VOL. 128, PART | 31 MAY, 2004 Transactions of the Royal Society of South Australia Incorporated INCORPORATING THE Records of the South Australian Museum Contents Berge, J. & Vader, W. Description of two new and unusual stegocephalid species (Crustacea: Peracarida: Amphipoda: Stegocephalidae) from Heard Island and the East ChinaSea - - - - - - = = | Jennings, J. T., Austin, A. D. & Stevens, N. B. Species of the Wasp genus Au/acus Jurine (Hymenoptera: Aulacidae) endemic to South Australia — - — 13 Turner, D. & Conran, J. G. The reproductive ecology of two naturalised Erica species (Ericaceae) in the Adelaide hills: the rise and fall of two ‘would-be’ weeds? - - - - - - 23 Zbik, M. & Pring, A. The Myrtle nee meteorite: a chondrite oy from South Australia — — - 3 Leppard, P. L., Tallis, G. M. & Pearce, C. E. M. Expected lifetime in South Australia IGE = bys Se ee Op ae lee ee 37 Riley, I. T. & Bertozzi, T. Variation in sex ratios in four Anguina (Nematoda: Anguinidae) species — - = - - 43 Smales, L. R. Syphacia (Syphacia) australasiensis sp. nov. (Nematoda: Oxyuridae) from Rattus leucopus (Muridae) from Papua New Guinea and Australia -— -— - - —- - - - = = 47 Jones, H. I. Gastric nematodes, including a new species of Abbreviata (Nematoda: Physalopteridae) from the mangrove monitor Faranus indicus (Reptilia: Varanidae) -— - - - - - = = = 53 Parsons, R. F. Enrichment-planting of the woody climbers Marsdenia australis and Rhynacharrhena Linearis in north-western Victoria. - -— — 61 Spratt, D. M. & Hobbs, R. P. Breinlia (Breinlia) ventricola sp. nov. (Nematoda: Filarioidea) from the heart of the red kangaroo, Macropus rufus, in Western Australia - - - - - = = = = = = 67 Phillott, A. D. & Elsmore, S. A. M. Black Noddies (Anous minutus) and Wedge-tailed Shearwaters (Puffinus pacificus) as potential hosts for fungi invading sea turtle nests at Heron Island, Queensland - —- — 73 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 INCORPORATING THE Records of the South Australian Museum VOL. 128, PART 1 EDITORIAL This is the first issue of the Transactions of the Royal Society of South Australia to incorporate the Records of the South Australian Museum. The Transactions have appeared uninterruptedly for 127 years, publishing good science from and for southern Australia and the world, and the Records have a similar record of achievement over 86 years. Why have the Society and the Museum merged these two successful journals, each with its proud record? The simple answer is that we believe that a single journal produced by the two institutions, which have many other beneficial linkages, can be better scientifically, can reach a bigger audience and can offer scientists a better platform for their research. In some fields, such as taxonomy, there has been much overlap, in others such as anthropology, little overlap in recent years, so that there is much complementary strength in the two journals. What we have done so far is simply to merge the journals and to plan to send the merged Transactions out to the subscribers to both journals. For this first year, there will be two issues. Future changes will depend particularly on contributors and readers, the two groups who make or break any journal. If we find strong authorial support, we shall need to consider more frequent publication, with all the benefits and costs inherent in such a change. Or we may find that there is a need to divide the journal into parts for readers with different interests, though for many (such as the current editor) this would be a step with the huge disadvantage of no longer providing a journal for generalist specialists with interests in all aspects of science as {l applies to southern Australia and related regions. The current issue, covering plant ecology, threatened species, parasitology, taxonomy, population science, and earth science, typifies this breadth and strength within the breadth. The future is in your hands. Support the merged Transactions as readers and contributors, and it will go from strength to strength. Oliver Mayo Editor, for the Editorial Board TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INC. CONTENTS, VOL. 128, 2004 PART 1, 31 MAY, 2004 Berge, J. & Vader, W. Description of two new and unusual stegocephalid species (Crustacea: Peracarida: Amphipoda: Stegocephalidae) from Heard Island and the East ChinaSea - - - - -—- = = = = = Jennings, J. T., Austin, A. D. & Stevens, N. B. Species of the Wasp genus Aulacus Jurine (Hymenoptera: Aulacidae) endemic to South Australia -— —- = = Turner, D. & Conran, J.G. The reproductive ecology of two naturalised Erica species (Ericaceae) in the Adelaide hills: the rise and fall of two ‘would-be’ weeds? Zbik, M. & Pring, A. The Myrtle Springs meteorite: a chondrite (H4) from South Australia — Leppard, P. I., Tallis, G. M. & Pearce, C. E. M. Expected lifetime in South Australia 1841 — 1996 Riley, I. T. & Bertozzi, T. Variation in sex ratios in four Anguina (Nematoda: Anguinidae) species Smales, L. R. Syphacia (Syphacia) australasiensis sp. nov. (Nematoda: Oxyuridae) from Rattus leucopus (Muridae) from Papua New Guinea and Australia - —- Jones, H. I. Gastric nematodes, including a new species of Abbreviata (Nematoda: Physalopteridae) from the mangrove monitor Varanus indicus (Reptilia: eg ty a Parsons, R. F. Enrichment-planting of the woody climbers Marsdenia australis and Rhyncharrhena Linearis in north-western Victoria - -— - - - = Spratt, D. M. & Hobbs, R. P. Breinlia (Breinlia) ventricola sp. nov. (Nematoda: Filarioidea) from the heart of the red kangaroo, Macropus rufus, in Western Australia -— — Phillott, A. D. & Elsmore, S. A. M. Black Noddies (Anous minutus) and Wedge-tailed Shearwaters (Puffinus pacificus) as potential hosts for fungi invading sea turtle nests at Heron Island, Queensland - - = PART 2, 30 NOVEMBER, 2004 Appan, A., Bergfeld, J. & Beveridge, I. New species of parasitic nematodes from macropodid marsupials in Western Australia - - - -= —~ = = = = = Hammer, M. P. & Walker, K. F. A catalogue of South Australian freshwater fishes, including new records, range extensions and translocations - - - - -— = Watts, C. H.S & Humphreys, W. F. Thirteen new Dytiscidae (Coleoptera) of the genera Boongurrus Larson 7firtudessus Watts & Humphreys and Nirripirti Watts & Humphreys, from underground waters in Australia — - - - Watts, C. H. S. Revision of Australian Scirtes Uliger and Ora Clark (Coleoptera: Scirtidae) — Hodda, M. Characteristics of an Australian population of Paraxonchium orthodon (Loof 1964) Altherr & Loof 1969, with a note on evolution and biogeography of the subfamily Paraxonchiinae (Nematoda, Dorylaimida, Aporcelaimidae) — Beveridge, I. & Johnson, P. M. Cestode parasites of tree kangaroos (Dendrolagus spp.: Marsupialia), with the description of two new species of Progamotaenia (Cestoda: Anoplocephalidae) - - - - - - ~~ - = Faucheux, M. J. Antennal sensilla of Sabatinca sterops Turner (Lepidoptera: Micropterigidae) Barker, S. Twelve new species of Australian Buprestidae (Coleoptera) and new synonymy - - ~- - = = = = = = = = = = Womersley, H. B. S. Additions to the Marine Algal Flora of southern Australia - -— -— = Smith, B. P. C. & Kokkinn, M. The use of emergence as an end-point for sediment toxicity tests using the Australian chironomid Chironomus maddeni — -— - = = Barton, P. S., Aberton, J. G. & Wishart, E. Spatial and temporal distribution of Culex australicus Dobrotworsky and Drummond and Culex globocoxitus Dobrotworsky (Diptera: Culicidae) at the Gippsland Lakes in eastern Victoria = — ~ = Williams, C. R., Snell, A. E. & Kokkinn, M. J. Studies of temporal host-seeking patterns of Culex Annulirostris (Diptera: Culicidae): a comparison of methods and populations tibbs, S. E., Kemper, C. M., Byard, R. W. & Long, M. Deaths of killer whales (Orcinus orca) in South Australia and implication of human interaction = — me Hinchcliffe, J. & Conran, J. G. The Tennyson sand dunes: vegetation structure and conservation status = — = 3 = Sie: ee eo a ass ee Smith, B. B. & Walker, K. F. Reproduction of common carp in South Australia, shown by young- of-the-year samples, gonadosomatic index and the histological staging of ovaries - - = = = = = = = = = = = = Brief Communications: Souter, N. J. A comparison among three artificial substrates for aquatic macroinvertebrate sampling - — =— RS Matthews, E. G. New synonymy and new names in Australian Tenebrionidae (Coleoptera) = — Insert to Transactions of the Royal Society of South Australia, Val. 128 Part 2, 30 November, 2004 77 85 99 131 169 249 259 261 DESCRIPTION OF TWO NEW AND UNUSUAL STEGOCEPHALID SPECIES (CRUSTACEA: PERACARIDA: AMPHIPODA: STEGOCEPHALIDAE) FROM HEARD ISLAND AND THE EAST CHINA SEA By J. BERGE! & W. VADER? Summary Berge, J. & Vader, W. Description of two new and unusual stegocephalid species (Crustacea: Peracarida: Amphipoda: Stegocephalidae) from Heard Island and the East China Sea. Trans. R. Soc. S. Aust. 128(1), 1-11, 31 May, 2004. Two new stegocephalid (Amphipoda) species are described: Mediterexis macho and Stegocephalina wolf. One of the species, Mediterexis macho, is the very first record of a stegocephalid from the China Sea, whereas Stegocephalina wolf was collected North of Heard Island in the Southern Ocean. Key Words: Amphipoda, Stegocephalidae, Mediterexis macho sp. _ nov., Stegocephalina wolf sp. nov. Transactions of the Royal Society of S. Aust. (2004), 128(1), 1-11. DESCRIPTION OF TWO NEW AND UNUSUAL STEGOCEPHALID SPECIES (CRUSTACEA: PERACARIDA: AMPHIPODA: STEGOCEPHALIDAE) FROM HEARD ISLAND AND THE EAST CHINA SEA. by J. BERGE! & W. VADER? Summary Berce, J. & VADER, W. Description of two new and unusual stegocephalid species (Crustacea: Peracarida: Amphipoda: Stegocephalidac) from Heard Island and the East China Sea. Trans. R. Soc. S. Aust. 128(1), 1-11, 31 May, 2004. Two new stegocephalid (Amphipoda) species are described: Mediterexis macho and Stegocephalina wolf. One of the species, Mediterexis macho, is the very first record of a stegocephalid from the China Sea, whereas Stegocephalina wolf was collected North of Heard Island in the Southern Ocean. Kry Worps: Amphipoda, Stegocephalidae, Mediterexis macho sp.nov., Stegocephalina wolf sp.nov. Introduction The Antarctic fauna of stegocephalid species was reviewed by Berge ef a/. in 2000, in which a total of 19 species were recognised. Since then two additional stegocephalid species have been recorded from the area, excluding the new species Stegocephalina wolf reported herein. Thus, the Antarctic stegocephalid fauna currently consists of 22 known species belonging to I! genera. The second new species herein described, is the first record of a stegocephalid species from the China Sea. A phylogenetic analysis of the family Stegocephalidae was presented as part of a recent revision of the group (Berge & Vader 2001). Based upon this analysis, a new classification at the generic level was outlined. As the classification at both generic and subfamily levels falls outside the scope of this paper, the two new species (Mediterexis macho sp.nov. and Stegocephalina wolf sp.nov.) described herein are treated within the framework of that classification. Especially for Mediterexis macho, the generic position may well be altered in the future (see also below). Material and Methods This study is based upon material from the South Australian Museum (SAM). All dissected appendages were mounted in polyvinyl-lactophenol and stained with rose-bengal. These appendages were drawn using a Leica compound microscope equipped with a drawing-tube, while the habitus-drawings were ‘UNIS, Department of Biology, Pb 156, 9170 Longyearbyen, NORWAY. Email: jorgen.berge@unis.no > Tromso Museum, Dept of Zoology, University of Tromso, 9037 Tromso, NORWAY. Email: wim@imy.uit.no made using a Leica dissecting microscope. Mature and immature females were distinguished from males by the presence of oostegites. The classification of setae and setae-groups follows that of Berge (2001). All scales attached to the figures are 0.1 mm unless otherwise stated. Symbols Al-2: Antenna 1-2; EP3: Epimeral plate 3; IP: Inner plate; L: Labium; LBR: Labrum; LMND: Left mandible; MX1: Maxilla 1; MX2: Maxilla 2; MXP: Maxilliped; OP: Outer plate; P1-7: Pereopods 1-7; PLP: palp; RMND: Right mandible; ST: Setal teeth on the first maxilla; T: Telson; U1-3: Uropods 1-3. Results and Taxonomy The present study is based entirely on material provided from the collections of the South Australian Museum. The material comprised, in addition to the two new species reported herein, a total of 3 species; Glorandaniotes sandroi Berge & Vader, 2003a, Parandania boecki Stebbing, 1888 and Tetradeion crassum (Chilton, 1883). The three above mentioned species were all found in Australia, whereas the two remaining, Mediterexis macho sp.nov. and Stegocephalinae wolf sp.nov., were collected from the East China Sea and Heard Island, respectively. Key to the 22 species known from the Antarctic and sub-Antarctic regions: 1. Pereopod 6 basis expanded, posterior margin convex (2) - Pereopod 6 basis weakly expanded, posterior margin straight or concave (17) - Pereopod 6 basis not expanded, about as broad as pereopod 5 basis (21) to i) Uropod 3 rami obsolescent or absent Stegocephalina pacis - Uropod 3 both rami well developed (3) 3. Telson entire (4) - Telson cleft (8) 4. Antennae subequal (5) - Antenna 2 elongate, longer than antenna | Parandania boecki 5. Antenna | flagellum with 5 articles Andaniexis ollii - Antenna | flagellum more than 10 articles (6) 6. Antennae elongate Parandania boecki (juveniles) - Antennae not elongate (7) 7. Labrum symmetrical, both reduced Parandania gigantea - Labrum asymmetrical, right lobe large, left lobe strongly reduced Parandania nonhiata 8. Rostrum weakly developed (9) - Rostrum large and distinct Stegocephalus rostrata 9. Coxa | anterior margin convex (10) - Coxa | anterior margin with a deep invagination Stegocephalus watlingi 10. Telson longer than broad, triangular and pointed (11) - Telson about as long as broad, not pointed (13) 11. Mouthparts not elongate and not forming a conical bundle (12) - Mouthparts elongate, forming a conical bundle projecting well below coxae Stegocephalina wolf sp.nov. 12. Epimeral plate 3 posteroventral corner acute, with one notch Stegocephalus kergueleni - Epimeral plate 3 posteroventral corner rounded, crenulated Genus Pseudo (see below) 13. Antenna | flagellum article | shorter or about as long as peduncle (14) - Antenna | flagellum article | distinctly longer than peduncle (15) 14. Coxae 1-3 broad and overlapping, coxa 4 posterior lobe exceeding pereon segment 6 Stegosoladidus antarcticus - Coxae 1-3 narrow, not overlapping, coxa 4 posterior lobe not exceeding pereon segment 6 Andaniotes linearis (immature) 15. Maxilla 2 gaping and geniculate, epimeral plate 3 posteroventral corner produced and with teeth Genus Pseudo (see below) - Maxilla 2 not gaping and geniculate, epimeral plate 3 posteroventral corner weakly produced, without teeth (16) 16. Antenna | flagellum with 4 articles Andaniotes pseudolinearis lobes strongly 'The validity of this genus is discussed in Berge & Vader (submitted), To avoid any nomenclatory complications, the classification according to Berge & Vader 2001 is utilised. J. BERGE & W. VADER - Antenna | flagellum with more than 10 articles Parandania boecki 17. Uropod 3 outer ramus 2-articulate (18) - Uropod 3. outer = ramus |-articulate Stegosoladidus ingens 18. Telson cleft (19) - Telson entire Andaniella integripes 19. Pereopod 6 basis posteromedially with 2-3 long plumose setae (20) - Pereopod 6 basis posteromedially with a row of short robust setae Andaniotes linearis 20. Coxae 1-3 broad and overlapping, maxilliped inner plate with 2 nodular setae Stegosoladidus debroyeri - Coxae 1-3 narrow, not overlapping, maxilliped inner plate with 4 nodular setae Andaniotes pooh 21. Telson entire (22) - Telson cleft (23) 22. Telson short, pereopod 4 subchelate, pereopod 7 well developed Parandaniexis dewitti - Telson long, pereopod 4 simple, pereopod 7 reduced 7etradeion crassum 23. Epistomal plate large, conspicuous Austrophippsia unihamata - Epistomal plate absent Schellenbergia! vanhoeffeni Remarks to the kev The key is rewritten and updated from that presented by Berge et a/. (2000) to include both the new Stegocephalina wolf sp.nov. (see below) and the two Pseudo Berge & Vader, 2001 species that were described, but not given a formal scientific name in Berge & Vader (submitted). Furthermore, all generic names in the key are updated according to the outlined classification of all stegocephalid taxa in the revision of the family by Berge & Vader (2001). Subfamily Andaniexinae Berge & Vader, 2001. Genus Mediterexis Berge & Vader, 2001. Mediterexis macho sp. n. (Figs 1-3) Holotype Male, 4 mm (SAM C6054), East China Sea (30° 06’ N, 130° 35’ E), 02.07.1966. Unique. Etymology The name refers to its densely setose peduncles of the antennae, compared to all other stegocephalid taxa. Diagnosis Antennae subequal, penduncles setose, accessory flagellum longer than antenna | flagellum article 1. Epistome laterally produced, epistomal plate present. Mandibles with transverse smooth incisors, left TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM Fig. |. Mediterexis macho sp.nov. Holotype. 4 J. BERGE & W. VADER Fig. 2. Mediterexis macho sp.nov. Holotype. TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM 5 Fig. 3. Mediterexis macho sp.nov. Holotype. lacinia mobilis weakly toothed. Maxilla 2 not gaping and geniculate. Labrum shorter than broad. Pereopod 6 basis posteriorly conspicuously expanded. Uropod outer ramus two-articulate. Telson entire. Description Head: Head retractable under pereonite 1. Rostrum rudimentary. Antennae: Accessory flagellum longer than flagellum article 1, articulation present. Antenna 2 peduncle article 3 short, about as long as broad. Peduncle article 4 shorter than article 5. Epistome: Epistome laterally produced and rectangular, with a long ridge on each side. Epistomal plate (medial keel) produced into a small elongate medial ridge covering the entire epistome. Mouthparts: Mouthparts not elongate. Mandibles with incisor transverse, smooth. Left lacinia mobilis present, weakly toothed and distally not produced. Maxilla 1 palp articulation absent, distally with long pappose setae. Outer plate distally rounded, ST in a 6/3 arrangement with two parallel rows: First row with ST 1-5 present, ST 6 absent and ST 7 present. Second row with ST A-C present. Inner plate with shoulder weakly developed, setae pappose. Maxilla 2 outer plate not gaping and geniculate, setae distally straight. Inner plate with setae-row A and B appressed. Row B with first 3 setae differentiated from the other setae. Row C present, row D absent. Maxilliped palp 4-articulate, article 2 not produced distally. Articles 1-3 with long slender setae. Inner plate distally with inner corner weakly produced, 2 nodular setae present. Outer plate with inner and outer setae-row present, both with robust slender setae. Distal setae-group absent. Labrum shorter than broad, both lobes reduced. Labium distally broad, distal finger absent. Pereopods and coxae: Coxal plates and basis of the pereopods smooth. Coxae 1-3 contiguous. Pereopod | coxal plate about as deep as basis, basis anterior margin weakly expanded. Pereopod 1 6 J. BERGE & W. VADER propodus subovate, posterior submarginal row of setae present. Pereopod 2 ischium elongate, distal posterior margin without plumose — setae. Propodus subrectangular, posterior submarginal row of robust setae present. Pereopod 4 coxa locking-structure absent. Basis with long setae on posterior margin, distally without plumose setae. Ischium with long plumose setae on distal posterior margin. Pereopod 6 basis posteriorly expanded, medially with a row of long plumose setae. Posterior margin of basis serrate. Pereopod 7 general appearance similar to pereopod 6. Basis medially with a row of short simple setae. Oostegites and gills: Gills present on pereopods 2 7. Oostegites unknown. Pleonites: Pleonites 1-3 dorsally smooth. Epimeral plate 3 weakly produced and rounded posteriorly, serrations absent. Urosome: Males with urosome not enlarged. Articulation between urosomites 2 and 3 present. Uropod | outer ramus with short robust setae on outer and inner margin, inner ramus with robust setae on inner margin, Uropod 2 outer ramus outer margin with robust setae, inner ramus inner margin with robust setae. Uropod 3 peduncle longer than telson, outer ramus 2-articulate. Rami without setae. Telson entire and rounded distally, submarginal setae on apex present. Females Unknown. Distribution Known only from the type locality, Remarks This species can easily be distinguished from all other stegocephalid taxa by the combination of 1) a laterally produced epistome, 2) the weakly differentiated last two pairs of pereopods, 3) entire telson and 4) the bi-articulate outer ramus of the third uropod. The presence of a not gaping and geniculate maxilla 2 and the smooth mandibular incisors indicate unequivocally that the species is a true member of the subfamily Andaniexinae Berge & Vader, 2001. However, it is not possible to assign it unequivocally to any of the genera of the family: its telson, mandibles and maxilliped resemble closely those found in the three genera Andaniexis Stebbing, 1906, Mediterexis Berge & Vader, 2001 and Parandaniexis Schellenberg, 1929. These three genera are also the only genera within the subfamily that do not possess a locking-process on the inner side anteriorly on the fourth coxa (see Berge & Vader 2003b), but the first maxilla and the weakly differentiated last two pairs of pereopods have close affinities to those found in the genus Andaniotes Stebbing, 1897. In addition, the second maxilla, with its slightly elongate outer plate and the arrangements of setae-rows A-D has closest affinities to the subfamily Andaniopsinae Berge & Vader, 2001. Thus, the assignment of this species to Mediterexis seems, at best, a temporary decision. However, as the phylogeny of the family and allocation of its species to genera fall outside the scope of this paper, this species is herein placed in Mediterexis based on the close affinities to that genus in the maxilliped, mandibles and epistome. This last character (see Fig 4, Mediterexis mimonectes (Ruffo, 1975) was used in the phylogenetic analysis (Berge & Vader 2001), but closer examination has revealed that the epistome shows a more complex set of characters than previously assumed (Berge, in prep.). As currently only one single specimen of this new species exists, its epistome was not available for SEM examination. However, examination under the light microscope gives reason to suspect that these two congeners possess a very similar epistome; that of M. mimonectes is herein illustrated. Subfamily Stegocephalinae Dana, 1852 Genus Stegocephalina Stephensen, 1925 Stegocephalina wolf sp. n. (Figs 5-7) Holotvpe Male 5mm (SAM C6055), Southern Ocean just North of Heard Island (52° 18'S, 73° 45’ E), 245 m, col. W. Zeidler, 13.06.1990, “Aurora Australis” st. 65, Sample taken from a sponge. Unique. Etvmology This species is named after the collector, Dr Wolfgang Zeidler, Adelaide. Diagnosis Rostrum rudimentary. Antennae — subequal, accessory flagellum shorter than antenna | flagellum article 1. Epistome laterally produced, epistomal plate present. Mouthparts forming a conical bundle. Mandibular incisors lateral, toothed. Maxilla 2 gaping and geniculate. Labrum elongate. Pereopod 6 basis posteriorly conspicuously expanded. Uropod 3 outer ramus two-articulate. Telson cleft. Description Head: Head retractable under pereonite 1. Rostrum rudimentary Antennae: Accessory flagellum shorter than flagellum article 1, biarticulate. Antenna 2 peduncle article 3 short, about as long as broad. Peduncle article 4 longer than article 5. Epistome: Epistome laterally produced, long and triangular. Epistomal plate (medial keel) produced TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM A Pig. 4. Mediterexis mimonectes Ruffo, 1975. SEM picture of the epistome. A: Epistomal plate, B: Epistome laterally produced, C: Left mandible, D: Labrum. into a small elongate medial ridge covering the entire epistome. Mouthparts: Mouthparts elongate. Mandibles with incisors lateral, toothed. Left lacinia mobilis present, strongly toothed and distally produced. Maxilla | palp articulation absent, distally with short simple setae. Outer plate distally subrectangular, ST in a 6/3 arrangement with two parallel rows: First row with ST 1-5 present, ST 6 absent and ST 7 present. Second row with ST A-C present. Inner plate with shoulder well developed, setae pappopectinate. Maxilla 2 outer plate gaping and geniculate, setae distally straight. Inner plate with setae-rows A and B appressed together. Rows C and D present. Maxilliped palp 3-articulate, dactylus absent. Articles | and 3 long and weakly setose, article 2 short. Inner plate elongate, 2 nodular setae present. Outer plate with inner and outer setae-row absent. Distal setae-group present, setae short and simple. Labrum conspicuously elongate, triangular. Left lobe reduced. Labium distally narrow, one distal finger present. Pereopods and coxae: Coxal plates and basis of the pereopods smooth. Pereopod | coxal plate about as deep as basis, basis anterior margin weakly expanded. Pereopod | propodus subovate, posteriorly without submarginal row of setae. Pereopod 2 ischium elongate, distal posterior margin with plumose — setae. — Propodus subrectangular. Pereopod 4 coxa locking-structure absent. Basis with long setae on posterior margin, distally with plumose setae on anterior and posterior margins. Ischium with long plumose setae posteriorly. Pereopod 6 basis posteriorly expanded, medially with a row of long plumose setae. Posterior margin of basis smooth. 8 J. BERGE & W. VADER Fig. 5. Stegocephalina wolf sp.nov. Holotype. TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM Fig. 6. Stegocephalina wolf sp.nov. Holotype. 10 J. BERGE & W. VADER Fig. 7. Stegocephalina wolf sp.nov. Holotype. Pereopod 7 conspicuously different from pereopod serrations absent. 6. Basis posteriorly smooth, medially with a row of Urosome: Articulation between urosomites 2 and 3 short simple setae. present. Uropod 1 outer ramus with robust setae on Oostegites and gills: Gills present on pereopods 2- outer margin, inner ramus with short robust setae on 7. Oostegites unknown. inner margin. Uropod 2 outer ramus with short Pleonites: Pleonites 1-3 dorsally smooth. Epimeral — robust setae on outer margin. Uropod 3 peduncle not plate 3 weakly produced and rounded posteriorly, as long as telson, outer ramus 2-articulate. TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM 1 Telson longer than broad, cleft and rounded distally, submarginal setae on apex present. Female Unknown. Remarks Stegocephalina wolf is separated from all other stegocephalid species by the combination of 1) the absence of dactylus on the palp of the maxilliped, 2) the fourth article of the peduncle of the second antenna longer than the fifth, and 3) the presence of only one distal finger on the labium. The present species has close affinities, both phylogenetically and morphologically, to the type species of the genus, S. ingo/fi Stephensen, 1925. Stephensen (1925:136) wrote that S. ingolfi is easily recognisable e.g. by the long and narrow mouthparts; the same is also true for S. wolf. The epistome and mouthparts are in both species conspicuously elongate, projecting well below the coxae. However, the present species differs from S. ingolfi in having a considerably shorter article one of the flagellum and only one distal finger on the apex of the labium. Acknowledgements We are grateful to Dr Wolgang Zeidler at the South Australian Museum for providing material. Also, we are indebted to Chris Jones and Prof. Geoff Boxshall at the Natural History Museum in London for their help and support in preparing the SEM pictures. The first author was funded by the Norwegian Research Council, project number 145384/432. References Berar, J. (2001) Revision of Stegosoladidus Barnard & Karaman, 1987 (Crustacea:Amphipoda: Stegocepha- lidae); redescription of two species and description of three new species. Journal of Natural History 35, 539-571. , De BRoyver, C. & VADER, W. (2000) Revision of the Antaretic and sub- Antarctic species of the family Stegocephalidae (Crustacea: =Amphipoda) — with description of two new species. Bulletin de l'Institut des Sciences Naturelles de Belgique 70, 217-233. — & VAber, W. (2001) Revision of the amphipod te rustacea) family Stegocephalidae. Zoological Journal of the Linnean Society 133, 531-592. & (2003a) Stegocephalid (Crustacea: ~ Amphipoda) species from Australia and New Zealand, with description of seven new species. Records of the Australian Museum 55, 85-112. = & (2003b) Metandania tordi n.sp. - anew stegocephalid (Crustacea: — Peracarida: Amphipoda) species from the Southern Ocean. Proceedings of the Biological Society of Washington 116(4), 1007-1016. & _ (submitted) Two new Antarctic stegocephalid (Amphipoda: Stegocephalidae: Stego- cephalinae) species, with implications for the phylogeny and classification of the two genera Pseudo and Schellenbergia. Deep-sea Research Part II. CHILTON, C. the New Zealand Crustacea. Transactions and Proceedings of the New Zealand Institute 15, 69-86. (1883) Further additions to our knowledge of DANA, J. D. (1852) Conspectus crustaceorum quae in orbis terrarum circumnavigatione, Carolo Wilkes e classe Reipublicae Faederatae Duce, lexit et descripsit. Jacosus D. DANA. Paris III. Proceedings of the American Academy of Arts and Sciences 2, 201-220. Haswett, W. A. (1879) On Australian Amphipoda. Proceedings of the Linnean Society of New South Wales 4(3), 245-279. Rurro, S. (1975) Nuovi Lisianassidi e Stegocefalidi del Mediterraneo (Crustacea, Amphipoda). Bollettino del Museo Civico di Storia Naturale di Verona 1 (1974), 439-453. SCHELLENBERG, A. (1929) Abyssale und Pelagische Gammariden. polio of the Museum of Comparative Zoology LXIX (9), 193-201. STEBBING, T. R. R. (1888) Report of the Amphipoda collected by H.M.S. Challenger during the years 1873- 76. Report on the Scientific Results of the Voyage of H.M.S. Challenger During the Years 1873-76 Zoology 29: 1737 pp. London: Eyre and Spottiswoode. — (1897) Amphipoda from the Copenhagen Museum and other sources. Transactions of the Linnean Society of London Zoology 7, 25-45. (1906) Amphipoda_ I. ~— Tierreich 21, 806p. STEPHENSEN, K. (1925) Crustacea Malacostraca. VI. (Amphipoda. II.). The Danish Ingolf-Expedition 3 (9), 101-178. Gammaridea. Das SPECIES OF THE WASP GENUS AULACUS JURINE (HYMENOPTERA: AULACIDAE) ENDEMIC TO SOUTH AUSTRALIA By J. T. JENNINGS’, A. D. AUSTIN? & N. B. STEVENS? Summary Jennings, J. T., Austin, A. D. & Stevens, N. B. Species of the Wasp genus Aulacus Jurine (Hymenoptera: Aulacidae) endemic to South Australia. Trans. R. Soc. S. Aust. 128(1), 13-21, 31 May, 2004. This study deals with the South Australian species of Aulacus Jurine, a fauna that comprises four geographically isolated, endemic species. Aulacus moerens Westwood is redescribed, and three new species (A. belairensis, A. flindersbaudini and A. grossi) are recognised. A diagnosis for the genus is presented, along with notes on taxonomic history and species diversity for Australia. Key Words: Evanioidea, Aulacidae, Aulacus, taxonomy, parasitic wasps. Transactions of the Royal Society of S. Aust. (2004), 128(1), 13-21. SPECIES OF THE WASP GENUS AULACUS JURINE (HYMENOPTERA: AULACIDAE) ENDEMIC TO SOUTH AUSTRALIA. by J. T. JENNINGS!, A. D, AUSTIN? & N. B. STEVENS? Summary JENNINGS, J. 'T., AusTIN, A. D. & Stevens, N. B. Species of the Wasp genus Au/acus Jurine (Hymenoptera: Aulacidae) endemic to South Australia. Trans. R. Soc. Aust. 128(1), 13-21, 31 May, 2004. This study deals with the South Australian species of Aulacus Jurine, a fauna that comprises four geographically isolated, endemic species. Aulacus moerens Westwood is redescribed, and three new species (A. belairensis, A. flindersbaudini and A. grossi) are recognised. A diagnosis for the genus is presented, along with notes on taxonomic history and species diversity for Australia. Kry Worbs: Evanioidea, Aulacidae, 4u/acus, taxonomy, parasitic wasps. Introduction The Aulacidae are a family of parasitic wasps that are endoparasitoids of wood-boring wasps (Xiphydriidae) and beetles (Cerambycidae and Buprestidae) (e.g. Carlson 1979; Gauld & Bolton 1988; Smith 2001). Although at various times the Aulacidae have been treated as either a subfamily or group of the Evaniidae s./, most recent studies regard them as a distinct family (e.g. Naumann 1991; Mason 1993; Gauld 1995; Jennings & Austin 2000; Smith 2001). Like other members of the Evanioidea, aulacids are characterised by the high insertion of the metasoma on the propodeum, but these wasps are also readily distinguished by the presence of fore wing vein 2m- cu (Gauld & Bolton 1988). Worldwide, the Aulacidae comprise 156 valid species (Smith 2001), but this is likely to be only one-third of the true size of the group. Smith (2001) has provided a catalogue of the world fauna, and included 48 species of Aulacus Jurine, 106 species of Pristaulacus Kieffer, and two species of Panaulix Benoit. Both Aulacus and Pristaulacus are worldwide in their distribution, but Panaulix is confined to sub-Saharan Africa. The Australian fauna comprises 34 described species (18 Aulacus and 16 Pristaulacus) (Smith 2001; Jennings 2001), but most taxa are inadequately described and date from the works of Kieffer (1911; 1912). No taxonomic studies have been undertaken on the Australian fauna since Crosskey (1953) described one Aulacus and two Pristaulacus species, even though modern collecting techniques, in particular Malaise trapping, have added significantly to the ' Centre for Evolutionary Biology and Biodiversity and School of Agriculture and Wine, The University of Adelaide, Glen Osmond, SA 5064, Australia. Email john.jennings@adelaide.edu.au “Centre for Evolutionary Biology and Biodiversit Farth and Environmental Science, The University of Adelaide, SA 5005, Australia. y, and School of number of specimens held in collections. As part of a project that aims to revise the aulacid fauna of Australia, this study deals with the unique and isolated Adlacus fauna of South Australia. Aulacus moerens Westwood is redescribed, and three new species from the Fleurieu Peninsula, Kangaroo Island, and the far north-west of South Australia are described. At the same time, the genus is redescribed and notes provided on taxonomic history and species diversity. Materials and Methods Specimens were observed under a Zeiss light microscope or using scanning electron microscopy (SEM). Specimens for SEM were first cleaned to remove obvious dirt and other debris and examined uncoated under a Phillips XL30 field emission SEM at Ikv and a spot size of three. Terms for general morphology follow Jennings and Austin (1994), and for wing venation follow the modified Comstock-Needham system, after Sharkey (1988), but with some modifications, and using the nomenclature of van Achterberg (1979) for cells. Terms for surface sculpturing follow Harris (1979). Where measurements are based on more than one specimen, data are presented as the mean followed by the range. The length of the ovipositor is measured from the tip of the metasoma. Abbreviations for institutions which are repositories of the specimens referred to in this paper are: Hope Entomological Collections, Oxford (OXUM), South Australian Museum, Adelaide (SAMA), and Waite Insect and Nematode Collection, The University of Adelaide (WINC). Systematics Aulacus Jurine, 1807 Aulacus Jurine 1801: 163. [nom. nud. | Aulacus Jurine 1807: 89. Type species: Aulacus 14 J.T. JENNINGS, A. D. AUSTIN & N. B. STEVENS striatus Jurine 1807, by monotypy (North America and Europe). — Blanchard 1840: 300; Schletterer 1889: 489; Kieffer 1903: 383, 453; Bradley 1908: 120; Kieffer 1912: 344, 370; Hedicke 1939: 17; Townes 1950: 113; Townes 1951: 659; Oehlke 1983: 441: Koslov 1988: 243: Alekseev 1995: 39; Konishi 1990: 638; Smith 2001: 268. Disphaeron Dahlbom 1837: 175. Type species: Aulacus arcticus Dahlbom 1837, by monotypy. — (syn. Hedicke 1939: 18). Aulacinus Westwood 1868: 331. Erected as a sub- genus of Aulacus. Type species: Aulacus (Aulacinus) moerens Westwood 1868, by monotypy. — Kieffer 1903: 481; Bradley 1908: 120; Kieffer 1912: 349; Hedicke 1939: 24. (syn. Townes 1950: 113). Pammegischia Provancher 1882: 302. Type species: Pammegischia burquei Provancher 1882, by monotypy. — Kieffer 1903: 383; Bradley 1908: 120; Kieffer 1912: 346; Townes 1938: 254; Hedicke 1939: 23. (syn. Kieffer 1902: 11). Parafoenus Kieffer 1910: 350. No_ species included. Type species: Parafoenus formosus Kieffer 1912, by subsequent monotypy. — Kieffer 1912: 345; Hedicke 1939: 26. (syn. Townes 1950: 113). Neuraulacinus Kieffer 1910: 350. Type species: Neuraulacinus vespiformis Kieffer 1910, by subsequent designation, see Kieffer 1912: 358. (designated from three species included by Kieffer 1911). — Kieffer 1912: 358; Hedicke 1939: 26. (syn. Townes 1950: 113). Micraulacinus Kieffer 1910: 350. Type species: Micraulacinus elegans Kieffer 1910, by subsequent monotypy, see Kieffer 1912: 348. — Kieffer 1912: 348; Hedicke 1939: 24. (syn. Townes 1950: 113). Disaulacinus Kieffer 1910: 350. Type species: Disaulacinus flavimanus Kieffer 1911, by subsequent monotypy, see Kieffer 1911: 224. — Kieffer 1912: 361; Hedicke 1939: 23. (syn. Crosskey 1953: 759). Pycnaulacus Cushman 1929: 17. Type species: Pycnaulacus brevicaudus Cushman 1929, by original designation. — (syn. Townes 1950: 113). Diagnosis based on Australian species Head with or without occipital carina; frons with or without a transverse carina above antennal sockets; antenna 14-segmented in female, 13-segmented in male; antennal insertions low on face, near lower margin of eyes; eyes small, circular or subcircular, remote from the mandibles; scape usually deeply convex in lateral view, much thicker than pedicel and flagellomeres; metapostnotum present between propodeum and = metanotum as a_ distinct sclerotisation; propodeum pyramidal, metasoma inserted high on the apex; metasomal first tergite (T1) and second tergite (T2) fused dorsally; hind coxa with (Fig. 16) or without a groove or notch on inner ventral surface, the apposed grooves or notches forming an ovipositor guide; hind trochanter with a transverse trochanteral groove; prefemur (trochantellus) present (Figs 11-12); each tarsal claw with one basal tooth (sometimes difficult to see); fore wings not plicate at rest; fore wing vein 2m-cu present, vein 2r-m often absent, largely spectral when present, vein 3r-m present, often largely spectral (Figs 1, 3, 5); ovipositor exserted, protruding well beyond apex of metasoma. Comments Aulacus was first erected as a genus by Jurine (1801), but as a nomen nudum since no species were included. In 1807, Jurine rectified this by describing Aulacus striatus. Early authors did not define the generic limits of Au/acus, so a number of genera were erected (see above) without due consideration of interspecific variation. This resulted in a proliferation of genera: Disphaeron Dahlbom, Disaulacinus Kieffer, and Micraulacinus Kieffer from Australia; Neuraulacinus Kieffer from Australia and South America; Parafoenus Kieffer from South America; and Pycnaulacus Cushman from the USA. By 1912, Kieffer had recognised some 41 species in six genera and the subgenus Aalacinus Westwood. Of these, 17 species were Australian. Although Kieffer (1902) had synonymised Pammegischia Provancher with Aulacus, the mainly North American Pammegischia were not transferred to Au/acus until much later (Townes 1938). In his 1939 catalogue, Hedicke synonymised Disphaeron and included a number of additional species in the other genera. Townes (1950) evaluated various characters such as wing venation, and in the process, defined the generic limits of Aulacus. This led him to synonymise Aulacinus, Micraulacinus, Neuraulacinus, Parafoenus, and Pycnaulacus with Aulacus. Townes erroneously synonymised Disaulacinus with Aulacostethus Philippi (now Pristaulacus). Disaulacinus was later synonymised with Aulacus by Crosskey (1953). At present, 48 species worldwide are included in Aulacus; 18 species are endemic to Australia (Smith 2001; Jennings 2001). Most of the latter are from the higher rainfall, forested areas of the east coast of Australia, including Tasmania. Here we treat the geographically isolated fauna of South Australia and distinguish them from the known Australian taxa. Because there are many undescribed species from south-western and eastern Australia, a key to Australian species would be premature. Aulacus belairensis sp. nov. (FIGS 1-2, 7, 11, 13, 16-17, 19) Material Examined Holotvpe ©, Belair N. P., I1-18.i1.1996, J.T. Jennings, SAMA. AULACUS OF SOUTH AUSTRALIA 15 Figs 1-6. Fig. 1. Right fore wing of Aulacus belairensis sp. nov. holotype female. The cells are: | = marginal, 2 = submarginal, 3 = discal, 4 = subdiscal, 5 = costal, 6 = basal, 7 = subbasal, 8 = plical. Fig. 2. Right hind wing of Au/acus belairensis sp. nov. holotype female. Fig. 3. Right fore wing of Aulacus flindersbaudini sp. nov. holotype male. Fig. 4. Right hind wing of Aulacus flindershaudini sp. nov. holotype male. Fig. 5. Right fore wing of Aulacus grossi sp. nov. holotype female. Fig. 6. Right hind wing of Aulacus grossi sp. nov. holotype female. Scale bars = | mm. 16 J, 1. JENNINGS, A. D. AUSTIN & N. B. STEVENS Kips 7-12. Dorsal views of head. Fig. 7. Aidacus belairensis sp. nov, holotype female. Pig. 8. lulacus flindershaudini sp- nov. holotype male. Pig. 9. du/acus grossi sp. nov. holotype female. Pig. 10. du/acus moerens holotype female. Lateral views of hind (rochanter and prefemur Pig. 11. flindershaudini sp. nov. holotype male. Scale bars = 200 pum 7. 9; 500 pum & 10; 50 jum 11, Paratypes SA: 1 @, Belair National Park, 10-17.i11,1996, J.T, Jennings, WINC; | 2, Belair National Park, xii. 1996, J.T. Jennings, WINC. Female Length: 5.0 mm (4.4 — 5.5 mm), excluding ovipositor. Colour: Head orange, with variable amounts of dark brown around ocelli; scape and pedicel orange with flagellomeres dark brown; propleuron dark tulacus belairensis sp. nov. holotype female. Pig. 12. dulacus 12 brown, body black except for variable amounts of orange on pronotum; legs predominantly light to dark brown; metasoma dark brown: ovipositor sheaths black, ovipositor orange; wings fuscous, darker apically on the marginal and submarginal cells. Tlead: 1.2 (1.0 — 1.3) x wider than long when viewed dorsally; face rugose, with a few large punctures near eye margin, pubescence short; frons without transverse carina above toruli, rugose, with a few punctures near eye margin, pubescence short; IWLACUS OV SOUTH AUSTRALIA \7 Figs 13-18. Fig. 13. Mandibles and elypeus of sidacus belairensis sp, nov. holotype female. Fig. [4, Dorsal view of mesoscutuim, scutellum and axillae of dadacus belairensis sp. nov. holotype female. Fig. 15, Dorsal view of mesoscutum. scutellum and axillae of dudacis flindersbaudini sp. nov. holotype male. Fig, 16. Ovipositor guide on hind coxae of Aulacuy helairensis sp. nov. holotype female. Fig. 17. Dorsal view of metasomal Tt and T2 of dalacuy hekrirensis sp. noy, holotype female. Fig. 18. Dorsal view of metasomal TH and T2 of -liddcus grossi sp. nov. holotype female, Scale bars = 200 pom 13,17, 18: 500 pam T4153 100 pam 16. vertex and gena punctate-imbricate, a few radiating striations near ocelli, with scattered short setae (Hig 7); posterior margin of head not concave in dorsal view; malar space 0.25 x height eye; clypeus 0.37 x as wide as high, margin sinuate with small medial process (Fig. 13); mandibles broad, with two large medial teeth (Fig. 13); distance from lateral ocellus to eye margin 0.83 (0.78 — 0.90) x distance between lateral ocelli; scape 1.6 (1.6 — 1.7) x length pedicel; first Magellomere 1.0 (0.9 — 1.1) x as long as scape, equal to length second flagellomere. Mesosoma. Propleuron rugulose, pubescence long, ventro-lateral carina weak; pronotum rugose; mesoseutum in lateral view rounded antero-dorsally, medial and lateral lobes rugose-punctate, with scattered setae, admedial lines present (Fig. 14): scutellum and axillae rugose-punctate (Fig. 14): mesepisternum reticulate, with short pubescence, mesepimeron broad, scrobiculate; metapleuron rugose, with short pubescence; propodeum rugose, Is J.T. JENNINGS, A. D. AUSTIN & N. B. STEVENS Autacus belairensis A, flindersbaudini A, grossi A. moerens oreo Fig. 19. Distribution of South Australian Au/acus spp. posterior margin scrobiculate; hind coxa with ovipositor guide ventrally on inner margin (Fig. 16); hind trochanter and femur imbricate, pubescence short; prefemur somewhat indistinct (Fig 11); hind tibia imbricate, pubescence short, with scattered emergent stout setae; hind femur 0.70 (0.68 — 0.72) x length hind tibia; hind tibia with ventro-apical pecten of short robust spines; hind tarsal segments 1-4 with ventro-apical pecten of short robust spines, segment 1, 2.8 x length segment 2; segment 2, 1.4 x length segment 3; segment 3, 1.6 x length segment 4; segment 4, 0.6 x length segment 5; hind tarsal claw 0.5 x length segment 5; fore wing vein 2r-m and 3r- m largely spectral (Fig. 1); hind wing with 3 hamuli (Fig. 2). Metasoma. Clavate, 1.67 (1.54 — 1.71) x length of mesosoma; Tl and T2 anteriorly narrow, smooth dorsally except for a few strigate wrinkles anteriorly on Tl; ovipositor 5.9 (5.0 — 6.5 mm) mm. Male Unknown. Remarks This species is named after the holotype locality, Belair National Park, South Australia (Fig. 19). All specimens were collected by Malaise trap, and nothing is known of its biology. This species can be separated from other South Australian species by the fuscous wings that are darker apically on the marginal and submarginal cells, and the strigate wrinkles anteriorly on metasomal Tl. Also, it is similar in size and appearance to 4. e/egans (Kieffer) which has been collected from several localities in New South Wales and Victoria. The two can be easily separated in that 4. elegans has a shiny head, an orange metasoma, and has different sculpturing patterns; for example, the vertex is largely smooth with just a few scattered shallow punctures. Aulacus flindersbaudini sp. nov. (FIGS 3-4, 8, 12, 15, 19) Material Examined Holotvpe 3d, West Bay, Flinders Chase Nat. Park, Kangaroo Is. S.A., 1.1986, A.D. Austin, SAMA. [Flagellomeres 2-11, right hind tarsal segments 4-5, and claw missing]. Male Length: 8.0 mm. Colour: Body red except metasomal TI dark brown dorsally; flagellomeres 2 and 3 black; wings hyaline; fore wing with brown spot apically on the marginal and submarginal cells. Head: |.6 x wider than long when viewed dorsally; face punctate-imbricate, pubescence short; frons without transverse carina above toruli, areolate- rugose, pubescence short; vertex areolate-rugose medially, with a few radiating striations near ocelli, with a few transverse striations posteriorly, punctate- imbricate laterally, pubescence short (Fig. 8); gena punctate-imbricate, with scattered short setae; posterior margin of head slightly concave in dorsal view; malar space 0.34 x height eye: clypeus 3.7 x as wide as high, margin sinuate with small medial process; mandibles broad, with two large medial teeth; distance from lateral ocellus to eye margin equal to distance between lateral ocelli; scape 1.8 x length pedicel; length first flagellomere equal to length scape, second flagellomere missing. Mesosoma. Propleuron smooth anteriorly to weakly rugulose posteriorly, with scattered long setae: ventro-lateral carina weak; pronotum rugulose; mesoscutum in lateral view rounded antero-dorsally, medial and lateral lobes strigate with scattered short setae, admedial lines present (Fig. 15); scutellum and axillae strigate (Fig. 15); mesepisternum —areolate-rugose, with — short pubescence; mesepimeron broad, scrobiculate: metapleuron areolate-rugose, with short pubescence; propodeum areolate-rugose, posterior margin scrobiculate; hind coxa without groove ventrally on inner margin, weakly rugulose to strigate dorsally, pubescence short laterally; hind trochanter imbricate, pubescence short; prefemur indistinct (Fig 12); hind femur imbricate, pubescence short; hind tibia imbricate, pubescence short, with scattered emergent stout setae; hind femur 0.77 x length hind tibia; hind tibia with ventro-apical pecten of short robust spines: hind tarsal segments 1-4 with ventro-apical pecten of AULACUS OF SOUTH AUSTRALIA 19 short robust spines, segment |, 2.0 x length segment 2; segment 2, 1.6 x length segment 3; segment 3, 2.4 x length segment 4; segment 4, 0.8 x length segment 5: hind tarsal claw 0.7 x length segment 5; fore wing vein 2r-m incomplete, evident only by a small node on 2r, 3r-m entirely spectral (Fig. 3); hind wing with 2 hamuli (Fig. 4). Metasoma. Clavate, 1.5 x length of mesosoma; T1 and T2 dorsally narrow, smooth; digitus about same length as basiparameres. Female Unknown. Remarks Aulacus flindershaudini is named to commemorate the meeting of Matthew Flinders and Nicolas Baudin at Encounter Bay, South Australia in 1802, This species is known only from the holotype location (Fig. 19), and nothing is known of its biology. This species is readily distinguished from other South Australian species by the presence of a brown spot apically on the marginal and submarginal cells of the fore wing (Fig. 3); this spot is absent in the other species. Aulacus flindersbaudini also lacks the eroove or ovipositor guide on the inner margin of the hind coxae. The brown spot on the fore wing is found in several species from south-western and eastern Australia, such as 4. pallicaudus (Cameron), but all of these species are largely black. Aulacus grossi sp. Nov. (FIGS 5-6, 9, 18-19) Material Examined Holotype 9°. nr. Victory Well, Everard Pk Stn, S. Aust. 2- 4.xi.1970. G. Gross, SAMA. Female Length: 4.5 mm, excluding ovipositor. Colour: Head black; mesosoma, antennae, and legs light brown; metasoma black except for basal part of metasomal T1; wings hyaline. Head: \.\ x wider than long when viewed dorsally: face, frons without transverse carina above toruli: frons and vertex rugose-punctate (Fig. 9), with scattered short setae, a little denser on face; gena smooth, almost glabrous; posterior margin of head not concave in dorsal view; malar space 0.16 x height eye; clypeus 4.0 x as wide as high, punctate, margin sinuate with small medial process; mandibles broad, with two large medial teeth; distance from lateral ocellus to eye margin 0.72 x distance between lateral ocelli; scape 1.8 x length pedicel; first flagellomere 0.72 x as long as scape, 0.52 x as long as second flagellomere. Mesosoma. Propleuron weakly rugulose-punctate, almost smooth, with a few scattered setae; ventro- lateral carina weak; pronotum rugose-punctate; mesoscutum in lateral view rounded antero-dorsally, medial and lateral lobes rugose, almost strigate, with scattered short setae, admedial lines distinct: scutellum and axillae rugose, almost strigate; mesepisternum rugose-punctate, with — short pubescence; mesepimeron broad, scrobiculate; metapleuron rugose-punctate, with short pubescence; propodeum rugose, with several pronounced carinae, posterior margin scrobiculate; hind coxa with groove ventrally on inner margin, weakly strigate dorsally, pubescence short; hind trochanter and hind femur imbricate, pubescence short; hind tibia imbricate, pubescence short, with scattered emergent stout setae; hind femur 0.8 x length hind tibia; hind tibia with ventro-apical pecten of short robust spines; hind tarsal segments 1-4 with ventro-apical pecten of short robust spines, segment 1, 2.9 x length segment 2; segment 2, 1.35 x length segment 3; segment 3, 2.0 x length segment 4; segment 4, 0.5 x length segment 5; hind tarsal claw 0.5 x length segment 5; fore wing veins 2r-m and 3r-m largely spectral (Fig. 5); hind wing with 2 hamuli (Fig. 6). Metasoma. Clavate, equal in length to mesosoma; Tl and T2 dorsally broad, smooth (Fig. 18); ovipositor 4.2 mm. Male Unknown. Remarks Aulacus grossi has a broad metasoma when viewed dorsally which distinguishes it from both 4. belairensis and A. flindersbaudini. It is also the smallest Australian species of Aulacus known, and can be distinguished from all other Australian species by its colour pattern, i.e. head and metasoma black, and mesosoma light brown. Au/acus grossi is known from a single specimen from Everard Park Station in the north-west of South Australia (Fig. 19), and is named after the collector, Dr Gordon Gross, formerly Curator of Entomology, South Australian Museum. Nothing is known about its biology. Aulacus moerens Westwood (FIGS 10, 19) Aulacus (Aulacinus) moerens Westwood 1868: 331. — Westwood 1874: 129; Schletterer 1889: 517. Aulacinus moerens — Kieffer 1902: 12; Kieffer 1903: 481: Kieffer 1904: 6; Kieffer 1912: 349, 356; Hedicke 1939; 25, Aulacus maerens — Dalla Torre (unnecessary emendation) Aulacus moerens — Smith 2001: 273. 1902: 106] 20 J.T. JENNINGS, A. D. AUSTIN & N. B. STEVENS Material Examined Holotype 2, Adelaide [illeg.] 1865, OXUM. Antennae missing, head glued to mesosoma, metasoma glued to card, ovipositor sheaths missing. Female Length: 10.5 mm, excluding ovipositor. Colour: Black, except tibiae and tarsi largely light brown, ovipositor light brown; wings hyaline. Head: |.18 wider than long when viewed dorsally; face rugulose, pubescence long; frons with pronounced transverse carina above toruli, rugulose, with long pubescence; vertex punctulate-imbricate, slight rugosity posteriorly near occiput, with short scattered setae (Fig. 10); gena punctulate-imbricate, with scattered short setae; posterior margin of head not concave in dorsal view; malar space ().2 x height eye; clypeus 4.0 x as wide as high, margin sinuate; mandibles broad, with two large medial teeth: distance from lateral ocellus to eye margin 0.9 x distance between lateral ocelli; antennae missing. Mesosoma. Propleuron rugulose, pubescence long, ventro-lateral carina present; pronotum without angular process, rugose; mesoscutum in lateral view angular antero-dorsally, medial and lateral lobes strigate, with scattered short setae, admedial lines present; scutellum — strigate; axillae rugose; mesepisternum rugose, with long pubescence; mesepimeron broad, scrobiculate; metapleuron rugose, with short pubescence; propodeum rugose, with medial dorso-ventral scrobiculate groove, posterior margin scrobiculate; hind coxa without groove ventrally on inner margin, strigate dorsally, pubescence long laterally; hind trochanter punctulate-imbricate, pubescence short; hind femur imbricate, pubescence short; hind tibia imbricate, pubescence short, with scattered emergent stout setae; hind femur 0.74 x length hind tibia; hind tibia without ventro-apical pecten of short robust spines; hind tarsal segments 1-4 with ventro-apical pecten of short robust spines, segment |, 2.7 x length segment 2; segment 2, 1.6 x length segment 3; segment 3, 2.0 x length segment 4; segment 4, 0.4 x length segment 5; hind tarsal claw 0.6 x length segment 5; fore wing vein 2r-m pale brown, tubular, 3r-m spectral in middle half; hind wing with 3 hamuli. Metasoma. Ovate, 1.25 x length of mesosoma; T1 and T2 smooth except slight rugosity medially; ovipositor 4.0 mm. Male Unknown. Remarks Although both 4. grossi and A. moerens have an ovate metasoma, the latter species is a much larger and differs in a number of ways, including its largely black colour, the lack of an ovipositor guide on the hind coxae, and the presence of a small medial process on the clypeus. It does not resemble any other Australian species. This species is known only from the holotype locality, Adelaide, South Australia (Fig. 19), and nothing is known of its biology. Acknowledgments We thank the curators of the collections listed above for the loan of specimens, and the staff of Adelaide Microscopy, The University of Adelaide, for access to the FESEM. This work was carried out with the assistance of an Australian Biological Resources Study grant and a Small ARC grant to JTJ and ADA. References ALEKSEEV, V. N. (1995) Evanioidea pp. 37-45 /n Lehr, P.A. (Ed.) [“Keys to Insects of Russian Far East in Six Volumes”. Vol. IV. Neuropteroidea, Mecoptera, Hymenoptera. Part 2. Hymenoptera] (Dal’nauka, Vladivostok) [in Russian]. BLANCHARD, E. (1840) “Histoire Naturelle des Animaux Articulés, Annelides, = Crustacés, — Arachnides, Myriapodes et Insectes”, Vol. 3. (Paris). BRADLEY, J. C. (1908) The Evaniidae, ensign-flies, an archaic family of Hymenoptera. Trans. Am. ent. Soc. 34, 101-194, plates V-XV. Carson, R. W. (1979) Aulacidae pp. IITI-I11S Jn Krombein, K.V., P.D. Hurd, Jr, D.R. Smith & B.D. Burks (Eds.) “Catalog of Hymenoptera in America North of Mexico”. Volume I. (Smithsonian Institution Press, Washington, D.C.). Crosskey, R. W. (1953) Two new species of Aulacostethus Philippi and a new species of Aulacus Jurine, from Australia, together with a key to the Australian species of Aulacostethus (Hymenoptera: Aulacidae). Ann. Mag. nat. Hist. (12) 6, 758-766. CusHMAN, R. A. (1929) New species of ichneumon-flies and taxonomic notes. Proc. U. S. natn. Mus. 76, 1-18. DAHLBOM, G. (1837) Species Aulaci Generis in Scandinavia habitantes. /sis. Jena 3, 173-177. DALLA Torre, C. GG. de (1902) “Catalogus Hymenopterorum hucusque descriptorum systematicus et synonymicus”, Volumen Ill: Trigonalidae, Megalyridae, Stephanidae, Ichneumonidae, Agrio- typidae, Evaniidae, Pelecinidae. Pars II., pp. 545-1141. (Lipsiae). AULACUS OF SOUTH AUSTRALIA 21 GAULD, I. D. (1995) Aulacidae pp. 192-193 /n Hanson, P.E. & 1.D. Gauld (Eds.) “The Hymenoptera of Costa Rica” (Oxford University Press, Oxford). & Boiron, B. (Eds.) (1988) Hymenoptera”. (Oxford University Press, Oxford). Harris, R.A. (1979) A glossary of surface sculpturing. Calif. Dept. Food Agric., Bureau Entomol., Occ. Papers “The 28, 1-28. Hepickre, H. (1939) Aulacidae /nm Hedicke H. (Ed.) “Hymenopterorum Catalogus”, Pars 10. (Dr. W. Junk, Gravenhage). JENNINGS, J.T. (2001) Aulacidae. [Checklist of Australian species] Australian Faunal Directory, Australian Biodiversity Information Facility, © Environment Australia. (http://www.environment.gov.au/abrs/online/r esources/abif-fauna). & Austin, A. D. (1994) Revision of the genus Crassifoenus Crosskey (Hymenoptera: Gasteruptiidae: Hyptiogastrinae), with a description of a new species from Western Australia. Rec. West. Aust. Mus. 16, 575- 591. (2000) Higher-level phylogeny of the Alagidae and Gasteruptiidae (Hymenoptera: Evanioidea) pp. 154-164 /n Austin, A.D. & M. Dowton (Eds.) “Hymenoptera. Evolution, — Biodiversity and Biological Control”. (CSIRO Publishing, Collingwood). JuRINE, LL. (1801) Nachricht) von einen Neuen Entomolischen Werks, des Hrn. Prof. Jurine in Geneve. Intellig. Litt.-Z., Erlangen 1, 161-165. : (1807) “Nouvelle Méthode de Classer les Hyménopteres ct les Dipteres. Hyménopteres”. Tome Premier. (Geneve). 319 pp. Kierrer, J.-J. (1902) Hymenoptera, Fam. Evaniidae /n Wytsman, P. (Ed.) “Genera Insectorum”, Fascicule 2, 13 pp., | plate. (Bruxelles). (1903) Les Evaniides pp. 347-482 /m André, E. “Species des Hyménopteres d’Europe & d’Algéric”. Volume 7, part 2 (Paris). (1904) Description de Stéphanides et ~ d@evaniides nouveaux. Bull. Soc. Hist. Nat. Metz (1, 1-30. (1910) Diagnoses de nouveaux genres ~ d’Aulacinae (Hym.). Bull. Soc. ent. Fr 1910, 350. (1911) Etude sur les Evaniides exotiques (Hym.) du British Museum de Londres. Ann. Soc. ent. Fr, 80, 151-231. (1912) Hymenoptera, ~ Evaniidae. Das Tierreich, 30, 1-431. Konisil, K. (1990) A revision of the Aulacidae of Japan (Hymenoptera, Evanioidea). Jap. J. Ent. 58, 637-655. Ichneumonidae, Kostov, M. A. (1988) Aulacidae pp. 242-244 /n Medvedev, G.S. (Ed.) [“Keys to the Insects of the European Part of the USSR”], Vol. II, Hymenoptera, Part VI Symphyta. Akademia Nauk SSSR, Zoologischkie Institut 267 pp. [Translation, 1994, E.J. Brill, Leiden]. MAson, W. R. M. (1993) Chapter 11. Superfamilies Evanioidea, Stephanoidea, Megalyroidea, and Trigo- nalyoidea pp. 510-520 /n Goulet, H. & J.T. Huber (Eds.) “Hymenoptera of the World: An Identification Guide to Families”. (Research Branch, Agriculture Canada). Naumann, I. D. (1991) Chapter 42. Hymenoptera (Wasps, bees, ants, sawflies) pp. 916-1000 / Naumann, I. D. (Ed.) “The Insects of Australia, A Textbook for Students and Research Workers”. Volume II. (Melbourne University Press). OriLKe, J. (1983) Revision der curopaischen Aulacidae (Hymenoptera — Evanioidea). Beitr, Ent., 33, 439-447. PROVANCHER, A. L. (1882) Faune Canadienne, Hyménoptéres, Additions et Corrections. Naturaliste can. 13, 289- 4 Il. SCHLETTERER, A. (1889) “Die Hymenopteren-Gruppe der Evaniiden”. II]. Abteilung. Analen des K. K. Naturhistorischen Hofmuseums, Separatabdruck aus band IV pp. 373-546. SHARKEY, M. (1988) Ichnews I, 2-12. SmtrH, D. R. (2001) World catalog of the family Aulacidae (Hymenoptera). Contrib. Ent., Int. 4, 263-319. Townes, H. K. (1938) Pammegischia and Trichofoenus discarded (aulacoid Hymenoptera). Can. Ent. 70, 254- 255. Ichneumonoid wing venation. (1950) The Nearetic species of Gasteruptiidae (Hymenoptera). Proc. U. S. natn. Mus. 100, 85-145. : (1951) Aulacinae pp. 657-660 /n Muesebeck, C.F. W., K. Vv. Krombein, & H. K. Townes (Eds.) “Hymenoptera of America North of Mexico, Synoptic Catalog.” (United States Department of Agriculture) Agriculture NonoErapy No. 2. VAN ACHTERBERG, C. (1979) A revision of the Subfamily Zelinae auct. (Hymenoptera, Braconidae). Tijdschr. Ent. 122, 241-479. Westwoop, J. O. (1868) XVIII. Descriptions of new genera and species of exotic Hymenoptera. Trans. ent. Soc. Lond. 1868, 327- 332. : (1874) “Thesaurus Entomologicus Oxoniensis; or, illustrations of new, rare, and interesting insects, for the most part contained in the collections “presented to the University of Oxford by the Rev. F. W. Hope, M. A., D.C.L., FL R.S. &c, with forty plates from drawings by the author” (Oxford). THE REPRODUCTIVE ECOLOGY OF TWO NATURALISED ERICA SPECIES (ERICACEAE) IN THE ADELAIDE HILLS: THE RISE AND FALL OF TWO ‘WOULD-BE’ WEEDS? By D. TURNER’? & J. G. CONRAN’? Summary Turner, D. & Conran, J. G. (2004). The reproductive ecology of two naturalised Erica species (Ericaceae) in the Adelaide Hills: the rise and fall of two ‘would-be’ weeds? Trans. R. Soc. S. Aust. 128(1), 23-31, 31 May, 2004. The pollination biology and reproductive ecology of the newly naturalised species Erica cinerea from Europe and E. glandulosa from South Africa (Ericaceae) are reported from the Adelaide Hills from surveys undertaken in 1995 and 2003. Examination of UV floral reflectance, nectar composition and floral visitors found that E. cinerea was honeybee pollinated (although the bees also practiced nectar theft by chewing through the corolla tube), whereas E. glandulosa was probably pollinated by New Holland Honeyeaters (Phylidonyris novaehollandiae: Meliphagidae) which filled the niche of the nectariniid Sunbirds which pollinate it in South Africa. Key Words: Erica, E. cinerea, E. glandulosa, Ericaceae, weed biology, reproductive ecology, pollination, fecundity, Australia. Transactions of the Roval Society of S. Aust. (2004), 128(1), 23-31. THE REPRODUCTIVE ECOLOGY OF TWO NATURALISED ERICA SPECIES (ERICACEAE) IN THE ADELAIDE HILLS: THE RISE AND FALL OF TWO ‘WOULD-BE’? WEEDS? by D. TurNER!? & J. G. CONRAN!? Summary TURNER, D. & Conran, J. G. (2004). The reproductive ecology of two naturalised Erica species (Ericaceae) in the Adelaide Hills: the rise and fall of two ‘would-be’ weeds? Trans. R. Soc. S. Aust. 128(1), 23-31, 31 May, 2004. The pollination biology and reproductive ecology of the newly naturalised species Erica cinerea from Europe and E. glandulosa from South Africa (Ericaceae) are reported from the Adelaide Hills from surveys undertaken in 1995 and 2003. Examination of UV floral reflectance, nectar composition and floral visitors found that EF. cinerea was honeybee pollinated (although the bees also practiced nectar theft by chewing through the corolla tube), whereas £. glandulosa was probably pollinated by New Holland Honeyeaters (Phylidonyris novaehollandiae: Meliphagidae) which filled the niche of the nectarintid Sunbirds which pollinate it in South Africa. Although fecundity data in 1995 suggested that there was considerable capacity for these plants to become weeds, the 2003 survey found that they had declined significantly, apparently as a result of competition or shading from taller, more aggressive weeds, mainly Bracken (Preridium esculentum) and Tree Erica (E. arborea). Nevertheless, as £. cinerea reseeds and £. glandulosa resprouts after fire, and £. cinerea can persist in the soil seed bank for over 30 years, their populations may recover and/or expand if the area where they are growing 1s opened up by fire. Key Woros: Erica, FE. cinerea, E. glandulosa, Ericaceae, weed biology, reproductive ecology, pollination, fecundity, Australia. Introduction Erica L. is a large genus of heaths which is centred around South Africa (Guthrie & Bolus 1905; Baker & Oliver 1967), with a second radiation in southern and western Europe (Webb & Rix 1972). Much of Southern Africa and Mediterranean Europe is characterised by cool winters and hot dry summers similar to South Australia and most members of the genus grow in nutrient poor, acidic soils. The European species, in particular, tend to be mycorrhizal, and the majority are calcifuges (Gimingham ef a/. 