VOL. 119, PARTS 1 & 2 31 MAY, 1995 Contents Transactions of the Royal Society of South Australia Incorporated Bourman, R. P. A review of laterite studies in southern South Australia - - Shiel, R. J. & Dickson, J. A. Cladocera recorded from Australia - - - - Kolesik, P. Skusemyia allocasuarinae, a new genus and species of Cecidomyiidae (Diptera) damaging lateral branch buds of drooping sheoak, Allocasuarina verticillata in Australia - - - - - Nicholas, W. L. & Stewart, A. C. New genera, species and a new subfamily of Xyalidae (Nematoda: Monhysterida) from ocean beaches in Australia and Pe a a ee a ee es Bradbury, J. H. & Williams, W. D. A new genus and species of crangonyctoid SERIES (Crustacea) from Western Australian fresh waters - - Fuller, M. K. & Jenkins, R. J. F. Arrowipora fromensis, a new genus and species of tabulate-like coral from the Early Cambrian Moorowie Formation, Flinders Ranges, South Australia - - - - - - Molina Ballesteros, E., Campbell, E. M., Bourne, J. A. & Twidale, C. R. Character and interpretation of the regolith exposed at Point Drummond, west coast of Eyre Peninsula, South Australia - - - - Smales, L. R. A revision of the genus Tikusnema (Nematoda: Acuarioidea) with the description of a new species from the false water-rat Xeromys myoides from Queensland - - - - - - - - - Brief Communications: Smales, L. R. Mastophorus muris (Nematoda: Spirocercidae) from the Musky Rat- kangaroo, Hypsiprymnodon moschatus - = = = 3 Vilizzi, L. & Walker, K. F. Otoliths as potential indicators of age in common ae Gee carpio L. (Cyprinidae: Teleostei) - - PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 29 4] 47 67 ih) 83 89 95 97 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 119, PART 1 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INC. CONTENTS, VOL. 119, 1995 PARTS | & 2, 31 MAY, 1995 Bourman, R. P. A review of laterite studies in southern South Australia - - - Shiel, R. J. & Dickson, J. A. Cladocera recorded from Australia- - - - - Kolesik, P. Skusemyia allocasuarinae, a new genus and species of Cecidomyiidae (Diptera) damaging lateral branch buds of drooping sheoak, Allocasuarina verticillata in Australia - - - - - - - - - - - Nicholas, W. L. & Stewart, A. C. New genera, species and a new subfamily of Xyalidae (Nematoda: Monhysterida) from ocean beaches in Australia and Thailand Bradbury, J. H. & Williams, M. D. A new genus and species of Eraneonscwid amphiged (Crustacea) from Western Australian fresh waters = - - Fuller, M. K. & Jenkins, R. J. F. Arrowipora fromensis, a new genus and species of tabulate- like coral from the Early Cambrian Moorowie Formation, Flinders Ranges, South Australia = - - . - - - - - - - Molina Ballesteros, E., Campbell, E. M., Bourne, J. A. & Twidale, C. R. Character and interpretation of the regolith exposed at Point Drummond, west coast of Eyre Peninsula, South Australia - - - - = = - Smales, L. R. A revision of the genus 7ikusnema (Nematoda: Acuarioidea) with the description of a new species from the false water-rat Xeromys myoides from Queensland Brief Communications: Smales, L. R. Mastophorus muris (Nematoda: Spirocercidae) from the aise Rat- kangaroo, Hypsiprymnodon moschatus - - - - - - Vilizzi, L. & Walker, K. F. Otoliths as potential indicators of age in common carp, Cyprinus carpio L. (Cyprinidae: Teleostei) — - - - - z > » 7S 83 89 95 97 PARTS 3 & 4, 30 NOVEMBER, 1995 Li, Q. & McGowran, B. Comments on some southern Australian foraminifera and description of the new genus Parredicta - - - - - - - - Cappo, M. The population biology of the temperate reef fish annie iin niga in an artificial reef environment - - - - Schluter, C., Bye, J. A. T. & Harbison, P. The most vigorous South Australian tide = - Bird, A. F. Studies on Eutobrilus heptapapillatus (Nematoda: Tobrilidae) the predominant nematode inhabiting the bottoms of Lakes Albert and Alexandrina, South Australia - - - - - - - - - - - Skerratt, L. F., Beveridge, I. & Durette-Desset, M. -C. Distribution of species of trichostrongyloid nematode parasites in the small intestine of the bush rat, Rattus fuscipes - - - - - - - - - - Barker, S. Eight new species of Australian Buprestidae (Insecta: Coleoptera) - - - Richards, S. J., Burton, T. C., Cunningham, M. J. & Dennis, A. J. A new species of Callulops from New Guinea and comments on the status of C. humicola comptus (Zweifel) (Anura: Microhylidae: Asterophryinae) - - Stewart, A. C. & Nicholas, W. L. Manunema pectenophora sp. nov. (Peresianidae, Leptolaimina), a nematode possessing unusual male supplementary organs - - - - - - - - - - - - Kolesik, P. Asphondylia dodonaeae, a new species of Cecidomyiidae (Diptera) damaging leaves and branches of hop-bush, Dodonaea viscosa (Sapindaceae) in Australia Kolesik, P. Contarinia bursariae, a new species of Cecidomyiidae (Diptera) infesting fruits of sweet bursaria, Bursaria spinosa (Pittosporaceae) in Australia - Williams, D, J. & Brookes, H, M. A review of the scale insect subtribe Andaspidina (Hemiptera: Coccoidea: Diaspididae) and a new genus, Notandaspis, for two Australian species - - - - - - - - - Connolly, R. M. Diet of juvenile King George whiting Sillaginodes punctata (Pisces: Sillaginidae) in the Barker Inlet - Port River estuary, South Australia - Insert ta Transactions of the Royal Society of South Australia, Vol. 119, parts 3 & 4, 30 November, 1995 99 113 123 133 143 149 157 163 171 177 183 19] A REVIEW OF LATERITE STUDIES IN SOUTHERN SOUTH AUSTRALIA By ROBERT P. BOURMAN* Summary Bourman, R. P. (1995) A review of laterite studies in southern South Australia. Trans. R. Soc. S. Aust. 119(1), 1-28, 31 May, 1995. Studies of laterite in southern South Australia are reviewed to throw light on the nature of laterite, its genesis, classification, its relationships to substrate materials and constraining sediments, its use as a morpho-stratigraphic marker and palaeoclimatic indicator, its relationships to deep weathering, and the timing of lateritisation. Evolving views of laterite as a rock unit, as an iron-rich horizon and as a weathering product are traced and processes attributed to laterite formation viz., capillarity, leaching, combinations of water table movements, leaching and capillarity, wetting and drying processes, weathering transformations of materials rich in ferrous iron, and as a lacustrine deposit are assessed. Fundamental to theories of laterite genesis are the roles of relative and absolute accumulation of iron and aluminium minerals. Key Words: laterite, laterite profiles, ferricrete, polygenetic profiles, peneplains, deep weathering, palaeoclimatic indicators, multicyclic landscapes, morpho-strati graphic markers, Transacrions of tt: Royal Society of S, Aust. 1998), WAU), 28. A REVIEW OF LATERITE STUDIES IN SOUTHERN SOUTH AUSTRALIA by ROBERT P. BOURMAN* Summary Bourman, R. F995) A review of laterite studies in southera South Australia. Tras. R. See S. Aust U9). 1-28, 31 May, 1995. Studies of kverite i southern South Australia ure reviewed! to (hrow light on (he nature of Jaterte, ils genesis, classification, its relationships to substrate materials ind constraming sediments, its ise as a morpha-stradigraphie marker and palacoclinnaitic indicator, its relationships to deep weathering, and the (ming of lateritisanon. Evolving views of laterite as at rock amit. as.an iron-rich horizon and as-a weathering product are duced iind processes attributed i laterne formation viz,, capillarity, leaching. combinatons of waler lable movenients, leaching and capillurity, wetting and. drying processes, weathering transformations of materials rich in ferrous iron, and as a lacustrine deposit.are assessed, Fundamenta, to theories of laterite genesis are the roles of relative and absolute accumulation of iran and aluniiniuin minerals, In Southern South Australia interpretations of lindscape evolution have depended heavily on recognition of parts of an original normal laterite profile, consisting of a pallid, bleached zone successively overlain by a mottled yong and latente, a ferruginous and/or alumioous crust, ‘Chis profile has been associaled with formation on a penepluin surface under a humid, bul seasonally dey, (ropical climate. The possible preservation oC a pristine lulerite surface of ereat anhquity in the modern landseape On uplifted peneplains has been entertained by some workers, but questioned by others. Alternatives ta this approach are provided by stratigraphic inyestigalions of polypenetic profiles, and a continual weathering model of laterite formation (hat results in lateral Variability in the distribution of pallid, mouted and daterite materials ord surface initially with regular Wpogruphy. Laterpretations of lateritiged landscapes include diflerential dissection of a complete laterite profile on an uplifted peneplain surface, mullicyelic landscapes successively lateritised and the formation and Jateritisalion of high level surtaves during uplift. Evidence of laterite formation under non-teopical conditions questions the climate-laterite correlation as does the lack of reliable minerals as climatic indicators of laterifsation. Furthermore, the recognition of laleritisation oveurring throughour the Mesozoic and Cainozoic restricts lhe usefulness of Iilerite asa palaeoctimiatic and morpho- straligraphic marker, Key Worps: laterite, laterite profiles. ferricrete, polygenetic profiles, peneplains, deep weathering, pakieochimane mdicators, mulsicyelic landscapes, morpho-straigraphic markers Introduction There iy a long history of research on materials called laterite in South Australia. The general distribution of lulwrilic materials in southern South Australia is shown in Fig.t. Specifically, these materials include ferruginous and aluminous crusts. yaniably described as ‘ortsteim, “Ferruginous duricrust (Lang 1965), ‘durierust’ (Woolnough 1927), ‘ironstone’ (Teale 1918), ‘ironstone cappings’ (Segnit 1937), “indurated zones) ‘ironstone gravels” (Prescot! 1931) and ‘fetricrete’ (Firman 9674; Bourman 1969'; Milnes et ul, 1985), 4 © School of Human und Environmental Sciences, University of South Australia, Holbrooks Road, Onderdale, South Australia, 5032. BotkMan, RP (1969) Landform Studies near Victor Harbour, B.A. (Huns) thesis, The University of Adelaide (unpubl). weathered bedrock. sediments and soils, variably lerruginised, mottled and/or bleached. This paper summarises and critically comments on these definitions, Issues addressed include the diversity of interpretations concerning the nature of laterite, processes of laterite development, laterite profiles, (he topographic and climatic requirements for its formation, its age, and reconstructions of: and interpretations drawn from, laleritie landscapes. The tert ‘laterite’ has been in the scientific iterature since the publivation of Buchanan (1807), David (1887) discussed the origin of laterite in the New England district of New South Wales. but did so without reference to Buchanan's work, and the term did not appear in.the South Australian literature until more than 100 years after its First usage (Teale 1918). Nevertheless. features. subsequently regarded as laterne were discussed by early workers under labels such as ‘Desert Sandstone’ (Woolnough 1927) and “Upland Mincenes’ (Tate 1879), to With few exceptions, the majority of investigators in South Australia have followed the view that laterite formed as a result of intense chemical weathering in a seasonally dry tropical climate on a peneplain surface, largely during the Tertiary. These conditions fayoured the development of a laterite profile comprising a leached sandy A-horizon successively underlain by laterite, a mottled zone and a pallid zone resting on unweathered bedrock (Fig.2). Generally, the present discontinuous distribution of these materials has been ascribed to differential erosion following tectonic uplift of the peneplain. Corrobinnie Depression Cleve Uplands | Neild ~ Glenville Hs 4 ad ee wit ROBERT P. BOURMAN Definition of laterite Laterite as a rock unit Early studies of laterite in southern South Australia were undertaken by geologists, who considered laterite to be a rock or sedimentary unit and equated it with ‘Desert Sandstone’ (silcrete) or with terrestrial deposits referred to as ‘Upland Miocenes. For example, Tate (1879. p, lix) regarded ‘evenly-bedded sandrock, mottled clayey sands and ironstone conglomerates, occupying flat-topped localities in the Adelaide foothills and within the ranges as ‘Upland Miocenes. He Flinders Mt : Olinthus f ? / 4 ara \ ve “Blue Range i -n rote 4 4) ” Turnby Bay. NN Flat ~ Pt Lincoln Cape {y AUSTRALIA 1000 Yallunda Taylor Wadella Springs Oy XO Cygnet - Snelling Fault Scarp mee Border, ae Parndana Fleurieu Peninsula ipa fe Banks Kangaroo Island 40 60 80 100 Km Fig. |. Map showing the localities of lateriiic areas in southern South Australia referred to in the text. Mig, 1A. (Opposite page). Inset in Figure 1. */ Ardrossan ( By me ea ee) s Ew n a aa) GO LATERITE IN SOUTHERN SOUTH AUSTRALIA Adelaide © ’ i} Hallett Cove Witton Bluff s Myponga Spring & ount Happy ~, s Valley _.: nf Kapund ue = aM Gawler “ & '® Highbury ‘ } & , Gun | Emplacement Mt Lofty ) Blackwood Mt Barker - Longwood a Clarendon a Meadows °, e . Macclestield Mt Compass Upper Cz «~ Hindmarsh Valley A < Peeralilla Hill Green Hills e pe a Parawa® e ca Wilson Hill «A Victor Harbor Bremer Basin \ a Millendeljg Fault “s. Harrogate nN ° 1 a’ Whalley Hill on! “> Lucernbrae ' ‘ *s~ Callington a) ‘ ‘ ‘ 4 ROBERT P BOURMAN loneinietsatey A lhyfiaian Lager tle — otewed oy Vu tealir varpurtrea Lap, peli, Mottled cw Titi tnt Melnice Wealpes PUN Bony Fig, 2, Sketch of the pedogenic model of the normal ur stayidard laterite profile incorporating a sandy. bleached A horizon above a laterite horizon, successively underliin by companion materials of mottled and bleached bedrock (Stephens 1946). considered to fave developed on a peneplain under humid tropical eanditions, considered hen to be correlative bout with terrestrial clays overlying fossiliferous limestone at Adelaide and Tertiary terrestrial sediments borderimy the ML Lolly Ranges. Later work has demonstrated that these sediments vary in age from Pleistocene ta Eocene. Furthermore, the limestone exposed in Adelaide 1s now known lo be Late Pliocene (LLudbrook 1980) rather than Miocene as assumed by Tate. The Desert Sandstone in northern South Australia, currently known as silcrete, was interpreted by Tate (1879) ds an extensive lacustrine deposit contemporary with fiver gravels and sands of the Upland Miocenes. Thus silcrete and laterite were noi distinguished and they were both considered to be sediments or rocks. Whereas Tate (1879) equated sediments within the fanges and on their flanks as Upland Miocenes, Benson (1906) separated them into iwo groups, with an older Miocene series cupping hills and a younger series flanking the Western escarpment of the Mt Lofty Ranges. More recent papers have also consideréd laterite to be a rock unit, For example, Major & Vitols (1973, p. 46) described laterite on Kangaroo Island as a ‘massive rock composed of pebble-sized pisolites of maghemite and limonite and fine-grained quartz sand cemented by limonite. This crust, up to 1 m thick, was overlain by white, fine-grained quartz sand and underlain by mottled yellow and red clay or rocks of the weathered Kanmantoo Group metasediments. The crust occurred as boulders where ripped up by ploughs and loose pisolites, mixed with white or yellow sand, were recorded on the margins of the inland plateau. 2 Bourne, £ A. (1974) Chronology of denudation of Northern Eyre Peninsula. M.A. thesis, The University of Adelaide (anpubl,). Lanerire as an jrat-rich harizoan Many geological studies have been concerned with laterite in only a very incidental fashion, and almost any iron-rich horizon has been regarded as laterite (e.4, Glaessner 1953a, Olliver 1964). At various locations within and on the margins of the Mt Lolly Ranges, Teruary sediments, variably weathered and ferruginised, have been reported to centain laterite. For example, at Happy Valley, Olliver (1964) described a sequence of Eocene marine Blanche Point Marls and North Maslin Sands overlain by Pliocene freshwater sands and clays capped and preserved by a lateritre horizon at about 200 m above sea level. The laterite vonsisted of a band of iron-impregnated sandy sediment. Similar occurrences were described tn many sand quarries in Tertiary sands in the Adelaide region by Harris & Olliver (i964) and Olliver & Weir (1967). Linerite as a weathering product The association of lateritic crusts with weathering, profiles (Walther 1889; Maclaren 1906) was iniroduced lo Australian studies by Simpson (1912) und Walther (1915). They espoused the view that laterite formed as iron dnd dluminium oxide effloresences were transported if solution from the water table by capillarity, Walther (915) assigned the term ‘laterite’ to the complete profile, However, laterite in South Australia has most commonly been considered to be an indurated ferruginous horizon ina Weathering profile (Stephens 1946, Hallsworth & Costin 1953, Connah & Hubble 1960), which is quite different from the original laterite, described by Buchanan (1807) as a low-level sedimentary deposit consisting of massive, unstratified iron-rich clay material, full of cavities and pores, which hardened once cut into blocks and exposed to |he atmosphere. Lang (1965) followed Hallsworth & Castin (1953), restricting the term ‘laterite’ to crusts associated with well-differentiated profiles apparently formed by in situ relative accumulation of iron oxides, “Ortsten’ was used by Lang (1965) to describe crusts developed by laterally derived ubsolute accumulations. Where ortstein crusts formed above weathered profiles and simulated in vita weathering profiles they were called ‘duricrusts. Lang considered that laterites on the oldest surfaces were only occasionally developed frojn materials recognisable in the pallid zone, and he assumed tat a discontinuous layer of Tertiary sediments overlay older rocks throughout the lateritic area. Maud (1972) also considered that only soils containing ironstones overlying mottled and pallid zones should be regarded as laterites. This definition has sometimes been ascribed the descriptor “ue laterite (e.g, Bourne TE), LATERITE IN SOUTHERN SOUTH AUSTRALIA 5 Classificanan of laterite There has. been litthe attempt to classify lateritic matenals m South Australia. Teale (1918) used the term ‘ironstone’ to describe ferruginous materials, which were noted to affect all materials except alluvium. They were categorised into four main types: Joose, concretionary gravels in deposits up to} m thick, iron- cemented sands and gravels, ferruginised slates and quarizites, and Jateritic ironstone forming hard sheets. Laterite was categorised by Forrest (19694) as "fossil or relict’, which referred to the complete normal laterite profile, ‘truncated, where the upper horizon was absent, ‘immature’, where the percentage of tron im the cupping was low and the underlying bedrock was only partially weathered, ‘derived’, when the cappiny, had ) Forresr, G. 1 (1969) Geomorphological evolution of the Bremer Valley, B.A. Hons thesis, The University of Adelaide (unpubl.), 4 BourmAn, R. P (1989) Investigations of ferricretes and weathered zones in paris of soufhern and southeastern Australia ~ A reassessment of the laterite concept. PD, | thesis, The University of Adelaide (unpubl), laterite crust. been derived from the reworking of higher crusts: and where this reworked capping rested on weathered. bedrock, and. ‘terricrete’ where an iron-rich crust incorporated partially-rounded quartz pebbles and overlay fresh bedrock. The use of the term ‘ferricrete. coined by Lamplugh (1902) 10 deseribe a ferruginous conglomerate, has been extended to apply to all iron-cemented and indurated continuous horizons and crusts in preference to laterite by some workers (e.g. Milnes et al. 1985. Bourman 19894). Perrierete was classified by Milnes ef al, (1987) and Bourman (19894) as simple types, which included ferruginised bedrock and clastic and organic sediments, and complex. types such as pisolitic. nodular, slabby and vermiform ferricrete, The diflerent forms of ferricrete were noted to display differences in micromorphology, mineralogy and chemistry that reflect the nature of the parent material, environmental conditions during iron impregnation and subsequent transformations during landform evolution. Mottled (Fig, 3) and bleached materials (Fig. 4) were regarded as having developed independently of the ferricretes by Bourman (19894). 6 ROBERT P. BOURMAN Processes of laterite formation It is necessary for the various potential processes of laterite formation to be understood so that more reliable interpretations of ages and relationships to underlying companion materials can be provided. For example, does laterite formation require a peneplain surface, as many workers in South Australia have claimed? Furthermore, with respect to pisoliths, is it possible to distinguish formation in place from transported origins? Many theories of laterite origins have concentrated on vertical translocations of minerals in the regolith that involve capillary rise, vertical leaching and fluctuating water tables. However, Milnes eral, (1985) and Bourman er al. (1987) demonstrated that ferricrete development in southern South Australia has been almost exclusively related to lateral physical and chemical transport to, and accumulation of iron and or aluminium minerals in, discrete preferred sites, Capillarity Teale (1918) favoured the role of capillarity in laterite formation. He concluded that laterite formation depended upon a ferruginous rock or subsoil for an a iron source, dissolution of iron, largely by organic acids, and a hot season to ‘pump the iron salts’ to the surface, causing oxidation and precipitation of limonite. Laterite development by prolonged chemical weathering during the late stages of the cycle of erosion (Davis 1909, 1920), on a Miocene continental peneplain with sluggish surface drainage, in a seasonally dry tropical climate that encouraged capillary rise of iron and aluminium in solution, was described by Woolnough (1927), who had widespread experience of duricrust in Australia. He considered the ‘Upland Miocenes’ of South Australia to be ‘veritable Duricrust albeit of somewhat aberrant type’ (p. 46). He noted similarities between ferruginous cappings in the Mt Lofty Ranges with examples in Western Australia, and regarded some of the ferruginous materials on highlands as ‘thoroughly typical lateritic crusts’ (p. 46) and that the ferruginous surface of much of the ‘Mount Lofty Plateau’ was underlain by highly decomposed arenaceous rocks, similar to those related to ‘Duricrust’ Laterite, ‘Upland Miocenes’ and ‘Desert Sandstone’ were thus considered as contemporary and equivalent duricrusts, resting on weathered rock materials (Woolnough 1927). Fig. 4. Bleached and kaolinised Precambrian Aldgate Sandstone exposed by quarrying at Longwood, in the South Mount Lofty Ranges at 400 m above sea level. The depth of the section is 30 m. LATERITE IN SOUTHERN SOUTIL AUSTRALIA 7 The capillary model of laterite lormation should result in the reversal of soil A and B-horizons, with the surface laterite being the illwyial B-horizou and the underlying iron-depleted clay being the eluyadl A- Honzon, Thus, this model requires. the eogenetic formation of the complete laterite profile. Leading The interpretation of laterite as the B-horizon of i tassi| podzolic soil was pursued by Preseoit (1931) in view of evidence of the dominance of leaching of bases in lmerite profiles, 45 opposed to capilary uplilt, evaponttion and surface precipilalion of iryn und Mumunum oxides. Tropical podzolisation became the most pervasive view on laterite formation im Sout Australia, with the laterite horizon being regarded as a fossil Tuvial B-horizon where laterite occurs in preity of andity. Johns (196la) proposed that poorly drained soils on Eyre Peninsula were Jeached (which appears ty be contradictory). during presumed humid pluvial conditions of the Pliocene, leading t0 the accumulation of iron oxides and the ir ara formation of Lalerile, Combinations of water table movements, leaching and eapillarity Both Whitehouse 0940) and Stephens (146) conturred with the general padzolie ortyin of laterite bul envisaged soufees of jron not only from the overlying leached Ashorizon but also trom iron- depleted, weathered bedrock by water lable Tuctuarions and cupillury rise, Hallsworth & Costin (1953) questioned that the upper podzolised layers of southern Australian laterites comprised parts of original laterite profiles, and suggested that they resulted from intense leaching after lateritisation. However, Prescott & Pendleton (1952) had pointed out that, in spite of current semt-aridity. relic podzolic soils with tronstone gravels in Western Australia remain acid, m-emphasising their hypothesis of the pedogenic origin of laterite, The interpretation of laterite as the indurated, iron- rich B-horizon of a fossil, podzalic soi) profile was favoured by Stephens (1946), who proposed a dynamic pedological model of soil Jormution, subsequent upon dissection of the Jateritic regions in South Australia. He regarded Jaterite us a pedogenic material and suggested that ferruginous concretionary gravels accumulated in the soil profile in the zone of oscillating seasonal water table as a result of alternating reducing > Ravine, F H_ (1959) The tegional geography of Kangaroo Island, Pl, D. thesis, Australian National Dbdversit, Canberra (unpubl, }, und oxidising conditions. He associated the water table Nuctuations with a low teltef und a humid eluate. Under these conditions the concretionary gravels were assumed to form an indurated horizon by their progressive enlargement and coalescence, Later uplift and dissection of the landscape was postulated to explain. the luterite mantling remnants of former peneplains. A major contribujon (6 pedologival studies wus made by Stephens (1946) who recognised the influences of soil development on both the im sip weathered bedrock and the eroded. transported debris. This model proved to be very productive for other pedologists (e-g. Northente 1946; Northcote & Tucker 48; Rix & Hutton 1953), Stephens (1971) later moditied some of his ews on laterite formavon when he investigated a possible co- genetic relauonship between sicrete and laterite. He considered that laterite formed by the aecumulation of hydrated oxides, Kaolinisation of mottled and pallid zones and the acidification of the whole profile, with pronounced leaching losses of silica and bases, Luterite was noted to form by hoth relative and absolute accumulation bat he believed that relative accumulation was predominant. He aisce concluded that although laterite formation was associated with a fluctuating water table. i was nol dependent on either pertect planation surfaces or opica) climates. yiews that have largely been ignured in the local literature. Wenting and drying processey Th opposition to Prescot & Pendleton (1952), Bauer (19595) favoured the view that laterite may currently be forming in southern Australia where the regolith is affected by wetting and drying. He postulated that under these conditions lerraus iron would iigrate upward during waterlogging and conver fo stable ferric iron in dry, oxygenating periods. He recognised a ready source of iron from decomposing country rock und i lemperalure regime warm enough to allow the reduction, jnigration and oxidation of iron. Weathering transformations vf materials rich in ferrous vn Mawson. (19072) described a large saucer-shaped body of bog iron ore, with a maximum thickness ol (Om, forming # flattish-epped hill about 200 m above sea level at Wadella Springs on Eyre Peninsula. He concluded that the deposit had originated from spring waters, with iron sulphate haying derived from the oxidation of underlying pyrite bodies. Thus Mawson, without confusing the occurrence with laterite, had observed and explained the fammanion ata distinetive type of ferricete, 8 ROBERT P BOURMAN The formation of ferruginous crusts, in such places as the Telford and Murray basins, by weathering transformations of minerals containing, ferrous ion such as glauconite, siderite, chamosite and pyrite to ferric iron minerals dominated by goethite has alsa been recorded (Bourman 1989, Bourman ef al. 1995). Lacustrine laterite The view of ironstone formation as lacustrine (€.4. Ferror IOI) or swamp deposits on a peneplain surface close to sea level, with the water table close to the vround surface was suggested lor South Australian samples by Segnit (1937). He also noted the occurrence of three types of fronstone cappings on high Icvel ground and Slopes in the Mt Lofty Ranges Vesicular ferricrete, fonned by iron oxide replacement of organic material in former swamp environments, has been recorded (Bourman 19894) in various landscape posiuons in the Mt Lofty Ranges (Fig. 5) and on Kangaroo Island. Relative and ubsolute accumulation: jn situ versus /ransported laterite Luatertte formation by relative (d7 yftv) and absolute (lateral) accumulation has long been recognised with different workers attributing differing significance to (hese processes. For example, Stephens (1971) attnbuted laterite formation dominantly to relauve accumulation, Crocker (1946) agreed with the in side formation of some ferruginous gravels but considered that some others have secondary origins, Milnes er a/ (1985) and Bourmun ef a/. (1987) presented evidence of dominant lateral transport in ferricrete and pisolith formation in the Mi Lofty Ranges, although the possibility uf int vite formation was not rejected (Fig. 6), Johns (1961a) also conjectured that most of the sediment deposited on the coastal plains and Central Basin of southern Eyre Peninsula was material resorted from lhe uplands and included pisolitic or massive ironstone gravels. Johns beheved that during Jatentisation the previously penepluined basement rocks underwent deep weathering, ferruginisauien and kuvolinisution, Lithologieal variations in the basement rocks were thought to have had no influence on the final weathering products. Maud (1972), following d’Hoore (1954) proposed absolute and relative sources of iron and aluminium oxides for the formation of laterite, The accumulation of iron and aluminium oxides was attributed either to the renioval of silica and bases or their accumulation from Outside sources Well-developed _ lateritic ironstones on Permian glacigene sediments were explained by the concentration of iron oxides from lateral sources, Whereas thinner crusts on pre-Permian rocks were ascribed to in situ weathering losses of silica and bases (Maud |972), Maud (1972) believed that following landscape rejuvenation and lowering of the water table, the zones of iron-enrichment irreversibly ae é — Fig. 5. View of bulldozer excavation om Peeralilla Hill showifg terruginous crust of vesicular ferricrete and light-coloured clays (bottom lef ot photograph) that melude calcite and barite. This deposit of ferricrete occurs on the summil surfitce bul below (he level of surrounding hills, Borehole evidence indicates that this deposit is underlain by sandy sediments, Excavation 2.5 m deep. LATERITE IN SOUTHERN SOUTH AUSTRALIA 9 hardened into lateritic ironstones. Brock (1964°) also agreed with d’Hoore (1954) that dissection of lateritic terrain, accompanied by lateral water movement, may have redeposited iron oxides on gentle slopes to form cappings. Wopfner (1972) carried out an analytical investigation of mottled materials that in other contexts have been referred to as lateritic. He discussed maghemite in mottled Cainozoic sediments at Hallett Cove, where both primary and reworked maghemite were identified. Maghemite was reported from two locations: small amounts (2%) of maghemite in conspicuous red mottles, within medium grained white sandstone, were used as evidence of in situ formation, whereas maghemitic sub-rounded ironstone pebbles in a conglomeratic horizon were considered to be indicators of reworking. The profiles and crusts were considered to be genetically related with the conglomerate forming by reworking of an original in situ crust. © Brock, E. J. (1964) The denudation chronology of Fleurieu Peninsula. M.A, thesis, The University of Adelaide (unpubl.). Many soils associated with uplifted peneplains in Australia have been noted to contain concretionary ironstone gravels, attributed by Prescott (1934) to former wetter periods when waterlogging of soils and shallow water tables were more common than at present. Chemical analyses of ferruginous gravels were interpreted by Prescott (1934) to demonstrate the concretionary character of the ironstone gravels. However, many pisoliths in southern South Australia appear to have formed dominantly by disintegration of ferruginous materials such as mottles, followed by physical transport and modifications in soils resulting in increases in iron content as well as a mineralogy dominated by hematite and maghemite (Milnes er al. 1987; Bourman er al. 1987), Transported pisoliths typically are associated with stone lines, have different chemical and mineralogical compositions to surrounding matrix materials and display multiple rinds. Milnes ef al. (1985) also considered that ferricretes in southern South Australia, as well as pisoliths, are dominantly remnants of iron impregnated sediments, originally formed in former valley bottoms and depressions, Fig. 6, Road cutting on the Victor Harbor-Cape Jervis road west of the Waitpinga road, exposing bands rich in pisoliths, at a depth of about 1 m, in ferruginous sandy sediments of probable Pliocene age and of aeolian origin. Other pisoliths occur at the ground surface and in the upper soil mantle. The pisoliths at depth contain only goethite, whereas those at the surface have higher iron contents and contain dominantly hematite and maghemite. Geological hammer 33 cm long. 1”) ROBERT P BOIRMAN Although there 1s general agreement that terruginous materials can form both by processes operating i sity and those related to transportation, there jis beer confusion in the use of the term fsiti. For example, some workers have considered that ferrerete, formed during landscupe dewynwasting. which involves both vere) and lateral movement ot clasts, fotmed in site. Such derrieretes may be benter regarded as residual witht ia sith weathering applying more strictly only to ovolumimous weathering (Bourman W94b), Laterite profiles The nermal laterite. profile Throughout the South Australian htenttane, following Stephens (1946), runs the thread of the normal laterite profile, which bay influenced many palaeo climatic and palaco-enyironmental reconstructions. Only rarely have stuclies departed Jrom this model. The norrial lateritic profile (Stephens 1946), was envisaged as essentially a podzol with A, Band C horizons of cluviation, Hluyidiocand weathering, with an accessory lalenle honzon usually above a clayey B-horizon, Ovcasionally several laterintic horizons were noted in one profile Stephens believed that the normal laterite profile was restricted to southern Australia, m Queenstand laterite awwas thought to occur as an horizon in red-eurth proliles (Arvin 1939: Whitehouse 1940), which vontined silicified zones. suggesting the incomplete removal of iissolved silica. The model presented by Stephens his considerable ment as emphasises the dynamic nature of landlorm change and pedogenesis. However. its dependence on the widespread occurrence of a former murnal laterite profile related to former regional water tuble Tluctuations, 1s unrealisuc and has led to siipliste explanations of landscape development, Furtherninre (here are various Objections (o the view thal (he onginal laterite is the illuvial horizon of a fossil podzotic soil Widespread Jateritic souls on the clevated pencplain of Kangaroo Island, the Mt Lofty Ranges. Yorke Peninsula and Eyre Peninsula were reported by Crocker (1946), who obseryed that they contained considerable percentages of logse and indurated lateritic tronstone gravels. Some of these gravels were considered to have formed in sine, but on dissected imaginal slopes secondary origins for them were suguested. Crocker (1946) followed the view of Prescott (O31) that laterite is the fossil illuvial horizen of a lfopical Pliocene podzohe soil, Thus he reiterated the then current Lhoughts about laterite and further promulgated. the association of laterite, peneplains, (rapical climates and the Pliocene (or Tertiary}, thereby setting the stage for jhe generation of circular urcuments. Spriggs (946) concurred with Prescott (1931) and Crocker (1946) vonceming laterite genesis, The pedogenic orgin of luterite was promoted hy Northente (1946) and Northcote & Tucker (1948) These workers mupped and described a relic normal laterite profile of Phoeenc age. the Eleanor Sand. on the lateriGc plateau of Kangaroo (sland. Cracker (1946) commented that latent residuals on Kangaroo Island were covered by grey and white siliceous sands derived Irom resorted A-honzons, originally developed on Pleistocene coastal ealeareous sand duncs. However. Northeot (1446) claimed thar the constant rate ol course to fine sand throughout the protile indicated that it had formed iy sita and that the surface lad oot received accessions oF wind Hlown sand, Consequently he regarded the Eleanor Sand as a relatively undisturbed fossil soil of Pliocene age. with a lateritic horizon developed i site and preserved Onan uplifted peneplain. However, Muleahy (960) has stugestod that such satidl ray TOL be fossil, but may have derived! from lunerite destrucnion, thus yielding a similar grain size analysis to that determined by Northcote. ‘Twidale (1983) considered that this sandy A-horizon provides evidence tor the preservation of an original Mesovzae jateme prolile, Rix & Hutton (953) regarded the suinmil surface in the south Mt Lofty Ranges as a block-faulted, uplifted and dissected peneplain. They follawed Sprigy (946), vorsidering that by Farly Tertiary times Precambrian rocks had been reduced ter 4 base surtice and subsequently buried by a Tertiary lacustrine and marine covermass. The soil pattern suggested to ther that a further cycle of erosion had removed the grealer part of the covermass. leaving isolated areas of varying extents thereby creating a new peneplam, with remnants of Tertiary deposits preserved in topognaphic lows. They postulated laterinsation of soils on the peneplain, prior to major faulting and dissection, concurring with Whitehouse (1940) that there had been comlemporanedus. laterite formation throughout! Australia in the Pliocene. Residual literitic soils were only mapped on hill summits and spurs 6a they suggested that erosion had removed most of a larcritic sandpluin following uplitt and dissection. One soil, the Yaroona Gravelly Sand. was regurdel #s an Original laterite profile and described as 3 massive band of laterite 22-30 cm thick. containing water- washed grits, gravels and sands, unconfurmably overlying kaolinised Precambrian shales (Pip. 7). Rix & Hutton (1953) deseribed other residual podzols un the area as exhibiting normal profiles of ferruginuns, motied und pallid zones, overlying unWweathered country rock, These workers were strongly influenced in their interpretations by the aormal laterite profile mode! of Stephens (1946) and presented a convoluted explanation of an anonialous laterite profile. preserved in a road cut south of Clarendon, in order to agcount LATERITE IN SOUTHERN SOUTH AUSTRALIA a for a mottled zone overlying a laterite zone. The section can also be interpreted as a geological sequence of Precambrian rocks weathered in pre-Tertiary times, and overlain by fluvial gravels, grits and sands of Eocene age (Mawson 1953). Subsequently these sediments were partially silicified and superficially stained red by small amounts of iron oxides in groundwaters. A thin grey soil with pisoliths occurs at the surface. The above example demonstrates how complex deductive arguments, within framework of the model of the normal laterite profile, were used to introduce events, for which there was no evidence, in order to explain apparently aberrant observations, Despite this, Rix & Hutton (1953) produced a detailed soil map. In southern South Australia the normal laterite profile of Stephens (1946) has been given excessive consideration, sometimes resulting in simplistic interpretations of landscape development. This has occurred despite observations indicating great variability in lateritic weathering profiles and despite the view of Stephens (1971) that the ‘normal profile’ is the exception rather than the rule, Bauer (19595) and Alley (1977) disagreed with the interpretation of laterite as a tropical fossil soil profile developed on a peneplain. However, their views have not been generally accepted. Polygenetic profiles - Alternatives to the normal laterite profile STRATIGRAPHIC APPROACH TO INVESTIGATIONS OF LATERITIC MATERIALS Firman (1967a, b, 1976, 1981, 1994) placed weathered zones and palaeosols within a stratigraphic framework. For example, he gave formal status to ferruginised clastic sediments and bedrock weathering profiles consisting of sesquioxides of iron and forming ironstone crusts, by introducing the name Yallunda Ferricrete. The Yallunda Ferricrete was reported to exceed | m in thickness in its type area at high levels over interfluves of the Lincoln Uplands (Eyre Peninsula) and on remnants of old high surfaces elsewhere. The term ‘ferricrete’ was used to describe ferruginous layers and crusts both independent of, and in association with, weathered profiles. Firman (1976) interpreted the various zones of the so-called normal laterite profile as having formed by different processes at different times, with the profile as old as the initial transformation of the parent material. Ferricretes in various stratigraphic situations, including ferricrete above and below Lower Tertiary sediments in the Barossa Valley, as well as ferricretes Fig. 7. Section in road cut near Clarendon, exposing the Yaroona Gravelly Sand of Rix & Hutton (1953), showing angular unconformity with ferruginised pebbles, grits and sands of Eocene age overlying weathered, bleached and partly kaolinised Precambrian meta-siltstones. Section is approximately 3 m high. 12 BERT & BOURMAN in the highlands of the Mit Lotty Ranges and the Linealo Uplinds were recorded by Firman (l987a). He alse suggested thal there were equivalents of upland lernieretes in the sedjmentary succession of the Murray Basin. These included oolite siderite-neh sediments and laterite in the Early Pliocene Bookpurnong Beds, ys well as ferruginous beds and cappings in the Late Plipcene Parills Sand near the Victoria-South Australia border, Some ol the lerricretes, however, have resulted from the relatively recent oxidation of pre-existing inon- nich sediments containing glauconite and siderite and cannot be used as reltable sor! stratigraphic markers. Firman (1976) consideyed that between Permaitn und Early Tertary times, some 200 Ma, the Mt Lofty Ranges region was a land mass experiencing protonged wealhering and erosion, so that by Barly Teruary noes a subdued and deeply weathered landscape had developed. Associated bleached profiles were considered 9 haye onginated in the Mesozoic, A range of different ages of weathering and lateritisation was reported, Decomposed, bleached or mottled bedrock underlying Eocene sediments was ascribed to pre- Tertigry weathering, a laterite profile developed in Bocene gravelly sands was used to indicate pos(-Hocene weathering: silicifiec and ferruginous zones in Early Pleistocene sediments, overlying older bleached zones were argued to have equivalents in laterite protiles in the adjoining uplands, and ferruginisation in carbonaceaous and pyriuc Encene sediments was wiribuled tu recent exposure and oxidation. The work of Firman 1s significant in attempting In establish stratigraphic ages for different weathering ieatures. Nevertheless, correlating weathering phases simply on shapes. sizes and colours of muottles may be unreliable, as similar weathering patlerns occur m profiles of different ages. Furthermore. Firman observed modificaGon of Some profiles by later weuthering, obscuring earlier weathering products. Firman apparently took no wecount of local environmental conditions, which may have favoured synchronous bleaching in one area and imotling in another Various questions remain unanswered, such us What happened to the iron derived from the bleaching of the Arckaringa Palacosol; where did the iron for the development of the San Murino Palaeosol come From; nd haw was it concentrated in diserete. bul more-or-less uniformly distributed mottles witht previously bleached bedrock? POL YGINIIe PROFILES AND CONTINUAL WRATIIDRING Mant op Ferricker FORM ation Milnes ef af. (1985, 1987) and Bourman e eal, (1947) combined investigations of the field relationships of ferricretes and weathered zones with wicra- morphological, chemical and mineralogical analyses and questioned the former development of noroul Jaierte profiles. These studies have suggested thar there was complex reworking and continuous Weathering of relic landscapes since the Early Mesozoic, and [hat ferricretes are dominantly remaants of iret impregnated sediments of ancient valleys or Jepressivns. Some ferricretes may be the culmination of processes beginning in the Mesozoic bul sill proceeding, resulting in the repeated dissolution. break up and neo-formation of lerricretes. as well as the ongoing and current formation of mottles and bleached 7ones, Some ferricretes may have developed as suggested by MePurlane (976). who postulated fern¢rete develupment by the surface aceumulation of ferruginous materials during landscape downwasting, the formation of gibbsitie-rich zones in near-surface situations and continued development of ferricretes and bleached zones. after uplifi. However, some other features of her model do not fit the observations in South Australia; there 7s evidence for some bicached zones and mottled zones being older than the ferricretes (Bourman 19894) and no evidence has been observed ol progressive development of profiles, with honzons having formed from progenitors resembling those currently beneath therm, An extension of this model is the continual weuthering hypothesis of Bourman (989%, 19934), which propases. Ongoing epigenetic transformations of ferricretes und weathered zones, with rates of change influenced by climate and events such as teetonism, sedimentary burial und submergence beneath Jakes and the sea Topographic requirements for laterite formation Feneplain concept in laterite development Many workers have associated laterite formation and preservation wath penepluined surfaces. However, Sprigg (1946) considered it unwise to associate laterite formation with peneplanation, which implied formation ower a very long peridd, since be believed thal lateritisation secupied only a relatively shor rime span, This interpretation has important implicanons for landscape evolution as laterite formation would first require the development of an extensive planation surface, The peneplain concept of laterne development begun early (Beason J8iL: Mawson J907b; Teale 1918: Woolnuugh 1927), and bas persisted (Campana Av Wilson 1954; Brock 19649; Ward 1966: Twidale 196b, 983, Maud 1972), For example, the summil surface ofthe MU Gofty Ranges was interpreted as un Early Tertiary differentially uplifted and dissected ‘peneplain, surmounted by monadnocks such as Mt Lofty. Mt Barker and Mt Gawler (Benson IIL). Tate (1879) attributed the discontimuous distribution of the “Lipland Miocenes’, separated by deep ravines, to extensive denudation after uplift of the ranges, LATERILE IN SOUTHERN SOUTH AUSTRALIA 12 Mawson (9076) extended the peneplain concept to Kyre Peninsula where he deseribed peneplains in the Port Lincoln are at about 00m ancl 6 m above sci jevel. He equated inottled clay beds underlying the lower surface with lreshwuater Miocene beds neat Adelaide, Some extremely perspicugious comments on the nature and formation of ferruginous materials i the south Mt Lofty Ranges were mathe by Teale (1918) and his work represents the niost comprehensive, detailed and objective discussion of iron -owides: mong all of the early mvestigators, particularly oo classification and theories of origin of ironstone crusts. ‘Teale (YTS) inlerpreted the surumil surface of the ranges as dislocated and eroded remnants of a former extensive pencplamn Woolnough (1927) sugvested thal the distribution of reninants of the terrestrial Upland Miocenes agreed completely with the physiygraphic vonditions postulated for duricrust formation ic, a peneplain with sluggish drainage, In the southern part of the Mt Lofty Ranges, Campans & Wilson (954) described a planiton surface at levels up to 420 m above sea level. as a pre- Tertiary peneplain, uplifted during Termiary and Quaternary times and deeply dissected by subsequent cyeles of ctosion and Brock (19645) identilied remnants Of an ancient lundsurface an the spine of Fleusicl Peninsila covering on aren of 25 krm*. the remnants Were described as having lithe relief und a capping of the normal kilerite profile of Stephens (1940). Ward (1966) also deserihed fal surlaces preserved on ihe crests and gentle back slopes of the western blocks of the Mt Lolty Ranges as relics of it pre-deformational ML Lotry peneplain, mantled by deep weathering and lalevite, Twidale (1968) described tie summit surface of the Mt Lofty Ranges as a lalerny ined peneplain, surmounted by a few residual remnants or monadnoeks, Today the tert “peneplain’ is rarely used in geomorphic literature because, among other things, 1 carries with itan undemonsteable, Highly thearetical mode of genesis. The terms ‘erosion surlice’ or ‘planation surface’ are preferted. Irregular surfaces Not all workers have considered peneplains necessary for laterite formation. Tf penepliin surtaces are necessary for laterite dormation, the unplieanou is that latertte formation follows peneplinaunon, although irregular surfaces suggest that laterite can form during, and as a result of Jandsecape downwasting. Some investigatory such ax Curmpana (1955) have demonsirated great complexities ard irregularities in Weathering and landscape evoly|ion, Working, near Gawler in the north Mi Lofty Ranges. Campana (1955) noted a leached lateritic soil overlying eneisses, schists and Tertiary Nuviatile deposits resting on it pre-Tertiary weathered erosion surface, He reported gravels and coarse sands cemented by iron oxides widiin the ‘Tertiary sediments. The mapping of the Tertiary (Early Eocene) strata m this area indicated thal deposition oceutred ina system of lakes gind rivers on a weathered surface of rodersle rebel, above which ridges of harder rocks projected, Campana (1955) considered that the non-marine strata and older tocks had been subjected to widespread lateritisation between the Early Eocene and the Miocene, The sequence outlined by Campana (1955) illustrates pre-Tertiary weathering, and bleaehing of basement racks. the deposition ol ‘Tertiary letrestrial sediments over a dissected landscape. differential ferrupinisation ob suilable host rocks aud Lhe likibitton of this by marine submergence, Bauer (19595) noted that repardless of elevauon, the laterite profile. the Eleanor Sand, oeeurs on reas of low reliel and poor drainage that would lave suited periodic waterlogging and drying (Mig. &). Thus he thought (hat toporraphy night buve been important in assisting the formation of 4 distinctive soil in two differing physiographic locations, that is, on a stepped topography with fat treads. but nol mecessanily peceplain. Lung (1965) reported on soils und geomorpholugy of the Yundi area within the south Mi Lofty Ranges, His work represents a departure from that of many eurlier workers us he invoked different types of weathering, erosional and sedimentary mtluences to explain the eurrent landscape and he envisaged lateritisation and duricrust formation as procecding over Jong periods of time, arn landscapes of variable relicf und positions abave seu level, In the same area, Maud (1972) noted Literitised surfaces occurring across infilled glacial yalleys and correlated therm with fhe summit surface despite their lower lindscupe posbons, The gentle non-teetonic tnelinations of jronstone coppings were reparded as original valley morphologies, aid Mand (1972) concluded that the original erosion surface was one of considerable: relief, Bourman (19895, [994a) presented a model of laterite formation involving an original landscape of some rebel that provided lateral local cnvironmentil variability This resultect in bleaching of higher parts of the landscape and iron accumulation on plateau margins, indepressions, swamps and valley bottoms. Primary iron minerals mobilised in sub-surface zones affected by water lable fuctuutions were concentrated in hemaftitic notes. Landscape downwasting concentrated dnd fragmented mottles at the sitface. further weathering modified them, formed pisoliths. cemented (rer lo Jor fertierete at the surface and further modilied the ferricrele, Porhons of (ie sult surface af the Mc Lofry Ranges have been continually uffccted by Weathering and erosion since the Permian crete = Miocene Jimestane c ferricréte Vermiform ferri Pisoliti Pisoliths pia Ie ROBERT P, BOURMAN salle This suggests ferricrete formation during landscape ace. “S evolution rather than being dependent on the presence ns of a planation surface, The continual weathering model postulates ongoing variable weathering, interrupted by tectomie activity, sedimentary bunal or marine or lacustrine submergence, Coastal dunes Climatic conditions required for laterite formation The vast majority of workers has equated laterite formation with a hot, seasonally dry tropical climate favouring the operation of intensive weathering processes. Low topographic relief and tropical climates were considered ideal for laterite formation, generating o many circular arguments related to the laterite-tropical climate-peneplain association, A tropical climate with pronounced wet and dry seasons, such as that of Darwin, was considered ideal for the formation of laterite by Walther (1915). This view has persisted. Stephens (1946), Sprizg (1946), Crocker (1946) and Northcote (1946) associated laterite formation with a pluvial period in the Pliocene und Johns (196la) believed that low relief and high tropical temperatures had favoured the removal of silica, with seasonal oscillations of the water table leading to the concentration of iron oxides, More recent workers such as Bourne (19742), Daily er al. (1974), Twidale & Bourne (1975a), McGowran (1979a), and Twidale (1976b, 1983) also favoured torrid, tropical conditions for Jateritisation. The tming of lateritisation has commonly been associated with independent evidence for tropical climates. For example, Twidale & Bourne (1975a) noted that palaeontological considerations favoured the Triassic as providing the most suitable humid, tropical climatic conditions for the formation of laterite in the Mt Lofty Ranges. In marked contrast. Firman (1981) proposed different climatic conditions for separate parts of the profiles, For example, the bleached zone of the Arckaringa Palaeosol was not considered to have been genetically associated with younger ferruginous zones but to have preceded the development of mottles and ferricretes, He ascribed bleaching to early cool climates and ferruginisation to tropical conditions. uo Other workers such as Bauer (19595) and Campana & Wilson (1954) considered that lateritic material might be forming at present jn southern Australia so that climate for lateritisation need be no different from that of today. Maud (1972) also argued that the process of Mt Stackdale 3 Mount Taylor Plain 15 130m ~ Kilometres Mt Taylor 10 surtace: ts Fig. &. Cross section of the Mount Taylor Plain, Kangaroo Island, showing the relauonships of identical vermiform ferricrete on a high pre-Miocene summil surface and a low & post-Miocene surface, wound 5 LATERITE IN SOUTHERN SOUTIT AUSTRALIA 1" iron oxide-envichment of sediments is curreatly proceeding. on broad valley floors in the Me Lofty Ranges under current climatic conditions. Furthermore, there is considerable evidence of miodern iron mobilisation and precipitation in southern South Australia (c.g. Bourman 19899, Ferguson ey al. 1984). «o thet a hemi tropicul climute need not be a pre requisate (or bleaching and iron enrichment, The role of climatic influences in the formation of ferruginous and siliceous duricrusts was. examined by Alley (977), who provided evidence that both laterite and silerete formed together, for at least some time during the Tertiary, on identical strata and under similar climatic conditions, suggesting that.some other factor(s) must have controlled the processes of weathering. Only the base levels of erosion differed hetween the silereted and [ateritised surfaces and stlerete developed on a surtace, the drainaye of which towed sluggishhy int) Tertiary lakes. Pulynological data were interpreted by Alley (1977) to demonstrate that the Kovene appeared to huve heen warm and temperate with a very high rainfall and that the Miocene was similar, with perhaps Slightly warmer lemperalures and a slightly lower precipitation. The concentration of silica at the landsurfaee Was attributed to high alkalinity, slow proundwater movement und a high water table chase to the lakes. Alley (1977) concluded (hat laterjte arid silorele covexisted for part of the Eurly Cainozoie in adjacent drainage basins, Consequently. laterite and silerete were got thought to form through the mechanism proposed by Stephens (1971) which invelved the formation of silerete by deposition of silice in Ury aones aller faving been derived trom Jateriue weathering elsewhere, Furthermore, the view that luterie is ussociated with tropical conditions snd silerete with aridiry was not supported because both fortned ia similar climatic and biotic regimes; anly the base levels. and groundwater conditions yaned. Using chemical (Auton 1977), palynological and suauigraphic evidence to suppart their argunient McGowran et a/, (978) disagreed with Alley (1977) that laterite and silerete formed concurrently on similar rocks and under broadly similar climatic conditions from Recene to Miocene times, However, Alley (978) couniered the arguments presented by McGowran ev al, (1978) and made a valuable contribution to the study of latente venesis by hay hlighitine the influence of local topographic and groundwater conditions in ats formation, a well as questioning Clioatic aiflucrices on laterite and silcrete development, Minerals as climatic indicators af lateritisation As noted above lateriasution is cotamonly associated with intensive weathering under tropical climatic conditions und certain minerals are suggested 4s indicators of climatic conditions. Kor cxaniple, Woptnes (1972) sugeesicd (hat miaghemite in motiles is a climatic indicator, oripinating by thermal dehydration of lepidoerowite Jormed by oxidtion onder fluctuate water table levels and warm climatic conditions, He voneluded that lepidocroente and goethite myy have formed as pels that were subsequently dehydrated and erystallised as maghemite and hematite under conditions of low telief, warm climate and heavy seasonal yalnfall, However, mayherte in laterite rootiles is very rare in southern South Australia as they are clominantly henatine (Bourman 19894). Moreover in lateritic arcs of the Mt Lofty Ranges, potentially-weatherable minerals mcluiding felspies, muscovite, vermiculite, chlorite and smectite have heen identified (Bourman 19894). In some cases there may have been neo- formation of these minerals bul itdoes seen anomalous that they should be so widespread in areas Considered to have been affected by lateritie weathering processes, Previously, Crawford (1965) had noled Iresh felspir gravel ip motded material at Ardrossanand sed this to argue Against literitic weathering Falaeoclimatic mdicators Depending on the climatic conditions considered essential for laterite formation, the presence of laterite has palacoenvironmental implications. There is little deubt that the operation of chemical processes ts aceclerated under hot moist conditions bat there 4 i growin body of eyidence suggesting that iron mobilisation and kaolindsation can occur under various climatic regimes - see Bourman (1993) for a summary - so that there ave considerable uncertainties linking Jaterite formation with a specilic climate. For example, theye are many reports of modern iran mobility from localities in the Mt Lotiy Ranges, Kangaroo Island and Fisherman Bay (Ferguson er a/. 984) under curren Mediterranean und semi-arid climatic conditions, These observations may also cust doubt on the rehability of correlating lerrestdal ferruginous crusts with evidence ol warm, hum chiates derived from marine climatic indivators (MeGowrin 19796), Interpretation of lateritic hindscapes Many different hypotheses have been presented to explain the distribution and evolution of laterite. These include the deVelopment of laterite on a surface of low relief, clase to sca level, followed by differential tectome uplift and dissection of thre lateritic surface, developmentof multiple erosion surfaces affected by episodic weathering anc Javerite formuttion, differential weatherme und laterite formation on a landscape formed by uplift and dissection of a surface orginally of low relief, and the weathering, erosion and scdimentation of a landscage before, durin: antl afer uplift. Lo ROBERT TP BOURMAN Reconsiruetion of lateritised landscapes ‘The models of landscape evolution presented to expliin the development of latente depend ou precomceptions of haw laterite forms. For example, il is often assumed that isolated occurrences of taterite represent dissection of a former continuous faterite surivce and that the differemt horizens of laterite profiles formed contemporaneously. In the past. many workers have tacitly assumed that the present day isolated and sporadic occurrences of baterite represent the erosional dissection of a former contiguous and unilonm Jaterite-mantled planation surface and that these remnants are excellent and reliable morpho- sauigraphic markers (eg. Twidale 1983). However, discontinuous distributions tay reflect only localised formation in favourable localines (Bourn |993a) where optimum topographic and climatic conditions did not penerally prevail, Henec, the oveurrenee of faler{le need hor necessarily Tndicate uv tormer extensive erosion Surhtee, Preservation af uplifted peneplains Interpretations of landscape evolution have commonly depended upan recognition of uplifted ane dissected former peneplains, and the preservation of parts of the onginal weathered surface, which can be used Wy reconstruc! the former surface. A review of the churacter and age of the sanumit high plain of the Mt Lofly Ranges was presented by Twidale (1976), who argued thal (he summit surface as of Mesozoic ue and has been preserved for some 200 Ma (Twidale, 97h), Recurrent uplift of the Mr Lofty Ranges, it was argued. postponed the ultimate degradation of the ranges by exposing new land to the area undergoing reduction, However, other workers have suggested that the best preservalinn of laterile is in relatively low lying points and least in arews of greatest uplift (Milnes eral. 1985). The development of the Mt Lofty Ranges on an unlicline, the Manaks of which are faulted, was one factor used by Twidale (19769) to explain the preservation of the Taterite-capped phiteau, {t was maintained that the bulk of the plateau is centrally located close to the resistant compressional zone of Ihe anoelime and remnants near the western margin are huttressed by sandstone and limestone outcrops, However, the folding. which occurred in the Cambrian. was very complex and did not result in the formation al a simple anhcline. Moreover, erosion of tis complex structure bas been so pronounced thal yertical and near- vertical rock structures are exposed. Furthermore, subsequent tensional faulting bas occurred within the runyes (Glaessner 19534), so that the core of the anges should not be considered to be in Compresston_ The preservation of the Mesozoic sandy A-horizon of the Jalgrite profile was thought iq haye assisted pilueosurface preservation by providing an absorbent cushion to protect the underlying lerruginous harizan from) rainfall (Twidale 19764). However, no evidence has been found by the present wuthor of 200 Ma old sundy A-horizons in the ranges. Conversely, sundy soils ure common, especially on Pertman glaevigene seditnents and oceur in landsurfaces. demonstrably ol posi-Mesozuic ages. A petmeable and porous ferruginous erust of the lalerile profile was also thought fo render this zone resistant to erosion, Howeyer, ferruginous crusts. are relauvely rare and discantinuous with the thickest crusts Ocedrring in positions well below the level of the postulated ancient surtace Gully gravure. involving the alteration of the locus of intense erosion through the protective influence ot vourse debris, was implied io reduce the rate of scarp retreat (Twidale \976a), However, no specific sites were disetissed and the present author has not observed the extensive operation of this process in the Mt Lofty Ranges. The unequal activity of rivers, whieh incise more rapidly than they erode laterally. was also suggested as a Contributury factor in summit surface preservation (Twidale |976a), While river incision may operune more rapidly than valley-side processes in some situations. che operation of the processes.of weatheriny, surface wash and pullyiny on valley slopes and bill summits for 200 Ma has Jed to considerable modihieation of the landscape (Milnes et al, 1985; Bourman 199%a), A wodel of landsvape evolution involving increasing: relief ammpliude inorder to account for the preservalion of these presumed ancient palaeoforms was presented. and evidence supporting this model for other areas, was discussed, However, itis unlikely thal the summit surface of the Mt Lofty Ranges has survived essentially unehunged for this enormous period of tine, Dixsection model The dissection model assumes ree mduralion at the top of former complete and continuous profiles and the Lack of preservation of complete profiles is often taken to imply dissection (Stephens 1946: Thomson & Horwitz 1962; Johns (96la, b; Maud 1972; Robertson 1974; Daily er af 1974; Twidale |983), Johns (196la) considered that much of the Lincaln Uplands of southem Eyre Peninsula is obscured by fossil laterite and lateritte gravels and conglomerates, a formerly continuous mantle now partly seripped following regwinal uplift, drainage rejuvenation and erosion. Johns (196/b) also interpreted the accurdanee of summit levely in the eastern Mt Lofty Ranges as a base-levelled lerrain of Phovene age, carrying sporadic cecurrences of terruginous grils and tatertles. He believed that onve-coutinuous ironstone cappings of Pliocene or post-Plioeene age have heen largely removed by crasion. The hest exposares of ironstones were reported from “Lucernbrae” where depustis abelut LATERITE [8 SOUTHERN SOUTH AUSTRALIA " Im thick were noted to mame Kanmantoo Group metasedimentary rocks. Mid (1972) now! thal although laterite profiles im the southery Mr Lofty Ranges are typically thick, with well developed ujottled and pallid zones, laterite horizons are rare. He ateributed this to erosional truncation of the profile, Robertson (1974) reported ironstone fragments and deeply weathered and kaolimised rocks in che central seetionof the Mc Lofty Ranges at about 450 m above sea level. He alse interpreted the weathered material as a remnant of a Tertiary laterite profile, Geologists and pgeomorphologists have been particularly interested in laterite, primanily to establish denudation chronologies. to estiblish the apes of particular landforms, to correlate widely spaced planation surlaces and to throw lighton the tetonic behaviour of upland areas. Examples of the use of latenue Weathering in interpreting landscape evolution are provided by the work of Sprigg (1945), Brock (19648), and Twidale & Bourne (1975a). Sprige (1946) considered Ialerrte formation in the Mt Lotly Ranges to be short-lived, correlated it with mottled Pleistocene sediments and believed that faulting and uplifr of a peneplain occurred after laterite formation, indicating land movernents of between 180 an ane 300 in during the Pleistocene Kosciusko epoch of block laulbing, This interpretation provides a very young age for lateritisation and faulting. whereas Brock (19645) interpreted the summit surface as a peneplain formed after prolonged subaerial weathering and erosion in the Palaeozoic, Culnmatag in a phase of crustal stability in the Mesozoic and Barly Tertiary, when latentisation occurred prior toupliit and dissection of the surlace, Even ereater antiquity of the Mt Lofty Ranges was proposed by ‘Twidale & Bourne (19754) who investigated the geomorphic evolution of the eastern Mc Lolly Ranges. A summit high plain (Tungkille Surfage) at 200) 300 im above sea level, ain etch surface, occasionally surmounted by scattered lateritic residuals up to 10 tm high (Whalley Surlace), was identified. The scattered Jateritic remnants were interpreted as remnants of a once-contiguous weathered surface of low reliet. The Whalley Surface and its agsiciated deep weathering were considered is be of Mesoroic age by extrapolation from Kangaroo Island (Daily er af, 1974), They also argued that it developed under a humid, tropical climate. Dislocation of the Whalley Surface by faulting was propesed although there was fy evidence of tured laterite on the downthrown side of the Milendella Fault. Tts absence, if i} ever existed, was explained by sub surilice dissolution of the iton oxides. Kennedy siodel of development of lmerttised surfer Twidale (INO8) accounted for jhe absenoc of dowataulted remnants of the lateritised erosion surface hy proposing an alternative to the traditional explanation of the suminit surface of the ML Lofty Ranges that it is an extensive literitised surface of erosion, develuped close bo regional base level in ihe Late Tertiary, and subsequently upthrust along ancient fault lines, after which it suffered dissection, While conceding that the ferruginous crusts of the postulated laterite profile might have been removed by sub-surface solution, the possible development of an extensive landsurface in relationship to loval base levels in the upper reaches, one of the possibilities suggested by (he work of Kennetly (962), was proposed, However, ou critical evidence was presented to show that the sumoniit surface of the Mt Lofty Ranges developed in thos fashion Alternatives te trancated larerite profiles While carrying oul regional geologeeal investigators on Yorke Peninsula. Crawford (1965) deseribed Pleistocene deposits, exposed in the sea-cliffs. at Ardrossan, as mottled dark red oi) olive green wrgillaccous sediments - The ‘Ardrossan Clays and Sandrock’ of Tepper (1879). He suggested that the mottling could be due to latentisation, with the upper indurated zone having heen removed by erosion and the pallid zone Oecurring sub-surface, However, fresh felspar gravel (i the mottled material argued against laterilic wealbering, Consequently, an ullermative non lateritic explanation of mottling produved by alternate wetting and drying in an environment of low relicf was also Suggested, Crawford (1965) obviously considered Iuterite within the framework of the standard laterite protile and attempted to fit his observations into it by postulating the erosional removal of an-upper indurated zone. He did, however, also consider an alternative non-la#teritic explanation for his observations, The validity of accounting for meomplete profiles by erosional truncation in landscape interpretation wits questioned (Bourman er ui, 1987; Bournan 1993a) by demonstrating great lateral variability in the spatial distribution of bleached, mottled and lerricreted zones, the development of which depended closely on local tmicne-environments (Bourman, 199338), Presurued remnants of lalerile crusts have heen shown lo be lags of ferruginous motiles waccumalating at the surface during landscape downwasting (Bourman 1894) and thus litterile crusts, as such, tay never have existed. Double planation theary af Fenner Fenner (1930, 1931) presented a double peneplanution hypothesis to account fof the evolution of the Mt Lofty Ranges, providing # geomorphic and (eectome (mevewurk lor the use of subsequent authors. The renter part of the Mi Lofty Ranges wis thought i 18 ROBERT 2 BOTIRMAN have been stripped of an easily eroded Miocene marine vovermass. He saw the double planation theory as necessury lo explain transverse drainage and exhumed surfaces in the ranges. He postulated chat a pre- Miocene peneplain blanketed with a Miocene marine covermass had been affected by block-fiulling. Wilting and ditlerential uplift in the Late Mincene or Barly Pliocene, Subsequently, this irregular surface was thought to have been peneplined, resurrecuing the oldec sueface ir places and developing a new peneplain on both Precambrian and Miocene rocks. Following this, Pieismacene (Koseiuska Epoch) teclomism renewed erosion. Some lault blocks retnained buried by Tertiary sediments and others, exhumed from beneuth the covermaiss, Were subjected to renewed weathering and erosion, Today it is generally agreed thal the Mt Lofiy Ranges Were not tulally immersed by the Miavene seas. so that the double planation theary cannot be decepted in its entirely, However, lirge reas of the Mt Lofty Ranges and Kangaroo Island (Milnes eral, 883) were covered! by Mincene seas ut heights in exvess of 200 m above sea level and there és evidence in the Bremer, Myponge und Upper Hindmarsh valleys that the shorelines were even higher than this (Bourman 19894). Moreover, ever hough the covermass of marine deposits may not hove totally covered the ranges, Tertiary lerrestritl sediments oceur eyrensively and at higher levels thar do the murme sediments. Consequently, the double plunation theory has considerable merit but it is still inadequate wo account lor all Of the geomorphic complexities of the ranges. which haye been variably exposed lo processes of weuthering, erosion and sedimentation for immense periods of ume, According to Sprigg (1945) laterite in the MI Lolly Runes formed on both a Precambrian an Cambrian hedrock undermass and a covermass ol Tertiary Nimestones and lacustrine sediments. Initially critical ol the double peneplanation theory, Sprigy (945) subsequently made the observation that the widespread ocenrrence Of laterite over the Mt Lofty Rangcs presented a polent argument in favour of this theory Lanwlscipes with multiple surfaces Landscapes with mulliple erosional surfaces have frequently been deseribed in South Austealia, with the suriaves beige marked by different weathering responses. Sonte examples follow, which illustrate varying interpretations of multicyelic landseapes. BuLkMAN, R. P. 11973) Georurpiic evelutin ol southeastern Fleurieu Peninsula, MA thesis. The Univernty of Adekide (unpubl. > Arey, NS, BE, (1960) The Camozote History of the Mid Nort of South Australia. MA, thesis, The Lmversiiy ol Adelaivee (uiypubh 4 Bourtman (1469). 