VOL. 121, PARTS | & 2 30 MAY, 1997 Transactions of the Royal Society of South Australia Incorporated Contents Smales, L. R. A revision of the Echinonematinae (Nematoda:Seuratidae) from bandicoots (Marsupialia: Peramelidae) - - - - - - Zeidler, W. A new species of freshwater amphipod, Austrochiltonia dalhousiensis sp. nov., (Crustacea: Amphipoda: Haale) from Dalhousie Springs, South Australia - - - - - - - - Bird, A. F. & McClure, S. G. Composition of the stylets of the aera Macrobiotus cf. pseudohufelandi - - - - - - - Bird, A. F. & McClure, S. G. Studies of the eggs of Macrobiotus cf. pace tnaie tis (Tardigrada) from wheat fields in South Australia - - - Kolesik, P. Two new species of Asphondylia (Diptera: Cecidomyiidae) from Halosarcia spp. (Chenopodiaceae) in South Australia - - - ~ - Jago, J. B., Tian-Rui, L., Davidson, G., Stevens, B. P. J. & Bentley, C. A Late Early Cambrian trilobite faunule from the Gnalta Group, Mt Wright, NSW Brief Communications: Jago, J. B. & Haines, P. W. Poorly preserved trilobites and brachiopods from the Kanmantoo Group, Fleurieu Peninsula - - - - - - Kemper, C., Dutton, J., Foster, B. & McGuire, R. Sightings and strandings of the pygmy right whale Caperea marginata near Port Lincoln, South Australia and a review of other Australasian sightings - - - PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 75 79 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 121, PART 1 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INC. CONTENTS, VOL. 121, 1997 PARTS I & 2, 30 MAY, 1997 Smales, L. R. A revision of the Echinonematinae (Nematoda: Seuratidae) from bandicoots (Marsupialia: Peramelidae) - = - 2 ~ “ Zeidler, W. A new species of freshwater amphipod, Austrochiltonia dalhousiensis sp. nov., (Crustacea! Amphipoda: Fopields) from Dalhousie Springs, South Australia - - - - Bird, A. F. & McClure, 8. G. Composition of the stylets of the tardigrade, Macrobiotus cf. pseudohufelandi - - - - - - - - Bird, A. F. & McClure, S. G. Studies of the eggs of Macrobiotus cf. pseudohufelandi (Tardigrada) from wheat fields in South Australia - - - Kolesik, P. Two new species of Asphondylia (Diptera: Cecidomyiidae) from Halosarcia spp. (Chenopodiaceae) in South Australia - ~ - Jago, J. B., Tian-Rui, L., Davidson, G., Stevens, B. P. J. & Bentley, C. A Late Early Cambrian trilobite faunule from the Gnalta Group, Mt Wright, NSW Brief Communications: Jago, J. B. & Haines, P. W. Poorly preserved trilobites and bende from the Kanmantoo Group, Fleurieu Peninsula - = - - Kemper, C., Dutton, J., Foster, B. & McGuire, R. Sightings and strandings of the pygmy right whale Caperea marginata near Port Lincoln, South Australia and a review of other Australasian sightings - - - 75 79 PARTS 3 & 4, 28 NOVEMBER, 1997 Bourman, R. P., Martinaitis, P., Prescott, J. R. & Belperio, A. P. The age of the Pooraka formation and its implications, with some preliminary results from luminescence dating - - - - - - - Vaucher, C. & Beveridge, I. New species of Potorvlepis Spasskii (Cestoda : Hymenolepididae) parasitic in dasyurid marsupials from New Guinea - “ : = 5 ? ; - - ~ + Littlejohn, M. J. & Wright, J. R. Structure of the acoustic signals of Crinia glauerti (Anura : Myobatrachidae) from south-western Australia, and comparison with those of C. signifera from South Australia - - Watson, G. F. & Gerhardt, H. C. The breeding biology and advertisement call of Litoria splendida Tyler, Davies & Martin - - - - - Barnett, E. J., Harvey, N., Belperio, A. P. & Bourman, R. P. Sea-level indicators from a Holocene, tide-dominated coastal succession, Port Pirie, South Australia - - - - - - - - - Gullan, P. J., Cranston, P. S, & Cook, L. G. The response of gall-inducing scale insects (Hemiptera : Eriococeidae: Apiomerpha Riibsaamen) to the fire history of mallee eucalypts in Danggali Conservation Park, South Australia - - - - - - - - - - - Olsen, A. M. An intensive monitoring study of two wetlands of the River Murray in South Australia; physico-chemical parameters and cyanobacteria concentrations. - - - - 2 . E - ? - Kolesik, P., Whittemore, R. & Stace, H. M. Asphondylia anthocercidis, a new species of Cecidomyiidae (Diptera) inducing fruit galls on Anthocercis littorea (Solanaceae) in Western Australia - - - - - Brief Communications: Wallman, J. F First record of the Oriental Latrine fly, Chrysomya megacephala (Fabricius) (Diptera : Calliphoridae), from South Australia - - Smales, L. R- The status of Cyclostrongylus medioannulatus Johnston & Mawson, 1940 - - - - - > = 4 . : < Invert to Transacnans af the Rayal Saciety of South Australia, Vol, 421, purts 3 & 4. 28 November, 1997 95 103 119 137 147 163 165 A REVISION OF THE ECHINONEMATINAE (NEMATODA: SEURATIDAE) FROM BANDICOOTS (MARSUPIALIA:PERAMELIDAE) By LESLEY R. SMALES* Summary Smales, L. R. (1997) A revision of the Echinonematinae (Nematoda: Seuratidae) from bandicoots (Marsupialia: Peramelidae). Trans. R. Soc. S. Aust. 121(1), 1-27, 30 May, 1997. The name Echinonema being preoccupied the genus here designated Linstowinema (nom. nov.) 1s redescribed. The type species L. cinctum comb. nov. is synonymous with E. meridionalis (sic) Chabaud, Seureau, Beveridge, Bain & Durette-Desset, 1980 but not with E. cinctum sensu Inglis, 1967, sensu Chabaud, Seureau, Beveridge, Bain & Durette-Desset, 1980. L. warringtoni sp. noy. is established for E. cinctum sensu Yorke & Maplestone, 1926 and sensu Chabaud, Seureau, Beveridge, Bain & Durette- Desset, 1980. Linstowinema inglisi comb. nov. 1s synonymous with E. cinctum sensu Inglis, 1967 and E. inglisi sensu Chabaud, Seureau, Beveridge, Bain & Durette- Desset, 1980 and is redescribed and four new species L. latens sp. nov., L. tasmaniense sp. nov., L. maplestonei sp. nov. and L. peramelis sp. nov. are described. Key Words: Linstowinema, Nematoda, Echinonematinae, Isoodon, Perameles, bandicoots, Marsupialia. Trasdetions Af the Ravel Sector cf S. Ame (P99T DAB 1-27 A REVISION OF THE ECHINONEMATINAE (NEMATODA:SEURATIDAE) FROM BANDICOOTS (MARSUPIALIA:PERAMELIDAE) by Lestrfy R. SMALES* Summary SMaAtrs. dR (1907) A revision of The Echinonematinae CSetratocdaSeuniidee) frony bandicones (Marsupiilie Perumehdaes. rans. Ro Sve. &. Ang VEC, 1-27. 30 May, 1997. The name &eiWienenia being preoccupied the genus here desiited Lins roveinenied (ner, nov) bs edeseribed, The type species L. cinciypr comb. ney. is synoayyious wath Eo prertdfeneléy (ate Chabaud, Seureau, Beveridge. Luin & Durelle-Desser, 1980 but nor with Ay eurecn seri Inglis, P9267, senaae Chbaud, Scurenu, Beveridee, Ram & Durette-Desset. l9RO. Lo werrinedoni sp. nuy. is established for &. oimenin sevisu Yorke & Muplesione, 1926 und sev Chabuud. Seureau. Beveridge. Buin & Duretie-Desset. 1980, Litisfeviiene ingtist comb. nay, 1s aYnonyMous with EL cient seas Inglis, 1967 and A. dedist seis Chubaud, Senreuu, Beveridec Bain & Diretle-Dessel, (SO umd ts redeserhed and four new species Lo fafeny spo nov, Eo feweuaitielise sp. nov, Le maplestane) sp. nov and 4. permimelis sp. nov, are described. All seven species can be differentiated as [ollows: fo crnetai 14-18 rows oF body hooks wilh unduluting edges, cesaphagus teeminacine atthe level of the sth tlth cows 4 ivahrinweni. 9-13 cows of body hooks with Gndidating edges, vesophagus terminating wt the Jevel of the Oh Wth row: 2. Jngliay, 1O-b rows of body hooks with undukiting edges, oesuphugus terminating at the level ob me Suh-Yih rows L. dares, YP fows of body hooks wih undulating cdes, otsephagis terminating posteriorly ta the Tbth row: L. daspreaniense, 1a1S rows hooks without undoliting cdes, oesophagus termimatiny at the level of the #th-LOth ew: 2 aaplestoel 12-13 rows of body hooks without undulating edges, LEXOpPhAZus termining posteriorly to Path tow ol heoks, 2, perdmeliy. 11-12 cows oF body hooks: without Unduliting cdees. oesophages termtinaung at level of the }2th row. A key ta the species ie given, Casiranarent AC OCs Lh Avadenobesilus, Lmaciaurus, Péruneles nasi and Bogut, Coidulio’ and 1. tasneditionae only ind obesitas, L. fates only inh nieeronens, boned and mec, bclesntis, dirty ak PB nevi, Loomupiestonen ind, digernrey and Poorer aod b peramelis onty in Ft howeaimytte Kei Wonns: Giafeinene, Newntoda, Eehinonenidnie, baade, Peraieles, handicuos Marsupiitia. Introduction Whet) Linstow (18984) described a thorny headed hematode occurring in uv bandicoat host, he called it Hoplocephatus cincius, He subsequently found thar Hoplocephalus was preoccupied. so he renanied the conus Eehinenem laer in the sume year (Linstow ISO8h) Eehineneaned has continued to be used for the genus anol the present. Kehineneme also is preoccupied, however, Having been used previcusly fol a penis of sponges by Carter in I88t) b now propose the name Lisroninema norm, nov The bandivoor hese orginally was identitied as Perneles abesilus, Linstow (1898a) deseribed and firured a pematode with seventeen rows of hooks on the cuticular dilation of the oesophagul region of the hody and Gesophuaus terminating on a level with rhe Oth row of hooks Yorke & Muaplestone (1926) desenbed a nematode with 12-13 rows of bocy hooks and a relatively short oesophagus which they identiiied us “AL einer”. They neither give meusurements nor provided a figure to show the relationship between the posterior end af the Hesaphacus and the rows of hody hooks af their Depertogor a) Hatayy, Cente! Oueonstornd Cniversiny Raekleanpren lel 4702 specimens. Yorke & Maplestone (126) assigned the type und, at that lime only speeies. 1. cient (Linstaw, 1898). te the spirurid family Rictulariidae Later Johnston & Mawson (1939) reported L. cinoma Tron a native cat, Dasvucks viverriiies, near Sydney and re-evaluated the available informiuvornon the host species, They coapeluded that Linstow's original descripuion wits based on muterial collected from Lseccton abesulus (Shaw. 1797) fron the Upper Burnett River in Queensland while Yorke & Maplestone’s redescription was bused on material from a bandicoat, posstbly fo maecreuriy (Gould, 1842). collected in the vicinity ef Townsville. Johnston & Mawson (1939) fierther concluded that both Linstow (7898a) and Yorke & Maplestone (1924) were describing material from the sume hast, namely. 7. mteerauray. Sinee 1 ebesuliy does not occur in the Burnett River region (Braithwatte 1955) but 4 meeronrus does (Gordon 1955), this would seen to be a reasonable conclusion, The first confirmed eccord of Liston inema frou the bundicoot genus Peraineley Geotlroy. 1803: ty by Johnston & Mawson (1940) Fron 2 nasite Cealiroy, 1804 collected near Sydney. These wathors noted differences. in the male tailor the specimens thev examined, Irom earher deseriptions by Linstow (18988) und Yorke & Maplestone (1926) but, nevertheless, assigned the specimens to Le cineniv, : Le. SMALLS OF the differences noled by Johnston & Mawson (1940). |he number ol papillae on the rule tail ts problemanc as neither they. Linstow 1 1898a) oi Yorke & Muplestone (1926) described the raniber and placement of papillae found on other specimens of Lrnsteowinemu, The expansion of the male badly annul the cloacal region described and figured by Johnston & Mawson (1940) ts alse) a Signitieant stctive, Although hot fentioned by Linstow (I898a). Yorke & Muplestone (1926) or Chabuud vr ot (1980) in their deseriptions of G2 eivernian. it ts computable to the nivale tail deseribed by Chabaud ev ul (LYS) for Echinenema sp. (fey occuring in # niatire Trem: ar unknawn touuhby, No further work was done on the genus until Inglis (1967) re-examined the relationships of the superfamily: Seuratoidey. He redeseribed L. e/neniay from material collected trom £ abestlus near Perth and placed the sole genus within a new subfamily. Ihe Eehinonemiatinge. lneated vwathir the seuratord family Schneidernematidae on the basis of the form of the moull opening. distribuljon of cephalic pipilae, long spicules and short guhernaculurn, Subseyuently. the allinties of the genus were clunfied by Quentin (1970) and the Echinonermistinie Included within the Seuratidac. Chabinud ef el, (1980) re-eximined all he avalible malerial. established two new venen) Seareokind Chabuud, Seureau. Beveridge. Bain & ure Desset, 1980 and Jielechind Chabaud, Serireaiy Beveridge, Buin & Duretle-Desset, [980 for wors from dasyurte marsupials and cedelined Echimeneeia tow Linstenvinena)y und £. einetune (te) Linstow. PSUS nee Inglis, 1967. In their desenpoon of LL. cineium they holed difheulnves in interpreting Linstow’s original Myure of 1R98a, bul decided that an oesophagus 18 mm dong agreed with their definition of a “long” oesophagus, that is, one terminating al the level of the most posterior body hooks. Linstow's (1898a) figure clearly shows. the oesophagus lerminanhy at the leyelol the th of 17 rows OF body hooks. Chabaud ef al. (1980) alse described three new species, namely FE. edinenlss (sic) trom a dasvurid, 2. navdronalts (ye) from obesuluy and Bo inglixi (ied = Eo ined seitsu luglis, 967.also from /, obésulay, These authors also indivated thal there were possibly addityonal species from the bandicool genus Peryimedes but they had insulficient material for detailed deseriptions. The bandicouts (subfiunily Peramelinge) ate rabbit-sized aminiveruus ofarsupiils with dau puinted heads unit compuce bodies. They forage by digging conteal holes with their short forelimbs and explote these holes with their pointed snouts (Gordon & Hulbert 1989), Perdmeles jieesata, the lome nosed bumdicom, 1s found alone te east cons! ol Australia, from rainforest in the rovth thposussty wetland and dry weodhand to areas walb little groupid cover in the south (Stoddart 1995). Irs distr7bution averlaps wilh A. drecieens. the turthern brow hundieaots which is found of the east coast north of the Hawkesbury River and veross the Morihenn Territory Lo the north ef Western Australia, in areas of tow ground cover weludios grassland. woodland and open forest (Gordon 1995). To the south, the distribution of P wayute overlaps thatol 1 vhesulies. the southern brown handicool which is found weress southern Austraha in Western Austria. South Australia and Victoria. sourhern coastal New South Vales. and Tasmania and prefers sandy soils with scrubby vevelition or low ground cover phit are burnt out fram Gime to ime (Braithwaite 1995), The cusiern barred handicoor, PR ennit Gray. Lo38. nw restricted to Tasmania, where vs disteibutfen overlaps with that of J, ebesulin, and a few peliet colonies in southern western Victoria, prefers open wrussland, bur may ulse forage in serub und heathland (Seebeck 1995) The western barred bandieool 2 bouceanville Quoy & Gaimurd, |s24. now exisung ently on Bernier and Dorre Islands Shurk Bay, Western Austiuia was previuusly found uvross mueh of the southern hallof Australia (Friend & Burbidge 1995), The only other bandigoat species sill extant, Loanredduy, the golden bindicoor, now survives only on Barron and Middle [slands off the coast bf Western Australia, the north western Kimberley region und sub-humid paris of the Northern Territory. having been previously recorded fram a much wider range of habitats (MeKenzie e i. 1995). In this study, all the available records nel rmtertal collected from peramelids, imeluding matey dissected [rom hosis beld in museum collections us Well ws live caught bandicours, have been examined. This hus provided sufficient material to reassess. the laxonomic Characters uvanable to use for species discriminaljon. redesignate and redescribe (he type species LE. einetum (Linstow. (898) comb, poy, = £: merdiengi« (vie) Chaband, Seureau, Beyenddye, Bain & Durette-Dessel, YSU. redeseribe und name fovorrargtae sp. nov. = L. carerune sensi Yorke & Muplestone, 1926; seni Chabuud. Seureuu, Beveridge, Bain & Durette-Desser, 1980 and 4, ingdiyé Chabaud, Seurcau, Beveridge: Buin & Duretle-Dessel, T982 and distingucsh the (oer additional new species front bandiceots that ane deseribed below, Materials and Methods Material and disseetion neeords from 719 Dandheoots were exananed, This matercal was derived from three sources. Firstly, the gisine intestinal tracts af $6 bandicools, catlected belween LINSTOWINEMA FROM BANDICOOTS Darwin OL. macrourus Mitchell Plateau a“ ———p, bousainville Perth Fig. | Melbourne = Cairns Townsuille Ponstel a Rockhampton AP eunnii PoaSttTe ah Hobart Present and former distributions of Australian bandicoots Gifter Gordon & Hulbert 1989), The symbols indieate the localities where bandicoots were collected between 1989 und 1996. Iscoden auras, L macrourus, @ Loobestlis i Peraneles bousiiiville, Po git, & Po nasiite, 1905 and LOSS and deposited in either the South Australian Museum (SAMA) or The Museuin of Victor (VM), were examined. These animals had probably been fixed in 5-10% formalin before being stored in 7O% ethanol, The nematodes dissected from these hosts were stored in 70% ethanol. Secondly. 79 animals were either collected us fresh roud kills or trapped alive. in spring-loaded wire box Wraps baited wath peanut butter or a peanut butter, honey and oats mixture, between 1989 and 1996. The trapped animals were killed by intraperitoneal inoculation of euthanasia solution pentobarbiione sodium (Nembutal @). The digestive tract of cach animal was examined under a dissecting microscope and any nematodes found were washed in normul saline, fixed in glacial acetic acid or hot or cold 10% formalin then stored in 70% ethanol. Alb the available specimens of Linsfowineme held in the Queensland Museum (QM). the Australian Helminthological collection of the SAMA (ALC). the Western Australian Museum (WAM), the CSIRO Division of Wildlife and Ecology (CSIRO) and The Natural History Museum, London (BM(NH)) were ulso examined. The preservation history of material from the former institutions is largely unknown but probably it was fixed in ethanol or formalin. Material from the CSIRO collection was fixed th hot lO% formalin. All material is now stored in 70% ethanol. Specimens were examined from all the extant bandicoot species (number af bandicoots in parentheses) from $1 localities across Australi fsooden auratus (5), 7 macrourus (1), 1 obestultes (85), Perumeles bougainville (13), Po guanii (6), P. nasuta (14), handivoot, no species given (9). Host 4 1... SMALES TABLE 1. Distribitian af bauadicaat species oxmmined fot Linstowinema species by State ec lecritary. Where iis specific Jacedin Hew bean given dale divseetion tecara dhe location is listed aw Austretia. Abibreviatinas Ay Auvtrelica, WAS Western Austria NT Northern Territory SA) denith Australi incduediny Kunseroo dvi und Pranklin Wleond> Q. Queensland: NSW New South Waless Vo Vietertay 2 Tasmenyre. Species A WA NT Dsencndenit cttrarns a - 3 Zo macrourus a 2 13 1, bbesulus 2 19 Peraneles baugainville 13 - : PL ounnir - - PL nasaret | - - bandicoat | 2| ri | lo Location SA Q v T - AU) \4 7 - 30) y i5 13 - _ - 2 4 - 7 4 2 - fl 8. n - - 30 65 27 V7 16 (istributions and the lowations of the 79 dtnimuls enllected since 1989 are given in Big. 1. Details ofall the localities where specimens were collected are listed in the deseriptions of species given below. Latitudes and Jongitudes are provided for all loculities that are listed in the Australian Gazeteer, The location, by stite, of all hosts species examined, is given in Table |, Specimens were examined after clearing im lactophenol or beeehwood creosote. Measurements were Wade with the aid of an ocular micrometer or drawing tube and map measurer, Meusnrements ire iver WT pas a rdnge from LO specimens followed by the inean in parentheses. unless otherwise stated, Alb the new material bas been deposited in the AHC. Comments on taxonomic characters Prior to L980 only one species was recognized in the genus Linstvavieme ie. Lehinonema enretine. Then Chabaud ef af (1980) described four species, three from perumelid hosts und one from the dausvurid, Deasvarns hallucatis Gould, 1842, The species aceurring in the dasyurid was distinguished from the others by having the first row of cephalic hooks longer than the second, The three species fron bandicoots were differentiated from each other on the basis of (he arrangement of hooks ahd spines on ihe body, the relulive sizes and positions of papillae op the clonal region. the relationship between the ventral spires aid pre-eloucal papillae, the extent of sind cuticular bosses surroundiig the cloud, Whe number of papillae on the tail oF the male and the length of the oesophagus relative 1 the hooks on the dilated cuticular partof the anterior body associated With the region of the oesophagus, Although Chabaud ef a (1980) recorded all the sensory organs on the mule tail tipas pairs of papillae tngls (1967) had noted pitirs of papillae and a pair ol phasmids. Thus Inglis (1967) reported 4 pairs of papillae and a parol phasinds on the tip of the tal of Eehinonema einer (sie) and Chabaud et al (1980) reported 4 pairs of pupilie for the sume species, Spicule morphology is uniform across the venus, differing onty iy total length and proportion of body length, Chabaud er ef. (1980) deseribed the oesephagus as enther “long. extending to the level of the last row al the body hooks or “short, terminating within the dilated cuticular region, Detailed examimilion ol specimens for this study has showe that alihough the lermdnation of the oesophagus relative to the surrounding hooks is consistent within each species, the aetual length of the oesophagus and its relationship to the body hooks may be difficult to determine, Speemens that are not camipletely straightened prior to or during fixation are problematic. [tis difficult to tell the extent te which the vesophigus may have contracted mie the neek reson and the cuticle surrounding (he oesophagus thay also be contracted, Furthermore, the range of lengihs of the oesephagus within populations ola species ean also he quite varnible. depending on tie sizes and maturity of the worms being measured. Both spines and hooks originate in the cuticle, but spines in this context, are defined ay being sal to Hiny und rootless, whereas hooks const of a thorn und a root anchoring the thorn within the cuticle, The smitllest hooks may be only slightly hanger than the largest spines, The relative lengths of the three rows oF hooks on the cephalic bulh ts a consistent character, but the dimensions af the hooks mity vary markedly between individuals of the same species from the same individual host, Therefore hook (limensions are noluselul for discriminating between species, The dimensions of the cephalic bulb are also LINSTOWINEMA VROM BANDICOOTS A unrelitble because of the potential variation caused by the extent ot relaxation of the specimens prior to fixation and the method of fixation. Other consisient chirueters at the species level, however, ure the number oF rows of hooks on the dthited anterior body and the morphology of these hooks. Ln seme species the roots of the hooks have undulating lateral edues giving them a “frilly” appearance: in others, the edves of the hooks are more or less plain, Key to the species of Linstowinema I, [st pow of cephalic hooks longer than 2nd row: PAUSES OP MUSYUPTUS oc ceercreernecen Leedmenielst And row of eephalie hooks longer than Pst row: purasitus OF bandheouts oe ee AY 2, Body hooks without undulating edges... 43) Poly Hooks willy Uniden CULES ce (OF ~ Ocsophugus terminutes posterior to hooks on cuticular dilation of oesephageal region: male will pairs of papillae, | pair ol phasmids on tail Ovsophigus fermimiles ih OF uNLeTIOr Lo pasteriar row OF body hooks on cutieular dikiion oF oesophageal reeion; male with 3) pairs ar pupilae, | pair of phasmids OF bil Upc et) Lo Mile with 8-10. demale with (0-12 body hooks: tnale with © pairs clowed! papillae all same size, body spines extend dong 800 of dorsal surface terminate at level of most anterior pair of lateral pre-vloweal papiihie on ventral surlaee 0. farens Male with (1-134. female wath 12-13 body hooks: male with 6 pairs of cloaeal papillae, | pair lateral ad-cloacal papillae lareer than other 5 pairs: body spines extend alone 75% of dorsal surface ferminate markedly anterior 16 clouval papillae on ventral surface 0 teplestenel 5. Male with alalike expansmons of body surroumding elowes with @ pats of cloweal papillae. 3 pairs of fateval cloacal papillae larger than 3 pains oF ventral clowcal papiliie. spicule leneth about 1/9 of body length, female wath tail longer than OQ. 0 cece be MANU te nse Male wilhoul alelike expansions of body, with 7 pairs clowcal papillae all same size, spreule leneth about Yio of body length; femule with tal shorter Un S00 va. perainelis 6. Male with 13-16. female with 14-18 rows of hody hooks; male with ala-like expansions of body surrounding cloaca, body spines terminiute atlevel of mostanterior pair of lareral pre-cloacal papillae on ventral SUmlaee cee L. einen Male with | 2. feniaie wilh FLL rows of body hooks; male without ali-like expansions oF body surrounding cloaca, body spines terminate inarkedly anterior to cloweal papillae on ventral SUIMPECE: scipernccuccupeesyesesece es: seereeecee crepes en reese 7) 7, Male with oesophagus shorter than 1570, termule With vesophagus shorter than L830. male with 6 pairs of clodeal papillae. purr of lateral ad-cloucal papiliae larger (haw other S pairs, 4 pairs of papillae, 1 pair of phusinids on Gal tip, spieule length ubour is of hody length: female with tail MMOMCT TAU OAD ccc seretecrensceeenese L, inglist Male with oesophagus longer than 1570, female Wilh gesophagus longer than |S60; rate with 6 pairs of cloucal papillae all sunve size, 4 pars of papillae. 1 pai of phasmids on tail tip. spicule lenvth about '/20 of body lengthy female with Kul Jonwer thin YSso Lo Warelietartt Systematics Order Aseurididy Super fhinily Searalores Family Seuratidae Sublimily Echingvemarinae Genus Lintowinesire Won, Way. Type species: LIST MEHL CHICTHA (Lap stow, L968 | comb, nov, Synonyons: Joplocepidus coictis Tinstow, USUs, Kehinonema mteriediovalis Chabad. Seurein, Beveridgs, Bain & Dureve-Desset, L980 nee Keinouena cinetunm send Yorke & Maplestone, 1920: Inelis, 1967; Chabaud, Seureau, Bain & Durette-Desset. 19X80 fin part), Liistowinema (\oln, Vey.) Genente HARMONIES Antenior end with cephalic bulb hearing 3 rows af I4-1 files of large hooks. Month opening Gtangulir i outline, without lips or lip-like struecures, with + pairs double cephalic papillae. | pair amphics, inner circle of sense organs on edge of mouth (see Inglis, (967. Figs 6. 7). Neck with 2-1] rows of very ting: spines. S-T8 rows wf 14-16 hooks surround an anterior culieultr dilution, assoeiuted with oesophageal region. Body with nurerous rows uf spines, miuhber of files of spines inereasing progressively towards mid body. decreasing Lawards posterior, continuing ta caudal ip of female, terminating at abou! '/3 of length dorsally, anterior to cloawwa ventrally on male. Short oesophagus simple, club shuped. surrounded by nerve cing at level of cephalic bulb. Deirids simple. conical. atlevel of [st row of cervieal books. Spicules lone. equal. identical, small gubernaculum present. Vulva at nvid- LINSTOWINEMA FROM BANDICOOTS 7 region of body; monodelphic ovejector directed anteriorly, Parasites of Australian dasyurid and peramelid marsupials, Linstowinema cinctum (Linstow, 1898) comb. nov, (FIGS 2-14) Synonyms: Hoeplocephalus cinctus Linstow, 1898a: pp. 469-471, Figs 3-11.) Echinonemia meridionalis Chabaud, Seureau, Beveridge, Bain & Durette-Desset, 1980: pp. 436-438, Figs 4, 5A, D; Spratt, Beveridge & Walter, 1991: p. 26. Echinenema cincta Linstow, 1898b: p, 672: Johnson & Mawson, 1940: pp. 473- 474, Fig 25: nee Yorke & Maplestone, 1926; nee Inglis, 1967: wee Chabaud, Seureau, Beveridge, Bain & Durette-Desset, 1980. 9 Echinonema vincta Mackerras, Mackerras & Sandars, 1953; p. 62, Echinonema sp. 2. Chabaud, Seureau, Beveridge, Bain & Durette-Desset, 1980: p. 438. Fig. SC, F: Spratt, Beveridge & Walter, 1991; pp, 23. 24 (in purt), Material exainined From /soodon obesilus New South Wales: 18, fragment Lismore, (28° 49'S, 153° 16'E), April, 1965, AHC 4413. 44, Timbillica State Forest, (37° 19'S, 149" 43'B), 15.xii1.1978, CSIRO N733; l4d4, 1922, Sidlings Swamp North, ‘Timbillica State Forest, (37° 17'S, 149° 45°F), 19.vii. 1994, 17.111,1994, 20.vii.1994, 23.11.1994, CSIRO N4213, N4074, N4228, N4075; 854d, 8822, Sidlings Swamp South, Timbillica State Forest, (37° 18'S, 149° 45'R), 14.vi1.1994, 20.vi1.1994, CSIRO N4212, N4230. From Victorias 26d, 149 2%, Melbourne, (37° 47'S, 136° 5Y’E), 91x.1991, AHC 30292; 229, Monash University, (37° 47'S, 136° 59’E), no date, AHC 30293, 30294; 26d, 229, Gorge Forest Road, (38° 21'S, 141° 36’E), Sept. 1962, AHC 30296; 26d, 11° 2, 7 fragments, no locality, no date, AHC 30295, 30298. From South Australia: 1d, 89 2, 6 anterior ends, Waitpinga, (32° 36'S, [38° 32'E), no date, AHC 4460; 33d 722, Scott Creek, (35° 04S, 138° 42'E), 5.x.1992, AHC 30291; 3¢d6, 399, Myponga, (35° 23'S, 138° 28'E), May 1966, AHC 4446; Kangaroo Island, South Australia: 1d. 29 2, Cape Willoughby, (35° 51'S 138° 08'E) 13.x.1990, AHC 30301; 29 @, Seal Bay, (36° 00'S, 137° 20'E), 41.1987, AHC 30304; 2d d, 22 2, Binowie, (37° 47'S, 136° 59'E), 5.viii. 1987, AHC 30303. From Perameles gunati Tasmania: 646 4, 142 9,4 fragments, Grove, (42° 59'S, 147° 07'E), AHC 30025; 334, 42°, Kingston, (42° 59'S, 147° 18’E), L1.vii.1992, AHC 30056, 30057, 30058. From Perameles nasuta: Queensland: 2° 9, Wongabel State Forest, (17° 20'S, 145° 30'E) 9.vii. 1982, CSIRO N1753; 12, Mt Nebo Road, (27° 23'S, 152° 47°F) tl.vini.1993, AHC 30316. New South Wales: 1c, Epping, (33° 46'S, 151° 05'E), 1 L.viii.1993, AHC 30316; 1& Epping, (33° 46'S, 151° 05’E). 14.vii. 1933, QM GL 12048; 344, 29, Sydney, (33° 50'S, 151° 15’), no collection data, AHC 1820; 19, Nadgee State Forest, (37° 26'S, 149° 54'E), 13.11.1978, CSIRO N493. Description Cephalic bulb with 3 rows of 14 (male) or 16 (female) files of hooks, 2nd row longest. 3rd row shortest (Fig. 3); neck with 5-9 rows of tiny spines; cuticular dilation of oesophageal region bearing 13- IS rows of 14 (male) (Fig. 2) or 16 (female) files of body hooks; Ist and last rows smallest, 4th-7th rows largest; roots of hooks with undulating edges (Figs 10, 13); remainder of body with up to 36 (male) or 54 (female) small spines at each annulation, over whole body of female; extending over */; of dorsal surface, terminating about 400 anterior to cloaca, almost reaching level of anterior pair of caudal papillae, on ventral surface of male body (Fig. 7), Ocsophagus simple, club shaped, terminating about level with 8th - 11th row of hooks, within the anterior cuticular dilation; '/s to '/j) body length (Fig. 2). Nerve ring surrounding oesophagus within cephalic bulb; secretory-excretory pore in neck; deirids conical, at level of Ist row of body hooks. Male: Length 12-22 (14.8) mm, width 460-730 (600). Cephalic bulb 260-490 (395) long by 325-420 (380) wide; cephalic hooks Ist row 145-225 (170), 2nd row 170-235 (200), 3rd row 104-145 (130) long. Oesophagus 1445-2040 (1790) long, cuticular dilation bearing 13-16 rows body hooks. Deirids 520-630 (590), nerve ring 320-420 (375), secretory- excretory pore 500-530 (550) (n=3) from anterior end. Spicules equal, similar, without alae, 935-1150 (1035) long, about '/i\4 body length. Gubernaculum short. simple, subtriangular 60-63 (n=5) long (Fig. 9). Nine pairs caudal papillae: 3 pairs ventral and immediately pre-, ad- and post-cloacal respectively, Figs 2-14. Linstowinema cinctum (Linstow. 1898). 2. Anterior end, optical section (lateral view). 3, Cephalic end (lateral view). 4. Cephalic end (en face view). 5. Cephalic end male. optical section at level of first row of hooks (en face view). 6. Female tail tip (lateral view). 7. Male posterior body spines (ventral view), 8, Vagina (lateral view), 9. Gubernaculum (lateral view). 10, Body hooks (lateral view). 11. Male tail (ventral view). 12. Male tail (lateral view), 13. Body hooks (lateral view). 14. Female tail (lateral view), Scale bars = 500 pm 2; 100 pm 3, 4. 5; 50 pm 6, 8, LL, 12; 25 um 7,9, LO, 13: 250 pm 14. x L. BR. SMALES | pair literal ad-eloacal, 2 pairs Jateral pre-eloacql: ill same size, 4 pais papillae, parr phasmids well posterior to cloaca. near tail tip (Fig, 1h), Cloacal resign with small cutieulur bosses; ahelike exprbsion OF body unterior and posterior to clone (iw, 11), Tail 310-450 (365) long (Fig. £2), Pemale: Length 16-22 (20) mun, wadtth 80021035 (RKO). Cephalic bulb 455-380 (505) lone hy 455-600 (500) wide: cephalic Hooks Ist row PRO-235 (210), Mel row 220-265 (245). 3rd maw 130-780 (155) long. Ocsophugus 1785-2125 (P9007) Jones eutieulyr dilation bearing 14-18 rows bedy beoks. Deirids W500 780 (G25), nerve cing 440-520 (470), seeretory- ckeretopy pee 520, S85 (n=2) from anterior end, Vulva 7140-10370 (S830) Prom aiterior end (Fug. 8), Vavinw aboul 300 long (i=1). Tail 985-1120 (1050) line t Figs 6. 14). Eves ovoid. 45-54 (48) log by 36- $1 (43) wide Type duane Perameles nase Geollroy, (X04 | ye ten “liny syeliey, Australia Sire an host Small intestine (ype specimen Neatype AHC 1%20 Remarks The present Ineation oF the material sleserbed by Linstow as Heydneephelns then renamed Lelvuey dent is inknown, Yorke & Maplestone (1926) and Chabad ef af (L980) neither gave a lecdtion tor the lype niuerial nor indicated whether they hie examined if, Linstow was working in Gottingen in 18908 but neither Mefinenema nor Maple lids ts fisted under specimens held in the Zeologishes Muscuin) der tlumbolt. Universitit, Berlin Ths Museum does, however, list holdings al other type specimens fran Linstaw, The specimens are not hela We (he parisite collections ef the BM UNH), the Inlernational Instinie of Parasitology, St Albans. or the US National Museunr Parasile Collection in Beltsville. The type host was given by Lamstow (1898a) as Perameles abexulus. one of iwo bandienat species collected by Richard Semon. These species were identitied hy Romer (1901), using the catagae of Thomas (1888) in the British Museurn, us Peveneler ohosulius, from Burnett River and Po pnete pure trom Couklown, with inedsurements being given for / meerura Perameles macrara is formdly listed as a synonym of fvacdon maerauins (Mahoney & Ride L988) but B obesilis is not. This is surprising hecuuse the taxongmie stitus of Po vbesalias was discussed by Maekerras & Maekerras (1960) who intheated that Perameles obese = Didelphix obesul = Tseddon abesulis, but that since the novtbern Laiutof disutbution af fseodon ubesalas is near Sydney, bumdicoals recorded iis A ebevuliy Gor Oneensland should be referred to us 2 meerouras, the species oecurring [rot north Queenstand te northern New South Wales Yorke & Maplestone (1996) Hat the type host as Peraineles abextla, dolinsten & Miwson (1930) decided that sinee Yorke & Maplestone tcl drawn cut original figure their material came From Townsville i) forthe Queensland and stared that Linatow's material from Jyaudon ebesday came tron Upper Burner River also in Queensland On that basis, Chubund een), (1980) voneluded that the host ot bath the Linstow atefiel and the Yorke & Maplestane material wis Dveeden meerouris, lhe northern brown and nit 4 abesilas: the southern brow buntigaut This does not, however, explant why Romer (M901) ia his denutication of the bandicoots eullecred: by Seynon Lists thent as two separate speeles rather thay us Perumeles macrira now bvoodon macrourus, The cHlalpeue of Thomas (LA88) however. does Tist 2 nasaia. so the Upper Burnen River bundicoots were probably 2. macrourus Chabaud ef ah (L980) noted That Les ion. the measurement given for te length of the oesophagus by Linstow (18980). was sinker to measurements ot specimens oxamincd by then that had heen collected fron) 2 mecrouras fron northern New Sotth Wakes (hroweh to nerthern Queensland. tn particular these specimens had the pesophagus lerminating wt uboul the level ol the posteriorend of the cuticular dilavon. 4 character they deseribed as a “long oesophagus’. Yorke & Maplestone (1926) did not indicate the lengil of (he oesophagus relative lo the rows of body hooks onthe cuticular dilation but they described 12 or 3 “eireles” OF 4 ip 16 rows of hooks, Two specimens eolleered by Nivoll in L915, currently held in the BM(NELY. which eouldl possibly be the fuiterial described by Yorke & Muplestone (1926) ahd vedeseribed below have the “long oesophagus” deseribed by Chabaud er al, (LOS) The specimens described hy Linstow (E898) ine drawn and libelled as haying (7 “circles” of hooks With Lhe Gesophages terminating awe the level of the Oth row of hooks. These churucters are consistent. net with LE einer seas Chabad eral (1980) but with specimens deseribedl by Chabaud er a/ (1980) as A. meridiaralis, occurring ta the southern browe bandicoot /xeaden obesuliy collected in South Australia, In a re-examination oF the material deseribed hy Chahaud ef al, (M980) as EL meridionativ., tovether alsa ' LINSTOWINEMA FROM BANDICOOTS 4 wilh speciinens collected fron long-nosed and hirred handicoots For this study. i) was found that all speciinens bad TetS rows ef body books aad) the ockoOphawos terminuted at the level of the Baie tl th row of hooks. Measurements of the oesophasts ramped front 1445 19 2041) for males amt 1785 to 2125) lor females, also consistent with the Measurements given by Linstow CUYosa). Johnston & Mawson (940) deseribed three forniles aid two ttales obtamied froin the intestine of the long oosed bandicaol Fo vex collected in Sydney and attibuted these to 2. cine They described ind figured differenves in the male tail. minely an expansion of the bady surroinding the Cloaca. similar to, bul not transparent, as are caudil ali Chabaud eral (1980) figured the posterior end ol aimale which they designated Fehinemema sp.2 from @ nasi registered in the ALC as (S20. They commented (hat their specimens were comparable wrth (hose described by Johnston & Miuwsen (1940), Neither sroup deseribed the anterior ends of the watts they cxaummed. The only material registered in he SAMA which might he the original Johnston de Miwsen specumehs is AHC [X20 The iorphology of the anterior. ends of these wornss,. |3- [8 body hooks, the oesophagus termining level will) the Ol) TLE row of books. is consistent with & mevidionalis seusit Chabaud, Searedu, Beveridge, Bain & Durelte-Desset, 1980, Chabaud erat (T9880) dit nol comment on the presence or ubsenee of an expansion te the Clowes on (hen Speconens bot contrast A. meridiomaliy with Cohinonemea sp, Disee Chahaud etal Vasu. py, 438, bie A) whiting thal Eefiinememe ap. 2. was comparable with the Tohnston & Mawsen (1240) desenpuun. Are exuniibition ob all the avartible specimens of Fe neridjenally (sie and Echinoneme sp. 2 (ie) Was fuled to shoy, aniy stntticunt dillerences beiweea them Sieh cifferenves us do eist can be anribited te the Rice chat Chabad en at L980) were doula Wil a mixed infeetion oF two species. namely L ciation and tL. weareiierane (described below) oveliceng Ta the material gollecred fron Wealtpinge ail Myponga, The male 2. cfietenn ask (ars popubiion were al the small end of the sive range und oesophageal length varies with work leneth and HHALLUDIy The déserption by Linstaw (PSUS of Ze cdots is congruent witht Hie revised deseripnog of muteyral desienaled L. merteivnlia by Chabiued of et. (TOR). The deseriplion by Yorke & Muplestone (1926) of specimens from h mecrinis und identified as &, cine. ts ebagrucnt with & carmen sensu Chabwud, Seurean. Beveridge, Buin & Durette-Desset, LOSO hee Linstow, PS9S, Theretore, | desienite a specimen fron ANC 1820 a the nearype of Lo efecnen Liiytowinemea cinch (Lanstow, 189Sh most closely resembles LZ. inylist (redeserihed below) in that the oesuphugus is celutively short in relation to the namber of rows of hooks. ending within the citicular dilation. The males of both species have 3 patry of caudal pupilae and spicules ‘ya - “/\= body length. Linsrmawinena cieenue con he distinguished frony 2. aiglive uy having 13-16 Gale) and [4-15 (female) body houks compared with 1-12 and 12-14 in L. frefisi. Although the ovsophugus ts cehibively short itis, however. longer than in 2. ing/isi, being fy body Jeggth in nmile Lo evenun compared with Wy, bady Jength in 4. delist. The pair ol lateral ael- cloacal papillae is bo Jarger than the other pains oF venteal pupillie in 2. eynedue but ws larger an 2. iuelis? (see Taglis WOT Pie, 9) The ale-like expansion of the body surrounding the cloaca af L, cliendn ys not found on 2. welsh (see Inglis 1Y67 Fig. 9). The body spines of L. eireiamn onty cover Ya ol the dorsal body surface bat cover |'7in of Lhe dorsal body surfiec ot 2, ineli’. Eggs of Lo emcaun ditfer from those of all other species in that they ure ovoid rather (hin almost spherical. The miternial (rom 7, oPeyalas from South Australia described by Chabaud ef ah (1980) as 2 meridionaliy was ound to be wniixed infection all. cincnint, Pemales with up to 18 rows of hedy hooks, odsaphagus terminaring at about the level of the 9th- Lith row)and 4 werrorgivn? Gemales with up to 3 rows of body hooks, the oesophagus termoiating about the level of the LOth -l3th row) deserihed helow. A conyparison OF the meusurentents piven by Chabaud ep af, (1980) ad {he speaiinens caamined for (his study reveal that the pnales (hey measured were smaller in size. the oesophagus, spicules: are ful were Shorter than in the specimens examined for this Study. ‘The females, however, were within the same size ringe ds for this study aud the comparative meusurements wre more consistent, The nuiterial dissceted from the scuthern brown hundicoot vollected it Seow Creek melided some females up to 46 mn Jong. These were larger than The specimens from Saul Australie studyed by Chabatider af (1980), (ip to 27h lowe), trap eustern Dirred bandicoots tram Tasmania (ap to 721nn long), und from long-nosed bandienols. from Tasmania. (30-32 mun fone) Other variations observed hetween feniale specimens collected in different hosts wid Jocalinjes in the present study and thase of Chabaud eb af Ch980) included the oesophagus fonvest in southern brown bundieaols Tram Seatt Creek (1853-23980), and shoriest in southern barred banediewots from Myponga and Waitpinga (F100): the tail Jongest in eastern brown bundivoots from ‘Tasmania (986-1122) and shortest in soothern brown bindicoors fram Seat Creek (S84-98O), Gd the vulva slightly mere Pastertar 10 L, k. SMALES (10,0380-14.450) in southern brown bandiceots from Seorr Creek than in eastern barred bandivoots fron Tasmanne (7140-10370) or southern brown handicnots From Wailpinga (10,800), The eggs from the Seott Creek specimens were smuller than those from Waitpinga or Tasmania, beng 33 by 36. 42 by 45, and 43 by 48 respectively. These differences could be either yvandhons befweer populations within the species. or the resull of contraction during: frxvation, Linsrowinema cinenun, originally deseribed as ovcurring iP. ehesuta tyre) Trony Queensland ts now recorded as also occurring inf ehevuluy, Pe nesuler and 2. guani, The geographical range new includes Vietoria, New South Wales. South Ausuibie meluding Kainigeoo Ishind, and Tasmania as well as northern and southertr Queensland. The record af f einem in L ebesulus from Lismore, New South Wales, ATIC 4413 1s a dubious bost record because the southern brown bundicoot ts net found newly as fur north as Lismore (Braithwaite 1995), In all probability the host was L wnererenrs. Linstowinema warringtoni sp nev. (PIGS 15-24) Synonyms: Lefironrene cineta sense Yotke & Mauplestone. 1926: pp. 347-348 nee Linstow, 1895; nee Trelis, 19672 yest Munday & Green. 1972: p. 1) (in party, sels Chabaud Seurcau, Beveridge. Buin & DuretleDessel, (980: p, 435-436 (in party, sensu Sprat, Beveridge & Walter, TU9T p. 25) Cin part, helinonema emer sensi Johnston & Mawson, 199) yy. a3 Material creniuned Prom dred aurains: ba 8, ne collection data ALC 30319. From (sondern meray Queenstands |. 17. Rollinestone. (19° 03'S, ld6° 24" Ra 24.1915. BM (NH) $450) 12.6.163-}665 Mossman to Daruotree Road. (16° 15'S, 457 19'B), 2.8 1991 AHO 30278: 2284, WV SY, Mossrrath, (16° 28'S, 149° 93'E 4, Wit b9S8, Sal T85s, Fand¥5h bait, PYAs, 12 PUSS. WAL ss. U4) 1958. OM GLI4351 GE 1436), GL43a63, GL.b6d. GLIAS65_ CL 14366. GL IS370, GL14372, GL14374, GL14377. GL.144s1 GL)4383; 264, 429, Caims to Mossman Read, (16° 58'S, 4S" 46H), 2a 1991 AHO 30274, 244, Gillies Highway near Lake Barring, (17° 15°S, 145° ARE) Ww 19901 AHO 30279 Goo, They, Atherton. (177 16'S. 145° 29"). 25,y, 1982. CSIRO Nigh 8s TES 2. Yorgubures to Atierton Road. (17> 1a’S. 145° 35", 29x 19YL AME 20278: Te of, 1} 2, Miviwinng. (17° 24'S. bbs 55 °b, 25.16 1957, OM GI1d368, ste, (892 2, Diddgee (17° 29'S 146° 00°R), QOGi,T8S& OM GE12653, GL4307; Res. 79 2, Millaa Millaa to Lonislail Read. (17> W'S. 14S" SPE) Shaw 1991, AHC 30276: Tod 2. 133-29) Innisbail, (177 32'S, 146" ONT). 16d 1959. 5.x 1953, 2ONTI9S6, 27-956, Ovi 1956. 19.yi 957. TG 1960, ALC 4528. OM GLI4457, GL14360, GLI4376, GLAS378, GL14379, GLI4382; 746 4,39 9. Atherion to Ravenshoe Road, (17° 46'S, 145° Q9'E), 3ELx 1901, AHO 30277; 16, 4%. Ingham, (18" 39'S. 146° 10"B), 31-x.199t, ALC 240; 25.0. 6% Y, Palmerston Aighway, 29,9. 1959. QM GL 14369277 °°, 60k south of Proserpine. (20° 14'S, 152° 357E) AHC 30266: 3© 4, Roekhumpton to Yeppoon Roud (23° US'S, 150° 44'E), Lo-viti 1990, Syii 1992. AHC 30271, 30267, 322. Yeppoon te Emu Park Road. (23° 10'S. 150° 46'R), 16i1x.1990 AHC 30270; 4a ¢, 15% 2. Rockhimptow to Emu Park Road, (23° 13'S, 150° 30°F) 17 411.1992, AHIC W272; 1, 24%. Rockhampton to Keppel Sands Road. (23° 20'S, 150° 48'R). Tait. 1992) AC A268: 69 9. Mt Glorious, (27° 21'S. 152° 54’E) 27.6. 1955. OM GL.L4R59: 692d, L204 F, Ashereve, (27° 27'S, 153°, 02H), 274.1956. OM GI14347, 1214 4. 20729. Paddington. Brisbane, (27° 28'S, 153° OV BR), 25.ni11 1955, 21,x%,1955, 14x, 1995, AHC 4371 QM GLI4350. GLI437l: 2945, Site Pp, Brisbane, (27° 24'S. 153° OIE). 28.48.1954: OM GL M4340, GLAda80: Ta a, P82. Mogeall, (27" 20'S, 152° 54’). 12.4.1967, CSIRO NIST, 2s, 705 Mt Nebo, (27° 3378. [52° STR), Tx. 1954, 23,y. 1994. OM GLI4359, AHC 30317. 3081S: Sef 139, Broukfield. (27> 30'S, 1S7° S5"b). 200.1973, My, 1967. AHO 19367, CSIRO NISI: (876,210 9, Rockles Crossing, Moy. 1867, CSIRO NIS52: New South Wales: 1. oa other ith, APIO 4462: Nu locality given: 97 4. 171 GS, SL 193s, Wax lG54, Mx 19Sd Bax ts. QW 45e. (2.795%, OM GL(4345. GLI4346. GL14352, GL14393, GLI4354, GLI4355. GL 14362, Pram /seadon obesitus New South Wiles: Te, Sidlings Swarnp North. Timbillica Stale Forest. (47 W'S. 149° 45’), TO) 1994 CSIRO N4228, Vielurias Tbe. JO 2 no eollectiow data, ALE 446|. ABC 30207: 66 4.39 9 Hulls Gap (377 08'S. (42° S1'hy ne date, AHO 30207) 94 49.32, Moura Reservoir, Grampians, (37° 148, 142° 30°), no dale. ANIC 30299. Sout Austrailia: 32 2. Mypemaa, (35 23'S. 13K° TS'R). no dae AHC +A. 395 | anterior end. Wuaitpinga. (35° A'S, 138° 42'B I nb date, AHO 4460. Kanvaroo Ishind, South Australie OF. Lot. Vivonne Buy, (25° 59°S, 137" 13°F), Lv.JO8S, AHC 30302; le 2, 39% 2, Parndana, (35 ATS. IAT? LSE 350986. ATIC 30307; 14d 4, 287%. Binnewie, (37° 47S. (36° 5S98'R), Soviit, 1987; ALC 30403, Prom Perankcles nasi Queénshunds bt 38S. Mareeba, (17° GO'S IAS” 26") 24a LOU). ALC 30036. LINSTOWINEMA FROM BANDICOOTS i] 4 * 4 ¥ * ¥ ¥ # Figs 15-24. Linstawimema warringtoni sp. nov. 15, Anterior end, (lateral view), 16. Cephalic end female, optical section at level of first row of cephalic hooks (en face view). 17. Female. hand cut transverse section through body hooks on cuticular dilation, 18. Body hooks (dorsal view), 19. Body books (lateral view), 20. Male tail (ventral view). 21. Female tail (lateral view). 22. Vagina (lateral view). 23. Gubernaculum (ventral view). 24. Female tail tip (lateral view). Scule bars = 200 por 15, 21: 100 pm 16, 17, PS. 19. 20, 22. 24: 50 pm 23. 12 LR, SMALES Deseription Cephalic bulb with 3 rows af 14 (male) ar 16 (enale) files of hooks (Pig. 16), 2nd row longest, 3rd row shortest; neck with a-7 rows tiny spines: cuticular dihition of oesophageal region bearing 9-13 rows OF J4 (mule) or L6 (lemale) files of body hooks (Pig, 17), Ist and last rows smallest, 6th - Sth rows largest, roots of hooks with undulating edges (Figs 18, 19): remuinderof body with up to 48 (male) or 54 (female) small spines at cach aonulation, over whole body af females extending over V/s of dorsal surface, terminating about S00, anterior to cloaca, not reuching Jevel of thost anterior pair of eaudal papillae, om ventral surface of mule body (Mig, 20). Ovsophagus, ‘io - Yin body length. simple, elub- shaped; terminating ul level of 9th- 13th row of body Hooks (Fig. (5) neat posterior end of anterior culicular dilation, Nerve ring surrounding. vesophagus within vephalie bulb; secretory- excretory pore On neeks deride conical, atlevel ot tsi row of body hooks. Males Length (5-20 (17.5) min, width 450-750 (505), Cephalic bulb 270-340 (295) long by 270-320 (205) wide; cephalic hooks Ist row 140-170 (155), 2nd row 160-190 (175). 3rd row 100-130-(1 10) lang Oesophagus 1575-1925 (1743) lone. colieular dilation bearing 9-11 rows of body hooks. deirids 490-550) (530); seerelory - excretory pore 360 (n=1; nerve ring 310 (n=l) fron: anterior end, Spicules similar. equal, without alae. G90-LO90 (850) long, about '/a body length, Gubernaculum short, simple, sub trangiilar, 50°75 (n=5) lone (Pig, 23), Teo pares viidol papillae: 3 pairs ventral and iinmediately pre-, ad- and posreloawal respeelively. | pair daleral ud- Cloaeul. 2 pains fateru! pre-eloucal all suime size: 4 pales papillie, pair phosmids posterior to cloaca near toil Vip (Fig. 20), Cloaeal region with small eutieu liar bosses, dlidike expansinns ihseal Mil 430-430 (90) long, Vemiale: Length 3241) (48) mm. widih 720-990) (edi) Cephalic hulls 450. 390 (470) long by 350-400 (280) wide, henring 32 rews al hooks, Ist rew 1 70- JHC 85), 2nd tow PHO-230 (95), 3rd pow P20. 150 (730) Jong, Qesophaiis TS70-2500 (2260) bans. vuteulie diladoo bearing |1-13 rows of bouly books. Deuirids S8O700 (O67): svererory-exerelory pore A7O-AOO) (390): nerve rings 200 360) (A301 drain uiferior end Vulva 12.5-05.6 (14.2) nim tron anterior end (Fig, 22). Tail 100-1900 (1265) long, (Figs 2), 24), Vagina about 300 Jong (n=l), Fags ilimost spherical 3044 (38) by 43-52 (40), Erynralagy This species is maned after Wirrnigton Yorke who with POA, Miplestane carried aut much af the early work on this ecnus, Type deeaulity Townsville, Queensland, Australi Type rest dyoedom macrourus (Ciguld, P42) Site in host Small intestine Type specimen Neutype BM (NM) 1950. 12.6. 163 Remarks The iwo worms, |. b¥. BM (NID 1950, 12.6.165-166 trom Perameles vbesula collected by Nicollin 1915 in Queenshind dre the only specimens registered i) Austalian or United Kingdon: parasite collections which could be the material deseribed by Yorke & Maptestone (1926). On examinution these worms were found to have the TL (imate) and ts (female) body hooks deseribed by Yorke & Maplestone (1926) for Behivenemea efor and te have the “long” oesophagus aiid all the other charaelers aliributed to 2 cineca by Chabad eat, (1980). J therefore designe the male speeiicn ul BM (N11) 1950, 12.6.165-166 us the notype, All the specimens ientiticd in this study as 6, warrington? conformed to the description siven by Chabaud ep al, (LORD) of Feiner. Adusrenvineniet warrington’ sp, nov, differs from. all other species i the genus in having Up to 9-1) rews of hooks (iale) or t1-12) (emile) on the oesophageal cunedta dilation amt a “long” oesophagus, thi as, the vesuphigus laemindtes: abo ee mew the fim tow ot body hooks. Male 4. weeengtoh can he distinguished fron L, civetiir by the combination of characters al (he posterior end. bey the terminating venteal body spies dir net extend te the most wotertor pare ol pupilhic in 2. wearing? buries orf Crean, whe disianve bepyeen spines yind clea is S00 pin not OO as tL. edaedan, the Tintited extent Of vuticular bosses surrounding the clowen in werrdigion’ compared with Le cinetan, bh, warrington? does net have an ulilike expansin surreundinyg (he clowea bul 2 cited does, and four pans of papillae vot three un the tal tip) The spietles of Lo werriitent foo body length ane shorter phan those of L. cinetane Vio body lenwih, “The opps al b. werrmeten; ire almost spherical hut tose ol 7. Curelae ure ovoid, Chabad ef af, (1960) in ther deseription of &, cineruny (sie) indicated (hab they had studied bumerous specimens from a range of localities including Woolwonga and Darwin in the Northern Territory. A re-cxuinioition of the imitertal from che Northern Territory has shown that ibrepresents a new LINSTOWINEMA FROM BANDICOOTS 13 species of Linstowinema, L. latens, described below. The measurements and figures of specimens reported by Chabaud ef al, (1980) however, are congruent with L. warringtoni rather those of L. latens, Spratt ef af, (1991) noted that there were no records of helminth parasites from /. auratus the golden bandicoot. The finding of L. warringtoni in one of five golden bandicoots dissected for this study is therefore the first record of a helminth from this host. Material registered in the QM as GL14345, 14346, 14352, 14353, 14354, 14355, 14362 was collected by Dr M, J, Mackerras. Therefore although no locality was given these specimens are probably from Queensland. The finding of L. warringtoni in 1. abesulus ts also a new host record. The specimens from Waitpinga, Myponga, Timbillica State Forest and in two of the hosts from Kangaroo Island were found in mixed infections with L. cinctum. The geographic distribution of L. warrington’ theretore has been shown, in this study, to extend from northern Queensland down the east coast of New South Wales to Victoria, South Australia and offshore to Kangaroo Island, A larger number of /. obesulus trom the southern states needs to be examined before ecographic distributions can be fully mapped. Populations of bandicoots in NSW, Vic. and SA, now have patchy distributions over a reduced range (Braithwaite 1995) and attempts to collect additional specimens of /. obesuluy for parasitological examination have been unsuccessful to date. Further work on the southern geographic distribution of L. warrington’ will be problematic, as bandicoots become more difficult to collect. The specimens of L. warringtoni found in a single P nasuta suggest either a natural low prevalence of infection, or an occasional, incidental infection of this host. Linstowinema latens sp.nov. (FIGS 25-36) Synonym: Echinonema cinctum sensu Chabaud, Seureau, Beveridge, Bain & Durette-Desset, 1980: pp. 435-436 (in part); sensu Spratt, Beveridge & Walter, 1991: p. 25 (in part). Material examined Type material: Holotype cd, allotype &, from lsoodon macrourus, Walsh Point, (15° 08'S, 125° 46'E), Mitchell Plateau, Western Australia, 23. vil. 1982; AHC 30322, 30323, 43d, 6592, paratypes AHC 13028, WAM 110-83, 116-83. Other material: From Isoodon mucrourus Western Australia: 1¢, 49 9, Mt Hart, Kimberley Ranges, (16° 48'S, 124° 55’E), 22.x,1993, 24.x.1993, AHC 30289, 30290. Northern Territory: 1634, 189°, Darwin, (12° 27'S, 130° 50'E), no date. 19.vi, 1993, 27.vi.1995, 28.vi.1995, AHC 4703, 30287, CSIRO N4413, N4414; 764d, 72 2, near Byers Rd, Stuart Highway turnoff, (12° 30'S, 130° 50'E), 18.vi.1993, AHC 30284, 30285; 7d d, 102 2, Adelaide River, Arnhem Highway, (12° 28'S, 131° 14°E), 20.vi.1993, AHC 30288: 13d d, 122 2, Bees Creck, off Stuart Highway, (12° 35'S. 131° 04'B), 17.vi.1993, AHC 30286; [1d d, 89 2. Jabiluka, August 1979, AHC 6421: 3° 9, Woolwonga, (12° 45'S, 132° 39'E), 19.x.1972, CSIRO N159. Queensland: 6d ¢, 112 2, Atherton (17° 15'S, 145° 29'E), 25.v.1928, CSIRO NI610: 11d d, 142 9, 14km north of Atherton (17° 15'S 145° 29'E), 15.41.1982, N1532: 1944, 129 9%, Yungaburra to Atherton Rd, (17° 15'S 145° 30'E), 29.x.1991, L.xi. 1991, AHC 30278, 30281; 1d, Gillies Highway near Lake Barrine (17° 16'S, 145° 35'E), 29.x.1991, AHC 30279; 64d. 82, Gillies Highway near Yungaburra, (17° 16'S, 145° 35’), 29.x. 1991, AHC 30283; 2344. 1169 2, Mareeba to Kuranda Rd (17° 00'S, 145° 26'B), 2.xi.1991, AHC 30282. Description Cephalic bulb with 3 rows of 14 (male) or 16 (female) files of hooks, 2nd row longest, 3rd row shortest (Figs 26, 27); neck with 2-9 rows tiny spines; cuticular dilation of oesophageal region bearing I4 (male) or 16 (female) files of body hooks, first and last rows very small, 6th - 7th rows largest: roots of hooks without undulating edges (Figs 29, 30); remainder of hody with up to 44 (male) or 60 (female) small spines at each annulation, over whole body of female; extending over °/jy of dorsal body surface, terminating about 300-400 anterior to cloaca, level with anterior pair of caudal papillae, on ventral surface of male body (Figs 32, 33). Oesophagus '/\y - 1/1) (male) ‘is = '/ie (female) body leneth, simple, club-shaped. terminating posterior to the oesophageal cuticular dilation (Fig. 25). Nerve ring surrounding oesophagus within cephalic bulb, seeretory-excretory pore in neck, deirids conical. at level of first row of body hooks. Male: (measurements of specimens from Western Australia, followed by measurements of specimens from Northern Territory), Length 15-18 (16). 15-21 (18.5) mm, width 425-625 (490), 515-715 (600), Cephalic bulb 280-350 (325) long by 280-380 (340) wide, 275-435 (350) long by 290-385 (340) wide; cephalic hooks, Ist row 150-180 (152), 105-165 (140), 2nd row 160-190 (168), 125-195 (155), 3rd row 100-110 (108), 72-117 (94) Jong (Fig. 26). Oesophagus 1200-1825 (1555), 1325-1990 (1715) long, cuticular dilation bearing 8-10 rows body hooks. Deirids 540-690 (610), 390-650 (530); L. R. SMALES AA’ “ean Figs 25-36, Linsrowinema latens sp. noy. 25. Anterior end. (lateral yiew). 26. Cephalic bulb (lateral view). 27. Cephalic end female, optical section at level of first row of hooks (en fuce view). 28. Vagina (lateral view). 29. Body hooks (lateral view), 30. Body hook (dorsal view), 31. Female tail (lateral view). 32. Male posterior body spines (ventral view), 33, Mate tail (ventral view), 34. Female tail tip (lateral view), 35. Male tail (lateral view). 36. Gubernaculum (ventral view), Scale bars = 200 pm 25, 26: 50 ym 27: 100 pm 28, 31, 33, 35; 25 um 29, 30, 32. 34. 36. LINSTOWINEMA FROM BANDICOOTS (5 secretory-excrerory pore not scen, 230-495 (305): nerve ring 300 (n=1), 225-365 (275) from anterior end, Spicules cqiil similar, without alae. 830-1090 (995), SAN-1100 (1005) Tong, about Yin to “is body leogih, Gubermaculum short. simple, sub-triangular. O1-68 Ure3). 66-79 (n=) long (ig. 36). LO pairs miudal papillies 3 pairs ventral and immediately pre-. fd and post-clodcal respeetively, | pair lateral ade chougal 2 pours Tateral pre-eloacil, all sume sizes 4 pairs papillae, pair phasmids. alaike expansions ibsent posterior to Clowes. near dul ip (Rig, 33). Clowcul region with siiall cuticular bosses. ala-like expansions absent, “hal 250-340 (31), 255-320 (305) long (ip, 35), Pemate, Length 20-38 (30), 25-41 (30) mim. widih 40) 750 1590). 665-1175 (840). Cephalic bulb 300- 380 (350), AOS TO (ASS) hone by 350-500 (385). S7TS-5A0 (430) wide, bearing 3 rows ol frooks, lst row LA0-T90 (175), P3S- (90 (155) 2nd row 170-190 CIRO) 40-195 (170). 3rd row TO0-1 200) EO, 80-1 35 (100) long. Oesophasus 1450-2075 (1790), 1S05- 3430) (J970) long: cuticuhir dilation hearing 9-17 holy hooks, Detrids 600-700 (660), 325-770 (405): seerelory-excrelory pore nol seen, 295-590) (420); nerve fing not seen, 205-405 (3455) (n=5) from anterior end (Pig. 28). Vulva 2-147 (Ee) ¥2- 17.2 (11.9) fin froin anterior end. Vagina about) 15 (n=2). Eves almost spherical 36-00 (50), 33-42 (35) by 34699 (34), Til THOT 210 (980), 920-1615 (1160) home (Fins 31. 