1979; Webb 1986). In many respects the soils of the South African Cape, Mediterranean Europe and South Australia are similar in being nutrient poor, often acidic, and support conspicuous sclerophyllous heath elements, represented in South Africa and Europe by the Ericaceae and in Australia by Epacridaceae (Specht 1979, 1981). There are three Erica spp. listed as naturalised in South Australia (Jessop & Toelken 1986): the European £. arborea L. and E. lusitanica Rudolphi, and the South African E. baccans L. Of these, the two European species, and £. arborea in particular, ' Centre for Evolutionary Biology and Biodiversity, Environmental Biology, School of Earth and Environmental Sciences, Darling Building DP418, The University of Adelaide, SA 5005. ° Present address: SARDI, SA Aquatic Sciences Centre, 2 Hamra Ave, West Beach, SA 5024. ‘Correspondence author: john.conran@adelaide.cdu.au. are widespread weeds on nutrient-poor, acidic soils with moderate to high rainfall (Cshures & Edwards 1998; Ojeda 1998). Erica spp. tend to coincide growth with water availability during winter and early spring (Gehrig 2000), flowering from late winter into spring and summer (Jessop & Toelken 1986). However, in 1995 populations of an additional two species (the European Bell Heather £. cinerea L. and South African £. glandulosa Thunb.) were found in a small area of disturbed remnant bushland bordering Mount Lofty House on the north- western boundary of Mount Lofty Botanic Gardens. Erica cinerea from temperate, western Europe has numerous ornamental cultivars grown in southern Australia (Spencer 1997) and is a temperate, severe cold-tolerant, generally summer-flowering species from low, open heaths on acidic soils (Rozé 1993; Ojeda et al. 1998; Leith et al. 1999). In contrast, E. glandulosa is a long-lived, frost-tolerant, year-round but mainly autumn-flowering, species from tall fynbos on well-drained soils in the southern Cape region of South Africa between Mosselbaai and Port Elizabeth (Baker & Oliver 1967). E. glandulosa is cultivated as an ornamental shrub in South Africa and Europe (Baker & Oliver 1967; Nelson 1997), and although not currently listed as grown in Australian gardens (Spencer 1997), is naturalised in Australia, but with no locality or weed status details (AFFA 2001). As these two species seem only to have naturalised recently in the Adelaide Hills, the populations were 24 Spencer Gulf ie ep art km / Fleurieru \ é Mount Lofty Botanic Gardens meee < wee a * v on e ¢ 250m Fig. 1. Mount Lofty Botanic Gardens site showing the locality of the Erica cinerea and E. glandulosa colonies. relatively small and confined to the one area. However, as they are morphologically dissimilar and come from different environments they may have different establishment strategies. Accordingly, the two aims of this study were: To compare the reproductive biology of the two species to assess how their strategies might relate to any success or failure to establish. To examine their population structure with particular emphasis on their potential fecundity, and to see if expansion had occurred after eight years. Methods The site Mt Lofty Botanic Gardens is a 97 ha reserve located in the Adelaide Hills 13 km SE of Adelaide (34° 58’ S, 138° 42’ E, 670 m altitude) combining ). TURNER & J. G. CONRAN extensive planted areas of exotics with a flora reserve of native Eucalyptus obliqua L’Hér. forest with sclerophyllous understorey (Haegi & Morley 1991). The area surveyed represents an approximately 10 x 100 m area of disturbed remnant eucalypt forest located on the NW side of the Gardens, close to the boundary fence with the grounds of Mount Lofty House (Fig. 1). The site understorey is a mixture of Epacridaceae, mainly Pink Ground Berry (Acrotriche fasciculiflora (Regel) Benth.) and Native Heath (Epacris impressa Labill.), other native shrubs, bracken fern (Preridium esculentum (G. Forst.) Cockayne) and introduced shrubby weeds including Spanish Broom (Cy/iszs scoparia (L.) Link) and Tree Erica (Erica arborea Lai Floral advertisement and rewards Advertisement for pollinators can involve several simultaneous stimuli including visual, olfactory, and short range tactile cues, and may be considered as any stimulus which attracts visitors to a flower (Facgri & van der Pijl 1979). Visual stimulus is a combined effect of the target colour (including combinations of colour and pattern) and the size shape of the target. Of primary importance is the pollinator’s visual spectrum (the visual spectrum of insects differs from that of humans) which affects the contrast between the flower and the background foliage (Dafni 1992; Dyer 1996). Flowers of the two species were photographed using Kodak P-3200 B&W film bracketted at f8-f22 with and without a UV transmitting filter (Hoya U360) by a Pentax MEII Super with macro lens and G30-rated flash. The film was developed following — manufacturer’s instructions for 10 minutes at 21°C, allowing visualisation of the flowers from the perspective of different potential pollinators. Because floral scent may be ephemeral, flowers were sealed in 30 mm glass vials for two hours in a warm, well-lit location and the vials were then opened and smelt (Bernhardt 1995). In addition, flowers were stained in 1% neutral red for 2 hours and rinsed in distilled water for 18 hours to detect possible osmophore-bearing regions (Datni 1992; Bernhardt 1995). Nectar volume was measured using micro-sampling pipettes and electronic balances, with ten flowers from each of the two species sampled at various times of the day to allow for temporal variation in nectar production. Nectar composition of the two species was examined using thin layer paper chromatography. Nectar from five flowers of each species was extracted by washing with 5 wl of deionised water, and spotted onto a sheet of Whatman’s No. | chromatography paper. Sugar standards were also added for glucose, ECOLOGY OF TWO ERICA SPP. IN THE ADELAIDE HILLS 25 sucrose, fructose, maltose, and galactose and the chromatogram run using a solution of 12:3:5 n- butanol: acetic acid: water. After 24 hours, the paper was removed from the solvent, dried, sprayed with an indicator solution of 1% aniline, 1% diphenylamine and 4% phosphoric acid in acetone and baked at 85°C for five minutes (Neimietz & Hawker 1988). The relative intensity of each nectar component was coded according to the nectar classifications of Percival (1961) and Baker & Baker (1983). Floral visitor behaviour was observed between 9 am and 4 pm (opening hours at Mt Lofty Gardens) over several days in Feb.-Mar. 1995, Sept. 2002 and Feb.-Mar. 2003 with observations mainly between 9 am and | pm, as pollinator activity was minimal in the afternoon. Birds were monitored from about 10 m using binoculars, and insects were netted, killed in jars with ethyl acetate fumes, examined under a binocular microscope and then washed briefly on a microscope slide with drops of absolute ethanol to remove any pollen. The resulting residue was mounted in a drop of 2.5% saturated aqueous Methyl Green and 2% saturated aqueous Phloxine in phenol glycerine jelly (Owezarzak 1952), and pollen identified by light microscopy. Population structure and fecundity Numbers of individuals of each of the two species were counted in order to obtain a total population count for the infested area. Twenty individuals of each species were then chosen at random, and the following information recorded for each: Number of shoots per plant Number of inflorescences on each shoot Number of flowers on each inflorescence Number of ovules per ovary Number of seeds per ovary Results Floral advertisement and rewards Erica glandulosa is an erect species >] m tall with large, orange-pink flowers in the leaf axils (Fig. 2B) which are classed as tubular-curved (Rebelo ef al. 1985) whereas FE. cinerea is a ground hugging shrub with inflorescences of small violet-purple flowers (Fig. 2C) which are classified as urceolate (Rebelo ef al. 1985). The violet-purple flowers of E. cinerea create an obvious contrast between the flower and_ its background (Fig. 3A). The corollas were classed as UV-light (pale) (Dafni 1992), with reflected shorter wavelength light making them high-contrast to the foliage. Reflectance was highest in buds and newly- opened flowers, with older (possibly no-longer receptive) flowers being less reflective and more like the foliage. Erica glandulosa has orange-pink flowers on a green background making for high visual contrast, but as the flowers reflect wavelengths outside the normal insect visual spectrum, they are likely to be low contrast to an insect (Richards 1986; Dyer 1996). The UV photographs showed low contrast between the flowers and foliage, although again newly-opened flowers were more reflective than older ones, and the lobes of recently-opened flowers (which visually showed a light greenish tinge), tended to be more UV reflective. No obvious scent or evidence of osmophores was found in E. glandulosa suggesting that the plant’s advertisement strategy does not involve scent. Although no obvious odour was detected in F. cinerea, two possible scent producing areas were identified. The edges of the sepals stained darkly and microscopic examination revealed specialised tissues of uncertain function, and the toothed anther appendages also stained heavily. Erica glandulosa contained a mean of 0.5+0.2 ul of nectar; whereas although minute droplets were noticed on the annular nectary below the ovary in E. cinerea, the volume was too minute to measure accurately. Chromatography of FE. glandulosa nectar revealed strong spots for glucose and fructose, with a weak spot for sucrose making the species SFG using the nectar composition coding system of Percival (1961) and hexose-rich following Baker & Baker (1983), whereas E. cinerea possessed all three sugars, but with a slight predominance of sucrose (SFG), thereby making it sucrose-rich. Flowers of £. glandulosa were visited in the mornings by New Holland Honeyeaters (Meliphagidae: Phylidonvris —novaehollandiae (Latham, 1790). These perched along the erect stems approaching the flowers from below, their curved beaks enabling them to access easily the slightly downward-curved, tubular flowers. The only observed visitors to E. cinerea were honey bees (Apis mellifera L., 1758)), and inspection of captured visiting bees found they carried only pollen from that species. Erica cinerea flowers were also observed to have had the corolla tubes chewed open from the side, suggesting that bees were also engaged in reward theft (Fig. 2D). Population structure and fecundity In the 1995 survey, there were 67 mature individuals of E. glandulosa aggregated as small groups of 2 — 5 plants and scattered across an area of about 1000 m’, but with no evidence of juvenile or seedling plants in the area surveyed. In addition, there were 1,436 mature (flowering) individuals of E. cinerea within an area of about 80 m2, as well as numerous seedlings. Within this, there was a smaller 26 D. TURNER & J. G. CONRAN ECOLOGY OF TWO ERICA SPP. IN THE ADELAIDE HILLS 27 27 m2? area of more open Acrotriche fasciculiflora and) Epacris impressa heath on the eastern (roadward) side which contained about 50 mature £. cinerea m?, and up to about 100 seedlings m? whereas the surrounding area contained only about four mature individuals and 10 — 20 seedlings m=. In contrast, by 2002, whereas there were 54 mature F. glandulosa plants still scattered over the same area, the numbers of EF. cinerea had fallen to only 34 adult plants in total, all within the original 27 m? area which had previously contained the bulk of the population (Table 1), and there was no evidence of seedlings. Data from the first survey suggested that the populations are highly fecund, with numerous seeds per individual and high total estimated annual seed set for both taxa (Table 1). Nevertheless, in 2003 not only had populations of both species declined significantly, but fecundity was also reduced, mainly through fewer flowers and inflorescences per plant, resulting in a dramatic decrease in estimated total seed set for both species at the colony level. Discussion The area that the two species occupy is in the north-western corner of the Botanic Gardens and although part of a native flora reserve, is directly below Mount Lofty House and was previously part of those gardens. Possible scenarios for their initial establishment are that these two Erica spp. were planted on the bank at some time in the past and have persisted as a remnant population, or that they spread from the original gardens into the surrounding bushland. Evidence from pollination studies in a number of ericaceous genera and Erica spp. in_ particular suggest that they are predominantly self-sterile (Anderson ef al. 2000; Ng & Corlett 2000) or with limited self-compatibility (Aparicio & Garcia- Martin 1996; Santandreu. & Lloret 1999). Nevertheless, F. cinerea is regarded as both self- and cross-pollinating (Knuth 1909; Bannister 1965), with the slightly protogynous flowers effecting pollen transfer before the buds open (Hagerup 1951). The pollination biology of the two species studied here shows the different strategies in insect- and bird-pollinated Ericaceae. That pollination is successful at Mount Lofty is reflected in the high levels of seed set, and the plants have clearly benefited from the adaptation of local pollinators to a new food source. This might also explain the scarcity of nectar in the SA flowers despite reports of copious nectar in European studies (Bannister 1965). Erica cinerea with relatively small, pale-purple, urceolate flowers falls into the entomophilous syndrome common in the genus (Knuth 1909; Rebelo ef al. 1984; 1985). Insect visitors to European Erica spp. include honeybees, bumblebees, flies, moths and butterflies (Knuth 1909; Aparicio & Garcia-Martin 1996; Santandreu & Lloret 1999) and Knuth (1909) listed a wide range of insects visiting £. cinerea, and reported corolla chewing and nectar theft by bumblebees. He also noted that pollination in this species was identical to that of E. tetralix L., where honeybees chew through the floral tubes because their c. 6 mm long mouthparts were too short to reach the nectaries of average-sized flowers (7 mm — the same as E. TABLE |. Summary of fecundity parameters for the two Erica spp. naturalised at Mt Lofty. E. cinerea E. glandulosa 1992 2003 1992 2003 Parameter n Mean+SD Mean+SD tor x? Mean+SD Mean+SD _ t or x? Plants per colony 1,436 34 1,428.0" 67 54 2.5 ns Main stems per plant 20 5.0+1.7 4.741.9 0.7 ns 5.2+6.9 5.0+£7.3 0.2 ns Inflorescences per plant 20) -18.8+8.2 10.3412.2 8.4 29.44+35.5 21.5+40.1 4.0" Inflorescences per main stem 20 = 3.9+1.5 2.6+1.8 3.2" 6.2+1.1 4141.8 33" Flowers per inflorescence 100 6.6+4.8 6.2+5.7 2.5" 3.2+1.0 3.041.2 1.3 ns Ovules per ovary 10 24.041.4 22.8+2.1 2.028 ns 708.0+122.3 692.9+173.1 1.7 ns Seeds per ovary 10 3.41.4 3.141.6 O5ns 215.0452.5 208.2+74.6 1.9 ns Seeds per plant (estimated) 422 153 n/a 20,227 13,430 n/a Total seeds per year (estimated) 605,992 5,202 n/a 1,355,209 725,220 n/a ‘counts compared using x? Fig. 2. Erica cinerea and E. glandulosa habitat and habit. A. Habitat of Eucalyptus obliqua forest with sclerophyllous shrub and bracken understorey. B. Erica glandulosa habit. C. E. cinerea habit. D. E. cinerea flower showing damage by nectar thieves. 28 D. TURNER & J. G. CONRAN Fig. 3. Photographs of the two Erica spp. under visible (A, C) and UV (B, D) light. A-B. Erica cinerea. C-D. E. glandulosa. cinerea). Nectar composition was similar to other European Erica species listed by Percival (1961) as visited extensively by honeybees, and the sucrose- rich nectar agrees with Baker & Baker’s (1983) observation that bee-flowers with floral tubes > 6 mm are mainly “long-tongued” bee-flowers and generally sucrose-rich. Most Erica spp. are generally considered to be scentless, although there is a number of species reported with marked honey scents (Rebelo e/ a/. 1985). The anther appendages in £. cinerea are thought to position the anthers for entomophilous pollen delivery (Knuth 1909; Rebelo et al. 1985) but the presence of apparently glandular structures which stain strongly with neutral red suggests that they may also serve as osmophores, and warrants further investigation. In contrast, £. glandulosa has orange-pink, tubular-curved, scentless flowers with copious nectar (Rebelo ef a/, 1984; 1985), and the hexose- dominated nectar in E. glandulosa reflected the results found by Barnes ef a/. (1995) for several ornithophilous species of Erica sect. Evanthe. The members of sect. Evanthe are thought to have co- evolved for pollination with nectariniid Sunbirds ECOLOGY OF TWO ERICA SPP, IN THE ADELAIDE HILLS 29 (Scott-Elliot 1890; Rebelo ef al. 1984; 1985), which fill the same ecological niche as the honeyeaters seen visiting it in our study, although there is evidence also of pollination of some Evanthe spp. in South Africa by long-proboscid Nemestrinid flies (Manning ef al. 1999), Although, the assertion by Baker & Baker (1983) that hexose-rich bird flowers are associated with passerines must be treated with some caution, given the dominance of sucrose-rich to sucrose-dominant Evanthe spp. reported by Barnes ef al. (1995), numerous South African passerine-pollinated flowers do produce high volumes of very dilute, hexose-dominated nectar (Nicolson 2002). The orange-pink flowers showed only limited UV-reflectance, suggesting that they are not strongly adapted for insect pollination (Richards 1986; Dyer 1996), although tubular red flowers can be attractive to long-proboscid flies (Manning ef a/. 1999). The glandular hairs which cover all parts of this species except the corolla possibly help to deter would-be insect nectar thieves, similar to those ornithophilous Erica spp. with externally viscid corollas (Scott-Elliot 1890; Rebelo ef al. 1985). Similarly, the presence of only anatomically rudimentary anther appendages in this species (Palser & Murty 1967; Hermann & Palser 2000) supports the assertion by Knuth (1909) and Rebelo e¢ a/. (1985) that they are involved in presenting the anthers primarily for entomophilous pollination. Nevertheless, if there is an anther appendage - osmophore connection, the loss of appendages in E. glandulosa would further support pollination by passerines with a poor sense of smell. What is most apparent from this study is that despite initial apparent success, with high levels of both numbers of individuals and/or annual fecundity, neither species has managed to spread; in fact both have declined in terms of above ground biomass. This is particularly the case for FE. cinerea where this previously abundant plant is now very scarce at the site, and the remaining plants are smaller and with lower fecundity than in 1995. The most likely explanation for this decline in £. cinerea is a combination of competition (Bannister 1965) and litter accumulation (Mallik ef a/. 1984). Shading can determine species composition in heaths and cause removal of understorey species (Vila & Sardans 1999), and crowding or shading by neighbours reduced fecundity in Erica multiflora L. (Vila & Terradas 1995, 1998). In 1995 E. cinerea was most abundant in a small area of relatively open epacrid-dominated heath, but by 2003 this, like the rest of the surveyed area, had been overgrown by Preridium and E. arborea, the latter of which outgrows Acrotriche fasciculiflora under dry conditions (Gehrig 2000), Erica cinerea prefers more temperate, mesic conditions than £. arborea (Ojeda er al. 1998) and would be disadvantaged under the hot dry summers seen in South Australia. Similarly, it grows mainly in low open heaths (Bannister 1965; Rozé 1993). whereas E. arborea generally grows better under forest cover or shade (Ojeda e¢ al. 2000). In contrast, E. glandulosa was apparently able to compete better for space, being of similar size to the invading bracken and shrubs, although crowding may have reduced fecundity, as seed set in resprouter Erica species is correlated with main shoot size and vigour (Riba 1998). Establishment in £. cinerea is mainly by seed, and flowering is improved by higher nitrogen levels (Leith ef al. 1999), although seed set and germination rate drops with population age (Mallik et al. 1984). Germination occurs in the presence of light, with 20 — 40% of the seeds germinating the season after they are shed, nevertheless, heat shock substantially increases germination rates (Bannister 1965), whereas litter accumulation inhibits germination and seedling survival (Mallik e7 al. 1984). Erica cinerea grows preferentially in short open heath, and recovers more slowly to from fire than taller heaths (Rozé 1993), although its seeds can survive for at least 30 — 40 years in the soil seedbank (Mallik ef a/. 1984; Thompson & Band 1997). Erica seeds in the upper 2 — 5em soil depth tend to survive fire better than many other Mediterranean woody heath species (Ferrandis ef al. 1999), but although £. cinerea can produce up to a million seeds m~, there is seedbank mortality of 20 — 40% after the first year, as well as low seedling survival (Webb 1986). In contrast, F. glandulosa is a resprouter (Ojeda 1998) — an unusual condition amongst Cape Erica species — although germination in many Cape Erica spp. is also smoke-triggered (Brown ev al. 1993), and it is possible that the absence of seedlings of this species, despite the large numbers of seeds produced, may be related to the lack of a suitable trigger, and/or the fact that many fire-dependent heath species do not germinate well under conditions of high litter and standing biomass (Rozé 1993: Lloret & Vila 1997). The breeding system of this species is as yet unknown, and further studies would be useful to determine whether it is cross-, self-pollinated or both, as this has implications for the ability of isolated individuals to set seed and spread further. In conclusion, the evidence suggests that although the populations of these two species appear to be in decline, despite their initial success and apparently high potential fecundity, were a fire or other disturbance to alter the conditions at the site, they may well be able to recover or expand, via the seedbank, representing a future weed problem awaiting an environmental trigger. However, as 30 D. TURNER & J. G. CONRAN bracken and £. arborea also recolonise quickly after fire or other disturbances, and frequent, repeated aerial biomass removal of the latter is needed to reduce its aggressive regrowth capacity (Riba 1998), it is possible that they may prevent further spread by the two new Erica spp. although this will require long term monitoring of the site, especially if the area were to be burnt in the future. Nevertheless, in either scenario, the real threat is to the native sclerophyllous understorey vegetation, as there are additional and apparently successful introduced competitors for the limited resources at the site. Acknowledgements The former Department of Botany, now Discipline of Environmental Biology within the School of Earth and Environmental Sciences, The University of Adelaide is thanked for the provision of resources for this project, the initial phase of which was undertaken by DT as part of a B.Sc. (Hons) degree. The director of the Adelaide Botanic Gardens and staff of the Mount Lofty Botanic Gardens are thanked for permission to undertake the research in the gardens. References AFFA (2001) “Annual Report 2000-2001" (Department of Agriculture, Fisheries and Forestry, Australia, Canberra). ANDERSON, G. J., BERNARDELLO, G., LOPEZ, P., SruESSyY, T. F. & Crawrorp, D. J. (2000) Dioecy and wind pollination in Pernettya rigida (Ericaceae) of the Juan Fernandez Islands. Bot. J. Linn. Soc. 132, 121-141. Aparicio, A. & GARCIA-MARTIN, F. (1996) The reproductive biology and breeding system of Erica andevalensis Cabezudo & Rivera (Ericaceae), an endangered edaphic endemic of southwestern Spain - implications for its conservation. Flora 191, 345-351. Baker, H. A. & Obtver, E. G. H. (1967) “Ericas in Southern Africa” (Purnell and Sons, Cape Town). Baker, H. G. & Baker, L. (1983) Floral nectar sugar constituents in relation to pollinator type pp. 117-141 /n Jones, C. E. & Little, R. J. 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(1984) Seed dynamics in Callina-Acrtostaphylos heath in north- eastern Scotland. J. Ecol. 72, 855-871. MAnnina, J. C., GoLpsiarr, P. & WINTER, P. (1999) Two new species of Gladiolus (Iridaceae: Ixioideae) from South Africa and notes on long-proboscid fly pollination in the genus. Bothalia 29, 217-223. Nemmurrz, C. & HAWKER, J. S. (1988) Sucrose accumulation in red beet vacuoles by UDP glucose-dependent group translocation - fact or artefact? Austral. J. Pl. Physiol. 15, 359-366. Newson, E. C. (1997) Erica pp. 485 /n Cullen, J., Walter, S. M.. Brady, A., Brickell, C. D., Green, P. S., Lewis, J., Matthews, V. A., Webb, D. A., Yeo, P. F. & Alexander, J. C. M. (Eds) “The European Garden Flora, Vol. V. Limnanathaceae to Oleaceae - Dicotyledons (Part III)” (Cambridge University Press, Cambridge). Na, S.C. & Cortetr, R. T. (2000) Genetic variation and structure in six Rhododendron species (Ericaceae) with contrasting local distribution patterns in Hong Kong, China. Molec. Ecol. 9, 959-969. 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New Phytol. 60, 235-281. Repeto, A. G., SIEGPRIED, W. R. & Crowe, A. A. (1984) Avian pollinators and the pollination syndromes of selected Mountain Fynbos plants. S. Afi: J. Bot. 3, 285-296. 4 & Oliver, E. G. H. (1985) Pollination syndromes of Erica species in the south- western Cape. S. Afr J. Bot. 51, 270-280. Ripa, M. (1998) Effects of intensity and frequency of crown damage on resprouting of Erica arborea L. (Ericaceae). Acta Oecol. 19, 9-16. RicHarps, A. J. (1986) “Plant Breeding Systems” (Allen and Unwin, London). Rozeé, F. (1993) Plant recolonisation after fire in Brittany littoral heathlands. Acta Oecol. 14, 529-538. SANTANDREU, M. & Lioret, F. (1999) Effect of flowering phenology and habitat on pollen limitation in Evica multiflora. Can. J. Bot. 77, 734-743. Scorl-ELLior, G. F. (1890) Ornithophilous flowers in South Africa. Ann. Bot. 4, 265-280. Specit, R. (1979) The sclerophyllous (heath) vegetation of Australia: the eastern and central states pp. 125-210 /n Specht, R. 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H., Burges, N. A., Moore, D. M., Valentine, D. H., Walter, S. M. & Webb, D. A. (Eds) “Flora Europaea, Vol. 3° (Cambridge University Press, Cambridge). Wess, N. R. (1986) “Heathlands” (Collins, London). THE MYRTLE SPRINGS METEORITE: A CHONDRITE (H4) FROM SOUTH AUSTRALIA’ By M. ZBIK* & A. PRING** Summary Zbik, M. & Pring, A. (2004) The Myrtle Springs meteorite: A (H4) chondrite from South Australia. Trans. R. Soc. S. Aust. 128(1), 33-36, 31 May, 2004. A single stone of 52.99 g was found 30 km west of the Old Myrtle Springs homestead on the Lake Torrens Plains, (30° 27° 13.8°° S, 137° 59’ 24.6” E). The chondrules in the meteorite are well defined and the dark matrix consists of fine olivine and pyroxene grains. Olivine (Fa,7, , 9), orthopyroxene (Fs,57. 97W0, 8,9), clinopyroxene (Wo,, . 29FS814 . 19), troilite and chromite are present. Radial pyroxene chondrules, porphyritic pyroxene chondrules, granular olivine-pyroxene porphyritic and olivine- pyroxene chondrules are abundant in the meteorite. Cryptocrystalline pyroxene chondrules and pyroxene-olivine chondrules are rare. Based on texture and mineral chemistry, the Myrtle Springs meteorite is classified as an H4 chondrite of shock stage S1-2, weathering grade W4. Key Words: Myrtle Springs, meteorite, chondrite. Transactions of the Royal Society of S. Aust. (2004), 128(1), 33-36 THE MYRTLE SPRINGS METEORITE: A CHONDRITE (H4) FROM SOUTH AUSTRALIA! by M. ZBik* & A. PRING™ Summary ZBIk, M. & PRING, A. (2004) The Myrtle Springs meteorite: A (H4) chondrite from South Australia. 7rans. K. Soc. S. Aust. 128(1), 33-36, 31 May, 2004. A single stone of 52.99 g was found 30 km west of the Old Myrtle Springs homestead on the Lake Torrens Plains, (0° 27' 13.8" S, 137° 59' 24.6" E). The chondrules in the meteorite are well defined and the dark matrix consists of fine olivine and pyroxene grains. Olivine (Fai72 : 06), orthopyroxene (Fsis7 + 07W01s:08), clinopyroxene (Wo i2.0Fsi4 +10), troilite and chromite are present. Radial pyroxene chondrules, porphyritic pyroxene chondrules, ‘granular olivine-pyroxene porphyritic and olivine-pyroxene chondrules are abundant in the meteorite. Cryptoerystalline pyroxene chondrules and pyroxenc-olivine chondrules are rare. Based on texture and mineral chemistry, the Myrtle Springs meteorite is classified as an H4 chondrite of shock stage S1-2, weathering grade W4. Key Worps: Myrtle Springs, meteorite, chondrite. Introduction The meteorite was found by Mr Don McColl of Glenside, South Australia on the 11th of July 2 while he was searching for tektites on Old Myrtle Springs homestead on the Lake Torrens Plain. The exact location as determined by GPS is 30° 27’ 13.8” S, 137° 59' 24.6” E. The site is some 21 km due west of the Old Myrtle Springs homestead and some 40 km WNW of Leigh Creek South township (Fig. 1). Mr McColl undertook a detailed search of the area for more pieces of the meteorite, but none was found. Given that there are few geographical place names in the area, we propose the name Myrtle Springs, the nearest geographical name. A report of the meteorite name and petrological details have been submitted to the Meteorite Nomenclature Committee and approval has been granted. Mr Don McColl surrendered the meteorite to the Museum in accordance with the South Australian Museum Act and was presented with a bronze medallion to commemorate the find. Macroscopic Description The meteorite is a 52.99 g single stone, shaped like a flattened rounded pebble about 4 cm in diameter and 2 cm thick and covered with fragments of a weathered brown crust (Fig. 2). The distinctive crust, 1 mm thick, is probably desert patina rather than a fusion crust. The silicate minerals throughout the interior of the stone are dark brown in colour and this ' This paper is dedicated to the late David Williams an intrepid tektite hunter who passed away 7th February 2003. “Tan Wark Research Institute, University of South Australia, Mawson Lake, Adelaide, South Australia 5095. “Department of Mineralogy, South Australian Museum, North Terrace, Adelaide, South Australia 5000. together with the desert patina, indicate that the meteorite has been exposed to the weather for a prolonged time. Metal is not present in the meteorite and it appears to have been oxidised to goethite and staining the matrix brown. A few unoxidized grains of troilite however, are still present, so the meteorite has not been completely oxidized. The extent of oxidation of the Myrtle Springs meteorite indicates some degree of antiquity for the fall, but it is rather difficult to judge the terrestrial age of the fall, possibly 20,000 to 35,000 years based on the weathering state (Wlotzka 1993). Jull e¢ al. (1993) found only a weak correlation between the degree of { coe er ees ae see L ot = aie | | Kittakittaooloo | Ke Lf aN) Witchelina $ coomna Bore ‘ Sone Springs sa | Leigh aes S | N 4 | 2 x Augusta | Fig. 1. Map showing the distribution of H4 chondrites in north eastern South Australia. 34 M. ZBIK & A. PRING Fig. 2. Upper and lower surfaces of the Myrtle Springs meteorite showing the shape of the stone and the weathered brown desert patina crust. The scale bar is marked in centimetres. Pig. 3. Different types of chondrules in the Myrtle Springs meteorite. (a) Barred olivine chondrule (BO), composed of the olivine bars and mesostasis in centre of micrograph and granular olivine pyroxene chondrule (GOP) to the left side (photo is 1.5 mm). (b) Elliptically shaped cryptocrystalline pyroxene olivine chondrules (C) display unusual wavy extinction probably due to shock metamorphism (photo is 1.5 mm). (c) Porphyritic pyroxene olivine (POP) chondrule with poikilitic structure (photo is 1.5 mm). weathering and the terrestrial age of meteorites in a detailed study of a group of meteorites from north western Texas, U.S.A. Sample and Analytical Procedures The polished thin section of the meteorite was used for both the petrographic examination and chemical analyses. Composition of the silicate minerals was determined using a CAMECA SX51_ electron microprobe, with a Moran analysis package, at Adelaide Microscopy, The University of Adelaide. Analyses were made using an accelerating voltage of 15 keV, a sample current of 20 nA, and beam width of 0.1 um. Mineralogy In thin section the meteorite is generally brown due to the goethite staining which permeates all of the section. The chondrules are clearly visible but they are extensively fractured. They are typically between 0.5 and | mm in diameter, but a few measuring more than 2 mm in diameter were noted. THE MYRTLE SPRINGS METEORITE: A CHONDRITE (H4) FROM SOUTH AUSTRALIA 35 The chondrules observed in the thin section are clearly defined which is typical in low petrologic type, but may have been additionally enhanced by the preferential weathering and iron oxide staining of the matrix. The chondrules and chondrule fragments are composed predominantly of olivine and pyroxene. Some chondrules display cryptocrystalline structure, Using the classification system of Wasson (1993), a number of distinct chondrule types can be identified in the meteorite. Barred olivine chondrules (BO). with pyroxene-plagioclase — mesostasis between the olivine bars, are common (Fig. 3a). Cryptocrystalline pyroxene and pyroxene-olivine chondrules (C) are rare; those few observed vary in size and shape and display strong wavy extinction. These C-type chondrules are mineralogically heterogeneous, with in some areas pyroxene and others olivine (Fig. 3b). Radial pyroxene chondrules (RP) are common, and display a range of crystal sizes but nucleation appears to have occurred at a single point on the rim of the chondrules. Abundant are porphyritic pyroxene (PP) chondrules, granular olivine-pyroxene chondrules (GOP) (Fig. 3a, on the lefi-hand side), and porphyritic olivine-pyroxene chondrules (POP). Porphyritic chondrules contain fine and coarse grained, euhedral olivine crystals and display a poikilitic texture (Fig. 3c). Chondrules vary in shape, whereas the small chondrules, those below 1 mm, are generally round, the larger are often more oval and less regular in outline. Turbid cryptocrystalline mesostasis between olivine and pyroxene crystals is present and has a plagioclase\pyroxene-like composition but the analyses were non-stoichiometric. — Distinct plagioclase grains were not found during petro- logical examination or microprobe analysis. The matrix of the meteorite does not appear to have been recrystallised and consists of fine-grained olivine and pyroxene fragments but it is greatly obscured by the goethite staining making tt difficult to study. Accessory chromite and troilite grains were detected in microprobe investigation. A number of quartz grains, aeolian in origin, were cemented into the secondary limonitic desert patina making crust around meteorite. Selected electron microprobe analyses for the Myrtle Springs meteorite are given in Table |. The olivine in the Myrtle Springs meteorite ts equilibrated, with a mean fayalite content of Fajz 9.0.6. (mean of 18 analyses). The Ca-poor pyroxene is uniform and shows only a narrow variation in composition with mean ferrosilite content of Fs)5 7.9.7, and wollastonite content of Wo; s,s (mean of 20 analyses). The Ca-rich pyroxene is somewhat variable in composition with mean ferrosilite content of Fsjyo-;o, and wollastonite content of Wor «i209 (mean of 15 analyses). Classification The Myrtle Springs meteorite has been classified as an H4 chondrite. The olivine (Faj75.9.) and Ca- poor pyroxene (Fsj5.7,9.7) compositions are within the range of the H chondrites (Keil & Fredriksson 1964). The equilibrated mineral compositions of the olivine and Ca-poor pyroxene, the presence of about 20% of the polysynthetically twinned Ca-rich pyroxenes, the lack of recrystallisation of the matrix, the well- defined chondrule boundaries and the glassy mesostasies all indicated that the Myrtle Springs meteorite belongs to petrological type 4 classification of Van Schmus & Wood (1967). The wollastonite content in the low-calcium pyroxene is within the range found in H4 chondrites (Scott e7 a/., 1986). Van Schmus and Wood acknowledge that in their classification scheme that petrological type 4 is defined by relatively ambiguous characteristics as it represents a transition between extremely variable olivine and pyroxene compositions (type 3) and the uniform compositions (type 5). We choose type 4 rather than type 5 because of the abundance of clinopyroxene; type 5 chondrites have only very minor clinopyroxene. Most olivine and pyroxene grains display sharp extinction and irregular fracture crystals in the Myrtle Springs chondrite. Some pyroxene display weak undulatory extinction. Both are indicative of the meteorite having been unshocked or only very weakly shocked after metamorphism. According to the classification scheme of Stoffler ef a/. (1991), the shock facies is estimated to be SI-2 (or i.e.) unshocked to very weakly shocked. The oxidation of approximately 95% of the original metal and troilite in the meteorite indicates weathering state W4 on the classification scheme of Wlotzka (1993). Other H4 chondrites from North Eastern South Australia. There are a number of other H4 group meteorites known from the north eastern region of South Australia (Fig. 1). The Kittakittaooloo meteorite was found in 1970 about I km NW _ of Lake Kittakittaooloo (28° 2’ S, 138° 8' E) and has an olivine composition of Fa,,, (Graham er al. 1985). The Coonana meteorite was found in 1962. near Coonana Bore (29° 51’ S, 140° 42’ E) and has an Fa,y olivine composition (Graham e/ al. 1985). Both of these meteorite localities are rather remote from Myrtle Springs being approximately 270 km north and 250 km ENE, respectively, of the site. The Witchelina meteorite was found in 1920 in the area of Witchelina homestead (30° S, 138° E), only 40 km north of the Myrtle Springs homestead, and it is possible that the two meteorites might be part of the 36 M. ZBIK & A. PRING TABLE |. Representative electron microprobe analyses (wt. %) of selected minerals in Myrtle Springs meteorite. OXIDE Olivine Ca-poor Ca-rich (No. Analyses) — (18) pyroxene pyroxene (20) (15) SiO, 39.26 56.42 49.60 TIO, 0.05 0.10 0.24 ALO, 0.04 0.41 3.41 FeO 16.13 10.38 8.83 MnO 0.48 0.52 0.47 MgO 43.41 30.58 22.86 CaO 0.04 0.94 10.77 Cr,O, 0.40 0.30 0.91 Total 99.76 99.05 97.09 same fall. Unfortunately the locality details for the original site for the Witchelina find are sketchy and only approximate. The Witchelina meteorite was a 3.6 kg single stone and is coated with a distinctive fusion crust. Analysis gave an olivine composition of Fa,,, (Graham ef al. 1985), slightly more iron-rich than Myrtle Springs. Witchelina, however is a much fresher stone. The metal has not been oxidized and metal and troilite are estimated to constitute around 3% of the meteorite. This indicates a weathering state of W1 or W2 and a terrestrial age of only a few thousand years. The ground mass colour has a distinctly greyish brown tone rather than reddish brown. While it might be expected, given the difference in size between Myrtle Springs and Witchelina, that the former would show the effects of weathering to a somewhat greater degree than the larger mass, but probably not to this extent. The difference in olivine composition, the presence of the fusion crust, and fresh metal in Witchelina all indicate that the two meteorites are not part of the same fall. Acknowledgments The authors wish to thank Mr Angus Netting and Mr John Terlet, of Adelaide Microscopy, The University of Adelaide, for assistance with the electron microprobe analyses. We wish to thank the editor and the two referees for their detailed and helpful comments on the manuscript. References GRAHAM, A. L. BEVAN, A. W. R. & HUTCHINSON, R. (1985) Catalogue of meteorites. 4th edition. British Museum (Natural History) London. 460pp. JuLL, A. J., DONAHUE, D. J., CIELASZYK, E. & WLOTZKA, F. (1993) Carbon-14 terrestrial ages and weathering of 27 meteorites from the southern high plains and adjacent area (USA) Meteoritics 28, 188-195. Keil, K. & FREDRIKSSON, K. (1964) The iron, magnesium and calcium distribution in coexisting olivines and rhombic pyroxenes of chondrites. J. Geophys. Res. 69, 3487-3515. Scott, E. R. D., TAYLor, G. J. & KEIL, K. (1986) Aceretion, metamorphism, and brecciation of ordinary chondrites: Evidence from petrologic studies of meteorites from Roosevelt County, New Mexico. Proc Lunar Planet. Sci. Conf. 17th, ELI5-E123. STOFFLER, D., KrIL, K. & Scort, E. R. D. (1991) Shock metamorphism of ordinary chondrites. Geochim. Cosmochim. Acta. 55, 3845-3867. VAN ScuMus, W. R. & Woon, J. A. (1967) A chemical- petrologic classification for the chondritic meteorites. Geochim. Cosmochim. Acta 31: 747-765. Wasson, J. T. (1993) Constraints on chondrule origins. Meteoritics 28, 14-28. WLOTZKA, F. (1993) A weathering scale for the ordinary chondrites. Mefeoritics 28, 460. EXPECTED LIFETIME IN SOUTH AUSTRALIA 1841-1996 By P. I. LEPPARD*, G. M. TALLIS & C. E. M. PEARCE Summary Leppard, P. I., Tallis, G. M. & Pearce, C. E. M. Expected lifetime in South Australia 1841 — 1996. Trans. R. Soc. S. Aust. 128(1), 37-42, 31 May, 2004. For each sex, population life tables have been calculated from mortality data associated with the age-specific population counts produced by each of the twenty- five population censuses that have been conducted in South Australia from 1841 until 1996. Estimates of expected lifetime have been derived separately for males and females for each census year. The computationally intensive statistical method of the bootstrap has been used to calculate a statistical sampling error for each estimate of expected lifetime. The results show generally increasing trends: from approximately 44 years and 48 years in 1841, to 75 years and 81 years in 1996, for males and females respectively. Key Words: Expected lifetime, life expectancy, bootstrap standard error, South Australia. Transactions of the Roval Society of S. Aust. (2004), 128(1), 37-42. EXPECTED LIFETIME IN SOUTH AUSTRALIA 1841 — 1996. by P. I. Lepparp’, G. M. TALLIS & C. E. M. PEARCI Summary Lepparb, P.1., TALLIS, G. M. & PEARCE, C.E.M. Expected lifetime in South Australia 1841 - 1996. Trans. R. Soc. S. Aust. 128(1), 37-42, 31 May, 2004. For each sex, population life tables have been calculated from mortality data associated with the age-specific population counts produced by each of the twenty-five population censuses that have been conducted in South Australia from 1841 until 1996, Estimates of expected lifetime have been derived separately for males and females for each census year. The computationally intensive statistical method of the bootstrap has been used to calculate a statistical sampling error for each estimate of expected lifetime. The results show generally increasing trends; from approximately 44 years and 48 years in 1841, to 75 years and 81 years in 1996, for males and females respectively. Kry Worps: Expected lifetime; life expectancy: bootstrap standard error; South Australia. Introduction Expected lifetime or life expectancy, meaning the average length of life from birth until death, is often used as a composite indicator of the health and social conditions of a human population. Expected lifetime is a single value that summarises the prevailing mortality experience of a defined population of individuals, and consequently provides a means by which the mortality patterns of different populations can be conveniently compared. The progress in the development of the general well-being of a particular population can also be assessed by a time sequence of relevant expected lifetime values. Alternatively, expected lifetime can be cautiously interpreted as a prediction of future average lifetime. Expected lifetime is conventionally calculated separately for males and females, and is one of the results of a (mortality or actuarial or population) life table analysis. This type of analysis uses age-specilic death rates calculated from age-specific population sizes and numbers of deaths. Hence life tables are usually calculated for census years only since accurate estimates of population sizes are required. Mortality data pertinent to the three consecutive years centred on and encompassing the census year are averaged in an attempt to smooth out random fluctuations in the observed numbers of deaths. The study of population mortality patterns is generally acknowledged as beginning with Graunt (1662), and subsequently the methodology of the life table has evolved to become a basic tool of demographers (eg Newell 1988) and actuaries (eg Benjamin & Haycocks 1970). There are many standard statistical texts covering the methodology of life table analysis (eg Elandt-Johnson & Johnson 1980). School of Mathematical Sciences, University of Adelaide, Adelaide, South Australia, S005, Australia, Email: pleppard@maths.adelaide.edu.au. Very tew official life tables and hence estimates of expected lifetime have been produced for South Australia. Invariably 19th century references to Australian mortality are based on New South Wales or Victorian data (Pell 1867, Burridge 1884). Some official South Australian life tables have been calculated for the period 1881-1910, but these tables were calculated from mortality data of ten-year periods. This methodology was discarded when the Commonwealth Statistician (Wilson 1936) decided to cease publication of separate life tables for each Australian State on the basis of the “.. time (thus) saved..”. In the 1970’s the Australian Bureau of Statistics recommenced producing life tables specifically for each State. Thus the complete mortality experience of South Australia, as measured by male and female expected lifetimes, has not been consistently or comprehensively investigated. Materials and Methods The results presented in this paper have been extracted from a thesis presented for an MSc degree at The University of Adelaide, Leppard (2003). The thesis can be accessed electronically at http://thesis.library.adelaide.edu.au/public/adt- SUA20030422.122816/. A very brief overview of the material covered in the thesis is given here. The essential data requirements to calculate a life table for a given census year are, first, the population size at each single year of age; and second, the number of deaths at each single year of age for that census year, and for the calendar years immediately preceding and following it. Population data were obtained from official sources for each of the 25 censuses carried out in South Australia since 1841: 12. colonial censuses and 13 post-Federation censuses. Sources for these data include the South tad Australian Parliamentary papers, the $A Government Gazette, the Statistical Register of South Australia (1850-1975), and various publications of the Commonwealth Bureau of Census and Statistics and its successor organisation, the Australian Bureau of Statistics. Age-specific numbers of deaths by sex are also available from these same sources, although the published data for the years 1841 to 1860 are in a statistically limiting form principally because of various types of grouping by age of the data. This deficiency of the official records has been overcome by tabulating the primary data contained in death certificates lodged with the Registry of Births, Deaths and Marriages, which are held in the archives of State Records of South Australia, Permission was obtained from the Registrar for access to the Registry’s records, and approximately 18,000 death certificates for relevant years of the above period were examined and the deaths they reported were classified according to year, sex and age at death. Indexes to the burial records of Holy Trinity church and the West Terrace cemetery were used to obtain data from which the age-specific number of deaths was estimated for each sex for 1840, It is impossible to provide within the space limitations of this paper a comprehensive description of the characteristics of all data sets that have been used to calculate expected lifetime. The reader is referred to Leppard /oc cit for a detailed description for each census year of the process of data collection, a discussion of the limitations of the data, estimates of expected lifetime calculated by the life table method, and estimates of sampling error calculated by using — the computationally intensive statistical bootstrap procedure (Efron & Tibshirani 1993). For each census year a robustness analysis 1s also presented, where an examination is made of the effect on expected lifetime of under-reporting in the age- specific population sizes and number of deaths. All data files and computer programs can be publicly accessed from the web site previously indicated. Expected lifetime is calculated here in the following way, conventionally for males and females separately. Let ,p, be the number of individuals aged between 0 and | years at a census. Let jd, be the average number of individuals who died between the ages of 0 and | years, determined from the number of deaths registered in the census year and the years immediately preceding and following the census year. Then the quantity ,q, is calculated from ,p, and iy as d, m0 170 ay ot te Py" ay It is sometimes recommended that, in circumstances where extremely high infant mortality prevails, a factor smaller than the factor 8 P. 1. LEPPARD, G. M. TALLIS & C. E. M. PEARCE of '/2 should be used in the calculation of ,g,. This has not been done in the calculation of expected lifetimes shown in Table 1, and consequently some of the values given there for the 19th century may be slightly larger than perhaps they otherwise would be. Newell /oc cif suggests that a factor of .3 be used instead of '/2 for “developing countries”. Recalculation of expected lifetime for selected census years for South Australia using this value for the factor has produced small differences that are less than the standard error of the expected lifetime. In an analogous manner, ,q, is calculated for age group |-2 years, ,q, for age group 2-3 years, and so on, for each yearly age group. Thus using standard demographic notation, ,g, is a mortality rate specific to the group aged between j years and (j+1) years. A distribution function for lifetime values, Fy is defined with this set of q-values as FOX) 1-1 a )- g )O> 9a) pg) for x = 1,2.3..., where F(0O) = 0 and here F(105) = | by definition. Expected lifetime is calculated from the tabular values of F’ by the trapezoidal method of numerical integration. It should be noted that neither / nor expected lifetime is directly applicable to the population of individuals in the census year from which the g- values are determined. They are concepts that prospectively apply to a hypothetical population of infants born in the census year, if it is assumed that the currently prevailing mortality rates, as measured by the g-values, eventuate for these infants throughout their lifetimes. There are mathematical arguments that suggest that expected lifetime is an underestimate of the unrealised average lifetime of this hypothesised population of infants. The potential effect of under-counting is examined in the following way. Population sizes are obtained following a census, and the numbers of deaths are compiled from the death certificates required by law. Both of these processes are not necessarily exact and some individuals could be excluded. Consider jp and jd) which are used to calculate jg», and suppose the “correct” values of these two quantities are larger by factors of O; and O02 respectively. Then the “correct” value of ;qy would be Ox ido = ido Or ipo + $O2 ido Ov ipo+ hido 3 OD. 3 iqy( “correct”’) = which is close to \qg) calculated by using the supposedly under-counted values jp) and jd) if ©. ~ |; that is, jgp is an appropriate substitute for 140 (“correct”) if the extent of under-counting is EXPECTED LIFETIME IN SOUTH AUSTRALIA 1841 - 1996 approximately the same for both the population size and the number of deaths. For each census year and sex, expected lifetimes are given in Leppard /oc cif for combinations of assumed levels of under- counting in both population size and numbers of deaths. Results and Discussion Estimates of expected lifetime (EL) for South Australian males and females for 1841-1996 are shown in Table |. A standard error, SE (EL), is also given for each estimate. Table 1 also shows the total population size and the average total number of deaths on which each estimate is based. The estimates of expected lifetime are also displayed graphically in Figure |, which is annotated to indicate 1. The first census at which the estimate of expected lifetime for females can be statistical ly claimed to be significantly greater than the estimate for males. 39 over this period in the generally increasing sequence of expected lifetime values. . Larger time intervals between successive estimates of expected lifetime as a consequence of the postponement of censuses during the Great Depression and World War II. The estimates of expected lifetime presented in Table | and Figure | have been derived from data from populations that have evolved from the initial British settlement of South Australia with subsequent migration from Western and Southern Europe. These estimates have been derived primarily from routinely recorded official population and mortality data from which Aborigines were excluded because of a government policy that remained in force until the second half of the 20th century. The two series of expected lifetimes that are presented here are the most extensive and comprehensive available for any Australian state or for the Commonwealth, with all estimates consistently calculated using a currently accepted standard methodology. The results delineate for the 2. The passing of the first South Australian Public first time the changes in mortality that have occurred Health Act, as a response to the deterioration of — in South Australia over 150 years of European social and public health conditions in the 1860’s settlement. The electronic provision of the basic data and 1870’s. The effect of this deterioration on files through the previously stated web address has population mortality is shown by the disruption — been made to assist demographic researchers. TABLE |. Expected lifetime with standard error for South Australia 1841-1996. Male Female Census Population Deaths EL SE(EL) Population Deaths EL SE(EL) 1841 8195 140 44.27 2.59 6345 102 48.17 1.88 1844 9525 172 41.95 1.56 7608 128 43.49 1.60 1846 12670 236 41.57 1.55 9650 163 46.67 2.08 1851 35309 544 45.73 1.24 27730 439 47.62 1.31 1855 43716 713 47.68 1.27 41470 620 50.00 1.27 1861 64640 111 48.55 1.06 61678 951 51.88 1.20 1866 85625 1434 47.39 0.86 77975 1178 50.34 0.88 187] 95288 1438 48.69 0.67 90164 1165 52.19 0.77 1876 110410 1972 43.57 0.58 102734 1659 46.09 0.64 1881] 149530 2283 47.62 0.50 130335 1819 50.76 0.62 189] 166801 2156 52.52 0.47 153630 1787 54.98 0.52 1901 184424 2169 54.56 0.47 178182 1821 57.72 0.50 1911 207358 2269 58.81 0.41 201200 1880 62.41 0.42 192] 248267 2663 60.01 0.38 246893 2221 63.53 0.39 1933 290429 2701 65.34 0.34 289546 2386 68.04 0.35 1947 320031 3478 67.17 0.27 326042 2995 71.28 0.26 1954 403903 3958 67.72 0.25 393191 3267 73.07 0.24 1961] 490225 4412 68.62 0.22 479115 3537 74.91 0.21 1966 548530 5030 68.57 0.21 543344 4034 13,22 0.20 1971 586451 5454 68.67 0.20 587656 4408 75.58 0.20 1976 619759 5402 70.06 0.19 624595 4488 76.93 0.18 1981 635695 5473 71.49 0.19 649467 4438 78.92 0.18 1986 666159 5626 72.98 O.18 679985 4824 79.48 0.18 199] 690805 5856 74.08 0.17 709802 5156 80.30 0.17 1996 698799 5989 75.33 0.17 722673 5504 81.34 0.16 40 P. I. LEPPARD, G. M. TALLIS & C. E. M. PEARCE pee _ _— es Expected lifetime in South Australia 1841-1996 | i oe | Great Depression | ; | bo | A \ 1 4 | 1) | 1 a ! | 7 j ae Stattically detectable gender diferences | ' | ' vferde 60 ‘ A } i | Vir ar | s ! | ne | 1 A 4 ] a t A / ot 1 A t i \ | \ 4 Public Health Act 1873 4 rs T ~~ T TO ee To {640 {860 {880 1900 12) 10 1860 1980 2000 Year Fig. 1. Expected lifetime in South Australia 1841-1996. EXPECTED LIFETIME IN SOUTH AUSTRALIA 1841 - 1996 TABLE 2. Comparison of expected lifetime for males. 4| SA Other Australia Britain US Sources (6) (7) (8) 1841 44.27 40.2 1855 47.68 1838-1854 39.9 1856-1861 45.58 (1) 1861 48.55 1871 48.69 1870-1881 46.47 (2) 1881 47.62 I881-1890 50.61 (3) 47.20 43.7 1891 2 ee 1891-1900 53.02 (3) 51.08 44.1 1901 54.56 48.0 1901-1910 56.76 (3) 55.20 48.5 1911 58.81 £033 65.34 63.48 58.7 62.7 1954 67.72 67.82 (4) 67.14 67.5 1976 70.06 70.27 (5) 69.56 69.9 69.9 1996 75.33 75.30 (5) 75.70 74.5 1S; (1) Pell loc cit (NSW). (2) Burridge loc cit (composite states, excludes SA). (3) Commonwealth Census 1911, Vol 3. (SA). (4) Wickens, Australian Journal of Statistics, 2, 1960. (SA). (5) Deaths SA, Australian Bureau of Statistics. (6) Australian Life Tables, Australian Government Actuary. (7) English Life Tables, UK Government Actuary’s Office. (England & Wales). (8) US National Vital Statistics Reports, Vol 51, 3. 2002. TABLE 3. Comparison of expected lifetime for females. SA Other Australia Britain US Sources 1841 48.17 42.2 1855 50.00 1838-1854 41.9 1856-1861 45.58 1861 51.88 1871 52.19 1870-1881 49.64 1881 50.76 1881-1890 53.8] 50.84 47.2 189] 54.98 1891-1900 56.10 54.76 47.8 1901 $7.72 51.0 1901-1910 60.39 58.84 52.4 191] 62.41 1933. 68.04 67.14 62.9 66.3 1954 73.07 73.09 72.75 cia: 1976 76.93 77.24 76.56 76.0 TFS 79.6 79.7 1996 81.34 81.34 81.40 42 P. 1. LEPPARD, G. M. TALLIS & C. E. M. PEARCE A selection of expected lifetimes presented here for South Australia in Table 1 are shown in Table 2 and Table 3 (column SA) for comparison with other estimates and other similar populations; with estimates for other colonies and for South Australia, by other authors (Other sources); with Australia, by the Australian Bureau of Statistics and by the Australian Actuary; with Britain, by the Government Actuary’s Office; and with the United States, by the National Vital Statistics Office. Generally, expected lifetimes for South Australia have compared more than favourably with the other estimates until the first third of the 20th century, after which time there has been a gradual lessening of the differences between expected lifetimes. This most probably reflects the effect of globalisation and standardisation in conditions of Western industrial societies. References BENJAMIN, B. & H. W. HAycocks (1970). The Analysis of Mortality and other Actuarial Statistics. Cambridge, Cambridge University Press. BurribGe, A. F. (1884). “On the rates of mortality in Australia.” Journal of the Institute of Actuaries 24: 333- 358. EFRON, B. & R. J. TIBSHIRANI (1993). An Introduction to the Bootstrap. New York, Chapman & Hall. ELANDT-JOHNSON, R. C. & N. L. JOHNSON (1980). Survival Models and Data Analysis. New York, John Wiley & Sons. GRAUNT, J. (1662). Natural and political observations made upon the bills of mortality (reprinted 1939). Baltimore, Johns Hopkins Press. Lepparn, P. 1. (2003). An analysis of population lifetime data for South Australia. 1841- 1996. MSc (Statistics) thesis, School of Applied Mathematics, The University of Adelaide http://thesis.library.adelaide.edu.au/public/adt- SUA20030422.122816/. NEWELL, C. (1988). Methods and Models in Demography. London, Belhaven Press. Pett, M. B. (1867). “On the rates of mortality and expectation of life in New South Wales as compared with England and other countries.” Transactions of the Royal Societv of New South Wales 1: 66-76. WILSON, R. (1936). Official year book of the Commonwealth of Australia, Commonwealth Bureau of Census and Statistics, Canberra. VARIATION IN SEX RATIOS IN FOUR ANGUINA (NEMATODA: ANGUINIDAE) SPECIES By I. T. RiLEy* & T. BERTOZZIf Summary Riley, I. T. & Bertozzi T. (200). Variation in sex ratios in four Anguina (Nematoda: Anguinidae) species. Trans. R. Soc. S. Aust. 128(1), 43-46, 31 May, 2004. Number and ratios of adult male and females nematodes in four Anguina species. A. australis from Ehrharta longiflora, A. funesta from Lolium rigidum, A. microlaenae from Microlaena_ stipoides and an _ undescribed species from Polypogon monspeliensis, were determined and analysed. The sex ratio in A. australis and A. funesta differed statistically from the expected 1:1, with 57% and 56% females, respectively. The ratios for A. microlaenae and Angina sp. ex Polypogon conformed to the expected 1:1. Galls initiated by single sex nematodes, both male and female, were found in the four species, except that no galls containing only males were found in A. australis. Reproduction in this group appears to be amphimitic, as single sex galls contained no progeny and the data point to the possiblity of a mechanism that favours female occupancy of galls. Key Words: Anguina australis, Anguina funesta, Anguina microlaenae, seed gall nematode, leaf gall nematode, gall initiation, sex ratios, reproduction, amphimixis. Transactions of the Roval Society of S. Aust. (2004), 128(1), 43-46. VARIATION IN SEX RATIOS IN FOUR ANGUINA (NEMATODA: ANGUINIDAE) SPECIES. by I. T. Ritey’ & T. Bertozzi' Summary Ritey LT. & BerrozziT. (200). Variation in sex ratios in four Anguina (Nematoda: Anguinidae) species. Trans. R. Soc. S. Aust. 128(1), 43-46, 31 May, 2004. Number and ratios of adult male and females nematodes in four Anguina species, A. australis trom Ehrharta longiflora, A. funesta trom Lolium rigidum, A. microlaenae trom Microlaena stipoides and an undescribed species from Polypogon monspeliensis, were determined and analysed. The sex ratio in A. australis and A. yA funesta differed statistically from the expected 1:1, with 57% and 56% females, respectively. The ratios for A, microlaenae and Angina sp. ex Polvpogon conformed to the expected 1:1. Galls initiated by single sex nematodes, both male and female, were found in the four species, except that no galls containing only males were found in 4. australis, Reproduction in this group appears to be amphimitic, as single sex galls contained no progeny and the data point to the possibility of a mechanism that favours female occupancy of galls. Key Worps: Anguina australis, Anguina funesta, Anguina microlaenae, seed gall nematode, leat gall nematode, gall initiation, sex ratios, reproduction, amphimixis. Introduction Species of the nematode genus Anguina form galls in plants, mostly in poaceous hosts. The type species, Anguina tritici, is considered to reproduce by amphimixis (Triantaphyllou and Hirschmann 1966). Galls are initiated by invasive stage juveniles (usually J2s), which, confined to the gall, develop to adults. So, mates can only be found amongst the relatively small number of individuals present in the gall. Therefore, a mechanism to increase the likelihood of both females and males occurring in individual galls would be advantageous. The mechanism would need to operate before or during gall initiation, unless sex in Anguina spp. can be environmentally determined. In two species, Anguina australis and Anguina microlaenae, most galls contain only small numbers of adults, with some galls containing only females (Riley ef al, 2001; de Silva and Riley 2002). It was suggested that this may indicate a mechanism whereby only genetically female juveniles could initiate galls or that sex was environmentally determined (de Silva and Riley 2002). In both cases, no progeny were found in the single sex galls, so there was no evidence of parthenogenetic reproduction. As sex ratios and adult populations in galls for only two leaf-gall forming species of the genus have been examined in detail, similar data were collected for two seed-gall forming species, Anguina funesta and “Plant and Pest Science, School of Agriculture and Wine, The University of Adelaide, PMB 1, Glen Osmond, South Australia 5064, Australia, Email: ian.riley(@adelaide.edu.au, ‘Evolutionary Biology Unit, South Australian Museum, North Terrace, Adelaide South Australia 5000, Australia. Anguina sp. ex Polypogon monspeliensis and further data for A. microlaenae. These data were examined for further evidence of mechanisms that determine sex ratios and first generation populations in Anguina galls. Materials and Methods Anguina australis. Data on sex of A. australis adults in galls in Ehrharta longiflora came from a published study (Riley ef a/. 2001) Anguina funesta. Lolium rigidum plants were grown (about 100 plants per pot) in a peat/sand mix (UC mix; Baker 1957) in 280 mm pots outdoors at Urrbrae, South Australia. Seed and nematode galls (about 100 per pot) were applied early in July 2001 and inflorescences harvested for collection of galls in late October 2001. Ten inflorescences were dissected and 203 galls collected. Gall positions (spikelet and floret) were recorded. Each gall was opened in water under a dissecting microscope to release and count adult female and male nematodes. The presence of eggs and juveniles was noted. Anguina microlaenae. A further eighty galls from each of two populations of Microlaena_ stipoides from Toowoomba, Queensland were assessed as for the A. fiesta galls from L. rigidiwm and combined with data previously obtained from these populations (de Silva and Riley 2002). Anguina sp. ex Polypogon. Anguina sp. galls from Polypogon monspeliensis were obtained from plants collected in 1992 and 1993 from two sites in south- eastern South Australia. Galls in these plants had formed in primary shoot meristems, pedicel meristems and ovaries and in this report are termed apical, pedicel and seed galls respectively. A total of 44 I. T. RILEY & T. BERTOZZI TABLE |. Gall types, sex and ratio of adults for populations of four Anguina species. A. australis* A, funesta A. microlaenae** Anguina sp. ex Polypogon Gall type Leaf Seed Leaf Apical Seed Pedicel Total Number examined 194 203 199 95 122 14 232 Females Range 1-5 0-13 0-6 0-7 0-8 1-3 0-8 eantSE 1.68+0.07 2.66 + 0.14 1.65 + 0.08 1.744014 1.3940.08 1.64+£0.23 1.55 +0.07 Median | 2 | I | | Males Range 0-3 0-9 0-8 0-8 0-11 0-3 0-11 ean+SE 1.96+0.04 = 1.98 +£0.10 1.75 + 0.10 1.904018 1.374010 1.214019 1.58 + 0.09 Median | 2 | | | | ] Adults Range 1-7 1-19 0-12 1-15 1-19 2-5 1-19 eantSE 2.88+0.08 4.6440.22 3.40+0.15 3.63 £0.29 2.754017 2.864033 3.12+0.15 Median 3 4 3 2 2 2 2 Proportion female Range 0.33 -1 0-1 0-1 0-1 0-1 0.4—1 0-1 ean+ SE 0.573 + 0.009 0.556 + 0.011 0.504 40.014 0.497+ 0.016 0.504 + 0.011 0.567 + 0.039 0.505+ 0.009 Median 0.50 0.57 0.50 0.50 0.50 0.50 0.5 *Data from Riley ef al. (2001) **Data from de Silva and Riley (2002) combined with further data collected on the same population 12] A. australis A. funesta °0C) 6 1% 5% 25% 50% 3 oe Gon | 42| A. microlaenae Males per gall Females per gall Fig. |. Proportion of galls formed by four Anguina spp. with various combinations adult nematodes (dotted line represents 1:1, area of the circle represents the proportion of the population at each combination and scale indicated by circles labelled 1, 5, 25 and 50%.) 