia or its relatives exist from studies of the cestode parasites of Australian fishes. Three Grillofia species have been described from larvae taken from Australian teleosts, one by Shahsarorn & Lester (1982) and two by Sakanari (1989), The genus Pseudogrillotia was not listed as occurring in the Australian region in a checklist of the parasites of fishes by Beumer ef a@/, (1982). A new species of Prendo grillotia trom a carcharinid shark reported herein therefore constitutes the first record of this taxon in the Australian region and a tiew species of Grillotia represents the first description of an adult of this genus from the region, Materials und Methods Cestode specimens were fixed in 10% formalin and lransferred to 70% ethanol for storage. Tentacles were dissected free and mounted in glycerine jelly or balsam to facilitate examination.. Whole mounted cestodes were stained with Celestine Blue, dehydrated in a graded ethanol series, cleared in clove oil and mounted in Canada Balsam. Terminology and numbering of tentacular hooks follows Dollfus (1942), Measurements are given in micrometers as the range followed by the mean in parentheses, unless otherwise * Department of Biology, University of Massachuserts Dartmouth. North Dartmouth, Mussachusctts 02747 U.S.A, ¢ Pppartanent of Veterinary Science, University of Melbourne, Parkville, Vic 3052, indicated, The number of measurements is indicated as (n). All specimens were measured, Type specimens have been deposited in the South Australian Museum. Adelaide (SAM). Drawings were made with a drawing attachment on an Olympus BH microscope. [n the figures, vitelllne follicles ure shown only along the lateral margiris of proglotides for the sake of clayity. Grillatia amblyrhynchos sp. nov. FIGS L9 Types; Holotype: from spiral valve uf Carcharhinus amblyrkyachos (Blecker, 1856}, Townsville, Queensland, 4,x7i,1985, coll, B, G. Robertson, SAM V4213; paratype: | specimen, same dare, SAM HC233372. Description: Moderate sized worms, wtal length up to 30 mun. Scolex (n=2) long and broad, 3.9, 4.2 mm long, 1.18 mm wide at level of bulbs. Two broadly oval, patelliform bothridia, 710-900 (780) long by 1070-1180 (1120) wide, notched on posterior border, margins only slighdly thickened and curved medizlly, posterior and lateral borders frec; adherent surface spinose with a barrow modian fissure c. 266 long, paralleled by four smaller fissures on either side al regular intervals (Fig. 1}. Pars vaginalis 2.56, 2.74 mm long: Tentacle sheaths spiral; prebulbar organs distimct. Bulbs 140-1180 long by 232-264 in diameter; retractor muscle originsies at midlength of bulb. Pare post-bulbosa lacking, Ratio of pars bulbosa to pars vaginalis 1:2.3 R. A, CAMPBELL & I, BEVERIDGE mt ee WF Aye yeaa BRO a 0. SONNY KY or , aa a ae a 7 7 Nant 4 ay Shey \ P Wy / i 4 VIA Ue tN OMI JOOS Db) SSS 4. Grillotia amblyrhynchos sp. nov; 1, scolex; 2, immature proglottis showing icstes; 3, mature proglottis, vitellaria shown only at margins; 4, bulb and prebulbar organs. Scale lines: 1-3, 1 mm; 4, 0.1 mm, Figs 1- NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTIA 39 to 1:2.4. Scolex ratio (pbo:py:phulb) 1:3.3:1.4- Tentacles short, length 1.40-1.46 min, tapering. tending to collapse; dianieter al base 110-114, diameter at mid- tentacle 95, diameter at tip 34, Armawwre heieroscambhous (sertsu Keveridge & Campbell 1989), heteromorphous, with band of small hooks in metabasal region; hooks hollow, Basal armature consists of numerous milcrohooks on extemal face between first four principal rows of hooks (Fig. 6); intemal face of basal armature consists of large books, reduced im size fram those of metabasal region. Principal hook rows alternate, consist of ascending half-spirals of seven hooks each; rows begin on imernal surface, merge with band of small hooks on external surface of tentacle (Fig, 8). Hooks of first proximal row (base of tentacle} reduced in size and number; hooks I(1') of first proximal row well separated but with transverse bases. Hooks 1(1") in all other rows, separated by distinct space, bases oriented longitudinally, rose-thom shaped, gradually increasing to maximum size at mid-tentacle then decreasing in size toward tip of tentacle, length 36-63 (52), base 94-46 (41), with distinct heed 4-1 (9) and me 4-13 (10), height 23-38 (32) (n=10). Hooks 2(2") rose-thorn shaped, similar to 1¢1 °) but smaller, length 36-49 (42), base 27-38 (31), heel 8 tne 9, height 19-29 (23), Hooks 3(3") erect, falciform, with extended heel 4-6 (5), toe absent, hook length 4-44 (39), base 19-23 (22), heaphi 22-30 (27), Hooks 4/4") and 5(5") falciform, erect slighily smaller, heel feduced, we backing: 444) length 29-42 (38), base 10-15 (13), heel 4, feng 17-23 (24); 5(5") Jength 23-25 (24), hase 11 heel 2, beight (7-24 (20), Hooks 6(6°) and 7(7°) spiniforn, smaller mn proximal rows 1-4, length 10-15, but with distinct toe 4-6; toe absent from Ihese hooks distal to row 4; 6(6") length 17-21 (19), hase 10, beight U-19 (15); 7(7') lengtir 15-21 (36}, base 8-10 (9), heawbt 13-18 (16). Hooks 6(6') ara 7(7") of principal rows much reduced and merge With band of hooks of extemal surface of tentacle. Viewed From bathridial or antibothridial surfaces of mmetahasal region, single row of 3-4 iuercalariny hooks between principal rows beginning with hooks 444’) of 3(5'). Second Intercalary row of two hooks may be presenl between principal rows in basal region of tentacle. Intercalary hooks spiniformn, length 1-19 (15), base 5-6 (5), beighn 6-'6 (10). Intercalary honk rows merge with irregularly arranged band of hooks on extemal surface, Band of hooks opposite proximal rows 1-10 small, wocinate, length 8-13 (10), base 4-6 (5), heignt 4, toe 4. Remainder of hooks forming band distal ra row 10 spiniform, length Le2t (8), base 4-4 (6), toe absent, heel 2, heaght 1-19 (17), First evidence of segmentation about 680 fron scolex, segments narrow, gradually increasing in length with maturity, becoming twice as long as wide when mature. Mature segments acraspedote, 1.92-2.36 11m (2.22 mm) by L40-1.50 ou (L4S mim}, terminal segmonts ¢. 2.50 mm by 1.44 mm Genital pores lateral, irregularly alternate, post-equatorial, 60-64 percent of segment length from anterior margin in mature segments, Cirrus lor possibly an hernaphnoditic) sac pyniform, 258-400 by 133-140, cortaining sinuous sperm duct. No armature visible. External seminal vesicle not observed. Testes subspherical, 200 by 180, numerous, about 376-532 (420) in mature segments, occupying all avatlable space medial to csmoregulatory canals including postovanan space. Vagina narrow, surrounded by gland cells, closely parallels posterior border of cirms sac, forming a bulbous dilatation ventral to proximal pole of cirrus sac. then tums posteriorly at Toidlime as chlated rube and cxicods to ovarian isthmus to form narrow fertilization dict, Ovary 144-176 by 656-728, subterminal, consisting of two small, transversely elongated asymmetrical lobes joined by ahort isthmus, poral Jobe smaller, Jobes subdivided into numerous lobules. Mehlis' gland immediately posterior to ovarian isthmus, c. 144 in diameter, Vitellarium follicular, forming layer encircling osmoreguiatory canals and Teprodactive organs. Uterns simple, median tbe. terminatiog in anterior one-fifih of segment. Ventral osmoregulatury canal largest, diameter 38; dorsal osmoregulatory canal extremely narro\y, sinuous, diameter 2. Brymology: The species is named afer its host, C. amblyrhyncitas. Remarks; Presently, (here are 20 valid species of Grillatia (see Sakanari 1989). G. amblyrhynchor sp. nov. has characlers of the subgenus Peragnfloria which Dollfus (19694) created to accommodate species of Grillotia in which the band of hooks could not be distinguished from tie adjacent intercalary rows (i.e. the inverealary rows merged wilh the band of hooks). Only two species have been aliributed to the subgenus Puraprillotla, G, (Paragrilloria) simmonsi Dollfus, 1969 and G, (Paragriliaia) rowel Campbell, 1977. Catra & Gavarrino (1999) provided a redescription of Rirntchobarkrlio situle Linton, 900 and showed that G. (F.} simmonsi Dolltus, 1969 was a junior synonym of it, They proposed the new combination G. stmilts but tid not discuss Dollfis’ (1969a) placement of the species, now recogmzed as G, similis, in tho subgenus Paragritlotia, Conceptually, the creation of Poragrilloria as a subgroup of Grillotia is useful except for the fact that @, simmons: does not fit the subgeneric definition. Griflata (P.) rowei was the first species described that met the subgeneric definition (Campbell 1977). G. amiblyritynehos sp, nov. is a second species with Uhis subgeneric character, The problem is not resolved by retuining G. simimenst as type of the subgenus Faragrivune Peragetiada, then, should be ciiminared on the basis of its chosen type species. R. A. CAMPBELL & i. BEVERIDGE Ute it bah ELE > Caw calG AG bs a a a pa S ue ae SSS DP (ec CO BaF So = c4 » (Qa oe ms Yew ts SP Gort. Of Zi VE fi 1 aS: f ise Ue hy ds fe wn gion, external face; 9, hooks | through 7 of principal row. Scale lines: rhynchos sp. nov.; tentacular armature: 5, basal region, internal face; 6, basal region, external face; vion, bothridial face; 8, metabasal re $ 5-9, Grillotia amblyi 5-8, 0.1 mm; 9, 0,05 mm. 7, metabasal re; Fig NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTIA 4 Both G. simifis and G rowel have six hooks per principal row untike G. amPlyrinwiches sp, nov. which has seven hooks per principal row. Nome of the remaining species of Grillotia has seven books per principal pow (Sakanari 1989). In G, similis und G. anbiyrtynchos there are four hooks pet intervalary row. There are two of threc interedlary hoaks per row in G, row? but the hooks have transversely elongated bases whereas those of G: amblyriynches are oval and have a longitudinal orientation. In G. similis, the first two intercalary hooks in each row (a and b) are twice the length of intercalary hooks c and d whereas in G, amblyriyiches the intercalary hooks are subequal, The proup of microhonks on the extemal face of the basal armature of G. amblyrhynchos is absent in G, similis and G, rewei. The presence of microhooks in the basal armature is typical of 13 species of Grillotia, Differences in the numbers of hooks across the bands uf these species is worthy of note, In G, similis only @ single hook file remains between the Lntercalary rows and poncipal rows on the external Face (See Fig, & Cara & Gavarrino 1999), In Fig. 19 of Dollfus (1969s) the band of hooks of G. simumwnsi is reduced to 1-3 hooks in width if one disregards all the principal hooks and |ntercalary hooks Therefore, the charactenstic feature for the subgenus Faragrilioria of having the intercalary rows Metge timpercepubly with the “band” of hooks is misleading because there is only a single file of hooks rather than # band exclusive of the intercalary and principal rows. In fact, this armature pattern demonstrates an intermediate pattern of symmewy bewween afypical betetoacanths and pocciloacanths with a single chainette: In G. rowel = single large hook and two small hooks are present on each side of the external face between the opposing Principal rows and in G. amblyrkynachos a band 6-8 spiniform hooks in width is present, The mature provketides of G, similis and G. umblyrlivnchos possess postovarian testes, a feature Dollfas (1969b) considered significant in separating Progrillaria (rom Grilloria. An heemaphrechtic sac was noted in G. eriniceus by Dollfus (942), and this feature has been conlinmed by us in additiumal material of the species, It was also shown to occur m G. similis by Cara & Guvarrine (1889) Beveridge & Campbell (3988) have pointed out the presence of an hermaphroditic sac in Numerous trypanorhynchs and have remarked upon the: generic and even familial significance of Ure genital terminalia, Pseudogeillotia spranti sp. nuv. FIGS 10-16 Types: Holotype: from spiral valve of Carehusrininues brachyurus (Guenther, |R7D), Tatham, WNS.W,, AL xtil984, coll, D.M, Spratt, SAM 4212; paratypes: four specimens, same data AHC HC23331. Description; Scolex craspedote, very Jong and slender, 62-84 mm (7.4 mmj n=5) Jong, maxinnunr width 088-116 mm (1.00 mm) an reginn of bulbs; two patelliform bothridia, 740-980 (830) wide, posterior margin indented, with a pair of fissures on lateral margins. Pars bolfridinlis 088-100 mm (0.95 mm); temacle sheaths regularly sinuous, pars vaginalis 367.1 mm (64mm); bulbs short and wide, 112-4.28 mm (1,21 mm) long by 220-340 (270) wide: prebulbar organs prominent; retractor muscle origin undetermined. Pars posibulbosa absent, Velum very shart, 30-80 (50). Scolex ratio (pb: pv: plulb) 1: 67 13. Tentacles up to 1.2 long, 70-140 (10) in diameter, basal swelling lacking. Armature hetercacagthous, heteromorphaus; heoks hollow. Principal rows of hooks alternate, ascending in half spirals of 9-10) hooks per row from external surface. Hooks )()) separated by a wide space, rose-thorn shaped, large, 70-100 (85) long. base length 60-70 (65), height 40-60 (50, n=10); hooks. 2(2') rose-thorn shaped, smaller, 60:75 (70) long, base 40-60 (45), height 55-80 (66). Hooks 3(3°) ty 9(9') elongate, slender, bases narrow, gradually diminishing in size, recurved at tip; 3(3') length and height 65-85 (77), base 15-35 (25); 4(4°) length und height 70-90 (84), base 15-30 (23); 5(5") length and height 65-80 (72); 6(6'} length and height 60-80 (65); 8(8") length and height 45-80 (62), 9(9°) length and helt 48-70 (62); 1009") length and height 44-56 (49). Single row of six intercalary hooks, 20-32 (23) long, present between principal rows alyned with principal hooks 5(5*) to 10010"). In basal region, principal hooks of bothridial surface larger than those: of antibothridial surface; ascending series of six hooks in file number 1, lengths 72, 55, 60, 70, 90, 95; hooks in file number 1’ fengihs 100, 90, 85, 90, 95. Internal surface of basal armature with band of smalt microhooks, 12-28 (16) jung, extending in closely spaced V-shaped rows along tentacle but eading at level of fifth principal rows; hand of amicrohooks diminishes from 12 hooks wide proxtmally at. first principal rows mp four hooks distally at fifth principal rows; band of hocks absent distal to lift principal row of hioks. Longest speciinen 50 tin, wilh about 65 seements; first mature segment about cumber 30. Segments acruspedote, largest mature segment 18 mni by 0,97 mm; genital pore m posterior half of segment, 980% from antenor margin; irregularfy alternate, Testes nuinerous (>300}, filling all available intervascalar medullary space, Linning thick pestovarian band, 40-60 (50, n=|0) in diameter; vss deferens visible at bevel of avanan isiiimus, coils extending just antenior to level of genital pore then laterally to enter cleus sec. Hermaphroditic sac weakly developed, abour 230 hy 0; external seminal vesicle present, weakly developed. Vagina mabiform, diameter about 20, joining R. A. CAMPBELL & I, BEVERIDGE 42 — Gas if Va ot ie rs Se a e Ak “| \ TS — Figs 10-1. Pseudogrillotia spraiti sp. nov.; adult worm: 10, scolex; 11, mature segment, Scale lines: 10, 1 mm; 11, 0.1 mm. NEW SPECIES OF GRILLOTIA AND PSEUDOGRIELOTIA 43 cirrus within hermaphroditic sac to form hermaphroditic dict. Vagina curves posteriorly bo ovarian isthmms, dilating to 40 but not forming distinct semittal receptacle. Ovary Gmmature, lobes 150 by #0 Mehlis’ gland posterior 03 iwvarian isthmus, 160 by 90; Uterine duct coils anteriorly nearly to level of hermaphrodittc sac; ierus tubular, ascends anterlorly nearly to antenor end of segment, utetime pore absent. Virelline. follicles form a circum-medullary sleeve around proglottis, Lateral osmoregulatory canals berely visible. Remarks; Dollfas (19699) created fhe perus Pseudegrillotie and the family Pseudogrillotiidae for F pleistacartha because of its combination af two bothridia, craspeiote scolex, metabasal armature with a band of nucnerous small hooks, Jack of a chainette and possession of a postlarval stage Carvasal et al. (1976) added a second species, P hasipinciata, from C. amblyrhynchos in Hawaii. P sprarti sp. nov. is similar lo F pleistacantha ia the number of hooks per Poincipal row and Jack of a basal tentacular swelling but can be distinguished from P pleistacantha by (1) Wick of 4 continucus band of small hooks im the metabasal armature and (2) lack of a pars post-bulbosa and velum. The new species can be distinguished fron P. basipwncrata by (1) the lack of a continuous band of hooks in the mnetabasal armature, (2) lack of a well developed basal armature, (3) absence of a basal swelling and (4) more hooks per principal now (9-1 vs. 6). P peruviune described by Escalante & Carvajal (1924) is most similar to P sprarti sp. noy. in having the bancl of hooks restricted tu the basal reglon of the temacle fut is readily distinguished from the new species by (1) possession of four intercalary hook rows between cach pair of principal rows compared with one intetcalary mow fy FP spraiti, and (2) differences in the stape of the large basal hooks (Fig. 14), Carvajal ev al. (1976) emended the family diagnosis of Pseudogrillotiidae with the description of the first species based upon an adult worm. The diagnosis is still in agreement with Bo aprotti and 2 peruvidna except that, 25 in species of Pterobothnidae, viz. 2 lurmaliensis Carvajal Combord & Campbell . 1976, the bard af hooks may be absent from chc metabasal armature. All other chanicters are in Agreement. Discussion ‘Trypanorhynch cestodes of the subfamily Grillotilixse Dollfus, 1942 and the family Pseudogrillotiidac Dolifus. 1969 have been reported from sharks ani skates in the Mediterranean, North Ajlantic, olf the west coast of buth North and South Amenvx, Hawaii and recently from Australian waters. Dollfus (1942) created the subfamily Grillotiinae within the family Lacistorhynchidae Guiart, 1927 distinguishing lt from other poeciloacanihs with two bothridia by (|) possession of a longitudinal band of hooke instead of a chainette and (2) presence of intercalary hooks between the principal rows. He further noted the possession of two notched patelliform bothridia, acraspedote scolex and lack of a postlaryal stage (without blastocyst), Ina sepunile paper, Doilfus (1946) created the subgenus Prygrillotia to accommoesiate x new species that lacked posturvarian testes and later (Doilfus 1969b) elevated Pragrilforie to generic status with an erended diagnoses, distinguished for Grilfotia by very long bulbs, no enigmatic prebulbar organs, absence of postovarian testes and a external band of hooks that 1s discontinuous but merges with the interpolated rows of hooks. Dollfus (19693) also created the subgenus Puragrilloria, and the familly Pseudogrillotiidue. Dollfus (19694) emphasized the distinction between the armatures of Grifloria and Lacistorlynchus by erecting a new famity Grillotildae because it separated genera with chainettes (Lacisarhynchns, Callitetrarhynchus, Flericeps, vide Reveridge & Campbell 1989) from genera with interculary rows and 4 longitudinal band of heoks on the external surface, i.e, Grilfotia, Dollfus (1969a) distinction stands m contradiction to the keys of Yarnaguti (1959) and Schmidt (1986). We agree with Dollfas (1969a) and recognize a further distinction of the armature possessed hy grilfotlids as being suitably accommodated by the definition of “atypical heveroacanthous" in which “there are more rows of hooks-on the external fice than on the internal face (of the tentacle)”. Our interpretation Of trypanorhynch armatures includes the Grillotiidae, Pierobothnidae, Molicolidae, Otobothriidae and Rhmoptericolidac as atypical hetercacanths thus separating those families with intercalary rows and longitudinal “bands” of hooks from theiz former classical inclusion with the pocciloacanthous families possessing chainettes, In our opinion the term “poeciloacanth" is best restricted to familics with chaiettes, as defined by Beveridge & Campbell (1989), and considered distinct froni those wilh longitudinal “hands”. Thus, the poeciloacanthaus (“chainette”) families are Dasyrhynchidae, Lacistarhynchidse, Hornelliellidae, Mustelicolidae, Gymnorhynchidae, and Mixodigmatidae (see Beverndge & Campbell 1989). Dollfus’ attempts (1969a) to subdivide species of Grillotia have been prompted by their rather homogeneous armatures and varied combinations of other scolex features, He created the subgenera Grillotie, Progrillotia, are| Peregrillotia, Poragrillotia, discussed and rejected above, remained 1 subgenus. Progrillotie Dollfus. 46 was elevated to generic stanis and o new genus and family, Pseudogrillotia a4 R_ A. CAMPBELL & |. BEVERIDGE | WRENS } WS i s Figs 12-16, Pseudogrillotia spratti sp, nov.; tentacle armature; 12, metabasal armature, external surface; 13, metabasal armature, external surface showing origin of principal rows; 14, basal region, internal surface on right side and bothridial surface at left; 15, basal region, transition between the basal armature and metabasal region on the extemal surface; 16, lower tiectabasal region, antibothridial surface showing single intercalary row of hooks. Scale lines: 12-16, 0.1mm. NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTTA 45 (Pseudogrillotiidae) were added by Dullfus (1969a). Schniidt (1986) included jhe genus Grillotia and Pseudogniloria in his key but omitted Progniitoria and the family Pseudogrillotidae. We concur with Schrult (F986) m considenng a craspedote scalex and postlarval Stage insufficient to warrant separation of Preudogrillotia into another family. We have found no Paper in which Dolifus ever formally defined the family Grillotiidae but he did comrast the Grillotidac as well as the subfamily Grilloliinae (Dollfus 1969a) in the justification for creating Psevdegritiotia and the Pseudogrillovidae. Despite the incrensing number of species described and the varlabiliy en the combination of characters now apparent, Dollfits' treatment of the group has remained basically unchanged since 1969. The diverse combinations of characters and taxonomic importance previously attributed to them must be questioned in view of species recently described. The various combinations include patelliform versus non- patelliform bothridia, craspedote ur acraspedete scolex, presence or sbsence of prebulbar organs, long or short bulbs, vanous origins of the retractor muscle, presence or absence of a postlarval stage between the blastocyst stage and adult, presence or absence of a basal armature, presence Or absence of a demarcation between the intercalary hooks and the longitudinal band of hooks on the external surface, and ¢ven the interpretation of the presence of a Jongitudinal “band of hooks for some species, For the reproductive system the details of the terminal genitalia are not well known but a hermaphroditic sac, combined with an accessory, external and internal seminal vesicles have been shown 10 occur in several species, the ovary may be terminal or subterminal in position, and postovarian testes are present in some species and facking in others. As proposed by Dollfus (1942) the sub-family Grillotiinae is justified as separate from thie Lacistorhynchinae in the Jack of a chainette, presence of a longitudinal ‘band™ of hooks in the metubssal armature, and présence of intercalary rows of hooks in the armature. These same characters ate implied as supporting elevation Ww family mink (Dollfus 19694). Complete revision of the family Grillotiidac should be made based upon examination af types of all species. The diversity of characters and combinations is not unique to Grillotia, but many of these same combinations can be found in the Gilquiniidse (see Beveridge 1990). We propose that they be accommodated by including them in a formai diagnosis of the family and recognised separately from the Lacistorhynchidae as folinws- Grillotiidae Dollfus, 1969 emcended Diagrasis; Scolex acraspedate or craspedote, Two wite bothndia, cordiform or patelliform, with or withomt thickened rims, posterior and lateral margins free. Principal hook rows alternate, Intercalary rows vf hooks present, irregular band of small hooks between termination of principal rows in metabagal armature. Basal armature preseal or absoiit, Hooks heleramorphous, hollow, Pars vaginalis long, tentacle aheaths sinuous. Bulbs long or shart, prebulbar organs present of sbsent. Retractor muscle originates in posterior half or bulb. Segments acraspedote, apolytic, Mature segments elangated, Genital pores marginal, irregularly alternate, Accessory seminal vesicle, internal seminal vesicle and bermaphomditic duct present in type species. Hermaphroditic duct, extemal amd intemal seminal vesicles reported for some species. Testes medullary, some usually postoyarian, Ovary usually separated from posterior segment margm. by a-space. Vitelline follicles form a continuous sleeve around the anternal organs. Uterus saccate with Lateral diverticular. Procercoid in copepods, plerocerci and postlarvae in teleosts, adults in elasmobranchs. Type genus: Grilloria Guiart, 1927 Diagnosis; Grillotiidac, acraspedote scolex; hothridia indented on posterior border, longitudinal “band” of hooks continuous, intercalary rows of houks may merge with “band”, special basal armature absent, prebulbur organs present, hennaphroditic sac present, imternal and external (or accessory) seminal vesicles present, ovary well forward, postovarian testes present. Other genera; Progriliotia Doltfus, 1969 Diagnosis. Grillouiidae, acraspedote scolex, bothridial margins not indented posteriorly, prebulbar organs lacking, longitudinal “band” of hooks interrupted Opposite each principal row, special basal armaature may be present, bulbs very long, prebulbar organs lacking. testes in longitudinal tows, accessory seminal vesicle absent, ovary at posterior extremity of segment, posiovarian testes absent. Type species: Pr. pastinacae Dollfus, 1946, Pseudogrillona Dollfus, 1969 Diagnosis: Grillotiidae, craspedote scolex, bothridia indented posteriorly, prebulbar organs preseril, longitudinal “band” of hooks conlinwous but may be restricted io basal region, special basal armature absent, hermaphraditic duct present, testes nicemose, ovary well forward of posterior margin, postyaginal testes present, posularval stage im life cycle. Type species: P pleistaccentha Dollfus, 1969. Acknowledgments Thanks are due to Mr B. G. Roberson aid Dy DB. M, Spratt for collecting specimens, and to te Australian Biological Resources Study for financial Suppor. 46 R. A. CAMPBELL & I, BEVERIDGE References Beumer, J. P., ASHBURNER, L. D,, Bursury, M, E., JETTE, E,, & Latuam, D, J, (1982) A checklist of the parasites of fishes from Australia and its adjacent Antarctic territories. Commonw. Inst. Parasitol. Techn. Comm. No. 48 (Commonwealth Agricultural Bureaux, Farnham Royal, England.) BEVERIDGE, I. (1990) Revision of the family Gilquiniidae Dollfus (Cestoda: Trypanorhyncha) from ¢lasmobranch fishes. Aust. J. Zool. 37, 481-520. & CampsBeLt, R. A. (1988) Cetorhinicala n.g., Shirleyrhynchus n.g. and Stragulorhynchus n.g., three new genera of trypanorhynch cestodes from elasmobranchs in Australian waters. Syst. Parasitol. 12, 47-60. —___ & __ (1989) Chimaerarhynchus ng. and Patellobothrium n.g., two new genera of trypanorhynch cestodes with unique poeciloacanthous armatures, and a reorganisation of the poeciloacanthous trypanorhynch families, Jbid, 14, 209-225, Cama, J. & Gavarrino (1989) Grillotia similis (Linton, 1908) comb, n. (Cestoda; Trypanorhyncha) from nurse aback in the Florida keys. J. Helminthol. Soc. Wash, 57, 15-20. CAMPBELL, R, A. (1977) New tetraphyllidean and trypanorhynchan cestodes from deep-sea skates in the western north Adantic. Proc. Helminthal. Soc.. Wash. 44, 191-7, CaRVAIAL, J,, CAMPBELL, R. A. & CORNFORD, E. (1976). Some trypanorhynch cestodes from Hawaiian fishes, with descriptions of four new species. J. Parasitol, 62, 70-7. Doirus, R. P. (1942), Etudes critiques sur les tetrarhynques du Museum de Paris. Arch. Mus, Natl. Hist, Nat. (Paris) 19, 1-466, (1946) Notes diverses sur les Tétrarhynques. Mém- Mus. Natl. Hist, Nat., 22, 179-220, (1969a) Quel ques espéces de cestodes tetrarhynques de la cote Atlantique des Etats Unis, dont l'une n’etait pas connue a l'état adulte. J, Fish, Res. Bd Can. 26, 1037-61. (1969b) De quelques cestodes tétrarhynques (heteracanthes et pécilacanthes) recoltes chez des poissons de la Mediterranee. Vie et Milieu, ser. A 20, 491-542. ESCALANTE, H. & CarvalaL, J. (1984) Larval trypanorhynch cestodes from Peruvian teleost fishes, with descriptions of two new species. Stud. Neotrop. Fauna Environm. 19, 185-194. SAKANARI, J, (1989) Grillotia heroniensis, sp. nov., and G. overstreeti, sp. nov., (Cestoda; Trypanorhyncha) from Great Barrier Reef Fishes. Aust. J. Zool. 37, 81-7. Scumipt, G. D. (1986) ‘CRC handbook of Tapeworm Identification’ (CRC Press Inc,, Florida.) SHAHAROM, F, M. & Lester, R. J. G. (1982) Description and observations on Grillotia branchi n- sp., a larval trypanorhynch from the branchial arches of the Spanish mackerel, Scomberomorus commersani. Syst. Parasitol. 4, 1-6. YAMAGUTI, 8, (1959) Systema Helminthum. IL. The Cestodes of Vertebrates, (Interscience Publishers, New York). ABUNDANCE AND LIFE HISTORY OF NATIVE AND INTRODUCED EARTHWORMS (ANNELIDA: MEGASCOLECIDAE AND LUMBRICIDAE) IN PASTURE SOILS IN THE MOUNT LOFTY RANGES, SOUTH AUSTRALIA BY G. H. BAKER*, V. J. BARRETT*, R. GREY-GARDNER** & J. C. BUCKERFIELD** Summary Seasonal abundance of the native earthworm, Gemascolex walkeri (Megascolecidae), and introduced earthworms, Aporrectodea rosea and Octolasion cyaneum (Lumbricidae), was monitored in three pastures in the Mt Lofty Ranges, South Australia. Highest numbers of earthworms were recorded in late winter and early spring. Densities of up to 108, 32 and 96 m” were recorded for G. walkeri, A. rosea and O. cyaneum respectively. All three species of earthworms occurred predominantly in the top 10 cm of soil for four to five months per year (autumn to spring), when soils were wettest. During other seasons, they were found lower in the soil profile. Distributions within pastures were patchy and could, in some cases, be explained by soil type (A. rosea) and soil moisture (G. walkeri). Although introduced species of earthworms generally predominate in pastures in southern Australia, native species such as G. walkeri can occasionally constitute a substantial proportion of the total earthworm population. KEY WORDS: Earthworms, Megascolecidae, Lumbricidae, pastures, South Australia, depth profile, life history. Transactions of the Royal Society of 5. Anst., (1993), Ut, 47-33, ABUNDANCE AND LIFE HISTORY OF NATIVE AND INTRODUCED EARTHWORMS (ANNELIDA; MEGASCOLECIDAE AND LUMBRICIDAE) IN PASTURE SOILS IN THE MOUNT LOFTY RANGES, SOUTH AUSTRALIA by G. H. BAKER* V. J. BARRETT* R. GREY-GARDNER* & J, C. BUCKERFIELD™ Summary Haxer, G. H., BARRETT, V. J, Grey-Garpner, R, & Buckerriecp, J, C. (1993) Abundance and lile history of native and introduced earthworms (Annelida ; Megascolecidae und Lumbricidae) in pasture soils in the Mount Lofty Ranges, South Australia Trans, R. Soc. S. Aust. 147(1), 47-53, 4 June, 1993. Seasonal abundance of the native earthworm, Gemascolex walke'rt (Megascolecidae), and introduced earthworms, Aporrectadea rosea and Octolasion cyaneum (Lumbricidae), was monitored in three pastures in the Mt Lofty Ranges, South Australia. Highest humbers of earthworms were fecorded in late winter and eurly spring. Densities of up to 108, 32. and 96 mr were recorded for G. walkeri, A. rosea and O: cyaneum respectively. All three species of earthworms occurred predominantly in the top 10cm of soil for four to five months per year (autumn to spring), when soils were wettest. During other seasons, they were found lower in the soil profile. Distributions. within pastures were patchy and could, in some cases, be explained by soil type (A. rosea) and soil moisture (G_ Wwalkeri). Although introduced species of carthwortns generally predominate in pastures in southern Australis, native species such us G, walker! canoccasionally constitute a substantial propartion of the total earthworm population, Key Worbs: Earthworms, Megascolecidae, Lumbricidae, pastures, South Australia, depth profile, lite history. Introduction Australia has a rich native earthworm fayna, dominated by the Megascolecidae (Jamieson 1981). Howeves, information on the life histories, distributions and abundances of these native species and the factors influencing them is scarce (Wood 1974; Jamieson 1931; Abbott e¢ al, 1985; Baker et al, 1991, 1992a). The distributions of introduced species in Australia (e.2. Lumbricidse), are better understood (Abbott 1922, 1985; Abbow & Parker 1980; Tisdull 1985; Kingston & Temple-Smith 1989; Baker ey af, 1991, 1992a), and there have been some studies of their life histories and population dynamics (Barley 1959a;, Tisdall 1985; Rovira er al, 1987; Kingston 1989; McCredie er al. 1992; Baker e¢ af 1992b). The influence of earthworms, cither mative or introduced, on soil structure ated fertility in Australia has received little study (Barley 1959b,c, Barley & Jennings 1959; Abbott & Parker 1981), Increased knowledge of the ecology und Functional importance of the resident communities of earlhworms in Australian soils will assigt in developing improved management strategies that may lead to increased plant production and help reduce soil degradation (Baker 19893,b). Baker et ai. (1992b} demonstrated the seasonal abundances and depth profiles of two introduced * Divisions of Entumology* and Sails“, CSIRO, P.M. Bag 2, Glen Osmond 5064, South Australia. species, Aporrectodea trapezoides ami A. caliginosa (Lumbricidae), in five permanent pastures in South Australia, A, trapezpides and A. caliginose were active and abundant in the rvot zone for three to seven months, from autumn to spring, when soi] moisture was highest. Mature adults were present from mid- winter to mid-spring, Patchy distributions of both species were found within some of the pastures and were partly explained by variation in soil type. This paper reports the seasonal abundances, life histories, lateral and vertical distributions and species associations of native and intrxiuced earthworm species that were found together with the mon abundant 4, trapezaides and A. caliginosa at three of the pasture sites sampled by Baker et af. (1992b). Materials and Methods Three permailent pastures in the Mt Lofty Ranges, South Australia were sampled for earthworms each month from March 1990 until November 1990 (Tungkillo) or January 1991 (Birdwood and Parawa). The climate of the region is Mediterranean, with cool moist winters and hot dry summers. Mean annual rainfall yaricd between sites (Tungkillo, 575 mm; Birdwood, 750 mm and Parawa, 900 mm) as did soil type (Tungkillo, unitorm sand [Ucl,4]; Birdwood, yellow and black duplex soils [Dy3.4 and Ddl.1) and Parawa, yellow duplex soil [Dy2.2}) (Northcote 1979), The yellow and black duplex soils are Palexeralfs and the uniform sand is a Quartzipsamment in the LIS Soil Taxonomy (Soil Survey Stalf 1987). Baker ef ail, G. H. BAKER, V. }. BARRETT, R. GREY-GARDNER & J. C. HUCKERFIELD (1992b) provide further information on chemical and physical properties of the soits af each site. At each site, soil samples were taken within « 50 x 30 m plot that was divided into five 10 m wide strips. Each month, three random samples were taken within each strip at Tungkillo and Birdwood and two samples were taken within each strip at Parawa. Each sample was 0.1m? x 30 cm deep. The samples were separated into three depths, 0-10 cm (within which the majority of plant roots occurred), 10-20. cm and 20-30 cm. The soil was hand-sorted for carthworms and cocoons which were preserved in 70% ethanol. Approximately 400 g of soil were collected From the 0-10 cm layer in one sample from each strip for gravimetric moisture determination, These soil samples were collected every month except March 1990, Curves relating matric suctions of soil water to gravimetric walet content wete calculated for the soils at each of the three sites using the filter paper method (Greacen et al, 1989), This enabled comparisons of soil moisture avatlabilily between sites. Species were identified within two weeks of collection, using the keys in Jamieson (1974) and Sims & Gerard (1985). Worms were separated into juveniles (no genital markings), subadults (with genital markings but no mature clitelhim) and adults (with mature. clitellum). Two soil types (yellow and black duplex soils) occurred within the plot at Birdwood, The distributions of these soils were mapped by taking 25 regularly spaced auger samples (each 3 cm wide x 50 cm deep) within the plot and supplementing these with additional samples where the soil boundaries were unclear. There were no obvious spatial variations in soil type wilhin the other two plots, The regular sampling at Tungkillo indicated that soil moisture varied markedly within the plot, being highest near the centre and éastern edge. This pattern was possibly related to the drainage characteristics of the site, the moistest soil belng found in the lowest lying areas Within the plot. To check this perceived pattern in soi] moisture, 22 auger samples (10 cr in diameter) ‘were taken at random co-ordinates throughout the plot during March 199], soil being collected from 0-10, 10-20, 20-30, 30-40 and 40-50 cm below the surface. Gravimetric soi] moistures were determined for each depth. Results Abundance and Life History Duta for the seasonal abundance and life history of three earthworm species that wers commonly collected a Tungkillo, Birdwood and Parawa are given in Figs 1 amd 2. Similar data for rarer populations of these and other species found at the three sites are given in Table 1. ari = f ia Numbat per square metre Number per square metre Bo ws 3 Numberper square metre i s oz = —— i ome Mer Apr May Jun Jul Aug) Sep) Gel ow Der day Manth Fig. 1.. Numbers (m~2) of juvenile (7 ——_1), sub-adult ( Cruces ) and adult ( —— ) Gemascolex walken’ (Tungkillo) (a), Aporrectodea rosea (Birdwood) (b) and Octolasian ¢yaneur (Patawa) (c) collected during 1990-91, Standard errors for the total earthworm numbers (all life cycle stages combined) are given for each month, * indicates the presence of cocoons. Gemascolex waikeri Jamieson (Megascolecidae) was common at Tungkillo, not found at Parawa and occurred in only negligible numbers at Birdwood (Fig. 1, Table 1). Aporrectadea rosea (Savigny) (Lumbricidae) was most common.at Birdwood and was also found at Tungkillo, Octolasion cyanewn (Savigny) (Lumbricidae) was found at Parawa but not at the other two sites. The abundances of G. walker, A. rosea and cyareum varied between months (Kruskal-Wallis H = 54.9, p < 0,001 for G, walkeri at Tungkillo; H = 42.3, p < {001 for A. rosea at Birdwood, H = 45,1, Pp < 0.001 for O cyaneum at Parawa), with highest numbers found from late winter to early spring (August to September or Octoher) (Fig. 1). Other species were penetally recorded in low numbers at the three sites (Table 1). Gemtascolex lateralis (Spencer) ABUNDANCE OF EARTHWORMS IN PASTURES 49 (Megascolecidae) was abundant at parawa in May and, June 1990 (Table 1), but most of the individuals of this species collected then were found in only one sample in each month (53% of the total collection in May and 69% in June), There was no significant variation between months in the abundance of G. lateralis at Parawa (H = 8&6, p > 0,05). Numbar per square metre a Motsture 20 & Ss Ss fe = Number per square matte ho a hy MbIStUTE S an ="\3 %y Moisture = o Number per square metre oe S we a 0 a _Teseait Mar Apr May Jun Jul Avg Sap Geb Noy Dec Jan Month Fig. 2, Numbers (m~*) of Gemascolex walkeri (Tungkillo) (3), Aperrectodea rosea (Birdwood) (b) and Octolasion eyaneurm (Parawa) (c) collected 0-10 cm ({[[__—_]), 10-20 cm (ea » and 20-30 cm ( MEE ) belo the soil surface during 1990-91, Standard errors for the total earthworm numbers (ail depths combined) are given for each month. Mean gravimetric soil moistures for 0-10 cin deep are also. indicated for each site (line). Adults of G. waikeri and A. rosea were mostly collected in early spring (Fig. 1), Adults of @. cyaneum were present on most sampling occasions, Earthworm cocoons were found at all three sites, during each month fram July to October (up to 39 cocoons per soil sample). Virtually all the cocoons (> 99%) were found in the top 10 cm of soil. The identity of these cocoons is unknown, Most of the cocoons probably belonged to Aporrectodea trapezoides, the most abundant species at these sites {Baker er al. 1992b). However, the breeding seasons of all species can be regarded as occurring within these months. Depth Profile G. walkeri, A, rosea and O. cyateum occurred mainly in the root zone (0-10 cndepth) for four to five months during autumn to spring, when soils were wettest (Fig. 2) (Pearson ty = 0.98, Ty = 0.86 and Ty = 0.94 for soil moistures and numbers of G. walkeri (Tungkillo), A. rosea (Birdwood) and QO. cyaneum (Parawa) respectively at 0-10 cm depth; p <0.01 in all cases). Earthworms becime more ‘common in the root zone compared with lower depths during May or June and less. so in October or Noverober. a OTunghllo O Birdwood 4 Parawa LN Suction (kPa) n Apr Way Jan dol fy Sop Nov Ber Jar Month Fig. 3. Matric suctions of soil water (kPa) at 0-10 cm depth at Tungkillo (©) ), Birdwood ( 0 ) and Parawa ( A ). The surface soil was moister in autumn (May) and drier in spring (October).at Parawa compared with the other two sites (Fig. 3). Proportionally more worms were present in the top 10 cm of soil at Parawa in May and less in October than at Tungkillo and Birdwood (Fig. 2, Table {) (comparing the total earthworm numbers at 0-10 cm and 10-30 cm depth at each site for all species included in Table 1 and Fig, 2,37 = 70.65, p < 0,001 and x? = 96.91, p < ODO for May and October respectively), Distribution Within Plots The distributions of some species at each of the three sites were patchy. At Birdwood, A, rosea was most common in the south-eastern comer of the plot whilst Gemascolex lateralis and G, stirlingi occurred predominantly in the north-eastern comer (Fig. 4), The distribution of A, rosea approximated that of the black duplex soils at Birdwood (Fig. 4) (x? = 16,25, p < 0901 where the samples with and without A. rosea 50) TABLE 1. G, H, BAKER, V. I. BARRETT, R. GREY-GARDNER & J. C. BUCKERFIELD Numbers of earthworms collected in low numbers at three pasture sites in the Mt Lofty Ranges, South Australia. Numbers in parentheses aré for earthworms in the top 10 cm of soil, * indicates adults were present and — that no samples were taken. Species Monti M A M I I A 8 oO N D J Total Aporrectodea rosea Tungkillo 3 2 26 2 6 4* I 6* 10* - — 60 (7) (6) (4) (1) (6) G) Microscolex dubius. Tungkillo 4* 14+ 8* - = 26 (4 (14) (8) Birdwood 2* 2 (2) {* 1 Parawa (1) Gemascolex stirlingt Tungkillo 4 ‘1 = ~ 5 (2) Birdwood 1 1 ad ] 6 1 M 1 i] 2 1 20 (5) (1) - G. lateralis Tungkillo 1 = a ] (ly Birdwood 5 4+ i 4 10 3 J 2 3 33 (1) (4) (10) (3) (I) Parawa 3 6 §3* 78% 17* 14+ QF 3* 21" 6* 19* 229 (53) (78) (IT) (4) (9) {1) G. walkert Birdwood 3* 3 6 _ (3) Native W Tungkillo * | 1 J 2 — — 6 ; 1) r)) (1) (2) Native J Birdwood ti i within the areas with yellow and black duplex soils were compared), but the distributions of the two Gemascolex spp. could not be explained by variation in soil type. — At Tungkillo, G. walkeri was most abundant near the centre and eastern end of the plot (Fig, 5a) (x? = 31.29, p < 0.001 where the frequencies of samples with 0, 1-5, 6-10 and > 10 earthworms inside and outside the area in Fig. 5a were compared), The soil within the area where G. walkeri was most common was moister in late summer than the soil outside it (Table 2). During the wettest months of the year, the same irend was also apparent. Mean soil moistures for the samples taken at 0-10 cm depth during the routine monitoring at Tungkillo from July to October 1990 - were 17.2% (where G. walkeri was most common) and 13.1% (elsewhere) (t = 2.23, p < 0.05). G. lateralis was mostly found in the south-eastern corner of the plot at Parawa (Fig. 5b) (x7 = 62.05, Pp < 0.001 where the frequencies of samples with 0, 15 and > 5 earthworms inside and outside the shaded area in Fig. 5b were compared). There was no obvious variation in habitat within the plor that might explain this patchiness in G lateralis. However, several Eucalyptus trees bordered the plot adjacent to the area where G. lateralis was found. Perhaps these remnants of natural vegetation ate responsible for local survival of this native earthworm. Discussion The life histories, patterns in seasonal abundance and depth profiles of G. walkeri, A, rosea and O. cyaneum in pastures in the Mt Lofty Ranges, South Australia are broadly similar to those reported previously for A, trapezoides and A. caliginosa (Baker et al, 1992). Earthworm activity in the pasture root zone. (0-10 cm depth) is mostly restricted to winter and early spring when soils are wettest. Adult earthworms are most commonly found in late winter-early spring, but for some species (e.g. O. cyaneuin and G. lateralis), adults can be found during most months of the year. The proportions of A_ rosea, O: cyaneum and Gemascolex spp. populations that survive summer are not known, but many of these earthworms have been ABUNDANCE OF EARTHWORMS IN PASTURES S| G.. lateralis A, rosea BIRDWOOD ‘ ® N ao}- a absent 20 a 71-5 + 6-10 a >10 0 10 ra] 30 AG 50 Fig, 4, Distributions of Aporrectadea rosea. Gemascolex lateralis, G. stirlingi and soil typés at Birdwood. Earthworm numbers are piven as0(0),1-5( Ml), 610 (+}and > 0( a ) per sample, Shaded areas indicate where all or the majority of eaathworms Were found, Soil rypes were classified as yellow duplex (Dy), black duplex (Dd) and intergrade of Dy and Dd (1). found in summer > 30 cm below the surface of the soi] (G, Baker, unpublished data). The increase in population numbers. that occurs in the surface layers of the soi] in autumn and easly winter must be explained by invasion of individuals from lower depths rather than by breeding, since no cocoons were found al that time. Whilst pastures in the Mt Lofty Ranges are dominated by introduced lumbricids, espevially Aporrectodea spp. (Baker ef al, 19924), some native megascolecids, such as G, walkeri, can be locally abundant and constitute a significant proportion of the tolai earthworm population. For example, G. wulkeri constituted 40.1% of the total population collected at Tungkillo in 1990 (Baker ef al, 1992b), [Most (54.8%) of the earthworms found at this site were A, trapezoides. | The contribution that native species make to soil structure and fertility has yet to be resolved, Experiments are currently being conducted to evaluate the role of G. walkers at Tungkillo in tairying surface- applied lime and nffsetting sail acidity: A, rosea is one of the most widespread and abundant earthworm species in South Australia and western Victoria {Baker e al. 1991, 1992a). Multiple regressions of the abundance of A. rosea against a variety of environmental variables have shown that soil particle size is statistically the most important regressor for sites in the Mt Lofty Ranges, as is the case for the other most common introduced species, A. trapezoides and A, caliginosa (Baker et al, 1992a)., In general, the abundance of these earthworms increases with % clay content, Variations in soil particle size helped explain differences in the abundance of A. trapezoides and A. caliginesa at several farms including Tungkillo, Birdwood and Parawa (Baker er al, 1992b). They may also explain the differences in abundance reported hete for 4. rosea at Tungkillo and Birdwood. A. rosea was more abundant in the black duplex soil (19% clay) than in the yellow duplex soil (7% clay) at Birdwood and the uniform sand at Tungkillo (also 7% clay), The absence of A, rosea from Parawa (where clay content was 14%) is, hawever, not explained by clay content. 32 ti, H, BAKER. V, 1, BARRETT, R. GREY-GARDNER & J. C. BUCKERFIELD TUNGKILLO G. walkerl PARAWA G. lateralis eered im 4s i a 10 20 a ay ay Fig. 5. Distributions of Gemascolex watkeri at Tangkillo and G. tateralis at Parawa. Eorthworm number. are given as 0 (O), 15 (M), 6-10 (+), 1-20 ( @ ) and > 204 A) per sample. Shaded areas indicate where the majority of eanhworms were found. ©} cyaneum was restricted to Parawa, the wettest site with an annual rainfall of 900 rom, compared with 575 and 750 mm at Tungkillo and Birdwood respectively. Previous surveys (Baker e7 al. 1991, 19923) have found O. evaneum at only a few sites in the highest rainfall regions of South Australia and western Victoria. There TABLE 2. Gravimetric moistures (%) for soils collected at Tangkilla in March 1991 Soils were collected at varying depths inside and outside the area in which Gemascolex walkeri was most abundant (sec Fig. 5}. Results of t tests for significant differences between data are included. Depth Soil Moastute- (em) Inside Outside i Prob. 0-10 1,48 0.93 §.20 < 0,001 10-20 2.60 iv? 4.65 < 0.001 20-30 3,83 1799 5.00 < 0001 30-40 474 2.55 3.78 < O01 40-50 642 2.82 2.80 < 0.05 ate a few records of O. cyanewm in Western Australia, all from the high rainfall region of the south-west (Abbott 1981), Sims & Gerard (1985) comment that O, cyaneum prefers moist habitats in Britain. The distributions of the native species within-the three sites were patchy, In some cases, possible reasons for the patchiness can be offered. For example, G. walker muy be restricted by soil moisture at Tungkillo. Baker ef al. (1992b) teported minor levels of parasitism af A. trapezvides and another introduced earthworm, Microscolex dubius, at Tungkillo during spring by an undescribed dipteran. G. walkeri was more abundant than these other two species in the top 10 cm of soi] at the same time, but was not parasilised, Acknowledgments This research was funded in part by a grant from the Australian Wool Research & Development Corporation, We especially thank D, Hobbs, A. Philp and V, Rathjen for the use of their land, Penny Carter for much assistance with the earthworm collections, Richard Merry for help with the pedology and soil chemistry and Peter Williams for help with the soil moistures, References Asner, 1. (198!) Twa species of lumbricid garthworm newly recorded from Western Austratin, Rec, West Aust. Mus. 9, 273-277. (1982) The distribution of earthworm in the Perih metropolitan area, Jbid. 10, 11-34, ____ (1985) Distribution of introduced earthworms in the northern jarrah forest of Western Australia. Aust, J, Soil Res. 23, 263-270, & Parker, C. A. (1980) The occurtence of carthworms. in the wheat-belt of Western Australia in relation to land use and. rainfall. Jbid. 18, 343-352. & __ (1981) Interactions between earthworms and their soil environment. Sail Biol. Biochem. 13, 191-197, . Ross, J. S. & Parwsn,C, A. (1985) Ecology of the lame indigenous earthworm Megascolex imparicystis in relation to agriculture near Lancelin, Western Australia- JR. Sac. W. Aust, 68, 3-15, Baker, G. H, (1989a) Role for earthworms in ameliorating soil acidity pp. 64-65, In K. J. Hutchinson (Ed.) “Biological Processes and the Fertility of Tem and Mediterranean Pastures.” (Aust. Wooi Corp. Tech. Publ.). ______ (1989) Potential for introduced earthworms. in pastures in southern Australia. Ibid. pp, 68-70, , Barrett, V. J., BUCKERFIELD, J. C., GREY Garpner, R, & Carrer, P. J. (1991) Distribution and abundance of earthworms in agncultural soils in South Australia and western Victoria pp. 30-51. /n Ri G, Faweet (Ed.) “Workshop on Tillage Systems, Rotations, Nutrition and Associaked Root Diseases.” (S, Aust. Dept. Agnc. Tech. Publ). ABUNDANCE OF EARTHWORMS IN PASTURES 53 , GREY-GARDNER, R, & BUCKERFIELD, J. C. (1992b) The life history and abundance of the introduced earthworms Aporrectodeq trapezvides and A. caliginosa (Annelida: Lumbricidae) in pasture soils in the Mount Lofty Ranges, South Australia, Aust. J. Ecol. 17, 177-188. ; BUCKERFIELD, J, C., GREY-GARDNER, R., Merry, R. & Douse, B. (1992a) The abundance and diversity of earthworms in pasture soils in the Fleurieu Peninsula, South Australia. Soil Biol. Biochem. (In press). Bar ey, K. P. (1959a) The influence of earthworms on soil fertility, I. Earthworm populations found in agricultural Jand near Adelaide. Aust. J, Agric: Res. 10, 171-178. (1959b) The influence of earthworms on soil fertility. Ti. Consumption of soil and organic matter by the earthworm Allolobophora caliginosa. Ibid. 10, 179-185, (1959c) Earthworms and soil fertility, IV. The influence of earthworms on the physical properties of a red- brown earth. Ibid. 10, 371-376. & Jennings, A. C. (1959) Earthworms and soil fertility, O10, The influence of earthworms on the availability of nitrogen, [bid. 10, 364-370. GREACEN, E. L., WALKER, G. R. & Cook, P. G. (1989) Procedure for the filter paper method of measuring soil ae suction. CSIRO Division of Soils Div. Report No. 08, 7 pp. JAMIESON, B. G. M. (1974) Earthworms (Oligochaeta: Megascolecidae) from South Australia. Trans. R, Soc, S. Aust. 98, 79-112. —— (1981) Historical biogeography of Australian Oligochaeta. pp. 887-921. In A. Keast (Bd.) “Ecological Biogeography of Australia.” Vol. 2 (W. Junk, The Hague) ‘pp. 887-921. Kinoston, T. J. (1989) Aporrectodea caliginosa and Ligmbricus rubellus populations under irrigated and dryland pastures in northern Tasmania. pp. 199-205. Jn P. P. Stahle (Ed.) Proc. V Australasian Conf Grassland Invertebrate Ecology (D & D Printing, Melbourne). & TEMPLE-SMITH, M. G. (1989) Earthworm populations under Tasmanian pastureland. bid. pp. 192-198. McCrepiz, T. A., Parker, C. A., & Asport, I. (1992) Population dynamics of the earthworm Aporrectodea trapezoides (Annelida: Lumbricidae) in a Western Australian pasture soil, Biol. Fertil. Soils 12, 285-289. Norricore, K. H. (1979) “A Factual Key for the Recognition of Australian Soils.” (Rellim Press, Adelaide), Rovira, A. D., SmerTem, K. R. J. & Lee, K. E. (1987) Effect of rotation and conservation tillage on earthworms in a red- brown earth under wheat. Aust. J. Agric. Res. 38, 829-834. Sims, R. W. & Gerarp, B. M. (1985) “Earthworms.” Syn, British Fauna No, 31. (Linnaean Soc. London, London). Sou. SURVEY Starr (1987) “Keys to Soil Taxonomy” Third Printing. SMSS Tech Mon. No. 6. (Ithaca, New York), TisDALL, J. M. (1985) Earthworm activity in irrigated red- brown earths used for annual crops in Victoria, Aust. J. Soil Res. 23, 291-299. Woop, T. G, (1974) The distribution of earthworms (Megascolecidae) in relation to soils, vegetation and altitude an he slopes of Mt Kosciusko, Australia. J. Anim. Ecol. , 87-106. TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 117, PART 2 THE EARLY CAMBRIAN VOLCANICS FROM RED CREEK, EASTERN MT LOFTY RANGES, SOUTH AUSTRALIA BY C. G. GATEHOUSE*, J. B. JABOT, B. J. CLOUGH*® & A, J. MCCULLOCH*} Summary In the Red Creek area of the eastern Mt Lofty Ranges, Early Cambrian lavas, tuffs and volcaniclastic sediments are interbedded with the top 270 m of the Heatherdale Shale, the top member of the Normanville Group. Tuffs and volcaniclastic siltstones extend up into the basal 60 m of the conformably overlying Carrickalinga Head Formation, the basal unit of the Kanmantoo Group. Several lava flows of trachybasalts, one of which exhibits pillows, occur about 100 m below the top of the Heatherdale Shale. The volcanics at Red Creek appear to be at a slightly higher stratigraphic level than the Truro Volcanics, the type section of which is 24 km to the NNW of Red Creek. It is proposed that the lavas at Red Creek represent a flow of hawaiite composition from a central volcanic complex, closely analogous to that of the Truro Volcanics but not necessarily from the same centre or erupted at exactly the same time. Geochemical data indicate that both the volcanics at Red Creek and the Truro Volcanics belong to the same alkaline Early Cambrian "within plate" volcanic province, which may be linked genetically with other Cambrian mafic alkaline provinces such as the Yumali/Coonalpyn area. It is suggested that the closest tectonic analogy for these provinces is that of a rifted continental margin as proposed for the Tertiary intraplate yolcanic province of eastern Australia. KEY WORDS: Early Cambrian volcanics, hawaiite, Mt Lofty Ranges, Normanville Group, Kanmantoo Group Jransuctions of the Koya! Society of S. Awst, 0993), 117(2), 57-66. THE EARLY CAMBRIAN VOLCANICS FROM RED CREEK, EASTERN MT LOFTY RANGES, SOUTH AUSTRALIA by C. G, GaTenouse*, J, B. Jacof, B. J. CLouGcH#* & A. J. MCCULLOCH*? Summary GateHouse, CG, Jaco, J. B., Clouex, B. I. & MeCuLvocn, A. J. (1993) The Early Cambrian volcanics from Red Creek, eastern Mi Lofty Ranges, South Australia. Trans. R. Soc. S. Aust. 117(2), 57-66 4 June, 1993, In the Red Creek area of the eastern Mr Lofty Ranges, Barly Cambrian lavas, tule and volcaniclastic sediments ame interbedded with the top 270 m of the Heatherdale Shale, the top member of the Normanville Group. Tufts and volcaniclastic siltscanes extend up ivito the basal 60 m ot the conformably overlying Carrickalinga Head Formation, the basal unit of the Kenmunioo Group. Several lava tlows of trachybasalts, ane of which exhitits pillows, occur about 100 m below the top of the Heatherdale Shale: The volcanics a) Red Creek appear to be al Slightly higher stratigraphic level than che Truro Volcanics, the type section of which is 24 kim in the NNW af Red Creek, It is proposed that the lavas at Red Creek represent u flow of hawaiite composition from a central volcanic complex, closely analogous to that of the Truro Volcanics but nol necessarily from the same centre or erupted at exactly the samne time, Geochemical data indicate that both the volcanics at Red Creek and the Truro Volcanics belong to the same alkaline Early Cambrian *within plate" volcanic province, which may be linked genetically with other Cambrian mafic alkaline provinces such a8 the Yurnali/Coonalpyn area, [t is suggested that the closest tectonic analogy for these provinces is that of a rifted continental margin as proposed for the ‘Vertiary intraplate volcanic province of eastern Australia. Key Woros: Early Cambrian volcanics, hawailte, Mt Lofty Ranges, Normanville Group, Kanmantoc Group Introduction This paper is a progress report on work an the Early Cambrian volcanics generally known as the Truro Volcanics of the eastern Mt Lofty Ranges. The Truro Volcanics are important in chat, although they are limited both in stratigraphic and geogmphic extem, they represent the most extensive outcrops of yoteanics in the Cambrian sequences of the Stansbury Basin/Kanmantoo Trough area. As described helow, voleanics in the Red Creek area extend From wuhin the Heatherdale Shale, the top member of the Normmanville Group, up into the Carrickalinga Hew Formation, the basal member of the Kanmantoo Group, Hence « study of the Truro Volcanics is important in the determination of the stratotectonic setting of the Kanmantoo Group transition which has been interpreted in various ways as set out below, You der Borch (i980) considered the Kanmantoa Group sédiments to be the initial phase of fully developed continental margin sedimentation which typified the easiern flank of cratonic Australia during most of the Palneazoic, Tamer & Foden (1990) suggested that this phase was marked by renewed mafic igneous activity in the form of the Truro Volcanics- The basal part of the Carrickalinga Head Formation ar Carrickalings Head lles north of Normanville where von der Borch (980) proposed that the Houghton Antictinal Zone acted a5 a separating feature between *S.A. Dept of Mines and Energy, P.O. Box 151. Eastwood, S.Aust, 5063 + Dept of Applied Geology, Gartrell School, University of South Australia, The Levels, S,Aust, S095, shallow-water sediments to the west and continental slope and rise environments to the cast, Scheibner (1986, Fig. 4) Implied that the Kanmantoa Group sediments were at least in part deposited on oceanic crust and that che “Kanmantoo Trough” was part of a marginal sea, extending eastwards to the Stavely volcanic belt of westem Victoria. Parker (1986) suggested that the Kammantoo Group sediments were probably deposited in an extensional tectonic regime on a marginal shelf reflecting tectonic activity to the west and a. continental marpin to the east. Powell (1990) regarded the “Kanmantoo Fold Belt’ as representing the westernmost part of the “Tasman Pold Belt”, He suggested that the Kanmantno Group represents passive margin sedimentation. The term Truro Volcanics was defined by Forbes er ai. (1972) from a type section in Levi Creek, UL km north of Truro (Fig. ), They: included two units within the Truro Volcanics (Fig, 2), with the lower unit comprising a 240 m thick sequence of interbedded volcgnictastic sediments, amygdaloidal volcanics and limestones. The upper unit as described by Forbes et al. comprises 60m of marble, shaly marble, metasiltstane and a thin {2 2) harizon of parphyritic andesite, Forbes et ef, (1972) noted that this sequence is overlain by the Hestherdale Shale which, near the base, contains clasts of volcanic rocks. However, the exact nature of the contact is unclear due to poor exposure. The Heatherdale Slrale, the txp unit of the Normanville Group, is overlain by the Carrickalings Head Formation, the basal wnit of the Kanmantoo Group. Furbes ef al. (1972) suggested thar che marble in the upper member of the Truro Volcanics is 3k C. G. GATEHOUSE, J. B. JAGO, B. J. CLOUGH & AJ. McCULLOCH FNL ABBEWEIT }™ “4 ia ret | at Mr ster 1 4 1 ag ! Trura e — if Cay Sa ca Hill sedan, e AADELMIDE a" fe SE Br’ O02 4 Ne SUBSURFACE DISTRIBUTION S \ Ye Metadalerités ye Ye NA Bosalt - andésite Waster’ limit) oF Murray Basi fe KILOMETRES LOCALITY \ — 5 he Sutherlands Maculia ~ “sea GASTON AQ _aedon Hill wt om ENLARGEMENT j Seen Fig, 1. Locality Map. The-type section of the Truro Volcanics is 2 km south of Mt Rufus 1. equivalent to the Fork Tree Limestone which at Sellick Hill. conformably underlies the Heatherdale Shale (Daily 1963). Rocks assigned to the Truro Volcanies by previous workers occur in outcrop only in the Karinya Syncline, north-east of Adelaide as shown by Cobb & Farrand (1984). A recent report which may extend the known distribution of the Truro Volcanics is that of Polomka (1988)! who reported float material of possible Truro Volcanics about 2.km NNE of Moculta, near the base of what he mapped as Carrickalinga Head Formation. However, the exact stratigraphic position of these rocks is doubtful because of the poor outcrop in the arca, Polomka described the rock as comprising fine to medium grained phenocryst pseudomorphs of epidate (2%), within a very fine actinolite (45 %) and epidote (30%) groundmass. 