34). Enyinaaleus The species nanie is derived tron the Latin fareny. mening Hidden. since ib wis nel found wher (he materi wits first examined, Type hese Jyoudor macrourus (Gould, e425 {ype lowelity Mitchell Plateau, Western Australia, Australia Nite tee dake Small iitestine (ype specimens Holotype mule. ALC 30322. allotype fentile. ATIC AQS23, parutypes ATIC 13028 Reimarks Linstowpreme fares sp. nove resembles L. WrPowonaise oceurriig inh daerouriy, in bein of siomihie sizes Qnales S21 mim in Lo deateny compured: with 15-20 nim dong in Lo werringieni), having the oesophagus of similar length (1575-1925 in Lo hens vounpared with 1200-1825 in L, warring males) and fodr pais of papillae Gn the inale Gul, Phe agsaphiaguis i 2. laren. however, terminates posteriorly to the hooks surrounding the oesophageal region whereas that of Zo wereiagron terniinittes at about the level of the Yih ~ 130h row of hooks. L. dates hus 8-10 Gauley or 9-12 (emale) rows Of hooks, roots without undulating cdees, while Lo werringtent bas 91) (male) or 11-13 (female) rows of hooks. wilh toots having cinching edges oo the dikited cutie region, The body spines on the dorsal surface of male 0. fates extend further Jowurds the posterior end, Giboul aor the body), than ond. warriveron’ (about “Vs ol the body) Ventrally the body spies extend to the sume level as the inost amerior pair of cundual papillae oud, laren, but de pot on Lo warhead The male tail is longer aL. waertigtent (A30-430) than in Le etsy (250- 340) The vagina of ho haens (115) ts shoner than thiol. terraced (300), Specimens were found ind prevonras Tron wortheryy Western Australia, the Northern “berritory vod northern Queensland, The population of nerthern hrawa bandicoots in Western Australia is salted from that of the Northern Territory and Queenslind (Gordon 1995), bib measurements of worms from hosts occurrii fh the Koinberley, Western Austrati. are consistent with those (rom Darwin. Northern Territory. The only niorphological difference observed between these pupulations was that the first row Of cephalic hooks of the Kimberley specimens Was alvest as long as Te second ri, ISQ-180, compared with 160-196 in males, bul in the Darwin specimens the difference in length between the Iwo rows of hooks was more murked. 105-163, compared with 126-195, This difference is got considered to be significant and us the Queenstind worms were similar to those from the Norther ‘Verritory. the material from all three localities ts coustdered to be vonspecilic. The northern brown handicoots colleeted From horthern Queensland were infected wath 7, Warn S lrosts, Lo laters. 4 hosts. ot bow specues, 3 hosts. Chabaud ef af, (1980) wleatitred all the material they examined from the Northern Territory and northert) Queensland as 2 edverane. now 2. Wwercinglonk Was not possible to determine from their paper which, Wo any, specimens fron northern Queensland, presently lowed in the OM, SAMA or CSIRO collections, they examined. The specimens they examined trom Woolwonga and Darwin in the Northern Territory lave been re- examined for this study, and ave all datens, The Queensland material exumnbed by Chabaud ey ef. (M980) could hive been either 2. Werrington 1, fatens, or both, L.R. SMALES 4| 40 LINSTOWINEMA ROM BANDICOOTS 7 Linstowinema inglixi (Chabaud. Seurcuu, Beveridge, Bain & Duretie-Desset, 1980) comb, mov (FIGS 37-47) Synonyms: Lehinonemed eine sensu Unglis. 1967: py. 122. (28, 131-134, Figs 8-10, Eehinonenne aeglist Chobaud, Seureau. Beveridge. Bain & Durette- Desset 980: pp. 437438. Spratt, Beveridge & Waller, (994) p. 26. Material examined From Jyoodan obesulus Western Austrailia: 447. 39 9 Murdoch, (ale 37'S. LIS* STR). 27vA8Sh, AHO 84901; 402-4, 10649, Perth. (34° 57'. 115! Sth, Jan. 1993, 95,1993, 311.1993, 1995. AMC 40257, 20258. 30259, 30260, 30262. 20263, 30264. 30005: Lhe ot. 44-29. Wartle Grove (3202'S, Elo" QO'E), 3.y, 1906. BM (NH) 1967. 616-626. WAM 25- 7; We 2. TYP, Glow Forest. (Oovitt.6@. ATIC 2u7A0, BM (NH) 1967, 473-523: 4¢ a. 109 9. Horesulule, LSI 1978, AHO 8885S; 40d 2. 108 Jorrahdile, (AZ) 30'S. 16° O7'L, 26.x.1993, ATIC 30261-44489 4, Albany, (357 00'S, 117° 52°B), oxi F944. TONE 9d CSIRO N4242_. N4243: Wad. 322%, Manjimup. (35° 15'S, 1160 09"). °7.vi. 1903. 16.6x.1993, AHC 30295, 30256: 625 4, IAS] 3, no locality given, 22.68.1977, AHO Sd8s Descriplton Cephalic bulb with 4 rows of 14 tale) (Figs 38. 49) or 16 (female) files of hooks 2nd row longest, 3rd row shortest, neck with 5-11 rows tiny spines: cuticular dilation of oesophageal region bearing |) i4 rows, 14 (mide) or 16 (emule) files ot body hooks. [st and last rows smallest, 7th - 9th rows hooks Jargest (Pigs 3, 7), roots of hooks with lindulating edges (Figs 40. 41). Remainder of body with up to 35 (inale), or 45 Gemaley small spines at euch annulation, over whole body of female, extending to 400 yim from tail tip Gn dorsal surlace and ubout 300-500 anterior to cloaca, not extending to anterior pair of caudal papillae, on ventral surtuce of ynule body (Pig. 47), Qesophagus '/11 (mile) to H/ja (female) bedy length, simple. club-shaped. lerminiting level with Bth-9th row of hooks. Nerve ring surrounding oesophagus within cephalic bulb. seeretury-exeretory: pore in neck. deirids comical. at level of Ist raw of body hooks. Male: Length (1-L8 (15) mm. width 325-625 (540), Cephalic bulb 266-325 (272) long by 247-350 (309) wide; cephalic hooks, Ist row 140-180 (155). Ind row 160-200 (180), 3rd row 115-140 (125) long (Pigs 3, 7). Oesophagus 1120-1565 (1400) longs culicular dikition bearing 10-12 rows body hooks, Deirids 410-650 (520); seerctory-caxcretory pore nol seen: nerve ring 412 (H=1) from) anterior end, Spicules similar. equal, without alice 700-7200 (985) long. about ‘y= body length. Guberniculum short, sub: tritngular. 90-58 (n=5) long. (Pig. 45). Nine pairs caudal papillae: 4 pairs ventral and inimediately pre-. ad- and prost-clowcal: | large pair lateral, ad-cloaeal. 2 pairs dateral. pre-cloacal; 3 pairs papillae, pair phasmids well posterior to elowea, near bul lip (Fig, 47). Cloucal region with small cuticular bosses, ala- like extensions of body absent. Tail 150-350 (205) lony (Pig. 460). Female: Length 15-30 (24) nim, width 500-935 (700). Cephalic bulb 312-357 (334) long by 293-422 (334) wide; cephali¢ hooks, Ist row 155-195 (180). 2nd row TR0-235 (205), ard row | 15-150 (125) long. Ocsophagus I385-1835 (1655) logs cuticular dilation bearing 12-14 vows body hooks. Deirids 485.570 (S45): secretory-exeretory pore 390 (n=l) frony anterior ends herve ring, sol seen. Vulva 9 &- (0.2 mm (n=2) (Pig. 4a) from anterior end. Vaya about 350 (n=L). yg almost spherical 40-48 (441) by 36-45 (40), Tail 610-920 (730) long (Pigs 42, 44) Type hose Jyvodon obesuluy (Shaw, 1797) Tyne foceltty Waitle Grove, pear Perth, Western Australia. Australia Site i hast Small intestine Type specimen Nestype BMINH) 1967 616 Remarks Material from /. e@hesifuy tron Wattle Grove. collected on 5.v.1966 und identified by Inglis ts Figs 37-47. Linafeavinemer taglist (Chabad. Senreau. Beveridge. Bain & Duretic-Desset, 1980), 37, Anteriun end, (lareral view), 38. Cephalic end (ea face view), 3 Cephalic end nile, optical section at level of first raw of hooks ter face view) 440, Body hook (dorsal view), +1 Body hook (lateral wiew), 42. Female tail (lateral view). 13. Vieina (lteral view). 44 Female tail tip (literal view). 45. Gubermaculum ¢yentral view). 46, Male tail (htteral view) 47, Male fil (ventral view) Seale bars = 200 yon 47, 42250 pm 38, 39.43, 44, 100 pin 40.41, do. 47. 25 pm +5. Is 1. ROSMALTS deposited in Wie WAM and BM (NE). This appears lo be the material deseribed as 2. cineuun by Inglis (1967), T therefore designite a specimen trom BM (NH) 1967. 616-626 as the neatype. Linnie dteliyi — (Chabauidl — Seureaut, Bevenidee, Bai & Duretic-Desset, 980) can be distinguished front L, werrtaetoud and Lo lens in having (he oesophagus terminating within’ the wileror coliculer dilution. Linstowineme inglisi has 1-12 (rate) and 124 (emule) rows of hody hooks compared with 81 and lbela rows for o., weeringwo) and SAO and 8-12 rows far do lererey. The spicules of £. felis, (Wis body length) ure relitively longer than for Lo weevingtond (oy body length) and 4. deren (iy body length), The male tuil al Lo jnglisi also differs fom both Lowarringions ane Hh larevis i being shorter 150-350 (265) compared With AXD-130 (390) ond 250-340 (407) respeetively vod in having only 3 pairs of papillae rather than 4, The female tal is also shorter (10-920 07300) in L. justivicompored with L. werriaatoi and Le hares (1000-1900) (1265)) and 7916S (L070). respectively, The pairof lateral pupitlic level with the cloacal ppentig is more prominent than the other pairs of pupillue surrounding aod anterior to the chouce of Z inglisi, Wi (es pespeect Lo dawlist resembles 2. edimendst trom dasyurid: nursapials but 2. daelisy differs trom &, edmeaney in having the 2nd. not the Ist row of eephulie hooks the longest, Lovstawinen cdinands? oceuts i Dayyerny falluveins trom ale Northern derrmtory while 2. daelivi oecurs ti 7, vheyuliy from southern Western Australia. Linstowinernes (iglisi can be distintished from L, cena, Which alse beours in abesulits and has the Oesophagns termining within the cuticular dilation OF the oesophageal region at about the 9th row al hooks. by (he number of rows of body hooks, 10-12 (mle) amd 12-14 female) compared with 13-16 and Id-I8 in Lh. eimenin. The differenees between J, clon anid O. ediai are discussed i) detail under /. CTCL, The measurenents of L. dagtiod trom this study are waiiuent with those give by Toglis (1967). Any differences belween the Iwo sets Ol imusurenents are because Inglis (1967) measured smaller woriis, 329-11 39 Fur the males compured with [1-18 inthis study and 11-184 for the females conipared wilh IS-AU in this siidy: Chabatid ep al. (1980) described L. Inglis) mules as having spines covering only 2 ol the body dorsally. Ao careful examination of specimens for this study, however bas shown tbat ibe dorsal spines, allhough tiny. extend alone about 2% vl hady, that is. further towards the tail tip than de the ventral spines. Linsewinemd inelisi has been found only in? Obesulas fron the south of Western Austeatia, Linsfrowineme taymantonsé sp.nov (PIGS dh-61) Synonyins) Echinoneme cinetuin sensu Munidiy & Green, M972: p21 Cin part, Aedinonemed tirelivi wi Sprit. Beveridge & Waller, 1991; p. 26 (in purty. Lelinonemea spt Chabad Seureau. Beveridge, Bai) & Duretie-Desset LOO: p. 453: Spratt, Beveridge & Waller. 199}: ps, 26. Material examined Type material: Lolotype oo, allotype '. tram Iynodon obesulios, Kingston (4h2" 50°78, JAN Rob, Tasnminits Thi (992, AMC 30820 40R71 Parutypes [209 EES WARE AOS10, 30411 Oren material: Fron lyovdemr obeadlus South Australia, Kangaroo Isdands bite, 499. Viviane Bay, (35° 59'S. 137" (312), no date. Lav LOSS. ABC 4458, 30302: 133 2. 2492. Hundred of Grosse, Tune 1983. ATIC 30305: S28 172), Kacatta, (a5° 59'S, 136° 56"), Sept, 1983, ALC 20300; [3.42 7- IS 9, Pulndanw (3S 47'S, 137° LOR U2iv, oud S.X1 1986, ANC 30300, 30307; bet. 12) Seal Bay, (36° 00'S, 1378 QO. 4atUs87s AH! 320304 Tusmania: $3 % 1529 Beuconsfield, (41% 12'S, 46" 49'E), 1991 AME 30315: 34 4, Glengarry, 1" 21'S, [46° 52°B), 1992, ATIC 30304 J. Hobart, (42" 53°S, }47° JOR) 25. yi 1982. CSIRO NI674;, Fad. 8. Gow Runge. (42° 50°S, (47! PRED, Suivi 1992. AH 303808; Bsa, 139 2, Margate tip, (43° 02'S. 147° 16°E). (Oj 1993, AC 30312: Loh. Upper Dromedary. ne date given, AHC 4530, 1244, Ye 9) hoe collection data, ATC 40313, 30344, Deseriplion Cephalic bulb with J rows ol [4 Ginaley av to (female) (Figs 49, 50) files of hooks. 2nd raw lonsest, id now shortest (Fig. 49). neek with Ss rows liny High 44-01, Aiaienydaenn femranieive sp. now. 48. Anterior end, optical section (literal yew). 10 Cyphalic bull cuter) view), 90, Cophili¢e end feniile. aptical scetion al level at Hist row of hooks (em fee view). 56. Fentale unl (ateral view). 52. Mile posterior body spines (ventral views 53, Body louks Cateral views, 54, Bony hooks (dorsal views S35. Vagina Hater view), 56, Male til (ventuil view). 57. Gubernaculunt (vental view), 58. Kenale (ail tip Uateral view), 54 Male Tai) Up vente view), 60. Male tail tip (ventral view), 61 Gubernactdun Uateral view). Scale burs = 300 yim 44. S12 200) Jr OOO, TOO pn SO, SO pan 92, 938, 54, 55. 5A: 75 pom SF. 58,50: 12 fH OL aly) I. RO SMALLS spines; Cuhecular dikition of oesophagenl region bearing 13-15 rows af L4 (male) or 16 (female) files ol body hooks Tstauel last rows smallest. 9th Oth rows lurgest: moots of hooks without undulating edpes (Pigs 34.54 neotainder of body wath up to 4h (mule) or 66 (emale) small spines at each annulation, over Whole body of female, extending over “/) Of dorsal surface, terminating about 350 unterior to cloaca, not reaching level of most anterior mae Gt caudil papilhie. on ventral surface of nile beady (Fig. 56). Oesephugus about Mois body leneth, simple, Clobeshaped, (cemimanng at level of Sth-TOth row OF body hooks (Fig. 48). Nerve ring sunounding cesophagis within cephalic bulb: seUreiory-e8crelury pore in neck: deinds conical at leweh or bat pow ot hedy hooks. Mile, Length [le T4 (12.6) jam, wilt 453-715 (605). Cephalic bulb 215-295 (255) long hy 270-390 (225) wide; bearip 3 rows of hooks, Ist raw L4U- 195 (170), Ind row 175-225 (200), 3rd row LOa-130 (120) long thie, 49), Gesophagis 1105-1580 (1380) Jong; cubeular ditaboo bearing 12-14 rows of boty hanks; deirids 355-480 (430). secretory-excrelory pore 300-440 (365) (n=O); nerve cing 240-300 (2801 (n=4) from anterior ene. Spicules snvilar equal. ayittioal alive 720-1030 C890) tors, about /)) body length. Gubermiculim short, simple, sublriangalar, S0-55 (n=3) Jong (Pigs. 57. 61). Ten paies caudal papillae: 4 puirs ventrul and immediately pre- . ad- and post-eloical respectively, Lo pair heer! ad Slowed, 2 pairs hicral pre-cloweal (Pig, 50): 3 hiteral pairs larger (hig, 56); 3 pairs papillae, pair phasmiads posterior to cloaca. near tail tip (Piz. 39), Cloueal region with small cuuucular bosses. ula-like expansion of body surrounding cloaca, Tail 235-325 (285) long (Fig. 60), Female: Lengthy 19-22 (21) moi, width 8TS-1155 1965). Cephalic bulb 195-325 (255) lang by 470-305 (460) wide; bewing 3 rows of hooks, Loi row 190- JNO (205), Ind rew 220265 (245). 3rd row 125-170 (150) long. Oesophugus 1445-1990 (615) lone. coliculur dilaGon beanie [4-75 rows body hooks, Deirids 355-450 (405); sceretury-excrelory pore 375 jn=1) [root anterior cna: nerve ring pot seen. Vulva 3525-7600 (63500) (n=3) long (Fig. 55), Vayina 175 (n=1) long. "Tail 715-935 (810) long (Pies SL, Sa). Leyes almost sphericil 36-48 (42) by 33-45 (48). Remarks Linstawineme wisnniieuse sp. nov. resembles 1. inelise and L, etmeram. alsa eecurring iL eheslas id hawviog three pairs af pupiiae on the male tual snd the Gesophagus terminating within the cuuculiu dihition af the vesophageal region, at about the sth - Lith (ow of hooks. Listawearenica Lasmeainiense can he differeritiited font 2. reelisr in having $2-13 (male) and 14-15 Gemaley raws ot body hooks Without Undulating edges, compared with Le} 2 (male) and 12-14 (female) rows of body: hooks wath undulating edges and fram 2. cine whieh has 13- 16 (nme) and Les (lemme) rows af body Hooks With undulating edues Linger fisinaiiedse huirther differs from Z. veglive baying all tiree pats Of papillae literal and anterior to the clowea, larger than those surrounding the elouca. ane in having the vloacal region with ula-like extensions of the body. Linsrowinema inelist has only one pair ol larger lateral ad- cloacal papillae and does pot have the ale like extensions, Linglewihesur cinetun, which does have vhelike extensions of the body, has ventral houdy spines extending (othe level oF the mosianterior parr of latent] caudal papillae bur ZL. lesiaiieise does nol. Liwlewinema cinetam hax wi spe patty ot papillae literal ynd anterior to the ¢loaca the sans: sive bul Lo tesmiunienve tus the three hater pairs larger The guberniculan ts more U-shaped in vental view in L, trsmatiense than th, iulisior f. ernein, Pemile Le tasmuniense Haye a shorter far TISA35 (S10) than Feiner 986-1122 (1050). The besophugus is whoue To body length in ob. fasmanienye males compared with Us in Le cient andl T/ib ny de dewdist The vulva is elosey to the anterior end im be tewmanense, about OO mm compared with 1O mn int 2. taedise and 12 mm in cei, The vagina of Lo teasmaniense. VS. ts shorter thin in Lo emenon 300 and Lo tiglive 350, Chabagd et ah ()980) fignred the (ail ofa male specimen, registered as AHIC 4530, tran A efesutinys Upper Dromedary. Tasmania, There is new only ane dimuged male specimen in the bottle avaiable lor Comparison, bub such characters us van be seen, Und (he drawiigs of Chabaud ef ad (L980) (Fig. S BLE, p. 438), are consistent witht L. faymaniense, Cinstowaiemae teesmaniease appears fo have vv googruphic range whieh extends weross Tasminma tmd Kangaroo Islund. Three hosts trom Kungaron Island were infected with two species at Linstowinend. one With Lo efter and be lannnunie@nye, und Iwo wilh Le wyweringdent and LL. tasmaniense. This stiggests past links hetween Kangaroo [sland hosts. dadvlind Australian hosts and Tasmania hosts. Bivinelouy The species ts named according ty a Tabet found ant AHC 1820, "EA. cinedam tasmaniensiy’, here determined to be ZL. civetum. which had appurenity been writen by Chabuud era when preparing their paper ol 1980. Type lavatiry Kingston. Tasmania. Austria LINSTOWINEMA FROM BANDICOOTS 2) Type host lsoodon obesulus (Shaw, 1797) Site in host Small intestine Type specimens Holotype male, AHC 30320, allotype female, AHC 30321, paratypes AHC 30310 Linstowinema peramelis sp. nov. (FIGS 62-66) Synonym: Echinonema cinetum Beveridge & Walters 1991: p. 22. SENSU Spratt. Material examined T\pe material: Holotype d, AHC 30023, allotype °, AHC 30097, from Perameles bougainville. Other material: From Perameles bougainville 1, 19. 1 anterior end, | posterior end, no collection data, AHC 30055, 30054. 30053: 1d. L. 3 anterior ends, 2 posterior ends, no collection data. AHC 4522, 23 5d. 1%. captive, University Adelaide, no date, AHC 13928. Description Cephalic bulb with 3 rows of 15 (male) or 16 (female) files of hooks, 2nd row longest, 3rd row much the smallest (Fig. 62); neck with about 6 rows 63 Figs 62-66. Linyfowinema peraneliy sp. noy. 62, Anterior end (lateral yiew). 63. Body hooks (lateral view), 64, Body hooks (dorsal view). 65. Female tail (lateral view). 66. Male tail (ventral view). Scale hars = 200 jim 62: 25 pm 63, 64: 100 jim 65, 66, a2 L. RE SMALES liny spines; cuticular dilition of oesophageal resin bear LO-12 rows, 14 Gmale) or 16 (femule) Files of boely hooks, first and last rows smallest. bth-8th rows lamest, roots of books without undulating edges (Pigs 63, 64); remainder of body with a row of small spines at each curieulan annulition (auinbers of spines nol counted), over whole body of female; extending over 7) dorsal surhiee. terminating about 300 unierior to elowea ventrully. reaching level of Wost uitenor pum ool cuudal pupillac, an verral surfave OF male bady (Pig. 66), Qesophagus simple Clubeshuaped, about Yine'/o body length, lermmunating al Vth lth row of hooks (Fig. 62) Sevretory- exerelun pare. derrids und nerve cing nut seen. Males in=2), Length 9 9 mim, wilth 475, 475. Cephalic bulb 235, 325 long by 234, 267 wide, cephihic hooks st pow 123, 129 2nd vow 117 150. Yl row 73, “3 tong Oesophagis FHS. 970 long, vuticuhe dilanon beuing b)-12 rows hody looks. Spreules stimilar, eqaal. without alice, 950, 1050 long, about 4) hedy length, Gubermeolin short, simple, subetyaedtir TT pairs candal papillae: 3° pairs veniral dnd dromediately pre-. ad-aid post-cloaval respeclively. | pair lateral ad-elowcal. 4 pairs Tatenal pre cloucak 7unterior pairs all about sume sive: 3 pairs papillac, pair phasmids well pasterion lo clowes, nea) Gal Up (big. 66) Cloacal region with small riticular bosses, alt-like expansion wbsenr, Til 210. 20 long. Ponale: (v2) Length |) inne. width 520, Cephalic haoks Ist row 2600, 145, 2nd row 140. 260, 3rd row 78, 140. Oesophuyus (190, 1250) longs cuticular dilation hearing If-12 rows bouy Hooks. Tait 460 louy (Fay. GS) Vulva not seen. kees more on less sphericul, 5t-S7 hy 60-63, Remurks Alihouph ther was only aosmall done ol Specrniens tind they were al in paar condigon, significant differcuces between these specimens und Oher species Of Lrwlaveiinemed Could be foe, Althouel) elements such as the vulva cold tol be seen, (he worrns appeared fo be mature. having fertilized eyes feamero. Since & homgativite is now extingbon meintand Austratia aud Tully protected an the istinds ti Shark Bay Western Australia, it is unlikely Cal any inure speeinens will becente avtilable, Accordingly this somewhat Taeemplete Jeseiplion is presented ae the best possible of the species under (he crreumstunces, AdaMeandieuy peramelly Sp, nov, most closely resembles 1 warrington’ and the speeies thar have been (istiniished a the discussion Ob L. wanvingtend, ty Unit there are no more than [1-12 (male) and 1O-12 Houle rows af body hooks compared with 9-11 Give) and Lh) 3 Gemaled and the oesophagus ts relalrvely lone, extending (othe Tlth or 12th row nt hody hooks. The awa species van be differentited hy sive. Linswinenet wareingiont Wales, 15-20 wie, und Jemules. 32-41 pm, ate much larger worms. than L perameliy, Y and Tt im, respectively, Lin Mewinwne peruned’s has plain edges on the roots of is body hooks bue 4. werriigroné fas undulaune edges. Phe numbers anil dnranvement of papillae on the posterior end af the male are alse different, Lid- SOwinend percwielis ts the only species of Lin Mewinema wilh Tour pairs of pupillae Taleral anid anterior tothe clones ull other spectes bave three. The spivules of 2. penumeliy (950-1030, Wa body length) ane relatively longer haa those of merrier (680.1090, 1/20 boy: demethy, Ltr sme perwmelis has three pairs of papillae mean the caudal op hur Lowesriigread has five. The lai eld. peramehs. (210, 240 (male, 460 Wena, is slorter thun that of 2. anaiefigtand (330-430 gid 1OaO 1900), The exss ob L, peranelis (51-57 by 60-63) are hirger than those oF 2. wurriagiont (3-44 by 3348), Liasnavineme perumely issiinilar ted. fredise anu 4. eet. in having three pairs of caudal pupae, Iican he differentiated front these (wo specs i having (relatively lod oesophagus, terminating: sl the level of the postenor rows ol liaoke van the cuticular dilation of the pesophipgea! rewion. and net WKH toad in hiwing up te 12 rows of body looks without undulating edges compared with Lo olelian Up to be anh Le crrera, up ter TS cows of bouly hooks with undulating estes, Linstewinena peraniel’s rescanbles L. rsnmeniense Wy diiving, Four pairs of caudal papillae eur the cuudal lip and body hooks without wiadilatine cdees, Kt differs from L, resmenienve tn having the oesophians terminating at the end of the eumeuli dilation and not within it, up ta C2 tows ol buds Hooks rather than [5 and larger eggs, 51-37 hy 60-63, compared with 36-45 by 33-45_ Althotigh the third tow of cephalig hooks js relatively much smaller than the first and second hows in Lh. perameliy Vii mn other species uf Liiiowinema, the cophalic hook siaes vary wreath berween jndivicuads und se this muy web he a consistent chiaraeter, The cight B. howedinuille, dissected to prnvide / peramelis tor this study Uhree ot whieh were ifeeted), Were all rewistered in the SAMA in 1936, hut wo other collection date were given in the museum revister. The collection data for AHC 452% and 13928 are equally sparse. the information Gn the labels giving Goly the laculity as possibly Soul Austad und captive mi the Zoology Department ot ihe Liniversity of Adeluide. Examination of the AHC records jiedicates what five additional bundicoots were dissected for helnioiis, (wo ol whieh were mfected with 4. pecamelis, This suggests that the prevalence LINSTOWINEMA FROM BANDICOOTS 21 al infection in Po beongeinitle by Lo permuneliy was ubout 48%, Epyiiolawy The specilic name ts taken from the label ot ALC 4522. the material originally registered as Fohiianemie cine perameles, here determined ts bef peranielis. Type lacality Unknown, Australiu Type hese Perameles bouvarwville Quoy & Gitimard, 124 Site tH dest Stall intestine Type specuuens Holotype mule, AHC 30023. allotype fenide, ABC 40007 Linstowinema maplestonei sp.nov. (FIGS 67-78) Material examined Ipe material: Ubolotype fd. allotype 9'. fan Perameles hasta Dinger Creek. (17> 26'S, 146 OO'T), Queensland, Ty.60. ATIC 30094, 30095: Paratypes 40. 1 9. 2 anterior ends. 1) posterior end OM 1446341, ALI 19763. Oren jnatental From — Perameley — mrsite Queensbind; 2/7 5, no locality given. Jax.57, OM GL M4S7 1d. 29 S214 fragment. Fmishial (17) 42'S. 146° OLR), 15.81.59. OM GL 4as56: 20 0. 2 southern Queensland, no diate, AHC 1726, bio fyeadon meeronrus Queensland 2° 5 , Mossman, (10° 28'S, 142° 23°). 41-1958, OM GT. 4303/2. 296. 299 Innistail, (17° 42'S, 146° QUE) 22.yi,. 79 ALIC 1726: 12, Brisbane, (27° 28'S, IS} OVE) no date, AHC 1738:54d 4,79. 1 antertor end, Paddington, (27° 28'S. 153° OLB) Aug, 195s, AHC 4371 from bandicom, io collection data, 3 3, ALLO 19667, Desc ripiis Cephalie bulb with three rows of Ib (vale) (Fig, 69) or 16 (female) files of large hooks, And pow laraest 3rd row sipallest (Fig. 68): neck with 5-8 rows ol tiny spines: cuhicular dilation of oesophageal rewion bearing 11-13 rows of 14 (male por 16 female) files-of body books, firstuod best 2 rows stallest. 7th - Oth tows hurgest: roots of hooks without undulations (Pigs 72. 73), reinainder of body with a row of up to 42 unaley or SO (female) small spines at eae wnndlation (bis. 71), aver whole body of female. extending over ‘4 dorsal surlace. termioaimy about 400 anterior to cloaca On ventral surface of (ale body. Oesophagus simple. club shaped uboul iii body Jength jerminuting posterior to oesophageul cuticular dilation (Fig. 67), Nerve ring surroundiny oesophagus within cephalie bulb; secretary -excretory pore in neck, deirids conical, at level of [st row of body hooks. Male; Length 13-23 (17.5) mm. width 475-560 (550). Cephulie bulb 285-350 (345) long by 260-310 (290) wide: cephalic hooks Ist row EE7-150 (130), And row 140175 (160). 3rd row 90-130 (120) long. Oesophagus [3602210 ()804) Tong, cuticular dilation bearing 11-13 rows body hoaks. Deirids 405- S30 (APS). nerve ring 255-300 (285). sevretory- excretory pore 325-405 (360) [rom anterior end, Spicules equal. similar without ale 820-7 100 (1030) long. about Yo body length. () pairs cauclal papihwe: 3 pairs ventral and iminediately pre ude and post- cloacal respectively, | large purr literal ad-clodeal, 2 pairs lateral pre-cloucal 4 pairs papillae, pair Phasmids well posterior ty clowea, near ual lip (Figs 74, 75), Clowcal region with small cultcular bosses. alu-fike expansions absent. Gubernaculun stort, simple. sub-triangulir in ventral view, 60 (n=l) long (Fig. 76) Tail 305-440 (375) long (Fig, 7X), Feniale; Length 18-35 (28) mm, width 560-765 (070), Cephatre bulb 325-425 (362) long by 274-425 (330) wide: vepbabe looks Ist row 150-176 (1604, 2nd row 170-225 (190), 3rd row 115-145 (125) Jong, Oesophigus lolS-3740 (2272) longs cuticular dikttion bearing 12-13 rows body hooks, Deirids 405- Sas (495) nerve ring 260-390 (3350), secretory- exeretory pore (n=2) 405. 470 trom wnterior ena. Vulva nob seen. Tail 565-970 (790), Eggs not measured, Remarks fiastowinenta maplestone’ spo now. resembles 7 warrington’ in having up tof fows of body Nooks. a rekilively Jong oesophagus and five pairs of caudul papillae on the male. TW differs from 2. weeringraad i having: FE-l4 (male) and 12-13 Gemale) compured with ® 11 (mated and 11-13 Cfenmiled rows of body books, The vesophagus of Lo mneaplestanes, extends beyond the cubeulur dilution of the Gesaphageal regan Of the hody whereas in L, warriteeront it termingtles at the levelof (he 9th - Tarh rows of body hooks. The hody hooks of L. maplestoved ave withour undulating cclves and thoseal Lo werrieren are with undulating edges; Mule body spies extend along Va of the hody dorsally and well above the lateral caudal papillae ventrally on Lo viaplesraner bur on L, Wario they extendatoig */s of the boudy dorsally undalmost to the lateral caudal pupitlae ventrally. All 6 pars of pre-, post- and ad-cloacal pupillac are the LINSTOWINEMA FROM BANDICOOTS a3 same sive on Le awarriteron’ bul L, anaplestenet las larger lileral al-cloucal papillae, The spicules of 2. maplestoner (/\ hody length) are relatively: longer (hit those oF Lo werraguiné (fn body length), The arrangement Of the papillae surrounding and unleror WO the elowca of 1, mreplestered inust closely resembles that for Lo img/id ie. wilh the haberal ad- clowcal parrthe lamest and the ventral body spines net extending to the level oF the cuudal papillie, Laistoyiineiie maplestone? differs. tram L. inedlisi im haying the oesophagos extend posterior to the body. hooks without undulating edges rather Wii an besbphagis which termites within the cubieular (ikiHon ait about the level af rows 4-9 of the boily hooks will undulating edges, Male 2. fy/isy hive only three pairs Of papillae on Gre caudal tip but /.. maplesione? have four, Male Lo taplestened on uverive huve longer tats (405-440 (A7S) than 2. inelish (150-350) (205)), the body spines al L, maplestone’ extend along oF the beady dorsally compared with almost the whole body (4a) on ZL. inelent. Linvtowinena meaplestone’ can he distinguished from £. litfeas. which also hus the Gesophagus extending posterior lo the body books, body hooks wilh undulating edges and four pairs of papillic on the caudal tip. dh having t)-13 Gmaley) and 12-134 (lemale) rows of Nooks compared with 8-10 (mule) and 912 (emake) rows ef body hooks ind. datens, Male Lo mapdéestaned have body spines extending Wong “of the body dorsally and well phoave the Kuteral couch papillie ventrally compared with / lateus. which hits body spines extending alan ia ot the body dorsally und level with the inmost anternar Lateral caudal papillae Vventtally, The literal ded clodeul pair Of papillae os largest On Le mraplestene! compared Willi wil 4 pairs of lateral elovedtl papi tlie heme the same ste oo L. tates. Memute 2, aplestane. Wuve on average shorter daily 65-970 (790)) tha 2. lereiis (790. TB ES CLOT). Linsionwineni ineplesiamet Gan be drflerentucded rom Forel in having bI-ta 0 (miley, T2-13 (emailer body hooks without undibaring edges vompured With }A-L6 (malen I-TS (female) hoy hooks with unduhiti cages: a ling oesophagus depminating posterior (6 (he pows af body houks, 10 A shor GesOphiigie lernmutine at the devel of te 8 Li rows ol hooks. Liason nuples(ones tas Tour pues af caudal papillae at the tail tip. awhile / crete has Three, the pair ot Lateral audbcloacal Figs G7 7S papillae is the largest on 2. deplestonter but all six pas of cloaci) papillie qe the same sive on L cine, Linslownrenre nreplesteme? does: not have avy alitike expansion surrounding the clowea bur le Cinetim dues: the yentul body spines oof 2. cieria extend dlmost tothe level ab the inost anterior lateral pairol pro-cloucal pagatlae but nL. aaydestener they do nots the female tail of 2. wuplestene’ (3035-970) (790)0 os shorter than thar of Le einer (986-1122 (1050). The other species of Lyostewimmena, lL perunntis wd 4. davaniense. which Nave body hooks without undulating edges have three pairs OF caudal papillae on the ual tip. These can be further differentiated from Lo dplesrone’ by the number ef rows of body hooks and the relative lengths of the oesophagus, Linsiowinema purametis has 10-)2 rows of buds books with the oesophagus terminating al the level of the L2ih rows... feayvmenmienve has (3-15 rows of body hooks with the oesophagus terminating at the level of the &th = (Oth row anid £. meplesroned his [l13 tows of body hooks with the pesophasus terminating: posteriorly to the (3th row. Lantenvinema peranelis hus four pairs OF kvteralad- and pre- cloacal papillae, compared with three pairs on L. aaplestoned and relatively shorter spicules ('/o of body lengthy compared with (jn) Linwtawinemer teasnaniense (ibs three large pairs of lateral cloaeal papillae compared with the one lange ad-cloueal pair of L. mreplestarrer. Beeuuse af the small number of female specimens wiluble, none was dissected, so allliourh enes Were seem, iL was diffieull to delermine which were mature and therefore suitable for measuring. As a result. no INGUSUreMenks were made. Linsnuinenid meiplestonel oeeurs in Poet and J mecroures (ron) Queenshuid and New South Wiles alhauvh (he examinition of mere hosts is required before the full extent of the geographic range wl (his Species cun be deterinined. In twa 1 retinitis. b. maplestaner occured ait iiwed mmfechons with J. warrt elon. Etyinndogy The species is nunmed ufler RAL Maplestone, whe together with W. Yorke. cured uut pioneering work on othe nematodes of Australian marsupials. Eelinenenne eplestone? Was Used on in tindited museum libel by Chibaud ef af (P98O) ton Liayioawineme specimens (rom Zo macrourus that were subsequently determined tobe Lo werringrean Lawton jee maplestened sp. nov. O72 Adnenor cid, optical section (litera view) 68 Cephalic bull Hater view) OY Cephitic end male. optical sectional tevel OF rst naw at hanks be ice SOW L720, Fenaite tal lateral view 1 1 Male, posterior houy spines (ventral View). 72. Body hook (later Wiew). 73. Bady haok (darsal yew). 7 Male gal Lip (ventral views 79. Mile dail (ventral view). 74. Gubermiculini (vented) view) 77. Flonule if ep cheteeal view TR. Mule till Hatera) view, Suale burs =S00 Li O72 200 pans. TOL S50 pin 69. 7577. 25 pm FD TITY Ta, Tee HOO i 7 ab LR SMALES Type liast Peranteles Waste Type locality Dinner Creek (17° 26'S. 146° OO). Queensland. Australia Site it feast Small intestine ‘Type speemnens Holotype tale, OMA0N95 QOM30094, (lotype female Discussiun Linshovinenae warrington’ appears ta be the domimint species of Livacewimenn, in eastern Austrian bindicoots. being found) from oorthern Queensland throweh to South Australia, including Kawigaroo tshiid, He ocedrs ib all extiit species: of Avoedon as well us one individual of Po acyl, indicating a low prevalence ii (his latter host species, Althourh ©. faery wes found) in northern Queensland populations oF the wurthemn brown bandicool, together with L. warringteni, it was the only species oecurrimg in northern brown bandicools fromthe Noriher Territory and (he north of Western Austialia, while L. fragilis? wis the only species ovcurtiiyg in southert, browa bandicouts m rhe south of Western Australia. Only one species, L, peramielis oecurtue a Ave or 13. 7. horganvilte examined, has been found exclusively in Perales spp. and three species 1 nasmanenye, L. larens and &. inwlist exclusively in hsondon spp. £. dates Oecurs in fe macrourus, L jasmiemicnse aid L, iedit oceur itl obesutis, OF the other three species. 2. dapleatoner has a north easter distribution occurring in 7 yee rouris and P. nosuid. Lo Cincvieea more south-custern distribution pecurring nL. ehesdlus. Po nesete and Boa, Further collections of material from south-custerti Austidia are needed before any typothesis on the distimbulion of species of Liasrowienur cun be Weveloped. tt does appear thar bandicoots (/ hess) Ot Kangaroo Iskimd, bur ot Tasmania, miy have been derived from stock in which all three species (2. weiinyrond. Lo einetiin, and b lusmentemse) Were prevent The continuing detrimental elects of Ruropean serlement Taye resulted imu putehy distribution off obesitas over a reduced yunge (Braithwaite 1995) This decline inthe host population may have affected the distribution and prevalence Of species ol Linstearinema on the iidinkind. The current prevalence of the three species of Linstawwiena in banweoots on Kangaroo Ishind may reflect past prevalences of these species i bandigoots on the mainland. The whility to trap bandicoots yaries with species their age. species and Ipeulity (Gordon & Hither 1989). Peranedes nasune is apparently more difficult lo trap tha species of /yoadun (Menkhorst & Seebeck 1995). This nay be the reason for the small number of Peraeles compated with Ssecdon collected in this study (see Table |) and in the amolntof material deposited in musciim collections froin each host genus. A farther coniplicuting lactor is the possible difference in prevalence of infection with Linsiowinena berveen the two, The records ol the SAMA and QM indicate that 90 2. nase have been examined for belnmth parasites and. of these. only eight were infected with LL. taaplestanel, six with Lo elven and one with Lo weannietens. Similarly, of SEA gem examined, only tour were infected with Lo cinenue and one with Ff werrtiwton’. dos unlikely chat Linvtowinena could have been Overlooked during dissection as (he worn are Taree nd cine be readily detected in the simill intestine. These low prevatences of infection contrast with Une prevalences found for species ol deeaden disseeted in this scudy. OF 72 hancicoots examines, 53 were infected with Linstowimenk spp, A working hypothesis would be that species af Lintomineme are dominant in the helminth communilies OF fseovdow but net in those ol Perameles. Wi some areas. species of bandicoot haye overlapping geographic ranves, do acronvuy and PB Hasite in the north-east, (oohbesi/is and P pasiited in (he soulheeast and 7 obesity atid Po gee ‘Tasmania, “Their habiti! preferences within each geographic region are different, and although they Inay nol be ii striek symypalry. opportunities foe Incidental infeedion and bost switching would Gx ise Observations trom this stidy suggest hit species at Linstowinenae nay switch trom 7. wheal be P gunn and PO eesues Veron Eo abesidige loa P honviwitte ond perhaps fron / onusutia ta | mrechnens, Cho mht aecaunit for rhe Geeurenee af Lingiwvinemea i Perameles. Additional collections of material fon Percmieles spp. across Australi and especially 7 ebesedtay trom south eastern Australia are needed to lest thes hypothesis, Acknowledaments Thanks are due to the lollowing collectors whe provided miatterial for (his study; [, Beyerndec, R. Beavis, T. Th Cribb, T. Dennis, M, Driessan, T. Foley. To Priend. Re Purer, Bo Gosslink P. Hayeoek. Po Hayward. O. Hiekiman, Bo Mundsy. R. Noking R, Norman, D. Obendorf, b. Owes D_ Spratt, the late M_ J. Mackerras, P. Presidente and LINSTOWINEMA FROM BANDICOOTS 27 the late A. J. Bearup. Thanks are due to-C. Kemper of the South Australian Museum, J. M. Dixon of the National Museum of Victona and T. Friend of the Department of Conservation and Land Management. Western Australia for allowing me to dissect bandicoots held in the mammal collections and EL. Smyth for the drawings, This work was supported by a grant from the Australian Biological Resources Study. References Brarhwarre, R, W. (1995) Southern Brown Bundivoot pp, W7O077 fa Strahan, Re (Rd) “The Miarnmals of Austrailia” (Reed Books, Chatswood). Chiabauo. A G,, SbukiaL, C.. Beyeripct, 1. BAIN, OL & Diunerve-Drsser M.-C, (1980) Sur les Nematodes Eehinonemiatinae. Amy. Parasol, lum, comp. 55, 427- da, Carink, OJ. CES81) Supplementary report on speeimens dredeed up from the Gull of Manaar together with others trom the seu in the vicinity of the Basse Rocks und from Buss's Strails respectrvely, presented to the Liverpool Free Museum by Capt, Th Cawne Warren. Aan, Afar, Nat. [livt. 7, 3601-385. Prinp, J, AL & Burpipdh, ASA, (1995) Western Barred Bandicoot pp. T7180) fe Striban, Re kaha Phe Miainimiils of Australia” (Reed Books, Chatswood). Gorpon. G, (1995) Northern Brown Bandicoot pp, | 74- 175 shied, & Unninien. A. TE (1989) Perumelidae pp. 603-624 fn Walton, D2 Wo & Richardson, B. J. (Rds) “Fauna of Austealia’ Vol 1B Maminlia (Australian Government Publishing Serviee, Canberra). Incuis, WG, (M967) ‘The relationships of the nematode superfamily Seuratoidea, J, Mefminthel, 41, 115-136, JouNstos, TFL & Mawsos, PM. (1939) Sundry nemiitodes From eastern Austealisin Marsupials. Trevis R. Sac §. Anst, 63, 204-209, R940) New and koown nematodes from Australian marsupials, Prac, Linn, See. NSW 65, 468- 476, & (1952) Some nematodes trom Australtae birds and mamoials. Tren Ro Sue, 8. Aust. 78, 30-37, Linstow, O. Vor (1898a) Nemathelminthen von -lerrn Richard Semon in Austrilien sesimmell, Denkschr vid: neathw Gesellseh. dene 8, 467-472. ()989h) Nemathelminthen. you Herm R. Semon in Australien gesummelt, 4eelogisehes Central Blan Letpziv 5, 672 MeKenvih. No1 Morris, KR. D.& Dickman, CR, 11995) Colden Bandicoot pp, 172-173 dn Strahan, R. (Rd.) “The Manimals of Australia” (Reed Rooks. Chatswood). MackErRAS, L Mo & Mackrreas, M. J (1960) Tuxonomy of the vommon short-nosed marsupial bundicoot of eastern Queensland. last do Ser, 23, 51-53. _ & Saspars, D. F. (1954) Punasies of the bandicoat, Proc R. Sec, Qld 63, 61-63 Mantowgy. J, A. & Ribk. W. DL, (1988) Peramelidie pp. 36-42 Jn Walton, D, W. (Lid.) “Zoological Catalogue of Australia” 5S Mammalia (Australian Government Publishing Service. Canberra), Menwnnorsr, PW. & Srapeen. J. A, (1995) Long-nosed Bandicoot pp. 77-78 fa Seebeck, J. bE (Ld.) “Mammals of Victorit Distribution. ecology aud conservation™ (Oxford University Press. Melbourne), Munday, Bo LL. & Grken, R. G. (1972) Patisites, of Tasmanian native aed feral fun Pert Uh blelminthes. Rec, Queen Vie, Muy. 44, 1-15, Qurntin, J.C. (1970) Sur le cycle evolinil de Searanin cvadarachense Desportes. 1947 et ses uffinites avee ecus des Nématodes Subulures (Ascuridia) ef Rictulires (Spirurikiy). Ann. Parasitel, non. conip, 45, 605-628, Romer, & (1901) Monotremuta und Marsupialta. Derdsclie med-naitew Gesellyeh, dena, 8. 153-160, Srepecn, J. G. (1905) Eustern Barred Bandicoot pp, bs82- 183 /n Strahan, Re (Rd.) “The Mammals of Australia” (Reed Books, Chatswood), Sprarh DL MM. Beveruogi. 1 & WaAuTiR. Eb, (hoor) A cutulogue of Australasian monotremes and oursupials and their recorded helminth parasites. Ree. 8, Aust. Muy, Manor ser t, 1-105. STODDART. B.( 1995) Long-nosed bandicaot pp. Us4-1K5 fv Struhan, Ro (id) “The Mammals of Australie” (Reed Books. Chatswood), Yorke, W. & Marcestonk. PA, (1926) "The nematode parasites Of vertebrates’ (Churchill, London), A NEW SPECIES OF FRESHWATER AMPHIPOD, AUSTROCHILTONIA DALHOUSIENSIS SP. NOV., (CRUSTACEA: AMPHIPODA: HYALELLIDAE) FROM DALHOUSIE SPRINGS, SOUTH AUSTRALIA By W. ZEIDLER* Summary Zeidler, W. (1997) A new species of freshwater amphipod Austrochiltonia dalhousiensis sp. nov. (Crustacea: Amphipoda: Hyalellidae) from Dalhousie Springs, South Australia. Trans. R. Soc. S. Aust. 121(1), 29-42, 30 May, 1997. A freshwater amphipod Austrochiltonia dalhousiensis sp. nov. is described and illustrated. It is endemic to a few artesian springs amongst the Dalhousie Springs complex in the north of South Australia. Morphologically it is very similar to other species of Austrochiltonia found in mound springs near Lake Eyre South but preliminary electrophoretic analysis of allozymes supports the recognition of a distinct species. It most closely resembles A. australis (Sayce, 1901) in that uropod 3 is two-articulate, but differs in a number of minor features, which collectively distinguish it from its congeners. Key Words: Austrochiltonia, dalhousiensis sp. nov., new species, amphipod, artesian springs, Australia, taxonomy. Tenscetions of the Rovet Suciely uf S Aust (1997), 12161), 29-42 A NEW SPECIES OF FRESHWATER AMPHIPOD, AUSTROCHILTONIA DALHOUSIENSIS SP. NOV., (CRUSTACEA: AMPHIPODA: HYALELLIDAE) FROM DALHOUSIE SPRINGS, SOUTH AUSTRALIA by W. ZEIDLER* Summary Zio, W. (L997) A new species of freshwiter amphipod Auvrechiltonna dalhousrensiy sp, ney. (Crustiven: Amphipoda: Hyaleilidaey fon Dalhousie Springs, South Australia. Frans. A. Soc So Aust T2101), 29-42. 30 May, 1997, A treshwiter amphipod Ansochienia duliousienas sp, noy, is deseribed and Ulusteated, Iis-endemie tua lew urfesian springs amongst he Dalhousie Sprigs comuples In the north of South Austealia. Morphologically itis very Similar to other species of Ansireeitonia found in mound springs near Lake Eyre South but preliminary cleetrophoretic analysis of allozymes suipports the revagnition of a distinet species. IL imost closely texernables \ aresdraliy (Sayce. 1901) 1h thal uropod 3 js lwo-urlieutale, bul differs in woumber of minor features, Which collectively distinguish 1 fromm ils congeners, Key Worps: dasieeditiena dallomyiensis sp. noy.. (Uxonomy. Introduction Aniphipod species of the genus Austrchitontesure among (he most common crustaceans Found in (he permanent freshwaters of soulhern Australia ranging from New South Wales to Western Australia and including ‘Tusmania. More recently Austrochilyn le Nasaudso been found in the inlaid water's of artesian Springs ii South Australia (Zeidler 1989) and at “Edgbaston” north-east of Aramac, Queenslond (personal colleehion. May 1988), When | re-estublished the genus Ausrochiltonia (Zeidler L988) it was my mlention to proceed with an Australian revision of the genus beginning with the description of species found in the mound sprigs Wear Luke Eyre South and at Dalhousie Springs im Northern South Australia, Since then | have exainmed a large number ol specimens from wide-ranging habitats in Southern Australia and have found them all tobe very similiar morphologically ond difficult to distinguish from the only previatisly-described species, A. anseralin (Sayee. LOO1) and Al subtests (Sayee, 1902). Williams (1962) revised the s¥slematics Of these bWwo species based om type inalerial and a range of specimens from New South Wales. Victoria, “Tasmania and Rotimest Istand. Western Australian and likewise found that, morphologically, specihe differences are minimal However, a preliminary andlysis af allozymes of specimens from the South Adstrahan mond springs usin electrophoresis, idiodtes that Austrochilionia in inost likely a yery speciose genus. Given its Soul Atiatailion Maseurie Sarthe Terie Aelia 8. Aust SUM) new species, Amphipod. artesian sponys, Australia, potentiul enormity, the project was abandoned duc to lack of resources. The species found at Dalhousie Springs is most similario A, ausirafiy Sayce. 1901 in that uropod 3 is two-arliculate. Tt has a very restricted distribution. oecurring at only three of about &0 active springs ti the region (Zeidler 1989). “Two of these springs are quite durge, with large outflows of warin water (P40°C) bul Auvtreciitenia is found only in the distant overflow where the waler is colder and close to ambient temperature. However. one isolated specimen Was colleeted from the edge of the pool of the main spring, which has w water temperature ot about 35°C. The other spring is a small relatively vold spring on the southern edge of the spring complex. In each case the animals were only found in the shallow edges of swamps or channels amongst the base of the sedge Cyperus laevivars 1. 1771 and sometimes alse the reed Phragmites australis (Cuv., 1AebL). The restricted and isolated distribution of this species of Austrachilionia makes i) vulnerable to habitat distarbance even (hough Dalhonsie Springs is within Wihira National Park, The purpose ob this paper is to establish the taxon so that park managers und visitors can appreeite tS significance and potential vulnerability, Materials ond Methods The Dulhousie Springs coniples (Pry. 1) consists oF about SOQ uclive springs all of which were sampled in 1985 (Zeidler & Ponder 1989) hut Ausrochilnania was found in only three springs (Fig. 2). The springs we coded following Zeidler & Ponder (1989, Fig. 2). a0 W. ZEIDLER Animals were collected from amongst sedges and reeds with a small hand sieve or picked off plant debris with forceps. A total of 424 specimens (230 90.174 oo, 20 juveniles) was collected and exumined. Physicochemical data for the sites sampled are limited but some measurements were made near the main source of the spring. These data are given in : . 1 LE SONS H : \ | | / { Northern Territory | Queensland ' \ j i} \ \ h \ | j kL = - -—=- Se — ——yPrdelle \ Dalhousie Springs* 4 | F i | ; 1 py Lediardatia if ! i \ ° | | oO i ) v \ iD RYH | Leaver Helps AW \ Matinee rv | South Australia ; | \ at) j ! j | / | i \ \ a) | 1 \ I ae j rig \ é \ gies nugeane | i) i : | i | . i i Borcelside | i ' WoT | ~ \ \ | 1 \ I 1 | ! | Saule \ a Fig. |. Lovation of Dalhousie Springs. South Australia, From Zeidler }O91, Table | and data on nearby springs are also available (Smith 1989). The new species was compared with the descriptions of Ausirochiltania given by Wilhams (1962) and with specimens of A. australis from Dandenong Creek. Victoria (SAMA (C3872) identified by Williams and used in the study by Smith & Williams (1983), Material reported here is deposited in the South Australian. Museum, Adelaide (SAMA) and. the Ausiralian Museum, Sydney (AM), All specimens are preserved in 75% ethanol or 2% formaldehyde/propylene-glycol solution. Of the types, only the holotype and allotype have been dissected (partially), with appendages removed from the left hand side of the animal unless otherwise indicated. Dissected appendages are preserved with the carcass or, in the case of the holotype, the mouthparts, uropods and telson are mounted in poly- vinyl lactophenol on a microscope slide, Specimen length is measured along a. lateral parabolic line drawn from the anterior extremity of the head through the mid-line of the body to the posterior limit of the telson using a pair of dividers and seule, The thoracic limbs are referred to as guathopod | and 2 followed by pereopods 3-7. Size comparisons of gnathopods exclude the coxa and dactylus, and of the pereopods, the coxa, with articles being measured along the mid-line, The following abbreviations are used in the text and figures. Al. A2 = first & second antenna, G1, G2 = first & second gnathopod: LL = lower lip: Md = mandible: Mxt, Mx2 = first. & second maailla: Mxp = maxilliped; O2-5 = oostegites fram pereopods 2-5; P3-7 = pereopods 3-7: PI = first pleopoda: T = telson; UL-3 = uropods 1-3: UL = upper lip; r= used as sufhx to indicate that appendage was taken from right hand side of the animal. TALE b. Temperate measurements and phystoachemical data (from Smith 1989) far springs fram which aniphipads were vallected at time of collection (except far Cal - data from [983 expedition) Field Chemistry Spring Temp. Temp. Aw Wiiter Temp. Cond. TDS pH bo Cc *C “C ASHE mgt! ppm siemens Cal - channel to main pool - - 43 1490 865 73 3.8 Cul - main pool x) 37 34 1780 1o00 7.9 67 Cal- main discharge channel = 25 36 33.5 2050 1150 7.7 47 Cd? - SW edge of pool 15 yu 32 1550 850 7.9 11.3 Cd2 - at or near swamp 15 11 ts 1650 - TAS 7.6 Gb 13 16 20 THO 4850 7d 45 A NEW SPECIES OF AUSTROCHILTONIA 4 st ANS | Ls .0 =I e | = St ‘ 5 ’ . 3 afl y , Hf ‘Dalhousie (ruins)} wo « Fiz. 2 Dalhousie Springs complex showing springs (coded) from which Austrochiliania dalhousiensis sp, nov. was col- lected. Collection sites for Cal and Cd2 are arrowed. Other major springs are shown as dots. Swamps [rom springs and ebeck heds are stippled (light stippling indicates ephemeral stream beds and heavier suppling areas oF permanent” walter). 32 W. 7EIDLER Systematics Austrochiltonia dalkousiensis sp. nov. (PIGS 3-9) Austoclifimoa sp. Zeidler 1989 83-K4, fig, 12.) B. 1YOLS 185 Holton &, Dulhousie Springs SA, from amongst reeds and sedees along eastern edge of swamp crealecl by outflow from main spring (Cal), 3.3 kin doth of edge of old airstrip. 26°23'07" S 135°30'26" hi. b2vi1985, W, Zeidler & K.L. Gowlett-Rolmes. SAMA C565]. Alfplypes Oviverous & SAMA C5652, Collected with holotype: Paratypes: AM P48840, 10 2-9 100" oT sume data as holotype. SAMA C5653, 24 29 (one oviserous), 17 cf sane data as holotype. SAMA C5054, 37 GY (three ovigerous), 11 oo, same data as holotype except 14-vi. 985. SAMA C5655, 24.9 DV OO Lsame data us holotype but ).) kim further north, 26°22°26" S L35°30°2A" EL Giher material examined: Al From Dalhousie Springs urea (Vig. 2). AM P4a841, 1 2 (damaged), sprit Cul, 26°25700" 8 135°29'53" BE. trom edge ol main pool, WF, Pander & BD. Winn, 3.vi.1985, SAMA ©5656, 2) GY (three ovigerous). 31 a, spring Cd2, fron -edges of swamp formed by outflow, approximately 0.9 km NW ob souree. 262433" S 13597845" E,W. Zeidler & K, lL. Gowlett-Holmies. 6,11 1985, SAMA C5657. 14 PO, 2 Aa, same data as previous lol except ori (O85. AM P48847, 10 2 2 (one ovigerous). 6 Oo" spring Cb1, from edges of swamp, 2604112" S 135°29726" EWE Ponder & D. Winn, 5, vi, L985. SAMA C5658, 9) 2 & (three Oviverous!, 61 O'S, 20 juveniles, Sume cla as previous lol except colleeted W. Zeidler & K.L Gowleti-Holmes. Descriprion of holotype male (Figs 3-6) Length 3,8 nin, Head wbout as Jong as deep. length Ulitoast equivalent! lo first two pereoniles, Antenna | about 3x length of bead; pedunculur article | length 15x width, uricles 2 and 3 subequal in length about 0.75% length of article 1: flagellum slightly longer than Lax peduncle, of ning articles with one ventral aesthetise dt base of each of last four articles. Antena 2 about O7x Jeapth of Al wath characterise ghind cone ab base; peduncular article L slightly wider thin long, article 2 wilth about 0,7 lenyih. 2 as long as article | and 0.78 length of ardele 4: Flagellum slightly longer than pedunele, of vight articles, Upper lip shahtly wider than Jonp. upieally rounded, bearing numerous short sete apieally. Lower lip With vestigial inner lobes; outer lobes sudowale Wil sctose distal abd (aver miuruias, Mancdibles without paps lel with inepsor ol sis teeth, acina mobilié of five teeth, spine row of three feathered spines und trituralive molar; rh with incisor of five teeth, lacina mobilis: of three teeth, spine row of two feathered spines and laiturauve molar with one long feathered seta. Maxilla | without palp, notched at palp's qorial posiion: outer plite with mnpe comb-hke spraes upically: inner plate very narrow with two feathered spines apicily, Maxilla 2; outer plate about !5y length of inner plate. setal row pesticted To apedt inner plate with one laree seta medially about 1.4 from apex. setul row apically and medially. almost to large seta. Maxilliped; inner plate Jarge, sub-rectanpulir, reaching end of merus, maximum width about 3« length of outer margin, wilh three apigal spine teeth, ihe inner one smaller. four plumose selae on inner marin and several apically: outer plate ovate, reaching midway alone inner margin of carpis, about. as wide dis inner plate. apical margin with three selue. Inher margin with several setae for distal hall, palp large, 4-articulale; merus proximally nurrovy, sub-triangular, outer margin about 2x length of inner margin with two selec on inner distal angle; curpais slightly broader than long. slightly expanded distally, distal Iwo-thirds of inner margin with row of setae, hwo sefae on outer distal angle and also pear inner distal angle: propodus slightly narrower and shorter thin carpus, distal margin with several strong series curved daetylus with strony ungurs. Coxul gills sausage shaped. present from G2 to Pb Gnathopod |; coxa shghdy longer than inaximun width, proximal width about (7x distal width, uNnicrion Margin concave, postenor Margin straight, distal murgin eyenly rounded with several evenly spaced sete: canes lridngular with large posterodistal tobe. with anterior nargin almost 2. length of postemor marine maximum width about 1.5% that of anterior twreit, posterior iigia with ¢lose-sel row of pine stout. pectinale spines: propodus subsrectungulap about 14a length ot curpus, slightly wider distally, widih O,6x lengit, posterodistal corner with two stout spines on elther side of ductylus. cluster of Jong setae on anteradistal corner, row of seven long sete medially. mixture Of tong and short setac near distal margins dactylus Shehtly shorter than width af propodus Htlnie neally wwarost palm. Gnathopod 2 length 1.6% Unit of Gi]. cosul gill length 2s wath, like shorler than cosa: coxa rectangular slightly longer than wide, abow O.88 length of basis. disud margin evenly rounded with several evealy-spaced Setic. merus with right angled bend: eurpos similar to Gl but withuut pecunate Apines; propodus slightly shorter than basis, length anterior margin LOX saaximum width, posteroproxrmad corner forming rounded lobe, palin oblique wilh numerous spines of varying lengihs on A NEW SPECIES OF AUSTROCHILTONIA 33 Fig. 3, Austrochiltonia dalhousiensis sp. noy., holotype Go. Scale bars = 1,0mim (whole animal), 0.2 mm (A,U.T). a4 W. ZEIDLER Wz Wey a SS Fiz. 4. Austrochiltonia dalhousiensis sp. nov., holotype co. Scale bar = 0.1 mm. A NEW SPECIES OF AUSTROCHILTONIA 35 Fig. 5. Austrochiltonia dalhousiensis sp. noy., holotype o. Scale bar = 0.2 mm. 36 W, ZEIDLER Fig. 6. Austrochiltonia dalhousiensis sp. nov., holotype 6". Scale bar = 0.2 mm. A NEW SPECIES OF AUSTROCTULTONIA either side of cutting edge followed by shallow wroove lor tip ol dactylus: dactylas claw-ltke, as long as anlerior margin of propodus. Pereopod 3 with part of propadus and dactylus inessing On right: coxal gill length almost 2x wrath, about 0.75x length of coxa: coxa like that of G2 but slightly larwer. shehtly shorter than basis; merus 5x uy long as basis, anterodistal comer produced, carpuy 78x length of merus. Pereopod 4 similar to P34: slightly longer than G2: cox with distinc posteroproximal excavation, maximuny width slightly more than length, shehuly longer thai bists, propodus slightly longer than merus; dactylus stout. length slightly less than 0.5x of propodus. Pereopod S slightly Jonger thu P4, coxal gill length ubout 2x witb, shghtly longer than basis; coxa width about 5x that of busts, anterior lobe slighty more than 05x length of basis, posterior lobe uboul 0.8% lengur of basis: basis slightly longer than wide with typical expanded posterior margin and posterodisial lobe veaching to about midway of ischjum: merus with posterodistal comer produced. length about (78 (hal of basis; carpus slightly shorter than mierus: propouus length bax that of carpus, ductylus stout. 