19737). identibed a multicvelic landscape marked hy (wo major erosion surfaces, the Spring Mount Plateau. developed during the time from the Mesozoic tothe Rovene and the Green Hills Surtice of Pliocene age on Pleuneu Peninsula. The former. underlain by a literitic weathering profile consistine of pallicl. mottled and ferrugious-rich zones, oecurred at ahout 400 my above sea level. The surtace was considered to have heen tilled to the southeas|. The second erosional surface 170-L00 m above sea level, was capped in places by ferricrete, 4 term used ty desenbe iron-cemerted crusts nor underlain by deep weathering profiles. The Green Hills fernereted surface Was thought to have formed fram reworking of laleniie material [rum the summil surface, Lismg stranded river enivels and civer profiles, Bourman (19737) sugyested thal base level durjag erosion of the Green Hills Surface in the Pliocene wats approximately 60 7 above sea level when fluvial action modified a resurrected pre Mineene erosion surlice, Forrest (1969) exarhined the geomorphic evolution of the Bremer Valley in the easlern ML Lofly Ranges and wWentilied two erosional surfaces of low relieh Which he cémsidered) had formed prior to a mayce murine trinseression in the Miscene Consequently. both the surfaces and their ussociated cappings of lateriie material were interpreted ws pre-Miacene im ave. The Miocene sea was presumed to five transgressed anurea with relict similar to thal of boday and lateritisation of the bedrock was presumed to have followed the development of the Whalley Hill and Lucembrie erosion surfaces prior to the Miocene, Another surfuce, an exhumed one wilh a remnant of derived ferricrete, and thought to have formed by stripping of the Mincene limestone cover was considered fu be al Phiovene age. In a study of landsurface development in lhe Mid North af South Australia, Alley (1969*. 1973, 1977) identified reinnants of u laterite surface, occurring high in the landscape but below resistant quartzite ridges, Remnants of the laterite surface were noted Lo he most cammon at stream-heads hut also to pecur an prominent hills that sland newrly 100m abeve modern valley Moors. ‘Pho luterite capping of angular quariz jraginents set in u matrix of iron oxides was observed lo overlie severely Weathered ind locally kaolinitic bedrock and (o be consistently thicker ory lower slopes Seeuons of the laterile surface were lhought \o have been dowe-lautled inthe Ear Tertiary and laler burial by Middle “Ternary sediments. A consistently lower silerete-capped landsurface was considered to be younger than the laterite surtice. 1) interpretations of mulficyelic landscapes and (he TecoysniGoan of the ages of difterent landsurluces (here ure inevitahly many disagreements and warkers in South Australid have not escaped these. An oxaniple follows, King (976) reenenised several of hes world LATERITE IN SOUTIIERN SOUTIT AUSTRALIA 1) wide erosional surfaces in the Mt Lofty Ranges of South Australia, He considered that south of the Willunya Fault, ‘laterite-enerasted tablelands’ represented the Moorlands planation (‘great Australian denudation cycle’) of Late Cretaceous to middle Cainozoic age, whereas north of the fault. the Mc Letty Ranges were thought to be surmounted by bis Rolling, landsurface of Miocene age that Jacked a true saterite, The Widespread landscape of Pliocene ape wis recoymised in broad valleys and basins accordant with 4 Pliocene coastal plain at about [80am nbove sea level and the Youngest Cycle was related lo deep valleys and gorges in the ranges, On the other hand, Twidale (1978) considered that Ihe summut surfaces both neh and south of the Willunga Fault were contiguous and of the same earls Mesozoic age, However, areas of laterite mapped by Twaidale (978) north and west ot the Willunga Fault onthe Eden and Clarendon Blocks are variably covered with weathered and ferruginised Eocene ty Pliocene sediments (Sprigg 1942, (940; Ward 1966), In places these have been eroded to expose an underlying: weathered pre Tertiary surface, eroded and tewedthered since exhumation (Sprige 1945). Consequently. the summit surface here cannot be ot carly Mesozoic age, Furthermore, there may be some support for King’s generalised scheme, as weathered zones have been stripped (rom large areas of Lhe summit sunface north of the Willanga Fault, espectally in the eastern Mr Lofty Ranges (Twidale & Bourne 19754), allowing further erosion and the potestial development of a younger surhice, possibly equivalent to King’s Rolling. surface. Age of laterite [lis very difficull to aseribe ages to laterine materials because they may have developed over logy Orne periods, samc inay have several possible modes of genesis, others. aré polygenetic having been considerably reworked and reweathered, and there are severe limitations on daling techniques applied to weathered materials (Bourman 19939), Furthermore, there ure fiely constraining sediments, These difficulties are apparent in South Australia, where Jaterttsation has been ascribed to periods fram the early Mesozoic to the present, There have been many assertions about the age of laterite in South Australia, offen withoul presentation ot convincing evidence. There has also heen a tendency to prescribe a single une of lateritisaon, When evidence of the timing has commnly been derived from limited study areas, front where there has often been widespread extrapolation. The following discussion of evidence presented by different workers in South Australia highlights the great yurtability | the ages attributed to lateritisation Early-Middle Tertiary Woolnough (1927) considered that faterttisarion eccurred during one period, in the Miocene, and many subsequent workers have generally supported the view of a ‘Yeruary age for lateritisation (e.g. Preseort & Peridleton 1952) but mot necessarily inthe Miocene. Aitchison et al (1953) reported Early Tertiary lacustrine motued sands, argillaccous sandstone and clays occurring sub-shorontally on a pre-Tertiary erosion surface in the Adelaide area, implying lateritisubon in the Teruary. und Campana (1955) favoured widespread lateritisation between the Iurly Bovene and the Miocene. Sections of a hiterite surface in the Mid North ot South Austalia were thought to have been down liulled in the Furly ‘Tertiary und later buried by Middle Tertiary sediments, so that Alley (1973) regarded the Jaterite surface to be of (?)Early to pre-Tertiary age and considered that it persisted until the Middle ‘Tertiary in the Barossa area. Lron-stained rounded quartz gratos and ferruginous pellets were reported from within a fossiliferous marine limestone of probable Upper Eocene age (Bourman & Lindsay 1973). intersected in a drill hole att 36 i underlying part of the Waitpinga Creek drainage basin, anid aban elevation of about 60 m above sea level This observution was. interpreted as indicauny the deyelopaient of fateriisation, or at least ferruginisation, prior te the Bocene, Phocene Many concurred with Prescott (193]) and Whitehouse (1940) that there had been widespread laterite formation throughout Australia in the Pliogene (e.g. Stephens 1946 Crocker 1946; Northoote 1946; Rix & Hotton 1953; Johns W6la, b). Fenner (1930, 1931) also implied a post-Miocene or Phincene age for laterite in the Mt Lofty Ranges. Working on Fleuneu Peninsula, Crawford (1959) identified laterite capping Wilson Hill at 320 m, around which af area Was mapped as the lower part of a katerite profile developed on Kanmantoo Groyp metisediment tary rocks. The occurrence of areas of hard laterite at lower elevations (100 m above sea level), on quartzose sediments. was interpreted as indigaunme a yery irregular original lateritised surface of Lale Tertiary ave. Subsequently, Bourman (1973) demonstrated thot the two decurrences were distinetive and probably of (wo difterent ages, with Miocene limestone separating the two types, The evidence presented by Horwilz (1960) for mayor lateritisation in the Pliocene is also equivocal, In the iritramontane Upper Hindmarsh Valley of Pleuricu Peninsula over 150 m of cross-bedded and mottled brown ferruginous sands, capped by crust of limonite- ROBERT P. BOURMAN ms cemented gravels, were reported, By extrapolation oe 8 8 8 these were considered to overlie fossiliferous Early 1 ee el Miocene limestones. The sands were thus tentatively assigned to the Pliocene. Horwitz also considered that by these lateritised Pliocene sands were continuous with ws limonite-cemented gravels on the high plateau (Fig. 9). ae z, Thus he assigned them to the same Pliocene age. Brock E © (1964°, 1971) questioned the contemporaneity of the Roe high-level and low-level crusts and Bourman (1969! , a o£ 19737) highlighted their different characters and m % & suggested that the higher crust was of pre-Miocene age a 2 E und the lower one of Pliocene age. G o 2 Harris & Olliver (1964) reported on palynologicul E 8 @ analysis of organic material preserved in “coal balls” cS @ exposed in Tertiary sands in the Barossa Valley. The basal Tertiary unit was described as a lJateritic sand and gravel overlain by laminated silty and sandy clays and was considered to be of Early Tertiary age. The clays were capped by an upper laterite. Previously the sands had been assigned to the Eocene (Glaessner 1955) or Pliocene (Hossfeld 1949) but Harris & Oliver (1964) suggested that the microfloras indicated a Miocene or possibly an Early Pliocene age for the sediments, Twidale (1968) concluded that the deep weathering in the Mt Lofty Ranges oceurred late in the Tertiary (Pliocene) and may have even conunued into the early part of the Pleistocene. Major & Vitols (1973) suggested that ferruginous pisolites on the western end of Kangaroo Island were of Late Pliocene or Early Pleistocene age as an aeolian calcarenite (Middle Pleistovene Bridgewater Formation) was thought to overlie pisolites, and elsewhere blocks of ferruginous " pisolite were noted to overlie marine limestone of probable Late Pliocene age. —? with the locality record indicates thar the taxon was listed with a “ef” and is pot positively identified Irom that State/Territory, Unless specifically noted. all records are from Australian inland waters, both fresh wod athailasie saline. Families are treated systematically in the sequence as given by Smirnoy & Tinims (1983), However, the /yoeryplus species are separated into the family Myveryplidae as proposed by Smirnoy (1992). For convenience genera and species within cach family are listed alphabetically. In the author citations, two authors who are sometimes confused are separated as follaws: (O,F.) Miiller, with urmlaui, who published in the late 1770s-80s and (P.E.) Mueller. with ve, who published in the 1840s. Kamily Sididae Baird, 1850 Diaphdnosoma Fischer, 1850 “PD. australiensis Korovehinsky, 1981; Qud (Korovehinsky 1981). Later finds all in Qld DPD excisumn Sars, 1885; Old (Sars 1885); NSW (Jolly 19646): SA (Shiel er al 1982): NT (Paice 1982'): WA (‘Timms 1988) D. sarsi Richard, 1894; Qld (Korovehinsky [981); NT (Tait eral. (984), WA (Timins 1988) “LE unguiculaium Gurney, 1927; Qld (Gurney 19271; Vic,, NSW (Walker & Hillman 1977); SA (Shiel et al, 1982): NT (Tait 1982!) WA (Brock & Shiel 1983); ?Papua-New Guinea (Korovehinsky 1992) D. vulzi Stingelin, 1905, NSW (Korovehinsky 1981) Tati, R, D. (1982) Plankton of Magela billabongs, No M)Sc, thesis, Macquarie University, unpubl, “Ds. BV. (1973) A comparilive study of the linmalogy ol three maar lakes m western Victorat. Ph.D. Thesis, Monash University, unpubl. Latonopsix Sars, 1888 L, australis Sars, (8&8: Old (Sars (888); Vic., NSW (Shel 1978); NT (fulli 986); WA (Timms 1988) °L. brehmi Petkoyski, 1973; WA, NSW (Petkovshi 1973b); NT (Juli (986); Qld (Timms 1986) Penilia Dana, 1852 P. avirostris Dana, \852; NSW (murine, coastal) (Dakin & Colefax 1940) Preudosida Herrick, 1884 *P susirafiensiy Smirnov & Timms. 1983: NSW (Korovehinsky, in Smirnov & Timms 1983) FP. szalauyt Daday 1898; Old, N'T, WA (‘Timmins 1988) Sarsilatena Koroyehinsky, 1985 3. papuana (Duday, O01); Pseudasida papuana Daday. 1901; Sursi/atona papuana: Korovehinsky (1985); NT (Korovchinsky 1985); Qld, WA (Timms 1988) Family Podonidae Mordukhai-Boltovskoi, 1968 Pleopsis Dana, 1852 P. polyphemoides (Leuckart, 1859): E. Australia (marine) (Dakin & Colefax 1940) Psendevadne (Claas, 1877) & tergestina (Claus, 1877); E, Australia (marine) (Dakin & Colefax 1940) Podon Lilljeborgz, 1853 P. intermedius Lilljeborg, 18532 View (estuarine) (Neale & Bayly 1974) Fvwidne Loven, 1836 &. spinifera Mueller, 867: E. Australia (murine) (Dakin & Colefax 1940) &. nordmannl Loyen. 1836, BE. Auswalia (marine) (Dakin & Colelax 1940) Family Chydoridae Stebbing, [802 Acroperts Baird, 1843 A. alanvides Hudendorff, 1876: NSW (Smirnow 971); Qld (Timms J988) A. Hlarpae (Baird, 1834); Lynceus harpae Baird, S342 deraperus herpee: Baird (1843). NSW (Smirnov 1971) AL neglects Liljeborg, 900; Acroperus avirestris Henry, 1919: Smirnoy & Timms (1983); NSW (Henry 1919) *4_ sinvarus Henry, 919: NSW (Henry 1919) Alon Baird. 1849 A archer’ Sars. 1888; Qld (Sars 1888) *4_ beverlevae Smirnov, 1989; Qld (Smirnov 1989n) A, cambiuei Guerne & Richard, 1893; NSW (Henry 1919), Vie (Shiel 1976); Qld, NT, WA (Tims 1988) %4, clathrata Sars. |888: Qld (Sars I8&88), NSW (Henry 1922) A, caslate Sats, 1862; NSW (Smirnov 1971); Vie (Vimar 19732): NT (Tait 1982!) CLADOCERA RECORDED FROM AUSTRALIA i A. crassicaudait Sars, (96; Qld, NT. WA (Timms 1958) A, diaphana kang, (853: Alonella diaphana (Kings: Sars (888); Alona david? Richard, 8S: Frey (199la); Alona david var thering? Richard, 97 Frey (19912), Alona punctaia Daday, 198, Prey (199la), see Prey (1991a) for comments on synonymy; NSW (King 1853); Qld (Sars 1888): View (Shiel 1976); SA (Shiel 19814); NT (Tait e¢ al. 1984); WA (Timms 1988) A. ynitand Sars, \R62; Alona mricrotata Henry, 1922: Smirnov & Timms (1983); NSW (Henry 1922); Vic. (Timms 19732); SA (Shiel et al, 1982); NT (Tait er al. (984): Qld. WA (Timms 1988) A. imreticalata Shen Chia-jui, Sung Ta-hsiang & Chen Kuo-hsiao, 64: Vie. (Morton & Bayly 1977); Tas, (Sniimey & Timms 1983) “ad, investix Smirnoy. & Timms, 1983; Vie. (Smirnov & ‘VYimins 1983) “4. laevivsima Sars, IS88; Qld (Sars 188K): NSW (Henry 1922) “4, tlaecracontia Synimnoy & Timms, 1983; NSW (Smirnov & Timms, 1983) AL mondedntia Sars. 1901; NY Guth t986), Old (Timms 1988) A, poppe? Richard, (897; Vie, (Shiel 19813) A, pulehella King, 1853, NSW (King 1853); Vie, (Shiel 19814), "Qld. NT, WA (Timmins 1988) A, quedrangularis (Miler, 1785); Lyaicens quadraneulars Miller, 78S: Alana quadrangularix: Smirnov (97D. Vie. WA (Smirnov & Timyns 1983) A, rectangula Sars, 62: Old SA (Smirnov 1971); Vic.. (Shiel 1976). NSW (Walker & Hillann 77). NT, WA (Timms 1988) “4. serdloides Snvirnov & Timms, 183; WA (Smirnov & ‘Tins 1983) A, sirtolater Sars, 1916; “tropical Australia” (Srnimov 1989a) ‘4, truncaia Smirnov, 1989; Old (Smirnov 1989) “A, unguiculaia Smimov, 1989; Old (Smirnov |98%a) Aleomella Sars, 1862 Al elathranda Sars, 1896; NSW (Sars 1896), SA (Shiel J981%): NT (Julli 1986); Qld (Timms 980): WA CLinms 1988) 4 excise (Fischer, 1854): Lynceus excisus Fischer. 1854: Alonella exeiva: Sars W62b; NSW (Henry 1922), Vie. (Shicl 1976); SA (Shiel 19817); Old (Hawkins I9KK) YWA (Bayly 1992) ‘Su, Ro (98t) Plankton of the Murray-Darling over system, with particular reference to the zooplanktor. Ph.D. Thesis, University of Adelaide, unpubl. ‘Mortons, 1D, W. (1973) Studies an some temporary Victorian witlers with special reference 10 the Microcrustaven, B.Sc, (Hons) Thesis, Monasl University, unpubl. A. exfeva (Lilljehorg, [853); Lyneeny exiguns Lilkjeborg, 1853; Alonella exigua: Mueller, 1867; NSW (Smirnov I97l): NT (Tait er al. 1984) tArchepleuroxus Smirnov & Timms, 1983 “4. beryl Smirnov & Tintms, 983; Vic., Tas.. WA (Sturmoy & Timms 1983) Adustralochvdorus Smirnov & Tininis, 83 44. gporus Smirnov & Timms, (983; Old, NSW (Smirnov & Timms (983), NT (Tit er al, 1984); WA (Timms 1988) Biapertura Smimov, 1971 *B, abbreviate (Sars, 1896), Alona abreviata (sc) Sars, 1896: Biapernira abreviare (sic): Smirnov & Timms 1983, NSW (Sars 1896) Comment! The spelling of the spectes name with d sige b as abreyiata in the original description (Sars 1896; 40) appears to be 4 typographical error, as 1 is later spelt (p.43 text: p. 79 fig. caption) as abbreviara. B. affinis (Leydig, 1860), Lynceus affinis Leydig. 1860: Alona whirelevgii Sars, 1896: Henry 1922. Alona affinis: Sars 1901, Alona longirasirts Henry, 1919: Smirnov 1971; Biapertira affinis: Smirnov & Timms 1983; NSW (Surs 1896), Vic. (Timms 19732), WA (Williamns 1979); SA (Shiti 1981"); NT (Tail 1982'); Old (Tinuns (986) *B. duoodonta (Heary, 1922); Alonella ducodonra Henry, 1922; Biapertura duoddoata: Smurnoy & Timms 1983; NSW (Henry 1922) INT (Tait ep al, 1984) *R imitateria Smirnov, 1989: WA (Smirnoy [98&9a) B. intermedia (Sars, \462), Alone intermedi Sars. 862: Biapertura intermedia: Smirnov 971, Qld (Gurney 1927); NSW (Smirnov 1971); Vic, (Shiet 1978); WA (Bayly 1962): NT (Timms 1938) B. karwa (King, 1853); Alona karua King, (853: Alonella kara. Sats, W888. Bapertara karna: Smirnov & Timms 1983; NSW (King 1853); Old (Sars, I88%); Wie. (Morton 19734), NT (Tait 1982'); WA (Timms 1988) Comment: King’s description is inadequate hy modern shindards. There are differences in the post-abdomen morphology of his species and that later hatched frean Qld mud by Sars (1888), although Sars considered the taxa identical, There ms now good evidence that & karua represents 2 species complex worldwide (Alonso & Pretus 1989). In our opinion the O00) km separation of the King and Sars taxa is sufficient to doubt conspecificity, hence their respective identities are not satisiactorily resolved at this time. The problent is compounded by eyrors in Smirnov & Timms (1983) (see Incertae sediy below). *B. kendallensis (Henry, 1919); Alona keridallensis Henry, 1919; Bianertira kendallensis! Smirnov 1971; NSW (Henry 1919), Qld (Smirnov 1971): Vie. (Timms 19732), NT (ulli 1986); WA (Grows ef al, W925 a2 RJ SMIEL & 1A “R. lonsiqua Smirnov, W871, NSW, Old (Smirnov 97h; Vie. (Shiel 19815). WA (Timms 1988) "B. macrecaper (Sirs, O94): Alona macrocoper Sars, 1894; Bianerrera macrecopa: Smirnov & Tinos 1983 (author dale given us 1895); Qld (Gurney 1927); Vic. (Morton & Bayly 1977); WA (Bayly 1982); NSW (Timms 1982) *B. rivuticandix Smirnov, OT, Alena intermedia Gumey 1927 (misidenuification): Biaperura rividicaudis, Smirnov 1971: Qld (Gurney 1927): Vie (Shiel (976); NSW (Timms. 976); SA (Shiel 1981), WA (Bayly 1982), NT (ull 1986) "Bo rusticaidey Smirnoy & Timms, 1983; Tas, (Smuirnoy & Timins 1983) RB. setigera (Brehm, 1941); Alona vattani seligore Brehin, 1931, Alana setizeni: Petkovski |97Au: Biaperiura setigera. Smarnoy & Timms 983; NSW (Bayly 1970): Vie, (Shiel 1976), SA (Shiel ORE): Old (Timms 1986) ?WA (Storey er al 1993) B. verracoase (Saes, WO, Alona verrucosa Sars. 901: Alone reclanwule pulehre Hellich. 1874 Smiron IW7l and Smirnov & Tirnms 1983; Biupernira verrnceusas Smupnoy 9895 Old (Smirnov 197: NT Gull 986): WA (Timms 1988) *B. willisi Smumdy, (989; Old (Smirnov 1989) Camptacercus Baird, 1843 *#C> australis Sars, 1896, NSW (Sirs 1896), Vic. (Shephard eral '918): Qld (Smirnov 17> NT (Tuit 1942"); WA (Timmins 1988) *Celsinonine Frey. 199] *C. Aypsilopham Frey, 1991, NSW (Frey 19910) °C. putraneusis Brey. 9) NSW (Frey 19914) *C. plataumentes Frey, 991; NSW (Frey, 1991) Chydorus Leach, |16 CC. eurvantus Sars. Ol: Qld (Timms 67); Vie (Walker & Hillraan 1977) C. herrmanni Bret, 1933; Qld (Timms 1967); Vie. (Shicl 19814) C. kallipyeos Brehm, 1933, NSW (Petkowskr 19738): Old (Hani (975%) Comment. Smienoy & Tints (983) regarded Perkoy - skhS record as a mistdentification of C. enryneuty, However Hann (1975°) independently recorded C. kal fipvges from NSW and Qld. Petkovskrs record should stand unlil.a thorough revision of the genus is mude- “CL ohscuiirosteis Brey, RT, NTL WA (Prey 1987) *C. opachy Frey, W87: NT. Qld, WA (Frey, 1987) ©. parvuy Daday, (898; “tropical Australia” (Smirnoy O89) © pubescets Sars. OL, NT Qld, WA (Timms 1988) C. rencufarns Daday, WWY8; “tropical Australia” (Smirnoy lk9a) “lass. BLD. 1975) Taxonomy ot Chydoridae m Ontara and genus Chvderis worldwide MS Thesis, Unversity af Watertoo Ontario, ripubl. DICKSON Comment: This specres was listed without comment by Smirnov (989), Ibis given as a synonym of © sphaericuy (Miller) in Flassner (1972), The relitton- ship of this taxon to the other ‘feavifarnislike relicu- jated taxa deseribed by Prey (1987) remains unresolved. We consider it unlikely to be Daday’s species. C. sphaeriens (Miller, 1783), Lynceny sptheertony Muller, 1785; Chyvdorius sphaeriens; Baird 1843, Chydarus cleland; Henry, 1919 was synonynnzed with Chydorus leonard? by Henry (i922), Chydarus leonard King, 1853: Smirnoy (1971) C, leonard’ was attributed to Sats. [896 by Smirnov & Timms (1983); NSW (King 1853); Vie. (Morton 1967), Qld (Tims 1967): NT (Tait 9829) SA (Shiel e7 a, 82), "WA (Bayly 1992) Comment: In view of the restricted distribution of Chydorus sphaericus s.sue Brey 1980), it rs likely that a complex of species accurs in Australia, none of whieh is Ihe nominate taxon (D.G. Frey pers, comm.) Dadava Sars, 1901 Do meacrops (Daday, 1898): Alans macropy Daduy, 1898; Daedeya macrops: Sars OL, Qld (Srainey OTD, NT (Tait-ez al 1984); WA (Timms 1988) Dispuralina Pryer, 65 D. weurvesrris (Birge. IN79); Plenroxus acalirostriy Birge, IK79. Akmelle acutirastris; Prey W59 Disparalona acutirestris, Fryer J971, “tropwal Australia” (Stairnov 1989a) Duntievedia King, 1853 D, erasyu King. 1853: NSW (King 1833); Qld (Sars la88); SA (Henry 1922); Tas. (Brehm IWS3a), Vic. (Marton 19734), NT (Tait eral, 1984): WA (Timms [9&8) Ephemeroparus Frey. (982 2. tridentatus (Bergamin, 193912 Chydonets tridemarus Bergamin, 1939: Chydorny barreiy Richard, (894, Fig, 329 jn Smirnov [97], Ephemeroporus trideniatus; Frey W8la; Qld (Smmienoy 1971): Vie. (Shicl 19814): NT (Tait ef al, 198A. WA (Timms 1988) Comment: see /neertue sediy tor other taxa referred to this genus in Australia. Eurvalona Sars, (901 F arientalis (Daday, (898); Alonapsis avientelis Daday, I8Y8. Evryalona acctdenralis Sars, OO: Smirnov I971; Auryelena erientalrs: Daday 05, Old (Sryirnew de Timms 19K3)) NT (Tait eral (984); WA (Timms 1988) Grapteleherts Sars, (62 GG lesindinaria Crhiseher, W48)2 Lincens texindinariuy Fischer, [848 (cited as IRSL in Smirnw & Timms |[1983]); Graploleberis lestucinarie, Kure. 1874; NSW (Henry 1919): Vie Shuel W76) NT (Tait 19824. Old WA Timmins aS) CLADOCERA RECORDED FROM AUSTRALIA 4 Kura Dybowski & Grochowski, 1894 A. latissime (Kurz, 1874), dlonopsis latissima Kurz, 1874; Kursia latixste Dybowski-& Grochowski R94, Vic. (Shiel 1976) RK. lungirostris (Daduy, 1898); Alona longirostrix Daday, 1898; Kursia longirostrix: Harding 1957; NSW (Timms 1972), NT (Tail ef al. 1984), Old, WA (Timms 1988) *Leberis Smirnov, 1989 “L. aenigmataya Survey, 1989; WA (Smirnov hb) Leydigia Kurz, 1874 L. acanthocercoides (Fischer, 1854), Lynteciey acaunthaeercoides Fischer, 1854; Levydleia acanthovercuides, Kurz, 174. NSW (Tins 1970), NT (ull 1986); Qla, WA (Timms 1988) {., australis Sars, BRS; Qld (Sars 1885); NSW (Shiel 1978). View, SA (Shicl 19813) L. ciliata Gauthier, 1939; NSW, Qld (Smirnow 1971); Vie, (Shiel 19814) *L. laevis Gurney, 1927; Qld (Gurney 1927): NSW (Snel 1981°): WA (Growns er ai. 992) L. fleydigi (Schoedler, 1863): Alena levdivit Schoedler, 1863; Leydivia leyiligi: Daday 1902; SA (Henry, 1922), Vie. (Shiel 1976); NT (Tait 19821), WA (Growns ¢f al. 1992), NSW (Kobayashi 1992) *Monope Smirnov & Timms, (983; Monoports Smirnoy. 1977: Smitnoy & Timms (1983: 34) *M. reticulata (Henry, 1922); Pleureaxus reticularis: Henry 1922: Monoparcs henrvyae Smirnov, 1977; Monope reticulata: Smirnov & Timms 1983; non Plenroxus reticulatus. Henry, 1918; Frey 199 1b: NSW (Henry 1922): WA (Bayly 1992) Coynment: Henry's (1918) taxon as figured is, according (o Frey (1991b). a species of Alonella, probably 4, cluthraruld Sars, 1896. Monospilus. Sars, 1862 “M, diporns Stainoy S ‘Turns, 983, SA (Shiel 1978) (as Monospilus sp. nov.), WA (Brock & Shiel 1983) (as Monaspilis sp); NSW (Shiel 1981') (as Monaypilas no. sp. Dy Vie, (Shiel & Croome, unpub), data) *M. vlongains Smirnov & Timms, 1983; SA (Shiel Wsl3) (as Monespiluy n. sp. 2) Comment: Neither of these taxa is referrable te Monaspilas s. str; indeed they are probably sor even vongeneric (DG, Frey, pers. camm.). Noralana Rajapaksa, 1986 N. vlobulasa (Daday, 1898); dlona globulosa Daduy, 1898: Notealona glebulosa: Rajapaksa & Fernando 1987. The nominate species iy not recorded from Australia, However a geographic subspecies ts kruwn: CW. globuloxa australiensis (Rajapaksa & Fernand, 1987); Indiqlona (Petkoveki, 966); Rajapaksa a Fetnando IS&T) NT (Smurnoy & Timms 95 as Indialona sp.); (ld (Rajapaksa & Fernando 1987; WA (Timms 98%) Comment: Indialona was reported {rom the NT by Smirnov & Timnis (1983), species not piven. Julli (L986) reported J. globulosa. also from the N.T As Rajapaksa & Fernando obtained their material frorn BY. Timms, who also collected the Smirnov & Timms material and identified the Julli material, it ts probable that all NT. records are N. globules austrelienscs. Oxyurella Dybowski & Grochowski. 1804 Q, singalensis (Daday, 1898); dlonepsis singalensis Daday, 1898; Oxyvurella singalensis: Smurnoy 1971, Old (Smirnov & Timms 1983). NT (Tolli 1986); WA (‘Timms 1988) OL tertuiceicdis (Sars, \862); Alona tentuicatdis Sars, 1862) Alona wallaciana Heaty, 919, Oxyurelle wallaciane: Sinimoy WT); Oxyurella tenuicauelis: Smirnoy & Timms i983; NSW (Henry 1919): Vic. (Timms 1973?) (as Oxyurella sp. *Planicirelas Frey. 1991 *P) alticarinatus Prey, 991; WA (Prey 199\b) *Plurisping Frey, 199] *P. chuulivdiy Frey. 991; WA (Brey 1991b) #2, multituberculata Frey, 991. WA (Prey 1991b) Pleuroxus Baird, 1843 *P foveatuy Frey. 1991: WA (Prey 1991) *P inermis Sars, 1896; Chydonis denticularus Henry, 1919; Frey (1991b); NSW (Sars: 1896). Vic. (Haase 1903); Qld (Gurney 1927): SA (Shiel 1981; WA (Bayly 1992) *P jugosuy (Henry, 1922); Chydorny jugosus Henry. 1922: Smirnov & Timms (1983); NSW (Henry 1922) Comment; Frey (J991h) states thal close study of the type specimen did not reveal cnough positive characters to towke 4 firm decision (regarding Pleureauy cf. jugosus), and hence this taxon, al least forthe present, must be regarded as a nomen dubiumn. Pleuroxus Jugesus io Smirnov & Timms (1983) ws not Chyderus Jugoses Henry, 1922: rather most of the description und all of the illustrations in this paper are lor Plarispina chauliodis Frey, 1991. *P. kakadidensis Smimay, QR NT (Smirmov 1989b) P laevis Sars, 1862; NT, Qld, WA (Timms 198%) PF similis Vavea, O00: Po ausralis Henry, 1922: Smimoy & Thrinis 1983: NSW (Henry 1922) Comment: Probably absent from Australia (Frey !991b) *P mocellarus Smunnov, 989. WA (Sutirtioy 1989b) Pseudochiydarus Fryer, 6% P. globosus (Raird, 1844); Chydoruy globosus Baird, 1843; °Chvdoerns aveustis King, 1853: Sars (888); Pseqdoclivdarus elobosus: Fryer 1968; NSW (King (854): Vie. (Shephard er af. 1918); Qid (Timms & Midgley 1969); SA (Smimoy 71, NT, WA (Timms (988) Ja R, J. SHIRL & J A. DICKSON Rak Smirnoy & Timms, 1983 +R. labrasus Smimov & Timms, 1983: SA, Tus. Vic WA. (Sotirnoy & Timms (983) *R. obtusus Smirnov & Timms. 1983, NSW, WA (Smirnoy & Thrans 1983); Qld (Timms 1948) Comment; Several new species of Kak from W.A, are included in an incomplete MS by the late D.G_ Frey. He also found Rak in South Africa The Rak MS will be completed by RIS. *Khynchochyderus Smirnoy & Tirniis, 983 *R, australiensis, Smirnov & Timms, 19823, Amblyaryachis: Bayly 1992 (nemen nude), NSW (Smirtiov & Timms 1983); WA (Bayly 1992) eSayera Sars, 04 “5. cnoki (King, 1866); Euryeereus cookl King, 1866; Saycia erbicularty Sars, 904: Smirnoy 1966; Suyela coekt: Smirnoy 966: NSW. Qld (King 1866); Vie. (Sars 1904) Comment: Alter examining a N.Z. population, Frey (1971) coneluded that it represented a new geographic subspecies, Saveia cook? novaezealandiue Prey, 1971, The Australian subspecies is designated Saycla ceokr cookt (King, 1866): Smirnov & Timo 1983 Family Tlyocryptidae Smirnayv, 1992 Hyveryptus Sars. \862 1. hrevidentainy Ekman, 905; Vie. (Shiel 19813), NT (Tait et al. J984) “1 raridemaris Smirnoy, 1989; WA (Smirnov !989b) 1, sordidus (Lievin, WAR), deanhocereus sondicus Lievin. I848: Mywerypiis serdidus: Sars. (896; NSW (Sars 1896): Vie. (Henry 1922): SA (Shiel OBIS). NT (Tart et uf, 1984) 1. spinifer Herrick, 1882) 1 dongtremis Sars, 1888" Smirnoy & Timing (983), 7) ahi Brady, (886 in Gurney (1927); Smirnov & Timms (1983): Old (Sars, 1888): Vie. [Timms 1973): WA (Walliams 1979), SA, NSW (Shiel 1981); NT (full 1986) Farnily Mactrothri¢idae Baird, 1843 Grimaldina Richard. 1892 G. braszat Richard, 1892: Qld. WT (Timms (988) Macrethrix Baird, 1843; Eehinisca Lievin, 48: Smirnov 1992 tM. breviseia Sinirnov, 1976; Qld (Smirnov 1976); Vie. (Shiel 19814); WA (Growns. er ai. 1992); INSW (Timms 1993) M. capensis (Sars, 1916); Behinisea capenstry Sars. 16; Smimov 1992; Vic (Smimney 1974). NSW (Shiel 19812), Tas., WA (Smirnov & Timms W&3). Old. NT (Timms 1988) SMITCHELL, B.D. 980) The coology of waste stubrliziation ponds. Phy, D. thesis, Liniversiry of Adelaide, unpubl ~M. carinata (Smirtiov, 1976); Echinisea carineara Smirnoy, 1976> Smirnov (1992), Qld (Smirnoy 1976), NSW, Tas., WA (Smirnov & Tintms 1983) *M. flabelligera Smirnov. 1992; Qld (Smirnov W992) +M_ flagellate (Smirnov & Timms. 1983): Eehinisea Hagellata Sinirnoy &¢ Timms, 1985; Sininev 1992; Tas: (Smirnov & Timms, 1983) 4M. harding? Petkovski. 1973: Echinisea hardingy- Smirnov 1976; WA (Petkoyskt 19736); NSW (Shiel 19814) M. hirsuticorniy Nortivan & Brady, 1867) Vie (Smirnow 1976); SA (Milehell 19804) 4M hystrix Gumey, 1927: Old (Gumey 1927): NT (Juli 1986) +M. indisrinceta Smirmev, 1992, NSW, WA (Smirnov 1992) *M. longixeta Smirnoy, 1976; Vic. (Smimov 1976); “tropreal Australia” “Tas, (Smirnoy 1992) M. mafavensis Idris & Fernando, 1981, Old (Tinwes, OBR) +M. pectinata (Smirnov. 1976); Eehinisca pectin Smirnov: 1976; Smirnoyw 1992; Old, Vie, (Smirnoy 1976); NSW (Smirnoy & Timms 1983) M. rosea (Laevin, IB48); Echinisea rosea Lacwit. i848: Smirnov 1992; Old (Smirnov & Timms 1983) M. schaunislandi Sars, 19035, Macrothriv bursialix Smith. 1909: Smirnoy & Timms (I9B3)2 Tits (Smith 1909), Vic. “tropical Australia” (Somrnoy (992) M. spinesa Ring, 1$S3; NSW (King, 1853); Vie, SA (Shiel l981*) AM. rimnist (Smirnov, 1976); Behinisea tinmnsi Sournoy, 1976: Smirnay 1992; NSW (Sunrney 1976); Qld (Tinwns 1986) M_ trisertalis Brady, 1886, Fohinisea triserialis Smirnov 1976; ?NSW (Henry 1922), Old, Vic,, SA (Smirnoy & Timms, 1983); NT Chulli 1986); WA (‘Tinims 1988) *M. williami (Smirnov & Timms, 1983); Behinise williams: Smirnov & Timms. 1983) Snvirnov 1992; Old (Smirnov & Timms. 1983); NT (oll 1986) Neothrix Gurney, 1927 WEN, armata Gurney, 1927; Old (Gurney 1997); Vie. (Morton 19734); WA (Bayly (982); NSW (Kobayashi 1992) +N paucisetosa Smirnov, W89hy Marrorhpiy paucisetosa Smirnov Y89b: WA (Smirnov 1289b) +N. superarmat Sinimov, 1989b, Qld (Sinimoy (989b) “*’Pyendemoina Sars, 1912 +P lemnad’ (King, 1853), Moina lemnae Katy. 1853; Psevdomoina lemneae Sars 19125 NSW (King 1853); Vie. (Shephard er al, 918); Tas. Smirnov & Tins. 1983); SA (Shiel & Koste 1985) CLADOCERA RECORDED FROM AUSTRALIA 34 Streblocerus Sars, (62 S. serricanmedatuy (Pischer 1849); Daphnia laticarnis Fischer, 1849: Srreblecerus Yerricaudurus Lilljeborg, 1900: Smirnov (976; Vie. (Smirnov 1976); Qld, Tas. (Smirnoy & Timms 1983) Family Moinidae Goulden, 1968 Moina Baird, 1850 “KML anstraliensis Sars. (896: NSW (Sars 1846); Vic. (Shiel W814); WA (Smirnov & Timis 1983), NT (Tait eral. J984) *M. baylyt Porro, 18S; Moina mongelica Daday, 9QL in Bayly (1976), Smirnoy (1976). Smirnov & ‘Yinuns (984) (misidenritied): SA (Bayly 1976); NSW (Williants 1986); Qld (Timms 1987) “M1 fletuosa Sars, 1897, WA (Sars 1897) M. micrura Kury, 174: Moina propingua Sars, 1845: Goulden (1968); Moina dubia Richard in Gurney (1927) (misidentified): Goulden 1965); Qlu (Sars 1885): NSW (Timms 1970): Vic. (Timms 1973°): SA (Shiel 1978); NT (Smirnov & Timms 1983) “M. lenuicorniy Sars, 1896, NSW (Sars. 896); Vic. (Henry 1922). Cominent. Possthly ulsa from South Africa (Unverified); Goulden (1968) Moinodaphnia Herrick, 1887 M inacleayi (King, 1853); Moina macleayi King, 1853; Moinodaphnia macteayr: Sars 1888; NSW (King 1853); Qld (Smirnov & 'Timms 1983); NT (lull) 1986); WA [Timms 1988]) Family Bosminidae Sars, 1865 Bosmina Baird, 1845 B. meridionaliy Sars, 903 (not 1904 as in Smimoy & Timms [1983]). For extensive synonymy. see Smirnoy & Tinuns (1983). See also Jncertae sediy below; Tas. (Smith 1909, as 8B rotundant); NSW (Jolly 1964); Qld (Timms & Midgley 1969); Vie, (Timms (973%); SA (Shiel et al. 1982); NT (Tait ev al, 1984); WA (Timuns 1988) Bosnunapsix Richard, 1895 B. dietersi Richard, 1897, NSW Uolly 1966); NT (‘Tait 1981); Old (Timms 1986); WA (Timms 198%) Family Daphniidae Straus, 1820 Ceriadaphnia Dana, 1852 C. coriuta Sars, 1885; Qld (Sars 1885); NSW (Henry 1922), Vic. (Shie! 1978); NT (Tait 1981); SA (Shiel ef.a/. 1982); Tas. (Koste & Shiel 1987): ‘A (Berner 1987) Comment: Evidently more than one small species of Ceriodaphnia with ai acute “beak” occurs in tropical Australia (cl Berner 1987), Unt a thorough revision of the genus has been made, these laxa should be referred w C. vortur 5.1 C. dubia Richard, 1894: Qld (Gurney 1927): Vie. (Shicl 1976); °NSW (Timms 1989) C. latieaudauis Mueller, 1867, ‘Vic, (Shiel 1978); 2QId (Timms. $988) C. quadrangila (Mover. 785; Cerindaphnia hakea Smith, 1909; Brehm (1953a)2 ?Ceriodaphnia planifrons Smith) Brehm (1953a); Tas. (Smith 1909); NSW (Jolly 1966): Vic. (Timms. (973+); SA (Shiel 1978) C. rotunda Sars. 1862, Vie (Shephard er al, 1918) Daphnia Miller. 1785 D. carinara King, 1853.5,/. For extensive synonymy. see Benzieé (J988: 136-139), NSW (King 1853): Vic. (Shephard 1898); Tes, (Shephard 1917), WA (Serventy 1929); Qld (Tinims 1968), SA (Mitchell 1978): NT (‘Timms & Morton 1988) DP, vephalata King, 1853; For synonymy, see Benzie (98S: 129); NSW (King 1853); Vic. (Sars 1914) *D, jolly’ Petkayski, 1973: WA (Petkavski 1973) D. lumhaltsi Sars, 1885: For synonymy, see Benzic (ORK: 113-14): Old (Surs 1885); NSW. Vic. SA (Shiel 19814); WA (‘Timms & Morton 1988) *2 nivalis Hebert, 1978; For synonymy, see Benzte (1988: 122); NSW (Hebert 1977) *D. aceidentalis Benzie, 1986; WA (Benzie &ha) Daphniopsis Sars, 1903 *P australis Sergeey & Williams. |S; Tas, (Sergeev & Williams 1985); SA, Vic. (Williams 1986) *D. pusilla Serventy. 1929; WA (Serventy 1929): Vic., SA (Bayly & Edward 1969), Tas, (Sergeev & Williams 1983) *D. quadranguluy Sergeev, 1990: Vic. (Sergeev 19904) *D. queenslandensis Sergeev. \990; Qld (Sergeev 1990h) Scapholeberis Schoedlen 1858 S. kingi Sars, 1903; Daplmia mucronata Miller, 1785: King 1853 (misidentification); Scapholeberis kingit: Sars, 1388 (namen nudem): Smirnay & Timms 1983; Scapholeberts kingt Sars, 1903; NSW (King 1853); Vic. (Henry 1922); NT (Sulli 1986); Qld (Timms 1988); WA (Halse el al. 1993) Simecephalus Schoedler, 1858 5. acutirasrramus (King, 1853); Daphnia elisabethae acutirastrata King, 1853; Simecephatus acutirostratus: Sars IRR; Stmoecephalus dilvertonensts Smith, 1909; Dumont in Smirnov & Tiinms 1983; NSW (King 1853); Vie, (Haase 1903}; Tas. (Smith 1909); NT (Tait et ad. 884); Qld (Timms 198%) 46 RO oT SHIEL & J A, DICKSON S exypunesus oustradienyis (Dana. (852); Daphnia anstrativeasis Dana, (52. Simwvephalas ausrraliensis! Sars, WRB, SS. exspelnasis austratiensis; Durnont in Snyrmov & Tipps (1983); Qld (Sars 1888), NSW (Surs. 1896); ?Tas, (Stith 1909) SA (Henry 1922): WA (Servenry 1929): Vie. (Morton 19734) S. Jatirastris Stingelin, 1906; ?S. theringi Richard, 1897; Dumont in Smirnov & Timms (984): NSW (Henry 1922), NT (Tait ef af 1984); Olid. WA (Tins 1988) S. serrulatus (Roch, Wal, NT ull x6): Qld (Timms (88s) Si vetulies (Miller, 1776), 2 ssp. recognized Fray Australia (see copiments by Dumont, in Smirnoy & Timms [I9834> 98-102]) S verlus elisabethue Using. B53), Daplinie elisahetliae King; Simmacephalus elixabethae. Sars (SSS; NSW (King 1853); Vic. (Shiel 197K) SA (Shiel 9815), Qld, NT (Tiims Tks) S. vetdlus gibbosus (Sars, 1896); Sirecephealis gihbosus Sars. 1896; Simecephalus ventas glhbusus! Dumont in Smirnov & Tinims 1983: NSW (Sars i896). Vic, (Shephard era/, 1918), SA (Shiel 1981+) *S. vicroriensis Dumont, 983: Vie (Dumont in Sroirooy & ‘Viontas 84) Ineertiae sedis Alona hairdi King, 1853, NSW (King Desenpuon inadequate. Alona karte King, 1853; NSW: Biapertura karva in Srirnoy & Tires (983) is erroneously referred to King, They list King’s dlona kare as incertae sedis. The species they have mistabelled is dlonella karua in Sars (1888), which is dpparently a misidentification of another species, not the nominate A, keruva, As tygured by Sars, itis clearly nol the taxon figured by King, und should be relocaled if King’s species is rediscovered. Alona mascufa King, (853. NSW: Inadequately deseribed, Rosina maritima Mueller, 1867; ~ off the Abrolhos, 300 miles north of Fremantle, Western Australia, nt November, 1910" (Searle 1936; 172). Not recorded again. or mentioned by Korinck in Smirnov & ‘Yirnms (1983). 8 maritimes is recorded as a synonym ol B longixping Leydig, 1860 in Plossner (1972) This is the only record of a murine bosminid from Australia and its identity is uncertain. Chydorus barroist (Richard, 1894): Qld (Smimov OT): = Ephemereporus burroisi, nomen dubium (See Frey 1982a), Frey noted (p, 234) that the figured specimens from Prospect Reservoir, NSW, in Srairnoy (1971 Figs 328, 330, 331, 332) are nat ISSA: conspecific with &. rrideemaiis (Fig. 329 in the same series). nor are they conspeciic with B harraist 5, Str. Chydonis livbridus Daday, 1905; Old, NSW (Sntnov & Timms 1983): Frey (92a) relocated ©, hybrids 5. str. tow new genus, Evhemeroporny, and the xen hecame &. dvbviday (Daday). The limited features of the Australian taxon assigned to °C. Ayhridus® as figured in Smirnov & Timms 983) are neither Chydorus nor Ephemeruporus, bul more correctly those of Rak (Frey, in MS). Clivdoras ovals Kurz, 1874: NSW (Henry 1922): No fipures or material are uvailuble of the tuxon identified by Henry and it has not been recorded again, Itis a Holarctic species and regarded as absent Irom Australia by Smirnov & Timms (1983). Daphnia honerata King, 153; NSW (King 1833): A speeles of Ceriadaphina, insdequately described anu figured. Sars (I888) vonsidered if close to the European CC. reffenlata (Jurine, 1820), but specifically distinct Dunhevedia podaera King, 1853; NSW: Not seen sinee Original deseription, which is inadequate. Euryeercus chnninghami King, t853: NSW (King, 1854). A chydorid, but not referable to Aunecercay, Auryeercns spinosus King, 1853, NSW (Ring. 1853) A chydorid, buat not relerable to Banwerciy. Plearoxas aduncus (urine, 1820) in Smimoy & Tins (1983), Aloriella nasuta Smith, W909) Chydorus denticulaius Henry, 1919, Chyderus anispinus Henry, 1922: southern Australia: Smirnov & Times (1983); These tixe were synooymized with the northern hemisphere Po aduncus by Smirnov & Timms (1983: 24). Frey (1991b), after examination of these and other extensive materials, considered that P udurcis does not occur in Australia, The entities of these various axa have yet to he resulved. P. denticulatus Birge, 1879) non Chydoris denticudanes Henry, 1919, NSW (Smirnov 1971); Vie. (Timms 19737); 7Old, ?NT, 7WA (Timms 1988). Comment: After examination of the available material in the Australian Museum labelled as P denciculatus, Frey (1991b) concluded that none of the specimens was of the nominated taxon and they were Certainly not conspecific. He considered that the species probably was absent from Australia. Zoopgeography Qur comments here must be considered preliminary. given the rapid changes in cladoceran taxonomy in reeent years. Widespread recognition of non: cosmupolitinism has provided a significant impetus to a more critical approach (cf. Frey 1982b), Ibis clear, particularly from some of Lhe last works of Frey (1991, b),. that a considerable degree of endemism is obscures! CLADOCERA RECORDED FROM AUSTRALIA 37 by cosmopolignn Warnes in the Austratian fauna [avr opiion, any cladoceran in Australia relerted to a species described from the ortiem hemisphere shoul be viewed with suspicion until critics! reviews of all fymilies, to the standard of Frey (991b), are achieved, On present evidence, Australia hay more cladoceran species: 165 vs ca. 120 (Frope) and 10 (U0S.A.) than are found inother Comparable areas Overall the level of endemicity Stands. al 43%, with five additional taxa also known trom New Zealund. ie Ausiralisjan endemicity is ev. 46%. To the endemic genera Neotlriv. Pseudamdina (Macrothricidae), Archepleurosus, Australochyderus, Monape, Rak, Rhyachovhydorns and Saveiae (Chydoridae) listed by Smirnov & Tinims 0983), Celsinonan, Leberis, Planicivclus snd Plurispina (Chydoridae) are added. Most radiation appears to have wecurred in the Chydoridae; 45 of the 94 recognized species (48%) are endemic. Australia may differ from other cegions in the Selvctive pressures which cause genetc divergence (4 Prey 19916). In any event there has been poarked specmbon iqarcas Where water 1 limited, c.g. southwestern WA, where the habitats are not those ‘normally Muicative or supportive of a diverse aguatic mivrolauna, @y:. rock pools. salinized wetlands. The aquatic microfauna of these habitats, iy cemimon wath those of billabongs and wetlands of the opposite side ol the Continent. have generally been jenored. We suspect (hat a diverse array of indigenons clacocerans is yet lo be discovered. Acknowledgments. Our eladoceran work continues to be wded by the umcomparable- observations dnd literature collection of (he late David G, Frey, Without his expertise and keen eye and support during 1982-42 [he study of Australian chydorids would have been overwhelming. Our gratetul thanks wy Libby Frey, who supported RJS jo travel to Bloomingjon in Dec, 1992 to help sort the incomplete manuscripts and recover Australian material from the Frey collection, For their willing assistance in his neophyte ‘cladocerai’ days (before being subverted to rovers), RJS thanks Nikolai Smirnov (Moscow) and Brian Tinims (Cooranbongy. Hendrik, Segers (Gent) kindly sought obscure references unavailable in Australia. Our thanks also lo Alice Wells (Canberra: taxononuc advice), Jan Bayly (Melbourme; distaibution records) and Jean Just and the Australian Biological Resources Study (Canberra: grant support) References Avossir, Mo& Prephs. hb (989) Alone tbericu, new species: first evidence of noncosmopoliunisen within the 4, kara complex (Clidovera: Chydoridae), Jo Creyt, Aral, 4) 459.476 Burp. W. (i8s4) List of Grtinosteaca tound a9 Rerwickshire. Hist, Berwickshire Nat. Cl, 95-100. _ (1843) The natura) histery of the British Eaomnostmcs VI. nih. 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(1900) Stisswasser-Cladoveren Ereehu Hambarger magathaensische Sanmmetreise 1892/93 2 Bani. Arthropoden.. Hamburg: I-25. Waker, Ko EF & HinumMan, TJ. (977) “Limnoloyial survey ol the River Murray i relation to Albury Wodony.” (A. W. D. C.. Albury). Wiiuiams, W. 2D. (1979) Notes of the freshwater fauna of northwestern Australia, espectally m the Kimberleys. Ree Wesr Aust Muy. 7. 213-227, SKUSEMYIA ALLOCASUARINAE, A NEW GENUS AND SPECIES OF CECIDOMYIIDAE (DIPTERA) DAMAGING LATERAL BRANCH BUDS OF DROOPING SHEOAK, ALLOCASUARINA VERTICILLATA IN AUSTRALIA By P. KOLESIK* Summary Kolesik, P. (1995) Skusemyia allocasuarinae, a new genus and species of Cecidomylidae (Diptera) damaging lateral branch buds of drooping sheoak, Allocasuarina verticillata in Australia. Trans. R. Soc. S. Aust. 119(1), 41-46, 31 May, 1995. A new gall midge genus Skusemyia and a new species S. allocasuarinae are described from South Australia. Detailed descriptions of the larva, pupa, male and female of the new species as well as its gall on drooping sheoak, Allocasuarina verticillata, are given. The new genus is placed in the subtribe Schizomyiina within the tribe Asphondyliini. Key Words: Cecidomyiidae, Cecidomyiinae, Cecidomyiidi, Asphondyliini, Schizomyiina, Skusemyia gen. nov., Skusemyia allocasuarinae sp. nov., Allocasuarina verticillata, South Australia. Transactions af the Royal Saeiery ofS. Aust. W998), TA}, Alaé SKUSEMYIA ALLOCASUARINAE, A NEW GENUS AND SPECLES OF CECIDOMYHDAE (DIPTERA) DAMAGING LATERAL BRANCH BUDS OF DROOPING SHEOAR, ALLOCASUARINA VERTICILLATA IN AUSTRALIA by P. KOLESIK* Summary Kovesi€, P1995). Skasernvin allecasuerinae, anew genus und species of Cecwdomyiidae (Diptera) damaging lareral branch buds of droopme sheoak, Allocasuarina vertieillake in Austealia. Trot. Bo Soe: 5. Aust. U9¢1). 4\-46, 41 May, 1995. A new gall midge genus Skusemyia and anew species & alfpeesmarinae are described fram South Austrulia. Detwiled descriptions ofthe lurve, pupa. mate and female oF the new species ds well as Tis pall en deoeping sheowk, Allocasuarina vertictlum, are given, The new genus is placed in the subtibe Schizomylina within the tribe Asphondy liar, Kiey Words: Cecidomyiidae, Ceedomylinac, Cevidomytidi, Asphondylini, Schizomy ina, Skavenoia gen, nov, Skuseniva altocusuarinae sp. now. Allocnsaurnny verticillata, South Austratia Introduction “Vhis paper is the second part of a study Gn the South Australian Cecidomytidae. Kolesik (in press) described the first South Ausiralian gall midge species, Becinetivarnia matarskii, trom Encalypies fasoiculosa. A new species is deseribed here that was found damaging the lateral branch buds of drooping sheoak . Altlocasuarina vertieillate (Lam.) L. Johnson (Casuarmaceae) in South Australia. The large nurubers of infested and ultimately killed branch buds al one site indicate that this species could become a serious pest (Pig. 31). The new gull midge has one generation per year in Adelaide, South Australia. A new genus is erected for the new species. It belongs to the subfamily Cecidomytinae and superinbe Ceeidomyiidi. tt is compared to other known genera of the subtribe Schizomyiina of the tribe Asphondytiinr. Materials and Methods A survey of the Cevidomyiidae associated With galls on plants in nature conservation parks around Adelaide was carried out beeween November 1992 and May 1993. All galls sampled were dissected and examined. Those which contained larvae of Cecidomyiidae were described, photographed and conserved for luter authoritative identification of the host plant species. The larvae from the gall kind described here were processed in lwo ways, A small number was preserved in 70% ethanol after notes were made on their colour The larger number was brought to the laboratory to rear to adults. Here the galls. were carefully dissected * Department of Horticulture, Viticulture and Oenology, Faculty of Agricultural and Natural Resource Scienves, The University of Adelaide, PMB L, Glen Osimond, South Austealta S064, Austeatia- and the larvae transferred with entornolagical tweezers into rearing pots containing sterilised, wet sand (Skubravil & Skuhravy 1960). Pots were examined daily and emerged adults preserved together with their papal skins in 70% ethanol after their colour had been noted. Microscope mounts of a series were prepared by miuverauion in 20% KOH. followed by processing through distilled water, 70 and 99% cthanol and xylene (o Canada balsam mountant for examination by interference-contras| and bright-light microscopy. Larvae, entire or dissected into two pieves, and entire pupae, were mounied doarso-venirally or laterally, Adults were dissected into tour (female) or five (male) pieces und the particular parts were mounted separately. Wing and head were mounted frontally. thoras laterally, abdomen dorso-ventrally or laterally and male genitalia dorso-yentrally, Measurements were made with an eyepiece reticule. Drawings were done with the uid of a camera lucida. The type series and other material retained in 70% ethanol are deposited in (he South Australian Museum, Adelaide [SAM] and United States National Museum. Washingtoo |USNM], Genus Skusemyia ven. noy. Adult Wings with R. joining C al wing apex, Rs weak, R, joining C near wing mid-length. M, 5 absent. M, weakly developed, Cu forked. Maxillary palpus with 4 sepments. Male antenna with 12 Nagellomeres. Female antenna with I! flapellomeres, the last three successively and progressively shorter, the last apparent Magellomere evidently a combination of the eleventh and twelfth, Flagellomeres cylindrical with necks, first and second not fused, with Jong and stout setae in two whorls, bearing closely appressed circumnftla. Tarsi with first segment substantially shorter than the secand. 43 rl KOLESIK first Lersomeres lacking yventrodistal spine, tarsal claws siinple. empodia mach shorter thun claws. Male jerminaha: gonovoxites Free ventrally, produced to form a roundly tiangular apie! process; gonosty)us situated dorsally on gonacoxtie, short and wide, with truncated apex bearing teerh of umtorm length: cerci simple, founded upically. hypopreet divided into two apical lnbes; claspettes large; aedeagus long. srout. tapering distally, Female abdominal sternite 7 longer than Stvrpile 6, Ovipositor: protructile, elongate, sclerolized, without basal lobes: verct fused. divided at apex, SEHOSL, Larva Head capsule with short posterolateral apodemes: Ameona short. Sternal spatula bilobate. Anus ventral. Thorucie and first through seventh abdominal segments with 6 dorsal, 2 pleuraland 4 ventral papillae. Biphth abdominal segment with 2 dorsal, 2 pleural and 2 ventral papillae. Terminal segment wilh § dorsal umd Aanal papillae. AlL collar Unoracie and abdominal papillae Usetose with exception of the dorsal pupillie onthe terminal seyment that are bearing very. short sche, Pupur Frans without projections, One of three lower fayial papillae with a seta, Lateral facil papillae absent, Cephalic selerite with two strongly chitinized swellings und two papillae wath long sete. First through seveoth abdominal seyments with 6 dorsal asctose pupillac and one pleural setose papilla. Sewend through eighth abdominal segments dorsally with 3 indistines (ramaverse rows of spines On anterior hall, Last abdominal seament with large pouch. emarginate medially but nat completely divided in io parts Whe species: Skusenvia allocasiarinad sp. wv. Hiyinology The genus 6 named after FAA. Skuse, author in I888/(890 of the first taxonomic studies on Australian Cee domy ndite, Remarks Skasemyiee lits in the tribe Asphondyliini of the supertribe Cecidomyiid) because the female seventh abdominal stermle 15 1.5 lines as long as the sixuh sternite and the eqhth tergite i wider than the seventh tergite, combined with the male genitalin having a ventroapical gonacoxal lobe and a dorsally situated fonostylus that is about as broad as long: it belongs lo tle subtribe Sehizamyiina becuse the first tarsomeres lack a ventrodistal spine. the male genitalia have claspettes, and the female lacks cere\-like lobes immediately posterior to the eighth tergue (Gagne 994), Within the subiribe Schizamyiina, Skusemyia resembles most closely Placerhela Rubsaamen. known from three European and one El Salvadorean species (Molin 1960, Skulirava 1986). The male aacennae of the iwo genera are the only ones in the subtribe with relatively simple cireumfila and with fagellomeres made ap ofa bulbous basal nodé and long neck an! resembling these of Desineure wid relalives (Oliotrophini: Lasiapteridi). The female antennae of Skasennvie und Placochela ure also similar except that the eleventh and twellth layellomeres of the new gerius are apparently amalgamated, which is unique mi Asphondyliini. The genitalia of both sexes are generally similar cso (Mohn 196))_ except in details of the oyipositer whieh differs in Skuse beeause the vere are diserete, at least attheir apices. and the distal setae are longer. The immature stages of Skusemyia are unique in Schizomyina, The pupal cephalic selente has two swellings thal are longer than the antermal homs. The larva has very reduced papillar sere. and the papillae of the terminal seyment are all situated at the end al separate lobes. Skusemyia allacasuarinae sp. nov- FIGS 13 Halorype: Oo, Black Hill Conservation Park, Sauth Australia [34°53'S., I38°44°E.|. 15.11.1993, P Kolesik. reared from larva tron lateral branch bud yall of Allpeasuarina vertiettaia (ham) L. Johnsen salnpled 25.1, 1993, 121270. [SAM]. Allvlype: 2, same dala bub emerged 16.11, 1894, (2127) [SAM]. Pararypes (all sampled with holotype): 4cr.cr and $09, emerged 13-19.111-1993; 7 pupae, emerged 13-18 71-1993; § larvae [SAM]. Other maternal 4 F 39 9 |SAM[|. 300,390 9 [USNM|..4 pupae [SAM] and 3 pupae |USNM], same dats us holorype but emerged 16-22-1717. 1993. 6 [SAM] and 3 larvae |USNM], same data bat sampled 25111993, 3 larvae, Wistow, South Auestratia [35°07 'S_. 158°53’E_], 23.41.1993, P. Kolesik, on A. vertierdlate. Description Male (Fig W) Colour: antenna prey, head black. thorax. brown. abdomen with sclerotized parts black and non sulerotized red, legs yellow, all setae black, Toul legeui of the body 3.24 mm (range 315 9 3.33-mm), Wing length 2,44 mom (2.34 - 2,52), width 0.98 mm (0,93 - 1,00). Wing membrane und especially veins densely vovered with selac, 50 - 60 xm. Antenna total lengeh 1.72 min (1.57 - 1.83); Mavellomeres with stout sete 54 - 160 am, longer at the distal whorl than at the basal whorl; closely appressed circumfila consisting of one transyetse and one longitudinal bands. Eye bridge + Ww 4 facets medially, eye facets rounded. Claws curves beyond the second third. 39 wm (37-41). Empodiuny I pm (8 — 14). A NEW GENUS AND SPECIES OF CECIDOMYIDAE 43 Fig. 1, Male of Skusemyia allocasuarinae gen. et sp. noy.: A. genitalia in dorsal view; B. wing, C. last tarsal segment with claw and empodium; D. head in frontal view; E. sixth flagellomere; F, last three flagellomeres. Scale bars = 100 pm. dd P. KOLESIK ra | r | 2 ry ( ) } 7 la B} in c-— D-——_ E Fig. 2. Female al Skusennia allacasuarinae gen, et sp. nov. A. ovipositer in ventral view, B. end of ovipositor im ven- tral View; C. first four antennal segments; D, fast five Mag- cllomeres, E. sixth (itgellornere, Seale bars = 100 wn, Terminalia: gonocoxite setose and setulose: zonostylus selose and setulose, bearing 17 - 20 sclerotized tapering teeth that are narrow and about 15 pm in length: hypoproct bearing one setaon each lobe. setulose; cerci deeply divided medially, setose and setulose; claspettes setulose Female (Fig, 2) ‘Total length of the body 3.63 mm (3.41 - 3,80). Wing length 2.65 mim (2.32 - 2.80), width 0.89 min (0,74 - 1.00). Antenna total length 112 mm (0,95 - 1.20), flagellomeres with setae, 5] - 15 jan, Last Mayellomere with a shallow constriction medially, Seventh abdominal sternite about 1.5 times longer than sternite 6. Ovipositor with one ventral sclerotized longitudinal band forked distally. distal hall of ovipositor with § -9 pairs of 6-9 jm Jong setae, cercus with 8 pairs of 5 ~ 48 pm tony setae. Colour.and other characters as an male. Mature larva (Fig. 3A-D) Colour red. Total length 3.70 mm (3,24 - 4.44). Integument smooth, ventrally with several transverse rows of spiculae on antenor half uF each segment. Head capsule width 61 jam (53-77). length 63 wm (S51 - 74), length of posterolateral apodemes 61 jum (51 - 64), Antenna 17 pn (15 - 20). Sternal spatula 268 yim (230 - 320) in length, with apical enlargement 86 am (77 - 105) in width and 42 yom (38-51) in length. Terminal segment dorsally with 8 lobes bearing papillae: @ with very short setue and 2 with corniform sete: Pupa (Pig. 3F-H) Colour, prothoracie spiracle, cephalic swellings and untennal horns dark brown, remaining: parts pale brown. Total length 3.03 mm (2.46 - 3.56). Integument covered with spiculae, ventrally 2-4 mand dorsally 4-6 um long. Antennal horns 33 ym (28 - 38) long. Cephalic swellings 77 yn) (74 - 80) in length. Cephalic pupilla with seta 76 aim (58 - 90), One of three lower facial papillae with seta 45 pm (38-51). Prothoracie spiracle [46 pm (140 - 151) long and 23 ym (20-28) wide across the base, with trachea ending at apex. Seta on pleural papilla 9 am (8 - 10), Dorsal spines of the first row [3 . 20 in pumber, 5 - 30 pm, spines of the second row 13 - 20 in pumber, 25 - 45 yam: spines of the (hird row 9 - 12 tn number, 35 - 65 jum. Gall (Fig. 31) Swollen lateral branch bud, forming spherical to spindleform rosette, 7 - 12> mm in diameter, polythalamous, pale brown in colour. One larva in cach of the 2 - 3 cells. Gulls appear in January - March Larvae eave galls to pupate in the soil. Eryinalogy Derived from the generic name of the host plant Acknowledgments I thank the South Australian Museum, Division of Natural Science, for providing the facilities that assisted this Work, Lam grateful to the museum collaborators Ms J, A. Forrest, Dr E. G. Matthews and Mr DB. Hirsi for their support. The Ministry of Environment and Planning, South Australia, kindly gave permission to collect in the nature conservation parks of Black Hill, Morialta and Cleland. I wish to thank Mr M, C. O'Leary, State Herbarium of South Australia, Adelaide, for the identification of the host plant species, | an grateful to Dr Raymond J. Gagné, Systematic Entomology Laboratory, U.S. National Museum, Washington, tor carelul review of the manuscript A NEW GENUS AND SPECIES OF CECIDOMYIIDAE viel, 4 f it TaN tem Fig. 3. Skusemyia allocasuarinae gen, et sp. nov.: A. - D. larva; E. - H. pupa; I. gall. A. stigma; B. head capsule in dorsal view; C. anal segment in dorsal view; D. sternal spatula; E. sixth abdominal segment in dorsal view; F. last abdominal segment in dorsal view: G. anterior end in ventral view; H. prothoracic spiracle, I. Allocasuarina verticillata - lateral branch bud galls caused by Skusentyia allocasuarinae gen, et sp. noy. Scale bars = 100 wm. A. - H.; 2 em I. 46 P. KOLESIK References GaGne. R. J. (1994) “The Gall Midges of the Neotropical Region.” (Cornell University Press, Ithaca, New York). KOLESIK, P. (in press) A new species of Eocincticornia (Diptera: Cecidomyiidae) on Eucalyptus fasciculosa in South Australia. J. Aust. ent. Soc. Moun, E. (1960) Gallmiicken (Diptera, Itonididae) aus El Salvador, 2. Teil. Senckenbergiana Biol. 41, 197-240. (1961) Gallmiicken (Diptera, Itonididae) aus El Salvador. 4. Zur Phylogenie der Asphondyliidi der neotropischen und holarktischen Region. /bid. 42, 131-330. SKUHRAVA, M. (1986) Family Cecidomytidae pp. 72-297 In Sods, A. (Ed.) “Catalogue of Palaearctic Diptera.” Vol. 4. Sciaridae: Anisopodidae (Akadémiai Kiad6, Budapest). & SKUHRAVY, V. (1960) “Bejlomorky.” (Statni zemedelské nakladatelstvi, Praha). TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 119, PART 2 NEW GENERA, SPECIES AND A NEW SUBFAMILY OF XYALIDAE (NEMATODA: MONHYSTERIDA) FROM OCEAN BEACHES IN AUSTRALIA AND THAILAND By WaRWICK L. NICHOLAS* & AIMORN C. STEWART* Summary Nicholas, W. L. & Stewart, A. C. (1995) New genera, species and a new subfamily of Xyalidae (Nematoda: Monhysterida) from ocean beaches in Australia and Thailand. Trans. R. Soc. S. Aust. 119(2), 47-66, 31 May, 1995. Five new species of Xyalidae are described from Australian ocean beaches and one from Thailand. Gullanema fragilis gen. nov., sp. nov., possesses many long cervical and somatic setae, but is distinguishable from other similarly hirsute Cobbiinae by a distinctive narrow peri-buccal region. Rhynchonema tomakinese sp. nov., R. collare, sp. noy. and Prorhynchonema gourbaultae sp. nov., in the Rhynchonematinae, differ from congeneric species by their spicules and amphidial regions. R. chiloense Lorenzen, 1975 and R. gerlachi Vitiello, 1967, and P. warwicki, Gourbault, 1982 are commented upon. Two new species with characters intermediate between the Cobbiinae and Rhynchonematinae are placed in a new _ subfamily, the Corononeminae. The cervical region, enclosing a cylindrical buccal cavity, is shorter and less attenuated than in Rhynchonematinae, longer and narrower than Cobbiinae. The labial region its set off from the buccal region by a narrow indented ring. Corononema parvum, gen. noy., sp. nov., is described from Australia and C. thai sp. noy., from Thailand, the two differing in the shape of the head. Key Words: Taxonomy, nematodes, Xyalidae, beaches, Gullanema, gen. noy., Rhynchonema, Prorhynchonema, Corononema, gen. nov. Transactions of the Royal Somety of S. Aust (1995). 1t9(2),47-06 NEW GENERA, SPECIES AND A NEW SUBFAMILY OF XYALIDAE (NEMATODA; MONHYSTERIDA) FROM OCEAN BEACHES IN AUSTRALIA AND THAILAND by WARWICK L. NICHOLAS* & AIMORN C, STEWART* Summary NicHouss. W, bo & Stewart, AC. (1995) New genera, species and u new subhimily of Syalidae (Nematoda, Monhysterida) [ror ocean beaches in Australia ancl Thailand, fins. Ro Soc. 8. dst 192), 47-66, 31 May, 1995, five new species of Xyalidac are described from Australian ocean beaches and one from Thailand, Gullaneme fragthy gen, low. sp, nov, possesses many long cervical and soralic sefae, but is distinguishable from other similarly hirsute Cobbiinae by a distinctive narrow peri-buceal region. Rivachanema tomakinese spo noy.. KR. collate, Sp. nov. and Proriyichoneme gourhantiae sp, nev.. in the Rhynchonematinay, ifler from congeneric species by their sprules and amphidial regions, R, chileense Lorenzen, 1975 and RK. gerfarhr Vitiello, 967. and P. warwteki, Gourbault, 982 are commented upon Two new species with characters intermediate between the Cobbiinae and Rhynehonematinae are placed ina new sublamily, the Corononeminae. The cervical rexion, enclosing a cylindrical buceal cavity. is xhorter and less attenuated than in Rhynehonematinae, longer and narrower than Cobbiinac, The labial region is set oll [rom the buceal repion by a narrow indented ring. Corwioiema parvnin, pen nov., ap nov, is described from Australia and. thai sp. nov., from Thailand, the twe differing in the shape of the head. Key Worns: Taxonomy, nematodes, Xyulidae, beaches, Gullanema. gen, Prorkvachonend, Caronanema, gen. nov. Introduction Neratodes, have been collected from sandy beaches on the southern, castern and northern coasts of Australia and southern Thailand. Previously we (Stewart and Nicholas 1994), described eight new species of Xyalidae Chitwood, 1951 belonging to well- known veners of Cobbiinae de Coninck, 1965, In this paper we describe 4 new species and genus of Cobbiinae, numely Gullanema fragilis gen. anv. . sp. nov., and threé new species from the other subfamily of Xyalidae, the Rhynchonematinae de Coninck, 1965, namely Rhyachonemu temukinense, sp, nov,, R. collare, sp. nov. and Prorhynchanema gourbanttae sp. nov. Species of Rhynchonente Cobb, 1920 are common in Australian sandy beaches, as in other parts of the world, and we cotiment on Australian specimens belonging to two previously deseribed species, The genus was comprehensively reviewed by Lorenzen (1975). We also erect a new subtamily, Corononeminuae, w hold Corononema parvum gen, nay:. sp. nov. and ©. thai sp. nov., with characters intermediate between Cobbtinae and Rhynchonematinae. The last named species is from Thailand. * Division of Botany and Zoology, Australian National University, 0200 Canberra. Australia, voy, Riivnelaneni, Materials and Methods Collections were made on the 90 Mile Beach ut Seuxpray, Vicloria (147° 23'E, 38 46'S); South Moruya, Broulec, Rosedale, Tomakin and Kioloa beaches, New South Wales (berween 150° 9'E, 35 55'S and 150 20'E,*35 32'S): Southport beach, Quéensland (153° 25'B, 27° 58'S) and Rapid Creek beach, a suburb of Darwin, Northern Territory: (30 50'F, 12° 23'S). Some specimens were also collected trom Pathaya beach. Chonburii. Thailand (100° 53’E, 12) 45'N). Specimens were collected in samples of sand dug up at low tide between the tidemarks to a depth of 40 em. Exceptions were samples of sub-tittoral sand wken from a boat in shallow water off Cronulla, New South Wales (151° 10'E; 34° 05'S), Nematedes were extracted from the sand by re- suspension in tap water, allowing the sand to seule and collecting the nematodes ona 50 jum. nylon mesh sieve. The nematodes. were washed olf the sieve imto sea water, then fixed in 5% tormalin in sea water. Specimens were picked up under the microscope with a fine pipette and transferred to 5% aqueous glycerol which was slowly dehydraled to anhydrous glycerol at 40°C. Permanent tnounts were made in anhydrous glycerol, and the cover slips ringed wiih Glyceel (Gurr). Glass beads (ballatini), selected under the microscope to approximate the diameter of the nematodes, were used to support the cover slips. ak WARWICK L, NICHOLAS & AIMORN C. STEWART Drawings and measurements were made using a caverta lucida, When mounted the nemilodes Jie ou their sides presenting a lateral view (except in Pig, 19) Our draWings show setae on one side only, that lying uppermost as mounted - Measurements are in pau front specimens fixed and mounted in this way, De Man's ratios are piven, i.e, a = body length divided hy greatest body width, b = length divided by length of pharynx, ¢ = length divided by tai) length, c = tail length divided by width at anus. V = anterior end to vulva as @ percentage of body Jength, and spicule measurements are arc length, For Rhynchonematinae, Lorenzen's (1975) formula has been used to sumumarise the characteristic features of each species described. In this formula a Jeter code, referring to drawings of characteristic structural leatares, is used to describe successively the form of body annulation, symmetry of the spicules, form ot the spicules, annulation surrounding the amptids, relative size of amphids in both. sexes, position of amphids relative to end of buccal tube, form of buccal cavity, and form of the vulva. An important character is the direction, either towards the anterior or towards the posterior, of the saw-tooih edge of culiculur annulation (reiferartie dick nit sageatiger Aussenkortur). In some species there is an abrupt mid- body change im direction. Where this is so, the distance from the head end to the inversion is expressed as a percentage of body length. In other species only the cervical and tail regions: have saw-tooth amulation, the inid region of the body having rounded annule profiles, Scanning electron micrographs (SEM) were made from specimens that had been postfixed in 14% aqueous osmium tetroxide, freeze-dried and coated with gold palladium. Type material is deposited im the South Australian Museum, Adelaide, and the museum's numbers are given im the text, Holotype numbers (all males) follow SAMA and the prefix V. Some paratypes are On the same slide, to give both a male and a female. Other paratypes are on slides labelled AHC followed by Oo nuniber. Taxonomic descriptions Tamily Nyalidac Chitwood, 1951, Annulated cuticle, ciccular eryptospiral amphids, female with single prodelphic ovary situated to left of gul. male with one or two testes, anterior situated to lefi GF gut, posterior if present to right of gut. Subfamily Cobbiinae de Coninck, 1965 With characters of family. Head and cervical region only slightly attenuated. base of buccal cavity conical, enclosed by pharyngeal musculature, 10 sensillu in second ring of cephalic sensilla, usually segmented. Genus Guilanema goa. nov. Type species: Gullanema frayiliy Diagnostic definiuten With characters of Cobbrinae, Cephalie region surrounding conival buccal! cavity eylindrteal. sec ol from wider cervical region. Strong mouth ring supporting six labial setae and six thin flexible lips. Very long angulated double and iriple sets of cervical setae and numerous long single and double sematir selae extending as far as the anus. Etymalogy Named after Dr Penelope Guillen im the School of Lite Sciences, Australian National University Gullanema fragilis sp nov (FIGS 1-12) Holotype; Male, SAMA V4259, Kioloa beach. NSW. 27. x1.J986. Pararypes; 4 males, AHC 24808 and 24813, Kiolog beach, NSW, 27.21.1986, 15.x7.1987 respectively: AHIC 24809 und 24812, Broulee beach, NSW, 3viii 1980. 