VOL. 113, PARTS 1 & 2 31 MAY, 1989 Contents. Transactions of the Royal Society of South Australia Incorporated Lee, D. C. & Birchby, C. M. Decoribatula gen. nov. from Singapore, with notes on allied Reticuloppia (Acarida: Cryptostigmata: Oribatulidae) from Tropical Australia - - - - - - - - - - - Hirst, D. A new genus of huntsman spider (Heteropodidae: Araneae) from south eastern Australia - - - - - - - - - - - Tyler, M. J. A new species of Lechriodus (Anura: Leptodactylidae) from the Tertiary of Queensland, with a redefinition of the ilial characteristics of the genus - - - - - - - - - - - - Boulton, A. J. Over-summering refuges of aquatic macroinvertebrates in two intermittent streams in central Victoria - - - - - - - - Southcott, R. V. The larva and nymph instars of Odontacarus (Leogonius) adelaideae (Womersley) (Acarina: Trombiculidae: Leeuwenhoekiinae) - - - Tiver, F., Sparrow, A. D. & Lange, R. T. The composition and distribution of the vegetation of north-west Eyre Peninsula - - - - - - - - Bourman, R. P. & Lindsay, J. M. Timing, extent and character of Late Cainzoic faulting on the eastern margin of the Mt Lofty Ranges, South Australia - - Ainslie, R. C, Johnston, D. A. & Offer, E. W. Intertidal communities of northern Spencer Gulf, South Australia - - - - - - - - - Koste, W. & Shiel, R. J. Rotifera from Australian inland waters. III. Euchlanidae, Mytilinidae and Trichotriidae (Rotifera: Monogononta) - - - Brief Communications: Davies, M. Rediscovery of Uperoleia orientalis (Parker) (Anura: Myobatrachinae) - - Nobbs, J. The occurrence of plant parasitic nematodes in the arid region of South Australia - - - - - - - - - - - PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 113, PART 1 a 2 zz a ss —- = = Se 4 —— S im =—— 8 ' TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INC. CONTENTS, VOL. 113, 1989 PARTS | & 2, 31 May Lee, D. C. & Birehby, C. M. Decoribatula gen. noy. from Singapore, with notes on allied Reticuloppia (Acarida: Cryptostigmata: Oribatulidae) from Tropical Australia - - + > = 6 = + > . " Hirst, D. A new genus of huntsman spider (Heteropodidae: Araneae) from south eastern Australia - - - - - - - ~ - - - Tyler, M. J. A new species of Lechriodus (Anura: Leptodactylidae) from the Tertiary of Queensland, with a redefinition of the ilial characteristics of the genus - - - - - - - - : - - Boulton, A. J. Over-summering refuges of aquatic macroinvertebrates in two intermittent streams in central Victoria - ~ - - - - - - Southcott, R. ¥. The larva and nymph instars of Odontacarus (Leogonius) adelaideae (Womersley) (Acarina: Trombiculidae: Leeuwenhoekiinae) - * Tiver, F., Sparrow, A. D. & Lange, R. T. The composition and distribution of the vegetation of north-west Eyre Peninsula - - - - - - - Bourman, R. P. & Lindsay, J. M. Timing, extent and character of Late Cainzoic faulting on the eastern margin of the Mt Lofty Ranges, South Australia - Ainslie, R. C., Johnston, D. A. & Offler, E. W. Intertidal communities of northern Spencer Gulf, South Australia 2 3 = e ; 3 e 2 Koste, W. & Shiel, R. J. Rotifera from Australian inJand waters, III, Euchlanidae, Mytilinidae and Trichotriidae (Rotifera: Monogononta) - - Brief Communications: Davies, M. Rediscovery of Upero/eia orientalis (Parker) (Anura: Myobatrachinae) + Nobbs, J. The occurrence of plant parasitic nematodes in the arid region of South Australia - - - - - - - - - - - 115 117 PARTS 3 & 4, 30 November Koste, W. & Shiel, R. J. Rotifera from Australian inland waters IV. Colurellidae roar: Monogononta) - - - - - - - Short, A. D., Buckley, R. C., & Fotheringham, D, G, Preliminary investigations of beach ridge progradation on Eyre Peninsula and Kangaroo Island - - Pledge, N.S. The occurrence of a diprotodontid marsupial referable to Euowenia grata De Vis in western New South Wales — - - - - - Austin, A. D, & Allen, G, R. Parasitoids of Uraba lugens Walker (Lepidoptera:Noctuidae) in South Australia, with description of two new species of Braconidae - - - - - - - - - - Barker, S. Contributions to the taxonomy of Australian Buprestidae (Coleoptera): New species of Astraeus and Stigmodera (Castiarina) and a key to Astraeus (s.s) - - - - - - - - - - > Jenkins, J. F. & Hasenohr, P. Trilobites and their trails in a Black Shale: Early Cambrian of The Fleurieu Peninsula, South Australia - - - - - Gowlett-Holmes, K. L. & Holmes, N. J. CG. Rediscovery of Primovula (P) heleneae Cate and description of a new species of Crenavolva from South Australia (Mollusca: Gastropoda: Ovulidae) - - - - - - Davies, M. Developmental biology of the Australopapuan hylid frog Litoria eucnemis (Anura: Hylidae) - - - - - - - - - Brief Communication: Beardsell, G. R. Hybridisation of Litoria chloris and L. xanthomera. (Anura: Hylidae) ; - = e = 3 = = = = Insert to Transactions of the Reval Society af Sauth Avstralia, Voi. 113, parts 2 .& 4, 30 November, 1989 19 145 163 169 185 195 215 221] DECORIBATULA GEN. NOV. FROM SINGAPORE, WITH NOTES ON ALLIED RETZCULOPPIA (ACARIDA: CRYPTOSTIGMATA: ORIBATULIDAE) FROM TROPICAL AUSTRALIA. BY DAVID C. LEE & CAROLYN M. BIRCHBY* Summary Decoribatula pustulata gen. nov., sp. nov. on an orchid from Singapore, intercepted at Adelaide Airport, is described. It is similar to Reticuloppia reticulata Balogh & Mahunka, 1966 from rainforest litter in Queensland, the description of which is extended. Both mites are unusual in having a deficient chaetotaxy on femora I and 11. KEY WORDS: Acarida, Oribatulidae, Decoribatula pustulata, new genus, new species, Reticuloppia reticulata Balogh & Mahunka, Singapore, Queensland, leg chaetotaxy. DECORIBATULA GEN. NOV. FROM SINGAPORE, WITH NOTES ON ALLIED RETICULOPPIA (ACARIDA: CRYPTOSTIGMATA: ORIBATULIDAE) FROM TROPICAL AUSTRALIA. by DAVID C. LEE & CAROLYN M. BircHay’ Summary Lee, D, C, & BircuBy,C.M, (1989) Deeoribunela gen- noy. from Singapore, with notes on allied Reticaloppia (Acarida: Cryptostigmata: Oribaiulidae) from tropical Australia, Trans. R. Soc. S. Aust, 13 (2), 1-5, 31 May, 1989. Decaribatila pustulata gen. nov., 5p. nov. on an orchid from Singapore, intercepted at Adelaide Airport, is described. It ix similar to Reticuloppia reticulata Balogh & Mahunka, 1966 from rainforest fitter in Queensland, the description of which {8 extended, Both mites are unusual in haying a deficient chaclotaxy on femora { and II. Key Worns Acarida, Ovibatulidae, Decaribaiula pustufate, new genus, new species, Reticuloppia reticulata Balogh & Mahunka, Singapore, Queensland, leg chactotaxy, Introduction The mites considered here have been studied because of their relevance to an ongoing study of sarcoptiform mites of South Australian soils, sampled from nine florally diverse sites. An introduction to relevant work on the advanced oribate mites (Planofissurae), with comments on the Oripodoidea, which include the Oribatulidae, is provided by Lee (1987), These mites and a group of seven species of Oribatula-like mites from the South Australian study aré atypical within the Oripodaidea in that they havea deficient chaetotaxy on femora I and []. Oripodoid leg chaetotaxy will be discussed more fully when the South Australian species are described (Lee & Birchby in preparation). The chaetotaxy for femur I and II on oripodoid mites is 0 anterior; 2 dorsal/2 ventral, I posterior setae (0,2/2,1), In the deficient chaetolaxy one or two setae are missing, resulting in one of three chsctotaxies as follows: (0,2/2,0; 0,2/1,0; 0,2/1,1. The Oribatula-like 5, Aust, mites differ superficially in having only short or medium length hysteronotal setae and they were collected from only the arid, semiarid or mallee sites; whilst the two species considered here have long hysteronotal setae and are from moist, tropical sites. The notation and methods of measurements follow Lee (1981) with modifications made by Lee {1987}. Measurements are in micrometers (jum). The type of the new specics is deposited in the South Australian Museum, Adelaide (SAMA) whilst rhe type of Reticufoppia reticulata is deposited in to. the Hungarian National Museum, Budapest, * South Australian Museum, North ‘Terrace, Adelaide. S. Aust. S000 etlewloppia Balogh & Mahunka Reticuloppio Balogh & Mahunka, 1966; p. 564 Type-species. Reticuloppia reticulata Balogh & Mahunka, 1966. Diagnosis; Oribatulinae. Hysteronoium with 14 pairs (SJ, 62, 3S) of long setae. Lamella absent. Hysteronotal lenticulus present near dorsosejugal furrow. Integument of hysteronotum with extensive reticulate sculpturing and cerotegument forming a thick, columnar refractile exudate. Femora I and L) with deficient chactotaxy, lacking posterior setae: I — 0,2/2,0; 1] — 0,271.0. Legs slim and long, with leg IV longest (Jeg If] ts lost). Pretarsal claws short (céntral claw {1 less than 0,33 x length of tarsus 11) and lateral claws much slimmer than central. cliew (depth less than 0.5 x depth of central claw I), Remarks: Reticuloppia is based on a single female, grouped in the Oribatulinae, and distinguished by the hysteronotum having a large number of Jong setae and teticulate sculpturing around a lenticulus (Balogh & Balogh 1984). In addition, the proteronotum lacks ridges, having only a weak sublamejla, and the leg chactotaxy is deficient in a way that is unique amongst esiablished oripodoid species. Reticuloppia reticulata Balogh & Mahunka FIG. t Reticulappia reviculata Balogh & Matunka, 1966; p. 564, Female: \digsomal length, 475 (470 in original description). Leg lengths (femur-tarsus); I — 270, M -— 254, 1f —?, (V—311. Tibial maximum heights: 1 = 25,5, 11 — 20.5, Hi — 7, LY — 18. Yellow brown colour. Thick (Maxitnum depth equal to distance between setae 2I-/2), whitish, columnar refractile i] DC, LEE & C.M. BIRCHBY Figs 1-2: Posterior aspects of femur-pretarsus of right legs I and 11, all setae on femora illustrated, p = posterior, = ventral. t, Retieuloppia reticulata Balogh & Muhunka. 2, Decoribatula pustulata sp. nov. cerotepument on proteronotum anterior to /2, thickest lateral to seta zl; also covering hysteronotum except around lenticulus and near pleural margins, with (hick patches posterolateral to seta Sl, lateral to $5 and between setae J6—J6. Proteronotum with faint cosrate sublamella. Sensory seta (32) with long stalk (length = x 3 caput), Margin of bothridium raised into turret-like structure, height subequal to length of <2 caput. Seta s2 short (0.5 x or less distance between setae j2-zi1), fine, smooth, without cilia. Hysteronotum with fine reticulate sculpturing under columnar exudate. Five multiporose foramina, foramen F°3 divided in F3a and F3b, Hysteronotal seta Sl confirmed similar to and nearly as long as 21. Podosternal chactotaxy: 34, 177, 3777, 37 with all third rank setae as long as /7/3 (illustrated in the wrong position by Balogh & Mahunka 1966, fig. 15) and 73 on pedotectum I, 7//3 posterior to pedotectum II (as for Decoribatula pustulosa sp, nov., Fig.4), /¥3 on discidium, Seta Sal shorter than Sa2 and Sa3. Opisthosternum with foveolate sculpturing, Four subcylindrigal eggs, exochorion . smooth, 80 x 230 (mean). Legs long (mcan femur- tarsus: 59% of soma) and slim (mean maximum tibial height: 27% of mean length). Material examined: Holotype female, rainforest litter, Mt Spec (18°S7’S, 146°1 VE), Townsville, Qld, DECORIBATULA, ORIBATLILID MITES a 19,17,1965, J. Balogh. Some appendages lost, no femur-tarsus Hl on either side, Remarks: The further description here of Reticuloppia reticulata adds to or corrects the original description, with a few important conlirmations. Reference to the relative dimensions of the legs is distorted by legs 11] being lost (usually # short, slits leg amongsr the oripadoids). Decoributula gen, nov. Tipe-species: Decoribatula pusrulata sp. nov. Diggnosixs: Oribatulinae. Hysteronotum with 14 pairs (5J, 6Z, 3S) of mainly long setae (51 is short). Lamella with only costate antenor part present, not reaching bothridium to seta 72, Smooth, paler mid- dotsal anterior area on hysteronotum, bot not clearly delineated as lenticulus, Integument of hysteronotum with extensive foveolate sculpturing, cerolegument inconspicuous, Femora | and U1 with deficient chactotaxy, but with posterior setac: | - 0,2/1,1; U—0,2/1,1, Legs of medium girth and long, with Icg TV longest. Pretarsal claws long (central claw {1 more than 0.33 « length of tarsus El) and tateral claws only slightly slimmer than central claw (depth mare (han 0.5 x depth of central claw 11), Remarks: Decoribatula is based on # single male, It is similar to Reticu/oppia in having # delicient femoral chaetotaxy, long hysreronotal setae, a divided anterior hysteronotal foramen (#3u, 43h) and a turret-hke bothridium to 22. It particularly differs, in having a lamella, a different type of ceTotegument, a different type of setal loss on femora | and I] and unusually large pretarsal claws. On the basis of this it is considered to be a sister- group to Reviculoppia bur different enough to be regarded a6 a Separate genus. Decortbatula pustulata sp. nov. FIGS 2-4 Male; Idiosomal length, 527, Leg lengths (femur- tarsus): 1-298, 11-280, LIL—285, [V —323. Tibial Maximum heights: | — 346, 1 — 31, IE — 23, IY - 23, Red brown colour, Incomspicucus (depth less than diameter of hysteronoral setal bases} cerplegument, coucentrated in round of uval pusiwes on hysteranotum, Proteronotum with partial costare lamelta, superficially inconspicuous bul backed by relractile infernal apodeme, not reaching back to turret-like bothridium (0 seta s2, Two multiporose foramina (#1, F2a) recognisable dorsally, Central setae (1 J2, 21) with 3 or 4 files of cilia, /2 longest. Sensory seta (22) with caput and exposed stalk subequal in length, Seta s2 short (0,6 % or less distance j2-21), fine, smooth, without cilia. Hysteronorum with smooth central triangulate area {rom anterior margin to level with seta J3, but no clearly delineated pale lenticulus, Tive multiporose foramina (F3a, £30, F4, FS, F6) each backed by a disc-Nhaped chamber i) integument. Hysteronotal setae smooth, S| short, subequal tu s2, whilst J5 and §5 very long with whip-like end. Podosternum with reticulate sculpturing. Subpodal ridge in three parts, pedotectum II and discidium large, extending laterally beyond level ol pedotectum |. Lateral setae (especially 2273) longer than central setae. Opisthosternum with three setuc (JZe4, /Z25, Se) represented only by bases on both sides and assumed broken off. Scia Sal Junyer than Sa? and Sa3. Shield dark, centrally horizontal and punctate with curving upwards, smooth marginal strip. Legs long (mean femur-tarsus: 56% of somah, with leg f second longest, and of mediuni-girth (mean maximum tibial height: 35% of mean length), Ventral flange on trochantera (11 and LV. Femora with ventral incrustation. Material examined: Holotype male (SAMA N1988474), on orchid from Singapore, intercepted at Adelaide International Airport, 13.iv. 1987, Grex Raker (S, Aust, Department of Agriculture), Remarks: The male of D. pustutata is described, although oripodoid species are usually based on females, because it is unlikely that more material will be collected and [his species is importan! to the classification af S. Aust, mites being studied (Lee & Birchby in preparation). There is a lack of sexual dimorphism amongst oripodoid character states that are considered important in distinguishing sptoes. The female is likely to be bigger and have the thickening around the genilal orifice separated from the ventrosejugal apodeme, Its relationships are considered under the remarks on Decoribatnda. Acknowledgments We thank the Australian Biological Resources Study for a grant to DCL. funding the salary of C.M.B., Ms Kathy Bowshall foi the notation and presentation of the figures and Mrs Debbie Lowery for typing the manuscript. 4 DC. LEE & C.M. BIRCHBY 100um Fig. 3: Decoribatula pustulaia sp. nov., notum of soma. DECORIBATULA, ORIBATULID MITES 5 Fig. 4: Decoribatula pustulata sp. nov, sternum of soma. 100um References BALOGH, J. & BALOGH, P. (1984) Review of the Oribatuloidea Thor, 1929 (Acari: Oribatei), Acta zool. hung. 30, 257-313. & MAHUNKA, S. (1966) New Oribatids (Acari) from Australian Soils. Folia ent. hung. 33, 553-568. Lee, D. C. (1981) Sarcoptiformes (Acari) of South Australian soils. I. Notation. 2. Bifemorata and Ptyctima (Cryptostigmata). Rec. 5S. Aust, Mus. 18, 199-222. _—__. (1987) Introductory study of advanced oribate mites (Acarida: Cryptostigmata: Planofissurae) and a redescription of the only valid species of Constrictobates (Oripodoidea). Ibid. 21, 35-42. A NEW GENUS OF HUNTSMAN SPIDER (HETEROPODIDAE:ARANEAE) FROM SOUTH EASTERN AUSTRALIA BY DAVID HIRST* Summary A new heteropodine genus, Keilira, gen. nov. and two new species, K. sparsornaculata sp. nov. and K. sokoli sp. nov. are described from South Australia and Victoria respectively. Keilira differs from other Australian genera in labium shape and the relatively short, thick embolus of the male. The arrangement of cheliceral teeth is similar to that of Heteropoda Latreille, 1804, but the general appearance resembles Australian species attributed to Olios Walckenaer, 1837, here considered as belonging in Neosparassus Hogg, 1903. A NEW GENUS OF LIIUNTSMAN SPIDER (HETEROPODIDAE:ARANEAE) FROM SOUTH EASTERN AUSTRALIA by Davin Hirst* Summary Hrest, D, (1989) 4 new genus of huntsman spider (Hereropedidae Araneae) trom south eastern Australia. Trans. R, Soo S Aust, (13, 7-23, FL May, 1989. A new heteropodine genus, Aeiliru, gen. nov. and twa new species, KL sparsarmmacnidta sp. nov. ad K, sokuli sp. nov. are duseribed from South Australia and Victoria respectively. Nef/ing differs from other Australian genera in labium shape and the relatively short, thick embolus of the male. The arrangement of cheliceral teerh is sumilar ro that of Heferopodg Latrenle, 1804, bit the general appearance resembles Australian species attributed to Olios Walvkenaer, 1837, here considered as belonging in Neosparassus Hage, 1903, KEY WorRDS, Arancae, Heteropodidae, Keslird gen. nov,, new taxa. Introduction This paper forms the first part of a laxonomic study on the Australian Heteropodidae A new heteropodine genus is defined for undescribed species previously mis-identified as Neosparassus (sublamily Deleninae Hogg, 1903), which they resemble in general colouring and shape, Neosparassus will be discussed fully later (Hirst in prep.), but it muy be mentioned here that despite its revalidation by Jarvi (1914) all relevant species hive been referred to Olfos in subsequent literature. Subfamilies of the Heteropodidae are poorly defined. Some useful characters of the Heteropodinae Keyserling, 1880 are referred to here in associating the new genus with that subfamily. Similarly, characters of the Deleninac are mentioned in comparing Neasparassus with Olios. Allhough limits of that subfamily are uncertain, it is retained here as separate from the Busparassinae (Jarvi 1912) on the basis of genitalic $uructure. Materlals and Methods All measurements are in millimetres, made with an eyepiece praricule. Eve diameters, interspaces and MOQ nieasurements are expressed as relative to the diameter of an AME. Drawings were done with the aid of a camera lucida on a Wild microscope. All abbreviations are standard for rhe Araneae. Bilateral variability of leg spine counts are indicated in parenthesis, Types are deposited in the South Australian Museum, Adelaide (SAMA), and the Australian Museum, Sydney (AM). Keilira pen- nov. Diagnosis: Labium twice as broad as long. Male embolus short, thick relatively strateht with a curved * South Australian Museum, North Terrace, Adelaide, S.Aust, 5000, tip. Female with small fossa and hood-like anterior marein. Definition: Carapace raised, flattish or convex in profile, highest in the fovea region, sloping wently to clypeus, Foveal groove weak or indistinct. Eyes subequal. Anterior eye row slightly recurved. Pusterior eye row straight to slightly recurved. Clypeus half width of an AME or slightly less. Cheliceral fang groove with three teeth on promargin, three or four leeth on retromargin and one of more median rows of minute, pointed teeth. Labium ai least. twice as broad as long, Sternum barely longer than broad or equal, slightly convex. Legs 2143. Anterior legs with four ventral pairs of spines on tibia, without an extreme distal pair. Reduced prolateral {cg spination, patellae withour prolateral spines. Female palp tarsal claw with four small teeth, Female genitalia with paired uncoiled spermathecae. Small fossa with weakly sclerotized hood-like anterior margin. Male embolus short, thick, relatively straight with a curved (ip. Palpal ubia with a small prodorsal distal extension and a triangular spur rising from a retrolateral apophysis, Type species: Keilira sparsomaculata sp. nov. Camments Although resembling Weosperassis in body shape and coloration, the presence of a distal prodorsal extension an the tibia of the male palp.and three teeth on the promargin of the chelicerae followed by one of more rows of median tecth, places the genus close lo Meteropoda and Pundercetes L, Koch, 1875, the Australian representatives of the Heteropodinac. Neosparassus and other genera af the Deleminae lack median teeth and a well defined extension on the male palp tbia. The retromarginal tecth of Aeslira differ trom those of Hereropoda and Panderceres in that both the latter have four long teeth which are evenly sized and close together with a small space between the two middle teeth. In Keilira greater spacing occurs DB. HIRST A NEW GENUS OF HUNTSMAN SPIDER 9 between the middle and distal teeth (ig. 8), This placement remains unchanged when a fourth tooth is present, this being minute and found al the proximal end of the row (Fig, 12). The teeth of the female palp tarsal claws of Fleferupoda and Puridercetes are elongated and curved, as Jong of Jonger than the curved claw tip, and comb-like. This condition is approached in at least one other Australian heteropodid, Oltus Aernitis Hoge, 1914, the correct generic status of which is unknown 4s no mature specimens have been scen. The clongated teeth on |he palpal claw are nut found in Xe/ire, but as all other Austratian genera narmally possess at least five teeth, it is unusual in having four. Keilira and Pandercetes have four ventral spine pairs on anterior ibiae but lack a distal pair, While four ventral spine pairs are found in Hererapoda, one pair is shorter and positioned distally near the articulation with the metatarsi, This character was used by Kaston (1980) to distinguish Heferoyada from Olies, the latter having two pairs and lacking an extreme distal pair. Neosparassus and most other Deleninac genera have three pairs with rhe distal pair near the articulation, The exceptions are some species of Pediana Simon, 1880 and juvenile specimens of at least Isapeda L. Koch, 1875 and Neasparassus (juveniles of other genera have not been examined). There, the distal spine 1s represented as 2 bristle barely distinguishable from these adjacent on the tibia margin, Keilira is similar to Heferopeda in carapace profile und in having leg 1 shorter than leg LU (Panderceres has a conesve carapace profile and leg T longer than lee tI), but differs in eye position and relative size of the anterian median eyes: The male bulb, in the upexpanded slate, resembles thar of Panderceres and, to a lesser extent, Heternpoda, Keilira lacks the long tapering embolus typical of other Australian heteropadid genera, having a relatively straight, short, thick embolus which tapers at the tip to a blunt point, For much of its length the embolus runs adjacent to the bulb In Pandercetes and Heteranada the embolus is well clear of the bulb for its entire length. It can be concluded that Keilira is referable to the Heteropodinge, yet is distiner from Hereropode and Pandercetes, However it is Known that Heteropodinge genera of uncertain status occur in Queensland with live or six ventral spine pairs on anterior tibia. These are not congeneric with Xeilire (unpubl, dala) but appear to be closely related and may prove to be w sister group. Keifira contains the smallest known Australian heteropodids. Its representation by a handful of specimens from the south-eastern parts of both South Australia and Victoria extends the known range of the Australian Heteropodinae, Little is known of their biology but they appear to have similar habits to Neosparassus (Main 1976), Several specimens ot Keilira sparsomaculata were collected whilst foraging at night on low flowering shrubs (Leptaspermum junipermum Sm.), which were attracting abundant jnsects, a habitat three species of Neasparassus have also been found sharing. One juvenile specimen of Keilire was foutid in litter at the base of shrubs during the day, Etymology: The generic name Ketilra is derbved from Keilira Station, Jt km N.E, of Kingston, SE, South Australia. Keilira sparsomacufata sp, Wo, FIGS 1-10 Holalype; SAMA NIYSTS, o. 13 km N, OF Keilira Station, 36° 37°S_ 140° 11° E, §. Aust, Feb. 1974, D. Hirst, Allowpe SAMA NIQR76, 9, same data as holotype, Paratypes: SAMA NI9877, 9, same dala as helotype: SAMA NI9S7B, 9, Cortina Station, Coorong, 369 20'S, 1399 46° FE. S. Aust. 201.1968, AW. Forbes. Other miaterial examined: SAMA NI9872, Juvenile, Cortina Station, same data as above; SAMA N1L9873, Juvenile, Kendall Station, Coorong, S. Aust., Mar. 1968, A.W. Forbes; SAMA N19874, Juvenile, AWinga Scrub, S, Aust., 16, ix. 1987, D. Hirst, Description of holotype (Figs 1-4) Carapace length 2.90, width 2.85, Abdomen length 3.40, width 2,20. Colour in alcohol: Yellow-brown carapace with pair of median black, shart linear markings with adjacent spots, anterior of fovea. Caput with darker suffusion in ocular region, eyes with black rims. Chelicerae reddish-yellow, Sternum yellowish, Legs; anterior pairs yellow-crange, posterior pairs yellow, palps yellow. Abdomen creamish-whire with small scattered spots of black pigment, fewer verctrally Carapace: Convex, highest between !! coxae sloping gradually tw. clypeus, steeply to posterior. Fovea indistin¢t, marked by darker pigment. Setae sparse, short (ca 0.2), moderately long (ca 0,4) bristles around ocular region, few