'Polomka, S. M. (1988) The geology of the Mt Karinya area, (South Australian Institute of Technology, Departinent of Applied Geology, Unpublished thesis). 2Cooper, B. 1. & Gatehouse, C. G. (1988) Sedan Hill, Carrickalinga Head Formation. Ia Gatehouse, C. G, (compiler) Kanmantoo Field Symposium Excursion Guide. S. Aust, Dept Mines & Energy Kept Bk 88/35, 3Gatehouse, C, G,, Jago, L B. & Clough, B. I. (1991a) 4 progress report on a measured reference section at Red Creek for the Kanmantoo Group in the Karinya Syncline. §. Aust. pert Mines & Energy Rept Bk 91/27. §MicCullech, A. J. (1990) The gealogy of the Towitta area. (South Australian Institute of Technology, Department of «Applied Geology, Unpublished thesis). 5Van der Stelt, B. (1990) ‘The geachemisiry, petrology and tectonic setting of the Truro Volcanics, (University of Adelaide, Unpublished B.Sc. (hons) thesis). 5Gaichouse, C. G., McCulloch, A. J., Clough, B. J. & Sarunic, W, (19910) Mi Rufus 1 Well Completion Report. 3° Aust. Dept Mines & Energy Rept Bk 91/25. TRankin, L.. R., Clough, B. J. & Gatehouse, C. G. (1991a) Mafic suites in basement beneath the Murray Basin: new data for the Early Palaeozoic history of the Tasman orogenic gprovince. S. Aust. Dept Mines & Energy Rept Bk 91/44 Rankin, L..R.. Clough, B. J,. Farrand, M. G., Bartek 8. R., Lablack, K. Gatehouse, C. G. & Hough, L . P. (1991b) Murray Basin’ basement transect project: 1990 well er i reports, §. dust. Dept Mines & Energy Rept BR 125, EARLY CAMBRIAN “TRURO VOLCANICS" 59 TRURO VOLCANICS TYPE SECTION DESCRIPTION CUMULATIVE THICKNESS (metres) ROCK UNIT Phyllite: Conglomerate and breccla: ... calcitio; of volcanic Iragments, scoriaceous, pebbles of porphyritic andesite. HEATHERDALE SHALE shaly in part, more massive near base with calcareous siltstone and thin interbeds ot dark volcanic rock with large feldspar phenocrysts (position of volcanics uncertain). FORK TREE LIMESTONE Volcanic rock: .. pinkish-grey. Weathered. Breccla: grey, amygdaloidal voicanic fragments, calcite matrix. Limestone; ..-- grey, cream - weathering. Voleanics: ..... green, phyliitic. Voleanics: ..... shaly. Exposure poor. VOLCANICS Volcanics: on top of calcareous greywacke? Green volcanics: with minor grey amygdaloidal volcanics, conglomerate at top of volcanic pebbles. Limestone: ,... pale graen-grey. Falds plunge approximately 50° to south. Volcanic rocks: . green apidote, calcite veined. Volcanic breccia: im thick of 120mm clasts of vesicular volcanics. Volcanic rocks: . metabasalt?, minor grey amygdaloidal volcanics. Volcanic rock: . . amygdaloidal, calcita vains; amygdales 7mm long, calcite-filled. Elongation trends 170°. Volcanic rock: . . fine grained. No outcrop. Green phyllite, gray limestone, chert, Silty phyllite, bioturbated. pale grey, medium-grained laminated quarizite. Fig. 2. Stratigraphic section al type section of Truro Volcanics, {1 km north of Truro (after Forbes er al, 1972), 60 C. G. GATEHOUSE, J. B, JAGO, B. J. CLOUGH & A.J. McCULLOCH At the extreme eastern edge of the Mt Lofty Ranges, Sutherlands (Forbes ef al. 1972), The best exposure Early Cambrian volcanics have been recorded at Sedan of Early Cambrian volcanics from the eastern Mt Lofty Hill (Cooper & Gatehouse 1988”; Gatehouse et al. Ranges are those mapped in Red Creek by Coats & 1990), at Red Creek (Gatehouse et al. 1991a*; Thomson (1959) as feldspar porphyrite. The remainder McCulloch 1990*; Van der Stelt 1990°), of this paper deals largely with the Red Creek area, Accomodation Hill (Forbes er al. 1972) and near north-west of Sedan (Fig. 1). Measured section (m} | 1 ' es S 3 KANMANTOO GROUP Carrickalinga Head Formation KANMANTOO GROUP Cartickalinga Head Fmn NORMANVILLE GROUP Heatherdtale Shale and Truro Volcanics wip 3 5 o of sc o> ce Oe TT ie do 2a = Fag o ® x acd © 2 a = COWG_LOMERATE Slistone Sandstone Shale Liensstone Fippies Velcanics Crans Inmination BHALE FINE SAND MEDIUN SAND f- COARSE SAND NE IMPAATE Pillow java Rack sampie ipcoton SILTY SADME 92-1044 Fig. 3. Stratigraphic section. at Red Creek. EARLY CAMBRIAN YOLCANICS 4it Stratigraphic setting Detailed measured sections at Sedan Hill (Cooper & Gatehouse 988) and at Red Creek (Gatehuuse er al, I99la*) show that volvanic rocks, generaily referted wo as the Truro Voleanics are interbedded within the Heatherdale Shale and cogiinue imo the basal paris of the gradationally overlying Carrickalinga Head Formation. The Red Creek section has been logged an some detail (Gatchouse et.al. 1991b%); only a summary of the section is given here (Fig, 3). About 300 m of Hextherdale Shale is exposed at Red Creek; the base is not exposed due io the presence of the Palmer Fault. As exposed at Red Creek the Heatherdale Shale is a pale grey larninated phyllite which becomes darker and less micaceous up-section. There are minor calcareous siltstone honzons, Within these units are thin tuffaceous and/or volcanogenic siltstone horizons which become increasingly abundant up-section About 200 m above the base of the measured section are several basaltic lava flows, one of which exhibits pillows (Fig. 4). The pillowed flow has an uneven base and cuts down into an underlying 0.2m thick tuff horizon; it appears to thin to the south. It contains several pillow strictures with chilled margins and triple junctions (Fig. 4). An intrusive igneous body seen between 145 and 150 m on the measured section may represent 2 feeder pipe to the lavas higher in the section, Above the pillow lavas the tuffaccous/ voleanogenic siltstone horizons continuc with reduced Frequency into the basal 60 m of the gradationally overlying Carrickalinga Head Formation. Detailed descriptions uf the complete stratigraphic section will be given in a later paper. In the north branch of Red Creek (Fig. 3), close to the buse of the Carrickalings Head Formation, there are several beds, and/or blocks, of crystal tuff comprising almost pure feldspar crystals clearly winnowed from-enclosing ash. Such beds may have Fi Wig. 4+. Pillow lavas al Red Creek. Note the triple junction al centre left. formed by current activity at the site of deposition or by differential sir-fall separation, Regional stratigraphic interpretation As noted above, tm Red Creek, lavas and voleatogesic sediments extend thorugh the exposed Heatherdale Shale up into the base of the Carrickalinga Head Formation. However, near the type section (Fig. 2) of the Trure Volcanics 24 kin NNW of Red Creek, the highest known volcunics are at least 200 m below the top of the Heatherdale Shale (see imag in Forbes et al, 1972. and Fig. 2 herein}, Indeed Forbes e ul. (1972) suggest that the bulk of the volcanics Lit the type area occurs below an equivalent of the Fork Tree Limestone, which on Fleuricu Peninsula ties conformably below the Heatherdale Shale, However, sinoe this correlation with the Fork Tree Limestone is made on lithological grounds only, and furthes, that there are calcareous horizons within the Heatherdale Shale both on Fleurieu Peninsula and at Red Creek, then there may be some doubt about the correlation. It could be argued (hat the marble of the cype area of the Truro Volcanics, described as Unit C4 by Forbes etal. (1972), is equivalent to the calcareous horizons found in the Heatherdale Shale at Red Creek rather than being equivalent to the Fork Tree Limestone, However, if the correlation of the Unu C4 marble in the type section to the Fork Tree Limestone is correct then it Sugpests that the volcanics at Red Cresk are in a higher stratigraphic position than those of the type section and should not be referred in the Tro Volcanics. In his discussion on the Stansbury Basin, Gravestock (is press) has included all these volcanics as Truro Voleanics. A further compheation is the almost complete Jack of exposure of the basal 150 a3 of the Heatherdale Shale near the type section of the Truro Voleanics. The recent drilling of Mt Rufus Na, 1 stratigraphic hole 2 fam north of the type section suggested that there may be several unrecugnised faults in the area (Gatehouse et al. '991b*). Until the position is clantied the volcanics described herein sre simply referred to as the volcanics fram Red Creek, Elsewhere in the Stansbury Basin, green tuff beds in the Parara Limestone may be correlated with the Truro Volcanics of the Karinya Syncline (Gravestock in press). A tuff bad from within the Heatherdale Shale at Sellick Hill has been dated at 526+4 Ma (Cooper «1 al. 1992), this tuff may also be equated with the Truro Yolcanics or the volcanics at Red Creek o¢ with neither, Petrography of volcanics at Red Creek In hand specimens the voleames in Red Creek have 4 grey fine vrained gtaund mass with distinctive large a2 C. G, GATEHOUSE, J. B. JAGO, B. J. CLOUGH & A.J. McCULLOCH (approximately 0.5 mm diameter) phenocrysts of feldspar (up to approximately 25%) and minor iron staining. The rocks show variable intensities of tectonic foliation. In thin section it appears that the phenocrysts are predominantly of zoned plagioclase although approximately 10% of the phenocrysts are of an alkali feldspar. The groundmass is predominantly of acicular feldspar laths producing a trachytic texture, The tectonic fabric of these rocks varies from slight to intense foliation, the feldspar phenocrysts having rotated parallel to the foliation, Alteration is pervasive with sericitisation of the feldspar and chlorite/iron- oxide replacement of mafic minerals. Sporadic veinlets of quartz and calcite cross-cut this tock, Petrologically the lavas classify as trachytic basalt. Geochemistry of volcanics at Red Creek Two samples (6728 RS 1632 and 6729 RS 1515) of massive pillow lava were taken from separate localities FLEURIEU TRUAO PENINSULA TYPE SECTION Dally 1983, ; Daily & Milnas 1972 Modified after Forbes et al, 1972 CAMPANA CREEK MEMBER BLOWHOLE CREEK SILTSTONE MEMBER MADIGAN CARRICKALINGA INLET HEAD MEMBER FORMATION HEAD CARRICKALINGA FORMATION 7 HEATHERDALE HEATHERDALE SHALE SHALE — ? FORK TREE LIMESTONE FORK TREE LIMESTONE SELLICK HILL FORMATION TRURO VOLCANICS WANGKONDA FORMATION 2. MOUNT TERRIBLE FMN > — MOUNT TERRIBLE FMN EQUIVALENT ? near Red Creek and geochemically analysed for a comprehensive suite of elements (Tables 1 and 2). Several analytical methods were used, including: ICP (acid digestion) — major elements XRF — As, Ba, Bi, Sb, Sn, V, Zr Atomic Absorption Spectrography — Ag, Cr, Cu, Ni, Pb, Zn Fire Assay — Au, Pt, Pd ICP Mass Spectrography — Ce, Dy, Nd, Er, La, Eu, Lu, Yb, ¥, Sm, Gd, U, Th, Sr, W, Ta, Mo, Nb, Ga, Co, Cs, Rb. The two geochemical analyses (Tables 1 and 2) of the pillow lava reflect their described lithology, as basaltic lavas. that have undergone preenschist facies metamorphism. Elevated loss on ignition (LOD values (average 11.5%) attest to alteration effects resulting in hydration. Clearly the present chemical composition of the lavas at Red Creek is not primary, as the volatile content is significantly higher than in analogous fresh rocks. This is to be expected from the presence of SEDAN RED HILL CREEK Altar Gatehouse (this paper) et af, 1990 CAMPANA GREEK MEMBER BLOWHOLE CREEK SILTSTONE MBR? MILENDELLA LST MBA MADIGAN INLET MEMBER CARRICKALINGA HEAD FORMATION CARRICKALINGA HEAD FORMATION MADIGAN INLET MEMBER HEATHERDALE SHALE with VOLCANICS HEATHERDALE VOLCANICS BASE CUT OFF BY FAULT BASE CUT OFF BY FAULT Fig. 5. Correlation diagram of the Cambrian sequences of the Red Creek-Truro-Sellick Hill areas EARLY CAMBRIAN VOLCANICS 62 Taste {. Geochemical analyses and CIPW norms, NB, CIPW weight % norms are calculated from analyses recalculated to 100% free of HO and CO,; ce is not recalculated, Fe data is recalculated nsing a Fe,: Total Fe cation cation of 0.20. Major Elements in Percent CIPW Weigh % Norms 6728 RS 6728 RS 163255 163255 SiO, 42.30 4230 ab 1885 19.92 TiO, 242 224 or 1971 13.69 ALO, 1580 1440 an 1847 1531 Fe,0, 875 BOS ne 985 13.29 FeO di W823. Mad 0.28 626 al 467 321 Meo 264 #332 mt 294 272 Cad 805 «885 il 532 4.95 NasO 378 450 O73 x6 288 (1.99 P.O, 0.78 60.66 Total 100.01 100.0) HO+ H,0 cb, Lol 10.80 12.30 Total 9848 0887 DE 4841 46.82 =not analysed, DI=Differentiation Inmlex hydrous secondary minerals in these lavas. Similar alteration of mafic lavas in the Victorian greenstone belts (Crawford & Keays 1978) is.considered to have caused hydration, along with slight addition of CO, and Na,O accompanied by leaching of SiO,, CaO, ALO, and K,O, However, the degree of chemical change was considered to be minimal, and magmatic trends Werte clearly visible. To minimise the effect of hydration dilution and related chemical mobility, plots using elements considered immobile during alteration are used and analyses are recalculated to 100% volatile- free prior to plotting. The lava at Red Creek plot on the border between phono-tephrite and tephrite-basanite close to the basallic-trachyandesite field in the SiO, versus Na,O+K,0 classification plot of Le Bas er al. (1986; Fig, 6), In the Nb/Y versus Zr/TiO, classification plot (Winchester & Floyd 1977; Fig. 7) using elements considered immobile during alteration, the lavas classify as alkali basalt. To further define the alkaline basalt a classification scheme devised for the Tertiary alkaline volcanics of eastern Australia (Johnson 1989) based on CIPW norms was utilised (Fig, 8). Using this classification scheme the division between sub-alkaline and alkaline mafic lavas is that alkaline lavas have <10% normative hypersthene; on this basis the basalts in Red Creek classify as alkaline and plot within the field of hawaiites. The presence of considerable levels (9.85 and 13.22%) of normative nepheline suggests these rocks are silica undersaturated. However, plots using immobile elements (Fig. 7) suggest these lavas have not attained silica undersaturation but show that the lavas at Red Creek and the Truro Volcamics at Mt Rufus 1 (Gatehouse et af. 1991b*) belong to 4a group of analyses that straddle the boundary between alkali basalt and silica undersaturated nepheline/basanite fields. The analyses agree with field evidenve suggesting that the lava at Red Creek represents a single thin submarine Java sequence from a common source, in that they plot close together on all classification plots. The lavas plot outside the tectonic discrimination fields of Pearce & Cann (1973; Fig. 9), although closest to the intraplate field; and in marked contrast to the MORE-telated metadolerites that occur in the Murray Basin basement (Rankin et al. 19912, b74) and ag sills and dykes in the Mt Lofty Ranges (Liu & Fleming 1990; Rankin ef af. 1991a”). On a MORRB nommalised spidergram (Fig, 10) the Red Creek lava shows a distinctive tread Of elemental enrichment relative to MORB values of the incompatible eletnents from Sr through to Ni, which is typical of the more silica TABLE 2. Trace elements in ppm. 6723 RS 6729 RS Detéction 4632 155 lienit (ppm) Ap <1,00 < 1,00 1 As 64.00 72.00 2 Aw 2,00B 7.008 iB Ba B80 610 id Bi 4,00 <4.00 + Ce 82.00 73.00 Ot Co 29.00 44,00 I Cr Ito LOS 4 Cs 1,30 t.20 2 Cu 6.00 $5.00 2 Dy 3.00 7.20 0.1 Er 3.90 3.20 O1 En 2.80 2.50 O.1 Ga 26,00 20.00 1 Ga 9.00 8 60 a4 La 38,00 36,00 a1 Lu 0,60 0.50 0,1 Mo 4.00 3,55 o.s Nb 76,00 66.00 os Nd 44.00 42.00 0.1 Ni 52,00 54.00 4 Pe 12.00 $2.00 4 Pa 1.008 <=1.00B LB Pt <5,00B <5.0)8 5B Rb 22,0 60.00 0.2 Sb <4.00 4.00 os Sm 9.40 a OL Sn <4.00 10.00 4 Sr 240 255 0) Ta 3.00 2,20 02 Th 4.80 3.60 0.1 U 2.60 2.20 aa v <5.00 <5,00 s WwW 3,00 3,00 ? Y 34.00 29.5 | Yb 3.70 3,00 OL Zn 22,00 76.00 3 Zr 380 320 + o undersalurated alkaline mafic layas of such provinces as the Tertiary eastern Australian volcanic province (Johnson 1989); note the marked contrast of the alkaline intraplate lavas with MORB related metadolerite from MB-12 in the Murray Basin basement (Rankin et al. 199la’ and b*). Other alkaline volcanic areas similar to Red Creek include the Yumali/Coonalpyn area, the Peebinga-1 16-00 12-BO a 60 6-40 Naz0 + K20% 320 Tt 4av0 82-40 35-00 BOG 580 7TDO SI02% A.,...,,.Hed Creek matic tava Fig. 6. Red Creek lavas on port of classification plot of Le Blas et al, (1986). Values calculated to 100% volatiles free. Zr/Tio; i] Rhyalite — ae Ahyodacite/Dacila 04 Tag A, Trachyandesite eS. Sub-alkaling Basalt 0.004 Nb/Y¥ 12] Mt Rufus’ 1 © Red Creek Fig. 7. Red Creek and Mt Rufus L lavas on part of classification plot of Winchester & Floyd (1977). Cc. G. GATEHOUSE, J. B. JAGO. B, J. CLOUGH & AJ. McCULLOCH (MBT-1) drillhole into the northwestern termination of the magnetically defined Mt Stavely Belt (Rankin ez al. 1991a’, b), and the Truro Volcanics as defined in Mt Rufus 1 drillhole (Gatehouse et a/. 1991b°: Fig. 10). Of the other volcanics the nearest are the Truro Volcanics that crop out on the west limb of the Karinya ‘Syncline 22 km NW of Red Creek; the lavas at Red Creek represent a more evolved lava type than the Truro Volcanics in their type section, resulting in a line more distant from’ MORB in Fig. 10. This chemical difference is seen in the petrology where the lavas at Red Creek are distinctively porphyritic with phenocrysts of alkali feldspar. The Red Creek analyses are compared with a classic continental intraplate voleanic suite in Fig. 11, where they are normalised to Karoo type basalts, the lavas at Red Creek and the other Cambrian alkaline volcanics noted above exhibit enrichments relative to Karoo basalt of the elements Sr‘to Ti, with Red Creek being one of the most chemically evolved suites. This supports the field relations which suggest that these lavas are not directly analogous to the thick piles of dominantly tholeiitic basalt fissure lavas of the Karoo Province, but rather are more closely analogous to the more localised central complexes seen in intraplate alkaline volcanic complexes (Johnson 1989). In summary, it is proposed that the lava at Red Creek represents a single flow of hawatite composition from a central volcanic complex, closely analogous to that of the Truro Volcanics but not necessarily from ihe DIFFERENTIATION INDEX 100 felsic focks 80. benmoreite 60 intermediate matic (.¢. alkali, basalt, basanite, hawaiila, nephelinite) TOCKS 40 mugearite hawaiite 0 20 40 60 80 NORMATIVE 100 x (an/(an + ab)) 100 a. .,Red Creek mafic lava Fig, 8, Red Creek lavas on CIPW normative classification plot for sub-alkaline intraplate lavas (otinson 1989). EARLY CAMBRIAN VOLCANICS 65 same centre or erupted at exactly the same time, However, both the laya at Red Creek and the Truro Volcanics undoubtedly belong to the same Early Cambrian alkaline ‘within plate’ volcanic province, which may be genetically linked to other Cambrian- mafic alkaline provinces such as the Yumali/Coonalpyn and Peebinga-1 areas (Rankin et a/. 199la’ and b’). The closest tectonic analogué for these provinces is that of a rifted continental margin-as proposed for the Ti / 100 zr Yu a Red Creek matic volcanics Fig. 9, Red Creek volcanics on tectonic discrimination plot of Pearce & Cann (1973). “Within plate” basalts plot in field D, Morb (ocean floor basalts) in field B, low Ktholeiites in field A and B, calc-alkaline basalts in fields C and B, NORMALISED VALUES. 0.1 t Sr kK Rb Ba Th Nb Ce P Zr Ti ELEMENTS ¥ Sc Ni Cr Red Graek volcanics Mt Rufus 1 lavas ——— MBT-1 lavas Yumali / Coonalpyn lavas MBY-12 matadolarie Fig. 10. MORB normalised spidergram for averages of possible Cambrian mafic suite from the Murray Basin basement and. the Cambrian lavas of the Mt Lofty Ranges. Tertiary intraplate volcanic province of eastern Australia (Johnson 1989); metadolerites which post- date the volcanics indicate 4 change with time from intraplate to MORB type composition (Liu & Fleming 1990) indicating an ensuing period of crustal thinning and major dyke emplacement, associated rifting and crustal extension. Acknowledgments The authors wish to thank Dr David Gravestock (S.A. Dept of Mines & Energy) for reviewing this paper and making some yaluable suggestions. Logistical support was provided both by the §.A. Dept of Mines & Energy and the University of South Australia, This work was supported by a University of South Australia Research Grant to JBJ, This paper is published with the permission of the Director General of the S.A, Dept of Mines & Energy, References Coats, R. P, & THomson, B. P. (1959) TRURO map sheet, Geological Atlas of South Australia, 1:63 260 Series. (Geol, Surv. S. Aust., Adelaide), Cops, M. A. & Farranb, M. G. (i984) A new occurrence of the Truro Volcanics. Quart. Geal. Notes Geol, Surv. S. Aust. 89, 8-10, Cooper, J. A., Jenkins, R. J, FE Comeston, W. & Witutams, I. 8. (1992) lon-probe zircon dating of a mid- Early Cambrian wf in South Australia. Geol, Soc. Lond. Journal 149, 185-192. Crawrorp, A. J, & Keays, R. R. (1978) Cambrian greenstone belts in Victoria; marginal sea-crust slices in the Lachlan Fold Belt of southeastern Australia, Earth & Planet, Sc. Lett. 41, 197-208. 100 NORMALISED VALUES Red Creek yolcanics > ‘Mt Rufug 1 lavas MBT-1 lavas Yumali / Coonaipyn lavas MBT-12 metadalerita Fig. ll. Karoo Basalt normalised spidergram for averages of possible Cambrian mafic suites from the Murray Basin basement and the Cambrian lavas of the Mt Lofty Ranges. 66 ©. G, GATEHOUSE, J. B. JAGO, B. J. CLOUGH & A. J. McCULLOCH Dany, B. (1963) The Fossiliferous Cambrian succession in Fleurieu Peninsula, South Australia, Rec, 5. Aust. Mus, 14, 579-601, & Mivnes, A. R. (1972) Revision of the stratigraphic nomenclature of the Cambrian Kanmantoo Group, South Australia. J; Geol. Sac. Aust. 19, 197-202. Forses, B. G., Coats, R. P. & Datiy, B. (1972) Truro Volcanics, Quart. Geol. Notes, Geol. Surv. S, Aust. 44, 1-5. GateHouse, C. G, (1988) TEPKO map sheet. Geological Aulas of South Australia , 1:50,000 Series, shect 6728-II!. (Geol. Surv, §. Aust., Adelaide), GateHouse, C. G., Jaco, J. B. & Cooper, B. J. (1990) Sedimentology and stratigraphy of the Carrickalinga Head Formation (low stand fan to high stand systems tract), Kanmantoo Group, South Australia, J J. B. Jago & P. J. Moore, (Eds) “The evolution of a Late Precambrian-Early Palaeozoic rift complex: The Adelaide Geosyncline” Geol, Soc, Aust. Spec. Pub,, 16, 351-368. Gravestocx , D, I. (in press.). Early and Middle Palaeozoic. In “Geology of South Australia.” (South Aust. Dept. Mines & Energy, Adclaide) Jounson, R. W. (Ed) (1989) “Intraplate volcanism in eastern Australia and New Zealand”. (Cambridge University Press, Cambridge). Le Bas, M. J., Le MAITRE, R. W., STRECKEISEN, A, & ZANETTIN, B, (1986) A chemical classification of voleanic rocks based on the total alkali-sili¢a diagram. J. Petrol, 27, 745-750. Lru, S. F & Fremia, P. D. (1990) Mafic dykes and their tectonic setting in the southern Adelaide Foldbelt, South Australia pp.401-413. Jn A. J, Parker, P. C. Rickwood & D. H. Tucker, (Eds). “Mafie Dykes and Emplacement Mechanisms”. (Balkema, Rotterdam). Parker, A. J. (1986) Tectonic development and metallogeny of the Kanmanton Trough in South Australia. Ore Geol. Rev. I, 203-212. PEARCE, J. A. & CANN, J. R, (1973) Tectonic setting of basic volcanic rocks determined using trace element analyses. Earth & Planet. Sc. Lett. 19, 290-300. PoweLL, C. Mc. A. (1990) Gondwanaland context of the Tasman Fold Belt. Geol. See. Aust. Abstracts, 25, 190-191. SCHEIBNER, E. (1986) Suspect terranes in the Tasman Fold Belt System, Easter Australia. Jn D. G. Howell (Ed.) *“Tectonostratigraphic terranes of the Circum-Pacific Region", Circum Pacific Council for Energy and Mineral Resources, Earth Science Series. 1, 493-514. TuRNER, S. P. & Fopen, J. D. (1990) The nature of mafic Magmatism through the development of the Adelaide Geosyncline and the subsequent Delamerian Orogeny, South Australia pp. 431-435 In A. J. Parker, P. C. Rickwood & D. H. Tucker (Eds) “Mafic Dykes and Emplacement Mechanisms”. (Balkema, Rotterdam). Von DER Borcu, C. C. (1980) Evolution of Late Proterozoic to Early Palaeozoic Adelaide Foldbelt, Australia. Comparison with post-Permian rifts and passive margins. Tectonophysics 70, 115-134. WINCHESTER, J. A. & Fuoyb, P. A. (1977) Geochemical discrimination of different magma series and their differentiation products using immobile elements. Chem. Geal, 20, 325-343, BIOLOGY OF ETIELLA BEHRII ZELLER (LEPIDOPTERA: PYRALIDAE): A PEST OF SEED LUCERNE IN SOUTH AUSTRALIA BY A. D. AUSTIN, T. C. R. WHITE, D. A. MAELZER & D. G. TAYLOR* Summary The general biology of the pyralid moth, Eviella behrii Zeller, a serious but sporadic pest of lucerne seed crops in South Australia, is documented. Larvae are recorded as feeding on six native and 18 introduced legume species. Head capsule widths of field-collected larvae show that E. behrii, like other pyralids, has five instars. Light-trap catches indicate the presence of at least four overlapping generations per year in the Adelaide area, with the majority of moths occurring during three generations from August to February. In lucerne crops in the Keith area, 150 km south-east of Adelaide, young and mature green pods suitable for oviposition are available from October to February. Two peaks in moth catches were recorded in late December and late January, respectively. Until mid-January, the number of eggs was strongly correlated with the numbers of moths caught, but thereafter the numbers of eggs and larvae decreased, with the second peak of moths giving rise to very few larvae. These observations are in agreement with the opinion of farmers that early lucerne crops are more heavily infested with E. behrii than late crops. Surveys of vegetation in the Keith area show that native host plants are very scarce, and that volunteer lucerne plants along road-sides and irrigation banks are the likely source of early infestations in Lucerne crops. Mortality agents recorded include ten species of parasitoids, two pathogens and a number of general heteropteran and spider predators. These and other possibilities are discussed as factors leading to the observed decrease in larval numbers in mid to late January, a time when lucerne crops still contain a large proportion of green pods suitable for larval development. KEY WORDS: Etiella behrii, Pyralidae, ecology, life history, lucerne, alfalfa, host plants, parasitoids, predators, pathogens. Transactions ef the Reval Society af S. sust. (1993), 117(2), 67-76. BIOLOGY OF ETIELLA BEHRII ZELLER (LEPIDOPTERA: PYRALIDAE)< A PEST OF SEED LUCERNE IN SOUTH AUSTRALIA by A, BD, Austin, T. C. R. WHiTE, D. A. MABLZER & D. G. TayvLor* Summary Ausiin, A. D.. Warre. 'T. C. R., Magvzer, D, A. & Taytor, D, G. (1999) Biology of Fella behrtt Zeller (Lepidopten; Pyralidae): a pest of seed lucerne in South Australia. Trans. R. Soe. 8. Aust. U7(2) 67-76, 4 June, 1993, The general biology of the pyralid moth, Eriella behrii Zeller, a serious but sporadic pest of lucerne seed crops in South Australia. is documented. Larvae ure recorded as feeding on six native and 18 introduced Jeguine species Head capsule widths of field-collected larvae show thal £. behrii, like other pyralids, has five instars. Ligh(-trap caiches indicale {he presence of at feust four overlapping generations per year in the Adelaide ares, will! the ninjority of yroths Occurring during three generations from August to February. In lucerne crups in the Keith area, 150 km south-east of Adelaide, young and mature green pods suitable for oviposition are available from October to February, Two peaks in moth catches were recorded in late December and late January, respectively. Until mnid- January, the number of eggs was strongly cocrelated with the numbers of moths caught, but thereafter the uuinbers of eggs and larvae decreased, with the second peak of moths giving rise to very few larvae. These observations an: in agreement with the opinion of farmers that early lucerne crops are more heuvily infested with £. bebtrit than late crops, Surveys of vegetation in the Keith area show that native host plants are very scarce, und than volunteer lucerne plants along road-sides and irrigalion banks are the likely source of curly infestations if Wweerne crops. Mortality agents recorded include ten species of parasitoids, two pathogens. and a number of general heteropteran and spider predators, These and other possibilities are discussed as factors leading to the observed deerease in larval nufubers in mid to lute January, a time when Jucerne crops still contain a large proportion of green pods suitable for larval development, Kev Worns. “ella behrii, Pyralidac, ccology, life history, lucerne, alfulfa, host plants, parasitoids, predators, pathogens. Introduction Ole of the most serious pesis which ean limit the production of lucerne seed in South Australia is Eriella behrit Zeller (lucerne sced web moth) (Maclzer ef al. 1982u, 1982b; Durhanr 1984; Austin ef al, 1986). It is One of seven species of Friella, the larvae of which feed on the seeds of legumes (Whalley 1973), The most witlespread and economically important species is the cosmopolitan &. zinckeneller (Treitschke), a serious pest of Jepunies in many parts of the world, which in Australia is known only from Queeasland (Abdul-Nasr & Awadalla 1957; Stone 1965; Singh & Dhooria 1971; Whalley 1973; Common 1990), The remaining six species are restricted to Australia, parts of south-east Asta and some Pacific islands, Only three species, E. chrysoporelia Meyrick, EB. hobsont (Butler) and E. behril, are recorded from South Australia; the former twoonly curely as adults, and only from the northern and western parts of the State, The multivoltine £. behrii is a widespread and common pest throughout Australia, and of economic significance because of its heavy but sporadic outhreaks in dryland and irrigated crops of seed lucerne. * Departmen of Crop Protection, Waite Ayricultural Reseurch Institute, University of Adelaide, Glen Osmond, 5. Aust. 5064, Little is known of the general biology or ecology of £. hehrii, possibly because of its sporadic occurrence. In most years it does little damage to lucerne seed crops in S.4., but occasionally it causes widespread damage. Atsuch times the species is also reported fo damage lupins, field peas and some clovers, e.g. in 1954 and 1971 (Austin eral, 1986; DAM and DGT, unpubl.). £. behrii is thought to live on native plants along road-sides and in patches of scrubland during spring and eatly summer before infesting lucerne seed crops in mid summer, but this has not been confirmed. The present study was therefore undertaken to elucidate this pari of its phenology and 10 document its genetal biology. Data have been included from a pest management program on Jucerne seed crops (see Maelzer et al. 1982a, 1982h, Durham 1984), information collected in 1959-60 (by DGT), and life history studies undertaken during 1980-82, Materials and Methods Location of study The project was centred on a property (‘Brecon’) 10 km south of Keith, S.A., inthe middle of a mayor area of ducerne seed production The area bas 4 Mediterranean-type climate with mostly cold, wet winters and hot, dry summers (Anon, 1987). The 68 A. D. AUSTIN, T. C. R. WHITE, D. A. MAELZER & D, G. TAYLOR average rainfall for the area is 471 mm per annum. The early (spring) dryland lucerne crops are watered naturally by rainfall, but the majority of the seed crops are managed to flower in mid-summer and are irrigated, mostly by flood irrigation rather than by sprinklers. Work in the Adelaide area, which has a similar climate to Keith, was carried out in 1959-60 and was centred at the Waite Institute campus of the University of Adelaide. Trap catches of adult moths Adults were caught at a light trap each night in two separate studies in widely different years: over the period 1.111.1959 to 18.11.1960 on the Waite Institute campus (by DGT) and from 1.xit.1981 to 20.11.1982 at ‘Brecon’ (by DAM). The second period of trapping was part of 4 general program of sampling insects in lucerne seed crops (Maelzer er al, 1982a, 1982b). The light trap in 1959-60 was a mercury-vapour type, while that used in 1981-82 was a 15 watt blacklight type (cf. Hardstack er al. 1979), located in the middle of a lucerne area containing both dryland (early) and irrigated (late) crops, Sampling eggs and larvae from lucerne crops Five sampling sites were selected at ‘Brecon’ around a field of lucerne which was to be irrigated later that summer. All were dryland areas (i.e. not irrigated) and were of the commonly grown Australian cultivar, Hunter River. Each had been closed off to grazing animals some weeks earlier and contained lucerne plants with flowers and pods. Samples were collected from these sites about once per week from the beginning of December 1981 until mid January 1982 and taken back to the laboratory. Ten small samples (each of about 50 pods) were also taken from one or two more distant, dryland sites (up to 30 km away). including fields of lupins with volunteer lucerne plants, Each sample was taken by walking through an area of 4-5 ha and picking stems of lucerne with flowers and pods from a number of widely spread points until 15-20 stems had been gathered. The stems were temporarily stored in a plastic bag on ice. In the laboratory each stem was searched, ftom the oldest raceme upwards, and each pod on each raceme was examined for eggs and larvae on the outside, and larvae inside. The procedure was continued until approximately 200 pods had been inspected. This meant searching from 1-12 stems (x = 5.0; $.D. = 1.9) bearing from 1-1] racemes each (X = 5.3; §.D, = 2,3). The racemes on any one stem were at any stage from unopened flowers to fully matured brown (dry) pods. All stages of flowers and pods were usually present on a single stem and, wherever possible, stems were selected that had some well-developed pods. For each raceme a record was made of the number of E. behrii eggs and larvae (classed as instars: 1 + 2,3 + 4, or 5) and the stage of development of the pods (classed as flowers, pods just forming, juvenile pods, young pods, mature green pods and mature brown pods — see Fig. 1). In earlier samples, no stages of E, behrii had been found on flowers, on pods that were just Fig. 1. Stages of pod development on lucerne plants: A, pods just forming; B, juvenile pods; C, young pods; D, mature green pods; E, mature brown (dry) pods, BIOLOGY OF ETIELLA BEHRII ZELLER (LEPIDOPTERA PYRALIDAE) 69 Fig, 2. Female Efiella behrii ovipositing onto a raceme of mature green lucerne pods. forming, on juvenile pods or on mature brown pods. Therefore, in later samples racemes bearing only these stages were excluded from samples, and only racemes bearing some young and mature green pods were examined. Later in the crop season, when the dryland crop had nearly all brown pods, similar samples were taken from an adjacent crop of irrigated lucerne, of the aphid resistant cultivar WL318, but counts were made of larvae only. Because of possible differences in numbers of larvae in young and in mature green pods, the numbers were expressed as stratified means per 1000 pods. Rearing techniques and laboratory culture Techniques for rearing field-collected E. behrii larvae through to the adult stage were developed so that a constant supply of each stage was at hand for life-history studies. Small larvae (Ist and 2nd instars) collected from lucerne pods were successfully reared on a medium based on either powdered peas or lucerne seed (cf. Shorey & Hale 1965), but the agar had to be boiled for twice the suggested time for the medium to set. Each larva was placed in a 2.5 cm’ vial containing 1.5 cm? of medium. Vials were sealed with hard plastic lids. The pupae were removed from the vials and placed in a dry container until the adults emerged. Both reared and field-collected adults were kept in gauze cages and given access to 10% sucrose solution. All stages were maintained in a dimly lit room held at 24-28°C and 75% R.H. in which the generation time was 25-35 days, Cooper (1979)! obtained large numbers of E. behrii eggs from females ovipositing into the crevices of a darkened wooden box. We had no success with this method, nor with a variety of other surfaces including perspex, cardboard, freshly crushed lucerne, and rearing medium with lucerne seeds embedded in it. However, we obtained eggs laid onto freshly shelled peas or fresh mature green lucerne pods. Peas proved most successful because eggs could be easily and safely removed from their smooth surface. Eggs were more difficult to remove from lucerne pods because of their more complex surface. They were collected with a fine paint brush and transferred to a dry glass vial, and the Ist instar larvae were then transferred to a second vial containing rearing medium. With this method we initially obtained 80 eggs and reared 90% through to the adult stage. Surveys of natural vegetation Extensive surveys for E. behrii larvae were carried out in the native vegetation surrounding the irrigated lucerne growing area south of Keith (for approximately 8 km in each direction) and along road-sides between Tintinara and Willalooka (35 km north-west of Keith) in October-November, 1981. Extensive surveys were also conducted by one of us (DGT) in 1959-60 around Keith and Tintinara, and this information was extracted from the unpublished records of the (then) Department of Entomology, Waite Agricultural Research Institute. 'Cooper, D.J. (1979) “The Pathogens of Heliothis punctigera Wallengren”. Ph.D, thesis, University of Adelaide, Unpubl. Ww AOU AUSTIN, TOC. RWI DA MAELZER & DUG. TAYLOR Paresiteidy and pathogens ef F. behvii To trap udults of known parasitonds of £. behrii, ten yellow pan traps fled with soupy water were placed armund the edges of the inigaled WLAI8 lucerne crop and ere CXanuned once per week, The traps were rut Inyin early December 1981 to mid-February 1982. In addition, 41 late-staze A. behrii larvae (4th and Sth instars) collected from both the drylind ung irrigated lucerne crops were reared on artificial deet in the laboratory to check for possible parasitoids, Similarly. one of Us (DCT) collected large numbers af &. beheti faryae From both native and crop plants in S0-Mb io obtain parasitoids. Also, other workers have bred parasitoids from £ behrii collected in the Keith area over the last 40 years. Voucher material of parasitoids collected in the above manner has been lodged in the insect collection at the Waite Institute, and has been used lo collate the duty presented here. Pathogens of 16 field-collected 4th and Sth instar larvae that died soon after in the laboratory wore screened for pathogens by DJ. Cooper at Waite Institute using the methods of Cooper (1979)! Results and Observations Life history stages At all stages of its life cycle & brheii parallels descnptions of bolt the behaviour aud the appearance ole, zinckenella, apart from being slightly smaller in size (Abul-Nasr & Awadulla 1957; Stone 1965; Singh & Dhoorit 1971). The eggs are ovoid, averaging 0.6 mim it length (8.0, = 0.03; n = 50) and 0.4 mm in widih ($.D, = 0,3), colourless when first laid, but Soon turn pale yellow or orange. A day before hatching the hiuck heat capsules and prothoracic shields of the Jatvac are clearly visible hrough the chorion, Eggs are laid on either young or mature green pods, and are usually placed between the pod and the calyx or between the coils of the pod. The frequency distribution of head capsule widths of field-collected larvae sugested tive peaks and hence Nive wstars (Fig. 3). but there was much vatisatien between peaks, The minge of cach instsr was beter differentiated hy the minima of a three-point mowing average peocess (Pig. 3), The mean widths for five inshirs were then calculated as 0.1%, 0.28, 1.44, 01,70, om 0.99 mim wall multipliers between means, for Dyar’s Jaw, of 1,54. 7,59, 1.60 and 141. Five instars were also confirmed from the rearing of £ behrii in ihe laboratory. The Ist instar has a pale yellow body and a very distinetive black head capsule and prothoracic shicld. Soon atier batching it constructs a fine fuinel-like silk tube around itself with one end ullached to the sortie of the pod, i then chews through the pod and into seed where it remains feeding, The entrance hole in the pod quickly heals over. the web tube soon disintegrawes, and no evidence remuing to indicate that the pod as pnlestes!, The larva docs not qonnally leave this firs seed until most of ity contents are devoured. It then atacks the next seed in the pod, although it is now too lange to get iaside it. By the late ard to carly 4th instar, We larva chews its way out of the origmal pod and enters one ar more new pods, eating some (and occasionally all) of the seeds. within Commonly, at this stage most seeds are only partly devoured, ‘fhe larva tow has a bluck head capsule, semi-circular prothoracic shield and spiracles, The pro- and mesothorax are golden-brown, the metathorax and abdomen pale green There are seven dorsal and later longitudinal pale: golden-hrown stripes on the body. By the late dil instar the larva attacks several pods, meshing these and its frass together with silk to form a protective lent. This webbing is the first readily observed outward sign that &. behrit has attacked o erop. By the 3th instar the larva has a pale golden- brown head with dark brown mandibles, The prothoracie shield 1s still black bul the rest of the body ix green wilh a pinkish tinge to the abdomen. The longitudinal bands are more diffuse and pinkish, This Howl instar becomes a steadily darker pynk, the green heing musked completely just prior (o pupation Pupation takes place a dew centimetres benwath the Surfitce of Ihe soil, inside a cocoun of silk, which incorporates particles ol soil. Adult moths are small (10-15 mm long), greyish and distinctive becatise of thew typical ‘inoul’ (formed by the forwurd-directed pulps), the white stripe along the full length of the leading edge of exch lore wing, und the transverse orange band near the buse of each fore wing (Fig. 2), At rest, moths have av ‘alert’ appearance with the head aod thorux raised and the abdometi and distal ends of the wings touching the surface. ‘The antennae of the female are simpoth and undifferentiated While those of the male have an enlarged basal segment and i “bush! of scales on the clonguted second segment The maxillary palps of ihe female diverge at their second yom fo form an-open ‘V" at the ip-of the wneat, These palps are parallel in the male moth, Hast plants and sources. of infesterien Table | lists all the plant species tor whieh & beled larvae have been recorded feeding on thelf seed [fi S.A, Fram 1959 to 1982, Larvae have also been rmeeorded on many of the listed exotic species in Other sbtles, as wellas on-soybeun (Glycine mvay (1L.) Merrill), peauues (Arachis hypegaea L.) and Acacia evelups AL Cun, ex G, Dan in Western Australia, Narihern Territory (Comunoa 1990) and Queenstind (Tumer 1980), all these hosts, lucerne is the dost commonly and severely attacked, but the species alse suiees sporauie damage lu lupins, Geld peas and some Govers (Austin eta, 1986; DAM & DGT, tunpulst BIOLOGY OF ETIELLA BEHRI ZELLER (LEPIDOPTERA PYRALIDAE) 7 8 Frequency 0.4 0.6 Head width (mm) ‘ | tHE } ia .O Fig. 3. Head capsule widths of larvae of Etella behrii collected randomly from the field at ‘Brecon’ (Keith) (n= 329), expressed as a frequency distribution (histogram) and three-point moving averages (MB). The survey of native vegetation in late 1981 revealed that there is. very little native scrub left in the Keith area, and what patches do remain are small, considerably disturbed and have very few native legumes, both in numbers of species and individual plants. Far more commonly encountered are volunteer plants of various exotic weed, crop and pasture legumes (especially lucerne) along road-sides, in clearings in scrubland, and in fields carrying other crops such as Jupins. Lupins are a potential source of recent significance which have been planted more frequently in the Keith area since 1980. Lupins are planted from early winter to early spring, the seed ripening and drying out up until late November, thus providing an ideal carly source of food for larvae, the adults of which emerge early in spring. Samples of 200 pods of green mature lupin pods taken from crops south of Keith and near Tintinara in early November, 1981 showed a small (<1%) infestation of £. behrii larvae. Seasonal phenology Observations in the laboratory indicate that females can lay more than 200 eggs and that eggs hatch within 24 h at 30-35°C, but usually hatch after 4-7 days in the field in spring-summer, Larval duration in the field during spring-summer varied from 10-28 days and the pupal period from 14-21 days. Adults survived 7-21 weeks in the laboratory when held at ambient temperature and supplied with 10% sucrose solution. The light-trap catches of adults at the Waite Institute campus in 14 day intervals for the period 1.iii.1959 to 18.11.1960 are given in Fig. 4. These catches suggest 72 A. D. AUSTIN, T. C. R. WHITE, D, A, MAELZER & D. G, TAYLOR TABLE 1. Known host plants of Etiella bebrii in South Australia (records collected 1959-82). Native Legumes Acacia victoriae Benth (elegant wattle) Daviesia brevifolia Lindley (leafless bitter-pea) Eutaxia microphylla (R. Br.) Black (Eutaxia, mallee bush-pea) Hardenbergia violacea (Schnee.) Stearn (native lilac) Pultenaea tenuifolia R. Br. ex Sims Pultenaea densifolia F. Muell. (dense-leayed bush-pea) Introduced Legumes Cytisus scoparius (L.) Link (English broom) Cytisus proliferus L.f. (tree lucerne) Genista monspessulana {L,) L. Johnson (Montpellict broom) Lupinus cosentinii Guss. (Western Australian blue lupin) Lupinus angustifolius L. (lupin) Lupinus uliginosus Schkyhr (birdfoot trefoil, greater loms) Medicago ciliaris (L.) Krocker Medicago littoralis Rohde ex. Lois (strand medic) Medicago polymorpha L. var, polymorpha (burt medic) Medicago rugosa Desr. (gamma medic) Medicago sativa L. (lucerne, alfalfa) Medicago truncatula Gaertner (barrel medic, snai) medic, caltrop medic) Pisum sativum U.. (field peas) Trifolium frapiferum L. (strawberry clover) Trifolium resupinatum.L, (shaftal clover) Trifolium subterraneum L. (subterranean clover) Vicia dasycarpa Ten. (woolly-pod. vetch) Vicia villosa Roth. (hairy yetch, winter vetch) at least four overlapping generations each year, with peak catches in late September (spring), late November and late December. There is then a generation with a peak in mid to late February and a hint of other small generations in March-May.-OQur trap catches for adults in December 1981 to January 1982 for the Keith area are in accordance with these earlier data by clearly detecting two large summer peaks (Fig, 6), even though they occurred at a slightly later time than the 1959-60 800 600 Ww x 2 wn Oo 400 #2 Lu a = 2 Zz 200 ty) peaks. When related to monthly mean temperatures (Anon. 1987) by iterative calculations, these observations suggest 4 mean generation time of about 210 day degrees above about 13°C. , South Australia has a hot dry summer (Anon, 1987) and as summer progresses, dryland lucerne should gradually becomes unfavourable for E. behrii. Our survey in dryland lucerne in the early sumimer of 1981-82 was planned to quantify this change; and in A 5 °o N D d F MONTH OF YEAR Fig. 4. Light-trap catches of adult Hriella behrii at the Waite Institute campus, Adelaide in 1959-60 accumulated in intervals of 14 days. BIOLOGY OF ETTELLA BEHRI] ZELLER (LEPIDOPTERA PYRALIDAE) Table 2 are given, for each sample (a) the numbers of juvenile, young and mature green pods , and (b) the total oumber of eggs and larvae of E. Sehrii in young and mature green pods (none were found in juvenile pods) The total numbers of E. behrii were obviously greater in mature green pods than in young ones from 61.82 onwards (Table 2}, but the numbers of different Stages were loo small to allow comparisons between the (wo sorts of pods, except for eggs and Ist+2nd instars on 30,xii.81 and 6.1.82. For egys on 30.xii.81 plus 6.1.82, there was a significantly greater number on young pods (51 on 1250 pods) than on mature green pods (31 on 1211 pods) (x? = 3.95, p <0.05). For eggs plus Ist and 2nd imstars, 78 on 1250 young pods were also significantly more frequent than 47 on 121! Mature green pods OC = 7.69; p <(0E). The comparison of these two sets. of data suggest that in the samples for these two days some young larvae were faillng to establish in the mature green pods. Obviously the larger number of all stages of EB. behrii on mature green pods after 6.1.82 was therefore due to larger numbers of older larvae, Because thé two types of pods could be considered as different sampling strata, the total numbers of £. behrii in each sample were Subsequently expressed as means per 1000 pods, corrected for the different. numbers of young and mature pods, Such means of all stages of E. behrii wre also given in Table 2, and separate means for eggs and for instars 1+2, 344 and 5 ate given in Fig, 5. In Table 2, the decline with time of the numbers of both juvenile and young pods illustrates the maturation of the crop, and the data in Table 2 and Fig. 5 indicate @ concomitant decrease in number's of eggs and Ist + 2nd instars of &. bekrii and an increase in numbers of older larvae. To try to make more: sense: of these trends, the. numbers of eggs and of all larvae were plotted against the numbers of adults of FE. beknii caught per week at the light trap at Keith (Fig. 6). ‘The dat Sugges! that the numbers of eggs laid up to and after the first peak of moths were strongly correlated with the numbers of moths each week. However, the 13 40 1 cays B inaiars 1 ST & owsarsse El invetar & Nurpbers per 1000 pede Dec, dan: Fig, 5. Corrected numbers per 1000 pods of Eriella behrii eggs, Ist + 2nd instar, 3rd + 4th instar, and Sth instar larvae in a dryland lucerne crop at ‘Brecon’ (Keith) in relation to calendar time from 1. xii.1981, numbers of eggs on 15.1.82 and 19,1,82 were much lower than expected in relation to the number of moths, which had increased to a second peak (Fiz, 6). Unfortunately, no counts of eggs were made after 19,i,82 because attention was then diverted to the main irrigated sced crop which was in full flower. The cause of the decline in numbers of eggs ii the dryland crop on 15.:.82 and 19,1.82 (Figs 5, 6) is not known, There scemed to be an abundance of suitable pods for oviposition on these two dates (Table 2), but it ts possible that the crop had dried out further so that even the young pods were subtly different from those on earlier dates and were- unsuitable for oviposition. So too there was an unexpected decrease in the numbers of young larvae on 19,1,82 (Fig, 5), possibly again because the pods were not as suitable for their establishment as they had been earlier. The numbers of larvae in irrigated luceme seed crops were of particular interest to us from the view point TaBLe 2. The numbers of juvenile (J) , young (¥) and matwre green (M) pods in each sample; and chi: numbers of ate sages af BE. behrli (Ab) in each category of pads. Numbers of pods Numbers of Z.b, in Mean & >, per Date of sample J Y¥ M Y feds M pods 1000 Y+M pods 09,12,81 276 a4 91 its) 4 156 16.12.81 362 637 207 7 6 Ls.4 22.12.81 267 573 330 6 3 10,0 30.12.81 113 650 S42 30 3 52.9 06.01.82 95 600 669 21 3? 615 15.01.82 Vv {87 463 2 24 40.0) 19.0.2 15 238 1165 i 44 32.1 Tutal 1145 3639 3467 7? 