8 females. AHC 24810 and 24813-15. Kioloa beach. NSW. 27.15.1986, 3.viii 1980, 26.yii.1978 and 15.01 1987 respectively; AHC 24812, Browlee beach, NSW, 3viii. 980; AHC 24816, Moruya beach, NSW, T.xuU.t98%, AHC 24817, Scaspray beach, Vic. 3.411 1988, Meayurementy: ‘Table | Deseriprion of Holarype male, Body cylindrical, cuticle finely annulaled, tail w long narrow tapering cylinder, Head widens sharply at buse of narrow peri-buceal ‘iret-lke’ region enclosing buceal cavity, Almost cylindrical buteal cavity, taperest al base, six longitudinal bars reinforce buccal cavity, strong circam-oral mouth ring, surmounted by six fexible very thin lips, about 6 gm high. and six 4m long inner labial setae. Six 18 wm outer labial, four 12 pin cephalic setae, in one ring, insert near hase of peri-buccul ‘turret. Complex hexaradiate array at cervical setae insert between base of buccal region and amphids: most anterior, six IL jan short setae; next, six longer unequal pairs of cervical setae, 36 and 14 pm; then six longer triplets of unequal length, 166, 102 and 50 am. The successive sels of cervical setae nut inserted in rings around the hody at the same level, but cach set slightly staggered relative to the long axis of body. Labial, cephalic and cervical setae segmented. Amphids, almost circular, 9 xin diameter, 31 yan behind mouth. Long unagual pairs and single setae inser irregularly between amphids and anus. longest. 209 pun, iN pharyngeal region. Also a few shart setae along hody continue posterior to anus. Two caudal setae, 168 and 145 wm long, at tip of 230 um tong tail, Pharynx cylindrical, cardia conical, rectum short, Two NEW XYALIDAE TABLE 1. Measurements of Gullanema fragilis gen. nov., sp. nov. Type Leneth Width Lips Inner labial setae Outer labial setae Cephalic setae Amphid diameter Mouth to amphid Buceal cavity Nerve ring Pharynx Mouth to vulva Mouth to anus Tail Anal Width Spicule De Man’s 4 " b " c " ul V% nn ttt tttEIttESESS * Excluding female with truncated tail (n=7) Figs 1 and 2. Head of male Gullanema fragilis sp. noy. Setae, which are annulated, are illustrated only on one side of the body. Holo Male 1443 36 =o w , oH HKHeraeam Male Paratypes n = 4 Range 1368 - 1482 24 - 36 3-8 8-10 22 - 26 13-19 7-8 27-35 9-18 68 - 90 262 - 330 1015 - 1293 17] - 225 21 - 33 37 --46 41-55 44 -5.4 65-9 6.8. -9.1 **Truncated tailed female Mean + SD 1444 258 29 4.85 a: 2.10 9 1,16 24 2,07 16 2.50 78 0,50 30 3.78 13 3.78 77 9.27 290 30 1200 126 199 24 26 5.25 42 4.43 SI 6.74 5 0,43 73 0,70 1.01 7.83 Female Paratypes n =8 Runge 125] - 2025 35 - 62 5-9 7-9 22-29 15 - 20 71-9 22-45 10-23 82 - 130 275 - 365 751 - 1279 JOL9 - 1645 30** - 380 21-48 26-43 3.9-S,5 41,7** - 8.6 L1**-11 74-81 Mean + SD 1580 261 47 8.82 6.9 0.58 8 0.93 25 2,06 17 0.53 7.8 0.70 32 7,28 1S 4.30 100 17 328 28 981 160 1332 213 279+ +. 347.91 34 7.36 AS 0.50 6t ¢ &.2* * 76 2,58 3) WARWICK 1 a : ranger wr Figs 3 and 4. Gullanema /rayilis sp. nov. 3, entive male, 4. spicules. outstretched lestes, antertor to left of gut, posterior to right, gonoduct filled with spermatozoa, spicules weak, hardly cephalated, uniformly curved, blunt bilid tips, 48 yum long. Puratypes, The numbers and location of somatic setae are rather variable, especially the oumber of caudal setae, which vary from one to three of unequal length, possibly due to breakage during preparation. The number dnd location of short cervical setae are varluble, 9-id ym long, Two sets of longer cervical setae are consistently present, the first set of six doublets of unequal length, the second of six triplets also of unequal length, but with (he points of insertion of each group staggered with respect to the longitudinal body axis. As an example of setal Jength in one female paratype doublet 58, 42, triplet 99, 62, 45, cervical 200), 55 pm. The post-amphidial setae apparently not inserted in regular rows or circles, the most anterior the Jongest, decreasing to short scattered setae behind the anus.. The distance the amphid lies behind the mouth is rather variable both in absolute terms and relative to pharyngeal length. Female paratypes differ significantly from males only in reproductive organs. Females are monodelphic and prodelphic. Most specimens of both sexes have long tails, but occasional specimens with much shorter tails haye been found: an extreme case is illustrated in Fig. 5 (on slide NICHOLAS & AIMORN ©. STEWART AHC 24810, which also contains a second female with a slightly longer tail), Because the degree of truncation is variable and found on infrequent individuals amongst numerous long-tailed individuals, the shert-tailed farm is not considered a separate taxon but an infrequent aberration, Because of the variation in tail length, the position of the vulva is more usefully related fo the percentage distance from head to anus rather than the more usual head to tip of tail, Differential diagnosis The new genus resembles Stemeria Mivoletzky, 1922. and less closely 7richorheristus Wieser. 1956, in possessing many very Jong setae. In Sremneria, as in Gullanema, but unlike Trichotheristus, groups of long cervical setae are inserted between the mouth and the amphid. Gullanema is quite unlike either genius in the formof the head and shape of the buccal cavity, In these two genera the head tapers gradually to the base of the large lips, whereas in Gullanema gen. nov. the head narrows sharply to a cylindrical ‘wrret-like’ region surrounding ihe buccal cavity, The outer labial und cephalic setae are inserted near the base of the turret, the inner labials at the mouth ring. The lips are smaller, supported by 4 very strong mouth rim. At the specific level, the weakly cephalated spicules with blunt bifid tips are an important character. Habitat Sandy ocean beaches. Dispributton New South Wales, Victoria, Subfamily Rhynehonematinae de Coninek. 1965, After Lorenzen, 1975: head und mouth region very attenusted, buccal cavity long and tubular, cuticle strongly annulated, first cephalic annule wider. amphids circular. tail conical. Uncertuinty regarding cephalic sensilla because of difficulty in resolving tiny head with hight microscope, inner Jabial sensilla undescribed, single ring of 6 or \O cephalic setae. Less than 900 um tong. Genus Rhynchenenia Cobb, 1920 Buccal cuvity in two parts; short anterior chamber, at level of cephalic setae, narrow tubular part at least 25 ym Jong., Amphids placed over or very close to end of buccal tube. Male with two testes. Rhynchonema collare sp. nov. (FIGS. 13-18, 25-30) Holotype: Male. SAMA. V4260, Rosedale beach, NSW. 5.11. 1986. Pararypés: 2 males, AHC 24818, Rosedale beuch, NSW, 5.111.1986 and AHC 24819a Broulee, NSW, 3yiii, 1980; 7 females. AHC 24819b-c, Broulee beach, NSW. 3.vili J986, NEW XYALIDAE Figs 5 and 6. Gullanema fragilis sp. nov. 5. female, infrequent short tail form. 6. common long tailed female. 51 52 WARWICK L. NICHOLAS & AIMORN C. STEWART _- OLS AM Figs 7-12. Gullanema fragilis sp. nov. 7. head to show buccal region. 8. showing amphid. 9. spicules. 10. SEM of buccal cavity. Il. entire nematode. 12. SEM cephalic and sub-cephalic setae. SCS sub-cephalic setae, OLS outer labial setae, AM amphid, SP, spicule, BC, buccal rim, LF, lip flaps, ILS inner labial setae, CeS, cephalic setae. NEW XYALIDAE 53 13,14,16,17,18 ———__________2's im 15 ———————. 50 pm Figs 13-18. Rhynchonema collare sp. nov. 13. female head. 14. male head. 15. entire female. 16. inversion of annules. 17. male tail and spicules. 18. female vulva and operculum. 54 WARWICK L. NICHOLAS & AIMORN C. STEWART TABLE 2, Measurements of Rhynchonema collare yp. mer. ‘Type Holo Male Male Length 476 412 Width 20 Y Body setae - 3 Ruveal eavity 53 ay Amphid. length 7 1k Amphid, width 10 10 Mouth to amphid 5? 40) Mouth to nerve ring - Pharyns 125 14 Mouth to vulva Mouth to inversion’? 36 55 Mouth to anus 399 428 Anal width 17 1k Tail 77 05 Spicule. long 16.8 18.7 Spicule, short 13.7 13,2 De Man's a 24 pad w bh ams) an Z e 6.2 63 £ c! 15 3.6 " VGE = Mate Paratypes Female Paratypes n= Male Range Mein + Sb 579 311-552 333 19 7 20.23 218 110 17-17 7 0.00 52 47-34 3] 2,63 19 6-95 8 (.32 12 6-7 67 0.45 a2 44-55 Ay 4.00 AS R499 90 AR? 162 124 - 152 138 |3 316 -39] 3S 35 38 56-73 60 5.07 515 439 - 483 465 iv 16 15-17 15.6 ORY 75 34-74 65 8.72 Ih2 - - - 12.8 - 34 22-26 24 1.52 3.6 36-41 34 0.27 a7 71-99 8,3 142 47 3.0 -4.9 4.2 O48 - 61-73 69 5.41 Measurements: Table 2. Description of Holotype mate Very small, with long attenuated cervical region. plump post-cervical region, rather long broad curved conical tail, recurved at hp. Cuticle strongly annulated, annule profiles sharply angled forward on Front half of body, backward on posterior, abruptly changed (inverted) at 56% body length; very thin somatic setae spaced uniformly between amphid and anus, Buccal cavity a shallow cup leading into long narrow parallel- sided tube with strongly cuticularised walls extending Jength of narrow cervical region to level of middle of amphid. Six cephalic setae at base of buccal cup, extremely large elongated amphids enclosing posterior 40% of narrow cervical region: no annulation between amphids. Pharynx cylindrical, cardia heart-shaped. Spicules unequal, weakly cephalated, lacking rectangular bend, tips turned up, Large gubernaculum encloses mid region of spicules, strong dorso-caudal apophysis. Paratypes: Amphids strongly dimorphic, in females relalively large, but much smaller than in males, separated by strong non-annulated cuticle. Vulva with operculum, on which annulation greatly reduced or absent. Terminal vaginal canal cuticular. Lorenzen’s formula:- bwaokg,+.2,0.w: “new letter because spicules do not correspond to any of those figured by Lorenzen (1975), Differential diagnosis The new species can be distinguished from some other species by possessing sharply angled annulation along the whole body and by the possession of sexually dimorphic amphids. It lacks the pre-anal supplements and strong somatic setae and equal spicules found in R. hirsutum Hopper, 1961, It more closely resembles R. chiloense Lorenzen, 1975, and R. scutatum Lorenzen, 197! but has quite differently shaped spicules. The male amphids are larger than either of these species, It differs from R. tomakinense sp. nov. in the larger male amphid, absence of annulation between amphids, and in possessing a cuticular vaginal canal. Habitat Sandy ocean beaches. Distribution New South Wales. Rhynchonema tomakinense sp. nov. (FIGS 19-24) Holotype: Male, SAMA V4261, Tomakin beach, NSW, 1. xi1986. Puratypes: 3 males and 5 females, additional male and 3 temales on holotype slide: male and female. AHC 24820a, Rosedale beach, NSW. 22.xi.1986 and female, AHC 24820b, Rosedale beach, 5.11.86; male, AAC 24821. Rapid Creek beach, NT. 31vii. 1986, NEW XYALIDAE 55 19,20,21, 22,23 24 € a, oy, Figs 19-24. Rhynchonema tomakinense sp. nov. 19. male head, dorsal view. 20. male head, lateral view. 21. female head. 22. vulva and operculum. 23. spicules. 24. entire female. 56 WARWICK L. NICHOLAS & AIMORN C. STEWART TABLE 3. Measurements of Rhynchonema tomakinense sp. nev. Type Holo Male Paratypes m= 3 Fermale Paratypes n ='5 Male Range Mean + SD Range Mean + SD Length 331 438 - 495 ARS a0 4Xy - 502 545 ay Width 22 19 25 AW 3.12 22-34 24 15 Body setac tl 9-18 12.6 - 13-16 [4.6 - Ruceal cavity 44 Ax 57 49 473 j4- 49 47 1,52 Amphid, length 4 9-12 2 L7s 6-9 72 04,10 Amphid. width iT) K-12 8 2A 6-38 6.4 O89 Mouth to amphid as AO - 43 3 158 48 - 44 420 20s Pharynx 121 15-110 10 LOS 120 - [ay 134 1) Mouth to vulva - - - - - 345-440 ARG 46 Mouth to inversion 54 S266 57 7457 5) - 58 55 2.86 Mouth to anus 474 382 - 434 413 29 424-324 476 18 Anal width Is 17-18 17) O.SR IS 18 16 ),52 Tail 77 36 GK 60 6.93 64-73 68 4.05 Spicule is I) 19 I4.00 0 4.04 - - - - Gubemaculurn 1] - - - - - De Man's a 25 22-24 23 1.00 20 26 a4 245 " bh 4.6 41 47 44 O30 35-45 41 O44 We c 7.2 TL RS 7.9 ORS 74 87 8.0 O63 a e 43 3.) - 4.0 Ad 0.52 42-47 4.2 O40) I Vo - - 66-734 71 285 Measurements: Table 3. Description of Holotype male Very small, cervical region long and attenuated, post- vervical region plump, tail curved, conical, rather long and broad with recurved tip. Strongly annulated, sharp border angled forward, especially in amphidial region. direction sharply inverted 54% from anterior end. Six very small cephalic setae inserted at base of buccal cup, from which narrow, parallel-sided cuticular buccal tube extends length of cervical region. Large circular umphids located over end of buccal tube, annulation continues between amphids. Cylindrical pharynx, heart-shaped cardia. Thin setae spaced along body from cervical region to anus. Spicules cephalated, asymmetric, slightly unequal size, without strong rectangular Curvature, tips turned up. Gubernaculum encloses middle of spicules, strong dorso-caudal apophysis. Paratypes: Amphids dimorphic, smaller in fenrale, vulva with operculum, vaginal canal not cuticularised. Lorenzen’s formula:- b,a,p*%r,+ .2.0,u. *new letter because spicules do not correspond to any of those figured by Lorenzen (1975). Differential diagnosis This species is very close to R. collare sp. nov. The two are sibling species from the same beaches bur can be clearly distinguished by several features. Annulation continues between the amphids in both sexes, whereas there is smooth cuticle in RK. collare sp, noy, The male amphids are not as large. The vaginal canal is not cuticular and the spicules show greater asymmetry, KR. tomakinense resembles R. chiloense Lorenzen, 1975, and R. scutatum Lorenzen, 1971, but has quite differently-shaped spicules. Habitat Sandy ocean beaches. Distriburion New South Wales, Northern Territory. Rhynchanema chiloense Lorenzen, 1975 Material examined One male and one female and juvenile, off-shore, Cronulla, NSW, one temale, Rapid Creek beach, NT, Description Lorenzen’s formula:- b.s,ijr,+,2,0,u, De Man's ratios:- a = 18-23, b = 3,2-3.6,¢ = 8-10, V = 73%; inversion of annulation 52%. Spicule are length 26, more than twice length given in Lorenzen’s paper, but that seems by comparison with the drawing to be chord length, and by comparing drawings our male's spicules are only slightly longer in a larger male. The cuticle and body setae are stronger than in the other species ol Rhynchonena we have found NEW XYALIDAE 4) 26 20m . "eg / } >Oum —_-__—_—_ : _ —_ Rigs 25-30, Rhynchonema collure sp. nov. by SEM. 25, male head, and cervienl region. 26. female head and cervical region. 27. annulation pharyngeal region: AAyichonema tomakinense sp. nov, 28, female head and cervival region 29. tale cervical region. 30. female posterior, AM amphid. OP operculum, AN anus. 58 Habitat Sandy ocean beach and shallow sub-littoral sand. Distribution New South Wales and Northern Territory. Rhynchonema gerlachi Vitiella, 1967 Material examined Three males and one female, off-shore Cronulla, NSW, Description Lorenzen’s formula‘- i,s,z,r,=.2,u. Agrees with Vitiello's (1967) description except that he did not illustrate the vulva. Our specimens possess an operculum but no strongly cuticular terminal duct corresponding tou in Lorenzen’s formula. According to Viticllo the spicules have bifid tips, but his igure shows sharply pointed tips with one spicule rotated on its long axis, The spicales in our specimens are just like his illustration. According to Vitiello there are three pre-anal papillae in males but these are not shown in his illustration. We observe one 1o three minute pre- anal bumps, but cannot determine whether they contain papillae because of the strong annulation, Our specimens are smaller than Vitiello’s adult specimens, L=390-632 compured with 742-793 but De Man’ rauos in our specimens:- a=20-26, b=3.5-4.8, c=6.340.2, c =2,8-3.9. V=75% are in agreement with Vitiello’s, Habitat Shallow sub-littoral and intertidal beach sand. Distribution New South Wales WARWICK L. NICHOLAS & AIMORN C. STEWART Prorhynchonema Gourbaull, 1982 Buccal cavity in two parts; anterior short, at level of cephalic setae, posterior tubular, not more than 15 um, Amphids circular, placed well posterior 10 end of buccal tube, Prorhynchonema gourbaultae sp. nov. (FIGS 31-35) Holotype; Male, SAMA V4262, Kivloa beach, NSW, 31.vii 1986, Paratypes: 3 males and 3 females, AHC 24822, Kioloa beach, NSW, 31.11.1976. Measurements: Table 4. Descriptian af Helarype male Body cylindrical, anterior atretiuated from level of amphids to smal! head, about 10% of body length: tail conical, curved ventrally, tip slightly reflexed. Cuticle weakly annulated, annules about 1 ym wide, first annule wider, inversion of annule direction 50%: sparse thin somatic setae. 11 ym Jong. Inner labial setae not visible. six short cephalic setae in one ring, less than 1 ym long, amphids circular, cryptospiral, 30% body width, situated much farther posterior, 9.5% of length, about 55 annules from mouth, well beyond end of buccal tube. Buceal cavity long narrow parallel-sided tube, 3.8% of body length, slightly expanded at level of cephulic setae; pharynx cylindrical, widens gradually as body widens behind level of amphid, cardia rounded. Spicules cephalated, smoothly curved, slightly attenuated narrow spoon-shaped rounded tip, gubernaculum simple plate with small caudo-dorsal apophysis, three past-anal caudal glands, Taare 4. Measurements of Prorhynchonema gourbaultae sp. nov, Type Holo Male Paratypes n = 3 Female Paratypes n = 3 Male Range Mean Range Meun Length 452 460 ~ 49| 476 439 - 453 445 Width 15 16-18 (7 19 - 20 19 Ruccal cavity 17 15-16 16 15-16 13 Amphid 2.6 25-3 28 3-35 3.3 Mouth to amphi 43 40 - 47 43 39 - 42 4\ Pharynx 144 1Q2 - [22 7 J 28 - 138 (32 Mouth to vulva - ---= - 30% - 322 414 Mauth to anus aK4 39) 6 420 4dOR 3K0 - 396 AMT. Anal breadth \4 l4 4 \4 14-14 i4 Tail 66 62-7] 68 53-62 57 Spieule 22 23-25 24.5 - - De Man's a ay 27-3 29 22-24 23 J b 3.1 3.8 - 4.4 4.) 4,.2-3.5 a4 us c 648 66-77 TA 71-84 AB ” a Aq 44-51 48 38-44 41 " V% - - - Ju-7) 7! 3 —_ NEW XYALIDAE 59 32 33 \ ae _— , 4 x ae ee 31,32 25 um a 33,35 25 um 34 50 nm Figs 31-35. Prorhynchonema gourbaultae sp. nov, 31. male head. 32. female head. 33. inversion of annules. 34. entire female. 35. male tail and spicules. 60 WARWICK L. NICHOLAS & AIMGRN C, STEWART Parutypes: Female paratypes resemble male holotype. apart from reproductive organs, but amphids slightly larger, 2.7-3.5 ym cf 2.6-3 yim. Single anterior ovary, vulva simple not cuticularised, no operculum. Habitat Intertidal sandy beaches. Distribution New South Wales. Differential diagnosis The new species differs from the other described species, 2 warwicki Gourbault, 1982, by having very different spicules and a longer cervical region and buccal tube. The relative length of the buccal tube is similarto R. brevituba Gerlach, 1953 but the spicules ure different and in R. gerlachi annulation does not show. inversion, Prorhynchonema warwicki Gourbault, 1982. Material examined Three males and 3 females from Rosedale and Tomakin beaches, New South Wales. New subfamily Corononeminae With characters of tamily, Buccal cavily cylindrical, deep and wide, not enclosed by pharyngeal musculature; amphid situated above base of buccul cavity, Cervical region only slightly attenuated. Six inner labial setae in one ring. six outer labial and four cephalic setae in second ring. Strong cuticular annulation begins at base of buccal region. TABLE 5. Measurements ef Corononema parvum sp. nov. Genus Carononema gen. nov Type species: Corononema parvunt Diagnostic definition Cuticle strongly annulated with cight longitudinal ridges. Lips high, incised. Deep groove around the head just posterior to the insertion of six inner labral setae, in [ront of outer labial and cephalic setae, sets off the lip region from the slightly tapered buccal region. Frymoalogy Corona, Latin for crown, because the head appears to be crowned by the incised lips, set off from the cylindrical buecal region by a deep groove, Coranonema parvum sp. nov. (FIGS 36-40, 50-53) flolotype. Male, SAMA V4263, Rapid Creek beach. NT, 31vii. 1986, Faratypes, 6 males, 5 AHC 24823a-c, Rapid Creck beach. NT, 31.vii.J986; | male, AHC 25824, Southport beach, Qld; 7 females, 6, AHC 24823, Rapid Creck beach, NT, 31.vii, 1986; 1, AHC 24824, Southpert beach, Qld, 31.vii.1986, Measurements: Table 5 Description of Haletype male. Cuticle strongly annulated, annules LS jm, with eight equidistant longitudinal ridges, weakly developed in cervical region, pronounced mid-body, which is polygonal in cross section, extending almost to blunt Type Holo Male Range Length RSS BON = [045 Width 27 22-26 Annulition 1.6 |.5-2 Lip height 3,5 45.6 Outer labial setac 5 4-6 Sub-eephalie setae \2 \}- 19 Body setae if 13-19 Amphid 5 4.48 Buecal cavity, length {3 Hh 46 Bovedl cavily, width 9 R10 Mouth t nerve ring 97 90 - 120 Pharynx 322 285- 360 Mouth to vulva - . 2. Mouth to anus FRI 714.473 Width at anus 2) 19.23 Tail y4 72-87 Spicule, are 2 25-31 De Man's a 32 32-44 " h 7 2.6 - 3.6 ml c LLG O34 - 14a by c a5 33-45 M V% - : Male Paratypes n= 6 Female Paratypes n = 6 Mean + SD Range Mean + SD 9 i) 840 - 1084 977 71 24 «1.46 22 - 38 28 4.3 18 O19 1.7-2 19 (12 34 059 4-55 5 050 46 (89 6-9 6.9 1.0L IS.4 0 2.97 {- 12 ia 1.00 16.2 2.39 95-2) a5 499 46 0.40 44-6 3.20 O82 I3.2 0 1.97 13-18 5.7 1.80 9 0.63 8-11 9.9 95 1o0 \] LNQ~ | 32 107 13 4 30 310-380 452 26 > ° ON7 ~ ¥70 799 63 843 = 107 TH - 975 900 74 2t 133 20-25 22 (A,27 79 5.92 68 - 105 80 12 278 2.56 - - - 7 SOI 26,0 - 40.0 33.0 At 29 0.36 2.4-3.1 24 0.27 1G 204 OB - 15.2 |2-4 191 AR 0.47 320 - 5,30 3,80 O7t = : 80-84 $I NEW XYALIDAE 6] 36 ——_———_______ 25 um Pe 3g: —_——_—_———__. 25 1m hy 37 —&- cm — 100 um 38 ——————————_ 50 um 40 ————————- 100 um aye 40 \ \ Trt prvi nitl yy vi pysa prey VUVHPVIV PET py yyy Vii Figs 36-40, Corononema parvum sp. nov, 36, male head. 37. head and pharyngeal region. 38. male tail. 39. spicules. 40. entire female. 62 WARWICK L, NICHOLAS & ALMORN C, STEWART cylindrical tip of tail, Cervical region enclosing buccal cavity nol annulated or ridged. slightly amenuated rowurds mouth. Deep groove, with less strong cuticle, surrounds buccal region just below the lips, Uwe scalloped cuticular rings: lie just within mouth. Six incised leal-like lips, six inner labial setae al base of hips. six outer labial and four cephalic setae insert posterior torhe groove: four strong cervical setae, |2 pen insert at base of nen-annulated buccal vegion on first annule, numerous body seiae, amphid fovea situated over base of buccal cavity, 5 wm diameter, 25% head width. Wide, deep buccal cavity with strong almost parallel-sided walls. Cylindrical pharynx, Two outstretched, inactive testes: spicules cephalated, rectangular curvature, simple pointed tips; gubernaculum surrounds spicule tips. Paratypes: The spicules ditter somewhat in the degree of curvature and may appear different because of partial rotation about their axes and the pointed lips may be turned ouiwards. Females. apart from the reproductive organs. closely resemble males, Females possess a single anterior gonad; the vulva has no operculum. The labial and cephalic setae are difficult ro measure by light microscopy because of their small size, but from scanning electron microscopy the inner labials are about 1.2 pm tong, the outer labials about 0.5 um and the cephalic about 2 wm long. Differential diagnosis The deep parallel-sided buccal cavity, without teeth, with lour strong setae at the base is distinctive, The indented weakly cuticular groove below the insertion of the lips is anlike that found in other Xyalidae. TABLE 6. Measurements af Corononema thai sp. nev. Habitat Sandy ocean beaches, Distribution Tropical and sub-tropical beaches in the Northern Territory and Queensland. Australia. Corononema thai sp. noy, (FIGS 41-49, 54-56) Holotype: Male. SAMA YV4264. Pathaya. beach, Chonburii, Thailand, 30.ix.1985. Pararypes: 4 males, AHC 24825, und 7 females, ABC 24826 Pathaya beach, Chonburii. Thailand, 30.ix 1985. Measurements; Table 6 Description of Holotype mate Strongly annulated cuticle, anmules. 1.6 jan wide, 8 longitudinal ridges from buccal region almost to (ip of tal, Buccal region not attenuated, only slightly widening [rom mouth to base, enclosing deep, wide unarmed buceal cavity. Lips high, leaf-like, fold over wide mouth, Lip region sepanuted from buccal region by deeply indented weakly cuticular circum-oral ring sandwiched between two circum-oral erenellated cuticular rings. Six inner labial setae inserted at base of lips. $1X outer labial and four cephalic setae inserted below indented ring. Long thin body setae spaced along bady, six cervieal setae inserted on first anoule at base ol buceal region, Cylindrical pharynx. Two outstretched testes. anterior to left of gut, posterior lo right. spicules cephalated, rectangular curvature, simple pointed tips; gubernaculum surrounds spicule Ups, Type Hole Male Runge Lenuth 9900) 642 - 95% Width 27 25-24 Annulation 1.6 16-17 Lip height 44 4-5 Outer labial sete 6 5-8 Sub-cephalle setae 4 yold Body setie 3 l0-16 | | Amphid + 7 -5.3 ie Buceul cavity. length 2 D168 Buceal eavity. width 2 K-11 Mouth to umphid 7 6-15 Mouth to nerve ring 76 8a - 118 Pharyns 264 24) - JKR Mouth to vulva - - "= Mouth to urus 19 AR - RIS Width at anus 2a 20 «24 Tail 7\ Hl -6% Spicule, ure 32 2h - 32 De Man's a an 24.0 - 384 " b An 2.5 -F.5 ls c 149 Ws IS2 4 c 0.4 2.6 3.) fi Vy Mule Parutypes ne 4 Female Purdivpes p= 6 Mean + Sd Runge Meun + SD 74 138 TO LOK 946 106 21 1,29 a) - 35 32 1st a) 0.05 14-14 1 OS ti DAS 5-75 7 At 64 144 h- 10 94 |.o3 \2 2.50 1O- 12 " 1 has «2.50 Ihe 14 12.4 152 46 O75 4-6 340 OH 125 2,89 1-13 1.7 (2 Ya 800 W- 12 \! ttl 1 3.74 3-74 58 LAT O5 16 7-42 h2 OMI 262 21 2K5 - 424 Alb SQ - m0- 871 TOR Kou OM] 13h 722 - O8k 858 1M) i V7 22 - 26 24.2 143 th 2.99 on = TOT BO \3 2s 263 isp 28.00 6.46 23,0 - 34,0 293 393 25 (AS 20-34 3.0) O34 HY 228 YO - 13.3 1.2 1.74 aN 24 1,10 ~ 4.00 60 Oda - nO - B+ at 1.60 NEW XYALIDAE 42 44 feames varia n “e fy Sie 2,8 ‘ » ne p~ C—O 43, ——_—————_ 100 pm 42 ———————. 50 jim Figs 41-44, Corononema thai sp. nov. 41. male head. 42. male tail. 43. head and pharyngeal region. 44. spicules. 64 WARWICK L. NICHOLAS & AIMORN C, STEWART 46 Figs 45-49. Corononema thai sp. noy. 45. cross section drawn from fractured specimen viewed by SEM. 46. spicules in ventral view. 47. female posterior end. 48, annulation by SEM successively near head, mid-body and near tail. 49. entire female. NEW XYALIDAE 65 30 pim Figs 50-56. Corononema yen. noy. by SEM, 50-53. C. parvum sp. nov. 54-56. C. thai sp. nov. AM amphid, LP lips, ILS inner labial setae, OLS/CeS outer labial and cephalic setae, SCS sub-cephalic setae, SP spicules, VL vulva, AN anus 66 WARWICK L_ NICAOLAS & AIMORN C, STEWART Paratype. Female paratypes possess a single anterior ovary, vulva without operculum, prominent post-vulyal hand. otherwise females closely resemble males. Differential diaynesis © thai sp. nov. diflers From C. parva sp.nov: by possessing a shorter broader head, Asan index of this ditlerence the ratio of length to breadth of the pre- annulated buccal region has been measured. In C. parviin the ratio ranged from 0.87 te 104 (=30), in C. that trom 0.64 to O75 (n=10), The annulation is shallower in C. thai. Both these properties are most vlearly seen in scanning electron micrographs of the hesel (Fig. 52). Habvitert Tropical sandy beaches. Distribution Vhuiland. Discussion The taxonomic position and rack of the Corononeminac presents difficulues. The circular eryptospiral amphids and single prodelphic female gonad are characteristic of Nyalidae. The strongly annulated cuticle and buceal cavity lacking teeth are also-shared with most Xyalidae, although some possess toott-like ridges in the buecal cavily, The deep unarmoured buccal cavity with strong cylindrical walls is found in other Xyalidae. such as Omicranemtee and all Rhynechonemutinae. but Whereas in Rhynchonemutinde the buccal cavity terminates as a narrow tube of varying length, the buecal cavity of Crrenoneme is relalively wide and short. The cephalic region isattenuated.. but not to the marked degree so chargoteristic of Rhynchonematinae. For these reasons the new genus, Caroyionemer, 18 best placed within the Xyulidae, but would stretch the definition of the nwa previously recozmsed subfamilies too far so that the erection of d new subtamily seems warranted, Lorenzen (1978) in tis review of the supertiumily Monhysteroiea does not recognise any subfamilies within the Xyalidae. noting, in this regard, that essential aspects of the phylogenetic relationships within the Monhystepojdea have not been cleared up. We do not find this sufficient reason to abandon the previously recognised division of the Xyalidae into two clearly separable subfamilies, namely the Cobbinue and Rhynchonematinae. Whether we are justified in creating another subfamily for two species with intermediate characters must be a matter of personal judgement. In Coronanenia the form of the cervical region and buccal cavity are intermediate between that tound Rhynchonematinae from Cobbiinue, but there is.no overlap.and clear diflerences remain between the three taxa. The deeply incised labial region and a flexible indented ring separating the cephalic region from the buceal region ure mm our view significint distinguishing attributes of the Corononeminae. Tt may well be when the very poorly known nematode faunae of Australia and South East Asia ure betier known more species will be found assignable to (he subfamily. The indented cireum-oral ring is probably flexible und may facilitate the mgestion of larger paruicles than would otherwise be possible. A flexible oral region has been Observed to lacilitate the ingestion of relatively large diatoms by other Xyalidae such as Dapronema- Acknowledgments We are grateful for a grant from the Australian Biological Survey which made this work possible. We thank Dr Russell Hanley, Northern Territory Museum of Arts and Sciences for making it possible lor us po collect on Darwin beaches, References Geriack. S. A. (1953) Die Nematodenbesiedlung des Sandsttandes an des Kiistengrundwassers an der iluhenischen Kliste. J-Systematischer Teil, arehe. Zo, fal 37, 517-640, GOURMALILT, §. (1982) Nematodes marins de Guadeloupe. I, Xyalidae nouveau des genres Rhynchonemea Cobb ol Provunchneme nov. gen Bull, Muy natn. Hist Nat, Paris HW ser, AL Sec, AL tas. 1-2 75-87. Hopper, BE. 961) Marine nematodes trom the coastline of the Gultal Mexico. Can. Zaol, 39, 359 365. Lokisven. oS (1971) Die Nematodentauna im Verklappungseebeit fir Industnieabwasser norehwestlieh von Heigoland. |. Araeolainida und Monhysterida. Zod/, Ane 187, 223-248, (TY75) Rhynchynema -Arien (Nematodes, Monhysterutae) aus Sidamerika und Europu. Mrcrafdiya des Meereybodeny $5, 225-251. _—____ W978) The system of the Monhysterdides (nerialodesy - A New Approach. Zant, Jb. Svs. 105, 515-536. STEWART, AC) & NieHolas, WL, (1994) New species of Xvalidge | Nemutoda: Monhysteridy) from Ausiralian ocean beaches, Invert, Texan, 8, 91-15, Viigo, PL 1967) Deaux nouvelles espetes du venre Rhynchonema (Nemittoda, Monhysteridae), Bull, Sac, cel, Fe 92, Al3-121. A NEW GENUS AND SPECIES OF CRANGONYCTOID AMPHIPOD (CRUSTACEA) FROM WESTERN AUSTRALIAN FRESH WATERS By J. H. BRADBURY* & W. D. WILLIAMS* Summary Bradbury, J. H. & Williams, W. D. (1995) A new genus of crangonyctoid amphipod (Crustacea) from Western Australian fresh waters. Trans. R. Soc. S. Aust. 119(2), 67- 74, 31 May, 1995. A new genus of crangonyctoid amphipod (Crustacea) from Western Australian fresh waters, Totgammarus, with a single species, T. eximius, is described. The species was collected from roadside pools in the south-west of Western Australia. Key Words: Amphipoda, Totgammarus, Western Australia, crangonyctoid, Paramelitidae. Transactions of the Reval Soerery of S. Ause (W9S), WH2). @T74 A NEW GENUS AND SPECIES OF CRANGONYCTOLD AMPHIPOD (CRUSTACEA) FROM WESTERN AUSTRALIAN FRESH WATERS by J. H. BRADRURY*® & W. D. WILLIAMS* Summary Reapnory. 1 H& Witaws, W. BD, (1995) A new genus of crangonycloid amphipod (Crustacea) from Western Australian fresh waters, Tras Ro Sec. 8S. dust. 92), O774, SL May, 1995. A new genus of crangonyetoid armphiphod (Crustacea) from Western Australian fresh waters. Jorguninaras , wilh a Siigle species, Fo cranius, 1s described) The species was onllected from roadside pools in the south-west if Western Australia Kiy WoRxps: Amphipoda. Jorgiminaris, Western Australia. cranganyeioid, Paramelilidae. Introduction All known crangonyctoid species of Australian fresh witlers [9 IY87 were comprehensively reviewed by Williams & Barnard (1988). In their review, knowh species were re-examined and redescribed, and some new species were described. They dealt with a total of 12 genera and 33 species. They considered the number of genera probably represented about half the number expected to oecur in Australia and noted that the number of species within genera was probably small. Purther work (Barnard & Williams, in press) supports this view; they deseribed two new genera. euch monotypic, ay well as a further new species of both Ansrogammaras and Uractena. This second review by Barnard & Williams (in press) described, ier alia, most material available to them from Western Australia, They did not deseribe, however, a taxon from thal State represented by only a single specimen, pending the collection of further material, Unfortunately, all attempts to obtain more specimens hive proven unsuccessful; exhaustive exammation of all the known collections from the area have yielded no further specimens and nor did collections made in 1994 on our behalf by A. J. Boulton. Since the single available specimen represents in our view u new genus, and to Jacilitne further studies of freshwater amphipods in Western Australia in particular and Australia in general. we now consider it appropriate to describe this single specimen. Methods oF dissection, description and potation follow those of Williams & Barnard (1988). To expedite the use of figures in the present publication, the abbreviutions are as follows; “Antennal sinus” refers wnly to the cephalic sings receiving umtenna 2. A - iumcnna Abd - abdomen; ace - acteessory; art -article; C - coxa, cox - coxalsd - dorsal; dact - dactylus; @ - eye; E- epimeron, fag - flagellum; g - gill; G - © Deparment of Zoology, University of Adelaide, South Australia SOOS. gnathopod: Hd - head) i- inner, | - lefts lac - tacinia miobilis; lat - lateral; LL - lower lip: MD - mandible; micd — medial; mol - molar: MP -maxilhped; MX - maxilla; o- outer; O- oostegie, p-palp: P - percopod: PC - prebueeal complex; pl - plate: post - posterior, Pp - pleopod: r - right; ret -retinaculum: st - sternal; 'T - telson; U - uropod: UL -upper lip: v - ventral; 1,2,3...7 - first, second, third. seventh, Genus Jotgammarus gen, nov, Eymtulogy Named for the combination of features of several gener, Type species: Totgenemerks extanus Diagnosis Pleon with sparse dorsal setation, pastrum weak Lateral cephalic lobes strongly. proyecting, antennal sinus moderate. eye not discernible in preserved specimen. Flagellum of first antenna lacking mayor armainents, moderately long. about 05x body length, iwive A2, Ratio of peduncular articles 2;2:1, Flagellum of second untenna and peduncle of sub-equal length, culecolt of type 9 present (Lincoln & Hurley 1981). Ratio of mandibular pulp urticles abour 2:9:6, article 2 moderately Setose. arucle 3 faleute. setae = ABDE (Barnard & Barnard 1983). Lubium lacking inner lobes, Maxillaeé 1-2 medially setose, inner plate wholly or marginally pubescent, Maxilla | outer plate ovate. medially ind laterally setose, palps asymmetric: left with thin apical spines. right wilh thick apical spines. Maxilla 2 inner plate with row of apico-medial weakly sub-muarginal setal spines. medial margin heavily setose. Muxillipedal palp arucles 2 ~3 with few lateral setae. article 3 with fine factal pubescence dorsally and a ventre-facial row of moderately long. curved setac. O8 1H, BRABBURY & WD. WILLIAMS Coxue 1- 4 willha cow of postenor spines. coxac ) 4-moderitely elongate. coxa (tapering below, cox d deeply emurginate post-dorsally, coxue 5 - 6 shorter thin 4, coxa 7 shorter than 5 - 6, Gnathopods unequal, soathopod | O.5x gnathopod 2. Carpus ol gnathapod | long, of gnathopod 2 short. Scythe spine absent from artele + of both gnathopous. each with a weak lobe, Propodus of both grathopeds rectangulir, palins weakly to moderately corivex, palmer corners prominent, lurned oul. First gaathopod lacking stron spines at palmar corner, second with G strony spines. Spines along palms of both gnathopods short, simple, without triggers, numerous. Perenpods 4 - 4owith posterior spifie sets on article 6 evenly spaced, PS - 7 moderately elynwute, urticle 2 broudly expanded, lobate Dueryls with | -3 sprites. Coxac 2 - 7 with sav-like gills. Thoracic segments 2-7 with dateral steraal gilts. Basumedial selae on inner rami al pleapods 1 - 3 plumuse, peduncles each with paired retingcula and paired (Hirst and seeond pleopod) or single (hind pleopod), plumose kecessory retinacula, Pleomtes With few dorsal setae and/or spines, Epimera with few ventral spines. posteriar qmargins Weakly setulite. Rami of uropods | - 2 extending sub- equally, each with 2 rowe of spines. Facial armament of Uropod Tweak, largely absent on uropod 2 which hears u strong, elongate upteo-medial spine Liropad J extended, magninumous, peduncle shart, ouler ramus 2 aniculine. artiele 2 short, Telson longer than broad, 100% eleft, not laverally turin, apically and disto-laterully weakly selose, bearing a single sub-apical spine on either lobe Additional déseription Flagellum of Al-2 lacking major armaments. Apical thargin of labrum. extended. Accessory blades (rakers) on mandibles with inter-raker plumase setae interspersed among rakers und additional Short plumose 4elle Iving between rakers and molar Matar trilurative, with plimose apical seta, Mandibular palp article 3 shorter than 2, palpurticle 2 Jacking baso-anterior setae with few median and apico-anierior setae. Both plates Of second maxillae with rows af long distal setae Maxillipedal palp inoderately long. Article 3 weakly produecd and finely pubescent at the upex which bears lone terminal setae; baso-medially hearing a sirle sub- marginal seta, medially Wilh a row of scythe selae extending to the base of the duetyly selae of [he venirul Jace constituting a comb row, aswell as u single long mid-lacial sela and 4 naw of shert setae basal te the comb row: jhe mid-distal dorsal face beanny fine pubescence DactyLof first gnathopod not reaching palmur comer, bearing a small, bent tiner tuoth-spine. Bucry) ut second ynuthopod reaching i end of palm, bearing Jsmallinner spines, Peeeopod 7 shorter thin 6. Article 2 of pereapods 5 - 7 equally setose pasteriorly. Sternal processes: fleshy sausage-shuped pills on thoracic segments 27, aniehed to mid-lateral argeins of segments, Puslero-ventral apex of epimera | - 3 blunt. as in Ausirogaminaras, Pleopods similar, except for numbers of retinaculae, rami approximately equal. Outer cunius of urupods |~ 2 slightly shorter than jnner ramus. Apidolateral corner of peduncle of uropod UL wilt 2 Spines, ram of both first and second uropods with > upical spines. Phe third weopod extending beyond the first and second in the entire animal, pedunenlar spines. apical. and Sub-apical, some medial sete of each taitius Plumose. Ventradistal spine on urosumite 1 ur base pl drypod | short, as in Auatragennaerin, Relarlonship This genus displays the eharicteristies ul crangonyetoid amphipods in possessing (Ly) sternal gills (2) un accessory Magellun of the first antenna with two or more drticles (3) ealeeoli of type 9 or linear (4) uropod | licking a basolacial spine on the pedunele (5) a lower lip without ianer lobes (6) a frst ynathopod thitt is not melita or mittenform in shape (7) 4 first enathopod that does not dominate the second, and (4) a inandibular palp of typical form (Williams & Barnard JORK). The new genus fits the essential criteria of the family Paramelitidue in possessing sausage-like sternal pills, dorsal setae on the telson, and linear or type-9 calceo|). VW differs front the Neoniphargidae in the ubsence of rugosities on the third article of the maxillipedal palp and gnathopeds. the form of the gnathopods (not srl! und mittentorm), the form of the carpi (not short and lobate), and non-dendritie or luip bearing sternal gills. It differs from the Perthiidae in thar the first antenna is significantly longer than the second. the mandibular molar is normally developed and triturative. rhe outer plate oF the maxilliped is nor very small die enathopods are not large. nor are the carpi short and deeply lobate. the curpi and propodi are not eusirid, dnd the sternal gills are pet dendritic, Joteannarns. bears atures in Common with alter paramelitid geuera, such as blindness, which occurs ie several, and m possession of an eclangate spine on fhe secand male uropod (as in-some Urvetend spp.) The combination of characteristies however, is uniguc. The genus varies from Austrogamtimarus, the most primuuve Australian paramelitid genus, in-several ways, In Jorgaminarns, dorsal setation of the pléonites: is weak, the lateral cephalic lobes project strongly, the antennal sims is Moderate, eyes ure absent, and the A NEW GENUS AND SPECIES OF CRANGONYCTOID AMPHIPOD 69 Fig. 1. Totgummarus eximius, sp. nov. holotype, male 10.6 mm. Whole animal, antennae and mouthparts, Scale bars: adult and antennae = | mm, mouthparts = 200 pam. WW | H. BRADBURY & W, D. WILLIAMS second urlicle af the mandibular pulp 1s relatively long will lew wpieo-unterion and no baso-anterior sete, Additionally. there is a extension of the apical margin of the labrun, 5 rather than 3 apical spines occur on the wher plate oF the maxilliped, cosae 1-4 are mderdely long ratber than elongule, the apex of coxa | Lapers, pereopods 5 - 7 are more even in length, and a seythe spre iy hol present on article four of the gnuthopods although a small lobe js present, The daetyls of legs 3 - 7 are mulliespinose, The peduncles (the pleopods are moderately setose. the apieo-lateral and apico-medial spines of peduncles of the Sivst uropod differ, usdo the relative lenpths of the urepod vam which also bear busu-ficial armaments. The third Uropod is miagniranois, Toteainmaruy eximiuy sp. nov, FIGS | - 3. Pryvinelaes! From eviniay. mieaning exceptional or alone. dype hacetiry lomporary roudside water in sands along the Scott River Road. south-western Western Australpt Dern With the charaateristies ab the gens (only trate Known, Material Evanined Hatoiwpe Western Australian Museum WAMI4-95. male 10.6 Min in rype series, Nooather specunens available. Beseripiton uf halvivpe (male) Body (Fig. t), pleanites 3 = 6 with Sparse transverse dorsal selalion and dorso-laieral spines on 3 - 6, firstamtenna (Pig, 1) primary Oagetlim sparsely s¢ioxe, Mugellunt of 35 articles. 18 x pedunele. No culeeoli, Accessory flayellunm 7 - 8 articuliiie. reachine, teanicle & of (he priury flagellum. Sccond antenna (hig. Ves Jength 0.25 x beady length, pedunecular articles 4S subequal, Magellum of 19 articles, setae sparse. Calecol) on urticles 1-13. Labrum (Fie 1: broadly rounded with apex slightly extended. harerully and apically pilose, Labium (Fig. 0: medially and laterally pilose with (0 vurved upico-medial spines on either lobe heh mandible trig. Ue palp article 2 senjtien 1A 2B - 15D - AL. article 2 with 2 Inedial »etic and Ob)QUe PAW EO apico-imedial setae, Incisor 6-tuelhed, lheinia mobilis +-roothed, 9 setose accessory: blades. 3 short plinose sete und | short bluntspine taward baseot molar, Anterior of molar densely pilise. Motar with) shork plumose seta. Right mandible (Pu 4. incisor 4-toothed, Jacinia mobilis bifid with 4 dentioukite weth on one side and 9 cuspate teeth and al blunt terminal tooth on the other, accessory blades of 3 toothed spines and 4 sctose inler-rakers, 3 short plumoseselae and | narrow blunt spine toward the buse ofthe molar, Molar with | sctose median, short. blunt Spine, and long plumose seta, Lefl first maxilla (Fig, 1): palp-article 2 with 10 thin upigal spines, otherwise naked, Outer plate medially setose. JO denteulate terminal spines, Inner plate ovate, laterally and medially with sparse straw-like pubescence, 5 upice-medial plumose setae, Right first maxilla (Pig, Ds palp article 2 with 6 thick apival spines. L disto-laterul moderately long spine und | luteral Sub-upical curved spine. Outer plate with 10 denticulute terminal spines abd 1 aatero-medial plurimose seta, Media area Witt long pubescence, Inner plate ws for left side. second maxilie (Pig. 1) syinnietrical, outer plate laterally setose, sub-terminal row of (0 curved spines. fertninal row ef many curved setae. Inner plate laterally seluse, SUb- imirginally pubescent; medial margin with fine sete proximally, row af setal spines distally. Masilliped (Pig. 1): palp article 3 with 9 inedial sevihe setae, l6-antero-frctal comb row see extending from MO4 to the sub-upex, 4 long terminal setae, 2 nid lateral setae and 1 median sew. Dacty! bearing [distil and | oanedial accessory spines and, dorsally, 4 short post-lacial setae basal to the comb row: outer plate laterally sctose. apically bearing sub- terminal row of 8 strong curved spines, a disto-medial sub-facial row of [O Tooth spines and 13 setae, selue distally sub- facial fo the teeth, proximally facial, Inner pliite terminating in 5 strony tooth spines and & plumose selves medially 6 long setal spines, the distal 4 plumose: basal to the inner plate g transverse row of % mediviy to long naked setae, First ynathopod (Fig. 2). coxa tuypered, 3 posteriar spines, weakly setose marginally: carpus moderately long. sub-equal to propadus, not lobed: propodus reclangulir, pulmar corner prominent, extended posteriorly, pal ucute, convex, daety! reaching cornet of palm, bearing small bent yner tooth, 5 phumose und | naked spines al palmer enrner: numerous short spines along palin Second gnathopod (Fig. 2)! larger than the first (Lett 2x, Right 1-78); coxa wilh row of 4 or 5 posterior spines, few snail distal setae; Carpus short, about 0.5% propodus: propodus reetanmular, longer than wide, palmar corner with 2 strong, naked spines and 4 pluniose spines. corner prominent and slightly extended posteriorly: dacty! not reaehing corner, but reaehing tothe second naked spine. bearimy 2 inner teeth at approximarely MOS; palin slightly convex with HUMETOUS SHORE Spins A NEW GENUS AND SPECIES OF CRANGONYCTOID AMPHIPOD 7) Fig. 2. Totgummarus eximius, sp. nov. holotype, male 10.6 mm, Gnathopods and pereopods. Scale bars: gnathopods and pereopods = | mm. dactylar enlargements = 200 pm. 72 J, H, BRADBURY & W. D. WILLIAMS — a a Fig. 3. Jotgammarus eximius, sp. nov, holotype, male 10.6 mm. Pleopods, uropods, telson, gills and abdomen, Scale bars: abdomen and gills = 1 mm, pleopods. uropods and telson = 200 pm. retinaculae = 50 pm, A SRW GENUS AND SPECIES OF CRANGONYCTIOD AMPHIPOD Va Pereopods (Fig. 2): coxa 3 with 9 posterior spines, cox + deeply emarginate, small setae and spmes below. no posterior spines. Coxa 5 bearing 3 posterior ventral spines, coxa O with 3 posterior ventral spines and 4 small posterior setules. Coxa 7 with 4 posterior spines Pereopods 3 - 4 length 1.2 x G2, subequal, article 2 ol both bearing long posterior setae, Article 5 of P3 apicu-posterior spine formula (proximal to distal): |-2-2-2 article 6) 1-3-3 -3-2-2-2 article Sof PA, 2-3 - 3- 4,artivle6;3-3-3-3-3-3 3. Pereopods 5. 7 of approximately equal length, Pereopod 5 articles 3 - 6 bearing long apico- posterior selue, upico-anterior spine formulae 2-3 ~2 and 3-4-4-4-3-3-4-4. Pereopod 6 article § bearing few long setae, article 6 many apico-pasterior setae. spine formulae 4 - 3-4-0 und 2-4-4 -5 - () - ~3 respectively, Pereopod. 7 upico-untenor spine formulde: article 5S! 4-6-6 and article 6: a-3-4-4-4-3, Gills (Pig.3)> coxall gill 5 slightly reduced, gills 5 -7 bi-lobed. Slernal gills 2-7 lateral, Epimera (Fig. 3): with few ventro-facial spines. - posterior margins with few small setules. Epimeron | slightly rounded posteriorly with single antero-ventral spine, Epimeron 2 with 3 srnall mid-ventral setae only. Epimeron. 3. naked ventrally. Pleon (Fig. 3): pleonies 3 - 6 with dorsal spines and/or sétac. Pleonite 5 with 5 spines in transverse groups of 2 and 3, Urosomite 6 with 1 dorsal spine on either side, Pleopods! pleopods | and 2 bearing paired, hooked retinaculae and paired aveessory retinaculae, pleopod Slacking second accessory retinacula, Uropods (Fig. 3): lirst uropod; peduncle length [2 x ramt, outer margin with | upico-facial spine, 2 medial spines, and strong row of 5 dorsal spines, without setae. Ram sub- equal, terminating in a cluster of S spines. Second roped; peduncle length equal to rami, lacking spine rows, bub-with a cluster OF T large and 4 short apico- facial spines. Inner medial angle with elongate spine 0.5 x length of peduncle. terminally spoon shaped. Inner ramus 1,3 x length of outer, lacking setae. Both tami terminating ina cluster of 5 spines. Third uropod: peduncle iength 0.35 x Jength of outer ramus, about the same length as. urosontite 3, bearing median transverse row of 5 spines. distal transverse row of 7 spines at the base of the outer ramus and u group of 4 apico-lateral spines, Outer ramus proximal article strongly setose baso-laterully with 4 disto-lateral clusters. of spines and setae, paired medial und single Jateral trigger spines apieally, medially a single sub- apical trigger spine and evenly spaced plumose selue. Small distal article, 0.13 x proximal, terminating in 3 short and 2 long setae, Inner ramus of 4 single article, equal to the length of the proximal article of the outer ramus, marginally setose. the medial setae plumose, 6 lateral and 5 medial trigger spines distally; 2 terminal spines and 4 setae. Telson (Fig. 3): 1.25 x urosomite 3, cleft 100% - Disto-lateral margins und apex. with sparse dorsal setauion, paired penicillate setules sub-muarginal al M.80, Single sub-apical spime on either lobe, Distribution Western Australia (south west), Scott River Road, sands in a roadside ditch coll, K. Davies, B. Knott, 03 Oct. OBL. Acknowledgments The authors wish to thank Dr A. J. Boulton (Universily of New England, NSW) lor his efforts to collect further samples of the species in January and February 1994. A.B.R.S. support during the finalization of the manuscript is gratefully acknowledged. References Bakar, tol & Barsarp, C, M, (1983) “Preshwarer amphipndi: of the world. 1, Evolutionary. patterns, IL. Handbook wd bibliagraphy” (Mayfield Associates. Mt. Vernon, Virginia. — & Wiiatis, W. Do fin press) The wixonomy of Preshwatsr Amphipoda (Crustacea) from Australian Fresh Waters Part 2. Ree. Aut. Mus. suppl. Lincorn, R, 1o& Huguey, D, E, (981) The calecolus, a sensory structure of gamimaridean amphipods (Amphipada: Gammuridea). Bull Brit. Mus, Nar Hist, (Zool,) 40, (03-116, Wituiams, W. BD. & Barnaros. J, L. (1988) The taxonomy of crangenyctoid Amphipoda (Crustacea) from Australian Fresh Walters; foundation studies. Ree. Aust Mus, suppl 10, L180 ARROWIPORA FROMENSIS A NEW GENUS AND SPECIES OF TABULATE-LIKE CORAL FROM THE EARLY CAMBRIAN MOOROWIE FORMATION, FLINDERS RANGES, SOUTH AUSTRALIA By MARGARET K. FULLER*, & RICHARD J. F. JENKINS* Summary Fuller, M. K., & Jenkins, R. J. F. (1995) Arrowipora fromensis, a new genus and species of tabulate-like coral from the Early Cambrian Moorowie Formation, Flinders Ranges, South Australia. Trans. R. Soc. S. Aust. 119(2), 75-82, 31 May, 1995. The recently discovered Early Cambrian tabulate-like coral Arrowipora fromensis gen. et. sp. nov. occurs in the Moorowie Formation of the eastern Flinders Ranges. It is found in an ancient reefal environment in association with Moorowipora chamberensis Fuller & Jenkins 1994 and Flindersipora bowmanii Lafuste 1991. Arrowipora fromensis has tabulate-like characteristics including the cerioid form of the corallum, wedge-shaped to spine-like septa and strongly developed dissepiment- like tabulae. Although unlike any other Early Cambrian coral, skeletal characteristics are similar to some micheliniids, which have a time range from the Late Silurian to the Late Permian. Arrowipora fromensis provides further evidence that the time range of the Subclass Tabulata possibly extended to the Early Cambrian. Key Words: Arrowipora fromensis, Early Cambrian, Moorowie Formation, tabulate- like coral, Flinders Ranges, South Australia. Transactions of the Raval Serer of 8, Aust (995), WA2), 15-82 ARROWIPORA FROMENSIS A NEW GENUS AND SPECIES OF TABULATE-LIKE CORAL FROM THE EARLY CAMBRIAN MOOROWIE FORMATION, FLINDERS RANGES, SOUTH AUSTRALIA by MARGARET K. FULLER & RICHARD J. F. JENKINS® Summary FULLER. Mo K., & Jenkin. ROLE (1995) airrowiporen fromensis. a new penus and species of tibulate-like cor) trom (he Burly Cambrien Moorowie Formation, Flinders Ranges, South Australia, Tran. R. Sac. 5. Anse, 19(2), 75-82. 31 Muy, 1995, Ihe recently discovered Barly Carnbrian tabulate-like coral Arrawipord fromensis: gen. et sp. nov. Occurs ITT ihe Moorowie Formation of the castern Flinders Ranges. tb ts found in ain ancient reebil environment in association With Mooravipare Mamberensix Fuller & Jenkins 994 and Flindersipora bawntanil Latuste 199), Arrowrpard fromenvis has whulate-like characteristics including the certo form ofthe corallum, wedge-shaped to spine-ke septa and strongly developed dissepiment-lke tabulue, Although unlike any other Barly Carnbrian coral, skeletal characteristics are silmikir to Some michelinnids, which have a tite range front the Late Silurian to the Late Permuan. drrawiporea fromenyis provides further evidence that the time range of the Subclass Tabulata possibly extended to the Barly Cambrian. Key Worbs: drrenvipura fromensis, burly Cambriitn, Moorowie Formation. tabulate-like coral, Flinders Ranges, Saute Australia. Introduction Arrowipora jromensiy gen. et sp. Noy. occurs in the Early Cambrian Mouvrowie Formation in the eastern Flinders Ranges of South Australia in association with Moarawipora chambercasts Fuller & Jenkins 1994, and Flindersipora bowmeni Latuste 199). tt is present in slumped réefal blocks within a megabreecia at a. site close to the disused Moorowie Mine (Fig. 1) described in Fuller & Jenkins (1994). The corals. aré preserved as upright coralla relative to bedding and clearly are in lite position within individual slump blocks. They occur in) association. with both fragmental and engrusting remains of the caleimicrobes Kenulcts Vologdin 1932, Girvanella Nicholson & Etheridge 1878 und Epiphvien Barneniin 1886, and current-deposiled archaeocyaths. The ancient reefal systern may have been established on a marginal fan comprising a coarse breeeia (Savarese eral. 1993). The high energy murine environment was responsible for the influxes of sediment preserved within the framework of the coral colonies. Arrewipyra fromens?s and the two previously described corals from Moorowie have few skeletal characteristics m common, MIDDLE CAMBRIAN WM Lower CAMBRIAN n Preservation SE adtutaiaw st , { The available material, collected many years.ago by =QUORN Ny Mr Brent Bowman, then a technical assistant al the University of Adelaide, shows parts of probably one Fig. 1. Location map showing fossil occurrence near the * Departinemt of Geology and Geophysics, University of Adelaide, South Australia 5005. Moorowie Mine and the distribution of Early and Middle Cambriin outcrops in the Flinders Ranges of South Australia. 76 MARGARET K. FULLER & RICHARD J. F JENKINS Fig. 2. Holotype SAM P34167 (complete specimen), illustrating rectangular shelves extending from a large colony (x1.0), THE EARLY CAMBRIAN TABULATE-LIKE CORAL ARROWIPORA 17 Fig. 3. Holotype SAM P34167 (reverse side of specimen shown in Fip. 2) with shell-like projections across adjacent sediment (x10). is MARGARET K coluny broken trom a large specimen (Pigs 2,3). Duling lite the colony appears to have been repeatedly hut partly covered by centimetre thick layers of Tine sediment which now fill large spaces between lateral expansions of the corallum. Many curallites: were sinothered. allowing only wlimited number to continue their growth, Subsequent corallites grew either inglined or spread faterully above the tenses of sediment. Transverse and oblique sections of small urchaeocyuths lying on their sides relative to hedding are evident ip eavities between extended shelves of the corallum (Fires 2,3}. The geapetal infilling of the urchaeucyaths lurther indicates that (hey were transported into the cavities with the sediment. Culcureaus sediments filling small cavities hetween the vorallites have generally been recrystallized, while the calyces (lopether with larger cavities) wre usually Mlled with very fine sand or silt, Laterally exvended shelves of the specimen SAM P34167 are irregularly reclingular or platy and project over the bioclastic and/or culcarenite matin (Figs 2,3), Coralhites also show unications of being eraded by rapid, energetic influxes of course sand. Caleite-filled fractures appar- endy related to post-diagenerie defornmution of the corillim oceur rarely (Figs 4B,C)_ Reerystallization has affected wll of the colony and sume of (he skeletal structures observed may be artifacts of diagenesis, There are, however, domains within the reerystullized fabric where sume evidence ol the primary structure of the skeleton appears (a be preserved, These relic, rather robust fihrous elements which evidently formed the sclerenchyme (ealcarcous skeleton of corallites), are seen as either lineations across the walls of corallites (in transverse section) {ind/or divergent bundles (in longitudinal section) (Figs 4&.5D). In longitudinal section, upturned spines along some corallite walls (Pig, $C). and spines situated on the upper surtuce of some tabulae (Fig. 4D) are represented by bundled tibres, giving both the wall and tabuiae a bumpy appearance. [n transverse section, Most septa uppear ty terminate in fan-shaped arrays of fibres, or similar arrays arise from. the walls (Fig. SC). The bundled fibres resemble primmlive trabeculae. However, fan-shaped tufts in carbonates often result from diugenesis (Oekentorp 1989). Systematic palacontology Phylum CNIDARLA Class: ANTHOZOA Subclass: 2TABULATA Family: uncertain Genus: Aprowipera gen_ nov, ‘Type speaes: Arrowipera Jronunaix sp, noy, PULLER & RICHARD LP. JENKINS Envriolony For the Arrowie Basin, an Early Cambrian shallow marine busin. extending over much of the area wl the present Flinders Ranges of South Australia. Digencyis Corallum large. massive cerioid, comprisiny polygonal corallites: corallites prismatic and irregularly cylindrical, walls separated by a medial plane, thick. wavy to crenitle. sometines almost straight; tibulie numerous, rarely complete, commonly dissepiment- like tabellaes septa nunieraus at absent, numbering up to 35.in each corallite; where present septa form short wedge- to spine-like projections into the lumens tinea pores whsent, Arrawipara fromensis sp. ny. FIGS 2-5 Etvmalogy Por nearby Lake Frome. DiaENOsIS As for genus, Ape specimens; The specimens described in dis paper ure held at the South Austrulian Museum (SAM), Holotype SAM P34167, a polished skib of a broken part of a corallum and thin sections SAM P34167 1. SAM P34167-2, Puratype SAM P31962-1. SAM P31962-2, counterparts comprising two triangular. large, cut, polished slabs approximately 34 cm normal to hedding and 28 em parallel to bedding. containing either two coralla or more likely the disjunct parts af one large corallum which formed numerous platy shelves, Thin seetion SAM P34168-1. The material wus collected from the Moorowie Formation, near the Moorowie Mine in the eastern Flinders Ranges.( Fig. 1). Description Colony Jarge. more than 24 cm tall and extending laterally Well in-excess of 23 em. In longitudinal section the corallum may broaden upward, or more cormmenty, forms wide shelves extending laterally over the adjacent sediment, Shelves are either irregularly rectangular in shape, with corallites tending to diverge slightly. or are plate-like. Individual shelyes measure up to 70mm high and {30 mm in width (Figs 2,3). The upper surface of the shelves is irregularly horizontal to concave, und culices may extend up to 7 mim above the uppermost tithellae. In transverse section (Figs 5A,B,C), the ceriid corallites. are seen as 5-8 (generally 6) sided polygons, varying belween 10 and l4 mm in diameter, Walls ure relalively thick. varying between 0.1 min and 1.0 mm, and are wavy to almost suuight. The inner Surfaces of the wally are irregular, due ti the msertion THE EARLY CAMBRIAN TABULATE-LIKE CORAL ARROWIPORA 79 Z wy ee Fig. 4. Longitudinal sections of Holotype SAM P34167 A,B. Adjoining sections illustrating general shape of the corallites, tabulae, vertical and basal corallite walls (x2.4). C. The irregular surface of the walls and upper surface of tabulae. Two fractures which post date growth are observed mid-to lower-right of figure, together with the recrystallized fabric within the corallite (x10.6). D, Enlarged section (x2.4) of corallite (lower right Fig. 4B) illustrating tabulae with possible septal spinules on the upper surface. E. Higher magnification (x40) of a corallite section illustrating diverging fibres of a vertical wall, a; and the similar structure of the basal wall, b; and tabulae, c. 80 MARGARE] K, PULLER & RICHARD J, F JENKINS Fig. 5A, Transverse section of Holotype SAM P34167 (x38), B, Transverse section of Paratype SAM P34168 (x45) showing variition in comtlite shape and septa, Tabulae are observed as irregular lines crossing the corallite: the midline of the wall (arrowed) may be seen it some adjoining corallites. C_ Enlarged section (x10,5) of SA illustrating septa. wall irregularities, Widline a und fabulae b, The reerystallized fabric is observed within the corallite. D. Corallite walls (x40) showing the bundles of fibres which cross the wall (irrowed) in seations of the specimen. TH RAKRLY CAMBRIAN TARULATE-LIRE CORAL ARRUB IPRA RI of numerous tabellae and septal spues. In thin sections. aw medal line divides the walls of adjoining eorallites (Figs 5A.C}. In transverse section al low magnifications (up to X 40) straight to slightly diverging fibres crossing the wills between adjacent corallites are Comunotily disruplead by the medial line (Figs 5A,B.C°,D). tu longitudinal section, fibrous elements diverge outward and Gpwards fron the medial line and often protrude inte the lumen giving the walls an irregular appeurance. The walls which truneate parent corallites and form the base of subsequent corallites, are composed of vertical lo slighuly inclined fibres. These partitions arise from the vertical walls and ure usually V-shaped, but may be undulating, horizontal or inclined (Figs 4C DE}, In longitudinal section (Pigs 2.3,4A,B,C), individual conallites are prismatic to irregularly cylindrical und upto 14 min Wide and 47.5 nun long, Corallites vary lithe in diameter and length, prior to the addition of new coralfites (increase). Increase is both lateral ani peripberal intracalicular. parricidal wilhin the established budy of the colony (Figs 2,44,B). ‘Tubolae are numerous, comuionly formed of invommplere. globose and dissepiment-like Labelle, Occasionally some wre Continuous weTOss Very TUTTOW cyrullites, Tabellae may arise from the wall. ot ftom adjacent tabellae, extending jmwand and curving downwards to pest upon other tabellae. They are very thin, yenerally less than 0.06 mm. often wayy wnd rarely strdight. Small projections often occur on Lhe upper surface of tabulae (Fig. 4D). In transverse section, tabellae are seen as wavy and crenate, srising from the walls and dnastomosing with adjacent fabellae (Figs 5A-C). At low magnification, the fibrous structure of fhe tabellag is sinmilar to that of the walls, with some bundles extending to give the srmull projections on the upper surface, In longitudimal section, the fibrous elements are normal to the byse of the tabellac. In transverse sectioyi, septal spines vary fram nuMeryus (about 35) to absent and are often diffieult to distinguish from other irnmegularities on the wall (Figs SA.B.C). Where present they arc short (up to c¢. 1.25 ith), generally equal in length, blunt triangular or spine-like in shape and equidistant trom each other (about 0.25 to 0.5 mm), They are commonly present on some walls while absent on others within a single corallitg, Septal spines appear to be continuations of bundles ot fibres of fhe fibrous wall, usually rerniinating us, or being seen as fan-shaped tults (sec above - Preservation), Lo longitudinal section, the generally upturned septal spines are observed lo oocasionally form short verlical rows on corallite: walls, Discussion 4. fromensis is unlike the bw previously desorbed Harly Cambrian carals from the same location, Flindersipora bowmarni Laluste 1991 (e.g. Lafuste of al. YA) and Movrawipara Chamberenyiv Puller & Jenkins 1994, 4. fromensis is distinguished from F bewmani by the size and general form of the colony, (he position and shape of tabulae and septa, as well ag the mode of inerease. In FE bonwwncn/, tabulae are mostly complete and concave proximally, there are 6-16 strongly devoloped slightly curved septa, the edges of which bear very short blunt spines; the walls are very short Segments between the sepra. Increase is by longitudinal fissien The main differences between 4. fremmensiy and M, chambereisis are in the size und form of the colanies, the size and shape of the corallites and the arrangement and shape of tabulae, Although bow are cerioid im colonial form. the former is much larger amd usually has parallel corallites, while those in M. charburinsix are generally divergent. Corallites are prisinatiy to cylindrical and up to 14 mim in diameter and 47.5 nim in length in 4. fromensis. bul much srualler tup to 5 mim in diameter and {9.5 mm in length) ine tuberond (yy integularly cylindrical in M. ckamberensis. "The presence or absence ul septal spines /s common ta hott corals: When present [hey are about the sane size and shape Tshulae differ greatly, being incomplete, globose and dissepimentlike (abellue) in) df. fremmensyis and coniplete, undulating and horizontal to concave upward in M, chamberensis, Although the microstructure bas not been studied at high magnification, there are sone sumilurities between lhe aboye corals at low mapnification. These toclude the purillel Abrus elements of the selerenchyme evident in trunsverse section, and the parallel to diverging fibrous elements in longitudinal section, Fan-like arrays of hres. are not present in M. chaniberensis, A medial Tine within walls of adjacent corallite occurs in both corals. Medial lines in the walls are common in iabulate corals. and represent the external epitheea (Hill JOT). A. froonensis 1s unlike ony of the previously deserined Cambrian corals suggested by Serutton (1979) 1o have tabulate affinities, but does have skeletal characteristics in common with some of the Late Silurian to Lite Permian michelininds, ‘The diagnestic characteristics for the genus Michelinia De Koninek [841 inelace thin ty moderately thick walls with & medial suture, short septal trabeculae, tbulie incomplete and globose sometimes with septal spinules on lhe upper surface. and large mural pores (Hil) 1981). The walls und fibulwe ure similar to those seen in 4. fromens/y hot ihe presen| taxon Jacks mural pores. 