on caput between eyes and fovea and along lower margin of clypeus. Eyes: In two rows, from above anterior row slightly recurved, posterior row straight, AME Rigs. 1-4, eilira sparsomeculata zen. ct 3p. nov. Holotype (hale) | bady alld (ght pal, dorsal; 2-4 Lelt palp. 2 ventral; 3 prolateral; 4 retrolateral. Scale line; = O.Smm, a) D, HIRST Figs, 5-10. Keilira sparsomaculata gen. et sp. nov. Allotype female, 58: 5 body, dorsal; 6 right leg [, prolateral; 7 tarsal claw; 8 chelicerae, maxillae and labium. Paratype female (SAMA N19878), 9-10: 9 epigyne; LO vulva, ventral, Scale lines: Figs 5 -— 6, 8 = 0.5mm; figs 7, 9-10 = 0.1mm, A NEW GENUS OF HUNTSMAN SPIDER i} diameter 0.20, AME - ALE: PME ; PLE = 1: 1: 0.9 = 0.0. Interspacess AME - AME 0.8 ; AME = ALE 0.5 ; PME - PME 1.55; PME — PLE 1.55 ; AME — PME |.1; ALE- PLE 1. MOQ: anterior width ; posterior width ; length = 2,75 53.4: 2.63, Chelicerae; Length 1,20, width 1.69, geniculate, almost glabrous, setae long (ca 0.6), shorter on retromargins. Three rows of tecth on fang groove; three teeth on promargin, middle toorh largest, three teeth on retromargin, middle and distal teeth about vqual, median row of about cleven minute pointed teeth, placed posterior ty middle tooth of promargin row. Labium: Length 0.30, widrh 1.70, emarginate posterio-laterally, six shart setae in recurved row across. anterior half with oné near anterior margin midway: towards each lateral edge. Longer setae on anterior margin, few setue on posterior half. Maxillac: Lengih 0.75, width 0.63, slightly Vonverging, setae long. Sternum: Lenzth 1.25, width 1,22, slightly convex, setae long. Legs: (Table 1), Scopula sparse. Splnes: Leg f, fe d2 p2 r3 (2 on right fe), ti rl v8, mt v4; Leg El, fe G2 p3 rl, ui v8, me v4; Lee IO, fe d2 p3, ti v4, mt p2 v4; Leg IV, fe d2 pl, ii pl v3. mr p2 v4; Palp, fe d2 pl all distal, cl p2. Claws: Teeth small with about 6 on praclaw, 4—5 on reiroclaw. Abdomen: Setac of shore to moderale length, Spinnercts: ALS basal segment 0.30, apical segment 0.08, PLS 0,20, 0,08. Palps: (Fies 2-4) Tibia retrolaterally with apophysis carrying a short triangular blunt-tipped spur. Embolus thick, short, straight for the most part along prolateral side of eymbium, then curved inwards with a short blunt (ip. Description of allatype (Figs 3-10): As in holotype but for the following: Carapace length 3.51, width 3.50, Abdomen length 5.40, width 3.80, Colour in alcohol: Abdomen creamish with larger black spots, Eyes: AME diameter 0.24. AME: ALE: PME: PLE = |: 1.04: 0,92: ().92 Interspaces: AME — AME 0,67; AME — ALE 0,5; PME — PME 1.3; PME - PLE 1.5; AME - PME 1; ALE - PLE 0,92. MOO: Anterior widti: posterior wideh: length = 28: 3,2 :.2,6, Chelicerae: Length 1.60, width 2.10, Lubium: Length 0.36, width 0.80. Maxillae: Leneth 0.94, width 0.71. Steroum: Length 1,80, width 1.80, Legs: (Table L). Spines: Lee |, fe (3 on right), ri r2; Leg IL, fe 2 on right); Leg TV, fe p2, mt pd el, Spinneretss ALS basal segment 0.36, apical segment 0.08, PLS 0.30, 0.10. Genitalia: Epigyvnum a low mound, highest posteriorly, small fossa with weakly sclerotiyed hood-like anterior margin (Fig. 9). Vulva paired, copwatory openings inside hood, spermatheca leading to a curved receptaculum shorily joined to a smaller receptaculum (Fig. 10), Etymology; The specific name (efers to the widely scattered spots an. the abdomen, Keilira sokali sp. nov. FIGS I-13 Holotypes AM KSI9141, o, Holey Plains State Park, 38° 13'S, 46° 51'E, Vict,, 29. ill. 1980, A. Sokol. Description of Holotype: Similar ta &. spursoruculata except as follows: Carapace length 3,44, width 3.22. Abdomen length 3.90, width 2,20, Colour in alcohol: With blackish markings laterally on carapace, Abdomen with numerous spots. Carapace: Raised byt flattish above, highest in foveul region, fovea distinct. Eyes; Both rows slightly recurved, AME diameter 0.20. AME: ALE: PME: PLE = 1-0.9; 0.8 - 0,9, Interspaces: AME —- AME 1; AME-ALE G9 > PME - PME 1.7; PME- PLE I.h! AME-PME 0.9; ALE — PLE 8. MOQ: Anterior width - posterior width ; length = 3.0 <> 3.4: 2,3. Chellcerae: Length 1.44, width 1,72, Relramargin with four teeth, irregular median rows ol & — 13 minute teeth arranged in triangular area with apex pointing distally, Labium: Length 0.36, width (.76. Short setae on posterior half, Sternom: Length 1,76, width 1.70. lees; (Table 2). Spines: Leg ] fe dl p2 3, ri 2 v8, tnt pl r2 v4; Leg IL, fe dl} p2 12, el vB, mi p2 r2 v4; Leg IL, fe d2 p2, ti p2 r2 v4, mt p2 12 v4; Leg IV, fe d) pl ti v4, nit pl 13 ¢2-0n right) v4; Palp, fe d2 pl, all disral, ti dl p2 rh, Palp: Tegulum positioned more basally in eymbium than in K. sparsomaculata and enlarged Tetrolaterally at proximal end, Embolus extending further io prolateral side, distal half straighter. Tibial spur longer and more upright on larger apophysis. Enyniotogy: The species is named after the collector, A, Sokol. Remarks. Known only from the holotype, K. sekali is separated from &, sparsomaculaia by the flatter carapace, four retrolateral teeth on the fang margin, several irregular rows of median teeth and abdomen with more numerous spots. Its habits are not known. Acknowledgments The complenon of this paper was supported by an Australian Biological Resources Sjudy grant, Special thanks go to Dr D, €, Lec for his helpful comments on manusctipt preparduon, Alsu io Dr M. Gray and C, Horseman (AM) for the loan of material and assistance. . D. HIRST Figs 11-15. Keilira sokoli gen. et sp. nov. Holotype maie: 1L Abdomen, dorsal; 12 cheliceral tecth, left retrolateral; 13-15 left palp; 13 ventral; 14 prolateral; 15 retrolateral. Scale lines: Figs 11, 13 = 15 = 0.5mm; Fig. 12 = 0.1mm, A NEW GENUS OF HUNTSMAN SPIDER 13 TaBLe |, Leg measurements of Keilira sparsomaculata gen. et sp. nov. values for holotype male with allotype female in parentheses. Leg Femur Patella Tibia Metatarsus Tarsus Total I 2.85 (3.32) 1.55 (1.84) 2.58 (2.82) 2.50 (2.92) 0.90 (1,12) 10.38 (12.02) II 3.21 (3.68) 1,69 (2.00) 2.92 (3.16) 2.92 (3.16) 0.98 (1.24) 11.72 (13.24) Ill 2.32 (2.56) 1,12 (1.38) 1.70 (1.96) 1.69 (1.81) 0.80 (0.95) 7.63 ( 8.86) IV 2,45 (2.84) 1.15 (1.36) 2.00 (2.24) 2.1 (2.40) 0.88 (1.00) 8.63 ( 9.84) Palp 1.00 (1.00) 0,52. (0.74) 0,49 (0,80) - - 1.50 (1.52) 3.51 ( 4.06) re TABLE 2. Leg measurements of Keilira sokoli gen. et sp. nov. of holotype male. Leg Femur Patella Tibia Metatarsus Tarsus Total { 3.48 1.83 3,30 3.20 1,04 12.85 II 4,20 1.95 3.84 3.58 1.14 14.71 Hl 2.88 1.38 2.18 2.04 0.84 9.32 IV 3.34 1.30 2.52 2.64 0.92 10,72 Palp 1.10 0.66 0.67 - 1.50 3.93 References Hoae, H. R, (1903) On the Australasian spiders of the Kocn, L. & KEYSERLING, E. (1871-1883) Die Arachniden subfamily Sparassinae. Proc. Zool. Soc. Lond. 1902 (2), Australiens. Vols 1 & 2. (Bauer & Raspe, Nurnberg). 414-466. . r X LATREILLE, P. A. (1804) Tableau methodique des Insects. Fadl bee Pe od CO oe NO Mslands. Proc. Nouv. Dict. Hist. Nat. 24, 129-200. Jarvi, T. H. (1912-1914) Das Vaginalsystem der Main, B. Y. (1976) “Spiders”. (Collins, Sydney). Sparassiden. Ann. Acad. Sci. Fenn. (A) 4, 1-235. at . J Kaston, B. J. (1980) “How to know the spiders”. 3rd Edin Pe ee. ) beh phe aa re Sperone (W. C. Brown & Co., Dubuque, lowa). : 4 is ? heel ° KEYSERLING, E. (1880) Die Spinnen aus Amerika. WALCKENAER, C, A, (1837) “Histoire Naturelle des Laterigradae, Nurnberg, 1880, 1, 1-283. Insects”, Apteres. I. (Paris), A NEW SPECIES OF LECHRZODUS (ANURA: LEPTODACTYLIDAE) FROM THE TERTIARY OF QUEENSLAND, WITH A REDEFINITION OF THE ILIAL CHARACTERISTICS OF THE GENUS BY MICHAEL J. TYLER Summary A large collection of frog ilia from the Tertiary site of Riversleigh Station in northwestern Queensland, includes 190 specimens of Lechriodus intergerivus sp. noy. The new species is described, compared with the four extant congeners, and shown to be the smallest member of the genus. A revised definition of the ilium of Lechriodus is provided, and the significance of the find as a link between the geographically isolated extant species is discussed. The rainforest habitat preferences of all extant species suggest a similar environment at Riversleigh Station in the Late Oligocene to Mid-Miocene. KEY WORDS: Lechriodus, ilia, Leptodactylidae, Tertiary, Queensland. A NEW SPECIES OF LECHRIODUS (ANURA; LEPTODACTYLIDAE) FROM THE TERTIARY OF QUEENSLAND, WITH A REDEFINITION OF THE ILIAL CHARACTERISTICS OF THE GENUS by MICHAEL J. TYLER* Summary Tyrer, M, J. (1989) A new species of Lechriodus (Anura: Leptodactylidae) from the Tertiary of Queensland, with a redefinition of the ilial characteristics of the genus. Truris, R. Soc, 8, Aust, 113, (5-21, 3) May, 1989. A large collection of frog ilia from the Tertiary site of Riversleigh Station in northwestern Queensland, includes 190 specimens of Lechriodus intergerivus sp. nov, The new species is described, compared with the four extant congeners, and shown to be the smallest member of (he genus, A revised definition of the ili of Lechriodus is provided, and the significance of the find asa link between the geographically ixolated extant species is discussed. The rainforest habitat preferences of all extant species suggest.a similar environment at Riversleigh Station in the Late Olizacene to Mid-Miocene: Kry Worps: Lechriodus, ilia, Leptodactylidae, [ertiary, Queensland, Introduction Lechriodus Boulenger comprises a group of vrownd-dwelling leptodactylid [rogs confined to the Australian Geographical Region. In addition to the morphological aticibures thal support its recognition, it is umigue in the nature of its distribution: of the five leptodactylid genera that have been reported to occur both m Australia and New Guinea, itis the only one which is represented by more species in New Guinea than in Australia (Zweifel 1972), McDonald & Miller (1982) clarified the geographic distribution of Léechriodus in Australia, demonstrated that a previous record from north (Queensland was in error, and confirmed that there is a major disjunction between the Australian and New Guinea populations. Thus, whereas other shared genera occupy the Cape York Peninsula of north Queensland (Tyler 1972), Lechriodus does not, and in Australia it is confined to the eastern seaboard along the Great Dividing Range between Qurimbah, N.SW, and Brisbane, Qld, All that can be interpreted about guch allopatry is that at some time in the past Lechriodus must have occurred In the intermediate area. Here | report a new species of Lechriodus from the Tertiary, The marerial comes from Riversleigh Station in northwest Queensland, which js intermediate between the distribution of the extant species, The finding is significant for several reasons: it constitutes the link in the distribution of the extant species; it is the first record of fossil Lechriodus, and it ts the first Occasion on which an Australian frog fossil has been reported from a site that js beyond the modern geographic range of its genus, * Pepartinent of Zoology, Universiry of Adelaide, G.P.O. Flax 498, Adelaide, S. Aust. SOQ, Material and Methods The material is deposited in museums and other collections abbreviated in the text as follows: Australian Museum, Sydney — AM; Department of Zoology, University of Adetaide —- AU?Z; Queensland Museum, Brisbane =~ QM; South Australian Museum, Adelaide - SAM, and American Museum of Natural History, New York — AMNH. Letters following the abbreviations are departmental identifications. Comparative studies were based on the osteological collections of thé Department of Zoology, University of Adelaide, supplemented by skeletal material of New Guinea taxa borrowed from the Amerivan Museum of Natural History. Osteological nomenclature follows Tyler (1974). The following measurements were taken with dial callipers: ilial length - measured from the superior extremity of the dorsal acetabular expansion to the distal end of the shaft; dorsal acelabular expansion to ventral acetabulat expansion (DAE-VAB) measured as the distance between their extremities, and acetabular fossa diameter measured at the proximal, external edge of (he peripheral acetabular rim. Fig, 1, Onemation of lium to horizontal plane durine preparation of descriptions. All measurements and deseriptive features are derived from the lateral surface, Expressions of relative extent of features are obtained with the \lium positioned in such a way that the proxwnal edge of the bone is maintained at 45° to the horizontal (Fig, lL). Concepts of relative size of 16 M. J. TYLER component features to one another are perceived with regard to their proportional size in other anuran taxa. Scanning electron micrographs were taken on an ETEC Autoscan SEM at 20 Kv. Systematics Family: Leptodactylidae Werner, 1896. Sub-family: Limnodynastinae Lynch, 1971 Genus: Lechriodus Boulenger, 1882 The definition of the generic characteristics of the ilium by Tyler (1976) was based solely on L. melanopyga (Doria). Now that representatives of all extant species and the fossil species have become available, the definition requires modification and expansion: Ilial shaft slightly to moderately curved, bearing large, fanlike dorsal crest extending along at least three-quarters its length. Maximum depth of dorsal crest approximately one-third from proximal end. Crest concave to varying extents proximally, flattened distally. Acetabular fossa diameter varies with size of individual from which it is derived: proportionally larger in larger species. Acetabular fossa bordered by rim in large species; rim confined to inferior half in smaller species. Location of dorsal margin of acetabulum ranges from base of ilial shaft to midway up shaft. Pre-acetabular zone narrow, meeting inferior border of ilial shaft in gentle curvature, commonly forming quadrant. Ventral acetabular expansion moderately developed, commonly truncate inferiorly. Dorsal prominence small, extending superiorly above superior margin of ilium, or not. Dorsal prominence moderately to well developed, oval and horizontal or vertical. Dorsal acetabular expansion acutely angled, its superior margin on a level with or above the maximum extent of the dorsal crest. Lechriodus intergerivus sp. nov. FIG. 2 Holotype: QM F16614. An almost entire left ilium collected at C.S. Site, Riversleigh Station, Queensland. Description of holotype: lial shaft slightly curved and bears enlarged, fanlike dorsal crest whose maximum depth is at position approximately one- third from proximal end of shaft. Proximal one- half of crest concave on lateral surface, progressively becoming less pronounced distally. Distal half of crest flat. Acetabular fossa small, deep and with prominent rim bounding inferior half. Dorsal margin of acetabular fossa situated slightly superior to inferior margin of ilial shaft. Fig. 2. S.E.M. of Lechriodus intergerivus (SAM P29771). A left ilium, but image reversed to aid comparison with Fig. 3. ANEW SPECIES OF LECHRIODUS (ANURA: LEPTODACTYLIBAR) " Pre-acetabular zone evenly rounded with narrow separation from acetabular fossa, Ventral acetabular expansion poorly developed and incomplete inferiorly, Dorsal acetabular expansion narrow bul well developed superiorly, extending to position approximately on a level With maximum extent of superior margin of (lia) crest Dorsal prominence poorly developed and scarcely deteclable: in profile visible as slight superior evlension upon dorsi! marein of illum, Dorsal protuberance ovoid, lateral, well developed and projecting: laterally, Anterior margin of dorsal prominence extends to position slightly anterior to antenor margin of acetabuluny, Length of ilium 13.6 mm; DAB-VAE 3.7 mm: acetabular fossa maximum diameter 1.4 mm, Paraivpes: Henk’s Hollow Site: SAM P29742: Two ‘Trees Sites AM F76951-32, AMNH 25351-53: Last Minute Site: SAM P29764-65; Gag Site AM F76957, AMNH 25355-58, SAM P27968; Upper Site; AM F76953-44, 76958, AMNH 25354, OM F14640-50_ 17036-39, SAM P29734, 29743~44, 29757-62; CS, Site : AM F76955-56, T6959. AMNH 25359-60, QM FL6615-18, 16674, 1703|-32, SAM P29746-50; Wayne's Wok Site; AM F 76960, OM Fl6636-39, 1703435, 17040-47, SAM P29756, 29766-67; Outasite Sites SAM P27929, 2Y751-55; R,S.O. Site: QM F16619-35, 17033, SAM P29735-41, 29745, 29771 (subject of SE,M.), Variation: The complete ilia range in length (rom 5.3 mm to 14.4 mm, A paratype is shown in Fig. 2. Nereis minimal difference in overall shape, and variation ts largely eenlined to the posttion of the dorsal prominence in relation to (he acetabulum. Thus some specimens agree with the holotype in having the anterior (nargin of the prominence on a level With the anterior margin of the acetabulum, i Others Lhe anterior half of the prominence pay project beyond rhe acetabulum. The extent and degree of the curvature of che dorsal crest varies in terms of its elevation, proximal limit and medial protrusion, Referred specimens: Small portions of an additional 79 specimens gre sufficiently complete co permit \denttlication, but $0 fragmentary that they do nor eontribute 10 an understanding of the nature op variability of (he species. For those reasons they are considered here “referred specimens” rather than Paratypes. «All of the referred specimens are lodged in the Queensland Museum; Henk’s Hollaw Site: Fl6652; Two Trees Sits Fl4688-89; Las) Minute Sites FL6673, 16675, 1647879, 16685; Gag Site; FL665|, F16676-77, 16680-83, LAGBA-R7, 1670-01; Upper Sites F16658-89, 16665-72, 170S0—53> 0,8, Sites F16660-64, 16684, 16690, 17048: R.S.O, Site: FL6653-57, 16691-99, 17049; Wayne's Wok Sire: Fl6702-14, 17054~683. Coniparison with other species: ia of all extant species have been examined; L. flescheri (Boulenger) (AUZ 8 uncat,): L. melunopyen (Doria) (AMNE 81223, AUZ 2 uneals; L. agenaposiv Zweitel (AMNH 74646), £, placveeps Parker (AMNH 74178). Representatives are shown in Fig, 3. The comparisons assume that {he sample size of & intergerivus is sufficielit ' reflect with reasonahle confidence the size of tha attvined by the species, With that assumption il is relevant te nore that the absolute sizes of the iia examined of each of the adults of New Guinea species is substantially lurger than any of the £, inrereerivus, The relationship between jlial length aod snour to vent length of extant species is almast Ifnear, Thus given ilial length alone it is possible to extrapolate snout co vent length, The lonvest iliues of L. intergerivus is 14.4 mm, The corresponding snout 16 vent length derived from the elosely lineal relationship ol congeners is approximately 37.5 wim (Fig. 4). This, the apparent maximum is fess Urea two-thirds of the size of the smallest adult £. aganoposis and L. platyceps, and is slightly below the range of L, meloriopigaand L. fletchert (Table 1). Clearly £. intergerivus is the smallest member af (he genus. As indicated by the genene definition, (he itinm an this genus is conservative and inferspeeitic variation is not pronouliced. For (hat reason barn reluctant to ascribe differences in ilial vharucters greater significanve than Ley inay merit. However, given the data on size, the integrity of £. ilergerivus is nol in question. Sraligraphy and lithology: The assumed statigraphic sequence of the sites at Riversleigh is that followed in the list of paratypes, OF the sipes named to dale those containing L, interger(yus form an almost unintercupted sequence, Ole of the fossil bearing sediments is described by Fland (1985) as fine-grained arenaveous freshwater limestones, and possibly a facies within the Carl Creck Limestone. The ages of the sediments containing £. dnterperivus are currently understood tu he between Late Oligocene and Middle Miovene (M. Areler pers. conim,), Elymefogy: Latin for “placed between”, $6 alluch ny to the geographic position of the fossil population belween those of the extant species, Phylogeny: Zweitel’s (1972) Proposed phyligenctic relationships is reproduced here ag Fig, 5. Giventhe ecographic lovation af L, /ntergerivus {Fig 6), the age of the Riversleigh Station deposits, aid the fact ihat the lower estimarlon of age is CONLeM POTN eoUs wih the emergence and subsequent uplift of ret M. J. TYLER Fig. 3. Ilia of extant species with or without ischium and pubis: A. Lechriodus fletcheri (AUZ uncat.); B. L. melanopyga (AMNH 81223); c. L. platyceps (AMNH 74178); D. L. aganoposis (AMNH 74646), A NEW SPECIES OF LECHRIODUS (ANURA: LEPTODACTYLIDAE) 19 tal) 7 / E E = a e c o | = 40 2S ae 3 Y = 4.48 + 2.98x 24 5 \ r 0.96 o c { | 30+ | | | | | | | | 20 { i 1 I | | fe) am | T 1 a —- — 5 10 15 20 25 30 tum Length (mm) Fie. 4, Length of ilia of Lechriadus species plotted against snout to. vent length, Assumed snout tu vent length of fossi! species indicated by broken lines, t-value for slope 10.891, p < .00L. For x = 4, y = 37.51 (95% confidenve limits = 33,75-44,25). Square = L. platyceps, circles = L. melanopyga; triangles =L. Fletcher, FLETCHER! MELANOPYGA ®LATYGEPS AGANOPOSIS fig. 5. Phylogenctic relationships of extant species of Lechriodus as postulated by Zweifel (1972), of New Guinea, L. intergerivus is a candidate as the ancestral stock from which the extant species evolyed_ With the existence of L. intergerivus, Zweifel’s. phenetic scheme would require two invasions into New Guinea: one deriving L. melanopyga from an intermediate ancestor that also gave rise to L. fletcheri, and the other leading to L. platyeeps and L. aganoposis. A slightly simpler scenario would derive L, fletcheri directly from L. intergerivus, with asecond stock giving rise to the three New Guinea species, Further systematic studies are required to test the validity of that option, Discussion The fossil frog fauna of Australia includes 22 species and nine genera from 10 sites of Quaternary ot Tertiary age (Tyler 1989). Riversleigh Station appears likely to be the richest fossil frog site yet discovered, in terms of the number and diversity of species. This trend is: common to other vertebrate classes, and Archer, Hand & Godthelp (1988) state thal more than 200 new species have been recovered there. The frog fauna will increase the total. Amongst the frog material from Riversleigh Station now available, L. intergerivus is the predominant species. OF 379 ilia now known from the site 190 (50%) represent that species. Extant Lechriodus aré predominantly inhabitants of temperate and sub-tropical rainforest and, in the absence of any data to the contrary, it can be inferred that the habitat of L. intergerivus would have been raintorest. A second point relevant to interpreting the mid-Miocene environment is the fact that a high proportion of the ilia of other frog species found in Riversleigh Station deposits are from small creatures. Because frogs lose water readily through the skin, the body mass/surface area ratio is such that small frogs are particularly vulnerable to dehydration. In consequence they are predominant only in areas which are moist throughout the year. In communities of frogs in Australia high frequency of small frogs occurs only in areas of high and seasonally reliable rainfall, such as the extreme southwest of Western Australia and the northern periphery of Arnhem Land in the Northern Territory (Tyler 1989), Acknowledgments This investigation was made possible as a result of a Suggestion from, and the subsequent encouragement-of, Dr Michael Archer. I am further indebted to him for provision of laboratory space, and yarious help from his colleagues Mr Henk Godthelp and Dr Suzanne Hand at the University of New South Wales. The research program was funded by the Australian Research Grants Scheme. Veronica Ward 20 M. J. TYLER | | \ \ | @} RIVERSLEIGH L, melanopyga L. aganoposis L. platyceps L. fletcher! >meOe® - Fig. 6. Geographic distribution of Lechriodus. A NEW SPECIES OF LECHRIODUS (ANURA: LEPTODACTYLIDAE) 21 TABLE I, Size af Lechriadus species in millimetres. Snout to vent length data mostly obtained from Zweifel (1972). Snout to vent length of donor L. aganoposis unknown, and maximum length of £, intergerivus extrapolated from Fig. 4. species snout-vent Jength ao GQ aganaposis 64-73 66-77 Jletcheri 42-48 45-54 intergerivus max. 41 melanopyga 38-47 46-60 platyceps 64-78 72-95 largest jlium examined size of ilium DAEF- acer. donor length VAE fossa diam = 24,2 8.1 3.5 AZ 16.3 42 1.9 237.5 14.4 40 1.7 §2 20.7 6.0 2,6 67 24.0 74.8 38 played a vital role in that she undertook the cataloguing and initial sorting of material, and prepared Figures. ]-2 and 4-6. Dr R. G, Zweifel and Dr C. W. Myers (American Museum of Natural History) lent ila of extant species and provided data about them, and Mr J. I, Menzies (National Museum & Ari Gallery, Konedobu) provided distribution records from Papua New Guinea. Mr P. Kempster prepared Figure 3. I am also indebted to Dr Keith Walker and Dr Margaret Davies for valuable discussions, and to the University of Adelaide for the provision of research facilities. The materials upon which this study was based were obtained through the support of the following funding bodies and organisations to M. Archer, S. Hand and H, Godthelp; Australian Research Grants Committee; Department of Arts, Sport, the Environment, Tourism and Territories; National Estate Program Grant Scheme; Wang Computers Pty Ltd; ICI Australia Pty Id; Australian Geographic Society Inc.; Mount Isa Mines Pty Ltd; the Queensland Museum; the Australian. Museum: the Royal Zoological Society of NSW; the Linnean Society of NSW; Ansett/Wridgways Pty Ltd; Mount Isa Shire Council; the Riversleigh Society and the Friends of Riversleigh, References Arcuer, M., Hanp, 8: & GoorHeip, H. (1988) A new Order of Tertiary zalamibdodont marsupials, Science 239, 1528-1531. Hann, S. J. (1985) New Miocene megadermatids (Chiroptera: Megadermatidae) from Australia with comments on megadermatid phylogenetics. Aust. Mamm. 8, 43-54. McDona.p, K. R, & MILLER, J. D, (1982) On the status of Lechriedus fletcheri (Boulenger) (Anura: Leptodactylidae) in northeast Queensland. Trans. R. Soc, S, Aust. 106, 220. Tyrer, M. J. (1972) An analysis of the lower vertebrate faunal relationships of Australia and New Guinea. In D. Walker (Ed,) “Bridge and barrier: the natural and culiural history of Torres Strait", Dept of Biogeography & Geomorphology, Research Schoo! of Pacific Studies, Australian National University, Canberra. (1976) Comparative osteology of the pelvic girdle of Australian frogs and description of a new ‘fossil genus. Trans. R, Soc. S. Aust. 100, 3-14. (1989) “Australian frogs’, (Viking O'Neil, Melbourne), ZWEIFEL, R, G. (1972) A review of the frog genus Lechriodus (Leptodactylidac) of New Guinea and Australia. Am, Mus, Navit, (2507), 1-41. OVER-SUMMERING REFUGES OF AQUATIC MACROINVERTEBMTES IN TWO INTERMITTENT STREAMS IN CENTRAL VICTORIA BY A. J. BOULTON* Summary Eight potential refuges for macroinvertebrates were sampled in two intermittent streams in central Victoria, Australia, during summer 1982-83 and summer 1983-84. Ninety-one aquatic taxa, mostly insects, were recorded. Receding pools harboured nearly three-quarters of these taxa; comparatively few were collected from the hyporheos or the water in crayfish burrows. Almost half the taxa were from refuges that did not hold free water. Macroinvertebrates persisted as desiccation-tolerant eggs (mayflies), larvae (chironomids and some beetles) or adults (beetles). There was remarkable similarity between the broad taxonomic representation in these refuges and those described for intermittent streams in Ontario, Canada. KEY WORDS: Intermittent streams, over-summering refuges, aquatic macroinvertebrates, Victoria, Australia, pholeteros, hyporheos. OVER-SUMMERING REFUGES OF AQUATIC MACROINVERTEDRATES IN TWO INTERMITTENT STREAMS IN CENTRAL VICTORIA by A. J. BouLToN* Summary Bouton, A. J, (1989) Over-summering refuges of aquatic macroinvertebrates in two intermitient streams in Vigloria, Trans. R, Soc. §. Aust, 113, 23-34, 31 May, 1989. Eight potential refuges for macroinvertebralws were sampled in two intermillent streams in central Victoria, Austraha, during summer 1982-83 and summer 1983-84, Ninery-one aquatic taxa, mosily insects, were recorded, Receding pools harboured nearly Uiree-quarters of these taxa; coniparatively few were collected from the hyporheos or the water in crayfish burrows. Almost half the taxa were from refuges thal did not hold tree water. Macroinvertebrates persisted as desigeation-tolerant eggs (mayflics}, larvae (chironomids and some beetles) or adults (beetles). There was remarkable similarity between the broad janonaniic representation.in these refuges and those described lor intermittent streams ip Ontario, Canada. KEY WORDS. Intermittent streams, over-summering tefuges, aquatic macromvertebrates, Victoria, Australia, pholeteros, hypotheos: Introduction Ephemeral (episodic) and intermittent rivers and streams drain over half of the Australian maintand (W,D, Williants 1983) but despite their ubiquily ard scientific interest, these systems have attracted little limnological attention (Boulton & Suter 1986; Boulton & Lake 1988), The situation is little better elsewhere (Williams £987), In intermittent streams, loss of water during the dry season is probably the most influential environmental parameter affecting the aquatic biota and has led to a wide range of physiological and behavioural adaptations (reviewed by Williams 1987). Bebavioural avoidance appears to play a major part in the survival of many stream invertebrates during drought. Williams & Hynes (1977) recognized eight distinct types of refuges that were used by the fauna of a tempvrary stream in Ontario during summer atid suggested that members of certain major taxonomic groups tended to over-suminer as similar stages in their life cyele. For example, Ephemeroptera and most Chironomidae over-summered us exgs whereas Gastropoda, some Odonata, Hemiptera. and Coleoptera survived the dry period as adults (Williams & Hynes 1977), In temperate Australian intermittent streams, tecolonization pathways and potential over- summiering tefuges have neyer been investigated and lidle is known about the physiological or behavioural adaptations exhibited by the aquatic * Centre for Stream Ecology, Department of Botany and Zoology, Monash University, Claytan, Vic. 3168. Present Address; Departinent of Zoology, University of Adelaide, G.PO. Box 498, Adelaide, S. Aust. 5004 biota (Boulton & Lake 1988), This study was aimed ar elucidating over-summering strategies of aquatic macroinvertebrates in two Intermitient streams in Victoria, 1 also was interested to see how closely the strategies used by biota in the intermittent Victorian streams matched those described by Williams & Hynes (1977) based on work done on intermittent streams in the northern hemisphere Materials und Methods Study Arey Two study sites were Jocated on the upper reaches of the Werribee River and two mare on its main tributary, the Lerdetderg River (Fig. 1). Both rivers arise on the southern edge of the Great Dividing Range approximately 100 km north-west of Melbourne andl flow south-east belore joining near Bacchus Marsh, north-west of Melbourne. Details of flow regime atid catchment vegetation are given in Boulton & Smith (1985); other physicochemical data are presented in Boulton & Suter (1986), ‘The two rivers differ in permanency: the Werribee River ceases flow almost annually whereas the Lerderderg River flows throughour sunimer for one year in three. On average, the Werribee River does not flow for nine weeks while the Lerderderg ceases flow for ax. The Werribee River did not Mow at all at one site (Spargo Creek, 8C) during the 1982 drought and only flowed for five mooths (late June ‘to Jate Novetnber) at the site downstream (Werribee Picnic Spot, WPS), The study pool at WPS dried up completely during the cnsuing summer, The following year, flow commenced in late June and continued for seven and a half months at both sites. Flow started in late May 1982 at both sites (Fireplace Ford, FF and Wheeler Road, WR) on the 24 A.J, BOULTON Lerderderg River and ceased in early December, At FF, the study. pool dried to a moist patch of leaves and water did nol appear in the depression until mid-March 1983. At WR, the study poal was dry for six weeks before it also refilled in mid-March; flow resumed at both sites in early May, continuing over the following summer. Physicochemical Monitoring of Potential Refiiges On 20.1.1983, diel ranges of ternperature in and around a receding pool at WPS were measured using 2 9-channel Miniature Intermittent Temper- alure Recorder (Grant Instruments, Cambridge, England). Thermistors were placed in shallow (5 cm) and deep (45 cm) water, below leat’ litter, beneath a large flat rock, in (he water of a crayfish (Engaeus sp.) burrow and in exposed grass in direct sunlight (regarded as. “air emperature” cf. normal meteorological practise). Recording commenced at 5,00 a.m, and ceased at midnight. At other times, spol walter temperatures (mercury: thermometer), dissolved oxygen (oxygen probe, Model 514A. Y.S.1., Yellow Springs, Ohio), To ’ Deyiestara ad “ Blakeville 4 ’ \ Fig, | Map of the study urea showing the locations of the four study sites: Werribee Picnic Spot (WPS) = | conductivity (conductiviry meter, Radiometer, Denmark). and pH (Metrohm pH meter, Model CH9100, England) were measured when potential refuges containing free water were sampled. Conductivity data were converted to values at (8°C (Ky3) (Bayly & Williams 1973) whereas dissolved oxygen was expressed as percentage saturation using the conversion table in Bayly & Williams (1973) and an appropriate correction factor for altitude, Biological Sampling af Potential Refuges A variety of collecting techniques was necessary to sample the diverse range of potential refuges: (a) An F.B.A. pond-net (300 zm mesh) was used to sample fauna in the receding pools. I vigorously shuffled along the bottom of the pool, sweeping the net from side to side across the disturbed path for 30 seconds for each sample. The size of the pool limited the number of samples that could be collected; while I was keen to ascertain the relative abundance of the fauna, | did not want to deplete the remnant populations, A nearby permanent lake (Shaws Lake, Fig. 1) was sampled similarly, To Trentham DIVIDING 37°27 5, Spargo Creck (SC) = 2, Fireplace Ford (FF) = 3, Wheeler Road (WR) = 4. OVER-SUMMERING REFUGES IN INTERMITTENT STREAMS 25 (b) Animals residing in the water in crayfish (Engaeus sp.) burrows (“pholeteros” sensu Lake 1977) were collected by carefully excavating the burrow opening and lowering a flexible plastic tube (6 mm internal diameter) into the burrow water which was sucked oul into a plastic bag, In. the taboratory, this waler was sieved (50 pm) with frequent washing to remove fine silt, (c) The hyporheas was sampled by digging holes inthe dry stream bed and sweeping a small hand- net (50 zm mesh) through the seepage. In some cases, it Was Necessary tO use a plastic lube to collect the water, Quantitative sampling of this habitat was not attempted. (d) Other potential refuges thar were qualitatively sampled while the streams were dry included the humid microhabitats beneath rocks, stumps and mats of leaf litter and dried filamentous algae, and among the roots of riparian plants (c.g, Cerex spp., Leptospermum flanigerum) and stranded Myriophyllum propinqguum (water milfoil) and Trelochin procera (water ribbon), Rotting wood was broken open and examined, and strips of bark were peeled from cxposed water-logged snags. Pieces.of decomposed wood were brought back to the laboratory for microscopic inspection. (e) Plastic bags were filled with surface (upper 10 cm) substrata and organic matter collected from the dry beds of riffles and pools. The water vontent of a subsample of the substratum was determined by subtracting the oven-dry weight (constant weight achieved after 48 h at 100°C) from the initial weight and expressing the value as a percentage. It was was tecorded. This process was repeated until no further species were seen. Subsequently, samples were taker) al wregular mlervals over the next fortnight, always returning the specimens to che tank. Some aquatia were marmraincd for several months to rear hatchlings through to adults to assist identification, Were identificd as far as practicable (sce Acknowledgments), Abundance was expressed qualitatively ias “present” {1-2 individuals), “common” (3-10) or “abundant™ (> 10); given the variety of collecting methods and the uneven sampling effort, more precise quantification was inappropriate. Results Physicochemical conditions in potential refuges Means atid ranges of spot water temperature, pH, dissolved oxygen and conductivity in two refuges that held free water when they were sampled are lisied in Table J, Not surprisingly, the ranges of these variables were grcater in {he teceding pools than in the burrow water of crayfish (Table 1), Hyparheic water was too disturbed during sampling to obtain reliable physicochemical data and dala trom Shaws Lake are too few to be useful, Continuous records of water temperature in a teceding poal al WPS illustrated the diel Nuctuation of temperatures in various refuges (Fiz, 2), Air temperature in direct suntight near the pool ranged from 3.5°C at dawn to 35°C early in the aflerncon (Fig. 2). The day was fine and clear with a light south-easterly breeze starting at 2.30 p.m. Sunrise TARE }. Means and ranges of waler lemperalure, pil, dissolved mxyzen and condueriviry in two. porentis! oversuntmering refuges, based upon n spot measurements Refuge Water Temperature PC) Receding x 3 pants range 7-25 n 25 Cray fist x 43 burnow range 7-12 waler 0 8 pH Dissolved Canduchivity Oxygen (Ky (% saturation) (Sem) 64 30.4 154 47-72 6-78 290 23 25 25 5.8 63 RR 5,2-6.2 45-79 66-J)0) 8 8 8 assuined that all weight lost during incubation was die to the evaporation of water. The test of the sample was emptied into an aquarium immediately upon return to Lhe laboratory and flooded with dechlorinated tap water, Within 30 minutes of immersion of the substrata, a hand-net (SOum mesh) was swept vigorously through the tank and the live contents examined using an Olympus stereomicroscope, and the rank. abundance of all invertebrates present was at 6.2) a,m, but the pool was shaded by surrounding forest until about 9.00 a.m. Sunset was at 8.41 p.m. Water temperature [i (he shallows (5 cm) lagged closely behind air Lemperature (Fig. 2a) while in deeper water (45 cm) the daily range was far less (Fig. 2b). Water in a crayfish burrow exhibited a diel range of less. than 3°C (Fig, 2b). The insulative capacity of several centimetres of eucalypt litter and dried filamentous algal mat approximated that of 26 AJ. ARULTON (a) Temperature (4G) ears ~~ Below leat ttter eee ee ee 0600 7200 1800 (b) t 4 | i. te an bans [Deepwater 1-7 Burrow water 410 Below rock 4. a ae _ oe 4 2400 0800 1200 1380 2400 Kig, 2 Variation in water temperature of several potential oyer-summering refuges in and around the study pool at WPS on 20,i.1983. See text tor details. a flar rock 1§ cm thick (cf. Figs 2a and b), During the day, conductivity remained at 270 »S/cm and pFI ranged from 5.5 to 6.1 (n = 6 determinations, mean pH = 5,6). Aquatic macroinvertebrates collected from potential refuges A total of 91 taxa was collected from the cight over-summering refuges sampled al four sites on the Werribee and Lerderderg Rivers (Table 2). Nearly three-quarters of these taxa over-summered in receding pools along the river beds. These pools also harboured tadpoles of Litoria ewingi (Dumeril & Bibron) (Hylidae) and Neobarrachus sudelli (Lamb) (Leptodactylidae); Jarge numbers of the latter perished when the pool at WPS dried up over the summer 1982-83. Several specimens of Galaxiux olidus Gunther (Pisces: Galaxiidac) were collected from the same pool. Few taxa were recorded from the hyporheos or crayfish burrow water (lable 2); individuals in Lhese habitats were usually tiny. Decomposed wood yielded smail oligochaetes, tipulids and boring chironomids (Orthocladiinae) while amphipods (Austrochiltonia australis) were found in curled up strips of Eucalyprus spp. bark near the margins of dried out pools. These habitats were not sampled intensively and it is likely that other taxa (e.g, helminthid beetles) may also use rotting, wood as a refuge over summer, Almost half of the taxa recorded were collected from refuges that did not hold free water when sampled (Table 2). Some of these (e.g. Nousia spp. and ?Dinoloperla (hwaitest) apparently hatched from desiccation-resistant eggs whereas others survived as larvae (chironomids, psephenids, helminthids and case-building caddisflies) or ss adults (dytiscids and helminthids) in moist microhabitats below rocks or mats of leaf litter and dried filamentous algae. Large numbers of Mmicrocrustacea (ostracods, harpacticoid and cyclopoid copepods) were recovered from dry substrata flooded in the laboratory. The water content of these samples was usually less than 10% by weight (range = 0,19 -—74,00%, mean = 7.71%, S.E. = 0.93%, m = 53) and did not differ significantly between samples collected from the beds of the pool and riffle habitats (Mann-Whitney U-test, p > 0.05). Comparisons among the refuges should be made cautiously because the collecting techniques and the numbers of samples taken from each habitat differed substantially. For this reason, J did not atlempt calculations of coefficients of similarity of faunal composition among refuges, Discussion The classification of over-summering refuges used by aquatic macroinvertebrates in temporary streams in Ontario (Fig. 3 in Williams & Hynes 1977) may be applied usefully to the data from the Werribee and Lerderderg Rivers. At a broad taxonomic level, the taunal elements over- summering in the various refuges are almost identival and du so al similar stages of their hfe cycle. This may seem unexpected given the differences in latitude, source of stream flow (snowmelt in Ontario, rainfall in this study) and the difference in the degree of species overlap with nearby permanent streams (Boulton & Suter 1986; OVER-SUMMERING REFUGES IN INTERMITTENT STREAMS 27 TABLE 2. Aquatic taxa recorded from potential refuges sampled over the 1982-1983 summer when the streams had ceased flow. Dead specimens were omitted. Abundance categories are qualitative (+ = present (1-)2 individuals), C = common (3-10 individuals), A = abundant (more than 10 individuals)) because of the different sampling frequency (nj and methodology (see text). * These individuals did not grow to identifiable size in the laboratory. ** (NMV sp.n) refers to the number of the specimen held in the voucher collection of the Museums of Victoria. Refuge Below Crayfish Receding Shaws Wood Dry substrata Hyporheos dried (Engaeus sp.) pools Lake and flooded in litter burrows bark laboratory riffle pool (n) (7) (12) (23) (2) (10) (47) (14) (5) TURBELLARIA Neorhabdocoela ?Mesostoma sp. + c Cc + Tricladida Cura pinguis (Weiss) Cc Cc NEMATODA Nematoda spp. + Cc A Cc A A Cc GASTROPODA Hydrobiidae Angrobia sp. A Cc + Glacidorbis hedleyi \redale - Ancylidae Ferrissia spp. Cc + Planorbidae Physastra gibbosa (Gould) A A BIVALVIA Sphaeriidae Sphaerium tasmanicum Cc (Tenison-Woods) OLIGOCHAETA Oligochaeta spp. Cc Cc A A o > > QD CRUSTACEA Janiridae Heterias sp. c + Ceinidae Austrochiltonia australis A A Cc Cc (Sayce) Atyidae Paratya australiensis Kemp Cc Parastacidae Cherax destructor Clark + Engaeus sp. C Cc Cc HYDRACARINA Limnesiidae Limnesia spp. + Cc Hygrobatidae Australiobates spp. Corticacarus spp. Mesostigmata Aquatic sp. ss a0 + > + + EPHEMEROPTERA Leptophlebiidae Nousia spp. Atalophlebia sp. ODONATA Lestidae Austrolestes Vio (Selys) 4 Corduliidae Hemicordulia 21au Setys + an > 28 A.J. BOULTON Refuge Below Crayfish Receding Shaws Wood Dry substrata Hyporheos dried (Engaeus sp.) pools Lake and flooded in litter burrows bark laboratory riffle pool (n) (7) (12) (23) (2) (0) (47) ~~ (14) (5) PLECOPTERA Austroperlidae Acruroperila utra (Samal) + Notonemouridae Austrocerca tasmanica (Tillyard) A Gripopterygidae ?Dinotoperla thwaitesi *Kimmins c C + HEMIPTERA Veliidae Micrevelia dubia Hale M. distincta Matipatil Notonectidac Anisops deanei Brooks A. Thackeri Brooks Corixidae Micronecta annae illiesi Cc Wroblewski M. a. tasmanica Wroblewski + O+ + ie COLEOPTERA Dytiscidae Antiporus blakei (Clark) A, femoralis (Boheman) Chostonectes johnsoni (Clark) Chostonectes spp. larvae Copelatus australiae Clark Hyderodes schuckardi Hope Lancetes lanceolatus (Clark) Liodessus shuckhardi (Clark) Necterosoma penicillatum (Clark) Necterosoma sp. larvae Platynectes decempunctatus c (Fabricius) Rhantus suturalis (Macleay) Sternopriscus mundanus Watts Gyrinidae Macrogyrus sp. + Hydraenidae Hydraena luridipennis Macleay + Hy, Vtricantha Zwick + Helodidae Helodidae sp. larvac ' Psephenidae Sclerocyphon striatus Lea larvae + Helminthidae Austrofimnius hebrus Hinton + A, maro Hinton ' A, “mermo” larvae (NMV sp, Ho)** Austrofimnius sp. larvae Simsonia tasmanica (Blackburn) larvae + DIPTERA Tipulidae Limonia sp. + Ormosia sp. + + + Psychodidae Psychoda sp. (NMV sp. 3)** p+ ©€ Q+ en 2) ++ C a“ ++ D+ 4+ + + A> FO AM at OVER-SUMMERING REFUGES IN INTERMITTENT STREAMS 29 Refuge Below Crayfish Receding Shaws Wood Dry substrata Hyporheos dricd (Engaeus sp.) pools Lake and _ flooded in litter burrows bark laboratory riffle pool (n) (7) (12) (23) (2) (10) (47) (14) {5) Culicidae Aedes spp. A Cc Anopheles annulipes Walker Cc Culex fatigans Weidmann Cc C. annulirostris Skuse + C. australicus Skuse + Chironomidae Ablabesmyia sp. 1 (NMV sp. 7E)** Cc + Ablabesmyia sp. 2 (NMV sp. 66E)** - Paramerina spp, (nr NMY sp. 32E)** Chironomus nr februarius (NMV sp, 136E)** Cc + Etnfeldia sp. + nr Dicrotendipes sp, (NMV sp. 34E)** Cc Riethia sp. (NMV sp, 5E)** + + Stenachironomus sp. (NMY sp. 3E)** + Calopsectra sp. (NMV sp. 22E)** Cc 4 +O Stempellina nr bausei or Monodiamesa sp. Cc + Orthocladius-Cricotopus complex (includes NMV spp. 12E and 160E). Heterotrissocladius sp. F Cc c Tiny chironomids* ‘ Cc A + + + Ceratopogonidae Bezzia sp. + Nilobezzia sp. + Stratiomyidae Stratiomyidae spp. + c + Cc Cc Empididae Empididae spp. + Cc 4 Dolichopodidae Dolichopodidae sp, 4 Muscidae Muscidae spp. Cc + + TRICHOPTERA Hydrobiosidae Ptychobiosis nigrita (Banks) + Hydroptilidae Hellyethira ?simplex (Mosely) + Calocidae Calocidae sp. Leptoceridae Leptorussa darlingtoni (Banks) Oecetis sp. Lectrides varians Mosely Cc Triplectides similis Mosely T. truncatus Mosely + Tiny leptocerids* + aaaanaa Total number of taxa 91 15 6 23 4 27 24,0 8 % of total number of taxa 16.5 6.6 747 25.3 44 297 26.4 8.8 boat ko] 50 ALT BOWLION Boulton & Lake 1988), However, physiological and behavioural adaptations employed by animals of common heritage that share gross morphological similarities and that are subjected io similar environmental selective pressures are likely to converge upon. a restricied miimber of solutions (parallel evolyition sensu Mayr 1963). In the Werribee and Lerderiderg Rivers, several commim laxa (Nousia sp, some geipoprerygid sloneflies) apparently hatched from desiccation. resistant eges (ef. Leboikuh! 1971; Snellen & Stewart 1979; Malicky 1982) and were among the fist invertebrates ta appear when Mow resumed, Similar finslings have heen reponed in other intermittenc streams (Harrison 1966, Churrer 1968; Hynes 1975; Ladle & Bass 1975; Williams & Hynes 1976, 1977; Abell 1984; Towns 1985}, Although simutiids (Diptera) were also common shortly after Mow resumed in the Werribee and Lerderderg Rivers. none emerged from the drv substrata flooded in the laboratory, implying chat epgs are laid by adults flying in when flow starts (ef. Hynes 1975; Abell (984). Mictucrustacea (ostracods, cyclopoid and harpactivaid copepods) emerged within hours of flooding dry substrata in the laboratory and harpacticoid copepods Were obscrved mating a day later. Morton & Bayly (F977) recovered ovigerous harpacticoid ferales only 24 hours after flooding some dricd mud from a temporary pool at Clayton, Victoria, Suypesting thal some species diapatise at an advanced stage of development (ef. Cole 1953). Another taxon, common shortly afler dry substrata were inundated, was a neorhabdocce] tcntatively identified as Mesostonta sp.. Bayly (1970) recorded Mesastoma from a temporary saline Jake in south- eastern Australia and observed thick-shelled exes in the uteri of some specimens |hat are apparently teleased when the animals die. A similar strategy for desiccation-tolerance has been observed in neorhabdocoels froma tentporary ditch m England (Cox & Young 1974). Gastropods in the Werribee abd Lerderdere Rivers survive drought enter by secreting a protective epiphmem (eg. Phvsasiza eibboso, Ferrissia spp.) (cf. Kenk 1949; Eckbtad 1973; Leger & Talin 1973) or by closing their operculum (eg, Angrolia sp, and Glacidorbis hedlevi) (Boulton & Smith 85), Most tended (o aestivate in moist intcrohahitats under stumps, dry algal mats and leaf litter (cf. Strandine 1941; Klekowski 1959; Casey & Ladle 1976). Whe bivalve SpAgerium sosmenicum probably minimizes water loss by closing its valves. Other members of the Sphaeriidae are oveyiviparous and broud their young while surface wirter is absent (Heard 1977; Horobach ef af. 1980, McKee & Mackie L981). Acstivating juveniles have been found buried in the subsisaium (Way er ul. 1Y80) bul I did not recover any from my study sites. Parastacid crayfish found in the Werribee River over-summer in their burrows where water temperatures remain quite constail, Surprisingly few other invertebrates (pholeteros sensw Lake 1977) appeur to use this refuge (ef. Creaser 1931; Williams eral 1974; Williams & Hynes 1976; Wigeins er ai, 1980). The pholeteros in other Victorian Engeets spp, burrows is also depauperate (Horwity ef ail, 1Y85), Nevertheless, this refuge seems to tw important for the survival of janirid isopods in (he Werribee River. Isopuds usually are absent from femporaty warers because they lack desiccation: resistant stages and are sedentary {Williams 1985). ‘The Water in burrows constructed by fish serves as sources Of recolonists after droughts in some sircams overseas (e.g. Tramer 1977; Glodck 1978} but this refuge was not evident in the Werribee and Lerderderg Rivers. Few taxa were abundant in the hyporheos of the study sites during summer. This paneity may reflect the crude sampling methods because ayuptiputls, janirid isopods, stoneflies, molluscs and oligachaetes have been recorded Fram the hyporheas of the intermittent Brachina River in South Australia (W.D. Williams 1983), Overseas, the hyporheus is. considered to be an important refuge from both droughts and floods in sorte temporary and permanent streams (Clifford 1966; Williams & Hynes 1977; Williams 1977, 1984). However, in desert streams whose beds ate mainly composed of unstable sand, hyporheic dormant stapes are rare because of high temperatures in the dry streambed and severe scouring of the channel during flash floods (Gray 1981, Fisher er a/. 