232 galls were assessed consisting of 95 apical, 122 seed and 14 pedicel galls. Galls were dissected and adults counted, as above. GENSTAT 5. (Lawes Agricultural — Trust, Rothamsted Experimental Station) was used to run statistical analyses. Results Table | gives details of the adults in the galls of each of the four nematode species. Infestation of the Lolium rigidum was heavy, nearly all florets in the spikelets examined containing galls. The contents of the galls from the two populations of A. microlaenae did not differ statistically and were combined. Differences between the three classes of gall produced by Anguina sp. in P. monspeliensis could not be reliably tested because of inequality of variance and no variance-stabilizing transformation was effective, so they are presented individually and as a total. Two galls with exceptionally large numbers of adults, one gall formed by 4. microlaenae with | female and 38 males and one apical gall formed by Anguina sp. in P. monspeliensis with 24 females and 31 males, were excluded from analysis as aberrant. The sex ratio for each species was tested for the whole sample by chi-square test (Table 2). The numbers of females per male in galls formed by both A. australis and A. funesta were greater than the expected ratio of 1:1. This was consistent with the mean proportion of females found in the galls of both species (Table 1) being more than half. The ratio for A. microlaenae and Anguina sp. ex Polypogon did not differ statistically from the expected 1:1. Despite the divergence of the sex ratio from 1:1 in two species, the only species with a median proportion of females that differed from this ratio was A. funesta with a median of 0.57 (i.e. 1.33:1), The proportion of galls with each combination of VARIATION IN SEX RATIOS IN FOUR ANGUINA (NEMATODA: ANGUINIDAE) SPECIES 45 TABLE 2. Chi-squared test of conformity of Anguina sex ratios to an expected 1:1. Anguina sp. Observed ratio Chi-squared Probability (female/male) (df=1) A. australis 1.41 15.91 < 0.001 A. funesta 1.36 22.01 <0.001 A, microlaenae 6.95 0.53 0.47 A. sp. ex Polypogon 0.98 0.07 0.79 adults for the four species is shown in Figure 1. The bias in female occupancy of galls can be seen for A. australis and A. funesta with 40% and 51% of galls having more females than males, respectively. By comparison, only 25% and 19% of A. microlaenae and Anguina sp. ex Polypogon galls had more females than males. Galls occupied by single adult males and females were found in the four nematode species except that no galls containing only males were found for A. australis. The greatest proportion of single sex galls was found for A. microlaenae (10%). No eggs or juveniles were found in any single sex gall. Analysis of gall position (spikelet and floret) for A. finesta in relation to numbers of adults and progeny and sex ratio revealed only one significant relationship (data not shown). A_ significant regression (P<0.001) was found between sex ratio and spikelet (the latter being the independent variable). The lower the position of the spikelet in the inflorescence the greater proportion of females in the galls. However, only about 6% of the variation in sex ratio was attributable to spikelet position. No effect of floret position within the spikelet was found. Discussion Reproduction in the species examined appears to be exclusively amphimitic, as no progeny were found in galls containing single adults. The sex ratio of 1:1 found in 4. microlaenae and Anguina sp. ex Polypogon is consistent with amphimitic reproduction. However, the overall sex ratio of about 1.4 females/male for A. australis and A. funesta points to the existence of mechanism that controls the sex of adults in galls. The data arising from the earlier, more limited, examination of A. microlaenae galls and the 4. australis galls led to postulation (de Silva and Riley 2002), based on single adults in galls being only female, that galls might only be initiated by genetically female juveniles or that environmental sex determination was involved. The additional "Chit, W. (1971) “Biology of Anguina mobilis n. sp. on capeweed (Cryptostemma calendulaceum)” PhD thesis, The University of Adelaide (Unpubl.). data presented here revealed that galls formed by single males occur in three of the species, including A. microlaenae and provide no support for either proposition. Therefore, the significantly higher number of females in A. australis and A. funesta points to a possible mechanism involving relative rather than absolute differences in behaviour between the sexes. Although both sexes can initiate galls, female occupancy is quantitatively more likely. Given that host invasion and gall initiation rates were high in the L. rigidum examined (mean of 4.6 adults per gall compared to 2.2 to 2.6 in field populations, McKay er al. 1981), it is possible that such a mechanism operates to maximise potential reproduction of the species when there is more competition for gall sites. This cannot be tested from the data obtained, as they were not derived from a controlled experiment. However, the tendency for more females in the lower galls in L. rigidum might be consistent with this proposal, if there was more competition for sites at the time these galls were formed. Chit! also found that galls formed by Mesoanguina mobilis in Arctotheca calendula had 2 to 7 first generation adults, most commonly with 3 females and 2 males, thus supporting a mechanism to favour females occurring more broadly in anguinid genera. A consistent pattern in the data is the low frequency of galls containing only single adults for all species and a strong bias towards galls containing two adults (one female and one male), in three species. It is possible that galls initiated by single nematodes largely fail to mature and or that gall initiation is favoured by the combined efforts of multiple juveniles. Juveniles must secrete plant growth regulators to induce galls, so it is possible that single juveniles might have difficulty achieving a critical concentration to effect the necessary change in the plant. The approximate balance of male and females in most galls also points to mechanism other than chance. The occurrence of galls containing multiple and variable numbers of adults in all species indicates that this process is not tightly regulated and must be to some degree a function of number of nematodes that invade the plant relative to the 46 I. T. RILEY & T. BERTOZZI number of potential gall sites. Examining gall occupancy under a range of inoculation levels could provide more evidence of the underlying pattern. Acknowledgments Tony Debicki helped with the culture 4. funesta in L. rigidum. Drs K. Owen and P. Williamson provided infested M. stipoides from Toowoomba. Ms M. Lorimer provided statistical advice. I. Riley’s position is in part funded by the Grains Research and Development Corporation. During the period the P monspeliensis galls were collected, T. Bertozzi was supported by the Australian Meat and Livestock Corporation. References Baker, K. F. (1957). The U.C. system for producing healthy container-grown plants through the use of clean soil, clean stock and sanitation. California Agricultural Experiment Station Manual 23 (University of California, Berkeley, USA.). bE SILVA, P. & Rivey, I. T. (2002). Aspects of the survival and reproduction of Anguina microlaenae (Nematoda: Anguinidae). Trans. R. Soc. S. Aust. 126: 45-49, McKay, A. C., Fisuer, J. M. & Dubé A. J. (1981). Ecological field studies on Anguina fiesta, the vector in annual ryegrass toxicity. Australian Journal of Agricultural Research 32, 917-926. Riey, 1. T., Scumirz, A. & DE SILVA, P. (2001). Anguina australis, a vector for Rathayibacter toxicus in Ehrharta longiflora. Australasian Plant Pathology 30, 361-364. TRIANTAPHYLLOU, A. C. & HIRSCHMANN, H. (1966). Gametogenesis and reproduction in the wheat nematode, Anguina tritici. Nematologica 12, 437-442. SYPHACIA (SYPHACIA) AUSTRALASIENSIS SP. NOV. (NEMATODA: OXYURIDAE) FROM RATTUS LEUCOPUS (MURIDAE) FROM PAPUA NEW GUINEA AND AUSTRALIA By L. R. SMALES* Summary Smales, L. R. (200) Syphacia (Syphacia) australasiensis sp. nov. (Nematoda: Oxyuridae) from Rattus leucopus (Muridae) from Papua New Guinea and Australia. Trans. R. Soc. S. Aust. 128(1), 47-51, 31 May, 2004. Syphacia (Syphacia) australasiensis sp. nov. is described from the caecum of Rattus leucopus (Gray) (Rodentia: Muridae) from Papua New Guinea and Queensland, Australia. The new species is distinguished from congeners with an oval cephalic plateau by the lack of lateral alae, a longitudinal ridge along the egg and a combination of measurements including spicule length, tail length, distance to anterior mamelon and size of eggs. The origins of the genus and the relationships of the species in the Australian region are discussed. Key Words: Nematoda, Muridae, Rattus, Syphacia, new species. Transactions of the Roval Society of S. Aust. (2004), 128(1), 47-51. SYPHACIA (SYPHACIA) AUSTRALASIENSTIS SP. NOV. (NEMATODA: OXYURIDAE) FROM RATTUS LEUCOPUS (MURIDAE) FROM PAPUA NEW GUINEA AND AUSTRALIA by L. R. SMALES* Summary SMALES, L.R. (200) Svphacia (Svphacia) australasiensis sp. noy. (Nematoda: Oxyuridae) from Rattus leucopus (Muridae) from Papua New Guinea and Australia. Trans. R. Soc. S. Aust. 128(1), 47-51, 31 May, 2004. Syphacia (Syphacia) australasiensis sp. nov. is described from the caecum of Rattus leucopus (Gray) (Rodentia: Muridae) from Papua New Guinea and Queensland, Australia. The new species is distinguished from congeners with an oval cephalic plateau by the lack of lateral alae, a longitudinal ridge along the egg and a combination of measurements including spicule length, tail length, distance to anterior mamelon and size of eggs. The origins of the genus and the relationships of the species in the Australian region are discussed. Key Worbs: Nematoda, Muridae, Rattus, Syphacia, new species. Introduction The pin worm genus Syphacia (Nematoda: Oxyuridae) is a cosmopolitan genus occurring in the rodent families Cricetidae and Muridae (Hugot 1988). All Australian rodents are contained within the family Muridae, either a few relatively recent arrivals, representative of the genus Ratrus (Murinae), or, the majority of — species (Hydromyinae), that arrived in Australia from Indonesia via New Guinea no more than 5-8 million years ago (Strahan 1995). The currently known pin worm species associated with the murids from Australia and the island of New Guinea are Syphacia muris Yamaguti, 1935, a cosmopolitan species; Syphacia darwini Quentin & Hugot, 1988 known only from Australian representatives of the hydromyine genus Melomys and Syphacia longaecauda Smales, 2001 known only from New Guinean representatives of the genus Melomys. Rattus leucopus (Gray) the Cape York rat is one of only six species of murids that occur in both Australia and Papua New Guinea (Flannery 1995; Strahan 1995), Examination of material collected from both locations revealed a new species of Syphacia that is described in this paper. Materials and Methods Nematodes, previously dissected from R. leucopus and stored in 70% ethanol in the South Australian Museum (SAM) or the CSIRO Wildlife Collection, Canberra (CSIRO), were cleared in lactophenol for examination. Specimens identified as Syphacia sp. “School of Biological and Environmental Sciences, Central Queensland University, Rockhampton Qld 4702, Australia. Email: Lwarner@cqu.edu.au were measured using an ocular micrometer and drawn with the aid of a drawing tube attached to an Olympus BH microscope. Measurements for 10 individuals of each sex are presented in zm as the range followed by the mean in parentheses. Syphacia (Syphacia) australasiensis (FIGS 1-11) Holotype 3 from caecum of Rattus leucopus (Gray) Brown River, Papua New Guinea, 25.vi.1968 coll. W, Ewers, SAM AHC 32142. Allotvpe 2 same data SAM AHC 32143. Paratypes Same data SAM AHC 5141. Other material examined From caecum of Rattus leucopus, E. Mcilwraith Ra., Cape York Peninsula, North Queensland, Australia 10.viii.1990, coll. P. Catling and P. Haycock, CSIRO N3325, 5 9°, 3 d6; Brown River, Papua New Guinea, 25.vi.1968, coll. W. Ewers, AM AHC 5201, 5152, 5149, 95 92,16. Site in host Caecum and colon. Description Small nematodes, typical oxyurid shape, with transverse cuticular striations. Cephalic inflation distinct. Cephalic plateau oval, elongated laterally; distance between amphids 30, amphids situated between cephalic papillae; cephalic papillae on lateral projections; six labial papillae; mouth 48 L. R. SMALES Figs 1-11 Syphacia (Syphacia) australasiensis sp. nov. 1. Male, anterior end, dorso-ventral view. 2. Male, lateral view. 3. Female, lateral view. 4. Anterior end, optical section, dorso-ventral view. 5. En face view. 6. Anterior end, optical section, lateral view. 7. Spicule and gubernaculum, lateral view. 8. Vagina, lateral view. 9. Male, posterior end, ventra view. 10. Excretory pore, lateral view. 11. Egg. Scale bars 1, 8,9, 10, 11 25m; 2, 502m; 3, 100pm; 4, 5, 6, 7, 10m. NEW NEMATODE FROM AUSTRALIA AND NEW GUINEA 49 opening simple, three small distinct pseudolabia. Oesophagus with isthmus, terminating in spherical bulb. Excretory pore posterior to oesophageal bulb. Nerve ring surrounding anterior oesophagus. Deirids not seen. Alae absent. Male Total length 690 — 1250 (810), maximum width 80 — 102 (95). Oesophagus 142 — 208 (192) long, oesophageal bulb 46 — 55 (51) in diameter. Nerve ring 75 — 120 (94), excretory pore 228 — 462 (259) from anterior end. Three mamelons with prominent annulations, spines not observed; first mamelon 297 583 from anterior end, second mamelon close behind first, third mamelon close behind second: mamelons all 35 — 40 long. Tail 142 — 181 (155) long. Spicule needle shaped 61 — 68 (65) long; gubernaculum 29 — 36 (31) long, ventral barb not observed. Two pairs preanal, one pair large post anal caudal papillae. Female Total length 1900 — 2900 (2400), maximum width 160 — 268 (170). Oesophagus 187 — 355 (298) long, oesophageal bulb 73 — 82.5 (78) in diameter. Nerve ring 107 — 135 (122), excretory pore 315 — 570 (460). Vulva with prominent knob, 436 — 804 (655) from anterior end. Tail 335 — 436 (385) long. Eggs with longitudinal ridge, 82.5 — 95 (87.5) by 29.5 — 36.3 (33). Etymology The species is named to reflect that it is found in both Papua New Guinea and Queensland. Discussion The nematode species described above belongs to the genus Syphacia Seurat, 1916 because it possesses the suite of characters proposed by Hugot (1988, see also figs 23, 24 and 39) to distinguish the genus. It belongs within the subgenus Syphacia Seurat, 1916 because it has neither the rectangular cephalic plateau and well developed triangular shaped lateral alae of the subgenus Crictoxyuris Hugot, 1988 nor the short conical tail characteristic of the subgenus Seuratoxvuris Hugot, 1988. Rather, it has an oval cephalic plateau, lacks cervical alae and well developed deirids. Syvphacia (Syphacia) australasiensis n. sp. in en fuce view most clearly resembles a cluster of species from murid hosts (Rattus species) from the Australasian biogeographic region, namely S. muris Yamaguti, 1935, 8. darwini Hugot & Quentin, 1985, S. fongaecauda Smales, 2001 and S. sulawesiensis Hasegawa & Tarore, 1996. It differs from each of these species in the form of the vulva, having a longitudinal ridge in the egg and males lacking lateral alae. It further differs from S. muris in having an oval rather than square cephalic plateau; from S. longaecauda in having a smaller cephalic plateau, 30u.m between the amphids rather than 45 — 50pm; from S. darwini in having three, not two, mamelons and from S. su/awesiensis in having the females lacking lateral alae. Syphacia australasiensis differs from S. /ophuromys Quentin, 1966 and S. megaloon Quentin, 1966, also lacking lateral alae in the morphology of the egg and vulva and in en face aspect. S. lophuromys and S. megaloon have a cephalic plateau elongated laterally with papillae and pseudolabia characterised as Group VIII by Quentin (1971). S. /ophuromys and S. megaloon are found only in African murids. Further, S. australasiensis differs from each of the above species in one or more measurements of oesophagus, tail, spicule gubernaculum or eggs (Table 1). The oxyurids, subfamily Syphaciinae are generally considered to have coevolutionary relationships with their hosts (Hugot, 1988). The similarities between S. australasiensis, S. darwini, S. longaecauda and S. sulawesiensis are therefore not surprising. Each has an oval cephalic plateau with distinct pseudolabia, “lips”, and mouth surrounded by 6 labial papillae; each occurs in an endemic murid host, Raflus species or Melomys species; each has evolved within the Australasian biogeographic region, Sulawesi being to the east of Wallace’s line, the boundary between the Asian and Australasian faunal regions (Raven, 1935). The relationship between these four species and S. muris also occurring in Rafts species hosts, but cosmopolitan in distribution, is not clear. Svyphacia muris is found in R. ratttus (Linnaeus) and R, norvegicus (Berkenhout) (see Smales, 1997) both of which are recent arrivals in Australia. Given that Hugot & Quentin (1985) found that S. mauris from endemic Rattus spp. in Australia, corresponded morphologically and morphometrically to the description from cosmopolitan Rattus spp., the suggestion of Hasegawa & Tarore (1996) that infections in Australian endemic murids have been acquired from the recent arrivals seems sound. The endemic Svphacia species may therefore have been derived from syphaciine populations introduced into the region as their rodent hosts invaded the Island of New Guinea and then Australia (Flannery, 1995). Subsequently speciation took place in both regions. Syphacia longaecauda has been found only in Me/omys spp. endemic to New Guinea while S. darwini is known only from Melomys cervinipes hosts endemic to Australia. This suggests that there has been no migration of Melomys between the two islands. Very few data are L. R. SMALES 50 ; 9€-6C ‘ Oe=Le ; 8C : Ie , 0€ . LV-6£ ej Ov winjnoewdeqn’) 7 89-19 7 O8-8L . 8Y - 19 i; SL : LO-9L : 08 anaidg 9£-6C 6C TC Ch-veEXx XS6-C8 7 X98-LL 7 O€X06 - O9XOS1 - 8PXOTT 7 001-76 SEX06 ; S854 9th I8T OL9 001 OS8 06r “SSE “Cr “brs “v8 oog SII 00c OI OI OVC “919 “OLE OOL LY TRL c8 gs cll LL Lol £6 EL “OF -96 369 0€ Or : : 06 09 al Gl, “09 001 OS WIPE Qing SeO Sse 807 6CY pOE Osh Sse “L81 “cr “BLT “19 Ocl = O91 00€ Z Ole 00 “807 “OVE OcE O&T psus| S90, e8s Ole! OS8 a “LOT : “S16 . OOS . OSY 3 OfY : -009 - O€8 UOPSUTBUL Js | vO8 I8Z 986 “OEP : “C99 . 09¢ - O8? : OSS - “SOL 3 0£6 i BANA OLS COV O9L Seo ceo 9eV “SIE “8CT “OSS -009 STC 06T O8C OCT Oce 00E “OLY “GLE OSL OO0r 40d AIOJOIOXO Sel 01 vs ell Lcl srl BULL OAIOU 0} “LOL “SL “SII “OC Se OL O01 08 Ol O01 “oll “HII 01 OIL pus Jue WOly 89C cOl eee col 09€ 091 “091 “08 “887 vel 06 SL OSC OST OLE Sil “Ole “001 OCC SOl upian Apog 006c ~—-OSTI Ors Olol O0rL OSOT “0061-069 “OL8E — -OS9I Ocrl Orit 000c —09ET Ococ —OOKET “000b — -0091 096€ S981 yisus| Apog SISUAISDIDAISND “S SISUAISDMDINS “S SLINUL “S UOO]DBIUL *S sAwosimnydo] “§ ppnvrapsuo] ‘S IUIMADP “5 3} £ fo) 2 3 £ 3} ‘3 3} 2 3} £ é ‘4 (966[ ‘2401D]. PUD DMDSAaSVT] ajduipxa 10f 2a8) 10.18 ul ag ADU SLINUA *S 4 4Of Sjuaua]a asay) 10f UAAIS SJUaUAINSHAU AY) JO DS.1AA adIA pu UUUNJOI & ay) Ul Y past) Aaysia aavy oj Avaddp yipia pun yj8ua] snSpydosao sp [Jam sp aiod K10jatdXa PUB BULI BAsIU YJ JO LOLIAJUD AY) WOU AIUDISIP AY) LOf UJUANG YW? JOsNpY Aq Uaass sp sjuauainsvayy “SISoY JUAapos UDI.AISNP WOAf (696 [) UNUANG 3 JO8NpY Aq Uadss asoy) aid SINUS AOf SuaUAINSDAPY [YT ‘SAJDUWS PUP 966] ‘24OAD] IP DMHBasvp] “CRG] ‘UNMANG Y? JOBNLT “1Z6T ‘996 ‘UIUaNG Aq vadrs sp SISUAISAMEINS *S puD soxtuOnYydo] *s ‘UOO|RBdUI “S “epNRooeBuUO] *S ‘TULALIEp “S ‘sLINW *S fo asouy YIM pasduios ApnyS siy) sisuaiseyeysne evioeydAs fo sjuamainspay “| TEL, NEW NEMATODE FROM AUSTRALIA AND NEW GUINEA 51 available from rodent hosts from New Guinea localities however, so there may be populations of S. darwini present there yet to be discovered. Rattus leucopus occurs in both regions and in both cases harbours S. australasiensis. This suggests that S. australasiensis migrated with R. leucopus into Australia. More data from murids across the region is needed before relationships can be derived and firm conclusions drawn. Acknowledgments Sincere thanks to Jan Forrest from the South Australian Museum and Dr. David Spratt from CSIRO Sustainable Ecosystems for lending me the material. References FLANNERY, T. (1995) Mammals of New Guinea (Reed Books, Chatswood). HAsecawa, H. & TARORE, D. (1996) Syphacia (Svphacia) sulawesiensis n. sp. and S. (S.) muris (Yamaguti, 1935) (Nematoda: Oxyuridae) collected from Rattus xanthurus (Graf, 1867) (Rodentia: Muridae) in Northern Sulawesi, Indonesia. Trop. Zool. 9, 165-173. Hucot, J.-P. (1988) Les nématodes Syphaciine parasites de rongeurs et de lagomorphes. Mem. Mus. Nat. Hist. Nat. Sér A 141, 1-148. ‘ & QUENTIN, J.-C. (1985) Etude morphologique de six espécies nouvelles ou peu connnues appartenant au genre Syphacia (Oxyuridae: Nematoda), parasites de Rongeurs Cricétidés et Muridés. Bull, Mus. Nat. Fist. Nat. Ser: 47, 383-400. QUENTIN, J.-C. (1966) Oxyures de Muridae africans. Ann.Parasit. 41, 443-452. _ (1971) Morphologie comparée des structures céphaliques et génitales des oxyures du genre Syphacia. Ann. Parasit. Hum. Comp. 46, 15-60. RAVEN, H. C. (1935). Wallace’s Line and the distribution of Indo- Australian mammals. Bull, Am. Mus. Nat. Hist. 68, Article 4. SMALES, L. R. (1997) A review of the helminth parasites of Australian rodents. Aust. J. Zool. 45, 505-521. — (2001) Svphacia longaecauda nn. sp. (Nematoda: Oxyuridae) Syphacinea from Me/omys spp. (Muridae: Hydrominae) from Papua New Guinea and Irian Jaya, Indonesia. Parasite 8, 39-43. STRAHAN, R. (1995) The Mammals of Australia (Reed Books, Chatswood). GASTRIC NEMATODES, INCLUDING A NEW SPECIES OF ABBREVIATA (NEMATODA: PHYSALOPTERIDAE) FROM THE MANGROVE MONITOR VARANUS INDICUS (REPTILIA: VARANIDAE) By H. I. JONES Summary Jones, H. I. (000) Gastric nematodes, including a new species of Abbreviata (Nematoda: Physalopteridae) from the mangrove monitor Varanus indicus (Reptilia: Varanidae). Trans. R. Soc. S. Aust. 128(1), 53-59, 31 May, 2004. Three species of nematode, Tanqua tiara (von Listow, 1879), Heliconema longissima (Ortlepp, 1922), and Abbreviata melanesiensis sp. nov., were recovered from the gastrointestinal tracts of 124 Varanus indicus (Daudin, 1802). Tanqua tiara occurred in two-thirds of the lizards, at a mean prevalence of 7.2 worms per infected host. It is suggested that the low intensity and geographically variable prevalence of this nematode may relate to the distribution of the invertebrate intermediate host in the discrete and discontinuous insular habit of the lizard. Abbreviata melanesiensis sp. nov. was recovered from two lizards, and is distinguished from other species in this genus principally by the enlarged tip of the right copulatory spicule, the thick sheath which envelops the retracted left spicule, the relatively conspicuous phasmids in females, and the elongated, thin-walled eggs. A single male Heliconema longissima was recorded. Key Words: Varanus indicus, Tanqua tiara, Abbreviata melanesiensis, Heliconema longissima, Solomon Islands, Papua New Guinea, Indonesia, Australia. Transactions of the Royal Society of S. Aust. (2004), 128(1), 53-59. GASTRIC NEMATODES, INCLUDING A NEW SPECIES OF ABBREVIATA (NEMATODA: PHYSALOPTERIDAE) FROM THE MANGROVE MONITOR VARANUS INDICUS (REPTILIA: VARANIDAE) by H. IL. JONES Summary Jonrs, H. 1. (000) Gastric nematodes, including a new species of Abbreviata (Nematoda: Physalopteridac) from the mangrove monitor Varanus indicus (Reptilia: Varanidae). Trans. R. Soc. S. Aust. 128(1), 53-59, 31 May, 2004. Three species of nematode, Zanqua tiara (von Linstow, 1879), Heliconema longissima (Ortlepp, 1922), and Abbreviata melanesiensis sp.noy., were recovered from the gastrointestinal tracts of 124 Varanus indicus (Daudin, 1802). Tangua tiara occurred in two-thirds of the lizards, at a mean prevalence of 7.2 worms per infected host. It is suggested that the low intensity and geographically variable prevalence of this nematode may relate to the distribution of the invertebrate intermediate host in the discrete and discontinuous insular habitat of the lizard. Abbreviata melanesiensis sp. nov. was recovered from two lizards, and is distinguished from other species in this genus principally by the enlarged tip of the right copulatory spicule, the thick sheath which envelops the retracted left spicule, the relatively conspicuous phasmids in females, and the elongated, thin- walled eggs. A single male //eliconema longissima was recorded. Key Worps: Varanus indicus, Tanqua tiara, Abbreviata melanesiensis, Heliconema longissima, Solomon Islands, Papua New Guinea, Indonesia, Australia. | Introduction The mangrove monitor, Varanus indicus (Daudin, 1802), has a wide and discontinuous distribution from Sulawesi in eastern Indonesia, east through Papua New Guinea and northern Australia to the Solomon, Caroline and Marshall Islands (De Lisle 1996). The taxonomy of the monitor lizards in the Varanus indicus complex has recently been revised (Boehme ef a/. 1994, Philipp ef al. 1999) to include several closely-related species. Varanus indicus sensu lato is found close to water, on beaches or riverbanks, and in swampy and mangrove areas (Cogger 1992). It is largely a terrestrial feeder, and its diet principally comprises crabs (especially grapsoid), smaller reptiles, mammals and birds, and occasionally turtles’ eggs (McCoy 1980). It is a diurnal lizard, and when alarmed will take to water (Cogger 1992), although in the Solomon Islands it will invariably climb a tree (McCoy 1980). Confusion in the nomenclature of Varanus lizards has resulted in a number of nematodes being attributed erroneously to Varanus indicus trom India (Ortlepp 1922, Mirza 1934, Sharief 1957, Deshmukh 1969, Ali & Ilyas 1969). Since these records are far to the west of the known geographical range of V. indicus, and as all state or imply that the lizards were locally obtained and were not exotic specimens held in captivity, it is probable that these nematode Microbiology, School of Biomedical and Chemical Science, University of Western Australia, Nedlands, Western Australia 6009, Phone: (08)9346-1270 or (08)9386-6 140. Email: hjones@cyllene.uwa.edu.au. records pertain to V. bengalensis (Daudin, 1802), or possibly V. flavescens (Hardwick and Gray, 1827) or V. griseus koniecznyi Mertens, 1954 (De Lisle 1996). Little is known of the internal parasites of Varanus indicus. The nematode Kalicephalus megacephalus Schad, 1962 was described from this host from Florida Island in the Solomon Islands by Schad (1962). Tangua tiara (Von Linstow, 1879) is primarily a nematode parasite of lizards in the genus Varanus, and has been recorded from a number of Varanus spp. from Africa to southeast Asia and northern Australia, in aquatic or coastal habitats, but has not hitherto been reported from /. indicus (see Gibbons and Keymer 1991). The present study was therefore undertaken to ascertain the nematode fauna of Varanus indicus sensu lato, as part of a study of the gastrointestinal nematodes of varanid lizards. Materials and Methods The results of dissections of 124 Varanus indicus are presented. Thirty seven Vi) indicus held in the collection of the Australian Museum, Sydney, and one from the collection of CSIRO Sustainable Ecosystems, Canberra, were examined. Dennis King forwarded nematodes recovered from stomach contents of 20 i. indicus collected from islands and small archipelagos of Bandaneira, Aru, Kai Besar, Yamdena and Selaru in the Moluccas in south and eastern Indonesia during Western Australian Museum/Museum Zoologicum Bogoriense expeditions in 1992/1993. In addition, nematodes from stomachs of sixty-six V. indicus collected at Maningrida in the Northern Territory by Tim Schultz 54 H. 1. JONES during 2001 were removed by Alain de Chambrier and identified by lan Beveridge and the author. The snout-vent length (SVL) of 58 hosts examined varied from 135 and 550 mm (mean = 335mm). Measurements were not available from lizards from Maningrida. The viscera were removed, and stomachs (and the intestines from the Australian Museum specimens) examined for helminths. Food residues were noted. All nematodes collected were cleaned, and stored in 70% alcohol with 10% glycerine. They were subsequently cleared in chlorolactophenol for examination. All specimens have been returned to the Australian Museum, Western Australian Museum, CSIRO and_ the Australian Helminth Collection respectively. Results Helminths recovered. Three species of nematode were recovered. The predominant species was Tanqua tiara, which was present in 75/124 lizards (60.5%, Fig. 1); specimens lodged as Australian Museum W 28660-W 28674; CSIRO Canberra N5294: Western Australian Museum V 4374 and V 4375, and in the Australian Helminth Collection. Prevalence was highest in the lizards from Maningrida (84.8%), and lowest from the Moluccas and Solomon Islands (32.8%). Total mean intensity of T. tiara was 7.2 worms/host. Four Vo indicus, collected from Adelaide River and Maningrida in the Northern Territory, contained >20 7: tiara (22 — 123); in the remaining 71 infected hosts 7. tiara numbers ranged from | — 16 (mean: 4.2). In four hosts some nematodes were attached with their heads buried in the stomach mucosa. A single male //eliconema longissima (Ortlepp, 1922) was recovered from a lizard from Maningrida in the Northern Territory. A new species of Abbreviata was recovered from two hosts collected from New Britain and from Woodlark Island, Papua New Guinea. There was no relation between the size of the host lizard, type of food residues in the stomach, and the presence or intensity of infection with 7) tiara. Both prevalence and intensity of this worm from lizards collected from the Moluccan islands in south-eastern Indonesia were lower (2 of 20 infected with one and two worms, respectively) than those collected from Papua New Guinea or the Solomon Islands, with highest prevalence and intensity in lizards from northern Australia. Neither of the two lizards with the new species of Abbreviata was concurrently infected with 7. tiara. One of the two lizards infected with this species of Abbreviata was one of the largest dissected (500 mm SVL). Food residues. In the 37 V. indicus from Papua New Guinea and the Solomon Islands, crustacea, in three I I 130 E 140 NORTHERN TERRITORY SOLOMON ISLANDS & Fig. 1. Map showing distribution of Zangua tiara in Varanus indicus. Open circle, lizard without infection; black circle, lizard with infection. M: Maningrida, where 56 of 66 lizards were infected. (No location data available for two lizards). PARASITIC NEMATODES IN V4ARANUS INDICUS cases identifiable as crabs. formed the most commonly recovered food source, and occurred in 16 lizards. In addition, a snake (Candoia sp.; Boidae), an agamid lizard skull, an unidentified lizard, a frog, reptile eggs, cicadas, a grasshopper, and a beetle were each recovered from a single host stomach, and vertebrae from an unidentified animal, and molluscs from two hosts each. Stomach contents from the 20 lizards from the Indonesian islands included 10 with insect remains, one with a gecko, one with a bird, one with reptile eggs, one with a crab, and four with plant residues. Twenty one of these fifty seven stomachs had no food residues; no fish remains were identified. Taxonomy Order Spirurida Superfamily Physalopteroidea Family Physalopteridae Genus Abbreviata Travassos, 1920. Abbreviata melanesiensis sp. Nov. Holotype Male, Australian Museum W 28675, from stomach of Varanus indicus, AM R129614, Amelei village, West New Britain, Papua New Guinea (06° 06’ S, 150° 37’ E), collected 27.x11. 1988. Allotvpe Female, AM W 28676, same data as for holotype. Paratypes 16 males, 9 females, 4 larvae/immatures, AM W 28677, same data as for Holotype. Additional non- type specimens: three males, four females and two immatures, AM W 28678, from /. indicus R124815, collected Guasopa, Woodlark Island, Milne Bay Province, Papua New Guinea (09° 15’ S, 152°, 56’ E), collected 09.viil.1987. Diagnosis With characteristics of the genus, viz. two large lateral pseudolabia, with externolateral tooth present on each pseudolabium, a bifid internolateral tooth and two double pairs of submedian teeth; male with wide ornamented caudal alae united anteriorly on ventral surface of body, supported by four pairs of pedunculate papillae, markedly unequal spicules, females with vulva in anterior portion of body. Mouth with dorsal and ventral corner denticles. Spicules well sclerotised, left four times length of right; left spicule in thick walled sheath, terminating in a very fine point, right spicule curved ventrally, with pointed, weakly-sclerotised enlargement at tip. Female tail slightly attenuated, vulva flush with body wall without extensions; eggs thin-shelled, elongate, length approximately twice width. Wn al Description (Fig. 2) Small to medium worms, males not greatly smaller than females, tapering at both ends, fine transverse cuticular striations. Mouth surrounded by two pseudolabia, each bearing large externolateral apical tooth and small bifid internolateral tooth. Bifid submedial tooth on dorsal and ventral border of each pseudolabium. Four to 5 small, regular denticles at dorsal and ventral median surface of each pseudolabium. Two sessile papillae and amphid on external surface of each pseudolabium. Cervical collar present. Nerve ring surrounds muscular oesophagus near its posterior end. Glandular oesophagus of uniform width, wider than muscular portion. Cervical papillae and excretory pore on external surface posterior to origin of glandular oesophagus. Male Caudal alae meet anteriorly, extend just beyond tip of tail posteriorly. Alae supported by four pairs of pedunculate pericloacal papillae and three pairs of very short pedunculate or sessile caudal papillae, of which the central pair is about one-third the distance between the first and second pair. Papillae on ventral surface sessile, 3 immediately anterior to the cloaca, and 2 pairs immediately posterior to the cloaca. Caudal tubercles arranged in rows, extending from anterior border of alae, converging towards cloaca, and running parallel to one another lateral to cloaca on ventral surface of tail and adjoining alae; diminishing in size posteriorly, terminating as small scattered tubercles at level of posteriormost caudal papillae. Spicules dissimilar, unequal. Left spicule approximately 4 times length of right, uniformly sclerotised, terminating in very fine tapering point, enclosed in conspicuous thick sheath. Right spicule thicker than left, heavily sclerotised, curved ventrally, and with a weakly- sclerotised pointed enlargement, slightly concave on one side, at the tip. Female Tail short, terminating in a slight attenuation; phasmids relatively conspicuous at two-thirds distance along tail. Vulva a transverse slit flush with body wall, without extensions or altered adjacent cuticle, posterior to commencement of intestine, about one-quarter of distance from anterior end of worm. Two ovaries, uterus with four branches, one of which often extends a variable distance anteriorly beside the posterior portion of glandular oesophagus. Eggs with smooth thin shells, elongate, embryos not visible, length almost twice width. Measurements (mm; range with mean in brackets) Males (N:6): length 18 — 24 (21.7); maximum width 0.36 — 0.54 (0.45); muscular oesophagus length 0.26 56 H. I. JONES Fig. 2. Abbreviata melanesiensis sp. noy. A, anterior end, male paratype, lateral view; B, anterior end, en face, female paratype; C, male tail, slightly-oblique ventral view; D and E, distal tip of right spicule, male paratype; F, tail, female paratype, lateral view; G, eggs in oviduct and vulva, female paratype, lateral view; H, egg. Scale bars, E and H: 100um, all other figures, 200um. PARASITIC NEMATODES IN VARANUS INDICUS 37 — 0.36 (0.33); muscular oesophagus width 0.10 0.14 (0.12); glandular oesophagus length 2.8 — 3.3 (3.2); glandular oesophagus width 0.24 — 0.28 (0.26); nerve ring* 0.26 — 0.38 (0.32); cervical papillae* 0.36 — 0.68 (0.53); excretory pore* 0.56 — 0.72 (0.61); tail 0.96 — 1.32 (1.16); distance between caudal papillae | and 2, 0.056 — 0.16 (0.11); distance between caudal papillae 2 and 3, 0.14 — 0.19 (0.16); left spicule 1.320 — 1.680 (1.470); right spicule 0.280 — 0.360 (0.320). Females (N: 6) length 26 — 33 (29.3); maximum width 0.62 — 0.74 (0.70); muscular oesophagus length 0.36 — 0.44 (0.41); muscular oesophagus width 0.14 — 0.18 (0.15): glandular oesophagus length 3.36 — 4.40 (3.93); glandular oesophagus width 0.26 — 0.41 (0.32); nerve ring* 0.34 — 0.38 (0.36); cervical papillae* 0.48 — 0.60 (0.54); excretory pore* 0.62 — 0.86 (0.70); tail 0.34 — 0.44 (0.38); vulvat 1.9 — 3.7 (2.78); 19.4% — 27.8% (23.3%) of distance from anterior end (* distance from anterior end; + distance posterior to oesophago- intestinal junction). Discussion Taxonomy Abbreviata melanesiensis sp. nov. is distinguished from other species of Abbreviata occurring in reptiles from the Australo-Papuan region by a suite of characters. The size, general form, characteristics of the anterior end and the male copulatory bursa are shared with a number of other species. However, the weakly-sclerotised enlargement at the tip of the right spicule 1s unique; a similar feature occurs in A, hastaspicula Jones, 1979, in which species however it is well sclerotised (Jones 1979). Furthermore, in that species the left spicule is considerably shorter (0.620 — 0.700 mm) and the right spicule longer (0.590 — 0.670 mm: Jones 1979) than in A. melanesiensis sp. nov., and the sclerotisation of the left spicule often appears discrete and discontinuous; in addition, in A. hastaspicula the female bears a tubular extension from the vulva. Eggs of A. me/anesiensis sp. nov. are elongate and thin walled; most species of Abhreviater in this region possess eggs with thicker and denser shells (Jones 1983b, 1988) except A. hastaspicula and 4. perenticola Jones, 1985, whose thin shelled eggs are subspherical. Other characteristics which differentiate this species include less disparity in length between males and females, the thick sheath that envelops the retracted left spicule, and the readily visible phasmids on tails of females. In other species of Ahbbreviata described from Australia and Papua New Guinea, except A. /evicauda Jones, 1983, from V. tristis (Schlegel, 1839) the penultimate caudal papillae are closer to the anterior papillae (Jones 1986) than in A. melanesiensis sp. noy. The enlargement at the tip of the right spicule distinguishes 4. melanesiensis sp. noy. from six other species of Abbreviata recorded from Papua New Guinea, viz. 4. oligopapillata (Kreis, 1940) (see Jones 1979), A. multipapillata (Kreis, 1940), A. natricis (Kreis, 1940), A. heterocephala (= Kreisiella) (Kreis, 1940), A. confusa Johnston & Mawson, 1942, (see Jones 1983a) and A. katilensis (Jones, 1979). A. borneensis Schad, 1959 from V. rudicollis Gray, 1843 in Sarawak also has the posterior portion of the tail free from tubercles, as in A. oligopapillata (Schad 1959). In A. melanesiensis sp. nov. the delicate enlargement at the tip of the right spicule is not visible unless this is extended through the cloaca, and if this spicule is retracted, dissection is necessary to ascertain its character; all specimens in the type host had the right spicules retracted, but in all four males in the second host the right spicules were extended and this character was clearly visible. The small denticles at the dorsal and ventral mouth margins are often not visible if obscured by the cervical collar. Biology The absence of a relationship between host size and numbers of 7. tiara contrasts with the findings of Shine e¢ al. (1998), who found that 7) tiara were present at a higher intensity in juvenile Mi sa/vator, They also found a significant difference in infection with this worm between two sites in Sumatra, though the intensity of worms was similar to findings from V. indicus in the present study (mean 6.1 worms per host). The arthropod intermediate host for 7. tiara is not known, but as Varanus specimens infected with this worm inhabit swampy, mangrove or riverine habitats, a crustacean intermediate host is possible. Although V. indicus feed on a wide range of prey, the fact that crustacean remains were found in 16 of 37 from Papua New Guinea and the Solomon Islands, and in only 2 of 20 from the Moluccas, (food data not available from lizards from Maningrida) illustrate local or regional differences in prey availability. The distribution of the worm in the present study may be related to the discontinuous nature of insular habitats. In the present study highest intensity of 7. tiara, 22 123 worms, occurred in indicus examined from northern) mainland Australia (Adelaide River and Maningrida), which may provide both more opportunities for the spread of worms and intermediate hosts than from small isolated habitats. Furthermore, 7: fiara have been reported from V. panoptes Storr, 1980, Vo gouldii (Gray, 1838) and Vo mertensi Glauert, 1951, from aquatic habitats in northern Australia at mean intensities of 18.9 (max. 120 worms), 20.5 (max. 50 worms) and 7.2 (max. 30 worms) respectively; an absence of correlation with Abbreviata spp. infection 58 H. L. JONES was also noted in that study (Jones 1988). The similar species 7) ophidis Johnston & Mawson, 1948, described from the aquatic file snake Acrochordus sp. was present in all eight Acrochordus sp. examined from north and northwest coastal Queensland, at numbers ranging from seven to 179 per host (mean, 51); (Jones 1978). As K. indicus, Acrochordus arafurae McDowell, 1979 and A. granulatus (Schneider, 1799) are sympatric over much of their range, it is possible that these two species of Zanqua are also sympatric. The single male #7. /ongissima reported in the present study was probably an accidental infection, taken with infected prey. The type host and type locality of Heliconema longissima (Ortlepp, 1922) are given as ‘snakes, Australia’, although all other specimens have been recorded from anguilliform fish, and Ogden (1969), considered that the type host identification is probably in error. However, H. longissima occurred in 3/5 specimens of the aquatic colubrid snake Fordonia leucobalia (1, 23 and 41 nematodes per host; Jones 1978). The scarcity of Abbreviata melanesiensis sp. nov. in Ki indicus contrasts with findings of prevalence and intensity of Abbreviata spp. in other species of Varanus and other large terrestrial reptiles in northern and arid Australia, in several species of which infection with Abbreviata spp. occurs at high prevalence and intensity (Jones 1983b, 1988). Epidemiological evidence suggests that termites might have a role to play in the life-cycles of species of Abbreviata from arid regions (Jones 1995), Since only two K. indicus were infected with Abbreviata sp., no conclusions can be drawn from the absence of concurrent infection with 7) tiara, Since Varanus indicus feeds on a wide variety of invertebrate (and, in the larger specimens, vertebrate prey; McCoy 1980) the nematodes present may relate to differences in diet; the predominance of aquatic prey in these lizards suggests that the intermediate hosts of T. tiara may be aquatic invertebrates. The fact that one of the two hosts infected with A. melanesiensis was particularly large may be significant. The larger of the two infected lizards contained a Candoia sp. snake prey item, and though it is possible that this Abbreviata infection was spurious, this is unlikely as several of the nematodes were already attached to and apparently feeding on the external surface of the snake. Conclusion This study demonstrates that despite the wide range of prey items consumed by indicus, this lizard supports gastric nematodes in low numbers, and at a moderate prevalence, with only three species recorded. No intestinal nematodes were recovered. It is possible that the large range of prey types may inhibit the development of parasite cycles, particularly in parasite species with narrow intermediate-host specificity. The low intensity of 7. fiara and geographically uneven distribution may relate to the insular and discontinuous nature of the region from which FV. indicus was examined; in this regard the higher intensities of this nematode recorded from other species of Varanus in northern mainland Australia may be significant. Further conclusions cannot be drawn without knowledge of the arthropod intermediate hosts required by this nematode to complete its life-cycle. Studies on the gastrointestinal parasites of other larger reptiles in this region may reveal a wider range of hosts for A. melanesiensis. Acknowledgements | thank Allen Greer for allowing me access to the lizards under his care in the Australian Museum, and for providing facilities, Ross Sadlier for practical assistance, David Spratt for allowing me to examine a lizard in the collection of CSIRO Sustainable Ecosystems, Canberra, and Brad Maryan for assistance at the Western Australian Museum. | am grateful to Alain de Chambrier for collecting nematodes from lizards collected at Maningrida; to lan Beveridge for identifying and forwarding these specimens, and to the late Dennis King for collecting nematodes during the course of his own studies, and to Mrs. Ruth King for making his papers available for my perusal. References Al, S. M. & Ityas, R. (1969) Neoxysomatium longicaudatum n, sp. from Varanus indicus in Marathwada, India. Marathwada Univ. J. Science 8, 73- 75. BOEHME, W., HorN, H. G. & ZIEGLER, T. (1994) On the taxonomy of the Pacific monitor lizards (Varanus indicus complex): resurrection of Varanus doreanus (A. B. Meyer, 1874) and description of a new species. Salamandra 30, 119-142. CoacGcpr, H. G. (1992) Reptiles and Amphibians of Australia. (Sth Edition, Reed, Chatswood, NSW 2067; I- 775). Dr Liste, H. F. (1996) The Natural History of Monitor Lizards. Krieger Publishing Company, Malabar, Florida, 1-201. DesHMukH, P. G, (1969) A new species of the genus Herpetostrongvlus (Nematoda) from Varanus indicus. Vest. Cesk. Spol. Zool. 33, 211-213. PARASITIC NEMATODES IN VARANUS INDICUS 59 Gippons, L. M. & Krymer, I. F. (1991) Redescription of Tanqua_ tiara (Nematoda, Gnathostomatidae), and associated lesions in the stomach of the Nile monitor lizard (Varanus niloticus). Zool. Scr. 20, 7-14. Jones, H. I. (1978) Gastrointestinal nematodes from aquatic Australian snakes. Mem. Queens! Mus.18, 243-254. (1979) Nematodes from Papua New Guinean snakes. Mem. Queensl. Mus.19, 393-397. (1983a) A collection of nematodes from snakes from Papua New Guinea. Syst. Parasitol. 5, 131-134. — (1983b) Abbreviata (Nematoda: Physalop- teroidea) in lizards of the Varanus gouldii Complex (Varanidae) in Western Australia. Aust. J. Zool. 31, 285- 298. : (1986) Differences in caudal morphology in male Abbreviata levicauda (Nematoda: Physalop- teridae)in two sympatric species of Varanus (Reptilia: Varanidae). J. Parasitol. 72,185-186. Varanus (Reptilia) from tropical northern Australia, with particular reference to the genus Abbreviata (Physalopteridae). Aust. J. Zool. 36, 691-708. (1995) Gastric nematode communities in lizards from the Great Victoria Desert, and an hypothesis for their evolution. Aust. J. Zool. 43, 141-164. (1988) Nematodes from nine species of MeCoy, M. (1980) Reptiles of the Solomon Islands. Wau Ecology Institute. Handbook 7. Wau Ecology Institute, Wau, Papua New Guinea. Mirza, M. B. (1934) Sciurus palmarum als ein interessante Wirt von Physaloptera sp. Z. f. ParasitKde 6, 638-641. OGpEN, C. G. (1969) A revision of the genus Heliconema Travassos, 1919, Physalopteridae (Nematoda). J. Nat. Hist. 3, 423-431. OrtLepp, R. J. (1922) The nematode genus Phvsaloptera Rud. Proc. Zool. Soc. Lond. 999-1107. Puiuipe, K. M., BorumMe, W. & ZEIGLER, T. (1999) The identity of Varanus indicus; redefinition of a sibling species coexisting at the type locality (Sauria, Varanidae, Varanus indicus group). Spixiana 22, 273-287. ScHAbD, G. A. (1962) Studies on the genus Kalicephalus (Nematoda: Diaphanocephalidae). Il. A taxonomic revision of the genus Kalicephalus Molin 1861. Canad. J. Zool. 40, 1035-1065. SHARIFF, A. (1957) On a new species of trichostrongylid nematode from Hyderabad, India. Ann. Mag. Nat. Hist. ser.12, 10, 705-709. SHINE, R., AMBARIYANTO, HARLOW, S. & MUNPUNI (1998) Ecological traits of commercially harvested water monitors, Varanus salvator, in northern Sumatra. Wildlife Res. 25, 437-447. ENRICHMENT-PLANTING OF THE WOODY CLIMBERS MARSDENIA AUSTRALIS AND RHYNCHARRHENA LINEARIS IN NORTH-WESTERN VICTORIA By R. F. PARSONS* Summary Parsons, R. F. (2000). Enrichment-planting of the woody climbers Marsdenia australis and Rhyncharrhena linearis in north-western Victoria. Trans. R. Soc. S. Aust. 128(1), 61-66, 31 May, 2004. Marsdenia australis and Rhyncharrhena linearis are woody climbers (family Asclepiadaceae) found in semi-arid to arid areas of all mainland states of Australia. Where studied, they are declining drastically, probably due to grazing. In Victoria, their conservation status is vulnerable. In 1991, in Hattah-Kulkyne National Park, a site was found carrying single plants of each species. At this site, 384 seedlings and 1,741 seeds of Marsdenia and 240 seedlings of Rhyncharrhena were planted underneath trees of Eucalyptus socialis in 1991-1992, half of the seedlings in mammal-proof cages. After high seedling mortality due to mammal grazing and especially to drought in the first summer, only six of the planted seedlings, all of Marsdenia, survived by the end of 1992. All of these, and a further 15 Marsdenia plants derived from seed sowing, have survived until present. Most of them have climbed up nets and ropes provided onto the stems of three Eucalyptus host plants and at least two of them have produced fruits. Marsdenia is likely to be an obligate outbreeder, in which case the genotypes introduced to the site will be vital in allowing future seed production there. Implications of the work for conservation and management are stated. Key Words: Drought, enrichment planting, kangaroo grazing, mallee vegetation, Marsdenia australis, Rhyncharrhena linearis, seedling mortality, woody climbers. Transactions of the Roval Society of S. Aust. (2004), 128(1), 61-66. ENRICHMENT-PLANTING OF THE WOODY CLIMBERS MARSDENIA AUSTRALIS AND RHYNCHARRHENA LINEARIS IN NORTH-WESTERN VICTORIA by R. F. PARSONS" Summary PARSONS, R. F. (2000). Enrichment-planting of the woody climbers Marsdenia australis and Rhyncharrhena linearis in north-western Victoria. Trans. R. Soc. S. Aust. 128(1 ), 61-66, 31 May, 2004. Marsdenia australis and Rhyncharrhena linearis are woody climbers (family Asclepiadaceae) found in semi- arid to arid areas of all mainland states of Australia. Where studied, they are declining drastically, probably due to grazing. In Victoria, their conservation status is vulnerable. In 1991, in Hattah-Kulkyne National Park, a site was found carrying single plants of each species. At this site, 384 seedlings and 1,741 seeds of Marsdenia and 240 seedlings of Rhvacharrhena were planted underneath trees of Eucalyptus socialis in 1991-1992, half of the seedlings in mammal-proof cages. Afier high seedling mortality due to mammal grazing and especially to drought in the first summer, only six of the planted seedlings, all of Marsdenia, survived by the end of 1992. All of these, and a further 15 Marsdenia plants derived from seed sowing, have survived until present. Most of them have climbed up nets and ropes provided onto the stems of three Eucalyptus host plants and at least two of them have produced fruits. Marsdenia is likely to be an obligate outbreeder, in which case the genotypes introduced to the site will be vital in allowing future seed production there. Implications of the work for conservation and management are stated. Kry Worps: Drought, enrichment-planting, kangaroo grazing, mallee vegetation, Marsdenia australis, Khyncharrhena linearis, seedling mortality, woody climbers. Introduction Two species of lianes (woody climbers) in the family Asclepiadaceae, Marsdenia australis and Rhyncharrhena linearis, occur in semi-arid and arid areas of all mainland states of Australia. In Victoria, both species are declining drastically, probably due to grazing. While small sucker shoots can be found when grazing pressure is low, no seedlings at all can be found, even in the absence of vertebrate grazing (Nichols, Browne & Parsons, 1991). The primary aim of the work was to investigate seed and seedling survival in the field to assist the conservation management of the species, both of which have a conservation status currently rated as vulnerable Victoria-wide (Victoria: Department of Natural Resources & Environment, unpublished). However, we hoped that any plants surviving from the experiments would be left to augment naturally- occurring populations. The work began as a B.Sc. (Honours) project from February to September 1991 (Smith, 1991). Methods Hattah-Kulkyne National Park was chosen for the work, being an extensive biological reserve carrying both species. At present, only 12 naturally-occurring Marsdenia plants are known there (Fiona Murdoch, " Botany Department, La Trobe University, Bundoora, VIC 3086, Australia. pers. comm., 20 December 2002). The work was carried out near one of these, which has a Rhyncharrhena plant nearby. The study site in Hattah-Kulkyne National Park is within the Mournpall Block, 0.6 km along Jasmine Track from its N end, then 0.2 km WSW of the Track (Fig. 1). The fenced Rhyncharrhena plot no. 9E is about 50 m away. The area is Eucalyptus socialis tall shrubland (‘mallee scrub’) on sandy loam topsoils. After various episodes of heavy grazing by stock and rabbits since 1847, stock grazing was stopped in 1974 (Parks Victoria records). Subsequent rabbit control meant that rabbit numbers were ‘very low’ during the first year of the project (D. Major personal communication). There was also grazing by a population of western grey kangaroos, estimated at 800 in the 5,700 ha Mournpall Block during 1991 (D. Major personal communication). All seed used was collected between 1988 and 1991 from the Red Cliffs area (50 km from the plots), Marsdenia from sites | and 3 and Rhyncharrhena from site | of Nichols, Browne & Parsons (1991), Germinability of seed lots was established in a growth cabinet using a 10 hour photoperiod at 25/15° C after plume removal. Seedlings to transplant into the field were produced by planting germinated seed into sandy loam, vermiculite and perlite (2:1:1) in pots in the same growth cabinet. At four weeks of age they were placed outside to harden and then planted at the field site. 62 R. F. PARSONS Hattah , Camping Ground HATTAH - KULKYNE NATIONAL x PARK + 34°50'S 142° 20E NS > eS . : vv, Stockyard Track HIGHWAY Fig. |. Location of study area in north-western Victoria. Study site shown as cross west of Jasmine Track. For both species, as fruiting seems only to occur in plants climbing in shrubs or trees (Nichols, Browne & Parsons, 1991), all seedlings were planted beneath low branches of mature Eucalyptus socialis trees. Four trees growing in an area of 50 m radius were used (Fig. 2) with four sites selected beneath each tree crown. For Marsdenia seedlings, at each of four planting dates from March to July 1991, one site under each tree was chosen at random and planted, providing four replicates. Each site consisted of a caged and an uncaged plot (for seedlings) side-by- side. Cages were 40 x 40 x 30 cm. Thus for Marsdenia the design was 4 trees x 4 planting times x 2 treatments (caged/uncaged) x 12 seedlings = a total of 384 seedlings planted. The Rhyncharrhena plantings were similar, but only 240 seedlings were available. Before planting, litter and understorey plants were removed and plots made level by rake. Seedlings were planted 10 cm apart, the species intermingled. The wire mesh cages were flanged at the base to deter rabbit entry via burrowing. The plots were watered to try to minimize seedling deaths from water deficiency. At each planting date, enough water to saturate the surface 5 cm of soil was added, both to the plots to be planted and to all plots planted previously. Water poured onto the sites was stopped from running off by infiltrometer rings put in place until infiltration was complete. At all planting and inspection dates, all herbs other than Marsdenia and Rhyncharrhena were identified, counted and removed from the plots by manual uprooting. The plots were monitored in September 1991, four times in 1992-4 and finally in May 2002 (Table 2). Eleven Marsdenia fruits containing a total of 1741 filled seeds were left over from the 1991 experiment. We removed their plumes and sowed them under a shallow soil covering in three of the cages on 2 March 1992 (rather than discard them). At the same time, surviving seedlings were marked with stakes to distinguish them from plants of seed origin. On 5 May 1994, nets or ropes made of natural fibres were positioned to provide all surviving plants with a pathway so that they could twine upwards into Eucalyptus crowns. This necessitated cutting the tops out of the cages. Results Survival of planted seedlings inside cages, after a winter of mild temperatures (coldest single night of 1° C) and before the severe water deficit of summer, was high (84 — 85%) for both species (Table 1). Judging from plant symptoms and climatic data, by far the main cause of mortality in the cages up until September 1991 was soil water deficit. Analysis of bite marks suggested that the much lower survival outside cages (49 — 52%) was due to either rabbit or kangaroo grazing. ENRICHMENT-PLANTING OF MARSDENIA 63 TABLE 1. Number of survivors on 3 September 1991 of seedlings planted from March to July 1991, Number Number Percentage planted surviving survival Marsdenia Caged 192 164 85 Uncaged 192 100 52 Rhyncharrhena Caged 96 81 84 Uncaged 144 71 49 TABLE 2. Height, leaf number and survival at five dates of 384 seedlings of Marsdenia australis planted from March to July, 1991. na = not available. Date Sep Mar Dee Dec May May 91 92 92 93 94 02 Age (yr) 3000 mg L"!), as these are common in the state (e.g. Williams 1967; EPA 1998; Hammer 2002a). Freshwater fishes in South Australia display a variety of physical forms and life histories. The dwarf galaxias Galaxiella pusilla is remarkable for its ability to survive dry periods in seasonal swamps, where it takes refuge in swamp-crayfish burrows (Geocharax: Beck 1985). Large species like the Murray-Darling golden perch Macquaria ambigua ambigua may cover long distances (for example, a tagged fish is known to have travelled 2300 km along the Murray and Darling rivers: Reynolds * Cooperative Research Centre for Freshwater Ecology, School of Earth and Environmental Sciences DP312, The University of Adelaide, Adelaide, SA 5005. Email: michael.hammer@adelaide.edu.au ' Evolutionary Biology Unit, South Australian Museum. ' HAMMER, M. (2001) Molecular systematics and conservation biology of the southern pygmy perch Nannoperca australis (Giinther, 1861) (Teleostei: Percichthyidae) in south-eastern Australia. Unpub. BSc(Hons) Thesis, Department of Environmental Biology, The University of Adelaide. 1983), whereas small species like the southern pygmy perch Nannoperca australis are much less vagile (Hammer'). Other species need to move between fresh water and marine habitats, although even diadromous species like the galaxiids Galaxias maculatus and G. brevipinnis may occur in ‘landlocked’ populations (Pierce ef al. 1985; Hammer 2002a; SKM 2002). In addition, there are euryhaline species like the small-mouthed hardyhead Atherinosoma microstoma, found in fresh or salt water (Molsher ef a/. 1994; Hammer 2002a). This catalogue lists 84 species in the freshwater fish fauna of South Australia. It updates earlier work (Waite 1923; Scott et al. 1974; Sim 2000), corrects and amends records of species and _ their distributions, and is designed to assist in research and planning for management and conservation. Methods Drainage divisions Five of the 13 principal drainage divisions in Australia (AWRC 1976) occur wholly or partly in South Australia, and provide a biogeographic framework (Fig. 1): * South East Coast (SEC), including the Millicent Coast and Glenelg River (part) river basins, * Murray Darling (MD), part of the Lower Murray River Basin, ¢ South Australian Gulf (SAG), the only division contained wholly within the state (the shared 86 M. PD. HAMMER & K. F. WALKER oe ees |< || om etres 0 LOO, 200 400 ~ AUSTRALIAN DRAINAGE DIVISIONS North-east Coast South-east Coast Tasmanian Murray-Darling South Australian Gulf South-west Coast Indian Ocean Timor Sea Gulf of Carpentaria Lake Eyre Bulloo-Bancannia Western Plateau Distant Islands (not shown) Fig. 1. Drainage divisions in Australia and South Australia (AWRC 1976). FRESHWATER FISH SPECIES RICHNESS IN SOUTH AUSTRALIA 87 boundary with MD is west of the Murray Mouth, but SAG includes the coastal streams of Fleurieu Peninsula), * Lake Eyre (LE), draining toward lakes Eyre and Frome, and * Western Plateau (WP), containing sparse coastal lakes and some ephemeral waters. Records of species were obtained from the literature and examination of specimens at the South Australian Museum, Adelaide (SAMA), including material from recent collections by the senior author, Information on rare or doubtful species was scrutinised with special care. Nomenclature The systematic framework and nomenclature employed here follow Eschmeyer (1998) and subsequent updates (see Californian Academy of Sciences on-line “Catalogue of Fishes”, March 2003"), except that the lamprey families Geotriidae and Mordaciidae replace Petromyzontidae (Strahan 1980), subspecific status is recognised for M. a. ambigua (after Musyl & Keenan 1992) and six informal taxa and a species complex are recognised. The informal taxa include dwarf flathead gudgeon Philypnodon sp. (Larson & Hoese 1996), Lake Eyre golden perch Macquaria sp. (Musyl & Keenan 1992) and western chanda perch, an undescribed species referred to in earlier literature as “Ambassis muelleri Klunzinger” (syn. A. agassizii), but lacking a formal name since “4. mueller” was invalidated by Allen et al, (2002). The carp gudgeon genus Hypseleotris awaits a formal review but, following Allen er al. (2002), this catalogue recognizes Midgley’s carp gudgeon H. “sp. 1” sensu Hoese et al. (1980) and Murray-Darling carp gudgeon H. “sp, 3” sensu Unmack (2000). In addition, a species complex of hybrids and possible semi-clonal hybridogenic forms are recognised (Bertozzi et a/. 2000), including Lake’s carp gudgeon 7. “sp. 2” sensu Hoese ef al. (1980). Following Allen and Jenkins (1999), prior records of northern purple-spotted gudgeon Mogurnda mogurnda (Richardson) in South Australia should be referred to Dalhousie purple- spotted gudgeon M. thermophila or Flinders Ranges purple-spotted gudgeon M. clivicola (these were described from within the range of M. mogurnda). Criteria for inclusion A “freshwater” species here includes obligate freshwater and diadromous species and _ select euryhaline taxa known to complete their lifecycle in fresh water. “Alien” species include exotic species (not native to Australia) and native Australian * http://www.calacademy.org/research/ichthyology/catalog/fishcatmain.asp species translocated outside their natural range. Alien fishes in natural waterways are regarded as established species if their populations are self- sustaining or if they are continually stocked, and as introduced species if records are few and isolated or confined to artificial waterbodies (and potentially could become established). The latter include interstate translocations within drainage divisions. Results Native fish richness A total of 58 native freshwater fish species in 15 families is recorded for South Australia (Table 1). All are shared with other states, except for seven endemics in isolated areas of LE. Mogurnda clivicola may be another endemic, as only small populations of uncertain affinity occur outside the state (Allen & Jenkins 1999; Wager & Unmack 2000). Forty-four native species are confined to fresh water. One of these, Australian smelt Retropinna semoni, may occasionally occur in the Coorong (Eckert & Robinson 1990), but is not strictly diadromous. Four euryhaline taxa meet the aforementioned criteria of “freshwater” species, namely A. microstoma, flathead gudgeon Philypnodon grandiceps, western bluespot goby Pseudogobius olorum and lagoon — goby Tasmanogobius lasti (e.g. Wedderburn & Hammer 2003). Thirteen of the 44 obligate freshwater species occur in more than one division, and none is common to all. Most obligate freshwater species occur in LE (24) and MD (24, plus 11 diadromous and euryhaline taxa). Diadromous and euryhaline species generally occur in more than one division. Remarkably, three diadromous species are recorded for WP, although data there are sparse (Table 1). New records for South Australia Fly-specked hardyhead Craterocephalus stercusmuscarum ?stercusmuscarum (Giinther) This taxon was identified in samples collected from the northern Flinders Ranges in 1994-95 (SAMA F7331, F9002, F9078). It is distinguished from the Lake Eyre hardyhead Craterocephalus eyresii (Steindachner), which occurs in the same region but not the same habitats, by fewer transverse scale rows (7-8 cf. 11-14 in C. eyresii) and dark lateral banding (Ivanstoff et a/. 1987; Crowley & Ivanstoff 1990a). Subspecific identification is tentative owing to taxonomic problems and_ the isolated nature of the population (the nearest known conspecifics are from Aramac Springs in the remote M. P.- HAMMER & K. F. WALKER xX xX yosad Awd elex (ZLQT “JoBuizunpy) Dinasqo p21adouunN’ xX xX xX yorod AwisAd wroynosg LOT JoyUNyH sip.ysnp po1adouunyN xX yosad uspjos a1hq ayeT i (paquosapun) ‘ds piwonbonpy xX xX yosod Arenysq (COQ ayUND) wnsoU0j]oOI bLuonbovpy q yorad otrenboeyy OE] olAng vorspjo.ysnp viuipnboopy x yoiad uspjos Surpreq-ABunyy (SPR ‘Uospreyory) oNsiquip pnsiquin vlApnbavyy x poo AvLiny (SEST ‘TeyoUN) aed wjaad vjjayoo]jno2vV a poo nor (6781 ‘AaIAND) sisuatuonbopu vjjayro]jnIavpy q x xX YSLOR[G JOATY BPg] ‘Uospieyory smpsowupul sisdopoyH oeprAtpyoio1od xX yoled epuryo UJa]sa\\, i (paqisosapun) “ds syssoqup q yosed epueyd LOT “Jeuyoepulars JZIsspsp sissoqup oepissequryy x peoyApsey poysods-A] 4 (L9g] “JeyUNyH) wiapssniusnoials;, UNADISnULsNIAaIS SNOYdaIO1IIVAD xX peoyApiey poyxsodsuy) L261 ‘UAILY 3 AopMoOID ‘Jyorsueay snapnf wniposnusnosays snjoydas0sayp.ay xX peoyApiey SJoao[H HO66I ‘ffoysueay 2 AopMOID Ladojs snppydas0dajvAy xX peoyApiey AeLinyy ZIOL ‘YoorNgopl solange snjpydasos1ajo.1D é xX 4 peoydpiey o1Aq oyeT H(EQQT “auyoepulars) usaia snypydasosa}o.1y Xx peoyApiey osisnoyeq HPL6| JOAO[D 2 Jforsueay sisuaisnoyppp snypydasosajo.1) xX xX xX xX peoyApsey poyyNoul-[[PUrs (1981 ayIUNH) VWoJso4so1W DULOSOULLAYIP aepluULioyly xX ysyMoqures 11989] (9681 ‘Z}91Z) 1210) DpIpuajds vluanjouvjapyy xX é ysyMoquies Aplin (QQ ‘neupased) syuolanpyf piuanjounjayy —- Awpltuaeyouryay x Yspnu ueluewisey, (PE6] “H09S) Iaapaj7 DUUBYI0aN xX SeIXE]es JVM (9€61 BW) MULISNd HIAIXD]OH x serxees payods Opg ‘SouUaloUg[eA sNaIDIINA] SVIXDIDD, q serxeyes ABLIN| ZLQ] ‘Aosulzunypy smjo.yso4 spixvjoH x Ni xX serxeyes uleyUNo| 99g] TouUND snp1jo spixnjpyH xX xX xX SeIxe]es UOWWWOD (Zpgl ‘suduosp) supojnapw spixojoyH 4 xX Serxe|es SUIQUIT[D 99g] “TouUNDH szuuidiaaig spixnjDy oeplixe[ey xX xX jjous uelensny (S681 JaqoM\) 1Mowas puudo.ay q SuljAvis uevlpensny POT JayUNH YUapidUl $A]I0.1J0JOLd oepruurdonoy ME é xX Suioy Auog (Q9Q| “ayUND) 1gasa VsSOpDDUIAN oeplodni) xX yslyqed JoyBMYSoL] BERT ‘TSU SnUpun] snunpuLy xX uepure} IOATIS (968 ‘ZIOIZ) Snajuasin snjiys010g xX ystpqeo s,AH LOT ‘Iouyoepulays Mpjudy snunjIsoaN’ xX ysyeo orsnoyyeqd HQ66[ “SIOQUIA] 3 UdT[Y lado]s sn.injisoaNy Xe ysyyeo 1adood 866] ‘BIOquIa 2 UdT[W sisuasadood saploanjisoaN depIsojo[d x x xX [99 PouULFOYS IPS] ‘Uospreyory sijp.usny pjnsup oeprypinsuy xX xX xX Aoiduuey poprsyoys (9pQ| ‘UOspIeYSIy) Xvpsoul DIgvpsopy oeplloepsloyyy x xX x Acide] poyonog ISR] ‘AoIn sippajsnv 11.1440aH) aeprinooy dM cel DVS d OAS UOISIAIC] oweu UOLUWOD uoxeL Ayrure J Jx9] 99S, “RITRNSNY YINOS 0} SIWpuU, ‘[snIEIs UreVaOUN = { ‘JouTyXe pouNsoid = q ‘pop10dei = X] “PI/P-usNy yinog fo suoisialp adpulp.sp ul SaYslf 1ajDMysad{ ALON “| AAV, 89 HNESS IN SOUTH AUSTRALIA FRESHWATER FISH SPECIES RIC £ v7 ol se 6l (8g [eI0], uray) sjezo], xX xX xX Agos u00seT 166] “8S80H USD) SnIgosoupUusYT xX xX xX x Aqo3 jodsan]q W1a}soA\, (O88I ‘aseanes) wn10jo snigosopnasg x Agos aisnoyjeq #661 WOsIe’y] 14aagjs snigosopAunjy) xX Aqos 1aseq H(Q68I ‘ZI9Z) sniuasa snigosopAuwjyD oepligoy é xX uosspns prouiey Jiemq (poquosapun) ‘ds uopoudajiyg xX xX xX uoespns peoyjeyy (POST “Wfery) sdaoppupas uopoudapiyg X uoaspns payods-ajdind arsnoyyeq 26661 ‘SUDUeL 2 uaTTy vjlydouiay) Dpuansopy xX uoaspns poyods-ojdind sasuey siopulpy 666] ‘SUbyUoL 3? Ud|[Y P/OI1A1]9 Dpu.nsopy q q uoaspns payods-aidind urayynog (SL8] ‘neujaise) vsiadspp ppuinsopy xX X (uoaspns dies s aye] “8'a) sutog pliqAp ,(xayduros sarsads) ‘dds sioajasday xX xX uoaspns dies Surpieq Aeunyy (poqiiosopun) ¢ ‘ds szuoajasday x xX uosspns dies s,Aa[spiyy (peqiiosapun) | “ds si.uoajasdapy xX x uoospns dies ulajsoyy (8681 ‘AQ[ISQ) Masuizuny siuoajasday aeplyoa[q X xX xX xX TJosuoD (TERI ‘SeuusTOUITeA) Mjjlau silaydopnasg oepnuydepnosg xX J9]UNIS COIR g (LIGI “UeEM WY YSOT[NDI) C0210g wNjL0Ig5 xX é x Jojunis poysueds (6S8I ‘JoyUND) “ojoo1un uodvsayjodoiaT X JOUNIS $,YoTo (LIGI “SUE WF YOT[NDIP|) 1y9/am snuvdpig xX yolad SATIS (SE8T “TTeyoUA) suupdApig snuvdpig 4 Jqunis papueg (P9OgT TeyWNDH) saplooiad pquvinluup oepnuodeiay xX yosed AwsdAd payeSarie,, 9861 “USTPV 2 JoUNy YINSaLIDA DoIadouuvN dM AT OVS GW OAS UOISIAIG, oueu UOUWOZ UOXeL Aqrure J 90 M. PD. HAMMER & K. F. WALKER upper reaches of Cooper Creek, Queensland). A molecular revision of Craterocephalus in progress indicates that sub-species within the C. stercusmuscarum species complex remain confused (P. Unmack, Arizona State University, pers. comm.) and further morphological and molecular analyses are required. Spotted galaxias Galaxias truttaceus Valenciennes This species was first reported in 1999 from karstic springs in coastal south-eastern South Australia (e.g. Ewens Ponds: Hammer ef a/. 