42 MARGARET K, PULLER & RICHARD 1 F IENKINS Micheline expanses White 1883 | Tabelleepliyiiun peculiare Sturm \948) (Stumm M48) from the Early Carboniferous of Arizona, is sintilar to A. frorerists with respect to the form of the colony, the size and shape of corallites and the arrangement of jabellae. Corallites are up ta [S mm in diameter in the larmer ane enim inthe latter, Corullites are wso of a-similar shape. being generally 4, 5 or 6 sided, but difter by the lack of septa in ML expanse. A most noticeable simukarity between the two is the placement, size and shape of the tabellae, They are incomplete and globose and ure arranged in similar manner in both taxa, arising from either the walls or adjacent tabellae. The tabellae in A. fromensis appear to be less. glohose, spaced slightly tirther apart, and havea more irregular and wavy surface. Aluiough 4. /ramensiy most closely. resembles some of the mivhelinids, beeatise of the long time separation between them (120 million years) itis highly unlikely that they ure relisted and more probable their skeletal similarities result from convergent evolution Conclusions The three described corals from the Moorowie Formation, A. framensis, Mo chamberensiy and F, Hewnani’, are very different in fori and arningement of the skeleton, The diverse nature of the corals from his ancient reefal environment indicates that during the Early Cambrisn, variubility in polyp form and skeletal morphology was well established. The genus Lichenurie has been recognized as the carhiest tibulate com, witha lime range from the base Of the Barly Ordovictan tothe eurly Late Ordovicuin (Serutton 1979, 1984. J992) Hull INST), Tt has beer described as primitive, cerioid, of simple morphology. usepture, but with tabulae and rare mural pores (Bassler 1950, Flower 96); MeLeod 1979, Serutton 984: Liah 1984). Although A. fromensis lacks mural pores, it hits asimilar skeletal structure (0 some micheliniids which post-datte Lichenaria. Most of the skeletal aspects ol A fromensis are characteristic of Palueozoie tabulate corals. These are (1) the cerioid form of the colony: (2) walls separated] by a medial line reflecting individual corallites (Scruton 1987); (3) the spine-like lo wedge-shaped seplt occasionally situated in longitudinal rows (Hill 1981); (4) individual corallites which Spread above the pockets of sediment within the colony, this. habit being usual tor enidarians following influxes of sediment (Serutton (979): (5) lateral increwse common, with peripheral intracalicular increase being described in some Favositidae (Hill W981). Although tabulae dre incomplete and dissepn ment-like, they are consistently and strongly developed both within individual and between adjacent vorallies in AL fromensiy. A. fromensis has anthozoan structural characteristics, most of which are evident in tabulate corals. It should thercfore prohably be included in the known group af tubulates, thus extending the tine range of this group to the late Early Canibrian, Acknowledgments The authors gratefully acknowledge Brent Bowman und John Hart who collected the studied material, We alsa thank Gino Snider for his phalographie work. References Bassinet. ROS (1950) Faunal fists and description of Palucovore Corals Mem. Geol, Sa Anr a4, 1Al5, Prowek, R.A (1961) Marleya and relitled colonial corals Nee Mec Inst. of Ter Mem. 7, 1-97, PoieR ML RK, & JesKiIne, RoE BE (00d) Moeprny pera chemberenisis. a comal tron the Burky Cambrian Moorawie Permation, Flinders Ranges, South Australia Trans, A Sea S. luyé W864), 227-235. Hi), D (ISAT) Rugosa and Tabulata pp, 1-762, fe Teicher. C (Ed) “Treatise of Inyertebrite Palaeontolowy. Part F, Coglenteritta, Supplement 1 (Geol. Soe Am. & Uno Kansits, Boulder, Coloradey- Larusie, J, Drogisxt, Bo Gaspin, Ao, & Gravesruck, 1, (991) The oldest tabulite goral and the associated Archucoeyalbit, Lower Cambrian. Flinders Runes, South Ausrnilta, Geublos 24, 697-718 Lape, R. 8, (i984) Liehendrty WINCHELL & SCHUCHERT, 1895! Lamotttia RAYMOND, 1923, and the carly history of the tabulate corals. IV Liternational Svinpasium on Fossil Cidaria Puluewmniovr am, Sd, 154-163 Oerkesmore, K (1989) Djagenesis in corals: syntirvl cements as evidence for post-mortem skeletal thickenings V Internavooual Symposium on Fossil Cnidaria. Massif Cnideiria 5, V9-177. Mereap. J BD, 1979) A Lower Ordewiciin (Cunadiin livhenarid Coral trom the Ozark uplift area, 2 Paleont, 33, §05-5U6. Savarisi, M_, Meine, JF, SoKave, dE & Bucks IN, I (1983) Paleobiological and paleoenvironmental context Of coral bearing Eyrly Cambrian reefs: implitations for Phanerozuie reef devclopment. Geology 24, 917-920. Scrotton, © T. (1979) Barty fossil Cnidarians pp. (6i-207 In House, M- R. (Bd ) “The Origin of major invertebrate groups” (Academic Press, London & New York). —— WOK) Origen and carly evelution of Tabulate Comets WV International Symposium on Possil Cnidaria Palavaniiolagr Ane Sd, WO-1R. —— (1992) Findersipora boven: Lafuste. ane the early evojulion of the tabulate comals: Fossi/ Cnidaria & Parifera 21, 29-33, Srunm, B. R (1946) Upper Devonian tetracorals trom the Manin Limestone, Felaennir 22, 40-47 CHARACTER AND INTERPRETATION OF THE REGOLITH EXPOSED AT POINT DRUMMOND, WEST COAST OF EYRE PENINSULA, SOUTH AUSTRALIA By E. MOLINA BALLESTEROS*, E. M. CAMPBELLYf, J. A. BOURNET & C. R. TWIDALET Summary Molina Ballesteros, E., Campbell, E. M., Bourne, J. A. & Twidale, C. R. (1995) Character and interpretation of the regolith exposed at Point Drummond, west coast of Eyre Peninsula, South Australia. Trans. R. Soc. 8. Aust. 119(2), 83-88, 31 May, 1995. The weathering mantle developed on granodiorite at Point Drummond, Eyre Peninsula, South Australia is examined using thin section and XRD analyses. Stages in the alteration of the granodiorite can be deduced by examination of the zonation of the regolith; release of oxides and hydroxides from the parent rock; removal of iron and kaolinisation; new concentrations of haematite in micropores; development of nodular structure and renewed removal of oxides and hydroxides. The possible age relationships of this profile with laterites and Plio-Pleistocene ferricretes from other South Australian sites are discussed. The age of weathering is uncertain but it predates the calcarenite (? Pleistocene) and is probably post Permian, with Plio- Pleistocene the most likely. Key Words: regolith, ferruginisation, Point Drummond. Transactions af the Reval Savielx af S. Aust, W995), WY) Ba-RR CHARACTER AND INTERPRETATION OF THE REGOLITH EXPOSED AT POINT DRUMMOND, WEST COAST OF EYRE PENINSULA, SOUTH AUSTRALIA by BE. MOLINA BALLESTEROS* E, M. CAMPBELL, J. A. BOURNEY & C. R. TWIDALEF Summary Mouna Bariisteros. E.. CAMpRrui. Bo M.. Bourne, JI A, & Twibsuby CR. (995) Character and interpretation of the regolith exposed ai Point Druinmond, west coast oF Fyre Peninsula, South Australia. Trans; Ro Noe. S. Aust, W9(2), 83-88, 31 Muy, 1995, The weathering ante developed on granodiorite at Point Drummond, Eyre Peninsula. South Australia is examined using thin section and XRD analyses, Stages in the alteration of the granodierite cam be deduced by examimation of the zonation of the regolith; release of oxides and hydroxides from the parent rock; remoyal of iron and kaolinisation; new concentrations of huemutite in micropores: development of nodular structure and renewed removal of oxides and hydroxides. The possible age relationships of this profile with Laterttes and. Plio- Pleistocene ferricretes from other South Austrahan sites are discussed, The age of weathering is uncertain but i predates the calearenité (2 Pleistocene) and is probably post Permian. with Plio-Pleistocene the most fikely. Kiey Worps: regolith. ferrivinisation, Point Drummond. Introduction The west Coast of Eyre Peninsula is characterised by high cliffs eroded in dune calearenite (also known as ueoliuniles see e.g. Crocker 1946) of Middle and Late Pleistocene age (Wilson 199] !). The calcarenite rests unconlormably on Precambrian rocks, mostly igneous and metamorphic. with granite and gneiss prominent, but including sandstone and conglomerie neur Talia. The uncontormity is uneven and the base of the calearenite commonly extends below sea-level, Elsewhere. the Precambrian basement is exposed m rather irregular shore plattorms and in the lower sections of the cliffs which. however, are composed mainly of the culcarenite, At several sites remnants of the pre-calcarenite regolith are developed on the Precambrian basement. One of the best exposures of the regolith, in terms of thickness, completeness and luleral extent, occurs at Point Drummond, a westerly projecting promontory located on the west coast of southern Eyre Peninsula, some 80 km north-west of Port Linvoln (Figs | & 2). The purpose of this short paper ts to describe the mineralogical variations between horizons within the regolith, and to discuss their genetic nnplieations, The site is on the southern portion of Point Drummand, * Department of Geology. University of Salamanca, 47008 Sulumarica, Spain, + Department of Geology and Geophysics, Unversity of Adelaide, South Australia, S005 1! Witsow, ©. C. (1991) Geology of the Quaternary Bridgewater Formation of southwest and central South Australia Ph.D, thesis, University of Adelaide, Adelaide (Unpubl). Two profiles. one from a south-projecting peninsula and one trom the cliff adjacent lo the access steps several hundred metres to the north, were examined. The samples were selected on the basis of colour and textural variations, The profiles are so similar both in appearance and upon analysis that they can be treated as one, Pont Augusla / Paint Brown . Streaky Bay EYRE PENINSULA Paint babhart Cowell Point N Drummond JS AUSTRALIA Port Lincoln TSA ADEE 0 bie) 100 Kms Uv Fig. 1. Locution Map. Description of the profile The profile, approximately 18 m thick, was sub- divided from the base to the top into five horizontal zones on the basis of their colour and texture (Pig. 3). Mineralogy and (exture have been studied in thin section (using. samples impregnated with a thinned aruldite to prevent any disturbance of the original structures) and by XRD analysis of bulk samples. na }, MOLINA BALLESTEROS. E M CAMPBELL. J. A. BOURNE & C. R. TWIDALE ee . os - Fig. 24, General view of Point Drummond, Eyre Peninsula, South Australia. 2B. The weathering profile sumple site, Point Drummond peninsula, showing the coastal plaviorm developed in granodiorite, the goethite-rich zone at the base of the cliff-and above this the white kaolinised zone, the haematite zone and the calcarenite remnant ut the surface REGOLITH AT POINT DRUMMOND RS Zone | The parent rock is a gneiss of granodtorilic composition consisting mainly of quartz, plagioclase, biotite and muscovite with some orthoclase. Secondary minerals resultiag from hydrothermal alteration of the rock prior to weathering mclude sericite, chlorite (from biotite), epidote-zoisite and calcite (from plagioclase - Sample |, Fig. 3, Munsell Rack Colour N&. white to. N3, dark gray - dry. The colours of other samples are Munsell Soil Colours - 1994). The granodiorite is intruded by amphibolitic and quartzitic veins. Schistuse shear zones are also present. These various rocks are all members of the Sleaford Complex, dated al 2,700-2,300 Ma and thus of late Archaean or Palaeo- proterozoic age (Flinter al, 1984; Parker er.a/, 1985) though the shear zones may resull from the Kimban Orogeny (~1700 Ma - Thomson 1969). A 25 a c 2 iss] 5 20> oO o = 10 — Ee a= 9 a2) 2 ) B + s F’s Old fissure \ Stages in the weathering of the gneiss can be traced by examination of the zonation of the regolith or weathered mantle, assuming that the weathering Front, or Jower limit of weathering, has descended into the rock mass from the surface. Hence, in these terms, the mitial stages of weathering are represented by the zone immediately above the weathering front and the most advanced by the near surface horizon. The coastal platform eroded in granodiorte and located between high and low tide levels, is. irregular with many blocky and bouldery rises and intervening clefis, Many of the outcrops are superficially altered, wilh rinds of ferruginoys oxides. and hydroxides developed at the margins of blocks, boulders and other exposures. The rinds are also found bordering fissures (Sample 2, 7.5YR 7/6, reddish yellow to 7.5YR 7/3, pink). In this zone the thickness of the rinds increases up the profile, but nowhere exceeds 5 cm. sainjonijs Mau jo Juawdojecsq sainjonis pjo By yO UOI}eEAIasUOD ¢ New void Fig. 3. Schematic diagram of the weathering profile on the peninsula, Point Drummond, A. Profile. -V, Zones of weathering (see teat for explanation); 1-11, sample numbers (sample 5 similar to 6). B. Principle processes: a, Unweathered parent rock. b. First stage of weathering (goethite-rich zone). C. Kaolinisation. d. Formation of new structures and conventration of haematite, @, Removal of iron and plasime separation. £. Development of peds and nodules. tia) LC. MOLINA BALLESTEROS, E M. CAMPBELL, J. A. BOURNE & C. R. TWIDALE The first stage of weathering discernible is the development of the rinds around the corestones. In these the epidote-zoisite minerals are dissolved and the alteration of plagioclase is manifested by the ‘ppeurance of zones of rundomly orjented clays. The clay minerals are prelerentidlly developed along fissures presumably beenuse the latter allow penetration of water. Biotite changes colour Irom green ty hrownish-yellow, reflecting a release of iron which, as goethite, is concentrated in cracks and lissures, Zone i Here. though the original rock structures are everywhere distinguishable. some of the blocks and boulders are entirely discoloured and the rinds ure Uneker than those in Zone L (Samples 3 & 4, IWYR 8/4, very pale brown to (OYR 7/6, yellow) with a white and (N8) developed on the outside, and a sirong brown (7.5YR 5/6) interior, Again the rind increases in thickness up the profile und, as in Zone f, ss also found along purtings. The contact between the weathered rind and the interior of the corestones low in the profile is sharp but is more diffuse at higher levels. ‘The rinds are pale in colour and voids are apparent in thin section. Plapioclase is progressively reduced higher in the profile and essentially isotropic clays, beheved to be mainly kaolinite, become doyniient. With the XRD method used, it is diffigult to dilterentiate any other polymorphs of kaolin. Resistarces such as quartz and muscovite are present, Fone Lit Almost the enure rock in this zone 1s white (Sample 6 N&) but most of the original textures and structures ure preserved and remnants of the yellow and red iron oxWes occur as spots in the upper pant of the zone At the top of Zone IH loss of material has led to the formation of voids. The weathering plasma (in the sense of Nahon 1991, p. 63), derived from weathering Of the parent materials, begins to appear anisotropic, especially im areas close to voids [the vosepic plasma separation ot Brewer (1964. 1976)]. Some, thuogh not all, oxide concentrations are related to voids. Zone IV The yellow and red spots present in Zone U1) here iierge to give a mixture of Oxides and hydroxides of iron in differing degrees of dehydration and erystallisation (Sample 7, SYR 4/4. reddish brown lo TSVR 6/8, reddish yellow io JOR 4/3, weak red; Sample & IOYR 8/8 yellow to IOR 4/3, weak red; Sumple 9, IOR 4/3. weak ted to IOR 5/6 ted). The zone is up to four metres thick. None of the original structures survives. XRD analysis shows that haematite is dominant in the weak red patches, kaolinite having been removed, On the other hand, where there is no iron oxide or hydroxide, the kaolinite altéroplasina is well preserved Muscovite remains. alber! weathered to varying degrees and quartz. is corroded and in some instances clearly disapgregated, At the lop of Zone LV voids are common, This zone is Similar to the “mottled clay™ horizon from a laterite profile deseribed by Nahon (1987), of which alumina (not analysed in this study) is a typieal component Zone V The lower part of this zone ts characterised by irtegulir Ul-defined fissures in the mottled clay, Also, anew structure occurs in the form of polyhedral peds some centimetres across, They bevorme stoaller and more rounded upwards, where they take the form of soft nodules 0,5-1.0 cr in diameter, reddish vellow in colour but with red oxide conceatrations 1m the interior (Sample !0, 75 YR 6/6, reddish yellow to 1OR 4/6, sed) (Fig, 4). The partings which define the polyhedral! pecs Pig. 4. Thnn section (crossed polars, plain light. x25) of upper partof Zone Vo Kaolinite plasma surrounds the ferruginous nodule (itrk ured). The light-coloured material in both the nodule wnbthe surrounding plasma is quan. are preferential zones of leaching. Removal of oxides and hydroxides has resulted in zones of concentration of skeletal grains (mainly quartz) fram the purent matenial, and in the appearance of 4 kaolinitic chiy plasma (a pedoplasma in the sense of Boulet 1974), It grades trom asepie, where oxyhydroxides are abundant, to skelsepie, vosepic and even masepic where the oxyhydroxides have disappeared (Brewer 64, 1976). The development of the nodules is a centripetal process involving the removal of oxides ond hydroxides from the margins of the peds and their concentmtion in the aodules, the redistribution of clays and the concentration of skeletal grains in the leached Zones. Both the promontory and the cliff profiles are over lain by a calcareous crust (calerete - Sample }t, 2.5Y 7/2, hyeht gray), located at the base of the calcurenite but developed on the weathered material and including nodules like those found ta Zone V CPi 4) REGOLITH AY POINT DRUMMOND gy Interpretation Various stages in the weathering of the granmtivrete pre eviderd; \. ‘The penetration of metearig walters trom. the surface und release of vaides and hydroxides fron minerals in the parent rock, especially biotite, ts the initial process in evidence The iron oxides are concentrated in fissures at Lhe weathering front, at the margins of blocks and houlders waned adjacent ty jount purtings. The hydrothermally altered minerals are dissolved and the plagioclases are weathered Lo clays. 2. Removal of ifon aad kaolinisation are represented by the appearance of white rinds. These processes are usually achieved in acid reducing solutions, which wppear to have leached most of the calcium, iron and sodium and some of the silica and produced the newly formed mass of isotropic to insepic materials. (see Brewer 64, p, 309), mamly kaolinite, Inside this isotropic plasmic material are resistates such ws quarks and muscevite {skeleton grains, of Brewer (1964, 1976)), On the whole, however, kaolinite is dominant, 3. Ferruginigation results from (he progressive development of & kaollnitic plasma, and a new porasity plus the destruction of the original features ancl structures. Micropores, especially, become the sites of new concentrations. of oxides (weak red spots), particulurly haematite (Pig. 4), due to the decreased mobility of the solutions in these pores (Dichter ef al. 1983; Tardy & Nahon 1985), This.is the origin of the ‘mottled clay horizon’ typical of protiles develaped under seusonally wet and dry climates, 4. The development of nodular structure is related to the removal of oxides and hydroxides by solutions probably emanating from an overlying soil, A new k#olinitic plasma is developed as the oxyhydrox ides are remoyed. The mobility of the materials 1s governed by the amount of oxyhydroxides: the less oxyhydroxide the more plasmic movernent and hence better ree organisation of the soil mass. 5, The removal of oxides and hydroxides requires nod salutions (i.e; those which are poor in carbemuales). so thit the processes described in paragraphs 1-3 inclusive predate the development of the calerete and the deposition of the dune culearenite The age of the profile, and the events ta whicl tt relales, are mainly problematic. 1is clearly younger than the Proterozoic rocks on which itis developed, and predates the calcarenite which, according to Wilson (1991). is Middle and Late Pleistocene i age (maxi- 2 Hossre.o, P. (1926) The pcology of portions of the Counties of Livht, Epre, Sturt and Adelaide, MSc thst, Univwmiry ob Adehide, Adeline (Unpubl), mum-¢. 700,000 years). But allocating at to an hiatus bl sore 1500-2000 Ma is neither precise nor inform ative, The extent of the hiatus can be reduced if two general arguments are accepted, First. although regoliths have survived the passape of ice shivets (see ey, Fogelman 1985), the profile, which has apparent equivalents al several points along the west coast of Fyre Peninsula (e.g. Point Brown, Point Labatt, Talia), is anlikely to have survived the Early Permian glaciation, which evidently affected most of the present state ol South Austratia (e.g, Ludbrook 1969), and subsequent erosion: for the reyolith has readily been eroded by marine agencies and by gullying, In these terms the regolith under debate is less than 250 Ma old. Second, the hiatus is farther reduced if il is conceded thin the revohith is most likely immediately to predate the cover material, that is the calcaremte, This last suggestion assumes that even if the development of the regolith began long before the deposition of the dune lunestone. it would have continued to evolve (see ee. McFarlane W986; Bourman 1993) up to (and even beyond) being covered. In these terms the young date for the regolith is of the order of 700,000 years, though, because ibmust have develgped over a long period, it ought to be assigned an age range ane could reasonably he labelled Plio-Pletsiocene. Broader considerations support this suggestion. First, with what other regoliths might the Point Druinniond profile be retites!? Perruginous regoliths are known from various parts of South Australia (e.g. Hossteld 1926-5 Northcote 146. Miles 1952; Horwitz & Daily 1958: Campana 1958; Cilaessner & Wade 1958; Horwatz 1960, Wopfner 1967; Daily et al. 1974; Twidale eral, 1976; Wright 1985; Milnes eral. 1985; Bourmanh er af. 87), They hive been variously defined and interpreted (see e.g. Bourman 1993 - but see also McFarlane c.g. 1986: Firman 1994), Some, characterised by » sandy Or silty Achoraton, a massive. commonly pisolific, ferrnginous horizon averlying a thick bleached zone, have been labelled lanerite, Others, consisting af cither a [erruginous crust aloe, of a onust resting ona thin bleached horizan, huye been lermed ferricretes (¢ 2, Lumplugh 1902; ‘Twidale 1976, p 196-197), The Point Driuniond profile does. not sit eusily with either of these, bul is perlaps closer to the ferricrete than to the Ialerile. particularly af it is considered logether with other stratigrapically comparable regoliths such ap that exposed at Point Brown. Ferricretes in southern Soulh Australia have been dated by various means. but oo Yorke Peninsula (Horwitz & Daily 1958) local stratigraphy indicates a Pliucene age. Equally, occurrences in the interior ol Eyre Peninsula have, on stithgtaphic grounds, also been attributed to the Teniary, some being considered Eocene but others clearly Pliocene or post-Pliocene (Rankin & Flint 199); Flint & Rankin 1991; Flint 1992), Given the Middle Late Pleistocene age of the overlying 4H bh MOLINA BALLESTEROS, B, M. CAMPBELL. J. A. BOURNE & © R. TWIDALE caleurenite. a later rather than an earlier Tertiary age scems appropriate for the Point Drummond exposure, und on balance a Phi. Pleistocene attribution is in keeping with the available evidence, Acknowledgments The authors thank Dr MJ. McFarlane for comments and suggestions during fleld work and Mr M. J. Wright und) Iwo referees for constructive suggestions on ar carly draft. Professor Molind’s-work in the Universily of Adelaide in the period August-December 1992 wats mide possible hy a grant trom the Direecion General de Investigacion Cientitica y ‘Teentea (DGICY'T). 92-176, Ministerio de Educacion y Ciencia, Spain References Boulit, R. (974) Toposcquences de sols tropicatin en Huute-Volti, Equilibre er désequilibre p&dobiochimatique. Mem, Office Rewh. Set. Teehn. Outre Mer 88, BouRMan, Ro P. (1993) Perennial problems in the study af laterite: A review, Anyt J Earth Sci, 40, 387-401 Mines. AJ Ro & Oapre, IM, (1987) Investigations of ferricretes und related ferruginous materials tn parts of southern and eastern Austtilia, 2. Genial. ME Suppl-Bi, 64, 1-24. BROWER, R_ (1964) “Pabpe und Mineral Analysis of Sails” (Wiley, New York). — — (976) Fabric and Minera) Analysis of Soils’ (Krieger New York) CAMPANA, Bo UY5S) The MU Lofty-Olary region and Kangaroo [sland pp. 3-27 In Cilaessner, MF & Parkin, 1 W (Fils) “Geology of South Australia” (Cambridge Universily Press, London), Cock, Re 1, (1946) Post-Mincene climatic and geologic History and its significance in relation to rhe genesis of the mayor soil types of Soulh Australia. CS LR Bull 193, DAY B. Twinaik, Co RO& MILKes, A, Ru (1974) The age ofthe literivized surfiee on Kangaroo Island anel ad jaccst areas of South Australia, Geol. See. dash Jets 21, 387-992 Dibiek, B, Free. B., Nation, BD & Taroy, Yo (1983) Pel kaolinites, Al-goethites. Al-hematites in wopical lernereles, Mom. Se. Géol. Strashoure Wh, 35-44. HM AN EB. (1999) Paleosols tn hulerite and silerete profiles: Beidence from the South Bast Margin of the Australien Prevambriun Shreld, Marth Sei Rev. 36, 149-179. bitnt, ROB. 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(i976) Ape and origin ab paleosurtaces on Eyre Peninsula und in the southern Gawler Ranges, South Australia, Z, Geomeorpl, 20, 28-55. WorrserR, He (1967) Some observations on Caingroir fundsurfaces in the Officer Basin, Geel Suri X tvse Ouarn Geol. Notes 23, 3-8, Wrist. Mod. (19835) Sails pp 77-87 Me Twidale ©. Tyler Mf & Davies. M. (Eds) “Nitural History al Bye Peninsula” (Royal Suctety of S. Aust. Adelnitey A REVISION OF THE GENUS TIKUSNEMA (NEMATODA: ACUARIOIDEA) WITH THE DESCRIPTION OF A NEW SPECIES FROM THE FALSE WATER-RAT XEROMYS MYOIDES FROM QUEENSLAND By LESLEY R. SMALES* Summary Smales, L. R. (1995) A revision of the genus Tikusnema (Nematoda: Acuarioidea) with the description of a new species from the false water-rat, Xeromys myoides from Queensland. Trans. R. Soc. S. Aust. 119(2), 89-94, 31 May, 1995. The genus Tikusnema Hasegawa, Shiraishi & Rochman, 1992 is redescribed. The species Molinacuaria indonesiensis Gibbons, Cranshaw & Rumpus, 1992 was found to be synonymous with Tikusnema javaense Hasegawa, Shiraishi & Rochman, 1992, the two species having been described almost simultaneously from the rice field rat, Rattus argentiventer. A new species of Tikusnema from the false water-rat Xeromys myoides is described. Tikusnema vandycki sp. noy. can be distinguished from T. javaense by the size of the adult male and female, the shape of the cuticular leaves on the pseudolabia, the length of the male tail and spicules, the length of the female tail and size of eggs. The implications of the presence of acuariid nematodes, normally found in birds, in a range of small mammalian hosts, are discussed. The significance of the presence of Tikusnema in Indonesian and Australian hosts cannot be determined until its presence or absence on the island of New Guinea is confirmed. Key Words: Tikusnema, Nematoda, Acuarioidea, Xeromys myoides, false water-rat, mammalian hosts. Transietions bf re Hewal Soler af A Maye (19057, TEC) 89.94 A REVISION OF THE GENUS TIKUSNEMA (NEMATODA: ACUARIOIDEA) WITH THE DESCRIPTION OF A NEW SPECIES FROM THE FALSE WATER-RAT XEROMYS MYOIDES FROM QUEENSLAND. by LESLEY R. SMALES* Summary Svares. dh. R995) A revision of the genus 7iaisnema (Nematodu: Acuarioided) witht the description of a new species Prom the false water-eat. Neronvs vryaides trom Queensland. Trans. Re Sue. &. Aust. E92), 894A, 41 May, t995, Nie genus Twsene Hasevawa. Shiraishi & Roehman, 192 is redeseribed, The species Melineenarta iIndonesiensis Gibbons, Crawshaw & Rumpus, 1992 was found lo be synunyrnous willl Tikisndvte Jd vdense Hasta, Shiraishi & Rochman, 1992, the two species having been desorbed almost simultuncously fremy the fiw (eld ray, Aaais urveriventen A new species of Tkusrena trom the false water-nt Aeron mveides ts deseriled, Tikusnema yandyeki sp. noy, can be distiawuished From 7 jeveense by the size of the adult male and lemale, he shape of the cUlicWar leaves on tie psetidolabia, the length of the male tail and spicules, the leagih of the jumule tailand size of egys. ‘The implieauons of the presence of acuariid nematodes, aormally found in birds, ina range of small mammalian hosts, are discussed, The significance of the presence of JrAwsnenta in Indonesian vind Austrian hosts cannot be determined until ts presence or abserice on the island of New Guinea 1s comiirmied. Key Worps: 7ikusnenta, Nematoda, Acudriodea, Xeraumyys nevetdes, false water-ral, mammalian hosts Introduction The lalse water-rat Xeremius myoides Thomas. 1889 isa small dark grey semi-aquatic rat whose preferred habitul 15 shallow coastal wetlands, such as swarnps, mangroves, forests, lagoons. or sedged lakes (Van Dyck 1994). ‘They are currently known from only six sites in north-centoul and north-eastern Australia. Their current Conservation status is valnerable und likely to progress lo endangered because of human proclivity to drain and develop swamps (Van Dyck 1992) They forage on the mud flats for food items including aquatic invertebrates, such as crabs, mud-lobsters, niussels. midrine pulmonates and polyclads (Van Dyek 1994). Nematodes dissected from specimens of X, yrvoides, collected by staff of the Queensland Museum were found to be species belonging to the Acuarioidea. The genus Tikusteme was erected for specimens from Rarus arveniventer (Robinson & Kloss, 1916), the rice field rat from West Java by Hasegawa ev al. (1992). Almost simultancously anew species of Molinaeuaria was described, also fram Ro argennivenrer from Java, by Gibbons er al, (1992). Comparison of material fram X. niyeides with type specimens of both species deseribed trom Autrity argentiventer suggest that the nematodes from X. wiveides are new species of Tikusheia while all the * Biology Department, Central Queensland University. Rockhampton 4702 Queensland, material from R, urgentiyenter ws cop-specific. Molinaeuaria indonesiensis therefore falls ay a synonym of Fikusnenta javaense. Materials and Methods Six lalse water-rats, Neronny anvoides. were collected from Myora Swamp, Stradbroke Island, Queensland during 1992, Fuecal pellets from two hosts were examined. The ulimentary tracts of the tour other hosts were dissected for helminth parasites alter the bodies had been fixed whole in 10% formalin. The nematodes so collected were cleared in lactophenol for microscopic examination. Figures were drawn with the aid of a drawing tube. Measurements, of 10 specimens in jem unless Otherwise slated. with the range followed by the mean. were made with the aid of an ocular micrometer, drawing tube und measuring wheel. Specimens of Molinacugria indonestensis Gibbons, Crawshaw & Rumpus, 1992 and Tikusnema javacnse Haseyawa. Shiraishi & Rochman, 1992 from Rattus areentiventer were also examined for comparison, The terminology used for morphological features in the deseriptions is that of Hird and Bird (1991) and the taxonomic system of Anderson (1992) is followed. Abbreviations. aré: Queensland Museum QM: Australian Helminthological Collection South Australian Museum SAM: United States National Museum Helminihologicul Collection USNM: International Institute for Parasitology 1" 90 LESLEY R Systematics Order Spirurida Suborder Spirarini Supertumily AcuarioiWed Family Acvariidae Sub family Seuratiinae Genus Tikusnenia Hasegawa. Shiraishi & Rochman, 1992 Type Species Tikusniemea javiense Hasegiwa, Shiraishi & Rocha, 1997, Tikusnema Hasegawa, Shiraishi & Rochman, 1942, Revived veneric diagnosis Cephalic end inflated and set off from body by consuicuon, Oral opening laterally compressed. Poeudolubta large, triangular in latery] view, each with two cephalic papillae and an amphid, Pseudolabia attached 10 each other apically. separated dorsally anc ventrally by cordoas and indented deeply at level of wimerior extremity of cordons. Posterior end of each pscudolabium forms two cuticular leaves euch sub- divided into teeth, Cordons small, net extending postenorly. Buccal cupsule long, culicular wall thick- nol striated. Pharynx divided into anterior muscular and posterior glandular portions. Deirids small, bicuspid. A pair of cuticular ornamentations: present laterally, posterior to deirids, Parasitic in the stomach or intestine of rodents. Tikusnema javaense Hasegawa, Shiraishi & Rachman, 1992, Synonym Molinacvaria’ indonesientsis Gibbons. Crawshaw & Rumpus, 1992; pp. [75-181 Material exaniined From R. atgenrivener: \o allotype Tikusnema javaense USNM 822235 Pusakanagata, West Java, Indonesia: 1 O° paratype IP BIOSSB from Sukamandi, West Java, Indonesia. Description As in Hasegawa ef al, (1992), From the combined Measurements of both Hasegawa ef al, (1992) and Gibbons ef al, (1992) the dimensions become as tollows: Male: length 9-2) mm, width 277-440, Buccal capsule 359-490 Jong, muscular portion of pharynx 410-560, glandolar portion 1130-1980 long, Deirids 296-440, nerve ring 450-560, excretory pore 525-830 from anterior end, Right spicule 190-210, left spicule 491-570 long; tail 556-990 long, Female: length 11.0-24.5 mm, width 293-510. Buccal capsule 330-490 long, muscular portion of pharynx 402-630, glandular portion 860-2030 long, Deirids 273-430, nerve ring 410-630, excretory pore 502-870 from anterior end, Vulya 5.02-12,21 mm from anterior end, Tail 230-520 long. Egys 28-31 by 38-44 SMALES Remarks Tikusnema javaense and M. indonesiensis were deseribed almost Simultsnewusly by Hasegawa et al (1992) and Gibbons e7 al. (1992). the descriptions appearing in different journals, Both cescriptions referred tw material collected on the island of Java from Rattus argentiverser, A caretul examination of (he deseriptions given by each group of authors, together with a comparison of the material they exawiined, has revealed that they gre of the same species. Any differences In measurements benween the two sets of materiul relate only to the fact thar the specimens described by Hasegawa ez.a/. (1992) were smaller thin those described by Gibbons eral, (1992). The lemales described by Gibbons er a/. (1992) for example ure lurger, gravid females containing embryonated epys while the smaller females described by Husegawa et af. ()992) contain unfertilized epgs. Hasegawa er a/. (1992) describe and figwee a pair of cuticular ornamentatiuns much larger then the deirids in the adult worms and even more prominent in the 4th sutge larva. Gibbons et al. (1992) did not mention this feature in their description of their more mature worms. Therefore it appears that the cuticular OTmamentavions may he 4 more prominent leature ol juvenile than mature worms, This would account for their upparent absenee in the specimens examined by Gibbons et al. (1992). The interpretation of the cephalic ends of the specimens, in particular the origins of the cuuicular leaves. by Hasegawa er al, (1992) appears consistent with both sets of muterial. Ay discussed in Hasevawa et al, (1992) the cuticular leaves of 7ikusnema originate directly tron’ the pseudolabia without separating furrows. Tikusnema also has small cordons nol extending posteriorly. By contrast the genus Molinacuaria, although characterized by the absence of pseudolabia (Wong & Lankaster 1985), does have Brooves. located immediately anterior to ptilina, Molinacuaria can be further differentiated from Tikusnema by a lack of cordons, Moalinacuaria indonesiensis lherelore falls ws a synonym of Tikusnenn Jovaense. The species name javaense has prioriry because it was published in October 1992, while indonesiensis did notappear until in November of that yeur. Tikusnema vandycki sp. nov. FIGS 1-16 Material examined From Xeremys myeides. 54 mmmature adults and fourth stage larvae, 31 anterior ends, 40 mature & cr, 26 mature 9 2 from Myora Swamp, Stradbroke Island. Queensland. TIKUSNEMA SP, NOY FROM THE FALSE WATER-RAT ww Pescriprian Long, Slender worms with tapered extremities, clticle thm, with fine annulations, Lateral alae absent. Cephalic culicular leaves each divided into 4-5 teeth, lateral rooth targest (Figs 2.3.5,16), Cordatis rod-like in dorsoventral view (Fig. 2). Cordons and dorsal and ventral rim of pseudolabia faintly striated (Figs 2,8). Muscular portion of pharynx narrower and shorter than glandular portion, pharynx aboot 1/7 body length (Pips 1,15) Nerve ring near anterior end of muscular portion, excraory pore posterior ta nerve ting. Derrids Liny, bifid, between nerve ring and exeretory pore (Fig, 1), A pair ot culictlar ornainentatieons, saruil. inconspicious at about mid level of plandular ponion of pharynx (Fig. 12). Mile: Length 27 (25. 30) mm, width at midkxidy 412 (317-470), Cephalic end 177 (156-245) long, 240 (215-260) wide, Posterior end of cuticular leat 220 (208-26) [rom anterior extremity. Buccal capsule 269 (260-287) long, muscular portion of pharynx 435 (370-680) long, glandular portion 3229 (2975-3872) long, Detrids 307 (186-325), nerye ring 377 (338-410). ° excretory pore 499 (442-559), cuticular ornamentation (one measurement only) 2685 from anterior end. Poslenor region curved ventrally. Caudal papillae arranged in 10 pairs, 4 pairs pre-anal, 6 pairs post anal, large pedunculate, Ist and 2nd pairs grouped logether, 3rd and 4ih pairs grouped together, Ist anc 3rd more lateral, Sth and 9th pairs grouped together Sth more lateral; th pair close to tail tip. Longitudinal cuticular ridges present anterior to cloaca (Pig. 19) Spicules dissimilar, right spicule short robust, rounded distal tip 241 (208-266) (Mig, 9); lett spicule, trifid distal tip 746 (682-813) long (Pigs 4.14), longest spicule about 1:36 body length, tail 721 (598-845) (Pip. I). Female: Length 34.8 (30-41) mm, width at mid body. 555 (510-629). Cephalic end 188 (156-201) long, 255 (240-273) wide. Posterior end of cuticular leaf 238 88-260) [rom anterior extrenuly. Buceal capsule 282 (266-292); muscular portion of pharynx 461 (325-650) long. glandular portion 4040 (3111-4675) Deirids 336 (273-383), nerve ry 399 (357-422), excretory pore 500 (455-546) from anterior end, Vulva circular. without Jips, 16.5 (3.6-19.4) mm from anterior extremity. Ovejector amphidelphic: vagina vera directed transversely, 550 (one measurement), vagina utering 250 (oné measurement) parallel to body wall (Pig. 6). Tail 621 (510-748) (Figs 7,10). Eggs thick shelled, 33.8 (32-34) by 48 (44-53). Etymology The Specific name vandyeki is given in recognition of Steve Van Dyck who first noticed the presence of these worms in the host, Host; Xeramyy onyeddes Location, Stomach Locality; Stradbroke Istaned, Queenstand ‘Type specimens: Holotype male. QMIL9I5; Allotype female, QM211926, Paratypes:! QM2\1927-30; SAM24832, Remarks Tikusnema vandycki can. be distinguished from T: javaense by the shape of the cuticular leaves. In T vandycki the leaves are subdivided at the edge into 45 teeth, but in 7 javaense Ute leaves have three teeth, the middle one being the most prominent. Jikusnema vundyek? can be further distinguished by its larger size; miles up to 30 mm tong, lemales up to 4t mim as compared with 21 and 24.5 in 7% javwense respeetively: The spicules of 7 vandweki are lonper (208-266 and 682-813) than those of 7) javeense (90210 and SOK)-570). However since 7 vardyek!is.a larger worm than 7. javiense the proportion of left sprcule to body length ts smaller for TL vandvek? (36) than tor 7 javeense (E18). Male @ vandveki have a sberter tail (598-845) than. do T javacnse (B40-990). The eyes of T. vandychi (44-53 x 32-34) are larger than those of T. javaense (38-44 x% 328-31), Comparative measurements of 7. javaense and T vandyeks are given in Table J. Since the specimens exanuned by Hasegawa er al. (1992) are smaller immature adults only the measurements from Gibbons ef al. (1992) of mature specimens are used. "This allows an easier comparison of the relative sizes of nouture adult specimens of cach species. The paired cuticular ornamentations at the level of the glandular portion of the pharynx are tiny and difficult to find in 7. vanevekt but more prominent in 7) javaense. The vagina vera of 7) vandycki appears to consist-of nwo parts, # globular heavily cuticularized part leading into a more tubular less cuficularized part, which in turn opens ilo vagina uterina (Fig, 6), The vagina vera of T jawaense is similarly figured in Gibbons er ad. (1992), Further investigation is needed to determine whether the vagina vera is actually bipartite or whether the distal, globular part is. actually an ¢laboration of the yulya, Fourth stage larvae and immature adults of 7 vandyeki show similar morphological features to. those of 7 javaense. A detailed comparison and anilysis, panticularly of the development of the cephalic structures, 38 being prepared for a subsequent paper Discussion The spirutid superfamily Acuarigides is usually found in birds. However an increasing: number of genera has now been reported from mammals, The genera Starimerinemd Osche, 1955, Antechiniella Quentin & Beveridge, 1986, Chandlerancma Lute & All, 1980, and now Jikusrtenta occur exclusively in 92 LESLEY R. SMALES TIKUSNEMA SP. NOV. FROM THE FALSE WATER-RAT 93 TABLE 1. Mean measurements, in wm unless otherwise idicated, of Tikusnema species. Measurements of T. javaense are from Gibbons et al. 1992. Standard deviations are given for the measurements of T. vandycki. T. vandycki T. javaense T. vandycki T. javaense Male Female No. of specimens measured 10 4 10 2 Length in mm 27+ 2.15 20.19 34.8 + 3.16 23.5 Width 412 + 43.39 410 555 + 43.67 445 Length buccal capsule 269 + 13.9] 465 282 + 31.55 515 Length muscular pharynx 435 + 95.09 540 461 + 89.07 585 Length glandular pharynx 3229 + 383.86 1870 4040 + 578.37 1935 Deirid to anterior end 307 + 48.85 390 336 + 33.15 410 Nerve ring to anterior end 377 + 22.09 550 399 + 24.96 585 Excretory pore to anterior end 499 + 37.45 765 500 + 27.58 825 Right spicule 241 + 19.73 200 - - Left spicule 746 + 53.62 535 - - Tail 721 + 67.32 915 621 + 65.41 505 Vulva to anterior end in mm (one specimen) - - 16.5 12.21 iS Figs 15,16. Photomicrographs of the anterior end of Tikusnema vandycki sp, nov. lateral aspects. Fig. 15. optical section. Fig. 16. showing the cuticular leaves of the pseudolabia. Scale bars =100nm. Arrows indicate cuticular leaves. Figs 1-14. Tikusnema vandycki sp. nov. Fig. 1 Anterior end, lateral view. Fig. 2. Cephalic region, dorsal view. Fig. 3. Cephalic region, lateral view. Fig. 4. Left spicule. Fig. 5. Cephalic region, enface view showing cuticular leaves. Fig. 6. Vulva, vagina and uteri, lateral view. Fig. 7. Posterior end female, lateral view. Fig. 8. Cephalic end, enface view, optical section showing cordons, Fig. 9. Right spicule. Fig. 10. Female tail tip. Fig. 11. Posterior end male, lateral view. Fig. 12. Cuticular ornamentation in pharyngeal region. Fig. 13. Posterior end male, ventral view showing cuticular ridges arising anterior to the cloaca. Fig, 14. Left spicule tip. Scale bars: Figs 1,6,4,=100 um; Figs 2,3,5,8,=50 wm: Figs 7,11,13,=200 am; Figs 9,10,12,=50 pm; Fig. 14,=25pm. 4 LESLEY R, SMALES mammils. (Gybbons er ul 192) while others Synhimantus Raatliet, Meory & Sissotf. 1912- Farcewuria Rao, 195) and Skt vebinuclave Sobolev. 1043 withourh primarily found in birds, ulso occur in mammals, Various arthropods and fish serve as intermediate hosts lor the life cycle stages of acwariids (Anderson 1992). The link between mamunalian Host and acuariid parasite therefore may be one of dietary habit (Stnales 1991), A particular set of dietury preterences and habits of a few mammals thus allows these odd oecurrences of iifechon by deusriids of byologicully unrelated host species. in geographically unrelated regions of the world. Slirews from Bulgaria, Isracl, Alaska. Burope: mice rats, raccoons, muskrats, from USA, rice Tield rats from Indonesia) pyreuean desmans from Spain: Anrecitinus species, water-rals and false water-rats from Australia are all able to be parasitized by acuariids under apptopriate cireunt stances (Quentin & Beveridge 1986; Hasegawa ef a. 1992; Alvurez eral, 1994; Anderson & Wony 1994), The precise nature wf the link would probably differ from one maromalhian host to another, For example the diet, including crustaceans, and semi-aquatic habits of WN. myeides appear to fit the required pattern, Australian rodents are all included within the farrily Moaridac Their ancestors are believed ty hive: evolved in South-east Asia about 25 million years ago (Watts & Astin 1980, Then some 15-20 million years aye members of the lineage colonized the Indonesian and possibly some Melanesian islands. Geological changes during this period :solated the islands Jor greater or lesser periods of tirae allowing further speciation to occur, By 5-10 million years ago Australia and New Guiiea had moved close enough to these islands to allow colonization by what has becore Koowe as the old endemic rodents. The Australian water rat group, the Pydromyint fony part of that old endente fiuna, The suggested perics! of divergence Within the group (Warts & Astin L981) would have the Australian and false water rats ovalvine along separate lineages belure cheat arrival in Australia Both genera have closer affinities. with Vavious New Guinean rat species, in body torn god ecoloyieal niche, than they do willyeach other, The fact that both genera have acuariid parasites can be seen as a refleerion of their aquatic fo semi-aquatic life-styles und the inelasion of crustaceans in their diet. However, the acuariids found mf chryogaster, Anrechinella and Swrhimantus are also found im Australian casyurid marsupials whilst Vikwsnenma from XN myoldes also vecurs in R. arvenrventer from Widonesia. "This 96 consistent With the scenario proposed by Watts & Astin (M981) that XL mvaides is a more reeent arrival om Australia than ff, chrysexester, How the radiation of the Hydrontyint is related to the murids of Southeast Asia is unknown (Watts & Keniper 1989). A survey of the prtrusites of the Papua New Guinean Hydromini needed to determine whith, if any, acuariid parasites are present. Conclusions may then be able to be crawn as to whelher (he appearonce of Tikusnema in RK. argentiverter antl X nryoidey has any sigmilicunt bearing on murid telauonships in South east Asia Papua New Guinea and Australia Acknowledgments My thanks to Steve Van Dyck and Lester Cannot of the Queensland Museum for giving ine access be the material from the false (water-nat, Dr Ralph Lichlentels of (he Biosystematic Parasitology Laboratory and Dr Lynda Gibbons of the International Institute for Parasitology for the loan of the material References Anderson, B.C. (1992) “Nematode parusites of vertebrates Their development and trapsinission” (CAB International, Cambridge) ~—— & Wong, PL. (1994) Skrjubinovlavay Kinsellai wsp. (Nematoda: Acuarioidea) front the clee wat Oryeanivy palustris. in Flonda, Syst, Fevasite!, 28, |-+ ALVAREZ, FB... Guor-Boreni A. WH. Quinieino. PB. Rey. Lb, Loprg-Romaw, Fo & Sanmartiiy, M. bL. (1994) Paracuaria hispanic asp. (Nematoda: Acuarildae), a stomach pariate of the pyrenean desman Galery Pyeenuicus Geotly, Cinseetivera:Tulpidae). with a oder of the genus Peracwanta Bao, 195i, did, 29, S412. Biko, A. Fo& Byrp, J. (1991) "The Structure of Nematodes* 2nd edtn, (Academie Press, San Dieu). Crmmons, J. M_. CRAwsHAW, MT. & Rumeus. A. 1. 11992) Malinieuaria indonesiensis tsp. (Nematoda Acuarioitesy Tron Rats arventiventer in Indonesia, Syst Aarasital 23, 175. 1BI. HasnGaws, H, Strats, S d& RocHiman (1992) Tidteseerne favaenve nosso rsp, (Nematoda: Acuarinidea) and other nematodes from Rattus araenrtvener collecred in Wess dasa J, Panastinl, TR. 800-404 Quentin, JC. & Brverince, L (86) Comparative morphogenesis of the cephalic structures of the aeuariid nematodes Starunerind sorties (Tiner, 951), datechinietls Snffodiax (Beveridge & Barker, 1975) pyen,, meomb and Shrjabineclava thapari (Teixeica de Freitas, 1953). dysr, Parasitol. 8, 63-171, Smaces, L. R_ (1991) A new species of dareciiniella Quentin & Reveridge, 1986 (Nemutoda:Acuarndac) trom the Australian water mit Jivdrgmus chiyyeyescen Geotlroy, (ROS Trans, R. Soe, 8, Ast, W5, 217-220, Van Dyex, & (1992) Parting the reeds on Myora'’s Nevrrnvy kibbutz, Wild Aust 29, 8-10, 11994) The rats at nepiune’s table. tase, Mar. His? 24, 30-37 Waris, CHS. & Astin, Ho F981) "The Rodents of Australia” (Angus & Robertson, Landon), —___— & Keweek, CM, (1989) Muridae pp, 949-957 Jn Walton, D. W. & Richardson. 8, J. (Rds) “Faun of Aasiratia” Vol 1B Mammalia (Ausr Cievi Publishing Service, Canberra) Wona, PL. & Lancaster, M. W. (985) Revision of the genus droyracanhopsts Dicking, 1861 and deseription of a new genus Molintucudria nawen. (Nematoda: Acwurtdoidea). Can, 4. Zool, 63, (S501564, MASTOPHORUS MURIS (NEMATODA: SPIROCERCIDAE) FROM THE MUSKY RAT-KANGAROO, HYPSIPRYMNODON MOSCHATUS BRIEF COMMUNICATION Summary The Musky Rat-kangaroo, Hypsiprymnodon moschatus Ramsay, 1876, the smallest and most primitive of the macropodoids, occurs exclusively in the rainforest of northern Queensland." It forages in the leaf litter of damp areas of the forest for fungi, fallen fruits and invertebrates. H. moschatus has a sacciform stomach intermediate in structure between the simple stomach of the phalangerids and the complex, compartmentalized stomachs of the potoroos and macropods.” Apparently the Musky Rat-kangaroo has not adapted to a diet with a high content of cellulose dependent on a fore-stomach fermentation chamber, but rather has retained an omnivorous diet of higher nutritive value.’ Tronsactions af tke Royo) Suctery of S Aust, (995), L9(2), 95~96, BRIEF COMMUNICATION MASTOPHORUS MURIS (NEMATODA: SPIROCERCIDAE) FROM THE MUSKY RAT-KANGAROO, HYPSIPRYMNODON MOSCHATUS The Musky Rat-kangarov, Aypsiprymnodon maschaius Ramsay, 1876, the smallest and most primitive of the macropadoids, occurs exclusively inthe rainforest of northern Queenshind,! It forages inthe leaf litter of damp areas of the forest for fungi, fallen fruits and invertebrates. H. rescharis has a sacciforn? stomach intermediate in structure between the simple stomach of the phalangends and the complex, compartmentalized stomachs of the potoroos and macropods.* Apparently the Musky Rat-kangaroo has not adapted to a dict with a high content of cellulose dependent on a fore-stomach fermentation chamber, but rather has retained vn emniverous diet of higher nutritive value!. A-small colony of H. maschatns has been maintained, for research purposes, at the Queensland National Parks und Wildlife facilities, Pollarenda, Townsyille, One of these animals died in its pen in 1994 and was subsequently made available for dissection. Twenty imale and 17 female Mastaphoras muris (Gmelin, 1790) were found in the stomach. The worms were fixed in 10% formalin, stored in 70% ethyl alcohol and then cleared in lactuphenol for microseopic examination. The males measured up to 25 mm and the females 63. mm Jong. The worms appeared maturé and healthy, the females being pravid, Mastopharas mariy is a nematode from the family Spiroeercidac, cosmopolitan in rodents of the families. Microtidae and Muridac*, [t has previously been recorded from rats, a “mouse” and eats in Australia. 2" A range of insect species including cockroaches has been finind lo be surtable intermediate hosts.” Both cock rmaches and rats are attracted to human food stores 'Jonnson, P.M. (1988) “Musky Rat-kangaroo” pp, 179-180 In Strahan, R, (Ed.) “Complete Book of Australian Mammals” (Collins, Angus & Robertsen, Ausiralia). “Hume, 1. D. (1982) “Digestive Physiology and Nutrition of Marsupials" (Cumbridge University Press, Cambridge), Vanderson, R. C. (1992) “Nematode Parasites of Vertebrates Their Development and Transmission” (CAB International, Wallingford), 4Mackerras, M. H. (1958) Proc, Linn. Soe. NSW 83/2), 101-160, S5Obendorf, D, L, (1979) Aust, J. Zool, 27, 867-879. 6Smales, L. R. (1992) System. Parasitol. 22, 73-80. 7Gordon, H. MeL. & Sommerville, R. 1. (1958) Aust, J, Science 21, 148-149, such as those kept.to feed the captive animals at Pallarenda, If M_ inuris were established in an infective cycle, including cockroaches and rats Itving in close proximity to the pens, then such cackroavhes when caten by Musky Rat-kangaroos could be the link between nomal mdent hosts and the accidental macropodoid hos, Once ingested M. muris ts apparently able to establish itself in the.simple non-fermenting Ral-kanuaroo stomach. The environment here could niore closely reseroble that of rodent stomach than that of the more complex fermentative stomach of macropods and potoroos. This is the first record of M. murixy occurring in a macropodoid marsupial. The ealy other records of M, mauris fron) marsupials are [rom a phalangeroid, the Broshtail possum, Trrehasyrus vidpecula (Kerr, 1792). Worms fram infected possums were: first described front Queensland)” and then reported from New South Wiles! as Propaspinera marsupialis Baylis, 1927. This material was subsequently reexamined and deterniined to be M. muriy'*, The diet of the Brushtail possum, including fruits and micat if offered.2-® is more similar to that of the Musky Rat- kangaroo than to that of other macropodois. The non- fermenting environment within the possums stomach is probably alse similar to that of A. meschatus. Therefore it is likely that, as with H, moschatus, To vulpecuia could also become accidentally infected wilh M. muris from time to time, 1 am indebted to P.M, Johnson for the opportunity to examine the Musky Rat-kangaroo. All the nematodes collected have been deposited in the Australian Helminthological Collection, South Australian Maseum, Adelatde. SQuentin, J. C. (1970) Ann. de Parasitol, 45(6), 839-845. °Baylis, H. A, (1927) Ann, Mag, Nat. Hist, (95)20, 214-225, Johnston, T. H. & Mawson, P. M, (1938) Ree. 3, Aust, Mus. (2), 187-198, "Baylis, H. A. (1434) Ann. Mag. Nat. Hist. (108)14, 142-153. 2 Johnstan, T. H. & Mawson, P. M. (1939) Trans. R, Soc, S. Aust. 63(2). 209-209. UWertheim, G. (1962) Trans. Am- Micros, Soc, 81(3), 274-279. Mew, R. A (19881 “Comnion Brushtail Possum” pp. 147-148 Jn Strahan, R. (Ed_) “Complete Book of Australian Mammals” (Collins. Angus & Robertson, Australia)- L. R. SMALES, Department of Biology, Central Queensland University, Qld 4702. OTOLITHS AS POTENTIAL INDICATORS OF AGE IN COMMON CARP, CYPRINUS CARPIO L. (CYPRINIDAE: TELEOSTED BRIEF COMMUNICATION Summary The common carp, Cyprinus carpio L., was among the first species of fish for which techniques of age estimation were developed’. The annuli of scales (seasonal zones of slow growth) have been used as growth indicators in carp from Asia, Europe and North America”®, Opercular bones®*, fin rays’ and spines'®"' have proven useful, and the eye lens may also have value” although it is unreliable for older fish’**, Otoliths have been used successfully for the cyprinid Phoxinus phoxinus’*”’, but not for carp!!! Transactions ef the Royal Society of S. Ausi., (1995), W942), 97-98 BRIEF COMMUNICATION OTOLITHS AS POTENTIAL INDICATORS OF AGE IN COMMON CARP, CYPRINUS CARPIO L. (CYPRINIDAE: TELEOSTED The common carp, Cyprinus carpio L., was among the lirst species of fish for which techniques of uge estimation were developed!. The annuli or seales (seasonal zones of slow growth) have been used as growth indicators in carp tron Asia. Europe and North Aimerica?>, Opercular bones’, fin vays “and spines!!! have proven useful. and the eve lens may also haye value! although iis unreliable for older fish. Otoliths have been used successfully forthe eyprinid Phavinis phoximes'® 2) bat not for carp! 2! It is not clear whether the authors considered the full complement of otoliths in these studies. Carp. and teleost fish general, have (hree pairs of ulricular. saccular and lagenar Otoliths, respecuively named the fapilli, sagittic and asterise! Given the peculiar morphology of The Jower part of the nmer ear of eyprinids (and other ostariophysan fish), the asterise ynd the lapilli are much larger than the comparatively thin. elongate sagittue?? Lapilli have been used to distinguish daily growth increments in the fllfsh Semrefitis corporaliy’ and the rose biterling Rhiadexs vecellarts necellatis. and adiurmi rhyihin of valeiim deposition has heen ceported in the asteriscr of youny poldfish Caragsyitey auras? Unspecified otoliths have been used to age tench, Titer tinea in Burope?’, hutitus-likely that these were asterised or lapilli mither (han sagittae, fn revent work on the cology of carp We have heen able consistently co recover well-formed onoliths with patterns thar upper to represent a chronolopieal record, As validation is necessunily 4b protmacted procedure we belweve that 4 prehmingry eommunivation i warranted This work hus special sizniticunce in Austruliy because carp are an Introduced species thutis widely believed responsible for the degradation of wetlands throughout the Murray-Darling Rasin2’. 1 the impact of carp is to be evaluated the ability to estimmite the ages of individual fish, providing for ireusurements of wrowth. feceuitment and other popularion parameters, Samples for thas study were obtained by wilbnetting backwaters of the River Murray al Swan Reach and at Gurr Gurra Lukes near Berri, from January 16 April 1994, Body weight (to within 0.1 wand fork leneth (can) were measured is oraentl heliire recovering the ofoliths und recording (he weights of the asterisc: (LE mu). Annuli (translucent bands) were counted on the distal side of whole asterise When more than three annull were present the earhescones, particularly (he first and second, were olen obscured by culaium deposition, When more than 5.6 annul Were present the outermost Ones were more eusily diseernible in transverse thin sections (hun in whole otoliths. The lapill proved Useful only when 2-3 annuli were present, and otherwise underestimaled the counts trom the asteriscr, The sigittae showed no recognisable pattern. For these redsons. aMensct appear to be potconully inere usetul us indicators of age in this carp population. In Figures live the asteriscus weight, fork length and body weight of 63 curp are plotted apainst the numbers of ammuh on the asteriscus, Strong correlations are evident in cuch plot (respeenvely, Spearman rank correlation coellicient r= 0.873, P 3 Kalimnaty z = . = a & Cheltennam € 4 Sspef se] & - as Papi = | 2a E Michallian . 2 eS ae ga 5 5 2 aJSa] bs 5 : = 2 E =e 5 2 & = Bs 2a a * 2 5 3 = z Barmsaahian [-4 © nu = Sa S 4 e a 5 eS bee £ 1" | Balcombian_ ! id | = Batefordian = 5 : 5 ; 4 6 ; = Longtordiary 4 a 4 [= a q an fis =a & Beri - Nossal ® & w “| . e = @ 5 2 FI & E = : a danjukian i ae ee a = ? g : 3 ah 2 = = = z = a” & 3 “Ee es 8 e ; — 2 3 & = 2 : = Willurgar £2 a a = : 2 eye eet += a : 2 :3 _— - Toe. = ¢ Hi : ~ Aldingan 1 (hat narned) Fiz. 2. Ranges of the taxa discussed. Geochronology after Cande & Kent (1992) and N/P zones after Berggren e/ ul, U98Sa, b). Correlation of local planktonic foraminiferal data and regional stages follows McGowran er al, (1971), Heath & McGowran (984), and McGowran & Li (1993, 1994), 12 0, LI & B, MeGOWRAN Quilty (982. p. 10) listed over 20 cibieidid species known from the Tertiary of southern Austmliiun and New Zculand, Together with the meonyex allied Cibr- cidiides. the planovonves venus Cibicidles averages 20% - 50% Of total fauna in most suniples, Typical vibicidid forms tnchide Cibieides thungia (Fiz. 4. 7, 8), GC mediverts (Fig. 4.9, 10), C. vortex (Fig. 4, 6a, bh), Cibicidoides perforatus (Fig. 4. Wand C. pyeud- nuverianus (= Cibicides neeperforams) (Fi. 4. 12. 1a). The evolution at ©. pseddoungerianus trom C peifardns was in the lite Bucene. by a reduction ot coarse perforations trom both sides (an CL perforarus,) und restriclion jo the spiral side of the test, The stranyraphically most useful species is Co karreriformix Hornibrouk, oecurring in the Oligoeene (Fig, 2), Other previously described species, such as C. subheidieert and C. opacis, are now placed in the genus Heterplepe (see below), Three cibteidids characterising the modern biolicies vin Lacepede Shelf, South Austalia, ure Crbivides refidvens (Fig. 4, 5), C. nweiovriv and Cibieidoides preudoungerianuy (Li etal, 195), Fig. 3. Ja, b. Crespinells umboniferd sketches of the holotype of 2Qperculing wanbonifera Howehin & Purr (1938) (see also the Sedrining inicrographs in Fig. 4. nes. ia, by. 2, b, Mastinella chapmani Gluessner & Wade (1959): sketehes of the holotype. Both types are deposited in the South Australian Museu, Note that the final chamber on both qests is missing, bul umbilical openings (uo) sre present. Family Epomuidae Hotker LWAl Genus Cresprned/ia Parr 1942 Synonvery arad Type species, see Luebhch & Tappary N87, p. S7Y, Remarks Pary (942) erected Ihe Gully Miocene taxon Oper culina unibonifera Howehin & Parras (he type species Of his genus Crespinedla. separating (his simple tarm from similurly planispirul bot internally comples Operculina Crespinella wax monospecihe until Quity (YS) udded to it another species. C. per’. with alow troch- ospiral (other than planispiral) coiling. The overall hrorpholugival similarity between C. uwnbentfera ind C. parri ted Quilty (980) to imply that both C. parri and C. umbenifeéra are phylogenetically related, with C. parri being the predecessor, Loeblich & Tappan (1987), however, rejected tis statetient on the basis ol the distinet trochoespiral coiling and supplementary suluinitl openings i Quilty’s species. Such contusien over the generic stats at CL pared needs (0 be clarified Li has Mspected the holotype of Co wabonifera, which was made availible from the Seuch Australian Museum, and found that it also possesses dn opening on the umbilical side (Fig. 3. la, hy Pig. 4. 4a. by. It is an incomplete specimen wath the final chamber missing, und a solall opening can be observed at the buse of the relic part of the missing, chamber, close ta the margin of the pronounced umbilical boss, Ne umbilical openings, however. were found related (0 ary previous chambers. We thus conclude thal the species C. parri is correctly assigned to Crespinella, a genus having species with a very low frochospiral to planispiral coiling and one or tigre supplementary openings on the umbilical side. Genus Hefkerinag Chapman & Parr, 193] Synonymy und Tipe species: see Loeblich & Tappan 1987, p. S3L, Remarks Geographically Hofkeriia semicrnara (Pix 9, Wh, 12 is similar to dimaena gippslandica. as both ure confined to the southeastern corner of southern continental margin (Carer (958. 1964) Almaena gippslandica is an earliest Miovene form und apparenuy has affinities with species trom Paratethys (see ubove). whereas Hofkerina semiornaiet seems i be entirely endemic ty the region wilh w pang: Prony the curly Miocene lo early nniddle Miocene. itis notewarthy that both Mofkering semiornata andl Crespinella vmbonifera, above. have a similarly thick wall, which mimics the wall inthe Eocene Mastinelle chapmani (see below). Unlike H. semiemata. however, C. uinbonifera and M. chapmani. have also been recorded from South Australia and Western Australia (Quilty 1980, 1981), [tis nol clear whether the thick wall in these endetme taxa signals a high CaCo, buildup in local waters during the warming phases of the later Eocene and eurly-iniddle Miocene. SOUTHERN AUSTRALIAN FORAMINIFERA 14 i4a wee, met 21)())1 omer 2001 Fig. 4. Scale bar = 100 pm, unless otherwise indicated, Ta, b, Almaena eippslandica Caner: two views of a single specimen, latest Oligocene, Lakes Entranee. Sample 156. 2, 3. Averiverinella adelaidensis (Howehin): two specimens, later middle Eocene. Tortuehilla Limestone, Maslin Bay. Sample AB-Tor, 4a. bo Aronnimannia haliotis (Heron-Allen & Eurland): single specimen frons the carly middle Miocene, Cadell Marl section, Sample C9. 5. Cibicidey refulgens de Montfort: Recent, Lacepede Shelt, Sample 89-1, water depth 171m. 60, b. Cibieides vertex Dorreen: single specimen, early Miocene, Lakes Eniqince, Sample $08, 7,8. Cibicides thangia Finlay, Wo specimens, carly Miocene, Lakes Entrance, Samples 992 and 732.9, 10. Cibicides medineris Finlay: two specimens, late Oligocene and carly Miocene. Lakes Entrance, Samples 828 and 1196, [L, Cthicideidey perforaius (Karrer); late Eocene, Blanche Point Formation, Maslin Bay, Sample A3-091 12, 13. Cibietdoides preudoungerianus (Cushman): two specimens, late Oligocene and eurly Miocene, Lakes Entrance, Samples 1196 and 956, Ida. b. Crespinella umbonifera: lwo views of the uncoated holotype of ?Operculina umbenifera Howehin & Parr, using a Philips &L20 scunning electron microscope at the University of Adelaide (CEMMSA), ht Q LI& B. McGOWRAN P49 es 5) Fig 5. Scale bar = 100 am, unless otherwise indicated. | Crespinina kingycotensis Wade! axial section, carly Ohgovene, Port Vincent Limestone, Yorke Peninsula. Sample BSI. 2) Lepideevelina howehini Chapman & Crespin: axial section, early middle Miocene, lower Morgan Limestone, Mannum. Sample Li/93-1. 3, 4. Exearnebaving cuvillieri (Poinant):; Wo specimens. carhest Oligocene, SADME bare Ad), western Murray Basin (3), F805, and SADME South Parklands Bore, Adelude (4), FP 808, both from Lindsay (I98L, pl. 44. figs 1, 3). Note that Lindsay’, 1994 (pers, comm.) considered the form inno. 4 nota typical specimen of that species, 5, 6, Halkyardia barrrumi Parts wo specunens, late Eocene, Custle Cove, Sample RIPI 19. 7a. b, Aelerolepa opaca (Carter): single specimen, carly Miocene, Lakes Entrance, Sample 724. 8-9. Heternlepa hrevoraliy (Carter); (wo specimens, early Miocene. Lakes Entrunee, Samples 788 (8a, b) and 984 (9). 10a. b. Heleralepe subhendingeri (Parr); single specimen, early Miocene. Lakes Entriince, Sample 852. 11, 12. Hajkerine semiornate (Mowehin): two specimens. curliest Miocene. WMC 703, Samples 45.35 m and 45.65 m SOUTHERN AUSTRALIAN FORAMINIFERA es family Chapmaninidae Thalmann. 1938 Genus Crespinine Wade. 1955 Synonymy and Type species. see Loeblich & Tappan 1987, p. 668. Remarks This genus, together with its only species C. kineseotensis (Fig, 5, 1), apparently represents ope of the numerous laxa endemic to southem Australia. It hay heen recorded in South Australia (Wade 1955; Ludbrook 1961). Victoria (Carter 1958) and Western Australia (Quilty 1981), Quilty (198i) also noted that lests of C. Aingscotensis became larger and more robust from cast to west, indicating a warmer temperature towards the western part of the southern continental margin. Crespinina Kugscotensis occurs mainly in the later middle Egeene to early Olipocene (Wade 1955). In the Port Vincent Limestone from Yorke Peninsula (Pig 1). Vis associated with some planktonic foraminiterst such as Guembelitria, an catly Oligocene marker m local biostraligraphy (McGowran & Beecroft 1985), ynd its last appearance precedes the first appearance of Amphistegina. The latter daium in the region was within the fate Oligocene (Lindsay 1985). Family Gavelinellidae Hofker, 1956 Genus Excornebovina Butt, 1966 Svnonviny and Type species: see Loeblicly & Tappan 1987, p. 633, Remarks Specimens referable to E. cuvilliert were found in the basal Ettrick Formation (Oligocene) from the western Murray Basin and eastern St Vincent Basin (Lindsay 1981'), but this record has never been made public These specimens were compared with the near- topotypes of FE. cuvilliert from Escornebéou, France. supplieu lo Lindsay by Professor C, W. Divoger (Utrecht), This record thus extends the peographic distribution of this taxon from Paratethys Lo southern Australia. Two of Lindsiy’s specimens are shown in Fig. 5, 4 4. Furmily Cymbaloporidae Cushman, 1927 Genus Hulkvardia Heron-Allen & Harland. 191% Svnonviny and Frpe species: see Loeblich & Tappan 987, p. 593. Remarks ‘The conical Halkyardia barirumi (Fig. 5, 3a, 6) has been widely recorded in New Zealand from where it ' Livpsay, J. M. (i981) Tertiary Stranyraphy and Foraminifera of the Adelaide City Area. St Vincent Basin, South Australia, Unpatl M.Sc. Thesis, The University of Adee, was unginally named (Homibrook er al. 1989). In southern Australia, Ludbrook (1961, as Halkverdia sp.) found similar forms in the western Murray Basin, and Quilty (1981) recorded it in the Nanarup Limestone near Albany, Western Australis (Pig. 1). On the castern margin of the St Vincent Basin, this species imakes Ewe brief appearances. in Ihe Tortachilla Limestone and the basal Port Willunga Formation (Lindsay 1967) McGowran et al. (1992) recently Correlated these twa intervals as from top PM to early PIS in the later middle Eocene, and upper PIS in the early Oligocene respectively. No record of this taxén has been reported to date from the eastern corner of southern Australia, Family Heterolepidue Gonzales-Donoso, 1969 Genus Heterolepa Franzenau, 1884 Synonway and Type species; see Loeblich & Tappan \987, p, O32. Renusrks Many species of Heicrolepa were previously recorded as Cibicides in southern Australia. The genus Heterolepa difters from the radially walled Cibicides in having a granular wall and an aperture which does not extend lar onto the spiral side (Loeblich & Tappan 1987). Cibicides breveralis (Fig, 5, 8-9), C. apacis (Fig. 5, 7a, b) and C, subheidingeri (Fig. 5. Wa, b) all appear to have these features, and are accordingly (ranslerved to the genus Hetérolepa. Also included in this genus is Cibreides victortensiv (see also Lindsay 1969, 1981), a species confined t> the middle Miocene, or Zones N9-N1I3 equivalents. Morphologically, H. victoriensts is similar to both A. brevoralis and H. subhaidingeri. but differs from the latter two in the strongly limbate sutures on the spiral side At the Morgan-Cadell section, western Murray Basin, Herervlepa decreases from the lower Morgan Limestone, disuppears in the Cadel! Marl, and reappears in the upper Morgan Limestone. The Cadell Marl is composed mainly of bioskeletons including abundant milinlid and discorbid foraminifera. and represents a restricted. but highly productive. environment, Phe marly sequence is dated at about }S Ma, in the Jater part of the Miocene chmatic optimum (Li & MeGawran 1995). Its absence froin the Cadell Marl indicates that Heierolepu may be an open marine genus only. in contrast to the ubiquitous Cihicides, Family Lepidocyvlindae Scheffen, 1932 Genus Lepidoeyelina Giimbel, 1870 Synonymy and Type species: see Loeblich & Tappan 1987, p. 614, Remarks The last oceurrenee of Lepidocyelina sensu lata was LOG Q, LI & B MeGOWRAN in the middle Miocene (Zone N9), if dor the late Miacene or early Pliocene (Adams 1992). This has heen apparently misquoted 10 he in the Aquitanian (Nd. eurliest Miocene) by Loeblich & Tappan (1987). The local representalive of this genus is L. hesvehini (Fig. 5, 2), a species widely reported fron various localities in southern Australia (Ludbmok 1961; Lindsay 1969: Lindsay & Giles 1973; McGowran [979; Quilty 1982, Chaproniere 1984; Lindsay 1985) Associated with many other larger forms. ib was conlined (a the litest early Miocene tu earliest nddle Miocene, or Zones N¥ and NO equivalents, Us occurrence in the region has been hailed as a sigrval of the Miocene climatic optimum (MeGowran 1979: Frakes e7 a. W987, McGowran & Li 1993, 1995), Fanvily Linderinidae Loeblich & Tappan, 1984 Genus Linderina Schlumberger. 