1982). One refuge not considered by Williams & Hynes (1977), possibly uncommon in Ontario, is chat provided by decomposing wood debris, abundant ajong the banks and stream beds al many Avust- falian intermittent streams, In my study, this refuge harboured oligochaetes, amphipods, tipulids and chironomids, More intensive sampling is likely ta vield further taxa because other workers haye recorded larze numbers of xylophitous taxa from permanent streams in Oregon (Afiderson ef a/, 1978; Dudley & Anderson 1978) und New Zealand (Anderson 1982), In the Werribee and Lerderderg Rivers, taxa aestivared under racks and mats of algae (where temperatures remained constant) as adults (e.g. amphipods, dytiscids} or larvae (e.g, chironomids, stratlomyids), Some latval stages seemed surprisingly tolerant of desiccation; for exutmple, a large water penny larva (Sclerocyphan stréatis), collected from the exposed surface of a flat rock on the bed of a riffle that had Wot Mowed for 19 OVER-SUMMERING REFUGES JN INTERMITTENT STREAMS 3) weeks, resumed activity immediately after ituimersion in water from 4 oearby pool. Sinllar tolerance of desiccation by larvae of Scteracyphor spp. has been described in Tasmania (Smith I98t) and Queensland (Smith & Pearson 1985). Adult aquatic Hemiptera {e¢. corixids, nolonectids) and Coleoptera (e.3, dyjiscids) prob- ably Ply in from nearby permanent waters. Such aerial recolonization of temporary pools and streams by these groups is commonplace {ee Fernando 1958, 1959; Fernando & Galbraith 1973: Williams & Hynes 1976, 1977; Wiggins ef al, 1980; Abell 1984; Williams 1985), | also found several species of adult dytiscids over-suminering below rocks and dry litter near the margins of pools at SC and WPS, This strategy seems less well-known for this group; only D.D. Witliams (1983), Boumezzough (1983) and McKaige (1980)' fin a temporary pond near Colac, western Viclonia) have reparted similar observations. Although the receding pools harboured most of the taxa thal over-summer at the study sites, they appeared to be the most physivochemically “harsh” refuge that [ sampled. The pools experienced a considerable diel range in water temperature, oxygen levels were Frequently below 20% saluralion and pH levels fell to Jess than 5. Conductivity: rises as the water evaporates and the pools are ofien stained dark brown with eucalypt leachate. Similar canditions have been observed in receding pools in Other intermittent streams in Australia (Towns 1983, 1985, Smith & Pearson 1987) and North America (Slack 1955; Larimore er af 1959; Clifford 1966; Harrel & Dorris 1968), The invertebrates that over-summer in hese pools have yarious adaptations that allow then to tolerate such physicochemical extremes, For example. larval dyliscids come up to the surface of the pool to oftain air through the terminal abdominal spiracle whereas adull Uytiscids store air beneath their elytra (Brition 1970), Some chironomid larvae (eg. Chironomus Wi febrwarivs) are particularly abiundanl in the receding pools and use haemoglobin to facilitate oxygen uptake (Colless & McAlpine 1970). MayNy nymphs (Asalophilehia sp.), also vommmna in this refuge, have large ills that are constantly oscillated to enhance respiration (Boulton & Lake 1988). Terrestrial oviposition by two species af leptacérid caddisfly common in the Lerderderz River (Leptorussa derlingioni and Lecirufes verians) may also be an adaptation to low or unpredictably fluctuating oxygen levels (Towns 1983}, Development of vulnerable juvenile stages ' MeKaige, MLE. (1980) Emergence and development of a@atatic invertebrate communities from dried awd after flooding. ASe Hens thesis, Department of Zovlogy, Manosh University, (Unpubl.). In a physicochemically harsh environment may be avoided or accelerated by ovovivipariry exemphilied by the stonefly Austrocerce tavmaniza, also recorded by Towns (1985) in a South Australian Intermittent stream. Although there is little known about the thermal tolerances of aquatic macrpinvertebrates in Australian intermittent strearms, it appears that many taxa can cope with short-term exposure to exiremes of warer teniperature Ibis likely that they remain near the bottom of the pool where the temperatures may be as much as 15°C cooler than those of |he surface water exemplified by the study pool at WPS. Such stratification is untcommon jn shallow pons {@2. Byars 1960; Butler 1963; Moore 1970; Hartland-Rowe 1972). However, Eriksen (1966) recorded surface-botiom differences of 9-16°C jn temporary turbid puddles less than SNe deep, and a shallow (10 em) rockpool fn a stream in the Pyrences had’a surface temperature of 29,09C while the bottom was 19°C (Chodorowska & Chadorowski 1966), Less marked stratification in pools in North American intermittent streams has been reported by Neel (1951) and Slack (1955), Isolated pools remaining in the stream bed are important oyer-sumimeting refuges for aquatic macroinvertebrates in intermittent streams elsewhere (e.g, Mack 1955; Paloumpis 1958; Larimore ef af 1959; Williams & Hynes 1976; Abell 1984), In temporary streams in Ontario, they also provide excellent breeding environments due to the ease with which they warn up and che abundanr plant food ihat develops within them, and they enable species with long-lived aquatic staves to complete their fife- cycles (Williams & Hynes 1976), Sintilarly, in the Werribee and Lerderderg Rivers, Ihese puuls support arich fauna stthough environmental conditons are harsh and predators are numerous (Boulton & Suter 1986). Taxa whose aquatic life-spans are brief enough to be completed while water 6 presen! are under less selective pressure to adop! Ufese stralewes than specles whose aquatic development takes longer than the period that water persists. Unfortunately, we lack information on the duration of aquatic stages Of many Australian macroinvertebrates and few generalizations may be drawn. Most crustaceans and moliuses listed tn Table 2 probably live longer than a year and this may account for their abilipy to survive in refuges other than the pools, Alchough many other raxa (eg. chironomids, culicids) can coinplete their aquatic stages in a matter of weeks. their survival in intermittent streams depends upon when their eggs hatch and how long water persists afterwards, Interpretation of the stenifieayce of many of these refuges will be possible when more information on the life historles of 32 AJ, BOULTON macroinvertebrates in Australian intermittent sireamns i4 available. In summiary, there appear to be five major over- simimerlog strategies enpploved by the fauna of these two rivers; i) rolerating extreme and variable enviroumen- tal conditions in the remaining pools, ii) surviving in moist microhabitats below stones, stuinps and mats of dried algae and leaf litter, and in rotting wood, iii) over-summering. jf flerohabitats where environmental conditions are relalively mild and comstan! such as in the hyporheos or the burrow water of crayfish, i+) surviving as desiccation-resistam! stitges Lit the dry subsira(um, and, v) living in nearby permanent water-bodies and flying in and ovipositing when flow resumes. The relative contributions from each of these tefuges reflects the nature of the substratum (ex, Clifford 1966; Gray 1981), the amount and pattern of discharge during the previous spring, and the severity of the summer, This last was illustrated by the 1982 drought when all the pools in the upper reaches of the Werribee River dried :completely, extinguishing several Commion taxa (eg. the shrimp Peratya oustratiensis) that | never recorded the following year. Differential survival of fauna in these various refuges undoubtedly influences the commurity composition of the stream and has a profound effect upon ils ecological succession during, the ensuing period of flow. Acknowledgments Itis a pleasure to acknowledge the enthusiastic eevouragement aud guidance fram De Sam Lake, Cenrre for Stream Ecology, Monush University where 1 did this research supported by i Commonwealth Postgraduate Research Award. Many taxonomists identi tied my macroinvertebrates; | thank Dr M. Harvey (Hydravarina), Dr C.MLS. Watts (Dytiscidae), Mr T. Weir (Heiminthidae), Dr J, Davis (Psephenidae}, Dr Ci. Theischinger and Ms C, Yule (Plecoptera), Dr P. Suler (Ephemerupiers), Dr A. Wells (Hydroptilidac), Ms R. St Clair (Platyhelminthes, Lepluceridac), Dr P. Horwitz (Parastacidac), Dr]. Lansbury (Hemiptera), Drs W. Pander and 8. Smith (Gastropoda) and Dr A. Sokol (Anura}, Dr. R. Marchant, Mr L. Metzeling and Mr, PL Lilywhite allowed me access to laboratory facilities and the vaucher collection at the National. Museum of Victoria, T was helped in the field by Leon Barmuta, Mark Harvey, Pierre Horwitz, Paul Huniplirles, Sabine Schreiber, Anthony Sokol, Gaye Spanguro, Alistair Suren and Cathy Yule and the publican at Spargo Creek deserves special mentun for his provision ol’ an over-sutimering refuge for semi-aquatic vertebrates, Mr Devo Mortun kimdly supplied me with aquaria and other equipment, Lam indebted to Drs Margaret Davies, Sam Lake and Keith Walker, Prof. Bill Williams and two telerees for constructive comments on the manuscript, References Anni, D. t. (1984) Benthic invertebrates of some California intermittent streams. pp. 46-80. Jn Jain, S- & Moyle, P, (Eds) “Vernal Pools and Intermittent Streams”, (University of California, Davis Institute of Leolugy, Calilornia). Annesson, MN, H. (1982) A survey of aquane insears associated With wood debris in New Zealand streams. Muuri Ora Wy, 2-45. Seen. J. 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Limaut 16, 7-514, Frasanoo, CH. (1958) The coloiisatio( of small freshwater halwrats by aquatic insects, [, General discussion, methods and cplonisarion Wi te aguatic Coleoptera. Ceyloe J Sh 1, 1F-154- _ (1959) The solowisatiun of small freshwater habiacs by aquatic imsects. 2. Hemiptera (the walerbugs). Shia. 2, 5-32 _ & GALURAITH, DL (1973) Seasonaliry god ~ dynaniigs of aquatic insects colonssing small habitats. Ferk. Interna. Verein, Limnal, 18, 1564-1875, Pisiie. SO. Gray, b, J, Gaiam, NB. & Busen, DE (1962) Temporal succession in & desert stream following, flush flooding, ee’, Monogr, 52, 93-110, Tiowek, GS (1978) The importance of catfish burrows Jn Maintaining fish populations of Wopical freshwater sleams in western Ecuador Frefdiana 13, 1-8, Gray. L, 1 (1981) Species composition aya life histories al aquatic insects in a lowtend Sonoran desert swream, Amer. Midl. Not, 106, 229-242. Harve, X, 1. & Dorms, T. 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M..& BouLron, A, (1985) The burrow habitat of two sympatric species of land erayfish, Eavoews uroserrers and &. rh ei (Dyespoda, Parastaridae), icy Not, 102, 88-197, HUNGERFORD, HB. (1920) The biclozy and ecology af aquatic and semlaquatic Hemiptera. Aansas Saiv. Sci. Bell, 1, 3-328. Tiyses f, D. 11975) Aneual cycles of macro-invertchrates of ariver in southernGhana. Freshwar, Bir 5, TI-83. Kiow, R, (1949) The animal life oT jemporary and Permanent pands in Southern Michigan. Mise. Pults, Afuy. Zool Univ, Mich 71, 1-66. Kuikowski, 8. 2. (1959) Survival of desiccating molluscs Planorbis planorhis | and dependence on some environmental Conditions, (English sumroary from Polish), Poh Arch. Ayadrabiel, 5e, 71-89, LapLe, M. & Bass. J, A. (L975) A new species of Metacnephia Crosskey (Diptera, Simulildac) from the south of England with notes an its habitat and biology. Hydrebiologia 47, 199-207. Lake, P. S. (1997) Pholeteros - the faunal assemblage found in crayfish burrows. Aust. Soc, Linnel, News! TS, 57-60, LARIMORE, R. W., Cuicoers W. Fo & Hecknome, C. (1959) Destruction and re-establishment of stream fish and invertebrates affected hy drought, Dons, ener Fish. Sac. 88, 21-285. Lecien, P. & Tavin, J, (1973) Comparison de rulsseaus permanenis é| fermporaires de la Provence Calcsine. Avin. Lbanol. 9, 273-292. LenmkunL, DM, (197) Stuneslies (Plecoprtera: Nemouridac) from temporary lenric habitats in Opegan. Aner Midl Nat. 85, 514-515, Macicky, H. (1982) Adaptations of Lwa stanefly species to intermittent streams in Crete, Aan, Mus: Goutundris 5, 179-182. Mays, F, (1963) "“Acimal Species and Evolitiow.” (Flarvard Universily Press, Massachusetts.) McKee, PM, & Mackie. G L. (1981) Life hestary adaptations of the fingernail clams Spfiyreriiyerr vccidentiale and Musewtiuh securis (a ephemeral habitats, Cy, J. Zool. 59, 2219-2229. Moore, W. GQ. (1970) Limnulogical snidies of tempuraty pount Tn sonihensrern Lousiana, Sonrawest. Nav. 15, F=110, Morton, Dh W & Barty i. AE. (1977) Syudies on the ecology of same¢ té¢mporary freshwater pools in Victoria with special reference ta micrgcrustaceans. Awsy. J, Afar, Fréeshwot, Res, 28, 439-454, NEEL, J, K. (1951) irterrelations of ceraiu physical and chemical features in a hardwater limestune stream. E&rofogy 32, 368-39). PaLoumeis, A. A. (1958) Responses of minnows td Mood and drought condivions in an intermittent stream, /awe Sraté Coll. ¢. Sci. 32, 547-56), Stack, K. V. (1955) A sindy of factors affecting stream roductivity by the comparative method, saves, wdianc Lekes Streams 4, 3-47. SmitH, J. A. (1981) Two Tasmanian species of Solerotyphoh Blacktum (Colegptera, Psephenidae) with notes On their life history and distribucian, J 4usy Entomol. Suc. 20, 277-288, Smitn, R. EW, & Pearsom, R. G. (1985) Survival of Seleroeyphan bivular Carter (Coleoptera, Pseyhenicac) in an intermittent stream in North Qucenstand. hic. 24, 10/-102, —_— (1987) The jnacro-invertebrale communities of fremporary pools i ao inlermittem stream in meopncal Queensland, Hydroticlogia 180, 45-61, SWELLEN, R. & Siewarr, K, W. (1979) Thee fife cycle of Perlesta placida (Plecoptens Perlidge} in an intermittent stream bn northern Texas. Ann. Entomal, Soc Arner 72, 659-666. STRANDINE, B. J. (1941) Effect of soil moisture anu ygae on the survival of a pond smi! during periods af relative dryness. Nuretifus 54, t28-030. Towns,.D. R, (1983) Terrestrial oviposition by two species of caddisfly in South Australia {frichoplera, Leploceridae). J. dust. Entomol Soc, 22, 113-118. —__— (1983) Limnological characteristics af a South Australian inrernuttent stream, Brown Hill Creek, 4isy, J. Mar, Fresiiw. Res, 36, S2t-837, TRAMER. E. 1. (1977) Catastrophic mortality ol stream bey Lrapped in shrinking pools, Aerie Aftal New. 97, 469-478. , 34 A.J, BOULTON Way, C. M., Hornsacu, D. J, & Burky, A, J. (1980) Comparative life history tactics of the sphaeriid clam, Musculium partumeium (Say), from a permanent and a temporary pond. Ibid, 104, 319-327, Wicains, G. B., Mackay, R. J. & Situ, I. M. (1980) Evolutionary and ecological strategies of animals in annual temporary pools, Arch. Hydrobiol. Suppl. 58, 97-206. WILLIAMS, D. D. (1977) Movements of benthos during the recolonization of temporary streams. Oikos 29, 306-312. (1983) The natural history of a Nearctic temporary pond in Ontario with remarks on continental variation in such habitats. Int. Rev. Gesamten Hydrobiol. 68, 239-53. _____ (1984) The hyporheic zone as a habitat for aquatic insects and associated arthropods. pp. 430-455 Jn V.H. Resh and D.M. Rosenberg (Eds) “The Ecology of Aquatic Insects.” (Praeger Scientific Publishers, New York.) (1987) “The Ecology of Temporary Waters.” (Croom Helm, England.) & Hynes, H. B. N. (1976) The ecology of temporary streams. I. The fauna of two Canadian streams. Int, Rev. Gesamten Hydrobiol. 61, 761-787. _____ (1977) The ecology of temporary streams. IL. General remarks on temporary streams. Ibid, 62, 53-61. » WILLIAMS, N. E. & Hynes, H. B. N. (1974) Observations on the life history and burrow construction of the crayfish Cambarus foliens (Cottle) in a temporary stream in southern Ontario, Can. J. Zool. 52, 365-370. WILLiAMs, W. D. (1983) “Life in Inland Waters.” (Blackwell Scientific Publishers, Carlton, Victoria.) _____ (1985) Biotic adaptions in temporary lentic waters, with special reference to those semi-arid and arid regions. Hydrobiologia 125, §5~110. THE LARVA AND NYMPH INSTARS OF ODONTACARUS (LEOGONIUS) ADELAIDEAE (WOMERSLEY) (ACARINA: TROMBICULIDAE: LEEUWENHOEKIINAE) BY R. V. SOUTHCOTT Summary The larva of Odontacarus (Leogonius) adelaideae (Womersley, 1944) is redescribed, and a lectotype designated. Larvae have been collected repeatedly on domestic cats in suburban Adelaide, South Australia, and reared experimentally to protonymphs and deutonymphs. The larva, protonymph and deutonymph are described. Experimental transfer of larvae from cats to man is described. A key is given to the known deutonymphs of Leogonius, and revisional notes on the classification of the larvae. KEY WORDS: Taxonomy, Odontacarus, Leogonius, South Australia, larva, protonymph, deutonymph, rearing, cat. THE LARVA AND NYMPH INSTARS OF ODONTACARUS (LEOGONIUS) ADELAIDEAE (WOMERSLEY) (ACARINA: TROMBICULIDAE: LEEUWENHOEKIINAE) by R, V. SouTHcotrT* Summary Sourncor, R, V, (1989) The larva and nymph instars:of Odontacarus (Leogontus) adelaideae (Wamersley) (Acarina: Trombiculidae: Leeuwenhoekiinue). Trams. R. Sac. S. Aust. 113(00), 35-45, 3) May, 1989, The larva of Odontacarus (Leogonius) adeloideue (Womersley, 1944) is redescribed, and a lectolype designated, Larvae have been collected repeatedly on domestic cats in suburhan Adelaide, South Australis, and reared experimentally to protonymphs and deutonymphs. The larva, prolonymph and deutonytnph are described, Experimental transfer of larvac from cats to man is described. A key is given tothe known deutonymphs of Leogerins, and revisional notes on the classification of the larvae, Key Worns: Taxonomy, Qdontecurus, Leogonius, South Australia, larva, protonyayph, deutonymph, rearing, cat. Introduction Womersley (1944) described Leenwenhoekia adelaideae from several larval trombidioid mites, collected from domestic cats in southern suburbs of Adelaide in 1931 and 1941, and three collected from rats, in Cairns, Queensland, in 1939, He had earlier (Womersley 1934) referred to some of the same (1931) larvae as “Lewwenhoekia (sic) australiense (sic) Hirst 1929" (sic), ie. as Leeuwenhoekia australiensis Hirst, 1925, originally described from larvae collected at Ashfield, Sydney, New South Wales, bul subsequently recorded more widely in Australia and the south-west Pacific area {Gunther 1952; Wharton & Fuller 1952), Following revisions of the taxonomy of the leeuwenhoekiine larval mites, (hose known from Australia, New Guinea and south-east Asia {previously referred to Leeuwenhoekia Oudemans, 1911 and Acomatacarus Ewing, 1942) are referred ty Odontacarus Ewing, 1929, subgenus Leogoriis Vercammen-Grandjcan, 1968 (see Southcott 1986a), In the present paper the larva of Oduntacarus adelnideae is redescribed, The first atternpt fo rear larvae was in 1941, but the protonymphs and deuronymphs described below were nol obtained until 1950-1952. A key is given for deutonymphs of Leogonius. The successful transfers of partly-fed larvae from cats (0 feeding on man ts described, Materials and Methods Shde-mounted mites in the South Australian Museum Adelaide (SAM) referred to O. adeluideae und collected by D. C. Swan in 193] and R. V. Southcott in 1941 were examined, They were Originally mounted in gum-chloval media, bar * 2 Taylors Rosd, Mitchum, S. Aust, 2062, showed evidence of extensive remounting to gum- chloral or polyvinyl alcohol-based media, with associated damage. Some specimens labelled as adelaideae from NSW. were excluded, The specimens recorded by Womersley (1944) from Cairns, Qld, 1939 were not located, Further larvae were collected topotypically in 194] and between 1946-1952 from the ears of domestic cats. Some were mounted in water-soluble media (generally gum-chloral based), whilst attempts were made to rear others to later instars. In 1941 larvae freshly collected from cats. were confined under a watch-glass attached to my forearm with adhesive strapping. Although the mites re-attached and increased in size, no successful transformation occurred, Mites were again collected from the ears of domestic cals [rom Ure same Site, during summer- autumn of 1946-1952, and placed on strips of damp blotting paper in sealed tubes at ambient temperatures. In une case a nearly-intact protonymph was obtained, In several cases transformation to deutonymph occurred. All specimens were then preserved in lactic acid, or dry, until studied, generally 35-40 years later. The devtonymphs and residual pieces of larval of protonymphal pelts were slide-mounted, through lactic acid to Hoyer's gum chloral medium (Baker & Wharton 1952), Microscopy was by a Leitz Orthelux microscope with phasc-contrast and polarizing facilities; its camera lucida was used to make the line drawings, Terminology and abbreviations are as in Southeort (1986a,b). All measurements are in um unless otherwise stated. Prefixes: ACB to aurhur’s registration numbers, N to those of SAM. Odoatacarus Ewing Odontacarus Ewing, 1929, p, 188 (for synonymy sce Southcotl, 19868, p, 171, aid contained references). 36 R. V. SGUTHCOTTE Fig. | Odontacerus adelaideae (Womerslcy), larva. A Dorsal view of lectotype, legs on left omitted heyond trovchanters: legs on right completed from specimens in same postures a5 lectotype, leg [ from specimen ACB520BA2, leg 11 from ACB270A, leg IIT from ACH270B. B Gnathosoma, dorsal scutuny and adjacent structures from paralectotype NIORR333 (ACR941), © Same for specimen ACBS20BA2. D. Dorsal idiosomal seta. F Ventral ihosomal seta, F Tibia 1 and tarus I, dorsal aspect, of paralectotype N1988333, (D, BE ww scale on right; otber figures to seale on left. THE LARVA. AND NYMPH INSTARS OF ODONTACARUS (LEUGON/US) ADELAIDEAE 37 Definition of larva; Trombidioidea (as defined by Southeott 1987) with anterior median dorsal seulum, with nasus, Wo anteromedian setae, two anterolateral setae, two posterolateral setae, and two sensilla, cach with non-expanded sensillary seta. Leg scemental formula 6 6, 4, Eyes 2 + 2, sessile, Defiatlion af deutornymph and wails: Trombidigidea with anterior median crista, expanded anteriorly to a blunt arrow-head-like shape (sagilla), bearing two nurmal setae, with two §énsilla towards ils posterior end, each bearing a slender, setulose sensillary seta. Leg segmental formula 7, 7, 7, Deutonymph with two pairs of suckers (acetabula) along each lateral penital yalye, adult with three pairs.. Dorsal idiosomal setae not leal-like, nor divided, bul may be present im two distinct forms. Eyes absent.. Type species: Trombicuia dentata Ewing, 1925, p. 257, Subgenis Leogonius Vercammen-Grandjcan, 1968 Definition of larva: as in Gulf (1979, p. 143). Type species: Leeuwenhoekia australiensis Hirst, 1925, p, 150, Odonracarus adelaideae (Womersley) FIGS |A-E, 3-6 Leuwentoekia (sic) austrriense: (sic) Womersley. 1934, p, 217 (partint). Leeiwenhvekia austrifiense; (sie) Womersley, 1937, p. 82 (purtiin). Leeuwenhoekia qusiraliensiss Womersley & Heaslip, 4943, p. 141 (partim); Thor & Willmann, 1947, p. 323 (partim). Leeuwenhoekia adelaidewe Womersley, 1944, p, 105; Gill, Moule & Riek, 1945, p, 29; Gill & Parrish, 1945, Plate 3; Taylor, 1946, p. 228. Acomutecnms adelaideae: Womersley, 1945, pp. 98, ILI; Southecott, 1957, p, 149, Acomatacarus (Acontatacerws) adelatdeuwe> Wharton & Fuller, 1952. p. 97. Odontacarus adelaldeae: Southcott, 1973, pp. 46, 103; 1976, p, 139; 1978, p. 16 (unnamed); Lee & Southeott, 979, p. 35; 1980, p. 7. Ordontacarus (Leogonius) adefaidese: Southcott, (96a, p. 180, Descripuon af larvar Lectatype: Colour in life orange red. Length of idiosuma (mounted on slide) 370, width 285; total length from tip of cheliceral fangs to posterior pole of idiosonia 463. Dorsal scutum wider than long (nasus in¢luded); nasus well developed, tongue-like, slightly pointed anteriorly, slightly waisted, meeting body of scunum ai approximately tight angles; anterolateral angles slightly obtuse, rounded; lateral borders slightly corvex; posterolateral angles obtuse, rounded; posterolateral borders sinuvus; posterior angle obtuse, rounded, Scutalac narrow, tapering, slightly bhutied at dip, with pointed, auistanding setules, Sensillary setae with 8-10 setules in distal half, Sensillary sockets level With PL seutalae bases and set slightly obliquely, Shield lightly porase, wilh Lwo larger pits near each PL angle. Standard and other data of scutum and legs of the lype series as in Table J. Eyes oval, curijoined, posterolateral to dorsal sculum; anterior eye with maximum diameter 16, posterior WW. Dorsal idiosomalae normal, liglily tapering, slightly blunted at tip, moderately setulose, arranged 2 (“humerats"), then 6, then th vague rows across dorsum; total abour 35, Ventral surface of idiosoma with a pair of pointed, setulose setae, 33 Jong, with centres of bases 34 apan, between coxae LIL. Behind coxae 11 Opisihovoma wih 42 setae, curved, setulose, pointed, but the more posterior setae longer, slightly blunted, resembling posterior dorsal jdiosomalae, Anus 24 long by 13 across; 16 setae anterior 1 level of middle of anus, and 26 posterior, Urstigma well chitinized, oval, 25 long by 16 wide. Coxalae 2, 1, 1, long, well setulose, tapering, pointed. Lateral coxala 1 44 long, medial coxala | 68, coxala I 53, coxala II) 46. Dorsal tracheal Opening present between posterolateral edge of gnathosoma and anterior border of coxa | (or overlying latter). Dorsal tracheal opening present between posterolateral edge of gnathosoma and anterior border of coxa I (or overlying latter). Gnathosuma normal, Combined chelicerae bases ce. 73 across, by 9L long from tip of cheliceral fangs to posterior edge of cheliccral bases. Fangs stout, curved, blunt-pointed, with 3-5 strong retrorse teeth along concave (flexor, dorsal) ede, and 5-7 blunted slenticles along convex (extensor, ventral) edec. Galeala 26 long, simple. Palpi normal, palpal seta! formula B, B, BNN, So + 7Ik Palpal tibial claw three-pronged, No supracoxala to gnathosoma or legs. Description of lees of larva (trom ACBS20BA2 and ACB270A,B); Leg lengths (including coxae and claws}: | 425, 11 355, II] 425. Scobalar formulae: trochanters 1, 1, 1; femora 6, 5, 4; genua 4, 4, 4; tibiae 8, 6, 6 (including 2 mastalae on uibia IN); tarsi 26, 16, U3 [including one mastala on Ul}. leg specialized seraé as folluws: Leg | {from ACBS20BA2y SaGel,38ad(21 long), VsCiel.66pul4), SeGel.7ipd(22), SoTil.59d(16), VsTil-88d¢2), Sofil.9Sad(10) (i.e slightly distal to Vs), Leg [1 (from ACB270A4): SoGell.3dpd(20), VsGell,64pd(), Solill 42pd(16), SaTiULS9padilo). Leg Wd (from ACB270B); SoGell.26pd(21), SoFilll.SOpd(27). 