0,5, length of propodus, Pereopod 6 length | 3x that of PS: vost! gill length about 2x width. about (75x length of basis; coxa almost as wide as basis, anterior lobe (3% Jength of basis. posterior lobe (5% length Of basiss renmining articles bike those of PS bat basis with siedight posteroproximal shoulder and carpus atihthy longer thay mers. Pereopod 7 longest, slightly exceeding PO, like PO but coxa semi-cinvalar ane lachkumge coxal gull widui ty length. about Od leneth oF basis: posterodistal lobe of basis more capanded reaching lo about midway of mesus. Pleopuds all onmoditied (vot as in Clete. Urepod [ whout |.5x length of (2; peduncle wiih spine on inner and outer distal comer, three large and one smaller spine On dorsal onter manga, one small spe On Inger margin: outer mumus SlighUy shorter than inner, length 0.7% that of peduncle, with twe yocutan and three terminal spimes: omer ranms with two sivalland three larver terminal spines and three medially. Ureped 2: peduncle with spine on inner andl ooter distal corner ind additional one on dorsal marpin; toner ramus |x length of outer ramus.and [3x that of peduncle, live liege spines clustered terminally and three spaced evenly medially: inner ramus wiih 20 spines of varying sizes gradually closer together lowiuds Up. Uropud 4 (vo-urienlate. imanially niece than O.Sk length ob rebyen: ranits 0.58 length of peduncle with three long sefae and one short setu revminally. Felson votire, subrectanpulin, shyltly wider than Jon. distal imurgin slighdly coneave with two snail Sete ut each vorner ~ a) Deseription of allotwpe female (Figs 7-9) Length 3.8 mm. ovigerous with 23 eggs in brood: pouch, same as male except for the fullawing, Antenna 2, Nagellum of seven articles. Gnathopod |; coxu relatively narrower and longer than for mule, width distally 0.8.x length: posterion margin of carpus with vlose-set row of LO peetinate spines: propodus relatively oarrawer than for male, slightly longer than carpus, Gnathopod 2 length 1.2% that of Gl: coxal onl eelatively smadler, less than 0.5x leneth of cox voxa with posterior margin produced to pout miccdilly, ratxamium width 0.5% length, as long as basis: remaining articles like those of Gt only relatively more slender. Pereopod 4 length about b.3% that ol G2; coxa similar in shape lu that of. G2, Pereopod 4 slightly shorter than P3; coxa without distinct proximal excavation, almost as ware as Jong. Pereopad 5 only marginally longer than P4; coxa width about 1.7% that of busts. merus, cirpus and propodus relatively shorter than for mite, Pereopod 6: basis with posterior miuruin rounded proximally: merus, carpus und propodus relatively shoner than for male, Pereopod 7 slightly shonet than Po: basis relatively narrower, ind merus, carpus an! propodiis suecessively slightly shorter than for male, Oostegites on Coxe 5, all with curled margins and numerous small hooks, together lorming ueht marsipium. First heartshaped, length 1.6. marion width. about O.7x lent of C2) second wapezoid, length almosr 0.5 that of P3, maximum width almost O.5x length; think oval-shaped of similar size to second: fourth sub-reeiangular with oblique distal margin. length anteriorly almost (55 (hat of PS. maximum width almost equal to length ol posteriur murern, Uropad | length 1.64 tht of U2. pedimele with five large and one stall Spine on Guler margin, miner margin With tWwer small spines proximally in addition {o lurge spine on distal corner; outer ramus as long as inner, length O.8 that of peduncle. with two lage and two smiller spines terminally and (wo medially; inner runius with three lirge dnd Iwo smaller spines terminally aed two medially. Uropod 2 peduncle with two large spines on oliter murgin, onter ramts slightly shorter than mner length tax that ol peduncle, one large and two smaller spines renuinally. three large spines medially; timer conus with wo terminal spines. cluster of fear near tip and another two tedially Telson with group of three small setae at wach corer, Evils token from the type locality in reeogmition of the restocted distribution of the species. 3K W. ZEIDLER Fig. 7. Austrochiltonia dalhousiensis sp. noy., allotype Q. Scale bars = 0.2 mm. SOLS Ws ut me-peee | \W A NEW SPECIES OF AUSTROCHILTONIA 39 Fig, 8. Austrochiltonia dalhousiensis sp, aoy., allotype 2. Oostegites on P3-5 not illustrated. Scale bar = 0.2 mm. 40 W. ZEIDLER Fig. 9. Austrochiltonia dalhousiensis sp. noy. Oostegites from allotype 2. G1-P4 from paratype @ , 4.8 mm, from SAMA C5653. Scale bar = 0.2 mm. A NEW SPECIES OF AUSTROCHILIONIA +| Variation Apart from minor variations due to size. paratype and other material @xaimined is very Similar ro either the holotype or allotype, The maximum recorded size of males is 5.2 mm and that of females 6.5 min bul Host specimens eXxamiied are around 4-0 Min long. Minor differences between specimens generally were noted us follows. The number of flagellar articles of Al varies fromm eight fo (en wath one small specimen having seven: A2 has from six to nine Flugellar articles but most specimens have only seven or eight. The number of gesthetses on AL ts remarkubly conmstunt will) only some larger specimens having an extra one Clive), ‘The nuanber of peetinate sete on the carpus OF GL varies from seven tonne Hiimales and ciaht to ten in females (similurly lor G2 of teniales). In the allarype the coxuc of G2- P+ ure of unusual shape, dilfering fro fyles and qoreovieerous females (Pig. 9) in thal the poster margin is produced ty a point medially und Pa is without a proximal excavation. In the hololwpe the basis of Po bus a relatively straight posteroproximal shoulder bur ii newly all other specimens examined the posterior margio is evenly rounded, Pereopod 7 ts usually longer than P6 but in the allotype i is slightly shorter, probably because of the relatively shorter propodus Which js normally longer (han the carprs The spinavion of 1 & 2 varies slightly with larger specimens Tavine one or two ext spies ob the peduncle and rami. Uropod 3 is usually two articulite and only one Specimen (a female trom 05654) had US with one article and then only on the jivht-hand side, Oostegites of females vary considerably in size but are expanded, as illustrated forthe allotype, in ovigerous specimens. The possibility (hat speciation may have occurred! between springs withoul any obvious morphulowical changes was considered ahd specimens for allozyme electrophoretic unalysix were collected fron all three loculives. A preliminary anilysis of this matertul using methods outlined hy Richardson ef ef. (1986) indivated fixed venelic differences of 10% or Tess (for Tt foci). thus supporting the morphological evidence of one species with hte yanation. Giyen these oresully, i more dehulled analysis was corstdered) (unecessary. Diseussiont The new species desenibed here closely resembles A, austratiy to What U3 is dwoeurticulite. However, a cuiiber of minor features collectively reudily distinzursh i from this species audats only other convenen AL syblenis, The nin distilpiishing features ure as follow. Remmles reach a larger sie (han les and the species is wenctally vot as tare as A. oiuntadiy (ales up te LO. gain, females up to 8,3 mm) or A. subiennis (ides up to 1,0 min, lemales up to.6.4 mm). Antenna | hax Jewer vesthetuses (5-7 in A, aitstralis). Both antennae have fewer flagellar articles (AT up to 17. A2 up te LL tn AL eistralis). The coxae of A. defhousiensiy sp. nov. are relatively wider and the excavanion on coxa 4 is nol us decp as in A, atstealis. or AQ subtennis. In both A ainvtrtlis and A. subteques, the lareral maven of the excavation of coxa 4 js at oghtangles to the posterior margin whereas in A. daliousiensis the aagte of the coxal excavation is much greater than 90°, In oviperous females of A. dalhousivrnss the coxac of G2-P4 have the posterior margin produced to a point medially und coxa 4 is without a chuwueterisue exeuvarion There are fewer peelinute spines on the exepus of Gt (iiales) and G! & 2 Vemales) than in AL australis (usually >>l0). For Gl (males) and G) & 2 (females) the carpus ts. Slightly shorter than the propodus whereas the reverse ys. truc for A, duspilis Other miso: differences between the new and other species no doubt exit but were not evident an the present study, Austrochilionia dealionstatsix is aly similar to Phrearschilroata unuplihalma Aeidler, V9, a phreatie species whieh also has a limited distriburion at Dajhousie Springs (Zeer L991). cypeciatly in that ovigerous femules of A. deliuisiensiy have coxa 4 without an exeavation, a feature chardeterishe of 2 qnophthalaa, Given the isolated habit of Dalhousie Springs, one would suspeet thut these two species would have Common aneesiy., Howeve), since electrophoretic analysis hay shown thal they diflerat about 80% of fhe 2} loct examined, (his does nol uppeur tu be the case, The closest relatives geoeraphically, apart fron? dmophititima, ace species ol Agatrchiltenia Fouad 10 the moand Springs newr Lake Eyre Sooth. Although A. dalhousiensis is morphologieally very similar to these apecies. eleetrophoretic analysis has shown thut it differs from thent at 73-80% of the 2b low exuynnmed. Clearly a more detailed morphological and genetic study of the genus is requited to defermine relationships. A single, damyzed female of Ay cdelhoninivnyts wis Jon in the pool of spring Cat (AM Pass4id hts record apmay be due to contaminated collecting equipment as the Watley Ceniperature at that locality: ts 37°C and freshwater amphipals preter cooler waters (Barnard & Barnard l9RA) Th therelore seeiis unlikely (hat A. de/Amsiensiy Occurs matupally por tn pool of Cal but is possible occurrence al this logulity warrants further ivvestigalivn. The factors determining the distrthatwou ol this species ure unknown, Us restricted distbution at Dalhousic Sprmgs ts puzelmy us riany appaccntly suitable habitats eaist im which this species was tet ound, Although restacted in its aistrbutran, the species is pchilively abundintat all of the collector sles, 42 W. ZEIDLER Like P. anophthalma, the presence of this species at Dalhousie Springs on the edge of the Simpson Desert suggests that it ts a remnant of a once more widespread fauna during a time when central Australia was much wetter than it is today (Krieg 1989), Acknowledgments lam inost grateful to Dr W. F. Ponder (AM) for his assistance in organising the 1985 expedition to Dulhousie Springs. He is also thanked for his assistance in the field and for collecting specimens. as is Ms D. Winn (AM). Ms K. L. Gowlett-Holmes (SAMA) assisted greatly with field work and the collection of specimens. Mr M. Adams Evolutionary Biology Unit SAMA conducted the electrophoretic analysis ound his expertise is — gratefully acknowledged, T also wish to thank Ms J. Thurmer (SAMA) for preparing bromides of the figures and Ms D, Churches who typed the manuscript, This study was supported by funds from the South Australian Museum. References Barware. J. L, & BakNARD, C. M, (1983) “Freshwater Amphipoda of the World. 1. Evolutionary Patterns” (layfield Associates. Mt Vernon. Virginia). Kaeo, GW. (1989) Geology pp. 19-26 Jn Zeidler, W. d& Ponder, W. F. (Eds) “Natural History of Dalhousie Springs” (SA Museum, Adelaide). RICHARDSON, 3. J.. BAVERSTOCK, P_R. & ADAMS, M, (1986) “Alloyyme Electrophoresis” (Acuderme Press. Sydney ). Sayer, O. A. (1901) Description of some new Victorian fresh-water Amphipoda. Proc, R. Sow. Viet. 13, 225-242. (1902) Description of some new Victorian lesh- waler Ainphipoda, No. 2. Lhid, 18, 47-58 Sairy, PC. (1989) Hydrogeology pp. 27-39 In Zeidler. W. & Ponder, W. Fo (Eds) “Natural History of Dalliousic Springs” (SA Museum, Adelaide). Sati. M. J, & WILLIAMS, W, D. (1983) Reproduction cycles ino some freshwater amphipods ino southern Australia. Mem. Aust. Mus. V8. 183-194. WILLIAMS, W, D, (1962) The Australian) freshwater amphipods 1, The genus Anstrechilronia (Crustacea, Amphipoda. Hyalellidae). Aus. /. Mar Freshw. Res. 13, 189-216. ZVIDLER, W. (1988) A redescriphon of Afracsiltonia capen- wis CK, H. Barnard, 1916) with a review of the genera ol the family Cemidae (Crustacea, Amphipoda). Ani. 8. Afi. Mus, 98, 105-119, (1989) Crustacea pp. 79-87 Ja Zetdler, W. & Ponder, W. F. (Eds) “Natural History of Dalhousie Springs” (SA Museum, Adelaide). (1991) A new genus and species of phreatic amphi- pod (Crustacea, Amphipoda) belonging in the ‘Chiltonia generic group, from Dalhousie Springs, South Australia Trans R. See. §. Aust, 115, 177-187. __& Powprr, WF (1989) Preface pp. ix-xi /7) AZeuller, W, & Ponder, W. F. (Eds) “Natural History of Dalhousie Springs” (SA Muscum, Adelaide), TRANSACTIONS OF THE | ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 121, PART 2 COMPOSITION OF THE STYLETS OF THE TARDIGRADE, MACROBIOTUS CF.PSEUDOHUFELANDI By ALAN F. BIRD* & STUART G. MCCLURE Summary Bird, A. F. & McClure, S. G. (1997) Composition of the stylets of the tardigrade, Macrobiotus cf. Pseudohufelandi. Trans. R. Soc, S. Aust. 121(2), 43-50, 30 May, 1997. The chemical composition of the two stylets of tardigrades has received little attention in the past, probably because they are so labile. We have studied these structures in the tardigrade, Macrobiotus cf. Pseudohufelandi, and have shown, using histochemical techniques and energy dispersive X-ray analysis, that these structures are composed of calcium carbonate. The response of M. cf. pseudohufelandi to the environmental stimuli of anoxia, differences in pH and temperature change demonstrate that this tardigrade can reabsorb its stylets under stress and reform them when the stress is removed. The possible evolutionary links between tardigrades and molluscs that have darts composed of calcium carbonate are discussed. Key Words: Tardigrade, stylets, Macrobiotus cf. pseudohufelandi, SEM, energy dispersive X-ray analysis, histochemistry, salivary glands. Transecttoas of tre Reval Seotery af 8 Anat (997), 122), 43-50, COMPOSITION OF THE STYLETS OF THE TARDIGRADE, MACROBIOTUS CF. PSEUDOHUFELANDI by ALAN F. Bikp* & STuarT G, MeCiure Summary Biko, A & Meci ure, SG. (1997) Composition of the stylets of the bordigmade, Muncohiotis cl preudohufelandi. Thins, KR. See. 8. Aust, 121(2), 43-50. 30 May, 1997, The chenucal composition of the two stylets of tardigrades has received little attention in the past. probubly because they are so Jabile. We have studied these structures in the tardigrade. Maereliuiis el psendohifelundi, und have shown, using histochemiesl techniques and energy dispersive X-ray analysis. that these structires are composed of calcium carbonate The response of M. ef. preadofufeland? to the environmentul stimuli of anoxia, differences in pLh und leniperature change demoustate that dis tardigrade can teabsorb its stylets under stress ane reform them when the Stress is removed, ‘he possible evolutionary links between tardigrades sind molluses that Have clierts composed Of calcium carbonate are discussed. Key Woks: Tardigrade, stylets, Macrahiotis cf pyendohufeland!, SEM, energy dispersive X-ray andlysis, histochemisiry, salivary whinds. Introduction Members of the Phylum Tardigrads (water bears) all feed using a pair of stylets Whose composition is such that they dissolve and break down on the death of the tardigrade and in some commonly-used lixa lives. Thus. they are frequently nor included in the camer lucida daiwings or photographs of the hucco- pharyngeal apparatus commonly used for taxonomic idemification. The general composition, shape and purpose af the twin stylets seems io be similir throughout the phylum and so the chemical compo- situ al stylets af one species probably holds for all Becuuse of theie transient nature, dissolving on deuth or fixation, turdigrade stylets have aot been anu lyzed, Th has been sugvested thi they ure calcareous in nature (Kaestner L968: Kristensen 1976; Wenck }O14. Kristensen unpub. both cited by Nielsen 1906) and that they resemble nematode stylets in forn) and fiunehion (Rigen 1962 cited by Kinchin 1994), The stylets of Macrebrolus cl psendohufelandi are curved, sabre-shaped structures about 400 pm in length that exhibit marked birelringence under poliar- ized light (Bird 1996), In this paper we examine the dissolution of the stylets in Various media under the polarizing micre= scope. Weir shut properties Under braght field microscopy and their elemental composiion using energy dispersive X-ray analysis in the scanning electron microscope, We alse examine the tetorma- bon of the stylets aller dissoluuon und disetiss these structures in the hehe of tardigrude evolution, 2 7havbord Row) Micha s. Aust Ste2 SSTRO bat and Willer, MH 2 Clon Osea 8 yust Sti Materials and Methods Locality, sot wee and extraction The tardigrudes were recovered from soil classified asa solonized brown earth from an experimental plot ona firm at Avon, South Australia (Bird 1996), After thorough mixing, 50 2 aliquots of the soil were pliced in a misting apparatus and processed as pre- viously deseribed (Bird 1996), The turdipricdes so collected were pieked out using a dental No. 3 nerve hroach. Walter which hid passed through the soil wis vollected from the misting upparatus and filtered through w 0.2 pin membime filter, The tardigrades were placed in a shallow layer oF this water ina ster ile Petri dish. Under these conditions, tardigrades remuined viable fora week or more without Feeding, Shlel dissetuiion The break down of the stylels was observed, on slides with vovershps sealed with nat varnish. i solutions with pH ranging: from 4-8 for different lengths of time (from the sturt of the experiment (a several days) and at teniperatures ranging From 15 - 30° C Observations were made with polarized light and differential interference contrast (Nomarski) aptics using a Vanex Olympus ABT research Microscope Sryler reformation Specimens were mounted in distilled water in scaled slides with pieees OF No. 1 hiss coverslips acling as spacers fo prevent crushing. These speci- mens were kept under observation. usiiig Nomarski optics. until the stylets could no longer be observed (usually about 3h), The vovership was then gently Al Ab BIRD & $.G, MeCLURE removed under a dissecting oicroseape and the motionless tardigrades phiced, together with some small nematodes to stimulate feeding activity, in shal- low distilled water ina Petri dish. After about | 2h the lardiaades, which bad resumed activity. were placed in sealed slides and re-examined under Nomarski optics, Prerzy dispersive X-ray analysis of the seylets under the scanning electron microscope (SEM) Because of the difficulties of obtaining intact stylets (rom tardigrades on sealed slides or after the use of acid fixatives such as PA 42), as used for the fixation of pemutades (Hooper 1986), a special lech- nique was udupted for these studies as follows. The fardigrades were transferred to a small drop of water which was then swamped with at excess of 6% gluteraldehyde in Sorensen’s buffer at pH 7,34, The lardigrades were then gently crushed ander a coyer- slip to permit rapid entry of the fixative without eaus- ing undue damage to them, After 2 h in the fixative the turdigrades were removed, washed four limes in 0.2 un membrane-filtered distilled water und placed ina sihall drop of the membrane-filtered distilled waler on the surface of a polished carbon stub. The lurdignides were manoeuvered close together and a second polished carbon stub was lowered on to the first so that the tardigrades were caught between the polished curbon surfaces, They were left in this posi- tion for 24 hto dry and then the uppermost stub was lifted direetly upwards, thus pulling the rardigrades apart and exposing their inner organs, including the buceo-pharyngeal region. This technique was also used on fresh. unfixed muterial. The stubs were kept iv a sealed dry container until examined and pho- togruphed in uw Cambridge § 250 Mk 3 SEM opera- ted at 20. KY using Ilford 120 roll film (FPS Plus), Energy dispersive X-ray (EDX) analysis was by means of a Link EDX system attached to the SEM. BDX analysis of characteristic lines with energies less [hun approximately O48 keV is pot possible with this delecior because of the absorption of the low energy X-rays by the deteetor’s beryllium window. Consequently, the direct identification of elements with alomic numbers less than that of sedium is not possible, Histochemistry: af the sedivary glanes daring sivler dissolution The unthraquincie dye alizarin red S’, sed as a stain for calcium. was made upas a 1% solution (50 me 5 ink!) either ima buffer solution at pH 4.0 or in distilled water (pil 5.0), The tardigrades were placed in a drop of the alizarin red S solution and gently Aldevh Chemical Compiny Fig, |. Head of living specimen of Maerebioniy cl preudo dufeland? immediately after beg placed in distilled water on a Sealed slide. Note the lwo eye spots in lower part of photograph and the two curved: stylets. Girrow heads), Norurski optics, Seale bur= 10 pm Fig. 2. Sume speeimen as in Piz. | but photographed just ailer the sivlets hud dissolved 166 rora later, Arrow heads tidicale the original location of the stylets. Nonrarsks opties. Seale bar = 10 pm. Fig. 3, The same as Fig. 2 but viewed under bright field optics. Arrow heads indicate the original Jocation of the stylets. This photograph illustrates the buccvo-pharyngeal structures used in the taxonomy of this genus. Seale hur = (0 um. TARDIGRADE STYLETS as squished. without causing gross damage to then, by placing a ghiss coverslip over the cop and with- drawing the stain sulation from under the coverslip with picees of Mer paper, The edges of the coverslip were then sealed with nail vartiish and the speeimens observed and photogriphed wider beight feldl optics. Results Disselatian af ite stylets The time tuken for diysolution ot the stylets uy cli tilled water. when westute of anoxia fas been induced by sealing the tardigrades under a coverslip on a side, is teriperatare dependent, Thus, a 30° 0) total stylet dissolution occurred within three hours whee us att 15° C the stylets were only partially dissolved afler this ume. The appearance oF the heud region ab Ww eb. prendohufeldid?, viewed under Noiiarski opues. as shown prion ty commencement of stylet dissolution we25° C (Vig. (und aller its completion three heurs later (lag. 2). The head region ts alse shown utter three hours. phatogriphed under nerinal bright feld Hplies (Fig hy, The sequence of sliges in the dissolution of the sty lens is ilustrated pos! cloudy ina seres ef pho lamicrieciphs taken under the polarvang miera- scope (Fig. 4\-H). Styler dissolution ty gridit aver the first (Wo hours (Pig. 4+ As) but aeeculerites over the third hour (Pig. 4 BH) so that warked ohunees were cletecled at approximately |S-iirmute intervals fron two hours onwards, Stylet dissolution Gecuered: more rapidly we ue solutions. than oy neutral ov alkaline solutions. took place within 50-90 tin ig O,0SM ACL and occurred immediately in. IM ACL and se is Similitr to Caco, inthis vespeet, The hirefringence exhibited by the touseles of the irivadiate pluiryox of Mech, pyewedolufelimet dil nor disappear over a period of three hours (Pig. +) ane the brighiness of these muscles, although mot as Warked as thut ol the siylets at the conmencemeat af the experiment remained cemstant througheut, Reformation of the spylets Turdivrales appear to huye the capacity to reform their stylets under favourable conditions in) what appears to be a reversal of the dissolution whieh ocvurs When they are placed in unfavourable situa tions, Speeimens that had been imduced to dissolve their stylets ri scaled slides and had hecome motran- less Were Observed to regan (heir Muscular activity and reproduce stylets. over w 12 h period, when placed in shallow distilled water inn uasealed Petri dish. in the company of several smull nematodes, This retormation did not always lead to precise realiznmtient of the stylets and the effieieney of this reformation and correct realignment may be depen dent on the state of health of the tardigrade at the commencement of the experiment. Enerey dispersive X-ray anitlysis ef the stylety Esxumination vader the SEM of the polished carhon surfaces bolding the dried. squished and disrupred lardivvades, cleatly shows whole stylets. or parts thereol (Fig, 5), By means of this technique, itas pos- sible to obluin suitable PDX analyses of the element tal composition OF the stents. Lecan beseen from the DX spectrin (Pig 6) thatthe sivlets ace rich in cal cium. wilh maximum X-ray robedsily Counts al erer sies matching the Cu pat 5.690 keV and the Ca Pal 4.012 KEV chinueroristic Nery lines. Hiyiochennyty of the salivary glenes divine siyfer dissolution When the jardigrades were venily squashed in a Ge salutian ob alc red Sut pH 3-0. rhe salivary glands surrounding the stvlers @raduully became: u deep red (Pigs 78) This reaction was bottle tnarked and Joculized, provided that the body of the turdi- wrade had been erushed co allow rapid penetration oF the stain. These vhinds. which are thought to be responsible for (he secretion uwod dissolution of the stylels (Rinchin 1994), ane apparently rich in-cul cium as indigated by their peachion to alizacin real S. Discussion Liformation on the nature of the tardigrade Siylets iS Sparse und hits received Tittle or no-satlention iA the two most recent general acequnls of (hese organisms, notably a most informative book on their general biuluey by Kinchin (1904) and a collection al papers published in the Zoological Journal of the Lrnnean Soviely i 1996 and edited by Mefines & Normiin. Tardigrade sivle(s were (hough! lotuye seine Sune ilarity with neniatode stylets by Riggin (1962 ) (cited hy Kinchin 1994). However. ibis clear that tardignade stylets (Bird (996) differ markedly from nemiutode stylets (Bird & Bird 1997) both in composition und structure, Thus, FA 4) }. a mixture of formaldehyde and avelic acid and & common lxalive for nema tudes. causes rupid dissolution of the tuntigrade atylets beauuse ol its low phl and the bigh tempera- ture used i the procedure, This fixative, however, does nol cause dissoluhon of nematode sbylets, Hoyers mediam. a moisture of guar arabic, chloral hydrate yond ghyeerol (Kinehin 1994) does nol dis- solve tardignide stylets and his the added advantage of functioning as a combined fixative and embedding medium. ininany illustrations of the bucea-pharyn- geal region of tardigrades, the stylets are not shown, presumably because they dissolved when fixed, The small size Ol tardigrade stylets (40 pin 1 AUR TARDIGRADE STYLETS 47 Vig. 5. Anterior region of dried and fractured Macrobiotuy cf. pseudohufelandi (see Materials and Methods) viewed under the SEM showing parts of the two stylets (arrow heads). The site of EDX analysis is circled. Scale bar = 10 pm. 12000 10000 Xray B000 Intensity 6000 (Counts) 4000 2000 .00 2.00 4.00 6.00 | 8.00 Energy (keV) Fig. 6. EDX spectrum of the stylet shown in Fig. 5, Fig. 4. Series of photomicrographs viewed under the polarizing microscope illustrating the stylets of a living specimen of Macrobiotus cf. pseudohufelandi exhibiting diminishing birefringence as they gradually dissolve in distilled water on a sealed slide as Shown in Fig. | - Figs 2 & 3. A. At commencement. B. After 56 min. C. After 96 min. D. After 113 min. LE. After 120 min. F. After 136 min. G. After 150 min. H. After 165 min. Arrows indicate parts of stylets. Note that the muscles in the pharynx do not lose their birefringence as the stylets dissolve. Scale bar = 10 am. 48 4.F BIRD & S.G, McCLURE length in Macrabiociy el preudolufeleard() and their lable nature, have made difieull determine theli composition. We fitye overcome this hy udupting it luchmique used to separate the layers tral uematode's cuticle (Bird & Deutsch 1957) whieh takes ante accuomil the adhesive qualities oF animal (issues fo the surfaces oo which they have been dried, In this instance. the Iwo surfaces were polished eurbon rods mounted on SEM stubs its described in Materials ane Methods. In some instiinces, this technique led Wa the exposure Of either (he whole or lure enough pices ol stylet to be recognizable (Fig, 5), These were then eusily photupruphed und subjected to chergy disper sive Neray analysis measurements 70 the SEM. These measirements clearly show that the elemental com pasion OF the stylets consists miiily of culenin felenents with omic numbers less (han that of sedi unture not detectable). "The rapid dissolution of the stylet in dilute acidsyand their unarked hrrelrireenuce tinder polivived light. indicite ahat they dre con posed of caletui curborate, ‘Thus, te copposa or ol durdigrade stylets differs fromm that of nematode stylet whichoare proicin in nariire, Hach tirdignide stylet Hes in the linen of a sali vary dha (Kinehin 1994). Kristensen (1976) cetted by Nielsen (996) has stveested that separate lobes ot (he salivinry whands are responsible for the Formation of the styles und their supports in the tondigradte, Ratillipes. Whether or not this applies to Mt ef. pacjalolufelanill pemiins to be determined, We have nor observed the Termation ov dissolution of the Sivlet supports oy Wis tindirade, iicating thal their chemist) compasioon is diflenent fom that of the wivlew. We have co inflienimbni on ihe origin and Cheiieal Compaspiion or the stylet Supports jy i, et parndolifelands Ou experinvnts indice that the seylets of Ae ef paren iifelandi are secreted and reubstithed by the salavy ekinds, We have shower that (Pies 78), wheo the Gindigrades ure gently squashed an a dilupe SHlivon of the stain alrzurin red Seal pit wl S20 ja sedleds the cell surrounding eagh stylet stairs as (Ire siylers break down, indicating (hat ealeturir is lifer tedhund diffuses throughour these two cells. Alizarin nl S is meoguived usa histuehemical rouge far ihe detoelony of caleium (Conn 1977) changing in colour from yellow toyed to purple over the pH tanve 3. 7-5.2, The gradual break down of the stylets ia slightly acidic aid anoxie covironinent, as demonstrated i) this paper, when photogruphed under polarized light (Fig. 4), closely resembles their step by step forma- tion us depicted in drawings by Marets (1929) (vited wn redrawn by Kinchin (994) showing stuges in their formation, Although stylets of caleiuin earboye ate ure urtusual in the animal kingdom und appear to be both unique and common (oall kudigrades, some- what similar structures are found in other graups at animals, Por instance, many of the terrestrial pub mone gastropods shoot durts of caleiuin curhomule into cach other prior te copulation. This behaviour is thoughr to actas a stimuiiie tothe sexo er The dart of Heliv dsypersa is about & - LO niin in lenge (Tompa 1982), at least ten times the si7e ot un entire tardivrade and some 200 Gyies larger (ha the 40 pe stylet of Macrobietiy cl) prendohufelandi, The cir, whieh takes ahout five days to fori. iy seecered Hr a dart sae that ts part af the reproductive system. The composition of (hese darts, wher subjected to energy lispersive X-ray analysis, is identical to the tardies gerade stylets with both Cy Kee uml Ca KE peaks (Lhint 1979). Phe evoluttonury srumheanee of these findings is Ohseure. However, they de lend same sup port to fhe phylogenctio position of the Nirdigryda proposed, as st result oF studies gia the 185 ribopornal gene sequence, by Moon & Kint (1990) These uuthurs shue “the tardigrade chide appear as an dependent lineave from the seimutode chide’ ana the culewreous niuire of the tardigrade stylet as Opposed to the protemmecous malice of the heitiatode stylet. lends siipport to this Hypothesis. Moon d& Kirn (1996) Jurther suggest Unat the turdligride clude is st sister group Of the pretostome cucoelomiale wssent blage (hat emnented belore the imolluses. animelids arthropods und sipunculids evolved. This would sug- gest independent evoluwon of the Culeareous str tires ja tarcdigrides and imialins. Somme deasere of The wags i Ou kiawledgee of the origins and relaGorships of the Hurdistada is bidicut: ed by the fact That hirdigrides are not mentioned ty a book or the origins and relationships among lower invertebrates. edited by Morris et val (1985), However sume priiitive Cambrniy fossils share some characteristics with both Onyehophora anid Tomigrada, ineluding poorly articulutect limbs ene ine in chiws (lobopodia, (eriiial ious abel the bis, 7, Whole specimen of Meerebrores ef, pseudehufetcd: That bas been crushed lo permit the entry of alizarn pou Scat pltS.0, The photograph was liken under bright fell ophes about 40 nia ater We conmnencement of stamiag. Showiny vye spots (small arrows) andl claws (iene ares). The area around the tidignide comlains material that fas been exid ed from the ruptarce animal and the two salivary glands Mar lie just anterion to the wuscular planyins are heavily stained, Seale bur = LOO ja Hig ¥ Rillitged portion of hig 7 showing eve spots (small arrows), the stamned salivary glinds (Sey che Wuscular pharyny (py amd material that has heen exuded by rupiuring to permit Gntey oF (he stiin Cire arawsy, Seale bar= 10 pm, TARDIGRADE STYLETS 49 @” 50 A.F. BIRD & S, G. McCLURE lust pair of legs merging with the caudal end of the body (Kinchin 1994). For these reasons it is specu- lated by Kinchin (1994) that the Tardigrada and the Onychophora have originated from a lobopodian line that diverged from the arthropod line in the Cambrian period. However, Nielsen (1996) consid- ers that Tardigrada are more closely related to Arthropoda than to Onychophora and suggests that all three phyla have originated from a group which he calls “Panarthropoda”, and that the ony- chophorans diverged first. It is clear that the origins und lineage of the Tardigrada remain obscure and that much remains to be discovered about them before they can be accurately (raced and defined. Acknowledgments We wish to thank J. Bird for constructive criticism of the manuscript and P. Kolesik for discussions on histochemistry. The senior author is grateful to Division of Soils (now Land and Water) CSIRO for accommodation and facilities and to the Australian Biological Resources Study for providing equipment that made this study possible. References Biro. A. F. (1996) Studies on the soil-inhabiting tardigrade, Maecrobiotus cf. pseudohufelandi, from South Australia, Traits, R. Soc. 8, Aust. 120, 147-154. & Birp, J. (1991) “The Structure of Nematodes” (Academic Press. San Diego). & Deusen, K. (1957) The structure of the cuticle of Ascaris lambricoides var. suis. Parasitology 47, 319- 32S, Coxn, H. J. (1977) Wilkins. Baltimore). Hoover, D. J. (1986) Handling, fixing, staining and mount- ing nematodes pp. 59-80 Ja Southey, J.P. (Ed_) “Laboratory methods for work with plant and soil nema- todes” (HMSO, London). Honr, 8. (1979) The structure and composition of the love dart (gypsobelum) in Helix pomatia. Tissue & Cell 11, 51-61. KAgSINER, A. (1968) “Invertebrate Zoology” Vol, TH (John Wiley & Sons Inc., New York). “Biological Stains” (Williams & Kinciin, I. M. (1994) “The Biology of Tardigrades™ (Portland Press, London). KRISTENSEN, R. M. (1976) On the fine structure of Batillipes noerrevangi Kristensen 1976.1. Tegument and moulting eyele. Zool. Anz. 197, 129-150. Mctwnnes, 8. J. & Noxman, D. B. Eds (1996) “Tardigrade Biology” Zool, J. Linn. Soe. 116, 1-243 (Academic Press, London). Moon, 8, Y. & Kim, W. (1996) Phylogenetic position of the Tardigrada based on the 18 S ribosomal RNA gene sequences. [hid. 61-69. Morris, S. C., Gtordk, J. D.. Gipson, R. & PLArT, TL M, Eds (1985) “The Origins and Relationships of Lower Invertebrates” (Clarendon Press, Oxford). NIELSEN, C. (1996) “Animal Evolution ; interrelationships of the living phyla” (Oxford University Press. Oxford). Tompa, A. (1982) X-ray radiographic examination of dirt formation in Helix aspersa. Neth, J, Zool. 32, 63-71. STUDIES OF THE EGGS OF MACROBIOTUS CF. PSEUDOHUFELANDI (TARDIGRADA) FROM WHEAT FIELDS IN SOUTH AUSTRALIA By ALAN F. BIRD*® & STUART G. MCCLURE Summary Bird, A. F. & McClure, S. G. (1997) Studies of the eggs of Macrobiotus cf. pseudohufelandi (Tardigrada) from wheat fields in South Australia. Trans. R. Soc. S. Aust, 121(2), 51-57, 30 May, 1997. The tardigrade, Macrobiotus cf. pseudohufelandi, and its eggs were isolated from soil from a wheat field at Avon in South Australia during the winter of 1996. The surface of the eggs was examined under the scanning electron microscope and was shown to be highly ornamented and reticulate with numerous “inverted goblet-shaped” projections. The dentate margins of the heads of these projections consist of clusters of coral-like globules. Statistical analysis of the size of the projections revealed two significantly different types of eggs in the samples. Examples of ornamented eggs were found in discarded exuyia, a condition which is not in accord with a recently proposed hypothesis on tardigrade egg evolution. Observations on feeding behaviour, population density and egg laying habits are presented. Key Words: Macrobiotus cf. pseudohufelandi, microscopy, tardigrades, eggs, morphology, evolution, soil. Treasaerous a the Rayal SocieiyafS, Mave, (L997), E212). S157 STUDIES OF THE EGGS OF MACROBIOTUS CF, PSEUDOHUFELANDI (TARDIGRADA) FROM WHEAT FIELDS IN SOUTH AUSTRALIA by ALAN FE. Bikn*® & STUART G. MCCLURE! Summary Binbo A, Bod Mec) ure $.0 (1997) Studies of the eges af Mocrbiotaeel. prevdolipfetind: VVardignada) fren Wheat Helds in Sour Australie Prams, A. See SAMs 12102) 5 1-57.30 May. 1997. The lindinnie, Mecoblotas eb pyendofmfetends, and lsc were isolated Hoi soil Prom a Wheat telat Avon in South Australia during the winter ol 1996. The surfice of Ihe eges was examined ender the seanning vleeiron microscope and was slieawin lo be iehty ornamented and redculate with numerous “verted gobler Shaped” projections, The dentate margins af the hends of these projections eansist of clusters oF coraldike globules. Statistical aniilysis of the sive of the projections reveuled (wo significantly dillerent types of eges in The sainples, EXaimples oF ormamented eges were Lound in discdrded exuyin, a condition whieh is Wor inadecart WITH recently proposed hypothesis on tandienile ese evellition. Observations on feedinu beh ioue, population density und exe layin habits are presented, Kby Woks: Macrae el, prendahujeland) iiteroseapy, tandimndes. eves. cenplology. eyaliinin sail 1 HY e. Introduetion Tivdigrades, also known as waiter bears ar mess piglets (Rinchin 1994), belong to a diserete phy lim ul vosmopoliiin distribution Tron diverse habitats ieluciig Waring. fresh water and semieterresuial cuvironiments. “The tardignides responsible for the was described jn this paper were identified as Mincrobiolus ef, preudohufeland: Iharos 1966 by S, Choxton (Aird 1996) and are semi-lerrestrivl, having heen isokited from sandy foun soil aia wheat fel ar Avon, South Australia. The tardigrade exe shell is ay tse taxonomic wile Co speeles identification, particuhurly in genera such as Minrbions where the shell is ornamented, Meacrohions et psendelufelund?, which are unty about SOO pm long hy [50 yim wide when fully sown, lay comparatively laree virculir eggs which hive brphly ommunented reliculited shell surlieces with numerous “inverted goblet-shaped” projections (Bird 1996), Ii tus been shown by Bertolini & Rebevehi (1993) What differences in exe shell morphology an Macrobiarus hufelindi, previously thought to be due fo Variability within this species, fall into seven iis- finel types that ure related to different animal hor photypes, Using ege shell morphology. along with other chiunteters. these workers have deseribed a num ber of new species from the M. fuife/andi group. bggs that belong to this eroup have pitted or reticulated shells with protruding processes shaped Jike inverted goblets, chulices. thread spools or “cooling lowers”, eu 2 Playtord oie Mitcham S. Aust, 3002 CSO taimd und Water, PMB 2 Glen Osmond S Aust 506d, Berlolanies af (1996) have suited that ornamented eps ire generally laid ree in soil or water ana smooth shelled egus ure laid in the moulted cuticle (exuviuiiy) These Workers inelide the tantly Muacrobiotidie, to whieh Mo ef. pweredeluefelumell belongs. in those families that hay tree, ornamented cees. Benalani ef al. (1996) Iawe put Torwand an hypothesis ta which they explait the evolution ot lurdigrade epes, In this pauper we examine the structure of fhe eve shell of At ef psendelvgetmed? und measure the processes provuding from the surlice of the ce shell. We also discuss its egg living habits in relation ly the Wypothesis of Bertolami et ad (L996) allel con) ment on feeding behaviour and population density. Materials und Methods The lardigeades were collected on ES July [996 fron sandy loam Soil and from (he same Tneulity at Avon, South Australia clatitude 34° 14S. longiude 138° 19° EB) us those collected previously. using the sumpling technique described by Bird (1996), The soi) samples were collected in mid-winter so that the sites were wet and the kirdigrades were feeding, reproducing und depositing czas. Vhe tardigrades und other metofauna, consisting predominantly of nematodes. were isolated from tits soil over a period of three divys using a misting appa- ritus (Yeates & Bird 1994). Tardigrades and nema- todes Were counted and tardigrade eggs were picked oul using a dental No. a nerve browch and examined. alive, in distilled water under a coverslip, with the light microscope begs to be examined under the seaming electron micrascope (SEM) were fixed in ta nm A. F. BIRD & 8. G. McCLURE EGGS OF MACROBIOTUS Ch. PSEUDOHUFELANDI 6% gluteraldehyde in Sprensen's phosphate buffer at pH 7.2 and at 5° Cand kept ui this Gxative for-sey- eral days. They were then washed three Himes in dis- tilled water and sonicated in an Elna T420 sonivator al adrequency of 35 kHz for W) see or unt shown by Microscopic observation to be free at debris, The eygs were freeze-dried by placing them between membrane lillers which were frozen rapidly by placing them tna slurry of freon cooled by liquid nitrogen. These filters with the eves were quhickly (ransterred to a [reeze driee and freeze-dried at - 70" C, The dried eges were picked up with double-sided tape whieh was ullached lo un SEM stub and couled with 30 nin of wold. This material was examined und photographed m u Cumbridge § 250 Mk 3 SEM operated yt 20 KV using PP4 Plus Tord roll filn. Results Numbers and feeding Atthe tine of collection the nutio of tardigrades Lo nematodes in SO gy of soil was 842297. Feeding on Hhenmatodes was also observed during the course of (this investigation. and the nematodes held by the tardigrades usually did not move although once a nematode broke Iree and moved uway. In one instanee, a tardigrade wus observed to be arching its back in the manner of a scorpion, curing feeding uclVily Buy levine ane eyes Leys thal were about 90-100 pm in diimeter in (he living unfixed stare (Pigs 1 2) and about 60-70 pun io the fised and dehydrited sue (Pig. 3), were Tad cither within exuvia (east cuticles) (Figs | 2) or free (Pigs Ai Figures 2. 3, 4. 5 clearly show that (hese eves buve a morphology of lhe MW frafelaimlé group with reticulited shells and charucteristicully uplurncd-chiilice-sNuped — protanding — provesses. i i Although ihe putrern of the reticulavions remains the same with the apertures on the reticulate surface of the cee shell being about 0.25 yar in chameter (Fig, 6), the shape of the protruding processes falls inio two distinct groups. Type tis shorter than Type IL is narrow at the base and has a wide distal head (Fig. 4). Conversely, Type His taller wider at the base and hus a narrower distal head than Type T (Pra. 5). For 10 processes af cach type. the differences in the mens ure statistically significumt with 95% confi- dence (Table 1) When the processes were examined under the higher magnification of the SEM (Fig. ©), the dentate cog-shaped margins of the head were shown fo con- sist OF clusters of globules with a structure resem bling madreporarian corals in uppearinee and were approsmmately 0.5 pin in diameter. The mudreporarian globules and reticulated shell surfaces wre simalar in both the shell types desenbed above. Diseussion Bertolani ep ai. (1996) have proposed an hypothe- sis fa expliin the evolution of tardigrade egys. According to these workers, the eggs of tardigrades have evolyed us a result of two events, the Urst bemg the wequisition of Ornamentation and the second the use of the shed exuviun as the site for exe laying. wilh the subsequent loss of ornamentauion, Thus, the ornamented eggs of the Macrobiotidae are thought te be laid free, Ole observations that the ornamented eees Of MO cl, psendolufelandi can either be band m exuvia (Pigs |. 2) or free (Pig. 3) do not appear to he th aceord with this hypothesis. The exuyia conning ortamented eggs were transparent and devoid of body contents and chu not appear to be females that had died before completing cep keying. A possible explanation Vor the laying of the e#gs in ihe exuyiiun by the Woch, pyendohufelundi fromthe solar Aven, Tahu) Le Meaviiements af the nee ipes vl prmceses prendiding fron the ene shell sifaces oe) Macrobioms of peuctohiufelanada, Type No. Heigl (int) Mean + 5D Ritiye i in a4 5 AAS Il 1 (ib (4+ 57-74 10.95] = 2.26 247 + Basal width (yim) Distal Width (lim) Menit +58D — Ringe Moan + ALD Range 43 0.2 3.048 5.3 2 3A? 5.3 2 405-4 2.1 ul Uy See WAAR 16,07 SD = Stundund Deviahon, | = Students (test Fie | Boe of Macroblons ch pyendelifelonads im cast cunele (esuviun) Arrows Tdicare posmon ofcust lees and ehiws. Bright field optics, scale hur = FOO yin, Nig, 2) Same specimen wt tgher magnification showing shell projectors (Pp vovered by cast cugele (OC), Bright Held opdes Scale bur = 20 pan, 54 A. F. BIRD & S$, G. McCLURE EGGS OF MACKROBIOTUS CE PSEUDONUFELANDI is that this may be an adaptation to the hol, dry: sum- ners experienced there. Our studies on the tardigrades and nematodes in soils froin Wheat fields at Avon in oidwinter when the soils were wel and (he populations of the mer: fauna could be expected to beat their peak. show that tardigrades make up) a SubstanGal component, although they are not as numerous us nematodes. The ratio-ot 84 tardigrades to 297 nematodes per 50 y soil found in these eXperiments varies at other sites where the tardigracde numbers per 50 ye soil may: be less ane nematode nutnbers greater (Bird 1996). However. i is Clewr hat the tardigrade presenve al Avon is wide- spread, The lardigrades isolated from soi at Avan feed on bematodes and can survive hot dry summers m an anhydrobiotic state (Bird 1996 ). Feeding on nemia- todes was ulso observed durmg the course of this mvestigation and the nematodes held) hy the tardi- erodes did not usually appear to be movitig. although they were coiled and therefore probably not dead. suygesoig tous that they might have heen paralyzed by some type of injeeted narcoue. Since there is pa Information on tardigrade diversity and geographical distribution in South Australia. frether studies are warranted. particularly on their feediug habits, is they may hive w role inthe biocontrol of the parasitic nematodes which have heen shown to oceur in the wheat felds at Avon (Yeates & Bird 1994), The line structure of the egy shell iy the genus Macrobioms is of greal iixonomice inportance, Prom our Observations on the ultrastructure of the exp shells of the Avon turdierades, wv would seem that there may be two populuions GF Ma pseudolufe- LA > lanai in the Avon soil or there may be two different Species, neither of which completely resembles those described so fur for (he Aufelaned: group (Bertolani & Rehecchi 1993: Biscroy 1996). The reticulated sur- fave of the shell und the structure of the glabules on the heads of the projections are Similar in the two torms of egus deseribed above but the diameter of the apertures on the reticulited surface of the ege shell is much less than that shown in the eggs of other members of the /iv/claned? eroup described by Bertoluni & Rebeechi (1993), Kinchin (1994) and Bixerov (1996). To our knowledge. the globules on the heads of the projections have not been described belore and might prove. together with the reticulated surfuee. fo be usetal taxonomic eriteria al the fade lend? group is further divided on the basis of egg ormumentahon. Clearly further studies ure required on the taxon omy of these lurdigrades und on their distribution in the semi-vrid aerivultural areas of South Australie dnd olfice Sika! regions of the Australian continent, Acknowledgments ‘The senion-autior is graterul for cogrant front the Australian Biologigal Resourges Study which pro- vided tuvilities that made this research possible anc 10 CSIRO Land and Water Adelaide: for aeeomntoda tion and equipment Dr WR. Miller, Department of Biology Southwestern College Winfield Kansas USA and Dr W.I,. Nicholas. Department of Botany und Zoology ANU Canberra ACT are thanked lor constractve eriticism of the manuseript References BeRror ANT, RB. & Reaecont £. (1992) A revision ol the Maeralyonrs Jrifelannli oroup (Vardigracta, Macrobiotidae), will same observations on the Gxogon we chunicters of ctitardigrides. fon, Sormtay 22. 127- 192, & Chaston, SK. 1996) Phylogenetic siz nifivance of epee shell variiion in tardigrades. Aol. Lin Soe. V6, | AY aR- Hien A. F (1996) Studies on the soilanhabiting taurdigrade, Macrobioms of prendahafelmd: trour South Australia Trams. Ro Soe S. Ausé, 120, 147-054. Riserov. Vo 1(1996) ‘Lordietades ot the Thinyr peniisuli with descriptions. of lwo new species. Aival I La Sea Mh. 215-257, Kawcnin, TOM, (194) “Phe Biology oof Tareligrades” (Portland Press. London), Yeares. GO. Wo& Birep. ALT (1994) Some observations on the influence of agriculiueal practices on the nematode fiumac of some South Australian soils bandane Apple Nematol. 17, 133-145, Fie 4, Scanning eleetron mieragruph of whole cegs mt Maerobferiy ef, psendoinfelind’ showing the inverted: goblet shaped prajections of Types F(T 1) and 112). Seale bar = 20 pin. Fig 4 Scanning eleetron micrograph of part of the egg shell surfice of a Type | eg showing projections with) hartower buses-wnd hurger dentate cow-shaped heads thunthose.ol Type WH, Note similarly rehendaled surfaces of ery: shells, Scale hours 3 on EGGS OF MACROBIOTUS CF. PSEUDOHUFELANDI 57 Fig. 5. Scanning electron micrograph of part of the egg shell surface of a Type Il egg showing projections with wider bases and smaller cog-shaped heads than those of Type I. Note similarly reticulated egg shell surfaces. Scale bar = 5 um. Fig. 6. Scanning electron micrograph of the head of one of the Type II projections at higher magnification showing the madreporarian globules (arrows). Note the size of the apertures on the reticulated surface of the egg shell on the top right hand side of the photomicrograph. Scale bar = | pm. TWO NEW SPECIES OF ASPHONDYLIA (DIPTERA: CECIDOMYITIDAE) FROM HALOSARCIA SPP. (CHENOPODIACEAE) IN SOUTH AUSTRALIA By PETER KOLESIK* Summary Kolesik, P. (1997) Two new species of Asphondylia (Diptera: Cecidomyiidae) from Halosarcia spp. (Chenopodiaceae) in South Australia. Trans. R. Soc. S. Aust. 121(2), 59-66, 30 May, 1997. Two new gall midge species are described from South Australia. Asphondylia inflata sp. nov. was found at Port Adelaide in swollen branches of Halosarcia pergranulata subsp. pergranulata. Asphondylia ericiformis sp. nov. was found at Lyndhurst, at the southern edge of the Strzelecki Desert, forming spherical, spiky galls on branches of H. indica subsp. leiostachya. Descriptions of the larvae, pupae, males, females and galls are given for both species. Key Words: Diptera, Cecidomyiidae, Adelaide, Strzelecki Desert, South Australia. Transactions of the Royal Society of S. Aust. (1997), 121(2), 59-66. TWO NEW SPECIES OF ASPHONDYLIA (DIPTERA: CECIDOMYIIDAE) FROM HALOSARCIA SPP. (CHENOPODIACEAE) IN SOUTH AUSTRALIA by PETER KOLEsIk* Summary Koesik, P. (1997) Two new species of Asphondylia (Diptera: Cecidomyiidae) from) Halosarcia spp. (Chenopodiaceae) in South Australia, Trans, R. Soc. S. Aust. 121(2), 59-66, 30 May, 1997. Two new gall midge species are described from South Australia. Asphondylia inflata sp. noy. was found at Port Adelaide in swollen branches of Halosarcia pergranulata subsp. pergranulata. Asphondylia ericiformis sp. nov. was found at Lyndhurst, at the southern edge of the Strzelecki Desert. forming spherical, spiky galls on branches of H. indica subsp. leiostachya. Descriptions of the larvae, pupae, males, females and galls are given for both species, Key Worbs: Diptera, Cecidomytidae, Adelaide, Strzelecki Desert, South Australia, Introduction Halosarcia indica subsp. leiostachya is a small, decumbent to erect shrub widespread along the coast Halosarcia 1s a plant genus comprising 23 species — and around inland salt lakes of mainland Australia commonly called samphires. The genus is endemic — (Wilson 1984). It is a common plant in the Strzelecki to Australia except for AH. indica (Willd.) Wilson which also occurs in Malaysia and other countries bordering the Indian Ocean (Wilson 1986). Two species of Halosarcia were found to be infested by two undescribed ~~ gall =midges — (Diptera: Cecidomyiidae) collected in South Australia during 1996. These gall midges are described in the present paper. Asphondylia inflata sp. nov. causes swellings of branch segments on H. pergranulata (Black) Wilson subsp. pergraniulata (Fig. 1) and A. erici- Jormis sp. noy. forms spherical, spiky galls on branch segments of H. indica subsp. /eiostachya (Benth.) Wilson (Fig. 2). Halosarcia pergranulata subsp. pergranulata is a ‘amar’ 8 shrub about 0.5 m high which grows in southern Australia (except Tasmania) associated with coast- Fig. 1. Gall of Asphondylia inflata sp, nov. on Halosarcia lines, estuaries, swamps and margins of inland lakes pereranulata (Black) Wilson subsp, pergranulata. Scale (Wilson 1984), The plant forms a substantial part of bar = 10 mm. the vegetation cover of saltmarsh flats north-west of Adelaide. These saltmarsh flats are areas covered by small, hardy bushes that grow on the landward side of the mangrove swamps. Areas that are regularly inundated by tides are typically dominated by Sclerostegia arbuscula and Sarcocornia quinqueflo- ra, while areas that are only occasionally flooded are dominated by Maireana oppositifolia and Halosarcia spp. In May 1996, a large number of galls caused by A. inflata sp. noy. was found on H. pergranulata subsp, pergranulata at Port Adelaide, about 400 m south of the Torrens Island bridge. Fig. 2. Gall of Asphondylia ericiformis sp, nov. on © Department of Horticulture, Viticulture and Oenology Faculty of : ‘ Agricultural and Natural Resource Sciences, University of Halosarcia indica subsp. leiostachya Wilson, Scale bar Adelaide PMB1 Glen Osmond S. Aust. 5064. = 10mm. P. KOLESIK 60 NEW GALL MIDGES FROM HALOSARCTA al Desert where it grows tna variety of habitats, includ- ing salt lake margins, open elay plains and gibber plains, Trois one of the dominant plants around Lyndhurst, where. in bebruary 1996, all examined shrubs exhibited a low (o moderate infestation by the gull midge Alericifarnity sp, ney, Material and Methods Galls were sampled trom Halosareia tndicu subsp. leiastuchye at Lyndhurst (15.16, L996) and Helosercte pergrdnuldi subsp. pereranulaca at Port Adelaide (5,y, 1996), The galls collected on both occasions were processed im one of two ways. A small nimber Was cur open and the larvae preserved in 70% ethanol A larger number of galls was kept in plastie bags ane the lirvae were reared to adults. Pupution look plaice within the galls. Pluste bags were exgurt- inca daily and emerged adulis preserved. together Will tein pupil skis, tn 706% ethanol, Canada bal- Sunk amounts of type speciinens for microscopic c\amination were prepared according to the teeh- mque outlined by Kelesik (l995a) ALL rmeasure- ments refer lo Lype series. The type series and other mate rekined in 70% ethanol are deposited in the South Australian Museum, Adelaide [SAMA and the Australian National Inscet Collection, Canberra [ANIC]. Genus Ayphandyite Loew, 1850 Loew. 1650; Dipterologische Beitriige. 1850: 21 and 37 (us subgenus Of Cecidopinia Meigen, 1803) Type species: Cecidumiia serothanny Loew. 13s. Jace 38 (des. Karseh. IS77): Revision der Gallmiieken: 15), Asphondvia is a worldwide genus thal currently Comprises some 260 deseribed species (Gagne 1904) Tt cantiins species that have a ventro-distal spine on the first tarsomere, the ovipositor with) lurve hasal lobes, female fMlagellomeres 9 - 12 progress- ively shortened. the ganovosite with aw ventro-apucil lobe anda dorsally situated gonostylus that is about as Wide as long and hears two basally merged teeth. Asphondylia inflata sp. nov. (FIGS |, 395, 7-9, 12-14, 1-18, 21. 24-27. 31, 44) Halotype: &, Port Adelaide. South Australia | 34°50" S. 138°30' Ef. cinerged 6.v.1996, P. Kolesik,, reared from braneh gall on HW. pergranudata (Black) Wilson subsp, pereranulini, gall collected 5.,]996, 12 | 283 |SAMA|. Paratypess 1a. 382, 2 pupae. 1 pupal shar [SAMA], be. 324. 1 pupa. | pupal skin [ANIC], all same dati but emerged 5.9.-13.vi.196: 7 lurve [SAMA], collected with holotype. Orher material: 29 2, 13 popil skins (SAMAJ, col fected with bolowpe, Mule (Pigs a-5, 7-9. 12, 14) Colour: selerotized parts of body dark brown, non- selerotized purty of abdomen orange. Head: Antenna: seape broadest distally. length 1.3 x breadifi ar distal end, 1G - 1-8 % length pedicel, pecicel about ay broads long. first Tagellomere 2,0) ~ 2.2% length of scape. flagellomeres evenly eylin- drical. eireunfila dense, equally distribured along scemients. Eye ficets close together. hexugunalid, cye blidge 6-7 facets long, Frons.with (2 - 18 setae per side, Lubella reduced in size. fused. Talerally wath 3 G setae. setulose. Masillary pulps 3 segmented, length of third segment. us well as total Jeng. vari able Thorax: Wing length 3.5 mm (range 3.4 ~ 3.5). width |b mm (14 - 15) Se cell pigmented proxt mally. Anepistermunr and anepimerou covered with scales. Claws of all legs subequal in size, similar in shape, as Tong as empocis, Abdomen: Genitalia: zonocosties short. with long veolo-apieal lobes gonostylus with 2 unequal apical frecth. ventral about 2 x length of dorsi: aedeimzus elongate and narrow. Fermele (Figs (4, lo-bS, 21. 24) Frons with 9 - TT setae per side: Twellth tagelly- mere sometimes fused with eleventh. Cireumliti comprising two longitudinal bunds connected by two short transverse bands.. Wine length 3.4 mun (2.7 - 37). wittth 1.3 min (11 - 14). Seventh ubdominul sternite 22 x (2.0 - 2:3) Jength of sixth. Genitalia: ovipositor 2.2 x (1.9 - 24) length of seventh sternite, cere) glabrous, Other characters-as i inale, Bins 3-140 4 Head of mule Asphomdylia inflata sp. acy un frontal View. 4. Genilatie of mule A yplieid\ a ciflatet sp. now, in dorsal view. 5. Gonostylis ol male Awpliwidvbie difatd sp wy in posterior view, 6. Gonostylus of male Asphonidy ta erieifarniin sp. (ov. Ti posterion View, 7. Last three Magellameres al malé Asphotdlia inflira sp.nov Bo Sixth Mazel lomere of pile Asphoiedviia fafiita sp. ney. 0. Male Asphendyia aafliid sp. nay, Last tirsomere with chiw and empodi-~ wi, LOd TL. Maoathiey palpus Of tale Asplandviierertedoriuy sp, doy. 12, First tursamere of male Asphondvlia miflo- He sp. NOY 13. Wing of male dyphondyiie vefer sp, now, Seale bars = LOO wor 4.4. 7-122 50 pint 5. @. S00 us 13, P. KOLESIK NEW GALL MIDGES FROM HALOSARCIA 3 Papa (Pigs 25-27, 31) Colour: abdomen orange. remaining parts dark brown Total length 3.0 mim (2.8 - 3.3). Antennal horns serrate medially. P83 pin (al - 191) long Upper and lower frontal horas simple. Two pairs of papillie on lower face. one of each pair with w seta Prothoracie spiracle broad at base. narrow on distil half curved beyond tracheal Opening at mid-length, Abdominal segments 2-8 with bwo pairs dorsal papillae. two pairs pleural papillae, and one pair yen- tral papithie. all papillte setose, Abdominal dorsal spines simple, prominent pairon last segment curved faterially, Lasse dasteee larva (Vig. 34) Coluur: orange. Integuiment covered with dense spietiiae. Length 3.0mm. Head capsule strongly pig- mented, postero-hitend extenmsiods shatter thar length of head capsule. Spatubie with two tong, pom ed anrerioe teeth. shaft narrowed pear mighdle. Widened again posteriorly, surrounded iiteriorly ind Jurerally by extensive pigmented, glabrous urea Papilhur pattern generally ay lor Asphondylte (Mahi 1955) exeepr only 2 lateral papillae on each side of sputulicaind ho terminal papillae visible au the avail- able specimen Lovnitoey: Phe name “hake” pst Latin adjeerive Tor i Fkated referring: 6 the appearince of the galled beanel, Galland hinlogy Brweh seaments of Milosareia poreraniulitte subsp, pererunilate infested by this gall midge are 2 - A times larger than normal in-votume, greyish green und Nard in eontrast to the vivid. green colour and sof texture of uninfested brunch segments (ig. 1) Bach wall has one to three chunthers, with one lurve in wich chamber, The chamber wall is fined with bard. pale-green, 0.25 - 0.33 mint thick tissue, Pupation takes plaice inside the gall A] cieular brown area appears on the top of the gall before the pupa cuts a opening wilh its antenoal horns, Ov 5 May. 1946, at Port Adelaide. the gulls appeared very, conimon in the host plant popalition covering sever al hundreds of me. Asphondylia ericiformis sp, Woy, (FIGS 2.6, 40, 11, 15. 