171 id 4. D AUSTIN, T CR, WHITE, D. A. MAELZER & DUG TAYLOR ia ~ 3 ' " a) a} 19 aii so fo wo oe, aan, Fen. Mig. & Number of eegs | A) ond larvae (MM) of Brielle betray per 1000 pods in samples taken at ‘Brecon’ (Keith) in a dryland lucerne crop inthe sunwoer of 1981-82; the number ‘of larvae ([2) similarly sampled in an adjacent irrigated lucerne-crop; and the number of moths (O) caught weekly ina light-trap in the same itrigated crop, in relation to calendar time from |.xii.1981. ol pest management, and the irrigated and unsprayed WLAl8 crop, which we sampled later in the season, had keen managed so that it flowered later than the adjacem dryland crap; and on 19,1.82 it was at about the same stage of development.as the dryland crop had been an 9, xi7,.82, The numbers of Jarvae in the irrigated crop are also giver in Fig. 6, On 15.1.82, they were about the same as those w the dryland crop on 15,1.82 and 19.i.82, but they similarly declined thereafter (Fig. 6) despite un abundance of seemingly suitable young and mature green pods for E. behri/. In particular, there wus no increase if the numbers of larvae afler the second peak of trapped moths on 19,1,82. Untirtunately, eggs were not recorded in these samples from the irrigated tucermne, ner were the larvae categorised into instars, It is not known, therefore. where the mortality occurred that caused the decreasing trend in Jarval numbers, The weather was not obviously unfavourable for moth survival and oviposition, so the mortality is likely to have been al the egg and/or larval stage, Two inain possibilities may be suggested for this decline. Firstly, it is possible that there was a very Jow establishment of young Larvae on the aphid resistant cultivar WL318, as Perrin (1978), for example, found in some varieties of cowpea attacked by Cydia ptychorr (Meyrick). The second possibility is that the mortaliry was duc to predators and pathogens. In particular, the decrease in larval numbers ocurred at the same time as a dramatic Increase tn the numbers of Campylonmmur lida Remer (dimpling bug), which is thought to be i, isn f ‘ ea ng ; = partly predacious, as is C. verbase? (Meyer), a well- known predatod of mites, aphids and psyillids in Canada (McMullen & Jong 1970). Finally, the second peak of moths in mid-January (Fig, 6) may be expected to be smaller than the first one because the area of dryland lucerne is muuch greater than thas of immgaled lucerm: and, as summer progresses, dryland lucerne is oo longer favourable for £. bekrii, So one might therefore expect fewer maths to be caught in a light-trap in February-March, as is partly illustrated in Fig. 6 for Keith. Other crops attacked by £. beh, such as lupins, would similarly cease to produce suitable green pods for young caterpillars in mid January unless they were irrigated. Therefore, in most localities there would be a reduction in numbers of larvae in late January to eurly February and a subsequent reduction im the numbers of moths at the next peak catch of maths, as shown in Fig. 4 for Adelaide in mid to late February, The few moths flying in late February would find even fewer suitable pods for wriposition at the end of the hot dry summer. So the species ts likely to be relatively rare in most localities after February and until the next flush of plant growth an spring, following Winter rains. During this imerval the species could either enter an obligate diapause or it could continue to breed in reduced numbers (on volunteer lucerne}, The light-trap data for Adelaide (Fig. 4) strongly support the latter alternative, because a few moths were caught in late April w early May. The lack of a diapause was also dentonstrated directly by ane of us (DGT) by successfully incubating to adult emergence, larvae collected! over the winter of 1959, Parasiteids, predators and pathogens The parasitoid species which bave been recorded trom &. beh are given in Table 3 along wath the silage from which they emerge, whether they are solitary or gregarious, and their status ay either primary or hyperparasitoids, The ichneumonid Femefucha oprnea Kerrick and the braconid Jconella alfalfae (Nixon) have been recorded as causing between 2% and 1% mortality 10 field populations in the Keith area in 1939-60 (DGT unpubl.). while the other species listod have heen only rarely encountered. Yellow pan-traps placed in the field at ‘Brecon’ in carly December 1982 first caught 7. cychea in early February. The numbers of this parasitoid appeared to mcrease through February aod March, but there were too few collected per trap to quantify diflerences between samples. Phanerotome sp, was also caught in late February but in very low numbers, Of 41 larvae of &. behirii taken from the field in late January to screen for parnsucids, 25 pupated, and three were parasitised (lwo by T. cycrea and one by Penerofoma 3p_j. T. cycred oviposits in) early stage larvae after the latter emerge from the pod and begin BIOLOGY OF ETIELLA BEHRIT ZELLER (LEPIDOPTERA) PYRALIDAE) 1S to build a webbed retreat, while Faneroiama sp. oViposits into the eggs of its host, Both species then complete their development after the final host instar has dropped 1o the soil and pupated. During the 1981-82 study we did not record any other parasitoid species, even though J. alfalfae has been recovered in moderate numbers in the past (DGT unpubl,). We did not directly observe any predation upon eggs und larvae of E. behrii, but several general predators wete. seen in lucerne crops, and they may feed on &, behrii. The predators included C. livida (see abave) and spiders, particularly of the families Oxyopidae, Araneidae, Lycosidue and Thomisidae. Of these species C. livida was by far the most common in irrigated crops during January-February 1982. Sixteen of the 41 screened larvae died before they pupated. After death they became soft, darker in colour and developed faint spots undet the integument. These larvae were later examined and found to contain large numbers of microsporidian spores and some bacteria (Bacillus sp. — possibly 8. thuringiensis Berlinger) in the haemolymph. Nothing is known of the mortality caused by these pathogens in the field, but B thuringiensis at least is known to be highly toxic to E, behrit \arvae- in the laboratory, having an LC,,, value of 311 spores/mm?, a figure 20 times lower chan that for Helicoverpa armigera (Hiibner) (Cooper 1983), Discussion Irrigated lucerne crops flower and form pods later than dryland lucerne and volunteer plants which grow around irrigated crops, along road-sides and in scrubland. So the number of moths available to attack irrigated lucerne crops each year should depend on the namber and suitability of pods available to.the earlier generations of £. behrii in that season, and on some Tunction of weather. However, the very low numbers of larvae in the irrigated crop south of Keith in Jamiary- February 1982, despite the presence of a large number of adult moths in the vicinity (Fig. 6) and suitable oviposition sites, suggests that some variable or variables, other than the number of moths, may be a key factor in the amount of crop damage that occurs from year to year, This variable might be predation or parasitism on eggs and young larvae, the effect of pathogens, or some property of the plant which prevents young larvae establishing and surviving in pods, either of certain cultivars or in certain years. Such a variable might also account for the unpredictability of attack of lucerne by &. behrii, In most years damage is so slight as to go unnoticed, but occasionally there is avery heavy infestation in one season, Such outbreak seasons appear from the available records to be infrequent, but when they do occur, outbreaks are widespread. To date, further research aimed at examining the factors responsible for heavy infestations of E. behrii in seed lucerne crops in South Australia, has been hindered by the infrequency of such events. In recent ycars there have been only low or moderate numbers of this pyralid pest reported in seed crops in the Keith area, possibly because of the more widespread and effective use of pesticides since the early 1980's. TABLE 3. Furasitolds kriown fo attack various stages of Etella behrii in South Australia (records collected 1959-82), Parasitnid Stage from which Primary (P) or hyperparasitoid (H) species parasitoid emerges solitary (S) or gregarious (G) Tchneumonidae Temelucha cycnea Kerrich pupa P;S Enynis sp. P;S Braconidae Iconeila alfalfae (Nixen) Sth instar larva P;S Phanerctoma sp. pupa PS Bassas sp. | Sth instar larve Ps Bassus 4p, 2 Sth instar larva PS Bracon sp. thaturé larva PS Elasmidae Elasmus: sp- pupa H:S or G Eurytomidac Eurytoma sp. pupa Ps Tachinidae ? Siphonini (gen. & sp. indet.) mature larva PSE tis) A. D. AUSTIN, T. C. R. WHITE, D. A, MAELZER & D G. TAYLOR Acknowledgments We wish to thank Mr A. Campbell for access to his property, ‘Brecon, Dr David Cooper for assaying pathogens of E&tiella, Dr Jenny Gardner for identification of native plants, and Drs Brian Cantrell and Mike Sharkey for identification of dipteran and hymenopteran parasitoids, respectively. We gratefully acknowledge the Commonwealth Special Research Grants Scheme and the lucerne seed growers through the Seed Section of the United Farmers and Stock Owners of South Australia Inc. for funding this project. We also thank Paul Dangerfield for providing technical assistance in the preparation of the manuscript. References Aput-Nasr, 8. & AWADALLA, A.M. (1957) External morphology and biology of the bean pod-borer, Friella zinckenella Treit. (Lepidoptera: Pyralidae). Bull. Sac. ent. Egypte Al, 591-620, AANon, (1987) Bicnnial Report of the Waite Agricultural Research Institute, The University of Adelaide, 1986-1987 (University of Adelaide, Adelai ie). Austin, A.D., DurHAM, M. & Bamey, P. (1986) Etiella Moth. Department of ‘Agriculture. South Australia, Fact Sheet, FS 51/85, 4pp. Common, £.F,B. (1990) "Moths of Australia.” (Melbourne University Press, Melbourne). Coorer, DJ. (1983) The susceptibility of Etiella behrii to Bacillus thuringiensis. J. Aust, ent. Soc. 22, 93-95. DurHam, M.S. (1984) Commercial monitoring of insect pests and beneficials of seed lucerne in south east South Australia. pp. 121-127 Jn Bailey, P. & Swincer D. (Eds) “Proceedings of the 4th Australian Applied Entomological Research Corference™ (South Australian Government Printer, Adelaide), Harpstack, A.W., Henpricks, D,E,, Lopez, J.D., STADELBACHER, E.A., Priuups,J.R, & Wirz, J.A, (1979) Adult sampling. pp. 105-131. In Sterling, W.L. (Ed.) “Economic Threshholds and ling of Heliothis Species on Cotton, Com, Soybean and Other Plants.” Southern Co- operative Services’ Bulletin No, 231. (Texas A & M University, College Station). Maenzer, D.A., Prynock; D,E., BAILEY, P-T, & MADGE, P.E. (1982a) Pollination and pest management of lucerne seed crops in South. Australia. pp. 121-141. /n Cameron, P.J., Wearing, C.H., & Kain, W.M. (Eds) “Proceedings. of the Australian Workshop on Development and Implementation of I.P.M.” (Government Printer, Auckland). Mapce, P.E, & Bailey, PT. (1982b) Pollination and arthropod pest management of lucerne seed crops in South Australia. pp. 385-392. J Lee, K.E. (Ed.) “Proceedings of the 3rd Australasian Conference on Grassland. Invertebrate Ecology.” (South Australian Government Printer, Adelaide), McMuLLen, R.D. & Jona, C (1970) The biology and influence of pesticides on Campylomma verbasci (Heteruptera: Miridac), Can. Ent. 102, 1390-1394. Perrin, R.M. (1978) Varietal differences in the susceptibility of cowpea to larvae of the seed moth, Cydia ptychora aeytick) (Lepidoptera; Tortricidae), Bufl. ent, Res. 68, Snorey, H.H. & HALE, D.L. (1965). Mass rearing of the larvae of nine noctuid spp. on a simple artificial medium. J. econ. Ent, 58, 522-524. SincH, H, & DuooriA, M,8, (1971) Bionomics of the pea ped Brees Fiiella zinckenella (Treitschke). Ind. J. Ent. 33, Stons, M. (1965). Biology and control of lima bean pod- purer in southern California. %ch. Bull. U.S. D.A. 1321, TuRNER, JW. (1980) Insect pests of peanuts in eastern Queensland. Qld Agric. J. 106, 172-176. Wuatiey, P.E.S, (1973) The genus. Etiella Zeller (Lepidoptera: Pyralidae); a zoogeographic and taxonomic study. Bull. Br. Mus. (Nat. Hist.) Ent. 28, 1-21. INCABATES HAMMER AND SETINCABATES GEN. NOV. (ACARIDA: CRYPTOSTIGMATA: HAPLOZETIDAE) FROM SOUTH AUSTRALIAN SOILS BY D, C, LEE* Summary Incubates Hammer is commented on and an allied genus, Setincabates gen. nov., established. Three new species are described from South Australian soils: . macronudus sp. nov., I. punctatus sp. nov. and S. hypersetosus sp. nov. (type). . angustus Hammer, previously known from New Zealand, 1s newly recorded from Australia and 1. medius Hammer is newly regarded as its junior synonym. A key is given for the adults of these four species. This is the first record of Incubates from Australia. An African species previously grouped in /ncabates is newly combined as Muliercula longisaccula (Mahunka). KEY WORDS: Incabates angustus Hammer, Incabates macronudus sp. nov., Incabates punctatus sp. nov., Setincabates hypersetosus gen. noy., sp. nov., Haplozetidae, Acarida, soils, South Australia. Treasuerions of the Rawal Society of S. Aiesr., (1993), IIW(23, 77-85, INCABATES HAMMER AND SETINCABATES GEN. NOV. (ACARIDA: CRYPIOSTIGMATA: HAPLOZETIDAE) FROM SOUTH AUSTRALIAN SOILS by D. C. LaB* Summary Lee, D. C, (1993) Incabates and Setincabates gen. mov. (Acurida: Cryptostigmata: Haplozetidae) from South Australian soils, Trans. R. Sac. S. Aust. 117(2), 77-85, 4 June, 1993. Incoubales Haxamer is commented on and an allied genus, Setincabates yen. nov., established. Three new species are described from South Australian soils; /. macronudus sp. nov,, 1. punctatus sp, nov. and S. Aypersetosus sp. nov. (type). I. angustus Hammer, previously known from New Zealand, is newly recorded from Australia and J. medius Hammer is newly regarded as its junior synonym. A key is given for the adults of these four species. This is the first record of Incabaies from Avstralia. An Affican species previously grouped in Incabates is newly combined as Muliercula longisaccula (Mahunka). Key Worns: Incabates angustus Hammer, Incabates macronudus sp. nov., Incabates punctatus sp. nov,, Setincabates hypersetosws gon. nov... sp. nov., Haplozetidae, Acarida, soils, South Australia. Intraduction The genus /ncabates Hammer, 1961 and a similar undescribed genus are examined as part of an ongoing study of sarcoptiform mites sampled from nine florally diverse South Australian sites, and for which Lee (1987) provided an introduction to the relevant work on the advanced oribate mites, The relevant mites wre from soil and plant litter or mass under either savannah woodJand, sclerophyll forest, mallee or coastal vegetation at only four of the sites. Incabutes is grouped in Haplozetidae Grandjean, 1936, which has been discussed by Lee & Shepherd (1990) in considering Magnobaies Hamamer, 1967. Relationships between Jncabares, the new genus and some other haplozetid genera are discussed. J. angiastate Hammer, 1967 is newly recorded from Australia, and two new species of Incabates and the new genus together with one new species are described. Material and Methods New material examined here, collected by the author, is deposited mostly in. the South Australian Museum (SAMA), but also in the. Natural History Museum, London (BMNH), the Field Museum, Chicago (FMNH) and the New Zealand Arthropod Collection, Lam! Care Research, Auckland (NZAC), whilst previously described material is deposited in te Zoological Museum, Copenhagen (ZMC). The morphological notional system follows Lee (1987), the somal chaetotaxy of which is.summarised in Figs * South Australian Museum, North Terrace. Adelaide, S. Aust_ 5000. 2 and 3, with the total setae present in cach file (e.g. 6Z) indicated by number Coming first, whilst a particular seta (e.g. Z6) would have the number last. The venter and legs have been described only for Incabates punctatus because of their uniformity within the genus, The abbreviations for zoogeographical regions follow Lee (1970, Fig. 427), Descriptions of eggs are based on those still within the female soma, All material was examined using a Nomarski interference contrast device. All measurements ate in micrometres (zm) and were made using un eyepiece micrometer at x250 magnification. Key to Australian Incabates and Setincabates species (adults) |. Thirteen pairs of hysteronotal seta. Hysteronotal foramen FS sacculate without narrow duct oo. eee ety seed, s2brneadinedbeas Setincabates hrypersetomis sp. nov, Ten pairs of hysteronotal seta or ulveoli. Hysteronotal foramen F3 sacculate with narrow duct ...., éncabates. Hysteronotal setae represented only by alyeoli. Tutorium ts absent. ......2..... saves pelle e's T, macronudus ap. nov, At Jeast one pair of hysteronotal setae ae Tutorium PRONE. Nyy ereesdthgtarateas wheartbeaeedtyaices 3, 3. Six posterior small hysteronctal fetal pat present (JS, J6; ZS, 26; $5, §6). Anterior soma punctate ose. Wirestewassg tig veuese T. punctatus sp. Nov. One posterior small hysteronotal seta] pair present (/6). Anterior soma not punctate -..... I, angustus Hammer. Systematics Incabates Hammer Incabates Hammer, 1961, p. 108 (type species by monotypy: Incabates nudus Hammer, 1961); Coeter, 1968, p. 25; Balogh ot ai 1984, p, 274: Loxton, 1985, p. 67; Corpuz-Raros, 1980; 174. 78 DE Definition: Haplozetidae. Hysteronmtem with |} pairs (2/, 62, 25) of short setze, microsetac or alveoli without thair setae. Hysteronotal forarnina with sarcule bag-like, either lapermg gradually to pore or with short narrow duct near pore, Dorsosejugal furrow entire. Translamella and prelamelia absent, costate or lineate tutorium (between setae jl-z2) sometimes present. Rostral eta {jl} directly in front of tamellar sera (21). Picromorph movable, with clear weakly sclerotized basal Hne. Discidium triangulate. Tibia | with large solenidium (s02) on tubercle. Tibia I without disodorsal spur. Genu I and 0 with two setae (vy ubsent), Trochanter [Y usually with distodorsal process exunded over femur IV (exception! [ncabares eugustus), distodorsal crown with only anterior angulate lobe, broad {lange extending along entire venter of caput, Pretarsus with three claws, slimmer fateral claws with pointed tip, General Morphology of Austratian species; Samal length range for adults: 239-445 (for all species 239-496). Sowal chaetotaxy: 2/, 22, ls; U, 62, 23; 32, WT, Bil, 31% AlZg, 18g. 2 Za, 3Sa. Leg chaetotaxy {solenidia tn parentheses): |— 1, 5, 2¢1), 442), 20(2); D — 1, $, 2(1), 40), 16(2); I — 2, 3, 10), 300), 15; TV — 1,2, 2, 3¢]), 12, Alveolrof somal setae bonnded by refracule ring, canal short and tapering without internal refractile ring. Pteromorph may lic close to pleural surface or be lifted away from it, Sub-bothridial flange present, Apodemes.1, ll, ventrosejugal and Il present, Ventrosejugal gap not wider than genital orifice, Subpedal and circumpedal ridges merged into single continous line. Slitdike pore Saf nearly longitudinal (less than 45° from longitudinal axis). Proximovectral spur on ferur f without caput collar. Conspicuous yentral flanges on femora I, Hi, and TV, wn femur Il margin curved, not angulate. Distrituaion: Peru (NTe}, Japan (Pc), Philippines (Om), Australia (Aa), New Zealand (An), Fiji (Ap), All species from outside South Australia are from moist localities, often in moss, cither in high altitude grassland or in forest or mangroves, and are sometimes arboreal. In South Australia ali three species. were collected from the svlerophyl! forest site, which bas the highest rainfall (annual mean in the range of 1150-1200 mm, moxily in winter}, but Jnowbates punctatus sp. nov, was also collected from the two mallee sites Which are dry (mean annual rainfall in the range of 350-500 man). Remarks: Incabates was considered allied to Protaribates Berlese, 1908 when it was established. More recently it has been allied to Scheloribates Berlese, 1908 (Corpuz-Raros 1980; Laxion 1985), which it is similar in having a tangulate discidium, hysteronotal sacculate foramina that gradually taper te 4 pore, and, for some species, in having no tutorium fin species wilh a cutarium, it only cosinte). In comtrast, Incabates tins a Gerived character state, a hinged rather jEE than a fixed pleramorpl as in scheloribatid genera, on the basis of which, Cnetter (1968) suggested that Incabates shwuld be transferred to the Haplovetidae Grandjean, 1936. Because /nonbates also has a derived trochanter LV (the distoventral crown with only an amterior lobe that extends along entire vemer af caput), and if there is a lateral proteronotal ndge it is a tulariam rather than a prelamelia ridge, this grouping in lhe Haplozetidae is followed here: ly current classifications of the Onpodoidea, the Haplozetidae are wit closely allied to the Protonbatidue Balogh & Bslogh, 1984, because of the importance given lo the former family haying derived sacculare hysteronutal foramina, tather than jultiporose foramina: out both Iacabates anG Protoribates have derived hinged pteromerphs. The significance of particular character states and so the accepted relationships amongst oripodoid taza, may well change considerably: In this paper it is considered that, since Incabates shares some character statés with mest genera of the more primitive Scheloribaudae Grandjean. 1933, characler states thal are derived in many members of ihe Haplozetidae (the discidium is réctangulate, the tutorium ia canspicuously laminate and hysteronotal sacculate foramina often have narrow ducts leading to their pores), it is more primitive than Magnobaies Hanmer, 967 and the genera closely allied to it (Lee & Shephard 1990). in distinguishing Incabates from other genera, the following scheloribatid genera must be considered: Muliercula Coetzer, 1968, Nennerlia Coetzer, 1968 and Styloribates Jacot, 1934. These three genera can be regarded as derived within the Schelortharidiae because: of character states of the proteronotum and trochanter IV. In general, members of the Scheloribatidae fold feg 1 against the soma so that the tarsus pomls downwards behind seta 1, lying behind the prelamella tidgs, also trochanter [V is simple, with the crown contined to the distal end, In contrast, haplozetids and the three scheloribatid genera listal above, have a tutorium of subtutorium. rather than a complete prelamells and, when the leg is folded, tarsus I can paint forward between setae zl-jh, whilst on truschanmer IV, the crown extends as a broad flange along the enture vemter of the caput. Therefore, these three scheloribatid genera are wnly distinguished from the faplozericd Jncabates by laving fixed pteromorplis, which ts sometimes a difficy]| character state to assess. Furiber studies may require that these four genera dre cither grouped into a new family or that the Scheloribatidae and Haplozetidae are merged. The naining of a species, that is very similar to Ihe type species of Mulierculz, as Uncabates longisaccutus Westrates this confusion. It is here combined as Muldiercida longisaccula (Matiunka, 1984) comb, now., despite its lack of a tutocium and the presence of 2 partial prelamellar ‘This is partly because these INCABATES AND SETINCABATES, HAPLOZETID MITES 7 character states are variable in Jncalenes, aud therefore the presence of a tutorium may not he diagnostic of Muliercula, but it is mainly because the pteromerph is only partially delineated from the hysteronotnm by a Clear furrow as described for Muliercula, where it is regarded as fixed, although ambiguously described by Coelzer (1968) as “pteromorphae immovably hinged”. Because of the similarity between Incahares and Muliercula, despite their grouping in: separate families, only the more extensive wing-like expansions of the lamellae of M. fongisaccula and its occurrence in Afriva support the pew grouping. Tn comparison te other haplozelid genera such as Haplozeres Willmnann, 935 and Magnabates Hammer, 1967, Incabates ts regarded as prinufive in having a triangulate disctdium, as on oribatulid and scheloribatid adults, Similarly it has only two setae on gen I apd the alyeolar canal of te somal setae is short and simple, facabates includes seven species: Lb angustus Hammer, 1967 (= 7. meafus Hammer, 1971, syn, nov.) (rom Australia (Aa), New Zealand [An) and Fiji (Ap); {. mactrenudus sp. nov. fron Australia (Aa); 1 major Aoki, 1970 from Japan (Pe): 2 mudus Hammer, 1961 from Peru (NTc) and Philippines (Om); |, pahabaeus Corpuz-Raros, 1980, from Phillppines (Om); / punctatus sp. noy. from Australia (Aa); J. strianas Cormiz-Raros, 1980, from Philippines (Ons). Incabates angustus Hammer fncabates angusnss Hammet, 1967, p. 43, 44, Fig. 57, S72. facutates medias Hanuner, (971, p. 42, Fig, 49. 49a, syn, noe, Type material examined: J, angustus, lectorype 9 {labelled “type” us vial of alcohol, three specimens recorded in Orginal description) examined (ZMC}. liverworts and sewall ferns, Gead tree trunk, native forest, Waitakere National Park. North Island, New Zealand, M. Hammer, 1962. f. madius, lectotype @ (labelled type in vial of alcohol, two specimens recorded |n original description) examined (ZMC), withered leaves, river bank, above mangroves, Corolevu, Vita Levu, Fiji Islands, M. Hamner, 962- Female: Soma oval, brown. Idiosomal length (original description from New Zealand: ‘about 0,35 mmm, from Fiji; ‘about 0305 mm), 335 (n = 1, New Zealand) of 318 (n = 25, Sclerophyll forest, 306-332), Leg lengths (femur-tarsus, Sclemphy!l forest. for 320}: 1-142, 0-127, 1-108, I'V-137. Tibial maximum heights (for 320): 1-22, (1-17, 1-14, [V-14, Proteronotum with mediuny width fostruin, Integument smocih, ‘Tuionum. present, sometimes indistinct level] with lameflar seta (21). Sensory seta (z2) with globular caput longer than exposed stalk. Sub- hothridial Mange inconspicuogs, Posterior margin of bothridium raised to form tooth-like spur. Imerlamellar seta (72) longer than distance j2 ~ zl, Hysteronotum with only one pair (/6) of noticeable small setze, nine pairs (J5, 21-6, S5 and 6) of alveoli, possibly with small Mucrosetae, Alveolus $5 posterior m ZS (South Australian specimens) or anterior to ZS (New Zealand Specimens). Foramen F3 conspicuously lazer and sometimes sacculus bilobed (South Australian specimens) or shghtly larger, mot bilobed (New Zealand specimens), Idjosternal setae fine and short, conite seta 3, J and J¥3 (not Ulustrated as present by Hammer, 1967: Fig. 37a) longest. Integument smooth excepi for indistinct reticulations around ventrosejugal apodeme and apodeme TIT. Discidium height less than 05x height of pedotectum MT, coxite seta JV3 néar discidium base, Eggs subellipsoidal, exochorion smooth, mean size 151 x 72 (n = 7), length 48% of somal length, eges per female — | (n = 3), or 2 (mn = 2). Legs short (median fermur-tmarsus length: 40% of somal lengeh) with stout girth (mean maximum tibial height 50% of mean length). Trochanter ['V anterior margin parallels posterior margin of femur fY¥ caput, angulate distodorsally but without process extended over femur I'V. Male: Similar to ferale but idigsoma shorter, mean length, 293 (7, Sclerophyll forest, 283-303) or 319 G1, Sayatinslt woodland) Referred material: 12 females (SAMA NIQ9iL — NI99112), seven males (SAMA NI99113 — NI99119), plant litter, sparse moss and sandy soil, under sclerophyllous shrub arnongst messmate stringybark (fucalyptus obliqua), dry scleropbyll forest, near summit of Mt Lofty (34°59°S, 138°45'B), Cleland Conservation Park, 9.v.J974, One male (SAMA N199120), grass, moss, plant litter and loamy soil, under manna gum tees (Eucalyptus vininalis), savannah woodland, Chambers Gully (34°S8’S, 138°41’E), Cleland Conservation Park, 12.¥i, 1974. Remarks; Incabates angustis was ane of four species that have only a single pair of bysteranotal seta (/6), but one of the other three species, | medius from Fiji, is here synonymised with it. The other two species, I. striatus with a stviated intégument and the stouter 1, nudus with a much broader lamella, are easily distinguishable as separate species, The slim /, angustus and f, medius are very similar with the new material from South Australia having intermediate character states, /. angustus is 350 wim long, has hysteromotal seta 96 anterior to $$ and the imerlamellar Seta (f2) & about 0.75% the length of the lamellar seta zi). £ medius is 305 xm long, has hysteronotal seta 56 level with $5 and the interlamellar seta is about O.5x the length of the lamellar seta. The specimens from South Australis are similar in size ta J, medins, have seta 96 posienor to 55, und the relative sizes of the interlamellar and lamellar sete are as J. angusuce. 80 D, C. LEE Fig. 1. Incabates macronudus sp. nov., female soma, notum. 1004um INCABATES AND SETINCABATES, HAPLOZETID MITES BL These differences are here regarded as intraspecific variations within a species distributed across Australia, New Zealand and Fiji, and comparable with differences within J nudus from Peru and. the Philippines (Corpus-Raros 1980). Incabates macronudus sp. nov. Fpe material Holotype female (SAMA NI99121), plant litter, Sparse moss and sandy soil, under sclerophyllous shrub amongst messmate stringybark (Eucalyptus obliqua), dry sclerophyll forest, neat summit of Mt Lofty (34°59’S, 138°45’E), Cleland Conservation Park, 9yv.1974. Female: Soma oblong, brown. Idiosomal length 446 (n = 1). Leg lengths (femur-tarsus for 446): 1-178, II-163, II-142, TV-166. Tibial maximum heights (for 446): 1-22, I-17, I-14, IV-l4. Proteronotum with narrow rostrum. Integument smooth, Sublamella obscured in dorsal aspect by laminar lamella, Tutorium absent. Sensory seta (z2) 3 100,.um Figs. 2, 3. Incabates punctatus sp. nov., female soma. 2, notum; 3, idiosternum Fig. 4. Incabates punciatus setae illustrated (vy = ventral). clavate (caput may appear globular if viewed end on), caput subequal in length to exposed stalk. Sub- bothridial flange conspicuous. Posterior margin of bothridium raised to form tooth-like spur. Interlamellar seta (j2) longer than distance j2-zi. Hysteronotyum with no obvious setae. Alveoli form clear refractile rings, mictoseta recognised in Z2, may be present elsewhere. Extra alveolus between Z2 and Z3 on right side only. Foramina with conspicuously refractile saccule, F3 oval and larger than other saccules. Idiosternal sctae fine and short, setae /3 and Sal the longest, setae 113 and I'V3 not located. Integument smooth except for indistinct reticulations around ventrosejugal apodeme and apodeme IIT. Genital seta JZg2 midway between JZg] and JZg3, rather than closer to JZg1, Discidium height less than 0.5x height of pedotectum Il, coxite seta /V3 not located. Egg . noy., female right legs I-IV, femur-pretarsus, also trochanter IV, posterior aspect. Only two subellipsoidal, exochorion smooth, mean size 166 x 70 (n = 4), length 42% of somal length, four eggs in single female. Legs short (median femur-tarsus length: 36% of somal length) with stout (mean maximum tibial height 39% of mean length). Trochanter IV with blunt process on angulate distodorsal margin which extends over femur LY. Remarks: The name macronudus is from the Greek for ‘large’ and the Latin for ‘naked’ and refers to its body size and the absence of recognisable setae on the hysteronotum. I. macronudus is the only species known in Incabates with no easily observable hysteronotal seta (I. nudus has one pair) and with J. major it has a somal length greater than 440 um. Because there is only one specimen, it was not dissected, which may explain why some coxite setae (7/3 and IV3) have not been located. INCABATES AND SETINCARATES, HAPLOZETID MITES &3 Incabates punctatus sp. nav. Fype material Types: Holotype female (SAMA N1987645), soil, plant litter ami sparse moss under ridge-fruited mallee (Eucalyptus éncrassata) clumps amongst broombush shrubs (Mefalewcu wneinata), Fetries-McDonald Conservation Park (35°1S’S, 139°O9'E), 20vi, 1974, Paraty pes, 45 females (SAMA NIG87646 — NIGRTES4, NI99122 — N199142, five — BMNH, fve — FMNH, five — NZAC) and 55 males (SAMA N1987655 — NIS87671, NI99I43 — NI99165, five — BMNH, five — FMNH, five — NZAC}), as holotype. Female: Sama oval, brown, Idiosomal lengta 259 (n = 25, Mallee broombush, 249-267), 257 (n = 2, Mallee heath, 256, 258) or 260 (n = 3, Sclerophyll forest, 252-267), Leg lengths (ferwir-tarsus, Mallee broombush, fioy 262): 1-113, 1-98, 111-82, PY-105, Tiblal maximum heights (for 262): 1-16.5, 1-13, M-11.5, TV-13, Proteronotum with medium breadth rostrum. Integument punctate anterior to sela 72, Sublamella mainly obscured in dorsal aspect by laminar lamella, Costale tutortum present. Sensory seta (22) clavate, caput subequal in length to exposed stalk. Sub- bothridial flange conspicuous. Posterior margin of bothridium raised to form small single or double tooth- like sour, Interlamellar seta (j2) shorter than distance 2-21. Hysteronorum without obvious setae anteriorly (Zl, Z2, Z3, ZA) but microsetae recognised for ZI and Z2, whilst posterior small setae (/5, J6, 25, Z6, 55, S56) present, Integuiient punctate anterior to fteromorplis. Foramina with conspicuously refractile saccule, £3 spherical and Larger than other saccules. Idiosterna! setac fine and short, variable relative lengths, but JA, (7fi, JZg3 always shortest. lotegument with indistinet reticulate sculpturing and punctate in coxistemnal region. Genilal seta JZg2 closer to JZyl than JZ23. Thscidom height more than 05x height of pedotectum I, coxite seta /¥3 level with, and close to apex, Egg subellipscidal, exochorion smooth, mean size 132 x f4(n = 13), length 49% of somal length, usually one egg per femats (n = 14), two egys in single fernale- Legs short (medion ferur-tarsus length: 38% of somal length) with very stout girth {mean maximum tibial height 51% of mean Jength). Trochanter ['Y with sharp process on angulale distodorsal margin which extends over femur IV, (dale: Sienilar to female but idiosomal sherter, mean length, 239 (25, Mallee broambush, 226-252), 249 (5, Mallee feath, 238-250), 251 (3, Coastal, 250-253) or 247 (3, Sclerophyll forest, 244-252), Referred material. Three females (SAMA NIOS7672, Ny99174, NIG9I7S) and five males (SAMA NIS9I76 — Ni99180), sand, plant litter, wnder Banksia shrubs (Banksig omata), amongst other sclerophyllous shrubs and sparse brown stringybark mallee (fucalypis bzxteri), Tainboore Homestead (35%57'S, 140°29'F), ar Mt Rescue Conservation Park, 4,vii.1974. Three males (SAMA NI99T71 — NI99173), soil, plant litter and sparse grass under coastal wattle (deacie.rophorae), Piccaninnie Ponds Conservation Park (39°03'S, 140°S7'B}, 3.vii.1974. Three females (SAMA NI99166 — NIY9168), two males (SAMA NI99169, NI99170), plant litter, sparse moss and sandy soi), under sclerophyllows shrub amongst messmate stringybark (Eucalypnes obliqua), dry sclerophyll forest, near sunnmit of Mt Lofty (34°S9'S, 138°45'E), Cleland Conservation Pazk, 9yv, 1974. Remarks: The name punctanes is From the Latin for ‘dotted’ and refers to what may be small pits on anserior Parts of the soma, which wre more extensive than in olher species, Also /. punctatus is unique within the genus m lacking four anterior purrs of frysteronatal setae (microsetae may be present), whilst the posterior six pairs have small setae, that are relatively long for Incabates. J. pahabacus Corpus-Raros, 1980 is similar but without such 4 clear difference in size between the two proups of setae. Also, on /. pahabeeus, the interlamellar setae are much longer and the hysterosotal alit-shaped pore A/S is short and not as long as seta J6, Genus Setincabates gen, mi, Type-species: Serincabates hypersetesus sp. now, Definition; Haplozetidae. Hysteronotum whh 3 pairs (4J, 6Z, 35) of miedium-sized setose setae, Hysteronotal foramina with saccule bap-like, gradually tapering (0 pore, Dorsosejugal furrow entire. Translamella, prelamella and tutorium ahserni_ Rostral seta {fl} directly in front of Jamellar seta (z)). Pteramorph movable, with clear weakly sclerotized basal Ine. Discidium triangulate, Tibia I with large solenidium {se2) on tubercle. Tibia Uf withour distodorsal spur. Genu | and Tl with peo setae {vy absent), Trochanter IV with distodorsal process extended over femur IV, distadorsal crown with only anterior angulale lohe, broad flange extending along entire venter of capul. Pretarsus with three claws, slinumer lateral claws with pointed tip. Distribution; Australia (Aa). Only recosd from South Australia, ai site which amongst those sampled has the highest rainfall, Remarks: The name Serincabates has the prefix Ser, an abbreviation of the Latin seta for ‘bristle or hair’, und refers to the presence of more hysteronatal setae on the single included species than on members of the samilar /ncabates and other baplozetine genera with short or medium-sized setae. The haplozetine Flagellobates Mahunka, 1973 with long hysteronotal seme has 14 ppits of hysteronmal setae. Some other subfamilies included an Haplozetidae by Balogh & Balogh (1984) have similarly numerous hysteronotal 84 D.C. LEE setae, Members of the Peloribatinae, all of which have long hysteronotal setae, have either 14 pairs or, in Acutozetes Balogh, 1970, 13 pairs of hystetonotal setae (possessing six not five setae in file J). The only other haplozetid species with more than 10 pairs of hysteronotal setae are members of the Pilobates Balogh, 1960 (Pilobatinae) with 14 paits and six pairs of genital setac and a straight, transverse complete ventrosejugal apodeme and one species of Rustrozeres Sellnick, 1925 (Rostrozetinae) with 74 pairs and a short fissure-like pore 43 similar in length to the hysteronotal foramina. The character states of Setincabates recognised here as distinguishing it from Mcabetes are probably primitive. Setincabates hypersetosus sp. Nov, Type material Types: Holotype female (SAMA NI99181), plant litter, sparse moss and sandy soil, under sclerophyllous shrub amongst messmate stringybark (Eucalyptus obliqua), dry sclerophyll forest, near summit of Mt Lofty (34°59'S, 138°45'E), Cleland Conservation Park, 9y.1974. Paratypes, 21 females (SAM NI99182 — Ni99196, two — BMNH, two — FMNH, two — NZAC) and 21 males (SAM NI99197 — NI99IIIL, two BMNH, two — FMNH, two — NZAC), as holotype. Female: Soma oval, brown. Idiosomal length 264 (22, 254-275). Leg lengths (femur-tarsus for 267); 1-22, 11-104, 111-86, [V-110. Tibial maximum heights (for 267): 1-19, 11-13, 0-13, [V-14. Proteronotum with broad rostrum. Integument smooth except for punctuations on gnathosternum between postoral setae. Sublamella mainly obscured in dorsal aspect by laminar lamella. Tutortum not present, although short ridge present around base of rostral setae (jl). Sensory seta (22) clavate. with long ellipsoidal caput more than twice length of exposed stalk, Sub-bothridial flange conspicuous. Posterior margin of bothridium rounded without tooth-like spur. Imerlamellar seta (j2) about x2 distance j2-zl. Hysteronotum with I4 pairs of small setae (5/, 62, 35), Foramina with small oval refractile saccule without parallel sided duct, all similar in size. Idiosternal setae fine and short, peripheral setae longer, 72 and JZa3 longest. Integument with indistinct reticulate sculpturing in coxistemnal region. Genital seta JZgi closer to JZgl than JZg3. Discidium height less than ).5x height of pedotectum I, coxite seta /V3 near, but posterior to apex. Egg subellipsoidal, exochorion smooth, mean size 132 x 65 (n = 4), length 49% of somal length, one egg per female. Legs short (median femur-tarsus length: 40% of somal length) with very short girth (mean maximum tibial height $4% of mean length). Trochanter I'V with short sharp process on angulate distodorsal margin which just extends over femur IV. Male: Similar to female: but idiosoma shorter, mean length, 239 (21, 244-285). Remarks; The name Aypersetosus is the Latin for ‘bristly’ or ‘setose’ with the Greek prefix for ‘beyond’ or ‘over’ and refers to the unusually extensive hysteronotal setation compared with members of genera similar to Sefincebates. 10940 Fig, 5. Setincabates kypersetosus gen, & sp. nov., female soma, notum, INCABATES AND. SETINCABATES, HAPLOZETID MITES 8S Acknowledgements I am indebted to Ms C. M. Birchby for the illustrations and to the Australian Biological Resources Study for funding her salary. Thanks are also due to Dr Henrick Enghoff (Zoological Museum, Copenhagen) for making types available, and Mrs D, Lowery for typing the manuscript. References Aokt, J. (1970) Description of Oribatids Mites collected by smoking of trees with insecticides. I. Mt Jshizuchi and Mt Odaigatiara. Bull. nam, Sci. Mus,, Tokyo 13, 585-602. BALoGu, J. (1960) Descriptions complementaires d’Oribates (Acari) d’Angola et du Congo Belge. Publ. cult. Co. Diam. Ang. Lishoa 51, 89-105. (1970) New oribatids (Acari). from New Guinea. II Acta zool. hung. 16, 291-344, —____ & BaLoau, P, (1984) Review of the Oribatuluidea Thor, 1929 (Acari; Oribatei). Acta zool. hung. 30, 257-313. ee FT: A, (1908) Elenco di generi e specie nouvi. Redia 15, Coerzer, A. (1968) New Oribatulidae Thor, 1929 (Oribatei, Acari) from South Africa, new combinations and a key to the genera of the family. Mems Inst. Invest. cient. Mocamb., Sér, A, 9, 15-126, Corpuz-Raros, L. A. (1980) Philippine Oribatei (Acarina) V. Scheloribates Berlese and related genera (Oribatulidae). Kalikasan, Phil. J. Biol. 9, 169-245, Granpigan, F, (1933) Etudes sur le development des Oribates. Bull. Soc. zool. Fr. 58, 30-61, ———— (1936) Observations sur les Oribates (10e serie). Bull. Mus, natin Hist. natur., Paris (2) 8 246-253. HAMMER, M, (1961). Investigations on the oribatid fauna of ue snes Mountains, Part Il. Peru. Biol. Skr, 13, 1-157, 43 pls. ——_— (1967) Investigations on the oribatid fauna of New Zealand, Part II. Ibid, 20, 1-70, 40 pls. (971) On some oribatids from Viti Levu, the Fiji Islands. Ibid. 16, 1-60, 35 pls. Jacot, A. P. (1934) Some Hawaiian Oribatoidea (Acarina), Bull. Bernice P. Bishop Mus. 121, 1-99, 16 pls. Lee, D. C, (1970) The Rhodacaridae (Acari: Mesostigmata); classification, external morphology and distribution of genera, Rec. S, Aust, Mus. 16, 1-219. (1987) Introductory study of advanced oribate mites (Acarida: Cryptostigmata: Planofissurac) and a redescription of the only valid species of Constrictobases (Oripodoidea), Ibid. 16, 35-42, ——— & SwEPHERD, K. J. (1990) Magnobates (Acarida: Cc igmata: lozetidae) from South Australian soils. Thons. Ri Soe St 14, 179-186. LuxTon. M. (1985) Cryptostigmata (Arachnida: Acari) — a concise review. Fauna N,Z. 7, 1-106. Mauunxa, §, (1978) Neue und interessante Milben aus dem Genter Museum XXXIV. A compendium of the Oribatid (Acari) Fauna of Mauritius, Renunion and the Seychelles Is. LL. Revue suisse Zool. 90, 709-724. ——— (1984) Oribatids of the Eastern Part of the Ethiopian Region (Acari). V, Acta zool. hung. 30, 87-136. WILLMANN, CC, (1935) — Faunistisch-kologische Untersuchungen im Anningergebiet IV. Die Milbenfauna. I, Oribatei. Zool. Jb. Syst. 66, 331-344. BRYOZOA IN COORONG - TYPE LAGOONS, SOUTHERN AUSTRALIA BY MARGARET SPRIGG & YVONNE BONE* Summary Unilaminar, multiserial colonies of living Bryozoa (Conopeum aciculata) are widespread throughout the Coorong Lagoon, South Australia. The bryozoan is an opportunistic species that is able to tolerate variable salinities. It establishes itself early in spring and reaches maturity prior to the drying-out of its nearshore environment in late summer, whereupon it dies. It is intimately associated with serpulid growth in the northern Coorong Lagoon. C. aciculata has also been found growing in the hyposaline waters of Lake Clifton, Western Australia, where it is found within clotted, calcareous thrombolites. The age of the Lake Clifton bryozoan colonies is unknown. Sub-Recent buildups of C. aciculata are extensive along the north-eastern side of the Coorong Lagoon, between Magrath Flat and 2.5 km south of Salt Creek. The colonies are multiserial and multilayered. The architecture of these buildups differs, with those in the northern lagoon being densely packed, flat to globose coalescing mounds whereas those in the more saline southern lagoon are loosely packed, highly contorted and convoluted "pavements". Serpulid association with these Sub-Recent bryozoans is minor. KEY WORDS: Bryozoa, Coorong, hypersaline lakes, buildups, serpulids. Transactions of lie Royal Society of S. Aust. (1993), 117(2), 87-95. BRYOZOA IN COORONG + TYPE LAGOONS, SOUTHERN AUSTRALIA by MARGARET SpPRIGG & YVONNE BONE* Summary Spricc, M. & Bone, Y. (1993) Bryozoa in Coorong-type lagoons, southern Australia. Trans. R, Soe 5. Aust. U7(2) 87-95, 4 June, 1993 Unilaminar, multiserial colonics of living Bryozna (Conopewmn aciculala) are widespread throughout the Coarang Lagoon, South Australia. The bryozoan is an opportunistic species that is able to tolerate variable sallnilies 1 establishes itself early in spring and reaches maturity pelor to the deying-~ut of its nearshore environment in late summet, whereupon it dies, It is intimately associated with serpulid growth in the northern Coorong Lagoon, C. aciculara bas alse been found growing in the hypogaline waters of Lake Clifton, Western Austrafia, where it is found within clotted, calcareous throtnbolites, The age of the Lake Clifton bryozoan colonies is unknown. Sub-Recent buildups of C. aciculata ure extensive along the north-eastern side of the Coorong, Lagoon, berween Magrath Flat and 2.5 km south of Salt Creek. The colonies are imulliserial and multilayeréd. The architecture of these buildups differs, with those in the northern lagoon being densely packed, flat to globose coalescing mounds whereas those in the more saline southern Jagoon are loosely packed, hiphly contorted and convoluted “pavements”, Serpulid association with these Sub-Recént bryozoans is minor, Ker Wonns. Bryozca, Coorong, hypersaline lakes, buildups, serpulids. Introduction The soothern Australtan continental shelf and its coastal inlets (Fig. la) have provided an tdeal environment for prolific bryozoan growth and preservation throughout the Cainozoic Era, Accumulations of their calcareous skeletons attain great thicknesses over tens of thousands of square kilometres of both open water and protected embayments. Their deposits extend from high-tide levels about protected and open cousts to beyond the edge of the continental slope (James & Bone 1992; Bone & James 1993). The extension of bryozoan growth into sub-coastal lagoons is less well known. Today, an encrusting chetlostome anascan, Coneperm aciculata (MacGillivray, 1891), ts found in South Australia's Coorong Lagoon (Fig, 1b), where it tolerates salinities that range from well below sea-water through to hypersaline. This bryozoan was formerly reported as Membranipora aciculata (Bone & Wass 1990: Bone 1991), but has now been confirmed as C. aricudata. A millenium ago it thrived in the Coorong waters when they were only marginally more saline than the sea. Another occurrence of whal is believed to be the same species (Bock pers. comm.) has recently been discovered in hypesaline portions of Lake Clifton, abour 100 km south of Perth, Western Australia (Fig, ley, The sub-Recent history of the Lake Clifton example is not known. Interestingly, both occurrences ure found in sub- coastal interdunal lagoons facing open oceans in mid-30 degree south latitudes, Both lagoonal systems ans * Department of Gellegy, and Goophysics, University of Adelide, Adelaide, S, Aust. 5005, subject to a degree of ground water drainage via highly porous and permeable, semi-consolidated , aealianitic, Pleistocene back-shore beach dunes, Both lagoons are similar in that significant salinity differences are manifested along their length, and these fluctuate according to seasonal water influxes. Holocene dolomite is found in both systems (Rosen & Coshell 1992), Lake Clifton was probably once a continuation of the present day Harvey Estuary (Moore et al. 1984) into Which River Murray waters from the adjacent Darling Ranges flow at its northern, Peel Inlet end, but it is now cut off from the sea. The Coorong still receives influxes of water directly {rom both the sea and its own Murray River, Although the Coorong and Lake Clifton environments possess these basic similarities, they alsnm have significant differences in detail and salinity fluctuations which affect bryozoan establishments and growth. Tn this paper, the Bryozoa and their environments are compared and summarised, with the emphasis placed on the Coorong example. Geological Settings The Coormng linear lagoons and Lake Clifion are products of repeated oscillation of Quaternary sea level and regional tectonic uplift. This has resulted in a sequence of abandoned sea beaches across a width suitable to create significant separation of respective sea coasts and their fossilised backshore dune deposits, both oo and off-shore. This is more apparent in the South Australian example. The present Coorong Lagoon lies ih the latest interdunal corridors of successive strandings of parallel Ocean-beach and backshore dunes of the Bridgewater 88 Formation. This is a highly calcareous cross-bedded medium to coarse grained aeolianite. (Sprigg 1952). Similarly, Lake Clifton lies between Jinear shore- parallel Pleistocene ridges of calcareous sandy Tamala Limestone. These ridges show large scale cross ! GREAT | BARRIER j i REEF AUSTRALIA \ F | | SHABK WESTERN : BAYS AUSTRALIA. — | | Sou AUSTRAL 1A LAKE CLIFTON “STUDY AREA eke \Prertoal Th Bunbaty Fig. 1. la: Map of Australia showing the locations of the Coorong Lagoon and the Lake Clifton area. Location Map. Ic; Lake Clifton Location Map, M. SPRIGG & Y.BONE bedding, and are composed of fine to coarse grained skeletal-fragment calcarenite with variable amounts of quartz sand (Playford & Leech 1977), In both the Coorong and Lake Clifton regions fossil soils with rhyzoliths have developed at various levels VQ aa fe Milang RIVER s ee Lake Alexandrina M Y ai ‘hag HINDMARSH ISLAND Pelican Pt, —\ Mark Pt A 4 a - S\Na MENINGIE a\t i 6 7" Bonney Oy Reserve , Magrath Hells Gate < 9 ua a Zz < é =: § > Pollcamans Pt. * Gemini Downs lisalt Creeh 2 aS a BUL BUL BASIN As a q J LACEPEDE x STROMATOLITE am «LAKE 5 SHELF _) et 2 SCALE: kms 0 5 10 15 mmm Barrage Ib; Coorong Lagoon BRYOZOA IN COORONG sy within the dunes. These dunes in places include calcretised layers, if the soil forming periods were of sufficient duration. The Lake Clifton Tamala Limestone has calc rete-lined solution pipes (Semeniuk é& Searle 1985) that are similar lo those in the Coorong (Bone & Wass 1990). These solution pipes are features of calerete profiles, and are formed tube-like, by calcium carbonate precipitation around regions of net vertical water percolation (Scholle et al. 1983) through limey dunes, In the Coorong area, erosion has removed much of the original dunes so that many of these large solution pipes are free standing by the water's edge, where they appear like the remains of petrified forests. The Coorong and Lake Clifton interdunal areas were both inundated by the sea during the Holocene. and in euch case, the consolidated aeolianites or their calcrete cappings provide excellent bases for bichermal buildups. (Fig, 2). The Lake Clifton example The rise in Sea Jevel during the Holocene caused some of these shore-parallel limestone ridges in the southwest of Western Australia to become inundated and form a lakeland system of linear barrier estuaries and lagoons, including Lake Clifton (Fig, le; Searle & Semenivk 1985), The ridges, which enclose the lagoon and separate it from the Indian Ocean 1.5 km awity, dip below sea level just north of the lake and run across the narmow bryozoan-rich sediments A BRYOZOA HORIZON (Memoranipara acicuiald) spray ore Aray zane rhyzalilhs splast) sone SuUPTANdal gone ertiaal Zone OW Water. he orey muddy sand fool clasts centoned layet highly foneyeamped of Bryozpa region (forming bionarns) ra ~s*, Puan of + ~ oS, Gemini Pont py Te pron ecaley i “a Jt at , i Q Pp Se a rg cantoites 74 y bryozoai bed a i) > 10mm thick limit of \ Bryomge \ rool moulas & chantels Sandy sol) Seg TF of the Rottnest Shelf (Collins J988) ro re-emerge at Rottnest Island (Playford & Leech 1977), The 21.5 km long, 1.0 km wide lake is replenished by winter mains falling directly on to it and by underground water [rom un extensive aquifer along its eastern shore. The lake is shallow, much of it being Jess than 1.0 m and a maximum of 3.0 m deep (Fig. 3). Water levels seasonally Muctuate up to a metre, periodically exposing and desiccating the large areas of thrombolites growing there. A bryozoan, tentatively identified as C. aciculata, intimately grows over and through the clotted caleareous thrombolites (Fig. 4), The largest colony found was 2 cm in diameter, The thrambolites (Burne & Moore 1987) outwardly appear similar to stromatolites and are constructed in part by the trapping and binding of detrital sediment by cyanobacteria. As Well as supporting Bryozoa. the lithoherms provide shelter for communities of fish, amphipods, isopods, decapods, nenid worms and various insects, The spaces between the clots often become filled with sediment rich in small gastropod (e.g. Cariella sp.) and ostracod shells. The salinity of Lake Clifton remains less than that of sea water throughout the year, though other lakes in the same system range from) hyposaline to hypersaline. Their salinity and water chemistry, and that of the regional ground water, were investigated by Moore (1987) to elucidate why thrombolites were restricted to Luke Clifton. She found the waters of x karst Suflace ‘ cn OHalky 2are endonline es nenens y speleathion HARUPAN - Plaly “calerete" ae xy) > an 4 £52 woh t shallow lagoon & sandspe ( 9990) ABN Quart’ caroonate Marert 7990) {shelly sands) ~% Fig. 2. Gemini Downs, Coorong Lagoon. Section A-A, through headland, showing relationship of bryooun bioherms to caleretised acolianites. 90 M, SPRIGG & Y.BONE Fig. 3. Lake Clifton shallow water, hyposaline environment - reeds at margins in contrast to Chenopodiaciae rimming the Coorong Lagoon. Figure (holding a thrombolite) for scale. all the lakes had ionic compositions proportionally similar to sea water, but that Lake Clifton, which receives virtually no surface run-off, obtains a large volume of its annual water from low salinity (1-2 %) ground water. This water is enriched in calcium and bicarbonate (Moore 1984) which locally modifies the chemical composition of the sediment-water interface. Moore (1987) found the calcified thrombolites were consistently associated with the aquifer’s discharge area along the lake's eastern shore, It follows that this, then, must also apply to the Bryozoa. Her study suggested the debouching ground waters provide a chemical environment conducive to the formation of the thrombolites that the Bryozoa encrust. The Coorong example Geological Setting The Coorong (Fig. Ib) is a modern artefact of an ongoing process of successive strandings of ocean- beach backshore dunes straddled across the northern limb of the regionally uplifting Mt Gambier Arch. This uplift resulted in Tertiary seas withdrawing from the Fig. 5. Sub-Recent laterally extensive buildup of Conopeum aciculata in the Coorong Lagoon. Width of photo is 3 m. Fig. 4. Clotted, calcareous thrombolite from Lake Clifton. The bryozoan Conopeum aciculata is most densely intertwined in the central portion of the thrombolite. Murray Basin. Relatively rapid sealevel oscillation in response to dramatic climatic change left sub-parallel coastlines which can be tied to Milankovitch insolation signatures (Sprigg 1979; Idnurm & Cook 1980). These roughly coincide with the perturbation cycles in the earth's orbit during the late-Quaternary. A sequence of “palaeo Coorongs” exists between ocean-beach backshore dunes to the east of the present Coorong system. These dunes have been preserved by uplift and low surface drainage over the Tertiary karst topography. Sediment cores taken from the Coorong suggest that initially the Coorong was a_ protected-marine environment, becoming progressively more restricted and subject to harsh salinity fluctuations (Gostin e¢ al. 1988). Stable 80 isotope studies of the sub-Recent buildups of C. aciculata in the Magrath Flat area indicate that the Coorong was only marginally more saline than sea water when the Bryozoa grew there around a millennium ago (Bone & Wass 1990). Later low stands of sealevel have resulted in the t ~~ as \ Fig. 6, Colonies of Conopeum aciculata encrusting bottle found at Magrath Flat in early 1990. BRYOZOA IN COORONG S| exposure of earlier formed carbonate flats (von der Borch 1976), These for terraces around both major lagoons and also around the extensively eroded and modified beach that forms an intervening string of shoals and islands in the southern Lagoun. These latier lie between the Younghusband Peninsula barner beach- dune system and the earlier consolidated backshore calcareous dunes. Hydrology The elongate nature of the Coorong estuary in an aca of low, winter raintall and high, sumujier evaporation, and the restriction of walertlow between its hwo major lagoons, results ina longitudinal salinity gradient, increasing southwards (Noye 1974). ‘This 1s compounded by slow mixing of less dense river water from the Murray, flowing over denser Coorong waters when levels between the sea and lagoon are appropriate (Geddes 1987). Tide, wind direction, currents, irreyzular bottom topography and barrage openings all contribute to the south-eastern portion remaining hypersaline year round (Noye 1974), Unlike Lake Clifton, however, groundwater seepage via aquifers to the Coorong, although probably significumt, is minor Compared to the high evaporation rate (Jensen et al, 1984; Noye 1974), Although drainage earlier this century arund Salt Creek introduced some freshwater into the Coorong for a short period. water quality in the carly 1980s was probably similar to pre- drainage (Jensen er al, 1984), from the time of early exploration of the ares through unui 1864. Tf so, it seems thut for at least a century salinities affecting bryozoan growth have been similar to those of today, and that even weir and barrage constniction (1920s and 1930s respectively) may not have radically altered conditions. Prior to this, in periods of drought when the Murray dried up, the Coorong would haye becume saline, and major floodwaters reaching the mouth would have freshened the lagoons, just as happens today, Salinity peaks have been noled to gradually. increase over a penod of years (Jensen er al, 1984). Such increases arc terminated by the release of large quantities Of Murray waters during the spring of years of heavy flooding upstream. Some of these waters reach aod freshen the southern reaches of the Coorong before the cycle begins again Vensen ef al. 1984), Bryezoa in 1987 Bone discovered laterally extensive sub- Recent multi-laminar buildups (Fig. 5) of encrusting anascan cheilostome Bryozoa. Individual buildups. are up to 40cm in diameter and 30 em high. The most extensive bioherm growth visible centres on Muaprath Flat (Fig, Ie). The bryozoan was identified as Membranipora agiculata (MacGillivray 1891), but 1 is now stivpested chat mt belongs w the genus Conapeutn. From miid-1987 to miil-1989 rare small scattered colonies (1 cm dianteler) were found living intergrown with the serpulid, Ficopamatus enigmaricus (Fauyel), along the casters margin of the Coorong from the Murray River to 2 km north of Policemans Point. The range of (he living serpulids, however, was found to extend ty Sait Creek (Bone & Wass 1990). In Spring. 