38 R. V. SOUTHCOTT Tig. 2 Odentacarus adelaideac (Womerstey), larva. Ventral view of lectotype, tegs on Jet omitted beyond trochanters, legs on right completed as in Fig. 1- THE LARVA AND NYMPH I[NSTARS OF ODONTACARUS (LEOGONIUS) ADELAIDEAE 9 TABLE |. Metric data for type series of Odontacarus adelaideae (Womersiey) larvae a N1988331 (ACB67) Character Lectotype n AW 66 5 PW 84 5 3B 24 5 ASB 50 5 PSE 28 5 L 78 5 LA 16 5 LB 62 5 LN 27 5 Ww 96 § AP 28 5 AM 36 5 AL 36 5 PL 49 4 PL/AL 1.36 4 AMB li 5 Sens 55 3 PW/LB 1.35 5 DS 28-55 4 Hum! 55 4 MDS 28-31 5 PDS 34-39 5 Gel 57 4 Til 60 3 Tal(L) 94 2 Tal(H} 27 2 Gell 44 5 Till 50 4 Tall(L) 78 5 Tall(H) 27 5 Gelll 51 2 Till 68 2 Talil() 95 2 Talii(H) 24 2 AW/AP 2.36 5 AW/TULT 0,97 2 PW/Tilll 1,24 2 PSB/SB 1.17 5 ) Humeral seta length @ For maxintum values Both tarsi I and II bear a large central dorsal solenoidala: SoTal.43d(15) and FaTal.4Gad(3) (from ACB270A); on tarsus IL (from ACB270B) 1s Sofall.40d(15), and FaTall.37ad(4) (ie. slightly proximal and anterior to Sofall (in usual convention of legs being considered stretched out horizontally at right angles to main longitudinal axis of mite), Pretarsal formula 1, 1, 0. Tarsal claws normal, slender, falciform, simple, neomedian longer than anterior and posterior, each with a fringe of minute onychotrichs, Description of Protonymph: (Fig. 3) from mounted specimen ACB522BA5) Colour in life not observed, presumably orange-red. Shape irregularly spheroidal, 630 long, maximum width 430, Skin without setae, thin, more-or less smooth but with many minor creases, possibly artefacts of mounting. mean s.d Observed Tange 70,80 £3.56 (66-74) 87.80 +2.77 (84~91) 28.20 43.03 (24-32) 51,00 +1.73 (30-54) 29.80 +1.79 (28-32) 80.80 +3.38 (78-86) 16.80 +110 (16-18) 64,00 +2.83 (62-68) 25.40 +2.30 (22-28) 95.60 +0.55 (95-96) 30,40 +1.82 (28-32) 37.20 +1.64 (36-40) 35.80 £0.45 (33-36) 53,25 +5.44 (49-61) 1,478 +015 (1.36—1.69) 10.60 +1.67 (9-13) 53.67 +3.21 (50-54) 1.374 +0,08 (1.28-1.47) 55.502 +1.00 (55-57) 55.502 +1.00 (55-57 }e 32.802 +1.30 (31-34)2 40.602 $3.13 (38—46)2 56.75 $1.26 (55-58) 62.00 +2.00 (60-64) 95,50 42,12 (94-97) 29,00 +2.83 (27-31) 47.40 +2.30 (44-50) 54.00 $2.35 (50-56) 84.00 +3.46 (78-86) 27.00 +0.71 (26-28) 53,00 +2.83 (51-55) 70,50 $3.54 (68-73) 101.50 $9.19 (95-108) 23.00 +1.41 (22-24) 2.334 £0.14 (2,13-2.52) 0.985 +002 (0.97-1.00) 1.215 +0.04 (1.19--1.24) 1.066 +0.12 (0.93-1,19) Developing limbs represented by finger-like processes; those of chelicerae and palpi still adnate to main mass of protonymph, those of legs becoming free. A few setae and a few fragments of larval tracheae adhering to protonymphal skin. Description of Deutonymph: (Figs 4-6) (from slide- mounted specimen ACB522AA/DI, supplemented by other specimens) Colour in life orange. Idiosoma ovoid, more pointed anteriorly and posteriorly, not waisted, flattened below; no division between propodosoma and metapodosoma. Propodosoma more or less conical; posterior pole of idiosoma rounded, Length of idiosoma (from tip of sagitta) 512, width 370; over-all length from chelicerae tips to posterior pole of idiosoma 634, 40 R, ¥. SOUTHCOTT Fig. 3 Odontacanss udelaideae (Womersley). Protonymph, entire, specimen ACBS22BAS_ Propodosoma bearing dorsally a short crista 151 long, with sagitta about I5 across, carrying [wo almost nude pointed setae 41 long, Sensillary area 46 across, wilh two large sensilla, each with a filiform sensillary seta 100 long, with faint setules in distal half. Crista produced posteriorad beyond sensilla, tapering to a blunt point 47 behind level of sensillary centres. Crista surrounded by normal setae similar to those of whole of dorsum of idiosoma. Dorsal idiosomal setae unexpanded, pointed, lightly setutose, curved, 18-46 long, the posterior setae the longer. There is no clear Separation into morphologically different types of setae in any arca of the dorsum. Ventral surface of idiosoma thickly covered with pointed, setulose setae, 15-34 long; setules more prominent than for dorsal setae; setae less serulose but longer towards posterior pole, where they ate similar to the posterior dorsal idiosomalac. Genital aperture 86 long, by 50 wide; valves lightly chitinized; cach media! valve with two or three setae 13-18 long at about mid-area, slender and Jess serulose than surrounding setac. Anus 49 long by 31 wide, with 2 or 3 curved setulose setae along each lateral valve, similar 10 surrounding setae. Fig, 4 Odontavarus adelaideae (Womersicy), Deutonymph, entire, dorsal aspect, specimen ACBS21AA/Di. Legs Jong, fairly thin; femoral to tibial segments more of less cylindrical, Lengths (including coxae and claws) I 960, I] $70, IIL 895, IV 925, Leg scobalae pointed, setulose; interspersed among them are many pointed sensory sctac (spinalae). A small vestigiala, 5-6 long, present on each of tibia I] and genua | and Ji, distally and dorsolaterally. Tarsus I outline and elongate oval, other tarsi more cylindrical (see fig, 4). Other morphometric data as in Table 2. Palpi normal, with generally lightly setulose setae (many appeating almost nudc), but along dorsal (extensor) edges of segments a number of unilaterally setulose setae (pectinalac), Palpal tibial claw stout, blunted, with four accessory thickened setae on dorso-medial aspect. Palpal tarsus clavate, ovoid, with several setulose setac, but carrying also a cluster of terminal smooth sensory sctae. Cheliceral fangs fairly robust, curved, blunt- pointed, with a row of about 10 minute denticles along flexor (concave) edge. THE LARVA AND NYMPH INSTARS OI’ ODONTACARUS (LEOGONIUS) ADELAIDEAE. 4l Fig. 5 Odontacarus adelaideae (Womersley). Deutonymph: gnathosoma and adjacent structures, dorsal aspect (partly in transparency), from specimen ACBS21AA/DI1, 42 R, V. SOUTHCOTT TABLE 2. Morphometric detq far deutonymphks of Odontacarus adelaideae (Womrersley). Character Specimen n ACBS22AA/D1L cL 104 8 5B 25 9 PDS 33-46 9 Gel 135 9 Til 187 9 Tal(L)* 244 9 ‘Tal(H) 67 9 Gell 75 i] Till 110 9 TalI(L.)4 124 9 TalI (FL) 29 9 Gelll 77 7 Tull 118 8 TalL(L)2 117 R Tall(H) 31 8 GelV 146 8 TiIV 198 8 Tal V(L)3 175 R Tal V(H) 36 8 Tal(L)/Til 1,30 9 mean sd. Observed Tange 115,38 +12.71 (95-135) 26.56 + 3.61 (20-31) 47,78 + 3.837 (44-456) 141.11 +14.00 (118-160) 206.1! $20.33 (178-226) 246.67 £18.26 (218-273) 68,56 + 4.45 (60-75) 79.63 + 5.58 (72-86) 113.00 +10,51 (96-128) 12.78 +13.27 (111-148) 30,00 + 343 (24-35) 86.86 + 811 (77-100) 129,88 $13.66 (1LI-146) 127,38 +11.33 (114-143 29.38 + 3.78 (25-34) 154.50 +11.65 (137-172) 219.63 +2423 (188-253) 182.00 +14,79 (162-205) 34.50 + 4.17 (29-41) 1.23 + 0.056 (1,15-1.30) a ) CL = cristal Jength, measured from anterior lip of sagitta to level of midpoints of sensillary sockets ® For maximum yalues 4 Omitting claws and pedicle. Identification of larvae of Odontacarus (Leogonius) in Australia Comments The recording of about 42 ventral opisthosomal setae in O. adelaideae larva invalidates caption number 12 of my earlier key to the larvae (Southcott 1986a pp. 179-180), which was based largely on previously published descriptions. Accordingly, captions 12~14 should be replaced by the follawing: 12 More than 45 ventral opisthosomal setae prESen fobs ew ghte pupelem att ber ec dale ta Be ELE 4 a pene 1 Fess than 45 (about 42) ventral opisthosamal setae present, AL seutalac in range 33-40 pm long. PAS 34-46 um long........ O. adelaideae (Womersicy) 13 More than 75 dorsal idiosomal setae present, .... O. austratiensis (Hirst) (including QO. pirsti{ Womersley, 1944)4) Less than 70 dorsal idiosomal setae present,.... 14 14 PL/AL <1.200000 200. ee eee O. novaguined (Womersley, 1944)? (including O, /ougipes (Womersley, 1945)4) PLVAL S20). epeitpa tte ole Sete 15 1 From Goff (1079), 2 Synonymized by Domrow (1956) and Golf (1979), 4 These two species (as larvaé) were synonymized by Goff (1979), However, see the comment of Southcott (1986a, p. 191). Key to deutonymph instars af Leogonius (Australia-SW Pacific area) 1 Dorsal idiosomalaé arising from small platelety and about 25 pm long! 9305 0000 0 QO. navaguinea (Womersley, 1944), Dorsal idiosomalae without the above combination of CHATACHETS~ y/-'.. ote/eteb aw edt tice mesetelits orice oe 2 2 Posterior dorsal idiosomalae over 5() zm Jong, Tal/Til mean 1,54 (tarsus | 210-228 ym long, tibia 1 130-154 1) a ar ©. australiensis (Hirst, 1925) (syn. O. Airsté (Womersley, 1944)?,3 Posterior dorsal idiosomalae generally loss then 50 ain POT sen sgee alo wf et ura aces - eh eo a h tetettht net 3 3 Tarsus I longer than 300 um {tarsus 1450 pm, tibia 1440 um, Tal/Til = 1.00, dorsal idiosomalae to 30 um longyl......... OL longipes (Womersley, 1945) Tarsus I less than 300 pm long.............. 205 4 4 Posterior dorsal idiosomalae 15-20 xm lony, ‘lal/‘Til 1.36 (tarsus T 185 pm tong, tibia I 136 um)4 Mey emery es pan pedictis ¥ O. undyi (Radford, 1946) Posterior dorsal idiosomalae generally to about 48 pm long. Tal(L)/Til 1.15-1,30 (tarsus 1. 218-273 ym long, tibia |! 178-226 pm............005. O. adelaideae (Wometsley, 1944) 1 Womersley (1945) 2 Domrow (1956) 4 Goff (1979) 4 Nadchatram (1963) ‘THE LARVA AND NYMPH INSTARS OF OUDUNTACARUS (LEOGONIUS) ADELAIDEAE 3 Fig. f Odontacarus adelaideae (Womersicy),. Deutonyniph, taternal genitaha aud anus, from specimen SCBS21AA/DI.. Metlerial examined of Odontacarus adelaideae: Type series: South Australia: Glen Osmond, Nov, 1931, D.C, Swan, from ears of cat, 3 paralectotypes (N1988333-N1983335, ACBK9$1-943). Unley Park {not Unley, as stated) by Womersley 1944), 171,194], RK V. Sguthvott. from ears of cat, one lectorype (NI988331. ACB67), one paralectotype (NI988332, ACE 66}. The lectotype was designated as N1988331 because it has not been remounted since 1943 {indicated by my. notes) and is, therefore, the most intact. Also it is in the same attitude as Womersley’s (1944) figure. I have excluded from the type series the three specimens recorded in the orginal description, from “Cairns, Queensland, 1939 W. G. H”[easlip], “from rats," as they can no longer be found in the collection of the South Australian Museum, and were presumably excluded by Womersley at some time after his 1944 paper. He did not reler to Chern in 1945. in his last publication on Odanfocerus. Other material: 5, Aust. Unlev Park (all trom ears of cats of topotype arva, cullewial RVG.) LiLIM4L, one specimen, ACH68; 9.1), 1941, several specimens, ACB72, 74, 928; LiitI941, 3, ACBT5-77; LLG.1941, 4, ACB 78,79 (S.ii1-3941, 2, ACBBO, 8); 29.711, 1941, lL, ACBS2; 30.11.1941, 1, ACBB3; 4iy.194], 1, ACBS4; 6,i¥.1941, 1, ACBRS; [1.[v.1941, 3, ACB86 BCE; I4ivi941, 1, ACHE7; 16,x7,1941, 3, ACBIAIH-102; 23.x7, 1941, 5, ACBIO3: 29.xi.1941, 5, ACBI04; 22. xi1.1946, several, ACB270; 16.41.1947, several, ACB319; 31-xiL.1950, several, ACB520; SLxii.1950, two batches of several larvae, from which deutonymphs were reared, ACB521, 922; 25.i1.19Si, several, ACB523; 1iii1951, several, ACBS24; 9.710951, 2, ACBS25; 61.1952, several, ACBS560 (deutonymphs rewred). Successful larva to nymph rearing experiements Lurvae were handled jn the 1950-]982 series of experiments as stated above. One protunymph (ACB522BA5) Was obtained, but the transformation was not observed, and discovered only when jhe tube was examined on 29.1195]. Earlier examination on 14.2.1951 revealed nothing unusual (however in these wet tube studies observation is at times difficult), Efeven deutonymphs were obtained from the 1980-1952 experiments. Nine of these came from experiments ACBS2) aid $22, with larvae collected On 31.xi/.1950, and nymphs having emereed over 15-29,1.1951, With batch ACBSG6O, collected on 6.11952, two newly emerged deutonymphs were found on 27.01.1952 The finding of an intact protonymph was lortunale, as this instar is generally unrecognizable alter the emergence of the deutonymphs From the above experiments, the larva to dewonymph {ransformations took 15-24 days Domprow (1956) recorded that in QO. qusiraliensis deutonymphs took 25-28 days to cmerge in his laboratory in Brisbane. Transfer of larvae of O. adclaidese from cats to malt On the afternoon of [1 April 194] | transferred seven attached larvae (batch ACB86) From the ears of @ cat to the skin of the dorsum of my forearm, under a watchglass kept in place by sticking plaster, On removing the watchglass 2.5 hr later ] Found two small larvae attached to my skin, Removal of the watchglass periodically was necessary, as condensation made detailed observations impossible, Next morning the underlying skin was sore, Three maculopapules with attached mites were observed at different sites; these maculopapules were 446 mim wide, and were itchy. That afternoon the area of erythema around the attachinent sites had grown to 10-11 mm wide, these had become conjoined (only two mites were now attached). Sinall while specks were seen at the summits of the papules (four bencath one mite, three beneath the other), 44 R, V. SOU PHCOTT presumably mite faeces, Small vesicles were present at the summit of each papule al (he mile attachment siles (20-22 hours after initial attachments (codes as § 4+ 20-22 hr). The tiching, redness and vesiculation increased, The attached mites increased slowly in size to “half- grown”, Ac‘S + 39 hr the whole area under the watchglass was itching, diffusely swollen and somewhat indurated; this area was 45 mm across, and by this stage the mites were considered ful) gmwn (as compared with the larvae that had been observed on the cat’s ears). The papules had become sO prominent as to press upon the covering watchglass. The mites continued to make amall faecal pellets beneath rhem. By S + ¢c, &) hr twa vesicles had broken and were coving serum, No lymphangitis or lymph- adenopathy had developed, After the miles had detached the swelling reduced rapidly, and scabs developed over the broken skin surfaces, By S + 8 days the papules had retrogressed to being almost Hat, and the erythema was fading. No general symptoms attributable Lo (he mite bitings or thelr lesions develuped, Unfortunalely, this attempt ro obtain mite dentonymphs by human feedings was \insuiccessful, A summarized account of this experiment was recorded earlier (Southcott, 1973), Acknowledgmeits [thank the South Australian Museum far access to material in their collection, This work was supported by the National Health and Medical Research Council and the Austrafian Biological Resources Study, References Basce, BE, W. & Wieaston, Ci Wo (1952) “An introduction to acarology.” {The Macniillan Campans, New York). Dewtrow, R, (1956) Three new Australian chigger nymphs {Acarina, Trombicutidae), Proc, Linn, Soc. NSW RI 42), 144-152, Ewing, H. B. (1925) A contributian to our Knowledge of the taxonomy of chiggers meluding the descriptions of &@ New genus, six new species aod a new Variely, 4 eer JA trop, Med, 5 (A), 251-265. (1929) A manual of external parasites. (Charles C, Thomas, Springheld, Mlinvis & Baltimore, Maryland} (1942) Remarks on the taxanomy of same American chiggers. (Tronthiculinae}, including the descriptions of new genera and species... Parasiral. 28 16), 455-499. Gi, DA, Moure. G. RK, & Rien, Ro EF (1945) Trombidiosis in sheep in Queenstand, Aust, ver, 21 12), 22-31. _, & Pawrisy, E. (1945) Microphotography as an aid to the identification of trombiculine larvae. J, Coune, Sek Ind, Res, V8 (4), 298-300, Gorr, M. L. (L979) The genus Odontacarys LAcati: Trombiculidwe> in New Guinea, with descriptions of four new species. nied, Entomerl. 1S (2), 144-154, GunTiter, C.F. ML (1982) A check list of the crombiculid lurvate Of Asin and Australasia. Proc. Linn Soc NSH, 77 (1-2), 1-60. Hikgst. S (1928) On a harvest bug (Leen wenhvekia oustraliensis, sp. tt.) attacking humun heings a1 Sydney, New South Wales. frams & Sar, frop, Mea, & Ave, 19 (3), 150-152. Lite. DC, & Souvucort, R, V, (1979) Spiders and ather arachnids of South Australia, Pp. 29-43, i Leonard, B. E. (ed.) “South Australian Year Book. No. 14: 1979" (Australian Bureau of Statistics, South Australian Office), (Reprinted 1980 as separate, pp I-15. 4South Australian Muscym, Adclaide)), NabcHarrkAM, M, (1963) The latva and nymph of Celantacaras audyi (Radtord) (Acarina; Trombiculidae). Pacific lasecis 3 (3), 535-440. OVOPMANS, AL. (IVI) Acarclogische aunteekeningen NAXVL, Ent. Ber, Amst, S458) 137-134, Ranrora ©. &. (1946) New species of larval mites (Acarina: Trombiculidae) from Manipur State, India, Proc Zool, Soe Londen 116 (2), 247-2165, Soutucomr R. V. (1957) The genus Acomiaracaries (Acarina: Trombiculidac. . Deseription of three new species fram Triniry Ray, orth Queensland, Trans, R. Soe, 9. Aus, BU, 146-155. — —— (1973) Survey of Injuries to Man by Australian Terrestrial Arthropods. {R. ¥. Souihcott, Mitcham, Suuth Australia (1976) Arachnid and allied syndromes in the Sustain region, Ree, Adel. Childs. dfosp. 1 (1), 97-187, ——__ (1978) Australian harmful arachnids and their alhes, 4R. ¥, Southeott, Mitcham, South Australia). (1986a) The genus Odontacarws (Acarina; Trombiculidite}, 1, Observations on the fife history and morphology of Odontacarus swan n. sp., and related forms. Ree. S. Awsi. Mus, (9 (12), 149-200, (1986b) Descripnon of Odantucuras veitoni sp. nov(Acarina: [rombiculidac). fbid. £9 (14), 213-217. ______ (1987) The classification of the mite families Trombellidge and Johnstonianidae and related groups, with 2 deseriptian of a new larva (Acarina: Frombcitidac: Notirotrombidium) from Nanh America, Trams. R. Soc. 5, Aust, Wt (1h, 2542, Tavion, FH. (6946) Section |. Description, pp. 7-234. (m Taylor, FH, & Murray, R. E. (Eds.) ‘Spiders, ticks and miles including the species harmful wm man in Australia and New Guinea" (Serviee Publication No. 6) School of Public Healt & Tropica) Medicine, Uuiversity of Sydney, Toor, S & Wittmann, Cy (1947) liombidiidae. Das Vierreich Tho, xxix-xxxvi & L&7-S41. VERCAMMEN-Ghas DEAN, Po A. (1968) “Phe (higgec Mites of the Far Basr (Avarjna: ‘Jrombivulidac & Lecuwenhoeklidae). At illustrated key and a synopsis; some new tribes, gener and subgenera" 4U, 5S. Army Med. Research and Development Command, Washingtan, DC.) Warton. G. W., & FULLER PL S (1952) A manual of he cluggers, The biolugy, classifiedtion, distriburcien, and importance to man of the larvae of the family Drombicriaas;Agetinad. Mem Ent. Sac. Washingion 4, [-185. THE LARVA AND NYMPH INSTARS OF ODONTACARUS (LEOGONIUS) ADELAIDEAE 45 WomeErRSLEY, H. (1934) A revision of the trombid and Australia and New Guinea. Trans. R. Soc. S. Aust. 68 erythraeid mites of Australia with descriptions of new (1), 82-112. genera and species. Rec. S$. Aust. Mus. 5 (2), 179-254. ______ (1945) Acarina of Australia and New Guinea. The _____ (1937) A revision of the Australian Trombidiidae family Leeuwenhoekiidae. Ibid. 69 (1), 96-113. (Acarina). /bid. 6 (1), 75-100. , & HEasip, W. G. (1943) The Trombiculinae _____—(1944) ~Notes on and additions to the (Acarina) or itch-mites of the Austro-Malayan and Trombiculinae and Leeuwenhoekiinae (Acarina) of Oriental Regions. /bid. 67 (1), 68-142. THE COMPOSITION AND DISTRIBUTION OF THE VEGETATION OF NORTH-WEST EYRE PENINSULA BY F. TIVER, A. D. SPARROW & R. T. LANGE* Summary The vegetation of inland north-western Eyre Peninsula, South Australia, is sampled for phytosociological analysis by transect quadratting. Using a combination of numerical classification (Bray-Curtis dissimilarity with average-linkage clustering) and ordination (detrended correspondence analysis), seven main vegetation associations are identified: Eucalyptus dumosa- Westringia rigida, Eucalyptus gracilis-Stipa, Eucalyptus gracilis-Eucalyptus oleosa, Eucalyptus oleosa-Melaleuca pauperiflora, Eucalyptus oleosa-Enchylaena tomentosa, Eucalyptus brachycalyx and Eucalyptus yumbarrana-Triodia irritans. The last occurs on the siliceous Moornaba sands, and the other six on the calcareous Woorinen Formation. Environmental parameters correlate only weakly with a floristic gradation over the calcareous soils, suggesting that historical influences may be important in determining the distributions of the vegetation types. KEY WORDS: Eyre Peninsula, vegetation survey, vegetation analysis, classification, ordination, plant associations. THE COMPOSITION AND DISTRIBUTION OF THE VEGETATION OF NORTH-WEST EYRE PENINSULA by F. Trver, A. D. SPARROW & R, T, LANGE* Summary Tiver, F,, SparRow, A. D. & Lance, R. T. (1989) The composition and distribution of the vewetation of north-west Eyre Peninsula. Trans. R. Soc. S. Aust. 113, 47-61, 31 May, 1989. The vegetation of inland north-western Eyre Peninsula, South Australia, is sampled for phytosociological analysis by transect quadratting. Using a combination of numerical classification (Bray-Curtis dissimilarity with average-linkage clustering) and ordination (detrended correspondence analysis), seven main vegetation associations are identified: Eucalyptus dumosa—Westringia rigida, Eucalypris gracilis~Stipa, Bucalyptus gracilis-Eucalypius oleasa, Eucalyptus oleosu—Melaleuca pauperiflora, Eucalpytus oleosa-Enchkylaena tomentosa, Eucalyptus brachycalyx and Eucalyptus yumbarrana-Triodia irritants. The last occurs on the siliceous Moornaba sands, and the other six on the calcareous Woorinen Formation. Environmental parameters correlate only weakly with a floristic gradation over the calcareous soils, suggesting that historical influences may be important in determining the distributions of the vegetation types. Key Woros: Fyre Peninsula, vegetation survey, vegetation analysis, classification, ordination,, plant associations, Introduction Until zecently, the vegetation of north-western Eyre Peninsula remained relatively unknown, The first detailed study was that of Crocker (1946). Unfortunately the accompanying map only extends northwest as far as Poochera, where the vegetation is described as Eucalyptus aleesa ~ E.gracilis - Edumosa edaphic complex. Specht (1972) also included only the southern-most portion of the area in his maps, describing the vegetation south of the Eyre Highway as Aucelypius socialis — E. gracilis open scrub and that north of it as Eucalyptus inerassata — Melaleuca uncinata open scrub. Boomsma & Lewis (1980) described 16 communities, defined by the tall dominants, which occur on north-west Eyre Peninsula. Their distribution maps are not detailed enough to be able to extract the relative importance of any of these communities. Descriptive surveys such as those mentioned above are not always capable of repetition by others (Nilsson 1986). For this reason, most modern Surveys use quantitative data of the whole flora, and various forms of computer-assisied multivariate analysis, which result in a higher degree of objectivity and repeatability, A mumerical classification of the vegetation of western Eyre Peninsula, one of the few such studies in South Australia, was undertaken by Margules & Nicholls (1987). Using complete species data from 104 plots each of 0.1 ha, cluster analysis was used to identify six vegetation communilies on western Eyre Peninsula. This survey did not extend northwards * Dept of Botany, University of Adelaide, GPO Bux 498, Adelaide, \S. Aust. S001 onto the sand dune formations, Statistical models were constructed to relate the occurrence of the six communities to three environmental variables: depth of carbonate layer, distance from the coast and latitude. Although allowing prediction of likely vegetation types in other remnant patches, these factors do not adequately describe the environmental factors important in determining the vegetation pattern. The aims of this study were to classify (he yegetation of north-western Eyre Peninsula (Fig. 1) using repeatable, numerical methods, and to provide some explanation of the main factors controlling the vegetation pattern, Fare r DB Geduna Yor) finale £ SP pa — ae 100 ?cO Kilomactes Big. §. Showing the location of the study area on Torth- western Eyre Peninsula, S.Aust. 48 F. TIVER, A, D. SPARROW & R. T, LANGE Geology, Landfarm and Soils of the Study Area Most of the surface geological formations of western Eyre Peninsula (see Fig. 2) are derived from calcareous sands, It has been suggested that these sands were blown in from the exposed marine continental shelf during periods of low sea level and aridity, which occurred during the Pleistocene ice ages (Crocker 1946; Twidale & Campbell 1985). Another possibiliry is that the sands represent palaco-dune systems that were formed by wave and wind action during periods of high relative sea level (Short ez al, 1986). The Bridgewater Limestone is composed of calearenite, grains of calcareous sand cemented together with calcite crystals. In the maritime zone itis exposed as coastal. cliffs but is otherwise covered by unconsolidated, deep, calcareous shelly sand of low water-holding capacity (Wright 1985), The Bakara Calcrete consists of calcarenite in which the shell fragments have been progressively dissalved and replaced by nodular and platy calcrete. To reduce the ecological complexity of the survey, these coastal limestone formations were excluded from the study. The Woorinen Formation closely corresponds to the Chandada Plain landform in which the dunes have consolidated to form an undulating plain with MOLINEAUX SANDS . Maltee no surface drainage (Twidale & Campbell 1985). Calcrete is usually present as a bed of nodules beneath the surface. The soils are calcareous throughout, and range in texture from sands to clay loams. They are usually shallow and reddish brown, but sandier grey varieties and redder varieties with higher clay content have been reported; they have been extensively cleared for agriculture (Wright 1985). Areas of the calcareous sands normally associated with the Bridgewater Limestone also occur on the Woorinen Formation. Near the coast these sands are coarse and shelly, but further inland they are firmer with less-apparent shell fragments, Wright (1985) reports deficiencies in a wide range of minerals in these sands, The Molineaux or Moornaba Sand is of extremely recent origin (c, 10.000 years bp. onwards) and composed of sand dunes aligned in a NW-SE direction. Bleached white siliceous sands with a yellowish B horizon occur on the dunes. The underlying scils of the inter-dune corridors contain calcareous carths typical of the Woorinen Formation. Methods Sampling Strategy A system of transects. was chosen as the best compromise between equal coyerage of the region ‘ e eee AUX SANDS 10 Kilometres ™, . Nunjikompita ™. ae, *Pimbaacla WOORINEN FORMATION §Wirrulla BaKARA GCALCRETE Fig. 2 The study area, showing major surface geological farmations (after Blisset 1969). VEGETATION OF NORTH-WEST EYRE PENINSULA 49 fo 14 20 25 Kilometres Nunjikompita © . s 3 . * g ° S my Pimbaacts . * Wirtulla «& Fig. 3 The study area, showing location of sites along seven iransecis, Shading represetuts uncleared vegetation. and efficiency of time spent on data collection. Transects were placed in suitable areas of native vegetation, running north and south from Ceduna, Maltee, Mudamuckla, Nunjikompita, Pimbaacla, Wirrulla and Cungena (Fig. 3). Transects were extended southwards