2000; SAMA F9217, F10111) representing a minor westward range extension into South Australia. Another single specimen from the same area occurred among specimens of G. maculatus collected in 1979 (SAMA F10109). Note that a prior report of G. truttaceus from SAG (Scott ef al. 1974) was based on misidentified specimens (SAMA F3094, F3188). Tasmanian mudfish Neochanna cleaveri (Scott) This species is known in South Australia only from a single specimen collected from Bool Lagoon in 1974, and previously registered as G. maculatus (SAMA F4919). Recent surveys have failed to locate others (Hammer 2002a). The new record is noteworthy as the species is cryptic, with an ability to survive extended dry periods by burrowing into mud or hiding under rocks and wood, and otherwise is native to Tasmania and Victoria (Fulton 1986; Koehn & Raadik 1991). A possible new terapontid A form of grunter (Terapontidae) resembling a deep-bodied Welch’s grunter Bidvanus welchi or a hybrid B. welchi x Barcoo grunter Scortum barcoo is known from Coongie Lakes (J. Puckridge, University of Adelaide, pers. comm. 2001). This form is listed as the ‘Cooper grunter’ by Sim (2000). It was also reported near Goyder Lagoon on the lower Warburton River in 2002 (Costelloe ef al. 2003). Range extensions Surveys in the Mount Lofty Ranges (Hammer') have provided three new drainage division records, namely a genetically distinct sub-population of Nannoperca australis from the Inman River Catchment (SAG), Hypseleotris sp. 3 from the same location, and Yarra pygmy perch Nannoperca obscura from Lake Alexandrina (MD). The review uncovered other, previously misidentified specimens of N. obscura in the museum collection dating from 1915 (SAMA F572), suggesting the species is native. The presence of mountain galaxias Galaxias olidus (a species complex presently under systematic review: Raadik 2001) recently was confirmed from the South Australian section of SEC (Mosquito Creek: Hammer 2002a). Despite its inclusion in a south east regional list by Glover (1983), no specimens of the species were previously known. In addition, Glover mistakenly referred to the Mosquito Creek population as G. maculatus. The presence of G. brevipinnis in MD is also confirmed (SAMA F153: Angas River, 1914; previously registered as G. maculatus), a record predating the Snowy Mountains Hydroelectric Scheme which appears to be the source of G. brevipinnis in the upper Murray catchment (Waters et a/. 2002). A report of R. semoni from SAG (SKM 2002) is suspect because voucher specimens are not available and no other records exist for the division (e.g. McDowall 1979; Unmack 2001). Other SAG reports of bony herring Nematalosa erebi and spangled grunter Leiopotherapon unicolor in the Lake Torrens catchment, and western carp gudgeon Hypseleotris klunzingeri as native to the Broughton River (Pierce et al. 2001) are also discounted in the absence of voucher specimens or other data, There is an uncertain report of fish resembling C. eyresii in the remote, isolated Durkin Swamp (WP), following exceptional rainfall (Ehmann & Tynan 1997). Finke goby Chlamydogobius japalpa Larson, Finke hardyhead Craterocephalus centralis Crowley & Ivanstoff and Finke purple-spotted gudgeon Mogurnda larapintae (Zeitz) potentially could colonise the ephemeral, lower reaches of the Finke River in South Australia, following floods from the headwaters in the Northern Territory, but they have not been formally recorded. Alien species There are records of 26 alien species in South Australia (Tables 2-3), although two may prove to be natives (Philypnodon sp. from the Onkaparinga River (SAG) (SAMA F10087, April 2002), and Murray rainbowfish Melanotaenia fluviatilis from SEC (SAMA F2409, dated 1903)). Most alien species records are for SAG (20 species, including 13 established alien species). There are high numbers also for MD and SEC, but few in the remote LE and WP (Table 2). Fourteen alien species are established in South Australia. These include seven exotic taxa and seven translocated native taxa. Another 12 alien species have been introduced, but are not established or present only in artificial waterways (Tables 2-3). These include barramundi Lates calcarifer in the River Torrens and Australian bass Macquaria 91 S RICHNESS IN SOUTH AUSTRALIA “TE FRESHWATER FISH SPE! 0 4 €1 L Ss (FI [R30], puvsy) poysipqeyso [eI], z L 07 LI el (97 [BJO], puv.aD) s[ejo], iX uosspns prouleyy jrema (peqtiosopun) ‘ds uopoudayiyd I poo Adaajsg (L98] “I9UYIepUID}S) PIDJOaUI] s1.110a]aAXE Vv Vv uosspns poyods-ajdind wroyynog (SL8] ‘neujsiseD) vsuadspp ppuansopy Xx uoospns divs Surpieq AvLnyy (poquosopun) ¢ ‘ds szyoajasdayy xX uoospns divs s,Ao]3pryy (poquiosopun) | ‘ds szoajasdtyy oeplnoo[y Vv Vv Vv I yosod JOATIS (SET ‘TPSYUP) snupApig snuvdpig ovpyuodeiay, Vv yosod Awi3Ad wsloyynos 19g] “eyUNyH sijp.ysnp vo1adouuvpy I sseq uel[eysny (998] “louyoepulays) Yipaynopuaaou viapnbovpy Vv xX V I yosad uapjos Surpieq-AeLunyy (SP8I Wospreyory) pndiquip pnsiquin viapnbovpy I xX Vv I poo Avr (EST MOY) 1J2ad mjaad yyjayso]jnoovjy Vv ysipyoryq JOARY 8PZl ‘UOspreyorY sipsousmu sisdopoH seplAYIYOIOIIg Vv yoiad epueyo LOT ‘Jouyepulars 1/Zisspsp sissoqup oepissequry I Ipunure.ieg (O6LI ‘Yoolg) afiunajn2 saloT oepruodo.nuad xX él ysymoquies AvLINY (SLI ‘neUpaseD) supp biuavjouvnjayy deplusejouRpoyy él SPIXERS JVM (9€61 “ORW) MyIsnd pyjaixnjvyH oeplixepey x Vv T Ysyyeo JoyeMYSoLJ (REST TOYS) snuppuny snuppuny depIsoj}o[d SHIDddS AAILVN NVITVULSOAV GALVOOTSNVUL I x xX Xx yoied uvadoing SSL] ‘snovuury syjpoianyf vuWag deplolog I xX x x x vIsnquiet) 6S8I “PARAL 1yOo!G]OY VISnquuvL oepitytoa0d I Noy Yoolg (FIST “HYyoUAD) sypunuos snurjaajog xX xX I jnoy UMOIg SSL] ‘snevuury py oupps I uowryes onuepy SSL] ‘snevuUry “pjps owns 4 xX I no Moquiey (Z6LI “wineg[e) sszydu snysudy.s02ucC aepruowyes I YOROPIOYIIM [RIUILIO (TPT IoweD) smopnvoyyinsuv snuinssipy oepnigod x x x youe (SSLT ‘snevuurq) youn pouty Vv xX x I dies uowod SSL] ‘snovuury ordups snuidta Vv xX xX xX xX YsyployH (SSLI ‘snoeuurT) smypunp snisspavy oeprunidda SHIOddS OILOXA dM cel OVS CW OS UOISIAIG oulvu UOUTUIO?) uOXe], Aqrure J 1X9] 096, ‘[Sme}s UTe}I0uN = { *(spuod JustINRAD OBeMOS “SLURP UULIey “D°d) SpEIIgRY [eloyNIe 0} poonponul = V ‘Sp1ooa1 Mo} “poonpo.Ul = | *poysl[qeiso Jo/pue psonponur Ajpenuyuos = x] ‘oyp.ysnp yinog fo suols}alp a8pulpip ul sjuauuo.saua saivM ysalf ul soyslf ual]P “7 AIGVL, M. P. HAMMER & K. F. 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UI UIseg “Y SuIpIe wo poxoysuely ¢ pure JIZISsNSsD sisspqup (0007) 7? 12 WIS (6661) TOLOLA VWVS ‘CY sesuy °3°9) pops0da1 A[[eUOIseddQ “UOUILUOD 990UQ e DOUI) DIULT addy, a01N0S s]lejoq p1092y soroadg M. P- HAMMER & K. F. WALKER SAMA F6199 (1986); Reid & Puckridge (1990) SAMA F7331, F9002, F9078 (1994/95) Glover (1985); Hammer pers. obs. 2002 Glover (1980); Pierce ef al. 2001 SAMA F5496 (1981), F7405 (1982) SAMA 2615 (1947), F4789 (1984) SAMA F1388 (1929) Wager & Unmack (2000) SAMA F10056 (1947) Pierce (1990) Glover (1979) Source Leigh Creek retention dam. Poisoning attempted, but still present in 1999 Pierce ef al. (2001) (Hammer pers. obs.) Cooper Creek near Innamincka. Population small, may not be viable. Stocked into Clayton Bore. Introduced to Moro George, Flinders Ranges. Now probably absent. WESTERN PROVINCE DRAINAGE DIVISION Several regional records (e.g. spring at L. Hamilton; L. Newland). Streaky Bay (not strictly freshwater habitat but included), Specimens from the North Flinders Ranges previously identified as most westerly record. C. eyresii. Taxonomic status requires further investigation. First record Neales R. (1984). Common there in 2002. Coongie Lakes/ Cooper Creek. Dams, reservoirs at Woomera (with Gambusia holbrooki). Davenport Creek near Ceduna and Laura Bay. Spring at L. Hamilton (extant?). 1 2 3 4 4 4 2 2 3 4 Record Details Type LAKE EYRE DRAINAGE DIVISION Macquaria ambigua ambigua and 4 Bidyanus bidyanus Craterocephalus stercusmuscarum Perca fluviatilis ?stercusmuscarum Maccullochella peelii peelii Atherinosoma microstoma Pseudaphritis urvillii Gambusia holbrooki Pseudogobius olorum Carassius auratus Amniataba percoides Carassius auratus Cyprinus carpio Species novemaculeata, sleepy cod Oxyeleotris lineolata and Atlantic salmon Sa/mo salar in the River Murray. Gambusia Gambusia holbrooki and goldfish Carassius auratus were recorded in all drainage divisions. Four large native MD species (silver perch Bidvanus bidyanus, Murray cod Maccullochella peelii peelii, freshwater catfish Tandanus tandanus, M. a. ambigua) are spawned in commercial hatcheries in other states and are commonly introduced to South Australia (Tables 2-3), including undocumented stockings in farm dams in MD and SAG. Translocations in drainage divisions within South Australia are not considered in detail here, but have reportedly included transportation of M. clivicola in the Flinders Ranges region and fish from Cooper Creek to a retention dam at Leigh Creek (see Pierce et al. 2001). Extirpations and species decline Museum records are not necessarily a_ true indication of range and abundance, but indications from all sources combined are that there have been significant declines in the range of several species. Records for some species may represent occasional stray individuals on the fringe of their geographic range, but these could not be distinguished from established species due to a paucity of detailed historic surveys and/or temporal replication. There is historical evidence (Table 3) that Murray galaxias Galaxias —_rostratus, trout — cod Maccullochella macquariensis and Macquarie perch Macquaria australasica formerly occurred in MD in South Australia. Ambassis agassizii was last recorded from the Marne River mouth (MD) in 1983 (Lloyd & Walker 1986), and state-wide extirpation appears confirmed for the southern purple-spotted gudgeon Mogurnda adspersa (last record in MD 1973: SAMA F3727; no sightings in SAG for >50 years). The river blackfish Gadopsis marmoratus can be considered extirpated from SAG (it may persist on Kangaroo Island, but the record is dubious: Table 3) and has undergone significant range contraction in MD (Sim ef al. 2000), exacerbated since 1997 by the loss to irrigation diversions of more than half of the spring-fed habitats in the Marne River, one of few remaining refuges (Hammer 2002b). Similarly, range contraction and on-going local extirpations have been recorded for N. australis (Hammer'). Estuary perch Macquaria colonorum was once more widespread in the lower Murray prior to the construction of barrages near to the Murray Mouth (Sim ef al. 2000). For SEC, N. cleaveri and the Australian grayling Prototroctes maraena have not been reported since 1974 and 1982 respectively and FRESHWATER FISH SPECIES RICHNESS IN SOUTH AUSTRALIA 95 other SEC species including G. pusilla have likely suffered large range reductions coinciding with massive loss of wetland habitat (Hammer 2002a). Other species are confined to small areas, including five endemic species in Dalhousie Springs (LE) (Wager & Unmack 2000), M. clivicola (recorded only from Balcanoona Creek in the Flinders Ranges (LE): e.g. SAMA ~ F3042), Murray — hardyhead Craterocephalus fluviatilis (very few sites in the lower Murray (MD): Lloyd & Walker 1986; Wedderburn & Hammer 2003); N. obscura (three habitat fragments in SEC and a small section of MD: Hammer 2002a; Wedderburn & Hammer 2003) and the variegated pygmy perch Nannoperca variegata (a 4-kim? spring-fed area in SEC: Hammer ef a/. 2000). Discussion This catalogue is a contribution toward an inventory of state and regional biodiversity. Well- maintained historic collections and voucher specimens are critical to record information, validate doubtful records and sustain progress in taxonomy, ecology and conservation. Ideally, this information should be updated frequently, as work progresses. Although surface waters in South Australia are limited (NLWRA 2001), the state harbours about one fifth of the continental freshwater fish fauna. As the state borders intersect, rather than enclose, some drainage divisions, and as most divisions allow access to the sea, the number of endemic species is comparatively low. Some ‘new’ records here arise from minor re-alignments of physiographic boundaries between drainage divisions or states (e.g. South Australian Gulf Drainage Division: N. australis; South East Coast Drainage Division: G. truttaceus), but others represent significant range extensions (e.g. Murray Darling Drainage Division: N. obscura; South East Coast Drainage Division: N. cleaveri; Lake Eyre Drainage Division: C. s, ?stercusmuscarum). Biodiversity assessments and monitoring should favour obligate freshwater fishes isolated within particular drainage divisions or regions, because they are most likely to have diverged (cf. Crowley & Ivanstoff 1990a,b; Musyl & Keenan 1992; Larson 1995; Allen & Jenkins 1996; Allen & Feinberg 1998; Hammer'). These studies may gain impetus from assessments of ecosystem ‘health’, as fishes are * LLoyp, L. N. (1987) Ecology and distribution of the small native fish of the lower River Murray, South Australia, and their interactions with the exotic mosquitofish, Gambusia affinis holbrooki. Unpub. MSc Thesis, Department of Zoology, The University of Adelaide. potential indicators (e.g. Harris 1995). Clarifications are needed in regard to the taxonomy of undescribed taxa, species complexes and the biogeographic status of some species, especially where there are few historical data. Fine-scale molecular markers may help to distinguish natural and translocated populations (e.g. Waters ef al. 2002). Alien freshwater fishes are ubiquitous in South Australia. They are most apparent in areas directly affected by human industry, particularly in the Murray Darling and South Australian Gulf drainage divisions. All such species are potential vectors for pathogens and parasites (e.g. Langdon & Humphrey 1987). Predators like brown trout Salmo trutta, rainbow trout Oncorhynchus mykiss and European perch Perca fluviatilis are implicated in the decline of small native fishes (e.g. Crowl ef al. 1992; Morgan ef al. 2002). Gambusia holbrooki is an aggressive, highly fecund competitor that undoubtedly has affected native species (e.g. Lloyd’). The feeding behaviour and high abundance of common carp Cyprinus carpio have contributed to destruction of wetlands associated with the River Murray (e.g. Sim ef al. 2000), and thereby affected native fishes. There is also some risk of genetic contamination of native stocks by translocated native species (Arthington 1991). The preservation of native biota is a management priority in South Australia (e.g. Kahrimanis ef al. 2001; EPA 2003), and avenues for the introduction of non-native fishes such as the government-sanctioned releases of salmonids, sales of fingerling angling species to the public, “conservation” stocking, releases of unwanted aquarium fishes and inter-basin transfers from the River Murray all need review within broadly-based programs of flow and habitat protection, particularly where small isolated populations of native fish occur, Acknowledgements We gratefully acknowledge access to the collections of the South Australian Museum, financial support to MPH from the Nature Foundation (SA) and technical help from S. Wedderburn. T. Raadik, Arthur Rylah Institute, Melbourne, kindly identified several galaxiids and assisted with information on G. rostratus and T. Trnski confirmed the identity of hardyheads at the Australian Museum, Sydney. Our thanks also to members of Native Fish Australia (SA) for field assistance and to A. George for advice on GIS data. Comments on a draft manuscript by two referees were greatly appreciated. 96 M. P. HAMMER & K. F. WALKER References ALLEN, G. R. & FEINBERG, M. N. (1998) Descriptions of a new genus and four new species of freshwater catfishes (Plotosidae) from Australia. Aqua Ichthyology and Aquatic Biology 3, 9-18. _ & Jenkins, A. P. (1999) A review of the Australian freshwater gudgeons, genus Mogurnda (Eleotridae) with description of three new species. Aqua — Journal of Ichthyology and Aquatic Biology 3, 141-156. 7 _, Mipairy, S. H. & ALLEN, M. (2002) “Field Guide to the Freshwater Fishes of Australia” (Western Australian Museum, Perth). ANON, (1996) Thorndon Park Reservoir bouncing back. Southern Fisheries 4, 4. ArtHINGTON, A. H. (1991) Ecological and genetic impacts of introduced and translocated freshwater fishes in Australia. 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Native Fish Australia (SA) Inc., Adelaide. 38 p. WILLIAMS, W. D. (1967) The chemical characteristics of lentic surface waters in Australia pp 18-77 /7 Weatherley, A. H. (ed.) “Australian Inland Waters and Their Fauna” (Australian National University Press, Canberra). Zietz, A. H. (1902) List of the edible fish of the lower Murray, 7rans. R. Soc. S. Aust. 26, 265-267. THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON, TJIRTUDESSUS WATTS & HUMPHREYS AND NIRRIPIRTI WATTS AND HUMPHREYS, FROM UNDERGROUND WATERS IN AUSTRALIA By C. H. 8S. WATTS* & W. F. HUMPHREYST Summary Watts, C. H. S. & Humphreys, W. F. 2004. Thirteen new Dytiscidae (Coleoptera) of the genera Boongurrus Larson Tjirtudessus Watts & Humphreys and Nirripirti Watts & Humphreys, from underground waters in Australia. Trans. R. Soc. S. Aust. 128(2), 99-129, 30 November, 2004. Thirteen new species of stygobitic Dytiscidae from inland Western Australia are described: Tjirtudessus hillviewensis sp. nov., T. microocula sp. nov., T. occidentalis sp. nov., T. padburyensis sp. nov., T. wogarthaensis sp. nov., Nirripirti arachnoides sp. nov. N. bulbus sp. nov., N. byroensis sp. nov., N. copidotibiae sp. nov., N. dingbatensis sp. nov. N. eurypleuron sp. nov. N. innouendyensis sp. nov., and N. verrucosus sp. nov. This brings the total of stygobitic Dytiscidae decribed from Australia to 55, derived from 33 discrete groundwater calcretes in 8 palaeodrainages. Key Words: Coleoptera, Dytiscidae, Stygobitic, Descriptions, New Species, Water chemistry. Transactions of the Royal Society of S. Aust. (2004), 128(2), 99-129. THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON, TJIRTUDESSUS WATTS & HUMPHREYS AND NIRRIPIRTI WATTS AND HUMPHREYS, FROM UNDERGROUND WATERS IN AUSTRALIA. by C. H. S. WatTs* & W. F. HUMPHREYS* Summary Warts, C. H. S. & Humpnreys, W. F. 2004. Thirteen new Dytiscidae (Coleoptera) of the genera Boongurrus Larson Tjirtudessus Watts & Humphreys and Nirripirti Watts & Humphreys, from underground waters in Australia. Trans. R. Soc. S. Aust. 128(2), 99-129, 30 November, 2004. Thirteen new species of stygobitic Dytiscidae from inland Western Australia are described: Tjirtudessus hillviewensis sp. nov., T: microocula sp. nov., T. occidentalis sp. nov., T. padburyensis sp. nov., T. wogarthaensis sp. nov., Nirripirti arachnoides sp. nov., N. bulbus sp. nov., N. byroensis sp. nov., N. copidotibiae sp. nov., N. dingbatensis sp. nov. N. eurypleuron sp. nov., N. innouendyensis sp. nov., and N. verrucosus sp. nov. This brings the total of stygobitic Dytiscidae described from Australia to 55, derived from 33 discrete groundwater calcretes in 8 palacodrainages. One species, 7. microocula sp. nov., has partial eyes and wings that are only slightly reduced. Boongurrus occidentalis sp. nov. was collected from both surface and underground water and is little modified for an underground existence. Geographically the new species extend the range of stygobitic Dytiscidae in Australia to the Murchison and Moore drainage systems both of which drain to the Indian Ocean. Chemical and physical characters of the groundwater in some of the calcretes in which the new species were found are given. Key Worbs: Coleoptera. Dytiscidae. Stygobitic. Descriptions. New species. Water chemistry. Introduction This is the sixth paper in our series describing the stygobitic Dytiscidae of Australia (Watts and Humphreys 1999, 2000, 2001, 2003; Balke e¢ al. 2004). In it we describe the new species found during fieldwork in Western Australia in winter 2002. Four new species in the Bidessine genus Tjirtudessus Watts & Humphreys, and 8 in the Hydroporine genus Nirripirti Watts and Humphreys, are described from the westerly-draining Murchison and Moore paeleodrainage systems. A new species of the genus Boongurrus Larson is described from material collected mainly from interstial sand/gravels in seasonal creeks in the Pilbara but also from three bore holes accessing two different underground calcretes. This new Boongurrus species is fully sighted and winged and shows little apparent physical adaptation to a hyporean existence yet was found together with a true stygobitic fauna in deep calcrete. For the first time in Australia two stygobitic species (only one described here due to lack of a male specimen in one species) were discovered with a true eye remnant, as distinct from the usual small sclerite or short suture line, and wings that, although rather small, still retain veins and folded tips “C. H. S. Warts, South Australian Museum, North Terrace, Adelaide, South Australia 5000. * W. F. Humpnreys, Western Australian Museum, Francis Street, Perth, Western Australia 6000. suggesting that they are at an earlier stage of adaptation to underground life than the other species so far discovered. Geographically the new finds extend the known distribution of stygal Dytiscidae some 300 kilometres to the west. We also recollected sites in the Northern Territory that yielded stygal Dytiscidae in 2001 without discovering any additional species. In addition, areas of groundwater calcretes near The Granites, Tennant Creek and in the Amadeus basin in the Northern Territory were sampled extensively without finding any Dytiscidae and only a sparse stygobitic fauna (Syncarids, Copepods, Amphipods) at the occasional site. Although too early to be sure the results suggest that the Australian stygobitic dytiscid fauna is restricted to inland Western Australian and the Ngalia basin in central Australia. The prime aim of this series of papers is to formally describe the dytiscid fauna and to provide a preliminary indication of the ground water characteristics in which they are found. Companion papers by and with co- workers are starting to address questions of phylogeny and evolution (Balke et al. 2003, Cooper et al. 2002, Leys et al. 2003) and the taxonomic composition of the rich fauna associated with the beetles (Taiti and Humphreys 2001, Karanovic and Marmonier 2002, Karanovic 2004). Materials and Methods The collection methods and measurements of 100 C.H.S. WATTS & W. F. HUMPHREYS physico-chemical parameters in the water largely follow those used previously (Watts and Humphreys 2000) except that the use of a Quanta-G (Hydrolab Corporation, Austin, Texas) water quality monitoring system attached to a 50 m cable permitted the measurement of various physico-chemical water quality parameters (temperature, specific conductance (or TDS), pH, dissolved oxygen (% saturation or mg L-'), oxidation-reduction potential (redox), and depth, the latter facilitating the determination of any vertical stratification present in the water column in some boreholes. The instrument was calibrated against the standards recommended for the instrument. Abbreviations used: BES Prefix for field numbers, WAM Biospeleology. SAMA South Australian Museum, Adelaide. WAM __ Western Australian Museum, Perth. MB Groundwater monitoring bore. Systematics Key to Australian species of stygobitic Dytiscidae 1 — Scutellum well developed; length 4.5 mm Copelatus abditus Balke et al. — Scutellum absent; length 1.0 to 4.9 mm..2 2(1) — Paramere one-segmented; metatibia approximately the same — width throughout; without pronotal plicae (Hydroporint) .......cseecesesseeeseeteeteeeenees 34 — Paramere two-segmented; metatibia narrow at base then strongly expanding towards apex; usually with pronotal plicae (Bidessini) «0.0... 3 Ba(Z)v "With: YES. ei, esceectenareeeeeprzenperisisneaseonsd + — Without eyes, may have a_ small chitinized plate or suture line where eyes Hortinally Ave ccs seass cet svececeegeeeeter sessed 5 4 (3) — Eyes of normal size; with elytral plicac .. pubs egies Boongurrus occidentalis sp. nov. — Eyes approximately one-fifth normal size; without elytral plicae........... ee Hants Tjirtudessus microocula sp. nov. 5 (4) — Body length approximately 1.0 mm; legs stout, without swimming-hairs on fore and midlegs ...........05 Kintingka kurutjutu Watts and Humphreys — Body length > 1.2 mm; legs normal, all with swimming-haifS ...........:00eceeeeeee 6 6 (5) — Mesofemur with spines on hind edge approximately the same strength as those on mesotrochanter; length > 3.0 mm...29 — Mesofemur with spines on hind edge much more robust than those on 7(6) — 11(9) — nny — 13 (8) — 14 (13) — 15 (14) — mesotrochanter; length 1.4 — 3.6 mm....7 Normal ventrites 1 and 2 without suture between them (ie. number of visible abdominal segments reduced to four) (Fig. 82); length 3.2 — 3.6 mm............0+ Sebwatboptanaces Tjirtudessus sweetwatersensis Watts and Humphreys Ventrites 1 and 2 with suture between them, at least in inner portion (Figs 83- 86); length 1.3 — 3.2 MM... eee 8 Pronotal plicae strong, well marked, excavated On INSIdC.........cceeeeeeeeeeees 9 Pronotal plicae weak, difficult to trace, may be absent, not excavated on inside . sia doneppfthepaen emencdey-=ocapenqer ania tore eaStoaT Me eter ke 13 Mesosternum with posterior portion triangular in midline (Figs 77, 79)....... 10 Mesosternum with posterior portion rounded in midline (Fig. 78)............+ 11 Prosternal process rounded at tip (Fig. 75); tip of metatrochanter pointed; lobe on apical segment of paramere short....... wineaap Phas Tjirtudessus morgani (Watts and Humphreys) Tip of prosternal process pointed (Fig. 76); apex of metatrochanter rounded; lobe on apical portion of paramere long.. angered Tjirtudessus bialveus Watts and Humphreys Head broad, deflexed, metatrochanter round; setae on mesofemur long.............- .Tjirtudessus silus Watts and Humphreys With none of above characters ............ 12 Combined length of first two segments of metatarsus > rest; eye remnant present as small oval or triangular structure; paramere with long apical lobe............... sia vie kag Mls saa ea bgeated he Bey Tjirtudessus pulpa (Watts and Humphreys) Combined length of first two segments of metatarsus approximately equal to rest; eye remnant reduced to single short suture; paramere with small apical lobe . Ho baie Settee Tjirtudessus cunyuensis Watts and Humphreys Elytron with row of large punctures adjacent to SULUTC 1.0... eee ete eteteees 26 Elytron without sutural punctures, other than a few weak ones near base .......... 14 Eye remnant present as a small oval or triangular Structure .........c cece 23 Eye remnant reduced to single short SUTLE ater teense aed dal Mt beeberedenedeeneeste i) Mesofemur with 5 to 7 spines on hind edge in basal half... cece 16 Mesofemur with 2 to 4 spines on hind edge in basal half ...........cceeeeeeteeeees 19 THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 16 (15) — 17 (16) — fea ee 19 (15) — 20 (19) — 21 (20)— 22 (21)— 23 (14)— Protibia thick (Fig. 73); protarsus moderately expanded, mesotarsus less so; mesotibia slightly angular......Bidessodes gutteridgei Watts and Humphreys Protibia thin (Fig. 74); protarsus and mesotarsus approximately the same size; mesotibia not angular... eee 17 Length 2.5 — 2.7 mm; suture between ventrites | and 2 complete (Fig. 83)........ susan hs Tjirtudessus padburvensis sp. nov. Length 1.6 — 2.0 mm; suture between ventrites | and 2 obliterated laterally (BISS:84-86). 2. cscarteevterieelaaeticcceieicte teed 18 Paramere with lobe as wide as rest of apical segment, flat on top, expanded slightly at tip .....7jirtudessus masonensis Watts and Humphreys Paramere with lobe shorter than rest of apical segment, rounded on top, tip pointed.......... Tjirtudessus yuinmeryensis Watts and Humphreys Mesofemur with four spines near base; antenna with segments | and 3 of similar length, segment 11 approximately 1.5x length of segment 10; length 2.1 — 2.4 160100 Eee ore eo Tjirtudessus cueensis (Watts and Humphreys) Mesofemur with two to three strong spines on hind edge near base; antenna with segment 2 large, oval; segment 3 much smaller and thinner, segment 11 approaching 2x length of segment 10; lengthrl3. 2 PSS es h.c.rPeves. ese sche 20 Mesofemur with two strong spines on hind edge near base; paramere with apical segment with two finger-like projections ein Tjirtudessus pinnaclesensis (Watts and Humphreys) Mesofemur with three strong spines on hind edge near base; paramere with apical segment with one finger-like projection.. SRAM URS fA sae Se, icc Rc 21 Mesofemur with three spines grouped together near base... Tjirtudessus Jridaywellensis (Watts and Humphreys) Mesofemur with two spines near base and One More distant... eects 22 Pro and mesotibia club-shaped; antenna with middle segments enlarged a little on INSIdE eee Tjirtudessus hinkleri (Watts and Humphreys) Pro and mesotibia elongate/triangular in shape; middle segments of antenna virtually symmetrical............ Tjirtudessus karalundiensis Watts and Humphreys Pronotum not constricted at base: prosternal process reaching or almost 24 (23) — Das Douay. 27 (26)— I06)—— 29 (6) — 30 (29) — 31 (30) — reaching mesosternum; 1.4 mm long....... dee calls Peas aes Tjirtudessus wilunaensis Watts and Humphreys Pronotum moderately constricted at base; prosternal process not reaching mesosternum; 2.3 — 3.2 mm long........ 24 Mesofemur with 6 spines close to base on hind edge.......... Tjirtudessus bigbellensis (Watts and Humphreys) Mesofemur with 3 to 6 spines spread out along basal half of hind edge............... 29 Suture line between ventrites | and 2 well marked (Fig. 83); aedeagus with medial lobe parallel sided, apex not upturned..... bee eweecasMenetearctee 5 Tjirtudessus challaensis (Watts and Humphreys) Suture lines between ventrites | and 2 weak (Fig. 84), usually obsolete in lateral half; aedeagus with medial lobe distinctly narrower in middle, apex upturned pies te teiat Ie 8 Tjirtudessus jundeeensis Watts and Humphreys Distinct oval eye remnant present ....... 27 Eye remnant reduced to single short SUC OM la aad meat ca. ce teeh tet oa Ateas ei retnal 28 Metatrochanter with tip slightly pointed (Fig. 11); sutural lines between ventrites 1 and 2 complete, distinct ....7jirtudessus hillviewensis sp. nov. Metatrochanter with tip rounded; suture between ventrites 1 and 2 obliterated laterally......... Tjirtudessus windarraensis (Watts and Humphreys) Mesofemur with 3 spines on hind edge (Fig. 28); suture between ventrites | and 2 complete; metatrochanter elongate (Fig. 29)...Tjirtudessus wogarthaensis sp. nov. Mesofemur with 5 to 6 spines on hind edge; suture between ventrites 1 and 2 obliterated laterally; metatrochanter rounded ............ Tjirtudessus lapostaae (Watts and Humphreys) Mesofemur with spines arranged in two comb-like rows along hind edge from base to apex; mesotibia thin, curved ....... De ee ee ere Bidessodes limestoneensis Watts and Humphreys Mesofemur with spines on hind edge spaced out, not dense and comb-like; mesotibia straight Pro and mesotarsus with basal segment much more expanded than other SCOMOMS techs. coer wetescecisy teh Mesenetton 3] Pro and mesotarsus with basal segment only moderately expanded compared with other segments Antenna with segments 8 to 11 noticeably 102 32 (30) — 33 (32) — 34 (2) — 35 (34) — 36 (35) — 37 (36) — 38 (37) — 39 (34) — C.H.S. WATTS & W. F. HUMPHREYS thinner than others, segment 3 longer than segment 2...Tjirtudessus magnificus Watts and Humphreys Antenna with segments 8 to 10 not noticeably thinner than others, segment 3 same length as segment 2.....7jirtudessus macrotarsus. Watts and Humphreys Pronotum a little narrower than elytra; length 3.5 — 5.0MmM.........cccseseereeeeeeees 33 Pronotum wider than elytra; length 3.2 — SST ssseecss seston Tjirtudessus eberhardi Watts and Humphreys Metatrochanter rounded at tip; aedeagus with central lobe straight, tip pointed; eye remnant small ..7jirtudessus raesideensis Watts and Humphreys Metatrochanter pointed at tip; aedeagus with central lobe twisted, tip knobbed; without eye remnant...7jirtudessus hahni Watts and Humphreys From the Northern Territory ... ‘ From Western Australia ...........0c 39 Head short, very broad, strongly deflexed; pronotum strongly narrowed at base; prosternal process anvil-shaped...... sdeaggiteartnendsteecesy Nirripirti macrocephalus Watts and Humphreys Head variably shaped, not deflexed, base of pronotum variably shaped; prosternal process “normally” shaped ...........++ 36 Protarsus with segment 3 not bilobed; pronotum not constricted at base; antenna thin, segments | and 2 subequal.............. FRR onseseysaned usncondecty Nirripirti pentameres Watts and Humphreys Protarsus with segment 3 bilobed; pronotum weakly to moderately constricted at base; antenna thick, segment 2 much broader than segment | fs To yng os tag dhagd Tas T EA FEE RaSaalca nosh nad nese agRRE? 37 Length 1.8 mm; body well-chitinized ..... suse bates civeee ty sep yes Nirripirti napperbyensis Watts and Humphreys Length 1.2 — 1.6 mm; body weakly CHILMIZEM «...ervsrerapsateaaderradacnssdcdernedicebiand 38 Length 1.2 mm; body only slightly constricted at junction of pronotum and SL yIPA NY, Aseetyyenovehaest Nirripirti wedgeensis Watts and Humphreys Length 1.5 mm; body quite strongly constricted at junction of pronotum and elytra wee Nirripirti newhavenensis Watts and Humphreys Pronotum (and head) about half width of elytra (Fig. 36) .....Nirripirti arachnoides sp. Nov. Pronotum > three quarters width of elytra 40 (39) — 41 (40) — 42 (41) — 43 (42) — 44 (43) — 45 (40) — 46 (45) — 47 (46) — 48 (47) — Elytron with visible ventral portion extensive except close to apex (eg. Fig. BS) its Aap eee egptterca ye lpanaed 41 Elytron with visible ventral portion narrow except in basal quarter (eg. Fig. BG) resscypedennteeditccatecaetussaenees Length 3.6 — 3.8 mm Nirripirti stegastos Watts and Humphreys Length 1.5 — 2.5 MM....... eee 42 Antenna with segments 6 to 8 greatly expanded, much broader than segments 9 and 10 (Fig. 42) hesberseeyitgnge theguszte Nirripirti bulbus sp. nov. Antenna with segments 6 to 10 of approximately equal SiZe ..........:ee 43 Meso and metatibia elongate triangular; body strongly boat-shaped, pronotum much narrower in front «2.0... 44 Meso and metatibia cylindrical (Figs 61, 62); front and rear of pronotum same width (Fig. 63) .....eccceeseseeeeees Nirripirti eurypleuron sp. nov. Length 2.1 to 2.3 mm, metatrochanter with tip sharply pointed ............ Nirripirti skaphites Watts and Humphreys sp. nov. Length 1.5 to 1.9 mm; metatrochanter with tip rounded ....Nirripirti killaraensis Watts and Humphreys Antenna with segment 2 larger and more oval than segment 1; 1.2 — 2.1 mm long. deseontuasnyatbernadnebeesiedabeoddeveanganuresieettesdey etd 52 Antenna with segment 2 more or less the same shape as segment | or smaller; 2.5 — BO mtn OME ete seessreadss de speasdonaterends 46 Mesofemur with row of about 20 closely placed small spines along hind edge (Fig. AB): seaeerdeosstete Nirripirti byroensis sp. nov. Mesofemur with 10 or fewer weak to very strong spines along hind edge .....47 Metasternal plate parallel sided (Fig. 80); mesofemur with 8 to 10 spines, closely placed, very strong; metatrochanter long and thin about 4 x as long as wide.......... Aiyssgigshdugeciccbes sbeupaeny rs Nirripirti fortisspina Watts and Humphreys Metasternal plate narrowing towards rear (eg. Fig. 81); mesofemur with 4 to 8 spines, weak to moderately strong; metatrochanter moderately elongate 2 to 2.5 x as long as wide Metatarsus with segment | as long as others combined, with confluent group of 5 strong spines in middle on outside (Fig. Si) ecsaetdene Nirripirti copidotibiae sp. nov. Metatarsus with segment | much shorter than others combined, without confluent THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 103 group of spines on outside... 49 49 (48) — Metasternal plate without wings (Fig. 81) eptqbiid tua peels Midgenad Nirripirti plutonicensis Watts and Humphreys — Metasternal wings obvious but short (eg. Eee OW) atc itscy eau dia dullest eit aad maida ng 50 50 (49) — Metafemur with 2 to 4 small lumps on top edge (Fig. 71)........ Nirripirti verrucosus sp. Nov. — Metafemur with smooth top edge........ 5] 51 (50) — Mesofemur with moderately strong spines; metacoxal plate nearly reaching MESOCOXAC. 0. eee eeeee Nirripirti hinzeae Watts and Humphreys — Mesofemur with thin spines; metacoxal plate at least the width of metafemur from MESOCOXAE.... eee Nirripirti darlotensis Watts and Humphreys 52 (45) — Elytron with shoulder flared outwards CPig. 86). «2. ciealaetereed Nirripirti hamoni Watts and Humphreys — Elytron with shoulder not flared (eg. Fig. OO) stasis wechecemntant lis upivalptensidinaaaeds 53 53 (52) — Metatrochanter produced into long strong Point. (Figs 68) eveseeseeeeeeder verses Nirripirti innouendyensis sp. nov. — Metatrochanter rounded, at most bluntly POSH seers ig eh IEA io ia cas Shae 54 54 (53) — Eye remnant absent; metatrochanter large, squat; hind leg stout; length 1.2 TMM, 0... eeeeeeeeeeeeeeees Nirripirti milgunensis Watts and Humphreys — Eye remnant represented by a short suture at side of head; metatrochanter elongate; hind leg elongate; length 1.8 - 2.0 mm.... a cMoshters concrete ners? Nirripirti melroseensis Watts and Humphreys Descriptions The following species descriptions are grouped in alphabetical order under genus which are placed in the order Boongurrus, Tjirtudessus, Nirripirti. Boongurrus Larson (Hydroporinae, Bidessini) Boongurrus occidentalis sp. nov. FIGS 13-18 Holotype m: ‘10 K NW Eerala Stn CHS Watts 23/5/01’, WAM 34222. Card mounted. Paratypes 25; 8, as for holotype, SAMA; 12, ‘Wittenoom Gorge Town Pool CHS & GA Watts 26/5/01’, 10 SAMA, 2, Lars Hendrich collection (Berlin); 1, ‘BES 9277 Killara Station north, unused water bore, 26° 03’ 55” S 118° 41’ 58” E, 6/6/2002 W. F. Humphreys and R Leys’, SAMA; 1, ‘BES 9318 Moorarie Stn nr calcrete quarry site 419, 25° 52’ 26S 117° 27’ O9E, 8/6/02, W. F. Humphreys & R. Leys’, SAMA; 2, ditto except ‘BES 9320’, | WAM 34177, | SAMA; 1, ‘BES 9246 Wagga Wagga Stn, mineral exploration bore 28° 26' 36” S 116° 38’ 9" E, 4/6/02, W. F. Humphreys & R. Leys’, SAMA. Description (number examined, 24) Habitus. Length 1.9 — 2.3 mm, relatively flat, weakly constricted at junction of pronotum/elytra; elongate oval; uniformly light testaceous, elytra uniformly darker; hindwing not reduced; eyes of normal size. Head. Narrower than elytra; smooth, shiny, weak reticulation towards rear, punctures small, sparse; without cervical stria, subparallel in posterior half, widest just behind eye, antenna stout, segments! and 2 cylindrical, segment 3 as long as segment 2, narrower, narrowing slightly towards base, segments 4 to 8 becoming progressively slightly broader, segments 9 and 10 a little narrower than segment 8, segment 11 about twice length of segment 10, each segment, except segment 1, with some very small setae on inside apically. Maxillary palpus elongate, segment 4 as long as segments | to 3 combined. Pronotum. A little narrower than elytra; anteriolateral angles projecting strongly forward; base weakly constricted, posteriolateral angles square, surface slightly rugose, with moderately dense, moderately sized punctures and a row of stronger punctures along front margin; basal plicae strong, excavated somewhat on inside, converging slightly towards front, reaching to about two-thirds way along pronotum. Elytra. Not fused, with weak inner ridges near apex (ligula), lacking in some; elongate, nearly parallel-sided in middle; rugose, quite densely covered with moderate sized punctures; plicae well impressed, straight, about as long as pronotal plicae. Epipleuron well differentiated from rest of elytra particularly anteriorly, lacking basal carina, relatively broad in anterior quarter then progressively narrowing to near apex. Ventral surface. Prosternal process strongly narrowed between coxae, reaching mesothorax, apical half elongate triangular, weakly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae separated. Metathorax strongly triangular in front in midline; wings short; broadly rounded in midline behind. Metacoxal plates large, quite strongly punctate, moderately rugose, metacoxal lines distinct, moderately widely spaced, reaching to metasternum, weakly diverging; closely adpressed to ventrite 1. Ventrites | and 2 fused, sutural lines distinct, ventrites 3 to 5 mobile, rugose, 104 C. H. S. WATTS & W. F. HUMPHREYS well covered with moderate sized seta-bearing punctures. Legs. Protibia triangular, outer edge bow-shaped, widest towards apex where it is about four times its basal width; protarsus weakly expanded, segment | as broad as long, segment 2 as wide as segment | and about half its length, segment 3 as long as segment 1| and a little narrower, deeply bifid, segment 4 very small, hidden within lobes of segment 3, segment 5 narrow, cylindrical, about 1.5x length of segment 3, segments | to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate-oval with a few relatively long, thin setae on inner edge; mesofemur with 4 to 5 moderately strong setae in basal half, only slightly stronger than those on other parts of the femur (Fig. 16), mesotarsus slightly less expanded than protarsus. Metatrochanter tip rounded (Fig. 17); metafemur relatively stout, lacking spines; metatibia narrow, moderately curved, widening towards apex; metatarsi relatively stout, segment | longest, segment 5 longer than segment 4, segments | and 2 in combination about as long as others; claws weak. Male Little external differentiation between the sexes. Median lobe of aedeagus broad in middle narrowing to sharp point; paramere broad, apical segment with relatively stout apical lobe well separated from rest of segment. Figs 13-14. Remarks This new species of Boongurrus closely resembles B. rivulus (Larson) from the Atherton tableland region of north Queensland differing externally only in the larger eyes, slightly less rugose body, slightly weaker tarsi and thinner antennae. The main differentiating characters are the size of the eyes which in the new species are of normal epigean size in contrast to those of B. rivulus which are about three quarters normal size and in the male genitalia. The aedeagus of B. occidentalis is broader medially and tapers to a longer and sharper point. The apical segment of the paramere is longer than in B. rivulus and the apical lobe smaller but much more distinct (Larson 1994), Figs 1-6. Tjirtudessus hillviewensis: |, lateral view of central lobe of aedeagus; 2, ditto dorsal view; 3, paramere; 4, mesotrochanter and mesofemur; 5 metatrochanter and metafemur; 6, dorsal view. Scale bar represents 1mm (habitus only). THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 105 Most of the specimens have been collected at the edge of pools in sandy riverbeds and interstitially to at least two meters from the water’s edge in an upstream direction. It has also been recorded as flying to light (in Australian Insect Collection CSIRO Canberra). A little surprisingly a few specimens were collected during this year’s fieldwork from boreholes in two separate calcretes together with a true stygobitic fauna. The Killara North site was an open unused water bore ca 3 m to water and a water depth of ca 9 m in a calcrete area and the associated stygofauna included Bathynellacea, crangonytoid and Ceinidae Amphipoda, Harpacticoida, Cyclopoida. The Moorarie site was a sealed monitoring bore in calcrete within 30 m of a calcrete quarry that penetrated the groundwater; the borehole contained a stygobitic fauna comprising Bathynellacea and Amphipoda. DNA studies (Remko Leys pers. com.) found no differences between B. occidentalis specimens collected from epigean and_ stygal habitats. Etymology Latin. ‘Occidental’ — western. A reference to its distribution. Tjirtudessus Watts & Humphreys (Hydroporinae, Bidessini) Tjirtudessus hillviewensis sp. nov. FIGS 1-6 Holotype m. BES 9399, Hillview Station, bore at Camel Well, 26° 58’ 20S 117° 27’ 09" E, 13/6/02, W. F. Humphreys & R. Leys’, WAM 34178. Slide mounted. Paratype I (partial), as for holotype except ‘BES 9398’, SAMA. Description (number examined, | + | partial) Habitus. Length 2.35 mm; relatively flat, weakly constricted at junction of pronotum/elytra; elongate oval; uniformly light testaceous; hindwing reduced, broad, about half length of elytron, tip slightly folded; weakly sclerotized. Head. A little narrower than elytra; smooth, reticulation weak, punctures sparse, very small; subparallel in posterior half, widest just behind eye remnant, eye remnant reduced to small triangular area. Antenna moderately stout, segments | and 2 broad, oval, segment 3 as long as segment 2, narrower, narrowing towards base, segments 4 to 10 subequal, segment 11 about twice length of segment 10, each segment, except segment 1, with some very small setae on inside apically. Maxillary palpus, elongate, segment 4 as long as segments | to 3 combined. Pronotum. About same width as elytra; anteriolateral angles projecting strongly forward; base weakly constricted, posteriolateral angles obtuse, overlying elytra somewhat; smooth, with sparse, very weak punctures and a row of stronger punctures along front margin; basal plicae moderately impressed, converging slightly towards front, reaching to about half way along pronotum; numerous long setae laterally in anterior half. Elytra. Not fused, tightly closed, lacking inner ridges; elongate, almost parallel sided, smooth, very weakly reticulate, sparsely covered with small punctures, a few widely spaced larger punctures close to inner edge; row of long setae near lateral edge, a few additional larger punctures with long setae, more frequent towards sides and apex. Epipleuron moderately differentiated from rest of elytra, moderately broad in anterior quarter, progressively thinner till near apex. Ventral surface. Prosternal process strongly narrowed between coxae, not reaching mesothorax, apical half broad, sides slowly converging to rounded apex, strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact at midline. Metathorax broadly triangular in front in midline; wings stout, narrow; narrowly rounded in midline behind. Metacoxal plates large, weakly reticulate, metacoxal lines indistinct, moderately widely spaced, reaching to about halfway to metasternum, quite strongly diverging; a few small setae-bearing punctures towards midline; closely adpressed to ventrite 1. Ventrites 1 and 2 fused, sutural lines distinct, ventrites 3 to 5 mobile, sparsely covered with small seta-bearing punctures, ventrites 3 and 4 with a long central seta or bunch of long setae. Legs. Protibia bow-shaped, relatively narrow, widest towards apex where it is about four times its basal width; protarsus weakly expanded, segment | as broad as long, segment 2 as wide as segment | and about halfits length, segment 3 as long as segment 1, narrower, deeply bifid, segment 4 very small, hidden within lobes of segment 3, segment 5 narrow, cylindrical, about 1.5x length of segment 3, segments | to 3 with very dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate/oval with a few thin setae on inner edge; mesofemur with 4 evenly spaced spines in basal half (Fig. 4) mesotarsus a little more elongate than protarsus. Metatrochanter tip angular (Fig. 5); metafemur relatively stout, lacking spines; metatibia narrow, moderately curved, widening towards apex; metatarsus elongate, segment | longest, segment 5 a little longer than segment 4, segments | and 2 in combination about as long as others; claws weak. 106 C.H. S. WATTS & W. F. HUMPHREYS Figs 7-12. Tjirtudessus microocula: 7, lateral view of central lobe of aedeagus; 8, ditto dorsal view; 9, paramere; 10, mesotrochanter and mesofemur; 11, metatrochanter and metafemur; 12, dorsal view. Scale bar represents 1mm (habitus only). Male Female not known. Median lobe of aedeagus with unusually long thin apical portion, slightly crenulated on top; paramere broad, apical segment with long, narrow, apical lobe separated from rest of segment except at its apex which overlaps rest of segment. Figs 1-2. Etymology Named after the pastoral station on which it was found. Remarks Most closely resembles 7) windarraensis in size, presence of an eye remnant, row of large punctures on elytron adjacent to the suture and pointed median lobe of the aedeagus, but differs in the shorter apical segment of the paramere and larger apical lobe, as well as having a more oval eye remnant, and a complete suture between the first and second ventrites which is obliterated laterally in 7: windarraensis. Tjirtudessus microocula sp. nov. FIGS 7-12 Holotype m. ‘BES 9223, Bunnawarra Station, bore nr. shearing shed, 28° 36’ 35” S 11° 34’ 25” E, 3/6/02, W. F. Humphreys & R. Leys’, WAM 34179. Slide mounted. Paratypes 37: 19 + 5 partial specimens, as for holotype, 8 + 5 partial specimens WAM 34180 - 34192, 11 SAMA; 4, as for holotype except ‘BES 9224, well near shearers quarters, 28° 36’ 36” S 116° 34’ 18” EB’, WAM 34193 - 34196; 4, as for holotype except ‘BES 9225’, SAMA. Description (number examined, 38) Habitus. Length 2.2 — 2.3 mm; relatively flat, weakly constricted at junction of pronotum/elytra; elongate oval; uniformly light testaceous; eyes reduced to about a fifth normal size; hindwing reduced, about length of elytron, tip folded. THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 107 17 Figs 13-18. Boongurrus occidentalis: 13, lateral view of central lobe of aedeagus; 14, ditto dorsal view; 15, paramere; 16, mesotrochanter and mesofemur; 17 metatrochanter and metafemur; 18, dorsal view. Scale bar represents Imm (habitus only). Head. Narrower than elytra; smooth, reticulation weak, punctures sparse, small; subparallel in posterior half, widest just behind eye; eye reduced to about a fifth normal size, lacking individual facets, darkly pigmented. Antenna relatively thin, segments 1 and 2 cylindrical, segment 3 as long as segment 2, narrower, narrowing towards base, segments 4 to 10 subequal, segment 11 elongate, a little less than twice length of segment 10, each segment, except segment 1, with some very small setae on inside apically. Maxillary palpus, elongate, segment 4 as long as segments | to 3 combined. Pronotum. A little narrower than elytra; anteriolateral angles projecting strongly forward; base moderately constricted, posteriolateral angles bluntly pointed, smooth, with sparse, very weak punctures and a few stronger punctures along front margin; basal plicae weak, strongly slanting inwards, reaching to about half way along pronotum; with row of long setae laterally in anterior half. Elytra. Not fused but tightly closed, lacking inner ridges; widest behind middle, smooth, very weakly reticulate, sparsely covered with very small punctures, row of long setae near lateral edge, a few additional larger punctures with long setae, more frequent towards sides. Epipleuron well differentiated from rest of elytra, moderately wide in anterior fifth, virtually absent along rest of elytron. Ventral surface. Prosternal process strongly narrowed between coxae, not reaching mesothorax, apical half elongate triangular, sides subparallel, rapidly narrowing to small blunt point posteriorly, strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact at midline. Metathorax triangular in front in midline; wings narrow; broadly rounded in midline behind. Metacoxal plates large, very weakly reticulate, metacoxal lines obsolete; a few small setae-bearing punctures towards midline; closely adpressed to ventrite |. Ventrites | and 2 fused, sutural lines distinct except in lateral fifth, ventrites 3 to 5 mobile, sparsely covered with small seta- bearing punctures, ventrites 3 and 4 with a long central seta or bunch of long setae. Legs. Protibia triangular, relatively narrow, widest 108 C.H.S. WATTS & W. F. HUMPHREYS Figs 19-24. Tjirtudessus padburyensis: 19, lateral view of central lobe of aedeagus; 20, ditto dorsal view; 21, paramere; 22, mesotrochanter and mesofemur; 23, metatrochanter and metafemur; 24, dorsal view. Scale bar represents 1mm (habitus only). near apex where it is about three times its basal width; protarsus expanded, segment | as broad as long, segment 2 as wide as segment | and about half its length, segment 3 as long as segment | much narrower, deeply bifid, segment 4 very small, hidden within lobes of segment 3, segment 5 narrow, cylindrical, about 1.5x length of segment 3, segments | to 3 with dense covering of adhesive setae; claws moderately long, — simple. Mesotrochanter elongate/oval with a few thin setae on inner edge; mesofemur with 5 to 6 spines in basal half, basal two close together (Fig.10); mesotarsus similar to protarsus. Metatrochanter weakly pointed (Fig.11); metafemur relatively thin, lacking spines; metatibia narrow, curved, widening towards apex; metatarsus elongate, segment | longest, segment 5 longer than segment 4, segments | and 2 in combination about as long as others; claws weak. Male Little external difference between sexes. Median lobe of aedeagus variable in width along shaft, tip bluntly pointed; paramere broad, apical segment with long, narrow, apical lobe well separated from rest of segment. Figs 7-8. Etymology Latin. ‘Oculus’- eye, ’micro’- small. A reference to the small eyes in this species. Remarks Tjirtudessus microocula appears to be in an earlier stage of adaptation to an underground environment than the other stygal Dytiscidae previously recorded in Australia (with the exception of Boongurrus occidentalis, which is certainly not an obligate stygobiotic). The eyes are only about a fifth the size of those in epigean members of the genus and do not seem to be organised into individual ommatidia. It is hard to envisage them as fully functional. As well as the presence of partial eyes 7) microocula has wings which are still large enough to require folding and still retain veins, the prosternal process is not deflexed and reaches the metathorax hence separating the mesocoxae as in epigean species. THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 109 Two other specimens, both female and because of that not described, of a much smaller species were collected from the same well. This species also has small remnant eyes, moderately developed wings and a prosternal process that reaches the metathorax. Biochemically they are sister species (Remko Leys pers com.). Tjirtudessus padburyensis sp. nov. FIGS 19-24 Holotype m. “BES 9329, Mt Padbury Station, Irrigation Well, site 412, 25° 41’ 54” S 118° 05’ 29” E, 8/6/02, W. F. Humphreys & R. Leys’, WAM 34197. Slide mounted. Paratypes 5; 1, as for holotype, SAMA; 4, as for holotype except ‘BES 9330’, 2 WAM 34198 - 34199, 2 SAMA. Description (number examined, 6) Habitus. Length 2.5 — 2.7 mm; relatively flat, moderately constricted at junction of pronotum/elytra; elongate oval; uniformly light testaceous; hindwing reduced, about length of elytron. Head. A little narrower than elytra; smooth, reticulation weak, punctures sparse, very small; subparallel in posterior half, widest just behind eye remnant; eye remnant reduced to small semicircular area. Antenna moderately stout, segments | and 2 cylindrical, segment 3 as long as segment 2, narrower, narrowing towards base, segments 4 to 10 approximately equal in shape, segment 11 similar to segment 10, each segment, except segment 1, with some very small setae on inside apically. Maxillary palpus, elongate, segment 4 as long as segments | to 3 combined. Pronotum. Same width as elytra; anteriolateral angles projecting strongly forward; base strongly constricted, posteriolateral angles bluntly pointed, overlying elytra somewhat; smooth, reticulation weak, punctures very weak, sparse, a row of stronger punctures along front margin; basal plicae moderate, straight, slightly excavated inwards, reaching to about half way along pronotum; with row of long setae laterally in anterior half. Elvira. Not fused, lacking inner ridges; elongate, widest behind middle, smooth, very weakly reticulate, sparsely covered with very small punctures, a few widely spaced larger punctures close to inner edge in apical third; a few additional larger punctures with long setae, more frequent towards sides. Epipleuron not differentiated from rest of elytron, that portion of elytron visible ventrally, relatively broad in anterior third, thin along rest of elytron. Ventral surface. Prosternal process strongly narrowed between coxae, not reaching mesothorax, sides subparallel, tip bluntly pointed, strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact at midline. Metathorax triangular in front in midline; wings narrow; rounded or slightly triangular in midline behind. Metacoxal plates large, shiny, virtually nonreticulate, metacoxal lines obsolete; closely adpressed to ventrite 1. Ventrites | and 2 fused, sutural lines distinct, ventrites 3 to 5 mobile, sparsely covered with small seta-bearing punctures, ventrites 3 and 4 with a long central seta or bunch of long setae. Legs. Protibia triangular, relatively narrow, widest near apex where it is about four times its basal width; protarsus expanded, segment | as broad as long, segment 2 as wide as segment | and about a third its length, segment 3 shorter than segment | much narrower, about as wide, bifid, segment 4 very small, hidden within lobes of segment 3, segment 5 narrow, cylindrical, about 1.5x length of segment 3, segments | to 3 with very dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate/rectangular with a few thin setae on inner edge; mesofemur with 6 spines in basal half (Fig. 22); mesotarsus similar to protarsus. Metatrochanter tip rounded (Fig. 23); metafemur relatively thin, lacking spines; metatibia narrow, moderately curved, widening towards apex; metatarsus elongate, segment | longest, segment 5 longer than segment 4, segments | and 2 in combination about as long as others; claws weak. Male Little external differences between the sexes. Median lobe of aedeagus relatively broad, gradually narrowing towards tip, tip rounded; paramere broad, apical segment with short apical lobe overlapping rest of segment. Figs 19-21. Etymology Named after the pastoral station on which it was found. Remarks A moderately sized very ‘average’ species with broad tarsi and a complete suture line between ventrites | and 2. The unusually small apical lobe to the paramere is shared only with 7) morgani an otherwise very different species. Tjirtudessus wogarthaensis sp. nov. FIGS 25-30 110 C.H.S. WATTS & W. F. HUMPHREYS () \) ‘4 : \) J 4 ' y ( * } 5 i 0 {/ Oo Ss x YX () \) "7, E SS 1% we So B 74] AS SS” aR, 25 6H mS x Sie LEE DP 29 30 Figs 25-30. Tjirtudessus wogarthaensis: 25, lateral view of central lobe of aedeagus; 26, ditto dorsal view; 27, paramere; 28, mesotrochanter and mesofemur; 29 metatrochanter and metafemur; 30, dorsal view. Scale bar represents Imm ZA (habitus only). Holotype m. ‘BES 9384, Moorarie Station, Wogartha Well, 25° 57' 58” S 117° 35’ 28” E, 12/6/02, W. F. Humphreys & R. Leys’, WAM 34200. Slide mounted. Paratype 1. as for holotype, SAMA. Description (number examined, 2) Habitus. Length 1.4 — 1.5 mm; weakly chitinized; relatively flat, moderately constricted at junction of pronotum/elytra; elongate oval; uniformly light testaceous; hindwing vestigial, about one eighth length of elytron. Head. Considerably narrower than elytra; smooth, reticulation strong, punctures sparse, very small; subparallel in posterior half, widest just behind eye remnant; eye remnant reduced to single small suture. Antenna stout, segment 1 cylindrical, segment 2 broader, oval, segment 3 shorter than segment 2, much narrower, narrowing towards base, segments 4 to 8 becoming progressively slightly broader, segments 9 and 10 a little narrower than segment 8, segment 11 about twice length of segment 10, each segment, except segment 1, with some very small setae on inside apically. Maxillary palpus, elongate, segment 4 as long as segments | to 3 combined. Pronotum. A _ little narrower than — elytra; anteriolateral angles projecting strongly forward; base weakly constricted, posteriolateral angles bluntly pointed, overlying elytra somewhat; smooth, reticulation strong, punctures sparse, very weak, a row of stronger punctures along front margin; basal plicae absent; with row of long setae laterally in anterior half. Elytra. Not fused, tightly closed, lacking inner ridges; elongate, widest in middle, smooth, strongly reticulate, sparsely covered with very small punctures, row of widely spaced larger punctures close to inner edge; row of long setae near lateral edge, a few additional larger punctures with long setae, more frequent towards sides. Epipleuron indistinctly differentiated from rest of elytra, that portion of elytron visible ventrally relatively narrow, relatively even width until near apex. THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 111 Figs 31-36. Nirridessus arachnoides: 31, lateral view of central lobe of aedeagus; 32, ditto dorsal view; 33, paramere; 34, mesotrochanter and mesofemur; 35 metatrochanter and metafemur; 36, dorsal view. Scale bar represents 1mm (habitus only). Ventral surface. Prosternal process strongly narrowed between coxae, not reaching mesothorax, apical half elongate triangular, strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact at midline. Metathorax triangular in front in midline; wings narrow; narrowly rounded in midline behind. Metacoxal plates large, strongly reticulate, metacoxal lines indistinct, widely spaced, reaching to about halfway to metasternum, not diverging; a few small setae- bearing punctures towards midline; closely adpressed to ventrite 1. Ventrites | and 2 fused, sutural lines distinct, ventrites 3 to 5 mobile, sparsely covered with small seta-bearing punctures, ventrites 3 and 4 with a long central seta or bunch of long setae. Legs. Protibia triangular, moderately broad, widest near apex where it is about five times its basal width; protarsus expanded, segment | as broad as long, segment 2 as wide as segment | and about half its length, segment 3 as long as segment 1, narrower, deeply bifid, segment 4 very small, hidden within lobes of segment 3, segment 5 narrow, cylindrical, about 1.5x length of segment 3, segments | to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate/oval with a few thin setae on inner edge; mesofemur with 3 spines in basal half, basal two close together (Fig. 28); mesotarsus more elongate than protarsus, individual segments about half as broad. Metatrochanter tip bluntly pointed (Fig. 29); metafemur relatively stout, lacking spines; metatibia narrow, weakly curved, widening towards apex; metatarsus elongate, segment | longest, segment 5 longer than segment 4, segments | and 2 in combination about as long as others; claws weak. Male Sexes externally similar. Median lobe of aedeagus slightly variable in width along shaft, narrowing to tip which is rounded and slightly twisted; paramere broad, apical segment with relatively short, narrow, apical lobe well separated from rest of segment. Figs 25-27. 