1989, conditions were upparently more advantageous lor the bryozoan. ft was found at Magrat Plat thriving on almost every hard surface, from the consolidated acolianite of relict sand dunes, to bottles (Fig. 6) and tyres. Colonies up to 6 cnr in diameter were found, bur only as unilaminar forms (Bone 1994). Later the same year, serpulids out-competed the Bryozoa to completely cover and conceal all trace of sub-Recent bryozoan buildups in this locality. The same occurred in 1991 with serpulid banks building on those of the previous year (Fig. 7). Thus, locatmp the sub-fossil bryozoan buildups in this area can be difficult even when their position is known because of the thick covering of serpulids, Searches during 1990-91 for new locations of the sub-Recent bioherms in the northern Lagoon proved fruitless because of the extent and thickness of these serpulid “banks” In 1991 in particular, for more than 20 km each side of Long Point, serpulids were prevalent im the shallows (Pig. 8). They were found in cauliflower-shaped proups of worm tubes attached to almost any potm ol hard substrate piercing the carbonate-rich muddy sands. They were even found colonising the handle attachment hole of an old pick, growing out from it in Opposite directions (Fig. 9). Modern serpulid mounds range to >LO m in diameter, with multi-seasonal growth making some up to 0.5 m high. They form biohernns (Figs 7 & 8) with many other animals additional to the Bryozou, such as crabs, isopods, copepods, amphipods, decapods, neriid worms (e.g, Cerdtonereis aequisetis Augener, 1913) and occasionally relatively large fish. Fip, 7. Living serpuhd mounds encrustng Sub-Recent bryozoan buildups in the northern Coorong Lagoon, 199}, Figure for scale. 7 M SPRIGG & Y.BONE Methods Mupping of the areal extent of sub-Recent buildups of the bryazoan, C acicalata in the Coorong was undertaken in concert with observation of its present day range, The easter side of the Coorong Lagoon was mapped in detail, fram Pelican Poin| to.8 km south of Salt Creek (Fig. tb). A boat was used on several occasions to cantinge mapping around reefs and islands inthe centeal part of the lagoon, and along the western shore, Salinity measurements and observations of colony. growth were made on a monthly schedule over two years, from pdints along the eastern coastline. Water saunples were collected for analyses at the same time. These were analysed for their concentrations of sodium, potassium, magnesium and calcium by the atumic absorptiun method The salinity of the same samples were tested by a Kent clevtrolyne conductivity meter, temperature corrected to 25°C, and converted ta % (total dissolved salts), Samples of very high sulinity were diluted and the salinities re-scaled . us the meter used was poorly scaled at very high values. Results The water salinity analyses confirm (he trends of earlier studies (Noye 1974; Geddes J984, 1987; Botting 1990). Data is biased, as sumple sites had to be along the eastern shoreline, Analyses of Na, K, Mg, and Ca showed the elements were roughly in proportion to seawater, but that usually Cas K. 4 water sample [rom the River Murray (S.A.) had Ca>Kk. Flooding of the Coorong by water released through the barrages stirred Up the fine carbonate mud on the floor of the Lagoon, reducing visibilty to almost zero. Ths, plus the greater water depth, hampered the mapping aspect of the study, Location of the Bryozaa was offer by touch and not by sight, so only a Small number were found. Howeyer, these were sullicient io indicate that the buildups are widespread in the southern Lagoon. Sub-tossil boildups were Jound at three separate levels, The highest and most altered level lies just below the high watermark, tiniming the terraces previously mentioned, The middle level is generally found 20-30 cm lower, These become exposed in summer, lying about IS cm above the lowest water level noted, ‘The lowest level is found approximately 50 cm lower down. In the noptherm Lagoon the sub-Reeent © aeieulata buildups form densely packed semi-flat layers (Fig. 10) The buildups inthe southern Lagoon haye layers which are convulute, which result in spaces occurring belween the more laminar layers (Pig. 11). These spaces often contain Coxiella confusa and other small gastropods, minute bivalves and many different ostracods, as well as foraminifera, The latler include Ammonia beceuril, Kiphidium articulamm and Discorbis dimidiarus, which aré marine fauna, and so may be midiecative ot much less saline water being present in this part of the Coorong at the time of bryozoan growth, Altermatively, they may have been re-worked fram sub- Recent sand dunes (Can pers. commn.), Qogonia, the fruiting bodies of non-niarine charophytes (c.g. Ruppia sp., Which is tolerant to a wide range of salinities from fresh to hypersaline in the Coorong today ~ Wamersley 1984) are alsa found within these spaces. In both 1989 and 1990 the bryazoan Cyphonaute larvae had settled and begun ta bud asexually by the beginning of Oetober. In 1990 at the end of the first week in Oclober colonies 10 cm mm diameter were common and widespread, growing in the central southern Coorong, both aleng the shore and around islands, and on “reefs” The most southerly discovery of hying €. acteulara was on the edge of the shallow hypersaline Bul Bul Basin, 4.5 km south of Salt Creek, These rare colonies Were. Small encrustations on the sides of consolidated terraces belonging to earlier carbonate flats of a higher water level (Table lL. Colony establishment time is not known for either year at che northern end of the study area at Pelican Point. At the torthern end of the Coorong, small colonies (Table 1) were common both years in the fresh waters adjacent to the barrages, where lake waters are known io seep through, A-sketch and cross-section over the Point adjacent tu the Gentini Downs Boat Ramp illustrate 4 typical southern Lagoon environment showing relationships batween features and the sub-fossil accurnulations (Pig. 2). These deposits are extensively intergrown around solution pipes. and some contorted bands (5-10 em thick) contam teepec-like structures and possible fractures. They are underlain by grey, semi-plastic mud, Tasie l. Size of colontey al madern Conopeum aciculats on north-eastern margin of Coareng Lagoon, Size differences were transitional between adjacent locations, “Warnecke Point’ is ath unnanwl] feature lying seth of Hells Gare, on the Warnecke property, Location Masiimum Sia (em) Bryazoan Colonies = — 989 1990) Pelican Paint 2 Mark Point 2. Lop Pot - Bonney Reserve — Magrath Plat 6 “Warnecke Point” 7 Poltcertans Povat 3 2uuwo We} Fp it] 0 Boat Runip Gemim Downs 2.0 trare) 2.75 (rave) Boat Ramp 4.5 km south of 16 Salt Creeks 15 (rare) BRYOZOA IN COORONG 93 Fig. 8. Close-up of living serpulid mounds, showing (heir similarity to stromatolite growth form, The destruction of the upper, central portion is caused by fishermen dragging their boats across the bioherms, Individual mounds average 1m in width Discussion Due to conditions prevailing in the northern Lagoon favouring prolific growth of thick serpulid colonies in 1989 and 1990, searches of this area did not reveal new locahions of sub-fossil Bryozoa- In the southern Lagoon, however, conditions were Jess beneficial to both serpulids and Bryyzoa. In 1990 serpulid growth was less abundant than in the northern Lagoon, but 1991 brought even poorer growth. Consequently ubiquitious sub-fossil sub-Recent bryozvan buildups were often exposed in the southern Lagoon. The convoluted nature of many of the layers. of the sub-fossil bryozoan buildups found jn the southern Lagoon were unusual in that instead of being parallel to each other like those found in the north, they were often highly contorted. Some twist upwards, only to turn sharply and grow back on themselves, thus leaving “holes”. Others were regularly laminated similar w those found in the northern Lagoon where one layer is laid down directly on the last, The growth of bryozoan colonies is largely affected by changes in their environment and this can cause changes in colony morphology (McKinney & Jackson 1989), ‘Thus it may be possible to trace parucular periods-of environmental change over Jarge areas of the lagoons through examination of the growth patterns of different buildups, if the contorted Iayers do indeed have a similar pattern at a particular time. The three separate levels of buildups found in the southern Lagoon may have grown concurrently, bul it is more likely that.as water Jevels are known to have changed in the lagoons that each might belong to a different time period. High water Jevels prevented clear mapping of these ditferent levels. Radiocarbon dating analyses currently being undertaken may elucidate these and other timing uncertainties. Microfossils found within the convoluted layers of bryozoan buildups include Oogonia from algae characteristic of marginal marine and non-marine saline lakes, The Foraminifera are all extant, benthic species, Cann (pets. comm.) suggested (i) that they could have become encased by bryozoan growth alter being washed in after eroding out of the surrounding aeolianite, or (ii) that having crawled there, the Foraminifera were trapped and died, as the fast growing, C. aciculaia entombed them. New colonies found a week into October 1990 were common and widespread in the southern Layoon. They were all about 1.0 cm in diameter. Colonies. 0.2 cm larger or smaller than this were not present, Thus, a switching signal for the beginning of asexual budding must have occurted almost concurrently over a large area. Actual growth rate was not determined, as a precise settlement timing is not known, but growth certainly appears to have been rapid compared to that Fig. 9, Serpulid colony growing through the handle-hale of a pick-axe, found in the northem Coorong Lagoun in 1991, O4 HRYOZOA IN COORONG Fig. 10. Common growth form of sub-Recent Conopeum aciculata in the northern Coorong Lagoon, The multilayered habirhas flat layers densely packed and relief is low, of most Bryozoa (McKinney & Jackson 1989). Growth ceases and the colony dies once it is exposed. The rate of multilaminar layer formation is also unknown. Only one example of multilaminar growth was found in present day colonies. The colony is located on the internal walls of a solution pipe, where iis well protected from high energy wave and storm damage. Tt may be that mullilayer growth occurred in (he one season, or that growth of the second layer occurred ina later season coincidentally on the earlier skeletal remains, Today, the living Bryozoa are only found growing on solid objects whereas many af the Sub-fossil buildups form pavements (5-8 cm thick) that appear to have grown out laterally from the terraces, over the soft carbonate muds. Other bands, up to 20 em thick, appear to have grown out laterally from consolidated acoljanite, so that now they are completely unsupported, other than for the hard rock they rim. ft appeurs. they avay have originally grown over unconsolidated sediment, since eroded away. Observations, coupled with analyses, show that very high salinities are lethal to C. aeiculara . These high Fig. 1. Common grown: foro: of sub-Recent Conopeum acied/ata in the southern Coorong Lagoon. The multilayered habit is convoluted and distorted, and results in many buildups becoming fragmented. salinities occur early in summer in the southern Lagoon, resulting in maximum colony size being relatively small, especially compared ta those in the northern Lagoon. This can be seen from Gemini Downs southwards to Salt Creek, where colony size ranges. from 2.75 cm down to 1.5 cm, suggesting 4 very Short growth period, particularly in 1990. The switch to the salinity level lethal to the organism could not be pinpointed, as samples were collected nearshore and thus did not always allow for local and duily salinity variations, due to slow mixing. However, even here daily variations are observed, confirmed by Botting & Associates (1990). Conclusions |. Sub-Recent buildups of the bryozoan Conopeum acioulata are widespread. in both the northern and southern lagoons of the Coorong. The buildups are up to 30 cm high and 40 cm in diameter. Growth form in the northern Lagoon is multiserial, noultilayered. gently arcuate and compact, Growth form in the southern Lagoon 1s multiserial and multilayered but the layers are highly convoluted. 2. Modern colonisation of hard. substrates in the Coorong by €. aciculaia is seasonal, occurring in the spring; and even more widespread than the sub-Recent occurrences. Colony size increases rapidly, utilising multiserial, unilaminar growth form. Maximom colony size 16 6 cm in diameter, occurring in the northern Lagoon, Serpulids frequently outcompete the Bryozoa, thereby concealing evidence of annual bryozoan presence. 3. Salinity in the Coorong increases southwards and increases seasonally up to 60%, due to high evaporation in summer. The higher salinities are lethal to the Bryozoa but not to the serpulids. Similarly, exposure duc to summer water-level fall kills the Bryozoan but not the serpulid colonies. Cyphonaute larvae are able to survive the high salinities, and re- colonise when conditions are optimal, Small bryozoan colonies occur in Coorong hyposuline waters adjacent to the fresh-water lakes at the northern reaches of the Coorong. 4. Modern C. aciculara occurs intimately associated with thrombolites in fresh water on the eastern margin of Lake Clifton, Western Australia. BRYOZOA IN COORONG us Acknowledgments We would like to acknowledge R. Sprnigg, A. Chivas and P. MacDonald for continued support and suggestions; P, Cook, P. Bock and the Jate S. Parker for assistance in identification of C. aciculata: I, Cann for identification of Foramintfera; L_ Moore for assistance with the Lake Clifion work; P. Hutchings for identification of serpulids and neriid worms; S.A. Dept of Fisheries for boat loan; S.A, Engineering and Water Supply (J. Whitbread and A. Jessup particularly) for their assistance; technical staff of the Dept of Geology and Geophysics, University of Adelaide. References Bane, ¥. (1991) Population explosion of the bryozoan Membranipora aciculaty in the Coorong Lagoon in late 1989, Aust. J. Earth Science 38, 121-123, —____. & Wass, R, (1990) Sub-Recent bryozoan-serpulid buildups in ihe Coorong Lagoon, South Australia. ?bid. 37_ 207-216. ——___ & James, N. P. (1993) Bryozouns as carbonate sedimem producers on the coul-water Lacepede Shelf, southern Australia. Sea, Geol. (in press), Borminc, J. & Associates (1990) Coonsng and Murray mouch working group project: Preliminary assessment of rok data base, (Engineering & Water Supply, aide) Burngé, R. Y. & Moors, L. S. (1987) Microbialites: Organosedimentary deposits of benthic microbial cofmumunities. Palatos 2, 241-254 Co.iins, L. B. (1988) Sedimems and history of the Rotinest Shelf, southwest Australia; A swell-dominated, non-tmpical carbonate margin. Sed. Geol. 60, 5-49, Esvewan, M, & Kiarea, C. F, (1983) Subserial exposure pp.l-5S4, in P. A, Scholle, D. G. Bebout & C. H. Moare (Eds), “Carbonate Depositional Environments’. 44?G Memoir 33. Geppes, M CC, (1987) Changes in salinity and the distribution.of macrophytes, macrobenthos and fish in the Courong Lagoons, South Australia, following a period af Rivee Murray flow. Trans. R. Soc. S. Aust. 111, 173-181. —____ & Burner, A. J. (1984) Physicochemical and biological studies on the Coorong Lagoons, South Australia, am) the effect of salinity on the distribution of the macrabynthos. Ibid. 108, 51-62. Gastin, V. A., Becperro, A. P & Cann, J, H. (195%) The Holocene tton-trepical coastal and shelf carbonate province of southern Austrilia. Sed. Geol. 60, 51-70. Tonukm, M. & Cook, P, J, (1980) Paleomagactism of beach ridges in South Australia and the Milunkovitch theory of ice ages, Nanue 286, 699-702. Jamas, N. P, & Bone, Y. (1989) Petrogenesis of Cenozair temperale wuler calcarenites, South Australia; A model for meteoric/shallow burial diagenesis of shallow water calcite sediinents, J. Sed. Pet, 59, 191-203. JENSEN, A., Hokv, P., Korii, Bh, Siemumep, &., ‘UiLL, M, & WEINER!, M. (1984) The effects of drainage on rovectesior behayiour in countries Cardwell and uckenghara and the effect on the Coorang: A report for ihe Minister of Water Resources 1981-1983. S.A. Dene Enviranment and Planning 66, 8-65 MacGituivray, PH, (1891) Description of new or Litnle known Polyzoa, Trans. R. Soc. 5. Aust. 3, 77.83. Moore, L. S. (1987) Water chentistry of the coastal saline lukes of the Clifton-Preston lakeland system, south-western Australia, and ils influence ou stronutolite formation. Aust: J. Marine & Freshwater Res. 38, 647-660, Kwort, B. & Stancey, N. (1984) The stromiatolites of Lake Clifton: Living structures ropresént the origins of lite. Search 14, 309-314 —__—& Turner, J, V. (1989) Stable isatopic, hydrogeochemical and nutriont aspects of [ake-groundwater relations at Lake Clifton, pp, 201-213, /n G, Lowe (Ed,) “Proceedings of Swan coastal plains groundwater Management conference” (W. Aust. Water Resources Council, Perth). Noye B,J, (1974) The Coorong - An introduction. pp.1'79 Jn J. Noye (Ed.) “The Coorong”, Dept. Adult Education, Onty, Adelaide Prob, 39, pote rh E. Besa R E. J uy) Geology and logy tinest Island, Geol. Sury, West, Aust, oe Rosen, M. R. & Cusner., L. (1992) A new location of Holocene dolomite foomation, Lake Hayward, Western Australia, Sedimentology 39, 161-166. SEARLE, D. J. & Semeniuk, V. (1985) The natiral sectors of the Inner Rottnest Shelf coast adjoining the Swan coustal plain Jown, RX, Soc. W Aust. 67, 109-115 SEMENIUK, V. & SEARLE. D. J, (1985) The Becher Sand, a new stnitigraphic unit for the Holocene of the Perth basin, thid, 67, 109-115. SpaiaG, R. C. (1952) The geology of the South-east Province,, South Australia, with special reference lo Quaternary coastline migrations and modem beach developments, Depr. of Mines, South Aust. Bull. 29. (1979) Stranded and submerged sea-beach systems of southeast South Australi and the aeolian desert cycle, Sed. Geol. 22, 53-96. voN DER Borcu, C. C. (1964) The distribution and preliminary geochemistry of modem carbonate sediments af the Coorong area, South Australia, Geochim. Cosmochim, Acta 29, 781-7199. (1976) Stratigraphy and formation of Holocenc dolomitic carbonate deposits of the Coorong area, South Australia, J. Sed. Per, 46, 952-966. WomersLey, H. B, 3, (1974) Plant life in the Coorong. Ja J. Noye (Ed,) “The Coorong.” Depr Adult Edunation, Univ, Adelaide Pub. 39, 81-88. FEMALES AND IMMATURES OF THE AUSTRALIAN CADDISFLY HYALOPSYCHE DISJUNCTA NEBOISS (TRICHOPTERA), AND A NEW FAMILY PLACEMENT BY A. WELLS* & D. CARTWRIGHT} Summary The female, larva and pupa of the northeastern Australian Hyalopsyche disjuncta Neboiss, 1980, are figured and described for the first time. Comparisons between Hyalopsyche disjuncta and several North American species of Phylocentropus Banks, 1907 (Family Dipseudopsidae Ulmer, 1904), show that both genera share apomorphous larval features. Transferral of Hyalopsyche Ulmer, 1904, from the family Hyalopsychidae Lestage, 1925, to the family Dipseudopsidae and supression of Hyalopsychidae are proposed. KEY WORDS: Trichoptera, Dipseudopsidae, Hyalopsyche, larvae, pupae, females. Transactians af the Royal Saciety of §. Aust. (1993), 417(2}, 97-104. FEMALES AND LIMMATURES OF THE AUSTRALIAN CADDISELY HYALOPSYCHE DISJUNCTA NEBOISS (TRICHOPTERA), AND A NEW FAMILY PLACEMENT by A. WELLS? & D. CARTWRIGHT} Summary Weuis, A. & Caxrwricur,, D. (1993) Females and immatures of the Australian caddisfly Hyalopsyche disjunen Neboiss (Trichoptera), aiid a new family placement, Trans. R. Sec, S. Aust, 107(2), 97-104, 4 June, 1993, The fernale, larva and pupa of the northeastern Australian Hyalopsyche dixjancta Neboiss, 1980, are figured and described for the first time, Comparisons between Hyalupsyche disjuncia and several North American species of Phylocentropus Banks, 1907 (Family Dipseudopsidac Ulmer, 1904), show that both genera. share apomorphous larval features. Transférral of Ayalopsyche Ulmer, 1904, from the family Hynlopsychidae Lestage, 1925, to the family Dipseudopsidae and supression of Hyalopsychidac are proposed. Key Woros: Trichoptera, Dipseudopsidae, Hyalopsycke, larvac, pupae, females. Introduction The history of unstable taxonomy of the group of polycentropodid-like caddisfly genera including Dipseudopsis Walker, 1852, Protodipseudopsis Ulmer, 1904_ Phylocentropus Banks, 1907, and Hyalopsyche Ulmer, 1904, is discussed in detail by Weaver & Msalicky (in ms), These authors:present strong support for the familial status of Dipseudopsidae, including the first three of the above-mentioned genera, and possibly also Hyalopsyche, an African-Oriental genus, which was placed by Schmid (1980) with Phylocentropmus in the family Hyslopsychidae. Convincing evidence for the monophyly of Dipseudopsidae Ulmer, 1904, sensu Weaver & Malicky, derives from a consideration of features of the female abdomens and larvae. Although reference bas been made in the past to females of Hyalopsyche (Ulmer (1915) identified three New Guinean female specimens as H. rivalis (Betten, 1909)). no descriptions of females are available, and unti] now no larvae have been associated, A single Australian species of Hyalopsyche, H. disjuncta Neboiss, was described hy Neboixs (I980) from two adult males from northern Queensland, Repeated efforts to collect more specimens, mcluding larvae, failed; H, disjuncta appeared to be rare, Recently, however, during an intensive collecting tip of two week's duration in the Jardine River urea of fr northern Cape York (as part of the Royal Geographic Society of Queensland’s “Wet Season” Cape York Peninsula Scientific Expedition), the authors collected * Northern Territory Museuin of Arts and Sciences, PO Box 4646, Darwin, NT: O8Ql. Present address: Australian: peabeie Resources Study, GPO Box 636, Canberra, ACT 260) a Bileey Laboratory, Werribee Treatrnent Complex, Private Rag 10, Werribee, Victoria 3050, numerous adults of both sexes, and just prior to departure, made the exciting find of several larvae and pupae. Female and Jarval and pupal features of this species aré here described and illustrated, and the larval niche as described. For comparisons, some larval features of Phylocenirams sp., and the female genitalia and head showing the tentorium of the North American P flucidus (Hagen) are also figured. Confirmation of sharing of the specialised larval niche and other synapomorphous larval character states, firmly supports the monophyly of Phylocentropus — Hyalopsyche, and we present an hypothesis for the derivation af the specialised female penilalia af Hyalopsyche from those of Phylocentropus, We argue that Ayelopsyche should be placed in the family Dipseudopsidae sensu Weaver & Malicky (in ms), and the name Hyalopsychidae is here-supressed, Ross (1965) and Ross & Gibbs (1973) discussed the: phylogenetic relationships of Dipseudopsinae (in the Psychomyiidse and subsequently in the Polycentropodidie), but with more data available, phylogenetic analyses of higher taxa of Trichoptera now in progress (Weaver & Frazer pers. comm.) should provide a more definitive assessment of the relationships of this family. Materials and Methods Adak Hyalopsyche disjunct were taken mainly at 12v or By black or UY lights; adults and larvae were collected into and stored in 70% ethanol. Comparative material of the North American Plyocentropus tuciduy (adults) waa provided by Dr J.S. Weaver, and adults of P placidus and Dipseudopsis indicus.and larvae of Phylocentropus sp. were provided on loan by Dr J. Motse. Specimens were prepared for study by treatment in caustic potash, or using a lacto-phenol preparation. Drawings were made using camera lucidas on a Wild 98 A. WELLS & D. CARTWRIGHT m 4 0.1 mm Figs 1-3. Hyalopsyche disjuncta Neboiss, female: 1, dorsal head showing tentorium; 2, mesothorax, dorsal view; 3, genitalia, lateral view. Figs 4-6. Phylocentropus lucidus (Hagen), female; 4, dorsal head showing tentorium; 5, mesothorax, dorsal view; 6, genitalia, lateral view. Scale bars = 1 mm, unless otherwise indicated. FEMALES AND (MMATURES OF THE AUSTRALIAN CADDISFLY 99 M5 dissecting microscope and a Wild M20 compound Microscope, Material ts deposited in the following institians: Australian National Insect Collection, Canberra (ANIC), Museum of Victotia, Melbourne, (NM); Northern Territory Museum of Ans and Sciences, Darwin, (4TM); and Queensland Museum, Brisbane, (QM). Systematics Hyaloprehe disiuncta Neboiss, 1980 PIGS 1-3, 7-13, '6, 18-21 Hyalopsyche disjuncoa Neboiss, WH 357-361; Figs 1-8, Holotype male, N, Quecnsland, 29.iv.1979, NMV. Material examined: N, Queensland: 1 o, it? 50°S 142° 30°F, Bertic Creek, at Telegraph Crossing, 5.11.1992, D, Cartwright ond A. Wells, NTM; 2 > or, 2 99, UP 50'S 142° 30'E, Dulhunty River, a Telegraph Crossing, 8-9,i7,1992, 1D, Cartwnght and A Wells, NMV; 2 oo, 1 9, I" 39°S 342° 28'E, McDonnell-Cockatoo Creek jn, 14,7i,1992, D, Cartwright and A. Wells, NTM; DB oo, § 9 9, i" 44'S 142° 29'E, Gunshot Creek, at Telegraph Crossing, 17.11.1992, D, Cartwrighi and A, Wells, (M- ioe,’ & 9, same loc. and collectors, 4-15.1L1992. QM; S oc, 15 9 ©, same loc. and collectors, 18-19,71, 1992, NMY; 21 ovo. 4 9, same loc., $-5,iV.1992, M. Crossland, ANIC: 2 oa, 19 9G, same loc, and collector. 10-11.1v.1992, NM; | pupa, same loc., D. Cartwright and A. Wells, 18.13.1992, NTM; 2 larvae, satne loc and collectors, NTM; 7 larvac (L reared to adult 9), J pupa, same Joc. und collectors, 19,7i,1992_ Neboiss (1980) illustrates the general body features of male Hyolopryche disjancia, and gives anterior wing length as 6 mm, but in our samples male anterior wing length ranges from 4.3-6:2 mm. Females conform with males in general features, including reduced maxillary Palpi.and warts on the mesothoraa (Fig. 2), but are considerably larger. An additional feature. the temtorum, is. figured (Fig. lt: its arms are well developed, with inner lateral Manges posteziarly, and the posterior bridge complete Female (Figs 1-3). Anterior wing length 0-80 mm. Genitalia im the form of a slender, elongate, non- Tetractile ovipesitor (Fig. 3). Sternité VIII not subdivided, exterrial gonopod ot segment VIII slender, laperead, reachmg almost in tip of segment [X. Postenoriy segment X is divided to form pair of distal lobes. each with apical cercus and one papilla partially fused on inner margin, the second projecting venteally from the outer Margin at about the level af division of segment X. Larva (mature) (Figs 7, 8 12. 15, 16, 38-20). Campodeiform, slender, elongate, (Fig. 7). Mend, pronotum, limbs and anal claws brown-yellow or pale brown, dorsal margin of prothorax and strengthening tidges of thoracic pleury black, rest of body pale. Body length 6.0-14.8 mm, head capsule width 0604.88 mm- Head round Ja dorsal view (Fig, 12); fronteclypeus sub-tnangular, extending to posterior of head capsule, anterior margin rounded, wntorial pits situated posteriorly; labrum broadly rounded with few setae, anterior margin finely divided. Maidibles (Fre. 16) stout, dissimilar, each with differing numbers of denticles apically and on inner distal margin, and with a dense penicillus medially om inner margin, antero- lateral margins crenulate; right mandible ragose antero- dorsally. On ventral head (Fig. §3), anterior ventral &potome (gula) short, forming an isosceles triangle; submenium rounded anteriorly, paired setae situated subapically; maxillary palps relatively slender, lobus intertius broad, obliquely truncate in dorsal view, and bearing a brush of haiz apically; labium modified to form a slender, elongate spinneret, about 34 as long as head capsule, labial palpi absent. Prathorax only with sclerotised tergite. Fore trochantin fused to pleuron (Fig. 18). Mesostemum (Fig. 8) with a pair of elongate sclerites posteriorly, Legs (Figs 18, 19) short, flattened, tarai with dense Fringes of setac, claw on fore limb strongly curved, Fore tarsus with an udpressed row of modified setae on the mesial margim. Abdomen elongate, with a lateral fringe of Fine setae, without gills, but with a group of anal papillae (Fig. 7}; anal prolegs clongate, with cluster of lang setae anterior to the slender strongly curved anal claw (Fig, 20). Pupa (Figs 9-11, 21). Length 7.5-10.3 sun Labrum (Fig, 9) short, with a smal) marginal papilla apico- mesally. Mandibles (Fig. 10), simple, slender, curved. Hook plates (Fig, 21) with 3-5 demicles, on abdominal segments IT-VIIT. posterior plates on segment V only, developed ints hand-like processes, Posterior abdomen (Fig, Lf) divided into two simple setose lobes. Biology. Larvae and pupae were extracted from fragmented silken tubes. The exact architecture of the larval dwelling is unknown, but the branched portions of rabeg appear to have all the features of the tubes of Phylocentrapys placidus (see Wallace er al 1976) and FP carofinus Carpenter (see Wiggins 1977, Fig, 19.4D). (Such features are also shared by Protedipsendopsis sp. (uh-named) as jllustrale! Sy Gibbs (1968, Fig, 3)). The architecture of the Hyalopsyche disjuncta dwelling is probably similar: a blunt-ended vertical branch with a small wpening Projected into the flaw, a series of connected cross tunnels, and a dilated net chamber. Habient, Larval tubes were collected from a sinall atream with a rather unstable base of coarse sand, The dwelling tuhes were found attached to logs amongst deeply packed sand and litter, usually in situations Where the Utter, trapped against logs embedded in the 100 A. WELLS & D, CARTWRIGHT Figs 7-8. Hyalopsyche disjuncta Neboiss, larva: 7, whole animal, lateral view, 8, mesosternum. Figs 9-11. Hyalopsyche disjuncta Neboiss, pupa: 9, labrum; 10, mandibles, 11, posterior abdomen. Scale bars = 1 mm, unless otherwise indicated. FEMALES AND IMMATURES OF THE AUSTRALIAN CADDISFLY 101 13 Figs 12, 13. Hyalopsyche disjuncta Neboiss, larval head, dorsal and ventral views. Figs 14, 15. Phylocentropus sp., larval head, dorsal and ventral views. Scale bars = 0.1 mm. 102 A. WELLS & D. CARTWRIGHT Figs 16, 17. Hyalopsyche disjuncta Neboiss and Phylocentropus sp., larval mandibles, dorsal view. Figs 18-20. Hyalopsyche disjuncta Neboiss, larva. 18, foreleg and proleuron; 19, midleg; 20, anal proleg. Fig. 21, Hyalopsyche disjuncta Neboiss, pupal hook plates, right side, Scale bars = 0.1 mm. FEMALES AND IMMATURES OF THE AUSTRALIAN CADDISFLY wy sand, was begmming to decompose. Collections of adults were obtained at lights beside smull and larger streams; the smaller streams generally were almost completely shaded by the canopies of riparian drees, the larger 4treams were open, Discussion Comparisons of features of latval Hyalepsyche disfuricra With those of Phyloceniropus placidas, P fecidies and P. carolinus (fram Ross & Gibbs 1973 Fig, 2; Wiggins 1977 Fig. 15.41, and this work, Figs 4, 15) reveal extraordmarily ¢lose similarities between species mm the two genera, In derived or specialised features, la general form the heads are similar (compare for example Figs (2, 13 and Phylacentrapus sp., Pigs 14, i): mandibles of all species are stout, with rugose sculpturing dorsally, albeit on only the right mandible in 4, disjurcta; ventral sclerites differ slightly, with the submentum of H. dis/uncie rounded rather than subquadrate; the iobus tjrternus of the maxillary palp of 4, disjuncta is broad and apically truncate rather than tapered and rounded: labia are modified te form virtually identical slender, elongate spinnerets. Thoracic features, too, show close resemblance and the highly specialised legs differ only in the exact shape of the specialised setae adpressed to the mesal margin of the fore tarsi. A. disfwreta Jacks gulls but conforms with Phylocentropus species in fonn of anal prolegs.and their claws, The larval niches appear to be wentical, Pupae are relatively uninformative: mandibles of Hi, disjuncte resemble those illustrated by Ross (1944, Fig. 204) for P. placidtes. Little information is available On pupye of Phulocentropus. Some of the above features decur in other caddis larvae. For example, specialised clongate spinnerets are seen in larvae of some Psychomyiidae and some Ecnomidae, although tn these families the spinnerets are proportionally shorter than in PhWoeentropas and Hyalopsyche. Simple anal claws also occur elsewhere, but are probably plesiomorphous. The highly tnodified, flattened limbs, the form of the mandibles and the larval niche are found only in Hyalapsyehe, Phylocentrepies, Protedipseudopsis and Dipsenedapsix. The probability of such a suile of resemblances evolving by converyence are slim. The simplest hypothesis is that these taxa anvse from the sante specialised ancestral stock. Adult characters do not concur with those of the larvae. No clear synapomorphies can be recugnised, which ms the very reason Hrat placement of these genera has been confused for so long. Aniong adult features several small differences in general body features can be observed benwecn Hyalopsioke and Phyocentropus, such as the arrangement of setac an the meta-scutellum (see Figs 2. 5) and the absence of several crossveins in the Wings, a8 well as other more major anes, such as shape of head, position of antennae, reduction of maxillary palpi and form of genitalia, These differetices probably represent autapomorphies for the particular genera: The wings of Hyalopsyche can be derived from those of Pkylocentrapus, which closely resemble polycentropodid wings, simply by loss of several crossveins; those of Phylocentropus probably represent the plesiamorphous form. Similarities such as form nf the tentormin (Figs |, 4) are probably also plesiomorphous, Superficially, the female genitalia of Hyaiopsyche disjurcta (Fig, 3) appear to be strikingly different from those. of Phylocentrapus placidas (Fig, 6), Yet upon close’ examination, similarities ase apparent, particularly in the cerct and papillae, Derivation of the Hyalapsyche torm from that of Pirylacensropus tequires only simple elongation of the arrangement stern in Phylocentropus, probably adapting Hyalopsyche females for oviposition into sand af crevices, Investigation of the female of Dipseudopsis indtoa McLachlan during this study, indicated that an the basis of female genitalia and wing form, Phylocensropus and Hyalopsyche are closer than Dipseudepsis- Hyalopsyche. Association of females and larvae of a species in Hyalapsyche has helped to stabilise the higher level taxonomy of this genus, placing it in the family Dipseudopsidae. On the basis of studies reported here and examination of tite literature, we suggest that Hyalopsycke and Phylacenrrapus tepresent sister groups. Ross & Gibbs (1973) suggested on the basta oi similarities between FPhylocentrepus and Polycentropodinae that Phylocensrapus is closer fo the ancestral stock than other Dipseudopsinae (=Dipseudopsidaz) and, given its occurrence in the Oligocene Baltic ambers. they suggested a Eurasian origin for ihe subfamily (=family). We have not found evidence with which to dispute this contention — the single north-eastern Australian species in Hyalopsyehe is undoubtedly of Oriental origin. Careful phylogenetic analyses in the future will elucidate tntrafimilial relutignships, bul al last ite family Dipseudupsidae is reunited, Acknowledgments We thank the Royal Geographical Society of Queensland for providmg the opportunity and financial support for field work at “Heathlands” and the surrounding Jardine River area of Cape York, and for the zenerous help and assistance given to us by members of the Socicty durimg aur stay ut “Heathlands" D, Cartwright’s work was undertaken with backing from his family, and A. Wells’ with backing from the Northern Territory Museum of Arts and Selences, Darwin. We are grateful w beh partics. 104 A, WELLS & D. CARTWRIGHT Dr J.S. Weaver of the University of New Hampshire, USA, kindly provided material of Phylocentropus lucidus and Dipseudopsis indica from his collection for comparisons. Adults of Phylocentropus placidus and larvae of Phylocentropus sp. were made available by Dr J. Morse, on loan from Clemson University, USA. References Banks, N. (1907) Descriptions of new Trichoptera. Proc. Ent. Soc. Washington 8, 117-133. Betten, C. (1909) Notes on the Trichoptera in the collection of the Indian Museum. Rec. Indian Mus. 3, 231-242. Gisss, DG. (1968) The larva, dwelling-tube and feeding of a species of Protodipseudopsis (Trichoptcra: Dipseudopsidae). Proc. Ent, Soc, London 43, 73-79. Lestace, J.A. (1925) Notes trichopterologiques. VIII, Etude du groupe polycentropidien et catalogue systématique des genres et espéces decrits depuis 1907 dans Genera insectorum. Bull, Ann. Soc. Entomol, Belg., 65: 93-120. Nesoiss, A. (1980) First record of the subfamily Hyalopsychinae from Australia (Trichoptera: Polycentropodidae). Arch. fiir Hydrobiologie , 357-361. Ross, H.H. (1965) The evolutionary history of Phylocentropus (Trichoptera; Psychomyiidae). J. Kansas Ent. Soc. 38, 398-400. ____ (1944) The Caddis Flies, or Trichoptera, of Mlinois. Bull. Illinois Nat. Hist. Survey 23, 1-326. ___ & Girss, DG. (1973) The subfamily relationships of the Dipseudopsinae (Trichoptera, Polycentropodidae). J. Georgia Ent. Soc. 8, 312-16. Scumip, F. (1980) “Les Insectes et Arachnides du Canada, partie 7”. (Agriculture Canada). 296 pp. ULMER, G. (1904) Uber Westafrikanische Trichopteren. Zool, Anz, 28, 353-359. _____ (1915) Trichopteren des Ostens, besonders von Ceylon und Neu Guinea. Disch. Entomol. Z., 1915, 41-75. WALKER, F. (1852) Catalogue of the specimens of neuropterous insects in the collections of the British Museum, Pt, 1, 192 pp. (London). WaLtace, J.B., WoopalL, W.R., & Staats, A.A. (1976) The larval dwelling-tube, capture net and food of Phylocentropus placidus (Trichoptera: Polycentropodidae). Ann. Entomol. Soc. America 69, 149-54. Weaver, 1S., WI., & Macicxy, H. (In ms) The genus Dipseudopsis from Asia (Trichoptera: Dipseudopsidac). Wiccins, G.B. (1977) “Larvae of the North American Caddisfly Gencra (Trichoptera)”. (University of Toronto Press: Toronto and Buffalo). OBSERVATIONS ON THE BREEDING BIOLOGY OF A MICROHYLID FROG (GENUS OREOPHRYNE) FROM NEW GUINEA BY G. R. JOHNSTON & S. J. RICHARDS Summary Transactions of the Royal Society of S. Aust. (1993), 117(2), 105-107. BRIEF COMMUNICATION OBSERVATIONS ON THE BREEDING BIOLOGY OF A MICROHYLID FROG (GENUS OREOPHRYNE) FROM NEW GUINEA The conquest of terrestrial environments is a major theme in vertebrate evolution. Classically the Amphibia have been viewed as an intermediate stage between the aquatic and terrestrial groups!?. However, the standard view of the amphibian life-cycle involving eggs hatching into aquatic tadpoles which subsequently metamorphose into terrestrial frogs belies the true diversity of reproductive modes in the Amphibia. The diversity of reproductive modes is greater among amphibians than in other vertebrates*. All orders of the Amphibia include species which deposit eggs in terrestrial environments, and terrestrial development has evolved several times within the order Anura®. Deposition of eggs out of water is a major step toward terrestriality among amphibians. Therefore, it is not surprising that most variation in reproductive modes among anurans occurs in tropical environments where there are high levels of atmospheric moisture’. A second major step toward terrestriality occurs in those groups which exhibit direct development. The evolution of direct development has been important in the successful invasion of montane environments by amphibians*. Direct development occurs in all Australopapuan microhylid frogs‘. Despite their abundance, little is known about the breeding biology of New Guinean microhylids. Small numbers of large, heavily yolked eggs are concealed in leaf axils, hollow stems, moss clumps or beneath the ground*. They may be attended by an adult frog, usually a male. These frogs generally do not form breeding aggregations (exceptions may be Asterophrys turpicula® and Sphenophryne mehelyi’) and may call spasmodically. Thus any observations are fortuitous and valuable. This note reports observations on the reproductive biology of an unidentified species of Oreophryne near Tabubil in the Western Province of Papua New Guinea (5°17'S, 141°12’E). This species was assigned to Oreophryne on the basis of its partially webbed feet and the presence of small clavicles®, Current confusion surrounding the taxonomy of this genus prevents reliable allocation of our specimens to any particular species, On the evening of 27.xi.1991 a single specimen was found 3.5 km SSW of Tabubil near a small creek, beneath a closed canopy mid-montane rainforest”. At the time of collection it was sitting over a clutch of ten eggs on the underside of a leaf 0.8m from the edge of the creek and 1.5m above the ground (Fig. 1). It was not heard calling, but its sex was determined later by dissection. Only two other species of frog were heard calling in the area (Rana grisea and an undescribed member of the Litoria eucnemiy species-group). The clutch was arranged in two vertical lines of four and six eggs each. The entire clutch was covered with a thin, transparent, membranous structure which yellowed slightly in alcohol, This layer was quite separate from the jelly-like capsule which surrounded the embryos and, although superficially different, may be homologous to the cord that joins the eggs of congeners for which the eggs are known®:"|_ The capsules were 3,8-4,0mm in diameter, The Fig. 1. A male Oreophryne (SAM R40884) attending a clutch of eggs on the underside of a leaf near Tabubil, Papua New Guinea, embryos were 3.7-3.8mm long. Each egg contained a very large, ovoid yolk body 3.1-3.2mm in maximum diameter. A single egg from this clutch was dissected (Fig. 2). The head (1.1X1.lmm) could be distinguished clearly from the laterally compressed body, which was 2.6mm long. Four limb buds were present, although no differentiation of the digits was apparent. Posterior to the hindlimb buds the tail was strongly compressed laterally, but no vascularisation was apparent in the preserved material. Three cephalic lobes and a pair of pigmented eye spots were clearly visible. A single pair of gill buds were present on each side of the head. Dorsally, two dark lines ran the length of the body, indicating that the neural groove had not yet closed. With the exception of the presence of eye pigmentation and the lack of neural groove closure, these microhylid embryos resemble stage five embryos of the South American leptodactylid, Eleuthero- dactylus coqui g, Price” illustrated a single, much older embryo of Oreophryne from Yapen Island, Irian Jaya. The tail was greatly enlarged and highly vascularised late in development, and presumably plays a role in respiratory gas exchange. 106 ¥ \ Re Aaa yy, . igen Fig. 2. Dorsal (A) and lateral (B) views of an Oreophryne embryo. Scale bar = Imm. Measurements of the adult frog were: snout-vent length 22.6mm; head length 7.5mm; head width 7.8mm; eye diameter 3.1mm; eye to naris distance 1.9mm; internarial span 1.8mm; tibia length 10.9mm; foot length 14.2mm. The frog and the clutch it attended have been registered in the South Australian Museum as R40884. A further two, morphologically identical, male Oreophryne were found calling from a cluster of large leaves 3m above the ground, 4m from a sago, swamp amongst mid-montane rainforest 6km SSW of Tabubil at night on 1.xii.1991. Their calls were recorded with a SONY Professional Walkman cassette recorder and an Electret Condenser microphone ECM 2200. Calls were analysed using ULTRASOUND v1.10". The call (Fig. 3) was a biphasic rattle, beginning with a short (0.3-0.4s) series of pulses at 3.2kHz which was followed by a longer (0.7-1.0s) series of pulses at the fundamental and dominant frequency of 2.8kHz, and at 3.2kHz. There were 26 pulses.s'. Eight calls uttered by two individuals gave a mean call rate of one per 63s. The call length was 1.0-1.4s (mean = 1.1, SD = 0.17). The breeding behaviour of thirteen of the 83 species of Australopapuan — microhylids!*5:!6!7 has been documented>-1,1618,19.20_ Four to 55 eggs are laid!'. This reduction in clutch size associated with large, yolked eggs is a common correlate of terrestrial reproduction in amphibians”! ‘ Males are known to attend eggs in two Phrynomantis s P.> two Cophixalus spp. and three Sphenophryne spp.>* Both males and females attend eggs in two Orcaghiryne spp. and one species of Cophixalus 10,20 Tyler? reported aggregations of Sphenophryne mehelyi attending their individual clutches of eggs. It is not known how long this association of adult frogs with their clutches may last. The association of frogs with their eggs may function in manipulation of eggs, to provide moisture for terrestrial eggs, in Protection. against predators or for removal of dead or infected eggs a0 This report is the first record of any direct developing species laying its eggs on the exposed surface of a leaf. A number of indirect developing frogs from other families are known to deposit res on vegetation overhanging water in the neotropics” Africa, Madasgascar’>, the Phillipines”® and New Guinea’, In all of these species the hatching tadpoles fall into the water below. The Oreophryne we observed breeding in New Guinea were also found in association with water. This may have been an artefact, however, as we concentrated our searches for frogs around water bodies and did not search the forest surrounding them as thoroughly. This work was funded in part by grants from the Peter Rankin Trust Fund for Herpetology and the James Cook University. Ok Tedi Mining Ltd, and Phil and Sue Gregory . ‘ . ‘ae . \ ' 1 | | Oe a ' ee FREQUENCY (kHz) TIME (seconds) Fig. 3. Sonagram of the call of an Oreophryne trom 6 km southeast of Tabubil, Papua New Guinea. ‘Carroll, R. L. (1988) “Vertebrate Paleontology and Evolution.” (Freeman, New York). “Romer, A. S. (1966) “Vertebrate Paleontology.” 3rd Edn (University of Chicago Press, Chicago). ‘Duellman, W. E. & Trueb, L. (1986) “Biology of the Amphibians.” (McGraw-Hill, New York). ‘Zweifel, R, G. (1972) Bull. Amer. Mus. Nat. Hist. 148, 411-546, 5Menzies, J. I. (1976) “Handbook of common New Guinea frogs". (Wau Ecology Institute, Wau). §§. Donnellan, pers. comm. {987. 7 Tyler, M. J. (1967) Trans. R. Soc. S. Aust. 91, 187-190. SParker, H. W. (1934) “A monograph of the frogs of the family Microhylidae.” (British Museum (Natural History), lon). "Byndioan, D. C. & Menzies, J. I, (1990) J. Biogeog. 17, 241-273. "Zweifel, R. G. (1956) Amer, Mus. Novit. 1766, 1-49. "Price, D. 8. (1992) Sci. New Guinea 18, 3-7. "Townsend, D. S. & Stewart, M. M. (1985) Copeia 1985, 423-436, BJordan, B. (1988) “Ultrasound V,1,10" (Queensland University, St Lucia). 107 assisted with logistic support and transport in New Guinca. Ken Sanderson allowed us to use his software to analyse the call, Nelson Kanem assisted in the field. Margaret Davies and Paula Dempsey made helpful comments on the manuscript, Our work in New Guinea could not have been done without the help of Guy Kula (Department for the Environment, PNG), Rosalyn Busava (Institute of Papua New Guinea Studies) and James Menzies (University of Papua New Guinea), “aweifel, R. G, & Tyler, M. J, (1982) pp. 789-801, Jn Grissitt, J. L. (Ed.), “Biogeography and ecology of New Guinea.” (Junk, The Hague). SBurton, T. C. (1990) Trans. R. Soc. S, Aust, 114, 87-93. “Blum, P. & Menzies, J. L. (1988) Alytes 7, 125-163. "Zweifel, R. G, & Parker, F. (1989) Am. Mus, Novit, 2954, 1-20. '’Mfehely, L. yon (1901) Termes. Fuzctek 24, 169-271. Tyler, M. J. (1963) Trans. R. Soc. S. Aust. 86, 11-29. imon, M. (1983) Behav, Ecol, Sociobiol. 14, 61-67, “ISalthe, S. N. & Duellman, W. E. (1973) pp. 229-249, In. Vial. J. L. (Ed.), “Evolutionary Biology of Anurans: Contemporary Research on Major Problems”. (Univ. Missouri Press, Columbia). ?2Bokerman, W. C. A. (1966) Ann. Acad. Brasil. Cien. 38, 335-344. 2Duellman, W. E. (1970) Monog. Mus, Nat. Hist. Univ. , Kansas 1, 1-753. ~ADews R. C. (1984) Oce. Pap. California Acad, Sci, 139, 1-70. 25Blommers-Schlosser, R, M, A. (1979) Beaufortia 29, 1-77. 26Aleala, A. C. (1962) Copeia 1962, 679-726. 21Tyler, M. J. (1963) Trans. R. Soc, 5. Aust, 86, 105-130. G. R. JOHNSTON, Schoo) of Biological Sciences, Flinders University, G.P.O. Box.2100, Adelaide, §. Aust. 5001, and S: J. RICHARDS, Zoology Department, James Cook University, Townsville, Qld 4811. THE TADPOLE OF THE AUSTRALOPAPUAN FROG LITORIA NIGROFRENATA (GUNTHER, 1867) (ANURA: HYLIDAE) BY MICHAEL R. CROSSLAND & STEPHEN J. RICHARDS Summary Transactions uf the Riyal Society af $ Aust. (1993), 117(2), 109-110, BRIEF COMMUNICATION THE TADPOLE OF THE AUSTRALOPAPUAN FROG LITORIA NIGROFRENATA (GUNTHER, 1867) (ANURA: HYLIDAE) The tervesirial liylid frog Liroria nigrofreneta occurs in seasonally dry forest and woodland habitats on Cape York Peninsula, the Torres Srrale Islands, and in southern New Guinea’. Little is known about its biology, and the tadpole is hitherto undescribed. We present observations om breeding activity of adalt frogy and describe the tadpole, fron Heaililunds Reserve between Weipa and Barmaya on Cape York Peninsula (U°45/S, 142°35°E), Reptoductive activity was observed during February and April 1992, when males called from the banks of temporary ralo-filled water bodies, Nome were observed tose elevated calling sites, and all called within $ m of the water’: edge: Ovipasition occurred in five cattle watering tanks between December 1991 and February 1992, and tadpoles were also observed in a large (l4.4 46 x 05 mJ but temporary dam in April 1992, A amplectant pair (male = 38.0 min SV, female = 46.0 mm $V) collected at the dam on 14.i7,1993 laid 445 eggs in a single clump 2-5 egpy deep. Reproductive activaly was not observed during September and November/December 101 when temporary water bodies were dry, Despite imensive searches, no L, migrofrenata tadpoles were collected from permanent crecks (Cockatoo, Gunshot, Cholmondeley, and Bertie Creeks) or from two permanent swamps during the study, suggesting that this species breeds predominantly in temporary water bodies, Tadpoles were collected from the dam and watering tanks in April 1992, Tadpoles (n = 288) were collected with a dip Nel, measured (body length) with vernier calipers, and ylaged?. Some individuals were immediaicly preserved in 10% formalin. Others were reared to metamorphosis for identification. Preserved specimens were revarned to the laboratory and standard measurements? were made using vernier calipers (foral length aud body length) and a dissecting mictoscope with an optical micrometer (all other tneasurements). Measurements involving apertures were taken fram: the centre of thé aperture, Table 1 shows the size distribution of L migrofrenata ai stages 25-39, A tadpole at stage 39 is illustrated in Pip. 1 and has the following dimensions (mm); 43,0 total length, 19.80 body Jength, 11.72 maximum body width, 1,34 maximum body depth, 4.41 basal tail muscle height, 3.53 basal tail muscle width, 4,54 maximum dorsal fin height located 857 from body term/nus, 3,91 maximum ventral fin height located 6,30 trom body terminus, 3.53 tail muscle height a1 maximum dorsal fin beight, 4.03 tail nuscle height at maximum ventral fin height, 4.91 distance bel\veen joining of ventral and dorsal Fi aspects to body, 10.46 body width at eyes, 10.08 width of outer eye surfaces, 8.19 interorbital distance, 2.90 eye diameter, 101 pupil diameter, 0.38 narinl diameter, 35 intertarial distance, 2.29 snaut-nariy, 5.80 snout-eye, 109 snout-spiracle, 5,17 transverse oral dixe diameter, The-oral dist of a tadpole at stage 39 is illustrated in Fig. 2. The mouth 16 aoteroventral (almost ventmil), and the Posterior tim of the labium is strongly folded. The marginal papillae are in two rows anterolaterally and 2-3 rows posteriarly with a narmow gap anteriorly. Submarginal papillae eccur in a latge patch laterally on the posterior half of the TaBLe 1, Body lenetk (mm) and Gosner (1960) stage of Litoria nigrofrenata tadpoles. Stage N Mean bédy length + SD 25. 253 9.80+2.94 26 8 16,74+1,59 27 3 18.9740.97 28 1 19.00 29 2 17,.6042.12 30 4 S,0140,17 31 3 19.624 1,36 32 Zz 19.)5+0.64 335 [ 20.40 37 t 20.90 38 5 20.86+0.98 39. i 19,80 Teg. |, Luneral view uly Litoria nigrofrenam tadpole fi stage 39. Scale bar = 5 mm: ty Fig. 2. Oral dise of a Litoria nigrofrenata twdpole at stage . Scale bar = 1 mm. labiumt, The papillae on the anterolateral margins are smaller than those.on the posterior margin, and the papillae in the ouler row are smaller than those in the inner row. The tooth row formula is 2(2)/3 and the AZ yap is very narrow. One specimen has. an additional, poorly developed tooth row located in the anterior gap in the marginal papillae. The keratinised jaw sheaths are narrow und serrated. The upper sheath is broadly curved (medial section almost straight), and the lower sheath s broadly V-shaped. The eyes are large and located dorsolaterally. The nares:are dorsal, oriented slightly anterolaterally, and open almost vertically- The body is oval and rather slender. The fins are high. arched, and terminate in a narrow tip or flagellum. The sinistral spiracle is located ventrolaterally and oriented posterodorsally. The lateral wall is Jonger than the medial wall and is unpigmented on the posterior two-thitds. The orifice is large, and is clearly visible in dorsal view, The vent tube is short, dextral, attached to the ventral tail fin, and opens 1.3 mm from the tail-body junction. In life the dorsum is dark brown. A cream stripe runs from the tip of the snout through the top one third of the eye- and terminates just posterior to the eye. The ventral surface: has a coppet pigmentation which extends one-third of the way up the lateral aspect of the body, Posteriorly, three pale stripes extend transversely trom the lateral surfaces onto the dorsum, bul do not connect dorsally. The pale stripes on the snout and body, are most conspicuous in smal! (stage 25) tadpoles, becoming indistinct in larger animals. The tail fins have a reddish coloration mottled with patches of dark pigment. The copper coloration of the ventral surface, the pale stripes and the reddish tinge of the mil fins disappear in preservative, but the patches of irregular, dark pigment on the tail fins are retained. The maximum body length atiained is 21.15 mm (Stage 38), The SV length ofa recently metamorphosed individual (stage 46) was 16.85 rm, and the postoculay stripe characteristic of this species was clearly visible. It has been suggested’* that Literia nigrofrenata, t. personata and L. wotjulumensis are more closely related tr each other than. to other Australian frogs. Similarities between the tadpoles of Litoria wotjulumensis®, and L. nigrofrenata support that proposed close relationship. Litoria nigrofrenata tadpoles share with L. wotjulumensis a tooth row formula of 2(2V3, and arched, heavily mottled tail fins. L, wotjuliumensis tadpoles apparently lack the patches of submarginal papillae on the lower labium and have a white venter® (copper in nigrofrenata). The tadpole of L. personuia’ differs from L. nigrofrenata and L. wotjulumensis in having a complete bordct of marginal papillae, gaps in the P! and P2 tooth rows, law fins, and distinct gold stripes. The larval morphology of this species does not support a close: relationship with the nigrofrenata species group. MRC: thanks the McCleod family and the Lyon family (Queensland National Parks and Wildlife Service) for their hospitality and assistance during field work at Heathlands, and L. Schwarzkopf for field assistance. Julie Martin drew the tadpole and oral disc, and R. Alford commented on the manuscript. Field work was funded by a CSIRO grant to R. A. Alford. \Copger, H. G, (1992) “Reptiles and Amphibians of Australia” (Reed Books Pty Ltd Sydney). ?Gosner, K. L. (1960) Herpetologica 16, 183-190. 