14. 20, 22, 23, 28-30, 32, 33, 35) Holotype: 2. Lyndhurst. South Australia {30°17' 8. 138921" Bj, 204.1996. PB Kolesik, reared front brinch gall on Malevarere indica subsp, leresracliya (Benth.) Wilson, gall collected 15.411,1996, 121284 |ISAMA], Porapes: V2 20) pupa. lb pupal skin [SAMA], IFS, bE pupae 7 pupal skin PANEC]. all suame dita but emerged 25-27,1),1996: 3 larvae [SAMA]. 2 Tar- var |ANIC], collected with holotype, Other material 7 pupae. + pupal skins [SAMA], col lected with holotype, Male (Figs 6, 10, 11) Frons with 6 - & selue per side, Wine length ab mm (2.1 - 4b). width bmn (9-16. Bye bridge 6-9 facets long, Ventral tooth on vonostylis as long ats dorsal, Otherwise as in AL laflane Female (Figs (5, 1Y. 20, 22. 23) Prous with 4-8 sete per side. Wing length 3 ny (2.3 3.9), width 1.2 nam (09 Tab. Seventh abdominal sternite 2.305 (2.0 - LA) length of sixth, Ovipositor 19% C14 - 2.0) length of seventl sternite, cere) tibrous, With line oF teeth dorsally. Otherwise ere, Hflata, Mipa (Figs 28-30, 32) fot length 4b a (4t - oh 7 Antennal horns 198 pre (1S4- 214) Jone. Upper frontal born simple, Hooother Hore presen. Dorsal spines of last abdomi- Hal segment bout same length, strarehe, Lusr instar larvae (Wigs 43,345) Total lengil 3.2 nin (2.4- 4.8). Spatula with long, pomled anterior eth, shaft stort broad, parallel sided. Tiree Literal pupitlae on eieh side of (horace segments. fo terminal papilkie visible on tevailubly specimens, Otherwise as an ch. iffede, hivinalogy The name “erieifecnos” isa composed Latin adieu tive formed from “erieius” (hedgehog) and “Tormnis’. referring to the the hedgehop-shuped gull, Gall und binlegy This species transforms brinch segments ol hias 14-24. 14, Female doydiendvia iillare sp. now, Busal lobes on pyipesitor in dorsal view. 15, Female Aspliondyliin eris olferinis ap. nov, Basal lobes of eyipositor in dorsal view, 16, Antenna al femiule Avplroweylica aiflata sp. voy. 17 & 14, Masillary palpus of female Asphondylic inflata-sp, nev. 19 & 20, Maxillary palpus of female Asylonedylia erroifarnits sponoy, 2), Sixth (hiwelomere af femiile Ayphendyvlie taflate sp. nov, 22. Female Asphondv lia evicifarmis sp. nov. bn of abdomen in laleral view, 23, Bemale Asphondy fie ericiformis sp. nov. End of ovipositor in lareral View. 24, Female Ayphondytia inflate sp. nov. End of ovipositor in duteral view. Seale bars = 100 pin 14-21-24, 24: S00 pm 22 64 P. KOLESIK NEW GALL MIDGES FROM HALOSARCIA 45 Halosarcia indiey subsp. leioytachye into spherical, spiky. monothalamous galls, each oceupied by one larva (Mig. 2). Outer diameter of gall @ - 12 mm- inner diumeter 2.0 - 25 mm, Chamber wall lincil with hard, brawn, 0,25 - 0.33 mim thick tissue. Pupation dukes place inside the gall, On 15 February, 1996.at Lyndhurst. (0 examined shrubs of the host pluat bore a total of about 200 gully of the new gall midge species, The galls contiined larvae or pupite. Remarks Four species Ol Aspliondylia have been prev ioushy knowe to occu i Australia (Gagné 1989: Kolesik 1995b). Asphondvlia dedenueue. a South Australian species commMan inthe Adelaide Hills, ntalforny term - inal branch stems und primary leat veins of Dedonuea Vixcase (Sapindaceae) (Rolesik 1995b). ‘Twa species, A dogwi and A. rubicinde, were described in the pre- vious century from adults caught in fightin Sydney, New South Wales. and their bialogy as unknown (Skuse PRSS, L890) The fourth, AL filli, was described froma feniles and pupae bred from an unknown plant in Darwin, Northern Territory (Edwards 1916). These last hice species, which were deserihed superficially and can not be conipured on then’ deseriptions, are not Considered in the present paper, but | plana review of Australian Asplonedytte spp dita hater stage. Morpholowical similarities helween the (vo new species and the fact that their réspeetive lost phuts belore to The saine wepilis, sugwest ac lose rehitivoship. They form a distincive group that excludes AL coderede Asplimedyvifa dodeitucue differs from the two few speciesin the following respects. Adulis hove prominent labetla and scapes is long as brow al the distal ends, The male hus u shoraeyentro-apreal lahe on the gomocoxite, srall lobes udjuwent ta iweth on the gonmostylus and ue wedeagus Wich shorter than the gonocusites. The seventh abdumrmal steiiie in the fenyale is three Himes longer than the sixth. The pupae has nie frontal bors. (be antennal horns ave trivnwelae and seimaled: Lamerdiy amb the prothapacie spiacte 7s Noe Considerably beodder atthe base The ared sur rounding the spatula inthe hurva 1s not przmented. Adults of the hwo new species diller from euch other most prominently in the shape of the gono- stylus and the end of the ovipositor, The yonu- stylus hears teeth of unequal length and the end of the ovipositor lacks external processes in A. iafla- tt. In contrast, the gonostylus of A. erieffermis bears teeth of equal length and the end of the oVvipositor is serrated. Mare differences are evi- dent in the earlier developmental stages. Uhe pupa ol A. inflate has both upper and frontal horns pre- sent and a prominent pair of dorsal spines on the last seginent is curved laterally: ind. ericifaraiy the pupa has the upper horn only and all dorsul spines on the last abdominal seginenr are equally stony and straight, The lacvae differ in the shape of the spatula and the sumber of lateral papillae two per side in A. daflata but three per side ii A. eriomiis. The wo new species resemble cach other in the shape of the antennal horns und pro thoracie spitucles in pupac. tbe antennal seaments in adults. the ventro-upicul lobes on the gomoeox- ites In midles and the relative lengths of the sixth und seventh wbdonimeal stertites mi females. The elongation of the yventro-apical lobe on the gone coxite 14 Unique to these bWo species und disuiog- wishes them from the other Ayala vlia spp. Acknowledgements Tamogratetul to PG Wilson, Western Australian Herbarium Comino for the wentification of Halayarore jidicu subsp. feisrarhya. Ro J. Chinnvek, Seuth Austrauhan Herburiuin Adeliide tor the identification Ol Halayetrcd pergranulie subsp. pergranntere, Tt, B Reardon who led a Sooth Avseatian Museun collecting trip during which Asplorndylie ericifearnis sp roy was discovered, A. Stark. Walle Germany for providing copies of Kasel’s aod Loew's papersaind J 0) Gray, Department of Hortieulture, Viti culture and Oengloey University of Adeltide and Ro} Gayne, Sysiemahe Entomology Laboriwory USBA Washinglom DC for ihrem coments on ay early dralt of the manuseript Pips 25-48. 25 Aqphaniviio dative ap naw Antenne parent pupa in ventral views 26. Ayydanediir inflate sp. new, Anteriot park el pupa rater stew. 27, Aphids tie (u/ledé sp, now, Prothoragie spiiaghe of pupa, 2S, daplonedyiar eeloiorays ap Hs, ARLEFLOL pare a parpa Or ventral yas. 24, lytic enietovins sp. roy Anterior parhol pupal in literal viesy 40, Aguiar lo iciloruits spony, Prothoraeicspiracle of pupa, 47 Pupa ol lsphendy tig taflala sp now. Last ybdtonn ind seanene iT dorsal view. 72. Pupaol Ataenedviie cercdor ssp now, Least abdoniina segment in dorsal view. 43 Larva ol Asplondylin eretiowiy spy, now Last (wo abdominal) segments in dorsal view. 442 Agterion part of tive ot Ayphornmdyiir ioleatiesp, toy, tn wut view, 35, Plead quid fist thorauie segment of larva ol lap hemidvdien vei ipernity op) noy, meventral view, Seale burs = TOO ya 66 P. KOLESIK References Epwarps, F. W. (1916) Two new Australian Diptera. Ayn, Mag, Nat. Hist, 103 (8th Series, Vol, 18), 498-502. Gaane. R. J. (1989) Family Cecidomyiidae pp. 152-163 /n Evenhuis, N, L. (Ed.) “Catalog of the Diptera of the Australasian and Oceanian Regions” (Bishop Museum Press and E.J. Brill, Honolulu). (1994) “The Gall Midges of the Neotropical Region” (Cornell University Press, Ithaca New York), KaArscH, F. A. F. (1877) “Revision der Gallmiicken” (E.C. Brunn, Miinster i. W.). Ko esik, P. (1995a) A new species of Eocincticornia Felt (Diptera: Cecidomytidae) on Eucalyptus fasciculosa in South Australia. /. Ausir. ent, Soc, 34, 147-152. (1995b) Asphondylia dodonaeae, a new species of Cecidomyiidae (Diptera) damaging leaves and branches of hop-bush, Dodonaea viscosa (Sapindaceac) in Australia. Trans. Ro Soc. §. Aust. 119, 171-176, (1890) Loew, H. (1850) “Dipterologische Beitrige. Vierter Theil. Die Gallmiicken. Zu der Offentlichen Priifung der Schiller” (Das Konigliche — Friedrich-Wilhelm- Gymnasium zu Posen, Posen). Moun, E. (1955) Beitriige zur Systematik der Larven der Idonididae (= Cecidomyiidae, Diptera). 1. Teil: Porricondylinae und [tonidinae Mitteleuropas. Zoologicc 105, 1-247, Skuse, PF. A, A. (1888) Diptera of Australia, Part 1. Proce. Linn, Sov. N.S.W, (2nd Series) 3, 17-145. Diptera of Australia, Nematocera Supplement 1, /hid. 5, 373-412. WILSON, P. G. (1984) Family Chenopodiaceae pp, 81-317 In George, A.S. (Ed.) “Flora of Australia.” Vol. 4 (Australian Government Publishing Service, Canberra). (1986) Family Chenopodiaceae (Dysphaniaceae) pp. 236-311 J Jessop, J. P. & Toelken, H. R. (Eds) “Flora ol South Australia”. Part | (South Australian Government Printing Division, Adelaide). A LATE EARLY CAMBRIAN TRILOBITE FAUNULE FROM THE GNALTA GROUP, MT WRIGHT, NSW By J. B. JAGOo*, LIN TIAN-RUIf,, G. DAVIDSONE, B. P. J. STEVENSY & C. BENTLEY* Summary Jago, J. B., Lin Tian-Rui, Davidson, G., Stevens, B. P. J. & Bentley, C. (1997) A Late Early Cambrian trilobite faunule from the Gnalta Group, Mt Wright, NSW. Trans. R. Soc. S. Aust. 121(2), 67-74, 30 May, 1997. Trilobites from a new locality within a siltstone of the Cymbric Vale Formation, western New South Wales, are described here as Redlichia cf. ziguiensis Lin 1978 and Hsuaspis cerastes (Opik 1975). The species described as Strenax cerastes Opik and Estaingia bilobata Pocock from nearby localities are included in a single redefined species, H. cerastes. The genera Pseudichangia Chu & Zhou in Lu et al. (1974) and Strenax Opik 1975 are placed in synonymy with Hsuaspis. The fauna described here is of late Early Cambrian (Late Botoman) age. Key Words: Cambrian, Trilobita, Australia, New South Wales, Hsuaspis, Cymbric Vale Formation. Tramsac rons af the Raval Soetwly of S, Aunt (1997), T28(2), 67-74 A LATE EARLY CAMBRIAN TRILOBITE FAUNULE FROM THE GNALTA GROUP, MT WRIGITT, NSW by J, Be JAGO*, LIN TIAN-RUU, G. DAVIDSON, B. PJ. STRVENS! & C. BENTLPY® Summary IAGO, 7. BL. LIN TARE, DAvIDSOM. GL STEVENS, BL POA Brainy, ©. (1997) A Late Early Chombrin (nobite Taunule from the Goalit Group, Mt Wrinht, NSW, Fras, Ro See S, Agst, 1202) 67 74, 30 May, 1997, Irilubites from a new Jogilily wilhin a siltstone of the Cyimbrie Vale Formation, western New South Wales, we deseribed Here as Aeedliohia el. ciguiensiy Din 1878 and Ayuwaypis cerustey (Opik 1975) The species deseribed us Sireniay cerates Opik und Exiinga blobare Pocock from nearby localities are included jn asingle redefined species. 7 coraytes, The gener Pyeudi tema Chu & Zhou mn Lier al (1974) wad Ate Opik ITS are phicecd! ia synonymy with Asay The funy described here is of hile barhy Cumbrian (hale Bolomun) ape, Kiy Worps: Cambrian. Trilobita, Aastedia. New South Wales. /Avneapis, Cymbrie Vale Formation. Introduction Davison’ discovered a small area of fossiliferous siltstone of the Cymbric Vale Formation in the Mt Wright urea, western New South Wales (Pig. 1). Davidson identified the trilobites as Avreren cerayles Opik and Myiiagia bilobala Pocock of Opik (19756). More samples were colleeted reeeutly and the results of the examinition OF the fossils are given below. ‘The Barly to carly Middle Carmbrian Gnalte Group comprises three formations, from bottom to top: (he Mount Wright Yoleanies, the Cymbrie Vale hormution and the Coonian Pormation, The distrih- ution af these formations us shown in Fig. | is ater Warris ind Rose (1968), The Gralla Croup crops oul along a broad valley west of Cymbrie Vale homestead and ts mostly con- Jined by the Mt Wright Fault on the east and the Lawrence Poult on the west A smaller urea of Cymbrc Vale Formation crops out in the core of an anticline west ot the Lawrence Fault, The stricture of the Gniltt Group is very imperfectly Known as a Deputmenel Append Gieolupy. Sehowbot haineernnge Tlityorsiy of Sant Atistedlie The bevels So Atist 3095 Department ol tgerth Sciences. Nanjing Uaiversiny 22 Hankew Bowl Nun 22008 People's Repablie of China, Geology Dopartinicit. Gliversity al isnt GPO Box 252 Hobart Tas TOOL | Cealogieal Survey of New South Wiles, 32 Sriphide Street Brokon HAL NSW 2880, Dowibsoa, GO (MOR A contibniron tothe geology at the Mi Wright area, BSe Clons) thesis, Austin ational Unversity (unpub) Wokiis. Bu (1967) The sfiiligriphy unit pakicontoloyy of Horthwestera Mew South Wales. PhD thesis. Sydney University (itu result Of geological complexity (faults and: folds), poor outcrop in much of the area and a lack of hed ding in the abound voleunies of the Mt Wrelit Voleunies, The siltstone to fine sandstone outerop from which the fossils were collected (Pig. 1) is tsoluted trom Oller outeraps by u vover of soil and cobbles repre- senting a Cainozoie lag deposit, Henee there ts nu outcrop continuity with any identified Gnali Group formation. The outerop also contains po diagnostic rock type. Phe structure of the Caialla Group is sulft- crently complex to render it impossible to vonfident- ly plive the ourerop ina formation, Davidson placed this and nearby oulerops im the Cyimbric Vale Formation, while Krise (1982) placed the pearby oulerops in the Coonigan Formation, As shown helow, the species deseribed by Opik (1975h) from the Cyinbric Vale Formation as Lyteiingian bilatrte and Strenay cerasfes are found wt the locality being considered here, Na such fossils ate known fan either the Mt Wright Voleanics or the Coonan Formation, Henge, the outerap is probably Cymbrie Vale Rormation. Previous Work ‘The teem Gialtt Group was mtroduced by Warris” who ussigned ao Early to early Middle Cambrian age to the group. Prom limestone in the Mt Wright Voleames, Warns identified algae and archacocy athids, including Tidlumoeyatiiy tracheatiy Taylor, From the Cynmibrice Vale Formation he identified Fetiingia biloba, Calodiseus sp. and Pagzerides sp, and assigned a mitddle or hue Barly Cambrian age, He identified the trilobites Redliehia idenea, Nyseridura saints. Pageria signtficais. FPeranopsis narmata. Orvetovephality sp. and J B.JAGO, LIN TIAN-RUL G, DAVIDSON, B. Pd, STEVENS & C, BENTLEY an Ow Vay 5 kilometres ais REFERENCE Ganazdic seuinierits, Willies Puplialy Go Ustune, sv ate Wyuatennity alluvin. Deyoran- Garlornlurous Rayanddle Punmatiin acifl Weetlerno sandstones, Mile - vate Davenian Snake Cave Sandetcne sarlatone, menor congloinemte wn a us ine = | Pally - Middle Ordeyician Rowena Fornitation, Salton, quanwte, sitslone, Dorvpyge sp. fromthe Coonigan Formation, indieat- ing an early Middle Cambrian age. In jerms of the Cymbric Vale Formuion, Pleteher (1964) first recorded archaeoeyatha from lenticular limestones near the base of the writ. Kertise (1982) noted four arehdeouyath Hinestone-bearing lenses: he considered two to be of Aldabanian or Lenan age and two to be of Lenan age, The uppermost! Cymbric Vile Formation contiuins well-bedded sandstones amd cur honutes with abundant fossils. Opik (1975b) record ed the Wilobites Divesuy alt, wraiidosius, Testeliipedal hilohata, Strenax verustey, 8. fletcheri, Serveadiscus deedalus. Meniscuchis tienen, Pinvenesttes fregumt und Pagetie, the monoplicephoran Scenely retieulta Ahe inarticulate brachiopods Borsfardia el. cdelutd, Livenlella and Nenbolus, eoerinoids ini sponge spicules, Age and Correlation As shown below we consider [hat the species described by Opils (LY75H) as Stremaey cenisfes anal Lying bilabrea tron nearby toualities should both be ineluded within w single redefined species Heispis cerastes. These: localities Opik eonsidered ju be equivalent to the Siemshtyk gol Horizon oF the Altay-Sayan region which, as noted by Zhuravlew und Oravestock (M94) can he roughly correlated with the Botonjun of Siberia. The archieoeyathia from the Cymbric Vale Formation whieh oecur below (he trilobites described by Opik (1975b) were correlated with the informal archaeoeyathid unit the Sviigecnema favus beds by “Aburaviey und Gravestack (1994) which wus considered fo he equivalent to the middle (o Lite Botoman by these aulhors, These authors considered that the Syrinvecnema fevus beds fall within the trilobite based Pararaie janue Zone of Jell (in Bengtson ep al. 1990). dell suggested that the Po jamae Zone can be Lal Garten Crifly Ghlidian Nootumbiilia Randslonae aafiletorn Getiolomerats, shale, limertore, correhded with the Botoniui and probably with the jute Botoman (see also brief discussion by tage MOOTWINGFE GROUP I i : Pp GWALTA GRA e) 7 =| Miditta anthem Conner F atnalior a Wnestons shale Cambrian Eyre it Yale Pay nation = Sele Paiee (terperiniitte: gelertnyes Sd Wie} Tar) Cambie Mb Wright Maluanios, Piormmciets snd del Volearies. rrierar alate. lenestanes < ARLES PAINT PURI fee ee Gai]! Ronit Ged. Te cuns TipMicHU ete ibbAbacelt Stl Peet HAS) Selene GIS, Het geo Sel iiss, PDR EA Letter | ee? ol bale Vrotermoyhe Early Garnbriiny’) Mitt fede, WOLD Stutle sthditinS Causal litnSti ties aan [sane Hihyenvetin betes Sryjron bait veth t kicks ol ant aal lliatite Tal oy) Lips ded Shi Trinh . Hui auleas --£ yey ° Walerloreu ——P——- Aplutyic ee fasulLowhly 1996), As noted above, the exact stratigraphic posi- tion of the present fuuna is a Hille unelear bul it prob- ably occurs high inthe Cymbric Vale Formation und Stratmraphically close fo the faunus deserthed by Opik (lY7Sh). The Coonipan Formation which over- lies the Cymbric Vale Formation has a very carly Middle Cambrian ave (Opik [975a, Jell (975; Shergold ef al, T98S), this providing an upper limil fo the age of the Tauna currently being desembed, Palmer and Rowell (1995) note that Heuuspia is Vig. 1. Geolowy of the Mt Wright siren shawing lovality of fossils described herein, Geolagy from Warris_ Rose (1968 Davidson! Kruse (1982), BL Stevens and A. Criwloul Cimpub. field data) EARLY CAMBRIAN TRILOBITES FROM WESTERN NSW nu elosely related to the Siberian genera Bergeriniel/us and Bergeromieaspiy which churaclerise the Botonjn of Stheria. In China Reedlichie cigviensis Gveurs tn the tate Lurly Cambrian Poldealenus Zone af the Yargtze Gorge This is approximutely equivalent to part of the Toyontin of the Siberns Platform, The above discussion suggests that the faunule deserihed herein is of late Barly Cumbriin age und probably of Late Botomuan age. Systematic Descriptions ‘Temmnvlogy essentially Follows Darrington es al (1959), Specimen miimbers refer to the palacontalag- ical colleertow of the South Austria Museum (SAMP). Family Redlichtidue Poulsen 1927 Sublimily Redhehime Poulsen 1927 Genus Reedlehia Cossman 1902 Redlichia ct. ciguiensix Lin V978 (Tig. 2A) W978 Redlichia cieuieasis Lin a Zhou & Lin 1978, p. 14S. PL 20, Fi. 8. Material Gne incomplete cramadium, SAMP35 344, Deseripliamt Criuntdium subrectinyular, about 35 mm long. Glahellw (including ocvipial ring) about 0.9 length OF crunidium. Ghibella tapers evenly forward to rounded glabellue anterior. Narrow shallow axial fur rows. Three pairs of lateral glabellar furrows; |p, moderately deep. directed slightly to posterior and meet at centre OF elahella to form ain evenly posteri- orly arched furrow: 2p pair, shallow, direeted slight- ly to posterior, 3p pair, represented by faint depressions on glabellar margins, Occipital furrow deepest later- ally: shallow medially where itas urched slightly to umeries, Anterior border furrow of moderate depth: anterior border very gently convex, and of uniform Jenvth (Sag), Very short (sagz_) preglabellur field. Small anterior areas of fiXigenae gently Conver; long wide palpeboul lobes extend from just lo anterior ol Sp furrows to level wilh occipital furrow. Narrow, distinct palpebral furrow shallows pasteriarly. Pulpebral areas of fixigenue very gently convex. Width of palpebral areas of fixivenie at their widest is about Gre-third that at Glabella. Presculir seetions Of fasial suture markedly divergent, Discussion A single incomplefe internal mould of cranidiumn makes a definite specific identification difficwlt However. 1s sive, outline, slender glibella, the posi- hon of the postenor of the palpebral lobes. the nature of the preglabellar area and the shape of (he preocu- lar seefion of the facial suture suggest Reelichia signiensiy Lin (ie Zhou & Lin 1978, p. 145. Pl. 20, Fig, $). However, it Jiffey's (a having a more tapered glabella, a forward arching at the centre oF the oceip- ital ting. and a more sharply rounded glabellur ante ion Superlamily Lllipsocephaloiden Matthew L887 Family Ichangiidae Zhu LO80 Genus Astiaypiy Chung 1957 Ayyayprs Chang (9572p, 45. Lu ed al, 96S. p. 85; Zhang ef al, JOSQ. p. 244; Sell in Benytson yr al. 1990. p. 310. Palmer aud Rowell. 1995, p. 16; Nedin. L995. p, 36. Lstaingia Pocock lU64. p. 462: Opik 1975h, p. 10 Peudichanwia Chu & “Zhou iy Lavet af, WTA, p, 93: “bute Zhane eral, TY p, 230. Syenax Opik 1975 p13. lotvietlee Zhang & Zhu in Zhang ef ul, LYS, p. 247. Type species: Lusiafops vitesis Chang, 1953, p. 12h. Pl. 2, Figs 1-16. Disettssron Jell (in Bengtson et af. 1990) placed Lsrafnera Pocock 1964 und Zhadella Zhang & Zhu (in Zharrg eluh I980)in synonymy with Asyaspisy Chang 1957, a move supported by Palmer & Rowell (1995). although the fitter authors expressed some doubt as ta the suprageneric position Ol Asuaspis In addition, Jel sugested that Strenar Opik 1975. might be regurded as a junior synonym of Pyeudichangid Chu & Zhou fn Lo er af, 1974 fron southwestern China. The present authors sopport Jell in placing Srremety in synonymy With Pretadicfangre bur alsy consider that Prendichangiir ty a junior synonym of Asaespis, thos also placing Saeaay in synonynry with Aswaspiy. The following species ot Pyeudiehangia as figured in Zhang eral 1980, P. daniianensvis (Chats) (PL 76, Figs 13. 14; PL 77. 1-3). PB ronwejfensis Zhu (PL 77, Figs 6-8). shuvessiy Zhang & Zhe (Ph 134, Fig. 2) and Pyeuelicfhangia (21 inwalite Zou. PL 77, Pigs 910, may all belong ina single species. we. Ayuasyrs domiaoeusis (Chang). 2haviella hubtensis Zlang & Zhu (see Zhang ev al, M80, PL 134. Pig. 3) also appeurs fo belong in H. demmianensis us far as. can be determined from the available figure although the glabelki of ckiiqeerses extends Curther forwards than dior the type species ol Ayuaspiy, A. vinenyiy, and the anterior sections of the facil sutures. of sinensiv are more divergent than those of daniteoen- wis. We would cegard these as specifie tather Un genenc differences, 7) J. B, JAGO, LIN TIAN-RUL, G. DAVIDSON, B. P. J. STEV | a : ENS & C. BENTLEY Fig. 2. A. Redlichia ef. ciquicnsiy Lin 1978, SAMP35344, cranidium, internal mould, x 1.5. B-L. Hsuaspis cerasyes (Opik 1975). B. SAMP35329_ cranidium, external mould. x2. C, SAMP35340, cranidium, internal mould, x5. D. SAMP35335, cranidium, internal mould, x2. E, SAMP35331, cranidium, external mould, x2. F. SAMP35343, crani- dium, internal mould, x2. G. SAMP35337, cranidium, internal mould, x2, H. Top. SAMP35327, cranidium, external mould, x2. Bottom. SAMP35328, cranidium, external mould, x2. 1. SAMP35342. cranidium, internal mould, »2. J. SAMP35333, pygidium, external mould, x6. K. SAMP35332, cranidium, external mould. x6, L. SAMP35341, par- tial thorax, external mould, x2. EARLY CAMBRIAN TRILOBITES FROM WESTERN NSW ra Jel in Bengison er af, 1990) quenced the taxcian- ic position of Estatigia bifobata trons western New South, Wales as described by Opik (1975b). 1h our view the specimens deseribed by Opih (19735b) doy Nol belong in Aywespis brloheta (Pocock 1964) hecause the vlabella oF Opik's Speenmens caxtends further to the anterior than does that of TL dflahena as deserthed by Pocock (L964). In addition, in three of the foureranidia figured by Opik. there is a marked forwards expansion of the glabelhke wheres si the pe inaterial of Ho bilibata figured by Poeork (1964) the glabella is either tapering forwards or has a slight waist, In our view the speciinens desenbed and fizwred hy Opik (l975h. po UL PL ot, Bigs 1-7) belong i the same species us those described by Opik (197Sb, po be PL 2) Figs 1-6) as Sirenen eefistes. Apart from the specimens fgured hereny. Numerous other specimens are available ad there is a coynplete gradation from specimens such as those figured here as Figs 2B.E whieh are quite similar tu the eranidium ilustated by Opik (lO7Sb, PL 2. Kaw. T) as the Holotype ob Siren cerastes get ck sp. Noy lospecnnens such as (hose figured herein as Pies 2G, 3C which are indistingdishable from those figurecl as Evtcinge biloba by Opis (WY75h. PL. 1). Hpik (L975, p. 16, PL 3, Pigs 1, 2 teu. Fig §) erected a subgenus of Srremay, el Syren (Semarscus) bused oo four crunidia with the oaly Hlustemed cranidium having a lengrh of only 3.8 nim. Hoos clearlyun ioinalure speennen which should net he the basis of a new taxon Jef (1990) erected Ayueapis eceipilasping which) is churacterised by a relatively long preelabellur Held crossed by a preglabellur median Hdwe and the pres- ence of a short slender oecipial spine. It is possible that the specimens described by Palmer & Rowell (1995) trom the Central Teansantaretie Mountains as Hsnaspis Cf Te bitohune (Pocock) belong in necimtospiia, becuse the Anturcue specimens hive a glabella of similir length to S72 accipitesyina as well as a osimihir preghibelkur fedian ridge. However. the Antarctic Specimens have an occipital node rather than an occipital spine. Hynaxpis ceraytes (Opik 1975) (Pig, 2B-L, Fig. 3) 1975 Stree cerasias Opik W975b, p. EPL 2, bises 1-6, leat. Mig + 1975 Exrainwid biloba Povoek. Opik 1975b. p_ 11, PL 1, Figs 1-7. Marevial Almost twenty crumdia, one partial free cheek (SAMPS5434), un ingoniplete thorax (SAMP35341) ancl an incomplete pygidium (SAMP35333). Dienst Species oF (/yvaxpis with distinedly expundedt anter- joy part of glabella: length of ghibella (excluding occipital ring) ubout O.75-0.8 that of cramadium fexcludipe oecipiul rine): occipital ring bears a sprog which in some speeimens is a siiall nodes i others ibis long und slender. Description Gently convex cramidivm with wichth slightly grester than length pag.) Deep wide uxtal furrows shallow anteriorly. Gently conver elubella of leneth fekcluding occipital jie) about O 7548 thar of cranidium, Cabell ticreuses tn width io antenor with sheht waist qear tp furrows: wlibella widest at eve ridges, broadly rounded yhabellar unteriar, hour pairs Jatera) glabellir fureiiws; Up fureows made rite jy deep and each extend abopr 1/3 of distinee aeross glabell and directed sliitly fo postertur, 2p tarrows shiutlow. extend about 1/3 distuice aenoys glabella. and directed very Slightly lo posterior, Ap furrows quite shallow and directed almost simight aeross vlubella; 4p furrows oeeur as small indistinct pits, Occipital ring longest medially; it bears a spine the length of which varies considerubly rom specimen Lo specimen. In Semi specimens a smull nade pre- sent; in others there is u lone slender spine which muy hive a leneth about W735 that of ghibellia (excluding oecipital ring) in muture specimens (Pig, 2D) and somewhat longer io immature specimens (2B\P). Occipital furrow shallows medially, Very short (sau,} venily conves preatabellie held, Anterior bor dey vently convey with similar tength (sue) be prewlubelkir field. Poorly-developed jalectrum pre- sent tr some specimens (Pre. 2G). Shallow anterior border furrow. Gently conver palpebral areas ob fy- igenue have width ubout sume us that ol vlabella, Prominent eentro-pasteriorly placed palpebral lobes have a length (exsug.) about 0.55 that of glabella. Postetior of palpebral lobes meet the broad postero- lateral border furrows, Well-developed eye ridges slightly narrower (han palpebral lobes, Shallow palpebral Surrows, Narrow posterolateral borders, Preocular sections of facial suture diverge forwaris atubout 30" to the lransverse, Short postacular sec- bons of ficial suture. Narrow postecoliteral border The single parbal librigenae fis a Wide border which extends mto an ineomplele genal spine. It Tits wreliculate ornamentation. Thorax with at Jeast nine segments. Convex axis has willl about 0.3 that of segment. Centrally places nodes on 4th and Sth segments of ivailuble thoras. Shallow pleural furrows; short bread pleural spines. The only available pysiditun as poorly preserved. [tis Small and transversely etliprieal. Axis has a width about 0.4 (hat ot pysidiuim, Axis extends almost to 72 J.B JAGO, LIN TIAN-RUI, G, DAVIDSON, B, PJ, STEVENS & C, BENTLEY Fiz. 3, Uyuspis cerires (Opik 1975). A. Left, SAMP35325, cranidium, internal moul. x3. Right. SAMP35326, eranid- ium, ioternal mould, x3. B. SAMP35338, evanidium, internal mould. x3. C. SAMP35330, eranidiain, internal mould. 2. D. SAMP35339, cranidium, internal mould, x2. Bb. SAMP35323, eranidium, internal mould, x3, F SAMP35336 cranidium, internal mould. x2. G, SAMP3S3344, librigena. external mould. x2. LARLY CAMBRIAN TRILOBITES FROM WESTERN NSW 7" posterior margin. Axis comprises Wwe uaal tings plus terminal axial piece. Two pleural furrows and one imMerpleural furcaw present. Two border spines can be seer in avatlible specimen. Morphogenesis The simallest ciaiditind is that figured as 2K. In ihis specnnen the length oF the glabell (exclading oceipiial ring) is 0.7 that of cramdium, tr larger specimens such as those figured in 3B,D the glabel- la has a length of O.8-0.85 that of eranidiam, The rel- auve sizeof the oegipilal spine decreases ~ in siull- er specimens (2B.B) the spine has a relatively thick~ er base than tv nore mature specimens (3B.D), HNC UANT OU The considerable variation in the shape of the whibella, the path oF the fei suttire. the shape itd length of the becipital ring, and the length of the vecipiall spine. of the speeuncns described and: tig ured here as AL cerastes raises the qhestian as to the validity of plieiig all these specimens within a single species, However, us noted above, there appears 1 be a-coniplete morphologeal yariation present in the uvailable specimens ind henve we feel the erection of more than one species can not be justified. A/siesypis cerasieys us desevibed herein differs tram UL daandoensix in having a relatively shorter glabellay the glabella of AL, dantldwensiy esther reaches or almost reaches the anterior border furrow, whereas that ol 1, cerasfes stops short of the border, However, its worth noling (hal wilh respect to the length al the occipital spine, JE damioensis, as Tyured in Zhang ef al (1980. PL 76. Higs 13, 14: PL 77. Figs 1-3) shows con- siderable variauior in length of occipital spine as dues A. ceraytes. The anterior of (he ghibella of H, cerisres is more expanded Unan is that ol A, diamtevenvis. Tsuaspix necipimsping at sell (1990) tas a small well-developed oecipital spine. The spine of H. cerastes arises From further back on the oecipital rng than thal of JE eceipiospinag: the spines on sone specimens of 4d. cerayfey ure much longer than the biggest spines on /7, eccipitosping. In Ayers the ratio of the length of the glabella (excluding oceipi tal ring) to the length of the cranidiam (excluding oceipilil ring) 1s O7S0.805 in A, ecelpitosping it ranges from 0.65-0.70. The glabella of AL stesiy extends further ter wards than does that oF (7. cerastey. The unterfor part of the dlabella of 7. cerdsres is more expanded thin H, sinensis, H. yinensiy bears only a sthall pedi node rather than a prominent spine as mn many spec- imens ol AL ceresies. The anterior purt of the glubella oF Z/. ceraytey ts more expanded than that of J. bitebate: the glubella of //, verastes t§ velanvely longer than that ofA. bile bette, The glabetla of (ef T. bilubitia as described hy Palmer & Rowell (1995) 0s shorter than that of A, rerester. Acknowledgments Sample collection was carried out under a licence Irom the New South Wales National Parks and Wildlife Service. This paper is published with the perinission of the Director-Geoeral, New South Wiles Department of Mineral Resources, Lin Tiun- Rut was supported by a visiting researcher's grant from the University of South Australia, Dr PA, Jell constructively reviewed an earlier version of this puper. Mr JS. Bau gave valuable technical assis- lance. Dr C. Jenkins supervised G. Davidson's hon- ours project References BiAdison, S. Cuinway Momus S.. Coun. Bod. Jee 2 AJ & Rowancak. BOS. (900) Lark Cambrian fossils from Sou Austratat, Meni. Assoc. Auxtretlas Patueuntal, 9, | 364 Foereniba. BLO) (1964) New Tilo shells tron the Lower Ondovicnin ind middle Palaeozore rovks of New Sonth Wales. Ree. Ana Muy, 26, 283-294, HargRINGTON, HLS etal (L959) Arthropoda l pp. F560 hr Moor. RO Co (kd) Treatise on Invertebrate Paleonialogy, Par OF eGeologieal Society al Amerivent nd Kansas Loviversity Press, Luwrenee, Kansas). Jago. 1B. (1Yo) Some comments on he Cambrian Hine scale in relation to pecen! radiometric dling of Australian Cambrian rocks opp. (OS-(06 fe Linan. E., Chimes Vintaned, | Ak Gozolo Bot Mds) TP Field Conference of the Cumbrian Staze Subdivision Working Groups”, Spain, 13-21 September 1996, Meld Trp Guide une Abstracts. Jett, PA I975) Australing Middle Cumbrian codiseaids wilh review of the superfamily. Paloemndographice A 150. [17 — (1990) Trilobite pp. 257-322 Jn Bengison, §,, Conway Morris, 5. Cooper, B,J. dell, PAL & Runnesar, BON. Barly Cambrian fossils trom South Australia, Mom. Asysoe. Maytriles Palueantol 9. Krviss, PD. (hYTHT New Archaeoevathi front the Burly Cuinbrian al the Mt Wrucht area, New South Wales, AMlcheringa 2. 27-47, (1982) Archucoeyathian bipstaiiweaphy of the Gniulae Group ait Mt Wright New South Wales Fealeeoutographicd A UT7. | 212 74 J.B. JAGO, LIN TIAN-RUI, G. DAVIDSON, B. P. J. STEVENS & C. BENTLEY Lu YANHOU, CHANG WENTANG, CINBN YIYUAN, CHU CHAOLING, Lin, H. L., ZHOU, Z. Y., QIAN, Y., ZHANG, S. G.& Yuan, J. L. (1974) Cambrian trilobites pp. 82-107 Jn “Handbook of stratigraphy and palaeontology of southwest China” (Science Press, Beijing). [Chinese. English summary]. Nurpin, C, (1995) The Emu Bay Shale, a Lower Cambrian fossil Lagerstiitten, Kangaroo Island, South Australia. Mem, Assoc, Australas, Palacontol, 18, 31-40. Opi, A.A. (1975a) Templetonian and Ordian xystridurid trilobites of Australia, Bur. Min. Res., Geol. Geoplrys., Aust., Bull, 121, 1-84. (1975b) Cymbric Vale fauna of New South Wales and Early Cambrian biostratigraphy. hid. 159, 1-78. Parmer, A. R, & RowskLe, A. J. (1995) Early Cambrian Trilobites from the Shackleton Limestone of the Central Transantarctic Mountains. The Paleontol. Soc, Mem, 45, 1-28. Pocock, K. J. (1964) Eytaingia, a new trilobite genus from the Lower Cambrian of South Australia. Palaeontology 7, 458-471, Rose, G. (1968) Broken Hill, 1:250,000 Geological Series, Sheet SH54-14, Preliminary Edition. (Department of Mines, NSW). SHERGOLD. J. H., SAGO, J, B., Cooper, R.A. & LAurIE, JR. (1985) The Cambrian System in Australia, Antarctica and New Zealand. Jat, Union Geol. Sci. Publ. 19, |-85. ZUANG WENTANG, Lu YAN Hou, ZHU ZHAOLING, QIAN YL. Lin, H., Zou Zatyt, ZHANG SANGUT & YUAN JINLIANG (1980) Cambrian tilobite faunas of southwestern China. Palaeontol, sin, ser. B 16, 1-497 [Chinese with English summary]. ZHOU, T. M. & LIN, T. R. (1978) Early Cambrian trilobites pp. 143-154 7/n “Sinan to Permian stratigraphy and palaeontology of castern Yangtze Gorges” (Geologicul Publishing House, Beijing). [Chinese]. ZUURAVLEY, A. Yu & GrRavestocK, D. 1. (1994) Archacocyaths from Yorke Peninsula. South Australia and = archaeocyathan Early Cambrian zonation, Alcheringa 18, 1-54. POORLY PRESERVED TRILOBITES AND BRACHIOPODS FROM THE KANMANTOO GROUP, FLEURIEU PENINSULA BRIEF COMMUNICATION Summary The Kanmantoo Group (Fig. 1) 1s a thick (8-10 km) succession of predominantly clastic metasedimentary rocks which crop out in an arcuate belt from near Australia Plains in the northeastern Mt Lofty Ranges to the western end of Kangaroo Island. On Fleurieu Peninsula, the Kanmantoo Group rests unconformably on the Normanville Group’, the uppermost formation of which, the Heatherdale Shale, contains a poorly preserved fauna of possible mid to late Botoman age’. The highest well-dated fauna from the Normanville Group are archaeocyathids from the Fork Tree Limestone at Sellick Hill which indicate an age close to the Atdabanian-Botoman boundary’. These are at least 400 m below the top of the Heatherdale Shale. The top of the Kanmantoo Group is not exposed but lower parts of the group are intruded by the Rathjen Gneiss which has an age of 516+ 4 Ma’, i.e. late Early Cambrian or early Middle Cambrian on recent Cambrian time scales*®. The stratigraphy of the Kanmantoo Group is described in detail by Daily & Milnes”. Dansacnons af te Raval Seciew af’ S. Aust 1997), PRU 7 BRIEF COMMUNICATION 5 POORLY PRESERVED TRILOBITES AND BRACHIOPODS FROM THE KANMANTOO GROUP, FLEURIEU PENT The Koomintoo Group (bine Fy) is a thiek (R-TO) key succession al predominantly ehistic metinedimentury rocks which crop oot avan urenate bell from near Australi Plains in the northeastern Mt Lofty Ranges to the western end af Kanvaroo Uslund On Fleurieu Peninswla, the Kanimnuntoo Group rests unconforimably on the Normianyille Group), the upperfoost formahoa af which. the Heulherdale Stale contitins a pourly preserved fiw of possible mid to lite Boroman age’, Phe highest well-dited aun fom the Nonnunville Group are aehieocyuthids frou the Pork Tree Limestone atSellick PHI whieh indice in age close to the Atdubatiin-Botoman boundary. These are at least 400) 1 below (he top ol the Heatherdate Shale. The top of the Kuumainloo Croup is hot exposed bet lawer parts af the croup are iitruded by the Rathjen Gneiss whieh hus an age oF Alé+ + Mat ne. dane Hark Cambrian or earhy Misdle Combruia recent Cybern time The alnitigraphy of the Kainminton Group is described in detail hy Daily & Milnes’ The Cambrian sequences as exposed on Fleurieu Peninstila. Kangaroo Istand aad) Yorke Peninsula show considerible Tateral aud vertical lithologies variation’. Currently, the lanited blosianivraphie inforinanc makes it dilficull to correlate accurately trom one region to. another jm hence derermine the Gining and sequence of Bork stmupraphie and tectomie events One of the problems tn fhis regard has been the lack of biostrutgruphicully uselul body fossils in (he Ryuommantot Group. The only stich fossils reperted to dite are the imarnculie brachiopod Lingiledée from about 10 ta above the base ob the baswl unit ob the Kanmiantoo Group, the Carrickuliiga Plead Formation, a Carrickalinga (head!) and ‘possible hyolithid fron acar the top of the Curriekalinga Head Formation near Blowhole Creek!!, Trace fossils have been reported from the Backstuirs Passage Formation al Accommoedaitlon Hill near Prue! This qule reports the presence of Vurther inarticulate brachiopods (Fig, 2D), from about 100m above the base ob the Carnckuliga Head Formation al Cumickulinga Head. They are sivalaind poorly preserved, but appearto represent uff lexist (Wo Species. In quarry near Parawa (Pig. 4). 01 627,551 (Torrens Vale, 1S04100 topomaphie map) on Cullovonua Creek Road, the vuthors have found the firsh known (rilobiles from the Kinmiantoo Group, The specimens come froma laminated iehisiltstone. probably part of the Tunkalilly Formation but possibly upper Tapinappi Formation, To dite we have found dhoot 20 specimens, allot whieh are almost coniplete, thus qidivaling low enerey depasitionul conditions As on scales SULA shown in Fe TAB.LC the speciinens are poorly preserved, leclanically deformed gn of no blostratigraphic use. dlihough, within the fimits of preservation, they seen to represent au single species, However, (herr presenve pndicules the possibility oof the eventual discovery of hiostraticnplieally riselil iilohites front the: Keamgatoo Group Middleton Sandstone Petrel Cove Formation Balquhidder Formation Group Tunkalilla Formation @ Tapanappa Formation Talisker Calc-siltstone Kanmantoo Backstairs Passage Formation Carrickalinga Head Fm (4) Heatherdale Shale Fork Tree Limestone Sellick Hill Formation Wangkonda Formation 2 =) 2 1) 2 > e G E tw 5 = Mount Terrible Formation Fig |. Stratigraphic outline oof Normanville anil Kaniewloo Groups, Fleurieu Peninsula, showing Ue levels of the fossils figured herein, MW Pip, 2, ACC, “Trilobites from) quarry fear Porawal AL Rubber cust of caxternal mould, SAMPS35445, 33. B- Internal mould, SAMP35446, x4. CL) Rubber cast of external mould, SAMP3S347, x3, DEL tnarticutate brachiopods from Carrickulinga Head, DBD. Internal moulds. SAMP35348a and SAMP3S448b0 x13, ob fuurtially extoliated specimen, SAMP35349, x10. Specimen numbers refer to the palaeontological collection of the South Australian Museum, AUSTRALIA a Adelaide Fleurieu U Peninsula area Gulf St Vincent Carrickalinga Southem Parawaarea Ocean 2a see inset Outcrop of Dies Kangarcd Kanmantoo Group Island 0 10 20 30km 138° E Tunkalilla Beach 36°S Southern Ocean Fig. 3, Locality map. ‘Jago, J.B. (1994) Aust. J. Earth Sci. 41, 445-453. Jago, J.B. (1996) Some comments on the Cambrian time- scale in relation to recent radiometric dating of Australian Cambrian rocks pp. 105-106 /n Linan, E., Gamez Vintaned, J.A. & Gozolo, R. “II Field Conference of the Cambrian Stage Subdivisions Working Groups” Field Trip Guide and Abstracts. ‘Thuravlev, A. Yu & Gravestock, D.I. (1994) Alcheringa 18, 1-64. ‘Farrand, M.G. & Preiss, W.V. (1995) Geol. Surv. S. Aust. Bull. 54, Vol. 2, 54-57. ‘Young, G.C. & Laurie, J.R. (Eds) (1996) “An Australian Phanerozoic Timescale” (Oxford University Press, Oxford). °Tucker, R.D. & McKerrow, W.S. (1995) Can. J. Earth Sci. 32, 368-379. Tel. Daily, B. & Milnes, A.R. (1971) Trans. R. Soc. S. Aust. 95, 199-214. ‘Daily, B. & Milnes, A.R. (1973) Ibid. 97, 213-251. °Gravestock, D.I. (1995) Geol. Surv. S. Aust. Bull. 54, Vol. 2, 3-61. Daily, B. (1963) Rec. S. Aust. Mus. 14, 579-601. "Gatehouse, C.G., Jago, J.B. & Cooper, B.J. (1990) Geol. Soc. Aust. Spec. Pub. 16, 351-368. Daily, B., Firman, J.B., Forbes, B.G. & Lindsay, J.M. (1976) Geology pp. 5-42 In Twidale, C.R., Tyler, M.J. & Webb, B.P. (Eds) “Natural History of the Adelaide Region” (Royal Society of South Australia, Adelaide). J.B. JAGO and P.W. HAINES, Department of Applied Geology, School of Engineering University of South Australia The Levels S. Aust. 5095. SIGHTINGS AND STRANDINGS OF THE PYGMY RIGHT WHALE CAPEREA MARGINATA NEAR PORT LINCOLN, SOUTH AUSTRALIA AND A REVIEW OF OTHER AUSTRALASIAN SIGHTINGS BRIEF COMMUNICATION Summary The pygmy right whale Caperea marginata is a rarely sighted species known primarily from strandings. The latter occur frequently in South Australia, especially along the north coast of Kangaroo Island and near Port Lincoln’. Migratory patterns are not known although it has been suggested that this species moves inshore in spring and summer™*. Sekiguchi et al.° postulated that such a movement off South Africa may coincide with an increase in the abundance of copepods, one of the presumed main prey of C. marginata. Spring and summer are also the seasons when juveniles most frequently strand**. There appears to be a broad mating, calving and weaning period between June and February’. This paper describes two recent sightings of live animals off Lincoln National Park, about 10 km east-south-east of Port Lincoln, South Australia and summarises past sightings and strandings in that area. Other known sightings in the Australasian region are also reviewed. Transactions of the Rayal Society ef S. Aust. (1997), 121(2), 79-82, BRIEF COMMUNICATION SIGHTINGS AND STRANDINGS OF THE PYGMY RIGHT WHALE CAPEREA MARGINATA N The pygmy right whale Caperea marginata is a rarely sighted species known primarily from strandings. The latter occur frequently in South Australia, especially along the north coast of Kangaroo Island and near Port Lincalna!? Migritory patterns are not known although it has been suggested thal this species moves inshore in spring. and summer’. Sekiguchi ey a/> postulated that such a move- ment off South Africa may coincide with an increase in the abundance of copepods, one of the presumed main prey of Co marginata. Spring and summer are also the seasons when juveniles most (requently strand’4. There appears to be a broad mating, calying and weaning period betweer June and February’. This paper describes two recent sight- ings of live animals off Lincoln National Park, about 10 km east-south-east of Port Lincoln, South Australia and sum- mirises past sizhtings and strandings in that area. Other known sightings in the Australisian region are alse reviewed, While sailing a 6-m yacht in Spalding Cove, off Lincoln National Park, two of the authors. J. D. and B, F.. observed lwo pyginy night whales, an adult about 6 m long, accom- panied by a calf about 2 m long. Paired blowholes (Fig. 1) confirmed that the adult was a baleen whale. Species iden- tification was bused on the curved jaw-line, medium-grey colour, a falcate dorsal fin placed well back on the body and the adults broad back (Figs 1-3). The animals were seen between 1030 and 1130 h on 4 January and 1300 and 1400 h the following day. On both occasions they were deep in the cove al the fur south-western side, about 300 m from shore (Fig. 2). Water depth (as determined by a depth- Fig. | - Adult Caperca marginata in Spalding Cove, South Australia, 4/5 January 1996. Arrows show the paired blowholes and indistinct white bar behind the head Note also the broad back. Photo: J. Dutton. (AR PORT LINCOLN, SOUTH AUSTRALIA AND A REVIEW OF OTHER AUSTRALASIAN SIGHTINGS sounder) was about Sm, water visibility 8 m, surface water temperature 20°C and the sea was calm. Spalding Cove ts a shallow, sloping bay with a sandy bolton and extensive sea- grass beds, Immediately prior to the sighting on 4 January, a large U- shaped swirl about 4-5 m diameter, followed by a curtain of bubbles, appeared 2 m from the boat. About 2-3 min later and 50 m away a large animal, the presumed adult female. surfaced and blew. This was followed by the blow of the calf near the adult. The calf swam slowly around the boat coming close to the bow and turning off. much as dolphins often do, It swam with an undulating motion, surfacing fre- quently for air (30-40 see or as long as 2 min). At this stage the boat was under motor at a speed of 2-3 knots (3.5-5.5 kph) and the calf showed no sign of being disturbed by this. The calf alternated between swiniming next to the boat and the nearby adult about 20-30 mi away and sometimes swam above the adult, a common position for dependent calves of the southern right whale Cubalaena australis (C. Kemper Fig. 2 - Juvenile Caperea marginata in Spalding Cove, South Australia, 4/5 January 1996. The arrow marks the prominent, falcate dorsal fin set well back on the body, Photo: J. Dutton. 40. pers. obs. The calls grey back und sometimes its head showed as WH suifieed und blew. Later ecamination of pho- tos showed (hal the almost white ventral colour extended well up the side of the hody and that a distinetive dark at cea pately tilerrupted this just behind the head (Pig 3). A similar colour pattern has been observed ony South Atricad juvenile C. margitata® and at recently stranded neonate in New Zealand (van Helden pers. comm.). The adult swam jn lirge circles in the general vicinity of Ihe bowl, somerimes voming very clase 16 i and at fimes turning on its side. [showed no signof being disturbed by jhe presence of the boat. The adult surfaced to breathe much less often than the call. ubout every 3 mi on more. Later exumination oF photos showed that the adult was a medium erey colour with a pale band or eheyron just behind the head (Pig, 1), The belly was lighter (han the hack. Matsiioka also deserihed and iMustrated light chevrons on the backs ofa large group of adult Cyanine observed use (Table 2), The afiiials were in the same part af the cove during the allenngon of the next day when JD) and BOR returned to sail there, A net fisher operiting in Spalding Cove during hilt laniiay and tiich of February 1996 reported seeing a smull Whale there on sevent oceysions. His deseription al jhe animal does not allow species identificution but it may hive been CL mearuiniie. possibly one ob the same ania ubserved in eirly January by J.D. and B. Ti The other recent South Australian sighting was made dur- ing the aternoon al 7 July 1996 by RM, who observed, with the aid af binoculars, a small whale from the shore new Cape Donington lighthouse, Lincoln National Park (Big, 4. Hi was about 50 m from shore and 60 m from the observer, The weather Was sunny and the sea calm, with a light southerly to south-westerly breeze of 1-5 koots (2-0 kph), The animal was moving slowly towards deeper water, i a south-south-casterly direction. At times it swam just — -_ below the surface with its mouth open, [twas nol possible to tell if the animal was teeding and no obvious signs of plankton swarms were visible ta the Observer Several low, thin blows were seen, The animal’s colour was dark grey on the back wad Tight grey underneath, The features which sug- gested thal this was a C marginaia were: 1) lighter patches between the mouth ind the Hippers. 2) whitish baleen plates which darkened towards the outer edges, and 3) a small, curved dorsal fin about -/) of the way along the back However, without observing the bowed jaw-line, the identi ficahon as Coren could jot be vonsidered confirmed since minke whites (Baldenaprent aeureraxtrai) share several of the above-mentioned features. Estitated: bey length of the animal ROM. observed was 45-40 mi. sue. gesting thal if was a post-weaning juvenile’, Nine stinunings of Co aaginata have been recorded i the Port Lineoln area from before 1948 to 1993 (Table 1) and, as suggested im 1964 by Hale! many more have prob ably aecurced without being reported. ALL the reportud strandings have been from Port Lincoln Proper and in, or al the entrance to. Spaulding Cove (Fig. 4). Both are shallow, scugriss-coyvered, sandy/mudllat bays with large (dal movements, All strandings involved single animals. abhough i some cuxes There may have heen a connection between certain events occurring within a short time ol cuch other, For example, u 3.38 fa juvenile was found recently dead on 2 February 1989 in Spalding Cove nd on | March 1989 a decomposed adult (unknown sex) was found on Bigkers Islind at (he entrance (othe cove (strand ing nos 6, 7. Fig. 4), On 6 April 1993 iin adult deme, with no evidence ol lactation, washed up tear on the south side of Port Lincaln Proper and six days liter an extremely emia cited 3.15 m juvenile stranded alive in Spalding Cove (stranding nos X, 9 Fig. 4+). Hts possible that both cases involved mother/ealf pairs, OF the five adults that have stranded, three have been fenmiles and two of unknown sex. Vig. 4 Juvenile Caperee inetrginata in Spalding Cove. South Australia, 4/5 January 1996, The line drawing helps to dis tinguish the bowed jawline. which is slightly distorted by a wave. The arrow marks a lateral colour pattern of white just behind the head and a-dark patch posterior to: this, Photos J. Dutton, — : ,y Pl dustim H pe | | Ane M 4 b , | va - beh " oe DO .t ) Fs ~ | ales 3 a F a Le } , f ertrt ESANHREN Pa a, ro WHORE ® g mss . OO _ a f ~ f ~~ } = oe fig. 4 - Map of Port Lincoln region showing positions of sightings (squares) und strandings (circles) of Caperce mergindta, Numbers reler to strandings in Tuble | and Sightings in Table 2. These stranding results suggest (hal (he region is Frequently used by females und calves. although such iformation can be misleading if there is. as in this ease, much human aetiv- iy inthe region and fherefore possibility of discovery CUPCUISSUS Reported sightings of ©. werginata, summarised in Table 2. are hor eoninion tn the Australasian region. To: our knowledge, the Spalding Cove sighting of Jumuary 1996 is the first time a cow and calf have been sighted and pho- lographed anywhere ip the seutherm hensphere. Nat included in Table 2 are two unconfirmed records found in the Australian Native Conservation Agency sightings data hase. one of three animals off Montague Ishind, NSW in June [992 (record no. 876) and the other from new Bernier Istand. WA in dune 1993 (io 1558). Since there was no sup- porting deseription ol the aninals to allow positive identiti- eulion and © angie can be confused with the minke Tanbh b st while, B aentorasirata, the records have not been included here. Many of the sightings listed in Tuble 2 and several Iran South Africa! were made inshore, suggesting thal ©) war ginara inhabits coastal waters. at least for some part of its life or annual eyele. Unpublished stranding data show that dependent young und recently weaned juyeniles (3.0-3,3 m) are more common ilong the central South Australian and westem Victormn coast (Remper vapubo, Buur of the sitht- ings listed in Table 2 involved dependent young or animals that, from ther sive. would appear lo tive been recently weaned. Three were from the Port Lincoln aren and ane wis from Portland tn western Vietoria. Larre. proteetend! bays with shallow, sandy bottoms and extensive seagrass beds may be importunt calving and weaning areus far CL ma vim. Some exanples are Porthind) Bay. Nepean Bay and (he Boston Bay region. Several strandings ol C2 ateargimater have been recorded in the Nepean Bay arcu! The sighting made in Cockburn Sound in }990- by B, and D, Parker (B. Parker pers. comm.) merits a specnl nove because ibrecords some holable behaviour as well as prob- able feeding by the 5-m animal they observed over a period of about one hour, The aninal was swimming very fas, Joaving oowake of water, and nodding its head poticeubly, The whale came to investigite the em bout, scraping itsell the first Gime aust the battond of the beat The second lime if approuched (he bout very quickly, almost in at charge. lifted the boat out at the water aod ainose caused The vecupants ta he thrown out! The authors wish ty thank B, Parker and D, Coughran tor supplying information on the sucht mm Cockburn Sound gad the Anstralian Nature Conservation Ageney for searely ing their records for Capered sightings, ©, Kemper thanks all those who have provided information on pyemy right whale strandings and sightings records, especially South Australian National Parks and Wildlife officers and Department ol Primary Industries South Australia (Fisheries) officers, J. Thurmer suggested and prepured the line drawing for Mig. 3, Revordy of Caperea mannake strandings iy tle Part Lincoln area. Stranding number (Fig, Ref.) croasy-metched lo Kino 4, “estimered length based on skeletal measurements. Museum no, M = specimen in South Australian Museu, Ss = he specimen in South Austeudlian Muse. Date Location ree Sex ore <1948 SW Port Lincoln \ 3738 Proper 26121955 Tulku ? M 3.05 1681960 Tulku 3 r dull 4.1ON4 Spalding Cove 4 - 6.05" IS.S.1985 Port Lincoln Proper 3 Ib 6,20 2.2 1089 Spalding Caye 6 M 3,38 1.3, 1989 Vickers Iskind North = 7 - 3,31* 64, 1993 Thin SE Horse Rock 8 F 6.08 42k SE Stamford Hill YA pous ¥ 12.4,1992 y MAIS Miisquir Comments ho, M5753 Mol It) seen ulive before stranding 50009 decomposed, not collected MI4580 decomposed M14465 washed up deud MI5024+ very fresh M15374 —syery decomposed M1I7362 washed up dead S0085 alive. very emaciated. returned to sea 82 TABLE 2. Sightings, including captures hut not sightings inumediately followed by strandings, of Caperea marginata in the Australasian region. Latitude and longinide given in dearees and minutes. Rel. is reliablitiy of tdentification (1 = certain, 2 = probable, 3 = doubtful). A =udult, S = subudult, J = juvenile. Date Location Lat. ! Habitat No. Rel. Size Comments Rel, Long. 1.1874 north end 46 508 - - \ S captured amongst 8 Stewart 1, NZ 168 00 B black fish , 911959 Bruny Island. Tas 43 178 bay, in - | S killed by fishers 9 147 18 EF 2-3m water 7.1960 Port Lincoln, SA - protected bay 2 2 A.J ne description of LO animals 4.1980 ~ 50 om SE 38 205 open oceun 5 | observed from I Cape Howe. NSW 150 20 E olf shell ship 4.) 985 Svela Seamount, 43508 open ocean, many 2/3 - feeding. no 12 120 nm SE Tas 15022 BE over sea mount description of animals 28.11.1986. Portland, Vie. 38218 sahdy, 1 | S shipping harbour 13 5.2, 1987 141 36E — protected bay possibly feeding 1.1489 Spalding Cove. 34475 shallow, 2 e A.J Vig. 4 ref. no, 2 this SA 13558 FE protected hay. stucly sengerasses 19.10.1990 Cockburn Sound, = 32 105 offshore | A photos surgest B. Parker WA IIS44E feeding behaviour — pers. comm, 26.11.1992 420 nm S Cape 41375 pen sea ~80) | A three groups near 6 Leeuwin WA 11S 38 E euch other 4.1.1996 Spalding Cove, 34478 shallow. 2 | A.J stayed in area at this SA 13558 E — protected bay. least two days, study SCUTUASses Fig. ref. no. 3 77.1996 Port Lincoln, SA 34.44 8 edge of 1 2 J Fig. 4 rel. no. 4 this 13600E Spencer Gulf study References ‘Kemper, C. M. & Ling, J. K. (1991) Trans. R. Soe. 3. ‘Kemper, C. M. /hid.accepted), Aust. 115, 37-S2. ‘Hector, J. (1875) Trans. Proce. N. %. Inst. 7, 251-265, Pavey, C. R. (1992) Aust. Mammal, 15, 1-6. CCiile: BROT Ry-Phe, Pais ane * $2,780. ‘Davies, J. L. & Guiler, E. R. (1957) Proc, Zool. Sov i tuiler, E.R. (1978) Pap, Proc. R. Soc. Tasm, 112. TSi- Lond, 129, 579-589. 213. ‘Ross, G. J. Bo. Best, PB. & Donnelly, B. G. (1975) J “Hale, H. M. (64) Ree. S.A. Mus. 14, 334-419. Fish. Res. Board Can, 32, 1005-1017. "'Cawthorne, M. W. (1981) Rep. Int. Whal. Conn. 31, “Sekiguchi, K., Best, P. B. & Bozena, Z. K. (1992) Mar, SC/32/Progress Report N, Z.. 201, Mam, Sci. 8. 288-293. ‘ . ' ; so a oy ‘Matsuoka, K., Fujise, Y. & Pastene L. (1996) (bid, 12, Blaber, 8. J. M. (1986) Emu 86, 239-24, 594-597. “Arnold, A. (1987) Aust. Nal. Hist. 22. 