1893 Swenvew dnd Wyre speciess see Loeblich & Tappan 1987, p. 645. Remarks The species Linderma glaessneri is large, discoid and internally complex with numerous chumberters (Pig. 6, 1). Like Halkyvardia bartrami, above, it wis restricted to the central and western parts of the region and has never been recorded from either Gippsland or Bass basins in the southeasrern corner. The stratigraphical oecurrence of Linderina wlaeysneri is also similar to that of A, partrumi in two short intervals; later middle Eocene (Zones top Pl4tower PIS) and earliest Oligocene (upper PI8). This record thus extends the range of that genus into the early Oligocene from the originally middle and late bocene (Quilty 1981). Family Elphidiidae Galloway, 1993 Genus Parrellinad Thalmann, 195) Synenvmy und Type species; see Loeblich & Tappan 1987, p. 677. Remarks Wade (957) emended Purrellina, a plunispiral elphidiid which appears to have been restricted to southern Australian waters during its early evalutionary history. It first appeared in the middle Oligocene, Zone P21] equivalents, abour 15 Ma ufter the evolution of its trochospiral ancestor Nereraralia Finlay, The New Zealand taxon, Discwrotalia, ts similar to Parrellina in many morphological aspects except the distinct evohite spiral side, and both sre believed to have evolved from the trochospiral Noreratalia (Eocene-Recent) in the late Oligocene. Ir ty difficult, however; to separate Parre/ling from Discoratalia, as same of our Oligocene-early Miocene specimens of Parrellina vrespinae and Poet imperdrety vend to be alse Tow-trochospiral (Fig. @& 5-6). ‘Typical planispical FP iinperdtrit (Fig. 6 7) seems to have decurred only from the curly Miocene to Recen|. Modern speciniens SUP Tmperairis Vor offshore southern Australia may tow at test> Dommi in diameter while is allied form F verriculata is much smaller and without peripheral spines. A large. typically plamispinal species existing in the early to middle Miocene (NG@-NIQ) is P eraticulatifornis (Fig. 6, 8), Fainily Siphonidwe Cushman, (927 Genus Siphonndes Cushman, 1927 Syaronyery aud Type species: see Loeblich & Tappan 1987. p. 572. Remarks A smooth form deserihed by Huwehin (1889) as Truncatulind echinaia vat. laevigata is appanently a Siphoninoiles (Pig. 6 4). Whether the smooth wall has been subject to the efter of cold waters js not known, This consistemt feattire gueanlees Ural the laxon 1s a distine| species, The genere description of Siphoninoides, as in Loeblich & Tappan (987), should be revised to embrace this feature. We found nurnerous specimens of S laewpare in suinples from the Cadell Marl section, western Murray Basin (Fig. 1). Theage-of these samples is wilhin Zanes top N& (o NY equivalenis, early middle Minvene- Quilly (994, pers: conuy.) mndicuted that uw similar form exists in the modern Swan River estutiry, Western Australia, Family Uvigerinidaé Haeckel, 1894 Genus Siphouvigerina Parr, 1950 Synonyms and Type species: see Lochlich & Tappan 187, p. 525. Remarks This genus was supposed to oecur only in the Holacene (Lochlich & Tappan 1987), However, we recently discovered torms similar to the type species S. fimbriata trom (he Lakes Enptranee section. Gippslind Basin, ina level correlated to the carlest Miovene,, One of the specimens is ilustrated in Fig. 6, 9). Our record thus extends the punge of this venus down te the early Mohocene, although the form was found only sporadically; Revels (1993) recently found the type specimen of 3. fimbriata ta be biserial throughout, a finding contrasting the conventional definition of the genus (e-g Parr 50, Loeblich & Tappan 1987), However, omany uvigerinid and angulogerinid forms are triserial intuilly and change to biserial a any later sage, The triserial part of the test would be difficult to define it carly chambers ure loosely epiled, a case most likely CXIStiNE iS. fimibriata, SOUTHERN AUSTRALIAN FORAMINIFERA {7 Fig. 6. Seale bar = 100 pn, unless otherwise indicated. |. Linderina glaessnert Quilty: late Eocene, Case Cove, Sample RIE! 19, 2, 3, Mastinelle chapmani Glaessner & Wade. (wo specinens, late Bocene, Adelaide area (Children’s Hospital). Sample 19.2-19.5 m, FI 955 und FI 956, both trom Lindsay (98T', pL 48, figs. 1, 4), 4. Siphoninoides laevigatus (Aowehin): later early Miocene. Lower Morgan Limestone, Sample LM2. Sa, b, Parrellina crespinae Cushman: single specimen, earliest Miocene, Lakes Entrance, Sample 140. 6a, b. Parrellina cl inperatrix (Brady): single specimen, eurly Miocene, Lakes Entrance, Sample 992. 7, Rarrelfina imperarrix (Brady): Recent, Lacepede Shelf, Sample 89-60. water depth 82m, 8, Parrediina crancularijormis Wade. hater early Miocene, Lower Morgan Limestone, Sample LM2. Y. Siphoyenerina fimbriata (Sidebolom): earliest Miocene, Lakes Entrunee, Sample N40, 10. Tbulogenerina ferax (Heron- Allen & Earland): later early Miocene, Lakes Entrance. Sample 700. 1k. Tibulogenerina mooraboolensts Cushman, later carly Mivcene, Lakes Entrance, Sample 868.12 Cifel/ia cestata (Heron-Allen & Barland); later early Miocene, Lukes Entrance, Sample 708. 13. Ficrortella convidea (Rutten): carliest Miocene. WMC 703, Sample 45,05 m. 14. Hadella hamiltonenyis (Glaessner & Wade); lite Eocene, Blanche Point Formation, Maslin Bay, Sample 099, 15a, b. Parredicta kalimnensts (Parr): single specimen, ater middle Miocene, Lakes Entrance, Sample 416, 108 O, Lt & Th Mectiwaan Family Siphogenerinoididae Suidova, 198i Genus Tubulogenerinag Cushman, {9297 Synanvery ane Type species: see Loeblich & Tappan 1987, p. $20. Remarks Gibson (1987. 1989) Gibson et al. 1991) conducted a series of studies-on the evolution and distribution of Titbulogenerina and related taxa. Two main conclusions from His studies are= (1) this genus ranged from early Encene to Plincene, with Europe being the site of its first evolution, and (2) species seem to haye migrated westward from Europe, through the Atlantic. to Pacific and Indian Oceans, According to Gibson (1989). mid-latitude Miocene species were largely confined to the laler early Miocene tw early middle Miocene. or Zones N6 to N& equivalents. Quilty (1977) reported T picerubealensix rum the carly Miovene in Tasrnania, In the Lakes Entrance oul shaft, we found Liree tubulogenerinines (Fiz. 6, 10-12): 1. ferax, T mooraboolensis and Cifellia costar. The combined range uf these species is fron 263 ry - 157 m inthe seetion, which is mid-N5 qo early NID in our cortelation (McGowran & Li 1993, 1995). We follow Gibson (1989) in considering & vosteater a tubulogenerimd without a toothplate. Revets (1994), however, classified Cifellia and Tubulogenerina into two different superfamilies, on the absence and occur rence of toothplates in these two genera respectively. Whether the touthplate ever exists in the early part of C. costiia is not known, and tittle evidence has heen found to resolve problents such as the developinent and reduction or function of foraminiferal toothplates (Revets 1993) Family Victoriellidae Chapman & Crespin, (930 Genus Muslinella Glaessner & Wade, 1959 Synonymy end Type species: see Loeblich & Tappan 1087, p. 596, Remarks Similar to several other endemic taxa, this zenus is also monospecific. Mastinella chapmani (Fig, 6, 2.3) isa large but internally siinple form ranging from the later middle Eocene to earliest Oligocene. Although not mentioned jn {he original description, sutural Openings occur on the umbilical side of some specimens. (Fig. 6, 2), possibly resulting from relic apertural extensions. This feature can be seen even in the holotype. sketched in Fig. 3 (compare Glaessner & Wade 1959, pl. 1, fig. 7). Crespinella parri Quilty, above, is morphologically similar to Masdinella chapmani at least in the following: (1) a large, low trochospiral test which tends 0 be plamispiral in (he final stage. (2) a. distinet peripheral keel. (3) sutural openings on the umbilical side, and (4) a thick. Jantinwtect wall theugl perlorations on M. chapmeani were much coarser. All these indivale that C, parr’ is marphologacally. if por phylovenctically, closely related 19 Maslinella. The peeurrence of C. parri im the late Gligocene is cryptogenic, and penuing sidies of its relationship with M. chapmuni are necessary. Genus Merariell¢ Chapman é& Crespin, 1930 Synovviny and Type species: see Locblich & Tappan 1987, p, 59. Remarks Glaessner & Wade (j¥59) emended this genus and discussed its affinities, They found the type species Fictoriella plecte io bea jumor synonym of Carpenteria convidea, now KF comnidea (Rutten) (Fig, 6, 14). The total range of K concidea in southern Australia is From the latest Bucene (P17) tw cartiest Miovene (M4). Ludbrook (1971, p. 64) noled the tansition of V/ convidea trom Carpenieria hamilionenyis (now Wad- eller hamiltonensis, see below), in the carliest Olio cence Glohiverina angiporoides dangiporoides Zone The Bocene-Oligocene record of (hal species, hiow- ever, is relatively rare. Only in the latest Oligocene and earliest Miocene did F conoidea become common und soulhern Ausiralia-wide, us well as fram north easiern Australia (Quilty 1993). Tt is conspicuous in the curbunate-chert association of the Gambjer Lime- stone in the Otway Basin (G. Moss, 1994, pers cont), Genus Wadella Srinivasan, 1966 Syronymny and Type species; see Loeblich & Tappan 1987, p. 596. Remarks The genera Wadella and Vieroriella are similarly large and high trochospiral, However. Mudella hwniltonensis (Fig, 6, 14) can be distinguished from F conotdea by its smooth test lacking pillars and less regular coiling. In the later middle to late Envene, Hiadelhe hamiltonensix was one of many large species endemic to southern Australia and New Zealand. Prior iw the lute Eocene, in southern Australia, MWadeflu Aaniltonensiy achieved a wider distribuuoa than ( convidea (Cooper 1979: Quilty 1981; Lindsay 1985) Inthe Maslin Bay seetion, W hami/tonensis was found in the Tortachilla Limestone and basal Blanche Point Formation. in an interval equivalent io Zones upper Pi4 i PIS (McGowran er al, 1942). Hiidella glebifarnus also evelved in the late Bocene, and ranged into the early Miocene, Unlike % humiltonensiy, W slobifarmis developed a law trochospiral test and globular chambers) SOUTHERN AUSTRALIAN FORAMINIFERA 109 Camiuy Bapeinidae Cushman, (927 Genus Parredicta gen. ov (FIG 7) Type species: Valvulinerta porifera Parr, 1950 2 3 —_— Fig. 7. Parredlicta porifera (Parr), Scale bar = 200 po, la- c, Scanning micrographs of the unvoated halotype of Vilvadineria paritera Purr, 2. 3. Twa specimens from Facepede Shelf, Sanyples 49-3 and 491. 7 water depths 123m (2) und I71 om. (3) respectively Enyirmulaey This geous is named in honour of W. J. Parr, who was one of the most influential and prolific foraminiferal students in southern Australia m the early part of this.century, and who originally described the species on which this few genus. is based: edrorue (Latin = proclamation or decree, Deseripiton Test medium to large, low trochospiral, biconvex chambers high, enlarging regularly, more than 6 1 the final whorl; 1% to 2% whorls in adult tests, surliace smooth, sutures radiate to strongly curved, depressed or flush on ventral side, flush and limbate on the dorsul (spiral) side; urnbilicus small, depressed or closed with shell material, but wilhoul a distinct umbilical boss, periphery narrowly rounded lo weakly keeled: wall culeareous hyaline, distinctly perforite excepra small area immediately above the apertural lip: aperture large, arched or slit-like, extending from periphery to marginal urea of the umbilicul depression. apernural lip distinet, regular or irregular: supplemeniary opembgs copimoen, resoliing trom cither irregular growth of the lip or relic extension of the aperture on the uinbjlical side. Rennttks This genus differs trom Walvwiineria mm haying an ovul test outline, angular periphery and supplementary openings, and lacking apertural Naps. Valvudineria Cushman has a pronounced apertural flap whieh proyects over the umbilicus (Loeblich & Tappan 1987, p. 547), Many species of Madvd/ineria are rounded in outline, with # distinctly lobate margin which ts broadly rounded in peripheral view, und have no supplementary openings On the umbilical side Parredichas introduced to accommodate two species which were originally considered as Planutina kalimnensis Parr (Fig. 6, \Sa, b) and valvalinerta povifera Parr (Pig, 7). Among others, Carter (1964) und Quilty (980) recorded Parredicra kalinmensis (both as Keladineria kelimnensis) in the Miocene of Victoria and Tasmania, Inthe Lakes Entrance oil shaft. it was found from 340 1m to the top of sampling level (63.6.m)...e. the earliest Miocene to late Miocene (Li & McGowran 1995). The younger oecurrence of P kalimmensis was veported by Quilty (1985) from the Pliocene in Flinders tsland, Bass Strail, Parredivte povifera (Parr), On the olber hand, seems to be a Quaternary species. On the Lacepede Shelf of South Australia, 2 poriferda occurs. frequently between 50 m and 200 m, and some specimens grow up to about 15 nin (height) x Tam (width), with over 15 chambers in the final whorl (Li er al. 1995). Quilty (980) sugpested that Crespinella parrt was the probable ancestor of both C. umbenifera and kalimnensis. His view is upheld here. The evolution of this lineage might have begun from C. puri in the later Oligocene. but the radiation of both Crespinella wrbonifera and Parredieta kalinuensis did not oceur until the carly Miocene. This was probably implemented by vy mearphological change Lron low trochosprral ty planispiral (Co purr? > C. umbonifera) and from keeled to weakly keeled or non-keeled (AY, parri > P. kalimnensis), The loss of the umbilical Hing (boss) alse took place in the early Miocene and subsequently became a diagnostic feature in younger specimens of P kalimnensis and, particularly, in the much younger 2 purifera (Fig. 7). Divirifution Southern Australia, carly Miocene to Recent Acknowledgments Amanda Beecroft compiled and scunned most of the Eocene species, J, M, Lindsay generously allowed the use of his unpublished data and scanning micrographs, and he.and P. G. Quilty read an carly draft and shared with as their Knowledge on the distribution of many species. The manuscript was reviewed by PG. Quilty and S$. A, Revet, whase econments are acknowledged We are indebted to S. Shafik Jor the Lakes Entrance Liu D&B MecGOWwRAN material aid ta Y. Bone for dredged samples fram the Lacepede Shell. G Trevelyan did thin seetuons and AL Shubber took several optical photographs BL McHelry arranged the loan af type material from the Scvath Australian Museum, J, Terletassisted in scaring the Uncoated Lype specimens, R. Barrett reproduced Figs. 47. This work was supported by an Austrafian Researeh Couneil crane. References AbaMs, CoG, 11992) Large fonmminitéra und the dating ol Neogene events pp. 221-235 InVsuchi, R. de Ingle 40, Jr. (Eds) “Pacific Neogene Environment, Evolution, and Events” (Univ, Tokya Press, Tokyo). Brecares. Wo A. Kent, DOV & Fryyn, 1d 98Say Paleogene geochronology and chmonostrangrphy pp, 41-195 fi Sretling, No (BdL) “The Chronology of the Geologie Recon! Geel. Sec, London Men. 10. : , & VAN Conviring. J. A. H88Sby Neawere Beochronology and chronostradgruphy pp. 2-260) fe Snelling. No J. (Ed. “The Chronology of che Geological Record” That Caspr, 8. Coo& Kent, D. (1992) A new geomagnene polirily Gime scale for ite Late Cretaceous and Cenozoic. Jone Geopliyss. Rey. 97, (3917-1395) Canrie. A. N. 1958) Tertiary forariiiiira tor the Aire district, Vieloria, Ball Geol Snes Bet 55, 1-76, _—____ (1964) Tertiary. (omminifers from Gippstind, Viera, and their stravigraphical sinifivance Men) Geel Surv. Vier 23, 1-154, ply 1-17, CHABRONIPRI, GC, A. (984) Qligncene and Muocene larger Foraminiterda rom Australia and New Zealand, BMR Geol, Gvophys, Bull, 188, 198, pls |-20. Cooper, B. J. (1979) Eocene ty Miavene sualigraphy of the Willunga Emboyment. Rept, Investigations. Geol. Surv 8 Alan 30. Faivey, DOA, & Mtrier, fC. (IORI) Regional ple lectomes and the evolurion of Australia’s passive continental margin. BMR June Aust Geal, Geapliyy. 6, b24. Frakes, L.A. McGoWRAN, B & Bowler, J. M (1087) Evolution of Australian environments pp, -16 /n Dyne, G, R, & Walton, DW. (Eds) “Fauna of Australia, General Aructes’, Vol JA (Australian Government Publishing Service, Canberra), Cinsow. TG. (1987) Morphologie changes and smugrahonal History of the genus Thulogenerind, Jone Foraminiferal Res. V7 227-2549, —.. (1989) Miocene eyulution of Tibilogenerina. in ime Indo-Pacitic and African areas, Maid. 19, 126-143. _ RARRIN. V, Potinany, A, & S/TRAKOS, K. (YI) Early evolutian of Tkbulogenerind during the Paleogene Of Burope, Ibid, 26, 299.312, GLAESSNER. M. F (1950) Walter Juries Pare dasi/. Jane Sey, 2, 2i1, & Warr, M, (1959) Revision of ihe foraminiferal family Vietoriellidae, Micrepaleoniuliey $, 199-212 Heath ROS, & MeGowtan. B, (984) Neogene datuin phines: lonuminilend successions in Austialia with reference sections from the Ninety-cast Ridge and the Online lava, Plateau pp. 187-192 In Ikebe, N& Tsuchj, R, (Ede) "Pacilie Neogene Datum Planes: Contribulions tu Brastratigtaphy and Chronology” (Uniy, Tokyo Press. Tokye)- Hormiprook. N, de B. Beawiek, RoC. & StRONG, C, P. (1949) Manual of New Zealand Permian to Pleistocene Foruminiteral Biostratigraphy, V7. Geol Surv Patent Bull, S6, (175, Howonin, Wo IRR9) The foraminifera of (he olter lectiny of Austria (No. Ll, Muddy Creek, Victorias. Viana A Soe. 8, Ause 12, 1-20, ———— (189]) The finaminifera of the older Terdury of Australia (No. 2, Kent Town Bore, Adeline). fd. 1, 350 350, br, QO & MoGowras, B 1995) Benthie formatter response 10 the Mincene osediation: on de nid-latituele: inarein) takes Bnirimec, southowstern Austeata Micropalennioloe’ Aa, vin press). <4 . Jamis. NP. Bont, Yo& Carre, JH (99S) Mixed forsminitéral Poficies an the Mesotrophic, WWid-latilude Lacepede Shelf, South Australia, Pulalos Wh Hin press), Liwnsay, 1M. (1969) Catnazoic foraminifera ane sinfierapby of (he Adelaide Plains Sub-Basin, Sonilt Austria, Ball, Geol Sam: & dur 42, 1-60. —— CYKS) Aspects nl South Australian Tertuary fominieileral biostratigripliy, with emphasis on suidies ot Mussina and Subhorina, SA. Dept Mines & Enerey, Spec. Pub S W721 ~—— & Gites. S. DB. 1973) Notes on the Lepidoeyel ine yone in Morgan Limestone ulvne ie Murray River, South Australia Geol, Sure 5. ust, Quart, Geol, Notey 45, | 7. CoLeuien, AL RL, di a TAPPAN, HL. (1987) “Foraminueral Genera and Their Classification”, pp. 1-970, pls. 847 (Vin, Nostrand Reinhold Company. New York), LuUpRROUR, N. He (1961) Struigriphy of (he Murray Basin in South Australia. Bull. Geel, Sur So Aust 30. 146s McGowan, B (1979) The Australian Tertiary: Jomminifem! overview, Mar Mierupal, &, 235-264, 980) Pitty sali -yeurs ago, dmmertcan Seatac 78, 40-99. (91) Maastricht and early Carnozore, southern Austeilia’ forminiltral biostruigraphy, Geel, Soe. Aust, Spec Pulz, 18, 79-98; & Beeckoby. A. (YRS Guembeliiria in Whe early Terhary af senifhern Australia and ils paleocednographic Signifeunce. S. 4, Dep Mines & Brera, Spee Pub 3. 137-261 & (1986) Nerire southern excratrpical foraininiterd ane the Terminal Bavene Event. Ailieoreasr, Faluvoclinaial . Palaeweeal 59, 234, & hy, 01993) Mincene planktonic foraminifera from Lakes Entries tn Gippsland: Midlatinde nerite signal Troy tranisionming Odean. Mei. as.. Auserilas, Palaennials 395-405, & (19494) The Mineene oseillatien in-southurn Australie Australain Vertebrate Evolution. Palaeontotagy und Systemes Ree So ayn Mae P7172, _ & — (1995) Mineene climatic oscillations: reuppraisal af the planktonic foramimierul record trom the Lakes Entrance of) shatt in Gippsland, southern Australi Mieropaleontilogy. AL, (in press). . Linnsay. SM. & Hanns, WOK, HY7b) Artcnpted teconciliadon of Tertiary biostratigraphic systems, OlWay Busin pp. 273-281 Ja Wopfiier, A. & Douglas, 1G. (Eds) “The Otway Basin in Southewstern Australia’, Geol, Sarvs, So ayl A vied Spee, Bull _. Mass. G. & Beecrorr, AL (1992) Late Hovene and early Ohwouene iesoulkern Austrilias local neritic signals of vlobal oceanic changes pp. 178-201 de Prothero, D. Rk. & Berggren, W, A, (Rds) “Kocene-Clivocene Climatic aml Biotic Kvolunun” (Piinceon Univ Press, Princetant, Pare, W I (1942) New genera of foriminifera trom the Tertiary of Victoria Min, Geol, Joun 2, 361-363 SOUTHERN AUSTRALIAN FORAMINIFERA —. (1950) Foruminifera pp. 237-392 Jn Johnston, T) H, (Fd_) "Reports BALN.Z. Antarctic Research Expedition 1929-193)", Series B (Zoology and Botany), 5, pls. IIENY (The Hassell Press. Adelaide), Query, P. G, (1974) Tasmanian Tertiary foraminifera, Part 1, Textulartina, Miltulina, Nedosariina, Pap, Proe, R. See; Kasmi. W8, 31-106, (1977) Tasmanian ‘Tertlary foranimifera. Part 2. chiefly Spirillinaeca to Glabratellidae. /bid, LL, 69-109, (1980) New. rotalid toraminiferids trom the Oligo- Miovene of Tasmania, Afeherime 4, 299-31b — (1981) Late Hovene benthic Foraminiferida, south coast, Western Australia. dawn A. Seo WE dies, G4, (3) 79-10). (982) Tasmanian Tertiary foraminifers, Part 3, Discorbuces (Eponididae) to Nonionwwea. Pap, Proc R. Sue Tasrn WO, 5-52, Appendix 1. A list of Almaena Samoylova A. gippslundicn Carter Amplusteginag dOrbigny Asterigerinella Bandy A. gallawayi Bundy A. adelaidensis (= Truncatilina marsaritifera vat, udelaidensis Howchin) Bronnimannia Bermudes B palmerae (= Discorbis pulmerae Bermiidez) BK hatiotiys (=Disearbis haliois Heron-Allen & Harland) Cifellia Gibson C. cast (= Chrysalidina costar Heron-Allen & Eurland) Cihicides de Monttort C. thungia Finlay C. medtoerty Finlay Cc. refulgens de Montlort C. vertex Dorreen Cihicidoides Thalinann C_ neoperforas Hornibroak C_ perforatus (=Roralie perforata Karrer) C. pseudounvertanus =Trauncarulina peeudoungerianus Cushman) C. karreriformis Hornibrook Crespinella Parr C. parri Quilty Co umbenifer (=? Operculina umbeanifera Howchin & Parr) Crespinina Wade C. Kingscotensis, Wade Discerotalia Hornibrook Eycornebovina Butt E. euvillieri (= Retalia cuvilliert Porgnant) Halkyardia Heron-Allen & Earland HA. bartrumi Part Reteralepa Franzenau H. brevoralis (=Cibicides brevoralis Carter) H, opatca (=Cihicides opacns Carter) H. subhaidinzert (—€. subhaidingen Parr) (1985) A Phocene foramin(lerid fauna from blinders Island, Bass Strait. /hid, 119, 89-91, — (1993) Tasmantid and Lorn! Howe seamounts: bioswatigraphy and paltcoceanographic significance Alcheringa V7, 27-53. Rivets. 8. A. (199]) The generic revision of the Reussellids (Foraminiferida). Jour Micropalacaniol, 10. }-15. (1993) The fonuminiferal toothplate, a review, /bud. 12. 195-168, Wane. M (1955) A new genus of the Chapmaniminac from southern Australia, Conte, Cushinun Bound. Foram, Res, 6, 45-44, (1957) Morphology and kixenomy of the foraminuteral family Elphidiidite. Jew: Washington Acad, Sei. 47, 330-339 genera and species H, victoriensis (=Cibieides \Vietoriensis. Chapman, Parr & Collins) Hofkerina Chapman & Parr H, sentiornara (= Pulvinilina semiornata Howehin) Lepidoecvelina Giinbel L. howehini Chapman & Crespin Linderina Schlumberger L. glaeysneri Quilty Maslinella Glaessner & Wade M. chapmani Glaessner & Wade Nororatalia Finlay Operculing COrbigny Parredictat Li & MeGowran P. kalimnensis (=Planulina kalimnensis Parr) P. porifera (= bilvudineria porifera Parr) Parrellina Thalmaun FP. craticulatifurmis Wade P. crespinae (= Elphidium crespinue Cushman) P. imperairix (=Polystomella iimperatrix Brady) P. verriculata (= Polystamella vericulata Brady) Siphominaides Cushman §. laevigata (= Truncatulina echinate var, laevigata Howchin) Siphouvigerina Parr S. fimbriata (= Uvigerina perrecta var. fimbriata Sidebottam) Jubnilogenerina Cushman T, ferox (=Bigenerina ferox Heron-Allen & Earland) Lo mooraboolensis Cushman Vicloriella Chapman & Crespin Ko conoidea (= Curpenteria vanovidea Rutten) Mictoriella plecte (=Carpenteria proteiformiy var. plecte Chapman) Wadella Srinwasan Meo heamiltonensiy (= Carpenteria Glaessner & Wade) Ke globifarmis (= Carpenteria globifarmis Chapman) hamilionensts THE POPULATION BIOLOGY OF THE TEMPERATE REEF FISH CHEILODACTYLUS NIGRIPES IN AN ARTIFICIAL REEF ENVIRONMENT By MICHAEL CAPPO* Summary Cappo, M. (1995) The population biology of the temperate reef fish Cheilodactylus nigripes in an artificial reef environment Trans. R. Soc. S. Aust. 119(3), 113-122, 30 November, 1995. Underwater surveys and observations of tagged fish were used to examine spatial distribution, temporal variation in abundance, habitat use and agonistic behaviour of a small population of Magpie Perch, Cheilodactylus nigripes, in a 1176 m* site beneath a pier over two winters. A marked decline in numbers of small fish in the population was observed in one year and the number of larger fish was more stable. The unstratified density of fish was between 1.6 and 3.4 fish 100 m* but locations of fish sightings were strongly positively correlated with two dimensional cover of hard substrata within the site. Cheilodactylus nigripes was a diurnally active micro- carnivore which used hard substrata for shelter and for feeding on benthic invertebrates. Movement patterns were measured or inferred from spatial patterns of distribution and were found to be restricted to small areas within the confines of the pier. Home range was estimated to be 26 m’ for one juvenile fish. Only juveniles < = 12 cm TL defended space aggessively against intrusion by conspecifics and fish >19 cm TL engaged in lateral displays with colour changes in agonistic encounters. These displays were considered to be related to maintenance of spatial patterns. Key Words: temperate reef fish, habitat use, agonistic behaviour, feeding, Cheilodactylus nigripes. Transeetionsuf the Rav Soeers ofS. Aust (985). EG, 1-2 THE POPULATION BIGLOGY OF THE TEMPERATE REEF FISH CHEILODACTYLUS NIGRIPES UN AN ARTIFICIAL REEF ENVIRONMENT by MICHAEL CAPPOF Summary Capen. M1. (1995), The population. biology of the teniperate reef fish Cheiledacius migeipes in an artificial reel envimanment Trans. Ro Sac, & Aur W984), 13-122, 30 November, 1995. Underwater Surveys and observations ol fagged {ish were used to examine spatial distribution, temporal variilion imabundance, habit use and agenisne behaviour of a Srull populition of Magpie Pereh, Chedtedecivtay nigripes, inv 176 m site beneath o pier over wwe winters: A marked decline in numbers of small tish in the population was observed in one year anu the number of larger fish was mere stable. The unstratified! density OF Tish) was berween [fr and 3.4 fish (00m? buf locations. of fish sightings were strongly positively correlated with Wwo- dimensional cover ob hard substrata within the site. Che/ldacilis nigripes was a diurna lly acuive mncro-curnivore Which usect hurd Substrata for shelter and for feeding om benthic invertehrates Movement patterns were measured or inferved tron: spatial patterns of disiribulion and were found to be pescicted to smal] areas within the confines ofthe pier, Home range! was estimated ta be 2h i) far ane juvenile fish, Only juveniles <= cm TL detended spave aggressively against inuslon by conspecifics and fish > 19 em TL engaged in lateral cisplays wilh colour chutes (aivonistic encounters. These displays were considercd to he related ty majitendnee of spithal patlenns. Replenishment of he pier population (as observed wy oceur in spring fran eeccuininent of Sem TL juveniles Vhese duta indicat’ the importance of refulively small marine protected dreds as refuges Tron spearfishing for CO. niripes. Key Worbs terapetate tect fish, habitat use, agonistic behawinur, feeding, Cheilodacr ius nigrines. Introduction Cheilodactyld fishes are a numerically important component of the cool temperate reel fish funas io Australia, New Zealand, South Africa, South America and Jupan (Lincoln Sinith ef ad, W989) Branden er al. 1986. Loum & Chout 980; van der Bist 981; Nielsen 1963; Sano & Moyer 1985). They are relatively linge, slow moving and can be casily approached underwater. making therm very popular targets for spearlishers, La south-eastern Australia (hey doniinate the catehes made in Spearfishing competitions Johnson 18S, Lincoln Smith ef al. J989) and there is evidence thitt spedrfishing is a major cause of lovalised depletion al chetlodactyhds in New Zealand (Cole et al 1990). Assessment of effects of recreational speattishing on cheilodactylid populations requires a knowledge of habitat use by the fish, (heir pallerns of movement and abundance und the sources of populuton replemshient, These data are essential for the inplementation of marine protected areas (Edy vane 993) at the proper sparial scale as uo means of managing temperate rect fisheries, A knowledge ot feeding habits is ulso desirable to determine (he role of cheilodaetylids ut trophie dynamics and fo predict r Australian Instijute af Marine Science, PMB 3. lownsville MC, Old. 4810 any Subsequent Chunges in benthic community structure which may result fram the effeets oF high fishing inortality. Their behavioural traits make these fish ideal subjects for underwater studies yet little is known of Ihe patterns of distribution und ecology of temperite vheiloductylids, especially those that inhabit the coastal reefs of southern Australia. Cheilodactylus tugripes is an abundant inhabitant ot shallow (<30 metres) limestone and basaltic reefs in southern New South Wales, Victona, Tasnkinia, South Australia and southern Western Australia (Hutchins & Swainston 1986). It is commonly Jound m assoetation wilh the hard subs(rata provided by artificial tyre reels. ship wrecks and piers. This species attains 4) em in length and rarely takes a baited hook although i comprises a major portion of the spearfishers’ catch in South Australia (Johnson 1985a). Limits to the speared catch are enforced by competition bag limits and legishited closure of marine reserves und all piers to spearfishing, With the exception of counts of C. muripes in surveys of reel faunas in the Great Australian Bight (Kuiter 1983; Branden er a/. 1986), and frequency in catches ul Spearfishing competitions (lohayon 185a-b). there have been no studies of the ecology of this species Mm Australia. The present study documents the patterns of spatial und temporal abundance, habitual use and agonistic behaviour in a protected populiion of Co migrpes Th below a South Australian pier. Specific aims of (he study Were to! (1) map the spatial and temporal patterns of abundance ob the population in relauou i ihe wricrohabitats provided by the pier, (2) deseribe the feeding morphulagy and diet of the species and (3) deseribe fhe agonistic behaviour amongst individuals and its importance in the maintenance of spatial patterns. Methods The study site Edithburgh pier is located on the woslern side of Gult St Vincent, South Australia at 35.5°S, 137.45°R (Fig. 1), The seabed below the pier sloped seaward tou maxinun depth ol approximately 4.5 m at low tide und consisted of a base of coarse sands, gravels und shell, The predominant hard substrata were artificially placed, inthe form of fallen pier niaterials, discarded ship's ballast and debris and w limestone shelf produced “T T atl) he Js 138 As | rae } \ | SS VaR Spencer Sul Gulf Vincent Adelaide Edithburgh Kangaroo Island — 35 N t o kin) tDQ ) N B. f outer half of pier pt oO) zo Fig, 1. Location and aspect of Edithburgh pier in Gulf St Vincent, South Australia, M CATO by dredyiny along the northern side. Large expanses Of seagrasses Surrounded (he structure. The configuration of pylons and major features vt the hard substrate below the pier were mapped (Fig. 2). The entire picr was 168 metres long bounded hy 53 rows of timber pylons in 4-5 columns, The study site Was under the outer half of the pier seaward of row 23 (Fig, 1). Tt ecomprised un area of 1176 my and was divided into 120 quadrats from 9-I] m° in area with relerence to the grid of pylon rows and colunms (Fig. 2). Topogtaphic complexity (Leum & Choat 980) was described for the site by estimating the (wo dimensional “cover” of hard substrata within eavh quadrat, using three ordinal categories of topographic complexity. A total of ROS min Sl quadrats was “simple” (cover =10%), 225 m° in 24 quadrats had “middle” level of cover (cover = 11-25%) and 147m in 15 quadrats were classified us “complex” (cover’> 25%). The seaward end and the southern side of the pier had the most hard substrata, in the form of boulders, slabs and blocks thal were usually less (han 0.8 m high, Disrrihurion Seven visual censuses of the population were nade during March-September 1980 using SCUBA, On each census (he whole habilal was searched und the positiun of each fish recorded. The total length (TL) of euch fish was estimated lo the nearest centimetre and a pote was thade if the fish were feeding wheo sighted, Fish were approached to within obe metre or less and length estimates Were frequently made in direct comparison to a 40 em plastic ruler, Four censuses of the size frequency under the pier were made by another diver in April-July 1981 and fish lengths were estimated to the nearest centimetre (K, Wehr unpubl. data), In analyses fish lengths (TL) were categorised as: R recruits (<6 em); Cl (6-12 em); C2 (13-19 em), C3 (20-26 om), C4 (>26 em). The choice of distinction between Cl and C2 was made to separate youny-ol* the-year from older fish. These were biologically meaning tul divisions Of the population, a8 colour and morphological differences occurred amongst them, The usual colour pattern of Co nferipes was three broad, verticul, dark bands on the white background of the body and caudal peduncle and a dark cheek stripe through the eye (Figs 3a, 4a), Recruits had a bright orange caudal fin and pale orange caudal peduncle with black tips on the catidal lobes. The mouth was more terminal than inferior and angled slightly apward giving these lish u shorter snout and deeper chin than larger fish, Size class Cl fish had the same caudal fin colouration but this faded with size to dusky or reddish in larger classes. There were protuberant crests on the preorbital bane of the two largest size classes which were not observed an sinaller fish, POPULATION BIOLOGY OF MAGPIE PERCH 115 Fiz. 3 Summary of colour changes displayed by large C nigripes. a. Normal banding pattern. b. Transitional banding pattern, c, Modified banding pattern - note dark midriff band and caudal pedunele, loss of posterior and cheek bands, white iris and white demarcation between dark bands. Feeding habits To avoid sacrificing the small population in the study site a sample of 21 Magpie Perch (TL 21-36 cm) was speared on 25 September 1980 under Port Giles pier 12 km NNE of Edithburgh pier (Fig. 1). Immediately afer capture the alimentary tracts were removed and preserved. The food bolus in each tract was dispersed evenly in water and taxa in four field views (7) were identified as far as possible, The yolume of each taxon in the pooled contents of the alimentary {racts was estimated by a water displacement method, Observation of mavement and beluaviour Underwater behavioural observations were made 1 60-minute periods at dawn, midday, dusk and midnight on five days during August-September 1980 from a single vantage point amongst limestone slabs at the seaward end of the pier in pylon row 53 (Fig. 2). Fig. 2. Comparison of pooled sightings of C. nigripes in 1980 and.an index of topographic complexity in quadrats (right) with the mapped study site (left), Pylon rows are numbered in ascending order seaward from row 23. “Simple” cover of hard substrata< =10% (no shading): “middle” cover 11-25% (mild shading); “complex” cover >25% (dark shading) 110 M. CAPPO q i 3 ] _ _ 1 _— = a ‘ . _ _ 2 7 : ] —. _. _ _ - is | ] Fig. 4. Photographs of colour change and Jateral displays. a. Normal banding pattern. b, Loss of posterior dark band. e. Transitional banding pattern, d, Circling and leaning displays. ¢. Anti-parallel orientation and modified banding. f Break in encounter and transitional banding. POPULATION BIOLOGY OF MAGPIE PERCH Wy Observations began belore sunrise and finished alter sunset lo (nelude crepuscular periods of activity. The movernents of four fish at this location were monitored simultaneously during the dawn observation periods. Each fish was recognised individually by tags or body markings. Further observations were made at a variety of locations on an opportunistic basis during 1981-1987. Data on the movement patterns of tagged individuals were collected during 1980, Seventeen fish were tagged with “T bar" anchor tags individually coded with coloured paints. Fish were captured with a large hand- et. tagged underwater, measured and released immediately at the site of capture, and a numbered stake was used to mark each release site. During subsequent dives the sightings of tagged fish were recorded on the site map, The larger fish in the population evaded capture and the tagged sample (TL 10.5-23,0 em) did not include C4 fish, Results Temporal changes in abundance and size composition The sightings of fish of the five size classes are shown for euch sampling date in Fig. 5. The mean numbers of total sightings for each census were similar for the two years with 24,85 +4.30 fish sighted in 1980 (95% CL = 19,32-40,37) and 30,504 1.55 fish in 1981 (95% Cl = 25.55-35.44), but seasonal declines. in sivhtings were different. In 1980 sightings declined from a Mareh high of 53 fish to an August low of 18 (Fig. 5). This was partly Tor! Langth C1 c4 326cm 8 Ea ca 20-26cm 50 g (T] G2 19-196 £ oa ‘wo 40 Gl 6-12¢em oo 2 ‘5 30 g E 20 3 c 10 census dale 198) Fig. 5. Changes amongst months in (he number of five size classes of Co onigripes at Edithburgh pier during Underwater visual censuses in 980 ant 198! due to the low visibility (< =0.75 m) encountered during (he August census, The decline was evident tor all size classes from March to April when both Cl and C2 declined by one half. The steady decline in Cl numbers may be partly accounted for by growth into the C2 size class which showed an increase in abun- dance. Recruits were first observed in early September 1980. and increased by late September (Figs 5, 6). Predation may play a role in size-specific mortality as a Southern Calamari squid Sepioteuthiy australis 20 March '80 25 April '80 a | | td June ‘80 os i 44 July '80 {2 August '80 2 Saptember '80 26 Saptembar '80 Number of fish sighted = = = — 4 6 8 10 12 14 16 18 20 22 24 26 26 30 32 M36 Pe | 1 | April ‘81 a ale oll obese f May '81 TIA eee a th. 0) ok bo” duly ‘81 4 6 B 10 12 14 16 18 2 22 24 26 28 30 32 M4 36 Total length (cm) Fiv, 6, Modal progression in estimated total length of C nigripey sighted at Edithburgh pier during WSO and 1981. I) MO CAPRO Was Observed to make several atlaeks On some Hewly recruited juveriles before capturing and consumme one in 1982 This patter was not seen in the 198] winter surveys wthough there was a slight decline in the proportion of class Cl fish from April to July. The tune low. was attributable purtly Wi low visibility (<=0,6 im) encountered during that census. In both years the winter papulation counts were al comparative levels in the study site, Changes in abundance were relatively sinall in the two largest size clusses amongst months, and between years (Fiz. 5). However, the sample sizes and numbers of fish were low, precludiag meaning ful stufishicul comparisons. Caution is therefore used i the interpretiGen of these data, Modal progression is evident in Fig. 6 for the smuller lish from. 1980 to 198] showing maximum growth rates of ybout 0.46 em month! for Cl fish and about 0.77 cm month’ for recruits. These are in general agreement with the measured rate of 0.46 cm month’ growth of a tagged fish from 13.7 em to 15.0 em over five months in 1980. However, the aveuraey und precision of length estimation Were noldetermined lor either observer so further estimation of growth from modal progression was not possible. Spatial distribution The overall density of fish in the study area was estimated al J.o4-3,.43 fish per 100 min 1980 using 95% confidence limits for the mean number of Jish sighted divided by the area surveyed. Localised densities were much higher and (he quinber of © figripes sighted in quadrats under the pier wus positively associated (Spearman R=0.5487, P= 0.000) with the amount of bard substrtta within quadrats using Spearnian’s rank correlation procedure (Zar J9R4)_ Fiflzen “complex” quadrats. comprised only 2.5% of the study aren but gave 5596 of Sighlings compared to 30% in “middle” quadrats and only 15% in the remaining 68% of the study area defined as “simple” (Pig. 2). Nearly 18%, ofall sightings were made in one quadrat on the scaward end of the prer where large limestone slabs and other hard substrata occurred These slabs. were the highest (0.6 - 1.2 m) afd aiost rugose hard substrata in the study area and provided extensive sheller (Fig. 2), Fish were rarely sighted in quadrats with 10% or less of bard substrai. with the exception of quadrats contaming tall three-dimensional structdres not well described by the two-dimensional method used to classify quadrats. For example, clusters of buffer pylons.on the seaward corners ol the pier were’ used together by enerusuing organisms and provided shelter siles, There was no clear relationship between water depth and fish size due to the shallow nature uf the habitat and the overriding infloence of three-dimensioral reel structure. However, the largest fish were generally restricted to the deepest Water and were nor seen in the area siloreward ol pier row St (Fig. 2) New recruils were seen thronghput the study wea. hut mainly dear the seu xurhive ont pylons at (he deepest seaward end, amongst arhoreseent bryozoans, Movement of fish in the smely area Qbservuuions throughout the day and in jhe night showed that individual Co nivrineys emerged Frew shelter sites just after daw and retreated to the sante sites during sunset. No fish were seen io he aetive al night. In five might dives ove of the largest fish ithe population were seen in the same sheler sie in a crevice beLween close pylons. resting morianless un vutstreiched pectoral fins with raised spinous dorsal fins. ILappesared thal largest fish emerged earlier and retreated later than smailer sive oclasses, The movements of three fish Were moyilofed in each of tie five dawn observation periods but the fourth and smallest (CV) sb was only seen for the first three days. The largest Fishy (C4) traversed an uverdge of 6.04 14) metres, a 23.0 om tagged fish moved 78+0.37 metres uid a 14.0 ein tagged fish nreved 2.5 40.55 metres nt the 61 minules afer eniergenee frum their individual shelter sites, The smallest individual moved only OSSIAN metres Only the langest fish moved Jar Irom the confines af the pier foraging amongst dow limestung shelves within JO menes of the seaward end Seven of the mags (41%) were nor seen after applivauion, Inree orbers persisted for unity 9 days. atid only five tips were sighted after Of divs The movement of lane Mayple Perch could fut fe assessed trom the tageing program because lhe mean length it release (ir ihese remaining tive fish was only 15.4 em (C1,€2) and the largest was 23.0 on (C3). Only four tagged fish were present at the end of the 1981) study period and euch of these fish was seen during evers cersusy Mllowing lagging. The loss of lags from the population was at least panty due to tag shedding On two oceasions fish were observed to ¥eruh the tag ol! agains! hard abjects, The sightings of all tagged tish were restricted fe the habytat underneath the pler within Timeines of the tagging site wilh the exceplion of We individuals. Several Weeks after Lage ing these two sly were fotind to be resident for |he remainder of Ihe stutly in Quads about 10 metoes away fon the tagping site. Tagged Nish were never seen to traverse thie seagrass beds on the siuthern Sidé of the pier which appedres! tu det as natural boundaries to the pier habirat Wilson (1975) defined “home range” to be the ares that an aairial lems thoroughly and habitually patrats, and “core red” to be the area of Ieaviest regular use within the home range Only a single tipged Tish was seen frequently enough to confidently estimate these Iwo areas. This smull C2 fish (13.7 cm) was sighted 18 umes Wethin 7 metres of the tagging site. The hame range of this fish was estimated to be only 26 ny hy Measuring the area ota polygon joining fhe outermost POPULATION BIOLOGY OF MAGEE PERCIT 1h% sighungs (Leum & Choar I980). Similarly, the corm fired Was estinwied at 17 nv ja be the area encompassing 50% of sightings. The focus of this area was a sidall shelter Site within a cavern amongst limestone blacks on the southern side of the pier in pylon row number 28 (Fig, 2). This fish was only sighted once outside the shaded confines of the pir and the furthest displacements were northward under the pier and westward along the pylons. Feeding morpholaay and diet Cheilodacrylus Wenpes bad a small moulli wilh tick fleshy lips anda siogle row of widely spaged peg-lpke teeth on the centary and premuxiilary. In the throat there were a paje of upper, and a single lower pharyngeal (ooth pauls covered with bands of villiform vecth. ‘There were IS tong and fine gillrakers on the Hirst will arch and the stomach was small with a large pylone region containing five short pyloric caeen, The fish were ohserved to inspeet closely poekets al sediment fa erevices or amongst fronds ol arborescent arganisms and fed jn a pecking mation The mouth was rapidly opened. lorming a suction with (he fleshy lips, and benthos was ingested with an audible clicking sound, By rapidly openime und closing jhe opercula, Fine silt way strained oul rhroaugh the gill chambers and larger particles were ejected froyn the moudt. Large polychactes were wrenched trom the substratum und vigorously shaken to break thent up inky pieces suitable for swallowing. When elose observations were thade, no evidence of prey was seen near the lish and they appeayed to select fecding substrata, but not the henthic organisms within, although fish directed repeated feeding “peeks” at large prey such us polychaetes once they were detected. The fish fed mainly on benthic invertebrate jaune wilh gammarid umphipeds about 4+ mm long predominuat in the pooled sample, Ol the 60'% of tood volume identitable, the fourteen major taxa were ranked as: gammuarids (25.6%): Polychaeta (10%); ostracods (7.5%), Bivalvia (6.6%), Brachyura (3.2% 1: Mysidacea (2.7%); and. Tammidacea, Arehaengastriy- pada. Polyplacaphora. Gastropoda, Isopoda, Ophiuroi- dea, and caprellid and tubiculous ampiipods (each Jess than 1%}. There did not appear to be any size-related diflerence in the feeding behaviour of C vigrpes with the except ion that small fish were observed to feed mure ofen than larger ones, OF all sightings made in |981 the Jollawing propertions were engaged in feeding when sighted: R79: Cl 76%; C2 79%, CI 49% C4 ASG Awonistic hehavianr Wilson (1975) defined agonistic behaviout to be any activiw related to fighting, whether ugeression or caneihation and retreal. Agonistie behaviour in © nigripes Was difeeted toward only conspectics of a similar Size and inree sizesspeciiic patterns were deseribed for the smallest and largest size classes. The grallest fish (< = 12 cm), elassified as recruits (R) ant Cl. aggressively defended space. Most common was a Short pursuit of incoming Ror! fish away trom a foraging area Also observed was the head-on spprouch of Cl fish te within about 4 em followed by sustained pursuit, darting in small circles with the dorsal fin fully raised. Biting was evident as audible sounds and tai! damage from broken caudal rays. This was termed “carausel fighting” by Chiszur (1978). Three stow lateral display palterns were observed amonest fish more than about (9 cm long (C3, C4) which did not involve such pursuit. ‘Uhe first involved pairs of 3 {ish which approached each other head- onand met head-to-tail in a parallel orientation, often within centimetres, and one or bath fish tilted upward slowly. The two fish then swam in slow circles in close proximity with seme lateral displays. but ne colour chanye, belore parting, At claburation of this theme occurred during (he approach of the large C4 fish when a characteristic colour Change occurred. The posterior band began to pale und the white midriff darkened with the development of a sharp white line between them (Pigs 3b.c, ded). When the fish met they assumed un anti- parallel orientation separated hy nly 4 few centimetres for abyut IS seconds, during which they slowly tilted 30 degrees (oa head-up, tail-down position (Fig. de) The posterior band blanched and the darkening of the midnlf and caudal peduncle deepened, Other colour changes were also striking. including blanching of the anal fin, eheck band, iris and peeroral fins. and definilion of a biack patch around the pectoral axil (Fig. 3h.c). When one fish withdrew the other followed and both began laceral “leaning displays” wilh lowered or raised dorsal spines, described by Chiszar (1978) as defensive posturing (Fig. 4d). The entire sequence lasted one lowe minates and once the fish broke off ihe encounter the midriff band quickly resumed its former white state tml the other bunds were slower to return (Fig. 4e.!). These displays were cleat) identified to occur When fish met and appeared &) relate to the position of the fish within the habilai. although it was not possible io define the home ranges and spatial boundaries involved in eliciting the agonistic behaviour. The sipnificance of a third type of hebaviour was more obscure and involved the loose aggregation of large fish into a slowly circling group above the seabed. Most of the circling fish had a pale posterior band and some of the fish exhibited a leaning display towards others, ‘There was insufficient intormation lo recognise charac teristics of “winners” or “losers” and no atlempt was rade t) identify the sex of the participams. but these lateral displivs were considered to relate to habia use. Ae) M CAPO Discussion The low. variability in-counts of C nivripes dimicnest most months in winters of two yeurs. the very close association bewween fish Sightings. and ropogruphic complexity and the restricled movements of ttsved fish were all evidence fora high degree of site specificity oC. nigripes under the Bdithburgh pier. Ruyose, hard substrara were used as sleeping und sheltering sites and us lecding substrata and the eopeept of a scall home minge may best desaibe the use Ob this Space by C nrertyqes These patterns of habitat ase indicute thal relatively small urtificial habitats such ay piers can aer as Important marine protected areas for this speeics. Murine protected! areas can full] a numberof important finetions in fisheries management including protection of “eriteal habiuty” and provision of areas for stock replenishment, lor fishery-independen| monitoring of stock Moctuations and for resolution of conflict amongst Competitors tor use of manne resources and hubitats. (Edyvane 1993), Planning the spatial scale and habilal composition of marine protected areas tor lemperale reef fishes. requires knowledge of sources of population replenishment, ontogenetic movement patterns, home hinge size, habitat requirements and natural habitat houndaries. Some of these can be inferred for © menpes fromthe simple observations presented here supported by comparison with other detailed studies ol {he eheiloduerylids associated with tees Pisiburion and mavement In translation of the pier observations to natural reef populations of C. nigripes in South Australia i is essential to recognise that major biological features at the habitat have been shown to alfeet the distribution oftenperate reef fish ata variety of spatial scales and (hese patterns have been maintained over lore line scales Jones 988: McCormick l989b), In cis regard the artiticht buture ot the pier habitat us considered to differ from nearby natural rects in two tain ways, Unlike the algal-dominated feels surveyed by Branden etal. (W986), the habitat beneath Edithburzh pier lacked macrophyte cover, perhaps because of shading, There Way thes be more suitable leeding substrata there tor C) nipripes. as Choat & Ayling 987) found that larger curmivoraus feel fishes, including cheilodaety lids, forage preferentially in open reef arcas which support greater densities of their invertebrate prey in cornparison to areas dominated by laminarivn algae. The amount of habit for feeding and refuge is further enhanced under the pier by the presence of the pylons und the Jouling communities that encrust them. Secondly, the density of C. rigripes beneath the pier (L6-3.4 per 100 m’) was six-fold higher than an estimate for unexploited reefs oF the Great Australian Bight (0.2 - 0.6 per WO mr) ealediated from the survey data in Branden eval. (986), The tome range Aves. overnent parieris ard agentistre hehaviewy oti tatu! ceers nay be diflerent sis a Consequence, The microhabliat requirements ot © aieripes. tor Shicller sites and feeding subsimara were nor deseribed wilh the simple habia vlassifieations used here bot co determined by studving assoenujene berween ubundunve lind habirat at syall spatial seates, MoCormick & Choat (987) strilified estimates of density of (he morwong Cheiladaetvis specrbitis in New Zealand. by ten habicar types and depih, un reprinted iverages af 0.25-2.09 fish per WOU nr. with the exeepion of the wpugraphieally complex Yumble boulderhank” habital where the density-wits [S87 Lish pet WOW There is wlsira clear role for ontogenetic movements lung onviranmentil gradients in establishing patterns of chejoddetylid abuindanee amd these should he consmered in selection of reef areas for protection, Atier First necruiuig in surge zones (Leum & Choat VSO) eheiuductylids are known te eve to prigressively deeper parts of the reet habitat as they grow (Sane de Moyer 985; McCormick 1989uch) Although the seagrass beds uround rhe pier appeurec fo wet us habrur boundaries whieh smaller © végripes did’ aot traverse. ik Was NOL pussihle to describe WaMigratiin snd eibiteation of fish with (he simple tech iydes Used in the study, Replenishrent ofthe pier Populishon was observed to occur only through the spring arrival of new reevults. The auttiniy decline in nulnbers of these young-ol the-vear io 1880 wis not abserved in I9K) censuses. und i was nol possible to tesolve the roles af size specific natural rortlitv. lageindueed mortuliry ar counting biases tn the decline af such a sil population. Properly replicated censuses stratified ley identify Individual fish and deleet counting biases ane diurnal and sedsonal diflerences in activity. sould help clarify these temporal changes. The pier map and census data presented here provide a buscline for future surveys Lo examine long-tertiy variations in pallerns of Ubundance oF Co aigripes al Edithburgh, iis possible that Home ranges and movements or natural reefs may be mure extensive amongst ull size Clusses of C. nigripes, und it unknown whether the Wistributions yeported here were more, ar less restvicted diirmg warmer months outside the study period, Leum & Choal (M980) uttribured signifivarit winter declines in oumbers of ©. spectabilis sighted to un extension of home ninge during cooler meanths The estimures of hone range Jor Co aleripes were relarively Strall in comparison to those constructed for C speerabihs by Leuny & Choat (IY80) an nalural reels and Qray depend on fish density aswell as habitat type Juvenile C) syectabiliy hud home ranges =< = 100 m_ which were abuut three times that estimated here for Co nripes ustig the same technique. Sinntirly the jurgest Co vcgnpey were always sighted jp i relanvede POPLLATION BIDLOGY OF MAGPIE PERCH (21 small atea atthe seaward end of tte Eaithbuesh pier but Leum & Chout (980) observed that larger C spectabilis moved large distances and had home ranges up to 50-70 thousaral tm’. Fevding leabits Cheladactvius nigeipes were diumally active, bentine cariivores feeding mainly on panunaridean amphipods and other small benthic invertebnites, Thetr mode of feeding is common ty ther reel-associnted morwongs tn the same genus, Bell (1979) reported (hat C. fuseus and C. speectabilis use ther thick fleshy lips to Wrench and suck aninmils off rhe substratum. mainly polychaeles, brachyutans, amphipods, gastrapaus, and bivalves. Sano & Moyer (1485) reported that the Japanese OC. cebra deeds mainly on epifouna. especially simmardean Jimphipods and decapods, while the syinpattic C. sonarus tends to wike both cpilauna (mainly gammaridean wnphipods, isopods, spanges und decapods) and intiunal polvchuetes. Agunistic beltavieu Tivenile CL agripes <= 12 em'TL were observed (defend space ageressively, but such defence muy bave been cnergelicully uneconomic lor laeer fish >t en TL oceupying lurger home ranges, Lh is proposed here that the lateral displays and colour-change during agonistic encounters amongst larger C. aigripes were related to the muinténance of some Undefined spatial pallernt of overlapping horne ranges, Such patterns were mapped by Lean: & Choat U980) for C speviabils Which directed agonistic behaviour only towards conspecifics of similar sve und only smaller size classes Vigerously defended space outside ol the spawning season. The habitat of C, spertabifis was described as a mosaic of exclusive lertofies occupied by sumaller fish through whieh the larger size classes foraged in larger, overlapping home ranges (Leam & Chout 980) and this may be a uselul model for turure studies of habitat use by ©, pleripes Using Chisear’s (1978) definitions of lateral displays i agonistic behaviour the description of “gariusel fighting” fits well the behaviour of CL nigripes javeniles Whereas the varions colour phases of larger fish cun be interpreted as varying degrees of threat in a typical “colour fight” The anti-parallel orientation adopted during these reciprocal lateral displays is widespread in fishes, and some species wilh long dopsal spines, such as the chaetudontids are reported to vilt the raised spines towards the uther lsh ina defetisive posture culled “rolling” or “leaning” if it approwches to closely (Chiszar 178) The possibihily ol a reproductive basis for same. ngonishc behaviours cannat be discounted for all observations of ©. vjeripes, as McCormick (9898) found that darge male C. apectabilis ipgressively defended territorwes during the spawning season by “rolling” dow) On to Totriders Jind restrained visiting fermales by “ight civeling, pursait and blocking” ot chasing and tail-nipping. Seme of the lamer C nigripes al Edithburgh pier wete observed to have fentacular protuberances on the prearbital hones which have been used in sthdying sexual dimorphism and separating the seacs of CL spectabiliy and C\ fiseus hy external char- acters in visual counts (MeCormick 19893; Schroeder et al. 1994), The high population densities of C. nigripes ac some deeper South Austrilism piers may provide the best chances of clarifying the spatial and sexual significance of agonistic behaviour of C. nigripes through observation aad tiorphometric studies. Fulnte research The resihence ol C. vigripes to spearfishing and recuvery Of depleted populations depend on growth rates and the soufees abd sate of population replenistiment, This study sugeests that widespread mavement of C vignpes amongst habitats is not an Important source of replenishment but further studies aL uppropriate scules are necessary to determine the contributions of recruilment and post-recruitment processes in determining spatial patterns of abundance. These data are needed to determine iF marine protected areas should include shoreline surge zones as recruitment sites with corridors of hard substrata linking them to adjacent deeper reef, or if isolated offshore habitats such ws artificial reets are adequate. The magnitude and frequeney of changes in population structure are likely ta be directly related the longevity of © aigripes as variable recruitment will have least elect and spearfishing the greatest effect on the population size of long lived species. When fishing mortality is absent in such cases age classes accumulate and terapore! consistency in population size may misk jin underlying instability in the age composition (Jones M988) Consequently. future studies of ©. nigripes population dynamics may require uualyses of age camposilions of unexploited populations im Conjunction with recrustment surveys (Doherty & Fowler 1994) and monitoring of the survival, growth and movement of individually recognisable (ist from ume of recruitment in permanent quadrats (Connell & Jones 1991). The results presented here forin a basis for such studies und for longer-term assessment ol temporal consistency in {he patterns of abundance of C_ nigripes in a mapped habilat, Acknowledgments Most of this study was undertaken whilst the author was a BSe(Hons) student in the Department of Zoology, University of Adelaide. A.J. Butler provided supervision, logistic and financial support and use ol the Coobowie Marine Research Station. The author also gratefully acknowledges the contributions of K, Weln for use of unpublished data. C, Proctor and A, 122 M, CAPPO Davis lor assistance with diving and revision of the pier map. and M, Keough, K. Branden and K. Handley for photography. The comments of M. McCormick, 1. Suthers, D. Booth and especially lwo anonymous reviewers greatly improved earlier drafts of the manuseript, References Brut, J. D. (1979) Observations on the diet of Red Morwong, Cheilodactylus fuscus Castelnau (Pisces Cheilodacty lidue), dase. J) Mar Fresinw. Res, 30, 129-133. Branpen. K. Lo Enoar, G, J. & Siepien, 8, A. (986) Rect fish populations of the Investigator Group, South Australia} a Comparison of wo census methods, Tran, R. Soe, S. Aust. W0(2), 69-76. CuHiszan, D. (1978) Lateral displays in the lower veriebrates : forms, functions, and origins p. 320 /n Reese. BE. 8. & Lighter, F. J. (Eds) “Contrasts in behaviour: Adaptations in the aquatic and terrestrial environments.” (Wiley and Sons, New York). Cuoar, J, H. & AveiG, A.M, (1987) The relationship between habitat structure and fish faunas on New Zealand reets. J) Exp. Mar Bial. Keel, WO, 257-284, Core, R.G., AYLING, T.M, & Creese. R. G, (1990) Effects of marine reserve protection at Goat Island, northern New Zealand, N.Z. J. Mar. Freshw, Res. 24, 197-210, ConsuLe, S. D. & Jones, G. P. (1991) The influence ot habitat complexity on postreeruitment processes in a temperate reef fish population. . Exp. Mar. Biel. Ecol. 181, 271-294, Doverty. PJ. & Fowrer, A, J, (1994) An empirical test of recruitment limitation in a coral reef fish Serence 263, 935-939. Epyvane. K. S. (1993) An ecosystem-based approach to murine fisheries management pp. 21-27 Jn Hancock, D. A. (Rd.) “Sustainable fisheries through sustaining fish habitat, Australian Society for Fish Biology Workshop. Victor Harbor, SA, 12-13 August, Bureau of Resource Sciences Proceedings.” (Aust. Goyt Publ, Service, Canberra). VAN DFR Etst. R. (1981) “A guide to the common sea fishes of southern Africa.” (C. Strunk Publishers. Cape Town). Hotcuiss. B. & Swamstox, R. (1986) “Sea fishes ol Southern Australia.” (Swainston Publishing, Perth), Jonsson, J. E. (1985a) Spearfishing competitions in South Australia (1983/84). 1. Shore and boat events. Fish Rey Pap. Dep, Fish. (S.Aust ). 12, 1-17. (I985b) Spearfishing competitions in South Australia (1983/84), 2. Australian skindiving convention Thid, 1A, V-15. Joxes, G. P, (1988) Ecology of rocky reef’ fish of north castern New Zealand : it review. NZ... Mar. Fresh Rey, 22. 445-462. Kurrer. R. (1983) An unnotated list of fishes of the Investigator Group, South Australia, Alv/i, Kes, Pap. Dep Fish, (S, Aust.) 7, (-12. Leum, L, L., & Cyoar. J, H. (980) Density und distribution patterns of the temperate murine fish Cheilodacrytis Spectabilis (Cheilodactylidae) ina reef environment. Marine Biology $7, 327-337. LincoLn Samim, M. P, Beir, J, D.. Porntarp, DB. A, & Russent, BoC. (1989) Catch and effort of competition spearfishermen in Southeastern Australi Aiyheries Research 811989), 45-461. McCormick, M. 1, (9894) Reproductive ecology of the temperate reef fish Cheilodeactylus speciahilis (Pisces : Cheilodactyliae). Mar Beal, Proer, Ser, 35, 113-120. _—— (1989b) Spatio-temporal panerns in the sbundance and population structure of a large temperate reef fish. Zbiel. $3, 215-225, & CuHoat. J. H. (1987) Estimating total abundance of 4 large temperate-reef fish using Visual strip-transects, Marine Binlogy 96, 469-478, NIELSEN, LG, (1963) On the development of Cheilodactyluy variegatuy Valenciennes 1833 (Cheilodaetylidae). Copeter 1963, 528-533. SANO, M. & Moyer, J. T. (1985) Bathymetric distribution and feeding habits of two sympatric Cheilodactylid fishes at Miyake-jima, Japan. Jap. J. Ichthyel, 32(2), 239-247, ScurRorper, A.. Lowry, M, & SuTHERS, [. (1994) Sexual dimorphism in the Red Morwong, Cheilodactylus fuscus Aust, J. Mar. Freshw, Res. 45(7), 1173-1180. Wison, E, QO (1975) “Sociobiology. The new synthesis,” (Harvard University Press, Cumbridge, Massachusetts), Zonk, J. H. (984) ° Blostatistical analysis.” (Prentice-Hall, New Jersey) THE MOST VIGOROUS SOUTH AUSTRALIAN TIDE By C. SCHLUTER*, J. A. T. BYE}, & P. HARBISONE Summary Schluter, C., Bye, J. A. T., & Harbison, P. (1995) The most vigorous South Australian tide. Trans. R. Soc. S. Aust. 119(3), 123-132, 30 November, 1995. Harmonic analysis of tidal records for the region between the city of Port Augusta and Yorkey Crossing in the upper Spencer Gulf indicates that the most vigorous South Australian tide probably occurs just north of the Whyalla Railway bridge and has a maximum range of about 4.1 m, just short of being classified as macrotidal. The special property of South Australian tides, that the semi-diurnal constituents (M, and S,) have about equal amplitudes, results in very interesting shallow water tidal interactions, in particular the generation of a large amplitude quarter-diurnal constituent (MS,). The intertidal environment of mangrove forests and especially the samphire flats of the upper Spencer Gulf is shown to be finely tuned to this shallow water tide. Key Words: tides, Spencer Gulf. Transactions of Me Kevul Seciery af) S, Atest (OOS), MICS), 52 THE MOST VIGOROUS SOLTH AUSTRALIAN TIDE by C. SCHLUTER*® J. A. T, Brey, & PL HARBISONT Summary Striloier. Co. Byrd QT. ke Haneison. P1995) The most vigorans South Ausvalian tide, Tras, Ro Soe 4. dist: V3), 123132, 30 November, 1995, Hurmonic amilysis of Gua) records for the region between the cy of Port AuLUSE amd Yorkey Crossing in the upper Spencer Gulf indicates thal fhe frost vigorwus South Austrtham nee probably oecurs just north: of the Whyallt Ruilway bdge and has a maximun mings bf about 1 un asi short of being classified as maerotidal, The special property of South Australian des. thal the semi-diurm! constituents (M, and Ss) have about equal aunplitudes, results in very interesting shallow water Hdal interactions. in particulue the generation of a large amplitude quarter-diurnal constituent (MS,). The interbdal enviroamene of mangrove forests and especially the samphire this of the upper Spencer Gulf isshown w be finely tuned (o (his shallawy water tide. Kry Worns: fides, Spencer Gull Introduction The tides of South Australia have altracted interest for aver 100 years (Chapman 1892; Easton 1970), However there appears 10 be no accaunt of (he region in whieh the lurgest tide oeeurs. This region. is oF interest lo dal theory because both Gulf St Vincent and Spencer Gull have large semi-diurnal (ides at their heads and alsa because the major lunar (M,) and solar ($3) constituents are of similar: magnitude. The diurnal (ide progresses from west to east along the Southern Shelf usa Kelvin wave which enters the South Australian seu where its amplitude and phase inerease regulatly and gradually towards the head of the gults. The semi-diurnul tide. oo the other hand. displays i tnuch iiore energetic response, Tidal characteristics in Gulf St Vincent and Spencer Gulf Ap-almost progressive waye enters Investigator Strait and becomes converted jnto & standing oseilauon within Gull St Vincent (Bye 1976), Bowers & Lennon (1990) investigated the tidal character using the classical model (Bowden 1983) in whieh an incoming waye ts refleeted at the head of the gulf in the presenve of a (rictional force linearly proportional to the tidul current velocity, Particular fitention. was given to the importance of Backstarrs Passage in this process. The + Department of Environmental Bayineerings Cenrre for Water Research. University of Western Australia, Nedlands WA. 6009, t The School of Barth Scyences, The Flinders Universily of South Australia, GPO Box 2100, S.Aust 5001, £ pH Govironment, 26 York Street, Adelaide S Aust. SOO, 1 Scuveter, ©, G, (1994) The Generation of Shallow Water Tides within at Mangrove Enyjronment, MSc Thesis. The Tlinders University of South Austrabia (unpubl, ) system can be best described as a quarter-wave resonance af [he open sea tide, In Spenver Gulf the tidal resonanee is more complex and lies closer tou three-quarter resoounce in which a tidal node oceurs between the head and the mouth of the gulf This behayiour results in a minimum semi- diurnal tidal amplitude near Wallaroo, beyond which there is a rapid inerease in amplitude towards the head of the pulf. Easton (1978) bas given an elegant mathematical demonstration of this resonance which also uses a frictional (er linearly proportional to the Gdal current velocity. Numerical models of the tides of Spencer Gulf bave been developed by Noye ef al. (981) and Bills and Noye (986), including fine resolution models of tidal eddies in upper Spencer Gulf (Noye 1984; Noye e7 al,, 1994), In both gulls mangrove forest und samphire flats are extensive, especially near Port Wakefield, Port Adelaide! (Schluter 1993), Franklin Harbour and Port Augusta. The des are-of great ecological sigmficunce to these areas, The most vigorous tidal system in South Australia occurs between Port Augustit and Yorkey Crossing in upper Spencer Gull This distinctive region has the character of a brine estuary (Bye & Harbison 1990). The action of tidal cureents is responsible for internal mixing of the waler column, Stabilising forces, sucli us Surface heating and horizontal salinity gradients, tend to result ina stratified water column, the lower stratum being denser than the upper, The dynamics Of the mixing process have been extensively studied, inilially in temperature stratified environments such as the shallow European seas (Simpson & Bowers 981). and more recently in the salinity stratified enviroment of the South Australian sea (Samarasinghe 1989). Stratification oceurs when the ratio of the horizontal density gridient multiplied by the depth and {i C SCHULUTER, JA divided by the density and the root mewn square of the slope Uf the wetter surtice die to the bde exceeds a critical valué (Bye 1990), In Upper Spencer Gull the tidal currents ovainian a vertically well mixed water column whereas in mid Spenver Gull a Very detailed ond extensive Observational programme fas shown that transient stratification occurs (Nunes & Lennon 1987), At the mouth af Spencer Gulf the hortzoncal density pradierit Maintains a siranfied exchange for about nine months ot tie year (Bye & Whitehead 1975; Lennon er ad. 1987) In the summer Months, however, Ite horizontal density geadien! 