only as far as the boundaries of the Bridgewater formation and its overlying coastal sands, and northwards as far as. was accessible by four-wheel-drive vehicle. In the northern part of the study area, sites were placed 1 km apart, but in the southern part of the area, which is widely cleared for agriculture, native vegetation was often restricted to roadside reninants, and some sites had to be located as much as 2 km apart. Field experience showed that quadrats of 0,1 ha as recommended by Whittaker (1978) and used by Margules & Nicholls (1987) were an appropriate size to achieve a good representation of species present without overlapping different vegetation types. One $0 x20 m rectangular quadrat was measured at each Site, vegetation Datu In each of the 162. sites sampled, all seed-plant Species were recorded, although only native perennials were used for analysis. Annual species were excluded because their occurrence depends on recent seasonal events; their inclusion would therefore lower the repeatability of the results. Introduced species were also excluded, since they do not form part of the natural vegetation pattern, Total cover was estimated for cach species present, and expressed as a semi-quantitative score, an adaption of the cover scale as Braun-Blanquet (1932); see Table }. Environmental Data Antivipating some influence on floristic composition by various disturbance factors. disturbance by erosion, grazing, animal tracks, undergrowth clearance and carthworks was visually assessed on a scale ranging from O (nil) to 4 (Severe). Sixty-two sites rated 2 (mild) or greater were excluded from analyses for definition of vegetation types, Landtorm was assessed in the field as dune-crest, dune-flank, rise or plain (including wide interdune corridors). A soil core was dug in the centre of each quadrat and the depth to limestone recorded, in intervals of 10 cm, Texture was determined using the field method described by Northcote (1971). Existing rainfall maps were not detailed enough for the purposes of this study, so a rainfall map was 50 F, TIVER, A. D. SPARROW & R. T. LANGE constructed, using complete Bureau of Meteorology rainfall records for 24 recording stations within or near the study area. This enabled the average rainfall at each site to be estimated and classed into 12 mm intervals, Vegetation Analysis The need to classify vegetation raises the question of whether vegetation types occur as discrete entities with distinct boundaries, or whether classifications are merely abstractions imposed by ecologists onto a continuously variable vegetation (Greig-Smith 1983). If vegetation consisted of random combinations of species a continuum would result and classification into discrete groups would be an entirely arbitrary process. If, on the other hand, variation was always discontinuous and no intergradation occurred, there would be a finite number of vegetation types in existence, and classification would be a simple matter. The actual case lies between these two extremes, ranging from the near discontinuous and distinct vegetation types to almost continuous ecotones (Webb 1954). GROUP DESIGNATION NUMAECR DF SITES iN GROUP 165 Consequently, a combination of two complementary analysis techniques is recommended when investigating vegetation patterns; a classification to define vegetation types, and an ordination to examine the relationships between the groups and the extent of ecotonal behaviour between them (Gauch 1982). For classification purposes, the dissimilarity between all pairs of sites was calculated with the Bray-Curtis distance coefficient, and then sites TABLE 1. Semi-quantifative scores expressing percentage cover (adapted from Braun-Blanquet 1932). Cover Score Percentage Cover absent 0-1 1-2 2-10 10 — 50 50 -— 100 UhwWheS o& BRAY-GURTIS GISTANGE COEFFICIENT +7Q +75 50 285 -90 | Eucalyptus dumgeg 5 Wastringie riglda W Eucalyptus gracilis 6 Stipa sp+ GUNGENA 293 | tH Eucalyptus gigogg 30 Eucalyptus grucilig Iv Eucalyptus vleoua 17 Melaleuca paupariflora Vv Eucalypius plegsa 15 Enchylaens tomentosa VI Eucalyptus brachycalys it Vil Eucatyptus yumbarrang 4 Triodia inritans T a a T art 1 ——— Fig. 4 Summary of dendrogram trom clustering analysis showing arrangement of groups at 0.65 distance level. VEGETATION OF NORTH-WEST EYRE PENINSULA iJ grouped from the least to the most dissimilar by the unweighted pair group method using arthmetic averages (UPGMA; Legendre & Legendre 1983) to produce a hierarchical dendrogram, the branches of which represént vegetation types. The complementary ordination method chosen was Wetrended correspondence analysis, DCA (Gauch 1982), Ordination methods such as DCA rank sites along a series of axes which describe the major floristic variation. The results are displayed as a scattergram from which continuity and disjunction of vegelation types can be determined and with which environmental factors can be correlated. The numerical taxonomy computer software package NTP (Belbin é7 a/, 1984) was used for both analyses. For the purpose of preparing vegetation maps, the sixty-two disturbed sites were returned to the data set, artd a further cluster analysis. performed. By the proximity of these sites on the dendrogram to sites of known vegetation type, most of the remaining sites were identified as belonging to one of the major vegetation types. Resuiis The classification dendrogram is summarised in Fig. 4. Field notes indicated that the groupings defined at the 0.65 distance level were the most readily interpreted, a higher level producing groups with large internal variation, and a lower level producing too many groups for consideration. Seven major vegetation types were identified at the 0.65 level, one of which, ne Vl, was widely AMIS = (Variation Explained 21 Gb? dissimilar from the other groups, only fusing with them at 0.90 level, A minor vegetation type was alsa identified. Represénted by only one site, Cungena 29, this site was dominated by Callitris preissii, rather than a eucalypt species. The relationships between the vegetation types can be seen from the ordination; a plot of the first two ordination axes is given in Fig. 5. The first axis explained 50% of the variation in the data set, and corresponds to the first major branching of the dendrogram, separating vegetation type VII from the others, The separation between the remaining six communities is largely supported by the ordifiation, although there is a certain degree of overlap, most marked between types [ and II, and types V and VI, The relationship of these six Vegetation types to each other obviously tends towards the “ecotone” rather than the distinct “association” end of the scale in vegetation pattern as described by Webb (1954). The species consistent in each of the vegetation types are presented in Table 2 and Fig. 6 summarises the associated landforms, soils and rainfall. Table 3 gives a full listing of species found in rhe seven vegetation types. Understorey species are equally important as shrubs and trees in delineating vegetation type. The vegetation types, their distributions (Fig, 7) and their environmental correlates are discussed below. The types have been named “associations” according to usual ecological practice, even though they do not fully fit the formal definition af “association” given by Braun-Blanquet (1932). AMIS § (Vatlatien Exptained SOM) Tig. 5 DCA ordination showing grelauioriships between the seven major vegetation types. Symbols denoting vegetation type are as follows: O , |; 4, 1h 4, 15 @, IV, @, Ve 0, Vis VW, VOL. 52 F. TIVER,, A, D, SPARROW & R. T. LANGE TABLE 2, Plant species characteristic of the seven major vegetation types, Marked species had u cover score of two or greaier when present, and could be considered daminants, VEGETATION CHARACTERISTIC PLANT SPECIES TYPE Key Species (75-100%) of sites) Usually Presenr (50-74% of sires) f Cussia nemophila Eucalyptus gracilis ® var. nemophila © Eucalyptus oleosa ¢ Eucalyptus dumosa* Dianella revaluia Geijera linearifalia ® Exocurpos aphyltus. Hestringia rigida * Pittosporum phylliraeoides Acacia colletioides Rhagodia preissti Eremophila glabra Melaleuea lanceolata Zygophyllum apiculaturm ul Eucalyptus gracilis * Acacia colletioides * Stipa sp. © Eucalyptus oleosa ® Enchyluena totnentosa Eremophila glahra Geijera lineorifolia Melaleuca lancedlata Sclerolgena obliquicuspis a Eucalyptus gracilis © Eremophila seoparia *® Eucalyptus oleosa * Triodia irritans * Westringta rigida ¢ Maireana eriaclade Zygophyllum aurantiacum Rhagodia crassifolia TV Eucalyptus oleosu * Rhagodia crassifolia Melalexca pauperiflora @ V Eucalyptus oleosa © Maireana erioclada ® Enchylaena tomeritosa * Rhagodia crassifolia * Geijera linearifolia Scleroleana diacantha Zygophyllum aurantiacum Sclerolaena obliquicuspis Threlkeldia diffusa Vi Fucalpptus brachycalyx * Melaleuca pauiperifiora « Maireana erioclada Enchvlaena tomentosa Rhegodia crassifolia Sclerolaena diacaniha Threlkeldia diffusa Zygophyllum aurantiacum Vil Eucalyptus yumbartana © Dianella revoluta Melaleuca eleuthrosiachya * Trivdia irritans * The Vegetation Associations Association f Eucalyptus dumosa-Westringia rigida The closest counterpart in the literature is the Melaleuca lanceolata association of Boomsma & Lewis (1972), which is said to occur with a number of species as co-dominants, including Eucalyptus dumosa and &. eracilis. Their classification inclhides a much wider range of vegetation associations than is here defined. It occupies a somewhat. irregular distribution throughout the study area, restricted to rising ground on shallaw calcrete, and the least sandy soils (sandy loams and Ioams). Association IT; Eucalyptus gracilis-Stipa This vegetation association could be described as a depauperate variation of Association I. Although some of the same species are present, Eucalyptus gracilis is the most important tall species uniling the group. The lowér diversity understorey Eremophila crassifolia Podolepis capillaris often includes a high percentage of perennial grass (Stipa), The relationship of Associations | and II is indicated by their proximity on the dendrogram (Fig. 4) and overlap on the ordination (Fig. 5). The association has not been previously recorded on western Eyre Peninsula, and &. gracilis has only ever been reported as a co-dominant with other mallee species for the whole statc (Boomsma & Lewis. 1980). lts distribution is also irregular, occurring on the dry central to northern section of the Cungena transect, but also scattered through the southern Pimbaacta and Wirrulla transects at higher rainfall, Very shallow, light soils on low lying ground are consistent with its occurrence. The resulting harsh moisture conditions probably reduce the number of understorey species present to. those which can tolerate them, such as the shallow-rnoted Stipa and Sclerolaena spp, VEGETATION OF NORTH-WEST EYRE PENINSULA 53 Association {i> Eucalyptus gracilis-oleosa Association UIT probably forms the core of the Eucabprs oleasa / £. gracilis association identified Ixy Crocker (1946) and Specht (1972), although Specht appears to misidentify £. ofeasa as &. Sociaifs. lt is the most common vegetation association in the northern part of the study area, but was not recorded by Margules & Nichatls (L987), Its ovcurrence in Lire southern section of the Nunjikompita transect at slightly higher rainfall seems to sugpest that its distritution is more restricied by soil than climatic factors. [t is not present in che south-western part of the study ares because the calcareous sands there favour other vegetation associations such as [VY and V1, or on the south-eastern section where calerete outcrops favour Associations | and JI. The occasional occurrence of Association JL) on dune flanks may be explained hy historical factors. [tis the most conimon inter-dune vegetation of the iorthern part of the study area, occurring on medium depth to shallow soils. Since the Molineaux sands have been deposited over only the lyst few thousand years, they are relatively unstable. In some instances sand from these dunes has drifted on to calcareous earths in the anterdunes already occupied iw Association 11 vegetation, and the long-lived species such as E. ofeos@ and & gracilis persist un a soil which would perhaps not at present favour their establishment. The understorey in these cases tends to include species less typical of the association, such as Triorlia irrifans, Association [¥> Lucalyptus cleosa-Melaleuca pauperiflora Association 1V contains a very species-poor and sparse understorey of chenopods such as Rhagedia crassifolia and Muireana erioclada, A known yegetation association including Eucalyptus aleosa, Rhegodia crassifolia, Zygophyllum aurantiacurn, Selerolevne diccantha and Melaleuca lanceolaio (Margules & Nicholls 1987), is probably equivalent to it, their Afelafeuca /anceoleta a possible misidentification of Af. pauperif{lora. Association LV occurs mainly in the southernwesiern part of the study area, on shelly calcareous sands, with scattered occurrences in the north, for cxample, on the Nunjikompita lransect, where Quilbers of this soil (ype occur on rises in the inler-dunes, These outlicrs are probably remnants of the old coastal dune environment which would have been present before the Molineaux sands werc deposited. Although the vegetation association appears to occur al slightly higher rainfall, this may simply be a result of the soil type which favours its occurrence being characteristic of the southern coastal higher rainfall areas. The low uiversity of tie association may be an indication that only a few understorey species are able lo persist an the coarse shelly sands, Association VO Eucalyptus oleosa-Enchylaena lomentoga Vegetation falling within Association V ts very variable and a large number of disturbed roadside siles were roost closely allied ro u, indicating rhat it may not bea naturally occurring association, but a result of disturbance. Mathy of the species are unpalatable, such as Geijere linearifolia, or tow growing chenopods, which suggests removal of more palatable understorey species by grazing. Further investigation would be required to ascertain if any of the sites represent @ naturally occurring Association V, Assuciation Vie Eucalyptus brachycalyx Vegetation Association V1 includes Eucalyptus brachyeniyx as the important upper-slorey species, together with a nange of low chenopods similar to that of Association V. The floristic similarity results in considerabje overlap between these two associations on the ordination (Pig. 5), [fis must common towards the coast, occurring in the southern central and western part of the study area on shallower soils than Association LV, Axsocivfien¥i; Eucalyptus yumbarrana-Triodia irtitans It is probable that kvcalppius yunebarrane was previousty recorded as Eucalyptus socials, and that the £. socutis/Melaieuca uncinata sandhill association of Specht (1972) corresponds to Association VII, as does Margules & Nicholls’ Community [af their Melalence adnvta is re- identified as M. eleuthernstachpa. Vegetation Association. VII is restricted to the sand-dunes of the northern, low rainfall part of the study area. The specles characterising it are entirely different from those found in the other vegetation associations, and include many species with distributions in higher rainfall parts of the state, such as Lomendra leucocephala and Guhriia lanigera (Jessop & Toelken 1986). This major Flonstic dasjunctinn corresponds to a similar observation in north-western Victoria, where the most marked discomtinuiy of mallee vegetation was between a Aucelypius incrossata/Albbertiad Lepitospermym association ‘with “southern temperate” affinities, and all the other associations, m which semi-succulent herbs, especially chenopods, were prominenr (Noy-Meir 1971). The disjunction observed is related to Le geology, since the heavier soils of the exposed Woorinen Formation present in the infer-dune corridors lypically support vegetation of Associations 4, 11 or I, The combined effect of coarse texture and 54 F, TIVER, A. D, SPARROW & R. T. LANGE TOPOGRAPHY RAINFALL SOIL DEPTH SOM TEXTURE r VEGETATION TYPE FREQUENCY PERCENT AGE VI Vil Fig.6. Histograms summarising observed environmental patterns for each of the vegetation types. For topography: P, plain; R, rise; F, dune flank; S, sandhill, For soil texture: L, loam; SL, sandy loam; LS, loamy sand; S, sand, increased depth of the sands of the dunes presumably creates a more favourable water relations environment, which allows the presence of a suite of species normally requiring higher rainfall, There is a higher percolation rate and deeper penetration of rainfall into sands than into clays, while evaporation dries the surface soil to a similar depth regardless of texture, so that at the end of along dry period, there would be more water available in the root zone of sandy soils than in clays (Walter & Stadelmann 1974). Clays also have a higher water holding capacity due to their smaller pore size (Ball 1986). This means that a larger amount of rainfall is required To bring air-dry clay to a range where water is available to plants than for sands. Discussion There is a clear vegetational discontinuity between the two predominating geological formations of north-western Eyre Penmsula, the siliceous Moornaba sand dunes and the caleareous Woorinen soils. Although it is possible to recognise plant associations within the vegetation of the Woorinen Formation and to relate their distributions to environmental features in a general way, sharp discontinuities do not exist. The strong gradation of associations is typical of the mallee on Woorinen Formation and its equivalents across the whole of S. Aust. (Sparrow unpublished data), There are two possible interpretations of these observations, Firstly, since there is no strong environmental discontinuity within the Woorinen Formation, the gradation may reflect combinations of species with independent environmental responses, which, further confounded by disturbance, results in almost continuous ecotones. Alternatively, the gradation of mallee associations could be caused by some degree of independence of the mallee and understorey strata, The mallees, VEGETATION OF NORTH-WEST EYRE PENINSULA 55 Fig. 7 Distribution maps of the seven major vegetation types. Shading represents uncleared vegetation and the broken line marks the Eyre Highway. 56 F. TIVER, A. D, SPARROW & R. T. LANGE TaBLe 3: Full species list (nomenclature following Jessop & Toelken 1986) showing occurrence in the seven major vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped in the final column. Introduced species are indicated with an asterisk. Species Vegetation Type i IL Ill IV Vv VI VII“ Other Acacia calamifolia + + + - - = + 7 Sweet ex Lindley Acacia colletioides + + + _ + + + os: Benth. Acacia hakeoides + - + = + + - ~ Cunn. ex Benth. Acacia ligulata + = = - _ = + + Cunn, ex Benth. Acacia merralli + + + + + + + - F. Muell, Acacia notabilis + = = = 2 _ _ . F. Muell. Acacia oswaldii + + + = + = = 7 F. Muell. Acacia rigens ~ = = - = _ 4 _ Cunn, ex Don Acacia sclerophylla . + + = + = _ = Lindley Acacia spinescens = - - - + = = = Benth. Alyxia buxifolia = = - _ + _ _ _ R. Br, Amyema melaleucae ~ — = pa ve “3 = + (Miq.) Tieghem Atriplex acutibractea + + + + 4 + + + R. Anderson Atriplex stipitata = + = e + 2 = = Benth. Atriplex vesicaria = + 4 + + A _ a Heward ex Benth. Beyeria lechenauttii + - = = = _ _ e (DC.) Baillon Beyeria opaca = - = + + _ rs = F. Muell. Billardiera cymosa = a _ + = 2s = _ F. Muell, Boronia coerulescens = = = = cs - = _ F. Muell. Callitris preissii 3 + = + = = - ssp. preissit Mig. Callitris preissi ~ = - - { - - n SSp. verrucosa (Cunn, ex Endl.) J. Garden Carpobrotus rossti - - = + + + > = (Haw.) Schwantes *Carrichtera annua - + - + + + = + {L.) DC. Cassia nemophila var coriacea + + a: 7 + _ “= & {Benth.) Symon Cassia nemophila var nemophila + + = + - _ 4 > Cunn. ex J. Vogel Cassia nemophila var platypoda = + = + fr = + + (R. Br.) Benth. Cassytha glabella + = = 4 J _ in! * R. Br. Cassytha melantha + = + + + + + _ R, Br. Casuarina cristata = = = L + = = + Mig. Chenopodium desertorum — = = = = = _ _ (J. Black) J. Black Cratystylis conocephala + a ; “ 4 , : P (F. Muell.) S. Moore VEGETATION OF NORTH-WEST EYRE PENINSULA 37 TABLE 3 cont. Full species list (nomenclature following Jessop & Toelken 1986) showing occurence in the seven major vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped in the final column. Introduced species are indicated with an asterisk. Species Vegetation Type I Il lt lV Vv val Vil Other Crypiandra leucophracta = rs -_ = _ _ + _ Schldl. Cryptandra tomentosa = = 7 = 2 s. + i. Lindley Danthonia spp. 4 + L. aS _ = 1 7 Lam, & DC, Daviesia asperula - = = = - - + _ Crisp Daviesia benthamii = : = 2 = - + = Meissner Dianella revoluta + + + + + + + ¥ R. Br. Dillwynia uncinata = = = = = = + _§ (Turez.) J, Black Dodonaea bursariifolia + - > ~ 2 5 + . F. Muell, Dodonaea stenozyga - 2 + * = = = + F. Muell, Dodonaea viscosa + = = = 7 5 rn = ssp. angustissima {DC.) J.G. West Enchylaena tomentosa + i + + + + + + R. Br. Eremophila crassifolia — - + + — _ + _ (F. Muell.} F. Mucll. Eremophila deserti + = is = + 23 = + (Cunn. ex Benth.) Chinn Eremophila glabra + + + + — + + _ (R. Br.) Ostenf. Eremophila scoparia + ef + + + + - 3 (R. Br) F. Muell. Eremoaphila weldii + “h + + i‘ + + _ F. Muell. Eucalyptus brachycalyx/concinna + + + + + + + + Blakely/Maiden & Blakely Eucalyptus dumosa + ' + 4s + + + + Cunn. ex Schauer Eucalyptus foecunda = = _ = _ - + _ Schauer Eucalyptus gracilis + + + ef + 4 < + F. Muell. Eucalyptus incrassate _ + = = = s + = Labill. Eucalyptus oleosa 4 + + + + + + 4 F. Muell. ex Mig. Eucalyptus socialis an — 4 = _ _ . _ F. Muell. ex Miq Eucalyptus striaticalyx ss + a _ - mn _ W. Fitzg. Eucalyptus yumbarrana s = + = - s + + Boomsma Eutaxia microphylla _ = = — ” = + - (R Br. in WT. Aiton) J. Black Execarpos aphyllus + 4 + + ' 4 + + R. Br. Exocarpos sparteus = = = - - - + = R. Br. Gehnia lanigera - - \ = _ = + _ (R. Br.) Benth. Geijera linearifolia + + + + + 4 + + (DC.) J. Black Goodenia varia — = = = = = 4 - R. Br. 58 F. TIVER, A. D. SPARROW & R, T. LANGE Tae_t 3 cont. Full species list (nomenclature following Jessop & Toelken 1986) showing occurence in the seven major vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped in the final column. Introduced species are indicated with an asterisk. Species Grammosolen truncatus (Ising) Haegi Grevillea huegelii Meissner Hakea francisiana Maconachie flalgania cyaneu Lindley Halgania lavandulacea Endl. Helichrysum bilobum Wakef, Hybanthus floribundus (Lindley) F, Muell. Txiolaena pluriseta Haegi Lasiopetalum behrii F. Muell. Lepidosperma laterale R. Br. Leptospermum coriaceum (F. Muell.) Cheel Leucopogon cordifolius Lindley Logania nuda F. Muell. Lomandra collina (R. Br.) Ewart Lomandra effusa (Lindley) Ewart Lomandra leucocephala (R. Br.) Ewart *Lycium ferocissimum Miers Maireana appressa Paul G, Wilson Matreana brevifolia (R. Br.) Paul G. Wilson Maireana enchylaenoides (F. Muell.) Paul G. Wilson Maireana erioclada {Benth.) Paul G, Wilson Maitreana pentairopis (Tate) Paul G. Wilson Maireana sedifolia (F. Muell.) Paul G, Wilson Maireana trichoptera (J, Black) Paul G. Wilson Melaleuca acuminata F. Muell. Meluleuca eleutherostachya F. Muell. Melaleuca lanceolata Otto Melaleuca pauperiflora . Meull. Microcybe muttiflora Turez. Muehlenbeckia gunnii (Hook. f.) Walp.) Myoporum platycarpum R. Br. Olearia brachyphylla (F. Muell. ex Sonder) Wakef. Vegetation Type af Il Ill IV Vv VI VIL VEGETATION OF NORTH-WEST EYRE PENINSULA 59 TABLE 3 cont. Full species list (nomenclature following Jessop & Toelken 1986) showing occurence in the seven major vegetation types. Occurrence in sites that could not be assigned to a major vegelation type are grouped in the final column, Introduced species are indicated with an asterisk, Species Olearia floribunda (Hook. f.) Benth. Olearia magniflora Vegetation Type I II Ill IV (F, Muell.) F. Muell, ex Benth.) Olearia muelleri (Sonder) Benth, Olearia pimetioides (DC.) Benth. Pimelea microcephala R. Br. Pittosporum phylliraeoides DC. Podolepis capillaris (Steetz) Diels Ptilotus obovatus (Gaudich.) F. Muell, Ptilotus spathulatus (R. Br.) Poiret Pultenaea elachista (F. Muell.) Crisp Rhagodia crussifolia R. Br. Rkagodia parabolica R. Br. Rhagodia preissii Mig. Rhagodia spinescens var. deltophylla F. Muell. *Salsola kali Santalum acuminatum (R. Br.) A. DC. Scaevola bursariifolia J. Black Scaevala spinescens R. Br. Schoenus subaphyllus Kiik Sclerolaena diacantha (Nees) Benth. Sclerolaena obliquicuspis (R. Anderson) Ulbr. Sclerolaena uniflora . Br. Sida corrugata Lindley “Solanum coactiliferum J. Black Stelligera endecaspinis A. J. Scott Stipa spp. I Templetonia battii F. Muell. Threlkeldia diffusa R. Br. Triodia irritans R. Br. Velleia connata F. Muell Vittadinia dissecta (Benth.) N. Burb. + + + + + + + + + + = - - = + + + + = + mJ - 4 a 4, ~ _- — — _ + + - = + = = 2 b = + + + + + - - - - + + + = + + + + = - + + - - + + + + + + + —- - - + + + + + - + + + a + + + + + + + = + 4 = _ + = = = it =. + + = + + + + + _ — — + + d + + + + - + - — VI VII Other + + = + = = + + - = 4 = + + + s 4 # “ + Pa + + + F + + - - + + + - = - + a + s + - + + - + - - + = + + - = + + + + = «= + ¢ + _ + = 60 J_ TIVER, A. D. SPARROW & RK. 1, LANGE TABLE 3 cout, Full species list (nomenclature following Jessop & Toclken 1986) showing oveurettce in the sever mgjor vegelatton types. Occurrence in sites that could not he assigned fo a mu/jor veperation type are grouped in the final coluinn, Intraduced species are indicated with an asterisk, Species Hestringia rigida = R ' . Br. Zygophylluny anienaphitum F. Muell, Zyguphvllum apiculatum ‘ F Muell. Zypophyllun) aurantiacuin/ovatunt 4 (Lindley) F. Muell./Ewart & ). White Zynophyllunt glaucum + F, Muell, ~ Verctation Type I i nt 1v V vi VIL Other ' ' ‘ + + - \ - - ' = - ca ~ + ‘ - _ + 4 + ‘ + = being very Jong-lived plants, may be indicating same historical influence, while the shorter-lived understorey species reflect current environmental conditions, Lange & Nicolson (1982) report mallees tecotding former extensions of a palacosol, and TWidale & Campbell (1985) and Short, ef al. (1986) provide ample evidence of dynamic changes in the surface geology of north-western Kyre Peninsula during the Holocene. A further issue is the occurrence of uncommon yepelalion associations in the area. One example is the Callitris preissii association represented by a single site on the dendrogram; another, towards the northern end ol the Mudamuckla transect and removed fram analysis due to disturbance, ~was unique lor the presence of Casuarina cristata. A regular sampling strategy inevitably Ieads 1a the omission of rare vegetation types, su there are undoubtedly other uncommon associations in the area, Acknowledgments We would like to rhank the staf? of the Vegetation Retention Branch of the S. Aust, Department of Environinent and Planning for practical assistance during the course of the study, The work was carried out as: part of the Middleback Ficld Centre programme for 1987, References Barr, DF (1986) Site and soils pp 215-284 Mn PB, D, Moore & $,B. Chapman, (Eds), “Methods in plant ecology” (Blackwell Scientific Publications, Oxford). Bri RN, 1, FAtrH. D. P. & Minchin, P. R. (1984) “Some wlgorithms contained in the Numerical Taxonomy Package NTP" (CSIRO Vivision of Water and Land Resources, Canberra). BLISseTT, A. H. (1969) STREAKY BAY preliminary map sheet, Geological Survey of Sourh Austratia, 1:250 000 series (Department of Mines and Energy, 5. Aust.). BoomsMaA. ©, D. & Lewis, N. B. (1980) “The native forest and woodland vegetarion of South Australia” (Woods and Vorests. Deparment, 5. Aust). BRAUN-BLANQUET, J. (1932) "Plant sociology” (MeCitaw- Hill, New York & London). Crocknr, R. 1, (1946) An introduction to the soils and vegetation ef Eyre Peninsula, South Australia. Trans. R. Sac. §. Aust. ™, 83-107. CiaucH, H, G. (1982) “Multivariate analysis in community ecology” (Cambridge University Press, Cambridge), GRrG-SMITH, P (1983) “Quantitative plant ecology" (Blackwell Scientific Publications, London). Jessop, J. P. & TOELKEN. Ef. R. (Eds) (1986) “Flora of South Australia” Parts I-(V (8. Ase. Gove. Printing Division, Adelaide), Lasou, R. T. & Nicorsos, K. PL (1982) Vegetation apparently recording former extensions of 4 paleosol. Trans. R, See, S, Aust. W6, 77-78. Licnynen L, & Lecenore, P. (1983) “Numerical eculogy" (Elsevier, Amsterdam). MARGULES, C.R.