112 C. H.S. WATTS & W. F. HUMPHREYS 41 ; ; < y * ’, YQ Jb x Sy A & i, MERE: SUE goss = = ST . Za. a | 5 IV \\ AN = My; / INN Ve \ Y Y' : i NN 42 | \ a Ye y 2 Figs 37-42. Nirripirti bulbus: 37, lateral view of central lobe of aedeagus; 38, ditto dorsal view; 39, paramere; 40, mesotrochanter and mesofemur; 41 metatrochanter and metafemur; 42, dorsal view. Scale bar represents !mm (habitus only). Etymology Named after the pastoral well in which it was found. Remarks A small species recognised by the three spines on the mesofemur, lack of pronotal plicae and with sutural punctures and pointed metatrochanters. The hind wings are the most reduced — to tiny flaps — yet seen in Tjirtudessus. Nirripirti Watts and Humphreys (Hydroporinae: Hydroporini). Nirripirti arachnoides sp. nov. FIGS 31-36 Holotype m. ‘BES 9367, Byro Station, Yalcallia Well, 25° 54’ 39 " § 115° 53’ 03 " E, 10/6/02, W. F. Humphreys & R. Leys’, WAM 34201. Slide mounted. Paratypes 21; 11, as for holotype, 5 WAM 34202 - 34206, 6 SAMA; 10 as for holotype except ‘BES 9368’, 4 WAM 34207 - 34210, 6 SAMA. Description (number examined, 22) Habitus. Length 2.2 — 2.3 mm; head and pronotum small compared with almost globular elytra, relatively flat, moderately constricted at junction of pronotum/elytra; uniformly light testaceous; hindwing vestigial, reduced to tiny flap. Head. Small, about a third width of elytra; smooth, moderately strong reticulation with small even meshes, a few scattered small punctures; sides parallel, weakly indented at eye remnant; eye remnant reduced to short suture. Antenna relatively thin, segments | and 2 cylindrical, segments 3 to 10 of roughly similar shape but segments 5 to 7 somewhat longer than others, segment 11 thin a little longer than segment 10, each segment with some very small setae on inside apically. Maxillary palpus elongate, segment 4 1.3x as long as segment 3. Pronotum. Much narrower than — elytra; THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 13 anteriolateral angles projecting strongly forward; sides sinuate, converging towards rear, posteriolateral angles obtuse; a few scattered minute punctures; long setae laterally, denser towards front; moderately strong reticulation. Elytra. Not fused, tightly closed, lacking inner ridges; widely oval, widest near shoulders, smooth; covered with fine reticulation; a few scattered small punctures; a few additional larger punctures with long setae, more frequent towards apex, near scutellum and sides. Epipleuron well marked, broad in anterior third, then gradually narrowing to near apex. Ventral surface. Prosternal process strongly narrowed between coxae, not reaching mesothorax, apical half spatulate, strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact in midline. Metathorax very sharply triangularly projecting forward in midline; wings short, very narrow; widely rounded in midline behind. Metacoxal plates large, metacoxal lines absent; virtually impunctate; closely adpressed to ventrite 1. Ventrites 1 and 2 fused, sutural lines distinct in inner two-thirds absent laterally, ventrites 3 to 5 mobile, virtually impunctate except for a few long central setae or bunch of long setae Legs. Protibia long, narrow, widest past middle where it is about 1.5 x its basal width; protarsus small, weakly expanded, segment I broadly triangular, segment 2 about one half length of segment 1, segment 3 longer than segment 1, very deeply bifid, segment 4 very small and hidden within lobes of segment 3, segment 5 cylindrical, about as long as segment 3, segments | to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate with a few fine setae at apex; mesofemur with row of 5 long spines along hind edge in basal half; mesotibia thin, slightly angular, front edge uneven with long stout setae; mesotarsus more elongate than protarsus (Fig. 34). Metatrochanter roughly oval; metafemur clongate, lacking spines (Fig. 35); metatibia straight, approximately the same width throughout; metatarsus elongate, segment | longest, segment 4 shortest; in combination segments | and 2 about same length as others, segments 2 to 5 without spines other than at apex; claws weak. Male Little external difference between sexes. Central lobe of aedeagus relatively broad, widening slightly towards front. Parameres of average width, tips with two long setae. Figs 31-33. Etymology Greek. ‘Arachne’ — spider, An allusion to its spider-like shape. Remarks A very distinctive species readily recognised by its pronounced spider-like shape - small head and pronotum together with large broad body and long thin legs - as well as the five long spines on the mesofemur, angular very spinose mesotibia, unusually thin tarsi, impunctate ventrites, wide epipleura and flanged elytra. Nirripirti bulbus sp. nov. FIGS 37-42. Holotype m. “BES 9324, Moorarie Station, bore nr.calcrete quarry, site 419, 25° 52’ 26” S 117° 27’ 09” E., 8/6/02, W. F. Humphreys & R. Leys’, WAM 34211. Slide mounted. Paratypes 11; 3, as for holotype, SAMA; 8 as for holotype except “BES 9325’, 5 SAMA, 3 WAM 34212 — 34214. Description (number examined, 12) Habitus. Length 2.1 — 2.5 mm; narrowly boat- shaped, relatively flat, slightly depressed in sutural region, base of pronotum not constricted; uniformly light testaceous; rather weakly sclerotized; hindwing vestigial, reduced to tiny flap. Head. Relatively small, much narrower than elytra; smooth, moderately strong reticulation with small even meshes, a few scattered small punctures; sides parallel; eye remnant reduced to short suture line. Antenna relatively thick, segments | and 2 cylindrical, about same length, segment 3 about same length but half width of segment 2, segment 4 same shape but a little shorter than segment 3, apex of segment 6 moderately expanded on inside, segments 7 and 8 expanded, almost globular, segments 9 and 10 narrower, segment I] about 1.3x as long as and thinner than segment 10, each segment with some very small setae on inside apically. Maxillary palpus elongate, segment 4 a little longer than segment 3. Pronotum. Narrower than elytra; anteriolateral angles projecting forward; sides almost parallel, posteriolateral angles right angles; a few scattered minute punctures and a few larger ones along front edge; moderately strongly reticulate. Elytra. Not fused, tightly locked, lacking inner ridges: elongate, parallel sided, smooth; covered with relatively strong, regular reticulation; a few scattered small punctures; a few additional larger punctures with long setae, more frequent towards apex and sides. Epipleuron not differentiated, that portion of elytron visible ventrally broad, of even width along most of elytron except close to apex. 114 C.H.S. WATTS & W. F. HUMPHREYS a Ooo We MI Ax Po, VW a \ oa ESS fj ay S 2, Figs 43-45. Nirripirti byroensis: 43, mesotrochanter and mesofemur; 44 metatrochanter and metafemur; 45, dorsal view. Scale bar represents 1mm (habitus only). Ventral surface. Prosternal process strongly narrowed between coxae, not reaching mesothorax, apical half spatulate, strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact in midline. Metathorax weakly projecting forward in midline; wings very narrow; widely triangular behind, apex blunt. Metacoxal plates large; metacoxal lines absent; virtually impunctate, evenly covered with moderately strong reticulation; closely adpressed to ventrite |. Ventrites 1 and 2 fused, sutural lines distinct in inner half, absent laterally, ventrites 3 to 5 mobile, virtually impunctate except for a few long central setae or bunch of long setae. Legs. Protibia narrow, widest past middle where it is about twice its basal width; protarsus weakly expanded, segment | broadly triangular, segment 2 about one half length of segment 1, segment 3 as long as segment 1, deeply bifid, segment 4 very small and hidden within lobes of segment 3, segment 5 stout, cylindrical, longer than segment 3, segments 1 to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate with a few fine setae at apex; mesofemur with row of 4 to 5 spines along hind edge in basal half, basal two close together, edge slightly indented between spines (Fig. 40); mesotarsus more elongate than protarsus. Metatrochanter relatively large, apex weakly pointed (Fig. 41); metafemur elongate, lacking spines; metatibia curved, approximately the same width throughout; metatarsus elongate, segment | longest, segment 4 shortest; in combination segments | and 2 shorter than others, segments 2 to 5 without spines other than at apex; claws weak Male Antenna with segments 6 to 8 more strongly expanded than the others and the ventral surface of segments 7 and 8 with a transverse grove at their bases. Tarsi similar in males and females. Central lobe of aedeagus narrow, apical quarter thin, tip rounded. Paramere relatively narrow, inner edge crenulated in basal half, tip with one long seta. Figs 37-39. Etymology Latin. ‘Bulbus’ - swollen. A reference to its swollen antennae. THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 115 Remarks A moderate sized, boat-shaped species with wrap- around elytral epipleurae and antennae with distinctive, swollen, mid-segments in both sexes but more pronounced in the males. Nirripirti byroensis sp. nov. FIGS 43-45 Holotype f. ‘BES 9365, Byro Station, Yalcallia Well, 25° 54’ 39" § 115° 53’ 03” E, 10/6/02, W. F. Humphreys & R. Leys’, WAM 34215. In spirit. Paratype 1; as for holotype except BES 9366, SAMA. Description (number examined, 2) Habitus. Length 3.9 — 4.1 mm; elongate, relatively flat, slightly depressed in sutural region, moderately constricted at junction of pronotum/elytra; uniformly light testaceous; hindwing vestigial, reduced to tiny flap. Head. Large, slightly narrower than elytra; smooth, moderately strong reticulation with small even meshes, moderately covered with scattered small punctures; sides sloping outwards backwards from antennal bases, then inwards to base; eye remnant reduced to short suture. Antenna moderately thin, segments 1 and 2 cylindrical, about same length, segments 3 and 4 half width and shorter than segment 2, segment 5 bit longer than segment 4, segments 6 to 10 larger, subequal, segment 11 a little longer than segment 10, each segment with some very small setae on inside apically. Maxillary palpus elongate, segment 4 a little longer than segment 3. Pronotum. Slightly narrower than — elytra; anteriolateral angles projecting strongly forward; sides weakly sinuate, converging towards rear, posteriolateral angles obtuse; a few scattered minute punctures and some stronger ones along front edge; reticulation relatively weak. Elytra. Not fused, tightly closed, lacking inner ridges; elongate, sides almost parallel; smooth; covered with weak fine reticulation; sparsely covered with small punctures; a few additional larger punctures with long setae, more frequent towards apex and sides. Epipleuron moderately differentiated, broad in anterior quarter, then rapidly narrowing to middle, virtually absent along rest of elytron. Ventral surface. Prosternal process very strongly narrowed between coxae, not reaching mesothorax, apical half spatulate, sharply pointed, strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact in midline. Metathorax sharply triangularly projecting forward in midline; wings relatively broad but short; moderately rounded in midline behind. Metacoxal plates relatively narrow; metacoxal lines obsolete, virtually impunctate, moderately reticulate; closely adpressed to ventrite |. Ventrites 1 and 2 fused, sutural lines distinct in inner half, indistinct laterally, ventrites 3 to 5 mobile, virtually impunctate except for scattered shallow punctures; each ventrite with a few long central setae or bunch of long setae; hind edge of ventrite 2 slightly sinuate. Legs. Protibia relatively narrow, relatively even width, about 3x its basal width; protarsus strongly expanded, segment | broad, wider than long, lobes slightly asymmetrical, segment 2 about one half length of segment 1, segment 3 as long as segment | but narrower, deeply bifid, segment 4 very small and hidden within lobes of segment 3, segment 5 stout, cylindrical, longer than segment 3, segments | to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate, laterally compressed, with a few fine setae at apex; mesofemur with row of about 20 short spines along hind edge in basal half (Fig. 43); mesotarsus similar shape but smaller than protarsus. Metatrochanter relatively large, elongate/oval, rounded at apex (Fig. 44); metafemur elongate, lacking spines; metatibia weakly curved, slightly expanded towards apex; metatarsus elongate, segment 1 longest, segment 4 shortest; in combination segments | and 2 a little longer than others, segments 2 to 5 without spines other than at apex; claws weak. Male Unknown. Etymology Named after the pastoral station on which it was found. Remarks A large, well-chitinized species with thin tibiae but greatly expanded pro and mesotarsi, particularly the basal segments. The expansions are asymmetrical. Although only two female specimens are known the almost straight, relatively even width of the metatibiae place it in the Hydroporini. DNA sequencing places it in a relatively isolated position within Nirripirti (Remko Leys pers com) Nirripirti copidotibiae sp. nov. FIGS 46-51 Holotype m. “BES 9335, Innouendy Station, mineral expl. bore, site 431, 25° 49’ 19" S 116° 11’ 29" E, 9/6/02, 116 C.H.S. WATTS & W. F. HUMPHREYS Figs 46-51. Nirripirti copidotibiae: 46, lateral view of central lobe of aedeagus; 47, ditto dorsal view; 48, paramere; 49, mesotrochanter and mesofemur; 50, metatrochanter and metafemur; 51, dorsal view. Scale bar represents [mm (habitus only). W. F. Humphreys & R. Leys’, WAM 34216. Slide mounted. Paratype 1; m. ditto except BES 8808, 11/4/03, SAMA Description (number examined, 2) Habitus. Length 3.2 mm; elongate, relatively flat, slightly depressed in sutural region, not constricted at junction of pronotum/elytra but base of pronotum narrower than elytra; uniformly light testaceous; hindwing vestigial, reduced to tiny flap. Head. Relatively broad, a little narrower than elytra; smooth, moderately strong reticulation with small even meshes, a few scattered small punctures; sides parallel; eye remnant reduced to short suture. Antenna relatively thin except for expanded segments 6 and 7, segment | cylindrical, segment 2 about same length, a little constricted basally, segments 3 and 4 half width and length of segment 2, segments 5 to 7 moderately expanded, segments 8 tol0 thinner, subequal, segment 11 about as long as segment 10, a little narrower, each segment with some very small setae on inside apically. Maxillary palpus elongate, segment 4 a little longer than segment 5. Pronotum. Narrower than elytra; anteriolateral angles projecting strongly forward; sides subparallel, posteriolateral angles obtuse; a few scattered minute punctures and some larger ones along front edge; reticulation moderately strong. Elytra. Properly not fused, lacking inner ridges; elongate, nearly parallel sided, smooth; covered with moderately strong fine reticulation; a few scattered small punctures, a few additional larger punctures with long setae, more frequent towards apex and sides. Epipleuron moderately distinct, broad at base rapidly narrowing in anterior quarter, virtually absent along rest of elytron. Ventral surface. Prosternal process strongly narrowed between coxae, not reaching mesothorax, apical half spatulate, apex pointed, very strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact in midline. Metathorax projecting forward in midline; wings virtually absent; widely rounded in midline THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 117 I p e) P, 4 Ss \ i 0 & hy _ 4 VF _ Fs he BW fA ye _ Ces ‘ J ~ @, () yg] p Se es 4 7 Ui a A Langh KGL TGR = —— ww en AA HH Amon 37 L SEs A Figs 52-57. Nirripirti dingbatensis: 52, lateral view of central lobe of aedeagus; 53, ditto dorsal view; 54, paramere; 55, mesotrochanter and mesofemur; 56, metatrochanter and metafemur; 57, dorsal view. Scale bar represents Imm (habitus only). behind. Metacoxal plates large, metacoxal lines short, indistinct, area between them and forward onto mesosternum depressed forming a large distinct midline groove; virtually impunctate, moderately reticulate; closely adpressed to ventrite |. Ventrites | and 2 fused, sutural lines distinct in inner two-thirds, absent laterally, ventrites 3 to 5 mobile, virtually impunctate except for a few long central setae or bunch of long setae. Legs. Protibia relatively broad, widest in middle where it is about 5x its basal width, laterally compressed, almost blade-like; profemur with distinct, thin, dorsal portion near apex covering base of protibia; protarsus moderately expanded, segment | broadly rectangular, segment 2 about one half length of segment 1, segment 3 as long as segment |, deeply bifid, segment 4 very small and hidden within lobes of segment 3, segment 5 stout, laterally compressed, about as long as segment 3, segments | to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate, laterally compressed with a few fine setae at apex; mesofemur with row of 6 strong spines along hind edge in basal half (Fig. 49); mesotibia with large blade-like structure on ventral side near apex; basal segment of mesotarsus strongly expanded, other segments a little more elongate than protarsus. Metatrochanter relatively large, basal two thirds narrow, parallel sided, apical portion narrowing to blunt point, deflexed dorsally (Fig. 50); metafemur elongate, lacking spines; metatibia relatively broad, weakly curved, approximately the same width throughout except much narrower near base; metatarsus relatively stout, basal segment very long, nearly as long as length of other segments combined, with distinctive comb of 4 to 5 spines on outside, segments 2 to 5 short, about same length, segments 2 and 3 much wider than others, all segments with very strong spines; claws weak. Male Female unknown. Central lobe of aedeagus relatively broad, slightly wider at tip. Paramere relatively broad, tip with two long setae. Figs 46- 48, 118 C.H.S. WATTS & W. F. HUMPHREYS Etymology Latin. ‘Copis’ — knife. In reference to the sharp raised ridge on the protibia. Remarks A medium sized species immediately recognised by the highly modified legs. The pro and mesotibiae are enlarged and angular, particularly the mesotibiae, with blade-like extensions on the inside. The metatarsi have a greatly elongate first segment which has the spines which are usually found evenly spaced along the outside edge grouped tightly together near the middle. Another unique character is the distinct midline groove on the ventral surface of the thorax. Although only one male is known it is unlikely that all of these distinctive characteristics are restricted to males, Nirripirti dingbatensis sp. nov. FIGS 52-57 Holotype m. ‘BES 9347, Innouendy Station, Dingbat Well, 25° 52’ 32” S 115° 53’ 43” E, 10/6/02, W. F. Humphreys & R. Leys’, WAM 34217. Slide mounted. Paratypes 3: 1, as for holotype, SAMA, 2, as for holotype except ‘BES 9346’, SAMA. Description (number examined, 4) Habitus. Length 2.0 — 2.2 mm; elongate, relatively flat, not constricted at junction of pronotum/elytra; uniformly light testaceous; hindwing vestigial, reduced to tiny flap. Head. Relatively small, much narrower than elytra; smooth, rather weak reticulation with small even meshes, a few scattered small punctures; sides parallel; eye remnant reduced to short suture. Antenna moderately thick, segments 1 and 2 cylindrical, segments 3 and 4 half width and length of segment 2, segment 5 bit longer than segment 4, segments 6 to 10 larger, subequal, segment 11 1.5x length of segment 10, each segment with some very small setae on inside apically. Maxillary palpus clongate, segment 4 a little longer than segments 2 and 3 combined. Pronotum, Narrower than elytra; anteriolateral angles projecting forward; sides diverging towards the rear, posteriolateral angles right angles; a few scattered minute punctures and some larger ones along front margin; long setae at sides particularly towards front; moderately strongly reticulate. Elytra. Not fused, tightly locked, lacking inner ridges; elongate, widest just behind middle; smooth; covered with moderately strong fine reticulation; evenly but sparsely covered with small punctures, a few slightly larger punctures with long setae, more frequent towards apex and_ sides. Epipleuron moderately differentiated, broad in anterior fifth, virtually absent along rest of elytron. Ventral surface. Prosternal process strongly narrowed between coxae, reaching mesothorax, apical half oval, strongly pointed behind, weakly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae not in contact in midline. Metathorax strongly projecting forward in midline; wings very short or absent; broadly rounded in midline behind. Metacoxal plates large, metacoxal lines weak, well separated, weakly diverging in front quarter, reaching to half way to mesosternum; sparsely covered with small punctures, moderately reticulate; closely adpressed to ventrite |. Ventrites 1 and 2 fused, sutural lines distinct, ventrites 3-5 mobile, strongly reticulate with scattered small punctures and a few long central setae or bunch of long setae. Legs. Protibia narrow, widest near apex where it is about twice its basal width; protarsus weakly expanded, segment | broadly triangular, segment 2 a little smaller, segment 3 as long as segment 1, deeply bifid, segment 4 very small and hidden within lobes of segment 3, segment 5 stout, cylindrical, longer than segment 3, segments | to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate with a few fine setae at apex; mesofemur with row of 4 to 5 spines along hind edge in basal half (Fig. 55); mesotarsus a little more elongate than protarsus. Metatrochanter with apex weakly pointed, tip close to metafemur (Fig. 56); metafemur relatively stout, lacking spines; metatibia weakly curved, weakly expanded towards apex; metatarsus elongate, segment | longest, segment 4 shortest, in combination segments | and 2 shorter than others, segments 2 to 5 without spines other than at apex, segments 2 to 4 weakly hour-glass shaped; claws weak. Male Antenna and protarsi slightly more expanded than in female. Central lobe of aedeagus broad, widening towards apex; paramere relatively narrow, tip with two long setae. Figs 52-54. Etvmology Named after the well in which it was found. Remarks A medium sized species with weakly expanded tarsi and strongly pointed but weakly arched prosternal process. In morphology close to N. THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 119 58 59 60 62 4 N ¢ #, 7 © ‘ ; 0 df « O EQ so lS, < = @, vi] As .) 3 SLY RK [a ey l we “ | J, Is eg. e f ) \ | Figs 58-63. Nirripirti eurypleuron: 58, lateral view of central lobe of aedeagus; 59, ditto dorsal view; 60, paramere; 61, mesotrochanter and mesofemur; 62, metatrochanter and metafemur; 63, dorsal view. Scale bar represents |mm (habitus only). innouendyensis and N. skaphites but with different prosternal process, different metatrochanters, thinner antennae and broad aedeagus. Nirripirti eurypleuron sp. nov. FIGS 58-63 Holotype m. “BES 9385, Moorarie Station, Wogartha Well, 25° 57’ 58” S 117° 35’ 28” E, 12/6/02, W. F. Humphreys & R. Leys’, WAM 34218. Slide mounted. Paratype m. ditto except BES 8856, 16/4/03, SAMA. Description (number examined, 2) Habitus. Length 2.3 mm; narrowly elongate, relatively flat, slightly depressed in sutural region, pronotum much narrower than elytra, not constricted at base; uniformly light testaceous; hindwing vestigial, reduced to tiny flap. Head. Relatively small, much narrower than elytra; smooth, moderately strong reticulation with small even meshes, a few scattered small punctures; sides weakly indented at position of eye remnants; eye remnant reduced to very short suture. Antenna relatively stout, segment | cylindrical, segment 2 about same length, more oval, segment 3 half width and shorter than segment 2, segments 4 to 6 subequal in length becoming progressively slightly wider, segments 6 to 10 subequal, segment 11 about twice length of segment 10, each segment with some very small setae on inside apically. Maxillary palpus elongate, segment 4 about twice as long as segment 5. Pronotum. Narrower than elytra; anteriolateral angles projecting strongly forward; sides parallel, posteriolateral angles square, overlapping base of elytra; a few scattered minute punctures and a few relatively large punctures along front edge; strongly reticulate. Elytra. Possibly fused, lacking inner ridges; strongly elongate, sides parallel, smooth; covered with strong reticulation; a few scattered small punctures, a few additional larger punctures with long setae, more frequent towards apex and sides. 120 C.H.S. WATTS & W. F. HUMPHREYS oy fan ee Oe oS Ot ey 69 Figs 64-69. Nirripirti innouendyensis: 64, lateral view of central lobe of aedeagus; 65, ditto dorsal view; 66, paramere; 67, mesotrochanter and mesofemur; 68, metatrochanter and metafemur; 69, dorsal view. Scale bar represents [mm (habitus only). Epipleuron not differentiated, that portion of elytron visible ventrally very broad, broadest at junction of ventrites 2 and 3. Ventral surface. Prosternal process strongly narrowed between coxae, not reaching mesothorax, apical half parallel sided, apex sharply pointed, strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact in midline. Metathorax projecting forward in midline; wings very narrow, short; moderately rounded in midline behind. Metacoxal plates relatively narrow, large, metacoxal lines absent; strongly reticulate, virtually impunctate; closely adpressed to ventrite1. Ventrites | and 2 fused, sutural lines distinct in inner two-thirds, absent laterally, ventrites 3 to 5 mobile; virtually impunctate except for a few long central setae or bunch of long setae. Legs. Protibia narrow, widest near apex where it is about twice its basal width; protarsus weakly expanded, segment 2 about one half length of segment 1, segment 3 as long as segment 1, bifid, segment 4 very small and hidden within lobes of segment 3, segment 5 stout, cylindrical about twice length of segment 3, segments 1 to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate with a few fine setae at apex; mesofemur with row of 5 spines along hind edge in basal half (Fig. 61); mesotarsus similar to protarsus. Metatrochanter with curved inner edge, apex bluntly pointed (Fig. 62); metafemur elongate, lacking spines; metatibia straight, approximately the same width throughout; metatarsus elongate, segment 1 longest, segment 4 shortest, in combination segments | and 2 about same length as others, segments 2 to 5 without spines other than at apex; claws weak. Male (Female unknown). Antenna and legs as above. Central lobe of aedeagus narrowing in apical third, tip bluntly pointed. Paramere rather narrow, tip with single seta. Figs 58-60. Etymology Greek. ‘Eurypleuron’ - wide ribbed. In reference to the broad epipleura in this species, THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 121 Remarks Occurs in the same calcrete as 7) wogarthaensis which apart from the generic differences is much smaller and has narrow elytral epipleurae. A number of species of Nirripirti are now known to have “wrap-around” elytra: N. stegastos, N. skaphites, N. killaraensis and N. bulbous. These can be separated by characters given in the key. Nirripirti innouendyensis sp. nov. FIGS 64-69 Holotype m. “BES 9339, Innouendy Station, mineral expl. bore site, 431m., 25° 49" 19” § 116° I1’ 29” E, 10/6/02, W. F. Humphreys & R. Leys’ WAM 34219. Slide mounted. Paratypes 5; 1, as for holotype, WAM 34220; 1, as for holotype except, ‘BES 9334’, SAMA; 1, as for holotype except ‘BES 9339’, SAMA; 1, as for holotype except ‘BES 9343, 10/6/03’, SAMA; 1, as for holotype except ‘BES 9342, 10/6/02’, SAMA. Description (number examined, 5) Habitus. Length 1.8 — 2.1 mm; elongate, relatively flat, not constricted at junction of pronotum/elytra; uniformly light testaceous; hindwing vestigial, reduced to tiny flap. Head Much narrower than elytra; smooth, moderate reticulation with small even meshes, a few scattered small punctures; sides parallel; eye remnant reduced to short suture. Antenna moderately thick, segment | cylindrical, segment 2 oval, about same length as segment |, segments 3 and 4 half width and length of segment 2, segment 5 bit longer than segment 4, segments 6 to 10 larger, subequal, segment 11 2x length of segment 10, each segment with some very small setae on inside apically. Maxillary palpus elongate, segment 4 a little longer than segment 3. Pronotum, Same width as elytra; anteriolateral angles projecting strongly forward; sides diverging slightly towards rear, posteriolateral angles right angles; a few scattered minute punctures and some larger ones alone front edge; moderately reticulate. Elytra. Not fused, but tightly closed, lacking inner ridges; broad, sides subparallel, smooth; covered with moderately fine reticulation; evenly but sparsely covered with small punctures; a few additional larger punctures with long setae, more frequent towards apex and sides. Epipleuron weakly differentiated, that portion of elytron visible ventrally broad in anterior fifth, virtually absent along rest of elytron. Ventral surface. Prosternal process strongly narrowed between coxae, reaching mesothorax, apical half suboval, tip sharply pointed, in same plane as rest of body. Mesocoxae not in contact in midline. Metathorax strongly projecting forward in midline; wings very short; moderately rounded in midline behind. Metacoxal plates large, metacoxal lines weak, relatively wide, diverging slightly in anterior quarter, reaching to about half way to mesosternum; sparsely covered with scattered very small punctures; closely adpressed to ventrite 1. Ventrites | and 2 fused, sutural lines distinct, ventrites 3 to 5 mobile, with scattered sparse small punctures and a few long central setae or bunch of long setae. Legs. Protibia narrow, widest near apex where it is about three times its basal width; protarsus weakly expanded, segment | broadly triangular, segment 2 about one half length of segment 1, segment 3 as long as segment | deeply bifid, segment 4 very small, hidden within lobes of segment 3, segment 5 stout, cylindrical, longer than segment 3, segments | to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate with a few fine setae at apex; mesofemur with row of 4 evenly spaced spines along hind edge in basal half (Fig. 67); mesotarsus a litthe more elongate than protarsus. Metatrochanter with basal half broad, apical half elongate produced into long thin point (Fig. 68); metafemur elongate, lacking spines; metatibia curved, approximately same width throughout; metatarsus elongate, segment | longest, segment 4 shortest, in combination segments | and 2 about same length as others, segments 2 to 5 without spines other than at apex; claws weak. Male Little external difference between the sexes. Central lobe of aedeagus broad, widening towards apex. Paramere moderately broad, tip with two long setae. Figs 64-66. Etymology Named after the pastoral station on which it was found. Remarks A small, parallel sided, species readily recognised by the long, sharply pointed metatrochanters. One of the small number of Nirripirti with the prosternal process not arched in ventral view and with the tip reaching the metathorax. Nirripirti verrucosus sp. nov. FIGS 70-72 122 C.H.S. WATTS & W. F. HUMPHREYS Figs 70-72. Nirripirti verrucosus: 70, mesotrochanter and mesofemur; 71 metatrochanter and metafemur; 72, dorsal view. Scale bar represents 1mm (habitus only). Holotype f. ‘BES 9386, Moorarie Station, Wogartha Well, 25° S758" § 1179935" "28" E,. 12/6/02, Wok: Humphreys & R. Leys’, WAM 34221. Slide mounted. Description (number examined, 1) Habitus. Length 3.2 mm; elongate, relatively flat, slightly depressed in sutural region, weakly constricted at junction of pronotum/elytra; uniformly light testaceous; hindwing vestigial, reduced to tiny flap. Head. Relatively large, a little narrower than elytra; smooth, moderate reticulation with very small even meshes, a few scattered small punctures; sides weakly concave behind eye remnant; eye remnant reduced to small suture. Antenna with segment 1 cylindrical, segment 2 more oval and about same length as segment 1, segment 3 same width but much narrower than segment 2, segments 4 to 8 progressively wider, segments 9 and 10 a little narrower than segment 8, segment 11 about 1.3x length of segment 10. Maxillary palpus elongate, segment 4 longer than segment 5. Pronotum. Slightly narrower than _ elytra, anteriolateral angles projecting strongly forward, sides narrowing towards rear, base weakly constricted, posteriolateral angles obtuse; a few scattered minute punctures and a few larger punctures near front edge; moderately reticulate with small meshes. Elytra. Not fused, lacking inner ridges; elongate, almost parallel sided, smooth; covered with rather weak fine reticulation; a moderate number of scattered small shallow punctures, a few additional larger punctures with long setae, more frequent towards apex and sides. Epipleuron weakly differentiated, that portion visible ventrally broad in anterior quarter, then gradually narrowing to middle, virtually absent along rest of elytron. Ventral surface. Prosternal process strongly narrowed between mesocoxae, not reaching mesothorax, apical half unusually elongate, apex bluntly pointed, strongly arched in lateral view with highest point (viewed ventrally) between coxae. Mesocoxae in contact in midline. Metathorax THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 123 1.0mm 0.imm 0.1mm / | 0.1mm Pe ~J ~ a \ 79 7 We, 0.imm 1.0mm MW Figs 73-86. Illustrations of characters used in the key to species. 73, protibia of Bidessodes gutteridgei. 74, ditto Tjirtudessus padburyensis. 75-76, Pronotal processes of T. morgani; 76 T. bialveus. 77-81, Mesosternal plates of; 77, 7. morgani; 78, T. pulpa; 79, T. bialveus; 80, Nirridessus fortisspina; 81, N. plutonicensis. 82-84, ventrites 1-2 or 1-3 of; 82, T. sweetwatersensis; 83, T. challaensis; 84, T. masonensis. 85, Ventral view of abdomen showing wide wrap-around” epipleura of N. stegastos. 86, ditto, N. hamoni showing flared shoulders and narrow epipleura. 86 1.0mm BO e 1.0mm 1.0mm 83 C.H.S. WATTS & W. F. HUMPHREYS 124 sadvulvipoaryed g sajaroyeo ¢¢ ds snssapngaily pure 6 LN ds mrditiny snipqn snypjadoy L.LN ds pndistin SISUASAAIDMJIAMS SNSSOPNJALLT eAre] CLIN ¢, 8 ds iuidissin nynfininy pysulury sisuaauoynyd YAldiidin solspsajs Nd ‘aou “ds sysuajpqsuip 1.11dL.tn ‘sou “ds snsoons.iad 1d ¢ ds snssapnp.ilT ds snssapnpanly saiaupjuad YAldidIN sisuadqdaddou 1Aldsdin SASADJOAIDUL SNSSAPNAALLT snyis snssapnpAly, SISUDUOSDUL SNSSapNJALT, puldssis0f ANGLIN, apazuly 111 SISUIASOAJIU 1ALALALIN, appjsodp] snssapnjally ‘ds snssapnjaulT 1UDSAOU SNSSAPNIALT, pdjnd snssapny.uly ds snssapnpul] SISUBUNS]IU NALGIALIN SISUIAUOISAUL] SAPOSSAPLY saylydoys NAIdsINy aou “ds s7suaotdg UAIIALIN ‘aou “ds s7suadpuanouut YA1d1ALIN saou “ds woinadaina YAldiadin ‘sou “ds snqjng 141dluN 7 ds snssapny.uly SISUATJaqsIg SnssapnpAll]T, STSUdATD STISSOPNPATL | SISUBABPAM NALGLALIN' SISUQUAADYMAU 1-111 AINE SNIDYAAIOAIDUL NALAIAALN, snaajvlg snssapny.AulyT, sIsuanaAUnd snssapnyaly sisuaazapunl snssapnyAnl$e SISUIAMOUUINA SNSSAPNBAUT, sIsuaapIsav.l SnssapnyAlT sisuasajopuuid snssapnyuly SISUA[JAMADPLY SNSsapnygALlT, SISUIJOPADP LAIGLAMIN SISUADLIDPUIM SASSapNgAUT, Llajyuly snssapny.ulyT 1UYPY SNssapnyauly, sisuapungiM snssapnyaAul], Ipapy.iaga snssapnyAlT ‘aou “ds pyno00491U snssapnyAuT, IUOUDY TAIIAAIN 195P113JJNB SaPOSsaplg SISUaIPUNDADY Snssapnpuly ‘aou “ds saplouyop.av Yd.) ‘aou “ds aviquopidos 14ididiN ‘sou “ds sysuapyjuvsom snssapnpuly ‘aou “ds sysuadingpod snssapnpuly ‘aou ‘ds sysuamarayyiy snssapnyAuly ‘aou “dS ST/D]MAPIOI0 STLINSUOOg ‘aou “ds ST/D]UaPTIIO STLINSUOOg | ds snssapnpuly SISUIDAD]]LY YAIGLAAIN SISuUaD]JDYI SnssapnyALlT, STSUDATD STISSAPNPAILT SNIYIUSDUL SNSsapnyAl] quasaid satsadg “LN culseg eleSN “LN culseg @1e5N “LN culseg el[eSN nioqqeN nioqqeN asouleD apisaey apisoey apisaey apisoey Aare Kaled Aared Aaled Aared Kare 00/\ auAoosen audoosery auAoosen UOSTYoINJA] uosIyoIny uOSTy SIN] uOsTyoINyA uOsTYyoINIA| uOsTysINA uosIyInyA UOSTysINA uosIysinyyy uosTyoINyy uOsTYpIN] UOSTYoINA uOsTysINJAL Aay[eAaoored aSpony WNOYPy [eNUDD “ECE USABYMON “TE Aqaodden ‘1 ¢ Ads :ndung ‘og slajemjaams :nAuny ‘67 sopuny “87 AIOUUINA “£7 uoseyy dP ‘OT WIS SOpORUUI “STZ sdulids yodaq ‘pz Qoyzeg oye7) UONRIS ISOI[III “ET BLIIBPUIA JUNO ‘77 TSM J9PTUTH “TT oye] yepinuresy “07 WIOLA PAP] “61 ore “8 SHOVNIVUC GNVINI eluemeuung ‘/ | uoneys 1opuer’y ‘9] Wong UNS] “C1 UONRIS SIDATY IIL ‘pT Ipunyerey “¢ | som OA TI Apuonouy] ‘| | d1IeIOOW “0 | Aingped WA “6 AMOIATIIH *8 YMON Prey “L ulg Ulg aLIB1ooyy “9 PLINUIPUI “¢ ele “P YWON PyPeYD *¢ sumoq unsny ‘7 on) ‘| SHOVNIVUC NYALSAM a1or9yea “9}919]B9 DUO UB} SOLU UL IND90 pauT].apun saloadg saded siy} Ul poyeo.zy ssoy} ae pyog Ul UMOYsS soldadg “payeorpul ore sosvUTLIP IOLIOVUL pur Uev9dQ UPIPU] oy} pue (7g “SIq) sula}SAs OSvUTeAPOoL]ed oyeIedos oY] “VI/D.USNP Ul Salpog ajasojp9 ajasgsp IsSuouly splosyAp fo saisads jvSAjs fo UOYNGLAISIP YL. 3 VIAN, THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 125 projecting forward in midline; wings very narrow; moderately rounded in midline behind. Metacoxal plates large, metacoxal lines absent; virtually impunctate; closely adpressed to ventrite 1. Ventrites 1 and 2 fused, sutural lines distinct in inner two- thirds, absent laterally, ventrites 3 to 5 mobile, virtually impunctate except for a few long central setae or bunch of long setae. Legs. Protibia distinctly club-shaped, widest towards apex where it is about four times its basal width; protarsus strongly expanded, segment | almost square, segment 2 about one half length of segment |, segment 3 longer than segment 1, deeply bifid, segment 4 very small and hidden within lobes of segment 3, segment 5 stout, cylindrical, about as long as segment 3, segments | to 3 with dense covering of adhesive setae; claws short and simple. Mesotrochanter elongate with a few fine setae at apex; mesofemur with row of 5 short spines along hind edge in basal half (Fig. 70), the basal ones closer together, anterior edge uneven with a number of small protuberances, metatibia somewhat club- shaped; mesotarsus considerably more elongate than protarsus. Metatrochanter moderately large, basal half parallel sided apical half triangular, apex blunt, well separated from femur (Fig. 71); metafemur elongate, anterior edge uneven, lacking spines, ventral surface with short setae; metatibia relatively thick, curved, approximately the same width throughout; metatarsus elongate, segment | longest, segment 4 shortest; in combination segments | and 2 the same length as the others, segments 2 to 5 without spines other than at apex; claws weak. Male Unknown. Etymology Latin. ‘Verrucosus’ — full of warts. In reference to the warty edges of the mesofemurs. Remarks A large species recognised by the long thin club-shaped pro and mesotibia and the noticeable bumps on the front edge of the mesofemur. In general morphology resembles N. hinzeae and N. darlotensis but as well as the club-shaped tibiae and warty mesofemurs it has more elongate metatarsi, the prosternal process is much less deflexed and, uniquely among the Australian stygal Dytiscidae, the surface of the metafemurs have a moderate covering of setae (Fig. 71). Although known only from a single female specimen the relatively straight, even-width, metatibiae place it in the Hydroporini. Mitochondrial DNA places it in an isolated place within Nirripirti (Remko Leys pers. com.). Discussion Distribution Species described herein represent stygal beetles from two palaeodrainage systems, the Murchison and the Moore, both of which drain to the Indian Ocean (Fig. 87); in earlier papers in the series (Watts and Humphreys 1999, 2000, 2001, 2003; Balke et al 2004) we also recorded beetles from palaeodrainage systems which drained to the inland. It brings the number of described stygal Dytiscidae for Australia to 55 species in five genera (Bidessodes, Copelatus, Kintingka, Nirripirti and Tjirtudessus). These are derived from 33 discrete groundwater calcrete deposits in eight palaeodrainages representing both coastal and interior drainages. In addition a number of other species that have not been formally described are recognised from larvae and females. Of the 55 described stygobitic species of Dytiscidae from Australia (‘bid.), T; microocula is the first in which the eyes are not entirely TaBLe 2. Water quality data for several of the stvgal water beetles. Values are presented for the upper layer of water only through which the beetles need to pass to breathe. The vertical stratification for some deeper bores is shown in Figs 88- 90. Species Temp. (°C) pH Specific Salinity DO% DO ORP(mV)_ Depth (m) conductance (TDS mg L') (mg L"') (mS cm!) T.. padburyensis 19.81 7.94 2.49 1.28 53.5 4.82 448 0.1 T. hillviewensis 21.07 7.87 L.8 0.91 88.1 7.72 295 0.2 N. dingbatensis 21.54 7.9 15 8.68 57.3 4.77 351 0.3 N. copidotibiae 27.12 7.39 2.54 1.31 82.4 6.5 371 0.3 N. innouendyensis 27.12 7.39 2.54 1.31 82.4 6.53 71 0.3 N. bulbus 25.17 8.53 2.6 1.34 93.8 7.64 346 0.3 N. arachnoides 19.87 8.1 4.88 2.6 72.9 6.51 228 0.1 N. byroensis 19.87 8.1 4.88 2.6 72.9 6.51 228 0.1 B. occidentalis 26.47 7.68 2.47 1.27 54.9 4.36 313 0.1 126 C.H.S. WATTS & W. F. HUMPHREYS GRE Lmestone. caterete | Surficial sediments Calcrete Map Fig. 87. The distribution of the species discussed in this paper in the groundwater calcrete aquifers of the Yilgarn area of Western Australia. The dark shading denotes groundwater calcrete bodies and the lighter shading the surficial sediments associated with the palaeodrainages incised into the Archaean basement. The calcrete bodies are referred to as 1: Byro; 2, Innouendy; 3, Bunnawarra; 4, Mt. Gould (the Wittenoom population of B. occidentalis sp. nov. is 400 km north of this site); 5, Moorarie; 6, Mt. Padbury; 7 Killara North; 8, Hillview. All sites are in the Murchison catchment except Bunnawarra which is in the Moore palaeodrainage. Map based on 1: Australia 1989 compiled by D.P. Commander. 4 : Temperature (° g 4 ra £ 2° ORP (mV) 10 a ; Be ue ae 3 2 E « = a & Dissolved oxygen 7 (% saturation) ts See ee a = 4 = Po) & oe fay Dissolved oxygen % Specific conductance (mS cm ~1 | (mg L-1) 2a? Fry a2 4a eR Fig. 88. Depth profiles of several physico-chemical parameters in the bore inhabited by Boongurrus occidentalis sp. nov. in an uncapped, never-used water bore in the Killara North calcrete. 2,500,000 Hydrogeological Map of Western reduced, being about one-fifth the size of those in epigean species and seemingly lacking discrete ommatidia. Most interestingly, an undescribed sister species (determined from DNA) from the same calcrete also retains eye remnants to a similar degree suggesting that this lineage is of more recent evolution to subterranean life than the other stygal members of the genus. Boongurrus occidentalis sp. nov. is the first clearly epigean species to be found in deep groundwater in Australia. However, other epigean species have been recorded from underground habitats: Copelatus australis (Clark) from hyporheic systems within river gravels in the Flinders Range, South Australia (Remko Leys, pers. com.) and C. irregularis Macleay from small pools of water in the furthest reaches of deep vertical caves in arid Cape Range, northwestern Australia (W.F.H. personal observation). THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 127 TABLE 3. Stygofauna recorded from the same calcrete bodies from which the Dytiscidae reported here were collected. Calcrete Associated stygal taxa Bunnawarra Badja Killara north Mt Padbury Mt Gould Innouendy Byro Moorarie Belele Amphipoda, Ostracoda, Harpacticoida, Cyclopaedia, Annelida Oniscidea (lsopoda), crangonytoid and Ceinidae Amphipoda, — Cyclopoida, Ostracoda Bathynellacea, crangonytoid and Ceinidae Amphipoda, Harpacticoida, Cyclopoida Bathynellacea, crangonytoid and Ceinidae Amphipoda, Cyclopoida, Ostracoda Bathynellacea, Amphipoda Microturbellara, crangonytoid Amphipoda, Cyclopoida, Ostracoda Ostracoda Amphipoda, Ostracoda, Cyclopoida Ostracoda _ Depth (m) ° & 20.8 fo) Depth (mm) & & r Depth (m) Temperature (°C) bp ‘i Temperature (°C) if E: iy, £ Ew Be ORP (mV) “ 16 ‘es Paneer me) ea 2 2 wo oe a 4 “ Py Ew 0 Dissolved oxygen y Bee vA pH (% saturation) Ow 78 ~ eee i Nae Dissolved oxygen (mg i-1) _ Depth (m) Specific conductance (mS cm 1) 3 4 5 4 $ é 7 ° Fig. 89. Depth profiles of several physico-chemical parameters in a capped bore inhabited by Nirripirti bulbus sp. nov. and Boongurrus occidentalis sp. nov. | | | ORP (mV) Specific conductance (mS cm-1 ) 21.2 21.4 216 218 22 1 1.5 295 296 297 Dissolved oxygen (% saturation) Dissolved oxygen (mg L -1 ) Fig. 90. Depth profiles of several physico-chemical parameters in Camel Well inhabited by Tjirtudessus hillviewensis sp. nov. in the Hillview calcrete. 128 C.H.S. WATTS & W. F. HUMPHREYS Environment Groundwater in the arid zone is sometimes markedly stratified in respect of salinity inter alia (Watts and Humphreys 2000). However, groundwater characteristics near the water table must be utilised by at least the larger species of stygal beetles, as they need to traverse it to reach free air to breathe. The physico-chemical characteristics of the superficial part of the groundwater at the sites of 9 of the 13 species described herein are given in Table 2. These waters are generally of rather low salinity for the Yilgarn (900-2600 mg L! TDS) with the exception of the site for N. dinghatensis sp. nov. (Table 2). The latter site, while being part of the Murchison palaeovalley, which now contains the large episodic Murchison drainage, now lies in the separate small drainage of the Wooramel River and showed a much greater salinity (8680 mg L"' TDS). Several sampling sites contained sufficiently deep water for profiling. Generally these showed little stratification and the values over which the variables ranged was small (Figs 88 and 90). Only the site occupied by N. bulbus sp. nov. exhibited a marked salinity gradient (Fig. 89) and this was accompanied by large changes in pH, redox and oxygen level. In other stratified systems a pronounced nadir in oxygen levels associated with the halocline and the reduction in pH have been associated with a cascade of nitrogen species and sulphur bacteria (Humphreys 1999). The typically high nitrate and sulphate contents of the Yilgarn aquifers potentially could similarly support chemoautotrophic sulphur bacteria, providing a source of energy for the ecosystem (Humphreys 2001), however, there is no indication from the ORP values that such is the case here and the changes in oxygen level may result from groundwater flow. Associated fauna The fauna associated with the stygal beetles is shown in Table 3 at a high taxonomic level. These are likely to represent substantial diversity and many short-range endemics, as has been found in some other taxa in calcretes containing stygal dytiscids where studies have been completed. For example, 31 species of copepods, including five new genera and 23 new species have been described for other calcretes in the Yilgarn (Karanovic 2003). Five new species of Candonopsis (Candoninae: Ostracoda) have been described from similar areas, all but one species restricted to a single calcrete area (Karanovic and Marmonier 2002). Finally, four new species of stygal Oniscidea (families Scyphacidae and Philosciidae) have been described from calcretes, three of the species from a single saline calcrete (Taiti and Humphreys 2001). Acknowledgments We greatly appreciate the support of Remko Leys in the field, Julianne Waldock in Perth, and the information and access provided by the managers of pastoral leases. The illustrations were very ably prepared by Howard Hamon. This work was partly supported by a grant from the Australian Biological Resources Study. References BALKE, M., Watts, C. H. S., Cooper, S. J. B., HUMPHREYS, W. FL & VoGier, A. P. (2004) A highly modified stygobitic diving beetle of the genus Copelatus (Coleoptera, Dytiscidae):; taxonomy and_ cladistic analysis based on mtDNA sequences. Sysfematic Entomology 29: 59-67. Cooper, S. J. B., Hinzr, S., Leys, R., WATTS C. H. S. & Humpureys, WF. (2002). Islands under the desert: molecular systematics and evolutionary origins of stygobitic water beetles (Coleoptera: Dytiscidae) from central Western Australia. Jnvertebrate Systematics 16: 589-598. Humpureys, W. F. (1999). Physico-chemical profile and energy fixation in Bundera Sinkhole, an anchialine remiped habitat in northwestern Australia. Journal of the Royal Society of Western Australia 82: 89-98. ee (2001). Groundwater calcrete aquifers in the Australian arid zone: the context to an unfolding plethora of stygal biodiversity. Pp 63-83 /n ‘Subterranean Biology in Australia 2000’. Eds. W. F. Humphreys and M. S. Harvey. Records of the Western Australian Museum, Supplement 64. KaARANovic, I. & MARMONIER, P. (2002). On the genus Candonopsis (Crustacea: Ostracoda: Candoninae) in Australia, with a key to the world recent species. Annals of Limnology 38: 199-240. KARANOVIC, T. (2004). Subterranean copepods (Crustacea: Copepoda) from arid Western Australia. Crustaceana Supplement 3: 1-366. Larson, D. J. (1994). Boongurrus rivulus, a new genus and species of water beetle (Coleoptera, Dytiscidae, Bidessini) from northern Queensland, Australia, Journal of the Australian Entomological Society 33: 217-221. Leys, R., Cooper, S. J. B., WATTS, C. H. S., & HUMPHREYS, W. F. (2003). Evolution of subterranean diving beetles (Coleoptera, Dytiscidae, Hydroporini, Bidessini) in the arid zone of Australia. Evolution 57: 2819-2834. Taitil, S. & Humpureys, W. F. (2001). New aquatic Oniscidea (Crustacea, Isopoda) from groundwater calcretes of Western Australia. Pp.133-151. In ‘Subterranean Biology in Australia 2000’. Eds. W. F. Humphreys and M. S. Harvey. Records of the Western Australian Museum, Supplement No. 64. THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON Warts, C. H. S. & Humpnreys, W. F. (1999). Three new genera and five new species of Dytiscidae (Coleoptera) from underground waters in Australia. Records of the South Australian Museum 32: 121-142. & Nirridessus and Tjirtudessus (Dytiscidae; Coleoptera) from underground waters in Australia. Records of the South Australian Museum 33: 127-144. (2000). Six new species of 129 os & (2001). A new genus and six new species of Dytiscidae (Coleoptera) from underground waters in the Yilgarn palaeodrainage system of Western Australia. Records of the South Australian Museum 34: 99-114. & (2003). Twenty-five new Dytiscidae (Coleoptera) of the genera Tjirtudessus Watts & Humphreys, Nirripirti Watts & Humphreys and Bidessodes Regimbart, from underground waters in Australia. Records of the South Australian Museum 36: 135-187. REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK (COLEOPTERA: SCIRTIDAE) By C. H. S. WatTs* Summary Watts, C. H. S. 2004. Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Trans. R. Soc. S. Aust. 128(2), 131-168, 30 November, 2004. The Australian members of the genera Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae) are revised. Thirty three species are recognised: Scirtes helmsi Blackburn, S. exoletus Waterhouse, S. brisbanensis Pic and 30 new species: S. alastairi, S. albamaculatus, S. auritus, S. baroalba, S. beccus, S. calmi, S. crassiantennae, S. cygnus, S. emmaae, S. interstinctus, S. kaytae, S. macroconcolor, S. microrotundus, S. musica, S. nalyerensis, S. nigerpalpus, S. orientalis, S. peniculus, S. podlussanyi, S. pygmaeus, S. pinjarraensis, S. ruforotundus, S. rivularis, S. spatula, S. storeyi, S. tindaleensis, S. triangulus, S. victoriaensis, Ora floccosus, O. improtectus, O. justafloccosus. The genus Ora is recorded from Australia for the first time. All species are described and the male genitalia illustrated. A key is provided to the species. Key Words: Coleoptera, Scirtidae, Australia, taxonomy, morphology, distribution. Transactions of the Royal Society of S. Aust. (2004), 128(1), 131-168. REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK (COLEOPTERA: SCIRTIDAE) by C. H. S. WaTTS*® Summary Watts, C.H.S. 2004. Revision of Australian Scirtes [lliger and Ora Clark (Coleoptera: Scirtidae). Trans. R. Soc. S. Aust. 128(2), 131-168, 30 November, 2004. The Australian members of the genera Scirtes Iliger and Ora Clark (Coleoptera: Scirtidae) are revised. Thirty three species are recognised: Scirtes helmsi Blackburn, S. exoletus Waterhouse, S. brisbanensis Pic and 30 new species; S. alastairi, S. albamaculatus, S. auritus, S. baroalba, S. beccus, S. calmi, S. crassiantennae, S. cygnus, S. emmaae, S. interstinctus, S. kaytae, S. macroconcolor, S. microrotundus, S. musica, S. nalyerensis, S. nigerpalpus, S. orientalis, S. peniculus, S. podlussanyi, S. pygmaeus, S. pinjarraensis, S. ruforotundus, S. rivularis, S. spatula, S. storeyi, S. tindaleensis, S. triangulus, S. victoriaensis, Ora floccosus, O. improtectus, O. justafloccosus. The genus Ora is recorded from Australia for the first time. All species are described and the male genitalia illustrated. A key is provided to the species. Key Worps: Coleoptera, Scirtidae, Australia, taxonomy, morphology, distribution. Introduction Australian Scirtidae (Marsh Beetles) are a common component of fresh water ecosystems. The larvae are detritus feeders living in shallow water near the edges of ponds, marshes and rivers, or in situations were they can readily reach the surface to breathe. Adults are terrestrial, sheltering in thick vegetation near water or feeding on nearby flowering shrubs. Very little is known about their natural history, in major part due to the inability to identify species, other than the tree-hole dwelling Prionocyphon niger Kitching & Allsopp (1987) from South-eastern Queensland extensively studied by Kitching (Kitching & Callaghan 1982). Blackburn (1891), Lea (1919), and Carter (1935) described a number of species (under the rubric Helodidae) but it was not until Jack Armstrong began his work on the group in the early 1950’s that much taxonomic attention was paid to the family. Unfortunately he completed and published only one paper in which he described 3 genera and 14 species (Armstrong 1953). When he stopped his taxonomic work he was close to the completion of manuscripts on the genera Cyphon and Scirtes; so close to completion, that specimens of these genera labelled as paratypes are scattered in a number of collections. The generic boundaries and placements of the Australian Scirtidae require close examination (Hiroyuki Yoshitomi pers. com.). This problem is less critical in the genera Ora and Scirtes, which, although both probably polyphyletic, are well characterised by the possession of greatly enlarged hind femurs that are used for jumping, presumably as * Division of Entomology, South Australian Museum. a predator avoidance mechanism. The genus Ora has not previously been recorded from Australia but three species described here from Northern Australia appear to belong in the genus. The genus Ora is typically found in tropical rainforests in America, Africa and Asia (Yoshitomi pers. com.). The few Australian specimens know have all come from tropical areas close to if not in rainforest. The genus Scirtes occurs most commonly in tropical northern Australia where species are often collected at light but also in southern Australia with one species, S. exoletus Waterhouse, reaching northern Tasmania. This is in direct contrast to the situation in the remaining Australian Scirtid genera which have a more southern distribution and are commonest in Tasmania, in both number of species and in number of individuals. Methods For the identification of many species dissection and examination of the male genitalia is required. The female genitalia, although varying somewhat between the species examined, are more uniform and offer few taxonomic characters. For dried specimens, specimens were sexed by examination of the apex of the abdomen. In many females (approximately 80%) the ovipositor was at least partially visible. In a few species the tips of the male genitalia are also often visible. Specimens without extruded ovipositors were softened in an ultrasonic water bath for 30-45 minutes and the genital complex (see later) was teased, approaching dorsally, from the rest of the abdomen with a pair of fine forceps. The penis and tegmen were separated from the modified abdominal segments 8 and 9 and mounted on card in a drop of P.V.A. wood-working glue, which is transparent 132 C. H.S. WATTS when dry, or on a microscope slide in a polyvinyl alcohol based mountant. Drawings were made from camera lucida tracings. For a number of species only a handful, or fewer, of male specimens were available, often old and hence difficult to soften and dissect, which resulted in less than perfect material to work from. The male genitalia are complex and can vary enormously in structure between species. In these circumstances the preparations were often adequate to clearly indicate that a taxa was new but not good enough to allow a confident interpretation of fine structural detail. Thus I expect that in some cases my interpretations of the detail of the male genitalia may need to be modified when fresh material becomes available. As well as the male genitalia, specimens of nearly all the small to medium sized species were also mounted on microscope slides. Source of specimens Specimens on which this revision was based were obtained from the following collections. AM, Australian Museum, Sydney; ANIC, Australian National Insect Collection, Canberra; QPIM, Queensland Department of Primary Industries, Mareeba; HUNG, Hungarian Natural History Museum, Budapest; NHM, Natural History Museum, London; NMV, Museum of Victoria, Melbourne; NTM, Northern Territory Museum and Art Gallery, Darwin; QM, Queensland Museum, Brisbane; SAMA, South Australian Museum, Adelaide; UQIC, University of Queensland Insect Collection, Brisbane. Notes on morphology Australian species of Ora and Scirtes, like their members in other areas, vary considerably in overall shape — oval, round, flanged, large, small — and colour — yellowish, reddish, black, mottled, spotted — but the differences are rarely discreet enough to differentiate species. Significant structural differences are few and I have only found two to be of much taxonomic use: the shape of the metacoxae and the male genitalia (penis and tegmen). The width of the metafemur, shape of the metatrochanter, grooving on the elytral epipleura and the relative size of the eyes are also useful to differentiate some species. Metacoxae. In Scirtes the metacoxal plates are extended backwards in the midline and cover the articulation of the metatrochanters with the coxae. The extensions of the metacoxal plates can be relatively wide and short or almost square, and the hind edge either straight (Fig.lb) or concave (Fig.lc). In Ora the ventral portion of the extension Fig. 1. Metacoxal plates, metatrochanters and metafemurs of; a) Scirtes auratus sp. nov.; b) S. nigerpalpus sp. nov.: c) S. tindaleensis sp. nov.; d) Ora floccosus sp. nov. Line = 1.0 mm. is lacking or virtually so, completely exposing the articulation of the trochanters (Fig.1d). Male genitalia. The basic male genitalia of Australian Ora and Scirfes consists of a penis and a bilobed tegmen situated immediately above it. Enclosing these are modified tergites and sternites of the 8th and 9th abdominal segments (Fig. 2). The nomenclature of the complex and very variable male genitalia of Scirtidae is unsettled. In this paper I follow Nyholm (1972). Penis. The most common structure for the penis (eg Figs 20-30) of the Australian species consists of an oval “basal piece”(“pala” of Nyholm) with a usually longer distal extension, the “trigonium”, which is articulated with it. The basal piece has either one or two parameroids arising from its sides. This ground plan is recognisable in most species but in some, i.e. O. floccosus (Fig. 4) and O. justafloccosus (Fig. 5), the morphology of the separate pieces is so altered as to obscure their REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 133 Fig. 2. Terminal abdominal segments of Scirtes emmaae sp. nov. a) tergite 7; b) sternite 7; c) tergite 8; d) sternite 8; e) tergite 9; f) sternite 9; g) Scirtes ruforotundus sp. nov. tergite 7. Line = 0.5 mm. homologies. In one, S. ruforotundus, the opposite appears to have happened and the structure has been greatly simplified into a unique (for Australian Scirtes), small, unstructured penis and a relatively simple tegmen (Fig.15). Tegmen. The tegmen lies above the penis and is closely attached to it basally. Apically the tegmen consists of two usually symmetrical lobes which can be broad and close together through to long, thin and well separated. These lobes are thought to — Fig. 3. Female genitalia of Scirtes emmaae. a) ovipositor; b) tergite 8; c) sternite 8. Line = 1.0 mm. originate from parameres which are variably and often intimately connected with the tegmen in Scirtidae (Nyholm 1972). In a number of the species described the lobes of the tegmen are broad and wrap around or enclose the penis (eg. Fig. 35), in most of these species portions of the outer edges of the lobes are strongly sclerotonized. In O. floccosus and O. justafloccosus as well as enclosing the penis one lobe is closely articulated with the penis and the other lobe is seamlessly fused to it C. H. S. WATTS (Figs 4, 5). Abdominal segments (Fig. 2). The 8th and 9th abdominal segments are modified and are only weakly sclerotonized. The 8th and 9th tergites are broad structures, each with lateral sclerotized strut- like structures (apodemes) extending backwards. The sternites are less strongly sclerotized, the 8th reduced to a small “U” shaped structure, with the arms pointing apically. The 9th tergite is broadly bilobed apically with a covering of short to medium length setae towards the apex. Between the species examined there is not a lot of difference in the structure of these segments and I have not used them taxonomically. Female genitalia (Fig. 3). Within the species examined there is some difference in the detail of the apex of the ovipositor (which is a modified 9th abdominal segment) but it is not great and I have not used it taxonomically in this paper. The ovipositor consists of a long thin basal portion (baculus) with a short two lobed apical piece (coxite) of variable length, with a small appendage at their tips (stylus). Closely attached to the Ovipositor is the 8th tergite which has two long, thin, lateral struts/rods. The 8th sternite is similarly shaped but with much shorter lateral struts/rods. Systematics Scirtidae with greatly enlarged hind femurs are currently placed into two genera, Ora and Scirtes, depending on the form of the hind coxal plates. In Ora the basal articulation of the metacoxae is exposed (Fig. 1d) whereas in Scirtes it is covered at least to some extent (Figs Ib, Ic). Within the Australian species of both genera the general form varies considerably, however the details of the external morphology are very similar which makes any attempt to key the species or to clearly define groupings difficult. In contrast, the form of the male genitalia varies a lot between species, even in those that appear otherwise identical. Because of this many of the new species are essentially defined by the form of the male genitalia. Nyholm (2002) has used the form of the male genitalia to propose a number of semiformal groups within the Northern Hemisphere members of Scirtes. 1 am not convinced that this approach is warranted at this stage of our knowledge of the Australian species and have not attempted either to use Nyholm’s groupings or to propose new groupings. Purely for the convenience of this paper I have used the phrase *S. helmsi species complex” for a group of species, including S. he/msi, which are virtually indistinguishable on external morphology. On the evidence of the male genitalia they appear unlikely to be monophyletic. REVISION OF AUSTRALIAN SCIRTES ILLIGER AND OR4A CLARK 10. Key to Australian species of Ora and Scirtes ly to Nn Metacoxal plates lacking ventral portion, completely exposing articulation of metatrochanters (Fig. 1d) (Ord)... 2 Metacoxal plates with ventral portion covering all or most of articulation of metatrochanters (Figs La-c) (Scirtes) cscs sessetceosaieeseeeeas 4 Dorsal surface mottled; pronotal punctures very SHTML Pet Rv sanstaagegeea tot teera Basiedocttan ogehocgesesiihte 3 Dorsal surface uniformly dark reddish-yellow; pronotal punctures StrOng.......ceeeeeseteeteeeeeeeees ng saevaheaMy na yenbut seer NTs O. improtectus sp. nov. Penis with three long thin pieces (Fig. 4)........... $1 Mcastaleba cu Pelt day cretiged aes O. floccosus sp. nov. Penis with two long thin pieces (Fig. 5) ............ batten, Mires Neer A nelle Mert O. justafloccosus sp. Nov. Each elytron with several large white areas; 2.0 — 2.3 mm long......S. a/bamaculatus sp. nov. Elytra without extensive white areas; Di = SVS ay TONG sass ssasesdosccesistecctnrspes teaseecezed 5 Hind edges of individual metacoxal plates CONCAVE) arscdaietaeiiadtoalde bones 6 Hind edges of metacoxal plates straight, slightly convex or slanted towards midline ................ 10 2.6 — 3.6 mm long; metatrochanter elongate, 2.5x as long as wide (Fig.la); uniformly golden VOM OWs psa duet asad eeateaioss S. auratus sp. nov. 3.2 — 5.5 mm long; metatrochanters 2.0x as long as wide (eg Figs 1b, Ic); reddish-yellow or Gathers ces deyetiregittideaternettecs tale alate Seating 7 Oval: body setae fine; colour dark chocolate red, hind edges of metacoxal plates weakly concave wehdhe elude sregtiabared aves S. ruforotundus sp. nov. Elongate-oval; body setae strong; colour reddish-yellow; hind edges of metacoxal plates Strongly CONCAVE.......:ccsecesceseetseseesseeesneeeseseeneeee 8 Front of frons flat, edge weakly to strongly upturned; antennal segments thick, weakly asymmetric with inner edge larger, particularly in males; elytra reddish-yellow often with darker sides and near scutellum; lobes of tegmen with spines on outside, tip of penis hooked. (Fig.12). Mahle Werepbasabia eee Ma ae S. kaytae sp. nov. Front surface of frons evenly rounded, front edge not upturned; antennal segments relatively thin, symmetrical; elytra uniformly reddish- yellow; lobes of tegmen smooth, tip of penis not hooked... Sides of elytra subparallel; dark reddish-yellow; tips of tegmen slightly bulbous, triangular extension near bases of lobes (Fig.16); tip of penis with three thin lobes, (Fig.16) ...... ee Npaarlianed Strbalnsctce beet bvdckaended es S. tindaleensis sp. nov. Oval; light reddish-yellow tending darker towards front; tegmen lobes with small triangular extension on outside near apex (Fig. Qik palit ga seaa ea ste pasa guaeiseseaaelcaah S. emmaae sp. nov. Il. 2.0mm long; almost round; elytra dark............... peciadlesekpasrpeates tht Vasctvades S. microrotundus sp. nov. 2.0 —5.1 mm long; oval rather than round; elytra variable 1m COLOUL...... cece see eeesseteeeteeeeeeenes 1] 4.5 — 5.0 mm long; uniformly light reddish- yellow (some specimens with vague darker pattern on pronotum); elytra flanged ............... diye Lelelertlegalspiea vice dgteecenaee dt S. macroconcolor sp. nov. 2.0 — 5.1 mm long; colour variable but never uniformly reddish-yellow in species > 4.0 mm long; elytra variably flanged ....... eee 12 . Head, pronotum and often elytra with dark/light colour pattern; epipleuron weakly grooved in front half; 3.5 — 5.0 mm long... eee 13 Without colour pattern (except occasionally on head and pronotum); 1.8 — 4.3 mm _ long; epipleuron flat except at extreme front............... (S. helmsi species complex. Only distinguishable on characters of male genitalia. See separate key below.) . Elytron with 3-4 disrupted darker stripes, elytron quite strongly flanged towards front... (bear Ter cee ek Ata Erect S. interstinctus sp. nov. Elytron without linear markings, either uniformly reddish yellow or with darker bases and sides; elytron at most weakly and narrowly HBT GEA vac. sc pe secoureenadiees S. exoletus Waterhouse Key to males of the S. felmsi species complex Le to Small (2.2 — 2.5 mm). Trigonium long and thin with apex bent at right angles (Fig. 32), single parameroid long, thin, sinuous; tegmen broad, lobes short and _— slightly asymmetric (Southwestern Western Australia) ........cccceeeece Eileandtarncaeiuaiaatis totuaatchiaa Scirtes pygmaeus sp. nov. Not with above combination of characters ....... 2 One lobe of tegmen extremely long and thin, closely associated with elongate trigonium and projecting well beyond apex of trigonium (Fig. 26) (Southwestern Western Australia) ...........00... hanantere tanesincaet¥azsardel Scirtes nalyerinensis sp. nov. Tegmen lobes symmetrical or nearly so, tips never projecting much beyond apex of trigonium Tegmen with two, well-separated finger—like lobes (eg. Fig. 24); trigonium of penis elongate > twice length of basal piece (eg. Fig. 40); one or two parameroids with the larger one elongate and usually with a hooked end (eg Fig. 40) .....4 Lacking above combination of characters: tegmen often with lobes broad and poorly separated (eg. Figs 31-33); trigonium of penis variably shaped, about same length or shorter than basal piece (eg. Figs 21, 29, 34): parameroid(s) variable but usually not elongate with hooked end. .......ececeeeeceeseeseeeeeeeeeeeeeeneeee 10 136 10. . Trigonium of C. H. S. WATTS With well developed second parameroid (right hand one) which is one quarter to one third length of main parameroid (left hand one) (Figs, DIOR MISTS 1p Ree PNT ATER POTEET A > Usually without second parameroid (Figs 20, 30, 24), if present, < a quarter length of main parameroid which is strongly hooked at apex (P1238 )et eh esa rpetesestelestittet ot lot emote te its eats te 8 Dark coloured, with dark antennae and palpi; elytron not flanged; trigonium of penis broad, tip rounded, slightly upturned. (Figs 27, 42) ........... Ratnam Treen rons ce S. nigerpalpus sp. nov. Light coloured with pale palpi and at least the basal segments of antennae pale; elytra weakly flanged towards front; trigonium of penis narrower (Figs 28, 37). ....eeeeeeseesseeeseeereeeeeeees 6 Main parameroid sinuate towards apex (Figs 37, 44), penis approximately 0.4x length of body.... pepreas nicest haste tataeete eet S. victoriaensis sp. Nov Main parameroid with apical hook, penis approximately 0.2x length of body ............0.. 7 Much of body dark. Largest parameroid with abrupt: hooks(Figs 33540 aires. staat dstctenees ae tuee cee seat ae tae rs Att Scirtes rivularis sp. nov. Body more or less uniformly yellowish. Largest parameroid hook not abrupt (Figs 28, 43) .......... Ba cet Ne RR Ee Miata te cohen S. orientalis sp. nov Penis golfclub-shaped; trigonium long and thin, tip sharp, (Figs 24, 40)........ S. helmsi Blackburn Penis with apex expanded dorsal/ventrally near tipi(Figs-20,:30;, 38. SO) tat. dss owesescenrueh eae 9 Usually with very small right hand parameroid, left hand (main) parameroid with well-defined hook at apex (Figs 20, 38)... 8. brisbanensis Pic Only one parameroid, with long hook (Figs 30, SOc Meceadcestaee mies S. pinjarraensis sp. nov. Trigonium of penis short 0.4x length of basal piece, strongly beak-shaped (Fig. 19)...........0. atts Se bet eee Seto cases ie seater ee se S. beccus sp. nov Trigonium of penis at least 0.75x length of basal piece, not beak-shaped 0.0.0... eee eens 1] penis asymmetrical, with prominent spine on inside (Fig. 25)... ples ih deteh terion: sunt t t Se eet es S. musica sp. nov. Trigonium of penis more or less symmetrical, without spine on inside (eg Fig. 23)... 12 . Penis with two distinct parameroids (Fig. 21).... ETT A RM eMac st atten S. calmi sp. nov. Penis with only one parameroid (eg. Fig. 23) .13 . Parameroid of penis swan-like (Fig. 23)... SET na Ce Ie Peat tats hee S. cygnus sp. nov. Parameroid of penis more elongate, not swan- TUG. aes semmeat omer oh a eRe th cases ce 14 Tegmen with lobes thin, well separated (Figs 18, 2D DOI BRR cece: Meaeees. Resta ite tes Beko Mc ec 15 Tegmen with lobes broad, partly separated, tending to wrap around and enclose penis (Figs 17,5343 D8. Ob Aaa ceves celevtaa yh Me BM acest os asa 18 15. Tegmen with lobes with distinct brush of setae near apex (Fig. 29)... S. peniculus sp. nov. Tegmen lobes without setae wo... 16 16. Penis complex, parameroid much larger than trigonium (Fig.18)........... S. baroalba sp. nov. Penis of more usual form, trigonium larger than parameroid:. (Pigs 23d) 22. .rmsn msnseon ae 17 17. Parameroid hooked at apex, about two-thirds length of trigonium; tegmen lobes with serrated inner edges near apex (Fig. 22)... eee aii rca arese sus ara rs S. crassiantennae sp. nov. Parameroid of penis not hooked, about same length as trigonium; lobes of tegmen smooth Ges Teen bate eee ee eee Et S. podlussanyi sp. nov. 18. Penis with trigonium broad (Fig. 34)... eoce aera sneha eesveeg Reon rs tee Rae S. spatula sp. nov. Penis with trigonium thin (Figs 17, 35, 36)....19 19. Penis with parameroid elongate, without apical hook, nearly as long as trigonium (Fig. 35)........ snarls cenit teen nesstiy, way, Maas S. storeyi sp. nov. Penis with parameroid short, either sinuate or hooked at apex (Figs 17, 36)... eee 20 Body uniformly light reddish-yellow except for darker head in some; parameroid of penis stout (Eas 2-25 22 GAMMA ON Gs sao sssilenstes atl guia a Geaete oases ye tne ate ates ne S. alastairi sp. nov. Dorsal surface chestnut, segments of antennae paler, shoulders of elytra and triangular patch at apex of elytra lighter, parameroid of penis more elongate (Fig. 36); 3.1 mm long wu... ST eR SRE I BE te , Ol S. triangulus sp. nov. 20. Ora Clark, 1865 (Species listed in alphabetic order.) Ora floccosus sp. nov. (Fig. 4) Types. Holotype male; “ AUSTRALIA 99.1.13 Queensland, Pinnocle village (camping) leg. A. Podlussany”, SAMA. Paratypes: 3; 1, “Cow Bay N of Daintree, N Qld. 27.xii.83 — 20.41.1984, I. C. Cunningham”, QPIM; 1, “Edge Hill Cairns, at light, 24/4/65, J.G.Brooks”, microscope — slide, ANIC; 1, “AUSTRALIA n Qld 15 km NNW of South Johnstone Light Trap Nov 1987 Fay & Halfpapp”, QPIM. Description (number of dissected males examined, 4) Habitus. Length 3.3 — 4.2 mm., relatively flat, broadly oval. Head. Light yellow-brown with darker patches; antennae light yellow-brown. Small, width between eyes about 2.3x dorsal width of eye; slightly REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 137 Figs 4-5. Dorsal views of tegmen and penis. Lines = 0.5mm. 4. Ora floccosus sp. nov. 5. O. justafloccosus sp. nov. depressed inwards from eyes. Punctures small, moderately dense, each with a prominent whitish seta. Frons with sides slightly diverging, front edge straight or very weakly concave, edges weakly beaded. Segment | of antenna large, barrel-shaped; segment 2 smaller, barrel-shaped; segment 3 a little longer but narrower; segments 4-10 longer, broader, particularly middle ones; segment 11 a little longer than segment 10; segments quite thickly covered with short whitish setae. Pronotum. Light yellow-brown with darker mottlings. Short, broad. Puncture small, even, moderately dense, each puncture with a whitish seta. Hind angles obtuse, front angles moderately extended forward, sides weakly beaded. Scutellum. Light yellow -— brown. Sides approximately equal length, lateral two weakly convex. Punctures small, weak. Elytron. Light yellow-brown with darker mottlings. Sides widely flanged in middle particularly over metafemurs. Densely punctured punctures of uneven sizes, each puncture with a short whitish seta. Epipleuron wide in front quarter, widest some distance from shoulder, then narrowing evenly to near apex, weakly concave particularly near front. Ventral surface. Light reddish-yellow. Pronotal process very narrow along whole length. Mesosternum with narrow groove for reception of pronotal process; tip just reaching to level of mesocoxae. Front triangular midline extension of metasternum narrowly triangular, strongly beaded; rear midline extension broad, more than twice as wide as long. Metacoxal plate a little wider than long, ventral part of plate virtually absent, completely exposing articulation of metatrochanter; anteriolateral angle extending narrowly some way along metasternum; sides weakly beaded; posteriolateral angles sharply pointed. Meta- trochanter small, about 2.0x as long as wide. Metafemur moderately swollen, widest in middle, hind edge with large notch near apex. Notch bordered with v-shaped raised ridge. Dorsal metatibial spine relatively long, a little longer than twice size of ventral spine and about three quarters length of segment 1 of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites with punctures small, dense, each with a short fine seta; reticulation moderate, fine, more pronounced towards rear; apex of apical ventrite truncated or weakly concave. 138 C. H.S. WATTS Male Little external difference between sexes. Penis complex (Fig. 4). The trigonium appears to consists of three long thin pieces joined to each other near their bases. Penis lacking basal piece. Tegmen lobes asymmetrical, complex, one roughly sickle-like with the blade-like top portion closely enclosing one of the trigonium pieces and the handle-like lower portion closely held to but not fused with the trigonium piece (Fig. 4). The other lobe of the tegmen has its base seamlessly fused to a piece of the trigonium and the upper portion closely enclosing another piece of the trigonium. (The figure illustrates the genital capsule slightly teased apart: in life the tegmen lobes tightly enclose the three pieces of the trigonium.) Variation The strength of the notch on the hind femur and the strength of the colour pattern are variable. Referred specimens — all female 1, Bramston Beach, near Innisfail, N Qld. 30 April 1976, D. H. Colless (open savanna), ANIC; 4, Cairns, 2/50 G B, J. G Brooks Bequest 1976, ANIC; 1, Russell R. at Bellenden Ker Landing NQ, 5m, 24 Oct ~ 9 Nov. 1981, EARTHWATCH/QLD MUSEUM, ANIC COLEOPTERA Voucher # 83-0588, QM Etymology Latin. “floccosus” — woolly. Notes This species and the very similar O. justafloccosus are distinctive species with widely flanged elytra and a generally ‘woolly’ appearance including the antennae and tarsi, These two species are separated most readily from O. improtectus by their mottled colouration and small pronotal punctures. Ora floccosus and O. justafloccosus can only be separated by the form of the male genitalia. Ora floccosus appears to be a more southern species from around Cairns and QO. justafloccosus more northern from around Iron Range. However too few male specimens are known to have much confidence in this geographic separation. Ora improtectus sp. nov. (Fig. 6) Types Holotype male; “Stuart Range Q Jan — Feb 1927 Hale & Tindale”, SAMA. Paratypes 42: . 1, “15 54S 163 32E Batten Point, 30 km NE by E Borroloola, NT, 30 Oct.1975, M. S. Upton”, ANIC; 2, “Cairns, 2/50, G.B”. “J. G. Brooks Bequest, 1976”, ANIC; 1, “Cairns 7-10-34" J. G. Brooks Bequest 1976”, ANIC; 2, “Cairns dist., E. Allen” “I 52 52”, SAMA; 2, “Halftide nr Mackay NEQ 8.ii.65 E. C. Dahms” QM; 1, “12.268 130.56E Holmes Jungle, Berrimah 10km S of Darwin, NT, 8.xi.72, at light, E. Britton”, ANIC; 5, “King R NT 10-1-16", NMV; 14, ditto, 7-1-16, 10 NMYV, 4 slides SAMA; 3, “King R NT Coll by W. McLemman esq and pres by H. L. White esq 14.10.16”, NMV; 1, “15 04S 145 O7E Mt Webb Nat. Pk. QLD, 28 — 30 Sept 1980, T. Weir’, ANIC: 1, “Nassau River NW Dunbar Stn., 18 Noy. 1983, A. Walford-Huggins”, ANIC; 6, “Thursday Island, 10/52, CM”, “J. G. Brooks Bequest 1976”, 5 QPIM, | slide SAMA. 2, “Thursday Is] NQ Oct-1952”, NMYV; 1,“ Smith Point NT L.viii.1982 C.Wilson & S. Collins”, “ex light trap”, NTM; 1, “Stuart Range Q Jan— Feb 1927 Hale & Tindale”, SAMA. Description (number examined, 43) Habitus. Length 2.5 — 3.6 mm., relatively flat, oval. Head. Reddish-yellow to dark reddish-yellow; antenna light reddish-yellow. Small, width between eyes about 2.0x dorsal width of eye. Moderately and evenly punctate, each puncture with a relatively long setae. Frons with sides slightly diverging, front edge straight or very weakly concave, edges beaded. Segment | of antenna large, cylindrical slightly curved at base; segment 2 much smaller, cylindrical; segment 3 as long as segment 2 but a little narrower; segments 4 — 10 long, narrow, cylindrical, becoming progressively slightly smaller; segment 11 slightly longer than segment 10, moderately setose. Pronotum. Reddish-yellow to dark reddish-yellow. Short, broad. Evenly covered with strong punctures, becoming almost confluent at sides, each puncture with a moderately long golden seta. Hind angles obtuse, front angles moderate produced forward, sides beaded. Scutellum. Reddish-yellow, tending to be lighter than elytra. Sides approximately equal length, lateral two convex; punctures large, well separated. Elvtron. Reddish-yellow to dark reddish-yellow on disc becoming darker laterally and towards apex. Sides weakly and narrowly flanged, more strongly towards front. Moderately dense strong punctures, each puncture with a short yellow seta. Epipleuron relatively wide in front quarter becoming much narrower over rest of elytron, front portion with wide shallow longitudinal depression in many. Ventral surface. Reddish-yellow with diffuse darker patches. Pronotal process very narrow between procoxae, apical portion not much wider. Mesosternum with narrow triangular groove for REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 139 Figs 6-11. Dorsal views of tegmen (left) and penis (right). Lines = 0.5 mm. 6. Ora improtectus sp. nov. 7. Scirtes albamaculatus sp. nov. 8. S. auritus sp. nov. 9. S. emmaae sp. nov. 10. S. exoletus Waterhouse. 11. S. interstinctus sp. nov. 140 C. H.S. WATTS reception of pronotal process; tip reaching just to level of front edge of mesocoxae. Front midline extension of metasternum narrowly triangular, beaded. Rear triangular midline extension of mesosternum broad, about twice as wide as long. Metacoxal plate a little wider than long, anterio- lateral corner extending some way along mesosternum, ventral portion of plate absent towards rear, exposing articulation of metatrochanter; sides beaded: posteriolateral angle sharp. Metatrochanter small, about 2.0x as long as wide. Metafemur greatly swollen, widest well before middle, hind edge with moderate notch near apex; reddish-yellow, diffusely darker in places. Dorsal metatibial spine moderately long, about twice length of ventral spine and nearly as long as segment | of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites reddish-yellow, often with diffuse darker areas; punctures rather sparse, very small, reticulation, strong, fine, more pronounced laterally; apex of apical ventrite truncated or weakly concave. Male Little external difference between sexes. Basal piece of penis short, rather square; trigonium much longer and for much of its length separated into two elongate parts with rounded tips, a broadly triangular thin structure lies on top of the trigonium for much of its length (Fig. 6). Tegmen longer than penis, lobes well separated, thin, with strong setae on inner edge in apical half (Fig. 6). Variation Colour varies from reddish-yellow to dark reddish- yellow. Etymology Latin. “Improtectus” — unprotected, a reference to the unprotected metatrochanters. Notes A small northern species separated from O. floccosus and O. justafloccosus by its smaller size, uniform rather than mottled dorsal surface and strong pronotal punctures. In contrast to S. floccosus and S. justafloccosus, the male genitalia are relatively normal. The tegmen is unique in having strong setae/spines on the inside towards the tips. Ora justafloccosus sp nov. (Fig. 5) Types Holotype male “12.438 143.18E, 11 km ENE of Mt Tozer QLD, 11-16 July 1986, T.Weir & A Calder’, ANIC. Paratypes 3, male; 1,“ Iron Range, Cape York Pen. N. Qld. 26 —31 May 1971. G. Monteith” “UQIC Reg# 53698”, UQIC; | slide, ditto, “11 — 17 May 1968 UQIC Reg# 53696”, UQIC; 1, “12.43S 143.17E 9km ENE Mt Tozer QLD 5 — 10 July 1986 T. Weir & A. Calder”, ANIC; 1, “15 03S 1435 09E 3 km NE Mt Webb QLD 30 April— 3 May 1981 A. Calder”, ANIC. Description (number examined, 4). As for O. floccosus except as follows. Male Penis consisting of two long thin pieces, slightly grooved and not as long as in O. floccosus (Fig. 5). One lobe of tegmen relatively similar to O. floccosus (Fig. 4) the other lobe fused to penis much nearer its apex than in O. floccosus (Fig. 5). Etymology. Latin. “Juxta” — near. A reference to its close appearance to S. floccosus. Notes See under O. floccosus. Referred specimens — all female 1, Captain Billy Creek Cape York Pen, N Qld, 142.50E 11.408, 9 — 13/7/75, G. B. Monteith, QM; |, Iron Range Cape York Pen. N. Qld. 11 — 17 May 1968, G. Monteith, UQIC Reg# 53695, UQIC; 3, Iron Range Cape York Pen. N. Qld. 16 — 23/11/65, G. Monteith, UQIC Reg# 53692, 88 — 89, UQIC; I, Iron Range Cape York Pen Qld 28 April — 5 May 1968, G. Monteith, UQIC Reg# 53697, UQIC. Scirtes Uliger, 1807 a) Species other than those in the S. Ae/msi species complex. Arranged in alphabetic order. Scirtes albamaculatus sp. nov. (Fig. 7) Types. Holotype female; “Cairns Queensland F.H.Taylor”, ANIC. Paratypes 4; 1, “AUSTRALIA, Qld Bramston Beach Eubenangee Swamp, 4.v.1987, J.K. Bulcjunas, collected on Melaleuca quinquenervia”, ANIC; 1, Cape Tribulation, 8.1.1983, R. Storey, At Light, QPIM; I, “Cape Tribulation, 1 Aug — 15 Sept 1987, A Walford - Huggins, coastal rainf. NQ, intercept trap, ANIC; | slide, Cow Bay N of Daintree, 25.1 — 7.2 1984, 1. C. Cunningham, SAMA. REVISION OF AUSTRALIAN SC/RTES ILLIGER AND ORA CLARK 141 Description (number examined, 5) Habitus. Length 2.0 — 2.3 mm., relatively flat, oval. Head. Reddish-yellow, antenna yellowish. Small, width between eyes about 3.2x dorsal width of eye. Moderately and evenly punctate. Frons with sides moderately converging, front edge slightly concave, edges weakly beaded. Segment | of antenna large, barrel-shaped; segment 2 about same shape and size, segment 3 about half as long and narrower; segments 4 — 10 same length as segment 2, cylindrical, progressively becoming slightly broader and flatter, segment 11 a little longer than segment 10. Pronotum. Reddish-yellow to dark reddish- yellow, disc somewhat darker, with slight darker markings. Short, broad. Evenly and moderately punctate, each puncture with a moderately long golden seta. Hind angles obtuse, front edge sinuate, front angles weakly projected forwards, sides weakly beaded. Scutellum. Reddish-yellow to dark reddish-yellow. Sides approximately equal length, lateral two slightly convex. Elytron. Reddish-yellow to dark reddish-yellow, with two large areas of dirty white. Sides beaded rather than flanged. Moderately and evenly punctate, each puncture with a short yellow seta. Epipleuron yellowish with colour varying to reflect elytron colour, moderately wide in front quarter, becoming much narrower over rest of elytron, central area of front portion widely depressed slightly. Ventral surface. Dark reddish-yellow with lighter areas, appendages tending lighter. Pronotal process very narrow. Mesosternum with relatively broad, triangular groove for reception of pronotal process; tip reaching past level of mesocoxae. Front extension of mesosternum small, in shape of equilateral triangle; rear midline extension in midline large about twice as wide as long. Metacoxal plate wider than long, anteriolateral corner extending some way along metasternum; hind edge slightly concave, slanting strongly to midline; length of midline short, about as long as midline of triangular backward extension of mesosternum; sides weakly beaded; posteriolateral angles bluntly pointed. Metatrochanter small, elongate, about 3x as long as wide. Metafemur greatly swollen, widest near middle. Dorsal metatibial spine moderately long, about twice size of ventral spine and a little over three quarters length of segment | of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites with punctures small, shallow; reticulation, moderate, fine, more pronounced towards rear; apex of apical ventrite truncated or weakly concave. Male Little external difference between the sexes. The only available preparation of the male genitalia is poor. From what can be made out the genitalia are comparatively small and complex (Fig. 7). There are at least two long (? trigonium) pieces to the penis reminiscent of species such as O. floccosus. The lobes of the tegmen are narrow, bluntly tipped and well separated (Fig. 7). Variation. Little variation within the five know specimens, except in the shape of the white areas on the elytra, with one specimen having the front patch broken up into two discreet patches on each elytron. Etymology Latin. “Albus”- white, “macula’”- spot, a reference to the white spots on the elytra. Notes A distinctive small species, with small eyes and several distinct white patches on its otherwise dark elytra, a large segment 2 of the antenna and elongate metatrochanters. The hind edges of the metacoxal plates are slightly concave and slant strongly backwards towards the midline resulting in the midline suture being relatively short compared to other species. Scirtes auratus sp. nov. (Figs 1, 8) Types Holotype male, “Qld. Townsville 10km NW 23/3/96 C. Watts”, SAMA. Paratypes 34; 1, “Homestead, Silver Plains Via Coen, N. Qld. I1.xl1, 1964 G. Monteith” “UQIC Reg# 53649”, UQIC; 25, “Ross R. Dam Spillway Townsville, NQ 1 Dec. 1986 T. Vernon ex Melaleuca leucodendra’’, ANIC; 1, “2k N Mt Molloy Qld. 5.2.97 C. Watts”, SAMA; 3, “Sk N.W. Mt Molloy Qld. 5.2.97 C. Watts”, SAMA; 2, “Nardello’s Lagoon Qld. 6.2.97 C.Watts”, SAMA; 1, “Qld. Bushland Beach 20km N Townsville A. J. Watts 23 — 30/12/97”, SAMA; 1,” Qld. Bushland Beach 20km N Townsville, at light, A. J. Watts 16 — 18 Jan 1998”, SAMA. Description (number examined, 110) Habitus. Length 2.6 —3.6 mm, relatively flat, oval. Head. Light reddish-yellow, antennae light reddish-yellow. Small, width between eyes about 2.8 x dorsal width of eye. Quite strongly and evenly punctate, well covered with long golden setae. Frons with sides diverging in front of antennal base which is deeply excised into side of head; front edge straight or very weakly concave, edges not beaded. 142 C. H.S. WATTS Segment | of antenna large, barrel-shaped; segment 2 smaller, barrel-shaped; segment 3 smaller and narrower; segments 4 — 10 long, rectangular, flattish; segment 11 a little longer than segment 10, quite strongly setose. Pronotum. Light reddish-yellow, sometimes with diffuse darker mottlings or patterns. Short, broad. Evenly and moderately punctate, stronger laterally, each puncture with a long golden seta. Hind angles obtuse, anterio- lateral angles moderately produced, sides weakly beaded. Scutellum. Light reddish-yellow. Relatively large, sides approximately equal length, lateral two weakly convex; punctured as on pronotum. Elytron. Light reddish-yellow with diffuse darker areas in some. Sides weakly and narrowly flanged in front third. Moderately and evenly punctate, each puncture with a short yellow seta. Epipleuron relatively wide in front quarter becoming much narrower over rest of elytron, front portion widely and shallowly grooved. Ventral surface. Light reddish-yellow, sometimes with diffuse darker areas. Pronotal process very narrow. Mesosternum with narrow groove for reception of pronotal process, tip reaching past level of front margin of mesocoxae. Front midline extension of metasternum relatively small, sharply triangular; rear midline extension of metasternum broad, at least twice as wide as long, apex rounded. Metacoxal plate a little longer than wide, with anterio-lateral corner extending some way along metasternum; hind edge moderately concave, posterior-lateral angles sharply pointed. Metatro- chanter narrow, elongate, about 2.5x as long as wide (Fig. 1). Metafemur greatly swollen, widest just before middle, moderate indentation on hind edge near base. Dorsal metatibial spine relatively short, about twice size of ventral spine and about half- length of segment | of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites with punctures small, shallow; with moderate fine reticulation, more pronounced towards rear; apex of apical ventrite strongly concave. Male Penis complex, two equal sized fleshy lateral lobes (? parameroids), trigonium trilobed with central lobe with small, strongly chitinized, serrated structure at its base (Fig. 8). (An alternative interpretation is a single lobed trigonium and upper and lower pairs of parameroids.). Tegmen lobes shorter than penis, elongate, thin, pointed, well separated, some quite strong spines on outside edge (Fig. 8). Variation Colour varies from light reddish-yellow to a golden yellow, with head and pronotum with diffuse darker areas or even distinct dark patterning in some. Etymology Latin. “Aurum” — gold, a reference to the colour of the beetle. Notes A relatively small, noticeably golden species with weakly concave hind edges to the metacoxal plates and relatively long, elongate, metatrochanters. Equally distinctive are the male genitalia which are unusually complex for Australian Scirfes, with fleshy parameroids and a small asymmetric, heavily chitinized, structure in the centre. I have reared the species from larvae, also relatively golden, collected from among 7yp/a in a shallow semi-permanent small lake. Adults were collected from the emergent rushes. Specimens examined Queensland. 2, Arriga via Mareeba, 16/10/85, K. N. Halfpapp, ex rice paddy, QPIM; 3, Cairns, 2/50, G. Brooks, ANIC; 1, Cardstone, 3 — 4/12/66, J. C. Brookes, ANIC; 1, Ingham, K. J. Sandery, 29/5/23, ANIC; 1, Christmas Creek, 15 K W of Fairview via Laura, 26 — 27/6/73, G. B. Monteith, QM; 1, Cow Bay N of Daintree, 18 — 25/1/84, I. C. Cunningham, QPIM:;: 1, 3mi SSW of Millaa Millaa, 30/10/68, R. J. Elder, ANIC; 2, Mossman, 8/1/84, at light, J. D. Brown, QPIM; 3, 2 mi SW of Mt Inkerman, 19 45S 147 30E, 11/12/68, S. Misko, ANIC; 7, 4mi W of Mourilyan, 5/11/66, sandy soil at light, E. Britton, ANIC; 5, Ditto, 11/66, G. Brooks, ANIC; 1, Pinnocle Village, 13/1/99, A. Podlussany, HUNG; 1, Rocky River via Coen, 10mi N, G. Monteith, UGIC Reg# 53654; 1,15km WNW _ South Johnstone, 9/5/86, at light, Fay & Halfpapp, QPIM; 2, Tolga, 10/1/86, at light, QPIM; 1,Walkamin, 15/3/84, at light, J D. Brown, QPIM. Northern Territory. |, Berry Springs, 30km SSE of Darwin, 11/11/72, at light, E. Britton, ANIC; 3, 7km NW by N of Cahills Crossing, East Alligator River, 12, 23S 132.56E, 27/5/73, E. G. Matthews, ANIC; 1, 5 km NNW of Cahills Crossing East Alligator River, 12, 23S 132,57E, 28/5/73, E. G. Matthews, ANIC; 1, Cahills Crossing East Alligator River, 12 26S 132 S58E, 29/5/73, at light, E. G. Matthews, ANIC; 1, Cannon Hill via Jim Jim, 18/8/71, T. Weir & A. Allwood, NTM; 1, Finniss River Station, 2/4/86, C. Wilson, ANIC; 2, Fogg Dam, 16/2/87, on Sida cordifolia, ANIC; 1, CSIRO HQ Kalpalga, 12.408 132 22E, 19/6/79 (my light), G. Monteith & D. Cook, QM; 1,10km N Jabiru, 21/9/82, R. I. Storey, at light, QPIM; 5, Jubiru, R. I. Storey, 17-20/9/82, QPIM, REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 143 Scirtes emmaae sp. nov. (Figs 2, 3, 9) Types Holotype male, “Cardstone QLD 15.xi.1966 J. G. Brooks”, ANIC. Paratypes 21; 1,“Qld. Bushland Beach 20km N Townsville, 23 — 30/12/97, A. J. Watts”, SAMA; 1, “Cairns Q Dec 50 J.G. Brooks, NMV; I, “Cape Tribulation, N. Qld. 24-29.xii.1980 R. I. Story & N. Gough Rainforest”, QPIM: 4, “Cape Tribulation, N Qld. 8.1.1983, R. I. Story, at light” QPIM; 1, “Cardstone QLD 16.xi.1966 J. G.Brooks”, ANIC; 1, “Cardstone QLD 19.xi.1966 J. G. Brooks”, ANIC; 2, “Cardstone, N.Q. xi.66. K. Hyde”, ANIC; 2, “Flying Fish Pt. 21/1/65 E. G. Dahms, QM; 3, “N.T. 12.358 131.20E Kemp Airstrip Rainfor. 24 — 25 July 1979, G. Monteith & D. Cook”, QM; 1, “Killymoon Ck. 25 k S Townsville Qld., 2.2.97, C. Watts”, SAMA; 1, “AUSTRALIA Northern Territory Mt Bundey, 144m” 13 13 582S 131 08 OI8E, 4 — 6 xi 2000, leg. A. Podlussany”, HUNG; 3, “AUSTRALIA 99.1.13 Queensland, Pinnocle village (camping) leg, A. Podlussany”, HUNG. Description (number examined, 107) Habitus. Length 3.3 — 4.3 mm., relatively flat, oval. Head. Reddish-yellow to dark reddish-yellow. Small, width between eyes about 2.2x dorsal width of eye. Strongly, densely and evenly punctate, cach puncture with a long golden seta. Frons with front angles bulging outwards and may be slightly bent upwards, front edge straight or very weakly concave, edge strongly beaded. Segment | of antenna large, barrel-shaped; segment 2 about half as long, cylindrical, segment 3 as long as segment 2, narrower; segments 4 — 10 long, narrow, cylindrical; segments 10 and 11 subequal. Pronotum. Reddish-yellow to dark reddish-yellow. Short, broad. Evenly and quite densely punctate, punctures moderately strong, confluent at sides, each with a long golden seta. Hind angles obtuse, front edge strongly sinuate, sides weakly beaded. Scutellum. Reddish-yellow; a little longer than broad, lateral sides weakly convex. Elytron. Reddish-yellow. Quite strongly and evenly punctate, each puncture with a yellow seta although often abraded off. Weakly flanged towards front. Epipleuron yellow, relatively wide in front quarter, becoming much narrower evenly over rest of elytron. Ventral surface. Light reddish-yellow. Pronotal process very narrow between procoxae, apical portion not much wider, strongly keeled. Mesosternum with narrowly elongate groove for reception of pronotal process; tip reaching level of mesocoxae. Rear triangular midline extension of metasternum narrower than front extension; front extension about twice as wide as long. Metacoxal plate broader than long with anterio-lateral corner extending some way along metasternum; hind edge strongly concave; sides beaded; posteriolateral angles rounded. Metatrochanter small, apex pointed. Metafemur greatly swollen, widest just before middle, quite strongly indented on hind edge near apex. Dorsal metatibial spine relatively long, more than twice size of ventral spine and about two thirds length of segment | of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites with punctures small, shallow; reticulation moderate, fine, more pronounced towards rear; apex of apical ventrite truncated or concave. Male No external difference between sexes. Basal piece of penis oval; trigonium a little shorter than basal piece, elongate, triangular, tip rounded; two similarly shaped parameroids, as long as trigonium (Fig. 9). Tegmen longer than penis, partially enclosing penis, lobes well separated with small triangular extensions on the outside near apex (Fig. 9). Variation Some specimens are uniformly reddish-yellow whereas the majority are darker towards the front. Etymology Named after my granddaughter Emma, whose father collected many of the specimens. Notes A relatively large species from tropical north and eastern Australia. Typical specimens are oval, reddish and with the colour becoming noticeably darker towards the front. Broadly sympatric with the other two large reddish-yellow, elongate-oval species with concave hind margins to the metacoxal plates, S. kKayfae and S. tindaleensis. Scirtes tindaleensis is more elongate and usually darker in colour; S. kaytae has upturned frons and asymmetric antennal segments and usually has dark regions towards the sides of the elytra. The male genitalia of all three species are very distinctive and even if only the tips are visible readily identify the species. Additional specimens examined Northern Territory. |, Black Point Coburg Pen., 11 09S 132 O9E, 15 — 23/2/77, T. A.Weir, ANIC; 1, 144 C. H. S. WATTS Berry Springs 30 km SSE Darwin 12 41S 130 58E, 11/11/72, at light, E. Britton, ANIC; 3, Casurina Beach 10km NNE Darwin 12 21S 130 42E, 22/10/72, E. Britton, ANIC; 4, Horn Isl, Pellew Group, 22 — 28 Feb.1968, B. Cantrell, UQIC Reg# 53670 — 1-2-3; 3, Howard Springs 24km S Darwin 12 28S 131 03E, 10/11/72, at light, E. Britton, ANIC; 3, Jim Jim Creek 19km WSW Mt Cahill 12 57S 132 33E, 24/10/72, at light, E. Britton, ANIC; 1, Kakadu np, Baroalba Springs 12 48’S 132 49E, 14.11.91, D.V. Wells, NTM; 1, Kakadu np Nourlangie Camp, 17-18/11/79, at light, M. B. Malipatil, NTM; 1, Katherine, 25/1/73, T. Angeles & N. Forresyter, NTM; 4, Lee Point Darwin, 7/3/67, M. S. Upton, ANIC; 1, 4mi SW Lee Point Darwin, 6/3/67, M. S. Upton, ANIC; 2,15km E by N Mt Cahill 12 49S 132 S51E, 29/ 10/72, at light, E. Britton, ANIC; 2, 16km E by N of Mt Cahill, 12 50’S 132 541E, 27/11/74, T. Weir & T. Angelas, NTM; 1, Smith Point. 3.8.82, C. Wilson & S. Collins, NTM; 2, Thorak Reserve via Berrimah, 28/11/74, A. Allwood, ex light trap, NTM. Queensland. 5, Annan R 3kmW by S Black Mountain 15 41S 145 12E, 26 — 17/4/81, A. Calder, ANIC; 1, Babinda, 9/37, ANIC; 1, Bucasia, K.J. Sandry, 20/2/93, ANIC; 1, Cairns, 12/1/35, ANIC; 2, ditto, 2/50, J. G. Brooks, ANIC; 1, Ditto, NMV; 1, 1km W Cooktown 15 28S 145 15E, 12 — 13/5/81, A. Calder, ANIC; 1, Cairns, 6/1966, A. MacQueen, UQIC Reg# 53727; 1, Edge Hill Cairns, 23 — 24/2/65, at light, J.G. Brooks, ANIC; 1, Ellis Beach 25km NNW Cairns 16 44S 145 39E, 19/5/76, E.B. Britton, ANIC; 4, Cow Bay N of Daintree, Jan-Feb 81, I. C. Cunningham, QPIM; Christmas Creek 15km W of Fairviews via Laura, 26 — 27/6/75, G. B. Monteith, QM; 1, Jardine R. Crossing 29km S Bamiga, 5/9/85, light trap, E. N. Marks, ANIC; 1, Innisfail, 25.12.59,V. Skablum, UQIC Reg# 53721; 1, Iron Range 11 — 17/5/68, G.Monteith, UQIC Reg# 53694; 2, ditto except 16 — 23.11.65,UQIC Reg# 53686/7; 1, Granite Gorge via Marreeba, 21/1/89, R I Storey, at light, QPIM; 1, Iron Range, 26 — 31/10/99, Wood, Dunn & Hasenpusch, QPIM; 1, Jullatten, 18/11/86, Malaise trap, A. Walford- Huggins, QPIM; 1, Lankelly Creek, MclIlwraith Rngs. nr. Coen, 28-32/10/69, B. Cantrell, UQIC Reg# 53661; 2, Marina Plains via Musgrave, 17/11/82, Storey, Brown & Jacobson, QPIM; 1, Mossman, 11/1/84, at light, J. D. Brown, QPIM; 1, Ikm S Mt Cook 15 30S 145 16E, 13/10/80, T.Weir, ANIC; 1, 2mi. SW Mt Inkerman 19 45S 14730E, 11/12/68, S. Misko, ANIC; 4, 9km ENE Mt Tozer 12 43S 143 17E, 5 — 10/ 7/ 86, T. Weir & A. Calder, ANIC; 3,11km ENE Mt Tozer 12 43S 143 18E, 11- 16/ 7/86, T. Weir & A. Calder, ANIC; 1, 3 km NE Mt Webb 15 03 S 145 09E, 3/5/81, A. Calder, ANIC; 1, 2km NE by E Mt Tozer 12 44S 143 13E, 1/7/86, A. Calder, ANIC; 2, 3km ENE Mt Tozer 12 44S 143 14E, 28/6/86, T. Weir & A. Calder, ANIC; 2, N Queensland, SAMA; 1,llkm WSW Petford, 21/8/1/88, at light, R. Storey, QPIM; 1, 32km S Ravenshoe 17 38S 145 29E, 13/2/66, K Hyde, ANIC; 2, Rocky River 10mi. N.17.2.46, G. Monteith, UQIC Reg# 53653/56; 7,15km WNW South Johnstone, 10/12/85, at light, Fay & Halfpapp, QPIM; |, South Johnstone, 12/79, at light, B. Pinese, QPIM; 2, Split Rock 14km S of Laura, 23 — 26/6/75, G. B. Monteith, QM; 1, Yorkeys Knob, 17/8/63, B. V. Timms, UQIC Reg# 53723; 3, 9km SE Yeppoon, 20-30/10/75, I.F.B.Commom, ANIC. Scirtes exoletus Waterhouse, 1880 (Fig. 10) Type Holotype female, “W Austral” “Scirtes exoletus (Type) C. Waterh.”, NHM. Seen. Description (number examined, 59) Habitus. Length 3.5 — 5.0 mm., relatively flat, oval. Head. Yellowish, rear and Y-shaped suture often brown; antennae light to dark reddish-yellow, distal portion of each segment lighter. Small, width between eyes about 2.4x dorsal width of eye. Evenly punctate, punctures relative large, each with a moderately sized golden seta. Frons with sides slightly diverging, front edge straight or very weakly concave, edges weakly beaded. Segment | of antenna large, curved; segment 2 smaller, barrel- shaped; segment 3 smaller and narrower; segments 4 - 10 long, narrow, cylindrical; segment 11 a little longer than segment 10, all segments setose. Pronotum. Yellowish with brown pattern. Short, broad. Evenly and moderately to quite strongly punctate, each puncture with a moderately long golden seta. Hind angles obtuse, front edge sinuate, sides weakly beaded. Scutellum. Yellowish, usually lighter than elytra, slightly longer than wide, lateral sides convex. Elytron. Yellowish to light reddish-yellow, often a little darker near base and side. Side weakly and narrowly flanged. Moderately to strongly and evenly punctate, each puncture with a yellow seta. Epipleuron relatively wide in front quarter, becoming narrower over rest of elytron, front portion weakly to moderately concave. Ventral surface. Light reddish-yellow. Pronotal process very narrow between procoxae, apical portion not much wider. Mesosternum with broad, flat, diamond shaped area in midline in front to receive prosternal process; rear tip reaching just reaching front of mesocoxae. Front extension of metasternum in midline, short, widely triangular; REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 145 rear midline extension of metasternum about twice as wide as long; approximately equilateral. Metacoxal plate about as long as wide, with anterio- lateral corner extending some way along mesosternum; hind edge straight or sightly convex, sloping towards midline; sides strongly beaded; posteriolateral angles rounded. Metatrochanter small, about 2.0x as long as wide. Metafemur moderately swollen, widest about middle, small notch on hind edge near apex. Dorsal metatibial spine relatively short, about twice size of ventral spine and about half length of segment 1 of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites slightly rugose with moderate punctures and fine reticulation, more pronounced towards rear; apex of apical ventrite truncated. Male Little external difference between the sexes. Basal piece of penis broad, rounded; trigonium about same length, triangular, apex rounded; single parameroid robust, simple, as long as trigonium (Fig.10). Tegmen about half-length of penis, with broad lobes, not or only slightly separated near apex, partially wrapping around penis, apical edges with short spines (Fig. 10). Notes A relatively large, flat, distinctive species, recognised by its yellow/grey colour with clear darker mottlings on the pronotum and head and a relatively short metatibial spine. It shares with S. macroconcolor and S. interstinctus the shallowly grooved elytral epipleura and straight edges to the metacoxal plates, but S. macroconcolor is uniformly coloured with a more strongly flanged elytra and S. interstinctus has dark stripes on the elytra, flanged elytra, usually stronger and less dense punctures and the hind edges of the metacoxal plates do not slope inwards as they do in S. exo/etus. The male genitalia of the three species are distinct. Scirtes exoletus has an unusually wide distribution, occurring in all States except South Australia. I have collected its large dark larvae from submerged leaf litter at the edge of drying pools in the Pilbara and from among emergent vegetation in farm dams in Victoria and northern Tasmania. Specimens examined Queensland. |, Boar Pocket rd. 8km N of Gillies Hwy., 21/2/70, J. G. Brooks, ANIC; 1, Colossum Creek, 10mi. S Miriam Vale, 20/12/66, B. Cantrell, UQIC Reg# 53665; 3, Davies Creek 22km WSW Mareeba, 6/11/84, Storey & Halfpapp, QPIM; 2, Danbulla via Yungaburra, 13/11/92, at light, Storey, De Faveri & Huwer, QPIM; 1, 7km N Hope Vale Mission, 4/10/80, T. Weir, ANIC; 1, Kenilworth State Forest, 1/4/69, B. Cantrell, UQIC Reg# 53660; 1, 7.5km NNW Kuranda, 20/11/82, Storey & Halfpapp, QPIM; 1, 13km W Kuranda, 7/12/82, J.T. Doyen, ANIC; 1, 8km W Kuranda, 28/12/86, H & A Howden, malaise trap, QPIM; 1, Lake Eacham, 16/12/82, J. T. Doyen, ANIC; 1, Mary Creek 16 33S 145 12E, 5/12/68, at light, Britton & Misko, ANIC; 1, Mt Tambourine, A. M. Lea, SAMA.;1, 3km ENE Mt Tozer, 28/6/86, D. H. Colless, malaise trap, ANIC; 1, 7km NE Tolga, 2/87, Storey, & De Faveri, QPIM; 2, Whitfield rd. 22km from Cairns, 21/10/71, J. G. Brooks, ANIC. New South Wales. 2, Collector, 2/61, C. Watts, SAMA; 1, Epping, 4/3/87, malaise trap, I. Buddie, ANIC; 1, Harrington, 8/9/83, G. Williams, ANIC; Tooloom Plateau Via Woodenbong, 30 — 31/12/66, G. Monteith, UQIC Reg# 53714; 1,Wingham Scrub 31 52S 152 22E, at light, 3/1/70, Britton, Holloway & Misko, ANIC. Northern Territory. |, McArthur River 16,47S 135 45E 14km S by W Cape Crawford, 25/10/75, M. S. Upton, ANIC; 1, Ditto, 6/11/73, ANIC; 1, Nourlangie Creek 8km E of Mt Cahill, 7/5/75, A. Allwood & T. Angeles, NTM. Tasmania. 4, 4 km W Port Latta, 27/11/00, C. Watts, SAMA. Victoria. 8, Healsville, 12/68, C. Watts, SAMA; 1, Thomson River Bells clearing, 8 March 1970, MV light, NMV; 2, Tullamarine, 4/9/75, SAMA; 1, 2 km E Warburton, 14/1/97, C. Watts, SAMA. Western Australia. 3, Crossing Pool Millstream 21 35S 117 O4E, E. B. Britton, ANIC; 1, Deep Reach Millstream 21 35S 117 04E, at light, E. B. Britton, ANIC; 1, Millstream Fitzroy Crossing area, 17/3/83, K. & E. Carnaby, ANIC; 5, Hammersly Range, W. D. Dodd, SAMA; 9, ditto, at light, 30/10/70, E. Britton, ANIC; 5, Millstream, Coll Ranger, Summer 2000, SAMA; 2, | km N Millstream, | — 4/71, M. S. Upton, ANIC. Scirtes interstinctus sp. nov. (Fig. 11) Types Holotype male: “15.30S 145.16E 5 km SEbyS Cooktown QLD 19 May 1977”. F.B. Common & E.D. Edwards”, ANIC. Paratypes 12; 4, “Dividing Range 15 km W of Captain Billy Creek Cape York Pen, N.Qld. 142 45E 11 40S 4-9 vii. 1975 G. B. Monteith”, QM; 1,” AUSTRALIA n QLD Davies Ck 22 km WSW of Mareeba Malaise T 2. xii. 1984 Storey & Titmarsh”, QPIM; 1, Ditto, 2. x.-6 x1 1984 Storey & Halfpapp”, QPIM; 1, Ditto “2 xil — 21.xii 1984 Storey & Brown”, QPIM; 1, “15.14S 145, O7E 7 Km N of Hope Vale Mission QLD 4 Oct 1980 T. Weir’, ANIC; 1, “8 Km W 146 C. H. S. WATTS Kuranda NQ 28 Dec.1986 H & A Howden Malaise trap”, ANIC; 1, “12.448 143.14E 3 K ENE Mt Tozer 28 June — 4 July 1986 D. H. Colless Malaise trap”, ANIC; 1, “NEQ 17 19S 145 37E Peeramon Scrub 750m 9 Dec 1995 G. Monteith Pyrethrum trees”, QM. Description (number examined, 13) Habitus. Length 3.8 — 4.7 mm., relatively flat, oval. Head. Reddish-yellow with darker markings; antenna reddish-yellow. Small, width between eyes about 2.5x dorsal width of eye. Evenly punctate, punctures relative large, each with a moderate sized golden seta. Frons with sides slightly diverging, front edge straight or very weakly concave, edges weakly beaded. Segment | of antenna large, curved; segment 2 smaller, barrel-shaped; segment 3 smaller and narrower; segments 4 - 10 long, narrow, cylindrical; segment 1] about same length as segment 10, all segments setose. Pronotum. Light reddish-yellow with dark brown pattern. Short, broad. Punctures large, relatively shallow, moderately dense, each puncture with a moderately long golden seta. Hind angles obtuse, front edge sinuate, sides weakly beaded. Scutellum. Yellowish, usually lighter than elytra, slightly wider than long, lateral sides convex. Elytron. Light reddish-yellow, with darker stripes sutural region narrowly yellow. Side moderately to quite strongly flanged in front half; evenly punctate with relatively large shallow punctures, each puncture with a yellow seta. Epipleuron relatively wide in front quarter becoming narrower over rest of elytron, front portion weakly to moderately concave. Ventral surface. Light reddish-yellow. Pronotal process very narrow between procoxae, apical portion not much wider. Mesosternum with broad, flat, diamond shaped area in midline in front to receive prosternal process; rear tip reaching past front of mesocoxae. Front extension of metasternum in midline, short, widely triangular; rear midline extension of metasternum about twice as wide as long. Metacoxal plate about as long as wide, with anteriolateral corner extending some way along metasternum; hind edge sightly sinuate; sides beaded; posteriolateral angles rounded. Metatro- chanter small, elongate, about 2.5x as long as wide. Metafemur moderately swollen, widest about middle, small notch on hind edge near apex. Dorsal metatibial spine about twice size of ventral spine and about half-length of segment 1 of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites slightly rugose with moderate punctures and fine reticulation, more pronounced towards rear; apex of apical ventrite truncated. Male Little external difference between the sexes. Basal piece of penis short, broad, rounded; trigonium about twice as long as broad, spatulate with raised ridge at base; single parameroid robust, a little longer than half length of trigonium, apex hooked (Fig. 11). Tegmen shorter then penis, lobes short, slightly asymmetrical, apical edges with short spines, strongly wrapped around penis (Fig. 11). Variation The extent of the brown stripes on the elytra is variable; in some they are reduced to the base and some scattered markings elsewhere on elytra. The colour of the antennal segments varies from nearly uniform yellowish to quite dark with much lighter distal portions. Etvmology Latin. “Interstinctus” — variegated, a reference to the dorsal colour. Notes A relatively large species recognised by its distinct darker mottlings on the pronotum and head and short linear markings on the elytra, relatively large shallow punctures and flanged elytra. It shares with S. exoletus the shallowly grooved elytral epipleurae, patterned head and pronotum and straight edges to the metacoxal plates but has linear markings on the elytra, more strongly flanged elytra, generally larger punctures and quite different male genitalia. Most specimens have been captured in Malaise traps, none at light, which is a different pattern than other Scirtes which may indicate a somewhat different natural history. Scirtes kaytae sp. nov. (Fig. 12) Types Holotype male, “Qld. Bushland Beach 20km N Townsville A. J. Watts 15 — 20/3/98”, SAMA. Paratypes 37; 21, “Qld. Bushland Beach 20km N Townsville A. J. Watts 23 — 30/12/97", SAMA; 10, ditto, “at light 16 — 18 Jan 1998”, SAMA; 3, ditto 6 — 11/98, SAMA; 2, ditto, 26-29 Feb1998, SAMA; 1, ditto, 28/3/98, SAMA. Description (number examined, 249) Habitus. Length 3.9 — 5.5 mm., relatively flat, oval. Head. Reddish-yellow. Small, width between eyes about 3x dorsal width of eye. Strongly, densely and REVISION OF AUSTRALIAN SC/RTES ILLIGER AND ORA CLARK 147 evenly punctate. Frons with front angles bulging outwards, front edge straight or very weakly concave, front bent upwards, edge beaded. Segment 1 of antenna large, barrel-shaped; segment 2 about half as long, oval; segment 3 as long as second but narrower; segments 4 — 10 long, narrow, tending to be more expanded on front edge, particularly central ones; segment 11 and segment 10 subequal. Pronotum. Reddish-yellow; short, broad. Evenly and quite densely punctate, punctures confluent at sides, each with a long golden seta. Hind angles obtuse, front edge strongly sinuate, sides weakly beaded. Seutellum. Reddish-yellow; about as long as wide, lateral sides weakly convex. Elytron. Reddish-yellow with diffuse darker areas towards sides and front. Quite strongly and evenly punctate, each puncture with a yellow seta although often abraded off. Weakly flanged towards front. Epipleuron yellow, relatively wide in front quarter, evenly becoming much narrower over rest of elytron. Ventral surface. Reddish-yellow, often with diffuse darker areas. Pronotal process very narrow between procoxae, apical portion not much wider, strongly keeled. Mesosternum with narrowly elongate groove for reception of pronotal process; tip reaching to level of mesocoxae. Rear triangular midline extension of metasternum approximately same size as front extension; about twice as wide as long, edge beaded. Metacoxal plate broader than long with anteriolateral corner extending some way along metasternum; hind edge strongly concave; sides beaded; posterior-lateral angles rounded. Metatrochanter very small, apex pointed. Metafemur greatly swollen, widest just before middle, quite strongly indented on hind edge near apex. Dorsal metatibial spine relatively long more than twice size of ventral spine and about two thirds length of segment | of metatarsus. Segment | of metatarsus a littke longer than other segments combined. Ventrites with punctures small, shallow; reticulation moderate fine, more pronounced towards rear; apex of apical ventrite truncated or concave. Male Basal piece of penis elongate oval; trigonium about as long as basal piece, a small sharp hook at apex; single parameroid a little shorter than trigonium, narrow, slightly curved towards tip which is sharply pointed (Fig.12). Tegmen about as long as penis, lobes narrow, well separated, with row of quite strong spines on outside edge (Fig.12). Front of frons more strongly upturned; antenna stouter with expansions on inside of antennal segments greater than in female. Variation There is considerable variation in the strength of the colour pattern on the elytra from almost uniformly reddish-yellow to reddish yellow with almost black markings on shoulders and each elytron with a broad dark stripe near but not quite reaching the sides. The strength of the shovel-like upturned frons of the male (and weakly in some females) is quite variable as are the internal expansions of the antennal segments. Etymology Named after my granddaughter Kayt whose father collected many of the specimens. Notes A large, common species in tropical north and eastern Australia, reddish but usually with noticeably darker areas on the elytra. The males are very distinctive with stout antennae with the inner portions of the individual segments enlarged slightly on the insides and frons with front edge projecting forwards and upwards. These characters are much less obvious in the females. The male genitalia are distinctive for the spines on the outer margin of the tegmen and hooked tip to the penis which is often visible in preserved specimens. Broadly sympatric with the two other large reddish-yellow oval species with concave hind margins to the metacoxal plates, S. emmaae and S. tindaleensis. Both of these are more uniform in colour without the lateral darker markings on the elytra and lack the asymmetric antennal segments and the upturned frons of S. kay/ae as well as clear differences in the male genitalia. Could also be confused with the rarer S. ruforotundus, a more rounded, dark reddish species, often with the disc of the elytra lighter and with normal frons and thin antennae. The male genitalia of S. ruforotundus are very different, and never protrude from the abdomen. The large black larvae occur among emergent vegetation in seasonal swamps. Specimens examined Queensland. 1, Archers Creek Mt Garnet Rd., 28/12/64, J. G. Brooks, ANIC; 1, Ayr 19 35S 147 24E, 30/11/70, W. B. Muir, ANIC; 1, Big Mitchell Ck. Mareeba-Molloy Road, 4/5/67, D. H. Colless, ANIC; 1, Bundaberg 24 51S 152 21E, 14/3/72, Frauca, ANIC; 1, Bundaberg, 20/2/72, H. Frauca, ANIC; 22, ditto, 14/4/63, C. Watts, SAMA; 1, Bundaberg, SAMA; 5, ditto, 25 — 26/ 3/ 84, K. H. Halfpapp, at light, QPIM; 5, Cairns, 2/50, G. Brooks, ANIC; 1, Calliope River 23km SE Gladstone 23 50S 152 13E, 23/1/70, light trap, S. Misko, ANIC; 2, Cape Pallarenda 19 14S 146 46E, 14 — 17/1/74, at light, R. A.Barrett, ANIC; 1, Carr Creek 18km NNW 148 C. H. S. WATTS 16 17 Figs 12-17. Dorsal views of tegmen (left) and penis (right). Lines = 0.5 mm. 12. Scirtes kaytae sp. nov. 13. S. macroconcolor sp. nov. 14. S. microrotundus sp. nov.15. S. ruforotundus sp. nov.16. S. tindaleensis sp. nov. 17. S. alastairi sp. nov. REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 149 Mareeba, 21/5/80, I.D. Nauman, ANIC; 10, Cemetry Point Veron via Maryborough, 24/12/70, I. F. B. Common, ANIC; 4, Gordonvale, 1/15/99, A. Podlussany, HUNG; |, Gin Gin, 2/4/62, C. Watts, SAMA; 5, Giru,18/4/81, B.B. Lowery, at light, ANIC; 1, Kalpower Crossing 75km NW Laura, 2/4/83, at light, R. 1. Storey, QPIM; 1: 1, 70km SW Greenvale, at light, 1-10/ 3/95, A. J. Watts, SAMA; 1, Mango Island off Millaroo, 13/3/74, J. H. Barrett, QPIM; 5, Marina Plains via Musgrave, 10/5/83, Storey & Brown, QPIM; 2, N Queensland, Blackburn Coll, SAMA; 22, 30km N Marlborough, at light, 24/11/81 Hangay, Herozeg & Vojnite, HUNG; 1, Mossman, 8/1/84, J. Brown, at light, QPIM; 6, Normanton, 4/5/63, at light, P. F. Aitkin & N.B. Tindale, SAMA; 2, Old Laura Station 28km N Laura, 3/4/83, at light, R.I. Storey, QPIM: 2, Ingham, 16/2/60, K. I. Harley, ANIC; 5, ditto, 6/3/84, K. H. Halfpapp, at light, QPIM; 1, ditto, SAMA; 1, Iron Range, 5/71, J. Brooks, ANIC; 16, Lansdown Station 7km S of Woodstock 19 40S 146 51E, 16.1.74, at light, RABarrett, ANIC; 2, Pistol Gap Byfield 22 50S 150 40E, 10/1/70, at light, Britton Holloway & Misko, dry sclerophyll, ANIC; 3, 2km S Ravenshoe 17 38S 145 2E, 13/2/66, K. Hyde, ANIC; 1, Tolga, 2 — 3/80, N. Gough & J. D. Brown, QPIM; 2, 7km N Tolga, 3/88, Storey & De Favers, QPIM; 1, ditto, 1/88, QPIM; 5, Townsville, F.H. Taylor, ANIC; 4, ditto, 12/3/58, K. L. Harley, ANIC; 9, ditto, 1/16/68, P. Ferrar, ANIC; 2, Stuart Range, | — 2/27, Hale & Tindale, SAMA; 1, Waterfall Creek 20ml N. of Rollingstone, 29.3.73, A. Allwood & T. Angeles, NTM; 2, Woodstock, 3/54, A. J. Brooks, ANIC; 1, Yeppoon, 14 — 18/12/64, I. F. Common & M.S. Upton, ANIC; 1, Yeppoon, 30/1/70, I. F.B. Common, ANIC. Northern Territory. 7, Bessie Springs 8km ESE Cape Crawford 16 40S 135 51 E, 12/4/76, at light, J. E. Feehan, ANIC; 7, Batten Point 30km NE by E Borroloola 15 548 136 32E, at light, 18/4/76, J. E. Feehan, ANIC; |, 22km WSW Borroloola, 16/4/76, at light, J.E. Feehan, ANIC; 1, Edge Hill, 4/64, J Brooks, ANIC; 1, Cahills Crossing East Alligator River 12 26S 132 S58E, 29/5/73, at light, E. G. Matthews, ANIC; 1, 5km NNW Cahills Crossing East Alligator river 12 23S 132 57E, 28/5/73, E. G. Matthews, ANIC; 1, Darwin River 16km SW by S$ of Noonamah 12 44S 130 58E, 16/5/74, T. Angeles & W. Mollah, NTM; 2, Jasper Gorge 54km NW of Victoria River Downs 16 02S 130 41E, 30.4.74, T. Weir & T. Angeles, NTM; 1, 10km SW Jabiru, 29/1/99, C. Watts, SAMA; 3, Katherine, at light, 6 — 10/2/68, J. A. L. Watson ANIC; 1, Katherine, 23/1/71, T. Weir & A. Allwood, NTM; 2, Koongarra 12 52S 132 SOE, 6 — 10/3/73, M. S. Upton, ANIC; 1, Lee Point Darwin, 8/3/67, M. S.Upton, ANIC; 1, Melville Isl., at light, 4/2/68, Matthews, ANIC; 5, McArthur River 14km SW Cape Crawford 16 47S 135 45E, 11/4/76, J. E. Feehan, ANIC; 2, Magela Creek 1km NNW Mudginberry HS 12 36S 132 52E, 25/5/73, Matthews & Upton, ANIC; 2, McArthur River 14km SW of Cape Crawford 16 47S 135 45E, 11/4/76, J. F. Feehan, ANIC; 1, October Creek on Borroloola Road, 7/4/76, T. Weir, NTM; 3, Roper River, 6/4/76, T. Weir, NTM; 1, Smith Point, 23/2/81, A. Allwood, NTM; 1, Tortilla flats, 3.3.82, J, Waldock, NTM; 1, Tindale 14 31S 132 22E, 20/12/67, W. J. M. Vestjens, ANIC; U.A.R. 21/2/67, C.S.Li, NTM; 1, Victoria River crossing 15 36S 131 O7E, 29/4/74, T. Weir & T. Angeles, NTM; 1, Wildman River Cashew project, 3/1/89, Malipatil & Houston, QPIM. Western Australia. 2, 8km S Cape Bertholet West Kimberley 17 19S 122 10E, 21/4/77, D. H. Colless, ANIC; 5, 3km S Coulomb Pt. West Kimberley 17 32 122 09E, 20/4/77, D. H. Colless, ANIC; 2, Fitzroy River, 11/4/84, at light, K. & E. Carnaby, ANIC; Kununurra, 27/12/82, R. I. Storey, QPIM; 6, Kununurra, 17-21/ 2/ 68, E. Matthews, ANIC; 1, 6km W Martin’s Well West Kimberley 16.08S 122 48E, 25/4/77, D. H. Colless, ANIC; 3, Ord River Valley Kimberley Res. Station, 9/3/82, E.S.C. Smith, ANIC. Scirtes macroconcolor sp. nov. (Fig.13) Types Holotype male; “14 49S 126 49E Carson escarpment W.A. 9 — 15 Aug.1975 I. F. Common and M.S. Upton”, ANIC. Paratypes 7; 1 slide, as for holotype, SAMA; I, “Bessie Springs 16 40S 135 51E 8 km ESE Cape Crawford NT. 26 Oct. 1975 M. S. Upton”, ANIC; 1, “nr. Katherine, NT 21 May 1992 P. S. Cranston & P. J. Gullan coll.”, ANIC; 1,°15 02S 126 55E Drysdale River, W.A. 3— 8 Aug.1975 I. F. B. Common and M. S. Upton”, ANIC; 3, “N.T. U.D.P. Falls 18 — 19 Jul 1980 M.V. Light M.B. Malipatill” NTM. Description (number examined, 8) Habitus. Length 4.5 — 4.8 mm., relatively flat, oval. Head. Light reddish-yellow. Eyes large, width between eyes about 2.3x dorsal width of eye. Moderately and evenly punctate. Frons with sides moderately diverging, front edge weakly concave, front corners slightly downturned, edges weakly beaded. Segment | of antenna large, barrel-shaped; segment 2 about two-thirds length of segment 1, oval; segment 3 about half length of segment 2, 150 C. H. S. WATTS narrower; segments 4 — 10 long, relatively wide, parallel sided, becoming progressively flatter; segment 1] about same size as segment 10. Pronotum. Light reddish-yellow. Short, broad. Evenly and moderately to quite strongly punctate, each puncture with a moderately long golden seta. Hind angles obtuse, front edge strongly sinuate, sides weakly beaded. Scutellum. Light reddish-yellow. A little longer than wide, lateral sides convex. Elytron. Light reddish-yellow to reddish-yellow, sutural region narrowly lighter. Sides quite broadly flanged, more strongly in middle. Moderately to strongly and evenly punctate, each puncture with a short yellow seta. Epipleuron relatively wide in front quarter becoming narrower over rest of elytron, front portion shallowly grooved. Ventral surface. Uniformly light reddish-yellow. Pronotal process very narrow between procoxae, apical portion a little wider. Mesosternum with diamond shaped groove in midline in front for reception of pronotal process; rear tip reaching just to front of mesocoxae. Front extension of metasternum in midline, small, short, triangular bordered behind; rear midline extension of metasternum about as wide as long; approximately equilateral, tip reaching to about middle of metacoxal plate. Metacoxal plate about as wide as long, with anterio-lateral corner extending some way along metasternum; hind edge straight, sloping slightly to midline; sides very weakly beaded; posteriolateral angles rounded. Metatrochanter small about 2.0x as long as wide. Metafemur greatly swollen, widest about middle, small notch on hind edge near apex. Dorsal metatibial spine about twice size of ventral spine and about two-thirds length of segment | of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites slightly rugose, moderately punctate, reticulation fine, more pronounced towards rear; apex of apical ventrite truncated. Male Little external difference between sexes. Basal piece of penis large, oval; trigonium shorter, squatly triangular, apex rounded; parameroid large, thumb- like, broad in lateral view, narrow in dorsal view (Fig. 13). Tegmen a bit shorter than penis, lobes short, conical, well separated (Fig. 13). Variation Some specimens have diffuse darker areas on the elytra and vague darker patterning on the pronotum. Etymology Latin. “Macro” — large; “concolor” — uniform colour. Notes A moderately sized species with uniform colour, straight hind edges to the metacoxal plates, weakly grooved epipleuron and a noticeably flanged elytra. Scirtes microrotundus sp. nov. (Fig. 14) Types Holotype Male; “Mossman Gorge, N. Qld. 23 Apr. 1967 D. H. Colless”, ANIC. Paratypes 3 slides, as for holotype, 2 ANIC, | SAMA. Description (number examined, 4) Habitus. Length 2.0 mm., flat, round. Head. Dark brown, Small, width between eyes about 2.7 x dorsal width of eye. Moderately and evenly punctate, each puncture with a moderate sized pale seta. Frons with sides strongly diverging, front edge concave, edges beaded. Antenna reddish- yellow, lighter towards base; segment | of antenna large, barrel-shaped; segment 2, barrel-shaped about three quarters size of segment | in both width and length; segment 3 smaller about two thirds length of segment 2 and narrower; segments 4 — 10 long, cylindrical; becoming a little flatter apically, segment! a little longer than segment 10. Pronotum. Dark chestnut, extreme margins lighter. Short, broad. Evenly and moderately punctate, each puncture with a moderately long golden seta. Front angles weakly extended; hind angles obtuse; sides weakly beaded, weakly upturned. Scutellum. Dark reddish-yellow; sides broad, slightly wider than long, lateral sides weakly convex. Elytron. Dark reddish-yellow. Sides weakly flanged towards front. Moderately to strongly and evenly punctate, each puncture with a short yellow seta. Epipleuron relatively wide in front quarter becoming narrower over rest of elytron, front portion widely and shallowly grooved. Ventral surface. Dark reddish-yellow, appendages tending lighter. Pronotal process very narrow. Mesosternum with short, shallow, relatively broad, triangular groove for reception of pronotal process; tip just reaching to level of mesocoxae. Front extension of metasternum relatively small, broadly triangular, rear midline extension of mesosternum relatively long, about 1.5x as wide as long. Metacoxal plate wider than long, with anterio-lateral corner extending some way along metasternum; hind edge straight or weakly sinuate, sloping towards midline; midline of coxae short, shorter than length of metasternal extension; sides beaded; posteriolateral angles rounded. Metatrochanter REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 151 small, elongate, about 2.4x as long as wide. Metafemur moderately swollen, widest about middle. Dorsal metatibial spine moderately long about twice size of ventral spine and about two- thirds length of segment | of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites with punctures small, shallow; reticulation moderate, fine, more pronounced towards rear; apex of apical ventrite truncated or weakly concave. Male Only males known. Basal piece of penis, oval: trigonium formed of two long thin pieces one broader than the other; single parameroid longer and thinner than trigonium lobes, apex pointed, small bifid structure near its base (Fig. 14). Tegmen about half length of penis, lobes broad, well separated (Fig. 14). Variation Little variation in the four known specimens. Etymology Greek. “Mikros” — small. round. Latin “rotundus” — Notes A small dark, almost round species with short metacoxal plates, weakly concave epipleura and distinctive male genitalia. Scirtes ruforotundus sp. nov. (Figs 2, 15) Types Holotype male, “15.30S 145 16E 1 km SE of Mt Cook Qld 13 Oct. 1980 T. Weir’, ANIC. Paratypes 28; 1, “Brandy Cr 8 mi. NE Proserpine Q (20 20S 148 41E) 1.xii.1968 at light Britton & Misko”, ANIC; 1, ‘Cairns 5/49 J. G. Brooks’, ANIC: 1, ‘16 03S to 16.058 QLD145 28E Cape Tribulation 21 — 28/3/84 A. Calder & T. Weir’, ANIC; 1 slide, Cape Tribulation NQ 14 — 17 Jul. 1982 S & J Peck coll, SAMA; 1, “Cardstone Qld 20/11/66 K. Hyde”, ANIC; 1, “Cooper Creek, I8ml. N. of Daintree River, N. Qld. 21 — 22.vi.1969 G.B.Monteith” “UQIC Reg# 53713”, UQIC; 1, “Crystal Ck. Q., 23 mi. SSE Ingham 19 58S 146 16E, 9/12/68 at light Britton & Misko”, ANIC; 2, “Gap Ck., 6ml. N of Bloomfield R. N. Qld. 13.xi. 1965 G. Monteith” “UQIC Reg# 53643/4”. UQIC; 1, “15 12S 143 52E Hann R. 73 km NW by W Laura Qld. 27 June 1986 T. Weir & A. Calder”, ANIC; 3, “15 16S 144 59E 14 km W by N of Hope Vale Mission Qld. 8 — 10 Oct 1980 T. Weir”, ANIC; 1, “15.14S 145.07 E7 km N of Hope Vale Mission Qld. 4 Oct 1980 T. Weir”, ANIC; 1, “Iron Range Cape York Pen. N. Qld. 11 — 17 May 1968 G. Monteith” “UQIC Reg# 53693”, UQIC; 2, “Iron Range Cape York Pen. N. Qld. 16 — 23.x11.1965 G. Monteith” “UQIC Reg# 53690/1”. UQIC; 1, “Mackay”, SAMA; 1, “Mitchell River Settlement, Qld. 4 iv.70 A. L. Dyce (From large open cavity 6’ above ground level in mango tree)” ANIC; 2, “9 km SW Madang PNG 1.1i.1988 bamboo internode B 18 R. Kitching”, ANIC; 1, “W of KOWI Madang PMG 3.1i1.1989 TH24/R Kitching”, ANIC; 3, “Miller’s Crossing, 30mls N. of Cooktown N.QId. 24-25.xi.1965, G. Monteith” UQIC Reg# 5384/4/5”; 2,15 30S 145 16E 1 km SE Mt Cook Qld 13 Oct 1980, T. Weir”, SAMA; 1, “N. Queensland”, SAMA; 3, “15 05S 145 07E Mt Webb Nat Pk. QLD 28 — 30 Sept 1980 T. Weir”, ANIC; 1, “12 44S 143 14E 3 km ENE Mt Tozer 28 Jun — 4 Jul. 1986, T. Weir & A. Calder” ANIC; 1, “15.038 145. O9E 3 km NE of Mt Webb QLD, | — 3 Oct 1980 T. Weir”, ANIC; 1, “Silver Plains Homestead Cape York Pen. Q 24 Dec.1962 J. L. Wassell”, ANIC; 1, “AUSTRALIA. n Qld.15 km NW of South Johnstone light trap 17. x. 1986, Fay & Halfpapp, “QPIM; 1,” “The Boulders” via Babinda, N. Qld 15.xi.1969 B. Cantrell.”, “UQIC Reg# 53669”, UQIC; 1, “Upper Mulgrave River N. Qld. 1-3.xi1.1965. Cent Qld. B. Cantrell”, “UQIC Reg# 53665”, UQIC. Description (number examined, 29) Habitus. Length 3.2 — 4.7 mm., relatively flat, broadly oval. Head. Dark reddish-yellow to nearly black, antenna light reddish-yellow to reddish-yellow. Small, width between eyes 2.2x dorsal width of eye. Moderately and evenly punctate. Frons with sides concave, front edge quite strongly concave, edges beaded. Segment | of antenna large, barrel-shaped; segment 2 smaller, cylindrical; segment 3 shorter, narrower; segments 4 — 10 long, relatively broad, front edge slightly concave, segment four longest; segment 11 a little longer than segment 10. Pronotum. Dark reddish- yellow to nearly black. Short, broad. Evenly and moderately densely punctate, each puncture with a moderately long golden seta. Hind angles obtuse, front edge sinuate, sides beaded. Scutellum. Dark reddish-yellow. Sides approx- imately equal length, lateral two weakly convex. Strongly and evenly punctate. Elytron. Dark reddish-yellow, disc tending lighter. Side narrowly flanged. Moderately and evenly punctate, each puncture with a short yellow seta. Epipleuron relatively wide in front gradually narrowing to near apex, front portion weakly and shallowly grooved. 152 C. H. 8. WATTS Ventral surface. Light reddish-yellow to reddish- yellow. Pronotal process very narrow between procoxae, apical portion not much wider, strongly keeled. Mesosternum with narrow, triangular groove for reception of pronotal process; tip reaching past level of front of mesocoxae. Rear triangular midline extension of metasternum as long as wide; front extension about twice as wide as long sometimes semicircular rather than triangular. Metacoxal plate about twice as wide as long, with anterio-lateral corner extending some way along metasternum; hind edge moderately concave; sides weakly beaded; posteriolateral angles rounded. Metatrochanter small, almost twice as long as wide. Metafemur greatly swollen, widest just before middle, with small notch on hind edge near base; reddish-yellow often darker on outside edge. Dorsal metatibial spine relatively long, broad, about twice size of ventral spine and approximately length of segment | of metatarsus. Segment | of metatarsus a little longer than other segments combined. Male Tergite 7 broadly triangular, with apodemes, with rectangular apical process (Fig. 2g). Penis very small, simple, not divided into basal piece and trigonium, elongate, tip rounded, without parameroids (Fig. 15). Tegmen about as long as penis, with wide transverse basal bit, lobes thin, sinuate, bulbous at tips (Fig. 15). Variation The colour is darker in some and the contrast between the lighter disc of the elytra and the darker sides is variable. The hind edge of the metacoxal plate varies from weakly to quite strongly concave. The relative lengths of the tegmen and penis vary a bit. Etymology Latin. “Rufus” — red, “rotundus” — round. Notes A relatively rare, moderately sized, shiny, dark reddish species readily recognised by its broadly oval, almost rounded, shape, weakly to moderately convex hind edges of the metacoxal plates, weakly grooved front portions of the elytral epipleura, long metatibial spine and the 7th tergite in the males with a pronounced apical process unique in Australian Scirtes. The male genitalia are unlike any other Australian Scirfes in having a small, weak, penis much shorter than the tegmen. The species is widespread in north Queensland and is also known from Madang on the north coast of New Guinea. The only habitat records are one from a tree hollow and one from bamboo internodes suggesting that the species may breed in such situations. Scirtes tindaleensis sp. nov. (Figs 1, 16) Types Holotype male, “Qld. Greenvale 70km SW at light 14 — 24 Mar 1995 A. J. Watts”, SAMA. Paratypes 65; 2, “NT. Kakadu NP c.lkm S of Arnhem Hwy on Pine Creek Rd. M.V. Light 25 — 30 Mar. 1980 M. B. Malipatil”, NTM; 52, “Tindale, N.T. 14.318 132.22E | — 20 Dec.1967 light trap W. J. M. Vestjens”, 47, ANIC, 5 NMV, 2 slides SAMA; 1, “Burrell’s Ck Stuart H’way, N.T. 25 Nov. 1972 D.H.Colless”, ANIC; 1, “Valley of lagoons via Areenvale Apr. 1988 n. Qld. K. H. Halfpapp”, QPIM; | “Iron Range Cape York Pen. N. Qld. 1- 4.v.1973 G. B. Monteith”, QM; 1, “Captain Billy Creek Cape York Pen, N, Qld.142 50E 11 40S 9-13 vii.1975 G. B. Monteith’, QM; 1, “3 mi S of Marmor, Q. 29 mi SSW of Rockhampton 23.43.58 150, 42E 13 xii.1968. at light Britton & Misko”, ANIC; 2, “NT Lake Bennett area c 25km SE of Manton Dam 25 — 30 Dec 1979 M. B. Malipatil” NTM; 11, “ULA.R. N.T. Feb.21.1967 Coll. C.S.L1”, NTM. Description (number examined, 66) Habitus. Length 3.5 — 4.9 mm., relatively flat, oval. Head. Reddish-yellow to dark reddish yellow; antennae light reddish-yellow. Relatively small; width between eyes about 2.5x dorsal width of eye. Strongly, quite densely and evenly punctate. Frons with sides diverging, front edge straight or weakly concave, edges beaded. Segment | of antenna large, barrel-shaped; segment 2 smaller, cylindrical, segment 3 as long as segment 2 but narrower; segments 4 -10 long, narrow, cylindrical; segment |] approximately the same length as segment 10. Pronotum. Reddish-yellow to dark reddish-yellow. Short, broad. Evenly and moderately punctate, each puncture with a moderately long golden seta. Hind angles obtuse, anteriolateral angles projected forward, sides weakly beaded. Scutellum. Reddish-yellow to dark reddish yellow, Sides approximately equal length, lateral two convex. Elytron. Reddish-yellow to dark reddish-yellow, with diffuse darker areas. Sides subparallel, weakly and narrowly flanged, more strongly towards front. Moderately and evenly punctate, each puncture with a short yellow seta. Epipleuron relatively wide at front, evenly narrowing to near base. REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 153 Figs 18-23. Dorsal views of tegmen (left) and penis (right). Lines = 0.5 mm. 18. Scirtes baroalba sp. nov. 19. S. beccus sp. nov. 20. S. brishanensis Pic. 21. 8. calmi sp. nov. 22. S. crassiantennae sp. nov. 23. S. cygnus sp. nov. 154 C.H. S. WATTS Ventral surface. Reddish-yellow occasionally with darker areas. Pronotal process very narrow for whole length, strongly keeled. Mesosternum short, with small elongated groove for reception of pronotal process, tip reaching level of front of mesocoxae. Rear triangular midline extension of metasternum wider than long, narrower than front extension. Metacoxal plate (Fig. 1) much wider than long, with anterio-lateral corner extending some way along metasternum; hind edge very strongly concave; sides beaded; posteriolateral angles rounded. Metatro- chanter small (Fig. 1). Metafemur greatly swollen, widest just before middle (Fig. 1), weakly notched on hind margin near apex. Dorsal metatibial spine relatively long. More than twice size of ventral spine and about two thirds length of segment | of metatarsus. Segment | of metatarsus a little longer than other segments combined. Ventrites with punctures small, shallow; reticulation moderate fine, more pronounced towards rear; apex of apical ventrite truncated or weakly concave. Male No external differences between sexes. Penis elongate, basal piece elongate, trigonium deeply bifid towards apex; single parameroid long, thin pointed (Fig. 16). Tegmen a little shorter than penis, with two well separated lobes partially enclosing penis, each lobe broadly triangular near base, thin towards apex which is weakly clubbed (Fig. 16). The distal portions of penis and tegmen often visible in preserved specimens. Variation Occasional specimens with lighter sutural region on elytra compared with rest of elytra. Etymology Named after the town where many of the specimens were collected. Notes A large, elongate, dark reddish species, broadly sympatric with the two other large species with concave edges to the metacoxal plates: S. kaytae and S. emmaae. Separated from S. kaytae by the lack of dark areas on the elytra, any hint of an upturned front edge to the frons and the antenna are thinner with the segments symmetrical. Separated from S. emmaae by the more elongate shape, darker colour and more strongly concave hind edges to the metacoxal plates. Scirtes emmaae and S. kaytae are most clearly separated from this species by the male genitalia (the tips of which are often visible in preserved specimens): S. emmaae has small triangular protrubances near the tegmen tips (Fig. 6); S. kaytae has a distinct hook on the tip of the penis (Fig. 12); S. tindaleensis has swollen tips to the tegmen lobes (Fig. 16). b) Scirtes helmsi species complex. The description of S. helmsi is placed first, the rest are in alphabetical order. Scirtes helmsi Blackburn, 1891 (Figs 24, 40) Type Holotype male; “T 3811 A7” Blackburn coll 1910-236” “Scirtes Helmsi, Blackb.” Specimen remounted with genitalia extracted and mounted on same card. NHM. The type locality is given as “Victoria; taken near Benalla” by Blackburn (1891). Description (number of dissected males examined, 9) Habitus. Length 2.8 — 4.1 mm., relatively flat, elongate. Head. Reddish-yellow to dark chocolate-brown, often with darker patches; antenna light reddish —yellow to dark brown, basal segments lighter. Eyes small, width between eyes about 3.0x dorsal width of eye. Moderately and evenly punctate, each puncture with a moderately long seta. Frons with sides weakly diverging, front edge straight or weakly concave, edges weakly beaded. Segment | of antenna barrel shaped; segment 2 about half as long and narrower, barrel-shaped; segment 3 about same size as segment 2, narrower, wider towards base; segment 4 about twice length of segment 3; segments 5 — 10 subequal, narrow, cylindrical, a little shorter than segment 4; segment 11 elongate/oval, a little longer and flatter than segment 10. Pronotum. Brown, margins narrowly lighter yellow-brown. Short, broad. Punctures moderately dense, small, each puncture with a moderately long golden seta. Hind angles obtuse, front edge sinuate, sides weakly beaded. Scutellum. Yellow-brown, usually lighter than elytra. Equilateral triangle or slightly wider than long, lateral sides weakly convex. Elytron. Chocolate- brown to dark brown. Sides weakly flanged in basal third. Moderately and evenly punctate, each puncture with a short yellow seta. Epipleuron relatively wide in front quarter, becoming narrower over rest of elytron, front portion flat. Ventral surface. Yellow-brown to dark reddish- yellow, often with diffuse darker and lighter areas. Pronotal process very narrow. Mesosternum with small elongate area in midline in front for reception of pronotal process; rear tip rounded, reaching past front REVISION OF AUSTRALIAN SC/RTES ILLIGER AND ORA CLARK 155 of mesocoxae. Front extension of metasternum in midline small, bounded behind by ridge; rear midline extension of metasternum short, about twice as wide as long. Metacoxal plate square, with anterio-lateral corner extending along metasternum; hind edge straight, sloping towards midline; sides beaded; posteriolateral angles rounded. Metatro-chanter small, relatively squat, about 1.7x as long as wide. Metafemur greatly swollen, widest a little before middle, weakly notched on rear margin near apex. Dorsal metatibial spine relatively short, about twice size of ventral spine and about half length of segment | of metatarsus; segment | of metatarsus a little longer than other segments combined. Ventrites with punctures small, shallow, reticulation, moderate, fine, more pronounced towards rear; apex of apical ventrite truncated or weakly concave. Male Little external difference between the sexes. Basal piece of penis short, round, orientated at right angles to trigonium (viewed laterally, Fig. 40); trigonium very long and thin, 3.8x length of basal piece, tip sharply pointed; single parameroid long, a little more than half length of trigonium, with well defined apical hook (Figs 24, 40). Tegmen about half length of penis, lobes thin, elongate, well separated (Fig. 24). Notes A moderately sized (for the group), elongate, dark coloured species, occasionally with vague darker areas on the head, with a very long, thin, golfclub- shaped penis, the tip of which often protrudes from the abdomen in preserved specimens. Generally larger and darker than the more common S. brisbanensis and S. orientalis from which it can be reliably separated only by the male genitalia. In spring the larvae are common in temporary streams in open forest around Forreston in the Mt Lofty Ranges of South Australia. Specimens examined New South Wales. 