3Altig, R. A. (1970) Herpetologica 26, 180-207 3 aon ML J. & Davies, M. (1978) Aust. J. Zool. Suppl. Ser, 1-47. Tyler, M, J, Davies, M. & Martin, A. A, (1978) Trans. R. Soc. 5. Aust, 102, 15-457. “Tyler, M. J., Crook, G. A. & Davies, M. (1983) Rec. 5. fuast. Mus. 18, 415-40. MICHAEL R. CROSSLAND and STEPHEN J. RICHARDS, Zoology Department, James Cook University, Townsville Qld 48i1. VOL. I17, PARTS 3 & 4 30 NOVEMBER, 1993 Transactions of the Royal Society of South Australia Incorporated Contents Shiel, R. J. & Koste, W. Rotifera from Australian inland waters. IX. Gastropodidae, Synchaetidae, Asplanchnidae (Rotifera: Monogononta) - - a UNI Tian-Rui, Lin & Jago, J. B. Xystridura and other early Middle Cambrian trilobites from Yaxian, Hainan Province, China - - - - - - - 44h Pell, S. D., McKirdy, D. M., Jansyn, J. & Jenkins, R. J. F. Ediacaran carbon isotope stratigraphy of South Australia — an initial study - = 5 - 153 Nicholas, W. L. Two new species of nematode (Nematoda: Enoplida: Thoracostomopsidae) from Lake Alexandrina, South Australia - - - - - 163 Hoste, H. & Beveridge, I. Interspecific and intergeneric relations between nematodes parasitic in the stomachs of kangaroos and wallabies - - - 17 Ryder, M. H. & Bird, A. F. Effect of Acrobeloides nanus (Nematoda: Cephalobidae) upon the survival of Pseudomonas corrugata (Eubacteria) in pasteurized soil from Kapunda, South Australia - - = - = =e 179 Zbik, M. The Big Rock Donga Meteorite: A new H5 Chondrite from South Australia 183 Tyler, M. J. & Godthelp, H. A new species of Lechriodus Boulenger (Anura: Leptodactylidae) from the Early Eocene of Queensland- - - 187 Barker, S. A new Australian species of Calodema (Coleoptera: Buprestidae) - - 191 Brief Communication: Wallman, J. F. First South Australian record of the carrion-breeding blowfly Calliphora nigrithorax Malloch (Diptera: Calliphoridae) - - - - 193 Errata, Corrigenda, Addenda - - - = : 5 a : E - 194 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. 117, PART 3 ROTIFERA FROM AUSTRALIAN INLAND WATERS. IX. GASTROPODIDAE, SYNCHAETIDAE, ASPLANCHNIDAE (ROTIFERA: MONOGONONTA) BY R. J. SHIEL* & W. KOSTE; Summary A brief history of this 12 part series of papers on the Australian Rotifera is given. Of 28 rotifer families, the Australian species of 14 families were revised in the first eight parts. In this part, keys are given to the genera and species of three further families: Gastropodidae (Gastropus, three spp.; Ascomorpha, three spp), Synchaetidae (Synchaeta, ten spp.; Polyarthra six spp.; Ploesoma, two spp.) and Asplanchnidae (Asplanchnopus, two spp.; Asplanchna, seven spp.) all of which are planktonic in Australian inland waters. A new species of Syachaeta is described from the Fitzroy R., Queensland. All species of these genera recorded from Australian waters are described and figured with known distribution data and ecological information. KEY WORDS: Rotifera, Monogononta, Gastropodidae, Synchaetidae, Synchaeta_ sp.nov., Asplanchnidae, Australia, descriptions, keys, trophi, biogeography. Transactions of iw Royal Seetery of & Aust (19935, 1703). 1-139. ROTIFERA FROM AUSTRALIAN INLAND WATERS. {X. GASTROPODIDAE, SYNCHAETIDAE, ASPLANCHNIDAL (ROTIFERA: MONOGONONTA) by R, J, SHIEL* & W, Kostet Summary SHieL, R. J. & Kosre, W. (1993) Rotifera from Australian infand waters IX. Gastropodidas, Synchuetidae, Asplanchnidac (Rovitera: Monogononta) Troms R. Sur. S. Aust. 117¢3), 111-139, 30 November, 1993 4 brief history of this 12 purt series of papers an the Australian Rotifera is given. Of 28 rotiler families, the Australian species of 14 families were revised in the Test eight parts. In this part, keys are-given tv (he senera and species of three Surther families: Gustrepadidac (Gartropus, three spp.) Ascomorpha, iree spp), Synchaetidse (Synchaeta, ton spp.; Polyarthra six. spp.; Ploesuma, two spp.) and Asplanchnidae (Asplanckaapte, (Wo spp.. Asplanchna, seven spp.) all of which are planktonic in Australian inlund waters. A new. species of Synehagra is described from the Fitzroy R., Queensland. All species of these genera récorded From Australian waters are described and figured with known distribution datt and ecological information. Key Worps: Rotifera, Monogononta. Gastropodidac, Syachustidae, Syrohete sp.nov.. Asplanchinidaa, Australia, descmptions. Keys, truphi. biageograptty. Introduction An earlier paper (Shiel & Koste 1979) collated the known records of Australian Rotifera froma widely scaticred literature. Av that time, 33] laxa were recognized from inland waters of the continent. Thereafter we proposed to accumulate additional recoTds for publication as short checklists, with descriptions of new taxa as they occurred (e.g. Koste & Shicl 1980). Our own samples accrued, and widely separated colleagues sent material from various parts of the mainland and Tasmania, Many of the rotifers we found could not be identified from the keys of Kutikova (1970) or Koste (]978a), the recognized authorities. Rotifers generally were considered cosmopolitan at that time, however, our samples indicated that the Australian rotifer fauna contained a distinct indigenous component. Taxonomic keys to them simply were not available, so to provide such keys becume our aim. The probable audjence for a taxonomic volume on Australian rotifers comparable to the revision by Koste (I97Ra) was considered too small to bear the production cost. Further, WK had taken some 12 years lo complete the European revision, and the prospect of another 12 on the Austraban fauna was daunting. A series of sthuller “manageable” papers seemed to be appropriate To pul keys to species in the hands of the scatvered collectors. We therefore started to drafi keys in the common planktonic species in May 1984 (WK was then 2), working by farnily or groups of families, The Editor of the CSIRO joirnals was approached, and agreed to take the first manuscript, on blelloid ” Murray-Darling Freshwater Research Centre, PO. Barn 921, Albury, N.Si, Jeet + Ludwiz-Brill Strasse 5, 49610 Quakenhriick, Gertany rotifers occasionally found in plankton, This wis published in The Austratian Journal of Marine and Freshwarer Research (Koste & Shiel 1986), The follawing year, the second paper of the series (the monogonont families Epiphanidae and Brachionidae) was published in a new CSIRO journal, davertebrate Taxonomy (Koste & Shiel 19874). Then followed a hiatus when papers IN and ['V were “lost” in the system in 1987-88 and subsequently withdrawn, withapt rancour, from JAvert. Taxon, To keep the series in Australia we approached the Editor of the Transactions af the Rayal Sociery of South Australia, wece accepted, and saw the Buchlanidae, Mytlinidae, Trichotriidae and Colurellidae appear in 1989 (Koste & Shiel 1989a, b), Subsequently, Lecanidae, Proalidae and Lindiidae (Koste & Shiel 1990a, bh), Netommatidae (Koste & Shiel 1991) and Trichocercidae (Shiel & Koste 1992) have continued the series, In the inten, the contents were expanded to include all known Australian rotifers, not only pelugic taxa, The littoral taxa are considerably more diverse, and determining whether some of them were in fact what they were named has been a slow process. The need for a complete global revision of rotifer systematics compounded our difficulties. We apologize for the rate- doerease. but plead that demands on both of us are BreateT as more rescurchers become intercsted in the Rotifera, and seek assistance. WK bears |he weight of the global conirunity of rotiter workers; RUS has only Australasia to deal with, hence has taken over production of the series, and is responsible for errors or omissions which may sneak through. So, for those avid readers who have agitated for keys to the fiarnilies yet 10 appear, the end is in sight! Three further familics are reviewed in this part (Gastropodi- dae, Synchaetidac, Asplanchmdac). Two remaining ploimate families (Dicranophondac, Microcodinidie, ne k J. SHIEL & W. KOSTE 32 spp., part X) should be cumpleted early In 1994, and the eaght families of Flosculariacea (73 spp, . part X1) and Collothycucea (I6 spp., part AJ 1) during 1994, By (his Ume a global systematic revision now in preparation stud he available, and the confused stirs of sume uf the taxa in nur earlier parts be resolved (see ls Rinweagraphy seetiom farcr), The first and intmductary vulumehas been published (Nogrady er al. 1993) und jhe systematic sections will be puliished in purts ty lamily. For recent research on rotifers, see also Gilbert et al, (1995), The three fainilies reviewed here contain about 33 species, all mire or leas exclusively pelagic in habit ‘They are less diverse than other commun pelagic families, 22. Bracluoudae or Trichocercidae, both with 45 taxa recurded (Koste & Shiel 1987a; Shicl & Koste 1992), bit species nF Gastropodidae and Synchaelilie appear W be numerically the most common plankters mm Australiair inland waters. Species frm the three families occur in virtually any standing freshwaters, where they may reach very high densuies, cy. in Nov, 8) >24,000 Syrehaesa spp. I! uceurred in Mi Bold Reservols, S.A. (Shiel er af, 1987). This represents 4 significant biomass, even though individual animals are small (most <200 gm). The same study teporied sequentivl Syrchagta species replacements (Five Laxa) in the filling Dartmouth Reservoir (|978-80), but otherwise rotiler limmoplankton composition aid conmuuniy siceession is poorly documented in Australia. Billabongs in particular may support a diverse assemblage of specics at any time, with two or more fava from cach family co-occurring. Temporal succession in response to seasonal Changes in pliysico- chemical and biological conditions may be very rapid, e.g. ina billabong near Wixlonga, species dontinants changed within days in an autumn series of daily plankton satnples (Tan & Shiel 1993). In the gastropadids and synchaetids, high population densities, coinbined with specialized feeding habits, produce profound grazing effects on preferred hactertal/algal populations, In contrast, asplanchnids do na reach such high densities, but may be significant predators in their aquutic food webs, making other rotifers and nlicrocrustaceans. Species of Asplanchna miaty: be the largest prodaiory zouplankters (> 2500 pm) in some frabitats, Details of feeding preferences ale given in the systemaric section. We siress (nel ites very likely that andeseribed taxa of chese rotifer families pecurin Australia, More than, 50% of the species in some of the genera we have reviewed (1) date jire indigenous, yct only 4 single usplanchiyd and a single synchacaid described here apparenily are endemic, While this. disparity may be real, if ils may reflect s poor level of taxonomic ¢isctimination in earlier studies, ic, a tendency tb ‘shovhorty taxa into the nearest described species, The lorthcoming glohal revision WUT resolve many of the extant avonialics, For 4 critique of some problenis pertaining to Australian microfauna see Green & Shiel (992). In this review the furmat of earlier parts is followed; for convenienve, genera andl species are treated alphabetically. Keys to rotifer families are included in Koste & Shiel (987a), which also contains brief descriptions of general morphology. A family level key in Wallace & Snell (199]) also may he useful, Known distribution and ccoloyical informalion. are given for the species we have encountered, Global distribution and ecology is given in Koste (1978). Type saterial generally was not designated in many early sturlies, nor type localities given. We have imvluded type locality if it is known, otherwise probable place of origin is given in parentheses. Some carly authors did nut specify origin of malertal, however we consider iL tikely that in the late 48th-carly 19th century their collections derived fein rcasumably close to home. Methods Indiving material, the rotifers reviewed here are place] readily into their appropriate families and yeners hy ihcir characteristic hody morphology (Fig. b)- Gaslropadidae (d4sewmerpha, Gastripus) (Pig 1;L, 1:2) are ovate-globular with firm cuticular loricae, ant distincuve dark ‘defascation vesicles’ or coloured chromutophares in the starch wall. Syochaetidae (Sinchaert, Palvarthea, Ploesonia) (Fig 1:3, 14, 15) ime more varied; Synchaeta species are iMoricate, pyriform or conical with distinctive umerodareml cilluned auricles (Fig. 1:3a). Polywrrra species are small cubes with dorsolateral and dorsoventral faliate appendages (Pig. 4a), Ploeserne is firmly loricate, with distinctive delineation of the lurica, and relatively large foo (Fig. b 5a), Asplanchnids (Axplanchknagacs, Asplanckna) are barge sacle forms (to 2.5 iim) wilt large pincer-like incudate trophi. Asplanchnids are superficially similar to, and likely. tc be confused with) large saccale epiphanids (Epighanes) (ct. Rosie & Shiel 19878), with which they may co-occr Preservation in alcohol, formalin, vluteraldebyde, ete, induces strong contraction af illaricate saccate forms, of retraction of amerolateral auricles in synchaetids. Resolution af species in the case of strongly contracted animals requires clearing in hypochlorite [NaOCl) and examination of the sclerotized masiax elements — the trophus. Trophii are generically distinctive; indeed, evidence to date suggesis that they are species specific. Specific detennination is mare difficult for those taxa with small meimbraneous Lophi, ¢.2. svachactids, than for taxa with large heavily sclerotized tropht, ¢.g. ayplianchnids. The difficulty of identifying preserved Sjrchaexe spp, , for example, was discussed hy Ruuteer-Koliske (1974). Details of trophi preparation for light- and electron ROTIFERA FROM AUSTRALIAN INLAND WATERS 3 Microscopy are given in Koste & Shiel (1989v) and Sanoamuang & McKenzie (1993), Particular care must be taken with Syechweta and Polyarthra species, with prolonged immersion, NaOC!] is likely to erode delicate trophi. Rapid replacement of NaOCl! with non- corrosive mountant (e.g. 10% glycerol-H,Q) is necessary. i A useful pictorial method has resulted from developments in computer software and videography, High resolution ‘videographs’ approaching light- Micrograph qualily can be achieved electronically for reasonable cost. The advantages include immediacy, electronic storage, and ability to print black and while oc coloured images on s range of laser- or video printers. Several of the photographs in this part were printed using a Sony CVP-G700 printer via @ muicrosgope-mounted Sony DXC-IO7AP video camera, Electronic images also may be captured by a frame trabber and printed via a laser printer. The resolution on 4 600 dpi printer approaches black and white photograph quality. The muin advantage here is immediacy. Samples received for identification can be checked, and some or all taxa “grabbed” onto disk storage, An electronic reference collection can be built up, disks can be exchanged much more readily than bottle samples, and printed images can be returned to collectors for the cost of a photocopy. Systematic section Rotifer classification has been based largely on morphology, and only in the past decade or so have there been advances in comparative biochemical and ultrastructural methods, ¢.g. electrophoresix, restriction fragment polymorphism, polymerase chain reactions, SEM, TEM (Koste & Shiel 1989¢; Nogrady er af 1993). Tig. 1. Habit (a) and troptus morphology (b) off , Gaseropus, 2, Ascomerphas 3, Syavkaeia: 4, Polvarthra: 5. Ploesoma: 6, Asplanchnu. Afrer Koste (19788), 4 R, J, SHTEL & W. KOSTE Yhese methods will resolve some of the problems observed with morphological criteria, bul they are not readily available to the majonty of workers. While there are still few workers globally wrth the necessary expertise, morphology will continue to be the principle classification foul. The classificution followed here is that detsiled in Koste & Shicl (L957a) based on the revision by Koste (19782). This also is the classification used by Wallace & Snell (1991) and Nogrady et al, (1993), excep! that bath these works follow the American practice of regarding the Rotifera as a Phylum rather than a Class of the Phylum Aschelminthes. A new classification system based solely on trophi structure has been proposed by Markevich (1990). If ultimately accepted as better than the classical system, it will nor affect the specific, generic or familia! placement of the Australian rotlfer fauna, but will change the placement above family. Taxonomic descriptions of al) the species tremed here are afer the revision of Koste (19784). Some wax have wditional desetiptive material fram the onginal authors where we felt it was needed. In yome cases further description has been added from the Australian matcnal where we were confident that the taxon was the sume as lhe nominate species, bue for which the onginal deseription was tMmadequate by modern standards, In some cases the original description lacked figures. trophus details, etc. We have tried to provide figures of general morphology, and to include trophi where possible, bul the process is fraught with difficulty when there is doubt that the Australia taxon is the nominate species. [n such cases we have included a comment, Abbreviations for morphometric measurements: BH=body height; BL=body leneth; BH=body width; FT=fom; FP=fin. Fl adult females unless otherwise noted, Family Gastropndidae Remane, 1939 Gustropodids are small, usually highly coloured rotifers common in inland waters Australia-wide They! are characteristically fast swinimers, abundar in spring inthe pelagic of reservoirs and billubongs, also in slow reaches of rivers (Shiel er al, 1982; Kobayashi & Shitd, in press). Two genera, Gasrrepus, with three spp. known and Ascomorpha, with six spp, known globally (Koste 1978a), Three species of each have been record- ed from Australia. It is not unusual to find more than one species of each genus co-necurring. In this event there 1s oftena size difference in the congeners. appar- ently Im ublize different resources. Features: rarely sermiplankionic: wel, stecate Or flask-shapeds laterally Natiened: loot present (Gastropus) or vbsent (Asco- moarpka): apical field has vemacles in some species; Mastax virgate, in Guayrapus with prepharyngesl cuticular tube, stomach lobed or with blind sacs. Key to penera 1. Foot absent, 1-4. dark brown to black defaecariun l=uceretion) vosivley,. 2.20. 22. Ascckmarpha Periy Foot present, anc ar two toos, no dark vesicles. -Casrropas Imhot dAscomorpha Pery Ascomorpha Perty 1850; 18, Ape: Ascamerpha ecaudis Perty W850, p.18. Body saccate ot ovoid, may be dorso-vencrally compressed, lacks font; cuticle thin, lightly stiffened or with dorsal and ventral plate (A. ovalis), apical field with membranelles, ciliary bundles, styli, palpar organs, Ingerlike tentacle used for holding prey cells; trophi virgate; unci chin, stilewo-Like: rami long, acute, elongate, right-angled dorsally (straight in A, ovatlis); stomach large (lobed or blind sacs}, filling almost entire body; intestine, cloaca and anus absent, wastes stored in 1-4 defaecation or accretion vesicles; large protonephridial bladder present; one cerebral cyespot. displaced to left in A. ecayais and A, saltans; dorsal and lateral antennae very small, difficult to see. Ascomorpha species teed by sucking chromatophores and cell contents from algal cells, or ingesting whole vells. These may be incorporated into the stomach wall, where they may survive and divide, before being digesied. Key to species of Ascomorpha Porty knawn fram Australia 1, Corona with finger-like palpar organ (Pie. 24, Ba) 2 Palpar organ abyent (Fig, hi). - (A, ecavedes. Petty 2, Lorica with distinct dorsal and ventral Plates... 5 vey eee, ovalin (Berpendall S\iffened cuticle may have. Ainac, hit od distinct plates ...--A salts Bartsch Ascumurpha ecaudis Perty PIC. 21 Ascomorpha tedudis Perty, 1850, p18. Sacculus viridis Gosse,, 151, p. 188. Type locality: (Switzerland). Descriptian: Saccate body widest in distal third; cuticle not striated; apical field with ciliary tufts and elongate styli; stomach Jobulate with yellow/green/brown contents; four lobes with dark defaecation vesicles, darker in older animals, vitellariuny with eight nuclei. ‘Trophi; tamivucute, elongate, with triangular alulae (Fig. 2:1b); manubria well developed, fenestrated, terminally straight or slightly curved: 1-2 SE carried w#tuched to cloveal opening; RE sp.nulate. ROTIFERA FROM AUSTRALIAN INLAND WATERS Ns TL 130-200 wm; TR 35 pm (FU 17: R 18; M 14) Ascomarpha avalis (Bergendal) Distribution: Perennial in plankton of ponds and lakes, FIG, 2;2 often with spring/autumn peaks. Probably panconti- nental. but not yet recorded from W.A. Abundant in rie tes ey aa cat ee P. us 4 R. Murray billabongs, also in lower R. Murray Spprigphe Welle: Carlene BAS: p,'34, plankton in summer (Shiel et af. 1982): 8.0-29.8°C, Type locality: (Greenland), pH 4.328, DO 5841.2 mg I, 16-551 pS cm', Description: Lorica consists of distinct dorsal and 68-400 NTU, alk. 1.9 mg I), TDS 24.9 ppm. ventral oval plates covered with thin membrane (Fig. Fig. 2. 1, Ascomerpha ecaudis Petty: (a) ventral: (b,c) irophus: (d) traphus, lateral: (e) lateral: (f) body section, 2, 4, ovglis (Bergendal); (a) ventral: (b) lateral: (c) trophus: (d) fulcrum, lateral: (c) sensilla; (f) using tentacle to hold Ceratium: (g.h) tentacle or pilpar organ, 3, A. saltans Bartsch: (a) lateral: (b) dorsal; (c) trophus, Jateral: (¢) trophus: (c.f) body sections: (g,h) morphs. from different populations, Wf after Donner (1943), (fa-e, 2, 3 after Wulfert (1960), Scale bars: adults 50 yim, tophi 10. ym. ts kK J, SHLEL & W. KOSTE 2:24, b), apical field with single main tentacle and smaller uovessury pulpar organ; median red cerebral eye. dorsal antenna small, lateral antenna vot reported. Troptn O) simple structure: rami occasionally asymmetric. with bung alulae; two suprarami above them; unci long. thin sods; manuhria distally double- crooked (Fig. 22c); stomach with four blind sacs, wastes accumulate as dark irregular balls (generally fout); RE spherical, covered with small ‘warts: TL (00-200 ym; male 70-80. am, TR 35 ym (FU 20, M 20, U My; RE 6460 pm, ME 44x40 pm. Distrituien: Convonon in plankion, particularly during Peridinium or Cerarivim blooms, A. qylls grasps dinoflagellate cells. with ats tentacle (Fig. 2:2f), bores. through the cell wall with the unci, and uses the mastax asa pump to suck oul the cell contents, Chromato- phores are digested intracellularly in the stomach wall. which Ts brownish-yellow us a consequence. Known from N.S-W,, N-T.. Qld, Tas. & View 14.0-26.0°C, pH 6.2-18, DO 8-94 mg I, 35-190 pS emy!, b-120 NTU. Ascamtorpha saleors Bartsch FIG. 2;3 Arcnimerpha selters Bartsch, 1870, p, 364 Tyne lacatitys thei Tibingen’, Germany. Description: Body oval, cuticle stiffened; dorsum with raised, flattened central section between parallel grooves (Fig. 2:32); head with many folds in contracted individual, apical tentacle commonly deflected dorsally, with two adjacent unusually long stylii (Fig. 23a, b); two bundles of stiff bristles form fine tubes each side of mouth: mastax with 2-3 sallvary glands; stomach not lobulate; gastric glonds no described. Trophi: rami with alulae and domed suprarami, manubrin rod-shaped; Fulcrum curved dorsally ar distal end, Ganglion with red cyespot (displaced to lefi); elongated retrocerebral sac behind ganglion; protone- phridia with ca. 3 flame cells; SE and RE sinoath; imle undescribed. TL. 100-165 pin, BW to’85 pm, height to BK um; ‘TR 28-34 wm. Distribution: YPancomtinental, not yet recorded from S.A. Summer occurrence in Murray-Darling feser- wits, e.g. 1. Burvinjuck. 12.0-27.0°C. pH 54-74, DO 58-30 met, 59-1900 4S cin!. 1-10 NTU. Comment; A single record of 4. saltany indice Trou L. Barracoota, Vic. (Berzins 1982) is unverified Smaller than the typical form (63 68 ymin contracted individuals), this Indian form has no appreciable trophi iiflerences from A. selnens and is here considered an ecotypic variant. Gasvrepus Imbhot Gastragus Imhof, 1891, p. 37. Body laterally compressed: cuticle stiffened: antersor mirgin undulate, corona a simple band of cilia, may have palps and ciliary bundles within the paracingulumn: hypodermis thickened, IMay have secretory function, c.g, gelatinous sheath in G, stylifer; dorsal antenna displaced caudally; lateral antennae asymmetric. difficult to see: Tastax with prepharyngeal tube (Fig. 3b, c);, wophi virgate;.rami stiletto-like or with multi- toothed forked apices; stomach saccate, contains oil droplets and coloured diciary lems, as in Ascomerpha, cells of stomach wall coloured green.or brown by intracellular ingested chromatophores; rectum reduced; anus barely visible, probably non-functional; vitelacium with $-24 nuclei; fotdistinetly annulated, inserted ventrally; single fused toc or two short acute toes, Males with rudimentary digestive tract, evertible penis, relatively large brain with conspicuous cerebral eye. Finely spihulate eggs are attached (o pelagic alpac. RE with short spines on outer shell, All three deseribedt Species are recorded from Australia, All are herbi- vorous, Their trophi picree algal eclls, particularly phytoflagellates. The mastax acts as suction puinp through the prepharyngeal tube Key to species of Gastrupus 1. Foor posteroventral with two lows 2G, stylifer Lmturf Foot niediwentral. wath single toe 20h). Anterior clangated (Fig. 2a); ‘occipital margin wnlbtting G. minor (Rousscleu Feud not Shonen octiplial marain smooth _G. Inpiopus (Ehrenberg) eve pees Gastropus hyptapus (Ehrenberg) FIG, 3:1 Notommata Ayptnpas Bhrenberg, 1838, p. 426 Fig. 50:6, Gastrapus hypiopns: Weber, 1898, po 752 Type locality: ‘Bei Berlin, Germany Deseriplion: Body Uansparent, amlerior cuticle stiffened, not tapered.and clongated, with longitudinal furrows; dorsally a short keel; foot short, 2-segmentexl in adults; stomach with large oi! droplets, contents mostly yellowish; brain with large dorsal cerebral eye; vilellarium with many nuclei. Trophi: rami apices with five prongs (Fig, 3:le); Jarge semicircular alulae on inner margin of manubria; males known; RE spiny (Fig, 3:12). TL 150-363 wim; T 25-30 pin; male 60-90 pam, TR 30-37 ym (FU 14 pm, M {5 pm), Distribution; Widely distributed in eastern Australia, from Qld to Tas., generally in stnaller waters, &.g. billabongs or stock dams. Apparently wider thermul tolerance here than in Europe, where it 1s reported as a winter farm. &0-23.5°C, DO 4.1-108 tg |, pH 4.47). 17-240 pS cm, 5-120 NTU, Literature: Evans (1953), Green (981), Russell (1961), Gestrapacs atime (Rousselet) FIG, 3:2 ROTIFERA FROM AUSTRALIAN INLAND WATERS 17 Fig. 3. 1, Gastropus hyptupus (Ehrenbeng): (4) lateral: (b) lorica with striae: (c) body section: (d) trophus: (e) trophus, lateral: (f) male-egg: (g) resting egg: (h) another individual, lateral: (i,j) dorsal and lateral views [rom another pupulation, 2, G minor (Rousselet): (a) lateral: (b) ttophus: (c) trophus, lateral. 3, G. sxylifer (imbof): (a) lateral; (b) trophus: (c) trophus, lateral; (d) ventral: (e) animal in gelatinous sheath; (f) body section; (g) ventral with foot extended. la-e, h, 2, 3.after Wulfert (1960). If, g after Nipkow (1961). Scale bars: adults 50 am, trophi 10m. 18 R. 1. SHIEL & W. KOSTE Norops mitor Rousselet. 1892. p. 359, Fig. 24:4-10. Gastrapus miitor: Weber (898, p 752. Type locality: (Burope}. Description; Head tapered, elongate: foot south, slightly clongated, Trophi: needle-like rami can be protruded through mouth (Fig, 3:26); long pointed alulae, almost symimeirival, at rami bases; single uncer, manubria bilaterally lamellate (asymmetrical), with sweeping curve to distal cnaok; fileruen rodlike. Animal generally colorless, although stomach contents may be brown and contain oll droplets; 4-8 nuclei in vitellarium. BL 82-140 am, FT 35 pm: T 20-23 pan. TR 24-30 um, Distribution. Kaown from Qld. N,S.W., N.T., Tas. in small leatic waters, with a single record from the Hopkins R. estuary, Vic > 80-22.0°C, pH 4,457.7, 14-60 nS cm. <10 NTU, Liferaiwre: Shiel & Kosice (0979), Keste (9811. Gastropuy stylifer Tmhot FIG. 3:3 Gostroputs srylifer tmhod. 189). p, 37, Sec Koste (19788) for extensive synonymy Tipe flocalin: Black Forest, Germany, Description; Notably brightly coloured;. hypodermis dtue, body fluid pink; gut conrents green/brown: red cerebral eye; occipital muargiti undulate; with tongitudinal striae. raised keel in cross-section; vitellariuin with 6-8 nuclei, Trophi: thin, long fulcrum, rami with fong pointed alulac; manubria and unci apparently fused into a cup-shape (Fig. 3:3b): long curved prepheryngeal tube present; SE almost smooth; RE. warty. TL 220-250 wm; W.56-70 am; FT 40 pm: SE 60%40 um; RE 60*44 «im; male 80 pm; newly-hatched juveniles 75-100 yum. Distribution; Lays eges in colonial phytoflagellates (Dinebryon, Uroglena),; leeds on Feridinium and other Pynophyceae, sucking out contents (Koste 197%). Most common species of the genus in our collections, occaswmally co-occurring with G. hyptopus; 9-26°C, pH 43-76, DO 4,1-10,7 my T, 13-490 gS enr!, 5-120. NTU, Family Synchaetidae Remane, 1933 Soft culicled to mipre or less loricate; body conical, pyriform, cup- bell-shaped, vasiform or saccate: corona of Asplanciina type with or without. ciliary aurickes; rigid bristles near mouth; elongate coronal sensillae curve inwards; mastax virgate with complex paired hypopharynx muscles; manubria closcly asso- ciated with margin of pumping chamber; foot and toes presemt, rudimentary or absent. Of four described genera, Ploesume, Polvarthra and Synchaens are known from Australias Pseadepleescamd is not Key to genera |. Dloricate body, cuticle thin bul maintains shape; corona with lateral ciliary auricles; foot more or less distinct with [Wo short toes, occasionally one ©. et oe -Ayrchaera Ehrenberg Auricles absewre. 0.2. eee 2 2(1), Body ioricate, saccute-cubpidal, cuticle thin but rigid; foot absent; lateral bundles of rigid serrated Whds whe ws iwssye ....Polvarthra Ehrenberg Body loricate, generally with ornamented surfaces (ridges, fillets, etc); fooi-opening or ventral aperture present, foot annular and distinct, two LOCTe se segressemecries Plogsama Hernck Ploesoma Herrick Ploesoma Herrick, 1885, p. 57, Body bean-shaped/saccate, distinctly loricate, variously ornamented: dorsal lorica anterior may have headshield, smooth or denticulate margin; ventral lorica closed, with foot-opening, or open with ventral aperture; fool in part or entirely annulaic, with robust toes; corona ol Asplanchra type, with two long digiiform palps ip apical ficld; dorsal antenna displaced caudally; lateral antennae ventral in last third of body; mastax virgate, large, can be extruded to grasp food items; oesophagus long, with longitudinal striae; stomach in distal third of body, Of 7-8 species Listed in Koste (1978a), two are recorded from Australia. All known species are planktonic or semiplanktonic carnivores, cating pelagi: and benthic rotifers. Cunnibalism is noted (Koste 1982), Key to species of Ploesoma recorded from Australia Anterior margin of headshield snivoth, straight to undulate P. muteata (Levandery Headshield margin with median, short, triangulur toathlike OXUMSIOM. oe. telat it 3 3 eee P. lenticulare Herrick Plocsoma tenticulare Herrick FIG. 4:1 Ploesoma lenticutore Herrick, 1885, p, 57, Fig. Sab For extensive synonymy, see Roste (978a), Type locality: (U.3.4.). Description: Occipital margin with smoothly rounded projection; lorica outline variable within and between populations; dorsal transverse Furrow contains opening of dorsal antenna, three longitudinal furrows. between this median furrow und occipital margin; extensive ornamentation as in Fig. 4:la; lorica surface coveted with small round knobs; yentral lorica with deep cup- shaped aperture; apical field with two digitiform palps. membranelles and sensillae, Trophii fulcrum long, planklike in literal view; rami without inner dentition, with lurge basal plates; manubria with lamellae, unci with two main teeth and siristed plate, Male unknown, ROTIPERA FROM AUSTRALIAN INLAND WATERS tia St Oes Gg DaBHS ee ie “” = Hind. 1. Ploesomer lenticulare Herrick: (i) dorsal: (b) dorsal, robust form; (¢) ventral: (d) lateral. swiniming: (e) trophus: (1) Waplus, lateral, 2. P rruncata (Levander): (a) lateral: (b) ventral; (¢) ventral, foot extended; (d) dorsal; (e) dorsal, contracted: (0 trophus, ventral: (2) trophus, dorsal, (h) trophus, lateral (B= brain; Da= dorsal antenna; E= eye; Fi= foot; Pus fulcrum, Gg= gastric gland; H= headshield; Hy= hypopharynx muscle: La= lateral antenna: M= mastax: Man/Mn= thanubriun MI= tanubrial lamella; Mo= mouth, Oes= oesophagus: Oesm= insertion of oesophagus; Ra= minus, SQ~ salivary glands Ste stomach: T= toe: V= vitellarmum, (1 aller Wulfert (1961), 2 after Koste (1982). Scale bars: adults 50 am. trophi 10 aim. 120 R. J. SHIEL & W. KOSTE BL 150-320 um; FT 70 um, T 18 wm; TR to 120 pm (FU 56 pm, M 34 pm, U 22 wm); RE 147x125 pm. A sinaller morph from the Amazon (135 xm) has larger foot (79 wm) and trophi (128 zm: FU 77, R 51, M 23, U 26 wm) (Koste 1978a). Distribution: Early record from N.S.W. (Whitelegge, 1889). Not seen again until Sept. 1991, when indi- viduals of the robust morph (Fig, 4:1b) were found in samples from McCartney’s Creck and Horseshoe Billabong in Barmah Forest (Vic,), persisting in the plankton until 12 Nov.: 15.0-21.2°C, pH 7.3-7.6, 47-70 uScm!. Ploesoma truncata (Levander) FIG. 4:2 Gastroschiza truncata Levander, 1894, p. 25, Fig. 1:9-10. Ploesoma truncatum: Weber, 1898, p. 740, 743, Fig. 24: 8-10. Type locality: Vicinity of Helsingfors (=Helsinki, Finland), Description: Occipital margin almost straight, with lateral indentations; lines and furrows on dorsum form characteristic triangular ornamentation (Fig. 4:2d, e); foot partially annulated: toe of medium length; male unknown, RE with dark brown-black short-spined outer shell. Trophi: fulcrum rodlike in anterior view (Fig. 4:22), broad, spatulate in lateral view (Fig. 4:2h), manubria crooked, almost 90° bend, both arms similar length, distal arm reaches only midpoint of fulcrum; large hypopharynx muscle for pumping. BL 130-300 um; FT 80 wm; TR: FU 364m, M 26 pm either side of ‘elbow’, U 8 pm; SE & RE 60-70 82-90 jem. Distribution: Single record from L. Ashwood, an acid dune lake near Strahan, W. Tasmania (Koste er al. 1988). 16.0°C, pH 4.1, 131 pS cm!. Polyarthra Ehrenberg Polyarthra Ehrenberg, 1834, p. 226, Body cylindrical, conical, saccate or cuboidal, in some species slightly flattened dorso-ventrally; cuticle transparent, suffened; foot absent, At base of head, dorsal and ventral, two groups of three blade- or sword- shaped serrated finlike processes (variously termed finlets, fins, paddles or rudders) (Fig. 5) which are variable in length and width between populations. They are absent in first generation hatchlings from resting eggs. P. dolichoptera and P. vulgaris also have two obvious but shorter serrated fins on the ventral side (Fig. 6:Ic), Corona of Asplanchna type: in apical field. two ciliated tentacles and sensillae; lateral antennae on distal third of body; trophi virgate (see Koste & Shiel 1987 for details of trophi structure); trophi musculature. striated; vitellarium with 4, 8 or 12 nuclei; dwarf males are vasiform, finless, with ciliated penis and foot. Three form series were distinguished by Ruttner- Kolisko (1974), viz: dolichoptera-vulgaris, remata- minor and major-euryptera. At present, six of ten taxa listed in Koste (1978a) have been recorded from Australia. Comment; The difficulty of specific determination from preserved contracted material was noted by Koste (1978a). He recommended calculation of indices of body:fin length and fin length:width, carmine staining of vitellaria, NaOCl-clearing for trophi analysis, etc. Comparative indices were used successfully by Guiset (1977) to separate Polyarthra species in Spanish reservoirs. Some intergrades of body and fin-lengths have been reported, so collection of all the relevant morphometric information for a particular population may not ensure specific placement. It is likely that trophi are species-specific, and will enable accurate species recognition. Key to species of Polyarthra known from Australia 1. Ventral fins present (Fig. 6:le).. 0.0... 0...0....2 Ventral fins absent. 2... 0.0... eee ec eee eee ed 2). FW >15 wm; trophi as Fig. 8:3b. 00.0.0... 0c ee .-......P vulgaris Carlin FW 120 jam; vitellarium with 8 nuclet; trophi as Fig. Gi2b rrr GFR. Pelt sore tay. Pattaten atthe. neers 4 BL <120 pm: vitellarium with 4 nuclei; trophi as Fig. Silb of S2bis uc. cag esess pees aagatsry cose ed 4(3). FW 8-Il am; trophi as Fig. 6:2b.....,P. longiremis Carlin FW 20-37 um: trophi as Fig. 6:3b. 0.0.0.0... 008, bocezcrmmt beh, yap able etataa P. major Burckhardt 5(4). BL. <70 pm; FW <5 pm; left dorsal fins notably longer than others; trophi as Fig. 8:Ib...... P. minor Voigt BL >80 um; FW >5 wm: all fins similar length; trophi as Fig.¢ Bi2Ds bese eara wels oe P. remata (Skorikov) Polyarthra dolichoptera Idelson FIGS 5, 6:1 Polyarthra dolichoptera \delson, 1925, p. 84. Type locality: Novaja Zemla, Russia. Description: Body an elongated cube, more slender Fig. 5. Polyarthra dolichoptera, L. Hume, N.S.W. Dried onto slide to show fin morphology. Sony CV P-G700 video print. Scale bar = 50 pm. ROUIPERA FROM AUSTRALIAN INLAND WATERS 12] than Fo yulgariy, fins long, slender, extend beyond posterior margin: ventral fins fine, bristle-like. oveasionally only a litte broadened, finely serrated: median rib of fins distinet, continues to.upex (Pig. 5) lateral ribs may be indistinct or absent; margins strongly serruled; lateral antennae al posterior corners, median eve: mule known: RE with spinulate inner shell und rodlet reintorcement between inner and outer shells. TR usyounetne: cuch nimus resembles a hatchet with a single tooth on the blade’ fitting a niche on the opposing bhide: a reverse barb on distal end of blade leads to long, slightly convex ‘handle’ (bulls of ramus); viewed ventilly, lamellar mandi form hemisphere (Fig. 6:Ib); long rodlike fulerurn, broader in laleral view. BL 90-140 jan, FL WO0-220 pam; FW 7-15 «am, ventral FL 40-72 pm: TR 60 pm: SE with large oil droplets; RE 56-72 %36-56 jan: index FL:BL >. Distrihurian: Pancomtinenral in billabongs or lakes. also in spring, in Jower R. Murray plankton, $.A~ (Shiel etal, 1982). More common/abundant wy cooler waters, with isolated occurrences above 15°C. 7,0-20°C, pH 4.8-8.2. DO 66-12.5 mye 1!. 91650 pS em!, 15-120 NTU, hie 6 1, Polyarthra delichoptera Welson (a) lateral, only Lsetol fins shown, (6) frophus, dorsal (fe fuleruin, be lanvetha in front olin matnubrium, r= ramus); (¢) bileral with both jaleral fin groups and shorter yentral finlets Gurrowed); (db fin morphology, b Hume specomen from Mig, 5. 2, PB lengeemis Carling (a) literal. only | ser of fins shows, (b) trophus, dorsal, (c) fin morphology: 3. Po ryyer Burckhardt: (a) Jateral, onty | set of fms shawn: (b) trophus, (ed) fin groups, Jorsal. lic. 2a. eu, 3a, ec. d after KRutikova (1962), 2b after Kutikova (1970). Seale bars: adults 50 pam, trophi 10 pom, 122 R. J. SHIEL & W. KOSTE Polyarthra longiremis Carlin FIG, 6:2 Polyarthra longiremis Carlin, 1943, p, 88, Fig, 1:3. T\pe locality; Motala River, Sweden. Description: Of similar body form to P. vulgaris, with wider caudal region; fins thin, commonly longer than body, reach considerably past posterior end; ventral appendages bristle-like. Vitellarium with eight nuclei; Jateral antennae in distal “4 of body, TR symmetric, superficially similar to that of P. dolichoptera in ramus structure, bul readily distinguished by three apical unci teeth opposing proximal to rami (arrowed in Fig. 6:2b), with serrate margins distal to unci; manubria curving rods with crescentic alulae on outer margin, meet subramal fossa at proximal end of fulcrum, almost at right angles to fulerum (Fig. 6:2b). BL 15-225 wm: FL 135-208 pm; FW 8-1 yn; ventral FL 61-66 pm: FL:BL >t; FL:FW>5. Distribution: Recorded from Qld (Russell 1961), but not seen again until a 1985 sample series in Tasmania (Koste & Shiel 1986), where it occurred in Hydro- Fig. 7. Polyarthra minor Voigt: a. lateral, elongate dorsal fin set arrowed, BL 72 pm, egg 5329 um. b. trophus, 28 um. Sony CVP-G700 video prints. Electric Commission impoundments. It was found subsequently in a 1990 sample collected from L. Otaro, N.Z. (Coll. M.R. James, Taupo). 13.2-19.0°C, pH 4,2-68, 21-215 nS cm". Comments: P. longiremis probably is more widely distributed in Australasia than sparse records indicate. Close examination of trophi structure is vital for any P. dolichoptera-like rotifer collected at >15°C. Polyarthra major Burckhardt FIG. 6:3 Polyarthra major Burckhardt, 1900, p. 414. Type locality: (Switzerland). Description: ins shorter than body, leaflike with midrib, feathered, weakly serrate; no ventral fins; lateral antennae inserted well before end of body; vitellarium with eight nuclei; male unknown; RE with outer colourless smooth shell and inner dark brown smooth shell. TR asymmetric: opposing rami teeth similar to those of P. dolichoptera. However, whereas ramus proximal to each tooth appears knoblike in dorsal view in P. dolichoptera, proximal ramus in P. mayor is serrated (cf. Fig. 6:3b); manubria with distinct ‘elbow’, ca. 120°. BL 126-197 ym; FL 102-188 wm; FW 20-37 jum; FL:FW >S5; FL:BL <1; RE 80-100X60-72 pm. Distribution: Only two localities, Coliban Res and L. Catani, Vic., (Berzins 1982), Not seen in our material, unverified. Polyarthra minor Voigt FIGS 7, 8:1 Polyarthra minor Voigt, 1904, p. 33. Type locality: Vicinity of Plon, Germany. Description: Body relatively small and broad; lateral antennae medial; fins very slender; fins of left dorsal fin bundle considerably longer than other fins; vitellarium with four nuclei; SE carried attached; RE and ME unknown. Trophi asymmetric, similar to those of P. remata, although smaller; single pair of proximal large rami teeth, with series of smaller teeth distally (6-8), all fitting complementary niches in opposing ramus (Figs 7b, 8:1b); fulcrum rodlike in front view, with broader head laterally, i.e. similar shape to axe- handle. BL 68-70 pm; FL 57-70/86-90 pm; FW 4 pum; FL:FW >5; FL:BL >1; TR 25-28 pm. Distribution; Only known from a single locality, a billabong on Magela Ck floodplain, N.T. (Koste 1981, Tait et al, 1984), until 02.1i1i.92, when a population was recorded in Ryan’s #2 Billabong on the R. Murray floodplain near Wodonga, Vic. 26°C, pH 7.7, 296 yS cm'!. Probably more widely distributed. ROTIFERA FROM AUSTRALIAN INLAND WATERS 3 Palvarthra remata (Skorikay) FIG, 8:2 P platyplera var, remata Skorikov, 1896, p. 71, Fiz. 7:3-4, Polyarthra remata: Rodewald 1938, p. i47. Type locality: Viemity of Kharkov, Ukraine, Description’ Fins Jonger than body; posterior rounded/lobed, lateral antennae just before posterior cormet, cyespot dark ted-black; fins slender with robust midrib, no laterals; vitellarium with four nuclei; RE with smooth outer shell and wrinkled immer shell. TR asymmetric: single pair of acute unci teeth oppose at proximal margin of rami in dorsal view (Fig. 8:2b, arrowed): inner rani margins with single lurge and several smaller teeth (Fig. 6:2c): external margins of rami curved, lamellate, similar to P dalichoptera; manubria rodlike, curved, extend slightly beyond rami lamellae margin. BL 80-120 wm; FL 80-10 wm; FW 7-8 gm; RE 48-60%32-44; FL:FW >5; FL:BL=b Distribution; Known only trom Vie, (Berzms 1963) and Tas. (Koste & Shiel 1987b), 13-16°C, Polyartha vulgaris Catlin FIG. 8:3 Polyarthra vulgaris Carlin, 1943, p, 87, Fig. bd. Type locality: Motala R., Sweden. Description: Fins lanceolate, with medial and lateral ribs, margins serrate, fins may extend past body margin; ventral fins slightly broadened. lightly serraled; RE reddish-brown with smooth outer shell, hooked inner shell and intermediate folded membrane: Fig. 8 1, Polyathra miner Voigt: (a) lateral; (b) trophus, dorsal, 2, Po remara (Slorikov): (a) lateral; (b) trophus, ventral; (c) rami dentition; (d) tip of fin. 3, P vulyaris Curlin: (a) lateral, trophus extruded, and carrying parthenogenetic. egg: (b) trophius, ventral (ftom Magela Creek, N\T. specimen) la, 2, 3b after Chengalath & Koste (1988). Scale bars: adults 50 ym, trophi 10 pm. 124 R. J. SHIEL & W. KOSTE SE lightly coloured to (ransparent, with a large oil droplet; ME (up to six) cartled attached; TR asymmetric; in dorsal view two large teeth on left ramus (Fiz, 8:3b); single tooth on cight ramus fits between them; rami borders distal to teeth hemi- spherical, complementary, left convex, right concave im dorsal view, BL 100-145 pm FL 8-160 pm. FW 16-20 pm; veniral PI, 30-70 yon, RE 78-88 52-60 am: SE 76*50 pm; ME 26%26 um, Distréburion: Most common Polyrrtira in Australian walers, pancontinental in lentic waters, also common in fener be Murray plankton (autumm) (Shiel ¢ al. 1982), cpa Se pH 4.9-8.7. DO 1.5-12.0 mg 1). 13-1100 5 em!, <1-NO NTU. Svachaeta Ehrenberz Syathaeta Ehrenberg. 1832, p: 135. Cuviele transparent, pliable; body conical-vasiform, depending on ingested food, size of vitellarium and egg development; foot shori, one-segmented; loes generally acute; corona an interrupted circumapi¢al band with widely projecting cihary auricles; apical field with four styli, clongated ciliated tentacles and sensillac, dorsal antenna in neck, lateral antennae th posterior 44 or at base of foot; mastax Large with distinct striated muscles; trophi virgats, lane, delivave: fuleram and manubria long, thin; in some taxa uner acule, with comblike sermated edge; internal orgun 18 as in Fig, 8:1, SE und RE appear to have species specific morphology; males are known for some species only; >30 species described globally, but taxonomic resolution imprecise. About 24) of these are from athallasic saline or marine waters (Ruttner- Kolisko 1974: Koste J978a), Marine rotiters, including synchaelils, have been neglected in Australasia. Only S. baltic has been reported off Port Jackson by Whitelegge (1889), and in Port Phillip Bay (Evans (951)_ tt is not inctuded in the key, but @ description is givett for convenience. It is likely that further marine species. of Synchaera, inter alta, will be found here. A list of known marine synchaetids and relevant bibliography is given by Ruttner-Koliske (1974), Ten Swichaer species have been reported from imland waters. Mncludinag a new endemic species described hete, Lt wats firsi recorded by the late C. R. Russell_ Christchurch N.Z. froma sample taken in 1959 in Warragamba Dam but apparently not described, Key to species of Synchaeta hnown from Australia ! Uneus of trophus with one main tooth, fo accessory teeth , rome, 8 Uncus ‘with main ‘and accessory teeth. sce. a 3/1). Lateral antennae in posterior third of hady.... 3 Laterul antennae nsar base of foot ae, __S. tremula (Moller) 3(2). Marked constriction below elliary aurieles Fig, 83a) . .. S. lakawirciana 1 paar No obvions constriction, , Body cylindrical; auricles small: ‘foes 9-10. yim; ai with 4-5 robust teeth, , 8. kivina Hood Body conform, auricles not small; toes 5 any uncy with 5-8 (cath. een Apical field tlaty uncus 6-8 waithed..-. 0002s se ..) 8S) eblonga Ehrenberg Apical [i field domed; uncus 5-4 toothed, : 5, tiroralis Rousselet Two large ciliated tentacles in apical field... ,..-- oo. 8. peerinata Farentery Apical field smooth or willt ciliated humps... .. ‘Trunk (nedially constricted, elongated (Fig, §:2a); BL PAM pm. occas S& grandis Zachtarias No obvious constriction, trunk convex at sides; BL to 320 pm -. BL <200 un: foat elongated. BL 200-315 pm: foot not elongated. cee SS stylata Wiernyyska BL > 150. am; fort and toes us Fir, 12:3, .-..- torch a tact trem gefose ps wth &. tangipes Gosse BL isn um, foot and wes ay Fig Md... ai ete w paetis alleen lec betlobe aloe: itll nn boost S. jal sp. “nv 43). St4), ee cs 6(1), (8) sb bebe 87). O18). Synchaeta baltica Rheenberg FIG, 9:3} Synchiger buttied Elivenberg, 1834, p, 220. Type localiry: (?Burape)- Description: Bell-shaped/canical; foot long, cy lindri- cal; trunk may be annulated in posterior; toes short; foot glunds short, indistinctly separaled; lateral apical sensillae on papillae; male not described; RE with projecting integument. TR not described. BL. 190-523 am. Distribution: Marine, estuarine, coastal \waters worldwide. Two records: one off Sydney (Whitclegge 1889) and Port Philip Bay (Evans 1951). Synchaeta grandis Zacharias FIG, 92 Spwrchacia grandis Zacharias, 1893, p. 23, Fig, 2 Type locality, Planer See, Germany. Description: Body very long, usually cylindrical behind medial constriction; colourless except for yellowish tint te-ciliary auricles and protruding apical field: foot and foot glands long; tocs very short; cye red or black, circular; ocsophagus very long, Trophi; wneus a broad plate with very fine denticles (Fig. 9:2c), dulerum long. slender; manubria with semicircular outer lamellac: male undesctibed,; SE, RE, ME with fine spines, BL.400-600 pam; head width 180-200 ym; SE & RE 80-92*70-80 pm; ME 56%54 pm. Distribution: Rare in our samples (5 of ca. 5000 to date): billabongs & mainstream R. Murray near Wodonga, Vic., Yamup Swamp, W.A. and & single record of a Syachaeta resembling & enandis from a humic stock dam-at Karanja, near Mc Field National ROTIFERA FROM AUSTRALIAN INLAND WATERS Ws Park, in Tasmania (28.14.87) (Koste ev al 1988): 10-16°C. pH 6.0-7,.6, DO. 10.7 tig 1!, 64-106 wS em |, S NTU. Synchaeta sp.nov. While # Visiting Researcher at Waikato University in May-June 1993, RUS chanced tpor the notes of the late ©, R, Russell held al the Canterbury Museum in Christehurch, NZ, Cecil Russell was the “Honorary Keeper of Rotifers” atv the Museum uncl his death in 1961, and had published extensively onthe N.Z. rotifers in-the period 1945-1961. with two papers: including Australian rotilers (Russell 1957, 1961), In one of his laboratory notebuoks, he listed “Svehaete n. spo frou a sumiple collected in Warragamba Dam, N.S.W,, by V. HL. Jolly, (W2 27.%,59). The brief description and pencil sketches. of animal and trophi in Russell's laboratory notebook (Fig. Vela. b) were not published prior to his death, and the taxon bad not been collected avin, We could nol recognise thas one ol the known Svichacta species reviewed by Koste (1978), Fig, sla, b are copied from ca. p. 49 of Russell's “Feb. | 1960 ~ Australian Roufers” notebook, and the description below from the following page. The text is verbatim, parentheses are used where a word is not clear and some punctuation has been inseried Riv. 92 1, Swreheera buftica Ehrenberg. dorsal, 2, 5. grandiy Zacharias: (a) dorsal, (b) encus and ramus; (e) ramus and fulcrum, (a) manubriuimn, Oyo views, 1, 2a aller Rousselet (1902); 2b after Stumberger (1979); 2c, datter Kutikova (1970)... Seale bars adults 50 jm, trophy lO wm, 126 R, J. SHIEL & W. KOSTE “Svachaeta nsp. Body conical, small; auricles small with weak cilia. Foot obsolete, toe single, with enigmatic dirty end, (immovable); antennae 4 short. Trophi with single tooth on each ramus, a modification of pectinata type having one tooth in each ramus; the rami are slightly triangular, Length of body 120-130 um. Length of toe 10 pm. Length of trophi 70-80 um. In the contracted animal has a marked curved anterior margin. This animal differs from other species of Synchaeta in its small size (these two words have been crossed oud) (unmistakable) toe, absence of foot, difference in trophi, particularly the plate.” No material could be found in the Russell slide collection held at the Canterbury Museum. To deter- mine the identity of this taxon, the assistance of Tsuyoshi Kobayashi at Australian Water Technologies, Science and Environment was sought. He found early collections (Nov. 1965) by Sydney Water Board from Warragamba Dam which contained four individuals of a Synchaeta, but Russell's species was not present. The species was therefore described as Incertae sedis in the first draft of this paper. Fortuitously, Fitzroy R. samples sent by Larelle Fabbro, from the University of Central Qld, arrived while the MS was with referees. The first specimens encountered (by RJS) were undoubtedly the same as those seen by Russell. Several were sent to WK, who verified that the taxon, although similar to S. longipes, appeared to be new. Synchaeta jollyi sp.nov. FIGS 10-12 Wpe locality: impoundment of Fitzroy R., near Ramsey Ck inlet, 70 km upstream of barrage at Rockhampton Qld, (approx. 23°03'S/150°00’E), 07.1.1993, Coll. L. Fabbro, Univ. Central Qld, Rockhampton. Holotype: Single female, mounted in’ glycerine- gelatine. South Australian Museum (SAM) V4244, Date and place of collection as above. Paratypes: Four females on slide V4245, SAM, Date and place of collection as above, Two slides, Synchaeta collection, MDFRC #4090, 30 ml plankton sample containing S$. jollyi from Fitzroy R., Qld, MDFRC #4090, Date and place of collection as above. Material examined: Ten females were examined and measured, Description: (from partially contracted individual). Small conical body; head slightly convex; auricles small, face forward, with slight lateral bulges in slightly contracted animal; body broad for two-thirds of length, tapers to rounded posterior (Fig. 10, 11:2a); single median crimson cerebral eye; lateral antennae at midline; distinctive long cylindrical foot, not retracted in preserved individuals, presumably not retractible. S. longipes has an elongated retractible foot, paired foot glands elongated, cylindrical; two minute toes, barely discernible. Trophi: large in relation to body; uncus single toothed (Fig. 12a) (cf. 8. pectinata); fulcrum straight, rodlike in anterior view, slightly curved in lateral view (Fig. 12b); manubria curved with hemispherical lamellae: male unknown; SE, RE, ME unknown. BL 107*7.8 pm; BW 9014.2 jum; F/T 23.6*4.5 wm; TR 70-82 pm (FU 54 pm, M 68 pun). Distribution: Only two localities known: type locality, the Fitzroy R. near Rockhampton, Qld., and Warra- gamba Dam (now L. Burragorang) N.S.W., one of Sydney's water-supply reservoirs. Probably more widespread. Etymology: This rotiter is named atter the late Dr Violet Hilary Jolly, one of Australasia’s first fresh-water ecologists, While with the Sydney Water Board in 1959, she collected the samples which ultimately led to the rediscovery of this species. Synachaeta lakowitziana Lucks FIG. 13:1 Syachaeta lakowitziana Lucks, 1930, p, 59, Figs A-F. Type locality: (?Europe). Description: Marked constriction in neck region noted in original description possibly artefact of cocaine narcotization and formalin preservation; plump elongate body; dorsally, shape of head pentagonal; lateral sensillae on short papillae; toes acute; vitellarium bilobed: foot glands small. Trophi: unci plates with 1-2 large dagger-like teeth and 6-7 accessory teeth (Fig. [3:lc); SE smooth, RE spinulate. Fig. 10: Syvachaera jollyi. sp.nov. Fitzroy R.. Qld. Coll. L. Fabbro, Uniy. Central Qld, Sony CVP-G700 video print. Scale bar = 50 pm. ROTIFERA FROM AUSTRALIAN INLAND WATERS 127 TL 350-300 pm; male HO pm; RE 72x64 (Fig. 13:1/) or 67x45 wm with 15 pm long spines (Fig. 13:1g). Distribution: In Europe. cold stenotherm in winter plankton of mountain lake hypolimnia and arctic waters. Three Australian localities known, bul in yiew of European habitat preferences of this species, all populations require detailed examination: two mainland rivers: Moorabool R, Vic. in 1954 (Berzins 1982), lower R. Murray in S.A. (Shiel & Koste 1985), and 1b Fig, IL, Syichaeta jollyi, sp.nov, 1, (a) contracted: (b) trophus. a humic roadside pool near L. Garcia in W. Tasmania (Koste er a/, 1988), 17.0-17.5°C. pH 3.1-7.0, 81-500 pS em!. <1-I50 NTU, Svichaeta littoraliy Rousselet FIG, 13:2 Syuchueta litteralis Rousselet. 1902, p. 398, Fig. 7:15. Type locality: (U.K.). Description: Resembles $8. oblonga, bul apical field more domed: two-part cerebral eyspot with stream of 2, (a) contracted, (b) trophus. | from original peace sketch by C.R. Russell, Canterbury Museum, Christchurch, N.%. Coll, Warragamba Dam. 27.X.59. V.H, Jolly, 2. Fitzroy &. specimen, Seale bars: adults 50 wm, trophi 10 am, 128 R. J, SHIEL & W. KOSTE red granules to large red cervical eyespot; pigment granules diffuse in cold period, increase in density in spring; foot trapezoid: toes very short. TR unde- scribed. TL 192-290 pm; toe S pm. Distribution: S. cf. littoralis was collected in a billa- bong at Wodonga, Vic. (winter) (Koste & Shiel 1980), 10.2°C, pH 7.2, DO 9.0 mg I', 154 pS cm!, 4 NTU. A few individuals were collected in L. Colongulac, Vic. (17.V.80). 