1s reduced due io the reversal of sign in the temperature yradicnt and vertical mixing can also occur ‘These considerations highlight the sigmilicance of the tidal regime jy the dispersion ol dissolved material introduced inte (he wafer column inn he cowstal provinces of upper Speneer Gulf, especially m the rine estuary, The Site Previously, meastutements af the ude in Spencer Gulf extended nly as lar north as Port Augusta. Recent tidal measurements (Bye & Harbison 1987, 1991, 1994) north of Port Augusta (Fig, 1), indicate thata very vigorous tidal regime exists. The site of the investigations was yoold wooden bridge on whieh an abandoned mineral railway Co a salt works crossed Over Spencer Gull) On (he caster shore of the gulf the salt bridge was Originally commected with an embankment which cuts (hrough the mangrove lorest and into the samphire Hats. The maximum span height of 4.4 m is just greater than high water springs, and atlow water springs, the water by confimed lou lew central spans where the max anit depth js about 30 em (Pig. 2a), The piers and spans we ideally sulted for instrument deployments ‘Tide fanges were secured to the piets and Current meters were suspended from the spans or mounted hy poles driver into the ground. On the western side, beyond 4 narrower fringe of mangroves, the bridpe toads directly to the salt works, The tidal observations vndertaken it (his site extended trom 28 February lo 2% March, 1986 (Bye & Harhisen 1987)- Tide gauges und current meters were also deployed for shorter petiods between the Central Australian Railway bridge and Yorkey Crossing. This station lies uppreximately 4 kin further north of the salt bridge and is bordered on both sides by samphire fats (Fig 2b) Which are covered at high waler springs and also Wuring the rare floodings which originate on the Pirie- Torrens plains north of Yorkey Crossing (Bye & Harbison 1994). Since the above observations were of limited rime spar, second mire extensive period was initiated! in 1993, The choice and location of tal equipment was HYF & P HARBISON based upon the pilot investigations. Aa Lnter-Ocean Sd current meter wits located within the town of Port Augusta at un abandoned road bridge using a cradle mooring device, The current meter is based upon an clectvomagnche ax measurement and thus has tea moving parts as do the conventional current roeters This makes the current meter immune to the effect of alval growth, The current meter obtained data between 27 July 1993 and 27 August 1993, A tide gauge located at this sie proved faulty, The tidal constants in Table T were obtained from the Port Augusta power statien tide gauge tot the same period as the salt bridge Jdeploviment. A bottom mounted pressure fide gauge was deployed atthe Whyalla Railway bridge north of Port August between 28 July and 30 October 1993_ Another bottom mhounted tide gauge was deployed at the salt bridge, but this gauge was destroyed by vandals. A botton mounted {ide palige Was deployed at the Central Australian Railway bridge between 18 September anc S| October 1993. This gauge cleurty indicates the unusual tide of the tegion. Tidal Analysis To distinguish benween a progressive und standing wave Situation the phase differences between the tidal corrents and clevations must be kfown. One of the most inportant aspects of tidal systems is the definition ob standing anid progressive waves. With standing waves. the phase difference between the tidal elevations and ciltrents is 90° while for 4 progressive wave the phase difference js O°. A standing wave may be considered as being the sum of two progressive waves, one directed landward and one of equal amplitude directed seaward. The Jandwatd energy Hux associated with the incident wave is, in this case, balanced by the seaward energy flux of the reflected wave and thus there is no pet energy flux, In dissipative situations the reflected wave will be frictionally attenuated and must be smaller in amplilude than the inbound wave and (hus a perfect standing wave is impossible, As the tide propagates from the open ocean. various nonlinear distortions occur in the tidal signal, These distortions ate primarily influenced by nonlinear mechanisms including frictional interactions and interactions with surrounding bathymetric features, as well as atmospheric effects and continuous freshwater discharges. The interactions of the primary astronomical tide discussed above may be represented by the growth of the shallow water tides whieh indicate the degree of nonlinear distartion af the primary signal. In the analysis of tidal signals, it is common practice to decompose the tidal signal into a harmonic series of amplitudes and phases, The total tidal elevation or current may thus be represented by the sum of the MOST VIGOROUS SOUTH AUSTRALIAN TIDE 125 BA Yorkey y Central Australian Crossing wf Railway bridge .*.*.) 1 bottom mounted— tide gauge salt bridge 1 bottom mounted tide gauge samphire salt bush Abandoned /‘.: } *% | mangroves Salt Worksalf 4". road railway Whyalla Railway bridge 1 bottom mounted tide gauge 1 bottom mounted tide gauge 1 InterOcean S4 current meter Fig. 1. Locality map of upper Spencer Gulf. 126 C SCHLUTER, | A. T BYE & P HARRISON Fig. 2. (a). View of the salt bridge looking westward Note un investigator (Pat Harbison) on the fringe of the mangrove forest, and the abandoned salt works in the background, (b), View lnoking southward trom just before Yorkey Crossing Note the samphire flats on (he westem side of the chinnel, and the Cental Australian Railway bridge in the backeround Both views were taken a short time alter low wurer MOST VIGOROUS SOL'TH AUSTRALIAN TIDE 137 Tante ( Vidal constituents for the upper Spencer Gulf. a = amplitude; g = phase in degrees. The tidal elevations are in vim and the currents are m ems. Currents are related 1 the magnetic bearing. The record lengths (sed with the analysis are given in the text. Elevations Currents Constituent Port Augusta Whyalla salt bridge Central Aust Port Augusta Port Augusta (986) Railway (186) Railway (Bast) (North) bridge bridge a a a £ 4 E u z a g a g O, 248 39 165 42 263 SI WO 75 UW.75 322 2.98 337 Ky 483 73 462 74 45) 80 308 90 U.68 | 2.14 4 M; 617 206 oll = 210 559 223 245 227 2.14 140 928 139 in td 257 104 267 AB 275 301 288 2.62 198 10.8 202 3M3S, 4a 24005 5 278 35 «WW G18 #3 O58 109 Meus x 224 1 400 92 30 33 1R9 0.28 218 (15 274 25M, 49 «93 107 78 Ih 12 330 4 52 42 245 ST 35M) 42 165 13 Is M 75 26 ORS O19 347 42 350 MO, v] 91 7 63 ld 175 28 284 003 79 138 172 SO, 3h at 200 Ol 242 Su 356 O45 200 345 219 SK, 3 94 13) 208 28 = 459 76 35% O1K 337 li 276 Mg 2 Sis 3 317 24 ST 50 132 25 59 118 U8 MSy 34 9 12 295 7 U4 1360 38 029 [42 I41 32 Sa 1 302 ta 523 5A 73 78 IRS O45 276 0.54 307 4MS,, 3 229 4 345 7 48 | 337 oos 7I 0,47 39 My Uf) 9g Q 497 5 (29 1 64 O13 355 0.26 349 2MS,, K 40 14 2 27 l49 2 44 O17 WY O46 45 25M, 18 139 Il 93 ad 2 \4 76 26 [68 163 169 Se 2 2K6 9 130 2 245 4 205 013 36 0,39 27 4SM, 4 242 6 35 4+ 408 4+ OW 202 O56 221 predicted harmonic series and (he residual signal, i.c. Results Galt) =] Cyveut) + Cyst) where ¢€om(t) is the observed tidal elevation or current, ¢re(t) is the predicted tidal elevation or current, (ri(t) is the difference between the ohserved and predicted tidal elevation und current, The predicted harmonic amplitude is given by i] Cult) = Lyacos(ort— 2.) =| where a) 1s the amplitude of the ith constituent, o is the frequency of the ith constituent, ¢ is the local time of |he dita and g; is the corresponding, phase lag, The residual signal includes all tidal frequencies which ure nol harmonically analysed as well as atmospheric storm surges. These storm surges usuully persist for 2 Wy 4 days depending, on almospheric conditions, The harmonic analysis of all tidal constituents (Table 1) utilising programs developed by the National Tidal Facility shows the importani aspects of the upper Speneer Gulf vide 2 Only part of py is 4 shallow water conshtuent, joy also is aw minor primary tide, The four major primary constituents (M,, S., Ky, und ©)) are the maim energy source for the region, und the interactions of the dominant semi-diurnal doublet (M, and $5) generate suites of quarter-diurnal (My, MS,, S,), frictional semi-diurnal (3M,85,452, 25M, and 38,M),) and frictional sixth-diurnal (4MS,. M,. 2MS,. 2SM,, 5,. and 48M,) shallow water constituents (Pugh 1987). Terdiurnal (MO,, SO,, and SK,) shallow water Constituents are also generated through inlerachions between the four mayor primary constituents. The MK, constituent, however, is not resolvable from the SO, constituent owing to the short length of our records, The tidal energy spectrum al the Central Australian Railway bridge (Fig, 3) clearly shows energy peaks lor cach of these bands, and also thal this energy is resolved by the harmomic analysis, Two prominent higher frequency bands not presented in Table |, are also shown Table | indicates that the primary constituents have a maximum amplitude between the Whyalla Railway bridge and the salt bridge and also that their phases increase between Port Augusta and the Central Australian Railway bridge. At Port Augusta the tidal currents lead the tidal elevation by about 60°, indicating a northwards propagation of energy. Each group of the shallow water constituents appears to behave differently, although there is large variability between the constituents within the groups. The J2% C_ SCHLUTER, J. A. T. clearest signal is shown by the quarter-diurnal con- stituents, the amplitudes of which ure far greater at the Central Australian Railway bridge. The amplitudes are approximately in the rauo of 1:2.) for niost stations in agreement with theoretical prediction (Gallagher & Munk 1971) and differ by about 180° between Port Augusta and the salt bridge (Table 1), This is consistent with a node occurring near the Whyalla Railway bridge where the quarter-diurnal amplitudes are a maximum. The phase of the terdiurnal constituents also differs by about 180" between Port Augusta and the Central Australian Railway bridge where (he amplitudes are about half of the quarter-diurnal amplitudes, The frictional sixth-diurnal constituents, on the other hand, tend to have inaximum amplitudes at the salt bndye und very variable phases. Finally, the fricuional semi-diurnal constituents show maximum amplitudes al Port Augusta, with the suggestion of secondary maxima at the salt bridge. Discussion The propagation of the primary tidal constituents (and also the frictional semi-diurnal shallow water constituents) into upper Spencer Gulf gives rise to the most vigorous Gide in South Australia which oceurs between the Whyalla Railway bridge and the sall bridge where there is a generation of frictional shallow water 0.01 » 0.001 Spectral Density (m*/cpd) 0.0001 Je-05 0 1 2 3 4 5 . BYE & P, HARBISON tidal encrgy. The position of this maximum tide coincides approximately with the node of the quarter diurnal tide. The maximum tidal range is detined us the suininiawion of the nieun spring semi-diurnal range (MSR) and the mean spring diurnal range (MDR) where MSR = 2(M, + 8,) and MDR = 2 (K, 4 Q,) (Easton 1978). Following the above definition the maximum recorded tidal range for South Australian Waters occurs at the Whyalla Ratlway bridge and corresponds to an clevation of 4.1m, compared to the range at Port Augusta of 3.9 mn. This tidal range identifies the tide as being al the very upper end of the mesotidal range (2.1 - 4.2 m) Friedrichs and Aubrey (1988) have classified estuaries in which the lunar semii-diurnal tide (M,) is dominant over the solar semi-diurnal ide ($2) int) ebb dominant and food dominant, In an ebb dorninant estuary, much wreater idal currents oceur during the ebb following the exposure of mudflats and the release of intertidal storage. To overcame these effects the duration of the ebb lide is far less than the food lide. The consequence of ebb dominance is to provide ii mechanism for the long-term outwelliny of sediments and pollution, In the reverse situation of Mood dominance, the estuary usually does not contain intertidal mudflats and thus the duration of the flood tide is less than the ebb and as a result the currents are greater on the flood ride. Flood dominant estuaries Residual Spectrum Observed Spectrum 6 7 8 9 10 11 12 Frequency (cycles/day) Fig, 3. Power spectrum of tidal energy at the Central Australian Railway bridge. MOST VIEOROLIS SOUTH AUSTRALIAN TIDE eau usually consist al more unstable zeometnes and wm (he lig teen become fled with fine sediments which arise from fhe net transport mtr dhe estuary. Chis Classification is based on the phase difference hetween the prinvary lunar tides (M5) and the major shallow water tide (My). In the ebb donwnant situation, (he mimiium M, current approximately cemeides wilh the maxiniunt of the ML, current, and ut the flood dowunant situaon (he maximum of (he My current approximulely coincides wilt the Maxirouny of the My current, We find thal this clissifeabon iy ot appropriate fy the brine estuary of upper Spenver Gull, which behaves either as hgh water oe low water dominant. High walter dominant conditions occur when the difflerenees between the High water level and mesn sea level are much grewler thar) those between the low water level and mean sea level: (he opposite oecurs for low water dominant conditions. fi high water dommant situations the maximum of the dominant shallow water conabluent approxiinately coineides. with the maxim of the primary Uidal amplitude and vice versa, For crample, arthe Central Australian Railway bridpe. the phase difference benween the MS, constituem and its corresponding astronumicul generating tide 1s (M,+S.)-MS8,=17~ The gh water danniuat enyironuient cansists of a well detined channel which 1s very shallow at low water, but Which can accommodate the propagation at the meoming-and oulyomy tides except yery near high water when overbank Now on the samptire hanks occurs. However, with the law waiter dorninunt environment, the channel is deep enough to have a negligible effect on low water levels, but.as high tide approaches, Turge yolumes ol] water spill mto the adjacent mangrove areas filling up the interfidal depressions und truncating the high water level. The node between these two environments where the most Vigorous tde ovcurs (iarks the position Where these tw opposite effects are in balance. These properties appear to be due to the almost equal amplitudes of M, and S$, (see below! High water Joniinant conditions are well developed atthe Central Australian Railway bridge such that the low water levels appeal to be “euler (Fig. db). There ure four interesting teatures of this pecerd. First, the low waler levels al /he peap lide are lower than ar the spring tide. It is believed that this is the result.at the formation of intertidal pools of gulf water trapped berween the Central Australian Railway bridge and Yorkey Crossing, Water is held in this region on the ebbing tide which ws slowly diridined until the ude tums bul during the neap pyele, less walter is able (o be stored io the upper reaches and subsequently the low weter level is less than the spring tidal level. Second, the ditrnul inequaliy of the ide produced hy the bewing of the diurnal dnd semi-ditrnal eon stituents 1S Signiticanily madulated ac the Central Australian Railway bridge relutive wo the Whyalla Railway bridge due to the generation of the ferdiurnal Udes. Third, the residual records show thal stornr sunes are usually greatly attenuated bewween Whyalla and Central Austratian Railway brilges, Fourth, the drainage (rom the sumphire flats retards the low tide much tore than the high tee ‘The approximate lags between Yorkey Crossing and Pant Augusta, determined direedy trom the short tidal records of the pilot study, were high water 35 min, low water 180 min, Similar results can be obtained from. Fig. 4a and b. The difference between the lags is promarily due ter the major quarter diurnal shallow water (We (MSy), 45 can be seen from Fig, Sa in which the three tidal constiluenls M5. S, and MS, are represented as well us the combinauon of the uirec. An interesting feature of Fig, 46 1s the form of the tidal Tiige curve during the ebb, which resembles an exponential drainage curve, and is clearly reproduced in Bip. Sa. ‘Thus the drainage from the samphire flats is explained harmonically by the existence al MSy,. This appears to be & unique property of the system in which M, and S, have approximately the same amplitude, The corresponding current record (Fig, 5th), which is consteucted hy differentiaing the tdal clevations with respect ta time. and scaling lo pave niaximum tidal velocities similar to the observatians (~U.A5 nis!) shows characteristic current spikes at (he beginning of both the ebb and flood tides which are of similar amplitude and also a slaw ehh (3 ems ') during low tide. This structure was observed in the Short period current meter deployments just south of Yorkey Crossing and at the salt bridge (Bye & Harbison 1987, 1994). Flattening of high water ain be seen inthe Whyalla Railway bridge reeurd (Pig. 4a) but the impurtanee of MS, (Table }) 0s much smaller here thar in the high lide demninant conditions at the Central Australian Railway bridge Conclusigns Following: at extensive field prograinine of (dal elevauions and currents, the laraest ude in South Australia § believed to exist in upper Spencer Gulf. north of Port Augusts. This tide is the peak of the three- quarter resonance in Spencer Gull which vives rise to arapid increase in amplitude northwards trom near Wallaroo, The jocalion of the Whyalla Railway bridge was observed 19 hitve the largest amplitudes of (he major astronomical tidal constituents (namely the M5 S.. K, and O, tides) giving acidal range whieh Ses within 4 to 4.5 m where typically the tidal range in both gulls 1 claser to 2.5 to 4a, Just heyond the bridee 130 C, SCHLUTER. J, A. T. BYE & P. HARBISON 10 T —T T —_ rt °° °»°»~=2~«' T € (a) ey 8 Qo = 2 6 2 o) 4 1 4 4 i 4 Predicted (m) lop) _| Residual (m) = ! ! 4/10/1993 6/10/1993 11/10/1993 16/10/1993 21/10/1993 26/10/1993 date 3 T T a > o———— eo < (b) He Oo 2 faa ne} (oe) 0 3 1 T £ o2 2 3 eit Ai o 0 = a ee La = é w =! 2 WwW) @ c =I 4 ——— ai ———|— 1/10/1993 6/10/1993 11/10/1993 16/10/1993 21/10/1993 26/10/1993 date Fig. 4, Tidal observations tor October 1993, (a). The Whyalla Railway bridge. (b) The Central Australian Railway bridge, The residual (Res) = the observed (Obs) - the predicted (Pred) tidal elevation. MOST VIGOROUS SOUTH AUSTRALIAN TIDE 131 Amplitude (m) - M, 06 + 3 Ms, Time (hours) Speed (m/s) Time (hours) Fig. 5. (a) Tidal elevations and (b) Tidal currents at the Central Australian Railway bridge, constructed using the tidal constituents M,, S5, and MS,. 132 C. SCHLUTER, EA a shallow water dal node occurs and we idennty Us location as the probable position of the largest tidal vurige. As in all tidal stadies, however, the lonver the length of tidal recards, the betler is the harmonic analysis, In this study we have relied on one-two month deployments ut several locations. Longer reeords would be necessary lo improve the decuracy of the Udal constants. The nonlinewr interaction between the major astronomical constituents and the surrounding bathymetric features leads (to the veneration of significant shallow water tides, especially the quarter diurnal MS, constituent which dominates because of the similar amplitudes of the M, and S, tides, These observations prompt the speculation that the samphire Mats and mangrove forest environment have evolved asa positive feedback t0 the shallow water tidal interactions. In other words, In the absence of the unusual South Australian tidal regime in which the major semi-diumnal tidal constituents (M. and §.) BYE & BP HARBISON have a similar amplitude, the intertidal environment would be quite different. tis also likely that the changes: in the intertidal envirooment of upper Spencer Gulf and its northward extension into the Pirie-Toprens plains hat have oecurred due tu sea level changes have been decisively influenced by shallow water ddal unteractinns. Acknowledgments The wurhors would like to Unk ihe seat of the National ‘Vidal Facility for the preparation of all the udul equipment used in this study und also for (he invaluable assistince in the lidal analyses. The invesigation was supported by a Marine Sciences and Technologies (MST) Grant 85/I0IS and a Flinders University Research Budget (URB) Grant, Helptul comments by the referees are alsy aeknowledged, References Biis, Po & Nove, Bod. (986) Tides of Spencer Gult, South Australia pp. 519-530 Jn Noye, BJ & May. Po (ids) “Computational Techniques & Applications CTAC-85" (Elsevier, Anisterdar). Bowers. DOG. & Lesson, GW. (1990) Tidal progression ig Tear-resonunt system a cuse study [rom Sourh Australia, Esteearine Coastal and Shelf Serenee 30, 17-34 Bri. J. A, T) 1976) Physicul oceanography of Gull St Vineent and Investigator Stra, Trans, KR. See. 5. Aust. 100, 143-160, (1990) Richardson Number profiles in laboratory experiments applicd to shallow seas. Geaplys, Alstropltys, Fluid Dyn. 52, 145-166, —. & Haruison, P (1987) “Hydrological Observations in Spencer Gulf and the Pirie Torrens Plains, South Australia during (986: Cruise report 14° (The Flinders Institute for Atmosphere and Marine Seienees, Adelaide), & (891) Transfer of inland salts to the marine enviromental the head_of Spencer Gull, South Austraha Palacoseour, Palacnclimatol, Palienecoul 84, AST-368, il BH (1994) “Hydrological Observahons in Spencer Gulf and the Pirie- Torrens Plains, South Australia during 1987 and 198%: Cruise report 15" (The Flinders Institute for Atmospheric and Marine Sciences, Adelaide), _ & Werrevieao. JR. Ine. (1975) A theoretical model ofthe flow in the mouth of Spencer Gull, Sourh Australia. Esinarine and Coastal Marine Science 3. 477-481, CHAPMAN, BR. Wo (1892) The tides of the coast of South Australia, duit Assen. ddvanee, Sci, Rep. Committee 2 baston, AK, (1970) The lides of the continent oF Australia Horace Lamb Centre for Oceanographival Research Res; Pap. 47, ——— (1978) A reappraisal of the lides i Spencer Gull Soutle Australian, Adsl J Mar, Freshy, Res, 3, 467-477. PRibbRICHS. CT & AnaREY, DG, (1988) Nonlinenr tical distortion in shatlow well-mixed estuaries a synthesis Faraanne Coastal and Shel} Seience 27 521-945. GALLAGHER. BOR Munk. W. CLO7L) Tides in Shallow Water Spectroscopy, JeHus 23, 446-463. Lennon. G. W., Bowers, DOG. NUKES, RL A., Sear, B. BD, AU, M., Boyre, 3, Wes, C, Weeeonip, Mo, Joransson, G,, Nine, S. Perrosevies, 2. StepHpson. P, Susmin, Av Ag dk Winkie. SB A, (887) Gravity currents and the polease of Sall from an inverse estuary, Nature 327, 695-697, Nove, Bo J. (984) Physical processes and pollution in (he waters of Spencer Gull, Mur. Geol. Gt, 197-220, —. May, R. 1, & Trunser M- DB. (O81) Three- dimensional numerival of tides mn Spencer Gull) Qeeu Management 6, TAR. —, Bins Po & Lewis. G (1894) Predionan afar Slick mayerment in Northert Spencer Goll pp, 320-328 tn Gardner, Ho, Singleton, DL, Stewart, OD. (Bas) “Computational Techniques & Applicatianss CTAC-93% (World Scientific, Sinuapere)- Nuwpes. Ro A, & Lenwon, G. W. (1987) Episodic strut cation and gravity currents iA a marine environment et modulated rurbulenee. .. Geapliys, Res. 93S, $465-S480, Puc, DT. 41987) “Tides, surges, and mean sew level” (Wiley, New York). SAMAR ASINGHE. LR. de sibva (1989) Transient salt wedges ia tidal poll’ A criterion for their formation, Esmarine Comite! and Shelf Science 28, (W148. Scubuper. C, CG, (1993) Tidal Modelling withia aw Mangrove Environment, preprints af the th Australasian Conference on Coastal and Ovoun Enymecring” (The Institution of Engineers, Australi). Simpson. J A & Beavers. Do 981) Morlels of stiarfcation and fron) moverient in shell seas Decp Seu Rey. IRA, 727-738, STUDIES ON EUTOBRILUS HEPTAPAPILLATUS (NEMATODA: TOBRILIDAE) THE PREDOMINANT NEMATODE INHABITING THE BOTTOMS OF LAKE ALBERT AND ALEXANDRINA, SOUTH AUSTRALIA By ALAN F. BIRD* Summary Bird, A. F. (1995) Studies on Eutobrilus heptapapillatus (Nematoda: Tobrilidae) the predominant nematode inhabiting the bottoms of lakes Albert and Alexandrina, South Australia. Trans. R. Soc. S. Aust. 119(3), 133-141, 30 November, 1995. Eutobrilus heptapapillatus, a cosmopolitan fresh water nematode has been isolated from the bottoms of Lakes Albert and Alexandrina where it comprises up to 87% and 85% of the nematode population respectively. The environment at the bottoms of the lakes in which these nematodes live is described and measurements of males and females from each of these environments are compared with those of a South African population. There are significant differences in tail length between the Australian and South African populations. Egg laying in the Australian population has been observed and is described. The presence of crystalloid structures in these nematodes has been noted and the possibility of their occurrence being associated with increased salinity is discussed. Key Words: Eutobrilus heptapapillatus, nematodes, Lake Albert, Lake Alexandrina, sediment, eggs, morphology, measurements, crystalloids. Hansaetiinis of te Raval Saciety af 8. Aust. (1995). W931. 19-141 STUDIES ON EUTOBRILUS HEPTAPAPILLATUS (NEMATODA: TOBRILIDAE) THE PREDOMINANT NEMATODE (NHABITING THE BOTTOMS OF LAKES ALBERT AND ALEXANDRINA, SOUTH AUSTRALIA by ALAN F. Birb* Summary Gino. A, F (1995) Studies on Fawhrilas heptepapiliaws (Nematoda , Tobrilidac) the predominant mematade inhabiting the bottoms of lakes Albert und Alexandrina, South Australia Jruny, Ro Soc S Aiest, 13), 133-141, 40) NoveTiber 1995. Eyinbriluy heptupepillais 3 cosmopolitan fresh water nematode has been iselarert from the bottoms ol Lakes Albert and Alesundrina where it comprises up lo 87% and 85% of the nematode population respectively. The cuviropment al the bollors of the lakes in Which these nematodes live is deseribed and measurements of inales and females from exe of these environments are compared with thase of a South Aftican population, There are significant differences in tail length berween the Australian and South African populations, Bee laying yn the Australian pepulalion has been observed and 7s deseribed, The presence of crystalloid structures in these nematides has been noted and (he possibility of their occurrence being associated with mereased salinity 1 discussed. Kiy Wanos: Buobrifus beplapapitiaius, pemulodes, lake Albert, Lake Alexandrina, sediment Morphology, measurements, crystal los Introduction The nematode Autobrilay heptapepillains (Joubert & Heyns, 1979) Tsalolikhin, O81 has a world-wide distribution in a range of freshwater habinuts. [n South Australia this nematode occurs at various sites on the shores of Lake Alexandrina and Hindmarsh [sland at the mouth of the Murray River (Nicholas eral. 1992) ind was the mast comin species extracted from a suiple dredged from a depth of 3 m at the southern end of Lake Alexandrina. To date no studies haye been published on measurements of this nematode nor of its presence or absence at the bottom of the adjacent Lake Albert. Th this paper | compare measurements of males and females oF E. heptapapillates: trom Lukes Albert and Alexandrina with those from South African populations (Swart & Heyns 1988). L also describe thetr habitats dnd their proportions to ather nematode species Jound in these habitats, as well as the percentage of F. heplapapillarus containing crystalloid inclusions. Materials and Methods Collection of material Samples were collected using a benthic grab from the bottoms of Lakes Albert and Alexandrina al the following localities. For Lake Alexandrina the cOlleeting site was at thé navigation marker No. 84 (Fig, 1 site [1)), The Lake Albert collecting site was 2-3 kin off shore from the town of Meningte with compass bearings 135" on the town’s water tnwer, 285° an trees. on the Coorong side of the lake, 205" on a *) Playford Road, Mitcham, S, Aust, 5062, apps, headland on the port side and KS” on a barren hill top on the starboard side (Fig. 1 site [2]), In each case the contents of the benthic grab were placed in a plastic hag and stored in u cooled msulated container, The dry weight of the sediment was determined by allowing it 10 grayitate from the water included in the benthic sample in a graduated cylinder. The supernatant was removed by suction and the sediment was then spooned into a weighed beaker whieh was placed i an incubator af 40°C. Dehydration was maintained until a constant weight was reached. Membrane (0,2 am) liltered walter sarnples from the Jakes were taken simultaneously with the sediment samples taken with the benthic grab. Salinity was calculated from. electrical conductivity (Nicholas et al, 1992) and a range of elements was analysed using the technique of Zarcinas and Cartwright (1983), Particle size of these samples was measured using various techniques as deseribed by Beech (Nicholas et al 1992), A large plastic container was filled with lake water from the sampling site and this: water was used lo dilute the samples during the sieving procedures used to separate the nematodes, This consisted of passing the samples through 2 mm, 850 pm. 710 zm, 250 pm, 120 wm and 90 pm sieves. Ln samples containing much sand, further sieving through 73 pm, 53 pm and 38 ji Sieves Was undertaken. However, in the case of Lake Albert samples, the sediment which passed through the 90 pm sieve would have blocked the remaining three sieves. Accordingly, the material obtained on the 120 zm and 90 win sieves was diluted to facilitate microscopic observation and aliquots were examined under the dissecting microscope. The nematodes were picked out alive on mounted eyelashes, their Movement indiculing their presence in the sample. 134 They were placed in a test tube in a small volume of filtered lake water and an equal volume of boiling double strength FA 4: | solution (20 ml 40% formaldehyde and 2 ml glacial acetic acid in 78 ml of distilled water) (Hooper 1986) was added to the shaken suspension of nematodes. These specimens were processed to pure glycerol using Seinhorst’s (1959) method and mounted in anhydrous glycerol on slides sealed to a coverslip by molten paraffin as described by De Maeseneer and D'Herde (1963). Nematodes fixed and processed into glycerol in this manner were photographed with Ilford Pan F film. Living nematodes, for example females laying eggs, were HINDMARSH ISLAND A, F. BIRD photographed using Uford Delta 400 film. These nematodes were observed and photographed using i Vanox AHBT research microscope equipped with bnght field and interference contrast (Nomarski) optics. Results The water enviranment Lake Albert is a relatively large body of water about 16 km x 10 km connected to Lake Alexandrina, which is approximately 30 km x 15 km, by 4 narrow channel of water (Fig. 1). LAKE ALEXANDRINA ~~ \ site (1) GOOLWA e << MURRAY MOUTH SOUTHERN OCEAN LAKE ALBERT site (2) MENINGIE Coorong Fig. 1 Map showing the location of collecting sites (1) and (2) in Lakes Alexandrina and Albert respectively. Taste |. Analyses of major seluble ions in water fram the shores of Lakes Alexandrina and Albert sampled en the sume day at a six monthly interval and from water sampled from the middle of Lake Albert ata later date. - mel Date Locality Na Cl Ca Mg K s B.C rss* (Site) ds m' oe 29 April 1993 Alexandrina 48 82 15 if 5 6 0.42 0.03 (shore) Albert 214 348 8 33 12 22 1.6 O10 22 Oct 1993 Alexandrina $4 105 ¢ 8 3 7 0.4 0,03 (shore) Albert 200 375 34 31 10 22 15 0.10 20 May 1994 Albert 188 205 34 28 10 16 1.23 0.06 (mid-lake) “EC. = electrical conductivity (deci-siemens mm) " TSS = total soluble salts (estimated percentage) A NEMATODE FROM LAKES ALBERL AND ALEXANDRINA li The results of (he analyses of water collected from Lakes Albert und Alexandrina are given in Table 1, From these results it can be seen that there is mostly a three-to four-fold difference in the tou soluble salts in water samples collected from the shores of the lwo lakes on the same day, These differences in the major ions persisted in water samples taken six months later (Table 1), The sediment environment The surface 15 cm of the soil ut the boule of Lake Albert consists of a slimy sediment, largely Composed of clay which made up 48-61% of samples of this surface sediment or slime laken from various pants ob the lake as the top component af core samples (Taylor & Poole 1931), {t Was estimated that only |/6th of the sediment from Lake Albert consisted of solid material. This material comprised 47% clay, 25% silt, 3% fine sand and less than 1% coarse sand. The nematade The most common nematode in the Lake Albert sediment, Extebrilus heptapupillatus, comprised up to 87% of the nematode population; the remainder mostly consisied of monhysterids. Similarly the benthic sample from Lake Alexandrina comprised up to 85% &, heptapapillatus. The ratios of larvae, males und females were similar in two different collections from Lake Albert, In one instance an aliquot containing 137 nematodes had 43% larvae, 16% males and 41% temales, In the other harvest the ralios were 39% larvae, 23% males and 38% females, In an aliquot cuntaming &,. Neprapapillaris trom the Lake Alexandrina benthic sumple, (he ratios were 51% larvae, 37% imiles and 12% lemales. An obvious difference beiween these |wo pupulations was the presence of crystalloid inclusions (Fig. 2) in 32% of the nematodes from Lake Albert whereas none was observed from the Lake Alexandrina sample. Comparison of populavons of E heptapupillatus, Specimens from both lakes were meusured and compared with each other and with those from South Africa (Swart & Heyns 1988), [rt can be seen (Table 2) thal the males of these three populauions are similar in many respects. For example. they are of similir length, have the same body width at the anus and have similarly-sized copulatory spicules. Diflerences in maximum body width and pharynx length could nol be analysed statistically due to the absenee of certain measurements of the South African population, There is, However, 4 significant difference in tail length (P.<0,001) between the Australian populations (Lake Albert with a mean of 179 yim and Lake Alexandrina with a mean of 173 ym) and the South African population (mean of 244 ym). This significant difference tetail lenge ys mot se pronounced (PF 0,05) between [he means ot coiled (14.3%) and uncoiled (also 14.3%) nematodes, Thus expressing distances between supplements as percentages rather than actual measurements when making comparisons between nematodes that are coiled into various shapes provides a standardised measure for differently-coiled nematodes. Measurements of the Australian populations were combined to obtain a pooled estimate of the means and standard deviations, Because these values uppeared normally distributed, standard deviations of the South Affican values were calculated assuming a normal distribution (Table 4). The positions of the Australian population male supplementary organs differ in some respects. from those of the South African population (Table 4) ullhough (hese differences are not significant except forthe $. and S,(P<001), These differences appear minor compared with the similarities that exist between the populations. Thus the females (Table 3) are of similar Jength and have a similar vulval position although the South Affican population appears narrower with a longer pharynx and a significantly longer Lail, The females of this species (Figs 5, 6) are didelphic and amphidelphic, The genital tract (Fig. 6) consists of ovary, short oviduct, pars dilulala and uterus that may contain oval-shaped sperm. Ege laying The laying process was observed in a specimen callected the previous day from Lake Alexandrina, Tt wis iia sitting drop shde in fillered Lake Alexandrina water (0.2 am membrane). Egg laying took place at 23'C and was very rapid, the actual emergence of the egg being completed in several seconds. The whole process was filmed (Fig. 7) (Mord XP L400 film), The egg is shown moving from the pars dilarata into the uterus (Fig, 7A, B) and from there into the vagina (Fig. 7@, DP). During the final stages of laying, the eg irioves Fron the vagina lo the exterior through the vulva (Fig. 7E, FG, H). The eps which is oval (ellipsoidal) within the nematode assumes a spherical shape soon afer laying (Fig. &), Tt has a mean diameter of 708 ym (42.9 SD) including the shell which has a mean thickness of &4 ym (+0.6 SD). This compares with in wero measdrements of (ixed material of 73.5 jn x 48.9 jun including an egg shell thickness of 5,5 un (Swart & Heyns 1988). Discussion Over 60 years ago Taylor and Poole of CS&IR Division of Soil Research (now CSIRO Division of Sails) published the results of a soil survey ofthe bed of Lake Albert (Taylor & Poole 1931), This work resulled from a request by the appropriate branches of both the State A NEMATODE FROM LAKES ALBERT AND ALEXANDRINA 137 TaBLe 4. Comparison of measurements of distances between supplementary organs, expressed as percentages of the sum of the distances between supplements, in different populations of males of Eutobrilus heptapapillatus. Parts measured Tsitsikama, Forest Lake Albert Lake Alexandrina Cape Province, South Africa South Australia South Australia (Swart & Heyns 1988) (present study) present study) Rinse Mean SD* Range Mean SD Range Mean SD Cloaca + ST 76-114 93 +1.6 4.7-11.8 8.5 +2.6 91-118 9.8 +1.1 Ss? + S6 7.6-9.2 3 +0,7 6.0-9.9 8.1 +1.4 8.7-12.3 10,3 +13 S6 * $5 88-113 10,2 +1.1 9,7-12.2 Wal 41.2 10.3-13.0 11.6 10.9 $5 » S4 16.2-17.8 17.4 40,7 16.5-23.1 20.1 42.8 17.0-21.7 19.1 +1.6 S4 + S3 14.4-17.6 15.6 +14 {3,2-15.3 14.3 +1.0 12,3-15,3 13.6 41.1 S3 + §2 19.9-21.6 20.8 40.7 15.3-19.7 Ti? +1.7 15,4-18.2 16.7 +1.0 82 + $i 16.4-20.3 18.4 +1.6 18.6-25.0 20.7 12,5 15.1-20.8 18.9 $2.0 *= estimated using sample size and range and assuming normal distribution Fig. 3. Photograph of an adult male Eutobrilus heptapapillatus from Lake Albert. Nomarski optics showing pharynx (p), intestine (i), pharyngeal glands (g), testis (t), retracted copulatory spicules (s). Scale bar = 100 pm. Fig. 4. Photograph of an enlarged portion of an adult male Eutobrilus heptapapillatus from Lake Albert. Nomarski optics showing the seven supplementary organs (small arrows) and the everted copulatory spicules (s), Note relatively short tail. Seale bar = 50 ym. 138 A, TDD Fig. 5, Photograph of un adult emule Euzobriles heptapapillatas Tran Lake Atbert. Norrshi optics showing pharynx (p), intestine (i) and vulva (¥). Scale bar = 100 pin Fig. 6 Photograph ofan entarged portivn ofan adult fernale Eutobrifics hemmapapiiawus tram Lake Albert. Nomarski opries showine (he didelphic reproductive syste voryisting on etch side of ovary Co} shore oviduct pod), pas dibatata (pal), HePus tur und vulva ty), Scale bar = 50 pm. A NEMATODE FROM LAKES ALBERT AND ALEXANDRINA 139 Fig, 7. Photographic sequence of egg laying in Eutobrilus heptapapillatus. Nomarski optics. Arrows indicate. vulval opening. A.. B. Movement of the egg from the pars dilatata to the uterus. C., D, Movement of the egg from uterus to vagina. E., F,, G, The process of laying as the egg passes from the vagina to the exterior via the vulva. H. The newly laid egg. Scale bar = 50 pm. 140 S & BIRD und Commonwealth Governments regarding the leasibilily of using the sail at the buttonr of the lake lor agriculture after it had been drained. Taylor and Poole (1931) showed that the drained lake would be unsuitable for agricullural purposes. Thus. at a time when the clearing of land was in full swing, these workers. were able to show, as a result of their snil survey, condueted under difficult conditions, that drainage of this lake would have been a costly mistake. Furthermore, their detailed results (Taylor & Poole 1931) provided valuable information on which to base further studies of the lake's benthos Taylor & Poole (1931) reported that lake Albert was once connected to the saline waters of the Coarang by an ancient river channel which persists today us 4 lagoon extending from the southern side of the lake (Fig. J). Thus, although Lake Albert is pot now flushed through by waters of the River Murray, as is Lake Alexandrina, it my once have been when the rivet was in flood The barrages at the mouth of the River Murray were not built until 1940 so that at the time of Taylor's and Poole’s survey ducing Match to April 1930 when the river was running law, Lake Alexandrina had hecome very saline as a result of incoming sea water, Thus water from Lake Alexandrina Was thaking (he water fron Lake Albert more saline and “was not potable for humans and taken uowillingly by stock until accustomed to 1” (Taylor & Poole 1931). Today, due to the presence of the barrages, Lake Alexandrina is much less saline than it was and although the quantities of soluble salts contained in its waler can and do vary from lime to time depending on river fushings, iL is clear from the samples collected on the same day fron both Jukes ata six-monthly interval (Table 1) that Lake Albert has a higher concentration of soluble salis than Lake Alexandrina during normal river flow. Nicholas vt ul (1992) have shown that the concentration of soluble salts in water collected from sites on the share of Luke Alexandrina varies from month to (ionth and hence iL was necessary. to colleet water from (he Iwo lakes on the same day $o that valid comparisons could be rhade, IC is interesting to speculate whether or net the presence of crystalloid bodies observed jin &, heplapapillates trom Lake Albert but not in specimens of the nematode collected on the same day from Lake Alexandrina when its soluble salt values were low. might he associated with increased salinity in these lakes. Crystalloid bodies were [ound in & heprapapitlatus collected (rom Lake Alexandrina both ut the water's edge and From the bottom of the lake in 97% of the nematodes examined (Bird ef al, 1991) but no correlation Was tade with the salinity of the lake at that lime. However, examination of these dara shows that the concentrations of sodium and chloride ions (Table 2 - Nicholas et ul. 1992) were greater than those Obtained Troi this lake during the present study (Table |), Furthermore, contrary te our bindings that nematodes nwintained in uquaria over ud pericd OF hwo months “appeared to be Free of erystalloids? b have found that there was un almost (hree- fold anerevse in both numbers of £, heptapepittalus and (heir erystalloid content when mud From Lake Albert was plaved inant aquarium tank, covered with Luke Albert water and left for four tnoaths with ovcayional aeration, Under these vonditions, the ratio of larvae (o lemales lo males was 85 512) 3 of which 93% contained erystalloids. The large number of larvae present would probubly be due lo relatively recent hatching from eps und the low number of udults to luck of food. The increase in the percentage of erystalloids present from 32% to 93% in nematodes kept iy a aquariaum tank for Jour months would probably be due lo an inerease in the cuncentration of soluble salts due ta evaporation From the lank. This partieular butch of Luke Albert water (eollected on 20 May 1994) had initial sodium and vhloride ion readings of IK8 and 265 mg 1! respectively (Table |) which are apparently high enough to induce the development of crystalloids. The vechrrenve and possible functions of erystalloids ina moinber of penera of aquatic free-living nematodes uve been mécanded hy various Workers (Bird ef ef. ee —— Fip 8. Phouaraph fy Of! treshhy laid cas of Enrabriles the yapapilaius Brighe held opoes showing vulva Cy) and Sphetical eves (ie) sett (herr refalively thick shelly (esp Scule bur = 20 pm a NEMATODE FROM LAKES ALBERT AND ALEXANDRINA ‘4 1941) hur as yet io clear-cut evidence for thetr funetion has been oblitined, Clearly fumher reseanch is required (9 lest whether these corystalloid struwtures ure produeed in’ the nematodes yp response to changes in thei enyironment and whether or fut they are a manifestation ala diseased! slate, since they appear ty be associated with small regular particles thal resemble idosahed ral viruses in some respeets (Aird er al. 199), Tthink that his nematode which predominates 1 the sediment at the bottom: of these Jukes is E heplapapiliaias altnough, ax will be discussed below, there are-some diflerences hebween Lie Australian anc South African populations. {has ull the characteristies of its Subfamily (Rutobrilingeé) deseribed by Tsalolikhin (1983) namely, hedgehow- like supplementary organs (Fig. 3), a muscular yagina and Well-differentiated female genital system (Fig. 5) and well-developed aod rounded pharyngeal whinds. Sintilarly, tis less than 3.5 mm in length ind it has the described species characteristics of a cuticle withant pronounced annulations, head bristles that do not exceed 14 pny and, most obvious of alf, mules with seven supplementary organs. Tt does nol quile Ail Tsalolikhin’s key in having females whose tails are not len lines greater than the body width at the anus (‘Table 3) but (his also upplics tothe South African spectiens (Swart & Heyns 1988). However. these various morphotogical Jiflerences ure likely to be reflections of variability hetween different populations of this species of nerhatode rather than suggesting thar the Australian and South African nematodes are different species. Another apparent difference is thal the egg. although oval-shaped when within the female. becomes round wher laid (Figs 7, &). However, it Seems likely that cue measurements in the past may have all been made while the eggs were within the female in fixed material. The transition from oval within the female to spherical an laying is, however, illustrated by Tsulolikhin (1983) in his book, Since eggs of these species can not be identified unless laid from an identified female, their shape for laxonomie purposes is fisted in the deseription of the genus Futobrilus as oval (Tsalolikhin 1983), Furthermore, since fixation leads to shrinkage, Measurements from fixed material will always be lower than those for unfixed, freshly laid eges. The combination of characters described above places (hese womis firmly in the Tobrilidae, Minor differences between the populations in South Australia dnd South Afnea ure nor considered significant and the worms are confidently referred to as &. heprapapillatas. Much remains to be learnt about this cosmopolitan nematode. Its feeding habils, rates of growth and longevity im the lakes wre unknown. Such information is needed af its value as-dn indicator of envirunmentil pollution jn these cnvjronments is te be determined. Acknowledgments ] wish to thank Angela Reid, CSIRO Biometrics Umit for the statistical analyses, the Soils Division, CSIRO for accommodation, facilities and expertise, including that of Adrian Beech, and the Zoology Department. University of Adeluide for facilities and equipment and especially for the help of lan MeGruith. | thunk Jean Bind for help with collecting and Jean Bird, Lester Cannon, Mary McHugh and Warwick Nicholas for constructive criticism of |he manuscript, This work Was made possible by a grant trom the Australian Biologival Resources Study for which L am most grateful. References Rinn, A, FE, MeCiiure, S G & Nicuor as, WoL (I99T) Observalions on crystalloid bodies in (he pseudocoelom OL kufobrilas heptapapillatus, J. Nemetal. 23, 39-47, Di MA sSeneeR, 1, & D'Herpe, J, (1963) Méthodes utilisees pour VPetude Wea anguillules libres duo sal. Revur de Cagrieuliure, Bruxelles We. 44-4 47_ Hooree, Do 4 O86) Handling. fring, staining und mounting nematodes pp. SY-80 i Southey. Jk (Ed) “Luboratory Methods tar Work wih Phint and Soil Nematodes” (HMSO, Londont, Nicnonas. W, L, Bian, A, Ry Bercy T, A, & Siewari, A.C. (1992) The nematode fauna of the Murry River egtuary, South Australias the etfeets of the barrages across iis mouth, Midroblelevie 234, 47-101, Spinniorst, J. W. (S90) 4 rapid methad for fhe transfer of nemitodes from fixative to anhydrous glycerin. Nematolovica & 67:69. Swart, A, & Hbyns, J, (988) Redescription of Eutebrilus heprapapillaris (Joubert & Heyns, 1979) Tsalolikhin, 1981 With notes on is morpholagy and possible exeretary system (Nematoda » Tobrilidae). Phytophylacica 20, lol-lok. TVavior, . K. & Poore HG. (1931) Report on the soils ofthe bed of Lake Albert, South Australia. Coun. setent inet, Res. Aust. 4, 83-95, TSALOLIEHIN, S.J, (983) “Nematode families Tobrilidae and Tripylidae world fiuna” (Nauka, Leningrad), Zarcinas, BA, & Cartwrioni, B. (1983) Analysis of soil und pliant material hy inductively coupled plasma-opticat! emission spectrometry, Division of Soils Tech, Paper No. 43. CSIRO, Ausinalia DISTRIBUTION OF SPECIES OF TRICHOSTRONGYLOID NEMATODE PARASITES IN THE SMALL INTESTINE OF THE BUSH RAT, RATTUS FUSCIPES By L. F. SKERRATT*, I. BEVERIDGE* & M.-C. DURETTE-DESSETT Summary Skerratt, L. F., Beveridge, I. & Durette-Desset, M.-C. (1995) Distribution of species of trichostrongyloid nematode parasites in the small intestine of the bush rat, Rattus fuscipes. Trans. R. Soc. S$. Aust. 119(3), 143-148, 30 November, 1995. The distribution of three trichostrongyloid nematodes, Nippostrongylus magnus (Mawson, 1961), Odilia bainae Beveridge & Durette-Desset, 1992 and Paraustrostrongylus ratti Obendorf, 1979, in the small intestine of bush rats, Rattus fuscipes, was investigated. Each of these species exhibited a significantly different longitudinal distribution within the small intestine. Interactions between the three species, identified by comparisons of the fundamental and realised overlaps in nematode distributions, were the probable cause of the differences in distribution between species. The different distributions, which are here interpreted as niches, occupied by the nematode species are consistent with the hypotheses that O. bainae was probably a parasite of hydromyine rodents which filled a vacant niche when it switched to R. fuscipes as a host, while P. ratti probably occupied another vacant niche when it switched to R. fuscipes from an original marsupial host. Key Words: Parasite, ecology, niche, Trichostrongyloidea, Rattus, interaction. Transactions if tie Reval Socters of 8 aus (1985) W193), a 148, DISTRIBUTION OF SPECIES OF TRICHOSTRONGYLOTD NEMATODE PARASITES IN THE SMALL INTESTINE OF THE BUSH KAT, RATTUS PUSCIPES by L. F Skerreary* |. BEVERIDGE* & M.-C. DuRWITh-DESSETT Suinuniary Skearar), Lh, BEYERIbeb 1, & DURE Dressel, MC (1995) Distribution of species of trighostrongyloid Hermulode parasites in the small antestine of the bush rat, Rats fascipes, lramy. Ro Sac. 8. Aust. W949). 45-148, AQ Novyerber (995 ‘The diamburion of three michostrngyloid nemalodes, Mippeytronuviay meinies (Mawson, 961), Odilies battae Beyerdwe & Durctlo Dosse( OZ und FRaraustrostrniies haiti Obendert, 979 in the small intestine of bush rals, Rares fieseipes, was investigated Buell of theso species exhibited a signifivantly difleren) fongiwdinal distribution Wilh the small iitesmme lntertictinnas berween the three species, identified by compansens of the fundimental and realised Ovceluprs je nematode dishibulions, were the probable cause of the differences in distribution between species. The different distributions. whieh are bere interpreted as niches, oceupied hy the nemutnde species ure consistent with the hypotheses (hal GO bamtee wis probably a parasite ol hydromyine rodents whieh filled a vacant Hiche when U swatehed te dk fseipesas a ost, while PB rane probably oceupied andther vacunt niche when W switened to Ro faserpes fron an urigenal inarsupnil host. Rey Wokos, Panuile, ceolawy, quohe, Trichostraaeyloideu, Ralls. iferactan Introduction One mode by which parasile evolution may occur is “host switvhing” (Chabaud 1965). This involves a break-down in host speciticity allowing the transfer ata parasite from its asaal host to an unrelated Host species necupying the same environment. The new host may be infected trough dre skin by free-living suiges af the parasie or may ingest the infective form of the parasitewith its food (Chabaud !965), The mechanism of host switching appears to be common among parasiie neniatodes (Chabaud 1982) and (§ based on the ussurmption that the invading nematode parasite is veeupying a previously vaedut niehe within the new host, In the case of intestinal parusites this is. usually defined as a restricted longitudinal of radial distribution within the gulof the host (Schad 1963), Host switching within the nematode superfamily Trichostrongyloidea is well documented (Durette-Desset 1985), yet few studies huve examined whether the invading nematode acrially occupies a separate or previously -vaeant niche. The {richostrongyloid nematode pitrasiles of the native bush rat, artes fieselpes, olfer the opportunity ro examine such an hypothesis. At one locality in Victoria (Blackwood) Obendart (1979) found that A fuyeipes wis parasitised by three species of trichostrongyloids, the heligmasomes Nippasrrangylas rhavnus (Mawson, 1961) and Crilia bainde Beveridge & Duretle-Desser. [942 and (he herpetostronpylid Paraustrosrrangyius rai Obendorl, 1979 (Obendart 1979; Beveridge & Durette-Desset 1992 a,b, 1993). P University of * Departmen) ot Vetenmary sciences Melbourne, Parkville, Vie. 3052. tT Laboratoire de Biologie purwvidure, Muscun nahanal U'Historre nitluretic, Pures, 75005, France rat! belongs to a genus which otherwise occurs exclusively in marsupials and which presumably has switched to its-current eutherian host (Obendorf 1979), Iris considered (Obendort 1979) 10 be one of only Wwe wuumples of trichostrongyloid nematodes switching from marsupials to native rodents, the other being Roolleva hydromyos in the water rat, Aydromv chryse- waster (see Mawson 1961. 1973). O. bainae belangs tow venus whieh is paraside primarily m hydromyine rodents and species of the xenus were considered by Durette-Desset (1985) to have switched secondarily toy murine rodents such as Ro faveipes. Only Nippo- strongylis magnus can be cunsidered an original parasite of this murine rodent (Beveridge & Duretie- Desset 1992). The current study was therefore undertaken to determine the ecological niches occupied by N, magnus, O. bainae and P raiti within the small intestine of R, fuseipes and to examine the extent of overlap between them to establish whether or not each mecupies a distinctive intestinal niche. Materials and Methods Ten bush rats, Rartus fiuscipes, were trapped front along the bunks of the Lerderderg River, Blackwood, Victoria, Australia (37° 29° 5, 144° (9° B) using collapsible aluminiuny traps baited with peanut butter, Immediately following euthanasia wilh chloroform, the small] intestine was removed and divided into sixteen equal parts. The rotal length of the snail intestine was measured. Gul segments were incubated in saline at 37°C for at least 2 hours and all nematodes which emerged fram the mucosa were fixed in hot 70% ethanol betare heing counted. P rat! was distinguished 144 L. F SKERRATY, |. BEVERIDGE & MO DURETTE-DESSET from other Species based on the deseriptinn of Obendorf (1979) using a stereomicroscope, Whereas NV Magnus and OL baihae were differentiated Using the Uescriphions of Beveridge & Durette-Desnet (1992 a,b) and a compound microscope following clearing i lactophenol, The rats were also routinely examined for other helminth parasites, The number of cach species of neniatodes. in individual sections of the small intestine was converted to a percentage of the total number of each species present. For cach species, the positions of the anterior, median and posterior nematodes were deterinined using the method of Bush & Holmes (1986), such (hat the section number in which nematodes oucurred was converted to a percentage of the total length of the smiul! intestine. It was assumed that nematodes were uniformly distributed within each seetion. Differences m distribution between species were tested Statistically using values calculated [rony a 2 % 5 contingency luble (species x sector of intestine) lor each species pair. Sections 5 to 16 of the intestine, where very few nematodes were found, were cojnb|ned to form av single cell in the table, The extent of niche overlap between the three species wus determined using the equation Cay = i- 1 CHIP — Pyjl) where Pri Py (Hurlherl t97s) lex , i= Yi gy such that Pxi and Pyi are the proportions of twa species, x and y, in different segments of the intestine This equation was used to calculate the *fundaniental overlap” belween species pairs, which 1s the extent of overlap (Cxy) io the mean disoributions af the wo nematode species and the “realised overlap” which ts obtained by determining the extent of overlap (Cxy) between two nematode species in individual rats and then computing the average of the individual overlaps Differences between fundamental and realised overlaps. that is when the realised overlap was substan- Hally Jess than the fundamental overlap, were used hy infer the presence Of competition between nematode species, Seven laboratory rats, RL norvegicus, were infected either subcutaneously or orally with 200 - 900 infecuve Tabre 1. Positiore af mean anterior median and postin individuals (+ slandard errr of mean) of Sippoxtrongy lis magnus, Odilia bainae and Paraustrostrongylus rally os percentage distances along the small intestine in ten narurally infected bush rats, Rattus fuscipes. Anterior Median Postenor _ No magnus 1.01+0.64 1[5,14+4.84 5663+ 10,17 O.bqinae = 2.1740.84 182349.07 4840+ 6.47 Porat 3,22 + 1.62 19.064 2,25 49.40 + 4.47 larvae Ol No srasitus ar (2 hartge (Table 3), The infective lame were oftained by culling a mixture ol faeces Ip naturally iniected rals with wolivalel Chareual on morsi filter paper and recovering developed larvae by sedimenkifion in waler Larvar wen Separated On the buss af morphological differenaes (shape of the tail) identifiable using a Sterermicraxcopa: (unpublished observations), Infected raty were killeal with chloroforn) 14 days after infeetion and the distribunon of nematodes in the sunull inrescine determined ina similar fashion to thal described dbove- Expertitental auteetns wilh Poreifh via oral. subcutaneous and perculaneaus, Toutes were uinsuceesshil Results Nemataite disrriherions in wile rts No magniity occurred in 100% nf rhe wild bush. rats examined), with a metin intensity of 67 uind.a range of L-+183, ©) fara was present in 00% of wild rats examined. with a mean tatensitv of 64 and a range of LTS. nat was present in 0% ot wild rats examined with a medn intenwity af I7L and a range of 11094 (Tadle 4), Oder parses found in the (en qaturally intbetee bash rats were very small numbers. of Capillarica sp, (Nematoda) in the first segment of the sanall intestine and Cupelfortid gusrrica (Baylis, 1926) within the squamous epithelium al the slomach in five rats. There was no patholowical reaction fo wlll Cepillaria or eee wiltiin the squamous epiihelium, Aerenikix spumoast (Schneider 1866) was found us the lange intestine of eight rats. The cestodes Chuaneaueniad ritticate (Sandurs, 1957), Hymenclepw cdininura (Rudalpby, (819) and Berfiella anapolvtine Baylis, 34 were found inthe small intestine ©. raifieela inhabtied the opening: of the bile duet. causing hyperplasia of the tile duet epubelum 4, diminutd and B anupolyned were found in segments 412 und SIG respectively. However, ditilhuia was distributed anteriorly te B. anapialyrieg when bath species oceurped jn rats. N, rhayruty occurred mainly (64%) in the anterior two segments of the ered! iresting in naturally infected rats. WIT Maximum intensity in the first segmenr, and its Numbers declined progressively in the rerwiming sections (Fig). The majority of O, buinae (82%) inhabited the five most alteriod segments in naturally jatected bush cats with a maximum intensity (30% | in the second seement (Fig. 1), The majority of PB raili (87%) were distributed throughout the five anterior segments af the sonal aitestine in muturally infected tats (Pig. 1) with maximum intensity deeurring it segments | ro 4. Simall intestine lengths ranged from 56-97 em (mewn 71 em)and hence the average length of cach segment Was 4.5 ei, TRICHOSTRONGYLOID NEMATODES OF THE BUSH RAT Nippostrongylius mugnius bainae Odilia ratlt Paraustrostrangylis ‘Mean percentage of nematodes in each section Section of small intestine Fig 1 Distribution (mean percentage + standard error [error bars|} of Nippostrenzylus maynus, Odilia bainae and Paraustrostrongvius radi in each segment (sixteenths) of the small intestine of ten naturally infected bush rats, Reis fuscipes N. magnus, O. bainae and P. rary were sequentially distributed along the small intestine of naturally infected bush rats but differed in the positions of [heir Inegn anterior, posterior and median individuals (‘Table 1). The mean anterior and median individuals of N, magnus Were anterior to those of OQ. hainae which were 7 TABLE Fundamental and realised overlap beiween niches 145 more untenor to those Po rari. However, the meun posterior individual of NV. mayniy was posterior to both O. bainae and Prati. This was due to Ny magnus inhabiting the posterior quarter of the sniall intestine in two rats, Whereas O. barnae and Po ratti were not found in this segment. The distributions of the three nematode species were significantly different. Chi- squared values obtained for pair-wise comparisons were N. magnus - O. bainae, y= 77.5 (p< 0.001). O. bainae - Prati, x 90.9 (p<0,001), N. magnus - Po raiti, %~ = 18.5 (p<0001), There was no correlation between mean posilions of nematodes with intensity of infechon, except in the case of the posterior position of QO. bainae (vr 0.68, p = 0,004). The varialion in numbers of trichostrongyloid nematodes in each segmentof the intestine of naturally infectéd bush rats was large (Table 4), with the standard deviation equal to or greater than the mean (sce standard error burs in Fig. 1). Despite this variation, the difference in distribution was greater in indryidual rats (1- realised overlap) than the difference in their mean distributions (I fundamental overlap). The realised overlap was lower than the fundamental overlap in 25 of 28 species interactions, The mean realised overlap between N. magnus and Po ratti wis lower thin the overlap between N. magnus and O. bainae and between O. bainae and P. ratti (Table 2). The total numbers of each species of nematode in individual rats and the realised overlaps in. natural infections varied between individuals but the two purameters were independent of one another (r< 0.3). Capillaria sp. occurred in such low numbers that of Nippostrongylus magnus, Odilia bainae cre Paraustrostrongylus ratti in the small inrestine af ten naturally infected bush raty, Rattus tuscipes. No. of rats infected Nematode species } C with both species pair Fundamental overlap Realised overlap (+ standard deviation) N. magnus — OQ. hainae 10 N. magnus — P. ratti 9 O. bainae — Prati 9 0.68 0.55 £0.14 0,61 0.40 + 0.27 0.82 0.55 + 0.23 TABLE 3% Percentage of Nippostrongylus magnus aad Odilia bainae vecurring in /6 segments of the small intestine of lahordtory reared Rattus norvegicus following oral or percutaneous infection with third stage larvae Segment no. of intestine Mean percentage of nematodes in segment (+ standard error) Nippostrongylus magnus Odilia bainae No. of rats 4 3 T Bt T3 OTOF3.8 2 2342.2 7.0 +6.7 3 25425 2.342.3 4 10.0 + 6.8 Q 5 16 0 U Ho L. fh SKERRATT, 1. BEVERIDGE & M.C. DURETTE-DESSET Pat d. Numbers of nematodes, Sippostrongylus magus. Odilia baynse ad Paraustrosicongylus ralli in aectiony (EG) uf thre sinalt testing ef ten materally pifected husk rats, Rattus Tuscipes, — Rat number 5 Secyon I 2 3 4 t 7 4 i) i) ) MWS ——s vs } 14.5.0 0.0, 21 5.01 wa 5.37 2.221000 43.2,2 37,31 49,4238. 2 (0,0 HOS 6140 14,3,5 16,0 8,310 A3136 21h) BATT AN, TG Fhe } 2.0,0 26,2 19.0 A432 336,27 O06 49,25 ALO 13.7.4) 11,2.228 4 230 11,2 w70 KIS 12,0 1.0.12 a7,56 FAB 312.26 9,5.133 4 U,2,0 0.0.0 61.0 35,18 (13.1 0.9.0 4216 10 31.34 9278 b 0.1.0 0.0.1 0.0.0) 144 24,0 0,6,1 0,1.22 O14 1,17,10 43,34 7 0.0.0 00,0 0,00 10,4,18, O10 07.0 O07 O54] 030,11 2.015 bs 0,0,0 0,0,0 0,0,0 214 25,0 0.12 oO1s 0.11 6.25.2 1.2 y) 0,0,0 (0.0 0,00 21.3 2.2.0 0.00 0.0L O01 aval 0,04) Tt) 0.0.0 0,0,0 0.0.0 int ina 0.04) 0.00) 00,0 (231 0,00 | 0,0,0 40,0 0.0.) 1a! Lhd 0,00 0,00 0,0,0 (2131 0,00 12 ),0,0 (0.0 (,0,0 10,1 1,10 0.00 0,00 (00 12.130 0,00 8 1,0,0 0.0,0 (1,00 10,0,0 00,0 0.00 0.00 0,0 10,0,0 0.0.0 \4 0,0,0 ),0,0 0,0,0 9.0.0 00,0 O00 0.0.0 0.00 10.0.0 0,00 5 U.00 0.0.0 6.0.0 90.0 0.0.0 0.0.4) O00 0.00) 9,0,0 0,00 if 0.0.0 0.0.0 00,0 90,0 0,00 00,0 0,00 (0,0 9.0.0 00,0 Total 20000 Uh1614 16.361 96.22.70 491033 147744 577378 99-6002 KITA 46 125,92, 1044 its passible interactions wath the trichostrongyloid pemmlodes Were not considered, Nemeterle distributions in experimentally infected reais No magnis occurred primarily in jhe anterior seemient of experimentally infected luboratory Fat, with a relatively small population of nematodes established in segments 2 to 4 (Table 3). Similarly, O. huinue became established primarily in the first segment of ihe miestine, wath small numbers of nematodes present in segments 2 to 4. The mean intensity of infection was WW lor NM meenus and i for QO. bainie. Discussion The significantly distinct sequential distabutions of (he three species of trichostrongyloid nematode along the small intestine i natural infections and experimental micetions suggest that each species oceupies a distinct niche. Furthermore, the fundamental ovetiaps in natural infechors between the species pairs \. negnus. - QO. bainae (68%) and N, magnus - Prati (6l%) were lower than the 70% yaluc suggested by several authors (Pianka er al, 1979: Holmes & Price 1980: Bull era/. (989) to indivate the eustence Of ecologically relevant differences. Only the fundamental overlap between the species pair. O. bainee - Po rani (82%) was greater than 70%. jlowever. the mean realised overlup (55%) between jhese two species was substantially less than 70%, imficating that these nwo species interact to separate their niches in individual rats, The distribunion of C, bainae in experimental infections (Table 3) differed when compared with natural infections (Pig. 1) in thur in MOnospecific infections i occurred in the mist walerior segment of the duodenuin and (his may be due ti Ihe absence of competition from NM) meninuays ort nati. However, other contributing factors may have been the different species of host, te smaller sample size in experimental infections or the smaller numbers of O; bainae in experimental infections. These observations sugwest that the nematode community in A. fiiseipes is an interactive one. Holmes and Price (1986) separated communities pf parasites into two categories, isolationist and interactive, based on their infrapopulations, that is, populations in individual hosts. They suggested that un interactive community has no vacant niches, purasiles are mot distributed independently and realised distributions of parasites are dependent on other guild members. Sotne of these features are present in the cause ol the (richostrongylou nematode parasites of the bush rar, since the realised overlap was less than the fundamental overlap in most interactions in natural infechons. However, (he small intestine of the bush pat does appear to have vacant niches in natural infections despite (he above evidence fur an interactive community. Although host immunity may reduce the size of apparent yvacunt niches (Noble e¢ af 1989) and low (riosmisyion milessniay prevent parasites filing all available niches (Price 1980), the distribution of the trichostrongyloids in individual rats wis independent of nematode numbers. Thus, assuming vacant niches over. the trichostrongyloids of the bush rat alse demonstrate one feature of an isolationist conmmunily (Holmes & Price I986). The “populatian concentration” and “individual response” hypotheses both explain why the species NV. magnus, O. hainee and P. rari should occupy distinct niches even when additional yvacunt niches are available (Holmes & Price 1986). The “population concentration” hypothesis bas two components, that narrow miche oeeupabon. ts exsentisl for the maintenance Of intraspecihe contact for mating purposes (Rohde 1979, 1982).and that the Qecupalion Of diserele miches is important as a TRICHOSTRONGYLOID NEMATODES OF THE BUSH RAL lAT reproductive isolating mechanism preventing hybridization (Sogandares-Bernal 950° Martin 1969), The “indiwidual response” hypothesis (Price 1984) argues that parasites adapt wo the environment they inhabit and consequently Mil nurrow niches. The distribution and overlap of trichostrongyloid nematodes jn individual bush rats Vary greatly from the means but are independent of nematode oumbers exceptin the case of the posterior extent of Ch byinue in the imestine. The varrability in these intracommiunities may be due (o the biological features of the nematodes and their interactions with one another but may also be caused by variability in the characleristioes of the host which influence parasite infraenmmunities such as host diet (Croll 1976) and blood supply (Croll & Ma 1977), The non-specific host response, pathological responses and acquired immune responses of the hast may alse influence the distribunen ol parasites. The three tichostrongyluid nematode specics found in RK. fiiscipes have different biogeographical origins. The genus Nippestrongvluy occurs primarily in Rarfees spp. in south-east Asia and in other rodents in Asia and the Middle-East. with a single species ul dermopterdns (Beveridge & Durette-Desset 1992). Since the endemic species of Ratrys in Australie probably reached the continent from south-east Asia (Waos & Astin 1981). it is likely that Nipposrrne vies reavhed Australia with them and that subsequent co- specuition led ta the evolution of N, magiis in R. fuseipes (see Beveridge & Duretic-Desset 9924). The genus Odilie occurs pomiurily in hydrogyine or “ole endemuc” rodents in Australia, principally in the genera Melumys and Uromys. Species occurring in Rares spp- have been interpreted as transfers From “old endemic" rodents, which probably evolved between S and 15 Million years ago, to the “new endemic’ Rariys spp. which have been presen! on the continent for about Loanihon years (Watts & Asin 1981). Species of Paraustrastrongylas occur in possums (Phalangeridiac. Petuuridae, Burramyidae) and rat kangaroos {Polorgidae) (Spratt ef a/, 199)), wilh single species, P rantina rodent. The transfer of Puraustrastrongyius, und probably Odilia, ss therefore presumed to be of revent origin (ess than | million years), Holmes (1973) supgested thar stable communities are older than imterachive ones and since most parasite comraunities dre stable, he coneluded thal they ure relatively old, The interietive component of the tnchostrongylod intracommunity in bush rats suupests that iC is a Comparatively young community: This 1 consistent with the hypothesis that 2 rans und possibly also O. haihae dre réevent invaders. Following their invasion of R. fiuscipes. PB raw and O bajnar have occupied distinctive niches within the new bost possibly duce te isolationist lorees and/or their interactions with the other trichostrongyloid nematode species present in the small intestine. Tt 1s possible that P raiti colonised & previously vacant mehe at the Lime of switching since No meagaas and ©. bainae do nol occupy this niche even when A rai is ubsent The wther parasites found in the small intestines ol bush rats. appear to occupy completely different niches from those inhabited by the (richostrongytoid nematodes or ta occur at a very low Tatensily and. prevalence (Cupillaria sp.) and therefore were pot vonsidered in the interactions of the trichastrongyloid com@munily in the present study, The distributions of A. dinijniuta and B. avapolytica overlapped, However. the Ko species were never found inthe same segment of the small intestine although only two fats were iMfected with both species, Beewuse of interactions which tay occur between ther, they may occupy sepurale niches (Holmes 1973), A larger sample ov rats would be needed to examine the extent of interaction between their cestode parasites. Acknowledgments Christine Anderson and Lee Bergerare thanked for thei help in tapping rats and collecting the nematodes. Robin Gasser is thanked for his adviec and help with trapping and Neil Chillon tor comments on drafts ol the paper, Rats were collected under permit no RP9I-095 from the Victorian Department of Conservation and Natural Resources and experimental procedures carried out under The University of Melbourne, Animal Experimentation Ethics Committee no, 270-056-0-89-077) The work was supported financially by the Australian Reseureh Council References Breyeribor. [ & DuremimDesser, M-C. (1992a) The morphology of Mippashartevlus magni. a parasite ol native Australian rodents. Tray, Ro Soc, 8) Awst. Uba. 109-115. & oo (1992b) A new species of trichostrongyloic nemude, Cuilin bere. from a vary rodent, Rurty Juacipes (Waterhouse) Thi 16, }23-128. & 484) Adult and tarval slapes of Puraasnostronentas maui (Nemiutidas Pichostrongylsideas trom: Karras juseipes, Ibid, WF, 27 46, Buu. ©. M., Bereacar, D. de Siargan. RB. (1989) No Competition fue resources between Wo fick species al the parupainic boundary. QGecvologia 79, 558562. Bris, A, O, & Nopstes. tC. 19867 lotesrinal helminths of lesser scHup.tlugks. an imeracnve sommunity, Can, £ Ave, 64 142-152. Chaaalo, AC (1965) Spécificité purusiture pp 548-557 tn Grassd, POP CE) "Trete die Zoologie Vol a, Chasm des Némuindes’ (Masson, Baris), 148 L. F. SKERRATT, I. BEVERIDGE & M.