266-270. CATHERINE KEMPER, South Australian Museum, North Terrace Adelaide S. Aust. 5000, JOHN DUTTON, 23 Mitcham Aye. Lower Mitcham S. Aust. 5062, BRIAN FOSTER, 51 Britainnia Road Nairne 8, Aust. 5252 and ROBERT McGUIRE, 83 Veran Terrace Port Lincoln S. Aust. 5606. VOL. 121, PARTS 3 & 4 28 NOVEM BER, 1997 Transactions of the Royal Society of South Australia Incorporated Contents. Brief Commu. Bourman, R. P., Martinaitis, P., Prescott, J. R. & Belperio, A. P. The age of the Pooraka formation and its implications, with some preliminary results from luminescence dating - - - Vaucher, C. & Beveridge, I. New species of Potorolepis Spasskii (Cestoda : Hymenolepididae) parasitic in pag hat soe from New Guinea - - - - - Littlejohn, M. J. & Wright, J. R. Structure of the acoustic Rane of C rinia glauerti (Anura : Myobatrachidae) from south-western Australia, and comparison with those of C. ste. from South Australia - - Watson, G. F. & Gerhardt, H. C. The breading Rivioes anti qavetiscenent cat of Litoria splendida Tyler, Davies & Martin - = Barnett, E. J., Harvey, N., Belperio, A. P. & Bourman, R. P. Sea- tees indicators from a Holocene, tide-dominated coastal succession, Port Pirie, South Australia - - = - Gullan, P. J., Cranston, P. S. & Cook, L. G. The response of gall- ee scale insects (Hemiptera : Eriococcidae: Apiomorpha Riibsaamen) to the fire history of mallee eucalypts in Danggali Conservation Park, South Australia - - - Olsen, A. M. An intensive monitoring study of two wetlands of the River Murray in South Australia; 2 eee a and cyanobacteria concentrations - —- - Kolesik, P., Whittemore, R. & Stace, H. M. Asphondylia pes idis, anew species of Cecidomyiidae (Diptera) inducing fruit galls on Anthocercis littorea (Solanaceae) in Western Australia - - nications: Wallman, J. F. First record of the Oriental Latrine fly, Chrysomya megacephala (Fabricius) (Diptera : Calliphoridae), from South Australia - - - - - - - - - Smales, L. R. The status of Cyclostrongylus medioannulatus Johnston & Mawson, 1940 - - - - - - - - - PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 83 95 137 147 157 165 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 121, PART 3 THE AGE OF THE POORAKA FORMATION AND ITS IMPLICATIONS, WITH SOME PRELIMINARY RESULTS FROM LUMINESCENCE DATING By R. P. BOURMAN*, P. MARTINAITIST, J. R. PRESCOTT? & A. P. BELPERIO£ Summary Bourman, R. P., Martinaitis, P., Prescott, J. R., & Belperio, A. P. (1997) The age of the Pooraka Formation and its implications, with some preliminary results from luminescence dating. Trans. R. Soc. S$. Aust. 121(3), 83-94, 28 November, 1997. Stratigraphic relationships, supported by luminescence dating, suggest that the Pooraka Formation spans a far greater time interval than previously recognised on the basis of radiocarbon dating and stratigraphic analysis of discrete sedimentary sections. It extends back as far as the Last Interglacial. Re-evaluation of the radiocarbon ages that indicate an interstadial age (1.e. Oxygen Isotope Stage 3; 45 to 30 ka BP) for the sediments is required. Alternatively, a considerable time interval for deposition of the Pooraka Formation would necessitate that the unit be diachronous across the landscape. An age extending back to the Last Interglacial (Oxygen Isotope Substage 5c; c. 125 ka BP) would provide the appropriate palaeo-climates and palaeo- environments for fluyial sedimentation. The revised age has implications for landscape evolution, archaeological and palaeomagnetic prospecting as well as the antiquity of the Diprotodon in the Adelaide area. Key Words: Pooraka Formation, Pleistocene stratigraphy, Last Interglacial, luminescence dating. Fresnel if de Raval Saetery ol S dase (1997), PEGA) BA OL THE AGE Ob THE POORAKA FORMATION AND ITS IMPLICATIONS, WITH SOME PRELIMINARY RESULTS FROM LUMINESCENCE DATING by RL PR BourmMan , P. Martinarns', J. R, Prescorr & A. PB Brrperny! Summary Bouin, ROP. MARINAS, By Prescorl J, Ry & BbipeRion A, P (L997) The age of the Poorake Pormanon and its plications, with some preliniinaey resulis from luminescence dating, Trai AL Sey Ao Att D2ECA), 83 4, 28 November, 1997, SmMighiphic relianships, supporwed hy liminescenve dling. sia@@est thatthe Poorake Rocnation spas alae vrewer (ime interval than previously recognised on the basis of ridiocarboo dating ant stratigraphic analysis af discrete sedentary sections. extends back is fir as the Last Tnterghieiil Re-evaluarion of the niuhecar bor ues That idicute an titerstudhtl age (ie, Oxygen (sotope Stige 3:45 10 30 ka BR) forthe sediments is required, Miernittively, a considerable time interval for deposition af the Pooraka Formation would necessitate that the wit he ditehronois weross the tindseape An age extending biek to the Last Niterglacial (Oxygen fsorope Substige 50.6, 125 ka BP) would provide the appropriate palieo-climates and paliea-enviconiients for (luyisl sedimentation, The revised age has impligaions for kindseape evolution, aehiwolowical and palaeomugnetic prospectiog as Wella the antiquity of thy Diprajeden inthe Adelitide area. Ky Wokbs: Poorake Portndtion, Plestucene stratiraphy, Last Witerglieial, Wimineseence dutine Introduction Linge areas of the Adelie Plans are underlain hy the Pooraka Pormation (Firman (67, (969, Callen eral, M95: Sheard & Bowman 1996) w reddish: brown coloured Pleistocene ahivial deposi with weakly developed calcareous pedogenic horivens that underlies river terraces and alluvial fans, Plies Unit is also Widespread beyond the Adgliide Plains. extending On to the Fleurieu Peninsula ane inte the mid-narth wl the state, where it Minky the Plinders whl Giwler Ranges. The red-coloured sediments that comprise the Widespread Pooraka Pormation have been aseribed different names by workers over time and indifferent areas, bor exaniple, they were originally referred to as the “nanimaliterots drift by Tate (1879) becuse skelotul remains of the extinet. giant marsupial, Vipratoden apitiin, were recoyered from thenr in areas Cothe west al the city of Adelaide, Ward (1966) relerred) to the sediments as the Christies Beneh Formation on the Noarlunga and Willunga sub- busing, Twrlule (1968) tumed them the Klemzig Sand during his investigation of the terraces of the Sehool or Bnviromnenal and Keeredion Mawar, (iene vl Eicineeriiye aid Lanwroniient: Ubpwersity ot Southey Avatedics Warrenty Ral Tlie bawels So Apst, S005 Dypatareat ed Pliysies vil Mathematical Miysies. Pl biivrgity cb Nolo lide nist, SOS Vonvierly Mines iid Eneniy doxgurees Soh Australian PO) Hos 19) Lastwrmd 5, Agst 3004, Cuerenthy Minti Gold, ta Cohalatore Ne Peron S. Atist Sue 4 Bev ADA RP OPYOU) Ladforn Siadies near Vietor Phorbour Bay (Homes thesis. The Cntogrsaey ou Adehttules ciapiity River ‘Torrens, and Bourn (1968, 19694) referred to them as the Adie Clay where they Mank the Rivers Hindiaesh and Taman in the Vietor Harbor aren (Localities shown on Pig. 1), The red-coloured Alu bears consistent stratigraphic relutignships food younger, greyeblivk alluviuin | Waldeita Formation of Ward (1966), Wilkerville Sand of ‘Twidiile (1968) und the Breckan Sand of Boutros (1968) ) whieh forms Tower terraees and Moodplains set within valleys carved out ol the red allivinina of the Pooraka Formation, Bstuuine shells collected from within the Walder Pormation in the lower reaches of the Onkaparinga River (Bourman 1972) retumed a ridigearbon uve of 4.580 + 160 veurs BAP. (Bourman 1979). During this Middle Holocene time the lower, dearecoustil reaches OF any streams were shillow, shellered estuaries as revealed by the presence of fossiiferous narine deposits at depth up valley, This interpretition ts Supported by evidence Worn a locality seyerul Kilometres rom the coast on the lower Onkaparinga River where ain uboriginal kitehen midden Containing estuarine shells citec at S820 + 90 yours B.PL(N.B, ‘Tindile pers, conom. in Twidale ey a 1967) is sired ona well dramed sand dline site at 20 mast ane adjaeentta the former, more eNLOMAIVE SULLY, Ages useribed io the Pooraka Formation and its equivalents The Pootuka Formaion tas generally been ascribed to the Late Pleistocene, with nest nuroerical ages, based on radioearben dating. falling within the ange nf S0.000-20,000 veurs (see review K+ R. BP. BOURMAN, P. MARTINAITIS, J. R. PRESCOTT & A. P. BELPERIO 9 AUSTRALIA SOUTH AUSTRALIA oe AB ADELAIDE \. d Beach /* 7 Hill Pym. “Ue p> . Weymouth Hill f Cape Jervis Fleuriel ( Waitp Bea tei “~ Tunkalllla mi Beach We RIVE Hallett Cove Embayrnent / ft ln Selly Sellicks \. o~ Myponga gay XN gawler River ee a / / f Oe pata SLA a Mt- Pleasant A: 3 . v rT 542 MAL Gawler : / ~ ADELAIDE |_| / Mt, 726 Loatty a £ 7 os rf vf Noarlunga Embayment An nse ie fe: Rone en arind® Willunga ~ Mt a oN Currency C4 Ty See SS vA Hindmarsh 7 Is. > ae eB i Victor } & Harbor inga” ch Pd Encounter Bay Study Sites OS ory Cc “ River Torrens, Walkerville Bie reek, Pooraka Ki Hindmarsh River., Victor Harbor 20 30 Scale in kilometres Fig. 1, Location map of sites. AGE OF POORAKA FORMATION 85 in Callen ev af. 1995). There has not been universal agreement on to which part of the Late Pleistocene the sediments should be allocated. Twidale (1968) assigned the Klemzig Sand to the Late Pleistocene and demonstrated that it must be older than 6,350 years BP. The distal end of the femur of a giant extinct marsupial, recovered from 3.6 m below the surface of a fllstrath terrace cut into the Adare Clay, in the Hindmarsh River, was dated at approximately 12,600 years BP (Gak - 2356) (Gill & Bourman 1972). Firman (1969) and Daily ef al. (1976) also considered the Pooraka Formation to be of Late Pleistocene age, but younger than the Anadara- bearing Glanville Formation, which is now widely accepted to be of last interglacial age (¢. 125,000 yr BP; Murray-Wallace ef al. 1988; Murray-Wallace & Belperio 1991; Murray-Wallace 1995; Belperio ef al. 1995), Confident separation between Late Pleistocene Pooraka Formation and earlier Pleistocene alluvial sediments is casily achieved in the coastal zone where they are separated by coastal facies, Inland, on the Adelaide Plains, the Pooraka Formation is readily distinguished from underlying Tertiary sands, the Keswick Clay and the Hindmarsh Clay. The Pooraka Formation is only weakly consolidated, carbonate impregnated and mottled in comparison with the underlying units (Sheard & Bowman 1996). Ward (1966) assigned the Christies Beach Formation to the Last Interglacial as he considered that the surface on it was graded to the last interglacial shoreline (his Epimonasterian high sea level) at approximately + 3 m above present sea level. However, at that time the Last Interglacial was thought to be considerably younger than the present 125,000 years BP. From the Dry Creek alluvial fan, Willams (1969) described reddish-brown clay overlying older grey- green and red mottled clay, now known to be the Keswick Clay (Sheard & Bowman 1987a,b; M. Sheard pers. comm. 1997), Williams (1969) also noted a calcareous red-brown earth developed within the sediments containing nodules and cylindroids of pedogenic calcium carbonate, A radiocarbon age of 34,600 + 2700 years BP on carbonised wood from sand 3 m below the land surface was obtained by Williams (1969), The carbonised wood was regarded as detrital in origin and thus was regarded as a rehable representation of the time of deposition. However, if the carbon were detrital, its age should predate the lime of sedimentation, which would mike the ''C date somewhat older than the time of deposition, The date was taken to indicate a last glacial (Wiirm) age for the sediments. Further radiocarbon dates supporting a last glacial (Wiirm) Guppy, DA, (1943) Geological reconnaissance of part of the Hundreds of Encounter Bay and Goolwa, BSe (Hons) thesis, The University of Adelaide (unpub.). age were derived from a study of alluvial fans on the western side of the Flinders Ranges (Williams 1973), Carbonised detrital wood recovered from depths of 8-9 m and 15 m within the Pooraka Formation provided radiocarbon ages of 33,270 (+ 2130 - 1680 years) BP and > 37,000 years BP respectively. Stratigraphic observations Critical evidence concerning the age of the Pooraka Formation occurs at Victor Harbor. Here the relationships between the last interglacial shoreline and the Pooraka equivalent unit, the Adare Clay, suggest that the unit is much older than 50,000 years. Bourman (1968, 1969!) established that red-coloured alluvium forms fill-top terraces along the Inman and Hindmarsh Rivers and grades to a shoreline at ¢. + 6 m ubove present sea level. The age of the shoreline is considered to be crucial with respect to the age of the terraces and the sediments which underhe them. Twelve species of shells have been identitied from this shoreline deposit (Guppy 19432) and it is significant that they contain the sub-fossil Anadara trapesia. Initially, Bourman (1968, 1969!) followed Sprigg (1952) and assigned these shells to the Holocene, Subsequently, the shells were radiocarbon dated returning ages of 33,170 + 3,180 -2,270 years BP (Gak-5561) and >30,320 years BP (GaK-6099), Although the above dates are compatible with those of Williams (1969, 1973) they are questionable because the period around 30,000 years BP was a time of low sea level. Furthermore, it is now generally accepted that materials whose true ages are beyond the range of radiocarbon dating (> 40 ka for most laboratories) may yield younger apparent ages, due to the diagenic incorporation of low levels of radiocarbon from modern activity. Thus, materials with an infinite age by radiocarbon dating techniques may yield an apparent age of 37 ka due to the incorporation of 1% MC with aw modern activity (Gupta & Polach 1985). Gill (1974) checked radiocarbon dates of this age against other dating techniques and concluded that radiocarbon dating may be reliable for young materials but older materials may return ages that are far loo young. Similar conclusions were reported by Bowman & Harvey (1983) and Belperio ef al. (1984). Not only do Anadera shells occur at the + 6 m shoreline at the coast at Victor Harbor, but extremely large Anadara shells were recovered trom a sewer trench c. 1.6 km upstream at a depth of 4m below the surface. within the Pooraka Formation equivalent unit and at the same absolute elevation of 6 mas al the shoreline (Fig. 2). A drilling programme (CSIRO Soils Division) further revealed the intimate association of Anadara shells with the Pooraka Formation equivalent unit, demonstrating that here the Anadara, last interglacial deposits (Glanville RG RP BOURMAK. FP MARTINATTIS. J. R. PRESCOTT & 4, Py BELPLERIO Formation) and the Pooruka Formation are a last interhienal age, intercalated coastal and terrestrial equivalents. The Anadara shells have been dated both by Uranium. Methods Thorium techniques (100.0006 (50,0000 yeuns BP) and by uming-acid racemisation studies (Kimber & The stratigraphic rehutionships between lust Milnes 1984). whieh provided results Consistent with interglacial molluses and the Pooraka Portion _ Anadara Shells («Glanville Formation} 15 WwW = 7 — ~ a S - > “ Y— > wt =~ = Ee j Cc o+-———— ---— —— eS ——— = Soa Level 3 . Pooraka Farmation —— ee ii 10 —— —— yo ~~—S—CS~;S Pormian Sediments 1S a Xt —————— —— a2 17] VOODIN Sen fio. 2. Sketch section showing the interfingeriny relationships af the Gkiille Formation aad the Pooruka Pornnaion i the lawer Hindmursh River at Victor Harbor. Fi 7 PHotosaph of 6m high ever bluff cut in Pooraka Pormution sediments downstreu of the Bridgeway Horelon Dry Creek, Powruke, Masinuin seetion exposed « 7m Red brown earth with assechkted calcium carbonate zones in upper purl of selon. AL the busc al the secvion The Pooraka Formation rests unconlormibly on ai older Plesitacene Unit tat Chisely resembles the Tartiipa Formation af Wank ted; bat whieh Sheard & Bown CM 87ah) consider is mure hkebs Ibe the Keswick Chay, AGE OF POORAKA FORMATION 87 equivalent, near the coast, at Victor Harbor provided strong indications that the Pooraka Formation might extend back to the time of the Last Interglacial. In order to test this hypothesis, and to determine whether the inland, terrestrial Pooraka Formation sediments were of an equivalent age, an attempt was made to obtain a numerical age for this formation using the technique of luminescence dating. A key site chosen for sample collection for luminescence dating was the Dry Creek (Pooraka) locality where Williams (1969) collected samples of detrital carbonised wood and carbonate for radiocarbon dating (Fig. 3). Unfortunately, the steep river bluff al this site has now been contoured and landscaped so that it was not possible to sample from exactly the same site as Williams (1969). A second site on the River Torrens at Walkerville, where a thick Pooraka section had previously been exposed, was selected as a subsidiary luminescence dating sampling site. As both sites haye suffered from human modification and landscaping of the former eroding river bluffs it was decided to collect samples for luminescence dating by drilling using an auger drill with an internal push cylinder. This method allowed sampling depths to be determined and the sample to be collected without exposure to light. Two holes were drilled at Pooraka and one at Walkerville, Samples were recovered at depths of 3,5 m, 4.5 m, 4.8 mand 7.5 m (for Pooraka) and 4.5 m, 5.5 m, 6.0 m, 7.5 m, 9.0 m and 9.3m (for Walkerville). The drilling site at Pooraka was located 7.74 m above the base of Dry Creek and the second site was 11.68 m above the River Torrens level at Walkerville. The same holes were used for both sample collection and scintillometry for dose rate determination, A summary of data collected for the Pooraka and Walkerville samples is shown in Table 1. Sample PKIS from a depth of 3.5 m is close to the level from which Williams (1969) collected detrital carbon for 4C dating. Luminescence dating methods Three methods for luminescence dating (LD) of the sediments were used: selective bleach thermoluminescence (TL) of coarse-grain quartz (Prescott & Mojarrabi 1993) and green light stimulated-luminescence (GLSL) of both coarse grain quartz and of fine grain separates (Aitken 1994; Duller 1996). In the dating of sediments it is assumed that exposure to sunlight is the agency that resets the luminescence clock and that the sample has been exposed to sunlight for a sufficiently long lime thal the stored energy giving rise to the luminescence has been reduced to a low, near zero, level. This is a reasonable assumption in open sites exposed to strong sunlight, but this may not be true where there is the possibility that the material was deposited by, or under, water, as in the present sites, or ina generally colluvial environment. Both the TL selective bleach and GLSL methods seek to overcome this uncertainty by making use of an easily-bleached component which can be reset to zero by short exposure to sunlight. It is assumed that this component has, in fact, been reset. Details of the methodology are presented in the appendix. Dating results Pilot TL runs were carried out on coarse grain quartz from all samples except PK2S/4.8, WV1S/7.5 and WYV1S/9.3. Such runs are designed to assess whether the sample is likely to be datable and, if so, TABLE Lo Summary of collected data for Pooraka and Walkerville samples. SAMPLE PPMU PPM Th PPM U PPM Th = =PPMU Sct. PPM Th Sct. =“%@K XRS TK Set GUO DNA DNA PKIS/3,5 1540.6 7042 1484011 7.23405 1.744017 7.124029 1.08+0.03 0864002 6840.07 PR2S/4.5 Ll+03 7841 1264011 731404 1.754017 7.684029 0.8940.03 0.82 +0.02 3.7 +0.06 PK2S/4,8 Ll+05 6341 E1O+010 6.24404 1.344016 6.76+0.28 0.894+0.03 0.9140.03 4.4+0.07 PKIS/7,5 12406 7142 1224010 691404 1.744017 9.694032 1.204004 1134003 1240.07 WVISM.S LR 403 1241 LSP 4012 17405 1.8940.24 1254044 1854006 1.604004 8.8 40.07 WVISIS.S 16406 1442 1934013 126406 2.334024 1054042 L8P+0.05 1324004 7.5 40.07 WVIS/,0 19405 Wel 1864012 103406 2474025 1214048 15940.05 1544004 8.3 +0.07 WVIS/TS 17405 142 1664012 1840.6 2.664026 IL1+047 1.71 40.05 149+0.04 15.7 + 0.07 WVISMO T8405 IS4t LI7+012 134406 2.954029 1344050 2.194007 171 +0.04 18.0 40.07 WVIS9.3 12405 1542 L88 +013 13.0406 2.954029 1344050 1.984006 1.714004 16.4 +0.07 The first two columns are the results derived from Thick Source Alpha Counting. A DNA was done for uranium only: ppm Th (DNA) were obtained by combining the count-rate from thick source alpha counting and the uranium concentration from DNA. ppin U Sct, ppm Th Sct & %K Sct were derived from the on-site gamma ray scintillometry data. 88 R. P, BOURMAN. P. MARTINAITIS, J. R. PRESCOTT & A. P, BELPERIO to give a limited-accuracy estimate of the acquired luminescence and the sensitivity to radiation, and hence plan the schedule for a complete dating procedure. A pilot run consists of eight dises: half of these are bleached, after which two discs euch of bleached and unbleached are given a radiation dose of 60 Gy. Por all samples except PK1IS/3.5. the shallowest of the Pooraka samples, the TL was clearly saturated and no further work on them was justified. Although PKIS/3.S was approaching saturation, a full dating procedure was carned out tor both TL and GLSL. Such procedures give the Equivalent Dose, D.. a measure in grays of the energy absorbed by the 3.0 4 9 Counts x10 1.0 100 200 300 400 500 600 Temperature (9C) Fig. 4a. Glow curves for sample PK 1LS/3.5, The figures next to each curve indicate the dose in Gy, Counts x10% 0 10 20 30 40 50 60. 70 80 Shine Time (in seconds) Vig. 4b. GLSL. shine-down curves for sample PKIS/3.5. The figures next to each curve indicate the dose in Gy, sample from radiation in the environment since il was last reset to zero. The age of the sample is found by dividing the equivalent dose by the dose rate in gray per kiloyear (Gy ka!). Equivalent doses, dose rates and ages TL glow curves ure shown In Figure 4a, shine- down curves in Figure 4b and corresponding dose curyes are shown in Figures Sa, b respectively, None of the curves is sealed. [It is evident that the dose curve of quartz (Fig. 5a) is close to saturation but that the growth curve for GLSL on fine grains (Fig. Sb) has a different shape and the curve continues to rise quasi-linearly for high doses. This is because the Counts «10° 0 0.0 200 400 600 800 Dose Gy Fig, Sa. Th growth curve for sample PKAS/3.9 for the [0 interval at 305° C. For Figures Sa and b, the curves are fitted by the “Australian slide” method: the (natural + dose) points are shown by crosses: the (bleached + dose) points are shown by cireles. There is an apparent sensitivity change for the Tl bleached curve bul the scaling factor does nat differ significantly from unity. Counts x1o* 9 200 400 600 800 Dose Gy Fiz. 5b. GLSL growth curve for sample PRIS/3.5 integrated over the first LOO s_ AGE OF POORAKA FORMATION Mel Panay 2, Bawls doses, dese ratey anlages far sample PR LS/35. COARSF GRAINED QUARTZ FINE GRAINS selective bleaeh Th GLSL GLSL IRSI 1), Gy 240+ 58 JK & 44 24445 > 150 dose mite TSAC/DNALRRS SCINT TSAC/DNAIARS SCINT Gy ku (90 #1, LG 1b 2544 (1,12 2542011 Weighted mec LUT © 1.03 2.54 = 1).08 ave kit 126 + 29 LOS + 22 llh+6 > 10 undifferentiated fine grafts consist of a mixture of minerals in which quartz and feldspar ure donmnant, Thus although the quite component saturates, the feldspar component does not, Equivalent dose values from all three methods appear in the first tine of Vuble 2. Ih will be noted that the two methods of finding D, give different values; this is characteristic ol the methods, Two distinet methods for finding ihe dose rules were used (described i the appendix). the aim being iO get two independent values for euch sample, to inprove the statistical precision and ws cheek on (he presence of radioactive disequilibrium (Prescott & Hutton 1995). "The values obtained are ineluded in Table 2. As with D., dose rates differ for course: and fine grains. The values are in excellent mufial agreement and this shows that radiouctive disequilibrium is absent, The weighted mean dose rates are shown on ling 3: line 4 shows the ages derived from the D. using the age equation from the uppendix. The weighted mean, 116+ 6 ku is dominated by the GLSL determination for Hie grains. Comment Allowing for stanstical Mtting uncertainties. all three ares are in good agreement. Bearing in mind (hal they are bused on different physical processes, allo? which assume that the luminescence sign was set to vero in the past it shows that this wis very likely the case and that the age being determined is indeed the tune of deposition, Th it is not so. the uppurent ages will be too large. As already inentioned, the dose growth curve of Figure Sa, which is lor quartz TL. shows: that the luminescence is close fo saturation. For this reason the estimate Of relative Gneertainty given by the ialysis is large. This is also true for GLSL, of quarts. For GLSL of fine grains, on the other hand, the presence of the Ieldspar component allows Dy to be found with significantly better precision, The best estimate of the the of deposition of sumple PKIS/3.5 is the weighted mewn age, 116 4 6 ka. In facet. this is dominated by the GLSL determination on fine grams. [is probably best to reaard the quartz determinations by selective bleach und GLSE, ws being supporuve of the GLSL fine grain age. Al the two standard deviation level the age exvecds LO ka, Consistent with these numbers. the saturation of the luminescence of all the other samples shows that they we as old or older than (his, Therefore, although only one sample has yielded a numerical age, there js enough evidence ly establish a lower limit to the age of the formation and itis in support Of the geomorphological and sedi- menlolugical evidence, In addition to the three methods alreiuly desembed, wilta-red stimulated luminescence (IRSL) of fine wrains (Wintle 1994) was tned. IRSL uses only feldspars and is less susceptible to saturation, although if may be subject to Tongeterm fading. Although (he growth curve was similiar im shiape ta the GLSL curve of Figure Sa, the stiustical fitting procedures did net salisly our eritena Por an accepluble D.. beyond showing that it was greater than LOO ka. Whether this is due to the sample or the methodology remains to he determined. However, the result is sufficiently cocouraging to suggest that it may be possible to find ages from the lower levels i Pooraka und for Walkerville where quitrtz methods were unsuccesstul: Discussion The location of the date derived from the upper section of the Pooraka Formation at a depth of 3.5 m below {he surface 1s i complete aecordinee with the Ot) RT ROURMAN. POMARTINAPTIS. LR. PRESCOTT & A. PL REL PERIO numerical age oblaned oh 176 + 6 ka avilh the Last Inlerglacal ranging from approximately (32-108 ke tsee Chen ero. Yt; Lumbeck & Nukuda 1902) Zhu et dil (99S: Sarling er al 1995) Ejsentner ep af 96). The GLSL technique olfers the possibility: of vbliining & Turninescence date for lower seetions al the Poorske Pormution thereby fucdiratiig the calculation of the rates of sedimentuion during the Taint Imerghiciah itmay be that sedimenhitton duping warmer, Weller, LIStTMberglaenu UimTes was quite rapid and this technique offers the opportunity to lest vis TV ‘These resulls Sugeest a far greater age range for the Poorgkii Formation thin has previotishy been recoemsed, In the past, the gee of the Poorakit formation fas heen poorly consiained we abou! 20- 50 ku BP pussibly heeause ol the now-known Hivikitions of padinearhon dating. Furthermore, 7 Sfould be poled ifjab other methods such as Th, GLASS, and IRS were not available when many vt the madiues bon dates were cured but. Many observutions and cores indicate that, an places, the Poorthie Farmition appedrs to be younet (hun ihe Just inberetacial Cilunyille Poroatien, especially in’ subsiling areas. For example. Ludhrook (1970) nthe Port Adelaide itrea described Pooraka Formation overlying calented Glanville Formation, whieh th Wut overlay Hindmarsh Clay. Relperin and Rive (WY89) shawed thal ol 62 cores from) the Gillman area. Eb record the sequence ul Holueene overlying calercted Glunville lormation Without Poorika Formation, and 5 record Halocem: overlying Pooritku whieh overlies calercted Glanville Vormiation, The calereted Surfaces itre inlerpreteel as pedogenic features developed during subuactial exposure and wuld have dormed prior to the deposition of the overlying sediments of the Pooraka Fornmdton, In contrast, four of the cores reeurd at sequence Of Holocene/Pooruka/Glanyville Porniation. withour 2 well-developed culerete oi the Glanville Formation, and one revords intermixcig of Pooruku und Glinyille. These Just cores miry ihdieate coeval Canville and Pooraka sedimentation or litle time Break between the (Wa, Extensive work by Sheard & Bowriun (1996), which involved drilling cores to depths of FO mr it 7H sites over the Adeliide Plains. intersected Pooruka Pormation in 80 cores. This work corroborites the Findings of Belperio & Rive (1989) for the voustal zone. More landward sites iniieated the sequences to be: Holoeene/Pooruka/Keswick Clay/Homdinarsh Clay and Holoverie/Poorakas Woon. Pi bat de Kose Pin Ply bid (18s) Appendix 3 Ceatechiea) Investaatiens lu Holiival Shores ti Banildlonstone Thomiboioh Giles Canaan) Werke dun Terry Wilark Bepiart ty Urhii Projects Aviheriiy at SA. Gunputts,) Keawiek Cluy/Tertiary sands orAdehddean rocks, At the MM) sites intersecting Pouraka Purnddon sediments. ‘Sheurd d& Brym (1996) rarely eXpefienecd diffloulties a adenufying Pooraks Formation from the overlying and underlying materiily (M Shearl pers. comm. 1997), Recent drilling work by Woodhead er al (Yas) for Baulderstone Hormibrook in the Haldfist Shares (Glenele Patawalonga Redevelopment urea) has demonstriee that depostis of Poorake Formation 2-4 om thick oeene well offshore frum the present eoust line. These deposits aire detavhei from more landward deposits either by vones Of fol-deposiuon or by carly Holocene erosion, Variuhle stratigraphic relationships are aipparent in this locality. Por exaniple. the tallow ing relariuinshipes Of Holovene/Pooruka/Glanvilles/Hindivirsh Chi. Holocene/Ghinvilley Hindinursh Chay ot Floloeenes Fulham Sand/Pooruka/Ghuryile/Minginarsh Clay! oveur within an S00 mM east-west section. Ie omuy be that the Pooruka Porniiion was Uepasiled dver i considerable period Of time This view 1 also supported by the oecurrence of palacosols within. Pooraka Pormation equivalent sediments is exposed it Sellicks Creck and Cobblers Creek. The luminescence data indicaie an ape approximately coincident with the Last Miler heh (125 ku BP), bul the resalts atany one place may be joflucneed by the terresirial/maripe interactions. For extmple. there is mut un achive Supply of alliwinin Othe coastline then marine deposition, exposure andl calerete formation will doittate, fh plices subsequently mintied by Pooraka or Holocene sediments. Th omay be fi only where there tsa sulfivient supply of terrestrial sediments to the coustline, sueh us where streams deboucl at the eoustine Th PelaGyely coystramed valleys (ee, Hindmarsh River at Vielar Harbor), that the Inrertinvering vhunacteristies af the two sediments ‘an be demonstrated. Et fsalso possible that Puorakat Formation sediments Jo not reueh the position of the Glanville Bormation uniibaflerit has been culereted, It should be noted that touch of the Pooraku Formation bas been deposited infind, well beyond the elevation and plinimetric position of the last inlerlaciel shoreline, so thar in these situations (here Ison clear siraneraphie rehkiionship between the two uns hnplicationy vf the Mraruka Loriiation exetidiin hack te the Last datercleavtel Wo the Trterpretation Uhl the Poorika Pormutien extends back to the Last Interelacnal is correct, wpe evaluation of rhe radiowarbon aves thar indicate glacial or (nterstadial ages for the sediments 14 required. Alternatively, the canflictuay results tiny suguest that there could be sediments, which AGE OF POOR AKA FORMATION 9] athoueh appearing similis Occupy a rune oF ages, Ati number of lowations, the Pouraka Formation his been noted to averlic the Glanville Bornution with « yariably developed calercle, Sugeesting surface exposure prior lo burial by younger Poortkit Fotmauon sediments, Elsewhere. such as in the Jower Hindnirsh Rivet. where there is uur interplay of coevail oustil anc terrestrial events. the strutipriphic relationships between We Pooraka and Glanville Formations may simply represent a hicies chunge and not ow geolagicul sucvession A considerable lime interval nay be requied for the deposition, OF the Poopake Pormition so that tts age conkl he diuchronie, with deposiion oecoriine during the rise in sea evel up to and beyond Lhat ef the hist imberchienl level, A lust intereletil awe privides appropriaie pulseu- chime and pulaeossea level conditions for exphiiniog the distribution oF the Pooruku Vormation, Close to the coast the unit rektes to a shoreline higher than the modern one, while inland the wetter. Woerelacial conditions would have hivourcel dwvradation of sediments w-opposed fo dpier ehicial conditions that would have (actlilaled dissecuen, This timing would alse ensure sufficient tine both for the build up of extensive deposits oF the Poerikit Portion aver large aveas ieluding much of the city of Adelaide. jind time during elicials and interstudials for erosion oF the Pooraka Parination to develop the extensive terruve system al the River Torrens (Twidile |968). A last intengiieial age for (he Pooraka Formation is al signiicunce Nrarchivalogicul prospecting. AL the Present Uine there ts debate concerning the untiyuiby of Humiins on the Australi continent. Substurrail chaes in Vesetition related to Aborginal burning pruchees faye been ilerpreted vis sesulling Yom human pmpach ural used bo infer the arrival of people in Australias lemme ics 140.000 yours (Rershaw eek WS. Kershuw 1993, 190d. 1995). These Claims have Beeb Questioned hy: varinus workers (eo Andersen Tad: Hope 1994; White 199d) Webb 19S). Tile Pouraka Formation is of bist tiberetactal He ny present ad prospecting opportunity to best i Aboriginal colonisadon had occurred in southern Auistalia prive to 125.000 years BP ducing the Gime ofthe penullanate ehiekil law sea level, A minot palacomugnetic event, the so-culled “Blake Event, his heen nle tied ro the northern hemisphere irr loess sequences (Llpneho ed af, LY95) “eapprosinnitely [20 ka. Given that the generat age of the Pooraky Fornvarion ts almost certainty of last iMerelienl uve, and that the upper part ob the formation has been useribed vu GEST aye of blo = 6 kit Uhere may be opportunities ta identity the Blake event in the southern hemisphere, A last Tnlerg ictal ave lor the Pooraka Formate has implications fev the ahquiyy ob the 2 peatertent in the Adelaide urea, Diprotodonlid remains. have been revurded well baek into Tertiary straui in the North Flinders-Cullabonna Phiins areas (Callen & Tedford 1976; Pledge & Tedford 1990; M. J. Sheard pers. comm. 1997.) but Hot se far, in the Adelaide vrea. Discoveries of Dipraloden remains have been mude i the Poorwku Formation (Twidale 1968) N, Pledge pers. comim, 1996) ab the Adelaide area und Winle the Diprarodon iray have predated and survived well beyond the age of this sedimentary unit, if was almost certainly roaming the swampy, iwprading Adelaide Phi some 125.000 years ago. The age has further implreations for landscape evolution as there appears ta have been a tajor erosional hiatus of some 120.000 years belween the deposition of the Pouraka Formation and the grey- black Wildeila Formation, which is oFimid-Holocene uve, There were ro sea levels hiuber than (he present in the iptervening period so that erosion would have dominated this interval of time. Alernitively, any sediments deposited during this (ime could: lve heen removed in bile stages of erosion. Canclusions The main conclusion from this work is that the Pooraka Formation must span a lar ereuter line period that previously recognised. probally extending as tar back as Lust Interglieiil dine te. 125 ku, Th vppedrs that the tectonic ane environmental setuag. whieb influences the supply of terrestrial Sediments to the coasting, sale sireme Importance in interpreting the stlatighaphie relavonships between the Pooruka Formation ane the Glunville Formation. Mueh mure luminescence CIN Work Is Lequiied fo coystrai dr document the ie ranges in diffgrent setlings ae. in the Wietor Lhurbor Setong where the bWwo qe jotermived. tir contest to the Port Adelide Setting where a tia]or calerety pulaeosol separctles thei, We mi expect luminescenve ating: techniques to document wes from list interelicnl tines. possibhy through te 30- SO aL uf the quitrty erains are sufficienily reset. This offers The possibility of providing ae pore reliible dating tool than radineurbon techniques. whieh Sill Stiwele do prowile mean dates pasty 4d) ka. The red-brewe alluvial sediments, referred ba is Pooraka Formation and equivalents, are eopisidered to extenu back toot least dhe Last Inleralocil of ¢, 125,000, vous BR This ts demonstrated by the Stratgfaphical relationships of the red-hrows sediments lo both younger and older sedinwits, the interdigitiing of dentonstrated last interghiectal miine depasits with then then grading tod + @ nM higher shoreline containing last interglacial marine ya R. PR BOURMAN, P, MARTINAITIS, J. R. PRESCOTT & A.) BRLPERIO fossils al Victor Farbor, and the fict that a higher sea level. together with associated warmer and wetter conditions would) favour ageradation. whereas colder, drier glacial conditions associated With lower seu levels would hive favoured erosion. AlLof these luclors Support a last interglacial aye. The luminescence data have demonstrated viability us un independent means of testing the hypothesis that the Pooruka Formation is of last interghieil age. Fluvial sediments present special problems for luminescence dating because the sediments being waler-deposited, may not hive been exposed to sunhieht for sufficiently long to zero the quart, grains, resulting in inherited saturation levels within the grains. A farther complication may be that in the Adehude area, the sediments may have not been transported sufficient distanees for the quartz grains to have been zeroed. Given these constraints, it is extremely gratilying Unt 1 was possible to achieve a rehuble lumineseence age for the Pooraka Formation at Dry Creek, offering the possibility of further advance in this area, An aim ot this projeck was (Oo establish the effectiveness Of luminescence as a citing fechmique for Quaternary: Mluviul sediments in order to resolve differing interpretations OF their ages and to (ucilitute correlation OF river terres jn different valley systems, Clearly, there is a peed for many further dates to be obtained from trver terrace and allawial fun deposits over wider-ranging afeas (o Verify the conclusions of the present work, However, the implications of un age foe the Pooraka Formation extending back to the Last Interglacial are so Sigmilicant that our preliminary results are presented here and provide the basis for further study. Acknowledgements Funding for the drilling of the Pooruka Founation und initial luminescence dating of the sediments wits provided by a University of South Australia Researeb Grant. The Th analyses were curried oul by 2. Marlinailis under (he supervision of Professor Prescott. We are grutehd ta Dr C. Murtiay-Wallace and Dr N, Alley for commenting on the text und te M, Bishop for assistinee with field work, We are upprecialive of the significant contributions of the referees Mr M, Sheard and De V, Gostin. Keferenves Atrbes, My ke (i904) Oprioul during: on nonespectilist feview. Ghat Geach. (Qual Set Rev) 1, 504-508, Anbrtson, At 1994) Comment on, Peter White's paper “Site H20 un the evidenee for early pecupation in Austialin’ Quel Mavtrnlavie V2, 0-31. Brrrecion aA. PB. SMinik B. Wo. Ponact. TL AL NP HOUR. COAG DeMasing. 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V7. 10% 12, (VOR9) Stritiwraphic investigations af Canby, ROAM Trppoke. Leb lata Sew Chanaziie rock veils and depositional environments, Like Frome urea, South Austtalia, frei. A See 8. Aust HL | 25- 167. S SHARD, MLL. Bike, MOC & Bhi phRiol ALP C1005) Allayial fins and piedmont slape deposits pp. 24243 Mit Drexel IT & Treiss. WAY, (Rds) The Geology of Soult Australia Val 2. Phe Phanerocon Sonth Aust. Geol, Surv Bull, 34. Chibi hE, CURIEAN, HE AL WHITE. Be A WASSE RIG, FF 1 C1997) Precise cliimateey of the last diterhieial pefiod, b= OMT clits fron fossil cork reeds i [he Rahamas. Creal, Sve Amen Aull, WA, 82-97, HANY Bo Flew As. EBL Borns, ROO & Linosay 1M (1976) Geology pp. SAD dy Twidale. C. Ro. Tylon M. J. & Webb, BAP) (Edsi “Natural History Of the Adehide Region” (Royal Society of South Australia, Adelpide) DUiLek. G. A, T. (996) Recent developments in laminescence dating of Quaternary Sediments. Pras Phiyy, Cieour 20. 127-145 EskNH anni AL ZW. 2. Re COLLINS. LB. 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(1992) Constraints on the age wid duration of the Just interglacial period uyd-on sea level vartations, Neture 457, 125-128. LuphrROok, NOEL (1976) he Glanville Formation at Port Adelaide. § Aus. Geral Surv. O. Geol. Netes ST. 4-7 Murray ober. CoV. (dae Aminostratighiphy of Qualerjur- Couslal sequences in southern Austeala «uy overview. Gaul, Hiteria?. 26. 09-86, — & Benengio Ao P.(199)) The List interglacial Shoreline in Australia Quer New Rev. LO 441-460. King. R. WoL Brion A. Po & Gostin. VA, (198%) Aminosthatteraphy oF the Last iterglacia) in Southern Australia. Senet D9, 44-36. Namal, KS. Vo & AlTREN. Mod. (19860 Annual dose vooversion fictors dor TL and ESR dune Am haeinerry 28, 202-205, Prebok, N.S, & Teprorp. &, H. (1990) Vertebrate Fossils pp. 99-209 In Tyler, M.J.. Twidile. CR. Davies ML & Wells, C. B. (Fils) “Natural History of the North lust Deserts” (Royal Society of South Austeatia, Advlaides, Preseor Jo Ro o& Hiertes, 2 To (LY9A) Dosmie ray contributions to dose rates for lumimeseence dnd ESR dating: large dep, and long-terin yariurions, Read. Meas, 23, 497-500, ie (1995) Frsirommitntal dose rates and ridivactive diseqailbrium from same Australis luminescence dating sites, Guar See Rew (Quart. Geoelron) $A, 4394s, KK MOARRABL BO (9999) Selective Bleach-An tinproved “partial bledch” method of finding equivalent doses for thermolumingscenve dating of quarts. Amelene THE. VA. 27-30). -Hestuey Do. & Wo rron. dT. (1993) Bstination of equivalent dose in (henmoluminesccnce doing - the “Austealtan stide” method, fai tS. REaDHEAD, M. L, (1988) Phetmoluminescence datine of qourty m deohan sediments from seutheastem Australis Quat, Set, Rev; 7. 257-264. Supako. M.S. & Bowman, GML OUR 7a) Definition of rhe Keswick Clay. Adclaide/Golden Grove Embayment. tani aind Eden Blocks, South Australian, Siar Austraalice, Geol. Surv. Quart. Geol Notes TA, ao. & (1987b) Redefinition ot the upper boundary oF the Hindmarsh Chiy. Adehude Plains Suh- Basin und Adelaide/Golden Grove Embayment, thie, 9- 16. N89) Sor, Stettignaphy and engineering geolpey of near surface amitenals of the Adelaide Plains. Mines cred Eneruy South Australie, Repart Book. 94/9 Spada. B,C, (1952) The geology of the Souit) boast Provingee, South Australia, with special reference by Quitlernury Coastline migragons and moder beach developments, Geal San. South Aust Kull 29, 1-120. Siimiuine, ©. H. Esan TM. MecCeiioon, M, oh & Tamuick. KR. (905) High precision U-series dating of corls from Wester Australia and implications for the mins und duratron af the Last Intermtuctal. Aurih Planet, Ser Lett. (35, W5-1 Md. Tare. R. (1879) The Anniversary Address ol the President Trans R. Sov, & Aust, 2, xxxix-texw. Twipack, C. R. (1968) “Geomorphology” CUhoiias Nelson (Aust) Lu Melnourne). __ , Datiy, By & Figmas, J.B. (1967) Bustitic and Climatic history of the Adelaide area, South Austiitio, A discussion. Jove Ceol TE 297-242. Wakp, W. 1. (1966) Geology. geomorphology and soils of the south-western part of Coury Adelaide. South Australin, CSIRO Soil Publication No 23. Wraps, RG, (1995) ODP Site 820 tal the itil colonigition of Subul. Que. Ausrealesia GRY3. (8 94 R. P. BOURMAN, P. MARTINAITIS, J. R. PRESCOTT & A. P. BELPERIO Wuite, J. P. (1994) Site 820 and the evidence for early occupation in Australia. /brd. 12, 21-23. WintLe, A. G, (1994) Infra-red stimulated luminescence dating of sediments. Rad, Meas, 23, 607-612. Wituiams, G. E. (1969) Glacial age of the piedmont alluvial deposits in the Adelaide area, South Australia. Aust. J. Sct. 32, 257. (1973) Late Quaternary piedmont sedimentation, sai] formation and palaeoclimates in arid South Australia. Z. Geomorph. 17, 102-125. ZHU, Z. R., WyROLL, K.-H., Coins, L. B., CHen, J. H., WaSSERBURG, GJ, & EISENHAUER, A, (1993) High- precision U-series dating of Last Interglacial events by mass spectrometry: Houtman Abrolhos Islands, Western Australia, Earth Planet, Sci, Lett, 118, 281-293. Appendix: Methodology for luminescence dating Quartz grains (90-120 um) were extracted as described in Huntley er al. (1993), Briefly, pure quartz of the right size was obtained by pretreating with HCI, followed by NaOH to break up clay aggregates, sieving, etching with 40% HF for 40 minutes at 20° C, magnetic separation and floating on heavy liquid at specific gravity 2.67. For measurement, 5.0 + 0.1 mg was deposited on stainless steel discs, Individual discs were post glow dose normalised with 6 Gy. Fine grains (undifferentiated as to mineralogy) were separated after the HCl and NaOH extractions by settling from aqueous 0.01N NaOH, the 4-11 um fraction being retained. This was then deposited on aluminium discs from acetone suspension, about | mg per disc. Individual discs were 0.5 s short-shine normalised, As stated jn the main text, both dating protocols make use of the easily bleached component of luminescence, In the case of TL this component is selected for by both temperature and wavelength of emission (Prescott & Mojarrabi 1993). For GLSL it is assumed that the stimulated emission comes from the easily-bleached component; optical filters also select for this component (Huntley et al, 1991). The output is expressed as intensity as a function of temperature for TL and as a function of shine-down time for GLSL, For both protocols the emitted intensity is measured for “natural” samples and for samples which have received additional doses from a calibrated laboratory beta-source (N+B). About half of these samples are exposed to laboratory bleaching by sunlight filtered by a 475 nm long- wavelength-pass filter (Chris James 101); this bleach removes the rapidly bleaching component completely. Some bleached discs are also irradiated (YB+8) to provide the shape of the “missing” part of the dose curve at doses less than the natural dose. The data analysis follows the so- called “Australian slide” procedure (Readhead 1988, Prescott ef al, 1993) and the data output is the equivalent dose D. expressed in grays. Such curves are known as “dose curves”. Two methods of dose rate determination were used: /n situ scintillometry (see e.g., Hutton & Prescott 1992) uses a sodium iodide scintillation crystal, 75 mm x 75 mm diameter in the auger hole from which the sample for dating is taken, The instrument is calibrated for K, U and Th and, independently, for total gamma ray dose. Scintillometry gives a completely self-contained measure of dose rate. Thick source alpha particle counting (TSAC) (Jensen & Prescott 1983: Huntley e7 al, 1986) gives a value for the contribution to the dose rate from U and Th together, and an estimate of the U and Th concentrations separately. In fact, the dose rate to the sample is effectively determined by the total alpha count and is almost independent of the relative amounts of U and Th. However, the measured ratio allows a (small) adjustment to be made to the dose rate. Combined with measurement of K, TSAC gives an independent measure of dose rate, X-ray fluorescence spectrometry is used to find K. In addition, U was found using delayed neutron activation (DNA). If this differs significantly from the other methods of assay for uranium, it gives an indication of radioactive disequilibrium, which was not the case here. It is most conveniently combined with the data from alpha counting to give the U concentration and hence a more accurate value for Th. These are the values shown in Table Table | includes the elemental analyses for all samples. The dose rates calculated for PK1S/3.5 using the conversion factors of Nambi & Aitken (1986) are shown in Table 2. The water content measured at the time of sampling was used in the dose rate calculations. Cosmic ray dose rates have been added in (Prescott & Hutton 1994), All data are included in Table 1, even though a numerical age was found for only one sample. The age calculation is conveniently expressed in terms of the so-called “Age Equation”: Age (ka) = TL of sample TL per unit dose (TL/Gy) x dose rate (Gy ka!) In this equation, “TL of sample” (which measures the accumulated energy) and “TL per unit dose” (which defines the sensitivity of the material to radiation) are measured in the laboratory on quartz or fine grains extracted from the sample; and “dose rate” is determined from measurements in the field and/or laboratory. Transactions of the Royal Saciety of §. Aust. (1997), 121(3), 95-102. NEW SPECIES OF POTOROLEPIS SPASSKII (CESTODA : HYMENOLEPIDIDAE) PARASITIC IN DASYURID MARSUPIALS FROM NEW GUINEA by C, VAUCHER® & I. BEVERIDGE' Summary Vaucuer, C. & Beveripaer, L. (1997) New species of Pororolepis Spasskii (Cestoda: Hymenolepididae) parasitic in dasyurid marsupials from New Guinea. Trans. R. Sac. S. Aust, 121(3), 95-102, 28 November, 1997, Potorolepis aruensis sp. nov. is described from the small intestine of Myoictis melas from the Aru Islands of Irian Jaya. It is most closely related to P. bradleyi from which it differs in mean hook number (17 in P. aruensis, 13 in P. bradleyi), size of cirrus sae (0,27-0,42 mm in P arvensis, 0.14-0.25 mm in PB bradleyi) and arrangement of testes. Cestodes tentatively allocated to P aruensis were also found in Antechinus naso. Pororolepis woolleyae sp. nov., from the small intestine of Murexia longicaudata from Morobe Province, Papua-New Guinea, differs from all congeners in having longer rostellar hooks (163-182 um), The generic diagnosis is re-assessed as well as the relationships between morphological sub-groups within the genus and the marsupial families they parasitise. A key to the species of hymenolepidid cestodes occurring in Australasian marsupials is given. Key Worps: Pororolepis, cestodes, Hymenolepididae, marsupials, Dasyuridae, New Guinea. Introduction Cestodes of the family Hymenolepididae Ariola, 1899 are common parasites of birds, rodents and insectivores in most regions of the world (Czaplinski & Vaucher 1994). Vaucher ef al. (1984) reviewed the species known from Australian marsupials, redescribing the three known species and erecting five new ones. All were allocated to the genus Hymenolepis Weinland, 1858 though it was noted that they formed a morphologically distinctive subgroup within this large genus. Subsequently, Jones & Anderson (1990) described a new species from a peramelid marsupial in New Guinea and transferred the other species occurring in marsupials to the closely-related genus Vampirolepis Spasskii, 1954, Spasskii (1994) erected a new genus, Potorolepis, to contain most of the species found in marsupials, although one, H. cercarteti, was transferred to the genus Rodentolepis Spasskii, 1954, with the implication that it was originally a parasite of rodents. Spasskii (1994) was apparently unaware of the species erected by Jones & Anderson (1990) and did not include it in his new genus. The hymenolepidid fauna of Australasian marsupials is relatively poorly known (Spratt et al. 1991) and its taxonomic and phylogenetic affinities are uncertain. In this paper, we describe new species of Potorolepis parasitic in dasyurid marsupials in New Guinea and re-evaluate the definition of the genus proposed by Spasskni (1994) as well as the host- Muséum d‘ Histoire Naturelle, Geneva Switzerland. Department of Veterinary Sciences, Universily of Melbourne Parkville Vie, 3052 parasite relationships between sub-groups within Potorolepis and families of marsupial hosts, a relationship first suggested by Vaucher er al. (1984). Materials and Methods Cestodes from Myeoictis melas were collected when the host animals were autopsied after a short period in captivity at La Trobe University, Melbourne. The cestodes were relaxed in water and fixed in AFA (Pritchard & Kruse 1982), Cestodes from other hosts were collected in New Guinea, Following the death of the host, the entire gastrointestinal tract was fixed in 10% formalin and the cestodes were subsequently removed and stored in 70% ethanol. Cestodes were stained in Celestine blue, dehydrated in ethanol, cleared in methyl salicylate and mounted in Canada balsam. In contracted specimens, the tegument and dorsal and ventral musculature were removed with a fine scalpel after clearing (Jones 1990) to improve the visibility of the internal organs. Some scoleces of each species were mounted in Berlese’s fluid. Serial sections were cut at a thickness of 9 ym in both longitudinal and transverse planes and stained with haematoxylin and eosin, Measurements are given in millimetres as the range, followed by the mean and the number of measurements made in parenthescs. Drawings were made using a drawing tube. All specimens studied have been deposited in the South Australian Museum, Adelaide (SAMA) or the Muséum d’ Histoire Naturelle, Geneva (MHNG). Host nomenclature follows Flannery (1995) and % C. VAUCHER & I. BEVERIDGE 3, Me ey De ee ee 2, Ply gy RSS Hes SOCAL PP MGV NEW CESTODES FROM DASYURID MARSUPIALS yy Sprall ef al (199). Tabulated morphological data were derived Irom Beveridge & Barker (1975). Vaucher erat (1984) and Jones & Anderson (1990), Host distributtan data were derived from Spratt er al. (1991). Potoralepis aruensis sp. wy. (PIGS 1-7) Tipexs Holotype fron srall intestine of Mivereriy melas wallace) Gray, 1858, Koboor ts., Aru goup (6° 128 134° 32’E). Idan Jaya. 16-vi. 1993. coll, PA, Woolley, SAMA AHC 77877: paritypes, 23 whale mounls,-8 scolcees mounted in Berlese’s fluid. serial sections. SAMA AHC 27878-27905; 2 whole mounts MANG 23407 INVER: addilional specimens; numerous specimens 2. xi. 1992. SAMA AHC S0SX6-305K7, 3 specimens, 3. sil 1992. SAMA ATIC 30588: numerous specimens Twi }993, SAMA AHIC 30589, Material examined: Vrom Myatens melas (Miijler.1840)) types. From Aarechinus — irever (lentink, HYPE): 5 speeniiens., Mt Kandi (7° 20'S. 146° 41°). Papua-New Guinea, SAMA AHC 27864-27876, SUSK4-30585. Deseriplion Based on types, Small cestodes up to 60 in length. Seotes globose. 0.52-0.65 (0.58.0 = 10) in diameter. Suckers sub-circular in Superficial views. unarmed: cup-shaped in section. with openings directed unteriarly, O.13-0.16 00, 15, 9= 10) & OL T5018 (O17. n= 10) Rostellum muscular, (180,26 (O21) x O.17-0.23 (0.21, n = 10): rostellar sae 0.26-0.38 (0.30, n= 10) x 0.26-0.31 (0.28 n= 10). Hooks 16- IK (17. = [0), arranged th Single ring with broad, curved blades; slender handles prominent: blunt cuards often with irregular surface, core of hook blade hollow. Hooks (.128-0.147 (138. 0 = 10) jong. Neck vartable, 0.74-2.05 (1.26.0 = 10) lony. Segments craspedote: mature seginents much wider chan long, 0.06-0-17 (O10, 0 = 10) long x 0.93-1,.63 (1.3). n= 10) wide: near gravid segments longer. but stivhtly narrower, 0.13-0.29 (0.22, n = 10) long. (.81-1.22 (104 mn = 10) wide. Genital pores Unihitteral Three testes arranged Imearly, one poral. two aporal: very little variation in testis distribuuon: single seement wilh + lestes: single segment wilh 2 testes: single segment with 2 poral, | aparal testes, ‘Testes oval. of similar size. (.10-0,17 (0,15. 1 = 10) long x O.08-0.1 1 (0.09, = LO) wide. Vasa efferenanit of antiporal testes run along dorsal margin of medulla (o elongate. pyriform external seminal vesicle (.06-0.16 (O10, 1 = 10) Tong x 0.04-0.09 (O07. n = 10) wide. anterior and dersal to poral testes: distal region of external seminal vesicle slender, sittwous, enters clongate Gurus suc 0.27-0.42 (0.35. 9 = 10) long x 0,04-0.05 (Q,0S. 4 = 1) wide Cirrus sac contains elongite internal seminal yesicle occupying two-thirds of volume of crerus sacs ne armature Seen On cirrus. Genital duets cioss osmoregulatory, canals dorsally, Ovary median, with 3-4 indistinct lobules, 0.06- O10 (0.08. n = 10) long. O10-0.19 (0.14. mn = 10) wide: vitellarium reniform, posterior to avury 0.03 0.06 (0.04. 2 = 10) long ¥ 0,05-0,07 (0.06, n= 10) wide. Vagina posterior and ventral to cirrus site. dilating to form sacciform seminal receptacle dorsal (i ovary; seminal receptacle 0.14-0.24 (0.18 n= 10) lany x O.O8-0.L6 (O0b w = LO) wide. Uterus origingles as Uansyerse sac on ventmil dspect ot medulla. extends to osmoregulialary canals. developing small number of diverticula, never becoming reticulate. No segments found with fully- developed eggs materi. Ventral osmoreguliutory: canal 0,03-0,05 (0,04. m = 10) i) diameter, dorsal canal narrower 0.01-0.023 (0.02. 0 = TO) in diimetes. Longitudinal strobilur museulature arranged in two concentric rings! outer ring Composed of numerous small bundles with only [3 libres per bonule: bundles of inner ring larger with 5-10) fibres. per bundle. Potorolepis woalleyae sp. nev, (FIGS #-14) Tivpes. Holotype from small intestine of Mirrexia lotigicaadaw (SMegel. 1866). Mount Missim (7 13'S. 146° 49'E), Morobe Province, Papua-New Guinea. voll. G. Gossek, 24.0.1084. SAMA ALHC 27906: paratypes. 9 fragmented specimens. | scoles mounted in Berlese’s Aluid. serial sections. SAMA AKC 27907-27919, 30590; 2 specimens MUNG 23408 INVEL Description Small cestodes, largest Imgment 35 long. Seales globose. 0.44-0.55 (0 50. n= 9) in diameter. Suckers Sub-circulin, unarmed, 0.15-0.21 (0.17. 9 = 10) tong x 0-70-4019 (0.15, n= 10) wide. Rosiellum inuseular 0,13-0,20(0,17, 9 = 9) long x O.19-0,28 (0.23. 1 = 4) Figs 1-7. Badeeilepis arueusis sp. nov. Types. 1. Scales with custellim everted. 2 Seoles wath rostetliany wrhdrawn. 3, Resiellar took ta profile. 4. Rostellar hook, view [rom posterior surface showing entarved hook guard, S. Mature Sscemenmts. 