13.0°C, no other ecological information. Synchaeta longipes Gosse FIG. 13:3 Synchaeta longipes Gosse, 1887, p. 5, Fig. 2:15. Type locality: * . . . near Dundee”. Lacustrine. Description: Broad, protruding triangular head with widely spaced ciliary auricles directed somewhat posteriorly; body broadest at level of lateral antennae; foot clearly demarcated from body, cylindrical, long, thin, with two small toes; cuticle transparent or with bluish tinge; mastax orange-red, occasionally with bluish flecks in trophus region. Trophus: unci with acute robust tooth; manubria slightly sigmoidal with small triangular alulae on proximal third of external margin; SE rounded ellipsoid, smooth shelled; RE with rodlets between shells; yellow RE contents contain red- orange oil droplets; male unknown. TL 164-204 wm: T 6-7 pm; SE 60X56 wm; RE 72-76 56-60 ym. Distribution: Probably pancontinental, not yet recorded from W.A. Rare, in billabongs and rivers, in winter- spring plankton of lower R. Murray, S.A. (Shiel ef al. 1982): 8.5-27.0°C, pH 6.2-8.5, DO 6.3-10.4, 27-400 1S em', <1-160 NTU. Synchaeta oblonga Ehrenberg FIG. 13:4 Synchaeta oblonga Ehrenberg, 1832, p. 135. Type locality: (Europe). Description: Variable morphology; trunk generally barrel-shaped, laterally convex, but may be bell-shaped or ovoid; cuticle with longitudinal striae, colourless or yellowish; foot conical, toes short, bulbous; eyespots of different size, generally separated, also with speckled pigment granules; some populations may have fused eyespots; dorsal antenna normal; lateral antennae minute. TR: unci 6-8 toothed, generally symmetrical with notch behind main tooth and second notch behind Fig. 12: Syachaeta jollyi sp.noy. Trophus (a) dorsal, (b) lateral. Sony CVP-G700 video prints. Scale bar = 1Oum. ROTIFERA FROM AUSTRALIAN INLAND. WATERS 129 group of accessory teeth (Fig, 13:4¢), Rami with rounded alulae; manubria with distal varlike flattening and semicircular alulae; SE carried only a short time; RE wath short spines on inner and outer shell; male known, TL 225-250 wm; male 95-102 wm; SE 62%58 unr, RE 56-64™.56-60 um. Distribution: In reservoirs, billabongs and rivers, must common of the smaller Syachaen species in our samples, offen with S. pectinata, NSW, Tas., Vic.. ao W.A, Probably more widely distributed im Australia than present limited records indicate. 9.0-23.0, pH 48-100, DO 6.2410 mg i’, 9-1650 pS cm', 2-150 NTU, Synachaela pectinara Ehrenberg FIG. 13:5 Synchacta pectinara Elrenborg, 1832, p. WS Type locality: (?Europe), Fig. B. 1, Synchaeta lackowitsitera’ Lacks: (a) dorsal, (6) irophus; (¢) uneus & ramus; (4) uncus, (@) manubrium; (fg) resting cogs. 2, §. litoralis Rousselet: dorsal. 3, 8. longipes-Gosse: dorsal. 4, 8. oblonga Ehrenberg: (a) dorsal, (h) trophus; (c) uncus & ramus, 5, Synchueta pectinata Ehvenberg. (a) dorsal; (bd) trophi; (c) uncus. Ia, d-g after Lucks (1930), 2, 3, da, Sa, d after Rousselet (1902); Tb, c, 4b, ¢ aller Stamberger (1979); Sc, d after Kutikova (1970), Scale bars: adults 50 pam, trophi 10 am, 130 R. J. SHIEL & W KOSTE ROTIFERA FROM AUSTRALIAN INLAND WATERS ba Deseripren: Two ciliated tentacles in apical field; foot short and wide; joc relatively small; fooc glands shorter than foot: eyespot dark red or purplish; lateral antennae ak beginning uf distal third of body. TR: unci platclike with gmoved facing margins, ram erescenlic. acute wt proxumal tips: fulcrum [ong, rodlike viewed dorsally. broader laterally; manubria rodlike with median shallow U-bend; broad crescentic lamellac along ca, % of outer manubrium (Fig. 13-Sb, d}; SE with oil droplets and gelatinous sheath; RE cither thin-shelled wath small hooks..or more robust and spiny. TI. 240-550 jam; toes 5-8 wim; male 160 um;TR 190 um, SE 75 xm; RE 80-90 pm, Dixtvitucion: Largest and most common Synchaeta in our samples; pancontinental in billabonys, stock dams. lakes and impoundments, also im lowland rivers. aulutnn-wanter occurrence in lower R. Murray (Shiel eral, 1982), 7,0-29,0°C, pH 3.9-8.7, DO 4.0-106 mg r, 9-1000 gS env! < 1-150 NTU, Syachaent srylara Wierzeiski FIG, 14:1 Syidhweta srplant Wierzeyski, 1893, p. Wd Tye localiry, Galicia, Polund. Description; Resembles S. lonipes; body tapers te base of foot, foot arises fram a broader base than in other species, (6 notas Clearly demyreated Irom trunk; toes very short; apical field mosily smooth. eyespot single, occasionally paired; lateral antennae in distal third of body; male known; SE and ME with relatively long, delicate bristles, RE with shorter bristles mito gelatinous sheath; TR: unci Gps curve tnwards, manubria lamellae margin serrated, TL 200-313 ym; FT 33-St) on, T 4 pm, Male 89 pum; SE 64-74x50 um; RE 80X68 am. Distribution: Uncommon in NSW, Qld, tl most common and perennial Syreherene in lowee R. Murray plankton. S.A. (Shiel er al, 1982), 8,0-27.0"C, pH 7.0-8.5, BO 70-118, 60-100 2S em! . 1-110 NTU. Synchaete tayina Hood FIG, 14:2 Aynehasta rivina Hood, 1893, p. 382, Fig. 17. Type localiry; (UK.). Description: Body almost cylindrical; auricles small; foot and toe short; vitellarium with 8-12 nuclei, foregut present; eyespots generally paired, occasionally single Jarger fused eyespot (may be -violet-red); lateral antennae deeply inserted: TR: unci with 4-5 rohust teeth; cami with triangular upcurving alulae: fulcrum todlike distally, laterally forms a steiate semicircle BL 176-254 jon; a 176 um specamen was 61 jan wide at the head with T 9-10 um; TR 56 j¢ FU 26 yum, R 38 pm, M 40 pm) Distribution: Rare, recorded from only nur localities: a flooded gravel pit nr Eildon, Vic., Cullen's L. and Little Coliban Res., Vic., single recom In lower R. Murray plankton, S.A. (spring) (Shiel er al, 1982) 180-23.0°C, pH 70-80, DO &3-11.0 mg |!, 70-602 aS em! 65 NTU. Syuehaera rremula (Miilter) FIG. 14:3 Worricella tremula Miller, 1786, p. 280, Fie. 614-7. Syacharia tremula: Bhrenberg 1832, p, 135. Type lovaliry, (Europe), Description: Bartel-to cup-shaped body, often yellow- ish in colour; cuticle with striae; Loes shurt, stoul; eyespuls sometimes with clusters of pigment granules, transitional forms with S, .chtenga are known, also seasonal variants and ecotypic morphs in brackish saline waters (Koste 19783); SE smooth-shelled, occasionally in gelatinous sheath; two RE forms; with fine bristles or with short spines. Male Known. TR: wuci asyormetric, each uncus with single main tooth, 4-6 accessory tecth, also smaller denticles, separated ly deep notches. Manubria slightly thickened medially. BI. 150-323 «m; 8-11 wm, Male HO wm: SE 93X76 yin. ME 62x51 jan; RE 75X68 ym- Mistribution: Rare. cleven records fram Barwon R,, Old and downstream Darling R.. N.SAWV., central Tasmania, Waranga Basin and upper Murray billabongs, Vic. 10.0-16.0°C, pH 4,9-9.2, DO §,-10.0 mg J! 19-355 pS cm, 4.0 NTU. Family Asplanchnidse Harring & Myers, 1926 Relatively large animals. (ta 2.5 mm); cuticle thin, transparent, but retains shape, which may be saccate, pear- or bartel-shaped, sometimes. with lateral protru- sions (cf. Fig. §7:3s), foot and toes are present in the swimming or creeping Harringia, rudimentary ih semiplanktonic Asplanchnopus, lost in Asplerecfertet, asi adaptation to a fully pelagic existence. Corona of Asplonchna-type (see Koste & Shiel 1987). All three genera have incudate trophi, that of Harringia does not have # suction function. Asplanchnopus and Asplanckna da not have intestine, cloaca or anus. Harringin is not Known from Australia, Two species of Atplanchnapus and seven of Asplanchie are known from Ausitalia; ome species, A. uxymmelnca, is endemic. Fig. 14) 1, Syehaete srylare Wierzejski: (a) dorsal; (b) trophus, (c) uncus & rons, 2, 5 tava Hood, dorsal, 3, 5. gremula Ehrenberg; (4) dorsul; (6) trophus; (c) uneus, ramus & manubrium; (d) uncus & ramus, (¢) uneus, 1a, 2, 3a, ¢ ofter Rousselet (1902); (b, ¢. Jb, d alter Stemberger (1979), Je after Kutikova (1970), Scale bars: adults 50 juin, trophi 10. am 32 K. 4. SHLEL & W. KOSTE Key to genera With rudimentry foot atid (wes 1 pttesmasppecas . Asplanchnopus De Gueme Foul and toes ubsent..... » Asplanchna Gosse, Asplanchnapus De Guerne. 1888 De Guerne IXBR, p. 57. Cuticle flexible, transparent; body saccate or pear- shaped, with or without protrusions: corona divided circumapical ciliary band; apical field with bundles of sensillae; one cerebral cyespor and two lateral ocelli on short papillae in circumapical band: paired dorsal antennae, retrocerebral organ «nd subcerebral glands small, vitellarium spherical, riboon- or horseshoe- shaped. with eightor many nuclei; foot short or long, x a ee Fig. 15. 1, Asplanchnopus hyalinus Harring: (a) lateral, (b) trophus. 2, A. multiceps Schrank: (a) lateral; (b) trophus. 3 unsegmented, annulated ot with a single distinct fp segment, toes conical, tiny or lamelliform: oviparous or viviparous. Key to species of Asplanchnopus known from Australia Dorsal antennae divided, widely separated; >50 pairs of protonephridia) flame cells; trophi >100 am... .. wtooeaelititeessss, (tips A. multiceps (Schrank) Dorsal antennae partly fused; 8-13 flame cells; trophus ARTE So steele sale oo leben A. hyalinus Harring Asplanchnepus hyalinus Harring, FIG. 15:1 Asplanchnupus hyalimes Harring, LU, p, 402, Fig. 32: 4 Asplanchna asymmetrica (Waste & Shiel). trophius. 4, A. brightwelli Gosse: (a) dorsal: (b) trophus; (¢) trophus, diagrammatic (up= apophysis on bulls of ramus; f= fulcrum; it= inner teeth on rami inner margins; la= tamella behind rami points: m= manubrium; r= ramus; rp= rami apices; sap— subapaphysis; sit= second ramus tooth; v,= Ist uncus; u,= 2nd ungus). latter Harting (1913), 2 ufler Weber (1898), 3\.4c after Shiel & Keste (1985), da after Hudson & Gosne’ (1886), 4h alter Haver (1952), KOTIPERA PROM AUSTRALIAN INLANID WATERS 133 Wype locatiiv: Foucmile Run. near Washington, B.C.. LSA Deseripnen: Body jnoderitely elongate, slight cons(riction between head aod trunk: foot ubout one- third body lengrh, segmented, distal jotot nice as long as proximal: toes lameildorm, similar length as first foot-joint; pedal glands as long as entire foot, corona with interrupted vireumapical hand, ventrally at mouth and laterally by two small papiiae bearing oeelli: cerebral eye present: mastax large, trophi incudare, distinguished tram ather species inthe family by lick Of inner teeth or reinforcing ribs; short oesophagas, large stomach with pale of gastiie glands opealiiye iato wnlenior region, etght pairs Ot protonephiridial Mame cells: ovary ribbon-shaped. “Oviparous, BL 520-660 jam: FT 90-120 pm, T 32-40 pan TR 63-75 jm. Disinlburion, Carnivore on sonall routers (e.g, Lecutie, Lepadella), Only four records, all Victoria: billabongs of Goulburn, Mitta Mitta and Murray. and Ovens R- mento L. Mulwatu: 15.0-17°C, pH 7.0-7.8, DO §.0-9.8 me b', 65 pS env! NTU, Ayplanchniopus nudticepy Schrank FIG, 15:2 Brachionuy mufiioeps. Schrank. 193, po 3th Pa. 32bh-18 Asplanchaopus vudticups: De Guerne IS88, p57, Wpe locality: (Germany). Deseripron: Saccute body. loot shurt, gradually menu into trunk; head margins sometimes redidish- voloured; vitellarium horseshoe-shaped with many! puch; 50 pairs ol protoncphridial fame cells: large bladder: TR: rani slender. apices occasionally slightly cleft, one cerebral eye. wo literal oeelliz RE yellow- coloured, spinulate. Viviparous. Male large with resnounts oF digestive tract, many Mamie cells. BL 445-1000 pron; male 400-500 pam: TR ty 190 am (PU 52 pm); RE 220-238 pm. Distribution; Carniwore on other rotifers and small microcrustacedns (Koste 1987). More widespread. than A, fiyatinus; N2SW.ONT., Qld, Vit: 10.0-28.0°C, pt 6.0-8.1. DO 69-130. 37-170 pS enr!, <1-24 NTU, Asplunehna Gosse ISSQ, po I Agplanctiut Gosse, Body. with thin transparent integument, saccile. tubular barrel) or bell-shaped. some species with lateral protrusions of the integument: corona an interrupted cilry wreath, upieal field large, more or less rounded; imestine and anus absent; ciliary bundles on relatively high papillae in apical field; one cerebral eye appended, to brain: lateral antennae large, at beginning of posterior third of body: dorsal antennae paired: TR incudate. horizontal in mustax with apices facing posteriorly, everted and extruded to sieze prey; oesophagus a wide extensible crop: kidney-shaped or spherical gastric ghind on oesuphiwus. For a review of research on Asplinehod, See Koste (L97Ru). Seven species of Asplanclii ate recorded Irom) Australia. Rey to species of Asplanchaw known from Australia I, Rami clearly usyitmetric under low rgnifieation, let rains With median inner (oath, laelin plate behine mms (ip, right rans wathoui either, 4 gaaviinetrce (Shel & ‘Kastet Rumi aymrivinie winder Tey Mga iCal ‘2 Vicellarium sphericsl ne PREIS. Pvae Vitellurtiim cibbar-lke- + a 32). Vitellarium with up to 8 nucle, rudimentary’ foul antds absent, fut pairs of protonepliridial Mane cells. - A, priodonia Gosse Vitelberimur with 12-15 nuclei; foorghinds present; 2-40 flume cells, oA hericki De Guerne Trophus without upaphyses, constunt 16 Aumne cells A, geal [De Guerne) Trophus with robust dupoplyses: 1-100 fame ceths +2) a ee Sis}, Rami inner margin wiltt distinet, darge towit tf Tnnet mirein Wwelh abscut oF rudinentury - : A. internidia Hudson Broad lamellae betind nana apices, Which are syiimetricul, weute: ca 32 suchen in vileHarum, 10-20 Name vetlsy resting exe with vesicular structure ; ooh Jjnghovelli (Crosse) Lamellue absent. apivey ay meet lofi bifureyte. rwht single > 50 queler im vilellariom:, 40-100 Mame eells: RE with pleated outer shell, OA), JA sieboldi (Leydig) Asplanchna asymmetrica Siiel & Koste comb. nov. FIG, 15:3 Asplanchria brivhiweltiisyminetriva Shel & Koste. ass pp. Ql. Figs 4a. bh. Ieonetype: Shiel & Koste (Yss) Fig, 44, b Paratypes: South Australian Museium (SAM) V3945 Type lewality: Solomon Daim, Palm Istand, Old. Descriptions Body saceate; horseshoe-shaped vitellarium; TR asymmetrical: left rumus with short medial ianer tooth, sobterminal lamella and. second umeus asin ad, brightwell’, nght ramus with gone of (hese. more (tapered anc arehed than that of A. brivhiwelli. BE, (4) dorsal: (b) traphns. 5,40 sieholedi thepelte: (a) dorgal; (b) trophus, lit after Wang (1961). Ib after De Beauchamp (195)), 2 after Wulfert (941), du, Sa after Hudson Gosse (886), 4b afler Haver (1952), 5b atier Hauer (1937), Seale bars: adults 30 pin, troaph 10 per 136 R, J. SHIEL & W. KOSTE %1.00K "30. Gum he . I AsrPplanchna » Priodonta: Cc go3dea 2.8 kV %1.80K 16. 4¥m Fig. 18. 1, Asplanchna brightwelli Gosse: trophus, ventral. 2, ophus, dorsal. 3, detail of tooth on inner ramus. 4, 4. priodonta Gosse: trophus, dorsal. Scanning electron micrographs, Kodak T-Max, Hitachi SEM, University of Waikato, Hamilton, N.Z. ROTIFERA FROM AUSTRALIAN INLAND WATERS 137 rami tips resemble those of A, srebolkdi, albcil less robust and Jess noticeable biliircale; well-developed apophyses; rami not lamellate, and no inner woth on scapus; RE with anastumosing ridyes. BL 580-900 pan: TR 104-156 pm; nrvale 20-439 piri, RE 136 ym, Distribution: in pelagial of ponds and lakes (Europe). Single eurly record trom Qld (Colledge (914), Not seen in Gir samples, needs verification. Cunmment; Some similarities with A. brighrwelli (apnphyses on (he rami) and A, gired/ (absent of reduced scapus teeth), however the specific distinction of this taxon was verified by the study of Gilbert ev al, (19795. Asplancina priuntonta Gusyse PIGS Wa, 17:4, 18:4 Asplanchna prindanta Gosse. (850, p 18, Figs 1, 2 Fywe tovality; Hyde Park. U.K. Descriptton. Body rounded io saceate, oRen with a single hump on one sidc at the posteriur; TR: distinctive sputulate proximal rami, with dentigulate inner Up (Fig. 16a, 18:4b); no tooth on scapus; rami external margins from hemispherical curve, tapering to-small, projecting (almost Tight angled} subapophysis; RE reported to be smooth-shelled. BL 254)-1500 jan, male 200-500 pm; Tr 60-80 pm; RE 127-150 ym. Distribution: Cosmopolitan, perennial in oligotrophic lakes, also in hrackish waler (Eurupe) Reasonably comnmen in bilabongs, reservoirs and rivers of eastern mainiant Australid, not secopded from Tasmaitia, N,T, or WA. 7,9-27.0°C, pH 74-82, DO 8.6-12.5 me |. 46-850 pS em!, pels bei Winkelothle. Arch. Fydrobicl. 56, IN-33% ____ (1961) Die Radertiere saunter Gewadsser der Dabener Heide I. Die Rormtorien des Presseler und ex Winkeliihler Teiches. [bid S8, 72-102 ZAcHARIAS. O. (893) Faunistische Und hrologieche Benbarhhongen am Gr Pliner Sec. Aeieh, Riel, O, Plt ¥, S82. XYSTRIDURA AND OTHER EARLY MIDDLE CAMBRIAN TRILOBITES FROM YAXIAN, HAINAN PROVINCE, CHINA BY R. J. SHIEL* & W. KOSTEF Summary Xystridurine trilobites are an important element of early Middle Cambrian faunas of Australia, China and Antarctica. An early Middle Cambrian trilobite fauna is described from the Damao phosphate mine of the Yaxian district, Hainan Province, People's Republic of China. These include Pagetia luoyacunensis sp. nov., Xystridura hainanensis Sun, 1963, X. yaxianensis Zhu & Lin, 1978, X. orientalis Zhu & Lin, 1978, Galahetes hainanensis Zhu & Lin, 1978 and Kootenia sp. The species of Xystridura are described in considerably more detail than previously possible due to the discovery of new material particularly pygidia. It is suggested that the three subgenera of Xystridura erected by Opik (1975) are of doubtful value. Galahetes pyrus Zhu & Lin, 1978 is placed in synonymy with G. hainanensis. KEY WORDS: Middle Cambrian trilobites, China, Australia, Xystridura, Galahetes. Thansacriony of the Rayel Sectery af S. Aust, 1993), U3), 141-182 XYSTRIDURA AND OTHER EARLY MIDDLE CAMBRIAN TRILOBITES FROM YAXIAN, HAINAN PROVINCE, CHINA by Lin TIAN-Rur* & J. B. JAGOt Sammary Lik, TIAN-RUr& Jaco, J, B, (1993) Xystridura and other early Middle Cambrian trilobites from Yaxian, Hainan Province, China. Trans, R. Soe. §, Aust, U7(3), IA1-152, 30 November, 1993. Xyatridurine trilobites arc an important clement of early Middle Cambrian faunas of Australia, China and Antarctica. Aq early Middle Cambrian trilobite fauna is described from the Damao phosphate otine of the Yaxian district, Hainan Province, People’s Republic of China, These include Paperia luoyacunensis sp, nov., Xystridura hatnanensis Sun, 1963, X_ yaxianensixs Zhu & Lin, 1978, X. orientalis Zhu & Lin, 1978, Galahetes hainanensis Zw & Lin, 1974 and Kootenie sp. The species of Xystriduraare described in considerably more detitil than previously possible due ip the discovery of new material pariculurly pygidia. Itis suggested that the three subgenera of Xverridura erected by Opik (1475) are-of doubtful value, Galaketes pyrus Zhu & Lin, 1978 is placed in synonymy with G, Aainunensis. Key Woros, Middle Cambrian trilobites, China, Australia, Xystridura, Galahetes. Introduction Xvrtridura is an important early Middle Cambrian trilobite genus Which is knowa from the Northern Territory (Opik 1975), Queensland (Whitehouse 1939: Opik 1975), South Australia (Gatehouse 1986), and western New South Wales (Wopfner 1966; Opik 1968, (975). Xvstridura templetonensis is one of the nominate species of the Xystridura templeronensis/Redlichia chinensiy Zone of the early Middle Cambrian of northern Australia. Xysrriduva was known only from Australia prior to its discovery on Hainan Island, Chena (Sun. 1963), It is now known from elsewhere in China fe.g. Zhou et al. 1982) as well as Antarctica (e.g, Palmer & Gatehouse. 1972). Opik (1975) described a second memiber of the Xystridurinae, Galaefes, Sram the early Middle Cambrian of Queensland; it has since been recorded from Hainan (Zhu & Lin 1978) and Gansui, northeast China (Zhou et al. 1982). McNamara (1981) discussed the evolution af the Middle Cambrian xystridurine trilobites from northern Australia. The purpose of this paper is to revise the work of Sun (1963) and Zhu & Lin (1978) and to describe the trilobites (including species of both Xystriduna and Galahetes) from the Xystndkra Zone of the early Middle Cambrian Damuo Formation from the Damao phoephate mine of the Yaxian district, Hainan Provirice, People’s Republic of China (Fig, 0), The area conbeins some of the must important phosphate deposits in China. As listed below the Damao Formation is richly = Degurtment of Earth Sciences, Nanjing University, Nanjing 210008, People’s brpweeiid of China. + Deparment of plicd Geology, Garrcll Schoo, Oniversity of South s Eeetralis, The Levels, South Australia, 5095, fossiliferuus and includes trilobites, bradoriids, brachiopods, bivalves, hyoliths, microfossils, etc- Data on the trilobites have been previously published by Sun (1963) and Zhu & Lin (1978), The present material, which was collected by Lin Tian-Rui in 1987, 1988 and 199) provides extra spécimens which give a better Understanding to the previously described species, In particular, it has allowed the pygidia of the various species of Xysiridura to be odentified, which is imporlant because the pygidium provides one of the key fealures in separating the different species of Aystridura, In addition this paper provides stratigraphic information onthe distribution of trilobites which was not given in the papers of Sun (1963) and Zhu & Lin (1978). The stratigraphic section at Damao as described by Zhang Niguang (1986), and revised in this paver, is piven in descending order as follows. Damao Formation (top and bottom not exposed) 12. Grey thin to medium-bedded siliceous rock intercalated with silty and muddy shale, conlainmg brachioped fragments. 12,1m ii. Grey and yellowish grey thin to medium-bedded micaceous siltstone iptercalaled with a few siliceous shale beds, yielding bradoriids: Houlongdongetla disuleata Lee, H. sphaerica Zhang, Ophiosema strumatum Zhang, 0. sicyedeum Zhang, Ophiosema (Sinophiosetma) chinense Zhang, O, (S.) transversun Zhang, OQ. (8.) deltuttum Zhang, O. (S.) paradaxuin Zhang, Guangdongeila obesu Zhang, and the braclnopod Lingulella lini Sun, 12m 10. Dark grey medium-bedded siliceous siltstone, bearing the trilobite Xjstridura sp. 6.1m 9. Greyish white and greyish yellow thits to mediuen- bedded siliceous siltstone intercalated with silty and 142 LIN TIAN-RUIN. & LB, JAGO Quatermary-Cretaceous sediments DS | Upper Cambtciun-thddle Sihurian seditnenrs jw | Middle Canibrian aéciiments 4 | Lower Cambrian sedimoma _& | Granite SF Ceemaco phosphate mine and fossil localiry —— Pauls Fig. 1, Looality map. In how A, B lidicates Beijing. muddy shales, containing the bradoriids Ophiosema subrotundan Zhang and Mannecasmia sp.; the trilobites Xystridura hainanensis Sun, X. yaxianensis Zhu & Lin, X. orenralis Zhu & Lin and Galahetes hainanensis Zhu & Lin; brachiopods Lingulella tangshihensiy Resser d& Endo and Acrothele sp. 25m 8. Greyish yellow siliceous shale intercalated with thin to mcdium-bedded siliceous siltstone and containing trilobiles Xyxtridura orientalis Zhu & Lin and Pagetia Iuoyacunensis-sp. nov. 5.3m Greyish while and grey medium-bedded siltstone and siliceous shale, containing the bradoriids: Houlonedongetla disutcata Lee, H. inflata Zhang, Yaxianelle sutcata Vin, ¥ punctata Zhang, Ophiosema strrumarum Zhang, QO. subrotundum Zhang, Indeta acuta Zhang. Guanedongella obese Zhang, Mannocoxsmia torquate Zhang, Plesidielymella sp., Braderia sp. \rilobites Xystridura hainanensis Sun, X, orientalis Zhu & Lin, Gulahetes hainanensis Zhuo & Lin, Kootenia sp. and Fagetia Inoyacunensis sp. nov.; brachiopods Lingulella tangshihensis Resser & Endo, and Homotreta sp. 5.210 =~ 6, Dark grey siliceous shale and phosphatic siliceous rock intercalated with dark grey phosphorite, containing the bradoriid Indota acuta Zhang; the trilobites Xystridiura hatnunensis Sun, X. orientalis Zhu & Lin, and Pagetia Inoyaetorensis sp. nov.9.6m Dark brows nodular manganese ore bed (thickness about 2m), with siliceous shale at he base, yielding brachiopod fragments. 2.4m 4. Greyoh white and dark grey medium to thick bedded siliceous siltstone and quartzose sandstone intercalated with dolomite and a few micaceous muddy shales, 42.2m 3. Greyish green and greenish red medium and thin bedded micuceous siltstonc intercalations. 55,.8m 2. Greyish white massive quurtzose sandstone and siliceous siltstone intercalated with a few medium to thick-bedded dolomite horizons. 8.3m 1, Greyish white massive medium-grained quartzose sandstone. 352m wv Ave and affinities of fauna The trilobites described herein comprise the following species Pagetia Ineyacunensis sp. TOV, XMstridura hainanensis Sun, X. yexianensis Zhu & Liv, X. orientalis Zhu & Lin, Galahetes hainanensis Zhu & Lin, and Kortenia sp. In-China Xystridura and Galahetes ave known from the Tianshan (Wang eral, 1985) and from Gansu (Zhou e af. 1982).. The aye here is equivalent to the Maozhuangian Stage of the North China biostnittigraphic scheme. In Austraba Xystriduca and Gulahetes are important members of the early Middle Cambrian fauna of Australia (Opik. 1975), The various species of Xytridura described by Opik (1975) are cither of Ordian or Templetonign age. In Australia Xystridura is known fromthe Northern Territory (Opik 1975), Queensland (Whitehouse. 1939; Opik 1975), wesiern New South Wales (Woptner 1966; Opik 1968, 1975) und from the Kalludeina Formation of the Warburton: Basin in northeastern South Australias (Gatehouse 1986). In Australia Ga/ahetes has been recorded from the Templetonian of the Duchess ace (Opik 1975). Xystridura has alsovbeen described from early Middle Cambrian rocks of Antarctica (Palmer & Gatehouse (972; Soloviev & Grikurov 1979). In summary members of the Xystriduridae are an important element of the very early Middle Cambrian faunas of Australia. Antarctica and China, The specimens described herem show some tectonic distortion. They are deposited al either the Nanjing Instiute of Geology and Palneontology, Academia Sinica (NIGP) or the Department of Earth Sciences, Nanjing University (NUESD), All figured specimens were whilened with magacsium oxida prior in photography. EARLY MIDDLE CAMBRIAN TRILODITES Ii Systemic descriptions Suborder KODISCINA Kobayashi, 1939 Family PAGE TIIDAE Kobayashi, 935 Genus PAGE TIA Walcott, M6 Svrionvins See Jell YTS, p, 30, Type Spevies, Pagetia hootes Waleott, 1916, py, 408, Ph, 7, Pigs 1, lat. Diagnosis: See Jel) WIS. p. 30. Pagetia lunyacunensis sp. nov, FIGS 2a-t. 6d. Dawsonia dawson Sun (963 p. G10, Pl 1, Pigs 7 Pa. Paxetia sp. Zu & Lin 1978 p. 439, Plt, Bip. 0. Material WW crianicia and nine pygidia. Enology: After the nearby town of Luoyacun. Holetipe: The cramilium (NUESD 210). Fig. 2b. is chosen us the holotype, Diagnosis: Meriber of Pageia with anteriorly tapered lubelfa, eranidial spine has Jeneth about two-thirds that ol glabella: shutlow trunsglabellar furrow close ty plibellar anterior; wide fixed cheeks; palpebral lobes eXtend fram opposite tainsalabellar furrow to opposite iintertor oF spine; well-developed palpebral furrows; short (say. strongly depressed preglabellar field; wide border; [6-20 vephulic border scrobicules stopr just short of margin. Pyzidiui with gently tapering wis Whigh Stops just short of posterior margin. Axis comprises four axial rings and terminus, Bach axial ring and terniims bears spine base. Pleural areas with (hree pairs of pleural lurrows: marrew border, Paintly: pustalose surface ornament. Dexseripiions Cranidiumn with anteriorly tapered glabetla. Jength about 0.6-0.63 that of cranidium, shatlow trinsghibelliir furrow placed close to well- rounded glabellaranterion Well-developed cranidial spine, length about we thins Ue of plibellia, Asxtal turrews deep, wides baculae absent, Wide fixed cheeks pulpebral lobes extend from. opposite transylabellur furrow to Opposite amerior Ob spine; well-developed palpebral furrows. narrow but distinet eye ridges meet wAtal Furrows just forwards of transelabellar furrow, Short, strongly-depressed preglabellar ticld: wide anterior border furrow not crossing lateral border. Wide border with 16-20 radial scrobicules which stop jst shart of margin, Wide. deep postenor border furrow Senmcirculir pygnhum, Axis tapers gradually reurwards; extends almost to posterioc margin. Axis comprises four axial rings and ternimus, On availuble specimens each axial ring bears spine base as docs tenmnitus, Semi-elliptical articulating half-ring, Pleural aureus contain three pairs of pleural furrows. Narrow shallow borer Turow. marrow border, Both eranidium id pygidium bave finely pustulose sunitce ornament. Discussion: Prior to the specimens described herein_ this species was Known from only the wo very poorly preserved crunicia figured in Sun (1963, PLL. Figs 7. Tay and Zhu & Lin (978, Ph 1, Fe. 1). Although somewhat distorted the available material indicates that it shoud be the basis of a new species. OF the Australian species of Pazera described hy Tell 1975), the new species 2 /ovcneris’s is prohubly closest WP howard Jel The cramdia of the twa species arc quite plinilur, although the cephalic border ol Juovacunensix is Wider than thar of diawerdi. The Pywicial axis of dveveewtensiy os wader Ui that ot Aowurdi, P lrovackuensis differs trom the type species ot fagetiu, Po bootes Wale (eg, see Rasett) 966, p 504, Pl. 59, Figs 1-13). in that PA heores basa wider and shallower preglabellar field. a longer and more slender crunidial spine, a4 narrower pyyidial axis with five axial rings plis a terminus bearing-a lone spine and a simeother pleural field. There are some sunilaciies between Po fvayacrnensis und P quebecensts Rusett (966, p. 306, PL SO Fivs 5-10) but PB faenvaeunensix ditfers from. Pqiehecensis. in having small palpebral lobes. i havin distinet baculac and in hot having pronounced spine bases on the pygidial axis. Order REDLICHIDA Richter. 1933 Suborder REDLICHUNA Harrington, ls Superfamily PARADONIDACE.A Hawle & Corda. 47 Family CENTROPLEURIDAE Angelin, [854 Genus XYSTRIDURA Whitehouse. 1936 Type Species: Xvstridura tenpleranensis (Chapraan) — Milesia lempletonensiy Chapnian, 1929, p, 214, PL 22, Fig. 19, refigured by Whitchouse (93%, Pl 2L. Fig, 10 (see also comments by Palmer & Calehouse 972, p, Ih, Diugnusis, See Opik 1975, p.20. AL. Discussion: Opik (1975) discussed Xyweridura in some detail, He suggested that there should be it least iree subgenera in Xwirhdard, we, NL iNyatridura), X (Polvdinetes) and X. (Inosacores) with the type of X. (XMstridura) being X. tenipletemensin. In his ditleresitial iugnosis Opik (1975, p. 31) states “In Jaesacares anid Polydiiotes the cephalic test ty strongly ornamented, the thorax expands rearwirds, the pyyietial axial lohe 1s fused with the thoras. and the pygidial margin is Visibly denticulate; furthermore in Polyedinetes whe pleural Ups are hispinose (sanple nm Nvstriddra and Inoxacdtes) and in lnesdecetes he anterior facial sutires are fused (funetional int Xyyrricdtera and Polyedinates) Ny should be nofed that species ingluded by Opik in X, (Xystridara) also show a thorax which expands rearwards, 0.8. N. milesé (see Opik 1975, Pl, 14, Pie. 2). and NX. curtert (see Opik 1975, Pl. 15, Fig, 1). The I44 LIN TIAN-RUI & J. B. JAGO EARLY MIDDLE CAMBRIAN TRILOBITES 4 present authors do not regard the caecal patterns Shown ur ibe species of Jeesacores and Polydinoiey as being of subpenetic taxonomic significance, This plus the variations allowed foc by Opik would suggest that his subpeneric groupings may be of doubtful value. Nysiridura Rainanensis Sun, W63 FIG, 3a-h Nertdura hainanensis Son, 1963 p. GOR, Pl 1, Figs 4d, Abu & Lins 978 p.dd0, Phd, Bigs 2,3 (ton da, 4b). Material. Twenty cranidia and eight pygidia. Lectorype (desiyaaled herein). The pygidium figured by San (1963, Pl. J, Figs 4. 4a) is selected us lectotype. Sun did not ereer a holotype. Diaxnosix: Species ol Xystridura in which glabella extends almost to anterior border furrow, glabella anlenor broadly rounded, Long palpebral lobes extend froin opposite anterior of occipital ring to just forwards of 4p furrows. Anterior end of palpebral lobe almost meets axial furrow, Pygidium with short axis: with four astal fines and a terminus, Two pairs of marginal spines; vety tony, slender anterior pair directed posterolateraly at an angle of abour45° second pair short. Deseriprion: Cranidium wide (tr), slightly convex, with a length about two-thirds the wadth between palpebral lobes, Glabella extends almost to amerior boarder fuerow, preglabellar field very short (nag.) but clearly separates preglabellar furraw from anterior border furrow; glabellar length about O.85 that of crinidum, Glabella has slight waist at Ip turrows: widest just forwards of 4p forrows; broadly rounded glubellar anterior. Narrow axial furrow of moderate depth. Three pairs of luteral glabellar furrows, Bagh Ip furrow. is directed slightly to posterior and extends whout 0.4 of distance aeross glabella; 2p furrows almost horizontal; 3p furrows slightly arcuate with convexity to-aoterior 3p furrows, slightly convex untenorly, set in from axial furrow and not connecting with i, Nurrow oceipital furtow deepest abaxially. Occipital Ting widest centrally; small median node. Long palpebral lobes extend from opposite anterior of Gcerpita) ring to just forwards of 3p furrows. Palpebral furrow very shallow, barely discernible on many specimens, Palpebril areas of tixigense wide and flat; at posterior end width 0.5-0.6 that of glabella: anterior end narrow where anterior of palpebral lobe almost Meets axial furrow, Anterior areas of fixigenue almost flat. Preoculur sections ol facul suture diverge markedly, Very short postocular sections of facial suture diverge slightly. Wide anterior border wath shallow terrace lines: narrow anterior berder tirrow Shallow posterior border furrow. Very narrow posterolateral border. Pygididm very wide (tn) With length about half width. Short axis extends just over ball length ol pygidium. Axis has four axial rims plus lerminus. Wide pleura) tield with in Jeast two pairs of pleural furrows und two pairs Ob interpleural lurrows. Two pairs of marginal spines, Anterior pair arise From first pleurae of pygadium; very long slender spines with wide base: directed posterolaterally at angle of ubout 45" Second pair short. poited with wide base. Ib ts possible that there isd very short third pair of spines hut this is difficult to determing on availuble miterial Discussion; The pygidia shown by “hu & Lin (978. Pl, |. Figs 4a, b) as belonging in X. hainanensis are now placed in_¥. orientalis. When compared wath the Australian species described hy Opik (1975) NV hainanensts is probably closest to X. fracte un that the cranidia are quite similar. The pygidia are similar in terms of lengths of pygidial axis but differ in the presence of the longer spines in_X. Aainantertsiy and in the nature of the pygidial pleural and interpleural furrows. The anterior pysidial spines ol faaimunensix ire longer than in any other species of Aystridara with the possible exception of X. gfe Palmer & Gatehouse, 1972, However, glacia has three relatively nartuw pygidial spines whereas Aainanensiy bis two spines, each with a large base, The anterior pygidial spine ol X. glacia at its distal end is directed approximately parallel to the axis; this is mot SU In XL Aainanersiy, even allowing for tectoniy distortion, AXvweriduna vaxianensis Zhu & Lin, (978 FIG. tae. Xywrridura yarianensis Zhu & Lin, 1978p, 440, PL 1, Figs 5-6, Nysrridura arientalis Zhu & Lin, 1978 p. 444. Pl. 1 Fig. § only. Material: One specimen comprising the cramidium, the right librigena, inmost of the thomex and the pygidiuimn: 14 cranidia and four pygidia, None is well preserved Holotype; The cranidium (NIGP 44903) figured by Zho & Lin (1978, Plt. Pig. 5) and refiwured here as Fig. 4u. Fiz, 2. af, Tagena leeyacunensis sp, nov. a. NUESD 207, eranidium »20; b, NUBSD 210, holotype cranidivn x202 ¢ NUBSD 212, cranidiini, »20; d, pygiiom, *20..c, NUESD 214. pygidium. x20; f cranidium (figured a5 Aayetia sp by Zu & Lin W978, Pld. Fig. 1). x20. gi, Keerna sp. g, NUBSD 218, partial pyeidium, x5) h, NIGP 44916, partial cranidium Cisured by “tu & Lin 1978 Pl 2, Pig. &); i, pygidium, NURSD 219, x# All specimens are internal moulds. 46 LIN TIAN-RUL & J, B. JAGO Fig. 3. Avstridura hainanensiy Sun, 1963. a. NUESD 220, eranidium, x4; b, NUESD 217, cranidium, x4: c, NUESD 222, cranidium, +4; d, NUESD 225, cranidium, x4; ¢, NUESD 228, pygidium, x2.5; f. NUESD 221, cranidium, x2: g, NUESD 223, cranidium. x3,5; h, NUESD 224, cranidium, x3, All specimens are internal moulds. EARLY MIDDLE CAMBRIAN TRILOBITES 147 Pig. 4. a-e Xyweridura yevranensiy Zhu & Lin, 1978. a, NUESD 229, almost complete specimen, rubber cast of external mould, x2) b, cranidium, rubber cast of external mould, x3; c. NUESD 234. cranidium. rubber cast of external mould, x3; d, NIGP, holotype cranidium (originally figured by Zhu & Lin 1978, Pl. 1, Fig. 5), internal mould, x2; e, NIGP 44906, pygidium, (originally figured by Zhu & Lin 1978, Pl. 1, Fig. 8 as belonging to X. orientalis), internal mould, x4, f, Kootenia sp.. NIGP 44917, pygidium. (originally figured by Zhu & Lin 1978, Pl. 2. Fig. 9), internal mould, x5. 14k LIN TIAN-RUIL & J. B, JAGO EARLY MIDDLE CAMBRIAN TRILOBITES 9 Diagnoses: Species of Xysrridura in which glabella extends tilmost to anterior border, Sharply rounded glabellar anterior. Short pygidial axis. Three pairs of pyzidial spines, Moderately long anterior pair with wide hase directed outwanls and backwards; second shorter pair extend inwards and backwards; third pair very short. Central posterior trargin of pygidium broadly curved to anterior. Description. Cephalon semi-circular. Glabella extends dimost fo anterior border furrow: tenerh about 0.85-0.9 that of cranidium. Glabella with slight waist at about Ip furrows; widest just anterior to 3p furrows. Glabellar anterior sharply rounded. Narrow axial furrow of moderate depih) Three pairs of fateral glubellar furrows. Each |p furtow dareeted slightly tn posterior: they almost ower abaxially, 2p furrws almost horizantals 3p furraws whieh do not meet the axial furrow slightly arcuate with Convexity lo anterior. Both palpebral and anterior sections of fixigenae almost flat. Narrow occipital furrow. Occipital ring widest abaxially; fam occipital node. Long palpebral lobes extend from opposite occipital ring just anteriorly of 3p furrows. Shallow palpebral fiirrow. Preocular sections of facial suture diverge markedly. Narrow shallow anterior border furrow: wide anterior border wilh funt terrace lines on some specimens. Shallow posterior border turrow, very narrow pasterolateral border. Poorly preserved libnigena with narrow shallaw horder furrow: wide border extends into eenal spine of indeterminate length. Thorax of 12 segments. Shallow axial furrows, Width of axis about 0.6 that of pleural regions, Axis paralle| sided from lirst to filth segment gradually narrowing posteriorly, Pleurae almost Mauls pleund furrows narrow ubaxially but continue as shallow furrow into well-developed posteriorly directed pleural spines, Available pygidia poorly preserved. Axis comprises four axial rings plus terminus. Wide pleural field with at least two pittrs of pleural furrows and two pairs of interpleural furrows. Wide Hat border with fajntly- developed terruve lines. Three pairs of rharginal spines. Anterior pair extending from first pleural segment of pyeidium; of moderate length with wide base and extending = posterolarerally, Second shorter pair extending postemumedially. Third pair very short; more ehange of slope in pygidial margin than distinet spines. Central posterior margin broudly curved to wnterior. Discussion: When compared with the species described by Opik. X. yentunensis is closest to X. templeromensis. However the cranium of yvevianensis has « ntore sharply rounded glabellar ameriov and longet palpebral lobes than rentpletanensis. X. yercianensiy differs tram X. hainanensisy in having a more sharply rounded glabellar anterior and a longer pyeidial axis. The pygidial spines in (he bwa species are quite different. The pygidinm (NIGP 44906) figured hore (Fi. 4e) as X. yaxianensis was originally assigned by Zhu & Lin (197K, Pl. 1, Fig 8) us Jf orievitalls, However, the discovery of the partially complete specimen figured as Fit. ht indicates that NIGP 44906 should be reassigned. Xvstridura drteatalis Zu & Lin, 78 FUG, Sa-e; FIC), Gish. tr Qwridura orientalis @tw & Line W978 p. 441, PL 1, Fig. 7. Pl. 2. Figs t-3. Marerial: One incomplete ecranidium wiih three incomplete thoracic segments, 15 incanyplete eranaia, four free cheeks and 12 incomplete pygidia Holotype: The eranidiun (NIGP 44905) figured by Zhu & Lin (978 (Pl, 1, Pie. 7) and refigured here as Fig. Sg. Diagnosis: Species al Xysrridura in whieh glabella extends almost co anterior border. Sharply rounded glabellar anterior, Short pyzidiu) axis comprising three axial rings plus terminus. Three pairs of pygidial spines, anterior pair long, wide and directed slightly laterally to slightly medially. second pair long, wide and directed very shehrly medially: third pair very short: posterior pygidial taargin minutely denticulate. Description: Glabella extends almost (o anterior border furrow; length about UY (hat of cranidium. Glubella has slight waist at Ip furrows. widest just anteriorly to 3p furrows. Glabellar anterior sharply rounded, Narrow shallow axial furrow. Three pairs of lateral glabellar furrows; Ip turrows directed posteramedially - 2p furrows directed slightly posteriurly: 3p furrows essentially elongated slightly anvettorly curved pits- Shallow occipital furrow deepest ahaxiully Decipital ring bears small low median node, Palpebral lobe extends from opposite anterior end of occipital ring to just anteriorly of 3p furrows, Very shallow palpebral furrows, Palpebral areas of fixigenae wide, Nar; narrow atwnterior Preoeular secuons of facial suture diverge markedly. Postocular sections of ficial suture nowhere well preserved. Wide anterior border with faint terrace Fi, 5, Aysiridura onentalis Zhu & Lin. 1978, a. NUESD 235, cranidium plus three anterior Uhoracic seyrnunts, internal mould, 2) bh, NUESD 238, pygidiuin, intemal mould, x4..c, NUESD 236, Jibrigena and partial cranidium. rubber cast of external tneuld. x2: d. NUESO 245, pveidlum, rubber vast of external mould, x2; e, NUEBSD, eranictium, intarnal nieuld teks S NIGP 44910, Whrigena, imernal mould, x2 Zhu & Lin 7H, Plt, Fw 8), internal mould, %3. 3, g, NIGP 44905, holotype eranidion (onyinally Figured by 150 LIN TIAN-RUI & J. B, JAGO EARLY MIDDLE CAMBRIAN TRILOBITES ba} lines; narrow anterior border furrow. Librigena has wide border with terrace lines, Border widens postenorly; extending into long genal spine. Pygidium with three axial rings plus terminus, Axis. length about 0.55 that of pygidium. Pleural field with two deep pleural furrows and two shallower interpleural furrows which extend almost all the way across. very wide pleural field, Pleural field has closely spaced terrace lines. Three pairs of pygidial spines; anterior pair long, extending posteriorly and slightly outwards to slightly inwards; second pair natrower and directed very stightly medially; thied pair very short; medially from the third paiy pygidial margin minutely denuculate; posterior margin deflected anteriorly at centre, Discussion: The cephalon ot X, orientalis is essentially the same as that of X. yarienensiy, However, the pygidia differ in that the anterior pair of spines of X, orientalis are much Jonget than.and have a broader base than those of X, yarienensts, The glabellar anterior of X. orienralis is more sharply rounded than that of X. kainanensis. The anterior pair of spines on the pygidium of X. Aainanensis is much longer than those of A. orteritalis. Genus GALAHETES Opik, 1975 Type Species. Galahetes fulcrasus Opik, 1975, p. 75, Pl. 16, Figs 4; Pl, 17; Pl. 18, Figs 1-3; Pl, 19, Figs 1-3; Pl. 20, Figs 1,2) text-figs 7c, 13. Diagnosis: See Opik 1975, 3. 75. Galahetes hainanensis Zhu & Lin, 1978 FIG, 6c,e.I,g. Galahetes hainanensiy Zha & Lin, 1978, ps. 44), Pl. 2, Fig. 4, Galahetes pyres Zhu & Lin, 1978, p. 442, Pl. 2, Figs 5-7. Meterial> Ten cranidta- Holotype: The cranidium (NIGP 44912) figured by Zhu & Lin 1978, Pl. 2, Fig. 4 and refigured here in Fig. Oc. Discuysion; The available specimens add nothing to the material figured in Zhu & Lin (1978). However, a re-examination of the matertal indicates that the cruntdia described by Zhu & Lin as Galahetes pyrux should be placed in the same species as that described us G. hainanensis and hence G. pyrus is a junior synonym of G. Aainanensis, The apparent differences between G. Rainanensts and G. pyrus are due to the effects of tectomc distortion. The holotype cranidium of G. hainanensis has been tectonically elongated while the cranidium figured as G. pyrusx by Zhu & Lin has been tectonically shortened. Order CORYNEXOCHIDA Kobayashi, 1935 Family DORYPYGIDAE Kohayashi, 1935 Genus KOOTENIA Walcott, 1889 Type Species: Barhyuriscus (Kootenia) davwsont Walcott, 1889, p, 446, Diagnosis: See Palmer 1968, p. 47; 1972, p, 38. Kantenia sp FIG. 2g-i, 46 Keotenia sp. Zhu & Lin, 1978 p. 442, Pl. 2, Figs 8,9. Material; Two incomplete cranidia and six incomplete pygidin, Remarks: The available material includes two pygidia which are better preserved than that figured in Zhu & Lin (1978, PL 2. Fig, 9). The pygidial axis comprises five axial rings and # terminus, The axis extends to the wide posterior border furrow. Narrow border. There appear to be al least five pairs of small border spines, There are four pairs of pleural furrows, Acknowledgments The uuthors wish to thank Dr Yu. Zi-yi (Nanjing Insutule of Geology and Palaeontology, Academia Sinica) for collecting some of the specimens, Mr Hu Shang-qing (Nanjing Institute of Geology and Palaeontology) and Mr Xu Fu-lin (Nanjing University) took the photographs. Bao Jin-song and Sun Xiao-wen provided valuable assistance in Adelaide, The University of South Australia provided financial support for Lin Tian-Rui during his visir to Australia. Fig. & a.b.h Xysrridura orientelis Zhv & Lin. 1978. a, NUESD 242. pygidium, x3; b, NUESD 241, pygidiumt, intemal mould, x3: h. NUESD 250, librigena, rubber cast of external mould, x2 ce, fg. Galahetes hainanensis Zhu & Lin, 1978. c, NIGP 44912, holotype cranidium of external mould, (originally figured in Zhu and Lin 1978, Pl. 2, Fig. 4), x3, ¢, NIGP 44914, cranidjum (originally figured in Zhu & Lin 1978, Pl 2, Fig, 6 as Gulahetes pyrus), x5; 1, NUESD 235, cranidiuin, internal mould, x3; 2g. NIGP 44915, cranidivin, rubber cast of external mould (originally figured in Zhu & Lin 197K, Pl 2, Fig. 7 as Galahetes pyrus), x4 d, Pagetia tunyacunensis sp, noy,, NUESD 208, pygidium, internal mould, x20). 152 LIN TIAN-RUI & J, B, JAGO References CHAPMAN, F. W. (1929) On some trilobites and brachiopods from the Mount Isa district, N-'W. Queensland. Proc. R. Soc. Vie. 41(2), 206-16. GateHouse, C. G. (986) The geology of the Warburton Basin in South Australia, Aust. J. Earth Sciences 33, 161-180. Jee, P. A. (1975) Australian Middle Cambrian Eodiscoids with a review of the Superfamily. Palaeontographica Abt. A 150, 1-97. McNamara, K. (1981) Paedomorphosis in Middle Cambrian xystridurine trilobites from northern Australia. Alcheringa , 5, 209-224. Orik, A. A. (1968) The Ordian Stage of the Cambnan and its Australian Metadoxididae. Bur. Miner. Resour. Aust. Bull, 92, 133-170. (1975) Templetonian and Ordian Xystridund trilobites of Australia, /bid. 121, Parmer, A. R, & GATEHOUSE, C, G. (1972) Early and Middle Cambrian trilobites from Antarctica. US. Geol. Survey Prof, Paper 456-D, 1-37. RASETTI, F. (1966) Revision of the North American Species of the Cambrian trilobite Genus Pagetia. J. Paleont. 40, 502-SIl. Socoviev, 1. A. & Grikurov. G. E. (1979) New findings about Cambrian trilobites in the Shackleton and Argentina Ranges. Antarktika 19, 54-73. Sun Yun-Cuu (1963) On the occurrence of Xystridura Fauna from Middle Cambrian of Hainan Island and_ its significance. Acta Palaeont. Sinica M1, 608-610. [In Chinese]. Watcott, C, D, (1889) Description of new genera and species of fossils from the Middle Cambrian. U.S. Nail Mus, Prac. 29, 1406. (1916) Cambrian trilubites, Smithsonian Misc. Colin 64, 303-456. WANG JING-BIN, CHENG SHOU-DE, XIANG LI WEN & ZHANG TAI-Ronc (1985) “Stratigraphy and trilobite faunas of the Cambrian in the western part of Northern Tianshan, Xinjiang)”. People’s Republic of China, Ministry of Geology and Mineral Resources, Geological Memairs Ser. 2, No. 4, Geological Publishing House, Beijing). 1-223 [Chinese text], 224-243 [English summary]. WHITEHOUSE, F, W, (1939) The Cambrian Faunas of North Eastern Australia, Part II, the polymerid trilobites. Mem. Qld, Mus, WU, 179-282. Worener, H. (1966) Cambro-Ordovician sediments from the northeastern margin of the Frome Embayment (Mt Arrowsmith, N.SW.)..2 Proc. R. Soc. N.S.W. 100, 163-179. ZHANG X1-GuaNG (1986) A comment on some Middle Cambrian bradoriids from Yaxian, Hainan Island, Guangdong. Acta Palaeont. Sinica 25, 63-76 [Chinese text], 73, 74 [English summary]. Zuou ZH QranG, Li Jin-Senc & Qu Xtn-Guo (1982) “Trilobita in Palaeontological Atlas of Northeast China, Shaanxi-Gansu-Ningxia Volume, Part 1, Precambrian and Early Paleazvic’, 215-294, 446-460. (Geological Publishing House, Beijing). [In Chinese], ZHU ZyAo-Linc & Lin Tran-Rut (1978) Some Middle Cambrian trilobites from Yaxian, Hainan Island. Acta Palaeont, Siniea 17, 439-442 [Chinese text], 443 [English summary]. TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. II7, PART 4 EDIACARAN CARBON ISOTOPE STRATIGRAPHY OF SOUTH AUSTRALIA - AN INITIAL STUDY BY S. D. PELL*}, D. M. MCKIRDY*, J. JANSYN & R. J. F. JENKINS* Summary Carbon isotopic data are presented for syndepositional marine carbonates in Ediacaran (late Neoproterozoic) sedimentary sequences from the eastern Officer Basin and northern Adelaide Fold Belt, South Australia. The sediments selected for whole-rock isotopic analysis contain in excess of 10% carbonate (mostly as calcite) and very little organic matter. Recently proposed lithostratigraphic and biostratigraphic correlations allow the construction of a composite '*C-curve which documents a pattern of pronounced secular variation in the isotopic signature of a terminal Proterozoic epicontinental sea (c. 590-545 Ma). Such isotopic data may reflect profound environmental changes immediately prior to the Precambrian-Cambrian transition and currently are being evaluated as a basis for worldwide chemostratigraphic correlation of sediments of this age. KEY WORDS: carbon isotope stratigraphy, marine carbonates, kerogen, chemostratigraphic correlation, Ediacaran, Officer Basin, Adelaide Fold Belt, Rodda Beds, Wonoka Formation, Billy Springs Formation Tearsecens of tie Royal Svelety of 3. Ausn 995). U4), 153 161 EDIACARAN CARBON ISOTOPE STRATIGRAPHY OF SOUTLL AUSTRALIA — AN INITIAL STUDY by S D. Pett’, D. M- Mckiroy’, J. JANSYN® & R. 1. F JENKINS" Sammary Pewt. §. D., McRuroy, D. M., Jansyn, J. & JENKINS. R. J. F., (1993) Ediacaran carbon isotope stratigraphy of South Australia — an imisial study, Trans: R. Soc. S. Aust, U4), 153-161, 30. November, (993. Carbon rsotnpic data are presesited for syadepasitional marine carbonates in Edidcuran (late Neoptera) sedimentary sequences Irani the caster Officer Basin and northern Adelaide Fold Belt, South Australia. The sediments selected for whole-rock isotopic analysis contain in excess of 10% carbonate (mostly as calcite) and very little organic matter Recemly proposed Ithostratigraphic and blosrrarigtaphic corretarions allow the constriction of a composite 68C-curve which documents a panem of pronounced secular variation an the Isotopic: signature ofa terminal Proterozuic epicontinental sea (c, 590545 Ma), Such isotopic data may reflect profound environmental changes immediately prior ta the Precambrian-Camorian transition and currently are being evaluated as a basis for worldwide chemostratigeaphic correlatiun of sediments of this age. Key Worps: carbo isotope siraligraphy, marine cargonates, kerogen, chomostratigraphic correlatiany Ediucaran, Officer Basin, Adelaide Pold Belt, Rodda Beds. Wonoka Formation, Billy Springs Fonmnistion Introduction Notwithstanding their perceived scarcity of fossil remuins, sedimentary rovks of Neoproterozuic age hold important clues to the foundations of the present biosphere and, in particular, Ihe urigins of megascupic life (Glaessner 1984; Jenkins 1991. 1992, 1993; Kaoll 199]; Knoll & Walter 1992). Worldwide multidisiplin- ary studies in stratigraphy, palseontology, biogeochem- istry and tectonics are now providing tantalising glimpses of the size of the crustal organic carbon reservoir during an incerval of enhanced global rifting and orogeny (Des Marais et al, 1992) and tunes of massive burial af organic matter in sediments flooring stagnant, stratified oceans (Donnelly et al, 1990). Sea levels Auctuated and thermohaline circulation was restored during two. or more ice ages (Jenkins I99I- Knoll 1991) involving Jow-latnude glaciation (Schinidt eral, 1991), Increasingly oxygenated surface warers (Knoll 199]: Derry ef af. 1992) inay be linked to the flourishing of giant microplankters (Zang & Walter 1989, 1992; Jenkins er al. 1992) whivh were a food source for the first, bizarre metazoans Jenkins 1992). The present study focusses on (wo thick, well preserved but widely separated Ediacaran (late Neoproterozoic) * Deparment of Geology and Geophysics, The Universicy af Adelaide, Adelaide, South Australia 5005 + Present address: Research School of Earth Sciences, The Australian Nazional University, G.P.O. Box 4, Canherrs, Australia Capital Territory 2601 We favour the designation Adelaide Fold Belt (e 9. Jenkins 1990) in preference to the older usage of “Adelaide Geosynctine’ because the genetic significance af the term ‘weosyncline” is urtclear with respect 10 the ensialic characteristics of the region of outcrop. Reid, # (1992) Edjacaran (Latest Proterozoic) strati- gruphig, isotopic and palacobivloyical studies in the Flinders Ranges B.Sc, Honours thesis, University of Adelaide, unpubl, tw sedimentary siccessions in South Australia (Fig. 1). One succession, located in the eastern Officer Basin, is largely subsurface bul has been penelraled by some sixteen stratigraphic and petroleum exploration wells from which core inaterial is available. Another is well exposed in the Flinders Ranges, part of the northern Adelaide Fold Belt'!. The latter area is the site of the nominated! Ediacaran siralotype Genkins 1981), The depositional histories of the: Flinders Ranges and eastern Officer Basin during Ediacaran time display stciking parallels. Northward thickening prisms of sediment accumulated in both arcas (Fig. 2), with southerly or centrally located shelfal regions of syn- depositional flexure zones trending more or less cast- wesl, and more rapidly subsidimg troughs positioned w the north (Preiss 1987; Haines 1990; Thomas 1990) Sukanta ef af. 199); Preiss & Kreig 1992). fm the Flinders Ranges the several kilometres of sediment present in the southern and central shelfal regions comprise, in upward succession, the Wonoka Fanration, Bonney Sandstone and Rawnsley Quarizite (Preiss 1987; Jenkins er al. 1992). Law inthe Rawnsley Quarizite, siltstones and thin sandstones of its Ediacara Member contain the well-known Ediacara fossil assemblage (Jenkins et al, 1983; Gehling 1988), The Bonney Sandstone and Rawnsley Quartzite together make up the Pound Subgraup (Jenkins 1975). North af the east-west trending hinge zone (Figs 1 & 2) the Wonoka Formation and Pound Subgroup increase in thickness. In the far northeastern part of the Ranges ac. 2.1 km-thick succession of siltstone with interbeds of silly sandstone and carbonate, collectively included in the Billy Springs Formation (Preiss [987), occupies a stratigraphic position above older parts of the Wonoka Formation and below a recently recognised condensed section of the Pound Subgroup (Reid 79927) We 1st restrict Ute usikge of the Billy Spnngs Formation to exclude these upper sandstones in which rare discoidal moulds, probably: representing metazoan remains, and a single fossil burrow have been observed. The Ediacuran sediments torm several depositional system tracts (van.der Borch etal. 1988; Christie-Blick eral, 990; DiBona er «/. 1990). The suggestion of glacigenic sediments within tbe lower member of the Billy Springs Formation (DiBona 1991) has. not been substantiated by the reporting of diagnostic characteristics such as faceted and striated clasts or till pellets. The Flinders Ranges underwent local extensional faulling during early Wonoka Formation time. ‘The subsequent Cambro-Ordovjcian Delamerian Orageny (Preiss 1987) shortened the sedimentary pile at several {hrust zones and tormed open to tight folds. In the eastern Officer Basin, the Rodda Beds (Brewer er al. 1987) include thin carbonates resembling the Wonoks Formation that extend across its southern stelfal region (Murnaroo, Platform: Thomas 1994); Sukanta et uf 1991). These carbonates lie above currelatives of the Bunyeroo Formation which include w Yi USGRAVE. avoo/Z) f, Sh, , yy “ 5. D PELL, D M, MCKIRDY, J, JANSYN & RJ, B JENKINS an important chronostratigraphic marker, the Acraman impact ejecta horizon (Wallace et al, 1989), tormed by the debris of a giant meteorite infall. ‘Ihe Wonoka- like carbonates of the Rodda Beds are host to large leiosphacrids. and process-bearing acritarchs, potentially of regional biosiratigraphic signilicance (Jenkins er al, 1992). Extensional faulting again characterises the start of the Ediacaran cycle which, in this instance, ts signalled by seismic reflector Fy (Thomas 1990; Sukanta ef a/. 