1, CSIRO Lab. Chiswick nr Armadale NSW, Jan.1966, B. Clydesdale, ANIC. Northern Territory. |, Birketts Woolshed, Mus Exp 1916 Central Australia, SAMA. South Australia. 5, 12km N Forreston, 5/3/03, C. Watts, SAMA; 2, Watts’s Gully Mt Crawford Forest, 20/11/99, C. Watts, SAMA; Victoria. |, near Benalla, Helms, BMNH. Scirtes alastairi sp. nov. (Fig. 17) Types Holotype male, “QLD Greenvale 70 Km SW. at light 17 — 28 Jan 96, A. Watts”, SAMA. Paratypes 10; 4, “QLD Greenvale 70 Km SW at light 14 — 23 Feb 96 A. J. Watts”, SAMA: 1, ditto, “28 Mar — 7 Apr 1995”, SAMA; 1, ditto, “6 — 15 Dec 95”, SAMA; 2, ditto, “1 — 10 Mar. 95”, SAMA; 2, ditto, “17 — 26 Jan 96”, SAMA. Description (number examined, 11) As for S. helmsi except as follows. Length 2.2 — 2.6 mm. Uniformly light reddish-yellow, head a bit darker in some. Interorbital width 2.4x dorsal width of eye. Male Basal piece of penis broad, oval; trigonium about as long or a little longer, curved in lateral view; one parameroid, stout, about half length of trigonium, sinuate (Fig. 17). Tegmen about half length of penis, lobes broad basally, finger-like apically, close together (Fig. 17). Etymology Named after my son who collected many of the specimens described in this paper. Notes A small yellow species resembling S. storeyi, recognised by the broad basal piece, thin trigonium and stout, sinuate parameroid of penis. Scirtes baroalba sp. nov. (Fig. 18) Type Holotype male, “12. 47S 132 .S1E Baroalba Creek Springs 19 km NE by E of Mt Cahill 28.x. 72, at light, E. Britton”, ANIC. Mounted on slide. Description (number examined, 1) As for S. he/msi except as follows. Length 2.8 mm. Interorbital width 2.5x dorsal width of eye. Reddish- yellow, head tending darker, basal segments of antenna lighter. Male Basal piece of penis very narrow; trigonium about 1.5x as long as basal piece, irregularly shaped with thin spine near apex; parameroid nearly as long as trigonium, relatively stout (Fig.18). Tegmen about as long as penis, lobes well separated, finger-like (Fig.18). Etymology Named after the type locality. Notes A small reddish-yellow species from coastal 156 C.H.S. WATTS Northern Territory with a very distinctive parameroid of the penis (Fig.18). Scirtes beccus sp. nov. (Fig. 19) Types Holotype male, “TE341 Tullamarine 4-9-75”, ANIC. Paratype male, as for holotype, mounted on slide, SAMA. Description (number examined, 2) As for S. he/msi except as follows. Length 3.5 mm. Head dark reddish-yellow, pronotum reddish-yellow, darker on disc, scutellum light reddish-yellow, lighter than elytra, elytron reddish-yellow, ventral surface reddish-yellow, antenna and palps a little lighter. Interorbital width 2.7x dorsal width of eye. Metasternal plates depressed in midline, posterior-lateral angles prominently rounded, hind edges slanting inwards. Male Basal piece of penis elongate-oval, trigonium small, with half its length comprising a thin beak- like portion with the curve upwards (Fig. 19). Tegmen much longer than penis, lobes well separate, tips projecting well beyond penis, apical half of lobes thin, basal half, wide, triangular, partially enclosing penis (Fig. 19). Etymology Latin. “Beccus” — beak, a reference to the beak- like trigonium. Notes The relatively large size, strongly depressed metasternal plates and the beak-like trigonium to the penis distinguish this species. The metasternal plates are reminiscent of O. improtectus but the lower surface is still present and still provides a cover, albeit a very short one, for the metatrochanter articulation (eg Fig. Ic). Scirtes brisbanensis Pic 1956 (Figs 20, 38) Type Holotype male, with genitalia extracted and mounted on same card, “N. Guinea Biro 1900” “Queensland Brisbane” “Monotype 1956 Scirtes brisbanensis Pic.” “Scirtes brisbanensis nm” “Scirtes brisbanensis Pic”, HUNG. Description (number of dissected males examined, 70) As for S. helmsi except as follows. Length 2.2 — 3.7 mm. Interorbital width 2.5x dorsal width of eye. Reddish-yellow to dark reddish-yellow, tending to be darker towards front; ventral surface lighter. Male Basal piece of penis small, oval; trigonium long and thin, a little more than twice length of basal, apex expanded somewhat in dorsal/ventral plane; one or two parameroids, larger (left hand one) about two thirds as long as trigonium, abruptly narrowing into apical hook; second parameroid little more than small oval knob, often absent (Figs 20, 38). Tegmen a bit longer than penis, lobes well-separated, finger- like, tips rounded (Fig. 20). Notes A common, widespread species, possible more northern in distribution than S. he/msi. Recognised by the finger-like lobes of the tegmen and penis with a thin trigonium with slight apical dorsal/ventral expansion. The second parameroid is very small or absent, the main parameroid is abruptly hooked. Specimens examined (dissected males only) Queensland. |, Big Mitchell Ck Mareeba-Molloy Road, 4 May 1967, D. H. Colless, ANIC; 1, Brisbane, 4/11/62, G. Monteith, UQIC Reg# 53678; 1, Bundaberg, 3/4/1975, at light, H. Frauca, ANIC; 1, Bushland Beach 20km N Townsville, A. J. Watts, 6- 11/2/98, SAMA; 1, Caincross Nat. Pk. Via Maieny, 7/4/66, G. Monteith, UQIC Reg# 53716; 3, Cairns Gordonvale, 15/1/99, leg. A. Podlussany, HUNG; 2, Colosseum Ck, 10mls S of Miriam Vale, 20/12/66, B.Cantrell, UQIC Reg# 53663; 1, Davis Ck Rd via Mareeba, 20/1/91, S. Defaveri, QPIM; 1, Deception Bay, 29/12/62, G. Monteith, UQIC Reg# 53645; 1, Gayndah, 11/1/64, H. A. Rose, UQIC Reg# 53722; 1, Greenbank, 8/1/63, G. Monteith, UQIC Reg# 53720; 1, Greenvale 70km SW, at light, 17 — 26 Jan 96, A. J. Watts, SAMA; 2, Homestead, Silver Plains, Via Coen, 11/2/64, G. Monteith, UQIC Reg# 53651/ 53647; 3, Knob lagoon, 30mi. NW of Doomadgee Mission, NW Qld., 22/5/72, G. Monteith, UQIC Reg# 53705; 1, Lawes, 6/4/63, A. MacQueen, UQIC Reg# 53699; 1, Ditto except, 18/12/62, G. Monteith, UQIC Reg# 53700; 2, 23km N of Mareeba, 12/11/89, R. I. Storey, at light, QPIM; Nardello’s Lagoon nr Mareeba, 29/3/96, C. Watts, SAMA; Tin Can Bay. 22/1/99, leg. A. Podlussany, HUNG; 1, 30km N Marlborough, 24/2/81, Hangay & Herozeg, No 244, HUNG; 1, 21km E Mareeba, 21/1/91, at light, R. I. Storey, QPIM; |, Moorehead R. N of Laura, 20/1/90, Fay & Halfpapp, QPIM:; 1, Paradise Falls Bunya Mts Nat. Pk. 26.528 151.35E, 6/10/84, REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 157 I. Naumann J. Cardale, ANIC; 1, 7km NE Tolga, Feb 1988, at light, Storey R. & D. E. Faveri, QPIM; 1, Tolga, 13 — 20/11/85, at light, J. D. Brown, QPIM; 1, Whileside Xing, N. Pine River, 12/7/63, G. Monteith, UQIC Reg# 53718. Northern Territory. 1, 4ml SW Alice Springs, 18/2/66, Britton, Upton & McInnes, ANIC; 4, Batten Creek 16.10S 136, 31km WSW Borroloola, 15/4/76, at light, J. E. Feehan, ANIC; 3, 22km WSW of Borroloola 16.08S 136 06E, 16/4/76, at light, J. E. Feehan, ANIC; 3, Bessie Springs, 16.40S 135.51E 8km ESE of Cape Crawford, 12/4/76, at light, J. E. Feehan, ANIC; 1, Ikm N of Boko Hill SW of Borroloola 12.26S 136.01E, 14/4/76, Key, Balderson er al, ANIC; 2, Calliope R 14ml SE Gladstone 23.508 151.13E, 23/1/70, at light, S. Misko, ANIC; 46km SSW of Borroloola 16.28S 136.09E, 23/4/76, at light, J. E. Feehan, ANIC; 1, Katherine Gorge, 26/10/75, at light, M. J. Muller, ANIC; 2, Mc Arther River 16.108 136.05E 48km SW by S of Borroloola, at light, 13/4/76, J. E. Feehan, ANIC; 2, McArthur River 16.47S 135. 45E 14km SW of Cape Crawford, 11/4/76, J. E. Feehan, ANIC; 1, Surprise Creek 16.258 136.05E 45km SW by S of Borroloola, 14/4/76, at light J. E. Feehan, ANIC; 2, Roe Creek 12km SW by W of Alice Springs 23.46S 133.46E, 27/9/87, M. S. Upton, ANIC; 1, Tindale 14.318 132.22E, | — 20/12/67, at light, W. J. M. Vestjens, ANIC. New South Wales. 4, Bogan River, SAMA; 2, New England, Glenrock, 5/11/97, leg. G. Hangay, HUNG; 1, Lachlan River 15km SW of Eublong, 28/12/75, Z. Liepa, ANIC; 2, Tamworth, Lea, SAMA; 2, Yuragir NP Station Creek, 20/11/82, J & E Doyen, at light, ANIC. South Australia. 1, 21km SE of Oodnadatta 30.40S 135.37E, 20/9/78, M. S. Upton, ANIC. Victoria. 1, Benalla, 18/2/67, G. Monteith, UQIC Reg# 53682. Western Australia. 5, 6km S Pinjarra, 23/10/96, C. Watts, SAMA Scirtes calmi sp. nov. (Fig. 21) Types Holotype male; “15.36S 125.15E CALM Site 28/3 4km W of King Cascade W.A. 12 — 16 June 1988, T. A. Weir”, “at light open forest”, ANIC. Paratypes 3; 2, as for Holotype, | ANIC, | SAMA; 1, “6.318 12516E CALM Site 25/1 Synnot Ck. W.A. 17 — 20 June 1988 T.A.Weir’, “at light open forest”, ANIC. Description (number examined, 4) As for S. helmsi except as follows. Length 2.7 mm. Light reddish-yellow, diffusely darker towards front; ventral surface and appendages lighter. Interorbital width 2.5 x dorsal width of eye. Punctures on dorsal surface relatively large. Male Basal piece of penis broad, oval; trigonium about as long as basal piece, triangular, apex rounded; two parameroids cach relatively stout, curved, about as long as trigonium, left hand one slightly hooked, right hand one with tip missing in only specimen (reconstructed in illustration) (Fig. 21). Tegmen about half-length of penis, lobes narrow, slightly notched on outside near apex (Fig. 21). Etvmology Named after CALM (Western Australian Department of Conservation and Land Management) who financially supported the collection of the species. Notes Most easily recognised by the broad aedeagus with two strong parameroids. Scirtes crassiantennae sp. nov. (Fig. 22) Types Holotype male, “AUSTRALIA, n. Qld. llkm WSW of Petford, 23.1.1988, R.L.Storey, at light”, QPIM. Paratype 1, male, “AUSTRALIA, n. Qld Petford-Irvinebank Rd, 6.iv.1992, Cunningham, DeFaveri”, QPIM. Description (number examined, 2) As for S. helmsi except as follows. Length 2.5 mm. Uniformly reddish-yellow. Interorbital width 2.5x dorsal width of eye. Segments of antenna relatively short, stout, segment 10 only a little longer than wide. Male Basal piece of penis broad, oval; trigonium narrowly triangular, about twice as long as basal piece, sharply pointed; single parameroid about as long as basal piece, moderately broad, with small but distinct hook at apex (Fig. 22). Tegmen longer than penis, lobes well separated, relatively narrow, broader in middle, inner edge serrated in apical half (Fig. 22). Etymology Latin. “Crassus” — thick. A reference to the thick antennae. Notes A small, uniformly yellow, species (one specimen 158 C. H. 8S. WATTS 24 28 Figs 24-29. Dorsal views of tegmen (left) and penis (right). Lines = 0.5mm. 24. Scirtes helmsi Blackburn. 25. S. musica sp. nov. 26. S. nalyerensis sp. nov. 27. S. nigerpalpus sp. nov. 28. S. orientalis sp. nov. 29. S. peniculus sp. nov. REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 159 has diffuse dark areas on head and pronotum) known only from the two type specimens. Recognised by the tegmen lobes serrated on their inner edges. The antennae (male) are noticeably stouter than in most other species in the S. he/msi group. Scirtes cygnus sp. nov. (Fig. 23) Types Holotype male, “Cow Bay, N. of Daintree, N. Qld.7- 20.11.1984, I. C. Cunningham”, QPIM. Paratypes 7; 3, as for holotype, 1, QPIM, 2 (1 slide), SAMA; 1, “Cow Bay, N of Daintree R., N. Qld, 18-25.1.1984, Storey & Cunningham”, QPIM; 1, Cow Bay, N of Daintree, N. Qld, 25.i-7. ii. 1984, I. C. Cunningham”, QPIM; 1,. “Iron Range, Cape York Pen. N. Qld. 1-4. v. 1973, G. B. Monteith”, QM.; 1, ditto except ‘26 May — 2 June 1971”, UQIC Reg# 53662”, UQIC. Description (number examined, 8) As for S. helmsi except as follows. Length 2.0 — 2.8 mm. Uniformly light reddish-yellow. Interorbital width 2.2x dorsal width of eye. Male Basal piece of penis relatively large, narrowly oval; trigonium narrow, about two thirds as long as basal, slightly expanded at apex; single parameroid as long as trigonium, broad, with hook at apex (Fig. 23). Tegmen longer than penis, lobes well-separated, relatively thin, with large triangular expansion in middle partially enclosing penis (Fig. 23). Etymology Latin. “Cygnus” — swan, a reference to the swan- like parameroid. Notes A small golden species recognised by the swan- shaped parameroid and triangularly shaped middle sections of the tegmen lobes which partial enclose the penis. Scirtes musica sp. nov. (Fig. 25) Types Holotype male, “15.03S 145.09E 3 km NE of Mt Web Qld., 1-3 Oct. 1980, T. A. Weir”, ANIC. Paratype male, as for Holotype, slide, SAMA. Description (number examined, 2) As for S. helmsi except as follows. Length 1.8 — 2.4 mm. Uniformly light reddish-yellow. Interorbital width 2.4x dorsal width of eye. Male Basal piece of penis short, round; trigonium relatively broad, twice as long as basal, asymmetric with a strong lateral spine; two parameroids, both nearly as long as trigonium, left hand one thin, widening towards apex, terminating in a small hook, right hand one bifid with two long finger-like pieces (Fig. 25). Tegmen about as long as penis, in shape of tunning-fork (Fig. 25). Etymology Latin. “Musica” — music, a reference to the tunning- fork shape of the tegmen. Notes A small very golden species recognised by the complex penis with a prominent spine and the tunning- fork shaped tegmen. Scirtes nalyerinensis sp. nov. (Fig. 26) Types Holotype male, “WA Lake Nalyerin 33 08S 116 22E CHS Watts 6/10/03”, SAMA. Paratypes 5 (3 slides), as for holotype, SAMA. Description (number examined, 6) As for S. helmsi except as follows. Length 3.5 — 4.2 mm, elongate oval. Light chestnut, head, disk of pronotum, antennae other than basal segments darker. Side of elytron weakly flanged in front half. Interorbital width 3.0x dorsal width of eye. Male Basal piece of penis very small, oval; trigonium long, relatively broad in basal half narrower in apical half, basal half with additional ‘flap’ adpressed to one side; single parameroid relatively short, thin, with abrupt apical hook (Fig. 26). Tegmen lobes thin, asymmetric, one longer than penis with tip sharply pointed and projecting well beyond tip of penis; shorter lobe thin and about two-thirds length of other, with small setae in basal half (Fig. 26). Etymology Named after the type locality. Notes A moderately sized species with relatively large 160 C. H. 8. WATTS eyes and an extremely long narrow lobe to the tegmen which projects well beyond the end of the penis which itself is relatively long. At first sight very reminiscent of S. he/msi but in that species it is the trigonium which is elongate and projecting. Known only from Lake Nalyerin in the Jarrah forest of Southwestern Western Australia. The holotype was collected from a flowering Hakea beside the lake. The other specimens were bred from larvae that were abundant in the shallow lake. Scirtes nigerpalpus sp. nov. (Figs 27, 42) Types Holotype male, “WA 2.5 Km W Serpentine 23/9/00 C. Watts”, SAMA. Paratypes 73, as for holotype, SAMA; 65, “12k W Serpentine WA, 24/10/96, C. Watts”, SAMA; 9, “6 km S Pinjarra, 23/10/96, C. Watts”, SAMA; 1, “Swan R Lea”, SAMA; 3, “32.238 115.59E 3km SE by S Serpentine WA 2 Oct 1981 I. D. Naumann J. C. Cardale”, ANIC; 1, “33,518 123.00E Thomas River 23 km NW by W of Mt Arid WA 4-7.xi.1977 J. F. Lawrence”, ANIC. Description (number examined, 74) As for S. helmsi except as follows. Length 2.6 — 3.7 mm. Head black or very dark brown; pronotum dark brown to black with narrow yellowish border; scutellum and elytra dark brown to black; ventral surface dark brown to black, trochanters and knees lighter. Interorbital width 3.0x dorsal width of eye. Metafemur relatively narrow (Fig. 1b). Male Basal piece of penis relatively short, oval; trigonium relatively broad, narrowing towards apex, about 2x length of basal piece; two parameroids, left hand one relatively broad, nearly as long as trigonium, narrowing abruptly near apex into thin, slightly curved, apical portion, right hand one short, thumb-like (Fig. 27). Tegmen about as long as penis, lobes well separated, narrow, anther-like, tips pointed (Fig. 27). Etymology Latin. “Niger” — black. Notes A moderately large almost black species recognised by its dark colour, including the base of the antennae and the palpi, moderately expanded metafemurs, relatively small eyes, short basal piece to the penis, broad spatulate trigonium and two parameroids, the larger, left hand one, with a relatively abrupt apical hook. Like S. pinjarraensis, S. nigerpalpus is only known from near Pinjarra in Western Australia. It can be separated from S. pinjarraensis, which is common in the same habitat, by its larger size, smaller eyes and darker colour, including antennal bases, palpi and scutellum. In a few specimens the pronotum is reddish-yellow in contrast to the dark head. In these the antenna and palpi are lighter than usual. Scirtes orientalis from Eastern Australia seems close but is much lighter coloured, has broader metafemurs, larger eyes and has a narrower trigonium to the penis. The larvae are common in shallow ditches in spring and the adults are common on nearby flowering shrubs. Scirtes orientalis sp. nov. (Figs 28, 43) Types Holotype male, “Russell R. at Belenden Ker Landing, N Q., 5m 24 Oct-9 Nov. 1981 EARTHWATCH/ QLD.MUSEUM Malaise trap, rainforest”, ANIC. Paratypes 24; 2, as for holotype, one of which bears additional label “A.N.I.C. COLEOPTERA Voucher No 83-0387”: 2, “35,16S 149.06E Black Mtn ACT, 600m., Dec 1987, M.E.Irwin, ex Malaise trap”, ANIC; 1, “Brandy Creek Qld. 18 km E Proserpine, 100m., 21 June — 10 Aug.1982, S & E Peck SBP43”, ANIC; 1, “Bruxner Park, Via Coff’s Harbour, 25.xi. 1967. NSW. G. Monteith” “UQIC Reg# 53712”, UQIC; 1, “Cann River, N. Vic, 28.1.1967 G. Monteith” “UQIC Reg# 53715”, UQIC; 1, “AUSTRALIA: n Qld. Danbulla S.F. 11km NE of Yungaburra, 21.12.1986, Storey & De Faveri”, QPIM; 1, “AUSTRALIA: N. QLD. Danbulla S.F. via Yungaburra, 13.11.1992, at light, Storey, De Faveri & Huwer’, QPIM; 1, “Kiola Forest Pk. NSW, 20m., 15km N Batemans Bay, 30 Aug.1982, S. & J. Peck SBP119” “wet sclerophyll litter’, ANIC; 1, AUSTRALIA. n Qld. 5 km NNW of Kuranda 1.y- 14.vi.l985 Storey & Halfpapp” “MDFI Intercept Trap Site No 24”, QPIM; 2, “AUSTRALIA Narrabeen” “NSW”, “22.2.1984, leg. G.Hangay”, HUNG; 1, ditto except 29.12.1984, SAMA; I, ditto except 25.12.1984, HUNG; I slide, ditto “23-xii- 1983” SAMA; 2, Russell R. at Bellenden Ker Landing, N.Q. 5m., 24 Oct-8 Nov 1981, EARTH WATCH/QLD.MUSEUM, Beating, rainforest”, QM; 6, “Russell R. at Belenden Ker Landing, N Q., 5m Nov | 1981 EARTHWATCH/QLD.MUSEUM” REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK lol “Q.M BERLESATE NO 361, 17.16.S, 145.57E, Palm swamp. Moss on tree trunks”, 5 QM, 1 slide SAMA. Description (number of dissected males examined, 23) As for S. helmsi except as follows. Length 2.4 — 3.2 mm. Dorsal surface reddish-yellow with darker markings on head and pronotum; ventral surface reddish-yellow, palpi and antennal bases lighter. Interorbital width 2.2x dorsal width of eye. Male Basal piece of penis small, oval; trigonium elongate triangular, a little more than twice length of basal piece; two parameroids, left hand one largest, about two-thirds as long as trigonium, relatively broad, with terminal hook, right hand one finger-like, about half length of other (Fig. 28). Tegmen about two-thirds the length of the penis, lobes moderately separated, thumb-like (Fig. 28). Etymology Latin. “Orient” — east, a reference to its distribution in Australia. Notes The extent of the dark markings on the head and pronotum are variable and in some specimens the base of the elytra is diffusely darker than the rest of the elytra. The male genitalia resemble S. nigerpalpus but have the trigonium consistently narrower and the tips of the tegmen lobes rounded rather than pointed. They also differ from S. nigerpalpus in their much lighter colour, broader metafemurs and larger eyes. Four specimens from the Northern Territory may belong to this species. The male genitalia (one specimen only) appear close but they are smaller (2.1 — 2.4 mm long) and the metafemurs are a little narrower: 2.1x as long as wide as against 1.8x. Associated specimens 2, 12.478 132. 51E Baroalba Creek, nr. source, rainforest, 19km NE by E of Mt Cahill, NT., 29/10/72 by sweeping, D. Colless”, ANIC; 2. 12.478 132. 51E Baroalba Creek, 19km NE by E of Mt Cahill, NT., 29/10/72, at light, E. Britton, ANIC. Scirtes peniculus sp. nov. (Fig. 29) Types Holotype male; 12.578 132.33E Jim Jim Creek, N.T. 19 km WSW of Mt Cahill, 24.10.72, at light, E. Britton”, ANIC. Paratypes 8: 5, as for holotype, 2 ANIC, 3 SAMA; | slide, “12. 47S 132.51E Baroalba Creek Springs, NT 19 km NE by E of MT Cahill 28.x.72, at light, E. Britton’, SAMA; 1,714.49S 126.49E Carson escarpment W.A., 9 — 15 Aug. 1975, I.F.B. Common and M. S. Upton”, ANIC; 1, “15.02S 126.55E Drysdale River WA., 3 — 8 Aug 1975, LF.B. Commom and M. S. Upton”, ANIC; 2,” 12.528 132.47E Nourlangie Creek, N.T. 8km E of Mt. Cahill, 27.x.72, at light, E. B. Britton”, ANIC; 1, “Jabiru, N.T. 17-20. 9.1982, R.[.Storey, at light”, QPIM; | slide, “12.46S 132 39E 12 km NNW of Mt Cahill, NT. 25.x.72, at light, E. Britton”, SAMA; 1 slide, “12. 50S, 132. 51E 15 km E by N of Mt Cahill, NT. 29.x.72, at light, E. Britton”, SAMA; 1, “12,50S 132,51E 16km E by N of Mt. Cahill.N.T. 16 x1.1972 T. Weir & A. Allwood”, NTM; 1, “12.528 132.46E Nourlangie Creek 6km E of Mt Cahill, N.T. 12.x.1972 T. Weir”, NTM. Description (number of dissected males examined, 6) As for S. helmsi except as follows. Length 1.8 — 2.5 mm. Light reddish-yellow to quite dark chestnut, head tending darker, antennae, palpi and extremities of legs lighter. Interorbital width 2.0x dorsal width of eye. Male Basal piece of penis rather narrowly oval; trigonium shorter, about half as long and wide, irregularly shaped, quite strongly curved in lateral view; single parameroid about as long as basal piece, relatively stout, sinuate or weakly hooked (Fig. 29). Tegmen a little longer than penis, lobes with very prominent oval area of strong setae a little below apex (Fig. 29). Etvmology Latin. “Peniculus” — brush/tuft, a reference to the brush-like groups of setae on the tegmen. Notes A small, relatively dark, northern species, recognised by the irregularly shaped trigonium and tegmen lobes with a strong bush of setae near apex. Scirtes pinjarraensis sp. nov. (Figs 30, 39) Types Holotype male, “12 K W Serpentine WA, 24/10/96, C. Watts”, SAMA. Paratypes 45; 40 (4 slides), “12 K W Serpentine WA, 162 C. H. S. WATTS Figs 30- 35. Dorsal views of tegmen (left) and penis (right). Lines = 0.5mm. 30. Scirtes pinjarraensis sp. nov. 31. S. podlussanyi sp. nov. 32. S. pygmaeus sp. nov. 33. S. rivularis sp. nov. 34. S. spatula sp. nov. 35. S. storeyi sp. nov. REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 163 24/10/96, C. Watts”, SAMA; 5, “6K S Pinjarra WA, 23/10/96, C. Watts”, SAMA. Description (number examined, 46) As for S. helmsi except as follows. Length 1.7 — 2.7 mm. Head dark reddish-yellow; pronotum dark reddish-yellow with yellowish borders of variable width; scutellum yellow; elytron reddish-yellow to dark reddish-yellow; ventral surface reddish-yellow with diffuse lighter areas; base of antenna lighter. Interorbital width 2.5x dorsal width of eye. Male Basal piece of penis short, oval; trigonium long and thin, 2.5 — 3.0 x as long as basal piece, slightly knobbed at tip; single parameroid nearly as long as trigonium, relatively narrow, rapidly narrowing in apical quarter to a thin curved portion (Fig. 30). Tegmen a bit shorter than penis, lobes well separated, narrow, tips pointed (Fig. 30). Variation There is considerable variation in the colour of the head and pronotum from completely reddish-yellow to having extensive areas of lighter colour. Etymology Named after the type locality. Notes A small species with a very limited known distribution, recognised by the short basal piece to the penis and the single parameroid with long, thin, curved apical portion (Fig. 30). In spring adults are common on flowering shrubs beside water in the area around Pinjarra in the Southwest of Western Australia. Here the larvae are common among dead leaves and debris in shallow ditches. Scirtes nigerpalpus is also abundant in the same places. Apart from differences in the penis S. nigerpalpus is larger and almost totally black including antennal bases and palpi. Scirtes podlussanyi sp. nov. (Fig. 31) Type Holotype male, AUSTRALIA Queensland Tin Can Bay, 99.1.22.leg. A. Podlussany” SAMA. Description (number examined, 1) As for S. helmsi except as follows. Length 2.7 mm. Light chestnut, scutellum, rear margin of pronotum, appendages and ventral surface reddish-yellow. Interorbital width 2.6x dorsal width of eye. Male Basal piece of penis large, oval; trigonium a little shorter, broad at base, evenly narrowing to rounded apex, curved upwards towards tip: single parameroid arising low on basal piece, relatively narrow, tip nearly reaching apex of trigonium (Fig. 31), viewed laterally basal half thick, apical half thinner and curved. Tegmen a bit shorter than penis, lobes well separated, narrowly finger-like (Fig. 31). Etymology Named after the collector. Notes A small dark species recognised by the large basal piece to the penis, parameroid nearly as long as trigonium and stirrup-like tegmen. Scirtes pygmaeus sp. nov. (Fig. 32) Types Holotype male, “WA | km W Kodjinup Swamp 34 24 03S 116 38 37E CHS Watts 1/10/03”, SAMA. Paratypes 8 (3 slides), as for holotype, SAMA. Description (number examined, 9) As for S. helmsi except as follows. Length 2.2 — 2.5 mm, oval. Reddish-brown, scutellum, narrow border of pronotum, basal segments of antennae, palpi, portions of head and ventral surface including legs yellowish. Side of elytron weakly flanged in front half. Interorbital width 3.2x dorsal width of eye. Male Basal piece of penis small, oval, trigonium long and narrow, 2.5x the length of the basal piece, tip sharply bent to one side; single parameroid arising low on the basal piece, long, thin, curved towards apex (Fig. 32). Tegmen a little shorter than penis, broad lobes relatively short, well separated, slightly asymmetric (Fig. 32). Etymology Latin. “Pygmaeus” — dwarf, a reference to its small size. Notes A small, dark, Western Australian species with pale bases to the antennae. The long trigonium is unique within Australian Scirtes in having the tip sharply bent to one side. All specimens were reared from larvae that were abundant in dead grass and other vegetation at the 164 C. H. S. WATTS edges of a large, seasonal, Melaleuca swamp. Scirtes rivularis sp. nov. (Figs 33, 41) Types Holotype male, “WA 2km SW North Dandalup, 2/10/2003, C. H.S. Watts”. Paratypes 3 (1 slide), as for holotype, SAMA. Description (number examined, 4) As for S. he/msi except as follows. Length 3.4 — 3.5 mm, elongate oval. Light to relatively dark chestnut, head, areas on disk of pronotum, middle and apical segments of antennae and much of ventral side darker. Side of elytron weakly flanged in front half. Interorbital width 4.5x dorsal width of eye. Male Basal piece of penis oval, trigonium relatively long and broad, about twice as long as basal piece, apex rounded; two parameroids, left hand one long, nearly as long as trigonium, narrow, abruptly hooked at apex, right hand one about half length of left hand one, finger-like. (Figs 33, 41). Tegmen a little longer than penis, lobes well separated, finger-like, narrowing towards tips, tips rounded (Fig. 33). Etymology Latin. “Rivularis’” — of a brook, a reference to the larval habitat. Notes A moderate sized species with relatively small eyes dark head, variegated pronotum and light coloured bases of legs. Close to the Eastern Australian S. orientalis but a little larger, darker and the parameroid crenulate on the bottom edge and the hook more abrupt (Fig. 41). The area of small spines on the top of the trigonium is only clearly visible on prepared slides. All the known specimens were bred from larvae collected from the headwaters of a small stream running off an escarpment in the Darling Ranges in forest country east of North Dandalup in South-west Western Australia. Scirtes spatula sp. nov. (Fig. 34) Types Holotype male, “Qld Greenvale 70km SW, at light, 29 Jan — 4 Feb 1997, A. J. Watts”, SAMA. Paratypes 2 males; 1, “Tolga N. Qld. 1-ii.1980, N.Gough, J. D. Brown”, QPIM. 1, “Katherine, N.T. 23.1. 1971 T. Weir & A. Allwood”, NTM. Description (number examined, 3) As for S. helmsi except as follows. Length 3.3 — 3.4 mm. Dull reddish-yellow, parts of head and pronotum slightly darker. Elytra moderately flanged in front half. Width between eyes about 3x dorsal width of eye. Male Basal piece of penis narrowly oval; trigonium a little longer, moderately broad, sides parallel until close to apex where they converge to rounded tip; single parameroid arising low on basal piece, about as long as trigonium, relatively narrow, narrowing slowly towards rounded tip (Fig. 34). Tegmen only about half length of penis, lobes broad, moderately enclosing penis, with raised, strongly chitinized, toothed, inner portion near middle (Fig. 34). Etymology Latin. “Spatula” — broad flat tool, a reference to the shape of the trigonium. Notes A moderately large species, recognised by the elytral edge moderately flanged, the spatulate trigonium, the single parameroid without a terminal hook and the enclosing, toothed tegmen. Scirtes storeyi sp. nov (Fig. 35) Types Holotype male “QLD Greenvale 70km SW at light, 12 — 21 Apr 1995 A. J. Watts”, SAMA. Paratypes 17; 1, as for holotype, SAMA; 1, “Biloela QLD 3 Apr, 1982 R. Howell”, ANIC; 1, AUSTRALIA n. Qld Granite Gorge 9.5 km SW of Mareeba, 9.12.1987, at light, Storey & de Faveri”, QPIM; 2, “Qld Greenvale, 70 km SW, at light, 14 — 23 Feb 96, A. J. Watts, SAMA; | slide, ditto, “Feb 96”, SAMA; 1 slide, ditto, “6 — 15 Dec 95”, SAMA; 1, “Iron Range Cape York Pen, N. Qld. 1-4.v.1973 G B Monteith”, QM; 1, “Homestead, Silver Plains Via Coen, N. Qld 20.x11.1964. G. Monteith”, “UQIC Reg # 53650”, UQIC; 1, “Australia, NT Humpty Doo 6km E, 9.2-4.3.1987, R. I. Story”, QPIM; 1, “Kalpower X-ing 75 km NW of Laura N. Qld. 2.1v.1983, at light R. I. Storey”, QPIM; 1, “4 miles S.W. of Lee Point Darwin NT, 6 Mar 1967, M. S. REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 165 37 \ Figs 36-37. Dorsal views of tegmen (left) and penis (right). Lines = 0.5mm. 36. Scirtes triangulus sp. nov, 37. S. Vicloriacensis Sp. Nav. Upton”, ANIC; 1 slide, “15.388 125.125E CALM Site 28/3 4km W of King Cascade WA, 12 — 16 June 1988 T. A. Weir’, “at light open forest”, ANIC, 1, “1510S 145. O7E 3.5km SW by S Mt. Baird QLD 3 5 May 1981 A. Calder”, ANIC; 2, “AUSTRALIA n. Qld tikm WSW of Petford, 23.1.1988, R. 1. Storey, at light”, QPIM; 1, “Russell R at Bellenden Ker Landing N.Q. 5m 24 Oct — 9 Nov 1981 EARTIHWATCH/QLD MUSEUM”, “A.N.LC. COLEOPTERA Voucher 83-0587”, QM; 1, “16.318 126.16E CALM Site 25/1 Synot Ck WA, 17 — 20 June 1988, T. A. Weir’, ANIC; 1, “Murgenella, NT 4.vili. 1982 C. Wilson & S. Collins”, NTM; 17, N.T., Darwin, Lee Pt on Eucalyptus blossom 28.1.1980 M.Malipatil”, NTM. Description (number examined, 17) As for S. helmsi except as follows. Length 2.0 — 2.4 mm. Light reddish-yellow, a bit darker or head. Inicrorbital distance 2.0x dorsal width of eye. Male Basal piece of penis narrowly oval, tigonium somewhat longer, thin, slightly sinuate in dorsal view, weakly curved in lateral view; single parameroid, thin, nearly as long as trigonium (Fig. 35). Tegmen as long as penis, lobes wide, partially enclosing acdeagus, with well-marked transverse, strongly chitinized ridge on inside near apex (Fig. 35). Etvmology Named after Ross Storey, the the specimens used in this revision including this species. Curator of Entomology at QPIM who kindly provided many of Notes A small, light coloured, northern species, recognised by the long, thin, single parameroid and the enclosing tegmen with chitinized ridge. Scirtes triangularis sp. nov. (Fig. 36) Types Holotype male, “Kuranda Range State Forest N. Qld 20 Apr. 1967 1D. TT. Colless”, ANIC. Paratypes 2; 1, as for holotype, ANIC; 1 slide, “Bamboo Ck., near Miallo N of Mossman, N. Qld. 25 Apr. 1967 D. H. Colless”, SAMA. Description (number examined, 3) As for S. felmsi except as follows. Length 3.1 mm. Head, pronotum, scutellum and ventral surface reddish-yellow, elytra dark chestnut, lighter on dise with small triangular area of yellow near suture at apex. Interorbital width 2.5x dorsal width of eye. Mate Basal piece of penis oval; trigonium almost as long, relatively narrow, with thin point at apex, sharply deflexed near base so that most of trigonium at right angles to rest of penis: single parameroid relatively narrow, as long as basal piece, weakly hooked at apex (Fig. 36). Tegmen about half length of penis, undivided except for apical fifth, lobes short, thumb-like (Fig. 36). 166 C.H.S. WATTS Figs 38-44. Lateral views of the penises of Scirtes helmsi-like species. Lincs = 0.5mm. 38. S. brisbanensis Pic. 39. S. pinjarraensis sp.nov. 40. S. helmsi Blackburn. 41. S. rivularis sp. nov. 42. S. nigerpalpus sp. nov. 43. S. orientalis sp nov. 44. S. victoriaensis sp. nov. Etymology Latin “Triangulus” having three angles, a reference to the triangular marking on the elytra. Notes A little known species with darkish elytra with distinctive triangular yellow area at apex in middle. Recognised by the sharply bent trigonium of the penis and short tegmen lobes. Scirtes victoriaensis sp. nov. (Figs 37, 44) Types Holotype male, “Dimboola, Vic. Caravan Park, light trap 18.x1.73, S Misko”, ANIC. Paratvpes 4: 1, “ Benalla, Vic. 18.11. 1967 G. Monteith” *UQIC Reg # 53681.” UQIC; 1, “Kiata, V. 20.ix.18 FL E. Wilson”, NMV; 1, “Noble Park, V. F. E. Wilson 28.6.19”, “Scirtes helmsi id by J. Armstrong”, NMV; 1, “Pakenham 31.12. 22 Vic C. Oke”. “Scirtes helmsi id by J. Armstrong”, NMV. REVISION OF AUSTRALIAN SCIRTES ILLIGER AND ORA CLARK 167 Description (number examined, 5) As for S. he/msi except as follows. Length 3.8 — 4.3 mm, elongate oval. Chestnut, base of antenna lighter, parts of ventral surface darker; covered with prominent silver setae. Side of elytron weakly flanged in front half. Interorbital width 2.7x dorsal width of eye. Male Basal piece of penis oval, trigonium longer, widely triangular at base, rapidly narrowing to narrow shaft which is slightly expanded dorsal/ventrally, sharp ventral ridge for much of length; two parameroids, left hand one as long as trigonium, narrow, sinuate, right hand one much shorter, thick, thumb-like (Fig. 37). Tegmen a little shorter than penis, lobes well separated, long, thin, tips pointed (Fig. 37). Etymology Named after the State in which the specimens were found. Notes A relatively narrow, large, dark species from Victoria, recognised by the uniform dorsal colour, pale antennae bases, weak elytral flanging and long, thin, sinuate left parameroid (Fig. 37). The penis is unusually large and robust. Acknowledgments I would like to thank the following people whose help greatly improved the paper. The curators of the collections noted earlier for allowing me to examine specimens in their care; Howard Hamer for preparing the illustrations and transferring them to computer files ready for publication; Debbie Churches for improving the layout of the manuscript and Hiroyuki Yoshitomi for improving the manuscript, drawing my attention to the presence of Ora in the Australian fauna, and sending me a pre publication copy of his important Doctoral work on the Scirtidae of Japan. References ARMSTRONG, J. W. T. 1953. On Australian Helodidae (Coleoptera). Proceedings of the Linnean Society of New South Wales 78, 19-32. BLACKBURN, T. 1891. Notes on Australian Coleoptera with descriptions of new species. Part X. Proceedings of the Linnean Society of New South Wales 6, 479-550. Carter, H. J. 1935. Australian Coleoptera. Notes and new species. Nol X. Proceedings of the Linnean Society of New South Wales 60, 179-193. KITCHENER, R. L. & ALLsopp, P. G. 1987 Prionocyphon niger sp n. (Coleoptera: Scirtidae) from water-filled tree holes in Australia. Journal of the Australian Entomological Society, 26, 73-79. _ & CALLAGHAN, C. 1982. The fauna of water- filled tree holes in box forest in Southeast Queensland. Australian Entomological magazine 8, 61-70. Lea, A. M. 1919. Notes on some miscellaneous Coleoptera, with descriptions of new species.-Part V. Transactions of the Royal Society of South Australia 43, 166-261. NyHoLm, T. 1972. Zur Morphologie und Function des Helodiden-Aedoeagus (Col.) Entomologica Scandinavica 3, 81-119. _ 2002. Scirtes japonicus Kiesenwetter and its allies, with descriptions of Scirtes ussuriensis n. sp. (Coleoptera, Scirtidae). Entomologische Blatter 98, 49- 60. Pic, M. 1956. Nouveaux Coleopteres de diverses families. Annales Historico-Naturales | Musei — Nationalis Hungarici 7, 71-92. WATERHOUSE, C. O. 1875-1882, Descriptions of new Coleoptera belonging to the families Psephenidae and Cyphonidae; Cistula Entomologica, 2, 563-573. CHARACTERISTICS OF AN AUSTRALIAN POPULATION OF PARAXONCHIUM ORTHODON (LOOF 1964) ALTHERR & LOOF 1969, WITH A NOTE ON EVOLUTION AND BIOGEOGRAPHY OF THE SUBFAMILY PARAXONCHIINAE (NEMATODA, DORYLAIMIDA, APORCELAIMIDAE) By M. Hoppa* Summary Hodda, M. (2004). Characteristics of an Australian population of Paraxonchium orthodon (Loof 1964) Altherr & Loof 1969, with a note on evolution and biogeography of the subfamily Paraxonchiinae (Nematoda, Dorylaimida, Aporcelaimidae). Trans. R. Soc. S. Aust. 128(2), 169-174, 30 November, 2004. The presence of Paraxonchium orthodon (Loof 1964) Altherr & Loof 1969 is recorded in Australia for the first time. The species has been found in NSW and Queensland to date. The morphometrics of specimens from the two localities show large differences, but the number of specimens is small, so these differences are inconclusive. The differences between the Australian and Venezuelan populations of P. orthodon are discussed and a cladistic analysis of the genera and species groups in the subfamily Paraxonchiinae is presented, resulting in Paraxonchium rhamphionus (Jairajpuri 1966) Altherr & Loof 1969 being placed in the “orthodon group” rather than, as previously, in the “laetificans group”. Some biogeographic trends and the relationship of the subfamily with Aporcelaimidae or Qudsianematinae are also suggested by this analysis. The subfamily is basically Gondwanan in origin, but the genera Parapalus and Paraxonchium may be Laurasian, with a secondary radiation in Gondwana. Key Words: first record, Paraxonchium rhamphionus, Tendinema, Gopalus, Aporcelaiminae. Transactions of the Royal Society of S. Aust. (2004), 128(2), 169-174. CHARACTERISTICS OF AN AUSTRALIAN POPULATION OF PARAXONCHIUM ORTHODON (LOOF 1964) ALTHERR & LOOF 1969, WITH A NOTE ON EVOLUTION AND BIOGEOGRAPHY OF THE SUBFAMILY PARAXONCHIINAE (NEMATODA, DORYLAIMIDA, APORCELAIMIDAE) by M. Hoppa* Summary Hoppa, M. (2004) Characteristics of an Australian population of Paraxonchium orthodon (Loof 1964) Altherr & Loof 1969, with a note on evolution and biogeography of the subfamily Paraxonchiinae (Nematoda, Dorylaimida, Aporcelaimidae). Trans. R. Soc. S. Aust. 128(2), 169-174, 30 November, 2004. The presence of Paraxonchium orthodon (Loof 1964) Altherr & Loof 1969 is recorded in Australia for the first time. The species has been found in NSW and Queensland to date. The morphometrics of specimens from the two localities show large differences, but the number of specimens is small, so these differences are inconclusive. The differences between the Australian and Venezuelan populations of P. orthodon are discussed, and a cladistic analysis of the genera and species groups in the subfamily Paraxonchiinae is presented, resulting in Paraxonchium rhamphionus (Jairajpuri 1966) Altherr & Loof 1969 being placed in the “orthodon group” rather than, as previously, in the “laetificans group”. Some biogeographic trends and the relationship of the subfamily with Aporcelaimidae or Qudsianematinae are also suggested by this analysis. The subfamily is basically Gondwanan in origin, but the genera Parapalus and Paraxonchium may be Laurasian, with a secondary radiation in Gondwana. Key Worps: first record, Paraxonchium rhamphionus, Tendinema, Gopalus, Aporcelaiminae. Introduction The subfamily Paraxonchiinae Dhanachand & Jairajpuri 1981 is a small, relatively homogeneous group of four genera and 17 species of free-living nematodes of the family Aporcelaimidae, order Dorylaimida. Five new species have been discovered and the subfamily reviewed within the last few years (Hodda et al. 1994, Siddiqi 1995). Loof & Zullini (2000) described a new genus from Costa Rica and transferred one genus containing a single species to the family Qudsianematidae. Andrassy (2001) transferred one species to the genus Aporcelaimellus. More recently, hitherto unknown specimens from Australia have been located and these appear to be from the known species Paraxonchium orthodon (Loof 1964) Altherr & Loof 1969. This paper presents a description of the Australian specimens, with a comparison with the other known population of P. orthodon. The specimens differ in a number of minor characters from the type population of P orthodon in Venezuela (Loof 1964). A cladistic analysis of the subfamily Paraxonchiinae, together with Aporcelaimidae and Qudsinematinae is also presented to clarify the relationships within Paraxonchiinae and with prospective sister groups. In the recent review of the subfamily, a number of evolutionary trends were noted (Hodda ef a/. 1994). Australian National Insect Collection, CSIRO Entomology, GPO Box 1700, Canberra ACT 2601 Australia. The location of P. orthodon in Australia allows a biogeographic interpretation of these evolutionary trends, with distinct Gondwana and Laurasian radiations. Such distinct biogeographic patterns have not been reported previously for nematodes. Materials and methods All measurements were along the midline, and used either an ocular micrometer (for straight structures less than 50,m) or a camera lucida drawing with map measuring wheel (for curved structures or greater than 50j1m). Abbreviations for morphometric indices (a, a’ etc) follow Hooper (1986). The cladogram was constructed using PAUP4.0b10, using a branch and bound search, maximum parsimony, equal weighting of characters, the tree unrooted, 11 characters as listed (Figure 2), Aporcelaiminae and Qudsianematinae as outgroups, and six generic or group level taxa within Paraxonchiinae. Four equally parsimonious trees were found, with the tree presented being the consensus tree. Bootstraps of 1000 replicates were used to assess the support for each group, with ties broken randomly, using the mean character difference as the distance measure with the Permutation Tail Probability (PTP) test (Faith & Cranston 1991) (1000 randomisations, Branch & Bound (=exact) searches). 170 M. HODDA B Fig. 1. Adult female of Australian population of Paraxonchium orthodon. A - whole body; B - anterior median lateral optical section; C - anterior lateral body surface; D - oesophageal region; E - reproductive system; F - tail. Structures indicated by arrows as follow: a.a. = amphidial aperture, a.f. = amphidial fovea, an = anus, g.r. = guide ring, int = intestine, n.t. = nerve ring, os = odontostyle, op = odontophore, ov = ovary, ut = uterus, va = vagina, vu = vulva. PARAXONCHIUM ORTHODON FROM AUSTRALIA 171 TABLE 1. Measurements of Australian specimens of Paraxonchium orthodon. measure in {um or morphometric index NSW Queensland specimen specimens (Hooper 1986) range number of specimens l 3 length of odontostyle 17 17-18 diameter of odontostyle 2 2 length of odontophore 33 30-37 Distance from anterior 3 3 to constriction Distance from anterior to guide ring = 12 11-13 length of oesophagus 225 191-242 diameter at base of lips 6 4-5 diameter at oesophago-intestinal junction 32 13-24 maximum diameter 37 14-24 distance from anterior to vulva 297 284-415 length of tail 33 28-30 diameter at anus/cloaca 17 10-14 Is 563 434-633 L 530 406-603 A 15.2 27.2-33.3 a 14.3 25.9-33.2 B 25 2.3-3.3 b’ 2.4 2.1-3.2 C 16.9 15.7-21.1 c 1.9 2.1-2.6 Vv 0.53 0.65-0.67 Vv 0.56 0.68-0.70 ratio of length to diameter of odontostyle 9 9 ratio of body diameters at lip base and oesophago-intestinal junction 5.3 4.2-4.4 Description of Australian specimens of P. orthodon Material examined QM (Queensland Museum, Brisbane) G 203090 (2 adult females), Camp Cable Qld Australia (27° 48’ S 153° 06' E), coll: R.C. Colbran, July 1967, soil around Casuarina sp. and Acacia sp.; QM G 203091 (1 adult female & one juvenile), Cooroy Qld Australia (26° 25’ S 152° 55’ E), coll: C. Banner, August 1967, soil; QM G 203092 (1 adult female), Maclean NSW Australia (29° 28’ S 153° 12’ E), coll: unknown, August 1963, soil. Method of fixation is unrecorded for all specimens, which are currently stored in anhydrous glycerol on standard microscope slides, having been remounted in 1992. Description Table |, Fig. 1. Body vermiform, small, greatly narrowed towards anterior extremity. Cuticle conspicuously layered, thick (about 1.5um), thicker towards ends of body, with fine transverse annulations. Sub-cuticule not annulated. Lateral hypodermal chords without conspicuous glands. Anterior face rounded. A rounded, shallow constriction about one body diameter (at the constriction) from anterior, circumference rounded convex anterior to constriction, without longitudinal — division. Amphidial aperture unsclerotized, close to level of constriction, slit-like, about half as wide as head. Amphidial fovea with inflexion, not sclerotized, without median support, not duplex. Odontostyle of medium thickness, strongly cuticularized, with aperture of about 3pm length or 25% of length of odontostyle, with broad central lumen and anterior ventral surface bent towards dorsal side of body. Odontophore arcuate, weakly sclerotized, rod-like. Guiding ring unsclerotized, single, not plicate. Oesophagus expanded in posterior 40%, muscular throughout, nerve ring located 30-50% of total oesophageal length from anterior, with 3 gland cells and conoid cardia at oesophago-intestinal junction. Prerectum short, about 2 anal body diameters or 50 um long. Rectum about | anal body diameter or 12 um long, straight, without rectal glands, with distinct cuticular indentation at anus. Female reproductive system didelphic, amphidephic, — branches approximately equal in length, reflexed for about half length to the left side for both ovaries, without sperm. Vulva a transverse slit, sclerotized, without flaps or disc-like structure between vagina and cuticle. Vagina sclerotized over most of length, straight, perpendicular to body wall, 10 — 14 um long, with narrow lumen at junction with uterus, without muscular sheath. Tail conoid, curved ventrally, terminus rounded, with cuticular thickening. Discussion Differences between Australian and Venezuelan populations The Australian specimens are similar to those from Venezuela and conform with the original description in most features except for the odontostyle and odontophore lengths and the morphometric indices a, b and V, which differ in range while still all overlapping. Differences between Australian specimens The observed wider range of the morphometric index “a” in the Australian population of P. orthodon compared to that for the Venezuelan population raises the possibility that the Australian population is in fact two separate and distinct taxa. One of these is probably P. orthodon, consisting of the three Queensland specimens which had values of the morphometric index “a” between 27 and 33. The 172 M. HODDA Laurasian Qu © 5 © m =, Q Cc Fa | © = © Ew Eo €5 © £ 22 58> 32 = E © of BOS Sin co) o € 3 OL C2. c & o 2) = cs 5c, = oo O a) ~ 3 © on) SS SL = Ss © oO c= go ae) (e) > Q = Le ve > a ® je) © oO coo OF oS SIO) — 6 Jal SOP SL Pat of ft ff oat a time (h) Fig. 2. Temporal host-seeking rhythm of Culex Fig. 3. Temporal host-seeking rhythm of Culex annulirostris from two geographically isolated populations as determined by human-biting catch. Values presented are mean proportions of the total catch each hour (+ 95% confidence intervals). SS = sunset. SR = sunrise. annulirostris from the Upper River Murray region as determined by hourly catches using carbon-dioxide baited traps. Values presented are mean proportion of the total catch each hour (+ 95% confidence intervals). SS = sunset, SR = sunrise. 228 C.R. WILLIAMS, A. E. SNELL & M. J. KOKKINN 09 08 4 S07 2° fos] ~ +> 25 Jan 01 E —*— 26a 01 goa 27a Ol Bos 603 z a2 Ol o4 Fig. 4. Temporal host-seeking rhythm of Culex annulirostris from northeast SA as determined by hourly catches using carbon-dioxide baited traps. Values presented are mean proportion of the total catch each hour (+ 95% confidence intervals). SS = sunset, SR = sunrise. Differences between the host-seeking rhythms in the two regions are apparent in the amount of activity in the hour after sunset (lower in northeast SA), and in the activity six hours after sunset (lower at the Upper River Murray) (Figure 2). These differences were confirmed by the GLM procedure, which detected significant region by hour interactions (F = 3.351, P = 0.044). Carbon-dioxide baited trapping The periodicity of Cx. annulirostris host-seeking at the Upper River Murray peaks at sunset, then steadily decreases throughout the night (Figure 3). All three sampling nights revealed a similar pattern. In northeast SA, a periodicity over the three sampling nights was not apparent, with one night showing a peak in activity at sunset, while another showed a peak at sunrise (Figure 4). Correlations between human-biting catch and carbon-dioxide baited trapping Statistically significant correlations between the numbers of mosquitoes captured by human-biting catch and serial carbon-dioxide baited trapping were detected for four of the five nights during which both methods were used concurrently (Table 1). Correlations returned r values from 0.539 at Mungerannie on 26/i/01, to 0.877 at Mundic Creek on 5/1/01. Discussion In the Upper River Murray region, human-biting catch revealed a nocturnal host-seeking pattern with crepuscular peaks in activity. This is consistent with the findings of previous studies (Standfast & Fieldhouse 1963; Russell 1987a). Hourly catches in TABLE 1. Results for linear correlations of hourly mosquito capture using two methods: human biting catch and carbon dioxide baited traps. Date Location r ris Jan 5,2001 Mundic Creek, 0.877 P<0.001 Upper River urray region Jan 6, 2001 undic Creek, 0.681 0.01 6 school of salmon Catching snapper 3 Smoky Bay, Eyre Peninsula Catching snapper and N/A Smoky Bay, Eyre Peninsula possibly seal 2 May 2000 SAM records * Ling (pers. comm. 2004) notes that he now doubts that these were indeed killer whales, in the absence of any other reports of such a spectacular event. 236 S. E. GIBBS, C. M. KEMPER, R, W BYARD & M. LONG increases the amount of time with other members of the pod. At about 2 years old, it enters the juvenile phase (2-6 years), characterized by great activity and curiosity, and their bold inquisitiveness often leads to them approaching boats. It is therefore possible that the juvenile killer whale that washed up near Emu Bay took butchered parts of dolphin from some human activity. The presence of undigested dolphin remains suggests that the interaction had taken place shortly before or at the time of the killer whale’s death. Piecing together the events around the time of its live stranding and subsequent death, we can hypothesise what might have happened. The adult female killer whale that was observed floating dead off Tumby Bay on 12 November may have been the mother of the juvenile that stranded and subsequently died. No live killer whales were seen when the juvenile stranded at Tulka yet one would expect that a lactating female would not normally desert her dependent calf (Haenel 1986). We conclude, therefore, that the juvenile’s mother may have died. Without its mother and perhaps separated from its natal pod, the juvenile would not have known how to hunt and may have taken dolphin pieces given to it by humans or raided fishing operations in which dolphin carcasses were being illegally used as bait. The presence of both digested and undigested dolphin remains in the juvenile’s stomach suggests that the dolphins or parts thereof were consumed over a period of time. Alternatively, if the dolphin pieces were in various states of decomposition at the time the killer whale consumed them, this suggests that the dolphins had died at different times and had been stored possibly for bait. The second scenario may account for the different species of dolphin, including the ‘unknown’ as they may have originated far from where they were consumed by the killer whale. The cause of death of both killer whales was not established. The carcass of the juvenile was decomposed, a veterinarian was not present and the dissection was carried out on the beach; thus making it impossible to check for many natural and unnatural conditions. The fragmentary nature of the stomach contents combined with incomplete preservation, also made assessments of the possible presence or absence of lethal or non-lethal injuries in the pieces of dolphin difficult. While analyses of samples of stomach contents did not reveal any evidence of poisoning only a limited range of poisons were screened. The body of the adult female was never recovered despite a public appeal to find it. Such a large animal in semi-enclosed Spencer Gulf should have been relatively easy to locate and even if sharks and other predators had started to eat it, the body would still float for weeks (SAM, unpublished data). The events surrounding the deaths of the killer whales point to suspicious human interactions involving both animals. Whatever the circumstances of death of these killer whales, the presence of dolphin pieces with incised edges from a sharp blade suggests that illegal activities occurred. All cetaceans and pinnipeds are protected in South Australian waters under the South Australian National Parks and Wildlife Act, 1972 (SA), Fisheries Act 1982 and federal Environment Protection and Biodiversity Conservation Act 1999. These Acts make it an offence to kill, harass, molest or injure marine mammals and, under the Fisheries Act, it is also an offence to be in possession of a dead marine mammal or part thereof (D. Kelly pers. comm. 2003). Violations are not uncommon and there are many instances of intentional killing of cetaceans and pinnipeds in the last 10 years (Kemper etal. in press). A successful conviction was obtained in the case of crayfish fishing operators shooting common dolphins and using the carcasses for bait in craypots during April 1990 off the south coast of Kangaroo Island. Public empathy for whales and dolphins has grown in recent times but there are still people who would harm and kill them despite legal protection of marine mammals. Killer whales, and other marine mammals, are especially disliked by persons who consider them a threat or nuisance to their activities or livelihood. More information is needed to prosecute offenders and this study demonstrates the importance of gathering rigorous data, including forensic studies, on carcasses that are found. Acknowledgments We would like to thank the following people for their contribution to this paper: Andy Warner National Parks and Wildlife SA, Mike Jones Royal Society for the Prevention of Cruelty to Animals for their assistance with collecting the Emu Bay juvenile; Marianne Stockholm for details and video of the live stranding; Neil Kopman for photos and information on the adult; Bob Hamilton-Bruce and volunteers who prepared the skeleton. Deb Kelly Department for Environment and Heritage for advice on legislation; J. Gardiner Forensic Science Centre for toxicological analyses, We thank Daniel Mau for producing images from the video. KILLER WHALE DEATHS AND HUMAN INTERACTION 237 References BAKER, A. (1990) “Whales and dolphins of Australia and New Zealand: An identification guide” (Victoria University Press, Wellington). BANNISTER, J. L. KEMPER, C. M. & WARNEKE, R. M. (1996) “The Action Plan for Australian Cetaceans. Wildlife Australia Endangered Species Program Project Number 380” (Environment Australia, Canberra). BerRZIN, A. A. & VLADIMIROV, V. L. (1983) A new species of killer whale (Cetacea, Delphinidae) from Antarctic waters. Zoologicheskii Zhurnal 62, 287-295. CHRISTENSEN, I. (1984) Growth and reproduction of killer whales, Orcinus orca, in Norwegian coastal waters. Reports of the International Whaling Commission (Special Issue 6): 253-258. Cotton, B. C. (1943) Killer whales in South Australia. 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F. & Myrick, A. C. (Eds) (1980) Growth of odontocetes and sirenians: problems in age determination. Reports of the International Whaling Commission (Special Issue 3). PITMAN, R. L. & ENsor, P. (2003) Three forms of killer whales (Orcinus orca) in Antarctic waters. Journal of Cetacean Research and Management 5(2), 131-139. VISSER, I. N. (2000) Killer whale (Orcinus orca) interactions with longline fisheries in New Zealand waters. Aquatic Mammals 26(3), 241-252. THE TENNYSON SAND DUNES: VEGETATION STRUCTURE AND CONSERVATION STATUS By J. HINCHCLIFFE!” & J. G. CONRAN! Summary Hinchcliffe, J. & Conran, J. G. (2004). The Tennyson sand dunes: vegetation structure and conservation status. Trans. R. Soc. S. Aust. 128(2), 239-248, 30 November, 1994. The Tennyson sand dunes represent one of the last largely natural dune communities along the Adelaide metropolitan coast, a region where there is a considerable effort being expended by local councils and community groups on revegetation for dune stability, habitat for native animals and for shoreline visual amenity. This study represents the first quantitative analysis of their vegetation. Fifty-two species were encountered, 22 of them exotic, and cluster analysis and ordination recovered three quadrat groups on the basis of species relative cover, representing fore-, mid-, and backdune quadrats. Key Words: Coastal foredunes, vegetation structure, conservation, ordination. Transactions of the Royal Society of S. Aust. (2004), 128(2), 239-248. THE TENNYSON SAND DUNES: VEGETATION STRUCTURE AND CONSERVATION STATUS by J. HINCHCLIFFE!” & J. G. CONRAN! Summary HINCHCLIFFE, J. & CONRAN J. G, (2004). The Tennyson sand dunes: vegetation structure and conservation status. Trans. R. Soc. S. Aust. 128(2), 239-248, 30 November, 2004. The Tennyson sand dunes represent one of the last largely natural dune communities along the Adelaide metropolitan coast, a region where there is considerable effort being expended by local councils and community groups on revegetation for dune stability, habitat for native animals and for shoreline visual amenity. This study represents the first quantitative analysis of their vegetation. Fifty-two species were encountered, 22 of them exotic, and cluster analysis and ordination recovered three quadrat groups on the basis of species relative cover, representing fore-, mid- and backdune quadrats. These correlated with distance from the shore and increasing litter cover, agreeing with other dune studies. Recruitment of Olearia axillaris was associated mainly with exposed foredune sands and the large numbers of juveniles at the site imply that ongoing self-regeneration of the vegetation is occurring. Nevertheless, the dominance of introduced species is a cause for concern, and our findings provide baseline information for longer term management of dune stabilisation, key species, weeds and rare to endangered taxa. Kry Worpbs: Coastal foredunes, vegetation structure, conservation, ordination. Introduction Vegetation is fundamental in the formation and stabilisation of coastal sand dunes, providing habitat for a diverse range of animals (Chapman 1976; Viles & Spencer 1995). Australia's recent history of intense human activity along its coastline has resulted in the reduction and modification of much of its native coastal vegetation. The most obvious and direct impacts have been foreshore erosion, the replacement of coastal vegetation by housing development, the introduction of exotic plants and animals and disturbance by beach users and vehicles. Integrated approaches to management, which are essential in order to address these issues (Morcom 2002; Morcom & Harvey 2002), require a detailed understanding of the current dune condition, vegetation patterns and trends. In many instances councils want to revegetate with native species but lack appropriate data on which to base selections. In the past, introduced species such as Ammophila arenaria (marram grass) and Elymus farctus (sea wheat-grass) were used widely for dune stabilisation, and dune stabilisation studies emphasised the advantages of introduced colonisers such as Elymus spp., Cakile spp. and Ammophila spp. Over native species, particularly Spinifex spp. (Hesp 1984; Heyligers 1984, 1985; Bird & Jones ' Centre for Evolutionary Biology and Biodiversity, Environmental Biology, School of Earth and Environmental Sciences, Darling Building, DP418, The University of Adelaide, SA 5005, Australia. Email: john.conran@adelaide.edu.au * present address: Spinifex Communications, 70 Chalfont St, Salisbury Qld 4107, Australia. Email: julichinchliffe@bigpond.com 1988). Nevertheless, the germination ecology of Beach Spinifex (Spinifex sericeus) in Australia was studied to improve techniques for its use in revegetation (Harty & McDonald 1973). Dune vegetation has long been recognised as a good model for long-term succession (Warming 1891), with early colonisation by sand-binding grasses followed with gradual displacement by shrubs or small trees (Olson 1958) creating characteristic zonation patterns (Boyce 1954; Gray 1985; Rozema ef a/. 1985; Partridge 1992). Walker e¢ al. (1981) suggested that disturbance in older dune systems is likely to increase the rate of nutrient loss by causing vegetation decline, however, few Australian studies address changes produced by both succession and disturbance. Early last century, Osborn (1914) noted that the visually dominant dune plants along the Adelaide metropolitan coast included Olearia axillaris, Scaevola crassifolia, Lepidosperma gladiatum as well as marram grass. Cleland (1932) and Fenner and Cleland (1932) surveyed the coastal vegetation between Outer Harbour and Sellick’s Hill, listing 62 species, of which nine were introduced, including boxthorn (Lycium ferocissimum), evening primrose (Oenothera stricta), prickly saltwort (Sa/sola kali) and couch grass (Cynodon dactylon). Specht (1972) studied the dunes between Grange and Semaphore (including what is now the Tennyson dunes remnant), recognising a number of species groupings and emphasising the importance of succession in terms of distance from the foredune edge. More recently, Opperman (1999) used multivariate techniques to define 52 floristically distinct coastal 240 J. HINCHCLIFFE & J. G. CONRAN 1 | Housing & Gulf St 4 Vincent 2 beach sands Adelaide City ! aa. BOO i fe are ' * \ 567891011 Tennyson Heights (housing) | Sse Fig. 1. Map of the Tennyson sand dunes showing the positions of the forty 50 x 2 m sample quadrats. Circled quadrats represent those with no vegetation cover at the time of study. THE TENNYSON SAND DUNES: VEGETATION STRUCTURE AND CONSERVATION STATUS 241 dune and clifftop plant communities across South Australia, relating them to ecological gradients and environmental parameters. Similarly, the vegetation of the Murray Mouth and Coorong was studied by Brandle (2002), where 11 floristic groups including several dune associations were recognised. However, although these studies represent invaluable resources for managers at the broader scale, because of the size and complexity of the areas covered, small-scale patterns within particular areas and their local management issues tend to be lost. The Tennyson sand dunes represent one of the last largely natural dune communities along the Adelaide metropolitan coast, a region where there is considerable effort being expended by local councils and community groups on revegetation for dune stability, habitat for native animals and for shoreline visual amenity (South Australian Coast Protection Board 1993). The dunes were surveyed by Deans ef a/. (1995) and the SA Coast Protection Board (1995), who recorded 39 introduced species, mainly garden escapes (e.g. Chrvsanthemum spp.), and recent surveys suggested that 57% of the species there are exotics (South Australian Coast Protection Board 2003). Three introduced foredune- building species, Ammophila arenaria, Cakile maritima and Elymus farctus, are regarded as so naturalised as to be beyond control (Heyligers 1985) and the ground-covering exotic African Daisy (Actotis stoechadifolia) is considered to suppress regeneration by native species at the site, and to alter dune topography and formation processes (South Australian Coast Protection Board 2003). In contrast, of the 40 native species listed, Kennedia prostrata is thought to be locally extinet, Lomandra leucocephala subsp robusta is rare to endangered and 11 others were reported as very reduced in the metropolitan area. However, these descriptions of the dune vegetation along the Adelaide metropolitan coast are mainly qualitative and there are no recent detailed, fine-scale quantitative studies the vegetation patterns in the metropolitan coastal dunes, or elsewhere in the state. Accordingly, our study applies an integrated multivariate numerical approach to one of Adelaide's last remnants of natural dune vegetation in order to provide baseline management information. The major aims of the study were to: 1. Quantify the floristic composition and vegetation patterns of the Tennyson sand dunes. 2. Correlate these patterns to site environmental features. 