& Nicnonts, A. 0 (1987) Assessing rhe conservation value of reninant habitat “islands”: Malle patches on the Western Eyre Peninsula, South Australia. pp 89-102 In D. A. Saunders, G, W, Arnold, A. A. Burbridge & A, J, M. Hopkins (Rds) “Nature Conservation; The Role of Remnants of Native Venetation” (Surrey Beatty & Sons with CSIRO, Perth) Ninsson, C. (1986) Methods of selecting Jake shorelines as Mane reserves, Biol, Cons. 35, 269-241. NORTHCOTE, K. R. (1971) SA bacmal Key for the Recognition of Australian Soils" (CSIRO, South Australia), Nov Meir, [, (971) Mullivariale analysis of rhe sermi- wid vegetation of souwth-casteriy Australia: nodul ordination by component analysis Proc, Ecol, Soc, AUS. 6, 159-193, SHorT, A. DB. Dotvekinguan, DG. & BUCKLEY, K.-C, (1986) “Morphodynamic, Hazarel aud Development Impact Assessment - Eyre Coast Protection Districi” (Department of Environment & Planning, Adelaide), SPecnt, R. L. (1972) “The Vezetation of South Australia” (S. Ausr. Crovt, Printer, Adelaide). TwiPALe, C, RB, & CAMPBELL LC. M. (1985) The form of the land surface. Jn C. R. Twidale, M. J. Tyler & M. Duvies (Eds) “Natural History of Eyre Peninsula” (R, Soe. S. Ayst., Adelaide). WatTeR, H, & STADELMANN, E. (1974) A new approach ta the water relations af desert plants. pp 213-310 fn G. W. Brown (Ed.) “Desert Biology: Special Topies.on the Physical and Biological Aspects of Arid Regions” (Acadentic Press, New York). VEGETATION OF NORTH-WEST EYRE PENINSULA 61 Wess, D. A. (1954) “Is the classification of plant “Ordination of Plant Communities” (W. Junk, The communities either possible or desirable?” Botanisk Hague). Tidsskrift 51, 359-380. WRIGHT, M. J. (1985) Soils. pp 77-87 In C. R. Twidale, WuitTaker, R. H. & Gaucu, H. G. (1978) Evaluation of M. J. Tyler & M. Davies (Eds) “Natural History of Eyre ordination techniques. pp 277-336 Jn R. H. Whittaker Peninsula” (R. Soc. S. Aust., Adelaide). TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 113, PART 2 TIMING, EXTENT AND CHARACTER OF LATE, CAINOZOIC FAULTING ON THE EASTERN MARGIN OF THE MT LOFTY RANGES, SOUTH AUSTRALIA. BY R. P, BOURMAN* & J, M. LINDSAY} Summary Stream erosion on the eastern flank of the Mt Lofty Ranges has exposed a reverse fault near Cambrai on the Milendella escarpment, indicating a compressive component to the uplift of the ranges. Cambrian rocks have been thrust over Pleistocene fanglomerates and Miocene limestone has been dragged up along the fault zone to an elevation of 160 m asl. This fault may indicate the reactivation of a compressive Palaeozoic fault during the Cainozoic. Diagnostic foraminifera have been identified in samples of the limestone, which is of the Early Miocene Mannum Formation (about 20 Ma), adding support to the unpublished view that the position of the limestone on the escarpment is due to tectonic uplift of 60-90 m since the Miocene. KEY WORDS: Tertiary limestone, foraminifera, reverse faulting, Mt Lofty Ranges, South Australia. TIMING, EXTENT AND CHARACTER OF LATE, CAINOZOIC FAULTING ON THE EASTERN MARGIN OF THE MT LOFTY RANGES, SOUTH AUSTRALIA, by R. P, BouRMAN* & J, M, LINDSAYT Summary Bouwmas, R. P. & Lainpsay, J. M. (1989) Timing, extent and character of late Cainozow faulting on the ewslern wnarein of the Mt Lofty Ranyes, South Australia. Trans. RX. Soc. 8. Aust. 13 ), 63-67, 31 May, 1989. Stream erosion on the eastern flank of the Mt Lofty Ranges has exposed a reverse fault near Cambrai an the Milendella escarpment, indivating «compressive component to the uplift of the ranges. Cumbrian rocks have been thrusi over Pleistocene fanglomerates and Miocene limestone has been dragged. up along ure fault zome to an elevation of 160m asl, This full may indicate the reactivalion of a compressive Palaeozoic fault during the Cainozoic., Diagnostic foraminifera have been identified in samples af the limestone, which is of che Early Miocene Mannom Formation (about 20 Ma), adding support to the unpublished view chal the positica of the limestone on the escarpment is due to tectonic uplift of 60-90 m since the Miacen& Key Worp® Tertiary limestone, foramunifera, reverse faulting, Mi Lofty Ranges, South Australia, Introduction The fault origin of the ML Lolty Ranges hus long been discussed (Benson 1911; Fenner 1930; Sprise 1945) and the majorivy of these workers considered that the faults were normal types. Glaessner (1953) presented a model for the tectonic evolution of the Mc Lofty Ranges, which involved basement complex flexuring, that resulted in reverse faulting on the margins of the ranges and norma! faulting within them, Campana (1955) preferred to explain the origin of rhe M1 Lofty Ranges by compressional doming, which led to minor fault disruption on their margins, Gibson (1963) noted that the Clarendon-Ochre Cove Fault displays a reverse habit where intersected by tunnelling operations during the construction of the Clarendon-Happy Valley pipeline, Recently Wellman & Greenhalgh (1988) favourtd the view thal compressive forces have heen important in the formation of the Mt Lofty Ranges, A re-evaluation of exposed fault contacts is required to resolve the nature of the faulting responsible for the uplift of the Mr Lofty Ranges. The Milendella Fault There is clear evidence. of reverse lauliing on the eastern escarpment of the Mt Lofty Ranges as first reported by Mills (1965)2, and it may indicate the * Gibson. A. A. (1963) Final geological report on tye Happy Valley Tunnels Project. S. Aust. Dept of Mines report 37/92 (unpubl) 2 Mills, K. J. (0965) “The stnictural peredlogy of ihe Milendella area of South Australia’. Ph.D. Thesis, University of Adelaide (unpubl) * Depr of Soi) Science, Waite Insitute, University of Adelaide, Urrbrac, S. Aust, 5064, | Cieolowival Survey, Soil Austvatian Departinent of Mines & Bnergy, Box 151, Eastwood Px. §. Aust, $063, re-activation of Palaeozoic ihrusi faulis, Stream exposures of the Milendella Fault occur west of Cambrai on ithe escarpment (Angaston MR 390/630) at 160 m asl (Fig. 1). The fault here |s of reverse type with brecciated Cambrian schists of the Kanmantoo Group of metasedimentary rocks thrust over Pleistocene fanglomerates. Two pods of Tertiary limestone have been dragged up at a high angle in the fault zone, which dips to the west aL 45° (Fig. 2). The Pleistocene fanglomerates affected hy the faulting are similar to Pleistocene fanglomerates tilted into a vertical position by movemen| along the Willinua Fault as Sellicks Beach (May & Bourman 1984), The sediments al Sellicks Beach were related to the Ochre Cove Formation, which May & Bourman (1984) regarded as of Middle Pleistocene age. If rhe uwo deposits are correlatives then faulting on the Milendella Scarp probably accurred until the late Middle Picistocene or the early Late Pleisiocene. No unequivocal evidence of dislocation of Late Pleistocéne sediments in the area was noted. It is suggested that during faulting, the Pleistocene fanglomerates were folded, and breeciated basemenr rocks broke from the hanging wall to tuinble on top of the fanglomerates. Calcrete mantles the convoluted fanglomerates, dying out al the laulr zone, so that the culerete may have derived fram solution and reprecipilation of the Tertiary limestones dragged to the surface along the fault zone and draped over the top of the fanglomerates. Calcareous pisoliths in this calerere are lypically laminaled around votes of locally derived bedrock. This supports the interpretation of in sine pisolith formation from the redistribuuon of Tertiary limestone, Alternatively, if the calerete has stratigraphic significance (eg. the upper Middle Pleistocene Bakara calerete of Firman 1967) then 64 R. P. BOURMAN & J. M. LINDSAY {= \ : ; E = ts 5 a z 5 z = Ww = re Oo =; ie < [oa a; = fs) Ww = fa z 4 = oO Cambrai = 4, e @ N < a) Qa A I e \ Ss / ©) Walker Flat + & i fa a \ yu DELAIDE ‘\. o 6 a Ms 3 MANNUM Teale Fiat SY, vo fe) 35° § & Ay , Caloote > & © ; = ad ri Sy aS J < Sr Sh. & s Sy x ed WESTERN MARGIN OF, FS MURRAY BASIN Ke Lake Alexandrina > , = c « VICTOR -% x, yy" HARBOR 97 CE we SS Lake , 10 20 30 40 50 Albert KILOMETRES Fig. | Darnton map showing major faulp zones; | site of fault exposure near Cambrai and © Lepidoeyellna-bearing limestone in the Bremer valley. the relationships may indicate fault disruption of the calcrete in the Late Pleistocene. It is not possible to determine the amount of offset accomplished during this postulated Pleistocene phase of faulting without detailed drilling. However, Mills (1965)? shows a thickness of some 30 m of Pleistocene sediments in section at the scarp (Fig. 3), arid as the limestone must have been dragged up trom below this level, this suggests an offset of less than 30 m. Dislocation of Early Pleistocene marine sediments across the Willunga Fault was reported by May & Bourman (1984) illustrating similar post- Farly Pleistocene faulting of about 40 m on the opposite side of the ranges. It is interesting to nole that although there is clear evidence of geologically recent faulting on the eastern margin of the Mt Lofty Ranges, there is no pronounced fault scarp associated with it, which may suggest thal the scarp formed largely in brecciated rock and rapidly degraded. Mills (1965)* suggested that displacement on the Milendella Fault since “pre-Tertiary peneplanation” was approximately 335-366 m, and he attributed 250 m of this to the Early Tertiary to Miocene, and 60-90 m to post-Miocene faulting. He noted Miocene sediments (?Morgan Limestone) at elevations up to 14() m on the scarp, correlated them with Miocene strata under the Murray Surface and LATE CAINOZOIC FAULTING IN THE MOUNT LOFTY RANGES 6 wn Fig. 2. Fault exposure near Cambrai. View south along the eastern escarpment of the Mt Lofty Ranges. Cambrian schists (K) of the Kanmantoo Group of metasedimentary rocks have been thrust over (?) Middle Pleistocene fanglomerates (F). Light coloured calcrete (C) mantles the fanglomerates and two pods of Miocene limestone (L) occur in the fault zone (arrowed), which dips to the west at approximately 45° attributed the difference in elevation to tectonic dislocation. Twidale & Bourne (1975) did not deny recent fault activity but preferred to interpret the perched limestone as essentially due to Tertiary eustatic influences, with the limestone under the Murray Basin being the older, lower Mannum Formation and that on the scarp as the younger and higher Morgan Limestone. Thus they discounted the view that there had been 60-90 m of dislocation on the Milendella Fault since the Miocene and argued for an erosional origin for the lower part of the scarp. The resolution of this conflict may be achieved only by an accurate age determination of the Miocene limestones on the Milendella scarp to compare with those from the Murray Basin. Palaeontological work reported here demonstrates that the limestone on the scarp, at least in this locality, is from the lower part of the Mannum Formation. Age and facies of the Tertiary limestone in the fault zone Four samples have been examined from the two pods of Tertiary limestone dragged up in the fault zone. Portions were crushed gently in a pestle and mortar to disaggregate partially, boiled in a dilute solution of sodium bicarbonate with a drop of detergent, washed free of mud through a very fine sieve, dried, and picked for foraminifera under a binocular microscope. Slides with the foraminiferal microfaunas are stored in the Biostratigraphy Branch collections of the S. Aust. Department of Mines and Energy. Lithology All four samples comprise quartzose fossiliferous sandy limestone (biogenic calcarenite), The rocks are cream to brown, hard to friable, somewhat recrystallised and, in part, leached. Quartz and minor lithic sand content is considerable and poorly sorted, ranging from very fine grained to very coarse grained. Quartz and lithic grains up to 10 mm are present. There are occasional pale green glauconite infillings, somewhat oxidised. Barnacle plates are frequent and notable; echinoid, molluscan and bryozoal fragments are frequent to common; foraminifera are common but mostly recrystallised; ostracods, Ditrupa tubes, fish and decapod fragments, and algal oncolites are rare. 66 R. P. BOURMAN & J. M. LINDSAY EVANS TRIG. age <- 397m 300 METAMORPHIC ROCKS Height in metres Pleistocene alluvium 100 . . Miocene limestone 200 Displaced Miocene limestone blocks Recent alluvium ee Eocene lignitic sands SEA LEVEL Kilometres Fig. 3. Cross section through the eastern margin of the Mt Lofty Ranges to the Murray Basin near Cambrai (after Mills 19652). Discussion of foraminifera, age and correlation The four samples have much in common and the microfaunas will be considered together (Table 1). Numerically, all the microfaunas are dominated by Elphidium chapmani and Notorotalia cf. miocenica, but these are not the most biostratigraphically significant. The overlap of the top of the range of Sherbornina atkinsoni and the bottom of the range of Operculina victoriensis in S. Aust. indicates an early Longfordian, Early Miocene age (Wade 1964; Lindsay 1985). For the Murray Basin proper, Ludbrook (1961) only overlapped these ranges in lower Mannum Formation, and this has been borne out by subsequent experience. Other foraminifera present in the four samples are generally consistent with Longfordian Stage and Early Miocene age. Notorotalia miocenica has been scarcely recorded previously from S. Aust., so the slightly earlier age here of a related form, compared with the range in Victoria (Carter 1964) is not regarded as incongruous. The solitary and small planktonic specimen identified as Globigerina sp. is non-diagnostic. TABLE 1, Foraminifera: Table of occurrences of selected species. i Foraminifera Samples F23/83 F24/83 F2/88 F3/88 Discorbis sp. from group D. balcombensis Chapman, Parr & Collins -D. cycloclypeus Howchin & Parr R R F F Ammonia sp. - Vv - R Pararotalia verriculata (Howchin & Parr) Vv Vv - Vv Sherbornina atkinsoni Chapman - - - Vv Elphidium chapmani Cushman ce, Cc A A E. crassatum Cushman F R Cc F E. parri Cushman = - - Vv Notorotalia cf. miocenica (Cushman) F F Cc A Operculina victoriensis (Chapman & Parr) Vv R Vv R? Globigerina sp. - - - Vv Amphistegina sp. _~ — R Cibicides vortex Dorreen - - Vv - Planolinderina plana (Heron-Allen & Earland) - - Vv - (Abundances: V = very rare; only one or two specimens recovered in thorough picking. R = rare; 3-5. F = frequent; 6-10. C = common; 11-25. A = abundant; more than 25). LATF CATNOZOIC FAULTING IN THE MOUNT LOFTY RANGES 67 The lithology of the saniples is consistent with, but not unique to, Mannum Formation as described by Ludbrook (1961) and observed subsequently, This Early Miocene Mannum Formation in the fault fone is distinctly older than the clevated, Lepidecyelina-bearing, Morgan limestone reported from the Bremer valley east of Kanmantoo (Lindsay 1986"); about 20 Ma compared with 16 Ma (Lindsay 1988), Environment af deposition The variety and nature of the fauna indicate marine conditions. ‘hree lines of evidence suggest & shallow-marine, littoral-nerific environment: the notable frequency of barnacle remains, the restricted Alphidium-rich foraminiferal facies almost lacking planktonic forms, and the quartzose and lithic clastic component of the limestone, with gravel-sived grains, This corroborates the sedimentological evidence of Mills (L965)*, who proposed a near-shore environment for the deposition of the limestone, Conclusion The evidence presented here demonstrates that most Of the elevational difference between the 3 Lindsay, J. M, (1986). New record and tectonic implications of a high level Lepivocyedinabearing linestorig, Bremer Valley, casrer) Mount Loft y Ranges. S.A. Dept of Mines and Energy cepore 86/66 (anpubl,), Muannum Formation limestones on (he escarpment. near Canibrai, and under the Murray Basin can be allributed to tectonic dislocation in the past 20 Mu as originally suggested by Mills (1965)2, Furthermore, it has also been demonstrated thal there has been at least 100 sm (and prabably much more) of tectoni¢ offsel of the early Middle Miocene / epidocyclina-bearing Morgan Limestone that occurs in the Bremer Valley (lindsay 1986)4 in the past 16 Ma, Consequently, evidence of considerable tectonic dislocation of Tertiary limestones on the eastern side of the Mr Lofty Ranges, separated by 50 km, together with faulted Pleistocene sediments provides strong support for pronounced tectonism since the Middle Miocene. Without denying the evidence for eustalic influences in the Tertiary (eg. Vail & Hardenbol 1979) iris clear thal post Middle Miocene faulting is far more significant in the evolution of the east Mt Lofty Ranges than suggested by Twidale & Bourne (1975), Furthermore, the clear exposure of a reverse fault on the eastern margin of the Mt Lofty Ranges supports the notion of a compressive component to the forces involved in the Tertiary uplift of the eastern Ranges Ackauwledgments This paper is publisted with permission of the Director-General of Mines and Energy, South Australia, References Besson, WN, (1911) Notes descriptive of a stereoursm at rene tale Lofty Ranges, Trans. R. Soc. S. Aust. 35, 103-111, CAMPANA, B. (1955) The structure of the Eastern South Austrahan Ranges; the Mount Lofty-Olury Ane. Gen. Soe. Aust, 2, 47-61. tArtrR, A, N. (1964) ‘Tertiary loraminilera trom Gippsland, Vicloria, and their stratizraphical significance. Mea. geol Surv. ier. 23. Penner, C, (3930) The major structural and physiographic features of South Australia. Trans. R. Soe. 8. Aust. 54, 1-36. Firman, J.B. (1967) Stratigraphy of Lare Cainozoic deposits in South Australia, 7bid, 91, 165-178, Gy) aessvpr, M, Fo (1953) Some problems of Tertiary geology in southern Australia, Jaun Proc. R. Sec, NSM RT, 31-45, Linpsay, MA (W985) Aspeers vot South Awatratian ‘Tertiary foraminiferal biostraligraphy, with emphasis on studies of Mussilina and Subbatina. Spee. Publ. S, Aust, Dept Mines and Energy, §, 187-232, LuprRook. 8. A. (1961) Stratigraphy of (he Murray Basin in South Australia. Bull, zeod Surv, S. Aust, 36, May, R, 1, & BouRMAn, R. P (1984) Coastal landslumping in Pleistocene sediments at Sellicks Beach, South Austratia, Trans KR. Soe. S dust. 108, 85-94 Spricic, KR, ©, (1945) Some aspects af the geomorphalaay of portion of the Mount Lofty Ranges. Jhid. 64, 277-302. TWibale, C. Re & Rovrnwe, FA, (1975) Cieomorphological eyolution of part of the eastern Mount Lofty Ranges, South Australia, shied. 99, 197-209, Van, PR, & Harpenren, J, (1979) Sea-level changes during the Tertiary. Oceanus, 22, 71-79. Want, M. (1964) Application of the lijeage concept ta biostratigraphic yoning based on planktonic foraminifera, Adieropyleontalagy WW, 273-290, Wertman, Po & GREENHALON, S. A. (1988) Flinders Mount Lolly Ranges, South Australia, theic uplift, crosion aud relationship te ccustal structure. Zraets, & Soc, S. elust. 192, W-19. INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF, SOUTH AUSTRALIA BY R. C. AINSLIE, D. A. JOHNSTON & E. W. OFFLER* Summary The fauna of the intertidal mudflats of northern Spencer Gulf was monitored between 1982 and 1986. Two main habitats were identified, the mid intertidal zone, and the seagrass fringing low intertidal zone. These habitats supported quantitatively differing faunas. There was no evidence of the species impoverishment reported elsewhere for the epizoic fauna of this region. Comparison of the characteristics of the fauna with that of another negative estuary, the Port River in Gulf St Vincent, has provided the basis of a monitoring programme to assess potential environmental stress imposed by power station development in northern Spencer Gulf. KEY WORDS: Intertidal habitats, benthic fauna, seagrass, Spencer Gulf. INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF, SOUTH AUSTRALIA by R.C AINSLIE. Dh A. JOHNSTON & EL W. OFFLER* Summary AtNsLip, RUC. JoHnstTon, DA, & OFFLeR, E, W, (1989) Intertidal communities of northern Spencer Goll, South Australia. Trans, R. Soc. S, Aust, 113( ), 69-83, 31 May, 1989. The fauna of the intertidal mudflats of northern Spencer Gulf-was moniored between 1982 and 1986. Two main hahitats were identified, the mid intertidal zone, and rhe seagrass fringing low intertidal zone: These habitats supported quantitatively differing faunas. There was no evidence of the species impoverishment reported elsewhere for the epiauic fuuna of thin region. Cormparisan of the characteristics of the fauna with that of another megarive estuary, the Port River in Gulf St Vincent, has provided the basis of a monitoring programme to assess potential environmental stress imposed by power station development in northero Spencer Gulf, Key Worps: Intertidal habitals, benthic fauna, seagrass, Spencer Gulf. Intraduction Northern Spencer Gulf can be defined as thar portion of Spencer Gulf, South Australia, north of the line from Ward Spit to Lowly Point, i.e. north of 33°S (Stefanson 1977). Tt is a unique, narrow body of water, projecting inland some 200km from the Southern Qcean coast (Fig. 0), The waters of northern Spencer Gulf are characterised by high summer salinities, up to 48 (Nones & Lennon 1986) and high summer temperatures with wide seasonal fluctuations, 11-25°C (Johnson 19811), Due to bigh evaporation, and low fresh water inflow to the northern Gulf, there 15 a longitudinal salinity gradient of about four in 20kin. The region has been described as a negative estuary, A limited number of studies has been carried out on the marine communities of the “middle Gulf” (SEA 19812; Ward & Young, 1982, §3) and concur- Johnson, J,E. (1981) Hydrological Data for Upper Spencer Gulf 1975=1978. Fisheries Res. Paper No. 3, 1-30, (Dept. fisheries, Adelaide), unpubl. Social and Ecological Assessment (J9R1) Vrati Environmental Impacr Statement for port and terminal facilities at Stony Point, South Australia. Prepared for SANTOS, unpubl. Kinhill (1987) Lipper Spencer Gulf Inlertidiat Survey, Final Repert, May 1987. unpubl. * Electricity Trust of South Australia (1977) Northero Power Station Enyironmental Impaet Statement. July: 1977, unpubl. Electricity Trust of South Australia (L985) Northern Power Station Environmental Impact Statement, Aupust 1985. Prepared by Kinhill Steartis, unpubl. 