1991). Farther north, in the Munyarai Trough (Fig. 1), presumed equivalents of the Wonoka Formation are represented by a prism of sediments as much as several kilometres. thick. Reflector F is an erosively downcutting surtace and marks the incision of large, canyon-like structures (Thomas 1990; Sukanta e¢ af. 1991) with basal pebbly beds and infilling siltstones that resemble the Billy Springs Formation. Subsequent reflectors or sequence boundaries. are speculatively equated with erosive events recorded in the Pound Subgroup, and the base of the Cambrian is a regional disconformity, surface & (Thomas 1990; Sukanta et al 1991). mm» Be Adelaide 3 Fold Belt at = 0 km 1000 ——st—S} v Fig. |. Map of Officer Basin, Adelaide Fold Belt and other ventral Australian basins contwining hick sequences of Ediacaras (ate Neoproterozoic) sedimentary rocks. Inset shows location of drillhole intersections in the Officer Basin, Specific outcrop lovalities: 1, Bunyeroo Gorge (site of Ediocaran type section) » 2, Mount Precling Syncline; 3, Black Range Springs. EDIACARAN CARBON ISOTOPE STRATIGRAPHY OF SCUTH AUSTRALIA 4 Materials and Methods A generalised synthesis of these stratigraphies (Fig, 2) prowides the regional framework within which we place our carbon isotopic measurements (6"C,..4 of whole-rock carbonate: Kaufman eral 1991) made on cores from four drillholes in the Officer Basin and on outerop specimens from surface sections in the ventral and northeastern Flinders Ranges (Hig. 3). The samples selected for isotopic analysis are fine-grained sedimentary rocks containing 12-12% carbonate carbon (CC; equivalent to 10-100% CaCO,). Their carbonate mineralogy as determined by XRD analysis is miaed (culcwe, dolomite), although culeite is the dominant component in more than 75% of the samples examined, Carbon iwotople composition in contiguous strala appears to be independent of mineralogy, sugvesting that dolomitisation was early and took place in a closed diagenetic system. ‘Total organic carbon (TOC) contents of the dolomicrites and calcareous and dolomitic siltstones from the Officer Basin) ure negligible to. moderate (0,03-0,87 %) and the dispersed organic matter is of relatively low thermal nmuaturity (kerogen atomic H/C = 0.80-L,15; Rock-Eval T= 419-~443°C. McKirdy er a7. 1992), Weak metanior- phism in the central Flinders Ranges (Lernon ef af. 1992) increases to greenschist facies (biotite zone) in (he northeastern region (McKirdy er al. (975; Preiss 1987). Fine-grained Jithofacies in both the Wonoka Formation and Billy Springs Formation ure organically very lean (TOC mostly <0.1%), In the Mount Freeling. Syneline there is some evidence of metamorphic loss of organic carbon from these units.. Results and Discussion ‘The oldest Ediacaran sediments in South Australia, offshore arenites and siltstones and a storm-dominated succession of ramp carbonates deposited below fairweather wave base, are represented by the lower to mid-Wonoka Formation in Bunyeroo Gorge (Hames 1990). Shallow-water (lagoongl to supratidal) carbonates of the upper Wonoka Formation were sampled here and at two other localities in the central and northern Flinders Ranges (Fig. 3), The resulting composite carbon isotope stratigraphy (Fig. 4) displays the following featurés: a low negative value (46°C = -2.4°%/nc) in the basal dolomicrite horizon (Unit 1); a shift to strongly negative 62°C values (-6 to -8.5 “/oz) which persist through 360 m of outer to middle shelf limestones (Units 3-7); and a steady positive trend From midway through Unit 7 (-4 foo} into Unit 9 (+4,5/on), a w 5 i i z ra 9 FLINDERS RANGES sg = B ec Zz S 3 g z :" va a NOMINAL SURFACE® oF TERMINAL * PROTEROZOIC EROSION © rs a vey QUARTZITE ; eoune __ mnie roan suaGRoUP > — a NO BILLY ’, " / REMAINING ar RECORD SPRINGS ae | FORMATION / \ thannprnnnnwe’s jaa — 7 ORNATE ! ‘ re i oan pane 8 5 ¥ p " BASIN 5 5 = 3 3 z a = 2 > g 2 & 8 5 3 PESEN SE> uw > BSF sunsace oe = NW ~.. 7 Bedoing . i Sm = a ay wo Sequence boundary 5 Ay (mainly paraconformable) a ‘oy Erosional sequence MURNAROO 82 boundary’ PLATFORM g EASTERN OFFICER BASIN MUNYARAI TROUGH 10 20 30 40 50 horizontal scale km Fig, 2 Comparison of generalised seismic section of the later Neoproterozoic of the eastern Officer Basin (based on the inlerpretations Gf Thomas 1990 and Sukanta er a/, 1991) and the correlative stratigraphy of the Flinders Ranges. IS6 Moderately positive 6°C values (+4 to +6'V/s0) are maintained through the Wonoka-like carbonates in the Officer Basin at Mutnaroo-1 and Observatory Hill-1 (Fig. 3). This interval includes giant process-bearing acritarchs in the Officer Basin Jenkins et al. 1992). Probable early indications of metazoans in the Flinders Ranges accur in Unit 10 of the Wonoka Formation (Jenkins 1984, 1993, Haines 1990). Sequence strati- graphic data from the northern Gammon Ranges (DiBona et al. 1990) suggest that Unit 10 predates the Billy Springs Formation. Our next segment of the Ediacaran carbon isotope profile comes from the casterm Mount Freeling Syncline where carbonates at the top of the lower Urlwin, B. (1992) Carbon isotope stratigraphy of the late Proterozoic Wonoka Formation of the Adelaide Fold Belt: diagenetic assessment and interpretation of isotopic signature, and correlations with previgusly measured isotopic curves. B.Sc. Honours thesis, University of Adelaide, unpubl. Ungo olya:i OFFICER BASIN ossenyarony) | HILE RM) © eral © | E 14394,2 % Meramangye-1 RODDA BEDS 7 = stromatolites Central Flinders Ranges | Condansicd Sedimentation 5. D. PELL, D, M. MCKIRDY, J. JANSYN & R. J, FE JENKINS member of the Billy Springs Formation. are exposed (Fig, 3). Here the record is fragmentary and incom- plete, largely because of structural complexity. and the scarcity of suitable carbonate lithologies. Nevertheless, the available data appear to delineate a second major positive excursion, 8°C values rise from -0.7 to +7/oo over a section of several hundred metres and (with one exception: +2.7°/co) remain relatively constant through the prominently outcropping carbonate that forms the ridge of Wildman Blu/f. We consider that the uppermost Wonoka Formation (Unit ll) may be coeval with some part of the Billy Springs Formation. Unit 11 at Black Range Springs (°C = +6.6/o0: Unwin 19923) also records a positive isotopic excursion. The more muted expression of this excursion in the same unit.at Bunyeroo Gorge (6°C = +2°/co: Fig. 3) may reflect increased oxidation and recycling of "C- depleted organic matter in restricted, shallow, lagoonaJ environments which contrast with the presumed open-matine conditions farther north (Haincs 1990). FLINDERS RANGES Mit Fraeling Syncling (Bast) ie . & be z 3 a= 5a ws w ae =e ge > a a Mt Frealing. Syncline jt Allachthonous 2024 68 §Mox, BOewte2eneaaaes ox, Fig. 3. Carbon isotope profiles of carbonate-bearing Ediacaran sequences in four drillholes from the eastern Officer Basin and three surface sections from the Flinders Ranges. Key: o 1-2% carbonate carbon; ¢ >2% carbonate carbon. EDIACARAN CARBON ISOTOPE STRATIGRAPHY OF SOUTH AUSTRALIA 157 OFFICER BASIN FLINDERS RANGES Ungoolya-1 Oo Q | coer 2 | Ol= 1281,2 i a < PARACHILNA| «5 > ro SEISMIC FORMATION 8 a: o|' gs oy 4s 1 fe Munyarai-i] oO & Observatory Hill-1 | 2 us z _ 1699 $ 7 6 \ = | \ 2896.6 pot EXTENSIONAL ; | ! EXTENSION FAULTING \ : | iA vit, + UNIT Fs SEQUENCE TECTONIC MOVEMENT BOUNDARY 864202468 BUNYEROO }=_ 816 % ADELAIDEAN MARINOAN Xx ronmarion 4 ACRAMAN IMPACT EJECTA REGIONAL SEQUENCE BOUNDARY Fig. 4. Summary of Ediacaran carbon tsotope stratigraphy of the Officer Basin and Flinders Ranges, South Australia. iss 8. DL PELL, DB. Mo McKIRDY. J. IANSYN & R. J. F JENKINS tn the central Flinders Ranges, sediments of mid and lale Ediwcanin uge (Bonney Sandstone and Rawnsley Quartzite) are devoid of carbonate. Fortunately, in the eastern Ofliccr Basin sugeesied currebsuves in the upper Rodda Beds (Sukanta ev al, (991; Jenkins et al. 1992) laclude calcareous and dolornitic siltstones (Pell 99%). Their detailed carbun isotope stratigraphy al Ungoolya-l and Meramangye-I is displayed in Fig. 3, Sciliments from) the 1720-1990 in depute interval of Ungoolya-1 are for the most part curbenate-poor (CC = 05-71%), The 68C values of such carbonate (mostly in the range -O.8 to 4f)25%s0) imust be anterpreted with caution (Lambert era. 987; Fairchild eral, \990) Kerogens tsulated frony the lower half of the succession provide a very coherent &"E signal (-28 to -2e°/s7 Moreoever. the moan difference between fhe §[C values of carbonate and organic carbou in these sediments (A"C = 25‘Yoc) coincides with the secund of twee modes in the frequency distribution of APC values for carbonaye-rich Rodda Beds (discusser bebow), We terefore suzwest thal the two tone positive 6"C values al 9646 m and 1304.5 m in Unguolya-l (+3.8 er aud 4.2 Yon: Fig 3) do aciuilly recur the jsotepic compasition of primary marine carbonate deposi a this. itd Edincaran highstand tract (Jenkins et ah 1992}, and thal shifty ia more negative values (in lhe range +0.2 to 0.8 S/on) represent diagenti¢ carbone resulting, from bacterial oxidation of organic matter, These slightly cibeureous andl dolomitic silistones are not notably pyritic. precluding bacterial sulphate reduction as a significam additional saurce of biogenic CO,. Equivalent late Rodda Beds sections at Ungoolya-t wid Meramaagye-l exhib similar carbonate 3’C trends (Pre. 3). In each instance many of the samples wnalysed are reasonably carbonate-rich (CC >2%) and therefore hkely In retain primary marine isotopic Signalutes, A sharp negative excursion from +1 to 1.5/0 al the tase Of the Rodda Beds penetrated by Meramangye-| appears to record detail not dncumented by tur sampling of the equivalent interval inunediaccly above the E, seismue reflector in Ungoulya-] (Fig. 3). The cnsuing steady rewem to positive 4"C values (peaking al +27 and +1.9°/m, respectively] in both well sections 7s followed by a second negative cacursion, although crostve loss of section attenuates this trend at Ungoolya-1_ Carbonate in the highest part of the Rodda Beds section at Ungoolya- is for the most part characicrised by moderately: positive 5°C values (+0.9 te +1.7 eo: Rig 3). Two anonijalously negative * Pell, 5. DB. (1989) Stable isotupe composition of onganic inatter and op-existine carbonate in the Late Precambnan af the Officer Basin: stratigraphic relatiomstvipys oritlh neighbouring basins and environmental significance. B Sc. Honoris thesis, University of Adelaide, mpubl. walties (-28 and -3,6 Yoo} usa are associnted with strata tw Which carbonate contents are relatively low wn hence may not be wholly of primary ungia The extent to which pnmary marine carbon isolopic compesitions hive been retained lFroughoul the Rodda Beds ts difficwL i asceetaum. One stuutegy adopted by previcus Workers td overcunié the problem of dui genetic alteration involves 6!3C measurement on carbonate and organic curbon in the same sample (Knoll et al. 1986). Primary processes (notably synthesis of organic matter by macioe aucotrophs aml Precipitation of cuthonute m equilibrium with CO, dissolyed in the water columm) tend to maintain a eonstuml isolupie dilference between carbonate anal organic carbon, whereas various secundury processes may independemly alter 65°C in either carbon phase (MckKirdy & Powell 1974, lrwinet al 1977. Hayes et af. 1983), Thermal maturation of the Rodda Beds bys not been severe enuugh to significantly miodify te primary isotopic composition of their kerogen. Even so, far the carbonate-rich strata examined. AVC ranges from ~24/oeto ~36°/ou; and the [recency distribution of these AVC values is distinctly bimodal (Fig, 5). Wt is likely that the damunant mode at 34 Yoo represents the primary isotopic separution between cuexisiing Ediacaran organic and inorganic carbon in the Officer Basin, This contrasts with the reported modal APC value of 28.5 “Yue for Upper Riplear and incomplete Vendian sections iw Svalbard and Easi Greenland (Knoll ep al. 1986). ue recoyuised cause of enhanced ismopic Fractionation between carbonate and (reaiic cavbon is Hiethandgerie diagenesis (irwiit eral 977). An altemative explanation is suggested by the results of recent stodies which have revealed that Neopraterozaic marine sediments containing well- preserved kerogens exhibit wide secular Nuctuutions in APC (Strauss ev al, 1992, Des Marais et a/, 1992). In particular. (he degree of fractionation hetween boexisting carbonate and aiganic carbon recorded in such sediments increases signiticamly helwesn AX aml 600 Ma ago, at a (ime When the atmospheric CO,/O. ratio was in decline privr jo the Varanger ghaciatiim, Oar ANC Vata are tn excellent agreement with (he terininal Proterozgic (-~-60G Ma) separation nf aC. and 6, reported by Des Manus et al (YY2, figs | & 2), Presumably there Was near optinumn isotopic fractionation during photosynthetic fixation af dissolved carbon dioxide in the Exhacaran seaway ol the Officer Basin (cf. Strauss er al. 1992) The APC distribution for Ediacaran (Rows Beds} carbonate — otganic carbon pairs has. a secondary mode at 25 Yoo (Fig_ 55. This manifestation af a low fractionation of the C isotopes tends i be associated with negative carbonate 6"C yalues. and hence probably reflects appreciable diagenetic precipitation of carbanate it response te in situ bacterial oxidation of organic inaterial with an isowpic composition (6C EDIACARAN CARBON ISOTOPE STRATIGRAPHY OF SOUTH AUSTRALIA = -23 to -28 ou) typical of planktic organte carbon from ocean waters with surface temperatures in the Tange of O-10°C (Sackel et al, 1974). Tt is CO, (or bicarbonate) availability, not temperature per ve, which ultimately controls the extent of discrimination against 5C during photosynthetic carbon fixation (Anderson & Arthur 1983). This in turn is a function of several variables, including pCO, in the external environment (Arthur et al, 1985), the development and operation of biochemical pumps for bicarbonate and carbon dioxide (Badger 1987); and evolutionary changes tothe kinetic properties of the principal CO_-fixing enzyme in prokaryotic and eukaryotic photoautotrophs (Badger & Andrews 1987), All these factors were in a state of considerable flux during the Neoproterozoic (Strauss eral. 1992) which witnessed iner alia the emergence and decline of giant phytoplankters (Zang & Walter 1989; Jenkins et a/, 1992; Knoll & Walter 1992), Hence, it is perhaps not surprising that contrasting microfossit assemblages (one dominated by spitiose acritarchs and the other by filamentous forms) from Widely separate stratigraphic levels of the Rodda Beds (Jenkins et al. 1992) should display markedly different §°C organic signatures (-29 to -31°/o0 and -25 to 26 “/ou, respectively). Conclusions Our composite carbon isotope trend (Fig. 4) shows that for several periods of its history the Ediacaran bal Ayliffe, D. J. (1992) Geological setting of the Late Prowrezoic Wonoka Formation at Pichi Richi Pass, southern Flinders Ranges: geochemical, stable isotope and diagenetic analysis. B.Sc. Honours thesis, University of Adelaide, unpub), No, of samples WK 23 15S scaway in South Australia was precipitating isotopically heavy primary carbonate (6°C = +0,5 to +6°/ov).. Similar "C-enriched carbonates have been docu- mented in other Neoproterozoic successions (e.g. Knoll eral. 1986; Aharon et al. 1987, Lambert er al, 1987; Brasier ef af. 1990):, Fairchild er al. 1990: Kaufman et al, 1991; Magaritz er al. 1991) and attributed to the maintenance of abnormally high rates of burial of organic mattet.in shallow cpicontinental seas at low palaeulutitudes (Donnelly et ad. 1990; Derry e7 al. 1992), Gonversely, the prominent negative excursion recorded in the Wonoka Formation may reflect recycling of organic matter from planktic and benthic primary: producers (Jenkins ef al, 1992) by (?)metazoan grazers and acrobic bacteria inhabiting the oxic water column of a storm-prone, deep water, ramp environment, Ongoing studies of this lithological unit at localities up to 200 km apart suggest that the markedly negative 65C values (-6 [0 -8°/o0) which characterise much of the older’ Wonoka Formation ate a basinwide phenomenon (Urlwin er af, 1993); and we draw attention to the unexcepuional isotopic signals recorded from Unit | and through Units 7, 8 and 9, A careful trace element and isotopic study by Ayliffe (19925) demonstrated that part of the Wonoka Formation filling a deep ‘submarine’ canyon ( >700 m of erosional relief) and displaying the same regional low 6"C signal (Urlwin e¢ al, 1993, Fig. 1) has never been in contact with meteoric waters. Morever, the ‘pristine’ (low) strontium isotope ratio of the canyon fill is strong evidence that there has been negligible alteration by epigenetic fluids. Sedimentological characteristics of the older Wonoka Formation are consistent with the precipitation of its calejum carbonate as whitings (Haines 1990), perhaps implying a low pCO, in the surface waters of the ambient sea, A13C Fig. 3 5 Frequency, distribulidn of APC valves in carbomale-rich sirata of the Rodda Beds, cusiémm Offiver Basin. Note: Coat —a ty {after Knoll ef al. 1986). (tH) 5. D. PELL, DM. McKIRDY, J. JANSYN & R.J, F JENKINS The uniformiy sparse TOC content of these sediments ulso is consistent with a lowered prinvary productivity, ind hence it is conceivuble thar their carbin isotope signature fags the developrient oF a ‘Strangelove ocean (see Hsu eral. 1985) in which biological productivity is totally pousoved by abnormally high concentrations of dissolved ©), (e.g. Jenkins 1991). Derry ev ul. (1992) concluded that, at about this time, the biosphere first entered [ls oxygenic Phanerozoic mode following a Garge and rapid flux of O, out of the carbon cycle. The younger part of our Ediacaran 6°C profile (Fig. 4) broadly paralicls the Isotope curve published by Lamben ev af. (1987) for the late Neopruterozaic earbonate-rich Doushantou and Dengying Formations of the Yangtze Platform, China, In both instances, isotopically heayy carbonate (6°C = +4 to +7 Yoc) is followed by a trend towands hghter although still mostly positive SUC values (+05 to +3%o0) in overlying strats. In this respect, the 65C trends sumimatised in Fig. 4 may also be compared with data Iron carbonate sections of the Nama Group, Namibia (Kaufman ev el, 1991) showing one marked positive excursion (fram —6 ty +6 /oe) in the lower part of the succession (Kuibis Formation) and relatively stable, moderately positive values (+1 to +2.5°/e9) through the overlying Schwarerand Formation, There 1s inde- pendent evidences, based on occurrences of Ediacaran soft-bodied metazoat fossil remains (Awsamik ef ul. 1985; Sun 1986; Jenkins 1992, 1993), chat |hese three successions in South Australia, China dnd Namibia are broadly cocval, The largety saliciclastic Rodda Beds are generally leaner in carbonate Hum the corresponi- ing Chinese and Namibian sections and hence vet isotope record is more susceptible to diapenetic alteration, Nevertheless, lwo minor negative excursions expressed in the Late Ediacaran profile from Meramangye-l (Fig. 3) appear lo be real and mu diagenetic artefacts. Their signifivance has yet to be evalualed, bul they may prave to be uselll in future inter-regional chemostratigraphic correlations (Brasier et al, (990; Maganiz et al, 1991; Kroll & Waller 1992), Acknowledgments This work was supported in part by a University of Adelaide Research Grant (D.M-MeK., RLF) and funding fram Esso Australia Limited (R.J,.P1), We thank K. R. Turnbull for assistance with the isotope amalyses; 8, Proferes for preparation of the figures; the South Australian Department of Mines and Energy far gooess ty drillepre; andl A, R. Chivas, D 1 Gravestock, W. V. Preiss and J. O. Stone for pertincm comuricnts of the piaiuseript. References Anaron, PB. SexipLowsei, M. & Sinch. §. B. (1987) Crnronosteatigraphic markers in the end-Precambrien carbon isotope tecurd of the Lasser Hinialaya. Nature 327, 699-702, Apbursun, T. F & Amur, M. A. (1983) Stable isotopes of oxygen and carhon and their application ta sedimen- tologic 2nd palecenyironmental problems. 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(compiler) (1987) "The Adelaide Geosyncline: Late Proteroyaic Stratigraphy, Sedimemarion, Pulazonmlogy and Tecwries" Geol Surv. 3. dust, Bull 53. ——_— & Kreic. G6. W. (1992) Stratigraphic drilling im the nomheaster Officers Basin Rioddit-2 well, & Aas. Dep Mines & Enerey Rey. 158, 48-51. SACKETT, W. M., Eapne, 8. J. & Exner, M. B. (1974) Stable isDt6pe Composition of ofgatie catbon ih Roceal Antarctic sediments, pp. 561-67], fr B, Tissot & F, Bienmer Eds) “Advances in Oreanic Geochemistry (973% (Edilions Technip: Parisi Senmiog, PW, WititAMs, & EB. & EMareion, Bod, (1991) Low palaeolutitude of Late Proterazoic glaciation carly nming of remanence in hacmatte al the latina Formation, Soutly Australia. Barth & Planen Se. den 105. 355-367, STRAUSS, HL, DES Manais, D.J., Haris, J) M, & Summons, R. E, (1992) Proterazoe organic carbon - js preservation and isciapic recon’, pp. 203-21) Jn M. Schidlowski_ S- Golubic, M. M, Kimberley, BD. M. McKindy & P A. 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(F988) Depositional sequence analysis applied to Late Proteroasic Wilpens Group, Adelaide Geosyncline. South Australia. Aust. J Fah Se, 35, 39-72 Wastace, M. W._-Gostrs, ¥. A. & Keays, R. R. (1989) Discovery of the Acramuan impact ejecta hlanitt in the iy ll and ¥15 stratigraphic signifivance. foi, 36, Zang, Wo & Watren, Mo R. (1989) Latest Proterozoic plankton from the Amadeus Basin in central Austfelia Matterey S37, 642-645. & (1992) Late Proterazon and Cambrian imicrafossils and biostratigraphy. Amadeus Basin. antral Austrailia. Assoc, Awstrelay. Palaeeniml Mew UL TWO NEW SPECIES OF NEMATODE (NEMATODA: ENOPLIDA: THORACOSTOMOPSIDAE) FROM LAKE ALEXANDRINA, SOUTH AUSTRALIA BY W. L. NICHOLAS* Summary Two new species of Thoracostomopsidae (Nematoda: Enoplida) are described from Lake Alexandrina near the mouth of the River Murray in South Australia. Mesacanthion alexandrinus sp. nov. belongs to the small group of congeners with unequal spicules, but differs in having only slightly unequal spicules and a simple gubernaculum. Enoploides stewarti sp. nov. is very similar to another freshwater species, E. fluviatilis from the River Volga, but differs in minor details of the spicules and supplementary organ. Enoploides and Mesacanthion are typically marine genera and implications of the presence of these two new species in freshwater Lake Alexandrina are discussed. KEY WORDS: Taxonomy, nematodes, distribution, Thoracostomopsidae, River Murray, South Australia, Mesacanthion alexandrinus sp. noy. Enoploides stewarti sp. nov. tranaqetions of the Royal Suciety af S. Ausi. (1993), U7(4), 163-170, TWO NEW SPECIES OF NEMATODE (NEMATODA; ENOPLIDA: THORACOSTOMOPSIDAE) FROM LAKE ALEXANDRINA, SOUTH AUSTRALIA by WL. NicHoLas* Summary NicHoLas, W. L, (1993) Two new species of nematode (Nemaroda; Enoplida; Thoracostomopsidue) front Lake Alexandrina, South Australia, Tras, R. Soe. S. Aust, 117(4), 163-170, 30. November, (993, ‘Two new species of Thoracostomopsidae (Nematoda; Enoplida) are dexenbed from Lake Alexandrina near the mouth of the River Murray in South Australia. Mesecarthion alexandrinus-sp. nov. belongs tothe smull greup of congeners with tinequal spicules, but differs in haying ody slightly unequal spicules and a simple gubernaculum, Enoptoies stewurtl sp. nov. is very similar to another treshwater species, &. fluviaiilis from the River Volga, but differs in minor details of the spicules and supplementary organ. Enoploides and Mesacarithion are typically mating genera and implications of ihe presence of these two new species in freshwater Luke Alexandrina are discussed, Key Woros: Taxonumy, nematodes, distribution, Thoracostomopsidae, River Murray, South Australia, Mesacanrhion alexandrinus sp, nov; Enopluides stewartiosp. nov. latraduction The numerous species of the genera Mesacanthion and Enoploides have worldwide distributions characteristically mbabiting occan beaches. Several species of both genera have been found in lreshwater (Riemann 1975). The two new species described in this paper. come from Lake Alexandrina, a large freshwater lake at the mouth of the River Murray, which has only comparatively recently (in 1940) been isalated From the sca by barrages built across the estuary. The teniatode faun of the estuary and of Lake Alexandrina has been described by Nicholas et al, (1992), together with observations. on hydrology and sedimentology. Thoracostomopsidae Filipjev, 1927, possess mandibles, one dorsal and two-sub-ventral, terminating in recurved hooks that project into the buccal cavity. In Mesacanthion Filipjey, 1927, the mandible is tormed from two Jateral rods joined by ab anterior curved bar, enclosing a roughened membrane, In Exoploides Ssaweljev, 1912, the lareral bars are fused for most of their length to form a solid rod, without any enclosed membrane, bifurcated anteriorly into curved hooks, In both genera a forwardly directed! tooth (onchium), with the opening of the duct of a pharyngeal gland at the tip, lies at the base of each mandible. The cephalic setve ure inserted further forward on the cephalic capsule in Mesacanthion than in Enoploldes. Materials and Methods The new species were collected during the investigation of the nematode fauna af Lake Alexandrina alseady referred to (Nicholas ef ai, 1992), * Department of Botany and Zoology, Australizn National University, GRO. Box 4. Canberra, ACT, 2601, Austraha in which 490 mi samples of sandy sediment were taken with a metal corer to a depth of 5 cm at the waters edge. Further details of procedure, sampling and a map locating sampling sites are given in the paper cited. Nentatodes were extracted from the sediments by differential sedimentation in tap water, collected ona nylon 75xm sieve, fixed in 8% fortialin, transferred lw 5% aqueous glycerol, which was evaporated to anhydrous glycerol al 37°C, After identification under the microscope, permanent mounts were made im anhydrous glycerol, Mesacanthion alexandrinus sp. novi FIGS 1. A-E. 2, A-D, Table 1 Types: Holotype: male, Lake Alexandrina, South Australin, I&vit. 989, coll, W. L. Nicholas and A. F Bird, in SAM 23875: paratypes: 2 males and 2 fenmates, Lake Alexandrina, South Australia, (6.vi.1989. 20Viii.1989, coll. WL. Nicholas and A. F Bird, in SAM 23876-79; mule and female, Lake Alexandrina, South Australis, 18.71.1989, coll, W. L. Nicholas and A. F Bird, (8.989 in W. L. Nicholas collection, slides 1220, 1221. Description of Holetype male. Cuucle smooth, not annulated, with sparse setac, up to 17 yarn long, along the cntire body, but fewer and shorter towards the posicrior end, Cephalic capsule high, 2.6-times width at base, dome-shaped, with three deeply incised lips, each carrying thin triangular |ip-flap, Six 12 ~m long labial setae insert at base of lip-flaps, Six 29 wm ianer cephalic setae and four adjacent 14 pum outer cephalic setae originate about half-way up cephalic capsule, Three broad mandibles (width % height), edged ‘by strong lateral and transverse bars, Jateral bars letinimating in divergent hueks, broad inter-bar Membrane externally rough; three onchia at base of 164 W. L. NICHOLAS Fig, 1. Male Mesacanthion alexandrinus sp. nov. A, head; B, mandible and onchium; C, head and cervical region; D. spicules and supplementary organ in lateral view; E, spicules and supplementary organ in dorsal view. TWO NEW SPECIES OF NEMATODE mandibles. Six pairs of cervical setae inserted immediately posterior to cephalic capsule, each pair with anterior short (10 pin) and postesior long (17 nim) seta. Pharynx long, cylindrical; cardia conical; rectum short. 1,3 times body width at cloaca. Testes, two, opposed, to right of intestine. Two long slightly unequal spicules, with straight shaft curving ventrally towards Lip. Gubernaculum flat plate with two terminal hooks, no apophysis; supplementary organ globular, inconspicuous, anterior to and 2.6 body widths from cloaca, Shorter spicule reaches level of supplement (78 pm, 3.1 body widths) longer just beyond it (86 gm, 3.4 body widths), Tail short, narrows sharply behind cloaca then cylindrical almost to tip. Additonal dimensions see Table 1. Puratypes! One male, SAM 23877, displays the spicules in dorsal view (Fig. 1 E) with right spicule longer, and gubernaculum with hooks. Females resemble males, apart From sexual organs, but lack six pairs of cervical setae and, unlike males, tail tapers uniformly to tip. Didelphic opposed gonads. Measurements in Table 1, Habitat: Freshwater, sand at waters edge of Lake Alexandrina, South Australia. Comparison with other species: Wieser (1953) published a key to Mesacanthion which he later updated (Wieser 1959). The spicules and gubernaculum provide the most useful specific characters, but because 1657 some species had been described only from females il was necessary to find characters applicable to females, The depth and form of the buccal cavity and length of the cephalic setae ure also useful, Species described only from juveniles were considered species inquirendae. Subsequently, Gerlach & Riemann (1974) listed 40 yalid species. Since then four more species have been described:- M. southerni Warwick, 1973, M, arabium Warwick, 1973, M. heterospiculum Sergeeva, 1974, and M. marisalba Galtsova, 1976, M. alexandrints sp. nov. belongs to the small group with unequal spicules. In M. diplechma (Southern, 1914), redescribed by Boucher (1977) and M. sautherni, the spicules are very unequal, whereas in M. alexandrinus sp. noy. they are both Jong and only slightly unequal. The gubernaculum in M, arahium is like that of M- alexandrinus, sp. nov, but {he former is much larger, has equal spicules, and possesses two rows of stout pre-cloacal setae, M. langispiculun Gerlach, 1954 resembles the new species, but has much longet and equal spicules. In M. Aeteraspiculum the spicules are uniformly curved, the tail is contcal and uniformly tapered, the cuticle annulated. In comparison with species described only from females, all included in Wieser (1959), M. alexandrinus sp. nov. belongs to the group.in which the buccal cavity is deep, conical, without basal pockets. It can be distinguished from all the others, either because it has very short labial setae or possesses very Jong cephalic setae, or has setae arising close to the base of the cephalic capsule, TABLE. |. Measurements of Mesacanthion alexandrinus. sp. nov Type holo para Sex Male Male Slide 23875 23876 Length 1630 1780 Max. width 38 Sq Labial setae 2 13 inner cephalic 29 27 Owler cephalic 4 13 Mandible hxw ox Id 8x10 Head, hxw 22%33 22x33 Norve ring, 126 141 Phurynx, 526 S04 Vulva - - Tuil 1D 131 Width at anus 25 29 Spicules, arc 78, 86 70; 79 Supplement 70 ‘Tail/anal breadth 4,32 4,52 Nerve ring/pharynx 0.24 0.28 1. ceph, setae/hw 0.42 0.39 Mandible h/w 0.64 0; 40 Spicule/anal width 3.12; 3.44 2.4); 2.72 De Man's ratio a 43 33 " b 3.1 3.5 " c 15 14 * c 43 45 - Va para para para para para Male Male Female Female Female 23877 1220 23878 23879 J221 1450 2400 2560 2270 2570 37 59 66 86 60 12 12 12 9 i2 28 29 27 27 28 12 16 1] 16 3 8x10 12x14 8x13 (2x14 14x15 20%27 28% 36 26x38 15X20 21x 114 153 151 180, 1é0 430 575 730 594 TVR - - 160K) 1242 1406 97 id 139 137 136 23 32 33 32 36 72;82 69;30 - - = 64 ed - - - 4.23 3.44 4,21 4.28 3.44 0.27 0.26 0.21 0.30 0.22 0.44 0.40 0.31 0,42 6,50 0,80 0.85 0.62 0.85 3.13; 3.56 9 2.15; 2.5 39 42 39 26 42 3.4 1.6 3.5 3.8 3.6 Ls 22 18 \7 19 42 34 43 4.3 34 63 35 55 166 W. L. NICHOLAS Mr Oumar yb PAO OVO) OO = = eee, D Cc 50 pm F ——_—_—_—_—150 pm, body hirsute, no Post anal organ, . : -E polysetusus Spicules <150 pm. not tirsive st atin eeu bat MR ve ome 2 2. Male with prominent post-anal organ..... E. eanpere Male without such organ, . 5 ae 3, Tail long (¢<(0, & =7),. no pos anal suite. | ct .E. " disparilis Tail Totes (c>10, c =5) itt re: or ~ post-anal setge or f+) 10)) |: Sn SS Rn eT 4 4, Tail medium c~l0-16, ¢ <5. ---2 eeu ee 5 tail very short, ¢>16, C<5,, 022.0022... 200-21 5, 7-8 pairs post-anal setae... 20... EL alexandrae 7 pairs pre-anal papillae. -E cirrhatus 6. Tail with two pester papillae, no terminal setae - an —- ay x fluviatilis terminal setae, no post-anal papillae ---- Tail with 3 wen greeter eteee dade © .......E. stewart sp. nov. Discussion Lake Alexandrina has been isolated from saline estuarine water for only 50 years, since the building of the barrages across the estuary of the River Murray: Possibly both species, have become adaptcd to freshwater in the very short time since they were trapped above the barrages. However, no similar species were found below the barrages (Nicholas et al. 1992), nor have they been found in extensive collecting on nearby ocean beaches subsequently. Before the building of the barrages the outflow from the Murray was subject to great variations, and in droughts the flow otf river water out of the estuary sometimes failed allowing sea water into the lake system. At times in the past when the climate was dner, these conditions may haye persisted long enough for the adaptation of marine species to tolerate wide variations in salinity and eventually freshwater. It is not knuwn whether any Thoracostomopsidae occur elsewhere in the Murray Darling system. It is interesting that E. stewarti sp, nov. yery closely resembles &. fluviatilis, trom the River Volga in Russia, and is closer to E. alexandrae and E. disparilis from the Black Sea than congeners. Perhaps it was carried to South Australia on the bottom of ships, although the Murray mouth was not a useful port because of its treacherous and unreliable entrance, In this context it is interesting to note that specimens of Ortcholaimus dyjardinii described from Plymouth in the British Isles, TABLE 3. Features separating Enoploides stewarli sp, nov. and four similar species. E. virrhatus E. aleandrae Cuticle annulated anoulated Spicule. arc 60 48/50 Spicule head rounded rounded c 16 12 rom ? Fe * Calculated from figures A. disparilis E. fleviaulis &. stewart annulated smooth smooth 35 112 107-121 0 cylindrical cylindrica] 9.8 25 2.1-3.0/1-7-21 7.2* 3* 1.8-2,7/2.5-3.0 TABLE 4. Comparison between measurements of Enoploides fluviatilis and E. stewarti sp. nov. Spicules, are Supplement jo cloaca Body Tail E. fluviatilis type 1610 Oo) range 1340-1900 54.61 Mule on loan 2219 67 E, stewurli sp, nav haletype 3080 134 2 paratype males 1930-2115 89-93 3 paratype 2587-2749 134-145 females * Calculated from figures. 112" 75+ ‘ol 60 12] 118 107-108 78-100 170 W. L. NICHOLAS together with several other nematode genera, have been identified in scrapings from the bottom of sailing ships entering the port of Sydney from the British [sles (for Bicentenary celebrations) as part of an unpublished study by the Australian Muscum. Studics of the as yet unexplored nematode fauna of Australian inland waters may throw fresh light on the natural history of the two new species, Acknowledgments 1 thank Dr Alan Bird, who took and processed most of the samples, and Dr A. Stewart, both of whom collaborated in a study of the nematode fauna of the Murray estuary, for their help, Tam grateful to Dr Tsalolikhin for loan specimens of E. fluviatilis from Russia. References Boucuer, G, (1977) Nématodes des sables fins infratittoraux de Ia Pierre Noire (Manche occidentale), [V. Enoplida. Bull. Mus, natn, Hist. nar. Paris, 3 ser., 468-752. Fiumey, LN, (1918) Free-living marine nematodes of the Sevastopol area, Trudy oseboi zoologicheskoi Laboratorti Sevastopol’ skoi bivlogicheskot Stantsii (2) 4, 1-350. (RAVEEH. M. 1968. Israel program for scientific transtations, Jerusalem, 1-255.) GA TSova. VV, (1976) Introduction to free-living nematodes as a component of the meiobenthos of the Chupe Inlet of the White Sea. In Nematodes and their role in the metobenthos. (Nauka, Leningrad.) pp 270. Gervacu, S. A. (1954) Nématodes marins libres des eaux souterraines littorales de Tunisie et d’Algérie. Vie Milieu 4, 221-237. —__— & RIEMANN, F. (1974) The Bremerhaven checklist of aquatic nematodes. Veraff. Inst. Meeresforsch, Bremerh. Suppl. 4, (2), 405-736. Jansen. P. (1986) The nematode fauna in the sulphide-rich brine seep and adjacent bottoms of the East Flower Garden, NW ASiaad of Mexico. II. Enoplida, Zoologica Ser. 13, 93-99. MICOLETZKY. H, (1923) Freilebende Nernatoden der Wolga, Ark, biel. Wolga-Staiion 7, 1-27. Nicnonas, W. L., Birp. A, E, Beecn, T. A. & Stewart, A. €, (1992) The nematode fauna of the Murray River estuary, South Australia; the. cffects of barrages across its tiouth. Aydrabtologia 233, 87-102. RIEMANN, F, (1966) Die interstitielen Fauna im Elbe-Astuar; Verbreitung und Systemalik, Arch. Aydrobiel,, Suppl. 31, 1-279. (1975) EBinwanderung mariner Nematoden in das Stisswasser, Untersuchungen in Kombien im Vergleich mit aussertropischen Regionen, Int. Revue ges, Hydrobiol, 60, 393-407, Sercrevs, N. G, (1974) New free-living nematodes order Enoplida from the Black Sea 2. Zool, Zh, 53, 120-125. SourHers, R, (1914) Nemathelmia, Kinorhyncha, and Chaetognatha (Clare Island survey, part 54). Proc. R, Irish Acad. 31, 1-80, Uzunov, J. (1974) Two new intersitial nemic species [rom Bulgarian Black Sea littoral. C. R. Acud. bulgare Sci: fi, 843-845. Warwick, R. M. (1973) Freeliving marine nematodes from the Indian ocean. Bull. Br, Mus, nat. Hist, (Zool.). 25, 87-117. Wieser, W. (1953) Free-living marine nematodes 1. Enpplaiiica: Lunas Universitets Arsskrift, N. FE Avd. 2, 49. (1959) Free-living nematodes and other small invertebrates of Puget Sound beaches, University of Washington Press, Seattle, pp 179. & Hoprer, B. (1967) Marine nematodes of the East Coast of North America, 1. Florida. Bull. Mus. comp. Zool. Harv. 135, 239-344. INTERSPECIFIC AND INTERGENERIC RELATIONS BETWEEN NEMATODES PARASITIC IN THE STOMACHS OF KANGAROOS AND WALLABIES BY H. HOSTE* & I. BEVERIDGE} Summary Associations between conspecific and confamilial nematodes co-occurring at high prevalences in the stomachs of the eastern grey kangaroo, Macropus giganteus, from Victoria, the red kangaroo, M. rufus from New South Wales and rock wallabies of the Petrogale assimilis species complex P. assimilis, P. sharmani and P. mareeba, from Queensland, were investigated using principal components analysis. A similar pattern of associations was found in each host species, consisting mainly of positive associations between nematode species. Negative associations were found with Rugopharynx australis the numerically dominant nematode in M. giganteus and M. rufus, and to a lesser extent with R. zeta in Petrogale spp. The complex nematode communities present in each host were shown to be stable, with few negative associations between members. KEY WORDS: Macropus giganteus, Macropus rufus, Petrogale assimilis, nematodes, communities, associations, multivariate analysis Transactions af the Royal Seelery of S. Aust. (993), LIT 4), ITE 17. INTERSPECIFIC AND INTERGENERIC RELATIONS BETWEEN NEMATODES PARASITIC IN THE STOMACHS OF KANGAROOS AND WALLABIES by H. HosTe* & |. BEVERIDGEY Summary Haste, H, & BrveRiodE, 1, (1993) Interspecific and intergeneric relations between nemalodes parasitic in the stomachs of kangaroos und wallabies, Zhuns, Ro Sac, S, Aust W7(4), 171-177, 30 November, 1993, Assvclations between conspecific and contitnilial nematodes co-Gecurring aC high prevalences in the stomiuchs of the custern grey kanyarod, Macropuy gigariteus, from Victoriz. the red Kangaroo, M. rufus from New South Whles and rock wallabics of the Perrevale assimilis species complex P ussinulis, P sharmant and P mareeba, fron Queensland, were investigated using principal components analysis, A similar pattern of assockitions was found in each host spectes, consisting mainly of positive associations between nenvalode species. Negative associations were found with Rugephuryns australis the numerically dominant nemalode in Af. gigumtews and M. rafts, and to-a lesser extent with R, zea. in Perrogale spp. The complex nematode communities present if each host were shown to be stable, with few negative associations between members. Kky Worps: Macropus xizqtteus, Macropus rufiis, Perrogale assimilis, nematodes, voramunilies, xssociatians,, muluvariate analysis Introduction Parasite communities in homeathermic animals arc frequently complex in nature, with many species of parasites occurring in an individual host or host organ (Bush et al, 1990; Kennedy & Bush 1992), Because of the complexity of the community and, frequently, the large numbers of helminth species involved, Ue question as to Whether community. members interact or have interacted in the past (o produce a stable, predictable structure or are non-interactive und result frum # random collection of independent, individual species, has received considerable attention (sce Holmes 1986; Proce 1986), Evidence for the existence of interactive communities has bean provided in the case of cestades of ducks (Bush & Holmes 1986 a,b) and trichostrongyloid nematodes in sheep (Diez-Banus etal. Y992; Hoste & Cabaret 1992) while at the other extreme. the complex assemblages of monogenean parasites present as ectopacasi(es of fishes appear to exhibit few interactions (Rhode 1979; Koskivaara & Valtonen 1992; Koskivaara ef al. 1992). Amongst nematode parasites, the best known examples of complex parasile communities existing within w single host organ are the oxyurid nematodes of tortoises (Schad (963; Petter 1966), and the strongyloid nematodes of elephants (Chabaud 1957), horses (Kennedy & Bush 1992) and kangaroos (Inglis 1971, Kennedy & Bush 1992), In the case of kangaraas, up to 40 species of nematodes. al! belonging to a single order, the Scrongylida, occur in the complex sitccular fore-stomachs of individual host species (Spratt er al. ™ Instiut national de la Recherche Agronomique, Nouzilly, France. T Deparment af Vetennury Scienve, Leniversity of Melhoume, Parkville, Vie. 3052 1990), with numbers of parasites reaching 300,000 or more (see Beveridge & Arande] 1979), Several studies (Mykytowycz 1964; Mykytowyez & Dudzinski 1965; Smales & Mawson [978h; Arundel e7 a/, 1979) have demonstrated that different species or genera of nematodes have different site preferences within the stomachs of kangaroos, as 1s the case with oxyuroid nematodes in tortoises (Schad 1963; Petter 1966), tut there have heen no studies undertaken to determine whether there is any evidence of interacnwas within these helminth communities, Recently, Hoste & Cabaret (1992) lave utilised a principal components analysis (PCA) and comparison with the model of Motomura 41947) in which the log, abundance of a species is correlated with ils rank in terms of abundance for examining the stability of nematode communities in sheep and the existence of interactions between the species ar genera of parisites present with a host. In this paper. we apply their techniques to examine whether there is evidence ol competitive interactions between the nematode parasites present in the stomachs of three taxa of macropodid marsuptals- Materials and Methods Parusitolagical data Dara unilised in this study were obtained trom earlier epidemiological studies on the parasites of macropodids. The data for Perrogale assimitis included 35 specimens of P assimilis as well as live specimens. of what Were lormerty known as the Mt Claro and Mareeba chromosomal races of tis species. They have recently been numed Po sharmant and P. mareeba respectively (Eldridge & Close 1992), Rock wallabies were collected overian extensive area of nonhery and Wm H_HOSTE & |. BEVERDIGE western Queensiand (see Beveridge e¢ al. 1989) during a study of the taxonomy’ of the hosts. Nematode taxa included in the study. their prevalence and mean imensity of infection are shawn in Table |. Any helminth species occurelog wurside the stomach, or at a prevalence of less than 10% ¢,u, Cloacina similis, Corerastrangylus coronatas, Labtostrongylus hancrofii, Macropastroneylus petrogale ec.) was not considered to be a core species (sensu Hanksi 1982) and was excluded from the statistical analyses. In addition, nematodes ‘such as Woodwardastrongylus obendoni for which intensity data were not available, were also excluded. Since similarity coefficients for the helminth communities in F. assimlis, P shanmani and P mareeba ure high (Beveridge ev ul. 1989), combination, of their data Was considered justifiable. Data trom Macropues gigenteus were derived from an opidemidlogical study conducted a1 Yin Yean, Victoria, in which wo adult and tour juvenile kangaroos were collecied al six-weekly intervals over Taste |. Prevalence and intensity ef jnfectian af the friecipal rematede parasites present in the stimach a] 39 Petrogale .assimilis. P sharmani und P marecba from Quevaslandd, Parasite species Prevalence Moan (%) — Tnvertsity Sugapharyna zeta (Johostin & Mawson, 1939) 33 2.190 Clnacina petrogale Johnston & Mawson, 1938 7Z 390 © pearsoni Mawson, 1971 ‘98 980 © parva Johnston & Mawson, (938 \od 780 . dydriformis Johnston & Mawson, 1938 AG 200 C1. sp. (undescribed) (Cap. | of Beveridge ef al, 1989) 54 420 Filarinema spp, 3 3 (£ disstmile (Wood, W931), FE australe (Wood, 1931), F varawewe Cassone de Baccam. 1985) Taste 2. Prevalence imél infensicy of infection of the Pracipal nematode pacasites presem in the stomach of 45 Mawropus giganteus fram Yen Wan. Victoria. Parusite species Prevalence Mean (%) — TAfensity Ruvgopharynx custralis (Moning, 1926) 100 R. rosemariae Beveridge & Presidente, 1978 27 Cloacina spp. 38 (C. vbtusa Johnston & Mavwsan, 1939, C. cf. dydeiformis Johnston & Mawson, 1938, C. cf. elegans Johnston & Mawson, 193%) Latsostrongylus spp 7A (. bipapillosus olusvon & Mawson, 1938), £. kungi Mawsen, 1955) Pheryngestrongylus kappa Mawson, 1965 46.250 2,210 7,630 2) Strmgploides sp: (undescribed) 39 835 a period of 10 months (Anindel « ai. 1990). The prevalence and mean intensity of infection for the principal nematode species in the stomachs of these hosts are shown. in Table 2. Any nematode species occurring in less than 10% ofthe host specimens (eg. Alocostoma clelandi) was excluded frean the snalysis, as were the species of intestinal cestodes. Data on the helminth parasites of Macrapus rufies were collected at a single localuy. Menindee, New South Wales (see Arundel ef al, 1979), with 12 animals collected every two months, ovet a perind of two years. Samples of kangaroos were collected within Kinchega National Park and on properties linmediately adjacent to the Park. The only difference in prevalence detected was in the case of the bile duct inhabiting cestode Pragamotaenia festiva (see Arundel ef al 1979). No differences were detected in the iniensity of infection with any parasite. Hence it was considered valid to use combined data trom. the two adjacenl collection localities (Table 3). Helminth parasites occurring af a low prevalence or intensity, such as the nematodes Macropostronyyloides spp.. and Hypadontus macropi and the cesiodes Progamoraenia reficola and Triplotaenia undosa were excluded from the analysis. Changes in the nomenclature of parasites since the publication of the original epidemuological papers have heen indicated in ‘Tubles 1-3. together with appropriate references. In M. siganteus, individuals of the genus Labiestrongylus and in M rufus and M. giganteus members of the genus Cloucina were not identified in a quantilative fashion to species level, because of inadequate informanon on the laxanomy of these genera, though the species present at cach loealiny were recorded. In M, gixanens, most Of the species of Cloucina present are urdeseribed. TAm& 3. Prevalence and intensity of infection of the principal nematode parasites present in the stomach of If) Macropus rutus from Menindee, New South Wales, Parasite specics Prevalence Mean (%) Intensity Rugopharyne australis (Manning, 1926) 99 51,850 Willabinesa cobbi (Kung, 1948) (syn. Zoniwaimay cobb) 84 9,055 Cloacina spp. (C. cf, Aydrifornis Johnsion & Mawson, (938, ¢. 71 expansa Johnston. & Mawson, 1939, CL maecrapadis Johnston & Mawson, 1938) Labtestrenaylus fongispicularis Woot, 1931 3,090 $8 332 Filarineme. spp. 92 a) (F flagrifer Monning, 1929, & australe (Wood, 1934)) Papillostrongylus sp. (undescribed) (= P, labians Johnston & Mawsan. 1939 sensu Arundel er ui, 1979) a! 1.5K} RELATIONS BETWEEN NEMATODES IN KANGAROOS Stulistical methods For each host species, a separate principal component analysis (PCA) was performed, using the STATITCF computer program (1988), on both the intensity data (sevsu Margolis er al,, 1982) and the frequency data defined as the number of nematodes of a particular species expressed as a percentage of the total number of worms within that host, The data were standardised prior to analysis as (actual valuc — mean value for the variable) + standard error for the variable. The data were arranged ina correlation matrix whose columns (variables) were the parasite species and rows were the individual animals, Component axes were defined from correlations between the variubles (parasite species numbers). The coordinates af each variable were then expressed in relationship to the new axes, Axes 1 to 3 were studied, the percentage of Taste 4. Percentage of total variability expressed by component axes 1, 2 and 3 following Principal Componenr Analysts (PCA) of the intensity and frequency of infection of Perrogale spp. Macropus piganteus and M,. rufus with nematade parasites, : Host species Axis | Axis 2 Axis 3 Total (lh +2 +3) Petrogule spp. (ntensity 44 #220 18.7 75.1 Frequency 316 23.8 192 74.6 Muacrepus giganteus Intensity 28.0 24,5 18.4 70.9 Frequency 33.6 213 183 732 Macropus rufus Intensity 37.0 19.7 168 73,5 Frequency 31.2 18, 18.0 67.8 3 variability accounted for by each axis being indicated for cach PCA in Table 4. For cach host species and each set of dala, ie, intensity and frequency, Buctidean distances between Parasite species were calculated by applying Pythagoras’ theorem to the coordinates of cach variable, j.¢. parasite species, was located within the three dimensional space constructed by axes I to 3 (Table 5), These distances characterised quantitatively the relation between nematode species. The minimum value for these distances was 0, the maximum 2. Distance values > 1.2 and < 0.8 are considered indicative of negative and positive interactions between species respectively (Hoste & Cabaret 1992). Tr each host species, the distances between each pair of nematode species were calculared in the three planes and the result was called D,. These D, distances were fitted to the logarithmic model of Motomura (1947) i.e. the decimal logarithms of distances were regressed on the rank of each species. pair. According to Motornuira’s (1947) model, the log,, of the abundance of a species is corretated with ‘its rank in the order of most abundant to least. abundant species. In addition, im order to represent the retation of one particular species to others present in the stomach, mean distances were calculated as the averages of the D, distances for each pair-of worm species including, ihe particular species of interest. These mean distances were called D, in both species of kangaroos as they were distances between a given parasite taxon and five others, and called D, in the case of the rock wallabies since the distance was from six other taxa, Taste 5. Coordinates of nematode speciey on component axes], Z and 3 following u Principal Components davlysis (PCA) performed on the intensit\ and frequency of infection of Petrogale spp., Macrequs gigunteus and M. rutus, with nematode purasites. Host Parasite Cloacina sp. 1 C.. hydriformis C. parve C. pearsont C. petrogale Rugopharvns eeta Filarinema spp. Ruyopharyrts austratis P. kappa Cloacina spp. Labiastrangylus: spp, R. rasemariae Strongyloides sp. fablostrongylus longisptenians Filarinema spp. Rugopharynx australis Cloacina spp. Hallabinema eabbl Papillostronyylus-sp. Petrogalé spp: Meecropuy sigantens Mueropeas mifas Imensity Data Frequency Data Axis Axis Axis Axis Axis Axis 1 2 =] ] 2 3 -0.625 0.311 0.497 0.382 0.070 0.772 218 -0,813 4064 +33R -O,7GR 860.124 “O.783 0.298 -O.448 -0.629 O.501 -0.453 0863 0.179 0.394 -0.743 0.319 = 0,409 “0.456 -0.785 0.137 -G,224 .0,.43K -0,24N 636 -0,081 (), 505 0,9)2 0.039 -0,.112 01.024 0.198 6.653 0.349 O120 -0.538 0.653 0.594 O<£.169 0.954 -0.244 0.05) A305 0,262 0.673 -D.109 O.876 = -0,220 0.374 O714 -0.294 -0.906 -0.248 0,039 1.370 -0.077 0.688 O<:.976 0.614 0,450 AUSI7 «=—-O.647) 41S -DO44d4 -9.120 0.582. 0.785 0.337) OF<:216 -0.264 -0.010 -0.710 C.004 o.a91 O87s D.162 -O0.088 0.622 4.686 0,060 0.014 -0.447 =(),. 444 0,141 O.900 -0.133 0.087 O<.988 0.037 -0.132 0.5035 0.627 -2:.019 -0.318 O419 0.578 O.3I9 -O.659 0.167 -0.050 0.259 =—0,386 0.138 -0.769 0.452 -0.728 0.393 -0.417 74 H, HOSTEL & 1. Results Average distances of individual species from remaining species (D, and D,) Comparison of the mean euclidean distances for each worm species with related species in the three different hosts showed several similarities (Fig. 1). Firstly, in the three host species, the mean distances cylculated from the frequency data were penérally higher than those calculated from the intensity data, Secondly, no mean distances calculated from intensity or frequency 1.2 DISTANCE b 3] Zz x a Cc Mw o 85 (=) 1.2) which suggests Negative interactions, This observation was: made in the three different host species, and among those pairs exhibiting the highest distance values, Rugopharyax spp. were usually present, These results suggest that Rugepharynis was negauvely associated with the othet worm species based on the frequency analysis. The D, distances calculated from the intensity daca (Fig. 3) were lower than those obtained from PCA performed on the frequency data, In the three different host species. the D, distances were less than 1.2, and usually less than 10, which tends to indicate the lack of any significant negative association between nemalode species. The rank distribution of patterns of these D. distances alse fitted Motomura’s model. When compared tm the distribution of the D, “ - a Las w = Oo 12 Ls Win soo 8o00 Heros b ae < bi a EOP ike Sea Sony 2 ERPs gee san5 c Tip. 3. Ranked distances (D,) between parasite species puirs hased wn intensities of infection in (a) Perroyale spp. ¢b) Macropus giganteuy and ic) M. ryfae valculatcd from rincipal components analysis. .cegend: as for Fig. 1, Ws distances pbtained ffoni the frequency analysis, the order of the different pairs in the intensity data was distinct, with a Jess well defined ranking of Rugapharyas australis. Discussion The results of unalysis both of [requency and intensity of infection data for al] three species of macropodid hosts can be fitted t¢ Motomura’s model (1947) for density associations between populations of similar species within the same biotope and suggest the existence of stable nematode communes in the stomachs of the macropodid species examined, Furthermore, most of the D, distances calculated on the intensity data were less than 1.