0 Neareeriivitl seumenl 7. Troasverse listological section of riiture segment: dorsal aspect towards top ol pape. Soule bars = Q.Tim L225 7) 0.0bon 24. Legend By, dorsal osmoregulatory canal CS. external seminal vesicles SR seminned reeeptacte: | pestis; UL ters Y, ventral osmoregul ory canul 98 C. VAUCHER & I. BEVERIDGE 09 #90 Qge0g 0) 09002 00.00" > s és & oo & EET = y4 co os - : . a ner Tau Onrert W en /)5) : (008100 89 FOS of 000 900 a0dd0 Oyo aus sor ay NEW CESTODES FROM DASYURID MARSUPIALS 99 wide; rostellar sac 0,26-0.29 (0.28, n = 9) long x 0.20-0.29 (0.26, n = 9) wide. Hooks number 15-19 (17, n = 6), arranged in single ring; hooks with elongate, falcate blades, slender handles and blunt guards; core of hook blade hollow. Hooks 0.163- 0.182 (0.172, n = 10) long. Neck variable, 0.90-1.41 (1.20, n = &) long. Segments craspedote; mature segments much wider than long, 0.05-0.06 (0.06, n = 5) long x 0.76-0.90 (0.85, n = 5) wide: gravid segments longer, of approximately the same width, 0.13-0.35 (0.20, n = 5) long, 0.57-1.22 (0.87, n = 5) wide. Genital pores unilateral. Three oval testes arranged in triangular array, with | poral and 2 aporal, of similar size, 0,08-0,13 (0,10, n = 5) long x 0.04-0.05 (0.04, n = 5) wide. Vasa efferentia from aporal testes run along dorsal margin of medulla to elongate, pyriform, external seminal vesicle 0.13- 0.18 (0.16, n= 5) long x 0.023-0.049 (0.043, n=5) wide, which extends along seminal receptacle, Cirrus sac pyriform, 0.14-0.17 (0.16, n=5) long x 0.036-0.042 (0,040, n=5) wide; cirrus sac contains prominent internal seminal vesicle; cirrus unarmed. Genital ducts cross osmoregulatory canals dorsally. Ovary median, with indistinct lobules, c. 0.06 long x 0.14 wide, on ventral surface of medulla; vitellarium reniform, posterior to ovary, 0.04-0.05 (0,05, n = 5) long x 0,02-0.03 (0.03, n = 5) wide. Vagina posterior and ventral to cirrus sac, dilating to form sacciform seminal receptacle dorsal to ovary; seminal receptacle 0.10-0.13 (0.12, n = 5) long x 0,05-0.07 (0.06, n =5) wide. Uterus originates as transverse sac on yentral aspect of medulla, extends to osmoregulatory canals, developing few diverticula; never becoming reticulate. Eggs spherical, 0.032- 0.045 (0.035, n = 5) in diameter. Ventral osmoregulatory canals 0.03-0.10 (0.05, n = 5) in diameter; dorsal osmoregulatory canals 0.01 in diameter. Longitudinal muscle arranged in two concentric rings; outer ring composed of numerous smull bundles with few fibres; inner ring composed of larger bundles with 10-20 fibres per bundle. Discussion Spasskii (1994) created the genus Potorolepis for several species of cestode from Australian marsupials which had previously been allocated to Hymenolepis (see Vaucher et al. 1984) or to Vampirolepis by Spasskii (1954) and Jones & Anderson (1990). Species included in the new genus by Spasskii (1994) were P antechini (Vaucher, Beveridge & Spratt, 1984), PR aklei (Beveridge & Barker, 1975), P. bettongiue (Vaucher, Beveridge & Spratt, 1984), A bradleyi (Beveridge & Barker, 1975), P. isoodontis (Vaucher, Beveridge & Spratt, 1984) and P peramelidarum (Nybelin, 1917). To these should be added P. peroryctis (Jones & Anderson 1990) comb. nov. (syn. Vampirolepis peroryctis (Jones & Anderson, 1990), the first species of the genus known from a New Guinean peramelid marsupial, Peroryctes raffrayanus. Potorolepis peroryctis is morphologically similar to P. peramelidarum and P. isoodontis, also from bandicoots, and clearly belongs within the genus Patorolepis, The species was presumably overlooked by Spasskii (1994) in erecting the new genus. Spasskii (1994) characterised his new genus as having a rostellum armed with more than 10 hooks each longer than 50 um, all with a well-developed, elongate blade, longer than the guard and with a tubular uterus which may develop diverticula. The new species described above possess these key morphological characters and are therefore allocated to the genus Potorolepis. They are readily distinguishable from the known species based on hook number and size as well as from the disposition of the testes (Table 1). Based on hook number, the species of Potorolepis fall into two distinct groups, those with 10-23 hooks, all of which are parasitic in dasyurid marsupials, and those with 24-40 hooks which occur in peramelid and potoroid marsupials. The two new species, both from dasyurid marsupials, have hook numbers in the range 15-19 and therefore most closely resemble P aklei, P. antechini and P. bradleyi. Both P. aruensis and P. woolleyae are distinguished from these species in having longer hooks. The lengths of hooks of P. woolleyae lie well outside the ranges of other species within this sub- group, though hook Icngths of P aruensis may overlap with those of P. bradleyi. Potorolepis aruensis Can be distinguished by mean hook number (Table 1) but also by the cirrus sac which is shorter in P. bradleyi (0.190 x 0.026 mm) than in P. aruensis (0.350 x 0.050 mm). In P. bradley, the central testis lies dorsal to the ovary (Beveridge & Barker 1975) rather than aporal to it as in P. aruensis. For these reasons, the specimens deseribed above from Myoictis melay are considered close to but distinct from P. bradleyi and warrant the erection of a new species. None of the specimens was fully gravid, the terminal segments instead having developing uteri which were only partly filled with eggs. This is surprising since the animals were ‘rausported to Melbourne following capture and maintgined in the Figs 8-14, Potorolepis woolleyae sp. nov. Types. 8. Scolex with rostellum everted. 9. Scolex with rostellum withdrawn, 10. Mature segments. 11, Gravid segment. 12-13. Rostellar hooks in profile. 14, Transverse histological section of mature. segment, dorsal aspect towards top of page. Scale bars = 0.lmm 8-9, 11; 0.2mm 10; 0.0lmm [2-13; 015mm 14. Legend: as for Figs 1-7, 100 C. VAUCTIER & I. BEVERIDGE Jaboratory unt] autopsy, providing adequate ime for cestodes to mature. The specimens trom Aateehinus wis are tentatively allocated to this species, They are poorly preserved and internal features are difficult to discern, Hook lengths are identical to specimens from Myaictiy, The number of huoks, 18-22 (20) (n = 5), is larger than in specimens from ML nels but the range overlaps. They may represent a distinct but very similar species, although the current evidence is equivocal, The cestodes trom Murexia lengicandata were quite severely contracted, limiting the morphological details which were visible in Whole mounts. Nevertheless. (hey represent a new species bused on the features of the rostellar hooks alone. Winle they resemble P. arvensis in terms of mean hook number, the size of hooks immediately distinguishes the material from all congeners. In having the three lestes arranged in a tnangular fashion, P. woollevue most closely resembles 2 uklei and PB. antechini, Apart trom adding to the hymenolepidid fauna known from mursupials in New Guinea. the new species described support the erection of the genus Potoralepixs by Spasskir (1994) in providing additional species which conform with the proposed diagnosis, Spasskii (1994) provisionally included in his diagnosis the character ‘genital ducts crossing osmoregulatory canals dorsally’. This is unequivocally the case in P aklei, P. brailleyi, P. aruensis, PL woolleyae and P. peroryetis and probably is similar in the remaining species of the genus, His generic definition (Spasskii 1994) also needs to be amended to allow for testes in cither a linear or triangular array and for cirri which are either armed or unarmed, Apart from these minor modifications. the generic definition provided by Spasskit (1994) appears to be reliable. The description of the new species also provides evidence in support of the suggestion first made by Vaucher ef ul. (1984) that cach family of marsupials Wis parasitised by a distinctive morphological group of hymenolepidid cestodes. Spratt e¢ al (1991) reported P. peramelidarum trom Antechinuys swainsont) which would represent a potential exception. However. the identification was tentative TABLE 1, Measurements and kev morphological features of species af Potorolepis fram marsupials. Species Species from dasyurid hosts Po aklei BF hlvadtev f antechini P aruensis P woollevae Species trom perumelid hosts P peramelidarum ft ispedontis BP perorvets Species fram potoroid hosts ES potornd P bettougiae Host(s) Amechinus sp. (indesertbed)* A. flavipes, Pseudantechinus bilarni, Sminthipsts lencepuas Anrechinus sp. (undescribed )* Antechinus swainsenit Myatets melas Murevia lanvicaidate Perameles Nana P gunnit Isoodon obesulus Iyoodon abexulus Perarvetes raffravanits Potorous twidaciylus Rettrongia gannardt Hook No, of Testis Length Hooks Distribution 83-1) (OT) }-17 (13) tiangular 103-128 (114) 1-15 (13) linear 56-59 (58) 23-24 Inungular (28-147 (138) l6-I8 (17) Iinear 163-182 (172) 15-19 (17) trrangular YA-101 (98) 35-38 linear 71-2 (79) 43-39 sub-triangulin 124-192 at) triangular 98-103 (102) 29.33 linew or triangular T9-91 (SO) 24-27 linear formerly ientited as Amechinus sumarrit (see Strahan 1995) NEW CESTODES [ROM DASYURID MARSUPIALS Wl and based on incomplete specimens. This dubious record has therefore been elimimated from considerabon ntl mare material is collected and the host record confirmed, The species found in dasyurtds. Fo aklei, Pooantechini, Po aruensis, P hradleyt and Po weallevae, belong tou group of species with & small number of rostellar hooks (10- 23) compared with 33-40 hooks tn Po tyeedentis, P. peramelidarum and Po pereryetis Yrom perameled hosts and 24-33 hooks in Po petoraé and B hettongice from potoroid hosts. The relative size of the ovary also separites the first Woogroups, We ratio of width of qwvary lo segment width being 9-15% in species trom dasyurid hosts compared with 26-39% in those from peramelids. In the first group the uterus contains relatively few egys. but itis clearly bilobed, More data are needed from hymenolepidids parasivsing peramelids and potoroids to confirm the ullity of this character. The observations of Jones & Anderson (1990) on a species froma New Guinean peramelid marsupial and the current desenpuons of new species from dasyurid marsupials from New Guinea provide additional support for the hypothesis advanved by Vaucher et al. (1984). The data alsa suggest that the hymenolepidids of dasyurids from Australia are similar ty those of New Gumea, as are the comparable cestodes of peramelid marsupials. Pinally. we agree with Spasski (1994) 1 allocating Hymenolepix cvercarteti Vaucher, Beveridge & Spratt. 1984 to the genus Kedentolepis, Key to the species of hymenolepidid cestodes occurring in Australasian marsupials based on rostellar hooks | Hooks small shorter than 30 yim Jeng, fraternoid in SHPO. eee vovenanl Hooks larger, Tength: ‘greater an in 30 yin “Tons. ht fralemmoid in shape - o scertasaxinc Oy srailverstreaiiey 2. Wvoks number 20-30, 14-18 pm long Tere TETT ESE a ET ESE Sty Tt ee etre aT Tee TT Ridenralepis Hatta Hooks 17-22. 172 2 pm Jong... Redentalepis cercartent 3. Fewer than 22 looks of if 22 hooks bragent, hooks >100 um long, More than 22 hooks or, if 22 hooks: presen. “hooks SU OWD gaty Neat sess sess ee assassin ea ee taasnaee ts 4+. Hooks shorter than or equal to 100 pin Potovolepus akles Hooks longer than 100 pin... 5. Hooks 163-182 pm Jang... Pintarolepis woallevae Hooks fess than (50 pam in length occ see eeeeee ere o 6. Hooks 103-)28 pm long. [0-15 in number prorssasshhbeaastedteinsabtbnoane (cethtooeeassante ee ofa Paroralepis bradleye Hooks 128-147 i all tong. 16-22 in number cece POLOrOlEpiy ATUCHSTS JT. Fewer than 28 WOKS, ccc gig se perseser neeesey ceteeeepay ere More thitty 28 HOOKS ccc sens besten ere 9 §. Hooks 56-59 jim long. 22-23 in gumber fo naire one bcobinsss bot .Patorolepis antechine Hooks 79-491 pire ton, 74 a7 id number eae ee ee, oe Patoralopis betngiae 9, Hooks 124-192 ym lonp, 40 in number Poderalepis PEPOINCTIS Hooks. less than ‘120. yin long. ‘few ér than 40 in number a mu) 10, Hooks shorter thi $3 WM. Pororalepis toodontis Hooks loner than §35 pmo 2 ee ee Wl Hooks number 29-34. Hooks number 35-38 ... at Potoralepis proton Patoralepis perametidarwn Acknowledgments We are grateful to Dr POA, Woolley who volleeted material used io this study and who also made specimens collected by others from New Guinea available to us. References Bevertoge. 1 & Barker. LK. (1975) Acuanid, capiliariid ind hymenolepidid parwsiles of the dasyurid marsupial, Mittechinus stiri Macleay. (8d. trom southeastern Australia. J. Meliiathal, 49, 211-227, CVAPUISSRE BO & Vancimr, ©. (199d) Family Hymenulepididae Anola. 1888 pp. 595-663 fn Khalil, b- F, Jones, A, & Bray, R.A. (Eds) “Reys to the Cestode Parasites Of Vertebrates” ICA Interminianal. Wallitigford) FLasnery, TOF 1995) “Maminals of New Giinea” (Reed Bas Chatswood ). Jonny. A, (1990) “Techniques Tor hand-sectioning thick- Viet Platyhelminths, Svat. Porusrtal, 15, 201-208, a & Asperson. TJ, C, (1990) Helminths at rodents and marsupils from Papuu-New. Guinea, with the description of two new species. Eelimastonu echymiperae usp. (Digenea Echinostomaticdae) und Manpiralepiy peranvctiy a ap (Cesteda bhymenolopididae ). Syst Pavonital. 15, 224-2497. Privciuaro. Mo HO & Keese. G. O | W. (19K2) “The Collechon and Preservation of Animal Parasites” (University of Nebraska Press. Lincoln), Srasskil. Al AL (1954) [Classification of the hymenolepidids of mammals.) Prdy Lab. tel mins 7, 120-167 (in Russian), 102 C. VAUCHER & 1. BEVERIDGE (1994) [On the systematic position of STRAHAN, R. (1995) “The Mammals of Australia” (Reed hymenolepidids from Australian — marsupials.} Books, Chatswood). Parasitologiya 28, 66-69 (In Russian). Spratt, D. M., BEVERIDGE, I. & WALTER, E. L. (1991) A VAUCHER, C., BEVERIDGE, I. & Spratt, D. M. (1984) catalogue of Australasian monotremes and marsupials Cestodes du genre Hymenolepis Weinland, 1858 (sensu and their recorded helminth parasites. Rec. S. Aust. Mus., lato) parasites de marsupiaux australiens et description Monogr. Ser. No. 1, 1-105. de cing espéces nouvelles. Rev. Suisse Zool. 91, 443-458. Transactions of the Royal Society of S, Aust, (1997), 121(3), 103-117. STRUCTURE OF THE ACOUSTIC SIGNALS OF CRINIA GLAUERTI (ANURA: MYOBATRACHIDAE) FROM SOUTH-WESTERN AUSTRALIA, AND COMPARISON WITH THOSE OF C. SIGNIFERA FROM SOUTH AUSTRALIA by Murray J. LITTLeEsJOHN® & JOHN R. WRIGHT® Summary Litr.eionn, M. J. & Wricut, J. R. (1997) Structure of the acoustic signals of Crinia glauerti (Anura: Myobatrachidae) from south-western Australia, and comparison with those of C. signifera trom South Australia. Trans R. Sac. S. Aust. 121(3), 103-117, 28 November, 1997. Advertisement calls of 51 males of the Australian myobatrachid frog, Crinia glauerti, from five sites, and 45 mules of the presumed cognate species C. signifera from four sites, were analysed and compared. Patterns of geographical variation in the advertisement calls of C. glauertr were explored and a cline was found in pulse rate. The structure and geographical variation in frequency of occurrence of another signal, the squelching call, of unknown function, in the acoustic repertoire of C. glauerti, were investigated. The findings are consistent with the earlier reports of more frequent occurrence of the squelching cal] in the south. Because of possible similarity in function, the squelching call of C. glauerti was compared with the encounter call of C. signifera. Key Worps: Crinia glayerti, Crinia signifera, advertisement cal), encounter call, acoustic analyses, pulse structure, spectral composition, geographical variation, Introduction The commonly produced call (= advertisement call, sensu Litthejohn 1977, Wells 1977) of the endemic south-western Australian species Crinia glauerti (Loyeridge) was subjectively described by Main (1957) as: “A prolonged rattling call.” Main (1957) also noted; “Adjacent to Perth the call of this species is constant and is predominantly a rattle, but occasionally a short squelching sound is made, On the south coast ... [away from related species], the squelching call is far more prominent .,. 2’ This observation of inter-populational variation in the calls of C. glauerti was considered by Brown & Wilson (1956) as a possible example of character displacement, with the squelching calls being more frequent in the populations that were allopatric to C. insignifera (Moore), a species which also has a squelch-like call (see Littlejohn 1959). Although some yalues were cited by Brown & Wilson (1956) from A.R. Main (in litt., based on andlyses by M.J, Littlejohn), the first published objective description, which included an oscillogram (= waveform) and an audiospectrogram, of the rattling call of C. glauerti was provided by Littlejohn (1959). This account was derived from the physical analysis of magnetic tape recordings of the calls of 12 males obtained at only one locality, South Perth (115°52' E, 31°59’ S), near the northern limit of * Deparment of Zoology, University of Melbourne Parkville Vic. 3052, | Hawe, §. M. (1970) Calling behaviour and Territoriality in Males of Two species of Crinia (Anura: Leptodactylidae), BSe (Hons) thesis, Department of Zoology, University of Melbourne (unpub.}. geographic distribution (Littlejohn unpub.), so that there was no consideration of geographic variation in call structure within this species. Littlejohn (1959) also noted “occasional call variation in the form of a compressed series of pulses” which may be equated to the squelching call of Main (1957) and an oscillogram of one of these pulse trains was presented. The disjunct allopatric south-eastern Australian taxon C. signifera (Girard) is presumed to be the cognate (sister) species of C. glauerti, with which it constitutes the signifera superspecies (Main 1957, Main et al. 1958). The advertisement call of this taxon is a short, rapidly repeated pulse train and the first published objective description, and an oscillogram, were provided by Littlejohn (1958). Littlejohn (1959, 1961) supported the proposed affinity of C. glauerti and C. signifera, on the basis of the much lower pulse rates of the advertisement calls, when compared with those in calls of the then recognized members of the related insignifera superspecies (Main 1957; Main ef al. 1958), Quantitative descriptions of the calls of C. signifera were also provided by Littlejohn (1964, 1970), Littlejohn & Martin (1965), Hawe', Littlejohn ef al. (1985) and Odendaal e? al. (1986). Interpopulational variation in advertisement calls of C. signifera was considered by Littlejohn (1959, 1964), and by Odendaal ef al, (1986), Straughan and Main (1966), through choice playback experiments in which tape- recorded advertisement calls of C. signifera and C. parinsignifera (Main) were offered as alternative stimuli, demonstrated that breeding females of C. signifera exhibited positive phonotaxis only to the conspecific calls. Encounter calls (sensu Wells 1977) 14 M. J. LITTLISOUS & J. BR, WRIGHT = territorial calls, Littlejohn ef al 1985) of ©. signifera) were identified through held) playback experiments by Hawe! and Littlejohn er al. (1985). The squelehing cally of C. glavertt may also have a territorial function but this hus yet be determined. In the absence of experimental documentation, the ternr “squelching call” (Main 1957) will be retained for the compressed series of pulses. Tape-recorded samples of advertisement calls of 51 individuals were obtained at five localities across the geographic range of C. glauerti (Pig, 1, Table 1) io provide a measure of geographical variation. Mundaring Weir (Site |) is the type locality for C. ulauverti (Cogger ef al, 1983), Squelching calls were 32° 118° 116° Fig. 1. Geograplic distribution (based on Tyler eral, 1994) of Crinia glaverti, and locations. of recording sites (see Table 1), TAREE 1. Denuls af recurring lavalites, produced in the recorded sequences OF 25 of these individuals, Sufficient squelching calls were obtained to provide a preliminary deseription of this type of signal for comparison with the conspecific advertisement call, and with the encounter calls of © signifera, and to-allow an estimation of geographical variation in the frequency of production relative Lo the advertisement call. Tape recordings of advertisement calls of 45 individuals from the closest populations of C. srgnifera were also obtained, namely from four localities in southern South Australia (Pig. 2. Table 1) at or near the western limits of the extensive geographic distribution of this species (See maps given by Brook 1983, 1984; Tyler 33° 37° 140° =. Vig, 2. Geographic distrburon (based on Brook 1984) 0 Crinta sivnifera in South Australia and locations of recording sites (see Table |). All sites are in Western Australia Cor Crinja elanverti aod in South Australia for OC) signifera: Species Sie Dates of recording Lacality Latitude Longitude (South) (East) ©. vlanerti I 19, 23.vil. 89 Helena River, at base of Mundaring Weir 3157" Hla! 2 O6.ViESG. 31 AV iSO 21 km south-south-west of Busselton 33°51 119! 7 DAVIE. WITVIRG Diamond Tree railway siding: 34°20” 1 bode 4 24..25.vii.86 3 kin cast of Nornalup 34°59° lin sa 5 OY HOw. 89 14 kim east-north-east of Albany 44°59" Lise © sivnifera 6 TS8ovili BOD PS.Vii 49 9.3 km south of Wanilla 34°36! )a5°4u° 7 03. O04 NY Yellowman Creek, 3.6 kit sonth-east of Melrose 32°51" yan 13" 8 28, 29.yii. 90 9 kim north-north-west of Victor Harbour 35°29! 138°36! 9 26.9190 4.4 km north-north-east of Millicent 37°31" ares CALLS OF CRINIA GLAUTERTIAND © SIGNILEKA rns 1985: Cogger 1992). The deseription of the encounter call (Sterritorial call) of Co signifera is bused on accodnts of Hawe!l and Lithejolin et al. (1985, unpub,). Materials and Methods Recording af calls Tape recordings were obtained at the breeding sites (Figs t, 2. Table 1) with an open-reel recorder (Nagra IV-S) and a cardinid dynamic microphone (Beyer M 88), Males of boty species gall from a variety of sites: om the banks adjacent to water, while sitting mn shallow water, or while floating and supported by emergent vegetation. The effective temperatures at the calling sites (surlace water, wet-bulb aie. depending on the calling position of the frog) were then measured with an eleetronie (thermistor) thermometer (Takura Digimula’ Model Doll, with sensor type SZL-64). The mean was used as. the elleclive Lemperature when an tndividudl was al the interluce between air und water. Where possible, the recorded’ mules of C. elawerti were colleeted. euthanased, preserved in Tylees fixative (Tyler 1962), stored in 70% ethanol and lodged with the Western Australian Museum. (f the specimen was preserved. the measurement of smout-urostyle length was liter made with dial calipers (to 0.1 tin; rounded 10 0.5 mm), Otherwise, a short plastic ruler (15 em) was placed along the dorsal surface of the living specimen held tn a thin, clear plastic bag. and the Smout-urostyle fend determined to te nearest 0.5 mm. Acoustic analysis of recordings For acoustic analysis, tape recordings were replayed on an Open-reel lape recorder (Revox B77 MkIT or Sony TC.S10-2). Advertisement calls were dnalysed by using a digilal audio-spectrograph (Ray Elemetrics Model DSP-5500 Sona-Graph), Where there were high levels of low-lrequency noise, a passive filter (Allison 2B. high puss, cut-off set at 512 Hz) was inserted between the output of the tape recorder and the input of the awudid-spectrogruph. Staustical procediires were carried out with SYSTAT, Version 5,03 (Systat Inc., Evanston). Characteristics af equipment used in recording and cUnaly8ix Tape transport speeds (1% cm 4!) of the recorders involved in recording and playback were periodically checked against a locally produced standard 1000 KUlz calibration tape and a frequency counter (Heath Schlumberger SM-LISA or Good Will GFC-8010G) (Revox, Seny). or against the 50 Hy AC mains Hequeney by a buill-in stroboseope (Nagra); overall viriations in Lipe speed through recording und playback are estimated to be less than +0 5%. The frequency responses of all Clectronic vomponents used in recording and analysis dre presumed ta he close lo Linear within the narrow range of frequencies of interest (c. 2.0 - 5.0 KHZ), based on manufacturers” specifications. The relatively high simpling rates used in the digital analyses (=44 kHz) préelude the produchon Ob artiliels from aliasing. Surucnire and acoustic anribines of calls The calls of the lwo species are of Simple structure ahd each consists of a group of discrete damped oscillations (Figs 3-5). Por convenience. und following previous usage, euch of these oscillations is referred ta as a “pulse.” and the group (i.e. the pulse train) as a call (Figs 3, 4). The number of pulses ina call was delermined by direct inspection of the displayed wayeforny. The depth of amplitude modulanon of the last two pulses in a cull was sometimes. less. than 100%: in sueh causes, separate pulses were recognized if the depth of modulation exceeded aboul 75%, The duration (to nearest ms) of acall Was taken as the fnterval frem the peak of the first pulse to the peak of the list pulse (= “peak-peuk duration”). The pulses in the advertisement calls arc produced in a yuaseperodic tashion, and the pulse rale (as pos!) wats calculated over a complete call as (n - 1 pulses) x 1000/peak-peak durition in ins, Dominant frequencies were measured as the peaks in # power spectrum of the whole call, To describe the temporal and spectral properlies of pulses in advertisement calls, a tape recording of one call of each of three individuals of cach species (CL glanerit, io > wh 2.1, cu. uth. 0 200 400 600 800 milliseconds Fig. 3. Waveforms of advertisement cally. Lipper, C rive vlawerni= Reterence R439-7, Sire 1s effective emperature, wWel-bulb air = PLAN? CO. Lower, ©. sfeiiferd, Referenee R408-2. Site 6. elfevtive temperatures. wet-bulh wie = He O.. wailer= 12.5 €, 106 M. J. LITTLEJOHN & J. R. WRIGHT Site 1; C. signifera, Site 6) was replayed into a digital sound card (Sound Blaster SB16, Creative Labs Milpitas), installed in a desktop computer (IBM-PC compatible). The sampling rate was set at 44,100 Hz and the sample size at 16 bits. Files were prepared in the WAVE format with the Creative WaveStudio Version 2.0 software (Creative Technology Singapore), and further processed with this package and with Spectra Plus Professional, Version 3.0 (Pioneer Hill Software Poulsbo). The figures of the waveforms and spectra (Figs 3-5) were prepared by the same procedure. One pulse (the middle pulse or next after the midpoint of a call if an even number) of each call was used for analysis. Conventional rise and decay times of pulses (from 10 - 90%, and 90-10%, respectively, of maximum amplitude) were not calculated because it would have been necessary to extrapolate between peaks of the carrier frequency (see below). Accordingly, the number of either positive or negative half cycles to reach maximum amplitude was counted and the time interval from background noise level to the peak Time (ms) Relative amplitude (dB) [\WWrerannne estimated. The pulse duration was measured as the interval from approximately 10% of the maximum amplitude (either positive or negative) at the start to the same level at the end of a pulse. Two dominant spectral peaks are present in most of the calls of both species (Fig. 4). There is a well-defined sinusoidal fundamental frequency within the wave train (Fig. 4) and this is referred to as the carrier frequency (CF), by analogy with amplitude modulation in electronics. Rates of production of advertisement call were determined by playback of original recordings on a Sony TC-510-2 recorder. For C. glauerti, the durations of five consecutive cycles of calls and intervals were measured; for C. signifera, 10 cycles were measured. The following protocols were employed to arrive at the sequences of calls selected for measurement: C. glauerti ~ the first six clear calls in the recorded sequence; C. signifera - the last 16 calls in the sequence were digitised, then the last five discarded. The interval from the end of the first call to the end of the sixth (C. glauverti) or tenth (C. Time (ms) 2 3 4 5 6 Frequency (kHz) d. a@ 30 2. o Gu z [-s FS 20 a oO 2 2 10 2 3 4 5 6 Frequency (kHz) Fig. 4, Structure of pulses in advertisement calls. Upper panels. (a). Expanded waveform. (b), Frequency spectrum for Crinia glauerti (Reference: R439-9, Site 1; effective temperature = 12.2° C). Lower panels. (c). Expanded waveform. (d). Frequency spectrum for C. signifera (Reference: R408-5, Site 6; effective temperatures, wet bulb air = 10,9° C, water = 12.7° C). Bo wn eo & onNn — CALLS OF CRINIA GLAUERTI AND C. SIGNIFERA 107 milliseconds Fig. 5. Waveforms of squelching calls of Crinia glauerti. (a). R441-5, Site 5, effective temperature = 11,8° C. (b). R44\-1, Site 5, effective temperature = 13.1°C. (c). R404-2, Site 2, effective temperature = 9.6° C. (d). R441- 4, Site 5, effective temperature = 13.1° C. (e). R407-6, Site 3, effective temperature = 11.1° C, (f). R404-3, Site 3, effective temperature = 9.8° C. (g), R407-7, Site 3, effective temperature = 10.7° C. signifera) call was then measured to the nearest millisecond, Numbers of advertisement calls of individuals analysed The number of advertisement calls of each individual to be analysed was determined as follows. Both species produce advertisement calls in long and regular sequences. In C. glauerti, production of the longer advertisement calls is slow - about 20% of the rate of C. signifera (see below). For C. glauerti, as the first step, four clear calls of each individual (n = 10) from Site 1 were chosen at random from the recorded sequence. If four clear calls could not be obtained, that individual was discarded from the analysis. For C. signifera, all individuals from Site 8 (n = 10) were used and data were obtained from three successive calls: the 12th, 13th and I4th (or the nearest clear call if there was an overlap) from the Start of a natural sequence, or after recording had commenced, For each attribute of the call and for each species, an analysis of variance was carried out with comparison between individuals. The variance was partitioned and the proportion of the variation due to within-individual effects and that due to between-individual effects was calculated. For both species and all variables, variation between individuals was far greater than that within individuals. For C. glawerti, the within-individual variation accounted for 13-20% of the variation in the data. For C. signifera, the within-individual variation accounted for 24-25% of the variation. Based on these results of the analyses of variance, it was decided that for C. glauerti, the average of two calls per individual would provide a representative sample for that individual. As the calls of C. signifera were slightly more variable, it was decided to use the average of three calls for each individual, Mean values for individuals are used in the subsequent treatment of these data. Effects of temperature Linear regression analyses of the full data set for advertisement calls (Table 2) indicated that for C. glauerti there was a significant (p <0,05) relationship between effective temperature and duration and between effective temperature and pulse rate. The linear regression analyses also indicated that there was a significant relationship between dominant frequency and effective temperature for advertisement calls of each species, Accordingly, values of the dependent variables were corrected to 11.0° C, the nearest integer to the pooled mean for effective temperature (C. glauerti, mean = 11.1° C: C. signifera, mean = 10.8° C); these yalues were used in subsequent calculations. Where the slope for the combined samples for each species was non-significant, the raw data were used in HOS M.S, LITT LISOVN & JR, WRIGHT Taber 2, Influence of effective temperature on four attributes of the advertisement cally of Crima vlauentt (9 = 91) end C signiters (n= 45) Resulls of analyses by linear regression. Sce Table 4 for ranges of temperatures. Atribute of call Species Slope Significance of Coellicwnl of Slope (pt determination (ie) Duration C. shaven “0.070 <0. ),291 C. sivnifera +0419 240 32 Sumber of pulses C glauerti O.151 OS516 0.009 C_ signiterdt +0,.290) (ddl 0.050 Pulse rite C_ vlauerty HAAS <0. (249 C. signifera r49o O.587 0,007 Upper dominant frequeney C elanerti ~ 1), 107 0.008 (34 C) sigmifera -156.566 0.001; n= 16, all specimens were alive when nieasured), A scattergram of the correlation of snout-urostyle length and carrier frequeney is presented in. Fig. 10. Site Fig. 9. Box plots Cor carrier Frequencies of advertisement calls at recording sites of Crint glaweri and C. sienifera. Values are corrected to 11> C where slopes are significant. See Fig. 6 for explanation, Tanne 5. Vales fin aun) for sneuturesivle lenuths of males of Crinta eliuerti one Cl signitera cellected after ther advertivement cally had been recorded. Species Site Sample size Mean C. vlanerti | 8 17.2 4 6 14.6 i 7 lo.2 | 3 (5.5 5 5 17.6 Total 249 16.31 C. signifera 8 7 23.4 os) 9 197 Toul 16 31 Range Standard Condition ol deviation specimen 16.5 - 19.0 0.80 preserved I4.0- 15.5 O58 preserveil 15.5- 17.0 O49 preserved IS.0- Lo.0 0.50 preserved 17.0= 16.0 0.55 live I40- 1o0 1.25 22,0 - 25.0 1.17 live 18.0 - 21.0 141 live 18.0 - 25.0 231 a ——————— 112 Geographical variation in advertisement cally af C. vlaueru Analysis of variance, with Site as the grouping factor, indicated that there ure no significant differences Jor number of pulses (p = 0,074) and curmer frequency (p = 0.411). There are. however, significant differences for the means (adjusted to 11° C) of duration (p< 0,001) and pulse rate (p < 0.001), Across the distance of about 450 km covered by the five sites (Fig. 1), there is no consistent trend in duration (Fig, 6, Table 4), although a multiple comparison (Tukey test) indicated that the means for Sites 1-3 are significantly lower than those at Site 4, and Site | also differed from Site 5. There is a cline of increasing values for pulse rates (Fig. 8, Table 4), A Tukey test showed that the following means for pulse rate differ significantly: Site 1 trom Sites 3, 4, and 5; Site 2 fram Sites 4 and 5; and Site 3 from Site a) Geographical variation in advertisement calls of C. signifera Sites 7. % und 9 wre within the continuous distribution of C_signifera (see maps given by Brook 1983: Tyler 1985) and are spaced at about 300 km intervals. Site 6 is in the isolate on Eyre Peninsula and about 300 km from Sites 7 and & (big, 2). 5000 4500 4000 3500 3000 Carrier frequency (Hz) 2500 2000 12 14 16 18 M.J. LITTLEJOHN & J-R. WRIGHT Analyses of variance, with Site as the grouping factor, indicated that there were significant differences for all four attributes of the calls (p < 0.003), Multiple comparisons (Tukey lest) revealed the following significant differences in meuns between sites; call duration - Site 6 Irom Sites 7-9: pulse number - Site 6 from Sites 8 and 9: dominant frequency (adjusted to 11° C) - Site 7 from Sites 6 and 9: pulse rate - Site 7 tram Sites 6, 8 and 9. Compearixon of advertixement calls of C. glauerti ane C. signifera Results of analyses of variance, with Species at the grouping factor, indicated that the advertisement calls of the two species differed significantly (a < 0.001) in all four attributes. From a consideration of the combined samples for each species (Table 4), culls of C. glaverti are much longer. with means of durations diflering by 3,4 times, and with no overlap in ranges of yariation (Pig. 6), The mean value for number of pulses in advertisement calls of C. glanerti is twice that of C. signifera but there is an overlap of ranges between 7.5 and 10.3 (Fig, 7. Table 4). For pulse rates, although the mean for C. glauerti is only 64% of that for C. sigaifera, the ranges overlap extensively (Pig. 8. Table 4). particularly for the closest sumples (Sites 5, 6). The mean for carrier 26 22 Snout-urostyle length (mm) 20 24 Fie, 1. Seattersram showing the correlation between carrier frequency ancl snout-urostyle length for mules of Crinia glaverti and C. signifera that were collected after thei’ calls had been recorded. CALLS OF CRINIA GLAUBRTEAND € SIGNIL ERA Iie TALL 6, Physical clnvacteristies af seven yquelching calls uf Crinia clauentiy fron the sue set ofcally presented it igure >. PAR Sn Individual ‘Temperature Duration = Number ot Low pulse High pulse Overall pulse Currier Gind tape (ey (ins) pulses rate (ps) (ps') rile (ps!) frequeney (He) feferenee) a i (R4+4E-S) LIB 475 av) 43,2 205,4 13.2 3704 bh (R441-1) 131 138 ce 31 794) 44,0 A7A6 & (RAQ42) %.6 324 a7 467 125.0 OR? 4392 dl. (RASTA) 13. S04 ‘I Abs Lift 52.7 AN40 & (RANT-0) Ih. 478 2\ S74 02,3 dis 4220 1 (R44 A} Os 604A a5 Hy O17 4.8 4263 gw. (R4Q7-7) h,7 #32 24 29,7 Wa 27,0" +048 Means Hat 359.0 40.0) ST XO 115.20 S455 4086 Runges 6-141 435% - X32 21-50 297. 46,7 617-2059 27. - 103.2 3703 43402 —— —— ———— ‘ower than low pulse fate becuse of the four long breaks in the tains frequencies is higher in C. glawerti, by L452 Hz, bul with an overlap of fabges between 3229 and 3568 Hz (Pig, ¥, Tablet), Srructire of the squelching call of C, lauerti These culls are highly variable, as is mdieated by ihe selection of wavetorms presented in Fig. 5. The pulses are sometimes in groups within weall and the pulse rate can vary greatly through a call (Fig. 5). Values for four attabutes of the seven calls presented in Fig, 3 ave given in Table 6, The pulses are of similar structure to those of the advertisement call and (here are also lwo peaks in the frequeney spectra, us in the advertisement call. For the seven calls presented in Fig. 5, the following data apply (see section on pulses of advertisement calls for methodology): duration - mean = 3.77 ms (range = 3.1 -5.1): carrier frequency - mean = 3955 He (range = 3481 - 4427); upper lrequency peak- mean = 3976 (range = 3703 - 4414); envelope modulinng frequency - meun = 877 Hz (n = 5: range = 735 - 1062); lower dominant frequency peak- mean = 3260 Hy (n = 5) range = 3100 - 3464), The difference between the means of peuks oF upper and lower Hrequencies of 716 Hz is consistent with the explanation advanced Jor the pulses in the advertisement calls - of the lower sideband of aa envelope modulating frequency, The pulses may be grouped within a call (Fig. 5) and the pulse rates can differ vonsiderably between groups mone call (Table 0). Geographical variation in frequency of vecurrence of the squelvhing call of ©, glauerti Reeorded sequences of calling by 25 of the SI individuals included squelching calls. Beeause of confusion fron: the calling by two or more close individuals, only those of 21 could: be reliably assessed. classified und counted (Table 7). A geographical trend is evident, with the frequency of production of squelching calls by individuals being ivher in the southern and south-castern sumples (Sites 4, 5, Table 8). No attempt was made to determine the extent of variation in the squelching culls of individuals. nor overtime for & population. Structure of the encatmiter calls of C. signifera Littlejohn ef ak (1985). by playbiek af advertisement calls of CL pariosiguifera and ©, vigniferd iat peak sound pressure levels (0 dB re 20 Pa) above €. YS und 104 dB respeetively. evoked encounter calls (= territorial culls sens Hawels Littlejohn 1977) from eight males of CL srenifera at Willowmavyin wi south central Vietoria (37°16" 8, I44°54" E), The encounter cull (Hig. 11) ts alse pulse train, with w regular pulse rite (Hawel, TAWA 7. Numbers af cedvertivemens calls. interiresiate (transitional) calls and squelching calls, and proparitanys if the fatter (ofall cally of (het iuadividiueal), produced by 21 moles of Cringe ghivert, Note that beeaise of overlap wilh neighbours. ealls of + Individuals Could not be unilysed, Site Individual Nauiber Number MuseDs Proportiitt ct) i “ 1) advertisement iterpwidiite squelching — sqielehing wills calls culls calls | | MI 0 ] (001 2 hit I2 2 7 (33 2 2 9 | I ()52 3 3 Ih 4 4 (15 3 3 27 j 4 (h?] 3 V alt | 4 (10 3 7 2| + I 4) 4 4 }3 ) 7 (48 + s in 2 | 0.20) 4 2 10 | 4 (1,22 a) lu Is : \2 (38 3 af in 5 4 (h2I 3 aT {5 6 4 ih 5 di 24 (| { 04 5 Su O} 0 if) vn 3 ( ‘I iQ) 13 WOU) 2 \h ' 5 0,27 2b i] 4 3 (105 5 s) 15 I } Q43 5 th It 0 | (34 5 Sh tu 5 \3 (Af Ld M.S, LITTLEIOUN & dR, WRIGHT Tabne §, Miaibens ind proportions of miley of Crinit glauert) prodietie yqueleline calls at each vite, ————kn— — Sie Number of nrales assessed Number of males producing squedchine calls Proportiun of males produemyge squelehing calls I i0 2 i 3 2 4 r 5 \ td 0.200 A (273 5 OAL? 1 0.500) it 1.000 ee COO OCC Litlejohn 1977), A wavelorn of an encounter call from Site 3 is presented in big, TL Values for means und ranges front the original date for advertisement cally and evoked encounter calls of the same five individuals discussed by Littlejohn er al (85) are presented in Table 4, u 700 200 300 400 500 milliseconds Mis Th. Waveform of an encounter call of OC) sieaifera (References: R4AO8-9) Sire 6. etfectiye lemperatures. wel bulb air 1 1.3° CL water 12.5" ©), TABLE 9. Compariven ef attributes of udvertvenenr cally cn eneounte? culls of Crnia signers from Willoynenin, Vater (frome date af Litdéjalay etal. 198s), Meuns and ranges (in parentheses) are given (n= 5) Datu are not cofrected Tar possible effects of temperuiure. Reflective lomperatures ranged frond 10.7-13.27 © (ean = 12.04), a Atirihute Advertisement call Encounter call Duration (ins) qa 192 (56-108) (| 79-2 ]4) Pulses/note 43 17s (4-5) (16-19) Pulse rate 40.6 RK_| ips!) (30),K-53.7) (742-1901) Upper dominant 4390 3217 frequeney (He) (3125-3642) (2425-4020) Production ef encaunter calls by males af C. stoner I wu subjective evaluation during playbuck of the lape recordings of ©. yfignifera, involving 53 individuids and some 3527 advertisement calls. three Interactions, presumed to jpvolye production of encounter Calls. were noted. Otherwise. the recorded sequences of 47 imdividuals consisted only of advertisement calls, Comparisan af squelching call af C. wlauerti and rerriorial call of C. signifera The variable squelehing call of C, glauerti (Fig. 5, Table 6) ts longer, contains more pulses, is of bhher average pulse rte, ine of higher dominant frequeney (han the tereitorial call oC. signifera (Fig 11, Table 9), Even so, they are both pulse trains whieh are ol similar cartier freqaeney bul of longer duration, contin more pulses, and are of bigher pulse rate than (heir respective advertisement calls (Pig. 3, Table 4), Discussion Siruchure of the advertisement culls uf hath species The advertisement calls of both species are or similar structure, cach consisting Of a quasi-periodic pulse Lain, The pulses are short damped oscillations, each with a sharp attack and an exponential decay. There are more pulses inthe calls of ©. eleueerti. und These are produced ut a lower repetition rate. The frequency Spectra are of sittilar shape. with most individuals having two peaks, the upper being aliribuited to the fundamental (= carrer frequency} while the lower 1s presumed to be produced as the side band of the envelope modulating frequency. The carrier frequency is higher in C, glawerti, and this may be correlated with the smaller size of males ol (his species ~ a usdal characteristic Of the calls of anurans (e.g. Robertson L986) There ts a (rend of Increasing pulse rate from north to south-east in the samples ol C. elawert and the durations are lower in the samples from Sites 4 and 5. Values for samples of the other two auributes display no obvious pattern, Advertisement calls from Site 6 in the geographical isulate of C. yégeifera on lower Eyre Peninsula are Jonger and contain more pulses than those to the cust in the main distribution of ©, Nignifera. Thus the westernmost sample of © Nignifera (from Site 6) is more similar to those of C giver than ace the others, The populations of © siumifera on Eyre Peninsula have probably been sepiraled from the main distribution singe the sea rose Lo its present level at the close of the last elacial period of the Pleistocene Epoch rome) 12.000 toe 6.000 years ago: sce Littlejohn er al 1993 for a summuary and references) and this isolation may have contributed to the divergence. CALLS OF CRINIA CE ALLERTE ANDO, SIGNIPLRA Ws Quendaal es at (LY86) recorded a sample of advenisenient als of Co vignifera from the same locution on Yellowmun Creek (ther Recording Site No. 5: our Sile No, 7) and aver a comparible range ol efiective (wierd temperatures (1K) 122° Cy, Yo - 41.2 © - this study). Por all four attributes, however, thei values ure Higher than those obtained in the present study. The use of other protocols anil abalytieal techniques may account for some of the differences. bur at This stage. no explanation can Pre offered, Frequency of varemaier cally iC, slavery) Moretecent observations by Litleyoha (unpub.) indicated that C. glaverd oecurs in syntopy willl © subinstgnifera (Littiejoln) near the eastern limit of Ws distributing (eg. Site S. Pig. ty) Crea swhneni/ercd bus an advertisement call that soureds ike u “eng low-pitebed squelch” (Lildeyohn 1957. 959) Bor two temporal utierbutes of the advertiseient calls of 37 individuals of C. whinsiguiferd conected to-an effective lenperalure wl 1 the mean curation rs 540 ms (range = 420 - O60) und the mean pulse rare ys 174 (range = 129 - 10) (Littlejohn M961). As these canges overlap those of the squelehing call of €. glawert? (Table 61. explunations alber (han repradiedve charieter Uisplicement aust How be sought! for the higher frequencies of becnmrenve of squelefing cally ip the soattraind south-east but none can be provided) at present. Clearly, there is a need for further investigwuons, including playback expeciments with ddvertisement culls and squelehing calls as stimuli tinder coatealled conditions. Such studies should be preceded hy jhe (locumentanou of frequency ol veeurrenee oof squelehing calls in natural ussembluges. und the context in which they are produced, The meusurement of the sound pressure levels of calls of conspecific neizhbours ts also required so that the appropriate stimuli can be applied (Linlejohn eral LYSS). By varying the levels OF sfinglation, thresholds could then be determine wd geographical patterns may he revealed. Relationships of C, glaverti Prom the presented data, appears that the nearest populations of CL vlawerti and C. yignifer CSites 5. 6) have the mose similarly structured! wdyertisement calls, Phe main difference helween the culls of the two Species is i cunier frequeney which may be devounted lor by the difference in sizes of the twe: tase (Table 3. Fig. (0). The similarities in call Structure ave Consistent with the postulatcd close relationship of the (Wo taxa suggested) by Viarin (W957). and subsequently supported by the multivariate numerical analyses of morphology ine fearures of Whe history carried oul by Bhike (19734 and Thompson (1981). Althoush consistent in showing a close ussechition between them. molecular studies do nol help ay resolving the relationships of Co efamerte and C. sigaifern, The albumin iimunolomed! wialysis of Daugherty and Maxson (1982) places © sfgnitera clusest aC riparia (1S WD units), followed by Co efadiert and ©. patinsigniferd Cooth 2+ 1D units) anu C. gearg iene (29 1D units). The clidisne analysis of allazvmes curried oft by Burendse (1984) offers several interpretations and appears incondlasive ahout the relauonships: in one scenumy Co glawert’ and C. signifera ure grouped with CL georgian. Burendse (1984) did not inelide CL niperia in his study Roberts & Watson (1994) have reviewed the recent litvrature on relationships within some eraups of Austrian frogs. Three of the specres of Crini described since the work of Main (1957) - ¢. biliveue Martin. Tyler & Davies (Murctin ef al WSO). Co renia ‘Tyler & Parker (Tyler & Parker PO74) amd €. rear Lidejathn & Martin (Littiejohn & Martin 1965) - have cleurly pulswile advertisement calls. Tyler & Parker (1974) noted the sinvilurity of the advertisement culls of © reniota and ©. efaierrn, buy they did not provide information whout the recording temperatures. As the recording of the call ob © remafa Was obtyined ut Morehead, Papua New Guinea in January. the ambient temperatures presumably were much higher than those apply iig te recordings analysed in the present study. Hence, divect comparison cannot be made woh the cally of C. glanert) obtained ab (emperiiores ob 13.4 C and lower. Blake (1974) noted thal Crit ripurte bucks y tympanum and columella and placed the taxon into a different species group ulong with Co fasmenitensis (Gunther), Because of the laek of a distinee tympanum in ©. remote (Tyler & Parker L974), 1 as suggested frat there may not be a close relationship between this species und ©. elanern As the name indicates, the advertisement calbal ce ilingue is stronaly biphasfe (Martner al W880), In this species, males commence a calling sequence with short calls (<545 ms) of high pulse rate t>76 ps!) and then gradually change over (a long calls (23580 ms) oF ow pulse rate < 54 ps!) Again, the high recording (amperatures (wet-bulb atir = 243.4 - 26.6° C) mean that it i not possible to make a proper comparison with the advertisement call of C clunerti. Evel so, the aidiospectrogrum of the Shor cull appears (ode simikic to some of the variations im the syuelching calls of Co elawerte. Martin er al (J980) considered the Functional significunee of the two disuoct culls of ©. bifimgite and rised the possibility thatthe short culls of higher pulse rade are milting calls, und that the long calls of tower pidse rile: are territorial walls, the converse af the situation Has Moa DIP IRMOTIN & TR. WRIGHHE for pulse rate in O) siuaiferea, The long call of €. Hilingue Comiuits about Iwiee as aiminy pulses. (227) as ub the advertisement call of iC) glanert tele TF allowance is made for the ditlerence my lemperalure (by Using QQ) Of 2), the pulse rare of C) ile ul HC woukl he about 20 pos band eould Overlap the pulse rates mm the advertisement calls of ©. e/awertt. Crinty bilineda possesses a Cyinpanuin bul possible close alfinities with ©. wanerti were nol considered by Martin etal (1980). Acknowledgements The field) stadies a7 Western Australia were carried gut under Licence Numbers 1274 (1986) and SF Lk6 (1989), issued by the Departinent of Conservation and Land Mingement, The field studies were wirried oul during fiekl research leave front the University of Melbourne. Equipment was provided by grunts from the Australian Researeh Council (ARC) and the Upiversity of Melbourne, Recordings of CL slenifera Von South Austrailia were abtained diving the tenure of a research grant from the ARC, Number A T8831316 (1989). Some of the ueouste and stitistical analyses. and preparation af parts af the manuscript, were carried out ducing the tenure al other grants tam the ARC, Small grants: Seheme (Gul Numbers SG 0935514, 8 OS94A7578) cod the procedures that were developed have provicee protocols Tor acoustic and statistieul analytics! techniques of Wider application, Mrs PG, Litllejohn assisted with recording al all af the sites iin Westem Australia and at wea of the South Australian localities. Dr T. CG. Litthejohn assisted with record ine at the other two South Austtalian localities, Mp bh. A, Smith, Western Austin Museuin arranged for ihe cunition ol The preserved specimens and kindly rite them available for subsequent exanminahion ane newsurement during a subsequent visit, Dy M, J, Keough provided jalvice Of stuistical aualyses and Dr Hh CL Benner-Chak provided an interpretation of the speetral amd temporal siete ot pulses i the ivertisement calls, References BaRESDsh, W. (19R4) Speciation in the genus Crude (Anurin Myobitrachidae) i souhern Australias A Phylwenciic ghulysiy ol qlovying dita supporting vhdemic speekiiii i Southwestern Australian Avelution 38, (248 1250 Biakh ALT DL 11979) Taxonomy and relativnships af myobutehing frogs (Leptodaetyfeieh A nonierical Approach, Anat fe Ave 21. 11-149 Brook, As od. (19KF) Ads a Australian Anuru’ (Depanrnent ait Zoology. University of Melbourne, Mirkville). (TU84) CATS ol Frogs oof South Ausualia’? (Department of Zoology, University of Melbourne, Parkville), Brows, Woot & Witsna., EO, (1956) Character ddsiphiceniont Swit 4and S$, 49-t. Conpk Th Cy (1992p Reptiles and Amphibians of Australie’ Sth gd (Reed Books, Chatswond —. CAMIHON, FL & Codern HW. My t)983a) “Aoolowion! Cuhowte of Australia. Vol}. Atmptibin aid Reptilia’ (Australian Government Publishing Service. Canberra) Davotbenn C) A & Maxson, be 982) 4 lehemical issessmenh oof ihe cvalutinn of tyohatrichine fregs. Herpetlagiia 38, 341-348 Leiba. MOP Ch9S7) A new species of frog of the penis Crile. West At. Net, 6, 1-23 (1958) \ new species of frag of the wenus (rine Tsehud) from soutlecaster) Austtatia Pree, Linn See, NSW 83, 227.226. (1959) Call differentiation ima complex ob seven speves of Crime i Anura, Leplodiery tlie), Avadenion 13, 452468, (MOL) Age dint origi af seme saathwestern Austin species ol Crigia (Amun Lepeodiety tidied yh S14 S36 fe Blair WO Chea Vertebrate Speesation” (Liversiby at Fesus Press, Aust), —-- HN Gedgraplie iition and nite eall Uifferentintion Wi Cefniy siaaifard Ebalytion UB, 262 Oty (1970) Chea rasanertitersiy (Ate: Leprodactylidae) Ceoyriphic distihution. mating call straciine. und rehitionships, Mroc. hl Sac. NSW Od, WY 127, ——— [IYT7T) Lone-tanee acoustie conmimuniealion ft anuriis: An iiteerated and evolutionary approdelt pp. 264-294 Jy Taylor D. TL & Cuttin Sb CRsy Phe Reproductive Binlogy af Amphibians’ (Plenuny Press, New Yorks, Tharrison, POA, & Mac wan RO © ¢7a85) Inferspecihe acouste interactions jn synpatrie populitions of Raidedke signifercd and Ay partistenifere (Ane Leptodaeciylidaes pp. 287 206 In Cirle Ciy Shine, R. & Bhan bh (eds) “Biglowy oF Austedasian Frogs and Reptiles” (Royal Zoolagieal Society of Nes South Wales. Syduey) & Surrey Boathy and Sans, Chipping Norton, & Manin, AL AW T9605) A new species of Crtiia (Aruna: Leyptoductyldae) fond South Austritia Crpere 1965, 319-424. ROKERES, JD. Warsos, G Boa Davies, M, (1993) hanily Myabatrachidue pp. 41-57 Ae Glasby. Co 1, Ross, GJ BO & Boesley, PL. (Eds) “Pau of Australia, Wal 2A Amphibia und Reptilia’ (Australian Cioverninent Publishing Seryiee, Canberra), Main. A. BR, (1957) Studies in Australian Amur I. The venus rine Tschudi in south-western Australia ane sung species from south-eastern Austila. Mak 2 “aol, 5, 30-55, J Lb ACK. & Lrereidonis, My ()958) Bvalution in three genera of Austratiun frou, Byoluiion 12, 224-233, Marin, A, AL Tyo, Mad, & Davis, M, ClYs0r A new species of Ranidelle CApura: Leprdacty lider trom Homhwestecn Austin, Capere TOBO, Yay CALLS OF CRINIA GLAUERTI AND C, SIGNIFERA 17 ODENDAAL, F. J., BULL. C. M. & TELFoRD, S. R. (1986) Influence of the acoustic environment on the distribution of the frog Ranidella riparia. Anim. Behav. 34, 1836- 1843. Roserrs, J. D, & Warson, G. F. (1993) Biogeography and phylogeny of the Anura pp. 35-40 I Glasby, C. J., Ross, G. J. B. & Beesley, P. L. (Eds) “Fauna of Australia. Vol. 2A Amphibia and Reptilia’ (Australian Government Publishing Service, Canberra), Ropertson, J. G. M. (1986) Female choice, male strategies and the role of vocalizations in the Australian frog Uperoleia rugosa. Anim. Behav. 34, 773-784. STRAUGHAN, LR. & MAIN, A. R. (1966) Speciation and polymorphism in the genus Crinia Tschudi (Anura, Leptodactylidae) in Queensland. Proc. R. Soc. Qld 78, 11-28, THOMPSON, M. B. (1981) The systematic status of the genus Australocrinia Heyer & Liem (Anura: Leptodactylidae). Aust. J. Zool. 29, 93-102. Ty er, M. J. (1962) On the preservation of anuran tadpoles. Aust. J. Sei. 25, 222. — (1985) Biogeography pp. 225-229 /n Twidale, C. R., TyLer, M. J. & Davirs, M. (Eds) ‘Natural History of Eyre Peninsula’ (Royal Society of South Australia, Adelaide). & PARKER, F (1974) New species of hylid and leptodactylid frogs from southern New Guinea. Trans. R. Soc. S, Aust. 98, 71-77. . SMITH, L. A. & JOHNSTONE, R. E. (1994) ‘Frogs of Western Australia’ (Western Australian Museum, Perth), WELLS, K. D. (1977) The social behaviour of anuran amphibians. Anim. Behay. 25, 666-693. THE BREEDING BIOLOGY AND ADVERTISEMENT CALL OF LITORIA SPLENDIDA TYLER, DAVIES & MARTIN By GRAEME F. WATSON* & H. CARL GERHARDTT Summary Watson, G. F. & Gerhardt, H. C. (1997) The breeding biology and advertisement call of Litoria splendida Tyler, Davies & Martin. Trans. R. Soc. S. Aust. 121(3), 119-124, 28 November, 1997. Breeding biology and description of the advertisement call of Litoria splendida are presented. Analysis of a call of L. caerulea is also provided and shows that these similar, syntopic species have structurally similar calls and thus presumably show significant acoustic interactions in mixed choruses. Key Words: Litoria splendida, Litoria caerulea, frogs, calls, breeding biology. Trascenods of the Royal Saucier at §, Awe (P9972, PIL VIM TI THE BREEDING BIOLOGY AND ADVERTISEMENT CALL OF LITORIA SPLENDIDA TYLER, DAVIES & MARTIN by GRABMP F. Watson & H. CARI GERIARDT: Summary Watsons, GP & Geniskor HC, (1997) The breeding biology and advertisement call ol Litnria splenditls Vy ler, Diyies & Martin. Trey. R, Sec. 8, ast T2104). 119-124, 28 November. 1997. Breeding biology and deseription of the advertisement call of (iroris splendida uve presented, Analyois of a calbol i. caerilen is also provided anid slows that these sitar, syalopre species have structurally siniikir calls wad thus presurmiibly show sinifegal yeoustc jolerachions in mixed choruses Key Worns; Lituria split, Laurie coerilec. logs. calls. breeding biology. Introduction Litaria splendida Tyler, Davies & Martin, L977, is one Of the largest fadult body length 82 - Loe ain (Hyler M82), und most beauaglul Australian frogs. The species is thought to be sparsely dystribuied throughout the widespread escarpment country at the Kimberley Division ot northern Western Australia (Tyler 1992). Literia splendida is notable for: hypertrophied parotid and rostral glands (Tyler & Davies 1993). a characteristic it shares with schescent members OF the closely related Litevte crerdteag an ability to ulilige ghindular seeretions as a Waterprool covermg (Tyler & Dayies 1993), and (he produciton of phirmacologically aetive caerins in (he skin secretions (Tyler & Davies 1993). What dite is known of the reproductive biology of L. splendida is based on reproduction by cuptiye mdividuals (Pyler 1994), fn wo aquarium, a female laid 2000 eggs, depositing them in discrete clumps ot up lo 200 eggs. which Tyler (1992) has suggested may reflect.an adaptation oF females m the wilil to lay batehes of ezes in several temporary ponds to maximize the likelihood Of at leas} some of (he offspring surveying the larval stage. Maximum length of the tadpoles was 54 mm. (Tyler 1992), Literia splendid is Known to call. and presumably breed, alter heavy rams in the early wet season (C. Done, Department of Conseryation and Land Management Kununurra unpub. obs.). During the numerous field trips to the Kimberley Division undertaken by one of us (GPW) and colleagues from the Liniversity of Adelaide over the past 20. years. however, calls of the species have never been heard nor has breedimy been observed. Jn fact, the speeres Departnivntiot Zoplogy, University of Melbourne Parkyille View Wd, Division of Biologie Science diners of Visser (abumabye Missohin USA GS5201 is rarely envountered except when associated with iuTificial habitats. for example ralet blocks. where CoOL, moist sites. iire readily available (Tylor 1992), Dining January. (997. we yistled the Kununurra urcu und en¢uuntered a breeding chorus of L. sendida, Here we deseribe the call of the species und provide brief notes Gn associated behawvieur, and alse a description of (he cull of syntopie L. ceerulec, au phylogeneticully closely related and ecologically similar species. Methods Recordings ot culls of Lo splendida were: ynade using a Sony TCD-5PRO cassette recorder (tape speed 4.76 ems!) and Beyer M-S& cardioid dymainic microphone. For comparative purposes, one call ota syilopic L. ceeruled was ohtamed trond a video sequence Of the breeding chorus (Canon Digital camcorder. Hi & mm tape). Air wet-bulb temperatures (the effective temperature of frogs sulling on land) were measured at the calling site of euch individual using an electronic thermistor thermometer (Takara Digimulti Model D6I1). Recordings were analysed on a DSP 5500. digital Sona-Graph (Kay Elemetries Corp.) using the in- built set-up #00 [Sampling rate (samples See!) 10240. requeney runge! 4+ ky] with playback on a Nakamichi Dragon cassette reeprder, Overall variations in tepe speed (i.e, from recording to playback) are estimated ap less than 0.5% and frequeney responses of all audio-eleetronic components ure close ta linear within the relevant lrequency range (bused on manufacturers’ specications) Por each call, theee prigmary attributes: were determined: (1) duration, ws the interval from the beginning of the first pulse to the end of the lust pulse (ms): (it) number of pulses per note (direct count). and (ii) dominant frequency (Hz). as the maximum 10 ti. hb. WATSON & HC. GERHARD vilue of the specteatn of power across the whale call, In addition, a derived characteristre, pulse repeliion tate (pulses 8). was caleulited as FOO) (irl pulses /duration ins, Levels oF resolution were less than | jis for tepporal eharacteristies and less than 40 Ve for Trequeney, Because there is no possibility of misidentifying (hese large vod distinclive Frogs und in the iiterests of their conservation, voucher specimens of recorded jnales were pot taken, Video ahd 35-mmn- photographic records OF (he culling niles were obtuined, Results and Discussion Breediiw vite and breeding behuavienn Breeding and calling were observed, following hreayy aftemoon fain, On the pight of P81. 