3. Relate these patterns to the overall health of the ecosystem and its possible long-term management. Methods Vegetation patterns The site consisted of an 1] ha area at Tennyson, near West Beach, 12.5 km NW of Adelaide, South Australia (34° 51’ S, 138° 42’ E), between Uriah Place and Bournemouth Avenue, representing approximately | km of foredunes (Fig. 1) which are subdivided into 18 areas by fenced beach access tracks. Twelve parallel transects were established 100 m apart, perpendicular to the beach and running inland starting 10 m behind the foredune edge. Quadrats were spaced 25 m along each transect, and positioned so that they did not traverse a walkway. The number of quadrats along each transect varied because of the presence of barriers (e.g. Escourt House, car parks etc...), but in total forty 50 x 2 m quadrats were surveyed (Fig. 1). Quadrat size was determined from species area-curve data and guidelines proposed by Kent and Coker (1992) for heterogeneous shrubby heathlands. Within each quadrat, 25 2 x 2 m cells were aligned parallel to the shore, and every species was given an average Braun-Blanquet (1932) cover score based on its cover within each of the 25 cells. Seven environmental parameters were assessed for each quadrat: distance from the foredune edge (m); topographic position, soil electrical conductivity, pH, surface sand exposure, surface sand disturbance, and litter cover. Topographic position was categorised using a nominal scale adapted from Tongway (1994) as follows: flat (1) lower slope (2) mid-slope- (3) upper slope (4) crest (5). Conductivity and pH were analysed using approximately 10 g of surface sand taken from the centre of each cell and dried at 100°C for 24 h. A 1:5 soil / water suspension was tested using an LC-84 electrical conductivity (EC) meter and a pH meter calibrated to pH 7, and the results averaged for the quadrat. Surface sand exposure (SSE) as the estimated percentage exposed sand in each quadrat, surface sand disturbance (SSD: the estimated percentage of sand showing mechanical disturbance), and litter cover were each measured using a six point scale derived from McDonald e7 al. (1990), ranging from 0 (undisturbed) to 5 (concentrated animal or human activity such as the creation of tracks, vegetation damage, removal or severe trampling). In addition, the numbers of Olearia axillaris plants within each of the three age classes (Juvenile, Mature and Senescent) defined by Hinchliffe and Conran (in press) were recorded within each quadrat to determine whether Olearia numbers and life classes correlated with overall site vegetation patterns, and/or with the site environmental parameters. 242 J. HINCHCLIFFE & J. G. CONRAN Data analysis After exclusion of six quadrats which lacked any vegetation, the quadrat x species matrix was subjected to cluster analysis by Sorensen association with flexible UPGMA clustering to produce a dendrogram using the program PC-ORD ver. 4.0 (McCune & Mefford 1999). Major divisions in the dendrogram branches were used to define groups. Species which were significantly (p < 0.05) associated with the resulting groups, were identified using indicator species analysis (Dufréne & Legendre 1997), which combines information on species abundance in a particular group and the faithfulness (% indicator value) of occurrence of a species in a particular group and its statistical significance, i.e. utility as a group indicator. Sites and species were ordinated in PC-ORD 4.0 using non-metric multidimensional scaling (NMS), canonical correspondence analysis (CCA) and principal component analysis (PCA). These different methods, which include both direct (CCA) and indirect (NMS and PCA) gradient analysis techniques, were used to assess the reliability of the resulting groups, i.e. if the same quadrat groups arose despite the analysis they are more likely to represent environmental response groups (Jongman et al. 1995). Dendrogram-derived quadrat groups and their indicator species were plotted into the ordination space, allowing the results of the classification and ordination to be viewed simultaneously and as a test of the integrity of the dendrogram groups (Kent & Coker 1992). In addition the correlation between the position of the quadrats on the ordination space and_ the environmental features recorded for the quadrats was calculated by correlating the scores for the environmental variables against the axis coordinates, plotting the direction of positive correlation as vectors onto the ordination by biplot analysis, where the length of the vector indicates the relative importance of that character in the ordination space (McCune & Mefford 1999). Results Fifty-two species were encountered in the 34 quadrats with vegetation cover, of which 30 (58%) were native and 22 (42%) were introduced. The natives with the highest cover were Olearia axillaris (present in 30 quadrats) and Spinifex sericeus (23), while the dominant introduced species were Euphorbia paralias (30 quadrats) and Lagurus ovatus (28), reflecting patterns seen elsewhere in South Australian dune systems (Opperman 1999; Brandle 2002). In contrast, other generally common to dominant southern Australian coastal dune species (sensu Opperman 1999) such as Nitraria billardiera, Mid-dune (2) Shoreward/ Back-dune (3) Seaward/ Fore-dune (1) [ Seaward/ | Fore-dune (1) | | 15 e Ely é mi *cosuire Ss * Dist Me a | gus ; Caki | 4 = Spin | 4 d | s Cyno (Mature) | a * pH | Litter 7 Mid-dune (2) s Distance 7! Shoreward/ » © = - Back-dune (3) Stress=14% 20 10 oo 10 NMS Axis 1 Fig. 2. A: Dendrogram of the quadrats using Serensen /UPGMA cluster analysis. B: Non-metric Multi- dimensional Scaling analysis (NMS) ordination of the quadrats, and biplot of significant indicator species and correlated environmental parameters. Vector lengths indicate importance and direction of positive increase for feature measured, parameters in parentheses represent non-significant trends. Leucopogon parviflorus and Senecio lautus were present in only two three quadrats, as were the locally rare natives Clematis microphylla (2 quadrats) and Pimelia serpyllifolia (1 quadrat). Surface sand salinities (NaCl equivalent derived from EC values) were relatively low, averaging 1,220 mgL-' and ranging from 632 mgL" to 2,556 mgL". The dendrogram could be divided into three quadrat clusters (Fig. 2A). Group | consisted of eight of the 12 foredune quadrats, with Cakile maritima and Elymus farctus as the significant indicator species (Fig. 3A; Table 1). These quadrats were typically flat, and similar to the Rolling Spinifex/Sea Fig. 3. A: Seaward/foredune community. B: Mid-dune association. C: Shoreward/backdune community. Wheat Grass Tussock Grassland of the Coorong dunes (Brandle 2002) and E/ymus-dominated beach meadow sites reported by Talbot and Talbot (1994). Group 2 represented 13 sites, most of which were located behind the foredune (Fig. 2A) and represent a mid-dune association (Fig. 3B). There were 9 significant indicator species for Group 2 (Table 1) of which the most strongly significant (p < 0.01), and thus most useful as group indicators, were Threlkeldia diffusa, Chrysanthemum coronarium, Rhagodia candolleana ssp. candolleana, Spinifex sericeus and Isolepis nodosa. Group 2 quadrats were also the only ones where the previously common (Cleland 1932) and now locally rare species Pimelea serpyllifolia ssp. serpyllifolia and Adriana klotzschli were recorded. The remaining 13 quadrats formed Group 3 (Fig. 2A), and were located behind the central walkway and on the west-facing backdune slope (Fig. 3C). There were nine significant indicator species (Table 1), seven of which are introduced weeds, the most significant (p < 0.01) being Ehrharta calycina and Oxalis pes-caprae. Group 3 was also the only area where Clematis microphylla, Calocephalus brownii and Melaleuca lanceolata were observed. 244 J. HINCHCLIFFE & J. G. CONRAN TABLE |. Significant indicator species, their codes and dendrogram group with which they are significantly associated (p < 0.05). Indicator values and probabilities derived using the indicator species analysis method of Dufréne and Legendre (1997). * = exotic species. Species nomenclature follows Jessop & Toelken (1986) and Carr & Horsfall (1995). Species Code Group Indicator P value (%) Seaward/Foredune *Cakile maritima CAKI 1 46.2 0.0237 *Elymus farctus ELYM 1 75.5 0.0001 Mid-dune Carpobrotus rossii CARP 2 44.1 0.0442 *Chrysanthemum coronarium CHRY 2 65.2 0.0001 Dianella brevicaulis DIAN 2 42.7 0.0347 *Euphorbia paralias EUPH 2 46.1 0.0326 Isolepis nodosa ISOL 2 57.8 0.0026 Meuhlenbeckea gunnii MEUH 2 47.6 0.0153 Rhagodia condolleana subsp. candolleana RHAG 2 56.4 0.0003 Spinifex sericeus SPIN 2 52.3 0.0006 Threlkeldia diffusa THRE 2 54.8 0.0001 Shoreward/Backdune *Cynodon dactylon CYNO 3 35.7 0.0211 *Ehrharta calycina EHRH 3 85.9 0.0001 *Lagurus ovata LAGU 3 48.2 0.0198 *Asparagus (=Myrsiphyllum) asparagoides ASPA 3 43.3 0.0299 *Oxalis pes-caprae OXAL 3 55.6 0.0030 *Poa annua POA 3 38.5 0.0130 Scaevola crassifolia SCAE 3 41.0 0.0187 Tetragonia implexicans TETR 3 43.6 0.0443 *Vicia monantha VICI 3 40.7 0.0212 All three of the ordination analyses showed essentially identical results, with the dendrogram groups discrete and arranged as a continuum. Because the patterns were the same, suggesting that the patterns reflect environmental responses, only those for the NMS are presented here (Fig. 2B) as they were both visually clearer than the others, and because NMS is considered to be generally more robust than most other ordination techniques (Minchin 1987). As expected, the significant indicator species for each group plotted into the same ordination space as the group with which they were associated. Correlation of the ordination axes with the environmental data showed that Group 1 was associated with proximity to the shoreward dune edge, high levels of exposed surface sand, low litter levels and a trend towards more numerous Olearia Juveniles. Group 2 quadrats were placed centrally in the ordination and were thus intermediate for all of the environmental parameters, although with a trend towards more abundant Mature Olearia. Group 3 correlated significantly with distance from the outer dune edge, high litter levels and reduced surface sand exposure, as well as a tendency towards fewer Juvenile Olearia. The Group 3 quadrats were also more spread out in the ordination plots, with quadrats dominated by Poa annua and Scaevola crassifolia associated with alkaline soils, while those with high cover scores for Cynodon dactylon, Oxalis pes-caprae and Lagurus ovatus located on more acid soils. Discussion The complexity of the dune structure in our survey was much lower than that reported by Specht (1972) for the Adelaide region, with many of his association-defining taxa rare or absent from our quadrats, although Olearia, Spinifex, Carpobrotus, Rhagodia and Cakile were still important. Olearia and Spinifex were similarly described as common along the dunes between Outer Harbour and Sellick's Beach (Cleland 1932), but in this survey the native perennials Adriana klotzschli and Pimelia serpyllifolia were also common, whereas these species are now rare. Similarly, Cleland (1932) recorded only 14 introduced taxa between Outer Harbour and Sellick's Beach, compared to the over 40 species now found just at Tennyson. Elevation and distance are used traditionally to classify coastal vegetation into zones, as both are assumed to reflect maritime influences, particularly exposure to sand abrasion, salt spray or storm waves (Brown & McLachlan 1990; DeOliveira 1994; Talbot & Talbot 1994). At Tennyson, the quadrat THE TENNYSON SAND DUNES: VEGETATION STRUCTURE AND CONSERVATION STATUS 245 groups also fell along a distance gradient, tending to support this view. However, other studies along the Adelaide coastline considered that species distributions relative to distance from the shore represented successional stages (Specht 1972, 1993, 1997; Specht & Specht 1999), possibly confounding the effects of environment versus time. Litter cover and vegetation patterns were correlated in our study, and litter is important in succession by modifying dune soils (Specht 1972; Enright 1978). Litter cover increases with successional age (Talbot & Talbot 1994), and at Tennyson, the positive correlation between distance and litter supports this. Higher litter levels towards the back of the dunes can increase soil organics and nutrients (Specht & Specht 1999), but can adversely affect seedling emergence and survival (Facelli & Pickett 1991). The origin (native or exotic) and physical structure of the litter can also affect community composition (Lenz ef al. 2003). Heavy backdune litter cover from Ehrharta and Lagurus may help to explain low recruitment by native species in this area, as Ehrharta was seen by Hilton & Harvey (2002) to suppress native plant recruitment on the Sir Richard Penin, SA, Nevertheless, correlation alone does not prove that grass litter is necessarily inhibiting native dune species recruitment, as Facelli & Pickett (1991) found that litter may favour establishment in harsh environments by retaining soil moisture and protecting seedlings from dessication. Specht (1972) also suggested that Olearia axillaris seedlings were associated with abundant litter (the opposite of our findings), and O. polita in New Zealand showed no spatial relationship between seedling density and litter (Willams & Courtney 1995), Similarly, although not significantly associated with quadrat groups at the site, Arctotis stoechadifolia was most abundant in the Group 3 backdune quadrats and is also considered to be a threat to the diversity and regeneration ability of indigenous taxa at Tennyson by its habit of suppressing seedling establishment and growth (South Australian Coast Protection Board 1995). pH can also affect coastal vegetation (e.g. Specht & Cleland 1961; Chapman 1976; Brown & McLachlan 1990), with higher pH = soils on calcareous dunes being more nutrient rich (Specht 1972). In our study, pH correlated with the second CCA Axis and may help to explain the floristic divergence between the quadrats of group III (the backdune), with Cynodon, Oxalis and Lagurus- dominated quadrats correlating with lower pH and those with greater Poa and Scaevola cover on higher pH soils. The relatively low surface sand salinities can be explained by rapid leaching of salt spray into the dunes, particularly once strand vegetation stabilises the sand (Specht 1972; Chapman 1976; Specht & Specht 1999). Low salinities have been reported in North American dunes (Oosting & Billings 1942; Boyce 1954; Barbour ef a/. 1973; Barbour & De Jong 1977), although arid shorelines are often more saline (Johnson 1977; Barbour et al, 1985) and closer to those seen at Tennyson. Nevertheless, salinity was not significantly associated with vegetation or other environmental variables in our study, except for pH (7 = 0.51). Boyce (1954) and Barbour ef al. (1973) were similarly unable to show correlations between — salinity and _— species distributions, with the latter reporting strong intra- site variation. The pattern may also be affected by seasonality, as salinities would be expected to be higher over the summer and lowest near the end of the winter rains, and this, combined with the timing of germination events may still have an effect on the subsequent vegetation, There is generally a strong relationship between dune elevation and vegetation (Barbour ef a/. 1973; Nakanishi & Fukumoto 1987; Talbot & Talbot 1994), as it affects the degree to which vegetation is affected by wind, salt spray, soil moisture and nutrients (Carter 1988). Nevertheless, at Tennyson topographic position was not associated with vegetation patterns, raising the question of whether smaller-scale changes in topography might be more important, but these would require sampling at a finer scale. Disturbed sand did not correlate with any other variable, suggesting that the dunes either show complex disturbance patterns or that disturbance is random, The latter seems unlikely in the light of human trampling patterns (Hylgaard 1980; Degouvenain 1996) and rabbit activity in dunes (Whatmough 1995), but does imply that disturbance does not reflect overall vegetation patterns at the scale sampled. The fact that those quadrats which lacked vegetation were all located close together in the middle of the site suggests that there may be some disturbance-related and/or sample scale effects which require further investigation if that part of the dune ts to remain stabilised or to be revegetated successfully. The species were distributed as a continuum, a phenomenon typical of coastal vegetation both in South Australia (Specht 1972) and more generally (Brown & McLachlan 1990; DeOliveira 1994), and correlated mainly with distance from the shore and litter, agreeing with studies of other dune systems (Talbot & Talbot 1994). Recruitment in the dominant shrub, Olearia axillaris, was associated with exposed sands on the seaward foredune, and not with high litter levels (c.f. Specht 1972), and the abundant juvenile plants at the site suggest that the dune vegetation 1s regenerating. 246 J. HINCHCLIFFE & J. G. CONRAN Nevertheless, the overall dominance of the site by exotic taxa and the possibility for future spread by more aggressive, potentially regeneration-preventing species, particularly in the backdune, suggests that overall site health is not optimum. Similarly, there were considerable numbers of dead shrubs, including Olearia across the site, and although recruitment was evident, the presence of dead shrubs in the “empty” quadrats implies that regeneration is not uniform. Placing these results in a management context, some caution is required since the patterns recovered are scale-dependant. The Tennyson dunes contain more than 80 plant species, ranging from grasses and herbs through to tall shrubs (Deans et a/. 1995), and the distribution and abundance of these species will vary at different sampling scales. This has _ practical implications, particularly if the same types of patterns are to be reproduced from survey data gathered in other manners or at different scales. Specht (1972) recognised six different vegetation associations for Adelaide dune systems over the same general distance from the shore as our study, implying that finer-scale sampling may reveal more localised patterning, although many of the key species for his zones were also significant indicator species within our study. In contrast, Opperman’s (1999) Olearia/Rhagodia shrubland was similar to our mid-dune association, but our fore- and backdune associations appear to be too dominated/degraded by weeds to fit easily into either Specht’s (1972), Opperman’s (1999) or Brandle’s (2002) vegetation classes. A major problem identified in this study was the dominance of introduced species. This dominance by exotic taxa indicates that the dunes are not comparable to natural systems and that management practices to encourage the reestablishment of more native taxa and to limit or reduce weeds across the site need to be implemented. The extent and potential for future spread by Elymus, Ehrharta and Arctotis are causes for concern, as are the reasons for the absence of native seedlings toward the backdunes. Furthermore, because sampling commenced 10 metres back from the foredune edge, but E/vmuis is considered to be most abundant just above the high tide mark or strandline (Heyligers 1985), its abundance at Tennyson, particularly in relation to Spinifex, may have been underestimated. It is also possible that the relative abundance of introduced species has been underestimated by the present survey, as the data were collected mainly across winter and exotic spring annuals or late-emerging geophytic weeds may have been missed. These findings provide baseline information about the structure and status of the dune vegetation at Tennyson from which strategies for more effective ongoing dune stabilisation can be developed. The management of weeds, key species and rare or locally endangered species at the site represent priority areas for further research. In particular, the role of disturbance, both natural and human on the biology and recruitment of the various weeds as well as the natives is essential if the dunes are to be managed successfully in the long term, Acknowledgements The study was part of an Honours degree by JH within the Department of Botany (now Discipline of Environmental Biology), School of Earth and Environmental Sciences, The University of Adelaide, who are thanked for the provision of resources. The Coastal Management Section of the Department of Environment and Natural Resources and the Hindmarsh Woodville Council are thanked for permission to undertake the research in the foreshore areas under their control. Ron Sandercock from DENR and the Tennyson Dunes Group are thanked for their ongoing support of the project and provision of resources and local expertise. 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(1891) De psammophile vormationer i endangered shrub Olearia polita (Wilson et Garnock- Danmark. Vid. Medd. Natur. For. Kjabenhavn 1891, 153. Jones), Nelson, New Zealand. N.Z. J. Bot. 33, 237-241. REPRODUCTION OF COMMON CARP IN SOUTH AUSTRALIA, SHOWN BY YOUNG-OF-THE-YEAR SAMPLES, GONADOSOMATIC INDEX AND THE HISTOLOGICAL STAGING OF OVARIES By B. B. SMITH*f# & K. F. WALKER* Summary Smith, B. B. & Walker, K. F. (2004) Reproduction of common carp in South Australia, shown by young-of-the-year samples, gonadosomatic index and the histological staging of ovaries. Trans. R. Soc. S. Aust. 128(2), 249-257, 30 November, 2004. Y oung-of-the-year (YOY) samples, gonadosomatic index (GSI) and the histological staging of ovaries were used to monitor the reproduction of common carp (Cyprinus carpio L.) in the lower River Murray, South Australia, from August 2001 to December 2002. Spawning occurred initially over 9 months from late September 2001 to May 2002, the longest period recorded in Australia. Key Words: Cyprinidae, Murray-Darling Basin, Australia, aging, cohorts, hatch dates, GSI, spawning. Transactions of the Royal Society of S. Aust. (2004), 128(2), 249-257. REPRODUCTION OF COMMON CARP IN SOUTH AUSTRALIA, SHOWN BY YOUNG-OF-THE-YEAR SAMPLES, GONADOSOMATIC INDEX AND THE HISTOLOGICAL STAGING OF OVARIES by B. B. SMITH**# & K. F. WALKER“ Summary SmiTH, B.B. & WALKER, K. F. (2004) Reproduction of common carp in South Australia, shown by young-of-the- year samples, gonadosomatic index and the histological staging of ovaries. Trans. R. Soc. S. Aust. 128(2), 249- 257, 30 November, 2004. Young-of-the-year (YOY) samples, gonadosomatic index (GSI) and the histological staging of ovaries were used to monitor the reproduction of common carp (Cyprinus carpio L.) in the lower River Murray, South Australia, from August 2001 to December 2002. Spawning occurred initially over 9 months from late September 2001 to May 2002, the longest period recorded in Australia. It recommenced in September 2002 and continued until at least December, when sampling ended. Contrary to previous reports, hatch-dates estimated from otolith analyses revealed that in each year, spawning was continuous from onset until completion, and that there were two peaks in YOY production between mid-October and December 2001 and mid-January and mid-March 2002. Over the entire period, there were at least 29 discrete spawning events at two locations about 30 river-km apart, most of them synchronous. GSI and histological evidence indicated spawning over seven months, including two months where the hatch-date data failed to identify any reproductive activity. Thus, the benefits of combining analyses of YOY and ovary samples are apparent here, where reproduction is protracted and there is potentially low YOY survivorship in some months, and where the local ecology of the target species is not well-understood. Key Worbs: Cyprinidae, Murray-Darling Basin, Australia, aging, cohorts, hatch dates, GSI, spawning. Introduction Studies of teleost reproduction employ two basic approaches. One monitors changes in sex steroid levels and/or the ovarian condition of mature fish via the gonadosomatic index (GSI) and macroscopic and histological staging. The other utilizes length- frequency distributions for cohorts and hatch- and spawning-date distributions estimated from otoliths, scales or other bony structures, derived from regular samples of young-of-the-year (YOY) fish. The two approaches are complementary. For example, gonad staging reveals recent or imminent spawning, but does not indicate the survivorship of the early life stages. On the other hand, YOY samples do indicate survivorship but not unsuccessful spawning due to infertility, fungal infection, starvation, predation or other environmental factors (Humphries and Lake * Cooperative Research Centre for Freshwater Ecology, School of Earth and Environmental Sciences, The University of Adelaide, Adelaide, South Australia 5005. ' Current address: Cooperative Research Centre for Pest Animal Control, SARDI Aquatic Sciences — Inland Waters, GPO Box 120, Henley Beach, South Australia 5022. Correspondence: Tel. +61 8 8207 5329; Fax +61 8 8207 5481; Email: smith.ben2@saugov.sa.gov.au ‘Hume, D. J., Fletcher, A. R. & Morison, A. K. 1983. Carp Program. Final Report. Arthur Rylah Institute for Environmental Research, Fisheries & Wildlife Division, Ministry for Conservation, Heidelberg, Victoria. Unpublished, 214 p. * Stuart, I. & Jones, M. 2002. Ecology and management of common carp in the Barmah-Millewa Forest. Final Report of the Point Source Management of Carp Project to Agriculture, Fisheries & Forestry Australia. Arthur Rylah Institute for Environmental Research, Heidelberg, Victoria. Unpublished, 214 p. 2000). Ideally, both approaches should be combined (Pinsent and Methven 1997), but this is uncommon in practice (e.g. Spranza and Stanley 2000). In Australia, published studies of reproduction in common carp (Cyprinidae: Cyprinus carpio L.) have produced incongruent results: YOY samples imply infrequent spawning over 2-4 months annually (Vilizzi 1998; Hume et al. 1983'; Stuart and Jones 2002’), whereas staging of gonads indicates regular spawning over 6-7 months (Sivikumaran ef al. 2003; Smith and Walker 2004). The disparity could reflect local variations in climate, but could also reflect the shortcomings of unilateral approaches to sampling and analysis. In this paper, part of an ongoing study (Smith and Walker 2003a, 2003b, 2004), we draw upon YOY samples and histological information to describe the reproductive biology of carp in the River Murray, South Australia. In this region, carp became a major pest following their introduction from Germany in the 1960s (e.g. Koehn et al. 2000). Materials and Methods Field sampling YOY sampling occurred every 4-6 weeks over 17 months (1 August 2001 to 31 December 2002) in backwaters at Punyelroo and Walker Flat (‘Walker Flat South’) on the River Murray downstream of Blanchetown (Weir 1), South Australia. At each location, six 20 m hauls of a seine net (5 x 1.5 mx 6 mm stretched mesh, 1.2 m cod-end) were made in 250 B. B. SMITH & K. F. WALKER likely nursery areas (water <60 cm depth, with abundant submerged vegetation: Sigler 1958; Smith 20043). This net was selective toward small fish (95% were between 10-45 mm SL; West and King 1996), but, as carp in this region attain 45 mm standard length (SL) about 6 weeks after hatching (Vilizzi 1998), the 4-6 week sampling interval was presumed to reveal all successful spawning events. Thus, carp of 10-45 mm SL were retained for analysis, and this also covered the size-range that is suitable for age estimation via the enumeration of daily otolith increments (Vilizzi 1998). If no YOY carp were captured in six hauls, sampling was abandoned; otherwise, it continued either until 50 fish had been obtained or 3 hours had elapsed. In total, 713 YOY carp were collected and samples were preserved in 70% ethanol (after Smith and Walker 2003a). Laboratory analysis All YOY carp were weighed using a digital scale (0.0001 g) after draining the mouth and gill cavities and blotting with absorbent paper (Treasurer 1992), and length (SL, 0.01 mm) was measured using digital calipers under a magnifying lamp. Subsequently, to correct for body shrinkage that occurs in alcohol, pre-calibrated regression equations were used to estimate ‘initial’ (pre- preservation) lengths and weights from those measurements made after preservation (Smith and Walker 2003a). Where possible, both the left and right otoliths (lapilli) were removed from each fish and mounted on microscope slides using Crystalbond" thermoplastic cement, and the one with the most distinct growth increments (validated as daily by Vilizzi 1998; Smith and Walker 2003b) was selected for processing and age estimation (post-hatch ages). To do this, lapilli were ground in the sagittal plane to thin-sections using a variable speed Gemmasta" 8” lapping machine with 12 um diamond polishing discs, and daily growth increments were enumerated under a 100-400x compound microscope linked to a high-resolution monitor. Two ‘blind’ counts were made (without knowledge of length/weight) and if there was any discrepancy between the age estimates, a third and final count was made. The final age estimate (in days) was reduced by -1 to account for there being, on average, one increment present at the time of hatching (Smith and Walker 2003b). In total, age estimates were made for 575 YOY carp. The remaining 138 fish were disregarded because 4 §mith, B. B. 2004. The state of the art: a synopsis of information on carp in Australia. South Australian Research and Development Institute, Primary Industries and Resources South Australia, Adelaide. Unpublished Technical Report, 70 p. both otoliths were lost (7 = 21) or rendered unusable in processing (over-grinding or cracking, = 55), or had indistinct increments (7 = 62). Estimated hatch-dates were calculated for each fish by subtracting the final age estimate from the date of capture (Equation 1), and hatch-date distributions were plotted separately for each backwater (Figs. la-b) and for the pooled data (Fig. Ic). Equation | Hatch date = date of capture — final age estimate Mean growth rates were also calculated for each fish (Equation 2). In this case, the mean length at hatch (5 mm TL) was subtracted from the ‘initial’ (pre-preservation) length at capture because only the post-hatch age was calculated, and all YOY carp are c.5 mm TL at hatching (Alikunhi 1966). If this was ignored, growth rate estimates would have been exaggerated, particularly with respect to very small/young fish (Smith 2004*). Equation 2 (‘initial’ length at capture Mean growth rate — mean length at hatch) (mm > a!) final age estimate Smith and Walker (2004) provide complementary information about the calculation of gonadosomatic index (GSI, equation 3) and the histological staging of ovaries, and include photographs and descriptions of each reproductive stage. Briefly, via observations of oocytes stages (named ‘whole-section inspection’) and inspections for migratory nuclei and post- ovulatory follicles, the ovaries of 111 female carp were histologically classified into one of six developmental stages: immature, early-developing, late-developing, ripe, spent and regressing. Ripe and spent ovaries were indicative of imminent or recent spawning, respectively. Mature females were sampled from November 2001 to October 2002. Equation 3 a 8 gonad mass : Gonadosomatic index — x 100% (GSI) total body mass Results The mean length and weight of the 575 fish for which an age estimate was made were, 22.7 mm SL (+ 6.3 S.D.; range 10.2-39.9 mm) and 0.43 g (+ 0.38, 0.017-2.13 g), respectively, and their mean estimated age and growth rate were 39.4 days NO n REPRODUCTION OF COMMON CARP IN SOUTH AUSTRALIA To (a) Frequency (YOY carp) 16 > (c) a Ss a Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan 2001 Estimated Hatch Date 2002 Fig. 1. Estimated hatch-date distributions of YOY carp sampled every 4-6 weeks from August 2001 to December 2002 at (a) Walker Flat South (7 = 317), (b) Punyelroo (7 = 258) and (c) Walker Flat South and Punyelroo combined (” = 575). No sampled fish were estimated to have hatched prior to 27 September 2001 or between 27 March and 4 October 2002. Minor ticks on abscissa are at 5-d intervals. 252 B. B. SMITH & K. F. WALKER 25, (a) November 2001 (b) 25 i December 2001 January 2002 February 2002 Frequency (YOY carp) March 2002 April 2002 May 2002 Initial length (SL, mm) Fig. 2. Modal progression of length frequencies of YOY carp sampled monthly from August 2001 to July 2002 at (a) Walker Flat South and (b) Punyelroo (*length and weight measurements precluded as fish were not preserved live on capture). REPRODUCTION OF COMMON CARP IN SOUTH AUSTRALIA 253 (+ 15.4, 12-94 d) and 0.5 mm-d! (+ 0.12, 0.22- 1.25 mm-d'), respectively. Seine net catches were consistent at Walker Flat South, where beds of ribbon weed (Vallisneria americana) provide year round habitat. At Punyelroo, however, few YOY carp (1 <5) were sampled in January and May 2002, when most submerged vegetation was exposed by low water levels (Table 1). The precision of estimates, indicated by the Index of Average Percent Error (IAPE = 2.08%: Beamish and Fournier 1981) and the Coefficient of Variation TABLE 1. Numbers of YOY carp per sample at Punyelroo and Walker Flat South from August 2001 to December 2002. No YOY carp were sampled between August 2001 and October 2001 or June 2002 and October 2002. Date Punyelroo Walker Flat South Total 17 Nov 01 28 24 52 16 Dec 01 50 50 00 19 Jan 02 4 50 54 26 Feb 02 30 50 80 20 Mar 02 50 50 00 16 Apr 02 49 49 98 25 May 02 2 50 52 13 Nov 02 2, 50 77 23 Dec 02 50 50 00 Grand Total 713 (CV = 2.94%: Chang 1982), exceeded the 95% target suggested by Campana (2001). Subjectively, 50 percent of estimated ages were considered to be within + 1-2 d of the true age, and the remainder within + 3 d. Cohort analyses Cohort analyses (after Brown et a/. 2004) indicate that 5-6 primary cohorts of YOY carp were spawned in 2001-2002 at both Walker Flat South and Punyelroo (Fig. 2). One and three cohorts, respectively, are apparent in data for the same localities in November and December 2002 (Fig. 3). Hatch-date distributions Figures la-c show estimated hatch-dates for YOY carp from August 2001 to December 2002. YOY carp hatched over at least 7 months, from late September 2001 to late March 2002, but not from April to September 2002. Hatching subsequently began in early October 2002, at the start of the 2002- 2003 season. Spawning was continuous, from onset to completion, at each location. Successful reproductive events lasted 2-10 days and were separated by periods of up to 2 weeks. Over the entire period, 29 successful events were apparent, 20 of them synchronous at the two locations. The distributions for each location are visually similar, 0 10 20 30 40 50 2 (a) [71 November 2002 (b) 0 2 [4 | 15 10 5 0 ry § > 9 ; Zod / December 2002 S 25 " S 20 1 a @ 15 ~ 40 Initial length (SL, mm) 0 10 20 30 40 50 Fig. 3. Modal progression of length frequencies of YOY carp sampled monthly from August 2002 to December 2002 at (a) Walker Flat South and (b) Punyelroo. 254 B. B. SMITH & K. F. WALKER n= 13 14 12 10 11 12 15 13 9 11 15 10 100 80 & 60 Hy | mmature S Early developing Ss SSSR Late developing 77) BSSes8g Ripe o 4 Spent a 40 Regressing n 2 oO § Le 20 7 0 Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct 2001 2002 Fig. 4. Relative numbers of female carp at each reproductive stage at the two sample locations, November 2001 to October 2002. 25 4 r 25 20 + 20 e iS D g Go 15 r 15 2 *< = 3 2 a = o 10 L 10 2 = 2 5 £ ° c 3 5 s 5 + 5 = oO = c oO {o) o 6 = 0 r T T T T T T T T T T 1 me 0 Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct 2001 2002 Fig. 5. Monthly changes in gonadosomatic index (GSI) for female carp at the two sample locations, November 2001 to October 2002. Females: @, solid line; Mean monthly temperature: A, dashed line; Approximate lower temperature limit for carp spawning (16°C), dotted line. Standard error bars included. REPRODUCTION OF COMMON CARP IN SOUTH AUSTRALIA 255 TABLE 2. Comparative evidence of spawning from gonad staging (GSI, ovary histology) and YOY samples (hatch-date distributions from otolith age estimates). ?: no mature females sampled. 2001 Aug Sep Oct Nov Dec Jan Evidence of Spawning Feb Mar 2002 Apr May Jun Jul Aug Sep Oct Nov Dec GSI ? 2? 2? \ Female 2? ? ? \ i ~ histology YOY V V \ V \ \ sampling and indicate that most surviving YOY carp arose from spawning events in November 2001, mid- January to late March 2002, and mid-October 2002 to at least early December 2002. Histological analyses and GSI Histological staging also indicated protracted spawning during 2001-2002. ‘Spent’ fish were observed in December, January (+1 fish classified as ‘ripe’), February, March and May. Later, spent fish were present in September, at the onset of spawning for 2002-2003 (Fig. 4). Thus, histological evidence suggests that spawning occurred initially over at least 6 months, from mid-December to mid-May in 2001-2002, and began again in mid-September 2002. There are no histological data for August, September and October 2001 or November and December 2002, as no mature fish were captured in these periods. Peaks in GSI (0.5-24.8%) occurred in November 2001 and October 2002, and lesser peaks in March and May 2001, suggesting that there was also spawning at these times (Fig. 5). Comparison of methods Table 2 compares the results of YOY samples and gonad staging (GSI, ovary histology), indicating months where there was evidence of spawning. Thus, spawning occurred in 13 of 17 months of observation, from September 2001 to May 2002 (9 months) and from September 2002 until at least December 2002 (4 months), when sampling ended. However, there is limited concordance between the two classes of methods. For example, in May and September 2002, histological examination of ovaries indicate some spawning, although no YOY carp were estimated to have hatched then. In November 2001 and October 2002, when many YOY carp were estimated to have hatched, there was no histological evidence of spawning. ‘ Smith, B. B. 1999. Observations on the early life history of carp, Cyprinus carpio: fecundity, spawning and tolerance of eggs to dehydration and salinity. BSc(Hons) Thesis, Department of Environmental Biology, The University of Adelaide. Unpublished, 32 p. V \ % ? V i 2 2 Vv Vv Vv Discussion Carp spawn in well-vegetated, slow-flowing or still water, when mean water temperatures exceed 15-16°C for >2-4 weeks (Swee and McCrimmon 1966; Crivelli 1981; Davies and Hanyu 1986; Brzuska 1989; Guha and Mukherjee 1991; Smith and Walker 2004). There are many suitable backwaters along the Murray in South Australia, and regional mean temperatures exceed 15°C for 6-7 months in 75% of all years (Smith and Walker 2004). In 2001- 2002, spawning was continuous over 9 months, from September to May, although most YOY carp were hatched in November and from mid-January to mid- March. Subsequently, spawning began in September 2002 and intensified in late October and November. The absence of YOY carp in samples at Punyelroo in January and May 2002 probably reflects seasonally low water levels and a lack of submerged vegetation rather than a lack of spawning. Where littoral vegetation is absent, or made inaccessible by falling water levels, YOY carp are forced into deeper, open water where they cannot be sampled adequately using a small seine. Other Australian studies report one short annual period of spawning. For example, the pioneering work of Hume ef a/. (1983') in Victoria suggested that carp spawned over 1-4 months, from mid- September to December. However, regular seine samples of YOY carp were obtained only from one lake, where there was ‘very sparse aquatic vegetation’ (Hume ef a/. 1983!, p. 10). Macroscopic staging of gonads suggested additional spawning in January and March, and potentially earlier (July, August), but this was apparently discounted (Hume et al. 1983', p. 57: Figure 3.6). In subsequent studies, adequate YOY sampling occurred only over the presumed spring/summer period until February (Vilizzi 1998, Smith 19994, Stuart and Jones 20027). In one study where YOY samples were obtained monthly over 4 years (Humphries et a/. 2002), and where a 1-4 month spawning period also was reported, the results may have been biased by concentrating most fishing effort in lotic environments, where carp spawning is minimal, and 256 B. B. SMITH & K. F. WALKER by using fishing methods (light traps, drift nets, plankton tow nets) that may be effective only when YOY carp are abundant (Smith and Walker 2004). More recent studies (Sivikumaran e/ a/. 2003; Smith and Walker 2004), plus observations by commercial fishers on the Murray (pers. comms: Keith Bell, K & C Fisheries; Henry Jones, Clayton; Shane Warrick, Walker Flat), argue for protracted spawning by carp, at least in south-eastern Australia. The patterns of spawning and YOY survival observed here are consistent with studies of carp in other parts of the world. That is, spawning begins in spring after the gonads mature in winter and, where spawning is protracted over 4-5 months or more, each female spawns more than once (Alikunhi 1966; Swee and McCrimmon 1966; Shikhshabekov 1972; Crivelli 1981; Horvath 1985; Davies ef al. 1986; Brzuska 1989; Guha and Mukherjee 1991). Where multiple spawnings occur, 3-4 months are required for the gonads to re-mature (Davies and Hanyu 1986; Davies ef al. 1986; Mills 1991). Thus, in this study, intense spawning was apparent from mid- October to December 2001 and again 3-4 months later, from mid-January to mid-March 2002. As the ‘window of opportunity’ for carp spawning in the lower Murray is about 6-7 months, based on their minimum temperature (15-16°C) and light (10-12) requirements (Smith and Walker 2004), most females probably spawn twice annually: once at onset and again after re-maturation of the gonads. The fact that spawning occurred over 9 months, rather than six, may be a consequence of variation in the timing of spawning in separated areas, mediated by pheromones which synchronise reproduction between cohabiting individuals (see Irvine and Sorensen 1993; Poling ef a/. 2001; Kobayashi ef al. 2002). Clearly, this is speculative and needs further investigation. Interpretations of spawning patterns for riverine fish may be confounded by larvae drifting in the main-channel, entrained out of upstream nursery habitat by high flows, if there is spatial variation in the timing of spawning. In this study, however, a closed population model is assumed because (a) the lower Murray is highly regulated by 9 weirs and their associated navigable locks, and is akin to a series of stepped pools, (b) there were no appreciable flows during the period of sampling and (c) carp larvae are not known to disperse via the main-channel during low flows. Thus, we are confident of our description of carp spawning pattern for the lower River Murray. Finally, this study argues for the utility of a dual approach to sampling, in that the shortfalls of one class of methods were offset by the merits of the other. Compared to the hatch-date estimates, data from gonad histology, GSI and cohort analyses failed to elucidate the timing or frequency of spawning. On the other hand, GSI data and ovary examinations indicated that spawning persisted until May 2002 and resumed in September 2002, where hatch-date data failed to identify reproductive activity. The former disparity might have been offset by increasing the number of monthly gonad samples; the latter is evidence for successful spawning but unsuccessful YOY survivorship. YOY survivorship may have been limited by the cooler conditions in late autumn (May) and early spring (September), as the food acquisition, metabolism and growth of larvae are strongly influenced by water temperatures (Houde 1987; Downing and Plante 1993; Hurst and Conover 1998; Hall and Rudstam 1999). Acknowledgements This paper is from PhD research by BBS, supported by grants from The University of Adelaide and the Cooperative Research Centre for Freshwater Ecology. We are grateful to Sandra Leigh, Kane Aldridge, Alex Ivey, Tamryn Herridge and Brenton Smith for assistance in sampling, and to Henry Jones (Clayton) and colleagues from state fishery agencies for collecting gonad samples. Gail Hermanis (Department of Anatomy & Histology, The University of Adelaide) and Sandra Leigh (South Australian Research & Development Institute, Aquatic Sciences Division) provided valuable advice on histological processing and_ interpretation. Finally, we thank three referees for providing helpful comments, which considerably strengthened the original manuscript. References ALIKUNHI, K. H. 1966. Synopsis of biological data on common carp Cyprinus carpio Linnaeus, 1758 (Asia and the Far East). Food and Agriculture Organisation of the United Nations. FAO Department of Fisheries, Rome. 114 p. BEAMISH, R. J. & FourNIER, D. A. 1981. A method for comparing the precision of a set of age determinations. Canadian Journal of Fisheries and Aquatic Sciences 38: 982-83, Brown, P., GREEN, C., SIVAKUMARAN, K. P., STOESSEL, D. & GiLes, A. 2004. Validating otolith annuli for annual age determination of common carp. Transactions of the American Fisheries Society 133: 190-96, BrzuskaA, E. 1989. Investigations on controlled reproduction of carp (Cyprinus carpio L.) in early spring. Acta Hydrobiologica 31 (1/2): 121-9. REPRODUCTION OF COMMON CARP IN SOUTH AUSTRALIA 257 Campana, S. E. 2001. Accuracy, precision and quality control in age determination, including a review of the use and abuse of age validation methods. Journal of Fish Biology 59: 197-242. CHANG, W. Y. B. 1982. A statistical method for evaluating the reproducibility of age determination. Canadian Journal of Fisheries and Aquatic Sciences 39: 1208-10. Crivetui, A. J. 1981. The biology of the common carp, Cyprinus carpio L. in the Carmargue, southern France. Journal of Fish Biology 18: 271-90. Davies, P. R. & HANYu, [. 1986. Effect of temperature and photoperiod on sexual maturation and spawning of the common carp I. Under conditions of high temperature. Aquaculture 51: 277-88. 4 , FURUKAWA, K. & Nomura, M. 1986. Effect of temperature and photoperiod on sexual maturation and spawning of the common carp II. Induction of spawning by manipulating photoperiod and temperature. Aguaculture 52: 137-44. DowninG, J. A. & PLANTER, C. 1993. Production of fish populations in lakes. Canadian Journal of Fisheries and Aquatic Sciences 50: 110-20. Guna, D. & MUKHERJEE, D, 1991. Seasonal cyclical changes in the gonadal activity of common carp, Cyprinus carpio Linn. Indian Journal of Fisheries 38 (4): 218-223. HALL, S. R. & Rupstam, L. G. 1999, Habitat use and recruitment: A comparison of long-term recruitment patterns among fish species in a shallow eutrophic lake, Oneida Lake, NY, U.S.A. Hydrobiologia 408/409: 101-113. HorvaTu, L. 1985. Egg development (oogenesis) in the common carp (Cyprinus carpio L.). In ‘Recent Advances in Aquaculture’. (Eds J. F. Muir & R. J. Roberts). pp. 31-77. (Croom Helm, London). Houpr, E. 1987. Fish early life dynamics and recruitment variability. American Fisheries Societv Symposium 2: 17-29. Humpuries, P. & LAKE, P. S. 2000, Fish larvae and the management of regulated rivers. Regulated Rivers: Research and Management 16: 421-432. , SERAFINI, L. G. & KinG, A. I. 2002. River regulation and fish larvae: variation through space and time. Freshwater Biology 47: 1307-31. Hurst, T. P. & Conover, D. O. 1998. Winter mortality of young-of-the-year Hudson River striped bass (Morone saxatilis): Size-dependent patterns and effects on recruitment. Canadian Journal of Fisheries and Aquatic Sciences 55: 1122-30. IRVINE, I. A. S. & SORENSEN, P. W. 1993. Acute olfactory sensitivity of wild common carp, Cyprinus carpio, to goldfish hormonal sex pheromones is influenced by gonadal maturity. Canadian Journal of Zoology 71: 2199-2210. KOBAYASHI, M., SORENSEN, P. W. & StAcEY, N. E. 2002. Hormonal and pheremonal control of spawning behaviour in the goldfish. Fish Physiology and Biochemistry 26: 71-84. KOEHN, J., BRUMLEY, A. & GEHRKE, P. 2000. Managing the Impacts of Carp. Bureau of Rural Sciences, Canberra. 250 p. Mitts, C. A. 1991. Reproduction and life history. /n ‘Cyprinid fishes: Systematics, Biology and Exploitation’. (Eds L. J. Winfield & J. S. Nelson). pp. 483-509. (Chapman and Hall, London). PINSENT, D. A. & METHVEN, D. A. 1997. Protracted spawning of Atlantic cod (Gadus morhua) in Newfoundland waters: evidence from _ otolith microstructure. Canadian Journal of Fisheries and Aquatic Sciences 54 (Suppl. 1): 19-24. POLING, K. R., FRASER, E. J. & SORENSEN, P. W. 2001. The three steroidal components of the goldfish pre-ovulatory pheromone signal evoke different behaviours in males. Comparative Biochemistry and Physiology Part B 129: 645-51. SHIKHSHABEKOV, M. M. 1972. The annual cycle of the gonads in wild carp (Cyprinus carpio L.) from the Terek Delta. Journal of Ichthyology 5: 855-9. SIGLER, W. F. 1958. The ecology and use of carp in Utah. Logan Agricultural Experiment Station Bulletin 405, Utah State University. Salt Lake City. 63 p. Situ, B. B. & WALKER, K. F. 2003a. Shrinkage of 0+ carp (Cyprinus carpio L.) after preservation in ethanol. Marine and Freshwater Research 54 (2): 113-6. & 2003b. Validation of the aging of 0+ carp (Cyprinus carpio L.). Marine and Freshwater Research 54: 1005-8. __& 2004. Spawning dynamics of common carp in the River Murray, South Australia, shown by macroscopic and histological staging of gonads. Journal of Fish Biology 64: 336-54. SPRANZA, J. J. & STANLEY, E. H. 2000. Condition, growth and reproductive styles of fishes exposed to different environmental regimes in a_ prairie drainage. Environmental Biology of Fishes 59: 99-109. Swee, U. B. & McCrimmon, H. R. 1966. Reproductive biology of the carp, Cyprinus carpio L., in Lake St. Lawrence, Ontario. Transactions of the American Fisheries Society 95 (4): 372-80. TREASURER, J. W. 1992. Length and weight changes in 0+ perch, Perca fluviatilis L., following fixation in formalin. Journal of Fish Biology 41: 1033-36. Vitizzi, L. 1998. Age, growth and cohort composition of 0+ carp in the River Murray, Australia. Journal of Fish Biology 52: 997-1013. A COMPARISON AMONG THREE ARTIFICIAL SUBSTRATES FOR AQUATIC MACROINVERTEBRATE SAMPLING BRIEF COMMUNICATION Summary Artificial substrate samplers (ASS) are a widely used method for collecting aquatic macroinvertebrates. They are most often used in large and deep rivers where other methods that directly sample the fauna may be difficult or impossible to apply'***. Different sampling methods may collect different subsets of the fauna from the same site. For example in several Australian lowland rivers, artificial substrates which collected colonising animals, airlift samples of the soft benthos and sweep samples of the edge habitat each collected different subsets of the total suite of macroinvertebrates within a site’. In south eastern Australia, a widely used ASS design is the onion-bag basket, which comprises a PVC ‘gutter-guard’ basket filled with two onion bags and a half a brick for ballast**. A variation on this design, the two onion bags being replaced by three citrus bags, was adopted by the Floodplain Ecology Group for the preparation of the Integrated Watering Strategy series of reports (Floodplain Ecology Group unpub.). Transactions of the Royal Society of South Australia (2004), 128(2), 259-260, BRIEF COMMUNICATION A COMPARISON AMONG THREE ARTIFICIAL SUBSTRATES FOR AQUATIC MACROINVERTEBRATE SAMPLING Artificial substrate samplers (ASS) are a widely used method for collecting aquatic macroinvertebrates. They are most often used in large and deep rivers where other methods that directly sample the fauna may be difficult or impossible to apply!?*4. Different sampling methods may collect different subsets of the fauna from the same site. For example in several Australian lowland rivers, artificial substrates which collected colonising animals, airlift samples of the soft benthos and sweep samples of the edge habitat each collected different subsets of the total suite of macroinvertebrates within a site’. In south eastern Australia, a widely used ASS design is the onion-bag basket, which comprises a PVC ‘gutter-guard’ basket filled with two onion bags and a half a brick for ballast’*, A variation on this design, the two onion bags being replaced by three citrus bags, was adopted by the Floodplain Ecology Group for the preparation of the Integrated Watering Strategy series of reports (Floodplain Ecology Group unpub.). The aim of this study was to determine if there was any significant difference in macroinvertebrate abundance, taxa richness and community composition between ASS with different internal substrates. A third filling, ‘flywire’ was also trialled to determine if it differed to the other two substrates in the aforementioned characteristics. ASS with the three types of substrate were deployed at two sites; one upland, the Broken River at Bridge Creek (36° 06’ 00” 145° 41’ 00"; altitude, 330 masl) and one lowland, the Broken Creek downstream of Nathalia (36° 02’ 30” 145° 06' 35”; altitude, 90 masl) in Victoria. The ASS were constructed using the same basic design as described in Humphries e¢ a/3 and Bennison ef al.°. “Gutterguard’ baskets (mesh size 10 x10 mm; 180 mm high x 180 mm base diameter) were filled with one of three fillings. The ‘onion’ bag samplers were filled with two double sided knitted nylon onion bags (each 350 x 770 mm, 2x2 mm mesh), the ‘citrus’ bag samplers were filled with three double sided nylon citrus bags (each 205 x 540 mm, 5x5 mm mesh) and the ‘flywire’ samplers were filled with three sheets of flyscreen (each 200 x 1000 mm, 1 xl mm mesh). Ten samplers of each type were deployed in the field for six weeks (22 February-6 April 1995) to be consistent with the monitoring carried out by Bennison et a/.5 and the Integrated Watering Strategy (Floodplain Ecology Group unpub.). Halfa house brick, or a similar sized rock was used as ballast, For the bag samplers, the brick was placed inside one of the bags and the bag was wrapped tightly around the brick. This bundle was then placed inside another bag which was wrapped around the bundle. This process was repeated for the remaining bag. The flywire sheets were wrapped tightly around the brick, one upon the other. The brick and filling bundle was placed in the basket and the top of the basket was sewn closed with the nylon cord used to anchor the ASS. The ASS were placed in pools at a depth of approximately | m, parallel to the bank, out of the main force of the current. Samplers were deployed by tying them to stream side trees or to stakes hammered into the edge of the stream. The ASS were retrieved by placing a 250 uum dipnet downstream of, and then under, the ASS as it was picked up and removed from the water. The basket was opened and cleaned in the net. Each piece of substrate (onion, citrus or flywire) was scrubbed thoroughly and examined for attached invertebrates before being removed from the net. Once all invertebrates had been removed from the basket, substrate and ballast, the sample was preserved in 70% ethanol. In the laboratory the invertebrates were sorted from the detritus, identified to the lowest taxonomic level practicable (usually genus or species) and counted. Differences in macroinvertebrate abundance and taxa richness were compared using a two factor ANOVA, with site as a random factor and substrate type as a fixed factor. Data were log transformed prior to analysis. Post hoc comparisons were undertaken using a Tukey HSD test. Differences in the macroinvertebrate communities collected from the two sites by the three substrates were analysed using a two way crossed ANOSIM procedure® in the PRIMER software package’. Species which had the greatest contributions to any significant ANOSIM sample divisions were identified using the similarity percentages program (SIMPER) in PRIMER. The Bray-Curtis association measure was used for log-transformed abundance data. A significantly greater mean number of macro- invertebrates were collected from the Broken River than from the Broken Creek (F, .,=15.92, p<0.001; Fig. la). Significant differences were also observed among mean invertebrate abundance collected from the different substrate types (F2s4=48.50, p<0.001). The onion substrates collected more invertebrates than either the citrus or flywire substrates, whilst the flywire collected more macro- invertebrates than the citrus substrates. There was no interaction between site and substrate type (F25s=2.108, p=0.131; Fig. la). Artificial substrates collected significantly greater numbers of macroinvertebrate taxa from the Broken River than from the Broken Creek (Fj ss=16.312, p<0.001). There was no significant difference in taxa richness among substrates (F254=0.846, p=0.435) nor a_ significant interaction effect (F25;=0.084, p=0.919; Fig. 1b), ANOSIM results revealed differences in macroinvertebrate community composition between sites (R=0.981, p<0.001) and substrate types (R=0.240, p<0.001). SIMPER analysis revealed fifty percent of the dissimilarity of samples between sites was explained by the empheropterans Atalophlebia sp 3/13 and Koorrnoonga sp A2, and the trichopteran Ecnomus cygnitus Neboiss being found only in the Broken River; whilst Ecnomus pansus Neboiss was found only in the Broken Creek. The chironomids Chironomus sp. and Procladius sp. and the yabbie Cherax sp. were found in higher average abundances in the Broken Creek than the Broken River. A range of invertebrates explained the dissimilarity 260 (a) Site (b) Site Fig. 1. Mean (+SE) macroinvertebrate abundance (a) and taxa richness (b) for the onion (clear bars), citrus (diagonal lines) and flywire (vertical lines) artificia substrate fillings collected from the Broken Creek and Broken River. between communities found in the different substrates. Those that explained more than fifty percent of the dissimilarity for all three substrates at the Broken Creek were the chironomids Chironomus sp., Parachironomus sp. and Dicrotendipes sp., for which greater average abundances were collected from the onion substrates than either the flywire or citrus substrates. Chironomus sp. was more abundant in the flywire than the citrus substrates whilst the opposite was the case for Parachironomus sp. A higher average abundance of the ephemeropteran Tasmanocoenis sp. was found in the citrus substrates than either the onion or flywire substrates, whilst the onion substrates had a higher abundance than the flywire substrates. At the Broken River, the flywire substrates collected greater average abundances of Chironomus sp., Oligochaeta and the water bug Anisops sp. than either the onion or citrus substrates. For all of these cases the citrus substrates had a higher average abundance than the onion substrates. Greater average abundances of Atalophlebia sp 3/13, immature leptophlebiids, Eenomus cygnitus, and the beetle Antiporus sp. were collected from the onion substrates than either of the other two. The citrus substrates collected more of these invertebrates than the flywire substrates except for Atalophlebia sp 3/13, where flywire collected more than citrus. More psphenids were collected from the citrus substrates than the onion or flywire substrates, whilst the onion substrates collected more than the flywire substrates. The three artificial substrate fillings collected distinct macroinvertebrate communities in different abundances from within separate sites. However differences between samplers were less than differences between sites. Differences in invertebrate abundance collected from the three substrate types are most likely due to differences in the surface area available for colonisation. For example the close weave and coarse fibres of the onion-bag, combined with its double surface (total surface area 1.07 m~), provided a greater area of habitat for colonisation than either the single sheets of flywire (0.42 m2) or the open fine weave of the citrus-bags (0.60 m*). That there was a difference in the macroinvertebrate communities found to colonise each of the three substrates was unexpected, as in a study of four Australian lowland rivers very similar communities were found to be collected using either onion-bag samplers or 0.5 m lengths of Red Gum (Eucalyptus camaldulensis Dehnh.)*, a sampling device much less similar to the onion-bag sampler than either of the two alternative substrates used in this study. Although the substrates collected distinct invertebrate communities within a site they were similar enough to be able to differentiate between sites. If invertebrate abundance or community composition is the parameter of interest then different substrates should not be used interchangeably. However invertebrate richness can be assessed with any of the substrate fillings. This project was funded as a part of EPV 1 Monitoring River Health Initiative Victorian Programme b) evaluation of artificial substrate sampling with rapid bioassessment. Keith Ward, Rhonda Butcher, Garry Bennison, Tracy Clark and Alex MecNee are thanked for their help and guidance. Richard Marchant and Leon Metzeling are thanked for their constructive comments on earlier drafts of this paper. 'De Pauw, N., Lambert, V., Van Kenhove, A. & Bij De Vaate, A. (1994) Environ. Monit. Assess. 30, 25-47. *Hellawell, J. M. (1978) “Biological Surveillance of Rivers. A Biological Monitoring Handbook” (Dorset Press, Dorchester). ‘Humphries, P., Growns, J. E., Serafini, L. G., Hawking, J. H., Chick, A.J. & Lake, P.S. (1998) Hydrobiologia 364, 209-218. ‘Rosenberg, D. M. & Resh, V. H. (1982) The use of artificial substrates in the study of freshwater benthic macroinvertebrates pp 175-235 Jn Cairns Jr. J. (Ed) ‘Artificial Substrates” (Ann Arbor Science Publishers Inc., Ann Arbour). ‘Bennison, G. L., Hillman, T. J. & Suter, P. J. (1989) “Macroinvertebrates of the River Murray (Survey and Monitoring: 1980-1985)" (Water Quality Report No 3. Murray-Darling Basin Commission, Canberra). °Clarke, K. R. (1993) Aust. J. Ecol. 18, 117-143. ’Clarke, K. R. & Warwick, R. M. (1994) “Change in marine communities: an approach to statistical analysis and interpretation” (Natural Environment Research Council, UK). N. J. SOUTER, Floodplain Ecology Group, Department of Conservation and Natural Resources, Kaiela Research Station, Shepparton, Victoria 3632. Current address, School of Biological Sciences, Flinders University of South Australia, Bedford Park, SA 5042. nsouter@adam.com.au NEW SYNONYMY AND NEW NAMES IN AUSTRALIAN TENEBRIONIDAE (COLEOPTERA) BRIEF COMMUNICATION Summary The following new synonymy is proposed: Paratoxicum Champion 1894 = Schizophthalmotribolium Kaszab 1940; Mithippia aurita Pascoe, 1869 = Tarpela catenulata Allard, 1877; Cheirodes sardous Gené, 1839 = Anemia caulobioides Carter 1920. New replacement names are proposed to for eight homonyms: Cyphaeleus carter! n.n. for C. formosus (Carter, 1927) not C. formosus Westwood, 1841; Cyphaleus georgemastersi n.n. for C. mastersi (Macleay, 1872) not C. mastersi Pascoe, 1871; Nyctozoilus macleayanus n.n. for N. macleayi Carter, 1926 not N. macleayi (Bates, 1872); Nyctozoilus carteri n.n. for N. marginicollis (Carter, 1909) not N. marginicollis (Bates, 1872); Nyctozoilus geomastersi n.n. for N. mastersi (Carter 1909) not N. mastersi Macleay, 1872; Nyctozoilus obesulus n.n. for N. obesus (Carter, 1933) not N. obesus Guérin, 1830; Nyctozoilus jaystepheni n.n. for N. stepheni (Carter, 1914) not N. stepheni (Carter 1909); Ulomoides carteri n.n. for U. macleayi (Carter 1926) not U. macleayi Carter, 1926. Transactions of the Royal Society of South Australia (2004), 128(2), 261. BRIEF COMMUNICATION NEW SYNONYMY AND NEW NAMES IN AUSTRALIAN TENEBRIONIDAE (COLEOPTERA) The following new synonymy is proposed: Paratoxicum Champion, 1894 = Schizophthalmotribolium Kaszab, 1940; Mithippia aurita Pascoe, 1869 = Tarpela catenulata Allard, 1877; Cheirodes sardous Gené, 1839 = Anemia caulobioides Carter, 1920. New replacement names are proposed to for eight homonyms: Cyphaleus carteri n.n. for C. formosus (Carter, 1927) not C. formosus Westwood, 1841; Cyphaleus georgemastersi nn. for C. mastersi (Macleay, 1872) not C. mastersi Pascoe, 1871; Nyctozoilus macleayanus nn. for N. macleayi Carter, 1926 not N. macleayi (Bates, 1872); Nyctozoilus carteri n.n. for N. marginicollis (Carter, 1909) not N. marginicollis (Bates, 1872); Nvetozoilus geomastersi n.n. for N. mastersi (Carter, 1909) not NV. mastersi Macleay, 1872; Nyctozoilus obesulus n.n. for N. obesus (Carter, 1933) not N. obesus Guérin, 1830; Nyctozoilus javstepheni nn. for N. stepheni (Carter, 1914) not VN. stepheni (Carter, 1909); Ulomoides carteri n.n. for U. macleayi (Carter, 1926) not U. macleayi Carter, 1926. While the author was preparing an online checklist of the Australian Tenebrionidae for the Australian Faunal Directory, Australian Biological Resources Study, eight homonyms were found to have resulted from previous taxonomic mergers of generic names, as listed below. At the same time, the opportunity is taken to propose one generic and two specific synonymies resulting from recent examination of types in Hungary and Germany. The four-letter codens below refer to institutions in which types are deposited, as follows: AMSA — Australian Museum, Sydney; BMNH — The Natural History Museum, London; HNHB — Hungarian Natural History Museum, Budapest; MVMA — Museum of Victoria, Melbourne; ZSSM — Zoologische Staatssammlung, Munich. The author thanks Dr Martin Baehr at ZSSM for financial assistance and the Australian Biological Resources Study for supporting the work on Tenebrionidae. Synonymy Paratoxicum Champion, 1894: 380. = Schizophthalmotribolium Kaszab, 1940: 173. New synonymy. Type species: Paratoxicum iridescens Champion, 1894, by monotypy. Syntypes in BMNH, not examined. Schizophthalmotribolium australiae Kaszab, 1940, by monotypy. Holotype in HNHB, examined. Mithippia Pascoe, 1869: 293. aurita Pascoe, 1869: 293. Adelaide, SA. Holotype in BMNH, examined. = catenulata (Allard, 1877: 97) (Tarpela). Australia. Holotype in ZSSM, examined. New synonymy. Cheirodes Gené, 1839: 73. sardous Gené, 1839: 73. Sardinia. Type not located. = caulobioides (Carter, 1920: 224) (Anemia). Swan R.., Perth, WA. Syntypes AMSA, MVMA, examined. New synonymy. Homonymy In the following list, homonyms in Cyphaleus and Nyctozoilus result from the synonymisation of Matthews! of a number of generic names in the subtribe Cyphaleina (Heleini), while the homonym in U/omoides is the result of the transfer of Platydema macleayi Carter to the genus Ulomoides by Doyen et al? Cyphaleus Westwood, 1841: 43. carteri new name for C. formosus (Carter, 1927: 33) (Oremasis) not C. formosus Westwood, 1841: 43. georgemastersi new name for C. mastersi (Macleay, 1872: 287) (Chartopteryx) not C. mastersi Pascoe, 1871: 357. Nyctozoilus Guérin-Méneville, 1830: pl. 4, fig. 2. macleayanus new name for N. macleayi Carter, 1926: 511 not NV. macleayi (Bates, 1872: 276) (Hypocilibe). carteri new name for N. marginicollis (Carter, 1909: 136) (Aethalides) not N. marginicollis (Bates, 1872: 277) (Onosterrhus). geomastersi new name for N. mastersi (Carter, 1909: 139) (Byallius) not N. mastersi Macleay, 1872: 284. obesulus new name for N. obesus (Carter, 1933: 169) (Onosterrhus) not N. obesus Guérin-Méneville, 1830: pl. 4, fig. 2. jaystepheni new name for N. stepheni (Carter, 1914: 379) (Aethalides) not N. stepheni (Carter, 1909: 133) (Agasthenes). Ulomoides Blackburn, 1887: 274. carteri new name for U. macleayi (Carter, 1926: 68) (Platydema) not U. macleavi Carter, 1926: 67. 'Matthews, E. G. (1992). Classification, relationships and distribution of the genera of Cyphaleini (Coleoptera: Tenebrionidae). /nvertebr. Taxon. 6: 437-522. *Doyen, J. T., Matthews, E. G. and Lawrence, J. F. (1990). Classification and annotated checklist of the Australian genera of Tenebrionidae (Coleoptera). Invertebr. Taxon. 3 [1989]: 229-260. E. G. MATTHEWS, South Australian Museum, North Terrace, Adelaide, South Australia 5000. 262 REFEREES The Editor wishes to thank the following colleagues who have scrupulously refereed papers for the journal. He apologises for any inadvertent omissions. Without peer review, scientific journals could not function, yet this essential role frequently goes unacknowledged. 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