() and even less than 8. As these distances are thought to reflect the intensity of parasitism wt the suprapopulation level or host population level, this fact provides additional evidence af the preponderince of positive associations between component species. and hence infers the extstence of stable communities, Positive assuciations are known Lo occur in the case of other host groups acquiring their parasite infections from grazing pastures oontaminaled with various species of infective: third stage larvae of nematodes such as ruminants’ (Diez- Ranos et al. 1992; Hoste & Cabaret 1992). On the other hand, the frequency-based distances mainly reflect the liifracomimunity present ln the individual host, and the fact that the frequency-based distances were generally greater (han those derived [rom intensity data suggests Lat additional regulatory factors are involved at the suprapopulation level, as oecurs also in the case of ruminants (Hoste & Cabaret 1992), The regulation of nematode populations in kangaroos is not well understood. Sotales & Mawson (1978) and Arundel ev vif, (1990) demonstrated that in the case of the Tammar wallaby, M. eavent? and eastern grey kangaroo M. gigantexs, in winter rainfall areas of South Australia and Victoria, there was an increase in the number of nematodes present in the slomach during the moist winter months which is the most favourable period of Whe year for Jarval development in the external environment. Arundeletal, (1990) also demonstrated an effect of host age on certain specics of nematodes (e.g. Rugepharynx rasemariae), with juvenile animals exhibiting, a higher prevalence and intensity of infection, while in other nematode genera (Cleacina, spp., A. austredis)intensity was tigher in adult hosts. By contrast, the study by Arundel et af. (1979) on the red kangaroo, AM. rufus, in the arid, non-seasonal rainfall region of westers: New South Wales indicated in the case of several of the dominant nematode species (& australis, Wallabinema cobbi) that there was no seasonal effect gr intensity of infection and thal jntensity oF infection increased linearly with host age. I all of these three Mucrapas species, Labiastrongylis 16 H. HOSTE & 1, spp. exhihiced an unique pattern of development, with lurval slages mafuring over a period of several months during the summer (Mykytowyez & Dudzinski 1965; Smales d& Mawson 19784; Arundel er al. 1979). Recause of the lack of detailed knowledge of the way tt which nematode populations in kangaroos are regulated, is difficult to expla what the udditional regulatory factors at the suprapopulation inferred by the present analysis might be. However, Petter (1966) also found an effect of host age and season on interactions between the oxyuroid nematodes of Lorigises, suggesting that these might be general phenomena, It contrast to other studies on gastro-intestinal strongylid nematodes (Haste & Caburet 1992, Diez- Banos et al, 1992), analysis of populations of the stomach-rhabiting nematodes parasitic in three species of macropodid hosts failed to teveal evidence of extensive negative interactions between nematode species, The only suggestion of negative associations involved Rugepharyrt australiy in red and grey kangaroos, and in both of these kangaroos, R. australis is tumerically the dominam stomach nematode (Arundel er ai. 1979, 1990; Beveridge & Arundel 1979). In the case of the rock wallabies no particular dominant apécies emerged, Rugepharyax zeta occurred at a Ingher mtensity in rock wallabies than other gastric nematodes, but occurred al a prevalence of only 50%. Cleatina pearsoni 2nd C parva by contrast occurred in 100% of the hosts examined but at a lower intensity. The differences in intensity and prevalence may counteract one another lo presenl a curmmunily in which there are few negative interactions, An addinonal consideration when comparing data from rock wallabies with that from the kangaroos is that in the former case, hosts were collected over 4 wide area of northern Queensland, compared with single localities for each of the kangaroo species. Negative associations are evident in the frequency dato only, which refers essentially to the infrapopulation at the individual host level; there is no such evidence from analysis of the raw intensity data, which relatcs to the parasite suprapopulation or the host population level, However, Holmes (1986) has cautioned that Imeructions are frequently ohscured when suprapop- ulutions are considered and that the optimum method fiw detection of interaction ts at the infrapopulation level, a conclusion which is clearly supported by the current results, Thus, the scmutode communities present in the three species af matrapodids examined here demonstrate similar features In feing stable, and probably mainly Tun-competilive communities, with the exception of Raugopharyrx australis. This lack of negative associations contrasts sharply with the results of Bush & Holmes (/986a) from studies on the cestodes of ducks and those of Hoste & Cabaret (1992) and Dicz- HEVERDICE Banos et uf, (1992) for runmnants, Robie (1979) suggested that in now-interactive cominumbes not all niches dre filled, there is no basis fur competition and differential localisation of parasites ovvurs becuse it facilitates repruduction rather than minimises competilion. Ratde's hypolfiesis (1979) is thereiore consistent with the data curremly available for nematode communtiies parasitic tn the stamachs of macropodids, The kangaroo sturmach is a relatively enormous organ in comparison with the size of Parasites, and dala From varjous sources (e.g, Beveridge & Acunde] 1979) suggests that healthy macropodids are capable of harbouring numbers of nematodes far in excess of those encountered in this study, Hence it is not unreasonable to assume that vacant niches are abundant within kangaroo stomachs, given the proviso tbat nematode pumbers only have been examined, without any consideration of their relative biomass. Whether the differential Jocalisations of nematodes observed are dye to reproductive sirategies or due to the presence of competition in the past. during the evolution of the parasite community structure, is net clear. Price (1986) predicted that considerable variation in community characteristics might be expected when a sufficiently wide range of communities was examined. The present results indicate that speciose nematode communities of homocothermic vertebrates, while stable in their structure, may nol necessarily display significant negative associations between their component members. Acknowledgments We wish to thank Drs J. H. Arundel, R. Close and §. ©. Barker for their involvenvent in (he collection of the original data und Drs D. M) Spratt und N. B Chitown and Two anonymous referees tor ber commenis on drafis of the manuscript. H, Hoste is 4 arvteful reciplent of a fellowship tréei the French Mimastry of Research and Space Tectinelngy: References ARUNDEL. | Hy. Bevoripor. L. & PRESIDENTE, P. J, A. (1979) Parasites and pathological findings in enclosed and Cree-ninging populations of Macropus nifies (Desmarest} (Marsupialia) at Menindee, New South Wales, Aust Wile! Res. 6, 361-379, , Dempster, K.J,, HARrIGAM, K, BE, & Black. R. (1990) Kpidemiological observations on the helminth paras of Macropus giganteus Shaw in Victoria. [bid 17, BeVeRiDGE, |. & AauNoeL J. H. (1979) Helminth parasioes of grey Kangaroos, Marropus eizanteus Shaw and re Sigiaron (Desmarest), in eastern Australia. Bid 6, , Spratt. D, M,, Barker, 5 C,, CLose, RL, & SuarmMan, G. B (1989) Helminth parasizes of rock wallabies, Petrogule spp. (Marsupialin) Grom Queensland thick 16, 273-2 RELATIONS BETWEEN NEMATODES IN KANGAROOS 177 Busn, A. O., AHO, J. J. & Kennepy, C. R, (1990) Ecological versus phylogenetic determinants of helminth parasite community richness, Eval. Ecol. 4, +20. & Howimes, J, C, (1986a) Intestinal helminths of lesser scaup ducks; patterns of association. Canad, J. Zool. 64, 132-141. & (1986b) Intestinal helminihs of lesser scaup ducks: an interactive community. /bid 64, 142-152. CHasaup, A. G, (1957) Revue critique des nématodes du genre Quilonia Lane, 1914 et du genre Murshidia Lane, 1914. Ann, parasitol. hum. comp. 31, 98-131. Diez-Banos, N,, Casaret, J, & Ditz-Banos, P, (1992) Interspecific interactions in naturally acquired nematode communities from sheep abomasum in relation to age of host and season in four areas of Leon (Spain). int, J. Parasitol. 22, 327-334. ELpRIDGE, M. D. B. & CLose, R. L (1992) Taxonomy of rock wallabies, Pefrogale (Marsupialia; Macropodidae). 1. A revision of the eastern Petragale with the description of three new species. Aust. J. Zool. 40, 605-625. HAnskt, T_ (1982) Dynamics of regional distribution: the core and satellite species hypothesis. Orkas 38, 210-221. Hovmes, J.C, (1986) The structure of helminth communities. pp. 203-208. Jn M. J. Howell (Ed.) “Parasitology-Quo Vadit?" (Australian Academy of Science, Canberra). Hoste, H. & Caparet, J, (1992) Intergeneric relations between nematodes of the digestive tract in lambs: a multivariate approach, Int. J Parasitol 22, 173-179. Inciis, W, G. (1971) apecanion in parasitic nematodes, Adyan, Parasital, 9, 201-223, Krnnepy, C, & Busu, A. O. (1992) Species richness in helminth communities: the importance of multiple congeners. Parasitalogy 104, 189-197, KOSKIVAARA, M, & VALTONEN, B, T. (1992) Dacnlogyrus (Monogenea) communities on the gills of roach in three lakes in Central Finland. Jord. 104, 263-272. amiga 5 & Vuorr, K-M. (1992) Microhabitat distribution and co-existence of Dactiagyrus species (Monogenca) on the gills of roach. /bid 104, 273-2381. Marcous, L. G., Escu, G. W,, Houmes, J. C., Kuris, A. M. & Scuan, G. A. (1982) The use of ecological terms in parasitology. J. Parasital. 68, 131-133. MorTomura, f. (1947) Further notes on the law of geometrical progression of the population.density in animal association. Sere’ Seiai 1, 55-62. Myrytowycz, R. (1964) 4 survey of the endoparasites of the ted kangaroo, Megaleia rufa (Desmarest) Parasitology 54, 677-693. & Dupzinskt, M. L. (1965) Sex ratio, weight, length and numbers of Labiastrangylus longispicularts (Wood), the large stomach worm of the red kangaroo (Megaleia rufa (Desmarest)), in relation to age of the host and season, {bic 55, 527-541. Price, P. W. (1986) Evolution in parasite communities. pp: 209-214. Jn M. J, Howell (2d.) “Parasitology-Quo Vadit?” (Australian Academy of Science, Canberra). Prtrer, A, (1966) Equilibre des espéces dans les populations de nématodes parasites du cdlon des lortues terrestres. Mém. Mus. nat. Hist. nat., Paris, Sér. A. Zael. 39, 1-252. Rowpe, K. (1979) A critical evaluation of intrinsic and extrinsic factors responsible for niche restriction in parasites. Amer. Nat. 114, 648-671. Scnap, G. A. (1963) Niche diversification in a parasitic species flock. Nature, Lond. 198, 404-406. SMALES, L. R. d& Mawson, P. M. (1978a) Nematode parasites of the Kangaroo Island Wallaby, Macropus eugenii (Desmaresl), I. Seasonal and geographical distribution. Trans. R, Sac. S. Aust. 2, 9-15, &__s(1978b) Nematodes and other helminth parasites of the Kangaroo Island Wallaby, Macropus eugenii (Desmarest), 2. Site selection within the stomach. [bid 102, 79-83, Spratt, D, M.. BEVERIDGE, i, & WALTER, E. L, (1990) A catalogue of Australasian monotremes and marsupials and their recorded helminth parasites. Rec. S. Aust, Mus., Monogr. Ser. 1, 1-105. Stav-itcr (1988) Manuel ‘Utilisation. (Institut Technique des Céréales et des Fourrages, Paris), EFFECT OF ACROBELOIDES NANUS (NEMATODA: CEPHALOBIDAE) UPON THE OF PSEUDOMONAS CORRUGATA (EUBACTERIA) IN PASTEURIZED SOIL FROM KAPUNDA, SOUTH AUSTRALIA BY MAARTEN H, RYDER*? & ALAN F. BIRD* Summary Acrobeloides nanus and Pseudomonas corrugata were co-inoculated into the same pasteurized field soil. The population of the nematode increased ten-fold in eight weeks, whereas the population of the bacteria decreased with time, in all cases. After eight weeks at 15°C, the population of the introduced bacteria was significantly lower where A. nanus had been added to the soil. At 22°C the population of P. corrugata was much lower irrespective of the presence or absence of A. nanus. The feeding of nematodes on bacteria in soil helps to explain the observed decrease in the population of microbial control agents introduced into the soil in field experiments. KEY WORDS: Acrobeloides nanus, bacteria, biological control, Cephalobidae, nematodes, Pseudomonas corrugata, soil, take-all. Transacitons bf te Reval Sariety af §. Aust 0993), 117(4), 179-182. EFFECT OF ACROBELOIDES NANUS (NEMATODA; CEPHALOBIDAE) UPON THE SURVIVAL OF PSEUDOMONAS CORRUGATA (EUBACTERIA) IN PASTEURIZED SOIL FROM KAPUNDA, SOUTH AUSTRALIA by MAARTEN H. RYpeR* > & ALAN F. Birp* Summary Ryper, M, H. & Biro, A. F (1993) Effect of Acrobeloides nanus (Nematada; Cephalobidae) upon the survival of Pseudomonas cerrugata (Bubacteria) in pasteurized soil from Kapunda, South Australia. Trans. R. Soc S. Aust, 117(4), 179-182, 30 November, 1993, Acrabeloides nanus and Pseudomonas corrugata were co-inoculated info the same pasteurized field soil, The population of the nematode increased ten-fold in eight weeks, whereas the population of ihe bacteria decreased with time, in all cases, After eight weeks ar 15°C, the population of the introduced bacteria was significantly Jower Where A, anus had been added to the soil, At 22°C the population of 2 cornegata was much lower irrespective of the presence or absence of 4. varus, The feeding of nematodes on bacteria in soil helps to cxplain the observed decrease in the population of microbial control ygents introduced inte the soil in field experiments. Key Wokos; Aerobeloides nonus, bacteria, biological conwel, Cephalobidae, nematodes, Pseudomonas enterpeiq, soil, bike-all- Intreduction With increased interest in the use of biological contro! agents as alternitives to chemical control measures has come an interest in the fate of these biocontrol agents in the soil, Ln particular, information on the effect. thal bacterial-feeding nematodes may have on bacterial biocontrol organisins is sparse. According- ly, We have isolated, cultured’ and microscopically examined a biocontrol bacterium and a nematode from the same agricultural soil and have studied their relationship both on agar plates and in the soil. The bacterium Pseudomonas corrugata isolate 240R is u biocontrol agent.against the. rowt-pathogenie fungus Gaeumannomyces graminis var. trittel (Ryder & Rovira 1993), This fungus causes the root disease take-all which is responsible for serious yicld losses in ceteal crops in southern, Ausitalia and elsewhere itvihe world. P. corrugata TI40R is a rifampicin-resistant derevative of the parent strain and is, therefore, easily identified. The nematode which we used was Acrobeloides nanus (Anderson 1968; Bostrom & Gydemo 1983; Nicholas & Stewart 1989, Bird er al. 1993), a cosnvopolitan, bucterial-feeding, free-living organism that is Widespread in agricultural and arid soils throughout Australia, The nematode A. nanus can use the bacterium in vitro as w nutrient source and is able to complete its life cycle and reproduce while feeding only upon FP corrugata (Bird & Ryder 1993), We show that the presence of 4. nonus can lead to a reduction of bacterial numbers when both organisms are Inoculated into pasteurized soil. “CSIRO Division af Soils and + Cooperative Research Centre for Soil and Land Management, Private Bug 2, Glen Osmond, South; Australia 5064, Australia. Materials and Methods Nematode Acrobeloides nanuy was isolated, by meyns of & misting apparatus, from the dry soil, collected six years previously (in 1986), from which P corrugata had originally been isolated (Bird & Ryder 1993). Bacteria Pseudomonas corrugate 2140 was isolated originally from the chizosphereé of wheat seedlings that had been grown in a field soi, a red-brown earth pH 6, collected fram Wagga Wagga, New South Wales (Ryder & Rovira 1993), The rifampicisi-resistant derivative P. corrugata 2H40R was isolated as a spontaneous mutant growing an Nutrient Agar, by J, Brackin, Monsanto Co., St Louis, USA. Sail ane pasteurization conditions Ficld soil, a sodic red-brown earth (Stace: er al, 1968),, more recently defined as a fine mixed thermic caleic natrixeralf (Soil Survey Staff 1990), from Kapunda, South Australia, that had been air-dried and sieved (<3 mm) was used in our experiments. Immediately prior 10 use, it was steam stenlized at 75°C for 1S myin in order to kill all nematodes but not all bacteria, fnaculatian of soil with bacteria and nematodes Treatments consisted of nematodes + bacteria (+N-+B), nematodes alone (N), bacteria alone (B) and controls without either, Fach treatment was replicated five times. The experiments were run ut 15°C and 22°C and were hatvested at eight weeks from contmence- ment. The nematodes, grown on P corrigaia 2140, the parent bacterial strain which is sensitive to tat] M.H. RYDER & A.F. BIRD rifampicin, on malt extract agar (Oxoid) were washed from the Petri dishes with sterile distilled water, centrifuged three times in sterile distilled water (200 »* g for 10 min) and added as aliquots to the soil samples. In the +N+B treatment, bacteria were added 40 the suspension of nematodes immediately poor tu indculation of the soil, At the commencement of the experiment, samples (20 g) front all four treatments were placed on 4 misting nematode-extraction apparatus und the nematodes in the eluate were counted, The -+-N+B treatment yielded four nematodes per g of dry soil, and +N alone five. No nematodes were found inthe soils with added hacteria (+B) or in the controls (C)- Bucteria (RP corrugara strain 2)40R) were grown on Nutrient Agar (Difeo), containing rifampicin (Boehringer) at 100 mg per litre, for two days at 25°C. The cells were harvested by washing the plate with 4m sterite distilled water. The cell density was adjusted to ca. 2 x 107 colony-forming units (cfu) per ml before inaculation of soil. The starting population of 2 corrugata was taken as the number of rifampiein- resistant bacteria recoverable from the soil 30 min aller inveulation. Five replicate samples were processed for dilution plating, The initial bacterial populations. were 15 x 30° cfu/g dry soil (+B) and LI7 x 10° cfu/g dry soil (+N-+B). The air-dried soil was wet initially to [2% (w/v) with distilled water, 20 2 portions of moist soil were dispensed into sterile containers (plastic-capped, 120 ml capacity), Either nematodes (+N) or bacteria (+B) alone were added to the soil in | ml, For addition of both organisms together (+N+B), the suspensions of nematodes and bacteria were mixed and then added to the soil in 1 mil. Sterile distilled water (| ml) was added to the control treatment. The organisms’ were mixed into the soi thoroughly using # glass rod that had been wiped with ethanol. but the mixing was done pently to preserve the nematodes, The tinal watet content of the soil was 19.4% (w/w dry soil). Incubation conditions The containers with treated sail were incubated in darkness at eather 18°C or 22°C. They were capped tightly, but once each week the lids were removed for several seconds in a laminar flaw cabinet, to allow gas exchange to oceur. The incubation period was eight weeks, fsolation anid enttmeratian of bacteria Sterile distilled water was added to each container se that the total volume of soil and liquid was 70 ml. The container was shaken gently for 30 sec to allaw the soil aggregates to disperse, One ml was taken for analysis of bacterial populations by dilution plating. The selective medium used wus Nutrient Agar conlaining cycloheximide (75 mg/l) and rifampicin (100 my/1), The dropict plating method (three replicate 10 jc droplets) was used except for the controls (Ch where 0.1 ml of the undiluted suspension was spread onthe plate. Bacterial colonies were counted after twa days at 25°C. [solation and contiting of nematodes The nematodes in the remaining 69 ml of sail suspension were isolated by means of Scinhorst’s two ask technique followed by sieving and counting (Hooper 1986). Statistics Populations of nemulndes and bacteria Were analysed by an analysis of variance of log-iransformed data (Genstat Version 5), Results The populations of both 4. nanus and PF corrugetts after eight weeks in pasteurized Kapunda soil at J5°C or 22°C are shown in Figs | & 2, There were no significant differences in populations of A. nanus at either femperature after cight weeks (bacterial treatment: F=0.18, p=0.68; temperature: F=1.29, p=0.27). Where nematodes. were added their numbers had increased at least ten-fold since the commencemem of the experiment, Addition of 4, nanus significantly | 0 weeks 8B weeks Zeeeeceend 60 (SE) : “| 40 [ts -aaneeon 30 Nerriatodes /g dry soil 20 10 Fig. i_ Histogram showing no signilicant difference in numbers of n¢matodes at either temperature after eight weeks irrespective of whether bacteria were udded or not Error burs represent standard error (SE) of the mean EFFECT UF A. NANUSON P CORRUGATA IN SOIL, (3! 0 WEEKS 8 weeks N+B row oO 2 Number of bacteria (log,, cfu /g dry soil) on Las] 5.4 5.2 5.0 Pt, Me 8 4,8 Ba | EA 15°C 22°C Fig. 2 Histogram showing a significant decrease in bacterial nuMnbers after cight weeks al 13°C when nematodes were present, reduced the population of P covitiguld aver eight woeks at 15°C (hematode treatment: F=15.49, p< 0,001; lempentiure: F=18,53, p< 0,001: interaction between nematode treatment and temperature: F=3,76, p=0.038). Numbers of P. corruyata decreased eight- Juld in the presence of A. nanas compared to two and three quarter-fold in the absence of the nematode. At 22°C over this period of time, bacterial nunibers were reduced. to approximutely one twenticth of the population «| the start of the experiment, irrespective of whether or not nematodes were present, Discussion The presence of A. anus m the soil significantly reduced the pupulauion of 2 corrugata after eight weeks at 15°C. This temperature closely resembles winter soil temperatures in the held in South Australia. when biological control of G graminis var. tritici by PF cormeata has been tested. Our results provide 3 possible explanation for the observed decrease in numbers of pseudomonads jn biocontrol experiments conducted in the field (Weller 1983; Ryder et af. 1990), After eight weeks at 22°C, the numbers of P currgata were much reduced. This vecurred irrespective of whether 4. manus was present or not, This suggests that competition with other bacteria. with Browth optima clase to 22°C, (nay be responsible for the decline In numbers of B cornigata in the soil. This Wecrease in numbers of P corrugen? is probably not due to increased temperature because its growth rate in vitro is higher at 20.30°C than at IS°C (Ross & Ryder, unpubl, ). Pasteurization of the soil resulted in the death of all som] nematodes and many, but not all, of the bacteria. This meant that the P. corrugate were introduced into 2 hacteriologically competitive soil environment, One such competitor, yet to be identified, proved to be a bacterium that formed visible white patches on the surface of the soil. These patches were considerably reduced in the presetice of the nematodes and were presumably fed upon by 4. manus. The bacterial-feeding nematode A. nanus is hoth Widespread in South Australian agricultural soils and is relatively abundant, forming 12-24% of the total nematode fauna in soil from field plots at Kupunda, SA, sampled in the spring of 1992 (Yeates & Bird in press}. This is equivalent to I-17 nemulodes per gram of dry bulk soil. Thus the levels of added A. rectnees in the experiment reported here, were representative of numbers found naturally in the field. The ability of 4. meriny, and other species of bacteria-feeding nematodes, ta graze on bacteria may affect tte biocuntro! performance of introduced bovteria, This may be particularly important for Wriduced bacteria in the rhizosphere. as there is evidence that bacteria-fecding nematodes can inhabit the rhizosphere in large numbers (Griffiths 1990). The relative populations of bacterivorous nematodes in the rhizosphere and bulk svil should also be of preat interest to those who wish to introduce specitic baclerial strains into the rhizosphere. Whether the effect of 4. nanus in decreasing populations of P corrugata can be large enough to decrease biocontrol performunce in a natural soil is not yet known. Interactions between rhizosphere or Soil bacteria and plant pathogenic nematodes have been studied by Bookbinder ef af (1982). Io their study, there were synergistic negative effects when the bycterial strains, including P corrugata isolated from symptomless alfalfa roots, were inoculated together with a range of plant pathogenic nematodes. However, as far as we are aware, outs is the first report of the relationship between a specific soil bactermm and a bacteria-teeding nematode isolated from the some soil, Further studies alany these lines with microbivorous nematodes and the bacteria Upon which they feed are necessary steps im understanding both the complexity of the interactions between these organisms in the soil environment and the limitations on the success of biocontrol agents in controlling disease, Further research on these |ines is also warranted because these nematodes are readily cultured, natural inhabitams of the soil whose tmundalive release in other situations may prove to be of immense benelil ti maintaining soil fertility while reducing the numbers of undesirable microorganisms. 182 M.H. RYDER & A.F. BIRD Acknowledgments The authors wish to thank Ms A. Reid, CSIRO Riometrics Unit, for performing the statistical analyses. References ANDERSON, R. V. (1968) Variation in taxonomic characters of a species of Acrobeloides (Cobb, 1924) Steiner and Buhrer, 1933. Can. J. Zool. 46, 309-320, Birp, A. F., De Ley, P. & Biro, J. (1993) Morphology, oviposition and embryogenesis in an Australian population of Acrobeloides nanus. J. Nematol., 25, 606-614. ___—s & Ryper, M, H. (1993) Feeding of the nematode Acrobeloides nanus an bacteria. Ibid. 25, 493-499, BooksinDER, M, G., Boom, J. R. & Luxkezic, F. L. (1982) Interactions among selected endoparasilic nematodes and three pseudomonads on alfalfa. /bid. 14, 105-109, Bostrom, 8, & Gypemo, R, (1983) Intraspecific variability in Acrobeloides nanus (de Man) Anderson (Nematoda, Cephalobidae) and a note on external morphology. Zool. Seripta 12, 245-255, GrieeitHs. B. §. (1990) A comparison of microbial-feeding nematodes and protozoa in the rhizosphere of different plants. Biology and Fertility of Soils 9, 83-88. Hoorer, D. J. (1986) Extraction of free-living stages from soil, pp. 5-30 Jn J.F. Southey (Ed.) “Laboratory Methods for Work with Plant and Soil Nematodes.” 6th Edm (HMS, London). Nicnouas, W, L. & Stewart, A. C. (1989) Experiments on anhydrobiosis in Acrobeloides nanus (de Man, 1880) Anderson, 1968 (Nematoda), Nematologica 35, 489-491. Ryper, M. H., BrisaAne, P. G. & Rovira, A- D_ (1990) Mechanisms in the biological control of take-all of wheat by rhizosphere bacteria. pp. 123-130 In D. Hornby (Ed.). “Biological Contral of Soil-borne Plant Pathogens’, (CAB International, Wallingford, UK). & Rovira, A. D. (1993) Biological control of take- all of glasshouse-grown wheat using strains of Pseudomonas corrugata isolated from wheat field soil. Soil Biol. and Biochem, 25, 31-320, Sor. Survey STAFF (1990) “Keys to Soil Taxonomy” 4th Edtn SMMS technical monograph no. 6 (Virginia Polytechnic Institute and State University, Blacksburg, Virginia, USA), Srack, H. C. T., Hupsie, G. D., BREWER, R., NORTHCOTE, K. H,, Steeman, J. R.. MuLcAHy, M, J. & HALLSwortH, E. G. (1968) “A handbook of Australian soils” (Rellim Technical Publications, Glenside, South Australia). Wetter, D, M. (1983) Colonization of wheat roots by a fluorescent pseudomonad suppressive to take-all. Phytopathology 73. 1548-1553. Yeates, G. W. & Biro, A. F (in press) Some observations on the influence of agricultural practices on the nematode faunae of some South Australian soils. Fund. and Appl. Nematol. THE BIG ROCK DONGA METEORITE: A NEW H5 CHONDRITE FROM SOUTH AUSTRALIA BY M. ZBIK* Summary The Big Rock Donga meteorite is a stone with a total weight of 11 kg found north of Fisher, South Australia, in 1989. It has been classified as a H5 chondrite and contains olivine (FAj 38), orthopyroxene (FSi69), plagioclase (Anj4Ore;Ab799), clinopyroxene, nickel-iron, troilite, and chromite. Mineral compositions and textures indicate that the Big Rock Donga meteorite was a metamorphosed part of the H-planetoid and was very weakly shocked before reaching Earth. KEY WORDS Big Rock Donga, meteorite, chondrite. Transacnons of the Roval Society of 8 Aust (99S), WV7(4) 183-185, THE BIG ROCK DONGA METEORITE: A NEW H5 CHONDRITE FROM SOUTH AUSTRALIA by M. Zpik* Summary Zuik, M. (1993) The Big Rock Donga meteorite: a new H5 chondrite (ror South Australia. Zrtas. RK. Soc 8. Aust. VI7(4), 183-185, 30 November, 1993 The Big Rock Donga meteorite is a stone with a total weight of U ky found north of Fisher, South Australia, ie 1989. Tt has been classified asa HS chondrite and contains olryine (FA), ,), orthopyroxend (Fsi_ 9), plagioclase (An Or, ,Abjyo) clinopyroxene, nickel-iron, troilite, and chromite. Mineral compositions and textares indivite tal thatthe Big Rock Donga meteorite was a metamorphosed part of the M-planetord and wes very weakly shocked belore reaching Earth. Kry Worns. Big Rock Donga. meteorite, chondrite, Introduction The Biz Rock Donga meteorite was found ip {989 by un unknown collector, north of Bisher on the South Australian part of the Nullarbor Plain. [t has a total weight of IL ke (South Australian Museum records) but is NOW in several pieces, The approximate co- ordinates lor the site ure 30°33°S, 120°58°R. The Meteorite Was collected illegally and exported to the United States of America, but we have obtained a 37 gram slice froman American mineral dealer, Mr David New, According to McCoy (1991, & pers. comm, 1993) the neleorite weighed 19 ky initially and was recovered asa single stone lo km north of Fisher by a rabbit trapper (who wanted t remain nameless). In recent years the Nullarbor Region has proved to be a productive urea for the recoyery of meteorites (Bevan 1992; Bevan & Binns 1989, b) and in the lust few years has attracted the attention of illegal meteorile collectors. Under legislation enacted by the Governments of Western Australia and South Australia all Ineteorites found in these stules ure the property of the Crown, An unfortunate consequence of the (legal trade to meteorites is the loss of important information On the exact date and location of the find. Physical description The stone is not heavily weathered but displays iron staining of the silicate minerals and. fractures filed with irom oxides especially hear the fusion crust. The interior of the meteorite 1s dark grey to black ip colour and ieditin to fine grained with heavy fine grained nicke-iron metallic inelusions, In thin section the meteorite 1s generally light coloured (Fig. 1). The chondrules are recrystallised and have poorly defined * Department of Mineralogy. South Australian Museum, North Terrace, Adelaide, South Australia SOOO, boundaries but are recognisable under crossed palars They are typically 0.5 mm in diameter. Metal and troilite occur as finely disseminated grains throughout the matrix, One piece of the slice wus cul and u polished thin section was prepared and used for petrographic examination and in electron microprobe analyses. Mineralogy Composiuons of the silicate minerals were determined with a JEOL electron microprobe at the University of Adelaide Centre for Electron Microscopy and Microbeam Analysis. Analyses were made using an accelerating voltage of 15 kV, a suumple current of 3 nA, and a bean width oF 5 am. Representative mineral analyses are presented in Table | Relic chondrules and chondrule fragiments are composed predominantly of olivine With lesser amounts of orthopyroxene. The majority of chondrules are porphyritie in texture comaining euhedral olivine crystals up to 0.2 mm in length set in a matrix ol Tashe |, dverave chemical compositions of niajar ainerals in the Big Rack Dongit meteorite. oxide weight olivine orthopyroxene clinopyroxene plagioclase SiO, 34.5 548 S14 62.7 TiO, 0.2 a4 (it ALO, (1 02 0.4 oft Fe 14,7 3 4.0 03 MnO 0.5 0.6 0.2 01 MgO 42.6 W8 16.5 - Cuo 0.04 0.6 22.4 Lo Na,Q - : 0.3 8.3 KO : - at 10 Cr,0, - 0.2 1.2 ‘Yotul 100.5 08.7 uso Oo 2 Fig. 1. Photomicrograph of the Big Rock Donga meteorite in thin section, showing a poorly defined porphyritic chondrule containing large euhedral olivine crystals in a matrix of subhedral prismatic bronzite crystals. subhedral prismatic bronzite crystals with interstitial growths of clinopyroxene and plagioclase. Barred chondrules occur occasionally and are composed of olivine. Radial chondrules composed of orthopyroxene are also present and contain thin lamellae of either clinopyroxene or plagioclase. The matrix of the meteorite consists of slightly fractured grains of olivine and orthopyroxene (bronzite) which display very weak undulate extinction and rare grains of clinopyroxene. Plagioclase is abundant throughout the matrix as small turbid grains which also display weak undulate extinction. Nickel-iron metal, troilite, and chromite occur as accessory minerals. Microprobe analyses show that the olivine in the Big Rock Donga meteorite is equilibrated with a mean fayalite content of Fa,,, (range Faj, , 9, 16 analyses). The orthopyroxene shows only a small variation in chemical composition with a mean ferrosilite content of Fs,,, (range Fsy¢ 517° 14 analyses) and a wollastonite content of 1.1 mol%. The composition of the clinopyroxene is Wo,,,En,,Fs,, (10 analyses) and that of the plagioclase is An,,Or, Ab.) (II analyses). The pyroxene geothermometers of Wells (1977) and Lindsley (1983) suggest that the Big Rock Donga meteorite was heated to temperatures of between 700° and 800°C during metamorphism while the meteorite was still part of the large H type asteroid. Classification Despite McCoy's initial interpretation of this meteorite as an H6 chondrite (McCoy 1991), the Big Rock Donga meteorite has been classified as an H5 chondrite. The olivine composition (Fa,,,) is within the range of the H chondrites (Keil & Fredriksson 1964), and the orthopyroxene composition (Fs ,, 9) shows that the meteorite belongs to the olivine-bronzite chondrite group. The highly equilibrated mineral compositions, crystalline matrix, poorly defined chondrule boundaries, suggest that Big Rock Donga THE BIG ROCK DONGA METEORITE 185 belongs to the type 5 classification of Van Schmus & Wood (1967}. The wollastonite content in the orthopyroxene is similar to that found in other HS chondrites (Scott e7 a/. 1986), The presence of weak undulatory extinction in olivine and plagioclase grains and the degree of crystal fracture all indicate that the meteorite has been weakly shocked after meta- morphism. According to the classification scheme of Stéffler er al. (1991), the shock facies is estimated to be $2, very weakly shocked. Acknowledgments The author wishes to thank Mr G. Horr for preparing a polished thin section of the meteorite and Mr Huw Rosser of CEMMA, University of Adelaide for assistance with the electron microprobe analyses. Mr B, McHenry is thanked for helpful suggestions to the manuscript. The financial support of the Friends of the South Australian Museum is gratefully acknowledged. References Bevan, A. W. R. (1992) Australian Meteorites. Rec. Aust. Mus. Suppl, 15, 1-27. —_— & Binns, R. A. (1989a) Meteorites from the Nullarbor Region, Western Australia: 1, A review of past recoveries and a procedure for naming new finds, Meteoritics 24, 127-133. & ____ (1989b) Meteorites from the Nullarbor Region, Western Australia: IT. Recovery and classification of 34 new meteorite finds from the Mundrabilla, Forrest, Reid and Deakin areas, Jbid. 24, 135-141. KeiL, K. & FREDRIKSSON. K. (1964) The iron, magnesium and calcium distributions in cocxisting olivines and rhombic pyroxenes of chondrites, J. Geophys. Res. 69. 3487-3515. Linostey. D. H. (1983) Pyroxene thermometry. Am, Mineral. 68, 477-493. pe T. J. (1991) The Meteoritical Bulletin. Mezeoritics 6, 256. Scott. E. R. D., Tayvtor, G. J. & Ket, K. (1986) Accretion, metamorphism, and. brecciation of ordinary chondrites: evidence trom petrologic studies of meteorites from Roosevelt County, New Mexico. Proceedings of the Lunar and Planetary Science Conference 17, El\5-E123. STOFFLER, D., Kett, K. & Scott, FE. R, D. (1991) Shock metamorphism of ordinary chondrites. Geochimica et Cosmochimica Acta 55, 3845-3867. VaN ScHMus, W. R. & Woon, J. A. (1967) A chemical- petrologic classification for the chondritic meteorites. [bid. 31, 747-765. WeLts, P. R. A. (1977) Pyroxene thermometry in simple and complex systems, Contr, Mineral. and Petrol. 62, 129-139, A NEW SPECIES OF LECHRIODUS BOULENGER (ANURA: LEPTODACTYLIDAE) FROM THE EARLY EOCENE OF QUEENSLAND BY M. J. TYLER* & H. GODTHELP} Summary Lechriodus casca sp. nov. numerically is the dominant species in a collection of ilia obtained at Murgon in South-east Queensland. Radiometric dating of illites that form part of the matrix of mammalian material taken there is 54.640.05 million years. Accordingly L. casca is the earliest frog fossil known from Australia. The principal significance of the discovery of this material is the demonstration that generic differentiation at least in part antedates the separation of Australia from Antarctica, and that the dominant anuran genus of the Oligo-Miocene also was dominant in the Eocene. KEY WORDS: Lechriodus, new species, ilia, Leptodactylidae, Tertiary, Queensland. Tranaenons of the Reval Soviwry of S. Ause (G94), MINA) 187 19. A NEW SPECIES OF LECHRIODUS BOULENGER (ANURA; LEPTODACTYLIDAE) FROM THE EARLY EQCENE OF QUEENSLAND by M. I. TYLer® & H. Gonrnecpt Summary Tyeer, Mod. & Goprunuy. H (1993) A new species of Lechriadas Boulenger (Anura: Leptodaceylidie) from (he Early Eocene of Queenstind Trans Ro See So Aus) 174), IBZ 1B9 30 Novernber 193, hechiiodis Cusea sp. nov. namerically is the dominant species in a collection of ilia obtained at Murgon in South-cast Queensland Radionetre dating of dltes that form part af the mati. of maniimalian onternal taken there 18 54.640,05 inillion years, Accordingly £2. cascer ts the earliest frog lossil knows (rom Australi The principal signiticanee of the diseovery of this material is the demonstration that generic differentiation it least in partantedates the yepaniion of Austmiba from Anturetica.ynd that te dominiat anuran ventisal the Oligo-Mioeene who was dominant in the Eocene. Kiy Woros Leehriodip, new species, itia, Leptodactylidae, Tertiary, Queensland. Introduction The lepodactylid (inyobatrachid of some authers) genos Leehriedus Boulenger comprises four extant species restricted to Australia and New Guiness. Amongst the 22 venera of Australopapuan leplodactylid lrogs revognised, four are represented on both landmasses and ace the only ones in New Guinea (Crinta, Lechriodusy, Limnodvaastes and Mixaphyes). Lechriodys is Unique amongst that group in having more species in New Guineas (three) than in Australia (one) (Awerfel 1972). The nature of the geographic distribution of Leedriedits ulso ts uousual, for in atl of the other genera common to both landmiasses, represenuitives exist on the adjacent shorelines, wheres the solitary species in Australia (L. fletcheri Boulenger) is confined 10 the eastern seabourd of New South Wales, and its northern limit is north of the Queenslind border. The northern lintit in Australia ts approximately 2000 km distant from the nearest congener in New Guinea (Zweilel 1972; McDonald & Miller 1982). The fossil record partially bridges the geographic gup in the distribution of extant species becuuse L. lntergerivus Tyler, 1989 js known fron Oligo-Miocene sites at Riversieigh Station south of the Gulf of Carpentaria, in northern Queensland. Here we report 4 further new species of Leeliriaduy from an curly Eveene site at Murgon, 160 km northwest of Brisbane, Lt represents the earliest trog fossil material yet found in) Australia. The biogeographic and systematic significance of the new species is discussed elsewhere (Tyler & Godthelp tn prep). Department of Zoology, ‘Uniyersity of Adelaide, $, Aust. S005. ¥ School of Biologies! Sciences, University of New South Wiles, Kensington, NSW 2006, Material and Methods The material is deposited in the Queensland Museum, Brisbane (QM). and the South Australian Museum, Adelaide (SAM) Letters following the ubbreviations are departmental qentilicauons. Comparitive Studies ure based an osteological eallections inthe Deparunent of Zoology. University of Adelaide. with fossil material from Riversleigh being processed at the time of these studies, and with scanning electron micrographs imelided in ‘Tyler (1989). Osteological nomenclature follows Tyler (1976. 1989), Scanning electron microgniphs were prepared on a Cambridge Autescan, Systematics Family: Leptoductylidae Werner, 1896 Sub-family: Limnodynastinae Tyler 1978 Genus: Leehriodus Boulenger. 1882 The definition of the generic characteristics of the Uium of Lechriedus of Tyler (1976) has been expanded hy Tyler (1979). A distinguishing feature of the yenus is the high dorsal crest extendiny for the entire length of the iltal shaft Lechriodus casea sp, nov FIGS |-3 Aololypes OM 024824, 4 lef ilium locuated at Main Quarry. Boat Mountain area. Murgon (Latitude 26°IS'S, Longitude 151°S7* E), southeast Queensland. Tingamarra Local Fauna. Description of holotype; The proximal portion of the ihum Jacks the greater portion of the acetabular fossa, and both the dorsal and ventral acetabular expansions (Fig, 1). The shaft bears a conspicuous concave dorsal M. J. TYLER & H. GODTHELP Fig. 1. (QM F24824) Holotype of Lechriodus casca sp. noy. 2, (QM F33558) Paratype of Lechriodus casca sp.noy.: lateral view. 3. medial view. A NEW SPECIES OF LECHRIODUS BOULENGER 184 crest proximally. The medial surface bears a conspic- uous, longitudinal indentation which is shallow and weakly rounded proxitnally and deeper and angular distally. The total length is 11.8 mm, Fararypes; 34 ilia - QM F20197, 20216, 20243, 20565, 20567, 20689-93, 20912, 21395, 23018, 23208, 23279, 24824-28; SAM P31700, 31703-06, 31710, 33558-65, All specimens were collected at Main Quarry, Boat Mountain, Murgon. Kiriation: All of the paratypes are fragmentary, but SAM P33558 (possessing a dorsal crest — Figs 2, 3) exhibits most of the features lacking in the holotype. The dorsal acetabular expansion is poorly developed whilst the pre-acetabular zone is evenly curved. The most complete shatt is that.of QM F24824 which is 1,8 mm, It is unlikely that any of the type series would have a complete ilial length greater than 15 mm. Comparison with other species! Lechrivdus intergerivus Tyler (1989) is the only known Tertiary meinber of the genus and completely dominates. the frog fauna at sites at Riversleigh Station during much of the Oligocene and Miocene (Tyler etal, 1990), From that species Z, casca is distinguished most readily hy the absence of any. proxinial tapering of the dorsal crest and by the very poor development of the dorsal acetabular expansion (vide Tyler 1989, Figs 2, 3), Etymology: L. “casca™, old. Stratigraphy and Lithology ‘The anutan and mammalian fossil remains have been recavered from authigenic illite/smectite clays which are part of a sequence of clays and dolomites, IIites from this horizon have been dated radiometrically and have revealed a minimum age of 54.6+0.05 Myr, A minimum age estimate of 29.0+0:2 Myr was obtained from Potassium/Argon dating of a superpositional basalt (Gadthelp ef al. 1992), Discussion Lechriodus is the dominant anuran genus in the Tertiary of Queensland, constituting 28-82% of individual ilia located at sites at Riverslewh Station (Tyler er ai. 1990) and with the lower percentages found at che more recem sites. This trend of an early dorninunce of Lechriedus has been maintained as work on the Riversicigh Station sites. continues and more ilia are recovered, As at 6.xi.92, Ill4 ilia had been recovered and identified and 562 (50'%) represent L, intergerivus, The recovery of a neW species of Lechriadus dominating the fauna at the early Eocene site at Murgon extends the known antiquity of the genus to almost 55 million years. The biogeographic and systematic significance of the find are discussed by Tyler & Godthelp (in preparation). Acknowledgments The excavations at Murgon have been supported by the Australian Research Council, the Bepartment of Arts, Spots, Envitonment, Tourism and Territories, National Estate Programme Grants (Qucensland}, the Queensland Museum and the University of New South Wales. Studies on the Anura have been supported by the Australian Research Council and the University of Adelaide. and invaluable assistance has been provided by Simon Bryars. The SEM photograph was taken by Stuart McClure (CSIRO Division of Soils, Glen Osmond). We ure grateful to Jim and Maureen Porter and Susan Maudsley for permission to work on their propetty. References GinoTHELP, H,, ARCHER, M., Crisis, R., HAND, & d,, & Gitkeson, C, (1992) Earliest Kaown Australian Tertiary mammal fauna, Netare 356, 514-516. McDonarn, K. R, & MILLER, J.D. (i982) On the status of Lechriodus fletcheri (Boulenger) (Anura: Leptodactylidae) in northeast Queensland. Thais, A. Sac, 5, Aust. 106, 220. TytpR, M. J, (1976) Cuniparalive osteology of the pelvic girdle of Australian frogs and description of a new fossil genus, Zbid. $00, 3-14, (1989) A new species of Lechrioduy (Anura: Leptodactylidae) from the Tertiary of Queensland, with «redefinition of the ilial characteristics of the genus, Abid, 413, 15-21. Hanp, 8. J. & Wao, S. J. (1990) Analysis of the frequency of Lechrivdus istergerivus ‘Tyler (Anura: Leptodactylidae) in Oligo-Miguene local faunas of Riversleigh Station, Queensland. Proc. Linn. Soc. NSW H2, 105-109. Zwuinn.,.R. G. (1972) A review of the frog genus Lechiriodus (Leptodactylidae) of New Guine und Australia. dan Mus, Noyit, (2507), 1-4. A NEW AUSTRALIAN SPECIES OF CALODEMA (COLEOPTERA: BUPRESTIDAE) BY S. BARKER Summary A new species of Calodema, C. rubrimarginatum sp.nov. is described from Queensland. The species is spectacularly coloured and increases to four the number of recognised species in Australia. KEY WORDS. Coleoptera, Buprestidae, new species, Calodema. Transactions of the Reval Society ef 8. Aust. (1993). WW7(4) 19}-192. A NEW AUSTRALIAN SPECIES OF CALODEMA (COLEOPTERA: BUPRESTIDAE) by S. BARKER Summary BarkER, S. (1993) A new species of Australian Caladema (Colcoptent: Buprestidae), Trans. R. Soc. S. Aust 714), 11-192, 30 November, 1993. A new species of Calodema, C rubrimarginatum sp.nov. is described from Queensland. The species is spectacularly coloured and increases to four the number of recognised species in Australia. Key Worps: Coleoptera, Buprestidae, new species, Caledema. Introduction The genus Calodema LaPorte & Gory. 1837, erected to accomodate C. regale, comprises a number of large and spectacular species found in rainforest in Australia and New Guinea and some adjacent islands. Adults feed on flowering trees high in the canopy of the forest and are particularly difficult to collect. The species described below has been collected recently in Queensland for the first time. A revision of the genus is In preparation. Calodema rubrimarginatum sp.nov. FIG. | Holotype: or, Rex Ra., Qld. 3.i1.1993, J. Hasenpusch, Queensland Museum 113006. Department of Zoology, University of Adelaide, S, Aust. 5005, LTH > Allotype: 9, same data as allotype, A. Hay, Queens- land Museum T13007. Colour, Head green with purple reflections. Antennae green with gold reflections. Pronotum and scutellum green with purple reflections. Elytra yellow with narrow black basal margin: triangular blue-green mark with truncated apex at spines: red margin commencing at humeral callus, thickening apically, Shape und sculpture. Head punctuation fine, even, dense; median impressed basal line; frons moderately hairy. Pronotum normal, L;W 0,65, punctuation fine, moderately dense, stronger and denser laterally; sides rounded from base to ‘4 distance to apex, then explanate and conyerging anteriorly; anterior margin bisinuate, basally feebly bisinuate: median lobe truncate, median impressed line from base to 14 distance tO anterior margin. Scutellum scutiform, anterior margin straight, concave, without punctures, Fig. la, Habitus illustration and b. Lateral view of Calodema rubrimarginatum sp.nov. holotype. Scale bar = 1 cm. 192 S. BARKER 0.16 width of elytron, Elytra slightly wider than thorax; elytral interneurs long, jointed basally, with fine, shallow, dense punctuations, scutellary striole weak; short secondary interneurs inserted between primaries 1 and 2, longer between primaries 3 and 4, 5 and 6, 7 and 8 with additional punctuations on shoulders; intervals flat, sides subparallel then tapering to pre- apical areas; apex bisinuate, both spines small, interval between sinuous and projecting close to medial spine, lateral spine anterior to medial spine. Prosternum hairy, smooth medially almost devoid of hair; hairy and finely punctuated at the sides. Conical process. with well defined anterior projection in female, smaller in male, proepisternum with coarser punctuations, dense medially, very sparse almost smooth laterally, with deep smooth fossa in posterior angle. Mesosternum and metasternum smooth medially, hairy with fine moderately dense punctuation laterally. Abdomen smooth and shiny, punctuations very fine; $7: male deeply concave; female truncate. Size: Male 34.0 x 13.5 mm (1). Female 36.3 x 14.3 mm (1). Remarks: This species differs from the three Australian congeners in the following ways: C. regale and C. wallacei, have round colour spots on the pronotum and neither has a red margin; C. plebeium has an entirely red pronotum and red elytra with black fascia. Etymology: Derived from rubrico L., red and margino L., margin. Acknowledgments 1 thank the following for assistance: Mr J. Hasenpusch, Innisfail and Mr A, Hay, Cairns for the donation of specimens; Mr P. Kempster, Department of Zoology, University of Adelaide, tor technical assistance, Reference Laporte, F. L. & Gory, H. (1837) “Histoire naturelle et iconographie des Insects Coleopteres” vol. 2 Suite aux Buprestides (P,.Dumenil, Paris). FIRST SOUTH AUSTRALIAN RECORD OF THE CARRION-BREEDING BLOWELY CALLIPHORA NIGRITHORAX MALLOCH (DIPTERA: CALLIPHORIDAE) BY J. F. WALLMAN Summary Transuctions of the Royal Soviety of S. Aust. (1993), W7(4), 193. BRIEF COMMUNICATION FIRST SOUTH AUSTRALIAN RECORD OF THE CARRION-BREEDING BLOWFLY CALLIPHORA NIGRITHORAX MALLOCH (DIPTERA: CALLIPHORIDAE) Carrian-breeding blowflies are important ecologically, medically and forensically’. In view of the difficulties encountered jn identifying them, it ts thefefore ¢rucial that their distributions should be fully documented. This paper records tor the first time the occurrence of one such species, Calliphora nizritherax Malloch, 19277, in South Australia. Recently, a single female of C. nigrirhorax was collected in 4 liver-baited trap on the North Terrace canmipus of the University of Adelaide (20.vit,1993), C nigritherax, together with its sister species C. ochracea Schiner, 1868°, is distinctive among Australian Calliphorg in having dense erect yellow hairs on the cyes and an entirely reddish abdomen, It is distinguished from C: pehracea by whitish pruinesence (dusting) on the mesonotum rather than yellowish. Publications defining the distributions of Australian Culliphora record C nigrithorax as restricted to Tasmania. These are clearly incorrect, on account of both the current record. and the holdings of the species in the Australian Nalional Insect Collection, Canberta, in Which there are specimens not only from Tasmania, but. also New South Wales, the Australian Capital Territory, and Victoria", The occurrence of C. nigrithorax in these regions was in fact first recognised in 1937’. Prior to my find the most westerly record of the species was from Myers Creek in Victoria ‘Heath, A. G. C. (1982) N. Z. Entomol, 7, 343-348. *Malloch, J. R. (1927) Prov, Linn, Soc. N.SW_ 52, 299-335, 4Schiner, I. R. (1868) Diptera, it “Reise der dsterreichischen Fregutie Novara um die Erde in den Jatiren 1857, 1858, 1859 unter den Befehlen des Commodore B. von Wiillerstorf- Urbair’. Zoologischer Theil, Zweiter Band, 1 (B). Abtheilung. (Keiserlich-Koniglichan Hol- und Staatsdruckerei, Wien), (36°23'S, 144.°16'B)®, 'The new record therefore extends its known range on the mainland westwards by about 500 km. The biology and ecology of C. nigrithorax are poorly known. It appears to prefer cool, moist localities. The fact that I have not previously collected this species in two years of widespread trapping of blowflics in southern South Australia suggests that it may have very specialised habitat or brecding requirements, [ts sister species is only locally common in the regions from which it is known, and while apparently never having been found in carcasses or live shee; in the field, has readily aviposited in thick fur in captivity”. C. nigrithorax may be similarly specialised. Alternatively, it may be a rate generalist. Whatever the case, now that C. rigrithorax is known from South Australia, all workers concerned with native blowflies. in particular forensic entomologists. and those studying the ecology of carrion, should henceforth consider the possibility of encountering this species in the mesic regions of the State. 1 gratefully acknowledge the (ucilities and financial support of the Department of Zoology, University of Adelaide. I thank Dr K. R, Norms for valuable discussion and guidance, and Dr A. Wells, Dr D, A. Duckhouse and Dr A. J. Boulton for their helpful comments on the manuscript, 4Kurahashi, H, (197i) Pac, Insects. 13, 141-204. *Kurahashi, H. (1989) Calliphoridae, pp.702-718 in N. L. Evenhuis (Ed.) “Catalog of the Diptera of the Australasian and Oveanian Regions”. (Bishop Muscum Press, Honolulu, and E. J, Brill, Leiden). 'Norris, K. R. (1993) Pers, comm. Hardy, G. H. (1937) Proc. Linn. Soe. N.SW. 62, 17-26. *Kuller, M. E, (1931) Proc. Linn. Soc. N.S.W. 56, 172-181. J. RK WALLMAN, Department of Zoology, University of Adelaide, §. Aust. S005. ERRATA, CORRIGENDA, ADDENDA Summary ya Errata, Corrigenda, Addenda Stee, Rt & Kose, W. (1992) Rotifera rom Australian \iland waters VII, Trichocereldae (Monogononta). Trans, R. Soe, S. Aust, WG), 1-27. Inthe Key {0 species of Trichocerca known {rom Australia, page 6, couplet 30 should lead to 31 und 32 respectively. Hip, 4:2, page 7, depicts Trichacercd bidens (Lacks) which also is shown in Fig. J, not Trichocerca cavia (Hudson & Gosye). The appraprivte figure of 1 cavie is as follows: Fig, Li, p. 21: fa, dh should be 2a; 2h (7) fevrua), 2 should be 3 (7. jenningst) and 3 should be 1 (7? gructis), Io the first paragraph of Other species of Trichacerca, page 26, we referred to other taxa of which we had only single Individuals. One of these, collected by John Green, was found ing shallow roadside drain near Wodonya, Vic. Tt was photographed, the trophus cleared in hypochlorite und drawn, and the animal provisionally called "7! Aaste’’’ in recognition pf Walter Koste’s contribution to rutifer research in Australian. The material was put aside until more specttiens could be found, Subsequently, a manuscript on Nigermin rotiters was received for. comment (by RIS) in Sept, 1992, in which anew trichacercid was described, Despite minor differences in traphus structure, it was apparently the same species ad te specimen from the Barnawartha-Wodonga Rd pool, and by remurkuble eoiicidence, it was named 7 keasreil ty can now be added tu the known VHehacerca species from Australia. Trichocerca kusiet Sepers Trichocercu kasiei Segers, 1993, p. 59-60, Fig. 24, Type lucaliry, Oguea Luke littoral, Imo State, Nigeria. Holotype: Female No, $21, Royal Museum for Central Africa (MRAC), Tervuren, Belgium. Paratypes: Two. temales (MRAC 822) and one female and one trophus preparation in the collection of the State University of Gheor, Belgium, Deseription; Body elongate, slightly curved, with dorsal keel lo ca, half body length (Fig. 1 a); broad lateral ciangular projection on tight side of head (dorsal), with ventral emargination: foot short; left toe 2X right, both curved; three substyll per toe; trophus (Fig, Ib, cl. the Victorian spécimen, ¢.) resembles that of 7 porcellus, T tigris, T uncinete; suongly usymmetric. with reduced right manubrinm and uncus; right ramus with distinctive (inger-like alula parallel to fulerum; left manubrivm large, terminally widened into booinerang shape, unici teeth fused, massive; fulcrum Jong, tertninutes in distal disc, BL. 178-196 pm; BW 46-54 am; LT to 108 pm, RT to 54 pm; TR 65-70 am. Distribation, Only two locations known tw date, the type locality in Nigeria and a Mowing (Zephemeral) roadside poal 10 km south of Wodonga on Wodonga-Bamawartha Rd, north- eastern Victoria (MDFRC Site #3194, 09.1X.91, Coll, JO Green), Vegetated littoral. 14.5°C, pH 88, 153 aS em! Comment: The single Victorian specimen has a longer teft manubrium (42 gm vs 40 pm). Vhis difference is accentuateil in the different perspectives dgawa below, ‘The apparent difference in the distal manubrium of the local specimen (Fig, le below), which is more sputulase phan thar of the type. mary alsu be a perspective urtefuct, Fig. {: Zhichocera bastela, ventral view: b, trophus, ventral, Nigerian specimen; c, trophus, dorsal, Victorian specimen. a, b after Seyors (1993), Acknowledginent For permission to use figures of T. kestei from Segers (1993), thanks to Henri Dumont, Bdilor, Aydrobjologia. Reference Sraers, H. (1993) Rotifera of some lakes in the floodplain of the River Niger (imo State. Nigeria). 1. New species wid other taxonomic considerations. Avdrobiolagia 250 6-61, ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED Patron: HER EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA DAME ROMA F. MITCHELL, D.B.E., QC., L.I B., DUniv. OFFICERS FOR 1993-94 President: W. D. WILLIAMS, B.Sc., Ph.D., D.Sc., Dip.Ed., F.A.I.Biol. Vice-Presidents: N. A. LOCKET, M.A., B.M., B.Ch., Ph.D., D.O. M. DAVIES, B.Sc., M.Sc., Ph.D. Secretary: Treasurer: J. F WALLMAN, B.Sc. R. H. WATERS, J.P., FA.I-M. Editor: Assistant Editor: J. BIRD, B.Sc. E. M. CAMPBELL, B.A.(Hons), M.A., Ph.D. Librarian: Programme Secretary: M. ANTHONY, A.L.A.A. R. G. SIMMS Minute Secretary: Membership Secretary: A. J. McARTHUR, B.E. G. A. CROOK, B.Sc. Members of Council: R. I. JAMIESON, B.Sc. P. G. STOTT, B.V.Sc., M.Sc., Dip.Ed. D. J. WILLIAMS M. J. WRIGHT, R.D.A. G. M. E. MAYO, B.Ag.Sc., Ph.D., D-Univ. Printed by Graphic Print Group, 10-14 Kingston Avenue, Richmond, S.A. 5033