1997 in an uren Of sandstone escarpment adjacent to a darge, temporary pond near the main ear park within the Hidden Valley National Park, Kununurra WA. The pond was formed within an ibdefined watercourse (hat ran along the base of the cliffs and Wis fed by ton-off from a number of temporary waterfalls that How down the clit face after heavy rainfall. Occasional cally of males al other nearby sites were heard but chorus behaviour was confined 10 this one sie, Four species of frogs. Litorta splenedida, 1. caertlea. Le rebella and Limnadynastes arnciis, were culling aruund the pond. Amplectunt pairs of 2. slendida and 1. caerulea were observed in and around the pond. No pairs of the other two species were seen, ullhough no exhaustive search wats curried out, The Foumy egy masses of /. arnatiy were scattered over the pond as were farge. floating, single-luvered sheets of hylid eges, presumubly (hose oF Le splendida and L. ceeruled, although we did netobserve any pairs of either species depositing cogs. Nevertheless. contrary to the suggestion of Tyler (1992), on this occasion all L. splendida would, Of necessity, Wave deposited their entire eps complement in the One pond because Ho other aquatic Habitits were available in the vicinity of the vhorus, Males of 4. vplenedida called [rom exposed positions either on (he pean vertion) eh face or on natin! ledges upon the rocky surtice. The tye recorded males were calling approximately: 1.5 und 2m, respectively, above the pond. Several other individuals and amplectant pairs ol L. splendid were Observed in simiku positions, Males of / cuertiea called from similar sites oi the foek face as well as from elevated positions i surrounding trees und on the ground near the pond, Liroria vahelle culled from ground-level sites near the pond and Limnodynustes ornaluy called whilst Moating in the water, Although we did not observe pair-lormation ut ego-laying. pairs Of L. splendid coun remain iw the (rating embrace tor prolonged periods, at least Wp to 94h, We observed several umplectant pairs silling in the open, or in cliff-faee ereviees, throughout daylight hours before the night of chorus wetivity deseribed above, Presumably these frogs had entered amplesxus during the previous night, Deseription af call A wave-lorm display and spectrogrun of the cull of L. splendida ave shown in Pig, 1, Table | lists the values of measured call allribules, The call is a fons, pulsed and apparently well-tunwd call that is rewulurly repeated (maximum call rate abserved wats 56 calls min’), The call is broad-band but has a tonal quitlily because its relatively high pulse cate (which exceeds the Lemporal resolution of the human auditary system and henee our ability to detect pulses at this rate) is perceived us a comples tone, The call is characterized by a very slow rise in amplitude witha rapid cut-oll after nuatnui intensity is reached, The calls of both individuals Jisphiyed a number of frequeney peaks in the power spectrum (Fig. 2, Table 1) with-an iaterpeak interval PARED LoSnmeary of cll attributes af Linnie splendida recorded i Hidden Valle National Fark, Kannativrea, Western Australia Values ure baead yp analysis of five calls of two individuals, Frogs were calling on a sapdstone chil tree between 1S and 2 ynabove a pond. Tempercitiires al the calling site were A,= 25.5" Cand Ay=25.1'C. Values for the first three attributes show the meant iid range (in parentheses), Call No. of Tridividuial Duration Pulses (mst Coe we bob, B28 (25-703) (79-89) Cin #2 THAG $22 (047-831) 74-94) Pulse Domiminil Other Notuble Repetition Frequeney Frequencies Rateips ) (Ha) (Ha) 1238 {280 400, S20, H4t) FRO (1204-1 25,2) 920, 1040, 1160, L440, 1560, L680 Has 1-H) 520, 640, TOU, BSU, (LOOK TTY. by HOO, 1260, 1520. 1O40, 1760, THRU CALL OF LITORIA SPLENDIDA 121 kHz 100 300 500 700 msec Vig. 1, Wave-form (upper) und audiospectrogram (lower) of the call of Litoria splendida recorded in Hidden Valley, Kununurra Western Australia, Wet-bulb air temperature at the calling site, 25.1" C. Note that the ordinate of the wave- form display is not labelled because it depicts a relative linear scale in volts. The apparent vertical discontinuity at around 600 ms on the audiospecirogram is an artifact of the printing process, Panne 2. Auribires of a veprescrnaive call ef Litoria caerulea recorded in Hidden Valley National Park, Kumunrra Western Australia Temperatures al the calling site were A= 25.5" C and Ay= 25.1° C. Call No. ol Pulse Dominant Other Notable Duration Pulses Repetition Frequency Frequencies (ms) Rute (ps!) (Hz) (Hz) 210) 32 147.6 1440 440, 580, 720. 1140. 1300. [S80 [22 ©. T. WATSON & H.C. GERHARDT 4 0 2 4 6 8 kHz Fig. 2. Power spectrit of the call of Liveria splendida aeross (wo ranges of Frequency; 0 - 4 KHZ, Wo stow details of energy peaks and 0) of upproximiately 120 Hz, Many of these frequency components result from amplitude modulation of at harmonic series generated by the vocal cords becuuse the interval belween components is nearly identical to the pulse repetition rate. Because of resonating and filtering characteristics of the sound- producing structures of the emitter, some of these frequency bands are emphasized, particularly those uround 520 (? the Fundamental frequency of the call), 640, YOO, 1280, 1420 and 1540 Hz (slight variations uround modal values occur because frequency values are measured in 20 Hz steps on the digital sonagraph), These spectral modifications make it difficult to determine confidently which components ure purt of the harmonic series and which are side bands arising from amplitude modulation. The frequeney band with most energy (dominant lrequency) dillered between the two recorded males (1280 for male #l and 1400 for male #2). with no within-individeal variation (within the resolution of this analysis) found in the five calls ofeach male that were analysed. Although the call includes speetutl energy peaks across a large frequency range (from around S00 ta 4000 Hy), little energy is present above 4 Ki (Mig. 2), Choris struenire Although only three other species were calling: in vhorus with J, splemdida, the chorus structure is of particular interest because the morphologically, bebaviourally and ecologically similur species. / 4 kH4, to show that there is relatively litle energy in the call above 4 kHz. cderiled, Was a conspicuous component (Fig. 3), Auributes Of a representative eall of syntopic 4. caerifea are listed in Table 2 and a wave-form ot “wo ot z = = s 3 P: 2 OG s Ee o a a] o g 2 5 4 @ 2 g x 3 3 a 2 = /E “ ~ a | i} kilz ~ secands Fig. 4, Audiospeetrogram of part of the ehorus ob toy species teeorded in Hidden Willey. Kununuira Western Australi. In (his reeardine Lira splendidi lis the loudest call, with backwound culls of f hibelhy and the very short cull of Linnie vinastes arn tay Eflective lemperatives for calling tales were: wet-bulb aie Lemiperatiine (2, splerutidia, th. ribettas 25.1'°C gnd water temperature (2. aca) 26.47 coeruled, coerulea Tt CALL OP LITORIA SPLENDIDA 134 kHz 100 300 msec 60 40 dB 20 1@) T T t 1 O 1 2 3 4 kiIz Pig. 4. Waveform, audiospectrogram and power speetrum of the call of Literia ederilea recorded in Hidden Valley Kununurra Western Australia. Wet-bulb air temperature at the calling site. 25. L" C. Note: (i) the different temporal seales on the wave-lorm ahd spectrographic displays; (i) that the ordinate of the wave-form display is not labelled because it depicts & relative linear scale in volts, display, spectrogram and power spectrum are shown in Pig. 4. The call has w similar dominant frequency and broad spread of peaks of energy as that of L. splendida, but it is considerably shorter, has fewer pulses. a much faster rise time and less abrupt cut- olf, as well as a higher call repetition rate (130 calls min'). To the human ear the call of 4. caerulea has a harsher, less well-tuned quality, Nevertheless, because of the broad spectral overlap between these two large species and their use of similar calling positions there is the potential for significant avousue interference between them, Our observations of calling and breeding in 1 splendide do not support the previous speculation that this species breeds only in the early wet season. Although our observations were made in’ mid- January, the wet season of 1996-97 was well established, two cyclones/rain depressions having already passed over the Kununurra area in the preceding four weeks (pers. obs.). During our visit, heavy afternoon rains fell on most days and this simulus appeared to trigger calling and breeding in L, splendida, Successtul reproduction also requires a continuous aquatic habitat for larvae to complete their development and it is likely that L, spleadide will suceessfully recruit new individuals to the 124 G. F. WATSON & H. C. GERHARDT population only during wet seasons that have sufficient regular rainfalls to maintain temporary ponds. Although we have no information on the ultimate fate of larvae from the breeding episode reported here, it is likely that the pond in which breeding took place remained in existence for much of this season, which was marked by substantial and regular rainfall. This outcome contrasts with the calling and possible breeding reported by C. Done from a nearby site in Hidden Valley. When this site was visited by one of us (GFW) a short time afterwards, no free water was present and larval development would have been impossible. From this experience of the unpredictable rainfall patterns of this area, even in the “wet” season, it is possible that successful reproduction in L, splendida is a relatively uncommon event, Acknowledgments We wish to thank D. Glanz and B. Watson for assistance in the field and J. Wright for analysing the calls. The study was funded by an Australian Research Council grant (S19711493). The work was undertaken under Department of Conservation and Land Management Licence No. SFO02003. References Tyzer, M. J. (1992) “Encyclopedia of Australian Animals. Frogs” (Collins Angus & Robertson Publishers Pty Ltd. Sydney). (1994) “Australian Frogs. A Natural History” (Reed Books Australia, Sydney). & Daviks, M. (1993) Family Hylidae pp. 58-63 In Glasby, C. J., Ross, G. J, B. & Beesley, P. L. (Eds) “Fauna of Australia, Vol. 2A Amphibia & Reptilia” (Australian Government Publishing Service, Canberra), & Martin, A. A. (1977) A new species of large, green tree frog from northern Western Australia. Trans. R. Soc, S, Aust. 101, 133-138. TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL, 121, PART 4 SEA-LEVEL INDICATORS FROM A HOLOCENE, TIDE-DOMINATED COASTAL SUCCESSION, PORT PIRIE, SOUTH AUSTRALIA By E. J. BARNETT*, N. HARVEY*, A. P. BELPERIOT & R. P. BOURMANE Summary Barnett, E. J., Harvey, N., Belperio, A. P. & Bourman, R. P. (1997) Sea-Leyel indicators from a Holocene, tide-dominated coastal succession, Port Pirie, South Australia. Trans. R. Soc. $8, Aust. 121(4), 125-135, 28 November, 1997. Peritidal Holocene sediments at Port Pirie in the northern Spencer Gulf of South Australia contain several indicators of sea-level change over the last 7,000 years BP. The elevations of present subtidal, intertidal and supratidal environments and corresponding sediment facies were surveyed in order to establish critical boundaries relative to the tidal spectrum. The subtidal Posidonia facies occurs at or below mean low water spring (MLWS) tide; intertidal sandflat, mangrove and samphire facies occur over specific intervals between MLWS tide and mean high water spring (MHWS) tide. Each facies is clearly identifiable in the subsurface, with intertidal sandflat facies particularly characterised by in situ articulated bivalves Anapella cycladae and Katelysia scalarina or K. peronii. A combination of several palaeosea- level indicators from different tidal facies best defines local sea-level change over the millennial timescale. Key Words: Holocene sea-level indicators, tidal zonation, prograding coastal sequence, facies boundaries. Transeo trons of ilte Revel Society ae S. Mast, (1997), F214), 125-155 SEA-LEVEL INDICATORS FROM A HOLOCENE, TIDE-DOMINATED COASTAL SUCCESSION, PORT PIRIE, SOUTH AUSTRALIA hy E.L BARNETT. N, HARVEY, A. Po Briprrig’ & RP. BouRMAN: Suramary BARN LEO. Haewiy, No Bhbrekio, AvP & Bouman, R.P(1997) SeieLeyel indicators froun i Holocene, tide daminwed coastal suceession, Port Pirie, Sour Australia Trans, A. Soo. S. Aust, W204), 125-135, 26 Novernber, 1997, Peritidal Holocene sediments at Port Pitie in the horthern Spencer Gull of South Australia comin several indieators of set-level change over (he Fast 7,000 years BP. “The clevahons of present subtidal, [arterial and supratidal envirvaments and eorresponding sediment facies were surveyed in order lo establish critical boundaries rehitive tothe tidal peer. The subtidal Posidonie ticles vecurs ator below meio low qatler spring (MEWS) Gide: intertidal sandflat, min grove dnd sumphire facies oecur aver specie miervals berweea MEWS Hide and mean high water spring (MHWS) ide. Back facies is clearly identifiable in the subsurface. with intertidal Sandflat facies particularly characterised by fa sifecarticulited bivalves Anepetla eyeleedan atid Katelysia scalar or K. perouit, A combination of several palucosea-level indicators from different lida Gigies best defines Weal sea-level change over the millennial timescale, Kay Wottos: Holocene sea level indicators. Uidal zonuticn. progridimg coastal sequence, fiers hoinilanes. Introduction Tide-diminued coustlines commonly generale prograding coastal sequences with excellent preservation of intertidal and shallow subtidal sedimentary facies (Belpeno ef af [988; de Boer er al PORK: Fleteher ef af. 1993), Such sequences can reveal Nigh-resolution records Of past sedimentation Often Containing a variety Ot palacosea-level iidicatoys — (Ferewsidt L988). A thorough understanding of the relationships of present Wdicators ak sea level, or invadition level, is required if correct iMlerpretations of pust relative sea levels ave to be achieved. With eritical appraisal of ihe present-day distribution of Titertudal facies, flora and fauna, pulawosea level history Tram subsurface steiraphy Gun be inove confidently interpreted, The northern Spencer Gull, South Australia (Pig. 1) provides an excellent example of a wide, prograding coastal sequence in uw mesolidul environment with an identifiable eonation of Higloceue coastal Uepesitional cnyionments, “A number af coastal stadies has been conducted previously in this area (Firman 1965: Bure 1982: Burne & Colwell 1982; Belperio et af l984a,b, 988: Gostin es al 1984. 198K: Norrish et ad. LOX6), Mowsoh Ciliates Cone fat Danieninenial Sduhes, GC iersily al Adlefnicle: And, SOUS, © Ponnerly Mines sid Gneegy Resouncus South Atserabi, POV Bos 1S) hasewotl So Ana 5064, Civendy Minpiir cole fa Ghitetone St Pallarton S. Mast 8004, Seftow) of Liv itorntinenhilaintl Reeruadiony Minjageent. baealiy: ol Hoeeeriy and Devitonmvnt Pieveryity af Seo Atetealie Winrerti ub Mike LevelaS. eMiiah 804s In particular, Burne (1982) identified several important palavosea-level indicators fray beach ridges, the lop ol the subtidal Pasidonia seagrass facies and base of the intertidal sandflar facies. and Belperio ef al, (M9Sdb) demonstrated the presence ol well-defined boundary between Pasideariia seagrass and intertidal sandtlat facies. Related stratigtaphic studies in nearby Gull St Vincent inclutle those by Cunn & Gostin (1985), Belperto ef al. (1986. LO88), and Belperio (1993, 1995), At Port Adelaide in Gull St Vincent. Belperio (1994) confitmed that the boundary between fhe intertidal sanme{fal ane nuingrove facies was a relhible palacosen-level mdicator, Prom all these studies, it is apparent thet there tre local and regional differences ins the reliability and distribution of various sea-level indicators. This paper provides a critical appraisal ol the dilferent palueosea-level maulivatars ina mesotidal environment, The wide progruding sedimentary sequence of the northern Speneer Gulf region. which forms the apes ula large relatively shallow inverse (or negative | estuary, is a direct response to the modern coastal environment, Warm temperatures ind law rainfall in ihe region promote high mites ob evaporation utd salinities whieh are often higher than iwerige for seiwialer, in cacess Of 40% and as nybel ats 48 (Bye T98T, Nunes & Lennon 1986). Seawater temperntires for the northern gull vary lypically berween (2° and 24°C (Nunes & Lennon 19k6), The tides are mostly seini-ditinal, with spring and peap hidal runges at Port Pirie of 3.8 im and 4m respectively. Die to the length of the gulf ana relatively show dea sealevel oscillations, wind stress can further jyercase the astronomical te 126 E. J. BARNETT, N. HARVEY, A. P. BELPERIO & R. P. BOURMAN b Port Augusta B * 32°39 Telowie Beach * AUSTRALIA ADELAIDE Weroona Island a | Le 4 tt _ _ : Km | ____.138°00' 7 : Fig. |. Location diagram of the study area in the upper Spencer Gulf, South Australia, showing the Port Pirie coastal zone, sampling sites marked by infilled circles, the tide gauge and Broken Hill Associated Smelters (BHAS). SIME. INDICATORS AT PORT PIRTE 17 sieniticandy. Th Speneer Gulf and much af the southern coastal Australia (Nunes & Lennon 1980), there is a hinge spring-Heap tidal movulition die te Khe dearhy equal lunar ane solar semidiarnal cunsditvents causing 4 dodge lide onee every lortght when With: tidal variation gees. Sehlater er atl, (1995) have postulated that this simile wmnplitude ool the major semiediuenal (idal constituents gives rise to particuhkie shallow qatter Hidal intereetions ti the upper gull, which promote sinphire and munyroave colonisation. Given the progradin Hdal sequence and (ie close prosimly of a tide piuge with reltible lony-term revords ut Port Pirie, this study was undertaken te deerme the vlevatiins OF goastal sedimentury vones velutive lo diode sea level and to jdeutit’y uppropriie jodern sediments asx anilogues ol Subsurface Holneene sedimentary lagies. This approich prayides sun opportunity to denuly the most reliable palawoseaslevel indicators iy a mesoudal environment aid ta develop a methodology for subsequent studies of relative land/sea movements at sites where bistorie clad lea valet. ‘Phe stidy is unigae tia that highhielits a Humber Of sedimentary facies, surveyed relative to modern sen level, ahd tlentifies critical equivalent iNnciwtOTS In the weologieul revord. Th does nev depend on one midieatar ti isolation, bul tises a suite of subtidal to supratidal indicaters 16 identify sea- level change. Methods Th Was vieeesstiry fo Survey the modern coastal covironments un detail to establish elevation Wilferences of the tidal zones relative (o local fidal datum. This wos done csi both basen and atomic spit levelling aisteumicnis. The first had ai ert ot Jess Tih | ennover 200 0), and neasivenients were kepl to within 400 in the, + 0,02 m). sine spint levelling. (he distance between each reading was less than 1OQ am, whieh weneraily kept vertical nehsurement errors to less than + O.O1 pi Taine based sites wathin the sandtlat, tmrave ane sabiphire Zajies were surveyed foo third-order Austin Plemht Datum CAHD) benchmirks. The supyeying wits conducted rnostly in the Pork Pirie eowstal Vicinity de Well ae Tuethee (athe noectheast within the Telowie Beaeh eoustal region. in order to decess all ol the modern-day tidal sethings (ig. 1), The present-day levels of the seagrass une sand thie gones al Port Pirie were veastred (ron water levels Mehabye to the Port Pie (de sauge., At Port Pine, ders Lidia dhithen (TI) is correlated! with (he lowest usiponumical Gide am related fo AID using i vorreelion of 1923 mi isurveyed | 7-03.10%4) Seaith Australie Ports Surberiny). Sea-level tdicators including seawrass. shell aid mengrove remains within und at the top of cach tidal zone were idenificd and recorded Tar later comparison with subsurhiee equivalents, In order to establish and simple the subsurface straliraphy, a totil of dhirty-live sites was selected within the broad coastal flats of Port Pirie Wig FA Vabrocorer was uscd to obkun cores 7A mul an diameter undp to 4m indepth At of the vibrocores were vorreeted Tur sediment eompuction by recording: penetranion depth versus core recovery length ane apply og correenon factor ta the thickness of the sedanents, Coring peripheral to anid Whi mangrove woodlands Was Carried cub Hsin a peut duger A baek-hoe wis used hy excavate scuimernts jit the sapratdal reson, Using this method, ny correchon for sanipling compuehon wits necessary. Surfieve and subsurface elevations in land- based vores anid excavations were surveyed to AMD, Murine-based vores were surveyed to TD uind taken Willi a few Kilometies of the tide giuiee to reduce the effects ol tidal lag and meteoralogical conditions, Samples were tiken back (othe laboratory, where the sediments were logeed with) particular attention being diven tothe elevations of facies boundaries and the presenee of sea-level tidreatlors. Modern depositional tidal environments in the Port Pirie aren The costal environment adjacent lo Port Pinte (Fig, 2) is a tidally-dominated lowland. Subtidal. Intertidhil and supeandal zones were distinguished hy the extent of marine influence or exposure and hy (heir vesetution assemblies and sedimentury ficies, Broad. shallow subtidal seagrass meidows pass laterally oshorewarnd frit ttertidal sands, miungrove woodlands, sarmphire-ulyal murshes una suplatidal evaparite Tat ehvitonnients, Chis association ab perhidal environments and their yegetition zones has. tow hinge extent, controlled the sliveessiae development al the coastal plain areund Por Pirie, A sehematie suminiy of the teal Zanes, assOciated vewetation and their pelaglopiships. to elevation OF (UNdaLON levels is given O) biguee 3, Tle Sublidtal zone Ii othe browd. shallow-nniae envicaninent northwest of Port Pirie township, seugriss meadiws, composed lairwely ol Pavidertia auivieedis grow (rent around amcan low water spring (MLWS) tide level vere TD) 1a LO below TD (igs 3, 4), Aayteatio daorradéy eat only survive liiited periodas ol emersence so that, al is upper growih limik a is generally patehy am restricted i below 0.25 m TD. At depths below 4 om PDL Pesidemia sins dominates the seavrass assemblage, Mawiderlea heal 126 bE, J. BARNETT, N. HARVEY. A, P. BELPERIO & R. P BOURMAN sheaths and rhizomes are resistant to decomposition, and seagrass fibres become incorporated into and bind sediments. A highly distinctive sediment facies results, producing calcareous mud and sand bound by masses of pale cellulose fibre. High sediment production and the binding and baffling action of seagrass. contribute to the rapid aceumulation of sediments in this environment, The intertidal zone The region extending (rom MLWS tide ta mean high water spring (MHWS) tide is defined as the intertidal zone (Fig. 3). This zone is characterised by periodic emergence and inundation during neap to spring high tides. At their most seaward boundary, broad sandflats have developed upon which some seayTusses cun grow ubove MLWS, Posidenia australis struggles to survive and is replaced by Zostera mnuelleri. Farther shoreward, bare sandflats are dominant, These sandflats are host to numerous epibenthic organisms including the intertidal molluses Barillarig sp., Veneridae sp.. Tellina sp.. Clanculus sp. Anapella cycladae and Katelysia scclarina or K. péronit and foraminitera that live on and beneath the sundtlat surtuce. These organisms occasionally accumulate in shallow tidal channels. Intertidal sandflats are replaced by mangrove woodlands above 1.32 m TD. Only one species of mangrove, Avicenna marina var. resinifera, has been recorded growing in South Australia (Butler e7 al, 1977; Gostin et al. 1984; Cann & Gostin [985) (Fig 5). Around Port Pirie, mangroves haye formed dense Fig. 2. Aerial photograph of the Port Pirie coastal zone. The subtidal and intertidal sandflats have been colonised by seagrasses (Seagrass meadows), Further inshore. mangroves form dense woodlands along the coastal margin and grow along dendritic tidal channels. Samphire communities occur more landward in the intertidal to supratidal zone. In the supratidal zone, the yexetation cover is sparse in between broad expanses of saltpans. The photograph covers an area approximately 10 km x 10 km. The aerial photograph as been reproduced with the permission of the Department ot Natural Resources, South Australia, Mapland, telephone (08) 8226 4946. SEA-LEVEL INDICATORS AT PORT PIRIE 129 communities within clearly defined tidal limits. While the lower limit of mangroves is close to mean sea level (1.75 m TD at Port Pirie), their actual lower limit, 1.32 m TD at Port Pirie, can be significantly different. Their distribution is — controlled fundamentally by their root system since the vertically protruding pneumatophores require both exposure to air and flushing of precipitated salts (Chapman 1975), A mangrove-algal association occurs at seaward levels of mangrove growth and along exposed tidal channels. Cyanobacterial mats also extend on to wide sandflats and into samphire areas in intertidal and supratidal zones. Numerous other organisms are associated with mangrove NW Subtidal Zone Intertidal Zone TD Elevation (m) woodlands, including the small mud crab, Helice haswellianus, which burrows into the substrate and promotes oxidation of the upper sediments. Gastropods, bivalves, polychaetes, decapods and other crustaceans, foraminifera and diatoms also occupy this zone. Landward of the mangroves at elevations above 2.6 m TD, are broad, flat, gently undulating plains upon which samphire-algal communities grow (Fig. 6). Sarcocernia — quinqueflora, — Sclerostegia arbuscula, Halosarcia halocnemoides and Suaeda australis are the main samphire communities present in the Port Pirie environment, followed by minor occurrences of Maireana oppositifolia and SE Supratidal Zone (WW) GHV Fig. 3. Present tidal and vegetation zonation along core transect relative to the Port Pirie tidal datum (TD) and Australian Height Datum (AHD). MLWS - mean low water spring tide; MSL - mean sea level: MHWS - mean high water spring tide Fig.4. Shallowly submerged seagrass meadow of Posidonia australis in the subtidal zone photographed during low tide, Width of field approximately 3 m, Fig. 5, Landward intertidal mangrove margin with intertidal samphire communities. Only one species of mangrove, Avicennia marina var. resinifera, grows in this southern temperate latitude. Dieback of mature trees along the landward margin can be observed, which generally indicates marine regression, The dead mangrove in left centre of the photograph is approximately 1.2 m tall. io Io, BARNETT, NOHARVEY A OP BELPERIO & Ro BOURMAN Hetlosareta indice toward the supratidal margin. The supraddal sane Above MEWS ude elevation (3.2 m TD). the supratidal zone (Pig, 3) 1s flooded on only the few vecusions when enher high or king tides combine with storm surge activity, predominantly from the soulliwest, or during and shortly alter extended periods of rainfall, Consequently. this zone is dominated hy evaporative processes und their associated sediments, In some ponded reas, algal mas we Well established und form cyanobacteria Hlats. Although this region consists miuainty of bare, poorly draining saline and gypsiferous flats. same samplires andsaltbushes survive (Fis, 7). OF these, Halosarcia heatloenemeides, Atriplex peliidese: Halosiucia welica wid Atriplex vesicciria are most thundint, Within this zone, variations in: elevation ive crewed by aeolian deflation and: formation vl eypsilerous dunes between remnunt tidal channels. hig, 6) totetidal samplire zone inchwing Sarencarnia quingqueflora, Sclerosteera crbuserla Heloseareica hadlocieiiaides ard Sudeda uatstrelis, The mangrove in the ripht foreground is gpprasinrately ooo lagh and the samphire bushes dre up to d0-enr in height, Mig 7, Supratthil samphire gone Tneluding oocascopal Hirlowipeda Tretlacnemotlen, \iriples Paluadase, Hiloyearier theica and Anipfes Yestera und hire exphinses of Salli, The simphire species are upto son iW herght Luneties hive also formed on the leeward margins of sabkha flats or salt lakes. The distribution of coustal environments around Port Pirie is shown in Figure &, Six distinetive tidal zones IWansecting the cost have heen identitied From seaward to landward, these are; 1) subudal acuprass micadows (not shown in figure), i) low intertidal bare or Zesrerg-covered sandflats. jd) intertidal mangrove woodlands, iv) high intertidal samphirealgal marshes, v) supratidal evaporative flats, and vi) supratidal ind extratidal clay and vypscous dunes and lunettes. Aerial photographic interpretation of (he mangrove woodland reveals that only minor change i its distribution ts apparent for the last 40 years or $0 (1957-1993), Mangroves have prograded seaward ito intertidal seaerags/sund (lal areas on the northwest peninsula of the Port Pirie River, between First and Sceond Creck and along the margins of he Port Pirie River itsell) This isin contrast to vapid seaward Mangrove colomsalton Ubi has occurred al Port Gawler (Cann & Gostin 1985) and landward colonisation in the Port Adelaide revsion (Burton 1982: Belperie 1993). Kvidence of depositional tidal sediments in the subsurface Much of the sedimentary stratigraphy at Part Pie represeats aggradation and progradution ol sedimems in peritidal environments since the hear stabilisation and slight fall in sea level trou) 7.000 yeurs BP lo present (Belperio 1995), Holovene sediments and Pleistocene alluvial sediments of the Pooraka bormation underlie most of the area, forming an undulating boundary with the overly tig tidal sequence, In some places, the upper sections of the Pooraka Formation show evidenee of being altered or gleyed by marine parewuters. The coastal sediments record an upward change jn sediment Facies that vorresponds with the lateral olamee i ike tidal zones The subtidal Pasfedenia facies is Wie Wiest extensive Holocene Udal facies in the region, Th consists of mostly grey, poorly sarted terrigenous. and caleareous sandy mud. with numerous fibres of Posidania australis and Vragmentary molluses (ee Spisule osp., Phasianella sp. Cantharidus sp, Dosinta sp. and Batillaria sp.) and foraminauera. Its thickness varies from greater Wan 4 man the present subudal zone but thins inland underlying intertidil and supratidal sediments to belween O und 2 m, depending on undulations in the surface of the underlying Pooraka Founavion. The landward extent of this Tietes indicates that much of the present coustil environment was a shallow marine environment during the early Wo mid Holocene The intertidal sundial fieies is warrey Co listht yrey, SEA LEVEL INDICAPORS AL PORT PIRIE WI poorly seried, terrigenous and calcareous shelly muddy sun fe occurs extensively inland beneath much ol the study area Having developed im response to upward showing of the subtidal sedimentary environment. Tn mest ool the Port Pirie region, the intertidal sandflal faeies is overlain by sanphire lucies. This is im contrast io The presentaiey: cidlal A0malion Where a transition fronn sandflit to Mangrove woodland generally oecurs, The intertidal mangrove facies consists of brown or hluish grey. mostly fonedleuredus sedinwnrs with fragments of roots. sheaths und fibres. tis largely resincled lo (he present diy distabution of mangrove wondlands, Le progriadational deyelapment and preservation of strata tive heen limited and Aviewiia marina sain resindere woodhinds upped to fave developed i telatively recent times, Where it is undeveloped. modern mangrove cools penetrate inte the underlying Facies, The sediment facies of the sample far Loris ut (hin veneer Over extensiveareas of sundtlac facies at fhe coastal plain, Ik consists of pale browat to yght 33°05" 137055! a Ht l 1 i i [ J henry Spencer Gulf SSS eee... = dt ST KILDA FORMATION. (UndiHerentialed Holocene marine and coastal marine sediments) Supratidal anc extratidal clay and gypsum dunes ond luneties WH Supratidal flats. Gyps@ous dlays JI Stranded beach ridges and coaslal dunes Shellgrit, shally sand and line sand Samphire-algal marsh Garhonale anc lerrigencus muds arey, Often Mottled caleaeous and tertyeenotis clay nich muds with occusional small gastropods, biyalyes wid foraminifera. Sow) plant fibres and thin tubular robles are upparent in some regions hut absent in others, dependiag of whether plot rater wats originally present ami/or preserved, Gypsuai content is variable. due largely to elevadion and evaporation History, With eypsurende dune sediments preserved atthe highest elevations of the supraddal cone. There is liltle clistinction between rarertidal and Sapa sumphire sediment facies, und the two are considered to form a single unit. While particule samphire species gan be identified grawing in wither the infertidl or supratidal gone, in the subsurface, sumphire roetlels und remains gunnel be dentitied te speeies level, In addition to the sediment facies ubove, several (HCPOCHVTOU MONS Of sUBLaeies Beele it the region (hal tive Conterponiian arilogues. In particular. pockets of cyanebacteral fieies are evident Vhroughout the intertidal ti supruodel ones. Wherever cyanobacterial Nuits ape present in the | 438°00' 3 Germein Bay Mangrove woodland. Organic, terrigenous mud Bare or Zostera-calanised law Intertidal sand or mud Hat, Mixed shell and quartz muddy sand = Organic/shelly sand/mud of mangrove woodland and sampnhire-algal marsh ABC Range Quartzite Pie} Cristal cofaey mpi ite Pon Preis eoriplivg) ron the South Avacaliie Geotoey Gutubuxe Mime anil [ney ay fesuurces South Aust ealit, [32 Ei. J. BARNETT, N. HARVEY, A. P BELPERIO & R. P ROURMAN intertidal or supratidal zone there are active sites of sediment aggradation, In the intertidal zone, storm ridge facies, or Cheniers when developed over muddy sediments, haye been formed during periods of combined high or king tides and storm events, Ridges are generally aligned parallel to the shoreline. Only one storm ridge is preserved in the western Port Pirie area, although several others occur to the cust. The northerly orientation of the coastline yenerally protects the area from dommiant southeasterly slovm- ridge forming events. Palaeosea-level indicators The specu significance of the northern Spencer Gulf is that the peritidal coastal suecession eantains a Well-preserved tevord of pulacoseu-level change, The sediments include various palueosea-level indicators that hive heen used, with uppropriate elevation dati, 16 reconstruct palacosea-levels, Although present-day tidal environments and equivalent sediment ficies may range over simificaot vertical elevations, the contuet between each sediment facies is generally more restricted, Subsurhice facies contacts can provide relatively precise estimales of pulaeosea-levels. given geourate Tidal Datum(m) ‘Supratidal flat facies Samphire-algal facies Mangrove facies Intertidal sandflat facies Subtidal Pasidonia facies Vio. Palaeoscu-level uriteria for the Port Pirte Coastal renin, D-E contact C-D contact B-D contact B-C contact A-B contact surveying: of the vertical extent of present sedimentary facies and thei’ contacts, Once the elevation range of a particular sedimentary contact is known, a height correetion for that contact can be made relative lo present sea level. This establishes the elevation at the time of deposition and indicates whether sea level has subsequently risen or fallen. We have established that, in the Port Pirie area, the boundary between Posidonia facies and overlying shelly intertidal sandfat facies provides a palaeosea- level datum corresponding to an upper limit of 0.25 + 0.25 m TD (Fig. 9). Consequently, the subsurface occurrence of distinctive, massed. fibrous Poyidonta facies in Jand-based sediments at elevations higher than 0.25 + 0.25 m TD implies that relative sea level was previously higher than at present, In a similar fashion, the intertidal sandflat facies generally occurs between 0.25 and 2.2 m ‘TI relative lo present-day sea level, However. a more precise palucosca-level estimate is provided by the sharp contact belween intertidal sandflat) facies and overlying miunyrove facies that cyuates to |.32-+-0.2 m TD (Fig. 9). At Port Pirie, the mangrove facies mostly occurs directly beneath the present mangrove woodland, und confidence in using Us contaee with the top of the sundflat factes is realest where Contemporary Elevation (Tidal Datum m) General Dascription of sediment contacts Change frorn calcareous clay to gypseous, 300.4 clay-pellet, structureless sediment Change trom organic, rooted, peaty clay 26+0.4 to cream calcareous clay, laminated to weakly rooted 2.2+0.5 Change from coarse shell sand or coquina ta cream, nalcareous clay 132+02 Sharp change from coarse shell sand or a coquina to organic, rooted, peaty clay Sharp change trom poorly sorted, fibrous, 0,25+0,.25 shelly sand and mud to cleaner, better sorted shelly sand SLA-LEVEL INDICATORS AT PORT PIRIE ae niussed wrteuliwd wadves of acelin credudie and Ketelivia sedlariic ork. perdi ure present, INUICALE frevete Post-norteny, preservation Where (he marwrove lictes 4s bseol, the contiel bemvcen the sundflat und samahine facies ts also stip, althoigh the present-day boundary herween the Heol sali gund Sgimphire Zones ds not well delined in the immediate wieiiity oF Pork Pirie, The upper lait ol the sindthial facies with samphire facies geeurs around 2.240.5 nm TO (Pig. 9), While the present Port Pirie tidal zones extit ia triosilion from mangrove woodlands to sample nirshes at 2-6 © On TD, this as nol commun Observed ih the subsurfuve sediments due tr the lack oF proseqdypondl development of this: strutiaraphic horizon, Consequently, the leyel at whieh the sunphire facies oecurs in the subsurface proves only an upproxmmite estamite of palacasea-level relayive Wits present elewauenr range aF 22 be A on TH, Although different samphite species are closely related Lo small elowition chanees, these are nyt bbserved at the mucro-level jn ihe subsurhice. Discussian Several factors must te adilressed when inturpretiny the evidence for palieusea-level change fron progetcding. peritidal sequences. In purteular, the relationship of ewel) indigalor ju sea level at the fime ol ils formation must be established. Al Port Virie. the subsurface presenee of iv sie fibros renwins OF the Scagrass Puyideie austradig indicates thar sea level was above this site ut the time of deposition, “Phe (ausitipn between Posie bacies und overlying intertidal sundllat facies is a more powerlulindicutor ol paulaeosea-level. vurrespondiig i) O95 +4 025 wy presentalay TD. Siilurly, mingroves grow will a Rurky broad: intertichal range, but their contagh well the imlertidal same facies provides durum of 132 = 0.2 m. Mangroves have previdusly heen cited as one al the more reliable figed, vie site palacosed-level indicutors (Hopley & Thom 183, Thom & Roy (983). For northern Spencer Gull, Burne (1982) reported: a range opm the elevihioo of seaward thmgrove colomsation from ES to 2.9 mM TR (0.4 ta 10m AHD) und us previously mentioned, we record a lower level ob mangrove volomisutionat 132+ 0.20 TD. Clearly, the level of seaward colomsaton of mangroves depends prmnwily on local coastal dynamics or coustal orientation, and will ovcur a a vuriely al clevulions relative to the tidal spectrum (Allen 1995). Therefore, i follows that the height of the contac between sundial und mangrove liwies will also vary. IL 1s apparent front the differences in elevation that the ise of mangroves as palilensed- fevel midicators can only be applied locally. where present-day clevalions uf mangrove seaward erowth we owell defined Even in’ this case. the palacoenviranment muy huve differed from the Moder environment, producing different teal ninges and mangrove distrbations. The boundary between samdflal and either Hngtave or samphine facies has the potenchil te define palacosea level. purtivularly singe its contiel in the subsurface is sharp. Hewever, the conmunelrtiin a Port Pirie is thik while the present-day zonahon from sandilar to mangreve waedland is exrernsive his (ransiian ts not easily observed in the subsurtace, Furthernore, while the present-day Lainsilinn belween sundial aml samphire zones is nol well tepresented iit Port Pirie, tis eowraee ii subsurface Sediments ts Widespread, Near Port Pirie, the present cleyalion ob the supdlflat/samphire houndary is 2.2 + 0.5 pm TD. To the northeast il Telowie Beuel, Unis boundury oewurs ue 2S 23m TD, a slightly higher elevation thaw for Port Pine possibly due to local geomorphic factors ane sedimentary processes in the lee of Weeroona tstand (Five. 1). The elevation of the top of the sand vanes depending on whether it is suceceded by mhwwroves of samphiee, Thus, athouely the conmuet can be used as a determinant of seci-level change, there ts wide range in ils elevation. This problem niuy be mimimised by curelul field surveying of the local region, Beach ridges and the lop ool Payidonuios seag rigs deposits are relatively good indicators oF pulaeossi- levels. However as wilh imungraves. beach ridge clovition dig) cunnot he used on a reg tonal busts since the elevations to whieh such ridges aire constructed ure highly dependent on local wave resimes, In regard to seagrass us a-sedevel indicator, Posidenta anstrals presently grows to 0.25 in TD (1.68 m AND) at Port Pirie, but elsewhere in northern Spencer Cull, on clevalion of nt mW TD (2-2 m AHD) fas beer observed (Burne (982), These differences muy he best explained by varying Comskul oriengiion, waye regime and coustal circulation patterns. Although a shore- parallel zonation of sediments and vegetition is common throughout the harthern sult, cach sediment and floral or fauna) community, whether Mosidenia- dominated seagrasses. Anapelia or Katelysig sp. bivalves, mangroves or hulophyles and silthushes, bas a broad regional range in elevation, Henees it ts impenttive that Toca! elevation eontrals und docu coniitions be used in assessing sei-level dala rather than applying regional values, Given that the cleyation range of bidi) facles atid sea-level indicators cur vary, vrealer aceurtey: on reconstruction of pulacesed-level is aehieved if several different indicators aire used. Each muicator. either relational or fixed, will provide evilence thut either Sapports or challenges andicators fram othe 134 E. J. BARNETT, N. HARVEY. A. P. BELPERIO & R. P. BOURMAN horizons. By using such an approach, some of the problems associated with tidal indicators, the elevations of which are influenced by local geomorphic and climatic variations, may be reduced. This study indicates that a combination of palaeosea- level indicators from the top of the Posidonia facies and the contact between sandflat and either mangrove or samphire facies is the most reliable method for establishing sea-level change in the Port Pirie area. A further factor to consider in the reconstruction of palaeosea-levels is whether tectonic activity or subsidence, due to sediment compaction, has occurred subsequent to deposition. There is little evidence of local tectonism in the northern Spencer Gulf during the Holocene, but rather, the region has been uplitted in response to isostatic adjustment of the Earth’s crust due to eustatic sea-level rise (Belperio 1995). The effects of sediment compaction in the region are less clear. While littke compaction has most probably occurred in either the thin veneer of samphire facies or within the sandflat facies, it is feasible that the extensive, muddy, Posidonia facies has undergone some compaction. If this has occurred, it would affect elevation corrections relative to present sea level, acting to decrease the apparent height of former palaeosea-levels. Conclusions The tide-dominated coastal plain around Port Pirie has resulted from sediment aggradation, coastal progradation and relative sea-level regression associated with slight sea-level fall following stabilisation around 7,000 years BP. It consists predominantly of subtidal Posidonia and intertidal sandflat facies. These facies occur throughout the coastal stratigraphy and underlie present-day intertidal mangrove and supratidal samphire zones. A tidal-vegetation-sediment relationship exists for each of the Holocene facies deposited within the coastal zone. In the upper subtidal zone, Posidonia australis dominates the seagrass community and binds the sediment. The intertidal zone is composed of bare or Zostera-covered sandflats that are replaced by Avicennia marina var, resinifera toward the shore. Further landward in the intertidal to supratidal samphire zone, Halosarcia, Sarcocernia and Atriplex communities have become established in between sabkha-like, bare supratidal flats, Associated cyanobacterial mats grow within mangrove, samphire and supratidal environments. For each sediment facies, biological palaeosea- level indicators are defined by their growth positions in relation to the tide. At Port Pirie, Posidonia australis represents the subtidal environment from just above mean low water spring (MLWS) tide (0.25 + 0.25 m TD) to depths greater than 4 m TD. Jn sin articulated shells such as Anapella cycladae and Katelysia scalarina or K. peronii are representative of the intertidal sandflat environment from 0.25 + 0.25 m to 1.32 + 0.5 m TD, and mangrove facies represent deposition between 1.32 + 0.2 and 2.6 + 0.4m TD. Good precision in palacosea-level interpretation can be obtained from peritidal sediments that reveal clear and consistent transitions and contacts from one facies to another. This study has established that the transition from Posidonia to sandflat facies and the sharp contact between sandflat and mangrove facies are the best palaeosea-level indicators in this environment. The contact between the sandflat and samphire facies can also be used to establish sea- level change, although only in areas where its present elevation can be established. Dangers are apparent in the broader, regional use of facies boundaries due to the often patchy and variable development of different facies along the coast. The use of tidally-dominated sediment contacts as palaeosea-level indicators depends primarily on an accurate determination of their present-day elevation ranges relative to tidal datum. Our research has demonstrated that in order best to define palacosea- level. fieldwork must be carried out at the local scale and take into account coastal processes that have been operating over the long or short-term in the region. Acknowledgments This research has been supported by a National Greenhouse Advisory Committee Grant. Field and technical support were provided by R. Rice, S. Rowe. B. Logan and J, Cann. D. Fotheringham assisted with surveying and vegetation identification. Access into Pasminco Metals/BHAS was granted by A. Gilbert. The aerial photograph was supplied by the Department of Lands, South Australia. The authors acknowledge K. Gowlett Holmes from the South Australian Museum for her assistance in the identification of shell material, and S. Proferes for drafting the figures. SEA-LEVEL INDICATORS AT PORT FIRIE 135 References Abbbs Oh WK. be (99S) Sall-rarsh growlle and: Huetuating sea levely Tmpleations of yo simulation model for Phindrian coastal stratigriphy and peat-based sea-level curves. Seek Gent WO, 2|-45 Brink, ALP. (1993) Land subsidence and searleyel rise inthe Port Adelaide estuary: implications for monitoring the greenhouse effeet, Aust. J. Barth Set 40, 359-368. 1 1995) The Quaternary pp. 219-280 Jr Drexel. J. & Preiss. W. V. (Eds) “The Geolowy of South Austiiha™ Vol. 2. The Phanerozoie. Geological Survey of South Australi Bulletin 54. (Mines and Energy, Adelaide), _. Swit Bo W, Pobaen, HLA. NitrRourk, C, A. DeMasiir., 1. J, PRrescory JR. Hans. I Re & Gustin, VooA ()O8day Chronolowical studies of the Quaterniry marine sediments of northern Spencer Gull. South Austtalia, Mian Geel, 1, 205-290, =_ Hans, J, R. Gostin, VA, & Powe HOA (19S84h) The stratigraphy of coastal curbonile banks and Holocene sea levels of northern Spencer Culh, South Australia, (bid, 297-313, - Cann. ib He & Costin, VA. (1986) Quaternary stratigraphic sedimentyry environments, northeusterny Gulf St Vineent, South Australia pp. 83-98 Jn Parker, A. J (Rue) "One diy excursions of the Adelaide region” Fighth Australitn Geovosicul Convention, Geologival Soelety of Australia (South Australian Division). —_, Gostin, VA, CAN. I. He. ke MURRAY-WALLACK, C, V. (1988) Sedimentorganisin zonation und the evolution OF Holocene tidal sequertees it southern Austalia pp, 475-97 di ile Boer, POL. van Gelder, AL & Nin SD. (lds) Tide-Influenced Sedimentary bovironmenty und) Pacies” (D2 Reidel Publishing Company, Dordrecho. Boenn, RV, 982) Relative tall of Holocene sea level and cousGu progradation, northeastenmy Speneer Gull, South Australia, BMRA Aust, Geol Geaplys. 7. 35-45. & Conwrin. J Bo (1982) Temperate carbonate sediments of Northern Spencer Gully South Australie a high salinity “lorumol’ provinee, Sedimeritel, 29, 223-238, BuRrow. TB, (1982) Mangrove changes recorded north of Adelaide 1935-1982. Trans. Ro See S. Aust 106, 183 1X9, BUTLER, A. J., Deepers, A.M. MeKILLUR Ss O.& THOMAS, Dp, PB (1977) Distribution and sediments of mangrove forests in South Austedian (id, POL, Fa-d4, Nye. J AT (1981) Exchange Processes for upper Spencer Gull, South Australia, did, LOS, 59-66, CANN, JH Gosvin, VAL (1985) Coustal sedimentary (acies und. foraminiferal biofacies of the St Kilda Formation at Pon Gawler, South Australia. (bid, 109, 121-142, Chapman, Vood. (1975) “Mangrove Vegehition” (J. Cranmier Germatyy }, bt Bork. PL, van Getbkk. A. & Nt. S. D. (Bids) (1988) “Tide-Influenced Sedimentary Environments and Facies” (D. Reidel Publishing Company, Dordrecht). Hiewas, J.B, (1965) Late Cainovoie sedimentation in narthern Spencer Gulf South Australia Trevi, AL Sao 4 Aust. 89, 125-131. Beerenne, CHL VAS PEEP E., BRUSH, GS. & SHPEM AVS, J. (1993) Tidal wetland record of Holocene sewlevel movements and climate history. Paldeapeoge Palacoctimatol, Paleeooecal WZ. 177-214. Gosiis, VoA. Hats. Ro & Benes, ALP elas) The sedimentary tramework of northern Spencer Gull, South Australian Mon Geol 61, 111-138. . Borrrrio, A. PB, & Cann. J. H. (1988) The Holocene non-lropical coustul and shell carbonate province of southern Ausiralia. Sed. Geel 60, 31-70, Hopuey, D. & Tron, B.G. (1983) Australian sea levels i the last 15,000 years: a review pp. 3-26 Jn Hlopley. 2. (Ed) “Australian Sea Levels in the Last 15,000 Years; A Review” James Cook University Department of Civounipliy Monogriph Series 3 Jimes Cook University, Townsville), Nokes, Ky Rosser, H, & Waren, Le hk (los) A geochemical study of metals present in sediments from Spencer Gulf, South Australia. Appl Geochim 1 LLP 130. Nowres. Ro ALK Lawson, GW, C1986) Physical property distributions tod seasonal trends in Spencer Gull, South Australias an inverse estuary. Aust 2 Meh [restin Res. 37, 30-53 Sener. C. Byt. J ALT. & Hagsison, Po (1995) The most vigorous South Austealian tide, Tras Re Soe. 4. Aust, W198, L23-142 Teewisdt J 1h ClO88) Palaeotidil reeonstructions of inshore tilal deposttional cnviraniniedts pp, 233-2603 de Boer, P. L.. vin Gelder, A, & Nia, S. 19. (eds) 'Tide- Influenced Sedimentary Environments and) Facies” (D, Reide} Publishing Company, Dordrecht), Thom, B. Gr. & Rov. Po S. (£983) Sea level change in New South Wales over the past 15 000 years pp, O4-84 Jn Hopley. D. (Rd.) “Australian Sea Levels in the Lust 15,000 Yeurs: A Review”. Department of Geourupliy. James Cook University of North Quecnslind Monograph Series 3 (James Cook University, Townsville), THE RESPONSE OF GALL-INDUCING SCALE INSECTS (HEMIPTERA: ERIOCOCCIDAE: APIOMORPHA RUBSAAMEN) TO THE FIRE HISTORY OF MALLEE EUCALYPTS IN DANGGALI CONSERVATION PARK, SOUTH AUSTRALIA By P, J. GULLAN* 7, P. S. CRANSTONT* & L. G. COOK* Summary Gullan, P. J., Cranston, P. S. & Cook, L. G. (1997) The response of gall-inducing scale insects (Hemiptera: Eriococcidae: Apiomorpha Riibsaamen) to the fire history of mallee eucalypts in Danggali Conservation Park, South Australia. Trans. R. Soc, 8. Aust. 121(4), 137-146, 28 November, 1997. Mallee communities, especially the plant components, are often considered to be fire- adapted but there is no information on how effectively any phytophagous insects re- establish their populations after a wildfire. We addressed this issue by studying the scale insect genus Apiomorpha Riibsaamen, in which species induce conspicuous, sexually dimorphic galls of species-specific morphology on Eucalyptus species. In early 1996 we surveyed the species richness and abundance of Apiomorpha galls in relation to fire history and species of host eucalypt in mallee vegetation at Danggali Conservation Park, South Australia. Half of the fourteen sites surveyed had been burnt by wildfire in late 1985, whereas the other seven sites had not been burnt for at least 45 years. Only the two commonest of nine Apiomorpha species showed little or no host-plant specificity. Long-unburnt sites did not differ in species richness nor in total abundance of Apiomorpha galls from sites burnt 1n 1985, although the two commonest Apiomorpha species differed in their responses to fire history. Key Words: Fire history, mallee, galls, Eucalyptus, Coccoidea, Apiomorpha. Tran neny afte Bowel Socrery ef. Avan 1997), ERTL LTT Ul, MALLEE EUCALYPTS IN DANGGAL] CONSE THE RESPONSE OF GALL-INDUCING SCALE INSECTS (HEMIPTERA: ERIOCOCCIDAE: APTOMORPHA RUBSAAMEN) TO TILE FIRE HISTORY OF RVATLON PARK, SOUTH AUSTRALIA by PL GuLLAN *. PLS. Cranston’ & LG. Cook" Suiinary Guitan. BI. Cragston. PS. & Cook. LG (007) The resprnke of gallanductiy scale ciseets (Mennytera Lrovoccnlie: Aprominphe Riibsaamen) to the lire history of mallee cuealypis in Dungwali Conservatron Park South Australian Tray. WN, S Ads. TBA), 137 P46. Ts Novernher 199? Mullee communities. espechilly the pliant components. dre often considered! to be fire-adapted bur there tne iiformating on how effectively any phytophigods pisces re-establish Then populations alter a wilttire We adilressad this issue hy studyie the sedle insect genus Aprmoernle: Baibsuamnen, in whieh species midltice CAfspiclious. SeSOdHY CIMA phe Balls Of species specitic morphology on Avcrvpnis species. Tt early (990 we surveyed Thespecios richness anid abundance of Apiearacenyaltes aaulls Wi relaticn ta fire history and species al hast encalypt in mathe veretilion at Pangea) Conservation Park, South Australia, bbullol the Fourteen stiles surseved laut heen burnt by waldfire in tate [9SS, whereas rhe orher seven sites Hate por heen burnt for al least 45 years, Only the iyo caummonest af mine Aviiimeepii spemes showeel little or no Host-plint specsienty. Leue-unburit sites id nat difler in speies rightiess ner in tani abundinee Of Apremanpa palls frond sites huent in Tess. clLodeh (he Avo Canmenest ieaiapla species differed in their responses to fire History. Gully of fb. metecucoly Gutlan were equally ubundgat ul long-unburdt and burnt stles. whereas valk of ovicelandtes Hepper) were on sverage four dimes more abundant al lome-unburnt (har al burnt sites; this differences nity relate ty Uiflerential dispersal steatewies of che (st cistur nymphs. We conclide that mallee willfires sb intervals al Hoare thi 10 yours wauld be unlikely to impact detrinentalby ba the long-lero survival ofa uliwerse A ptirneipilis assemblage provided That some longer Onburn’ areas (rerigi) remain to serve as souree’s far the calonising ny tiplis hy Worps: Dare history, nmaitles gulls, Eaenhipria, Cyceailew, Aprenoryltia: Introduction Tire is i significant factor i Australi ecolory and inmuch of the continent the flora is fire-adapted (Barlow JRL) Many ophints, tiecluding inst Bicalyprs Lo He, species (Myttaecic), possess fire- protected structures (epicormic buds or Nenotubers) from whieh new growih sprouts, or have seeds thal serminite aller fires (CH MY8 tab: Hodgkinson & Griffin 1982: Noble 1982) Sueh regeneration ubihties ure purteularhy characteristic ab pints i mallee vevetulion, thal is. woodland eormmunities Uonhimited by imultiestenmead cuealypts, which are themselves also called mallee (Noble 1982) Kor inuny cugulypls, espeerlly millees. fire cua fer itate seed germination (Wellington 1989) or elimiale phytophigous tisects (Noble 1482) une parasitic Inisileloes (Call (Yb 1a) Gucalypts dominate most vl Australis forested ecosystems ind have cormples rehwonships with mative ands. ineluding many iisects (Greensinde & New 1991). Habind MaaeeMent using five whether Comastent wath History (ee. lo reerese purported past Aboriginil lind mitnuwement practice) or far havard reduetion, has effects (hat are rehitiyvely well understood fo Iyision of Motu aid Avcloy. The Suxtrahiin Mater Ciel Cadberna ACT 0200, ' Suinath Rewuntoes amb Biewliversiey Programe Bevisiony at Dtoiintagy CSEROEGIPOD Hes DITO irbotie wel Tne vepetulion bub the implicutions foe many other portant Organisms, especially invertehrites. wre largely unknown (Bradstock ey af, LYYS: Friend & Williams 1896). The few Australian Studies of the effects of fire on invertebrates have coneentrated on soil and littsmdwelling organisms (ee. Campbell & ‘Tanton TY§ 1; Neumanis & Tolhurst 991; York 1994) or have sampled a wider invertebrate ussemblige usm only pitfall traps: (ag. Prend & Williuns 1006), Arboredl insects probubly survive fites less well tha epigcie and Hypogeie speeies (Whelar 1995) und, provideck sampling biases can be addressed. should be good candidates For study ine the cflects of fire on dnveriehrate populivons. tn the presehe sty we examine the effects of fire history on cndemic Agsttahan scale aiseets (denmiplera Coceoiden) that five only fa the caopy oF cuealy pts Seale insects of the genus Apiemearpha Rilbsaumen (hriococeidie) Tive within galls That they induce on iheir eucalypt hosts (Gullan 1984acb) | tn Apioniuphe. couch adult female resides ia large. oflen symmetrical und woody gall usually on the stem but sometimes on leaves, buds or traits of the host cuealypt (Guatht 2984), These are easily recognised as Cocco walls because there ts a small apical orifice through whieh the female can chiminidte her waste honeydew and also mate with the rate Her offspring or firstinstur nymphs. eulled crawlers. inake thei exit from) the muternal gall dhrough: this sine apeniny and then disperse fo inithite new eulls {3K PUCOLLLAN. I SOURANSTON & LG, COOK on sullble foliage, The sadls of tales ave much smiuler than (hose of the females, rarely more than une centimetre Tong. and are tobolae with an apioul orifice unl sometimes un Gulwardly-dirceted Tange ab the apex, The shape of the gall of the female ts pecullir to lhe Apiemoreia species thal induces revardless ofthe identity of the host eucalypt, Most Aplomorpha specics show sane degree ol speemieny Wea restriged range Of Encalyptis species (Galhin ital Apiamorphe hus 39 deserthed species (Cjullan 1984a; Gullan & Jones 1989), each ol whieh Haims Chavaeterisnic., sexudly-dimorphie galls, Generally. Apdomarphe can be identitied Lo species Jevel in the field, even by Hotespeehilists with minal Grass, Seale inseels are otlen claimed lo have low yagility (lor reviewosee Hinks & Denno M99) heeause the wingless adult females usually spend) thei entire lives ona side host phint and oviposit thece, adult males are short-lived and weuk-fMving. and all dispersal is duce to the movement of the vrawlers, Which dsually remain on their natal host (conditions une Havourible (Gireathead (990: Nested eral, 1995), Ih Apiomer phe. our held observations ih sclerophyt wowthinds have sugeested (bal there is lower diversity and abuidanee of galls io aes sabypeetcd te Hequent or severe burning, This reduetion seers Linrehited to the suitability OF post-fire enealypts for gall development, since the epicormic Push oF foliage that follows a fire resembles the preferred plant material utilised by Apromenphe in leony, inburnt areas. His more Tikely that afer a fire kills the galls and their occupants, i takes time for reenlenisation by erawlers fe oecur and if fires are irequent, OF potential sources of colonists ate distant, locul populitions Wiiv Wet resestiblish. This hypothesis. renwuis speculitive at the ubscnee of quuntilication of ny differences in species diversily wid ubundidmee between bunt wad) lange cnibernt ureus, We fovestinated re-colonsation alter lire hy surveying the species rivhness and uhundunee ol Apromaepha it rclatien to both the fire histary ane the species OF host eucalypt in mallee vewetution in South Australia, Thos study was undertaken tn mmilee for several reasons. Firstly. mutlee is a) typicully Australian vegetation that has beer ti seclous deeline for the ust 150 yeurs threugh kd elewranee und other forms of degradation (Land Conservation Council 1987: Cheul JO8G Harris 1990) Secondly. it his heen hypothesised that Mallee plant aud) animal commniies are muintiined by episodic fire (Noble 1982. 