-C. DURETTE-DESSET (1982) Spectre dhdtes et Evoluiion des nématodes parasites de vertébrés. Mém. Mus, natn, Hist. nar., Paris 123, 73-16. Croui, N. A. (1976) The locaton of parasites within their hosts: the influence of host feeding and diet on the dispersion of adults of Nippostrongylus brasiliensis in the intestine of the rat. Jar. J. Parasitol. f, 441-448, & Ma, K. (1977) The location of parasites within their hosts: the influence of surgical manipulation of the intestine and mesenteric blood supply on the dispersion of Nippostrongylus brasiliensiy in the rat. Ibid, 7, 21-26, Durette-Desset. M.-C. (985) Trichostrongyloid nematodes and their vertebrate hosts: reconstruction of the phylogeny of a parasitic group. Adyan. Parasitol. 24, 239-306, Homes, J. C. (1973) Site selection by parasitic helminths: interspecific interactions, site seyregalion, and their importance to the development of helminth communities. Can. J. Zool. 54, 333-347, ____ & Price, P. W. (1980) Parasite communities: the roles of phylogeny and ecology, Syst. Zool, 29, 203-213, & (1986) Communities of parasites pp 187-213 dn Anderson, D. J. & Kikkawa, J. (Eds) “Community Ecology; Patlern and Process” (Blackwell Scientific Publications, Oxford), Hureert, 8, H. (1978) The measurement of niche overlap and some relatives. Ecology 59, 67-77. Martin, D. R. (1969) Lectthodendriid trematodes from the hat Peropteryx kappleri in Colombia, including discussions of allometric growth and significance of ecological isolauon. Proc. Helminthol. Soc. Wash. 36, 250-260. Mawson, P. M. (1961) Trichostrongyles from rodents in Queensland with comments on the genus Longistriata (Nematoda: Heligmosomatidae). Aust. J. Zool. 9, 791-826. (1973) Amidostomatinae (Nematoda: Trichostrongyloidea) from Australian marsupials and monotremes. Trans. R. Soc. 5S. Aust, 97, 257-279, Nope, E, R., Nogie, G. A.. ScHab, G. A. & Maclnnis, A.J, (1989) “Parasitology: the biology of animal parasites.” (6th edn.) (Lea & Febiger, Philadelphia), Orenporr, D, L, (1979) The helminth parasites of Rectus Jiscipes (Waterhouse) from Victoria, including description of two new nematode species, dust, J, Zool, 27, 867-879. Pianka, E. R,, Huey, R. B. & Lawnor, L, R. (1979) Niche segregation in desert lizards pp. 67-5 Jn Horn, D. J., Stairs, G. R. & Mitchell, R. D. (Eds) “Analysis of Ecological Systems” (Ohio State University Press, Columbus). Price, P. W. (1980) “Evolutionary biology of parasites” (Princeton University Press, Princeton, New Jersey), _______ (1984) Communities of specialists: yacant niches in ecological and evolutionary time pp, 510-523 /n Strong. D.R., Sinberloff, D. S., Abele, L. & Thistle, A, B. (Bus) “Ecological communities: conceptual issues and the evidence” (Princeton University Press, Princeton, New Jersey). Rouwpr, K. (1979) A critical evaluation of intrinsic and extrinsic factors responsible for niche restriction in parasites. Amer. Nar. 114, 648-671. (1982) “Ecology of marine parasites.” (University of Queensland Press, St. Lucia). Scuap, G. A. (1963) Niche diversification in a parasitic species Nock, Natyre (London) 198, 404-406, SOGANDARES-BERNAL, F. (1959) Digenetic trematodes of marine fishes from the Gulf of Panama and Bimini, British West Indies. Tulane Stud Zool, 7, 69-117. Spratt, D, M., Beveripor, I. & Warrier, E. L. (1991) A catalogue of Australian monotremes and marsupials and their helminth parasites, Rec. §, Aust. Mus., Monogr. Ser. 1, 1-105, Warts. C. HS. & Asus, H. J. (981) “° The Rodents of Australia” (Angus & Robertson, Australia). TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 19, PART 4 EIGHT NEW SPECIES OF AUSTRALIAN BUPRESTIDAE (INSECTA: COLEOPTERA) By §. BARKER* Summary Barker, S. (1995) Eight new species of Australian Buprestidae (Insecta: Coleoptera). Trans. R. Soc. S. Aust. 119(4), 149-156, 30 November, 1995. Six new species of Castiarina namely C. corallina sp. nov., C. ernestadamsi sp. nov., C. euknema sp. noy., C. octopunctata sp. nov., C. oedemerida, sp. nov. and C. prolata sp. nov., one new species of Themognatha, T. viridescens sp. nov., and one species of Astraeus, A, powelli sp. nov. are described. Key Words: Coleoptera, Buprestidae, New species, Castiarina, Themognatha, Astraeus. Traicdertons af the Reval Soerety of So Ast (99S). WY 149-456 EIGHT NEW SPECIES OF AUSTRALIAN BUPRESTIDAE (INSECTA: COLEOPTERA) by §. BARKER* Summary BaAkkeR, 5. (1995) Fight new speeies of Aostrahan Buprestidae (Insects, Coleapten) Trans. R. Sav, S. Aus, 119(4), 149-156, 30 November, 1995, Sik new apecies of Casriarina namely C. cerallina sp. nov, C. ernestadamsi sp. noy,, Co enkriertia sp tov, Co sctupunctaa sp. poy,, C eedemerida, sp. nov. and C. prelara sp. noy,, one new species of Themognailia, L viridescrns sp. nov,, and one species of Asiraews, dy powell sp, nov. are deseribed- Key Worps: Coleuptera, Buprestidae, New spevies, Confaeiia, Memoynatha, Astraeus Tatroduction Despite the increase in specialist collecting of Australian Buprestidae undertaken by amateur entomologists over the lust twenty years, new species are still being found. This reflects the vastness of the conlinen( as well as the cyclic nature of the lle histories of many of the arid area species, OF the species deseribed herein, specimens of Astrecus powelli have been available since 1970 but the specimens im the South Australian Museum collection were madvertemly sorted into a tray full of A, aberrany ¥, de Poll, the closes! species, und it was only through the alermess of Me M_ Powell thal this species has heen recognised. Jemounathe virtdescens was collected many years ago in inaccessible country ut Jron Range, Cape York Peninsula, A serics bas now become available from the Jack Macqueen collection, lodged with the Australian National Inseet Collection (ANIC), Three species. have only reeently been collected. Castiavina pedemerida and C. pralata in Queensland und C. corallina in Western Australia, Castiarind euknema was known lor some years from two female specimens and a male has only been located recently in the collection of ANIC. Castrarina vrtapuncrstia specimens have been collected frequently but until now have not been distinguished from C. paralleta. Until recently C, ernestadamsi has been confused wilh C. srramined Macleay, Materials and Methods Male genitalia were prepared and displayed by the method described by Barker (1987), The holotype ts ilustrated in all species. Measurements given are mean total body length and width with standard error, except Where there are insufficient speeiniens availible to niake the last calculation, Codens used in the texr for muscuntand private collections following the four letter * Departinent of Zoology, CU niverstry of Adelaide 5, Aust SOS. system of Watt (1979) and Arnett et al, (1993) ure: AIMS: Australian Tnstitate of Marine Science, Townsville; ANIC: Australian National Insect Cullection. CSIRO. Canberra: SAMA: South Australian Museum, Adelaide: WAMA: Western Australian Museum, Perth: SHIQ: Mr J, Hasenpusch, Innisiail; MASA: Mr T. M, S, Hanlon, Sydney; MPWA: Mr M. Powell. Meiville: GWQA: Mr G. Wood, Athertan. Castiarina corallina sp. cw. (FIGS 1D, 2C) Holawpe> oro \44 kin NW Wittenoom, W.A.. 25.11.1994, M. Golding, M. Puwell, WAMA. Allorype: 2, IO km NW Wiltenoom. W,A,, 25.11).1994, M_. Golding, M. Powell. WAMA. Faratypes; 2-2 9, Millstream Stn, W.A., 25,171.1994, M. Golding, M, Powell. MPWA: 1 co’. same data as allotype, MPWA: 4 cr cr. 145 kin NW Wittenoom. MPWA, SAMA, Colour Head. antennae dark blue Pronotum red-brown, in some specimens with medial smudge of dark blue along basal margin, Scutellum dark blue. Elytra red- brown with the following markings: dark blue along busal margin, in some Specimens with mark extended fosurround scutellum) large post-medial spot on each elytron, apical mark. Ventral supface: pre-sternum red- brown, meso- and meti-sternum dark blue: abdomen red-brown except laterally at base variably dark Blue, legs dark blue. Shape and sculpture Head closely punctured, broad median sulcus, short muzzle. Antennae. antennameres |-3 obconic, 4-11 wwothed, Pronotum closely punctured. narrow basal fovea, apical margin projecting medially, basal margin bisinuate; laterally pardllel-sided at base, ungled outwards, rounded to widest pre-medially, tapered to apex. Scutellum cordiform, glabrous, tat. Elytra punctale-sirtale, intervals conver, punctured: laterally 150 5. BARKER paralfel-sided al base, rounded ut humeral callus. coneuve rounded post-mediully. narrowed to spineless apex, apices hardly diverging, apical margin subser- rate, Ventral surface, shallowly punctured, edges of abdominal segments glabrous, withoul hairs, $,: upically rounded in both sexes, Stee Males. 4.3 + O48 4 5.4 + 0.14 mm (6). Females, 16.8 x 6.5 mim (3), Aedeavus (Fig. ID) Parameres angled outwards from basal piece, rounded apically, Penis pointed. sides acutely angled away. Apophysis of basal piece medium width, tapered, rounded apically, Remerrks The dorsal tnarkings of this species resemble those of Cusdarina quadriplagiata (Carter) which has only heen collected in Queensland. However, C. cerallina in 4 narrower species, it is a paler red than C. yuudriplagiata and the male genitalia differ (Pig. IC). Epymolagy Derived from L. eeraflinas, coral coloured. Castiarina vedemerida sp. nov (FIGS 1H, 2F) Holonpe- oo, Georgetown, Old, 284).1993, | Hasenpusch, SAMA I 21285, Paratype: Qld: lor. Georgetown, 10.1993, J Husenpusch, JHIJA. Colour Head black. Antennae dark blac. Pronotum brow with the following black markings: medial spot, smaller spot on cach side. narrow basal border, expanded anteriorly on each side. Seuteluin black, Elyva yellow-brown with the following black markings; M-shuped mark along anterior margin, ats covering humeral callus; post-medial fascia reaching margin. expanded anteriorly and posteriorly along suture, apical) mark. Ventrul surface: prostermum yellow-brown) meso-sternum and imeta-sternum and coxaue dark blue; abdemen yellow-brown, 8. with lateral black spol, §, with lateral black bar, 8, black apical edge, edges of S.. S,, S., testaceous: legs dirk blue. Hairs silver. Fig. 1. Photmicrographs of tnale acdeagi ot the following Casriarina, Astraeus and Themognarha species. A. Themognathe viridescens, B. Castturina acuminata Kerrenians, C. Custiarina yuadriplagiahs Carter. D. Casdurind corallina sp. nov. F. Castiarina prolata sp. noy. F Castiaring eciopunctaia sp pov, G. Astraeus powelli sp. nov. HL Castiarina cedemerida sp. nov, Lb Casdarina ernestedamsl sp. nov, J. Casnerina srraminca Saunders. K, Castiering eukneme sp, nov, |. Castaring rufipes MacLeay. Photomicrographs of procigers of types of Castiarina ectapanetata sp, poy, M. Holotype mile, N. Allotype temale. EIGHT NEW SPECIES OF AUSTRALIAN BUPRESTIDAE 151 Shape und sculpture Punctured over entire dorsal surfiee, Ted closely punctured, median suleus, short muzzle, Antennae. intennomeres +3 obconic. 4 Vs toothed, Sl toothed. Pronotum closely punctured, glabrous urea surrounding basal fovea, fovea at each angle; apical margin stright, basal margin barely bisinuate: laterally rounded from base to apex, widest pre- ~~ ~ i . f i {2 a " A ‘a | { i- . x Fig. 2. species, A.C ernesiadamsi sp. nov holoiype. B.C. yirigeta Habitus ilustrations of the following Crsiarinia Macleay, C. C caralling sp. nov, holotype. BO & Herp tit Sp, nov, holotype. b. ©. enknen sp. nov, holotype Fe) wedemerida sp. now holviype CC prolate sp. tov. holotype medially. Seutellum scutiform, punctured, excavate. Elytra punctate-striate, intervals convex, heavily punctured; laterally angled out from base, rounded at humeral callus, concave. rounded post-medially at widest part to bispinose apex; small marginal spine, sinall sutural spine, margin rounded and indented belween spines, apices slightly diverging. Ventral surface closely punctured, edges of abdominal sezments glabrous, few short hairs, S,: males truncate, indented medially; females unknown. Legs: in male tarsomeres 1-3 without pulvilli, replaced by median longitudinal ridge. Sice Males, 10.0 x 3.5 mm (2), Aededazus (Fig, |) Parameres angled outwards [rom basal piece. rounded at apex. Pems blunt, sides acutely angled away. Apophysis of basal piece medium width, tapered. rounded apically, Remarks This species appears to be an oedemerid mimic as its colour and pattern are similar to known oedemerid Species; the model is unknown. It is not close to any other known species. Etymology The name is derived from that of the beetle family Oedemeridae. Castiarina euknema sp. nov. (FIGS IK, 2B) Holotype: o , Surveyar’s Pool, W.A., iS.vili.i983, I, PD, Naumann, J. ©. Cardale, ANIC, Allotype: 9, 2 kin N Jabiluka, N.TL. 21111981, M, Cappo. SAMA I 21286, Furatype. WA.: 9, 34 kin 5S Roebuck, W.A,, Bwii. 984. M. Powell, M, Golding, MPWA, Colour Head black wilh yellow reflections, elongate yellow frontal spot. Antennae blue-green. Pronotum: anterior and basal margins black with yellow reflections: yellow laterally: medial black mark with yellow reflections in the shape of sleeveless T-shirt. the neck enclosing avery small yellow spot. Scutellum black with yellow reflections. Elytra yellow with the following black markings: narrow busal margin; pre-medial fascia with ends expanded anteriorly over humeral callus reaching anterior margin and enclosing yellow basal spot, posteriorly reaching margin and enclosing yellow spot on margin; post-medial fascia reaching margin and Necting posterior extension of pre-medial fascia and with iL enclosing yellow medial spot; mark covering whole apex, clongate yellow mark reaching margin, but nol suture, lying between this and post-medial 152 & HARKER fuscia, In allutype black marks less prominent and upial tnark encloses a small yellow apical spot., Ventrul surlace yellow, edges of sutures black with green reflections, 5,, 5. 8, with lateral spots coalesced with marks along edges, Legs: femora and tibia blue; larsomeres durk blue, Hairs silver, Shupe and seulplire Head shallowly punctured, shallow median sulcus, medi Jength muzzle. Antennae compressed, antennomeres |-4 obconic, S-Il toothed, Pronotum Shullowly punctured, basal fovea extending forwards to middle as glabrous line, basal notches on each side closer to margin than middle; apical margin straight, basil margin barely hisinuates laterally rounded from base. widest piv-medially, rounded and natrowed tu apes. Seutelleny seutiform, without punctures, excavate along anterior margin, BLyWa pufctale-striate, intervals flat anteriorly, convex apically; laterally angled oul from base, rounded wt humeral callus, concave, rounded post-mediully and narrowed to Irispinose apex; marginal and suturul spines equal, medial spine Slightly larger, margin rounded und indented between spines, apices shightly diverging, Ventral surlace with shallow punctures, edges of abdominal segments wlubrous, few short hairs. 8, truncate both sexes. Size Male. 10.9 x 4.2 mn (1). Female, 1.9 x 425 mm (2). Avdvuyus (Fig, WK) Parameres parallel-sided from basal piece, rounded medially then purallel-sided, rounded at apex. Penis pointed, sides angled away. Apophysis of busal piece medium width, lapered, rounded apically. Remarks This species has similar markings to those of C. rifipes Macleay, except that the legs are blie whereas they are ted in the other species, The male penttala ure quite different (Pig. 1L), Eryvinology Derived from Gr. enknemes, beautiful legs. Castiarina prolata sp. nov. (FIGS IE, 2G) Holotype, Of, Cardwell Ra, Qld. 23, 9,1993, J, Husenpusch, SAMA 1 21287. Alloipe: &. sume data as holotype, SAMA | 21288, Pararypes: | or, | 9 same data as holotype, JHOA, Colour Head and antennae black with green and gold reflections. Pronuotum black with green reflections ut hase. Scutellum) black with blue reflections. Flytra yellow with black inarkings coalesced forming ao elongate yellow basal mark. a round yellow pre-medial mark and a preduminantly red subeapical mark, louching Margin not reaching suture and merginy ute yellow closest to it, Ventral surface green with yellow reflections, Legs; femora, dorsal surface deep blue, ventral surlace green wilh old reflections; tibia and tarsi dark blue. Hairs silver. Shape and sculpiure Head shallowly punetured.. broad median suleus, shor muzzle, Antennae, antennomeres 1-3 obconic, 4-11 toothed. Pronotum. shallowly punctured, glabrous, small basal fovea, larger fovea at each basal angle; apical margin projecting slightly medially, basal margin bisinuule; rounded from base, narrowed to apex. Seutellum tulipiform, few punctures, glabrous. fat. Elytra anteriorly punctate-striate, intervals raised postenorly, intervals convex; laterally parallel-sided at base, rounded at humeral callus, rounded and tapered posteriorly, then attenuated, rounded at apex (o pointed marginal spine, deeply indented and rounded to minute sharp sutural spine. apiwes diverging. Ventral surface with shallow punctures, edges of abdontinal segments glabrous, Sparse medium length hairs. Mesosternal process inflated, Elytra. cuntlevered oyer last visible abdominal sternite, Legs: femora and tibia long and thin: tarsomeres with enlarged pulvilli. $5: truncate both sexes. Sine Males, 10-9 x 3.4 mm (2). Females, 12.4 x 3.6.mm (2) Aedeugus (Fig. 1E) Purameres elongate, angled outwards front basal piece, apically rounded, Penis pointed, sides obtusely angled away, Apophysis of basal medium width, elongate. tapered. rounded apically. Remarks This species 6 a 'C. produvta’ group mume. lt resembles C. acuminata Saunders which also has urdatly attenuated apical spines but is heavily costate Che apical spines of C prelata are both obvious wherews in C. geuninara the marginal spine is absent. Male venilalia differ (Pig 1B)- Erymalogy Derived foun L. predatus, elongate. Castiarina ernestadamsi sp. nov (PIGS (1, 2A) Halatype: oO. Mourangee. Edungalba, Qld, 26.41.1979, E. E. Adams, SAMA T 21289. Pararypes: Qld: 3 o, Mourangee. Edungalba. 8. A Adanis & BF. b Adan, SAMA. Colour Heud maroon at base blending into blue-green, blue muzzle. Antennac, antennomeres [-2 blue-green. 3-1 BIGHT NEW SPECIES OF AUSTRALIAN BUPRESTIDAE, In bronve green. Pranotum maroon, Scutellum blue or blug-gecen, Elyira pale yellow with lollawmy dark blue nuirkings: darrow basal margin; small mark over cach humeral callus; remnant post-medial fascia touching jatcral margin reaching hall way to suture, small remnant murk ofsuture, small apical mark. Ventral surhice: Slernum muroon, abdarien teskiceous, lees blue. Nias silver Shapy and sculplure Head shallowly punglured, medium sulcus, short mazele. Anteunac, antennomeres LF obeonic, 4-11 toothed. Pronotum shallowly punctured, basal foves extending forwards lo middle us globrous line, basal notetes represented by glabrous area on each side closer to mhargin (han quddle; apical margin projecung mediilly, basal margin barely bisinuate: Taterally parillel-sided at base. angled outwards and rounded ta widest pre medially, rounded and narrowed to apes, Sculetlum scutiform, glabrous, flat, Flytra punctate- striate. intervals convex. lightly punctured; laterally angled outwards from buase, rounded at humeral cullus, voneave, rounded post-medially and narrowed to bispinose apex; both spines small and sharp. margin rounded and indented between spines, apices hardly Jiverging, apical margin subserfate, Ventral surface with shallow punctures, edges of abdormmal seetnents glabrems, stermum with few medium length hairs, abdomen almost hajrless. S,: males truncate, female unknown, Site Males, 24 + 0.09 x 4.8 + 0.04 mm (4), Aedeagus (Fig. 1) Lightly tanned. Purameres parallel-sided [rom busi! piece. pre-medially rounded, parallel-sided then rounded lo apex. Penis pointed, sides obtusely angled away. Apophysis of basal piece narrow, rounded ayneatly Renuirks This species was previously confused with ©. siraiminea MacLeay. However it ig smaller, it has i smooth pronotum with small punctures whereas C straminea bas heavy punctuation. I has darker elytral markinys than C. straniinea and the male genitalia differ (Fig. Wi. A form of C. strigara MacLeay (Pig. 2B) occurs mm the type locality, sinnilar in size and pattern but not in colour tr has very light elyeral inarkings, the post-medial fiscia being considerably reduced. The head and pronotum are green with reddish reflections in some specimens: Ltymolouy In honour of Mr BF, ©. Aduims. octogenarian. Edungalba, Queensland who has assisted my research for many yeurs, Castiarina oclopunectata sp. nov. (FLGS IF, 2D) Holonpe: o 9 km NNE Zuothus, WA. 21x 1986, M. Powell, WAMA Allorpe: G. Wialka, W.A., 2L.1%.1970, S. Barker, SAMA J 21290. Paratypes: W.A.. | co SAMA: | 9. summit Mt Cooke. 10.%7,1956, S. Barker, SAMA, 1 9. Wialkiv 1k.ix.1957, 8. Barker, SAMA; 2 cor, Kalbarri N.P.- 23.4x.)969, FH, Uther Baker, SAMA, 4 orot, 2 9 @, Beverly T. 0.. Brookton Hwy, 19,ix.1970, 8, Barker. SAMA; 2 Oc, sume data as allmype, SAMA, 1, 6 kin S.Tammin, § x1.1970, §. Barker. SAMA: 3 oo, 3.0 9, 1/2 way between Glen Eagles and Brooktou Hway. 13.x 1980. 8, Barker. SAMA: 1 9, & kin 6 Woolgangie. 22.%.1980, S, Barker, P. G. Kempster, SAMA: | oy. 10 kan E Merredin. 12.4, 1990, §, Barker. SAMA? | ot, 4 km W Zanthus, 2!.4.19%6, M- Powell, MPWA, I of, same data as holotype, MPWA; 1 4). Moora, WAMA: | ©, MeDermd Rock 27.1%/3.%.)978. 1. F Houston, WAMA; 2 oor, 1 @. Lake Ningham, WAMA; | 9, Merredin. WAMA; | o, Karlgarin, W. Duboulay, WAMA: | 9, Dedari, 7.x.1978, T. M.S. Hanlon. WAMA; I @. 21 km W York. 4.xi.1978, T, M.S. Hanlon, WAMA; 1 oF, 1.5 kin S Mt Jackson, 5/11.34.1979, TF. Houston ef al. , WAMA; |oc, | 9, 12 km NNE Bungalbin Hill, H/tB.ax1979, T. B Holston eral, WAMA: 1 oo, | @, 14km NNE Bungalbin Hill. WiS in 979, T. F, Lluuston ec: al. WAMA: 3 oo, | oD, 18 kin NNE Bungalbin Hill, W/18.1%.1979. Tl) Fo Houston ef al), WAMA; 2 © Q, Dedari, 21.in.1979. 1. M.S, Hunton WAMA; | c. Mt Dale, 20x, 1980, 'T. M.S. Hanlon. MHSA; | o, Muckinbuddin, 10,x,1978, RB. P MeMillan. WAMA: | ot, Mt Walker. 34 kin b Narembeen, 23.«, 1980, R. PB. MeMillan. WAMA: 3 co. | @, Mt Observation, 2bx.1987. Ro P McMillan, WAMA: | 9. Beacon, 20.%.1981, R. PL MeMillan, WAMA; L Q, 20.1%, 1990, Bonnie Rock, S. Barker. SAMA: | o&, 10 km E Mervedin, 12, 7.1990, S. Barker, SAMA: 3 cr ol 2 9 @. Dedari, 22.«.1991, TM. S. Hanlon. MHSA. Colour Head. antennae. pronotum dark coppery-purple. Seutellum dark blue with coppery-purple teflections. Elytra yellow with coalesced dark blue markings with coppery-blue reflections forming the following yellow marks: 4 large medial spots in a row on each elytron, the basal, pre-medial and post-medial roundish, the pre-upical clongate: narrow margin from base. not reaching apex. Ventral surface and legs brown with coppery-purple reflections. Hairs silver. Shape and sculpture Head closely punctured, broad median sulcus, short muzzle. Antennae, antennomeres 1-3 obconic, 4-Il ist S BARKER louthed. Pronofum) closely punctured, basal levea estending, forwards to middle as glubrous impressed Hiessupleal margin projceting broadly medially, basal Htrgin almost straight, laterally parallelsided al base, slightly rounded, widest medially. slightly rounded to apes, Sculellum scutiform. wabrous, excavate. elytra puneRde-striate, intervals Convex, smooth medially, punclured and rough laterally: luterally angled out from base. rounded at humeral callus, concave, rounded Post-medialhy ond arrowed t spineless apex, last iMerval indented and straight, apices slightly diverging, Eniire ventral surface covered in dense. flat hairs, alse present uround Jaterul margins of pronotum and in SOME specimens cnervaching onto dorsal surtiwe, §,; Males truncites females rounded und slightly pointed, Size Miles U9 + 016% 4.0 +4 0.06 min (35), Females, 12,7 = Ut x 4.2 + 0.07 mm (25) Adedeagus (Pig. UF) Parumeres angled oulwards and gradually widened from basal piece, rounded at apex. Penis pointed. angled away ebtusely, Apophysis of basal piece medium width, rounded apically. Proctiger with apex bluntly bilobed, the two projections variable (Fig, IM). Female terminatia (Fig, IN) Proctige?; apex with Iwo narrow, pointed lobes. Remarks A species complex exists in Western Australia whieh includes C) parallela (White) with a more or less continuous gradation in size from the smallest (C parallel) to larger species. C. paratlela also occurs mousiorn Australia, Until now T have not been able jo separate the individdal species in the western conmiplex, ©) vawihepilosa Hope and C yin Saunders dre closely related species but only oecur in castern Australia. From examination of the terminal abdominal SOHNE, it now appears as if males and females of Hoth castern and western specimens of C) parallels have a rounded proctiger and can be distinguished on (hal basis and on (heir colour size and structure of male genitalia, Both sexes of the larger western species huve ao ormamented proctiger CC. vetpunetata 1% distinguishable on the basis of the structure of male genitalia and the lack of apical spines on the elytra, Further work ts required to delimit the remaining species Ervnnligy! The name is derived from b eeto. enht und 1, PuUHCTATO spontea, Themognatha viridescens sp. nov, (FIGS 1A, 3) Holotype. , Tron Ran, SAMA I 2129), Old, 20.vi 1980. Wood. Allotype: 2. Tron Ra., Qld, 30.1.1966. J, Kerr, ANIC, Fararypes: Qld: 29 @, bran Ra., 3/9.V.1966, J, Kert, ANIC, 1 a. Iron Ra. 24.v,1974, M. Walford Huggins. MHSA; | &, tron Ra., Qld, J9v1978. G. Wood, GWOA; [sex indeterm. 5.1966. J. Macqueen. J. Keer, ANIC. Colour Head, antennae. pronotum and scutellumn bright green with yellow reflections. Elytra: yellow with following dark green markings: narrow basal margin; medial fascta not reaching margin, apieitl mark connected along sulure to fasent. Ventral surface muinty bright green with yellow refleetions, male with tesiaceous patches on S, and S., abdomen all green in female; legs bright green with yellow reflections. Shape and sculprare Head punenition fine, even, dense, median impressed basal line, trons moderately hairy; labrum longitudinally divided and pointed. Pronotum narrower than elytra, LeW 0.6, punetation fine moderately dense; sides rounded from base to 1/3 distance to upex, then explanate and converging anteriorly, laterally flattened; untenior margin bisinuate, posterior margin almost straight median glabrous line from base to near apex Scutellum scutiform, anterior margin straight, concave without punctures, 1/8 width of elytra. Elytra slightly wider than thorax; elytral interneyrs long and with scutellary striole, strongly marked with heavy punctition and additional punctation on shoulders: intervals flit, sides sub-parallel then tapering to pre- Apical areas; apex bisinuate, both spines prominent, interval hetween sinuous, lateral spine anterior 1 medial spine, Ventral sulface: prostemum hairy, finely punctured, with a defimte forward medial projection; pro-episternum finely punctured with deep smooth fossa in posterior angle; mesusternum und meta- sternum smooth medially with coarser punctation laterally; hairy, hair long and fine medially, shorter Mv ; inal ‘ f omm Fig. 3. Habitus Hustation af Themownatha varidesceny sp. nov. Holotype EIGHT NEW SPRCIES OF AUSTRALIAN BUPRES TIDAL 155 and coarser laterally: abdomen smooth and shiny in nine, punctation very fine, apieally huiry in female, S.: male deeply concave, fernale rounded. Feet: tarsal claws without a notch, Size Male, 31.4 x U4 mm (2). Ferales, 30.7 « ILS mm (4), Acdeauus (Fig. 1A) Parameres parallel-sided trom basal piece, ungled outwards premedially, rounded al apex. Penis pointed, sides acutely angled away, Apoptiysis of basal piece medium width, tapered, rounded at apex. Remarks This is un unusuul species as ft shows two characters found in the related genus Caledema C&G, I hits a small medial projection on the anterior margin of (he prosterum. but not as large as those found in, Cule- dema. The seutellum ts wider thar in obher Thenig- athe in relauon to the elytral width, although not as wide as any of the known Calodema species, However, the body is not sinuous in lateral profile, the pronotum is nous wide as the elytra and the elytral interneurs ure cleurly defined as in Themognadie. The tarsal claws ure not notched, bul this is a variable character found in some Themognatha and not in others, Etymology Derived from L. viridis, green. Astraeus (Depallus) powell sp. ny, (FIGS. 1G, 4B) Holotype: , Quaitading, 7,X1,1970. S, Barker, SAMA I 21292, Allorype: © , Quairading. 26,1 1991. M- Gniding, M. Powell. WAMA, Paratypes: WA: 1 or, Tammiin, §.xi1970, S, Barker, SAMA; | co, 17 km E Dowerin, 21.4.1989. M. Golding, M. Powell, MPWA; 1 cr, 6 km SE Tanunin, 15.xii.1990, M. Golding, M, Powell, MPWA, | 0”, 43 kin E Merredin, 26.8199], M. Golding, M. Powell, MPWA;: 2 oct, 32 km EF Yellowdine, 21/22,x.1991, T. M.S, Hanlon; 2 9 9, Quairading. 27x11). 1991, K K.. MPWA: 4 & 9. Quairading, 11.1992, M. Golding. K, K.. MPWA. Colour Head. antennae. pronolum black with blue and purple reflections. Elytra black with following yellow murkings: irregular marks ulong width of one strite, roainly concentrated along 4th and 8th intervals from sulure and along the murgin on basal half. Ventral surface and legs purple: lateral yellow spots on S,,'5,, S$, in MUS! Specimens last one absent in holotype. Hairs silver. Shape and xculptnre Head punctures small medially, larger hiterally, small glabrous median keel near apex merging into impressed hig, 4. Habitus illustrations of the following Asiraens species A.A, dbberans \ le Poll. B. 4, panel/esp, noy, holotype line basally, Pronotum punctures smaller medially, larger and in form of shallow fovea laterally, glabrous median urea; anterior maunsin projecting medially, basal murgin strongly bistnuate, laterally fhajry. Elytra intervals between striae Convex and smooth, [aterully parallel-sided from base, rounded post-medially und narrowed to apex, small pre-apiecal notch. on margin. hroad outwardly curving satural spine, hairy overall, Ventral surface and Jegs punctured und Hairy. Size Males, 12.6 + 0.22%44 + 0.11 min (7). Females, 15.3 + 0.38 x 5.6 4 (12 mm (7). Aedeagus (Fig. 1G) Parameres parallel-sided from basal piece, gradually widening until rounded lopointed upex, Apophysis ol basal piece medium width, rounded apteally. Remarks This species has been contased With 4a, uberranys \ de Poll (Fig. 4A). Ut differs fron) that species by being narrower, haying most of the yellow elytral marks medial and post-medial whereas ind. aberrears they are more evenly distributed, by having single outeurving sutural spines on the elytra while im A. vberrans the sutural spines dre small and there is a definite small marginal spine, Envnolowy In honour of Mr M. Powell of Melville, W.A. who has assisted my research for many years. With the addition of the above new speutes the key to Astraeus (Depollas) (Barker W978, p.j07) requires the following replacement: Replace 4. uberrans van de Poll with; 4a, small, ulmost straight marginal spine Sibert eye pees ey yey ye tberrans Vary de Poll 4b. broad, oulcurving marginal spine Meg OR me eT ..pawelli Barker With the addition uf this species the sub-genus Depalliis now contains nine species, 156 S. BARKER Acknowledgments I thank the following for their assistance: Dr T. F. Houston, WAMA; Mr T. A. Weir, ANIC; Mr M. Cappo, AIMS; Ms H. Vanderwoude, Department of Zoology, University of Adelaide. I thank the following collectors for the loan of specimens: Mr E. E. Adams, Edungalba; Mr T. M. F. Hanlon, Hunters Hill; Mr J. Hasenpusch, Innisfail; Mr M. Powell, Melville; Mr G. Wood, Atherton. References ARNETT, R. H. Jr, SAMUELSON, G. A. & NisHtpA, G. M. (1993) ‘The Insect and Spider collections of the world’ 2nd ed. (Sandhill Crane Press, Gainsville). BarRKER, S. (1975) Revision of the genus Astraeus LaPorte & Gory (Coleoptera: Buprestidae), Trans. R. Soc. S. Aust. 99, 105-142. (1987) Eighteen new species of Stigmodera (Castiarina) (Coleoptera: Buprestidae). Ibid 11, 133-146. Watt, J. C. (1979) Abbreviations for entomological collections, N.Z. Zool. 6, 519-520. A NEW SPECIES OF CALLULOPS FROM NEW GUINEA AND COMMENTS ON THE STATUS OF C. HUMICOLA COMPTUS (ZWEIFEL) (ANURA: MICROHYLIDAE: ASTEROPHRYINAE) By STEPHEN J. RICHARDS*, THOMAS C. BURTONT, MICHAEL J. CUNNINGHAME & ANDREW J. DENNIS* Summary Richards, S. J., Burton, T. C., Cunningham, M. J. & Dennis, A. J. (1995) A new species of Callulops from New Guinea and comments on the status of C. humicola comptus (Zweifel) (Anura: Microhylidae: Asterophryinae). Trans. R. Soc. S. Aust. 119(4), 157-162, 30 November, 1995. Callulops sagittatus sp. nov. from the summit of Mt. Binnie, Western Province, Papua New Guinea is described. It is a moderately large species (males 44.1-47.8 mm, a female 56.3 mm S-YV) distinguished from congeners by the presence of an orange stripe from the tip of the snout dorsally across each eyelid, forming an arrow-shaped mark on the crown. The advertisement call is a series of 11-12 deep croaks uttered from the entrance to, or deep within, crevices between rocks. The female paratype contains large (4.5 mm diameter) unpigmented eggs indicating that, like other Australopapuan microhylids, larval development is completed with the egg capsule. Among the Asterophryinae, Callulops sagittatus and C. h, humicola share a unique condition of the mandibular branch of the trigeminal nerve suggesting that C. sagittatus and not C. h. comptus is the closest relative of C. h, humicola. This and a number of other consistent morphological differences indicate that C. h. comptus warrants elevation to specific status. Key Words: Anura, Microhylidae, Asterophryinae, frog, new species, Callulops sagittatus sp. nov., Callulops humicola, Callulops comptus, New Guinea. Ieansactiuns of the Roval Sovivey of 8. Aust. (1995~ 194) [STAB A NEW SPECIES OF CALLULOPS FROM NEW GUINEA AND COMMENTS ON ‘THE STATUS OF G HUMICOLA COMPTUS (ZWEIFEL) (ANURA: MICROHYLIDAE: ASTEROPHRYINAE) by STEPHEN [. RICHARDS* THOMAS C. BURTON]. MICHAEL J, CUNNINGHAM & ANDREW J.. DENNIS* Summary RICHARDS, SoJ.. Burron.T, C., CuNsiGiam. M. J & Onis. A, J. (995) Anew speeres oF Calladaps tem New Guinga and comments-on the stars of C. fnivote complus (Aweilel) (Anurat Microhytidue: Asterophiry inte). Trans. R. Soe S, Aust O94) 1587162, 30 November 1995 Callalaps sagivatas sp. poy. fron) the sununit pF Mi Binnie, Western Provinee, Pupusa New Guinea iydescribed. Iisa moderately large specios (males 44-478 Wing a temale 56.4 mm S-V) distinguished Mont eongeners hy the presence of an orange stipe from the tp of the snout dorsally aeroxs each eyelid, forming ari arroweshaped mark onthe crown, The advertisement call is a series af 1-12 deep croaks uttered from the enitianee to, or deep within, crevices beiween rocks, ‘The female paratype contains large (4.5 mm diameter) Unpigmented eggs mdicating that Tike other Australopapuan microhylids, larval development is completed within the egg capsule, Among the Asterophry imac Cedlalops sawitatas and ©.) huwnicola-share a unique condition of Ihe mand ibulur branch of the trigeminal nerve Suggesting that ©. sagi(aias and not © Je Compras isthe elosest relulive of Coy diarealia This and a number of other consistent morpholoieul differences tulieate That CL A. errngrtiey Warrants elevation tO Specific stitus, Key Worps: Amira, Microhylidac, Astérophryimac. fray, new species, Cilludops sequiniotas sp now, Ceaflatepes humicola, Calluleps comptis, New Guinea. Introduction Microhylid trogs of the subfamily Asterophryinac ure restricted to the New Guinea. miumtiland and nearby islands (Zweilel & Tyler 1982). This ecologivgally und morphologically diverse graup occurs from sea level to subalpine meadows hyzh in the central cordillera (Zweite! 1972), Ln a review of the Asterophryinae Burton (1986) recognised eight penera and 43 species. Blum & Menzies (1988) subsequently deseribed nine new species of Xenebairachus and Xenorhina, and Richards ef a/, (994) described a new species of Axteroplrys, bringing the total to 53. Additional undescribed species. uccur in useui collections, and field work continues to reveal Unnamed taxa. Durning a survey of the fauna Of Mt Binnie, Western Provinec, Papua New Guinea (Dennis ef al 1995)! three of us (SR, AD, MC) collected an undeseribed species of the asteruphryine genus Callulops. The discovery of this new species peeessitates a reassessment of the relationships of the taxa currently recognised as subspecies of Callulops humicola, Here “ Zoology Department, James Cook University, Lownsville, Qld 481. { Division of Biological and Chemical Scienves, La Trobe University, Bendigo, PO Box 199, Bendigo. Vie. 3550. + Zoology Deparment. University of Queensland, Qld 4072. 1 Dennis, A., RICHARDS. S. & CUNNINGTIAM. M. (1995) Preliminary survey of nyarimals, birds, reptiles and frogs on the summit of Mt Binnie, Western Province, PNG, 20-23 November, 194. Report to Ok Tedi Mining Limited (unpubl, ) we deseribe the new species wad demonstrate: thal Callnlops lunmicola camptus warrants elevation i specific status. Materials and Methods Specumiens are deposited in the Biology Departinent. Universily of Papua New Cjuinea, Port Moresby: (UPNG) and the Queensland Museum, Brishane (QM). Recordings of mating calls were made in the field with a Sony Professional Walkman tape recorder with aw Eleetret Condenser Microphone ECM-Z200 und were analysed using the sound dnalysis progrun ~Canary” (Cornell Grnithology Laboratory, 1994). Measurements were made to the pearest 0.05 mm with cial callipers or to the nearest 1 mm using a binocular oneroscope with aq ocular muicromeler, Methods of measurement follow Zwejfel (1985) except the snout-naris measurement, laken from the tip of the snoul to the cenire of the nurs, Measurements. (min) were: snoul-vent length (SV); tibia length (TL); eye diameter (EYER): eve-naris distance (EN); internarial distance (IN); sneut-naris distance (SN): head width atungle of the jaws (HW): head length from tip of snout to angle of the juws (HL); horizontal diameter of tympanum (EAR), hand Ieogth (HD); foot length (PT). The rather featureless palmar and plantar surfaces of the bands und feet. and the poorly defined tympanic annulus made measurement of the hands, feet and tympanum difficull, and the measurements should be trealed with caution, Ope of us (TCB) dissected the superficial throat and jaw musculature under a Wild M3Z microscope with the aid of topical application 13% *& J RICHARDS, T. C, BURTON, M J CUINNINGITAM & Ad, WINES of the jodime-potassium iodide solution of Bock & Shear (1972), Phe squamosal bone was also examined, Systematics Callilops Boulenger \s applied Ww asterophryine microhylid frogs formerly referred to Phryiwmeietiy Peters, following the recommendation of Dubois (L988). Callulops is distinguished by two skull characters: two supplementary slips lo the M, intermandibularis arise from the dentary: one via a tendon and the other directly, and run together. more or less parallel to the mandible, to insert upon the ventral fascia of the M, submentalis and semetiines also upon the adjacent medial aponeurosis of the M intermandibularis (Burton 1986), The second character is (hat the otic ramus of the squamosal bone is about the same length as the zyomutiv ramus, and itis not twisted 1.c. the postero-lateral surface of the otic ramus rs.continuous with the lateral Surface of the zygomatic ramus (Burton 1986). Tr his revision of the asterophryines. Zweitel (1972) described Callulops li. humicola and Ch. compres. These taxa resemble each other superticially apart from relatively longer legs in Ch. fumicela, and an orange postocular stripe in adult C. A. compius which is only “somewhat developed in young humicola” (Zwerlel 1972 p. 476). The geographic ranges of these taxa abut. Zweite) reported only one instance of symputry, and was reluctant to assign the taxa fo species status in the absence of evidence of reproductive isolation. Burton (1986) added two further characters (o distinguish the taxa. First. in Cf Aumieala the M depressor mandibulae arises from the dorsal fascia, with some fibres from the otic ramus of the squamosil and the posterior surface of the adjacent prootie, in Ch. comptas and all other Calluleps, additional fibres arise from the entire posterior and ventral surfaces of the tympanic ring. Second, in CA. Juomivola the mundibular branch of the trigeringl berve passes directly ventro-laterally through the M, adductor mandibulae posterior longus on its way tw the THandibWear musculature; in CA, comprus and all oiher asterophryines this nerve passes antero-laterally between the M. a. m. posterior longus and the M.m anterior longus, and then postero-ventrilly aeross the lateral Surface of the M. a. i, posterior longus belore plunging towards the mandibular musculature, Burton (1986) made no taxonomic recommendation regardiny the status of these taxa. Callulops sagittatus sp. nov. (FIGS 1-5) Holotype: UPNG 90ST an adult male collected by 8, J, Richards, M. Cunningham and A, Dennis on Albay 1994 at an altitude of 2200 mon the summit of Mt Binmte. Western Province, Papua New Guinea 4b" 7 MY"E, S° 12'S). Fararypes, UPNG 9052 (adult female), OMI 6231 (udule indie), sume daig as holotype Definition A muderely large and robust species (males 44,147.86 itn SY, a fermale 56.3 mm S-V) distinguished froyn congeners hy a combinalion of the following charuclers; fingers and toes witout expanded dives. 4 dISTINeT ohinge stripe dorsally wn the head from the snol extending deross cach eyelid, lympanurn Indistinel, adverisenient call a senes of deep “crawks" wilhatslominiml frequency of 609 He, a pote reperitiin rate of LRS-2.2/s and a pulse rate of 189-14 02/ms Fig, 1 Crtudaps sagiindnas ap now. in life (SV 569 pi) Deycriplion of halornpe Body robust, almost pear shaped (Mig. 1), head brouder than long (MW/HL 1-23) with nares-closer to tip of snout than to eye (SN/EN 0.6) and directed Jaierally, Internarial distance greater than distanee from eye to nares (EN/IN 0.78), eyes large (EY E/S-V 0.119). Snout blunt, hrowlly rounded in dorsal wiew and rounded if: lateral view Cig, 2), Cunttius rostralis ruunded, lorcul region steep, slightly coneave. Tympunum indistiner. wnnulus barely visible. Bursal and ventral surfives tminutely gtanular. a weak supralympamne fold. Anterior palutal ridge long, smooth, posterior palatal ridge with 1) digtinet dennicles. Limbs short (TL/S-V 0.38), relative lengihs of fingers 3>4>2>1), fourth finger only marginally longer (han second, Fingers unwebbed, tips without expunded discs, subunicular tubercles low, rounded, Palm smooth exeepl for a low toner metacarpal oiberele, Relutive lengths of mes 4>3>5>2> |. Toes A NEW MICROHYLID FROG FROM NEW GUINEA 159 unwebbed, lips without expanded discs, subarticular lubercles low, rounded, A low, oval inner metatarsal tubercle; no outer tubercle (Fig. 3), Colour in hfe uniform deep red-brown dorsally on body and limbs, grading laterally into a uniform lighter brown ventral surface. Throat slightly darker brown than rest of venter. Head deep red-brown with an orange stripe dorsally from lip of snoul along eanthus and over eye, forming distinet arrow shape on crown. Slight orange tinge on upper surface of thigh. No other inarkings dorsally or ventrally, In preservative brown with a mauve Hinge dorsally, brown ventrally, stripes on head very pale pink. Fig, 2. Views of head of Callulops sagittanus sp. noy. holotype (UPNG 9051), A, Dorsal view, B. Lateral view, Seale bir 10 mm, Fig. 3. Hand and loot of Callulipy saginarss sp. nov. holotype (UPNG 9051), A, Plantar view of foot. B. Palmar view ol fand. Seale bar = 10 mm, Dimensions of holotype S-V 47.8; TL 18.2: EN 3.3; SN 2.0; IN 4.2: BYE 5.7: HW 17,0; HL. 13.8; HD 12.5: PT 19.3; EN/IN 0.785; TL/S-V 0.38; HW/HL 1.23, EYE/S-V 0.119; HW/S-V 0.355; width of toe tip on fourth loe 1.0 (width of penultimate phalanx 0.8), width of toe tip on third finger O.9 (1.0). Musculature The superficial throat musculature and squamosal form conform to the definition of Callulops. The M. depressor mandibulae arises predominantly from the dorsal fascia, but also reccives substantial contributions from the otic ramus und the posterior and ventral margins of the tympanic ring. The mandibular branch of N. trigeminalis passes directly yentro-laterally from the brain. case and penetrates the M, adductor mandibulae posterior longus on its way to the mandibular musculature (Fig. 4). Advertisement call We recorded two call sequences butonly one of these is of sufficient quality for detailed analysis. The mating call is a series of deep, guttural croaks “erawk, crawk, crawk.... The recordings contained Il and 12 notes lasting a total of 5.57 and 5.186 seconds respectively (note repetition rate = 1.85/s and 2.2/s). Both calls had a dominant frequency of 609Hz. Individual notes in the Il-note call lasted 118.9-182.3 ms (mean = 154.2), contained 10-13 pulses (mean = 11.63) at a rate of 11.89-14.02/ms (mean = 13,23). Fig. 5 illustrates the first four notes of an H-note call recorded at the type locality on 20.xi-94 at an air temperature ol} 13.35°C., 1a) S.J RICHARDS, T, C. BURTON, M Fiy 4 Dotso-fateral view of musculature of right jaw of Culluleps wagicanis sp, nov, A.M. adduct imandibuliue anteriog Tongus: D. M-_ depressor mandibulde; PM ahluctir mandibulae posterior longus: 8. M. adducror niandibulie extemus superticialis (severed); Vo mandibular branch of trveminal nerve. Scale bar = 5 mm. The calls were uttered at ipregular intervals, willt long periods (often over five minutes) between calls. On several occasions we heard 3 melodious, dove-like “coo cow..." vocalisalion utlered immediately following, or from the same vicinity as. one of the call types deserihed ubove but we were unable to confirm whether tL was part of the vocal repertoire of this Species, Natural history The type series was collected in disturbed rainforest and secontary regrowth at altitudes over 2000 m alung jhe access road to the summit of Mt Binnic. One male was calling from the entrance to.a deep crevice between rocks in a vertical road cutting, and the other was culling (ror deep within a labyrinth of crevices among large rocks in the road cutting, The female was collected on the surface and appeared to be approaching the latter male. Additional males were heard calling within crevices adjacent to the road but we were unable to trace them due to the Sporadic nature of calling and their subterrancan habitats. The female contains large, Unpigmented egps indicating that development occurs within the egg capsule like other Australopapuan microhylids (Zweitel & Tyler 1982), Two mature eggs measured in the ovary were 4.5 mm in diameter | CUNNINGITAM & A. 4. DENNIS The skin of this species ys thick and glandular (Fig. 4) and the animals exuded a slimy mucus wher handled, apparemly as a defensive mechanism hariatian The colour pattern is Consistent im the three available specimens, all of which exhibit the distinetiye oranpe stripes on the head. Some of the fingers and toes of each specimen aré dehydrated making accurate measurement difficult, but none has expanded dises: some fingers and loes have faint, vestigial grooves an the Ups, Measurements and propertions of tHe teu paratypes (PNG 9052/OMJ60231) are: S-V 56,3/44 |; TL 1:8/14.6, EN 3.4/3.3; SN 2.1/2.0; IN 4.6/3.9: EVE 6.0/5.2; AW 19,0/17-7: HL (6.0/6.1: FT 2.0480. HD 13.2/11.7: EN/TN 0.76/0.846; TL/SV 0.35/0.33; HW/HL 118/109) BY E/S-V 0106/0, 018; HW/S-V 0 337/040); width of toe tip on fourth toe (width of penultimate phalanx) 1.1 (0.99/08 (0.7). width of toe tip om thin finger $0 (1.00/10 (10), Comparison with ather species Callulopy (sensu Dubois (988)) now includes IS species, Morphologically they are rather conservative, und there is extensive overlap in most body proportions among species (Zweifel (972 Table 6), The comparisons belaw are based lurgely on the detailed descriptions of taxa presented by Zwetfel (1972) The presence of orange stripes dorsally on the heal distinguishes C sagiftaney Jrom knowh congeners. Callulaps boetteer{?, C. eursdactylus and C slateri further differ from C. sagitiatus in having greatly expanded linger and toe discs (vs no dises). Callulops doriae, © dubius, C fusens, Co humicola nomicoli and Ch. compins, C. kopsteini, C. persenatus and €. robustas have smal) grooved disus on the fingers and toes, Callulupy derive (00 mm), Co perxename (72,5 mm) and €. rabuses (73 mm) are much larver species and C. dubius appears to be 4 much smaller species (maximum S-V = 24 mim). Although sample size is small, there appear to be differences between the mating calls of C saginartus and those of both C. persenarys.and C. robustus, Two calls of C, personas have a dominant frequency of about 1000-4500 Hz (vs 609 Hz) and contained 5 notes (vs I-12), Calls of C. rohastus from Misirna Islnd (the type locality) have a dominant frequency of about 8O0H? (J, Menzies unpubl, data). Callulops h. compiuy is the only other Species in which adults have orange stipes on the head. but in this species the orange markings are restricted toa Short litera post-ocular stripe, Callulops sagittaine shares with C. ft. Awmicsla the condition of the mandibular branch of the trigeminal nerve. As this condition is unique umong the asterophiryines 1 appears to be a synupomorphy indicating u close phylogenetic relationship between these taxa. However, ©. h, Aumicila differs (rom C. saeinatis in the possession A NEW MICROHYLID FROG FROM NEW GUINEA tol 2000 HZ 1000 -1000 Time (ms) Fig. 5. Audiospectrogram (top) and wave form (bottom) of first four netes of an TH-note call sequence of Callulaps sagutaus sp nov recorded it the type loealny. Air temperature 13,5°C. of grooved dises on the fingers, and lack of orange stripes on the head. Three species, namely ©, glundulosus, C. stictogaster and C. wilhelmanus share with C. sagifiatus the lack of finger and toc dises, Callulapy vlandulosus. differs from Co sagittatus im having a coarsely mottled ventral surface (vs uniform) and in having a well-developed glandular area behind the car, whereas C. stictogaster is a larger species (lo 80 mm) and has a distinet tubercle between the eye und the nostril (lacking in C. sagitrarus), Callulaps wilhelmanuys closely resembles C. sagiftalus and has a similar call (J. Menzies unpubl, data), size and colour pattern. It is distinguished predominamly by the absence of orange markings on the crown and the condition of the mandibular branch of the trigeminal nerve. Zweilel (1972) discussed four specimens of Callulops trom Busilmin on the northern slopes of the Star Mountains that he tentatively assigned to C. robusius. This population is geographically close to the type locality and is ata similar altitude. The frogs are withm the size range of C. sagittalus but none exhibits the orange stripes typical of this species and their identification remains uncertain, Srarus of Callulops h. humicola and C. h. compitus Although we still lack calls or other reproductive data for these taxa, their classification as a single species is no longer tenuble, given the evidence that the axon most closely related lo C. fr, Auimicela is not Ch. comptus but C. sewitarus, In hight of this and previously reported consistent morphological differences (Burton 1986; Zweifel 1972) we propose that C. humicola compris be elevated to specific status as Callulops compius (Zweitel) new combination. Zweilel (1972) presented a thorough description of these two taxa and a detailed comparison with each other and all other Callulops except C. seagitianes, with which they are compared above. Erymology From the L. Sagitta (= arrow) with reference to the arfow-shaped orange markings on the crown. Acknowledgments Field work in Papua New Guinea by SR, MC & AD was generously supported by Ok Tedi Mining Limited. We are particularly grateful to lan Wood, Andrew Storey and Jan Roderick of the Environment Department for their support, and to Paul Weldon who provided shelter on (he summt of Mt Binnie. The Gregory family of Tabubil and James Menzies (University of Papua New Guinea) assisted us in numerous ways and we are extremely grateful for their hospitality. Roselyn Busasa (Institute of Papua New Guinea Studies) and Dr Navu Kwapena (Department of Conservation and Environment) facilitated the processing of our research visas and export permits respectively. Mare Hero kindly provided access to his “Canary” program, and Lucy Smith produced figures 2 & 3, James Menzies and Richard Zweifel provided uselul Comments On (he manuscript. 162 S. J. RICHARDS, T. C. BURTON, M. J. CUNNINGHAM & A. J. DENNIS References Bium, J. P. & Menzies, J, I. (1988) Notes on Xenobatrachus and Xenorhina (Amphibia: Microhylidae) from New Guinea with descriptions of nine new species. Alytes 7, 125-163. Bock, W. J. & SHear, C. R. (1972) A staining method for gross dissection of vertebrate muscles. Anat. Anz. 130, 222-227. Burton, T. C. (1986) A reassessment of the Papuan subfamily Asterophryinae (Anura: Microhylidae). Rec. S. Aust. Mus. 19, 405-450. Dusois, A. (1988) Miscellanea nomenclatorica batracho- logica (XVII). Alytes 7, 1-5. RICHARDS, S. J., JOHNSTON, G. R. & Burton, T. C. (1994) A remarkable new asterophryine microhylid frog from the mountains of New Guinea. Mem. Qld Mus. 37, 281-286. ZWEIFEL, R. G. (1972) Results of the Archbold expeditions. No. 97. A revision of the frogs of the subfamily Asterophryinae, family Microhylidae. Bull. Am. Mus. Nat. Hist. 148, 411-546. (1985) Australian frogs of the family Microhylidae. Ibid. 182, 265-388. —_____ & Ty Er, M. J. (1982) Amphibia of New Guinea pp. 759-781 In Gressitt, J. L. (Ed.) “Biogeography and Ecology of New Guinea” (Dr W. Junk, The Hague). MANUNEMA PECTENOPHORA SP. NOV. (PERESIANIDAE, LEPTOLAIMINA), A NEMATODE POSSESSING UNUSUAL MALE SUPPLEMENTARY ORGANS By AIMORN C. STEWART & WARWICK L. NICHOLAS* Summary Stewart, A. C. & Nicholas, W. L. (1995) Manunema pectenophora, sp. nov. (Peresianidae, Leptolaimina), a nematode possessing unusual male supplementary organs. Trans. R. Soc. S. Aust. 119(4), 163-169, 30 November, 1995. Manunema pectenophora, sp. nov., with three unique pre-anal male supplementary organs, is described. These are comb-like organs held clear of the body on short rods. Two previously described species of Manunema, the sole genus in the Peresianidae, possess tubular supplements. M. pectenophora also differs from the other species in that the single testis is anterior. All Manunema species possess four long cephalic setae, no labial setae or papillae, circular amphids, a strongly annulated cuticle, a narrow tubular buccal tube, a narrow cervical region expanding to accommodate the strongly muscular pharynx, two outstretched ovaries ventral to the gut and simple curved spicules. The taxonomic placement of the Peresianidae is difficult but the conclusion of other taxonomists that it belongs within the Leptolaimina is supported. Key Words: Taxonomy, marine nematodes, Peresianidae, Manunema. Transactions of the Reval Seeterv of S dust (995), L941, 163-169, MANUNEMA PECTENOPHORA SP. NOV. (PERESIANIDAE, LEPTOLAIMINA), A NEMATODE POSSESSING UNUSUAL MALE SUPPLEMENTARY ORGANS by AIMORN C. STEWART & WARWICK L. NICHOLAS* Summary Srmwarr, A.C, & NICHOLAS, W, L. (995) Manunema pectenophorn, sp. noy. (Peresianidae, Leptolisnina), dnemulode possessing unusual ale supplementary organs. Trans. Ro Soc. §, Alest, W9(4), 163-169, 40 Novernber, 1995. Manurnema peclenephera. sp. noy.. with three unique pre-anal male supplementary organs, is deseribed, These are comb-like organs held clear of the body on short rods, Two previously-described specios of Mamurreme, the sole genus inthe Peresianidac, possess tubular supplements. M. pectenopliara also diflers from the other species in that the single tests ts anterior, All Manunente species possess four long cephalic setae. no labial setac or papillae, circular amphids, a strongly annulated cuticle, a narrow (ubular buccal lube, a narrow cervical region expanding lo avcommoadate the strongly muscular pharynx, (Wo outstretched ovaries ventral to the wut and simple curved spicules. The taxonomic plucement af the Peresvaradae ts difficult but the conclusion of oiher taxonomists that jt belongs within the Leptolaimina is supported, Key Wokbs! Taxonomy, murine nematodes, Perekianiduc, Munanenns. Introduction Mununema pectenoplara sp. nov, possesses prominent male supplementary organs. i.e. ventral pre- anal organs found in many male nematodes, but in the new species they are unlike those described previously, The Peresianidie contains a single genus, Manunema, comprising only two previously-described species. namely M. proboscidis Gerlach, 1957, and M. catnilata (Vitiello & de Coninck 1968) Riemann, et al. 197i, The taxonomic placement of the Peresianidae has proved a problem. Some characters. suggest placing the farnily in the Leptolaimina (Chromadorida), others are closer io the Desmoscolecoidea (Monhysterida). Materials and Methods Specimens were collected fram the intertidal zone ol beaches at Darwin NT. Samples of about 2 kp of sand were dug up at low tide and the meiofauna present was briefly suspended in 5 litres of tap Water with vigorous stirring. As soon as the sund had settled, the water was passed through a 60 pm nylon sieve and the fauna retained on the sieve back-washed into a beuker with sea water. They were immediately fixed by adding formalin to give a final concentration of 5%- Later, the mejofauna Was examined in petri dishes under a binocular microscope, The new species Was isolated by pipette from the many hundreds of other nematodes collected and the nematodes mounted on microscope slides in anhydrous glycerol. Cover slips were + Division of Botany and Zoolayy, Australian National University, Canberra ACT 0200, supported by glass beads (Ballatinr) selected under the microscope to be slightly wider than the nematodes and the cover slips were ringed with Glyceel. (Gurr). Measurements are in jem from specimens fixed and mounted in this way. De Man's indexes (ratios) (Portuner 1990) are given, i.e, a= body length divided by greatest body width. b=length divided by length of pharynx, c=length divided by tail length, c'= tail length divided by width at anus, V =anterior end to vulva as a percentage of body length, and spicule measurements are arc length, Drawings and measurements were made using a cameta lucida. When mounted, the nematodes lie on their sides presenting a lateral view, and our drawings, with the exception of all four cephalic setae. show setae on one side only, those lymy uppermost as mounted, For scanning electron microscopy, some specimens in 5% formalin were washed in phosphate buffer, pH 7, containing 3% sucrose. post-fixed by the addinon of 2% osmium tetroxide, washed, sonicated and finally freeze-dried. The specimens were mounted on metal stubs and coated with gold/palladium before examination in the microscope. Type specimens are deposited in The South Australian Museum, SAMA, Adelaide. and their numbers tn the Museum's Australian Helminth Collection, AHC, are given in the text. Manuneéma pectenophora sp, nov (FIGS I-ll) Holotype: Male, Rapid Creek beach, Darwin, NT, 19.x,1992, SAMA, AHC 30000. Measurenients: Table | 164 A.C, STEWART & TABLE | Measurements of Manunema peetenophora sp, ier W. L, NICHOLAS Type Holo Male paratypes j= 4 Female paratypes n= 4 Male Range Mean +5) Range Mean +8D Length 448 466-506 490 ) 475-50%. 497 7 Maximum width ly I-12 a OS 16-20 18 183 Cephalic setae 13 10-16 \3 3.06 12s {5 2.50 Body. setae i} Ila I2 1.73 W-14 Q [.83 Mouth to amphid 12 10-11 10 O58 9-9 {) 150 Amphid diameter 47 303.2 3.1 0.12 3.6-3.6 36 0.00 Width at amphid 6.5 5.0-6.5 5.8 O76 5.6-5.6 5.0 0.05 Bucea! cavity Bie) 32-46 3a 2.08 Waa a3 150 Width at buccal cavity 12 Y-lt {0 145 942 iT] 1.50 Muuth to nerve ping 5S} 34-60 a7 3,06 53-00) a4 2.08 Width wt nerve ring i) 13-14 13 O58 Ik 16 173 Pharynx 83 77-83 80 i §2-88 85 2.58 Width at cardia I+ 1-17 (4 2 52 MW) 1k 14 249 Mouth to vulva . - - - 252-273 265 10 Width at vulva - - - - 15-20 18 [83 figy : 4479 57 20 Mouth 10 utlus 341 391-434 42 22 407-448 426 4 Tail 3 72-84 78 6 67-74 71 3.77 Width up anus i] I-11 il u 8-11 0 1,29 Spicule, are length 4 23-25 24 | Gubernaculurn ll 10-11 W 0,99 Anus jo Ist supplement” 68 4,5-7,3 64 LA - Anus to 2nd supplement* 1B 1-15 3 2:25; - Anus to ard supplement’ a4 3-34 33 162 - - - De Man's a 33 3H 39 ai] 2.63 25-30 1k 26 De Man's h 5.9 5.6-6.6 44 O47 57-62 5.8 (1.24 De Mans ¢ 6 5,9-7.0 63 0.63 69735 70 (122 De Mars ¢° 74 6.57.6 7A 55 68S 75 O96 De Mun’s V Ye - - 5153 33 148 AAs pereentage of body length Deseriprion of Holotype male Small, body when fixed strongly curved, head and cervical region folded back along body, tail curled. Cuticle strongly anmulated: lateral ridges from oid pharyngeal region to mid tail, wavy in register with annules: four rows of prominent body setae, arising from pronounced cuticular hemispherical swellings, dorso-lateral setae alternate with ventro-lateral setac. Four long cephalic setae arising form sackets; labial setae absent: amphid circular, Buecal cavity, woah minute ridges around mouth, mitially narrowly conicul extending posteriorly us a narrow parallel-sided tube Pharynx, in vervieul region (35% of pharynx length) narrow purallel-sided, encloses buccal tube, then a wider muscular cylinder, somewhat constricted hy prominent nerve ring, two cytoplasmic clefts between herve ring and expansion; cardia short, eylindrical Intestine simple tube, anus and rectum project slightly from body contour; caudal glands not observed (probably obscured by strong annulation), Single testis lo left of intestine; spicules cephalated, smoothly curved, lips pointed; gubernaculum slightly curved plate. Three pre-anal supplementary organs, most anterior one about mid-way between cardia and anus, the other twe close to anus. Each supplement resembles an outwardly and slightly forwardly directed comb, with about I prongs. mounted on a cuticular rod arising deep in the body wall. FParutypes; SAMA. AHC 300017. Measurements of three males and four females are given in Table |. In paratype males, as in the holotype, anterior supplement about 33% of body length in front of anus, second and third supplemenis, closer to anus, apparently more variable in position, probably due to different degrees of body curvature, Long testis, to left of intestine. with many developing sperm, begins just anterior to mud body, continues as long sperm duct. SEM of another nile, Figs @ and 7. shows a tenous transparent film overlapping the base of a supplement and adjacent cuticle. We interpret this as mucus. present oyer the surfave of treshly fixed specimens and preserved hy freeze-drying but lost when specimens are transferred lo glycerol for light microcopy. Females (Fig. 5) similar t males apart from reproductive organs and absence of supplementary orguns. Didelphic, two very short ovaries outstretched , ventral fo gul. Three females each have single large egy, 43, 48 and 79 ppm long, respectively, overlapping the vulva, 10 left of intestine. ‘The largest is probably at an early stage of the first cleavage division. MANUNEMA PECTENOPHORA SP. NOV, 165 3,4 25 um Figs 1-4. Manunema pectenophora sp. noy. 1. Male head. 2. Entire male. 3, Supplementary organ. 4, Spicules and gubernaculum. 166 a a x soam % * om wt yy ey < sien ieee Fig. 5. Female Maninema pectenophara sp vow Differential diugnosis. The form ob the Supplement distinguishes MM. pectenophera sp. Noy, Irom the other described species of Manunema, none of which possesses camb-like structures mounted oo rods, The new species differs From Af. annulate in the orientation of the sinule testis. Habitat Sandy ocean beach. Distribution So far known only trom Rapid Creek beach, a suburb of Darwin, Northern Territory. Epymology Named from L., pecien, a comb. Discussion Supplementiry organs are common in many farnilies of Adenophorea, where they are associated with sensilla, and are generally believed to play a part in copulation, They may be tubular, setose or papilliform und are often ussociated with cuticalar ornamentation but none like the organs described here has previously. been reported. They do pot appear ty be associated wilh sensilla and conceivably serve some mechanivul role in copulation, M. proboseieis possesses two pres unal tubular male supplementary organs (Gerlach 1957). Viticllo & de Coninck (1968) claimed. that supplements were lacking in M. annii/ata, but Riemann etal, (\97L) redeseribed M. annufara, reporting two pre-anal tubular supplements. Neither Gerlach (1957) nor Vitiello & de Coninck (968) comment on he buceal cavity, We agree with Riemann eve, (971) thar the buceal cavily 1s long and tubular, Lorenzen (98h) includes a long tubular buccal cavity us one of the diaynostic characters of the Peresianidae, The taxonomic position of the Peresianidac, ta which C. STEWART & W. L, NICHIOLAS Manurnena belongs, has been the subject of sore doubr. partly because their small size hus led lu some uncertainty abour taxonomically iniportant characters, His significwnt that scanning electron microscopy Uoes not show either Guter Libial papillae or setae, nor any external Wiandfestation of janer labial sensilla. Ail (he described species Haye four Jong sub-median cephalic setae inserted in sovkets. In Lorenzen’s (1981) phylogenetic classification of the Adenophorea, ovaries ventral te (he intestine and a Single posterior lesus are significant characters in Manuneny, consistent with the placement of ihe Peresianidae in the Leptoluimina but. while the location und Jorm of the ovaries in M. peetenaphora are the game as in. M. proboseldis, we have observed a single anterior testis. in three Males OFM. pectenpphere, The form of the umphids, the long narrow buccal lube and jubular supplementary organs are consistent with Leprolaiminas ventral outstretched ovaries ure not (Lorenzen 1981). In the possession of four cephalic setae, the absence of outer jablal setae, the possession ob four Sub-median rows of alternating body selite arising trom peduncles and the anus on a protrusion from the body cavity, Manunema resembles the Desmouscolecoidea, within the Monhysterida, rather than the Leptolaimina. In Vitteilo and de Coninck’s (1968) view. the similarities between Peresiane unnulata. now renamed Mantimema annulate Rienvann et el (1971), and Melia spinosa Gerlach 1956 jndicated a phylogeneue link between the Hal\plectidae (Leptolaimina in Lorenzen’s clissificalion) and the Desmnoscolecida, in which they placed the new species. The similarities to which they drew attenuon were the four cephalic setae and the position of the non-vesicvular amphids, but in other respeets the species are unalike, differing in the structure of the cuticle, buccal cavity, pedunculate setae and (he location of the anus. In fact. as Riemann ¢7 al. (971) point out, there ure sinmlarities between Mertiw'me and other Desmoscolecoidea, for example with Tricoma niirabiliy. Timm 1961, although Mann- vema shows greater similarity with such Leptolaimina as dnomenema haplostema Hopper 1963 and Leplolaimus tritubulatus Boucher and Helleouét 1977. Although M. pectenophora does nut possess tubular supplementary organs Ord posterior testis (leptolanind characters of Maniinema probuscidiy aad Mf. annulaia) we cancir with the placement Of Peresianidae in (he Leptolaimina, with a possible link between Leptolaimina and Desmoscolecoidea Acknowledgments We thank Dr Russell Hanley for providing facilities in The Northern Territory Museuniagd the Australian Biological Survey for financial support for one of sy to work in Darwir, MANUNEMA PECTENOPHORA SP. NOV In/ oW — _ 7 _ _ 4 8 Figs © and 7. Seunning electron microscopy of Manunemea nectenophora ap: nov. 6, Entire male. 7. Enlargement supplementary organs. SQ supplementary organ to show 168 A. C. STEWART & W. L. NICHOLAS 2 _ Figs 8-IL. 8. Scanning electron microscopy of female Manunema pectenophora sp. nov. CeS cephalic seta, AM amphid. 9. Female by light microscopy. 10, SEM of female head. Il. Male by light microscopy. SO supplementary organ. MANUNEMA PECTENOPHORA SP. NOV. 169 References FortTUNER, R. (1990) Ratios and indexes in nematode taxonomy. Nematologica 36, 205-216. GERLACH, S. A. (1957) Die Nematoden fauna des Sandstrandes an der Kuste von Mittelbrasilien. Mitt. Zool. Mus. Berlin 33, 411-459. LoreENZEN, S. (1981) Entwurf eines phylogenetischen Systems der freilebenden Nematoden. Verojff. Inst. Meeresforsch. Bremerh. Suppl. 7, \-472. RIEMANN, F., VON THUN, W. & LORENZEN, S. (1971) Uber den phylogenetischen Zusammenhang zwischen Desmoscolecidae un Leptolaimidae (freilebende Nematoden). Veroff Idst. Meeresforsch. Bremerh. 13, 147-152. VITIELLO, P. & DE CONINCK,. L. (1968) Peresiana annulata n. gen., n. sp., type interessant de Desmoscolecida. Rapp. Comm. int. Mer. Medit. 19, 201-204. ASPHONDYLIA DODONAEAE, A NEW SPECIES OF CECIDOMYITIIDAE (DIPTERA) DAMAGING LEAVES AND BRANCHES OF HOP-BUSH, DODONAEA VISCOSA (SAPINDACEAE) IN AUSTRALIA By P. KOLESIK* Summary Kolesik, P. (1995) Asphondylia dodonaeae, a new species of Cecidomyiidae (Diptera) damaging leaves and branches of hop-bush, Dodonaea viscosa (Sapindaceae) in Australia. Tran. R. Soc. S. Aust. 119(4), 171-176, 30 November, 1995. A new gall midge species Asphondylia dodonaeae, is described from South Australia. Detailed descriptions of the larva, pupa, male and female as well as the infestation symptoms on leaves and branches of hop-bush, Dodonaea viscosa Jacq. subsp. spathulata (Smith) J. G. West (Sapindaceae), are given. The new species is diagnosed and compared to other species of the genus Asphondylia. Key Words: Cecidomyiidae, Asphondylia dodonaeae sp. nov., Dodonaea viscosa, South Australia. ‘Dunsactions of the Revel Society ofS Aast (1995), 194), T7L17O. ASPHONDYLIA DODONAEAE, A NEW SPECIES OF CECIDOMYLIDAE (DIPTERA) DAMAGING LEAVES AND BRANCHES OF HOP-BUSH, DODONAEA VISCOSA (SAPINDACEAE) IN AUSTRALIA. by P. KOLESIK* Summary Korpsik, P. (1995) Asphomiviia dedonaeac, a new species of Cecidomviduy (Dipteru) darhaging leaves anid branches of hop-bush, Dedeneee viveuse (Sapindaccae) in Australia, Trang, R, Soe, S. Anse, W9(4), 17L17A. 30 Navermber 1995. A new pull midge species dsphondylia dodonaese, is-described trom South Australia, Detailed deseripions of ihe larva, pupa, nude and female as well as dhe intestation symplonis on leaves and branches ot bop-bush- Dodinaec viscosa Jacq, subsp. spatiudata (Smith) 1. G. West (Sapindaveae), are given. The new species is diagnosed and compandd te other species of the penus Ayphandylia- Kiy Worns! Cecilamyidie, Aiphandylia dediaeae sp. nov, Dadonuea visvesa. South Australia. Introduction The new gall midge species described here was found festing leaves and terminal branches of hop- bush. Dodonaea viscosa Jacq. subsp. spaihutate (Smith) 1. G. West (Sapmndaceue) in South Australia. Dodemnaed viscosa Jacg. is a shrub of tree up ta & mvtall It occurs throughout Australia. und extends intd tropical Asia, America and Africa and inte temperate southern Alricu, New Zealand and Pacific islands (Reynolds & West 1985). [ts leaves arc used in various parts of the world in folk medicine to control fever, colic, inflammation, swellings, rheumatism and pain (West 1984: Ahmad er al, 1987; Wagner et al. 1987; Mata er af, 1991), In several countries it is used as firewood. material for tool handles and for reclamation of unused or degraded landscape areas such as sand dunes, nmurshlands und mine wastes (Norem ef al. 1982; Reynolds & West 1985). In Australia a purple- leaved form is grown widely. in gardens ard the foliage is valued for its decorative appearance, The hop-bush is «common shrub in remnants of the original flora around Adelaide where it forms a substantial part of the medium-high vegetation cover if the pature conservatyon. parks. During, 1992-1993 large numbers of galls were found onalinest all shrubs surveyed in Morialla and Cleland Conservation parks. The new gull midge appears to have two generations in the Adelaide area, the first from January to February, (he sevond from September to October. Shrubs bearing galls from two -suceessive generations of the gall midge cau otlen be found, * Depirtment of Horticulture Viticulture and Oenology, Faculty bf Agricuiturt! and Natural Resource Seicnees, University of Adelaide PMB | Glen-Osmonds, Aust. 5064, Materials and Methods. Leal and branch stem. valls of Dodenaed wscuse subsp. sparhulara were sampled in Morialta (27 xi, 1992 and 26.ix.1993) und Cleland Conservation Parks (3.7.1993). The parks are adjacent and located about 13 kin north-east of Adelaide, The galls obtained on 26.ix.1993 were processed in two ways. A small number was dissected and the larvae (along with one larva from 27.xi,1992) and pupae. were preserved in 70% ethanol atter notes were made on their colour, A larger number, with larvae and pupae retained within galls. was brought to the labortory to rear to adults, Branches with galls were kept in plastic bags. Larvae pupated in their galls. Plastic bags were examined daily and emerged adults preserved together with their pupal skins in 70% ethanol after their colour had been noted, Canada balsant mounts of a series for microseopic examination were prepared according to the technique outlined by Kolesik (1995). The type series and other milenals retained in 70% ethanol together with dried examples of the galls are deposited in the South Australian Museum. Adelaide |SAMJ. Australian National Insect Collection, CSIRO, Canberra [ANIC] und United States National Museum |USNM|. Washington DC USA. Asphendylia dodunaeae sp. noy. (FIGS 119) Holetype: of, Marjalta Conservation Park, South Austnilia (34°54'S, 138°44°&), 29,1x,1993. P. Kolesik. reared From larva trom leal gall of Dadenaca vivewsa Jacq. subsp, spathulara (Smith) J, G. West, sampled 26.14.1993. 121272 [SAM]. 72 P KOL EStK Altpinnes 2. sume dita, 121273 [SAM], Paranpes: 3%. 39 9, 4+ Jarvae, 4 pupal skins |SAM|. 20 4 29 9.2 larvae, 2 pupal skins [ANEC]. ull same dau Lb larva. sunmpled 27.41.1992 [SAM]. Other marerial (OS a, WG SISAM|, Sa a,59 9, JWSNM], 10 pupal skins [SAM|. 5S pupal skins [USNM |. 10 pupae, all same dita as holotype. 3 karvae [SAM|, 5 Jarvace [LISNM], all collected with holotype Dreeneasis Wings with R, jounny C at wing apex, Rs absent, kK joining C at wing mid-length. M, ,. absent, M, weakly develuped. Cu forked, Sc cell opaque Plagellameres 12 in number, cylindrical wilh short geeks, firstand second not fiised, with short and stout setae and bearing anustomosing slightly appressed vircumfila, Male flagellomeres ull uboul same length, feinale ones, espevially the apical three. successively and progressively shorter “Tarsus: first segment substantially shorter than second, bearing ventroapical spine; claws simple; empodia longer than claws, Male rerinalia: gonocoxiles free ventrally, short. with small apiual lobe, gonostylus situated dorsclly on ganacox ite. short, bearing Wwe teeth merged basally* hypopruet and vere? bilobate: aedeagus long. stout, tapering distally Femule abdominal sternite 7 about (hree times jonger than sternite 6. Ovipositor: clongate, sclerotized. wilh large basal lobes; cerci fused. glabrous. bearing tew microsetae Male (Figs '-7) Colour: selerouzed parts of body dark brown. selae and Scdles black, non-sclerotized parts of abdomen orange, Wing length 2.4 mm (range 2.2 - 2.6), width fmm (10-12). Wing membrane and veins densely vovered with setae, 55 - 120 pm, microtrichia dense, about 0.5 pm long. Flagellomeres with sloul setae, 34- 38 jan, more or less equally positioned on the segments, Cireumfila: two long and two shart longitudinal bunds with long bands connected to each other by transverse circular bands on both ends; each of the short bands attached on batti ends to one of (he long ones by shert wansverse arch; the transverse virculur bunds on the distal end of the flaugellornere arched strongly. Eye faeets rounded, eve bridge 8-¥ facets Jong. Maxillary palpus 4 or 4 segmented. offen specimens with different number of segments in lett und right maxillary palpus can be founds however, lolul length of beth palpi about the same. Palpiger weakly developed. Legs vovered with sete and seales_ the Jatter serrated ut distal end. Female (Figs 8-12) Wing length 2.6 mim (2.6 — 2.7), width 1,2 mn (1,2 1.4). Flagellomeres with stout setae, 30 - 35 am Circumfila comprising two Lransverse bands eotinected by two Short longitudinal bunds. Chiws somewhat stronger than tr male, Abdomjnal sternite seven 3,2 times (3.1 - 3,3) longer (har sternile sia. Setae of eeret 6-8 iy sumber and fess than | yn in length, Orher ehuructers as in miule Matare larva (elas 13-15) Colour pale Orange Tot length 19 mini (17-221 Flead capsule Width 91 jan (90-92), length 29 am (26 31), length of posterolateral extensions 10 yan (9 - (0). Antenna $4 pm (13 - (5). Sterna) spatulu bilobale, tt) pm (08-116) in length, with apival enlurgenrene 6% pn (04 - 74) ie wrth and. inepsian Fh pat (Al - Aa) in depth. Arew around spatula not scleyotized. Ags Jorsul, One pair of stemal papillae qa thoracic aud first to seventh abdominal segments. One pair of ventral pupillae on collar, seeond und third thoracic and first to cizhth ubdontinal sezments. ‘I'wo pairs ef lateral papillae on tharacic segments. Pleural pupillies Test wnd third (heracté and list to enehtl) abdern|ial sepmenis with one parr Second thoreic seemenk wilh Iwo pairs. Dorsul papillie: voller, Third thoruere and last two abdominal segments with one pair. first bye thoracie and first to seventh ahdomindl segmenis with lwo pairs. The setae on lateral papillae 3-5 iy long, those on other papillae 4 - 20 um tong, Pupit (Figs 16-18) Colours antennal horns, prothoracie spiracles anil Jorsal spines. dark brown, remaining paris pale brown. Total Jength 3.0 mm (27 0 4.2) Attemnal hianns triangular, serrated, (ol pom (147 - 182) tn Jereth Cephalic papillae with seta 34 yan (36 - 44). Upper and lower frontal horns absent. Two pairs of lower facial papillae. each consisting of one seluse (5 - 15 jm) and one asetuse papilla. Two triplets af lareral fadial papillde, cach consistmy Of two Setose labour 2 pint papillae and one asetose papilla, Prothoracie born with trachea ending al is mid-length, 96 pam (44 103) long. Second to eighth abdominal segments with two pairs of dorsal papillae (length of setae 8 - 14 juni two pairs of pleural papillae (9 ~ 31 ym) and one pair of ventral papilive (12 - 1 am), Dorsal spines simple, 46 - 104 in number und 8 40 jan in teneth, woth leqwtt and number increasing from second to ninth segments, Gall (Fig. W) This species forms subglobular monothalumous walls onrerminal branch stems and leaf main veins. glabrous, 4 mm Jeng and 3 mm in diameter, green in calour One larva oveupies cuch gall. Puputiow takes place inside the pall, Circular necrotized tissue area, Hrown jn colour, Uppears.on the top of the gall before the pups Cols a circular openiow with its dutennal horns: by moving ils body up und down, The lid to this opening remains attached nthe gall by a thin strip of uncut lissue, The pupa raises two thirds of its body Quiside the gall shortly before emergence ay adult. On 24 A NEW SPECIES OF CECIDOMYIIDAE FROM DODONAEA VISCOSA 173 Figs 1-7. Male of Asphondylia dodonaeae sp. nov. |. Last three flagellomeres, 2. Sixth flagellornere. 3, Wing. 4. First tarsomere. 5. Last tarsomere with claw and empodium. 6, Head in frontal view. 7, Genitalia in dorsal view. Scale bars = 100 pm, 174 P, KOLESIK 10°— Figs 8-12. Female of Asphondylia dodonaeae sp. nov. 8. Sixth flagellomere. 9. Mouth parts in frontal view. 10. Antenna. Il. End of abdomen in lateral view. 12, End of ovipositor in ventro-lateral view. Scale bars = 100 pm. A NEW SPECIES OF CECIDOMYHDAE FROM DODONAEA VISCOSA 175 Figs 13-18, Asphondvlia dodonacae sp. nov. 13-15 larva. 16-18 pupa. 13. Sternal spatula. 14. Head capsule and collar segment in dorsal view. 15. Last two abdominal segments in dorso-lateral view. 16. Anterior part in ventral view. 17. Last two abdominal segments in dorsal view. 18. Prothoracic spiracle, Scale bars = 100 jm, Ly P BOLESIK a) Pus. WoGalhs of typi wWieedodemacae sp. now. on Davlonaert Wycosd Jacq, subsp. sperfalare (Smith) J. G. West. Seule hur > 2 em. November 1992. the vast majority of the galls were dried, Only a few of them sul contarnmed larvae. On 3 January 1993, the galls were fresh and contained inimalure larvae, On 26 September 1993, thost of the galls were occupied by pupae, with a few vecupied by larvae una few already empty. Empty galls retained pupal skins in openings. On cach ol the latter oecasiins a few dried galls from fhe previous generation were presenton the stiribs, The species seems to hive (wo generations in the area surveyed — udulis of the first generation appear possibly from January to February and those of the second generation [rom September to October. Etymology The species name is derived from the generic nanye of the host plant. Remarks The new species can be assigned to the genus Asphondylid because the female seventh ubdominal sternite is more than £5 times longer than the sixth. the male genitalia have a ventroapical gonocoxal lobe and dorsally situated gonostylus that is about us brouwd as lone, Combined with the first (arsomeres having u ventrodistal sping, the gonostylus hearing two basally merged teeth and the uvipositor having large basal lobes (Gagne 1994). Within the venus Anpfrenedydia it 1s disdnguished fram other species by lacking both upper and lower Jromtal horns im the pupa, Acknowledgments The Ministry of Environment und Planning, South Australia, kindly permitted collection within the nature conservation parks of Moriulta and Clekind. Martin C, O'Leary, State Herbarium of South Australia, Adelaide, courteously identified the host plant species, lam grateful to David B, Hirst, South Australian Museum. Natur! Serenve. Jolin DB, Gray. University of Adelaide. Depurtnient of Horticulture, Viticulture and Oenology and Raymond J Gagné, Systematic Enlomology Laboratory; USDA, Washington DC USA, lor their comments on wy early draft of the manuscript References Auman, Vo U. Parma, to & Faia. 4, (987) The saponins trom Dedended visewsd. [iuenaptia 38 36) 362 Grom. Ryd. (804) “The Gall Midges of ihe Neotropical Keaion” (Cornell University Press, [duet New York). KOLESIK. P1995) A new species al EZoemercornia Felt (Diptera: Cecidomyidae) on Raculypiis feaseiculasa vn South Australia. 2 Aw ent Soe 34, 197-152 Maia, BR, Cosiketas. J L., Orisanto, DL, Preis Mikanoa, R., Castanppa, BP, Rie, Fort & Dri-Rin, fh 991) Chemicul studies on Mexicin pints used jn tradinional medicine, XVIL. New secondiry metabolites trom Poduneee vacesa. 2 Nur Prod, $4, 912-917, Rokpm, M, A,, Day. A, BD, & Libeke, KR. Lb, (982) An evaluation of shrub and tree species used for revegetuting copper ming wastes inthe south-western Linited States, /, drt bnviran & 290404. Revxoius. 8. T) & West 4, G, W085) Sapmdavene jy 4-1n4 Ji George. A. S. (Bd) “Flora of Austrtlia’ Vol. 25, (Australian Government Printing Serviee. Canbernit), Wacwer, Ho Lupwie. C_, GRotIAHN, L. & KHAB, Mo Y (JOR7) Biologieully active saponins Han Liaderianes viveosa, Phytochen, 26, 697-701. Wesi, JG, 984d) AD revision ol Dadondqen (Sapindaceae) im Austrailia, Branovia 7, 1194, Mills CONTARINIA BURSARIAE, A NEW SPECIES OF CECIDOMYTIIDAE (DIPTERA) INFESTING FRUITS OF SWEET BURSARIA, BURSARIA SPINOSA (PITTOSPORACEAE) IN AUSTRALIA By P. KOLESIK* Summary Kolesik, P. (1995) Contarinia bursariae, a new species of Cecidomytiidae (Diptera) infesting fruits of sweet bursaria, Bursaria spinosa (Pittosporaceae) in Australia. Trans. R. Soc. S. Aust. 119(4), 177-181, 30 November, 1995. A new gall midge species, Contarinia bursariae, (Diptera: Cecidomylidae) is described and illustrated. Larvae found inside fruits of Bursaria spinosa Cav. (Pittosporaceae) prevent formation of the seeds. Detailed descriptions of the larva, pupa, male and female and the infestation symptoms are given. Key Words: Cecidomyiidae, Contarinia bursariae sp. nov., Bursaria spinosa, South Australia. Transeetions of the Royal Saeiwty uf 8. Aust, (W999), MOC) PSL CONTARINIA BURSARIAE, A NEW SPECIES OF CECIDOMYIIDAE (DIPTERA) INFESTING FRUITS OF SWEET BURSARIA, BURSARIA SPINOSA (PITTOSPORACEAE) IN AUSTRALIA. by PB KOLESIK* Summary Ro.esi, P.(1995) Comurinia bursarian, a new species of Cecidomyiidae (Dipleny) infesting fruits. of sweet bursuria, Barsaric spinosa (Pittosporaceae) in Australia Trurs, R) Soe 8. Auusr 11904), 177-181 30 November, 1995. A new gall midge species, Conmarinia hursariae, (Diptera; Cecidomyiidael 1s described and illustrated, Larvae found mnside Fruus al Rursarie spinosa Cav. (Pitosparaceae) preven! formation of (he seeds, Detajled descriptions of the furva, pupa, oiile aind female and the infestation symploms are viven ki Wows! Cecidomyiidae, Coarina barsariae sp, noy., Bursaria spiaosa, South Australi, Introduction Burseria spinosa Cay, sweet bursaria or Christmas bush, is 4 shrub usually 1-3 mt call. The genus is endeimi¢ to Australia. Bursarid spinosa can be tound in South Australia, Queensland, New South Wales, Vielona and Tasmania where it is common in woodland vepetation (Bennett 1986). Voluminans clusters of white flowers make the Shrub a useful honey plant (Cunningham ere. (981), The gall midge species described here was found to prevent seed production in A sprmosa in Morialta Conservation Park. near Adelaide. Materials and Methods Fruil capsules of Bursarta spinosu were surveyed in Morialts Conservation Park (3 km north-east of Adelaide) on 19 February 1995, Those which contained Jurvac of the new species were brought to the laboratory where the fruits were cut oped aid the extracted Jarvae processed in two ways. A small number was preserved in 70% ethanol after their colour had been noted. The remainder were transferred with entomological forceps into pots containing sterilised, wet.sand and reared to the adult stage. Pots were examined daily and emerged adulls preserved together with (heir pupal skins i 70% ethanol afler their colour had been noted, For microscopic exumination adults, laryae and pupac were mounted on. slides in Canada balsam according ta the technique ootlined by Kolesik (1995). The type series and other material retained in 70% ethanol are deposited in the South Australian Museum, Adelaide [SAM| and Australian National Insect Collection. CSIRO, Canberra [ANIC]. > Deparment ‘nt Horticulture, Viticulture: and Oenology, Facahy of Agricultural and Natural Resource Sciences, University of Adcluide PMB | Glen Osmond S$. Aust. 5064- Contarinia bursariae sp. nov. (FIGS. 1-13) Holotype: @- Morialta Conservation Park, South Australia [34°54'S, 138°44’E], 2.i01.1995, P. Kolesik, reared from larva (rom fruit of Burseria spinosa Cay. , sumpled 19,i1.1995, [21274 [SAM], Allonype; 2, same data, 121293 [SAM]. Paratypes: 2 & 29 9,1 pupal skin [SAM]. 20°", 29 %, | pupal skin [ANIC], all same data but emerged 2.111995 - 61,1995, 4 larvae [SAM], 2 larvae [ANIC]. sampled with holotype. Ovher material: 21 larvae, sampled with holotype [SAM]. Male (Figs |-6) Colour: anienna grey. head black, thorax brown, abdomen with scleronzed parts browm and non sclerotized parts yellow, legs grey with black scale strips alang segments, Wing length 1-26 nim (1.19 - 1,31), width 0.47 mm (0.44-0.51), Vein C broken at juneture with R.. M, in form of stripe of setae, Ry selerouzed on base only: Wing membrane covered with setae, 17 - 22 pm long. Abdominal tergites 2 - 6 with caudal setae only. Head with postvertical peak present. Eye facets rounded, eye bridge 8 - 10 facets long med: ially. Eight fronte-clypeal setae tn all specimens. Antenna total length 1.43 mm (1.32 - 154), Length measurements of third Hagellomere (zm): proximal niode 30 (28 - 32). proximal neck 17 (14 - 18), distal node 36 (34-38), distal neck 28 (24 - 31). Circumfilar loops reaching the mid-length of the next node, Tarsal claws curved at mid-length, about as long as-empod ium. Genitalia: gonocoxite setose and setulose, gono- stylus with strongly sclerotized claw and un array of plates below it, sparsely setose with densely setulose pouch ut base: hypoproct deeply divided mediully, with one Seta on each lobe, setulose; cerci deeply divided medially, setose and setulose, aedéagus us long as hypoproet and cerci, 178 P. KOLESIK Figs 1-6. Male of Contarinia bursariae sp. nov. 1. Head of frontal view. 2. Last flagellomere. 3. Fourth flagellomere. 4 Genitalia in dorsal view, 5. Wing. 6. Last tarsomere with claw and empodium. Scale bars = 100 pm. A NEW SPECIES OF CECIDOMYIIDAE FROM BURSARIA SPINOSA 179 Female (Figs 7-9) Wing length 1.28 mm (1.23 - 1.37), width 0.48 mm (0.43 - 0.53). Antenna total length 0.79 mm (0,69 - 0.83). Third flagellomere with node 45 wm (43 -49) and neck 6 pm (5 -8) long. Circumfila appressed, consisting of two transverse rings connected by two longitudinal bands. Other characters as in male. Mature larva (Figs 12-13) Colour yellow. Total length 2.44 mm (2.20 - 2.75), diameter 0.18 mm (0.14 - 0.21). Integument smooth, ventrally with several transverse rows of spiculae on anterior half of each segment as well as with longitudinal rows around anus. All ventral, pleural. lateral and dorsal papillae with minute setae, sternal Figs 7-9. Female of Contarinia bursariae sp. nov. 7. Last three flagellomeres. 8. End of ovipositor in dorsal view. 9. Abdomen in lateral view. Scale bars = 100 pm in 7 & 9: 10 pm in 8. 180 P. KOLESIK 13: Figs 10-13. Contarinia bursariae sp. nov. 10. Larvae inside fruit capsule of Bursaria spinosa Cav. (left loculus with fruit, right one infested). 11. Anterior part of pupa in dorsal view. 12. Sternal spatula of larva. 13. Terminal segment of larva in dorsal view. Scale bars = 5 mm in 10; 100 pm in I-13. A NEW SPECIES OF CECIDOMYIDAR FROM BURSARIA SPINOSA LSy papillae asetose. Terminal segment with one pair of stublike. asetose papillae and three pairs al setose papillae, with one of the three pairs having longer setae than the ouler two, Head capsule width ST aa (43-54), length 40 ym (37-45), Jength of posterolateral apodemes 42 aor (35-46), Sternal spatula 152. jn (139-175) in Jength, with apical enlargement 44 pm (41-47) in width and 20 ym (J9-2)) in length, Larva cain jump short distances hy arching its body and inserting, its posterior end between the spatula enlurgement und the integument and by subsequent quick teleusing of the posterior end. Pupa (Pig. Wb Head with sinall, angular, slightly sclerotized antennal hortis. Cephalie papillae with seta 223 - 250 win long. Two pairs of lower facial papillae, one of cach pair setese (Ml - 23 ym) and one usctose, Two toplels of lateral facial papillae, one of each triplet setuse (about 5 pm) and two asetose. Prothoracic spiracle with trachea ending at its dpex, 133 to 168 aim long, Second to eighth abdominal segments wath strongly selerolized, simple dorsul spines, 5 - 15 in number and 4 - 25 pm in length, Infestation syimiproms (Fig, 10) The miestation of Bursarta spinesa by Crnuarieie hursarige can easily be overlooked because there is no apparent mulformation of the fruit capsules. However, in transmitted sunlight several larvae can be recognised inside the capsule, The larvae occupy ane or both locules of the capsule, preventing the development of seeds. Up to eight larvae were obsetved within individual fruits. Despite the absence of seed in intested fruit no siymificant decrease in the total seed production per plant was observed due to the low infestation incidence in comparison lo the enormous number of fruit per plant, Etymology Derived from the generic name of the host plant. Remarks The genus Comarinia is one of the largest genera if Cecidomyiidae represented in all zdogeagraphical rewions. Larvae of all known species are phylophayous, most live gregariously in Mowers, buds and fruits which are often mallormed (9 galls. Others are found in Tnalformed leaves und stems. Almost all known species are hosi-specific., sometimes with different species living on the same plant. The genus Cunteriaia in the context Of this paper is defined is below, Larva: terminal segment with 6 sctose papillae and 2 jarge. stublike, asetose papillae. Adults: maxillary palpus with 4 segments, antenna with 12 flagellomeres: wings with R, joining C beyond wing apex: tarsal claws simple on all legs. Male: flagellomeres binadal, with a single series of circumfilar loops on each node: genitalia with stout, unlobed gongcoxile, slightly tapered gonustylus, bilabed hypoproct and simple, short, destally tapering vedeagus, emule: ovipositor very long, retractable, the cerci tiny, dorso-ventrally flattened, and closely approximated mesally, The genus Conturinia is known im Austratia from 12 species, all of them from inflorescences and seed heads of Gramimue und Cyperaceac (Harris 1979). The species described here differs morphologically from the previously-desertbed Australian species in the number of setae on lermale cerer and the relative length of male circumfilar loops: female eerci bear eight setae in C_ bursariae; those in all the other specics bear more than 14: male circumfilar loops reach the mid-length of the next node in C bursarae; those in the other species never extend beyond the base of the next node, Acknowledgments The Ministry of Environment and Planning, South Australia, kindly permitted collection in the Morialta Conservation Park, Martin €. O'Leary. State Herbarium of South Australia. Adelaide, courtcously identified the host plant species. Special thanks go to John D. Gray, Deparment of Horticulture, Viticuluire and Oenology, University of Adelaide wnd Rayrnand J, Gagné, Systematic Entomology Laboratory, USDA, Washington DC USA, for commenting on am carly draft of the manuscript. References Bensptt, & M. (1986) Pamily Pitosporaceae pp. 429-436 In Jessop, 4. Poand Toelken, H.R. (Eds) “Flora of South Australia, Part 1 (Lycopodiaceaé - Rosaceaey” (South Australian Government Printing Division, Adeluide), CUNNINGHAM, G. M_, MULHam, W, E,, MiitHorre, PL... & Leck, L. H, Y8t) “Plants of Western New South Wales” (New South Wales Government Printinye ONice, Sydney}. Hareis, & M_ (1979) Deseriptions and host ranges of the sorghum midge, Conzarinia.sorghicola (Coquillet) (Diptera: Cedidomy udae), and of eleven new species of Contarinia reared from Graminae and Cyperaceae in Auswatia, Bull, em. Rex, 69. 164-182, Kovesik, P. (1995) A new species of Luninesicarina Felt (Diptera: Cecidumyrudae) on Auvalypins farciculasa in Sauth Aostralia. “. aux ent. Soe, 34, (47 182. A REVIEW OF THE SCALE INSECT SUBTRIBE ANDASPIDINA (HEMIPTERA: COCCOIDEA: DIASPIDIDAE) AND A NEW GENUS, NOTANDASPIS, FOR TWO AUSTRALIAN SPECIES By D. J. WILLIAMS* & H. M. BROOKESt Summary Williams, D. J. & Brookes, H. M. (1995) A review of the scale insect subtribe Andaspidina (Hemiptera: Coccoidea: Diaspididae) and a new genus, Notandaspis, for two Australian species. Trans. R. Soc. S. Aust. 119(4), 183-189, 30 November, 1995. The subtribe Andaspidina is recognised as one of three subtribes of the scale insect tribe Lepidosaphini. A review of the literature is presented and diagnostic keys are given to subtribes and to genera of the subtribe Andaspidina. Notandaspis gen. nov. is described for Mytilaspis (Coccomytilus) hymenantherae Green, a species described originally from Victoria and presently included in Andaspis and for a new species Notandaspis oodnadattae sp. nov. from South Australia. The new species is unusually large for the subtribe. Key Words: Coccoidea, Diaspididae, Andaspidina, Notandaspis gen. nov., Notandaspis hymenantherae (Green), Notandaspis oodnadattae sp. nov., scale insects, Australia. Tranwactions of the Reval Seevery at & aise 99S), MP9), PRS TEE, A REVIEW OF THE SCALE INSECT SUBTRIBE ANDASPIDINA (MEMIPTERA: COCCOIDEA;: DIASPIDIDAE) AND A NEW GENUS, NOTANDASPIS, FOR TWO AUSTRALIAN SPECIES by DoS. WILLIAMS*® & H. M. BROOKES+ Summary Withiams. D.. & Broones, HM. (995) A review of the scule inseer subtribe Andaspidina (Hemiptera: Coccoulen, Diaspididie) und it new wenus, Nelondaspes, lor two Australian species. Trams, R, See. 8, Aust, M94), ISS-1K4. GU November Ys, The subtriby Antlispidina ty recognised as ne of tree subtibes of the scale mseet tribe Lepidosaphini. A review ol the literature is presented and diagnostic Keys are given to subtibes and to genera of the subtriby Andaspidind, Norandaspis ser nov is described tor Murildspis (Coccomvitusy hymenditherie Green, a species Jeseribed originally trom Victoria and presendy ineluded in andaspts and for anew species Notanelespis-oodnadattae spy gov, from Sourh Australia, Phe new species is unusually large dor the subtribe, Kiy Worns: Coccoides, Dyaspilidue, Andaspdina, Nefandasis gen nov, Nelendaspis Aymenantherae (Green), Netandaspiy ccdnadattte sp. nov,, scale insects, Ausinaliin, Introduction Although neurly 250 species of Australian armoured scale insects (family Diaspididae) Have so tar been deseribed, most of the endemic species cannot be recognised from the original deseriptions without referring Wo authentic specimens in collections. A few species have been redeseribed as parl of revisions ol veneru but there js a pressing need for a complete revision Gf all the named species. Since a catalogue of world species was published by Borchsenius (1966) ihwould be fairly easy to extract most of the pertinent literature on Australian species. However. the work involved in alse describing the new species already in collections, dnd those still to be discovered, estimated al inuny hundreds, could take many years, Numerous exotic Species have also become established in Australia, some eausing damage to cultivated crops und irees and these alse need revision. In the present work two species are described in the subtribe Anduspidina, Australian species at present ussigned (oO this group are damduapis Aynienantherae (Green). do inelser (Green). AL ninterete Brimblecontbe and Menindaspis recurvate (Froggatt). A. livmenantherae is assigned to a new genus in which a new species with an unusually large adult female is also jnelided, * Deparment ol Fntoratogy, The Natural History Museum, Cromwell Road, London SW7 SBD ¥ Department of Crop Protection, Waite Campus, University of Adelaide, Glen Osmond, S. Aust. S04. Current address: § Yeo Avenue Hivhgate, S. Aust. $063 Materials and Methods The species are deseribed from slide-mounted speei- mens of the adult female and the illustrations show the dorsal aspeet On the left and the yentral aspect on the nght. Morphological (erminology is the samme as that used in Williams & Watson (1988) where reference may also be mude to a generalised illustration of the adult female. Further specimens have been prepared on mivrascape slides for this study using the techniques discussed by Williams & Watson (1988). The term megaduct was adapted by Takagi (1992) from the term meygapore proposed originally by Balachowsky (1954). These duets. when present, numbering 2-7 on each side of the pygidial margin. are enlarged and are much larger than any others on the dorsum of the pygidium. The orifice of each megaduet ts longitudinally cliptical and surrounded by a heavily sclerotised rim. Abbrewiations of the depastlories are aus follows: ANIC, Australian National Insect Collection, CSIRO, Canberra, Australia. BMNH. The Natural History Museum, London, U.K, Historical Review of Andaspiy and related genera Ih the present work two Uibes, Diaspidini and Lepidosaphin) are recognised in the subfamily Diaspidinae. Based on the works of Borehsenius (1966) and Balkichowsky (1968) the subtribes Andaspidina, Lepidosaphina and Cocesmytilina are available in the tribe Lepidosaphini and are here accepted. Genera of the subtribe Andaspidina include Andassiy MaeGillivray, Cafe Williams, Péretedespis Mamet. Metandaspis Williams, Saotemaspiy Balachowsky und the new genus Norandaspis gen. nav. here deseribed. Ié4 4 WILLIAMS & H M. BROOKES The names Lepidosaphini and Lepidosaphing ate sed here walhout inflection formed {pom the nomjnal genus Lepidesaphes Shimer despite the varicus spellings Lepidosuphedion, Lepidusaphidin, Lepidosphedina tind Lepidosaphidinu. The genus Anedaspis was tamed ny MacGillivray (921) with Myrilaspis flava var hawaifensis Maskell 4s type species, MacGillivray also ineluded the Australian species Lepidasaphey ineisor Green, Hall (1946) accepted the venus and ineluded the African spocies Lepidusephes punicue Laing. Rua & Ferrys (1952) revised Andanpiy and meluded 10 species, eight af which were trom Asia. Brimblecombe (196) described the pew specs A. numeriia Frovy Queensland. Takagi & Kawai (1966) deseribed four new species of dadaspiv trom Japan and added further records of previausly Ueserthed species. In a detailed study of adult mules, Ghaur (1962) uccepted the subtribe Lepidesaphidina to inelude Lepidosaphes Shiner and Andaspis, Lepidosaphidina was accorded equal rank to the Diuspiding of the tribe Diaspidini, Williams (1963), in a review of Anidaspis, accepted 22 species and provided 4 key, Alsu imeluded in the review were the new genera Caia, With C quermea Williams from Pakisian as type species, und Metindaspiy with Mytilaspis recurvara broggatt deseribed from New South Wiles. us type species. He also included Meiandaypis javanensis Williams from Jaya and stated thal both new genera were related ta Andaspis. Ina catalogue of o-Galled Diaspidoidea of the world, Borehsenius (1966) recoynised (he Uribe Lepidosuphidini Shimer and the two subtribes Leposuphidina and Coccomytilina Borchsemus, He Included -dadaypiy and Caie in’ the subtribe Lepwdosaphidiag and Meferdaspls in the subtribe Coccomytilina and transferred the Australian species Mynlaspis (Corcemytiles) hymenantherae Green to Andespis. Mamet (1967) described the new genus Purunilaspia with 2 vinson’ Mamet fron Maurititié as type species Barchsenius (1967) desetibed the genera Ravaspis Borchsenius with Andispis mori Ferris as type species, Pararaaispis Borchsenius with Lepidesaphes meliae Green as lype species and Reenwalespis Borehsenius with type species Reonwalaspis guercicnla Borchsenius, The new species Ravaspis tnedica Borehsenius, &. rae) Borchsenius and Roomvulaspis quercicala dese cibed in the sane paper were purpaned io be Indian in origin but Danzig (1968) indicwed that the localities on all the original labels were in China. Takagi (1970), diseussing the Diaspididae of Tarwan, synonymised the names Ravespis, Paruraoaspis and Roonwalaypis with Andaypiy bul sugpested shat the genent may be valid in some degree ys species-zrupss, All Uhpce genera desertbed by Borchsenilis possess pygidial mepaduets Baluchowsky (196%), unaware of Mamet's Parandaspis, described the new vents Prrandusper with P castelbrance’ Balachowsky as type species, He also discussed the tribe Lepidosaphedini and erected anew subtribe Andaspidina to include Andaspiy, Cute, Metandaspis and his new genus FParandaspis, He provided a key to the three subtribes Lepidosaphedinas, Coccomynlina and Andaspidina and a key to the gepers of the sublribe Andaspidima, Raluchowsky (1973), realising (hat the nume Purundaspiy Balachowsky was a junior bomonym of Parandaspis Mamet, proposed the name Savromespiy Balachowsky to replace Paranedaspis Balachowsky with S. cuylelhrancyi as type species, Williams (1980) synonymised the name @, dase Willaims, deseribed from India. with 4. mummers Brimblecombe and commented an its distribution th Australia and the Pacifie region and its association with jhe symbtorie tungus Sepiebayidiam sp. Willitms & Watson (988) discussed the Pacific species of 4rdespis including two new species from Papua New Guinea. Takagi (992) commented on some Unusudl gener of (he Lepidosuphedini ay a tribe of the subtiinily Diaspidinae and suggested thar Merandaspiv Javanensis, based ona study ob the first mister and adult femule, was 4 Somewhat odd forny’ bul could beloms: ty the (ribe, Danzig (1993) recently accepted only the tribe Lepidosaphini without subtribes. Systemativs Superlamily Cocevidea Fullén, 184, Family Diaspididae Targion) Tozzetti. 168. Sublamily Diaspidinae Targioni Vozzeite, 168. Tribe Lepidosaphini Shimer, |68. Most peniera of the family Diaspididae or armoured scules are included jn the Wo subfitmilies Aspidiotinae and Diaspidimae. ‘The subfamily Aspidiotinae, based oo characters of the adult temiule, contains genera with peetinac or plates and lohes that ure never buobed. fe the subfamily Dinspidinae the plates are replaced by aland spines and (he lobes anterior (o the median lobes are often bilobed. The ODiaspidinge are usually subdivided inte (he tribes Dtaspidint and Lepidosaphini. Major characters of the Lepidosaphim, mostly defined by Takagi (1969) and never found ip the Diaspidini, include mepaducis. a pair of gland spines between the median lobes and abdominul segments HEY with either lateral tubercles ur spurs. One op tore of these chameters may be absent. NEW GENUS GF AUSTRALIAN ANDASPLIDINA Ih In the present work the Subtribe Andaspidina 1s recognised and cao be separated from the Iwo alter subiribes al the tribe Lepidosaphin by the following Key achipted from Balachowsky (96%) Sume genera ind species assigned ta the tribe Lepidosaphini are difficult to place in any of the subtribes. Merverspis calligent Borchsenius. lor Instunee, ducks lobes une gland spines but possesses ineguduets. The species i) nevertheless related to other species of Mereenuspis Gomez-Menor possessing glint spines and well-developed Or reduced lobes (Danzig 1993). Phaulonytihis Levnardi, an Australian genes. has small conical lobes. lacks gland spines hut PHssesses Megaducts. 1 was included in the subtribe Leptdosaphina by Borchsenius (1966). Another Aastrilian genus, a/laironwrilas Leonardi, has small Inumgular lobes bul licks megauducts. Borchseniiys (1966) jmeluded this genous in the subtribe Coceomyulina. According lo Takayi (1992), Mitulaypis MacGillivray, with more or less triangular lobes, is a primitive genus of the tribe Lepidesaphini, probably of the subtribe Coccomytilina. Howardia Berlese & Leonardi also belongs to the tribe Lepidesaphini but its position remains obscure. The yenus possesses incdian lobes similar to those of Anedaspiv. Eaeh median lobe af Howard has a narrow, tninsverse puraphysis at each basal corner and. in addition. a large club-shaped sclerosis urising froor the inner basal corner, Although Takagi (1492) tentatively included Hewerdia in the subtribe Coccomytilina, the name Howardiing Borchsenius is available for it but this subthibe was creeted originally to include other genera alsi. presently inthe tribe Diaspidini. [the follow ing key to Subtribes. amily those genera. possessing well- develuped median lobes in the adult female are included, omitting the genus Howeredia lor the present, The correct assignment of muny genera must await more delailed research possibly of first and seeond instr nymphs, Key to subtribes of the tribe Lepidosaphini with well-developed median lobes (adult females} 1. Meditin lobes with parallel or subparatlel sides, euch lohe either without notches or with a single ouler noich, Dorsul Math ital Aewucduels on the pygidiwm present arubsent,2 Meditn lobes now wirh pandlel sides. each lobe with ager WANN staBAL, diverging slightly curving round toi long oblique ouler marin, the naitgin ether smooth or serrated. Dorsal mirginal megadiets on the pypidiiay either present on ahsent Andaspiding Balaehowstey megaulucts always present on the Ton euch side, . A epidasaphina Shimer Doral, ra meaduels always absent front pygicie ra _-Cocceamytiling Birelsenius Dorsal marginal pygidtum, numbering 2 ms Genus Notandaspis gen. nov Type species: Mythaspiy (Coecommtilin) frynrenaniire rae Green Dienst Adult female on microscope slide clongate aval. scementation of thorax and prepygidial segments distinet. Spiracles with quinguelocular pores. Anrennae each usually with 3 long setac. Pygidium rounded with mediin lobes proniment, sct close together. teiamygular or oval, inner edges short and diverging, outer edges lofi, Second. third and fourtt lobes small, represented by sclerotised pornts, Megaduets absent. Muacroducts of pygidium, including muacginal duets. all ubout same size. Gland spines short between median lobes; anteriorly about saine length as thedian lobes. Venter with microduects and glind tubercles present as far forward as head. Discussion This genus is erected for the type species described fron) Vietaria ulda pew species Irom South Australia, In Jacking megaducts and possessing dorsal pygidial macroducts all about the same size. the new genus is related to Sualamaspis, a anomalous genus without gland spines in the adult female but with all the other characters of the subtribe Andaspidina, Fivonvey The name Noniiduspis is based on the Greek word Hetos, Theaning south, combined with the present eeneric name Andaspis, The new genuy Neraredaspiy can be separaled trom other genera of the subtribe by the following key: Key to genera of the sabtribe Andaspidina (adult females) |. Pyeidiumpalways with +7 dorsal marginal megaduers on each side, these much larger than other dorsal ducty Pygidiamn always without dursal marginal megsducts, any marginal ducts present always bout sane Seas other desu) ditets . 28. of + 2. Mediundobes euch with sinalis mish un Ober margin. Anal opening situated towards upex of pygidiurn ; Cufa Williaris Meuiiin lobes ‘each with Luter Marvin smooth Or finely serrated, Anal APO situated lowards base of pyvidiun i, Gland tubercles present on ventral suriace of head Purunduspis Mamet Gland tubercles absent trom ventral surface of head Andaspis MaoGillivtay ) Dorsal duets oF pygidium. tncliding any moruinal pypidirl ducts, always in the farm of microduers’ only Metandiispis Williains Dorsal dacts of pyetdiun notin the form of mictaducts diways inthe formeot macroduicts and all about sane size 5 UkO OL, WILLIAMS & 1, M. BROOKES S Gli spines absent fram pygidial margins. .—, Santonuispny Baldehowsky Gihind spines present on pyeidil murine ANeneidaapiy Williams & Brookes gem nov, Nontndaspis hvmenantherae (Greeny vamb. ney. (PIG, |) Myrilaspis (Coccomytilus) hymenaniherae Green 05: 5. Lectotype &, Victoria. Myrniong, on stems and wwies Ol Avmenantheru hanksii (BMNIL) (here designated) fexanniied |, Lepidesaphes iymenantherae (Green), Sanders 906: VW. Coeconytlas hymenantherae (Green). MacGilliy ney 2b 293, Andaspis liymenadntherde (Green), Borchsenius 66: 7\ Adult fentale Seale deseribed orivinally as teddish-brown, more ur tess covered by fibres of the bark pan which it rests! Adult female on microscope slide clongalte-oval, about }.8anin long and 11 mm wide, widest at about first abdominal seyment; body membranous to lightly selerotised, pygidium moderately sclerorised. Abdominal segitients strongly lobed literally. Lateral spursabsent, Anterior spiracles each with a group ol 47 quinguelocular pores; posterior spiracles each with 2 or 3 quinguclovular pores. Antennae cach wath 4 setuc all about Same length. Pyzidium rounded. Median lobes prominent, set close together, almost triangular, cach with rounded upex. outer edge fincly serrated and longer than inner edge: ashort, blunt paraphysis arising from umer and outer basal angles. Second, third and fourth lobes represented by short, sclerotised projections, Gland spines minute and barely perceptible belween median lobes: a short pair present between cach median und second lobe and groups of three gland spines about us long as median lobes present between cuch second and third lobe and euch third and fourth lobe. Anal opening sittared towards base of pygidium. Vulva present near middle of pygidiuin. Pertvulvar pores ubsent, Dorsal ducts of pygidiuin all about sume size, ravh about 20 jam long, arranged in loose marginal bo submedian groups onmeach segment, Other dorsal ducts on abdomen ubout same size as pyzidial ducts, preseot around margins and in submedial groups of 6-10 An sexment V, subjedial groups of 4-9 on segment TV and usually submedial groups ob b3 ducts on segment HH) Duets around margins becominay progressively srnaller to miesothoras, Ventral surtiee With oncginal gland spites as far ferward as abdontinal sewmenct IW. Gland tuberelos present on thorax und first abdominal segment Submarginal microducts present on prothurinx mesotlras and lateral lobes of abdominal seenents Small ducts situated on anurgins of thorax aud tirse abdomingl segment. Pragiosis The presenee of alinost thiangular median lobes on (he pyadriniasa good distinguishing character ot this species. Each outer edge of a median lobe ts, neverthe- less, longer than the inner edge. The lectolype designated ts one ol six specimens on a single slide labelled ‘Myrilaspis hymenanidirn Green, Type. from Mymenanitera dentatit. Victoria. Austedia, coll, J Lidgett No, 63° and is clearly marked in-red tok. Tas lurther located on a diagram showing, the posinons of all six specimens or a separate label fixed to the baek of the slide. The other five specimens are here desivmated puralectoypes (BMNE). Notandaypiy pidnadattae sy). wey, (PG. 2) Marerial examined Hololype, @, ANIC, South Australia, 70) kn west of Oodnudulta, on stems ol dcucia uneura, | s1970, FD. Morgan. Paralypes: suine daktas Holotype & SO (ANIC). So 9 (BMNH). Aduli female Seale dull white, 4 mm long, exuviae upreul, pale white, cork layer of phint in some iiatances growing in strands over scale caver, Adult female on microscope slide, elongate oval, largest available specimen 3,2 mim long, 1.2 min wade, widest at metathorax. moderately aclerotised throughout, pale brown, pygidium light brawn segments well constricted behind head and prothurax und hetween thenicic and prepygidial segnrents Anterior spiracles euch wilh uw group of tht quinquelocular pores. posterior spiracles each with | ur 2 quingueloculin pores, oevusionally absent Antennae cuch with 3 setae. one thicker ind longer than others, Pygidium rounded, Median lobes. prominent. each almost oval, the short Inner edge and long Outer edge linely serrated, A pair of slender puraphyses present, each arising from inner und outer basal angles, directed antero-medialy or almost transversely but not meeting. Second, third und fourth lobes represented by soll svlerotised points. Gland spines short and minute between median lobes, a subeqodal pair present between cuch median und second lohe, a group of three, all about us longas median lobes, situated helween each second gnd third lobe, Anal opening lying near middle of pygidium. Vulyie sittated anterior to position of anal opening, at dbout one third length al pygidium from buse. Pervulvar pores ubsent. Dorsal duets ob pyran NEW GENUS OF AUSTRALIAN ANDASPIDINA 187 Fig. 1. Notandaspis hymenantherae (Green) comb. nov, A. Adult female, general aspect. B. Pygidium. C. Dorsal margin of pygidium. D. Ventral margin of pygidium, E. Antenna. F. Anterior spiracle. 188 D. J. WILLIAMS & H. M. BROOKES Notandaspis oodnadattae sp. nov. A. Adult female, general aspect. B. Pygidium. C, Dorsal margin of pygidium, Fig. 2. D, Ventral margin of pygidium, E. Antenna. F. Anterior spiracle. NEW GENUS OF AUSTRALIAN ANDASPIDINA isu all about same size, cach approximately 20 am long, numerous ulong margins and arranged in ill-defined rows to middle of pygidjum except on segments IL-V where they form distinct submarginal raws and submedian groups of 710, Ducts around thargins becoming, progressively smaller anteriorly as far forward as mesothoray. Ventral surface wilh submarginal microducts of two types. An elongate type. each about 15 pm long, present in submurginal groups on abdominal segments FV and V. A shorter type, each about 10 aim long and with area surrounding opening selerotised; presen! in marginal groups on head, thoracic segments and second ubdonunal segment, and others present instill groups near libium and medial area of head. Gland spines present in groups on prepygidial margins and minute, Weuineare gland tubercles present subijarginally on prothorax and neur inner edges of groups of microducts. Diagnasis This 18 a lurge species compared with others in the subtribe Andaspidina with the scale cover reaching 4 tam long and the adult female 3 mm long, The scale cover of most other species searcely exceeds 2 mn long and the adult female is rarely more than | mm long. At first sight the seule of NV. oodnadatee resembles an ovisae of many species of FErlacacens (Eriocoecidae), Although each of (he median lobes is almost oval there is a distinct, short inner edge and a long outer edge as in all species of the subtribe. The shape of the median lobes distinguishes the species from N, hynenaitherac which possesses almost (ruingular median lobes, The positions of the anal opening and vulva are reversed ja both species, the anal opening of N. evdnadattae lying posterior lo the position of the vulva and mV, Aveenantherae the anal opening lying anterior w (he position of the vulva. Etymology The name ts based on the place namie ‘Oodnadauue. References Bat acniowsaky. A. S. (1954) “Les cochenilles. paleareniques de fa tribu des Diuypidini” (institut Pasteur, Paris). (68) Sur une apuvelle sous-tribu de Lepidesaplicdini (Coeemdes-Diaspididae) cereéee par ly découverte d'un nouveau vere nuisible wu Culéier @Arabie ly Sao-lome, Rew. Zivk Bor. Afr TS. 54-63. (1973) Nouveau nam de genre ee un Diaspididae de Suo-Tomeé. Ball, Soe. ent. Fy 78, 2 BorRCHSENIIS, NOS. (MG) YA catalagye vm the armoured scale inseets (Diaspidoides) of the world”. (Nauka, Moseoyy, Leningrad), (967) Materials on the fauna of seale insects (Homoptera, Coecoidea) from India, Ub Andespiy Maece with (three new allied genera (Didaspididuc) Ln, Oborr 46. 724734 BrimaiecomBe, AOR. (960) Studies of the Cocooiden (0, New species of Diaspididae, Qe. A. agree. Sei, 16, 381-407, Danaic, E, M. (1964) Ga the types af species deseribed hy NS. Borchsenius in the article “Materials on the Runa Of scule inseets (Homoptera, Covcoideu) fromy india, £1. Ent Obvcr Ah, 8434. —____ ( (993) Fauna of Russia and neighbouring countries, Rhynchota Volume X, Sele Insects (Coccinea) Farnlies Phoentoococerdie and Diaspididae. Hawa Russia ONS) 4, 452 GHavRt. M, 8. K. (1962) “The morphology and taxonomy of male seule dascets (Homupleru: Coeconlea)” (British Museum |Natural History|, Landon). Green, F, BF. (1905) Some new Victorian Covcidae Fieiorian Nat, 22, 3-8, Wank, W, J, 946) On (he Euiiopian Diaspidini. Trans, K, wat, Sow Lame, 97, 497-502, MacGiniiveay, A,B. (921) “The Coceidae. Tables Jor the identification of the subtamilies and some of the more IMporlant penera and specics together wilh Uiscusstons ot their anatomy and lite history’ (Scarab Company, Urbana, lings). Mamet, 1 R, (1967) New wener and species of Coccmden from the Masvurene Eslunds (Homoptera). Mauris Dest, Bull. 6, 89-102 Rao Vo Po& Perris, G. & (952) The genus Aneaspris MacGillivray (insectt: Hormeptera: Coccoiden), Micrventumatauy U7. \7-32. Sanpers, J G, (1906) Catalogue «af recently described Cocewae, Tech. Sen Bur Ent, U8. W218 Tawagl, S. (1970) Diaspididae of Taiwan biased on material collected (7) conaeection with the JapunelS. cosopenilive selence programune, 1965 (Homoptera: Coveoides) Part U1. Jnyeent matsum, 33, 1-146, (1992) Mitulaypis and Scluperaspis: their distributions and (axanomie positions (Homoptera; Cavcoideu Diaspididie) /bid, 47, 34-90. —__ & KaAwal, §. (1986) Some Diaspididae of Japan (Homoptera; Coccoideas, (hid, 28, 93-119, Wil tidms, D. 4, 1963) Synoptic revisions of |. Lindinyespis and [1], Anduspis with pwo new allied gener (Hemiptera, Coceowea.) Ball Br Mus. nat, Hist Eni WS. 31. (980) Aadesply dist Williams idenieal with ol. mumerad Brimblecombe (Wemiprera: Diaspididaey, a species found on toy and absecnted with the fungus Seprobasidivn. Ball, ent, Res. 70, 259-260 —— & Wasson, G. W, (1988) “The scale insects of the ~ tropical South Pacific Kegion Part L The armoured scales (C.A,B. Imernational, Wallingford, ), DIET OF JUVENILE KING GEORGE WHITING SILLAGINODES PUNCTATA (PISCES: SILLAGINIDAE) IN THE BARKER INLET — PORT RIVER ESTUARY, SOUTH AUSTRALIA By Rop M. CONNOLLY* Summary Connolly, R. M. (1995) Diet of juvenile King George whiting, Sillaginodes punctata, in the Barker Inlet — Port River estuary, South Australia. Trans. R. Soc. S. Aust. 119(4), 191-198, 30 November, 1995. The diet of juvenile King George whiting, Sillaginodes punctata (Cuvier & Valenciennes), was determined by examining the stomach contents of fish collected over two years from shallow eelgrass and unvegetated habitats in the Barker Inlet — Port River estuary, South Australia. Estimates of weight of prey actually ingested by fish were made by combining abundances and sizes of prey found in stomachs with data on the size — weight relationship of potential prey items collected separately. Fish ate epifaunal invertebrates exclusively. A range of crustaceans formed the main prey, with smaller fish taking mostly harpacticoid copepods. Amphipods were more prominent in the diet of larger fish, which also fed upon polychaete worms. Fish fed mainly during the day. Fish collected at night typically had very little food in their stomachs, as measured by a fullness index (ratio of estimated ash-free dry weight of ingested prey to dry weight of fish). Relatively few fish were caught over unvegetated habitat, but where comparisons could be made, polychaetes rather than crustaceans predominated. Key Words: Sillaginodes punctata, fish diet, predation, crustacea, seagrass, Zostera. Transaenons af the Royal Sacre af & Wish W995, TAL. [0] tum DIET OF JUVENILE KRING GEORGE WHITING SILLAGINODES PUNCTATA (PISCES: SILLAGINIDAE) IN THE BARKER INLET - PORT RIVER ESTUARY, SOUTH AUSTRALIA by Rob M. CONNOLLY* Sumanary Cossoriy, RM (1995) Diet ol juvenile King George whiting Sillveimedes panei, inthe Barker Liley - Port River estuary, South Australian, frase, RoSoe Stash W904), IY ETM JO Novernber, 1995- The dit ul pivenile King George whiting. Siwides plana (Cuvier & VYulencienies), was determined by examining the stonieh contents of fish valleeted over Uwerveun Cron) shallow eelgrass and unvegetitod habyals Wthe Barker Tiler ParORiver estiivry, South Australi. Ustiiiates ob weitt of prey actually ingested by fish were made by combing abirtlinees anil sives ol prev foul mpstoniachs with dan on the size - weight relationship ul potential prey items cullecied separately, Bish ale epifiunal irvertebrates exclusively, A range of crustucedns formed the man prey, WI sriatler Gal taking mostly harpacheoid copepods. Aniphipods were more prominent ihe det yr larger Fish. whith alvo fed upon polychaere worniss Wish ted mainly during the day, Mish collected a{ might typically had very Hitle food in their stomuchs, as mewsorcd by a lallness inde (ratio of estimated ssh- tree div weight al ingested prey ta diy weight ot fishy. Relatively few fish were cought over unvegelted habitat, Bit where cormpurisons could be rude. polyelietes nulier than erustieeins preduniimied. REY Worbs Slawinedes panei Web diet, predanon, cvustucea. scudrass, Zagter, futraduction Fish from shallow, soft-substratum habitats are lypicully carnivorous, preying mainly on small, moule invertebrates, Invertebrates associated with the seagrass vunopy Hr sediment sorties (epiaini) are more Woportint than invertebrates from wathin sediment (inhuuing) (KTanpp er ad, 1988), Despite the high levels ol primary production sustained by shallow seagrass meadows (Hillman er af. 1989). lew fish aerually COOSUING scaghitss I Temperate walters (Klan ed al. LORY), The diets of small tish trom seaziass habitats fA Soully Australia haye not heen reported. The diets of lish in Western Port, Victoria, an enclosed waterway with vegetation similar to that in the Barker Lilet Port River region, have been studied by Robertson (1984) ahd Bdgar er a/, 1993)! who confirm (he importance of epifaunal invertebrates, especially crustaceans, The diet ol juvenile King George whiling (Sillieinodes puneraia) in Western Port is deseribed i Robertson W977). Fish oof tis spectes: fed ain erustaeeans (harpacucoul copepods, mysids and amphipods) ufter setting from a planktonic larval stage into eelgrass beds. Larger juveniles (>40 mm length) fed upen ehost prawn (Calliamasse) larvae and polychaetes. * Department of Zoology, University af Adefuide. S. Aust S005, Present address: Raiculty of Enviranmental Sciences, Griftith Universiy Qld 41 Engak, GJ. HAMMOND, LS. kc WialrsoN, GF rgb) Consequences for commercial fisheries of loss of seayeriten beds if southern Australia. Report ty Fishing Industry Research & Development Committee (unpubl) primarily in vovepetated patches adjacent to eelgrass. A wide variety of measures and indices involving £ul upalyses his been used i attempts to quantify the relative Tmportince of fvod categories to Tish (Berg 9792 Hyslop 1980). Mrequeney of oceurrence, abundance. weight and volume have been used butany one of these may he misleading (Bere 1979). Indives COMPS TH Varies Ways the basic measures listed above have been devised (e.g, Pinkas ef al, 1971) but no index is advantugeous in all situaricns. Different oud Culegories gui prominence depending on the Weighting piven to the different variables in the index. Bere (1979) recommends thal where an index combining abundance, weight or volume, and perhaps frequeney of oeeurrence, is used, values for the separate varubles should also be shown. | consider that it may be as informative to forego the index, given that it is influenced by the weighting given to each variable. and simply present results based on. for example, abundances and weights, Decisions about the importanve Ob food categories lo lish are best bused not on the weight or volume of prey remaining inthe gut but on the weight or volume of prey ingested. The ideal way of calculating weight or voluine Of food intake for prey such as motile inverlebrales is lo determine the weight - size (e.g. length) relationship lor all prey categories using whole unimals. and then to estimare the weight or valume of rngested prey based on the number and size of individual items found in fish, Edgar et al. (1993)! deseribe un upproximate method for estimating weights in Which prey items are allocated to asize-class known lo pepresent the runge of sizes retained on it particular 192 kM siewe (West size Within a siiek OF |ieratehieally arninged sieves, The size of invertebrates is then used Ie esuimnute then weight (Edgar 1990), In calculating the relative importance Of food types by. dividing the number or weight of i food type by the number or weight of all food ina particular fish, no distinction is made between a fish having in its stomach one harpacticoid copepod and one calanoid vopepot and a fish having 50 harpacticoids aid 50 calaneids, Many studies therefore include some estimate of gut fullness, The most common method has heen to assizn puts to one ol several subjective vategories of fullness (Berg 1979). A measure more repeatable by other workers ts (' indice de repletion of Hureau (1969, described by Berg 1979) in which the weight of ingested food is presented as i proportion of the total weight of the fish, The primary aim of the present study was to record the dhet of juvenile 8. panetee m the Burker Inlet - Port River estuary. This estuary has been declared an uquaie reserve im recognition of its jntportance in providing, habital for juvenile fish, especially S$. punciaia Gones 1984), the most important species veonomicaully in both the conmmercial and teereational fisheries of South Austrafia. A secondary aint was to compare diets of fish fron eelgrass (Zostera muellert) and unvegeluled habitats, Materials and Methods Juvenile Silaginedes punctaii were collected over two years during surveys comparing the fish fauna of eelgrass and tnyeyetated habitats im the Barker Inlet - Port River region (1387 30° FB. 34° 48° S) (for deseriptions of the estuary and the Surveys, see Connolly 19940), At each sampling period, the stomachs and oesophagi ofall fish (or of 10 randomly selected tish where more than JO fish were caughy from cach site were removed, and the contents examingd. The number of sites and total number of lish examined ateach period are shown in Table |. The most satisfactory way of determining whut fish fal is Lo esurmine items only from the anterior part of the trace This is because food ilems from (he oesophagus and stomach are more Likely to be inter and are more easily recognised than items further along (he gut, the blas caused by differential gur passage rates or digestion rates of different food items is likely to be reduced (Berg (979) aud items towards the anterior end of (ire tract give a more reliable guide to the diet of fish just prior (o capture. This is an advantage for the secondary ain of this study. namely comparison of the diet of fish caught over eelgrass and unvegetated habitat. The truct af 8. punerete less than about 25 min jong isa simple, uncoiled tube, narrowing posteriorly: contents of these smaller fish were examined from the sechion anterior lo the marrowing. CONNOLLY Tair 4 Sillaginades punctata evaniied for stomach COMA (uiniber ubsites ac which So Punelibe were canels, number uf fish esandnedd, wed nivelion tener of fish separa for cach habitat ar cach sampling pertod. All iste were caleered citrine the eevmine exconT these dnarked Nigh Habis: Fo = eelgriss (7 = Unvevetated, Sampling period Habitat Nuuiber Nuipher ot Malian of sites fish examined length Gand January 90 E 8 tie RU ul 2 tl (a April 1990 b g 48 1 i ) | 1) August 1990) be i 45 44 t Z 2. ah) Outaher (W490 iD K 48 nS L | 7 AS February 194] b K VW 4 v ‘ ih Wn June 194] hr 4 Ad 4 Uv 4 7? a4 Tine WU) Night id 7 a4 y U | 4 ya Crther (94 k x fv 4 v 4 0 1y Oel. OL Night t J a) th u | | 7 Individual ems were mostly cither intact or nearly so, und were identified to major taxa and counted Animals were measured using a grattcule in the microscope eye-piece and were assigned to wu sieve mesh site-class so that weights of ingested prey could be estimated usimy the length - weight relationship deseribed by Edgar (990). The miajority of prey items were crustaceans and where individuals were ngt whole, sizes were estimated by roughly piccing logether parts of un animal (in (he case of large crustaceans such as amphipods and mystds) or by using other individuals of the same taxon as 4 guide (for copepods), The only tuxa recorded other than erustacea were polychaetes dnd chironomid lurve. Chironomid larvae were rare and Were always whole Although polychaetes were olten whole, they were soniclinies im pieces; estimating sizes Of polychaetes chopped jnto pieces was the most problematic part of this method. In these cases the number of anterior ends was counted and lengths were estimated to ry to lake account of the general size of individuals. Euch prey item was assigned to a size-category reluling to the range of lengthy Of thal taxon retained oo different mesh sizes. These size ranges were DIE OQ) WUVENILE KING GEORGE WHITING 103 determined. by measuring the length ol numerous speeimens of eaeh taxon from epifaunal samples taken ithe time of sh collections. Por cach taxon, relative length - frequency histogrants were plotted for each imesh size, and a range of lengths wus chosen as representative of a imesh size by seleeting upper and lower lintiis where bistogrants from adjacent mesh sizes crossed, Size ranges for each taxon are shown in Vuble 2, The meun ash-free dry weight (AP DW) of invertebrates can be related to sieve mesh size using Halgur’s (990) equation, log B= a + blog § (where B= AFDW (mg), S = Sieve Size (mmm) and a and b vary depending on broad taxonomic category}, Since each sieve size retains animals ranging fron thal sieve size lo the Heal. 5S is eXpressed asa geametne ntean calculated using the equation. log S = (log 8, + Jog $),))/2, in whieh S$) = mesh size of the 1 sieve and $,,) = mesh size of the next size up |bdgar 190), For each fish, the percentage abundance of each food category was culculuted as n/N. where nis the number of individuals of the foud category and Noa the total nomiber of id) viduiads of albeategories in that fish, The same calculation was made lor each category based on estimated weight (AF DW). The average percentage ubundunce and weight af euch food culegory were culeulated for cach site. The average percentaze TABLE 2. Sige ranges fran of prey types matching mest AIZEN, Blank wells indicate that prey type was not found on that mesh, Prey Mesh size (mi) - type 4 2 I OS (has QA25 0.075 Hary ails SUORUR PSS AOR ecu SY Mau Shh <()h Cyl SHH9 WSS4.09 ocIS9 Cala a! Wil In UML AAU t Vat cas Myst as A a Tana e3 Sled lita <2 Chr Shs 4-64 <4 Gin 245 (4a )- Sumpling Habitat Empty Empty Pullness Fullness period (nt) 1%) index fal jndes (bj dianuary W9O EF 3 5 AAR USI) 4.01 (0.50) L 0 it} ART (Rd) AKO 11 Rd) April 00 k 1) () 2.44 (()44) J i) a 204 (tal August 90 de () iY AAT UHL \ if) i 255 ith ln} Outober WT 2 3 ATV {OTH 420 Ou) \ 0 0 260 (30) LOO (36) Febroary OL 7 KN) 4.23 (124) 46 (1 2A) u 0 i 3,60 (006%) 3:00 (0.63) June 9 [ f) ) B52 (073) 252 rs) u | \4 AS (OKO) SAP Oa) Tan 19) I: a | 114 (aT) ST CY Night ul 2 50 OM6 july) OAR pay Ovtober ULE 1 5 444 (84) 472 78) ul () () 4.35 (76) 345 T6) Weber WY) TE Is (od 299 (0.4L LO (2b Night U u i] O98 tava) OUR (vad More than hall the lish caught at might indune and Oetober 1991 had empty stomachs. In fish caught at night with food in their stomachs. the types of food wert similar io those in fish caught during the cay, Fora given period, the quantity of food tn fish caught al night was significantly less than in fish caught duringy the day When fish With enipty stomachs were iachided (Mann-Whilney U-test results: June 1997, p = 0,047; Ovlober 199) pp = 0.014), but was not significantly different when fish with empty stomachs were excluded (June 1991, pp = 0.186; October 1991, p = 0,221). Comparisons of the dict ot fish caught over celgruss and unyegeliled hibital are limited by the small nurmber OFS. punrraia caught over invegetated habitat and the small gumber of unvevetated sites al whieh lish were caupht Over all periods, polychaetes seemed tiv predominate iy fish frotn unvegetated habitat, Ih October 1990 and 1991. when fish were cuueht at 4 unyegeluled sites. Increasing the vhance that the data are representative ol the hubitat more yenerally. ooly lish from uwevetated sites had taken polychaetes. Fish from celerass sites tended (6 contain a preater Guage Of crustaceans, Cuprellid amphipods, for example, were recorded only from fish caught over eelgrass at both periods, Discussion The diet ot Siflaginodes punerata fits within the typical diet for fish ftom shallow, sott-substratuiy habitats, Stomach contents at the periods sampled give ny indication of feeding on anything ofher than motile invertebrates, Juvenile Sv panera caught at periods inthe first half of the your were large enough to be able to take snl) individuals of other fish species but there was no evidence of (his. Although gastropods are culen by some fish species. none was tound in the present study. The prominence of harpucticoids and aniphipods in the diet of smaller juveniles and aa increased prominence af polyehuctes. in older juveniles matches the pattern Wd. punctate frou Western Port, Vicloria (Robertson 1977), Bruce (1995) has suggested that (he shift in diet towards Jarger crustuceans and polychuetes with in¢rousing fish size may be rcluted to the iming of gut coiling. Bruce’s study of larval and postlarval & puncrata trom South Australian waters shows {har coiling ofthe gut tube and tigration of the anus begin in fish 2b = 24 mm long, and are complete in fish of 26 inm. Most fish caught in October 1990 and 1991 during the present study were > 26 mm long and did have called puts. These fish bad a predominance ot smull crustaceans such as harpacticoids, however. midicating that the shift towards larger crustaceans and polychaetes does not happen uni] after gut coiling. DIET UP JUVENILE KING GEORGE WILITING (97 Results sugvest that $ pierare feed on a narrower ringe of prey umd include more polychaetes in iherr Wich when over unvegetuted fabri, Lubbers er al. W490) have also reperted that for juveniles of several species Of [ish from an estuaty in Chesapeake Bay, USA, diets of fish collected frum Unvegetated areas ticluded a much ereater proportion of polychaetes than diets oF fish collected from vegerted ures, Evidence trom the present siudy is, however, obtained [rom only i small qumber of fish from very few sites, The sinall number of fish examined from unyegetaled sites could account Jor the failure to find food types such as tardean slirimps recorded infrequently tm fish trom CCIANINS Sites, Ividence from the ive night sampling. periods. Sumeests that juvenile §, punctate feed manly durin the diay. The stormnwehs of fish collected at night were ofien vither empty or eentiuned only a snrall quantivy ol loud. Either fish fecd in a limited way at night or food tn the stomach of fish collected af night remarned from feeding during daylight hours. The time between sundowt und collection oF fish at night ranged from fouria seven hours. The rare at which food is evacuated by juvenile. Guritvorous. murine lish ofa similar size 0 the fist Siddivd bere haus heen shown to range yatously from 2.7 lo 4.8 h (Rosenthal & Paflenhoter 1972), 6 bh (Archumbault & Feller 1991) and fram 10 jo 40 h fRyer & Boehlect 1983), These laboratory estimates Of wit evacuation times, however. lend to be overestimates |Loekwood 80). Food is presumably clew of the stomach belore it is fully evacuated from (he gut, so stomach empiying Himes could be shorier than these mentioned aboye. On the other Hand. wut paussape tiles wre much slower in colder water (Durbin ard W83; Ryer & Boehler 1983) and, in the evening watt Ieiiperatures of June and October 991 of about 14°C, food fay have remamed in guts much longer li is therefore impossible to distinguish berween the possibiliies of limited nocturnal feeding and food remulining in stomachs from daytine feeding. The ratio of ingested food to lotal fish weight did not seen Ly vary consistently with the size of fish taken al different periods. This contrasts wath the study of silver hake (Merlncciuy bilinearis) aod Atlantic ead (Gadus nora) by Durhin et af, (983). using the sae measure, in which it was found that the ratio was greater in larger fish, Durbin epa/ (983), however, used a much larger size range, includiny juvenile and adult fish. Diflerences in (ae ratio for 8. paren might occur in larger fish, Any differences in gut passage rates or rates of digestion tor different food types-could have affected the apparent relative importinee of lond types, These biases were not determined during the present study but should hive been liniited by examining, food onby from the besophigos and stomach of fish. Dilterential digestion rales tend to underestimate the importince of solt-bodied invertebrates (Scholz era/ 1991) and, lor juvenile S. prnicrette. this meus hut polychaetes are the liaxon most likely to be anderestimited. This study confirms that juvenile S$ punerare within the Barker Inter - Port River estuacy feed on epifaunal invertebrates. Experiments in the same estuary have shown that removal of eelgrass canopy reduces epiluunal invertebrate productivity (Connolly 1995)- Abundances of juvenile A) punctate are not reduced directly by removal of eelgrass canopy but are correlaled with levels of jovertebrile productey ity (Connolly (994b). The ongoing threat to the health of eelgrass in the estuary from Huntin activities such os treated sewage und stormwater discharge should therelore he viewed as a polentilly detrimental influence on S. pureuiea populations Acknowledgments 1 thank Dr Alan Butler for fas belp and encouragement, Barry Bruce for ideas about dietary shift. and Drs Gree Jenkins and Tony Fowler tor helptul comments on the manuscript. This work rests In part upon financial dssistanee towards research Costs provided by the Royal Sociely of South Australia Tne. and the Mark Mitchell Foundation, The work was done while | was supported by an Australian Postgraduate Research Award. 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