1989) hut no research has been done te investigate how efecuvely any phytophapous msecey peestablish their populations in matice aller fire. Thitdhy, itis easier to count the galls on mallee ehealypts Wan on the Giller forest ang woodkind species. Seale disects of Apiomearpla are good canchdites for studying the effeets of wihtfire because, unlike more mobile wsects (Whelan ered 1980). they are fob ubleold move From the thair host plants to aver the Tames and. although the thigk Wall Ob the galls Hits been suggested loa be an adaphition for fine protection (Koteju L986). the bit intensity of mullee wildlires. usially hills the overstorey lobiage (Bradsivek 1990), Ty Gontrist ta the wellek iow) banes of pilhalling (rapping and other methods ol sampling soil-or fitter Hain (see Whelan 1995). our subvey method lor prone potentially assesses Ihe tanh nuinher oF galls on each tree Further Advan tiges of the Use ab Aponte galls bs that their ubundinee vires Tile wallh season of survey singe (he Temules of most speaies probably live tor much longer Thun a year (L.A. Cook unpub.) une walls can remain on the trees for several years: (ue the death of the occupant This providing a recon ol the presence of (he species ul a site. Lastly. in order to elucidale | eausul relationships berween iwerlvhrate abundanee patterns and: fine, scleetoel invertebrate groups need to be examined at a tines level OF taxonomic resolution (han the order oy family level that ts used inmost studies (Pricid: & Williaits 1996). Foom this perspective. apioneeplre is a ideal study genus because gulls can he identified readily to speeies level, ever ny We Geld, Methods The work Wis curled out ih Danggal) Conservation Park whieh wits estiblished tn 1976, heciine Australia's Mist Biosphere Reserve in (077 andis now parcul ihe Bookmark Biosphere Reserve. This quarter of 4 million heewire reserve js about 90 kin north oF Renmark in South Australia and tes in the northern halPor the Murry biasin, adjacent (or the New South Wales border (Pig, UT), tb inelides hol arid ind mallee lam systems and allows aevess (4 some speclicnlar old-growih wallee. OF purtientar sivnifieance ty our study is the doeumented (re History of Danggali Conservation Park (rom National Parks and Wildlife Service, Seutlh Aust. File Tis Hol been tiscd as a puapennent tool th the inaintenwnee of tis mallee for egoservation: recorded wildlires either have heen mulurally-ovcucring, Tollowing lightning strikes. or the result oF burma acuents, Many areas have niu evidence of burning for u considerable perid perhaps for over a century, Or ul least noLsinge either JOP? or [951 when extensive wildtifes burn muel ol the region, (loweyer, of special releyange loo the prescnl stinty isu inajor lighining-induced tire thin burnta huge eential area of the reserve i December GALL-INDUCING INSECTS AND MALLEE FIRE HISTORY \39 no SECU HO ce eee eer) PU te “ we ~ os "Morgan Vale" Ruwis eT Mt Morgan AUSTRALIA WALES UU PUREVINUULEEUEADO LAAN Hain ne SOUTH SOUTH NEW HUTTE “Cangpus" Homesicad FT Re 7h fn, “Aypurna th, 4 “eg Homestead mn Park boundary — Road or track @ Landmark Bayi c0d 10 OHH UTU OHNO EELAAU PRAHA LANA nan @ Study site Extent of 1974 fire i Extent of 1979 fire . “mn 0 5 1a 20 SEE | fea Extent of 1985 Fire Fig. 1. Map of Danggali Conservation Park, South Australia, showing the extent of recent wildfires and the locations of the 14 study sites where galls of Apiomorpha were surveyed, Inset shows location of Danggali Conservation Park. (Figure based on map and aerial photographs from the National Parks and Wildlife Service. South Australian. eH) 1985 (Fig. 1). Since then, (he burnt eucalypts live regeneruled from (heir mates Hanotubers, although the dead stags of past sdbstuntial limbs still protrude from the now-lMourishing mallee teerowth, We made conparisons belween the Apienerplee galls associited with eucalyprs in the old-prowth mallee (“long-unburnit’ sites) and in the mallee that had regenerated after the L985 fire Churn” sites). All Fieldwork Wits conducted in April 1996. Fourteen sites were chosen (ig, Ty seven iy cach treatment type (burnt versus long-unburnt), with the site co- ordinales as given in Table 1. All enealypts at the burnt sies had been burnt completely inthe 185 fire tind sites 6-9 were completely surrounded by post- fire reveneraled mallee. We only used burnt sites 1 the eentral aren uf the reserve hecause Wwe were vertuin dhit vegetation in this area had been vompletely destroyed in 1985, whereas the Veeeuilion around “Tipperary Dany and “Morgan fe Ruts and in the owe smaller areas also burnt i F985 on the periphery of the reserve (Fis, 1) appeared 16 hive heen wore patehily burnt Five hiirntand five longeunburnt sites had an understorey Wilh porcupine gruss, Trindia irrirany Re Be. (Ciramineae), sometimes ay the dominant ground- laver plant. Che ater four sites either had reasonably hire Shon Ora mixtire of shrub species, The jiallee cuculypl species al the study sites were Hucelyplas incraysata Sieber ex DC. (syn. 1. canine Pm Muell, & Bel ex fo Muell.), & didmayea Cun, ex Oxley, 2 gracilin T Moell,, Bo aleosa P Muell, ex Mig, dnd E. seciadéiy b Muell, ex Mig, Ideniilication Wis Indde using Costermans (1994), except tat the name E. (erassetr (syn. 2. cayiefe) is retained (MLL. Brooker pers. comm), The prapertions: of each species varied aecording to locality but usually three ol the five mallee enealypis were present at ull sites (Table 1). We chose sites i dong-unburnit or post fire regenerating patches to provide 10 multi-stemmed 12, PT, GULLAN. BS. CRANSTON & LC. COOK inallee eucalypts. of between 3 and 6 min height, per site These 10 trees were chosen us representative al the proportions of caul speevies inthe local ares. "he height was inposed by the need to inspect (he total foliage hoth front the ground and from a 3 in ladder pluced in the centre of the tree, For cach tree we vounted the number of walls conlaining live and dead Apiomerpha females, Tie surveyed galls varied in size fron a few iim Cyouns or aborted) to more (han 4 em lone. und in shape from eylindrical and bud-like to urn-shaped or ovend, Galls were revorded us vootaining dew females i they were old and brown or showed signs of atuck by predators Or parasitoids, Usually, the presence of u living coecoid Was Confirmed by (he presence of White powdery wax atthe gull orifice, Only voucher material and galls of uncertain identity were vollected. so the survey Wos ?elatively non destroetive, Voucher specimens. of galls: and slide mMotinted sects ol Apiomarplia have been deposited ithe Austratiain National Insect Cullection (ANIC), CSIRO. Canberra. All ditt analyses pertain to galls contdinine five plus dead female jaseets, unless otherwise stuled The gills of dead inseels were jneluded in eounts becuuse any sucvesstul initiadon ol uw gall was evidenve thar josect had reached the site and that the Wee Wis at Suitable hast, Host speciieiy within Apioniorpha was examined by calenliting the percentige of (he surveyed teees Of euch eucalypt Species ial supported galls al femules of quel Apromorplia species. The response of Apromerpli Species 10 fire Was evaluated using one way analysis of varianee (ANOVA) Lo compare burnt and Jong Unburnt sites in terms of the total numbers of palls of females (of all Species summed), total numbers of gulls af the seven least common species (Le. walls of AL mallecacote Gullan and Ay ovientaides (Vepper exchided), and lolal Apiomerpha species found aut PANE 1 Site localities (omm GPS reclining) with qire histany flan unit daa) perses Dit in LORS CBE) aed Hee umes af tives af each species sampled ar euel site. Ko dy — E. dumusu. E. go> bk. gracilis, Boi = BE, inerussata, Ban 9K, Oleost, Bos, = D. soetalis, Nawihery en the fant of each spectes ealunin “hr tor the fatal Fniien Of ees warwyveu fie (het specter, Site History Latinde Longitide oad, Eee Mai, E. 0. EL | tu 33°17 13" moras’ a " a it] 4 2 lai ABT" 14oras'uy" 4 [ {) 4 2. x 6 a ee Issa" {) 2 () Oo 2 + ob 17" 140) a5 08" a 3 oi 4 it} 5S leu Aa Luray [4() 38°27" 6 it] 0 () + o b 33°16" lath ya ye 3 | I | 4 7 4b AM Gd" 140 43'08" 6 u (1 () + yooh FAA Ay" Ltd's” 4 | 0 {) 3 ae 33 lesa" 1g s7” | 7 {) {) + Wh dew 38°34" 30" ln arsy” i) 5 \) ‘) 5 Woodew 34°05'02" l4or42'40" 3 + ‘) i) 4 12 lr 33 17! 0" 4040's" a a 4 if] 2 Mi lu 43074" LAPS! 4 J i 2 ) I4 ob 33° 19/20" Ly 35" ay" h 2 0 0 2 q 5 7 a w GALI-INDUCING INSECTS AND MALLEE FIRE HISTORY M41 each site (14 units total), Analysis was carried out at the site level, not the tree level. because there were Iwo or more different (ree species per site and many trees had no oor few galls. Two species. A. malleedcola aod AL eviceloides. were common enough fo examine their mdividual responses to fire al the site level using ANOVA but the data were transformed [ly (x+1)] fo correct for skewness, The responses of AL malleeacela and A, ovicoloides to fire also were evaluated using individual trees as the Units Of analysis. In this cuse. binary presences and absences were analysed using Chi Square tests because gully were not abundant or widespread enough to satisfy underlying statistieal assumptions wt this level, ATL amilyses were carried oul using IMP™ (SAS Institute Ine., © 1989-91), Results Apiomorphia species recorded A total of nine species of Apiemerila was recorded fron) the sites surveyed, ‘These were: A. calyoing (Tepper) A. densispinese Gullan, A. karsehi Ritbsaumnen. A. mMalleeavolu, Ay miinita matleensis Gullun, A. avieolaides. Al regilaris (Tepper). A. strombylose (Tepper) and AL arnedis (Tepper). All of these species have been collected previously from mallee vegetation in southern Australia (Tepper 1893: Gullan 198). Host-plant specificiy af Apiomorphit One species, A. densixpinesa, was recorded fron just three sites and solely on E. dsmose but only six gulls were found and none of these contained a live coccoid. Six other of the nine species ol Apiomarplia showed some degree of host-plant specificity (Fig. 2), Galls of A. cahycina were found on four of the seven surveyed trees of BL prerassete as well as ou two other cucalypt species, Galls oF AL Kerselii alse occurred on three eucalypt species. whereas A. munita malleensis, A. regularis, A. strombylosa and Ao urnalis each were reeorded from only two cucalypt species, However, 13 of the 14 trees that supported galls of A, drnalix belonged to E. gracilis and nine of the 10 trees with galls of A, strembylove were EL socidlis, Only one species. AL ovicoloides, was recorded on all five species of eucalypl it was the commonest species, occurring On 52 of the 140 surveyed irees, The next mast common species wits A. malleeacola which was found on four species ol eucalypt and on 37 of the 140 surveyed trees. Species richness and abundance of Apiomorpha in relation ta fire history All nine Apiomorphe species were found at both burnt and Jong-unburnt sites, Burnt and long-unburnt sites did not differ significantly in the number of Apiomorpha species recorded on survey trees (F, 9=3.57. p=0.08) (Table 2), ANOVA of the total Vania 2. Meany per site # ESD, orwith range in parentheses, and significance of differences due to fire hisrary tiediversity of Apomorpha species, number af galls ef-all Apiamorpha species, monber of galls of A. malleeacols, mune of galls ofA, ovicoloides aif number of gally of all species excluding \. malleeacola ave A, ovicoloides, (Foy A. mallecacola ful Av avicoloides, means and rnges ure from the caw deta bur F and yp values cre from transformed cette, as indicated by!) Long-upburnt Burnt Fis Value p Value i (sites) 7 Mean number of Apiomerphe 5.34.1 Species per site Mean number of Apiamrearpeltt gully per Site 374495 Mean miunber of A mullecacula galls per site 9,4 (2-12) Mean number of Av nvicolotdes falls per site 13.3 (6-30) Mean number of Apiamoarpha gills per site with A, wielecacald and AL eniealatides excluded 16,6 £544 12, 7.6 (0-21) BAH) 40414 AST 0.08 24.34 148 3.96 07 40! 0.54! av OU! Id. ().34 (3a “o Of surveyed trees with galls present °s of surveyed trees with galls present “e of Surveyed trees wilh galls present % of surveyed trees with galls present 60 50 P. J. GULLAN, P. S. CRANSTON & L. G. COOK A, calycina Ea€&9 Fi Eo, &s, Eucalyptus species A. malleeacola Ee, Eg El BG ES Eucalyptus species A, ovicoloides E.d Eg Ej} Eo Es. Eucalyptus species A, strombylosa Ed Eg Ei Eo Es. Eucalyptus species So of surveyed Irees wilh galls present “s al surveyed trees with galls present % of surveyed lrees with galls present % of surveyed trees with galls present A. karschi Ed Eg El Eo, Es, Eucalyptus species A, munita mallensis Ed &g9 Ei Eo Es. Eucalyptus species A. reguilaris Ed Eg EF Eo. Es, Eucalyptus species A, umaills Ed &.g Ei E.0. Es. Eucalyptus species OALLSINDUCING INSECTS AND MALLE FIRE HISTORY | b5 nuibers of galls of females (live anel Jeu) recorded from) cach site todiedied that the fire history of the afles did fot affect ull abundinees af Apiomerplng specigs Ub) =3,.96, pe0.07) (lable 2), Burnt and fons unburolsiles did not dilferin the percentage of live fo dead Apemnarplin 230) oF galls at lone- unhurt sites and 24% OF gulls i burnt sites were estimated to contain live coveaids. EXcept fur A, mofleewcala and A, oavienlondes lhere were insulficien| daly lyr siarstieal analyses bused on individuail Species. although for the therd most ubuman) species. AL nrauliy there were compuruble populations at both burnt and lone -unbuy mt sites, bor Al maleeucol, ANQVA of the trinsformed ubundunes [I (xb bd} ah cach site indivated that numbers ab galls did not differ benween burnk and lutig-ubburnt sues (FP) =0.40, p=). 34) (Table 2), Chi Squive atilyscs OF presenee-ubsence dale for A, malleracela on individial trees alse indicated that Tire History did notafieer the likelihood of finding A, mallecaola YaMls QA) \,=0.04, p=0.84), with walls being present ai 27% ob all (ees sanpled in dong: Gabum) sites awd 260% OF all trees sumpled al burnt Stes. TP trees oF EL irerassara. on whiel A, maflecacale was never Tound, were exeluded, the resuills renmained very similar Cg?) =U,214, prst,04: gulls presenton 30% ob lone unburnt trees and 26% 4) burol (rees), Burnt ind long-unburnt sites differed onty slightly in the percenniwe of dive to dead A, matlecacolen Ad OF galls ut king-unburnt sites nid S90. co) 4ills ah burnt Sites were estimated to contain live coccoids, Along the burnt sites surveyed there wits no evidence that galls of A, medleedonle were mone abundiunt we sites close to (range 1- Lb wah mean OfG-7 gulls per site), compared with distut from (range M21 with mean of 4.43 gulls per siter, long Unburnt aresis, Ii contrast, for A, orteulerdes, ANOVA of the transformed abundance [h, (k+1)) al eich site indicuwed (hit numbers ef galls were significantly lower at burge sites (PF) = 114A 7, ps0,01) (Tuble 2), Burnt and long-unburnt sites differed only slightly in the pereentive of live to dead. avicalordess (O06 OF gulls ut Jong-ontuent sites and 1% oF gallsat burnt Sites Were estitmied Lo contin live coecoids. Chi Square amilyses ol presence-ubsence dati for 21. ovicoleides on individual trees also tdjcateal that fire Instory ullected the likelihood of finding A ericolofiles, gully Qe) .=9.04, p=0.003), with gatks being present on 44% of all trees sampled in long- Unhurnt sites, hur on only 24%) of hurnt trees. Among (he burnt sites surveyed, walls of A. auiediaidey were Slightly nore suuree at sites FOU-200 ni trom unburn Veachition (range 0-3 with mean ol LO gulls per sites than atosites several kin distant trom lone-unburnt areas (rime 1-8 with mew of 4.5 galls per site). IWohoth A. mallemeeda and Al ovicaliides were excloded tron the anulysis GF total numbers ol galls, ANOVA of the gall abundance at cach site indicated that numbers uf galls of the other species combined did net differ hetween burtand long-unburnt sites (FP) =0.39) pets) (Tuhle 2). Martealiny fuctors The orttaal occupants of imny of the Apioanaples gulls (hal we recorded during our survey erther had been Killed by parusituids. probubly wasps. vl removed by preditars, prohibly cockatoos aie a Aid ! : > wah | J ¢ i nly i Thy ear 3 ' iy! i , | = } i fA * oy a / ' a oo By toi ood \ Tr yyy Nou Ome ior Fab Mer Ag Muy dun Ju) Ang Sep Onl Now Gb von Fel Mar Anh May. die of ' a5 ' ab Man) Yew Fig. 7, Dissolved ammonia. Banrock and Loch Luna, November 1994-June 1946, = kook Lunn © teantinil ah CAAA leuid coded pte nitrate (tral! + ao pire © oe Coe an 17 ay he fv Hen dare Fete dan Fee mar App Muy: dun a oa edie Monin Year Rig &. Dissolved reactive nitrate, Banrock and Loch Luni, November 1994-June 1996, PHYSICO-CHEMICAL MONITORING OF TWO WETLANDS 153 * Loch Luna Acanroch | ail i | ae Gow." Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jur 95 ith Nov bec Jan Feb or) | i Month - Year Fig. 9. Total phosphate. Banrock and Loch Luna, November 1994-June 1996. \ | «| ||, : ih * || a ara | a w | } oy + | | | Lf fs | iN a pene ytel RR eh eo ating” Tasso ae | is 7 pened T Nov Dec Jan Feb Mar Apr May Jun JU! Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun a | a5 | 96 Month ~ Year Fig. 10. Cyanobacteria (blue green algae). Banrock and och Luna, November 1994-June 1996. g. L three Anabaena species were predominant through the rise and subsequent decline of high cell multiplication (incipient blue green algal “bloom”). During November 1994 the number of cyanobacterial cells reached a low of 41 cells ml! but after 29 November their numbers increased rapidly to peak at 23,700 cells mF! on 27 December 1994 before declining to 97 cells ml! two months later (Fig. 10, Table 1). Anabaena coiled species was predominant until 3 January 1995 after which Anabaena circinalis displaced it until mid - February. The coiled species again became predominant until June 1995 when all three Anabaena species were present in low numbers until late November 1995 when multiplication of Anabaena coiled species began again. This species peaked at 2530 cells mb! in early February and a month later cell numbers fell to 63 cells ml! (Table 1). Cell numbers rose slightly to 444 cells ml! on 19 March 1996 but no cells were detected on 9 June 1996, Other cyanobacterial species identified in the water samples, although occurring only in low numbers, were Anabaenopsis elenkini (6 December 1994 - 20 February 1995), Aphanizomenon sp. (22 November 1994 - 13 June 1995, 23 January - 19 March 1996), Oscillatoria sp. (3 November 1994 - 18 July 1995, 9 January - 19 March 1996). Cylindrospermopsis raciborski, Planktothrix spp., Arthrospira_ spp.. Microcystis aeruginosa and Pseudoanabaena spp. were identified from time to time. Discussion Surface water temperatures, pH, turbidity and conductivity levels followed similar trends in both Banrock and Loch Luna wetlands and were comparable with the values recorded between 1990 - 1993 for the eight floodplain wetlands of the River Murray in South Australia by Suter ef a/. (1993). River Murray turbidity values were highest (70 NT units) on 14 May 1996 and lowest (20 NT units) on 26 December 1995. Mixed waters of River Murray and Darling River (Lock 3 Pool) registered 48 NT units on 9 June 1996. In the 10-year period 1978-88 Lock 3 Pool water averaged 60 NT units (Mackay & Eastburn 1990), The high conductivity value of 1898 EC units in Banrock on 3 November 1994 was caused by a blocked inlet pipe into Banrock; with the clearing of the blockage conductivity values in one week dropped to 1507 EC units. Turbidity in Banrock increased from 104 to 155 NT units between 5 and & November 1994 due to turbulence from the rush of water following clearing of the blockage. Seven wecks for conductivity values and five weeks for turbidity values were required to reach equivalence with Loch Luna values. In Loch Luna conductivity levels rose from 327 to 2720 EC units on 3 October 1995 after spring rains and strong runoff. At Barmera, 13 mm of rain fell on 31 August, 9.6 mm on 5 September, 9.8 mm on 25 September, 10 mm on 3 October and 53 mm on 23 October 1995. Seiche effects in Lake Bonney and added runoffs caused the high conductivity water from Lake Bonney to flow through Chambers Creek into Loch Luna wetland. The outward movement of the high conductivity water (2970 EC units) from Lake Bonney was traced from data recorded on 6 October 1995 at position POISI7 in Nockburra Creek, a tributary of Chambers Creek, Six weeks elapsed before the high conductivity water from Lake Bonney had been diluted to 721 EC units (14 November 1995). The range of DRN concentrations in Banrock was 0 - 7.9 mg}! and in Loch Luna 0 - 7.48 mgt!. The high registrations occurred mainly after heavy rains but there are also nitrate contributions from time to Ja4 ‘AM OLSEN tite Tony water tron the River Murry. and fron avriculliral dritos, town effluents amt sewage Jischurzes us well as Jovaleed autolvie breakdown of nitragen - fixing ble green alee (Anahecine spp. There isin tnknewr DRN puteent lapae fron huge Hoeks oF pelicans (> LOOQO birds) swans. vormorants and ducks and lesser nunibers of ather water birds resident in Baoraek and Laeh Lane Wetlands und on the banks of the River Murty, Nitvates are also prodiwed by lightaine (Sith 1996), The highest TP level (0.95 nib ) recorded in Lach Luna veeurred on 24 Outober 1995 the day aller 53 ni at yar felhat Barrera and Sk mm at Banrovk Stabe (Co Lb, Rohliehl pers. comm. 1995), Such heavy rain aml consequent runol cause Potton Jisturhinces in shallow wethinds whieh redistribute Jisselyed unaite flosphorus compounds und iWortatie phospheris bound to suspended ar Jistivbed bolo) organic putheutates: 1) the wiler coh, Brees ef a, (1985) have alse drawn dllenion to the complexity of chemival relauiashipes withon werkinds aml the elects of weather fietors. SUCH os Wands and feinpemdure, On Water chemistry No blooltis of eyanobacteria were observed in Banrock wetland, “The multiplication of eyaiobueterial wells je Baaroek during December IWS wis hulled by the inercased witer inflow ollowiig removal of the blockage Hi the titer pipe. Whe eyunobucleriie were (Mlushed oul preventing any further development ofa eyarmobacterial bloone in Birroch wetland that scar IW Loch Lune evinobacterial cells reached a utr of 23,700 cells tal! oon 27) December helore declining wozere by 1+ Mareh 1905, After the oollapse of the cyanobacterial populariain ia Lach Lunn December (YO. a small bub visible blue areen abil bloam developed dowtstreaaiy i Februaey vind Mareh 1995 along the eastern hank ol Ihe River Murray wujacent te TLoek 3 weir Nis thely (hil this bloor had gis genesis in the November - December 1004 cell mulfiplionion te loch) Land Upstreain und pn the same side af the River Murry. Small blue ereen olval hloors Had ocetrred mH this sales THcadon Lr previous yes (hock 3 staft pers, cop, LOO8 |), Bowling (1994) repurted the occurrenee ane possille eases OF a severe Anefaen etre inulin blow tit Lake Corellign NSW oir fate 190-0 | wher cell numbers exceeded (004100 vellsiml! The physicu-chenneal levels i the lake fi 1990) had nines close to the 1994-4 values i Loeb Lun far Wailer lemperiures, pil, turbidity and conduetiviry hil lower Wasi in concentrations af TP and DRS fan were Toupd dor Loch Lane. Bowling (1o94) capressedt The view suboul the Like Cirgelleo bloent hat “wiilnueh severe! underlying eanses ol this bloom are probable, the elevuled nuirient concentrations, espechilly of tatal phosphorus. were major Taeturs that contributed ta an He drew aiention to the Taek that mase physicoeehemical stidics af cyanobacteria blooms were slanted ufler the blooms had oceurred. To this study of the physive-chemeal properties of The two Wwetlinds a seach far blot - Torning tosie cyanobacteria species Was started betere any cell MuUplication Hac commenced jimh a seasonal pullern is) described. Cynnobacterid: celhy may romain dormant Ti eolk waters cand grow: best ut wuller lerpperitures excecding 13°C. with Gptinnl erawth rales a) 25° Cor higher (kobarts & Zolury JOSS), The efleek of wiler lempernitures on eyunobuclerial cell nurmbers in Loch Lune is shove wilh dita front Table} which shaw that Blears only occurred between November une Pebruary, Nurmhers Ot Aikimeecha spp. tose suddenly fran cero on $5 November 1994 (20.1 ©) toy 23,700 nmil-l on 27 Decensher 1904 424.2" OC) and thea began subsidioy, ever Uneil the wiatttr teniperauores were high, perhaps because of OxHUstian OF WUUPENES. The rise (und fall fa cell numbers iy aso be rehiled to the concentrations of the two mitrieitts, DRN aml TP recorded dutiog the vrowth and decline ol the }994-5 vind 1995-6 blue ween ulgal outbreaks Mable 1 However, sivee the concennyiot af dissolycd reactive phosphorus is nol known, (hese relationships must be teeatod with caurion, There were Jour ceeusians (20 December 199d, 4 and 31 Jani 1995 and F& April 1995) when TP concentrations in Loch Luniowere at or above 0.46 mel! oun DRN was present even tn law cnneenreations (Pr - 2.2 mel. Aten vaeh ol these events there was a vise in the nunbers ar cyanobacterial cells. hy October 1995 (mean 16,9 °C) (here were four PP coneen(nitions between (44 and 1.95 ggl! hur DRN was low or absent and ng vel mulrephiearion developed, A patrern rehued ta totrivat availability anu lomperniture Gin be seen in curly 1996, Alter an Wnerewise ty 3190 cells on 23 tunuary 1996 the cell Hmbers began to deelhe possibly die te very lov eonecntrations Of DRS aed Mretuiting values of TP Wothe walter colin, Phe eyanobieterie populiion reughed vero on 14 May (Y9G. "The mean wattere lemperiture onthad chite was 14 oC whieh is dear the MIT MAT Temperate range Tor eeowel al many free living Gyanubucterin species, Prom (he data obrincds i this study ibis stimested Nhat outhreaks af cyanobacterial bloenis id jet secur in the supers of [9945 gad [995-6 beeuuse {here were muadequitle concentrtions of LP and DRN ni (he wiler columim during the period of lavourable growth for eyanobacteria Dane fram this study indivare that LP coneentradin abowe O.S6 gt! PHYSICO-CHEMICAL MONITORING OF TWO WETLANDS 155 and DRN concentration at or above 4.0 mgt! im the welland may provide for continuous growth in the Anabaena species. Acknowledgments This study was funded by the Murray - Darling Commission through the Natural Resources Management Strategy, Project 53128. 1 am extremely grateful for the assistance [rom the following contractors: Mr M, C. Schultz, Glossop High School for the physico-chemical analyses, Mr P. Christy and staff Australian Centre for Waiter Quality Research, PMB Salsbury SA for identification and enumeration of cyanobacterial species and Messrs R, [. Gropler and J, A, Pillar for their regular and supportive help with Held sampling from a powered dinghy, My thanks to Mrs B. Page. Tilley Murphy and Hughes, for accounting assistance and payment of accounts, Mrs J. A. Rourke, Adelaide for preparing the manuscript and Mr G, Wright, SARDI West Beach for the preparation of the figures. Mr B. T, and Mrs E. L. Engel former owners of Banrock Station and BRL Hardy Ltd, the present owner ire thanked for allowing access to the wetland. Mr C. L. Rohrlach, Manager Banrock Station and his staff gave valuable assisiance throughout the study. The author thanks the editor, Mrs J. Bird and the two unnamed referees for their constructive comments On an earlier dratt, References AUSTRALIAN NaALURE CONSERVATION AgENCY (1996) “A dictionary of important wetlands in Australia’ 2nd edo (ANCA, Canberra). Bowe ina, L. (1994) Occurrence and possible causes of a severe cyanobacterial bloom in Luke Cargellico. New South Wales, Aust. J. Mar Fresh. Res, 45, 737-745. Brows, S. V., MAuer. M.'T. & Carpenter, S. M. (1985) Limnological studies of waterfowl habitat in south- western N.S-W. 1. Water chemistry. /hrel. 36, 59-67. Goonan, P.M. Ber. J, A. Tompson, T. B. & Surer, P. J.(1992) Wetlands of the River Murray floodplain, South Australia, | Preliminary survey of the biota and physico- chemistry of ten wetlands from Chowilla to Mannum, Trans, R. Soe, 8. Aust. 116, 81-94. Hach (1984) “Instrunient Mannal” Ist edn (Hach, Colorado), (1902) “Water analysis handbook” 2nd edn (Hach, Colorada). Haiso (1990) “Enumeration of algae. estimation of cell volume. and use in bioussays 1990" In the series; Methods for the examination of waters and associated materials (HMSO, London). Mackay, N, & EAsTBuRN, D. (Eds) (1990) “The Murray” (Murray - Darling Basin Commission, Canberra). Roparts. R. D, & Zamsrty, T. (1987) Temperature effects on photosynthetic capacity. respiration and growth rates of bloom-forming cyanobacteria. NZJ. Mar Freshw, Res 21, 391-399. Sarre. R.L. (1996) “Ecology and Field Biology” Sth edn (Harper Collins College Publishers Ine., New York), Sut. P. J... Goowas, P. M.. Beer, J. A. & THomeson. T. B- (1993) “A biological and physico-chemical monitoring study of wetlands from the River Murray floodplain in South Australia” Murray - Darling Natural Resources Management Strategy Wetlands Management Monitoring Program. Project No. S002 Final Report. Australian Centre for Water Quality Research Report No, 7/93. ASPHONDYLIA ANTHOCERCIDIS, A NEW SPECIES OF CECIDOMYTIDAE (DIPTERA) INDUCING FRUIT GALLS ON ANTHOCERCIS LITTOREA (SOLANACEAE) IN WESTERN AUSTRALIA By PETER KOLESIK*, REBECCA WHITTEMORE & HELEN M. STACET Summary Kolesik, P., Whittemore, R., & Stace, H. M. (1997) Asphondylia anthocercidis, a new species of Cecidomyiidae (Diptera) inducing fruit galls on Anthocercis littorea (Solanaceae) in Western Australia. Trans. R. Soc. S. Aust. 121(4), 157-161, 28 November, 1997. The fruit galls on the Western Australian yellow tailflower, Anthocercis littorea (Solanaceae), reduce the reproductive potential of this plant, but their causative agent has, until now, been unknown. Our research has shown that a new gall midge species, Asphondylia anthocercidis, induces these galls. The larva, pupa, male and female of the new species are described and illustrated. Key Words: Diptera, Cecidomyiidae, Asphondylia anthocercidis, Anthocercis littorea, Western Australia. Line tans ap the Ravel Seerene ap. Aust, (F907, Pad, TS7-1Od, ASPHONDYLIA ANTHOCERCIDIS, A NEW SPECIES OF CECIDOMYIIDAE (DIPTERA) INDUCING FRUIT GALLS ON ANTHOCERCIS LITTOREA (SOLANACEAE) IN WESTERN AUSTRALIA by Prerer KoLesin®, REBECCA WiirrreMoRb? & Helen M. Sracet Summary Rolbsnk. Py Wireromort. Ro & Sixvck Te Me (1Q97) Ayyhondvlia qaimthevercilis, a new species of Cecidomyndae (Diprera) ihducing (rait galls on AnMiocerciy fitferca (Solunaceue) ty Wester Australi, Zens, RK. Soe, So Aust. D2, US T-161, 28 November, 1907. The (unt walls onthe Western Australian yellow tatlower, Admocercis inored Labill. (Sokindeeae). reditee the reproductive porential rf this plant, bub their etusative agent his uatiT now, been inkrown. Ourresedrelt his shown that ai new gill mide species. Asplondyiitmrhocerchdis. muuces these walls The farva, pupa, Male aid lemuleol the new species are deseribed and dlestrated. Key Won: Diptera, Cecidomyridae. Asyplomivia aamoceraidis, Anthacercis litte. Western Australi I b / : Iniroduction The gill midge fauna of Western Auswahl is poorly known, with only (We species having been desoribed previously (Gagné 1989), One of them, Tponnit barnemissca’ Colless, is a species which presumably feeds on fingi growing inthe seiland jn jeal Witter (Colless 1965), The biology of the second species, Eveinelicornia austrdfasian Felt is unknown (Pelt 1915). although (his species is tikely to be a phint leeder considering that its congener £ Malweskii Kolesik, causes walls on Eirealyprs faycieulosaan South Australia (Rolesik }99Sa), The yellow tailllower Aathecercis fittorea Labill (Sohkinaecae). a shrub which grows lo 3 im. 1s endemic 16 the south-west coust of Western Austialia, primarily on calcareous sands in disturbed habits such as recently burnt areas, roadsides. Fire breaks ind cleared Jots (Purdie ef al. 1982 Whittemore!) The froi valls on A. /itere have been known foe some time (Purdie ey af 1982 ) bul their causative agent has remained unknown, I uly 1996, one of us OR. W.) collected froit galls from A, dittorce containing larvae and pupae, from which adults were reared, The salleinducer proved to be a new speeies of adh midge whieh is described below. The development of the galls and their impact on the reproduction of A, fittarese are deseribed by Whitlemore!. The genus Ayplandyiie in the context of this paper DepAP enn ob Phen clita. Vili Tite ane Erertalopy, Miwulty oe Avrioulluni ond Satin Resodree Scieheds. The Lineversity ob Adelie PMB Glen Ounomd S Suet Sherk Departiaent ot Beraiy. Greversity al Western Austitlie Neotunds W Aust 6407 Warr sokn RC hQ86) Aspects of Ihe dnscetendiowd fri) galls and repraddetve: biahhgy al anoeoreds fiterec: (Sulimacaae BSe (Hone) thesis. bfiversity of Western Aastratia (anus) is defined by Kolesik (1997) "The new species ty lo he attributed to PAK. Material and Methods Stems ol Aniiocerciy littarea bearing fruit galls were collected al Hillarys, about 20 km nortiecast ot Perth, on 23.971,1996, A small number of galls was dissected and the larvae und pupae preserved in 70% clhanol, Larvae and pupae retained within galls were reared lo adults on stems avhieh were kept in plastic viitls. Larvae pupated within (he galls, Emerged gall midges were preserved tovether with pupal skins in 70% ethanol, Canada balsam mounts of the type series tur microscopic exalnination were prepared according to the technique outlined by Kolesik (19S). All measurements refer Lo the type series, The type specimens and other material retained in 70% -elniel are deposited in the South Australian Museum. Adeéluide (SAMA) and Australian Nutional Inseet Collection, Canberra (ANIC). Asphondylia anthocercidis sp. nay, (PIGS 1-15) Haloryper 2. Hillarys, Western Australia [41 48" S, 115°45' Ej, emerged 28.vir L996. Ro Whitlemore. reared from larva rom fruit gall on Anidiecerers littorea Labi. gull colleated 29 vii L996, 121435 |SAMA}. Paraivypes: 2 8S. 3 99 5 pupal skins (SAMA, 3 29.3 F F.4 pupal shins [ANIC], all same date but emerged 28-30.v11, 1996; 2 larvae [SAMA]. 2 larvae {ANIC], all collected with holotype, Other mererial [all SAMA|: 2.6 8.5 2 9,2 pupal shins, 4 pupac, wll same chit as paratypes: 2 larvae, collected wilh holotype 158 P, KOLESIK. R, WHITTEMORE & H, M. STACE Figs 1-8. Asphondylia anthocercidis sp. nov. | - 4 male, 5— 8 female. |. Head in frontal view. 2. Last three flagellomeres. 3, Genitalia in dorsal view. 4. Gonostylus in posterior view, 5. Basal lobes on ovipositor in dorsal view. 6. End of ovipositor in lateral view. 7. Last five flagellomeres. 8. End of abdomen in lateral view. Scale bars = 100 jim 1-3. 5-7: 50 um 4; 500 fim &. A NEW SPECIES OF ASPHONDYLIA 159 Figs 9-15. Asphondylia dnthocercidis sp. nov. 9 - 11, 13 pupa; 12, 14 larva; 15 infestation..9, Anterior part in ventral view. 10. Anterior part in lateral view, 11. Prothoracie spiracle, 12, Head and first thoracic segment in ventral view. 13. Last abdominal segment in ventral view, 14. Last two abdominal segments in dorsal view, 15, Fruit gall on Anthocercis literea Labill. [redrawn from Rippey & Rowland (1995)|. Scale bars = 500 wm 9,10; 50 pm 11; 100 pm 12-14: 10mm 15, 1Ov) PO ROLESIK. 2. WHITTEMORE & WM, STAC Description Mule (Vigs 1-4) Colour; sclerotized parts of body reddish-brown, noselerotized parts of abdomen grey, Head. Antennin scape broadest distally, 17-20 x breadth at distal end, 2.62.7 x length af pedicel: pedicel width 12-14 x length; first Magellomere Ls Joy Tengith of scape. fagellomeres evenly evlindrical: citoumfita dense, equally distributed along flagellomeres. Eye favets hexagonoids eye bridge K-48 facets Wide, Prons with WO-17 sete per side. Lubella prominent, laterally with 7-10) setae. sclulose. Manithury palpus 3-seamented, seenjents successively and progressively longer: Thorax, Wing length 2.4 tim (range 3.13.7). widih 13 mm (hd-l4) Se cell pigmented proximally, Claws of all legs subequal ih size and similar in shape. ws long as empodii, Abdomen. Genitalia: gonostylus with two fare, apical (eth of same lengths aedeazus elongate and narrow, reachine tniddle of gonosty tits. Femiule (Pigs 5-8) Frons with 9-20 sere per side. labella with 7-4) selue laterally, Circunfila comprising twat longitidinal and two short transverse bands. Wing Janeth 3.6 mim (2.3-3.9). width bd mm (i235), Seventh abdominal sternite 18 ().0-2)2) 8 leneth ol sith. Genital ovipositor 1.9.8 C8 -2.0) length of seventh sternite; basal Jobes an ovipositor broud i dorsal view, divided in posterior thitd) pecially, fused vere: phibrous, Pupa (Pigs Sell. 14) Colour brawn. Total length 4.00 nn (3.64.5). Anennal horns not serrated, 242 pri (237-247) long. Obe Upper and three lower frontal horns. Profhoracie horn slightly curved, basal part about 2 x width of lenmminal third, terminal third setose. Abdoeniml dorsal spines sitiple, straight, with 2-7 pitirs Gn list sexment curved laterally, Maiure larva (Figs 12. 14) Colour: yellowishayhile, Total length 3.4 min (2.6- 41). Mead cupsule strongly pigmented. postero- huteral extensions not developed, Spatula with four witerion teeth, miner pair smaller Thun outer shit narrow, broadened both at mid-lengih and) base, suroudded anteriorly and daterally by extensive pigmented area, Hach side of spatula with triplecane pair OF fiteral papillae, all setose. Six terminal papillic present, one pair corpiform, 2 pairs with short setae, other papillae as for Ayplaned ylrer (Molin 1955) Gall and biology This gall nidge induces delornyution oF Fruits of Anthavercix lillaren, The upiearpellite: Ovaries are transtormed into glabrous, spherical to ovare, nippled galls, 7-18 mim long and 7-13 mm wide (Pig. 45) and bright green to purple in colour, Inside the gall a chamber. about 3 mm Jong and 2 mm wide, ts oveupied by one larva, The chamber is always lined with fungal mycelia. Although the fungus was abundant in the many gals examined, no sexual stages were observed und the fungtis remains unidentified, Visible seeds are rarely produced in alls although pollination is esscatatd) to vetain the oull on the plant. The numbers of galls ine (ilerea populations are ofien very high. with the galls outnumbering the Hormuallysdeveloped fruits by tp tt 38 limes (Whittemore), Pupution takes place within the gall, AC the end ot iis development the pupa cutsan opening. in the gall and Lifts mast af its body outside the gall. Phe pupal skin then splits open ahd the adall enierges, At Hillarys in 1996, the adults emerged throughout the entire bost plant lowering period, Le, fom April to September. Distribution Ayphandylia anthocercilis sp. nay. ws syonpaten with A. filtered aeross (he entire geographic distribution of the host plant, whieh tanges tron Kalbarres [27"50' &. LIASO7 KE] it the north te Israelite Bay [3427° S. 119923" Ey) tn the soul (Whittemore), Envimology The fame is derived from the generic name of the host plant, Remarks Asphondvlia isa worldwide genus wilh six spectes previously described [rom Austratia. The file history of three ol thent is known: AL dedonecae Kolesik induces walls on leaves of Dodonaea viveose Jacq, subsp, apadiudade (Sin West. and AL m/tever Kolesik and A, ericiformiy Kolestk Induce galls on branch segments of Malosarcia peruranulate (Black } Wilson subsp. pevgeaidane amd A. indica subsp, leiosmchyer (Benth.) Wilson. tespectively (Xolestk 1995b, 1997). Life histories of (wo other species, A loewe Skuse and A, evbienndd Skuse. are unknawn (Skuse PSS8, 1890), The remaining species. A. fill Ldwards, has been reported to indice galls on the stent of an unidentified pling (Bdwards 1916) Aspondvlie Ai, As loewi and AL rebierrda ape: (ot considered in the present paper. The deseriptians of these three species were superlietal and therefore itis A NEW SPECIES OF ASPHONDYLIA lol Hol possible Lo compare them with each other or with A, dedonaeae, A. inflata, A. eriviformis or A, wuhocercidis, A review of the Austrahan species of this venus is planned by PL K. The new species differs from Asphondylte dedonacae in the longer adult scape. the wider teeth on the gonostylus. the unserrated antennal horns and the presence of both upper and Jower frontal horns in the pupa. the shafted spatula and the presence of a piginented area around the larval spatula. The new species can be distinguished from both A. inflata and A. ericiformis by several characters, In A. cithocercidis. the wedeagus reaches the middle of sonostylus, pupae have three lower frontal horns, the prothoracic horn is setose at the distal third and is about twice as Wide al the base as is the distal (hird. At least two of the dorsal spines on the last pupal segment are curved literally and the spatula has four anterior teeth. In both AL faflare and A. erictfarmis, the uwedeagus extends beyond the middle of the gonostylus. The pupa ol A, inflata has one lower frontal horn and that of A. erfciformiy has none. In both species. the prothoracic horns are asetose and about four times wider at the base than at the (erminal third. In the pupa of A. inflata, only the prominent pair of abdominal dorsal spines on the last segment is curved laterally: in A. erieiformis all spines are straight. In both species the spatula bas two unterior teeth. Acknowledgments The field work Was supported by the University ol Western Australia while R.W. was completing bet Honours degree, We are grateful to J. D. Gray. Department of Horticulture. Viticulture and Oenology University of Adelaide and R, J. Gagné. Systematic Entomology — Laboratory USDA Washington DC for their comments on an carly dratt of the manuscript. References Coctess, DL bh (1965) An extraordinary Australian Cecidomyiid, possibly related i the Nymphomyiidae (Diptera, Nematocern). Pro Reni. Sac. Lond. (B) 34, 145-149, Eowakos, FW. (19760 Two new Austaihan Diptera Ann. Mea. Nut. Aist. 103, 498-507. herr, EL P.(1915) New genera and species of gall nidyes. Proe, US nam. Mrs, 48, 195-211. GaAGNE. RoI. ( 1989) Family Cecidomyiidae pp. 152-104 In Hvenhuis, NW, L. (Ed.) “Catalog of the Diptera ol the Austrulasiain and Oveanian Regions” (Bishop Museum Press and Rod, Brill, Honolulu). KoLbstk. P.(1995a) A new species of Fovineticoria Fell (Diptera: Cecidomyiidae) on Eucalyptus fuseieulase in South Australi, A Ads ent Soe. 34, 147-152, — (1995b) Asphonidvita ledenieae, a new species of Cecidomyiiae (Diptera) damaging leaves and branches of hop-bush, Dodonaea viveosa (Sapindacene) in Australia. Tran. Ro See. 8. Ads EIQ 17 L176. (1997) Two new species of Asphamnidylia (Diptera: Cecidomyitdue) fram Hilosareie spp. (Chenopodiaceae) m South Australia, (hie. 121. 59-66. Monn, E. (1955) Beitrige zur Systematik der Larven der Honididae = (=Cevidomytidae, Diptera) 1. Teil: Porricondylinue und Itonidinae Mitleleuropas. Zedlovien 105, 1-247. Purpiz. RoW... Symon, D, B, & Hara Lb. (1982) Family Solanaceae pp, |-208 fn George, AS, (hd Flora of Australia” Vol. 29 (Australian Government Publishing Service, Canberra). Rirrey. E, & Row asp, B. (1995) “Phints of (he Perth Coast and Islands” (University ol Western Australia Press. Nedlands). Skuse. FA. A. (1888) Diptera af Australia, Part 1, Proe, Linn. Soc. NSW. (2nd Series) 3, 17-145, (1890) Diptera of Australia, Supplement |. /bid. 5, 374-412, Nemitocera - FIRST RECORD OF THE ORIENTAL LATRINE FLY, CHRYSOMYA MEGACEPHALA (FABRICIUS) (DIPTERA: CALLIPHORIDAE), FROM SOUTH AUSTRALIA BRIEF COMMUNICATION Summary Blowflies are well known for their ecological, veterinary and forensic importance’ but they are also significant medically as mechanical vectors of dangerous pathogens’. The Oriental Latrine Fly, Chrysomya megacephala Fabricius, 1794’, is notorious in this regard. Adults of the synanthropic form of this species (see below) are attracted to foodstuffs, human and animal faeces and carrion* and have been implicated in the transmission of viruses”, protozoans’, enteric bacteria’ and helminths*. The larvae are also known as facultative parasites in traumatic lesions in humans and other animals’. Tis tans af tlre Royal Socteny aS) Avast (1997), Wy, 164 The, BRIEF COMMUNICATION FIRST RECORD OF THE ORIENTAL LATRINE FLY, CHRYSOMYA MEGACKPHALA (FABRICIUS) (DIPTERA; CALLIPHORIDAE), FROM SOUTH AUSTRALIA ny Blow/lies are well Koowir Cor their eeologiesl vetering dod Jorensie importance! hut (hey are alsa significant medically us tneclanical vectors af dangerous pathoweny: The Oriental Lutrine Ply, Cheywonwd ievecepheata Pubrighis. [P94 is fotorigus mn this regard, Adulls of rhe aynanthirapic Lorn ol this species (see helow) unc attracted fo Toodstulls, Hunan and ainiinel Gieees and carrion! and have been Tipheaed in (he (ansmission of viruses’. protozoa! entero haeler iy and belts. The larvae see WO KAQWI TS fHcuTLative parasites i triumatie lesions i HUM shel other guiiagels:, TPrepart lure the first record oo eueeey ede in South Australia, It was eaueli ai lire numbers ina liver-baited Lap on The Nort Terrie campus or tie University ol Adib in April. 997. 1 had Wot previously encounluredt i uP this sie. despite periodic upping of blowHies over the preceding seven years. Neither hac | encountered it elsewhere TH South Austadin There are pe specimens uf © Hievecephale Ton this sake in the eitoiiologival collection abthe Soult Australian Miscuimauil the closest record vil iy fo Soul Avsttuha oo that ool three fentiles at Murrcinbatenian, New South Wales (s4 58'S, taued2" be), approvimalehy 950 kaye the east al Adulane UK, R. Norris Divisional Entomol CSIRO pers, comm, [Y7). Ulsewhere in Aust G. megerephaliy occurs weross Ue fie north ol the eentinent. down the Gast codsts of Oueenshid und New South Wales and in south-wese Westen Alisttalia, W lias not beee recorded frei Vietoria or Rist! Clinyyomye iegueephuli is probably a HEPOUCTON to South Austria, as il is actively ex pamting ies Fe iT GHer parts AP Wark Sittce the TOTS HL hue invaded New Aeahana!!, several parts of Afhied und arens HH South. Cantal! and North America! Ut is alse fined i) lapin and ds widespread throughout the Oriental reign sod (he Ausiro-Mahiyan and Polyresin subrewions al the AUSTiiasian reson, (Cheyanne negaveplidto Oceuis 1 Lwe harms which are Horpliglowiehy duel ecolusicdlly different. "They hive been defined by Kurahishil cs the “norm aid “derive lt’ Faris The Horni) (arn ts resteicred to tropredh faresis on Sout Pacific hands from the Bismurek Atehipelaust to Suro, Has believed tebe the plestonarphie Lorne eb the species, The derived fori is synantiivapic and dispersive HHO is Thouwwht Wo have orieinated jit Papua New Guiness ain Ween farzinelioglu, AZ, (06) Glowlles iRichiinwad Publishiie Co Lik Shout, ‘Greenberg, Bo (1971 "Flies aid Disehae. vol 2. Bialowy und Discuse Transmission” (Priicetaa University Press, Pringen Wabrictus, fh ©. (1794) “Enromuleia systematics crendiliy ch aueGi Secunda elisses, ardiney. wenera, species unheeds sy munis. loci bservationibus, descriptiomibus’. Tome +00. G, Pratl. Maroiaes, the westerh bodndary of the species! ynewseal Uieuiburion! “The individuals eolleeied from Adebnide are ofthis (orm, ‘The derved form can be distinguished tram the normal form hy the greutly-enkirged onimatiche in the upper (wee (hinds wf the eve Th males. “Phe noenal form hus aaly shighlly enhurged onamuidia in this vesion % Sine the two fornis differ so markedly in there eeologicul preferences and WH SOME AApeets Of their merphology. they may be sufligivutly divergent genenediy lo warnint subspeeitie vr “yen specitic status. This miehl berevweuled by buwhenicul analysis. bs creed aut recently for ather closely-rclaled blow(lios af the genus Culliphera'. although a detailed Morpholosicul comparison would abso be required betore a conelusion about tverr status could he teached. Boil forms af) megacephete can be distineiasted fron ullathee known Austin species of Chryyannya hy the following Motphotogicul characters: Hairs on prevlar kaob black. loner thio height ef Roob: anterior thoracic spineless bhickish brawny legs bhick) subyibrmsal sealae bhicks eyes Hr males willl oniauitidiae ig (he upper Uwo-thirds enlaived and sharply demuncuted from the small ones in the lower thirds [rons in females wider tn the middlehie tl, However, the species is similar physiologically (ae rithet species al Chrysomved in that it is Mermophilie, ant (herefore, in South Australian will presumably be active OOTY ducing the waemer months of the year that is benween Oetober and April. Because al its habits, Co itegdcepheald deserves serious ubention front it publie health perspective and its distribution ih Atistralig: should (herefore be monirered, tts larvae could also be cneountered iy forensic cases in the Adeltide revion, but because their moarphalagy bears Superficial similarity to that ol seme other commun euriion- hreeding species al blowlly. they eGuld be iistaken bor them. Sinee the pile of development al Co mega lela ita gWen temperature differs significantly from these orbper species? misidentificdiion Of this important fly could Ted to serious errors TH extinates of (he tiie singe death ot DUT Corpses Uihank Drs Do A. Duekhotise, RoR. Nortis ane ALB Bird for theiy user commenr on the munserph, Tho ahi also thamks Dr Norris for his helphil advice and casisiince iouttaigry the loa of specimens. F7rimpt, 8 (1905) “Mybisis in Mao and Animals in thet World” (Butlerworths. London), Purtanetto. S. We PL Campos, Mba Gy Thaesi, C2 Ma, Burulll, G. M, & Ishihata, G, Ky ()984d) Rey, Microbiol, 45, 170-174 ‘Harris, A. & Down. Ho A. (1440) Ani Prop, Med, 26. 7AY-ROG, ‘Lima, M.1. POS. & Baie, KOO) Achy Biol, Manan, 20) Oless 164 *Monzon, R. B., Sanchez, A. R., Tadaiman, B, M., Najos, O. A., Valencia, E. G., De Rueda, R. R. & Ventura, J. V. M. (1991) Southeast Asian J. Trop. Med. Public Health 22, 222-228. °’Spradbery, J. P. (1991) “A Manual for the Diagnosis of Screw-worm Fly” (CSIRO, Canberra). Herman, T. J. B. (1990) Weta 13, 12-13. “Braack, L. E. O. (1991) Onderstepoort J. of Vet. Res. 58, 311-312. Guimaraes, J. H., do Prado, A. P. & Linhares, A. X. (1979) Rev. Bras. Entomol, 22, 53-60. “Kurahashi, H., Wells, J. D. & Ogino, K. (1994) Jap. J. Entomol. 62, 860. Wells, J. D. (1991) J. Med. Entomol, 28, 471-473. Kurahashi, H. (1984) Dispersal of Filth Flies through Natural and Human Agencies: Origin and Immigration of a Synanthropic Form of Chrysomya megacephala pp. 37-63 In Laird, M. (Ed.) “Commerce and the Spread of Pests and Disease Vectors” (Praeger Publishers, New York). '6Kurahashi, H. (1982) Monogr. Biol. 42, 689-698. '7Wallman, J. F. & Adams, M. (1997) Aust. J. Zool. 45, 337-356, 'SDear, J. P. (1986) Fauna N. Z. 8, 1-86. “Wells, J. D. & Kurahashi, H. (1996) Med. Entomol. Zool. 47, 131-138. ~»Wijesundara, D. P. (1957) Ceylon J. Sci. 25, 169-185. 210’Flynn, M. A. (1983) J. Aust. ent. Soc. 22, 137-148. J. RF WALLMAN. Department of Zoology, The University of Adelaide Aust. 5005. THE STATUS OF CYCLOSTRONGYLUS MEDIOANNULATUS JOHNSTON & MAWSON, 1940 BRIEF COMMUNICATION Summary Cyclostrongylus medioannulatus Johnston & Mawson, 1940 was originally described by Johnston & Mawson’ from three females found in the stomach of Macropus thetis (sic) now Thylogale thetis collected from the Burnett River district in Queensland. The authors commented that the worms differed from other species of Cyclostrongylus Johnston & Mawson, 1939 only in having a narrow supporting ring around the buccal cavity. Mawson’ revised the genus and considered its relationships with related genera. She found the type species of Cyclostrongylus which had been erected in 1939°, to be identical with that of the type species of Oecsophagonastes (Johnston & Mawson, 1942) and placed Oesophagonastes in synonymy with Cyclostrongylus. Treason of the Royal Seeter: aS. Aust, (1997), T2165, BRIEF COMMUNICATION THE STATUS OF CYCLOSTRONGYLUS MEDIOANNULATUS JOUNSTON & MAWSON, 1940 Cy lostonevtis oedivcmilunis Johnsen & Mawsen, IM40 was onwingtly deserbed by Johnston & Miewsion! from three females found int the stemmed af Waeropas rents (ae) now Clivlvvale thediy collected trom the Burnett River distviel in Queenshund. The authors commented that the words differed trom other speeies of Cyelosmene vis fohinston & Mawson. 1939 only in having a iareosy Supporting ring dround the bucest! cuyily, Mawson? revised the genus and considered tts relationships with relitedt gener, She found the type species of Cyelasmenowes which lid been erected in 1439 to be mdenteal with that ot the type specivs of Gesephavornaytes Sobroston d& Mawson 1942) and placed Cesoplemanasies in syaonyinw with Cy lusenevios, Maawsoie listed the valid species ol Cyclosiomeyliy us €) wellahiae Johnson & Mawson, [949 (ype species) eecurring in Wallahia biculor. Co gallerélr Johnston ok Mawsan. 1939 Geeurrine Tn Mew reps rufourtsens, ©, kertune (Miowson, }YS5) occurring in Ay, euxont ull ME rifeerisnin CC heprtons iMawsen, 65) oeourrm im MW. dieseliy and Co parina Joliston & Mawson, 1939 uccurring in Ad, pervs. Ol the other species previously Cyelosironeviis, ©. déysanilis, lohrston & Miawsen. 1958 veeurring in We Bivelor owas referred tw Manmpostoneylnides (Yaron, T6b and Co cledeeds Johnston & Muwson. 1939 oecurmmg in A defer now A, yivdHels Waa new gents later deseribed as Alecesromd Mawson, P974° Subscquedthy ©. valle: wos eedesenbed is Wellabmema gallard!, othe case oF OC) medina Mawson! Pound (har the specimen libelled as the Pype was i femule Rageplicaryins qtralis (Monning. 126), abyiously placed there in error. Sinee the original material could Hat be found she declared the species a speeles frnynivencde, Mawson? also revised the genus Macrapesrang lie Yorke & Maplostone, 1926 and ereeted Hiree new fener Including — Pepevestrongylis, — Pupovesmronwuytiis yas chuetensec by an oesaphawus With eV lindrical Corpus. a qurrow isthiusindh ending ina bolh; anda buces! capsule whieh is thickest ar midength and has an anterior border aussie tn ‘Johnston, TAH & Mawson. FV (1940 broc, Linn. See. NSW 65, 468-476, ‘Mawson, PM. (1977) Trans. BR. Soe. S. Aust, LOD, 19) 20. Vohnston, TH. & Mawson, PML (193) Prac. Linn, Soe NSW 64, 514-536, ‘Beveridge, b. (984) Aust. Zool. Suppl. Ser YL. 1-8. Without projections, The species icluded the type species P wellabiee Jolston & Mawsen, 1930), accurting iM, puifegrisens and Wo hivalen Po peaevani ovcurring in Petrowale turevalis., M. vageml and Me puilieriseas and 2B iomecoccurring in ML tema, Bevenndye® then uiended the coneriv diagnosis, extunied the host niige ut FE pearvens to inchile M. fidivinesis wind described two new species. 7! mocrupodiy ocvurriny 1 ML pigeniteis, ME cufies und ME. robustus, und FL diviegele oveurring ww Thivlogale vinmatica, UT hrunii and Petrogale peryephom Beveridge’ commented thal Modivlevele Oecurred commonly in 7 vigimetica ti northern Queevishin, anid i Perweale penephone tram central Queensland hit had mut heen found in 7 artgeninien or 2 thers in southem Oueenshind, Spratt ev af! disted ©) tedioanaildtus as a species ol Popovastronglis. Reeendy a slide (HC 2660) labelled Cyclosmennlies mrecdiodneataiies to PL ME Mawson’s handwriting, was found in the South Australian Museum. Adelaide. The slide COUsISDs Ol pitees Of bWwo Tenmale worms Henibed uy resin und has allkached to jl another band-wrillen note by Mawson Indicating that the specimens are two ol the anly three females ever vollected. The charueters of the oesophiyiis. buccal vapsule, female tail and ovejector thal can be Jistinguished in these two femules are consistent with the characters al Hapovestinteyiins, Purther. the bea! capsule has an unnulirihiekening around the middle as 15 lound in Potlivlagete. The eesophagiis (O07) Mn is sharin in O. medivaniilaius than in Polivlawale (09-094) mim. Phe measurements OF the posterior end of CL nreetiocmiaiiy Ure OL Cong rue WHE those of A iivtowle. Vulva te fall up 0.35 compared to 0.7240, 80. nd Wil 0.24 compared to 0 45- 0.59 mim respectively, The viawini vera oof ©. mediocmoulatin. however. ts Ure sie tenth as that ot Je Nvlegale. Cvelostronuyluy medioannidanie. Weretore. is leary wspectes of Fapovestueyils anel is mest similar to Po thyfowale. Addition materiah including: male specimens. iA needed betore un exact determunition can be thade. No other specimens of Popovasinertio, Towever have been Found in 7 iferty bo date Mawson, PM, (1977) Trays, R, Soe S Aust TOL 51-62. "Beveridge, 1. (1956) Bull, Mus, Nati Plist. Nal. Parts 4 Sen 8 (4, 2), 257-265, ‘Spratt, D.M., Beveridge, £., & Walter, ILL. (1091) Rec, So Aust. Mus. Motion Ser 1} 105. LAR SMALES. Departinent of Biology, Facully of Applied Scienve Central Queenstind University Rockhampton Old 472. ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED Patron: HIS EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA SIR ERIC NEAL, AC, CVO OFFICERS FOR 1997-98 President: T. C. R. WHITE, BSc, BSc(For), PhD Vice-Presidents: M. DAVIES, BSc, MSc, PhD M. A. J. WILLIAMS, BA(Hons), MA, PhD, DSc Secretary: Treasurer: O. W. WIEBKIN, BSc, PhD J. H. BRADBURY, BSc, MSc Editor: Assistant Editor: J. BIRD, BSc N. F ALLEY, BA(Hons), MA, PhD Librarian: Programme Secretary: S. BARKER, BSc(Hons), PhD Minutes Secretary: Membership Secretary: A. J. McARTHUR, BE Members of Council: P. KOLESIK, BSc, PhD A. F. BIRD, BSc, MSc, PhD, DSc S. PICHELIN, BSc (Hons), PhD M. J. WRIGHT, RDA R. D. SHARRAD, BSc(Hons), PhD, DipT(Sec) Printed by Graphic Print Group, 10-14 Kingston Avenue, Richmond, S.A. 5033