2 e Environmental Sciemes an Tnineering Branch, Electricity Trust of Sourh Ausiralia, PO. Box. fi, Eastwood &, Aust. S063 rently with this study, Kinhill (1987)9 have examined intertidal communities wt Ward Spit and Lowly Point. Shepherd (1983) provides the only published description of benthic communities of the narrow reaches of the northern Gulf. Shepherd found a marked reduction in species number in the epizaic ‘communities on Pinna bicolor in northern Spencer Gulf compared to that of the same habitat in Gulf St Vincent, raising the possibility of a general species impoverishment in the hyper-saline waters of northern Spencer Crulf, He emphasised rhe need for further detailed study of other “component parts” of the biological system to determine its capacity fo accept additional stresses of industrial wastes and discharges, Singe the mid 1950's Playford Power Station las been operating on the castern shore of the northern Gulf, discharging warmed cooling water to the Gulf. The discharge from Playford Power Station, operaling at it’s Maximum nominal capacity of 330MW, resulted in surface water temperatures ubout 6°C wbove ambient near the power stalion, with only occasional incursions of warmed, watet (about 3° above ambient) to nearby intertidal zones (ETSA 19774). In 1977 and 1985 rhe Electricity Trust of South Australia issued Environmental Impact Statements for three further power station units, with a total gencrating capacily of TSOMW, to be built in this area, This Uevelopment forms the Northern Power Station. Thermal discharges were predicted (a more than uouble {ETSA 19774, 19855), The effects of warm water discharge fram power stations on benthic fauna have been the subject at a Tiumber of reviews (Coutant & Talmage 1975; Talmage & Coulant 1980; Craven e/ af 1983; Langford 1983). Effects vary from site to sile asa aL A.C. ATNSLIE, DD A. JOHNSTON & E, Wo OFMLER function af the climatic, hydrological and biolugicul features (Crema & Paglial 1980}. Greatest effects are Ghserved jiear the outfall, ac the hotcest times of the year (Langford 1983) and include changes in the biological characteristics of the benthic tummunilies within the zone of influence of the warm water plume (Robinson e7 af 1983; Robinson 1985; ‘Thomas ef a/ (986), In particular, Barber & Spenver (1984) and Thomas ef af (1986) have described the decline in populations of particular bivalve mullusce species and the establishment of dense popwations of opportunistic worm species as characteristic of \hermal disturbance in soft substrates, The present study began in 1982, three years prior tu the commencement of operation of the new power station. Intermittent operation of the first 250M W unit commenced in the suntmer of [9RS, with SOMW operating by summer 1986. The aim of the study was lo investigate spatial and Lemporatl nallerns of the benthic communities in the negative estuary of nocthern Spencer Gulf. It was anticipated that detailed descriptions of these communities would identify whether or not there were any biplogical impacts from the existing power station development (and ¢arly operation of the new development), and establish bascline data for long Lerin assessment of the eflects of increased thermal discharge; This study concentrated on the intertidal and shallow sublittoral fringe, a5 the ouly other published South Ausiralian study on the impacts uf thermal discharge on the marine environment had deseribed well defined paticrns of change in the communities af this yone (Thomas ef a! 1986), The thfluéice on communihes of sediment Lype, degree of emersion, and presence of scavrass are investigated in this study. Stalion focniiens, elevatians, and sedintents. Near Port Augusta the narrow channel basin of northern Spencer Gull is bordered by shallow sublittoral seagrass meadows and extensive intertidal mudflats backed by stands of the grey nangeove, Avicennia marina var restniferd (Fig. 1), Seven transects were established along the eastern shore-of the gulf, bath adjacent to and mmote from, exisling and proposed power station developments (Fig. 1). Three stations were established at dilferent intertidal zones un each transect, Hach station hud an area of about 25m?, On each transect, station 1 was located 30m scaward from the mangrove fringe, station 3 was loyaled al the seaprass verge, (emergent at extreme low tide} and station 2 was located midway between | and 3. The locations and elevations af each station were confirmed relative to gn esmblished Porc Aneusta Power Station darum, using an BDM theodnlite. Sediment samples were collected from cach siation on all transects and analysed for pruin size according to the Weni(wurth scale (Folk 1968) with injnor modifications (Thomas ef af 1986). Graphic means, Mz, (Folk 1968) were determined for the samples. The Bray-Curtis classitication technique was used Lo determine homogeneity groups bases on the percentage distribution of particular sediment size classes. in each sample (Miedeke & Stephenson 1977; Thomas ef af 1986). Werer ane oir temperature. For the study period 1982-1986, a continuous record of Gulf water temperature was made at the cooling water intakes of Playford Power Station. A continuous three hourly record of ambient uit fremperature was mnade at rhe SA Weather Buredi’s Port Augusts weather station No, (19066. Fauna Each station was sampled using a hand operat: ed voret collecting 3 sediment sample of JAG cme, surface area 80 ern? 10a depth of 20 em. The sam- ples were taken during daylight hours. The pallern of species accumulaiion with repent- ed tandom coring within a station area was deter- mined. By subsample 20, about 90% of the spe- cies recorded in 40 subsamples had been found, and those species which occurred after subsample 20) comprised an extremely small number of the total number af individuals sampled. The decision was made to proceed using 20 subsamples per station, exceeuing the species-area requitements proposed by Cain & Castro (1959) and mecting the stringent propusaly of Wemberg (1978), Each Station was sampled twice yearly (winter and summer) From June 1982 10 Sune 1986. Samples were treated as desyribed in Thomas er .a/(1986). Faunal relation- ships (berween stations, transects, tidal zones, aid samipling mes) were interpreted using Bray-Curtis classificu(iun aller rool-reot transformation of the data, and dendrograms were constructed with group dverage sorting (Swariz 1978; Bield. ed al 1982), Species constancy, abundance rank scores, aid average rank scores were assigned us in Hitilstone (1976) for 160 spectes totalling 34,312 individuals. Species associations were detentined using Bray Curtis classification analysis an non-transformed data, considering the 40 top ranked species for the period 1982-1986. Results Seation elevations and sediments Tidal Puciuations relative to station zones result in dilferent periods of emergence for statins 1, 2 and 3 om all 1ransects. Considcring February 1984, Fig. 1. Sampling station locations. Northern S denoted. INTERT(DAL COMMUNITIES OF NORTHERN SPENCER GULF 3Guik ausStAacia SOUTHE AN OCEAN INSHORE 7] SEAGRASS FRINGE PA= Posidonis PS- Petigania stnuona AA—Ampnibalis antarctica Caaliing Water Ourret Channa PORT PATERSON pencer Gulf, South Australia. Inshore fringing seagrass species also 72 RB. C. AINSLIE, DA. JOHNSTON & E, W, OFFLER the station | zone is emergent for over 200 hours/mionth, station 2 for about 80 hours/month, and station 3 for about 5 hours/month, or less than LY of the time, In winter even shorter penods of emergence oveur at the low intertidal zone. Local meteorological canditions, such as wind velocity, also affect the degrce of exposure on low spring Tides, Thus stations ) and 2 are clearly mid interti- dal stations while station 3 is in the low intertidal zone. The mid intertidal stations ate devoid of seagrass cover. The seagrass species al stations 3 include bath Posidonia australis and Posidonia sinuosa, without clear cui zonution in the shallow water, Some Amphibalus antarctica is present at transect 7. The hierarchical classification of sediments from all stations gives two major groups ol stations, proups (a) and (b) Fig. 2. The sediments of statiuns forming group (a) have Mz values Which gencrally range from medium sand (1.50) to coarse sand (0,250). The exceptions are station 1.1 which is clas- sifted ax fine sand (Mz = 2.5}, and the subgroup formed by sediments of stations 6,1 and 6.2. These last two stations have Mz values between 0 ancl — 25 (ie. at the lower end of the very coarse - sand scale), Fhe second major group of sediments (b) have Mz values ranging from very coarse sand [—259) hough to granules (1.7), with stations 7.1 and 7.2 having the coarsest sediments recorded int the survey, Although sediments of station 7,3 have un Mr? value of 1,256 (medium sand), this station forms its own subgroup on the dendrogram because of the restricted distribution in grain size around the median, unlike all other samples which had mire OF less even ranges of grain sizes. Aur and water temperatures Air and water temperatures 1982, 1984 and I98S show that seasonal water temperatures follow air temperatures, Ambient average nonthhy water tem- peratures range from [1-12,5°C in winter to 24-28°C in summer (Fig. 3). here is no indication of change in the overall umbient cemperature from 1982-1985, as a result of intermittent cooling water discharge fram the existing pawer station development. Fauna A total of 160 species were recorded ; polychaetes were the most Numerous with 52 species, includ- inw several new records for S, Aust. (Table 1). Mol- iuses: with 50 species and crustaceans, 40 species, were next i atuindance, with 18species from var ious ofher taxa (lable L), = 19 “0 30 = eo « = 70 J vn 80 w 90 A i is io, i a a a ee ee a STeTiom b Fig. 2. Dewdfograincl northern Spencer Gull stations by sediment prades. Fig, 4 presents the station faunal homogeneity dendrograms for the first two seasonal surveys (winter 1982, summer 1983), and for the last two surveys (summer and winter 1986). [0 all cases, at about the 25-30% level the homogeneity dendro- grams consistently show two major collection clusters with high intra-group homogeneily (Pig. 4 (1-4), Groups A and B). Generally the faunas of stations Land 2 on all transects comprise Group Aon each dendrogram, and station 3 faunas lorm Group B. In two of the dendrograms presented (winter 1982 and simmer 1986) there are exceptions to this generalisation, but there is no seasonal or spatial paticrn to. the exceptions, and the major trend persists, There is no evidence of consistent subgrouping within the major observed associations {Group A and Group B, Fig. 4). This tendency for faunas of mid intertidal stations | and 2 to group more closely together than to those of station 3 has been consistent throughout the our year study peri- od and is also reflected in the the tidal zone analy- sis, (Fig. 5) for each survey 1982-1986. The species homogeneity dendrogram cleter mined using the 40 most common species.is.present- ed in Pig. 6. Accepting the arbitrary cut-off level of 25% similarity, this dendrogram shows two Major groups of species (Al and Bl, Fig. 6). Although all but two of the forty most common species occur over all stations, those which form Group Al always contnbute a greater proportion of the total number of individuals in the samples of stations | ard 2 and those of Group BI are al- ways more abundant in the station 3. faunas, Together these analyses (Figs. 4, 5 & 6) indicate, thar the two habitats represented by the mid inter lidal region (stations | and 2) and the low interti- INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF 73 TABLE 1. Species recorded throughout the study, New records for South Australia are indicated by an asterisk. eee ee en ee EE PEE a Exclusively Exclusively mid intertidal low intertidal POLYCHAETES Olganereis edmondsi (Hartman) Neanthes vaalii Kinberg Neanthes cricognatha (Ehlers) Ceratonereis transversa Hutchings & Turvey + Ceratonereis mirabilis Kinberg Perinereis nuntia (Grube) Perinereis amblyodonta (Schmarda) + Perinereis sp Nereis trianguluris Hutchings & Turvey Nereis sp 4 Nematonereis unicornis (Grube)* + Marphysa sp* Arabellidae 1* + Arabellidae 2* + Lysidice sp Eunice sp + Schistomeringas sp Lumbrinereis sp Nephtys austratiensis Fauchald Glycera americana Leidy 7 Phylo sp* + Scoloplos cylindrifer Ehlers Leitoscoloplos normalis Day* Leitoscoloplos sp Naineris grubei australis Hartman Phyllodocidae 1* + Phyllodocidae 2* + Phyllodocidae 3* + Cirriformia filigera (Grube) Cirratulidae sp Boccardia chilensis Blake & Kudenoy Harmothoe sp 14 Harmothoe sp 2* Harmoathoe sp 3* Lysilla apheles Hutchings* Liomia medusa (Savigny)* Eupolymnia nebulosa (Montague) Streblosoma sp* Terebellinae 1 Terebellinae 2 + Terebellinae 3 Terebellinae 4 Polycirrinae* +++ +++ A - species seperated on basis of anterior elytrae. Barantolla lepte Hutchings Notomastus sp + Paraonidae Pherusa sp ae Armandia sp Galeolaria caespitosa (Savigny) Scalibregma sp* Maldanidae* Maeelona sp* NEMERTEAN ++ Nemertean spp + SIPUNCULID Golfingia margaritacea adelaidensis Edmonds 74 R. C, AINSLIE, D. A. JOHNSTON & E. W, GFFLER BIVALVES Tellina deltoidalis Lamarck Laternula recta (Reeve) Katelysia peronii Lamarck Katelysia scalarina (Lamarck) Brachidontes erosus (Lamarck) Xenostrobus inconstans (Lamarck) Ostrea angasi Sowerby Pinna bicotor Gmelin Malleus meridianus Cotton Anomia trigonopsis Hutton Circe scripta Cotton Electroma georgiana (Quoy & Gaimard) Trichomya hirsuta Lamarck Chlamys bifrons Lamarck Musculus paulucciae Crosse Corbula flindersi Cotton Lucilinidae GASTROPODS Salinator fragilis (Lamarck) Nassarius burchardi (Philippa) Nassarius pauperatus (Lamarck) Monodonta constricta Lamarck Bembicium melanostomum (Gmelin) Batillaria estuarina (Tate) Batillaria diemenensis (Quoy & Gaimard) Bedeva paivae (Crosse) Cominella erburnea (Reeve) Cominella sp Austrotiotia densilineata (Tate) Diata lauta A. Adams Quibulla tenuissima Sowerby Lepsiella vinosa Lamarck Conus anemone Lamarck Cantharidus irisodontes (Quoy & Gaimard) Clanculus plebejus (Philippi) Clanculus weedingi (Cotton) Phasianella australis (Gmelin) Hydrabia sp Microcolus sp Sophismalepas nigrita (Sowerby) Pterynotus triformis (Reeve) Nudibranchiata Opisthobranchia Netomella sp Elegidion occiduus Cotton Asteracniea sp | Asteracmea sp 2 Stomatella auricula (Lamarck) CHITONS ischnochiton variegatus Adams & Angus Ischnochiton contractus Reeve Heterozona sp CEPHALOPODS Hapalochlaena maculosa Quoy & Gaimard Exclusively mid intertidal Exclusively low intertidal + ++ +444 +++ INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF Exclusively mid intertidal 75 Exclusively low intertidal DECAPODS Penaeus latisuicatus (Kishinonye) Processa gracilis (Baker) Leander serenus (Heller) Leander intermedius (Stimpson) Paguristes frontalis (M. Edwards) Paguristes brevirostris (Baker) Callianassa aequimana (Baker) Callianassa ceramica (Fulton & Grant) Crangon socialis (Heller) Pilumnus fissifrons (Stimpson) Philyra laevis (Bell) Cryptocnemus vincentianus Hale Polyonyx transversus (Haswell) Gomeza bicornis (Gray) Halicareinus ovatus (Stimpson) Litocheira bispinosa (Kinahan) Portunus pelagicus (Linnaeus) Helograpsus haswellianus (Whitelegge) Ceratoplax punctata (Baker) AMPHIPODS Maera mastersii (Haswell) Birubius panamunus Barnard & Drummond Elasmopus bampo Barnard Allorchestes compressa (Dana) Ceradocus serratus (Bate) Cymadusa sp Gammaridae | Gammaridae 2 Gammaridae 3 ISOPODS Cymodoce longicaudata (Baker) Zuzara venosa (Stebbing) Exosphaeroma sp Cirolana woodjonesi (Hale) Cymothoidae Anthuridae 1 Anthuridae 2 TANAIDS Apsuedes australis (Haswell) Paratanais ignotus (Chilton) Tanaidac CIRRIPEDS Batlanus amphitrite Darwin Elminius modestus Darwin CNIDARIANS Anthothoe albocincta (Stuckey) Cricophorus nutrix (Stuckey) PISCES Nesogobius hinshyi (McCulloch & Ogilby) Gymnapistes marmoratus (Cuvier & Valenciennes) Plotosidae ++ + +4444 76 R_. AINSLIE, D, A. JOHNSTON & GB. W. OFFLER ECHINODERMS Leplosynapta dolabrifera (Stimpson) Taeniogyrus roehucki (Joshua) Amblypreustes sp Trachodotu sp Ophiuroidea | Ophiuroidea 2 Dendrochirotida Evhinoidea ASCIDIANS Ascidia sp DIPTERA Dolichopodidue Exclusively mid intertidal Exclusively low intertidal dal fringe (stations 3) can be considered to have quantitatively differing faunas. fig. 7 compares mean nuniber of species/m?, and mean numbers of individuals/m? for these two hahitats over the period 1982-1986, Significanily more spectes/m? are found in the low intertidal sta- ions. Reference to the species list for all surveys reveals that of the 160 species collected, 86 (54%) oceur over the whole tidal range. This 55% includes 37 of the 40 most common species, Fifty-nine spe- cies, or 38% of the total, occur exclusively in the low intertidal zone, and only 8% occur exclusively in the mid intertidal zone occupied by stations 1 and 2 (Table 1), In contrast to species abundance, the communities of mid intertidal stations | and 2 consistently have significantly higher mean num- bers of individuals (Fig. 7) than those of the scagrass fringing stations 3, Nine of the 10 most common species are More abundant at stations | and 2 (Table 2, Fig. 6); conversely, of the 61 spe- cies exclusive to the station 3 community, only two appewr in the “top 40” list (Table 2). Nevertheless the less common, but exclusively station 3 species, contribute to the consisient separation on the sta- tion and tidal zone faunal homogeneity dendro- grams (Figs. 4 and 5). Fig. 8 shows {he intersurvey time dendrograms comparing whole survey collections for each of the two communities for the period 1982-1986, Discussion Community-habiat relationships of the mid inter- tidal and low intertidal zones. In many studies scdiment characteristics have been shown to be a significant factor in determin- ing, benthic community patterns (Gray 1974; Saenger ez uf 1982" Jones e¢ af 1986). In intertidal regions particularly, there is the potential for wave action to grade sediments, resulting in coarser par- ticles on the upper shore line. Such a graded series of sediments is potentially reflected im the nature of the benthic communities (Gage 1974; Robinson 1985). On the. low energy shore of northern Spencer Gulf there is no sediment gradation, with random groupings of station sediments irrespective of the zones in Which the stations occur. There is a high overall similarity in the sediments, the two main groups discerned (medium to coarse sand, and coarse sand to granules) showing about 70% similarity on the homogeneity dendrogram (Fig, 2)- In common with other studies of the South Aus- tralian Gulfs where a limited range of sediments is found (Kinhill 19874; Thomas er af 1986) there is no evidence that this factor influences benthic community distributions in the northern Spencer Gulf, The mid intertidal zone is characterised by spe- vies which, although more common in this zone, oveur over the whole tidal range (Table 1), The most abundant species is the mussel, Bruchidontes erosus, Which forms dense mats in this zone. Such mussel mats are characterislic of many low energy sand and mud flat substrata in 8, Aust. (Womers- ley & Thomas 1976; Thomas ef a/ 1986), Other bivalves, although less abundant thai the mussels, are also characteristic of the mid intertidal zone. These include Je/ling deltoidalis, a species belong- ing to a genus known to have a behavioural response to light direction which contributes to tts zone maintenance (Newell 1979), Katelysia scala- rina, and Laternula recta. These bivalves contrib- ute to the abundance of many other species as they provide the substratum for the settlement of the barnacles Elminius modextus and Balanus amphi- INTERTIDAL. COMMUNITIES OF NORTHERN SPENCER GULF 77 PORT AUGUST& POWER STATIONS AIR-SESWATER TEMPERATURE COMPARISONS welee Temri me rose MEeSURED AT BLaT FORO Baste Cur pce? v 36 w 25 2 = ~ 29 < i f& wood # s = WoMFHLe AW, OST w 19 ~~" wake Thue, 7 montatr ay, 5 TT waren Teme wom Fee WAN APR dey UR 4uL AUG Bte SCH Woy Dec a o E ro —_ a = we a 2 w - eS FER be BPR mk) Gu Jul 4uG sae pot Mey cer Fig. 3. Monthly averages of daily maximum air temperatures recorded at SA Weather Bureau Station No 019066, Playford Power Station, Pt Augusta (Fig 1), Monthly average ambient Gulf water temperature derived from vontinuous hourly records at the ciyculating water intakes, Playtord Power Station. trite, and the sea anemone Anthothoe albocincta, There also appears to be a close relationship be- tween the large bivalve species and the omnivorous worm species Nephiys australiensis, Neanthes vaalit, Olganereis edmondsi, and Nereis trian. gularis, The last species seems generally to be as- sociated with mussel beds (Hutchings & Murray 1984). Several species of scavenging and car- hivorous snails (Nassarius pauperatus, Bedeva pai- vat, Batillaria estuarina, and Batillaria diemenen- sis) also Occur in the community dominated by the mussel beds, as does the scavenging erab PAilyra faevis, characteristic of the mid intertidal zone on low energy coastlines of S, Aust, (Womersley & Thomas 1976). The herbivorous top shell, Monodonta consiricia is common among the dense mussel beds which provide a substrate for algal growth, particularly Enteromorpha spp. In the extreme weather conditions of the north- erm Spencer Gulf it is clear that a number of ubig- uitous intertidal species are restricted to the low in- tertidal zone. These include the polychaetes Nematonereis unicornis, Ceratonereis transyversa, Barantolla lepte, Cirriformia filipera and Leltoscoloplos normalis, for which a range of habitats has been documented (Hutchings & Mut- ray 1984) and the gastropods Nassarius burchardi, and Ausfrolioiia densilineaia, also found elsewhere over broad tidal ranges (Thomas er al 1986), Can- versely, there are a number of species which are generally characteristic of the subtidal zone. These include such species as the hammer oyster Malleus meridianus, the scallop Chlamys bifrons, the blue swimmer crab Portunus pelagicus, and several other species primarily, or only, found in associa- uon with seagrass (Electroma georgiana (Ludbrock 1984), Golfingia margaritacea adelaidensis (Ea- monds 1980) and Amblypneustes sp. (Shepherd & Sprigg 1976)). The major environmental factor which appears to separate the habitats described in this study is the “degree of exposure”. This is a function not only of the length of the periods of emersion (in a region of high summer air tempera- tures) but also the protection from environmental extremes provided by the presence of seagrass in the low intertidal zone, Comparisons with other areos, Are the intertidal communities of norchern Spencer Gulf impoverished as evidence may sug- yest for epizoic fauna on Pinna bicelor? There are inherent difficulties in comparing coramunities of various marine and estuarine areas. Generalisalions from one area to another are often of limited value beyond the broad observation that marine domi- nated sites generally have more specics than estua- rine sites (Collett et a/ 1984), Accepting this timi- tation it can nevertheless be seen that the total nurn- ber of species found in the present study (160) is within the range of specles number recorded in several studies of eastern Austratian “estuarine” regions (Rainer & Fitzhardinge 1981; Jones e7 a/ 1986), and is comparable 10 the number recorded in # recent study of the intertidal mudflat fauna of the Port River estuary, Gulf St Vincent (Tha- mas er a! 1986), About one and a half times as 78 % SIMILARITY % SIMILARITY % SIMILARITY % SIMILARITY R. C, AINSLIE, DB. A, JOHNSTON & E. W. OFFLER Fig. 4. Station faunal homogeneity dendrograms, (1) winter 1982, (2) summer 1989, (3) summer 1986, (4) winter 1986. many species are recorded in the present study as were recorded by Kinhill (1987)8 for intertidal mud- flats about 50 km further south in Spencer Gulf Where salinities are four to five lower (Nunes & Lennon 1986). Kinhill (1987)4 did nal sample at the seagrass fringe (D Evans pers. comm.). Disregarding the 38% of the total number of specics which are exclusive to the seagrass (ringe in the present study leaves a species number very similar to that found by Kinhill 987)". Therefore this study provides no evidence of species impoverishment in the intertidal zone, related to high salinity or other environmental factors characteristic of the area. Natural vartations in the intertidal communities of northern. Spencer Gulf For.a baseline study to be of use in post develop- ment monitoring an. understanding of natural var- iations in the species matrix is necessary, The faunal homogeneity dendrograms for both stations and tidal zones demonstrate the consistent relationship between the mid intertidal and tow intertidal com- munities over several seasons. Between 1982 and 1986 the common core species characterising these two habitats remained constant (Table 2). Perhaps surprisingly no seasonal pattern emerged from the intersurvey comparisons. The over-riding pattern INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF 79 which showed for both habitats was the trend for sequential subgrouping ol surveys, with survey fau- nas within each of the two habitats showing high similarity, grouping at the 60%-70% similarity ley- el in each case, Sequential grouping of survey fau- nas does not appear to be unusual. Saenger et al (1980) interpreted such a pattern as demonstrating progressive recolonisation following environmen- tal disturbance, Buchanan ef a/ (1974, 1978), however, observed similar sequential subgrouping of survey faunas from undisturbed marine environ- ments and interpreted this pattern as indicating a more or less continual change in the presence and abundance of occasional or rare species whilc the core species remain relatively constant; Given the overali similarity in survey faunas over time this seems the most likely explanation here. The slightly lower intersurvey similarities of the low intertidal communities, compared ro those of the mid inter- tidal communities, isa consequence of the fact that the former are characterised by more rare or occa- sional species. Monitoring Jor changes associated with environ- menial disturbance Changes in estuarine faunas in response to or- ganic or thermal disturbance have been well documented (Gray 1976; Parker 1980; Bamber & Spencer 1984; Thomas er a/ 1986). In particular, in the negative estuary of the Port River, Gulf St TABLE 2. Cornman species of the intertidal fauna-of Northern Spencer Gulf, determined by reference to constancy, and abundance rank scores (Hailstone 1976). Note that the top 40 species ranked in this table includes all 21 species® which would be selected on Field e7 ai’s (1982) arbitrary cutoff for dominant species, namely species which contribute 4% or more to the total papulation of any one survey, % ABUNDANCE % CONSTANCY RANK CODE SPECIES (Refer Fig 6) ia | Wi Oleanereis edmondsi >} wi Nephtvs australiensis °3 B7 Brachidoniés erasus 4 BI Tellina deltoidalis oF RIS Nassarius pauperalas *g, 6 C4 Paguristes frontalis 7 Bia Batillaria diemenensis °8 B4 Katelvsia scalurina 9g C7 Elminius modestus "10 W6 Scatoplas ¢ylindrifer IT w2 Neanthes vaalii 12 BY Monodonta constricta "12 C2 Callianassa ceraniica 14 B3 Salinator fragilis 14% w7 Cirriformia filigera °16 CR2 Balanus amphitrite °17* Wwi0 Baraniolla lepte F1RY CIR Apsuedes australis 19* W30 Nemeriean spp 20 Wil Nereis (riangitlaris 20° BS Nassarius burchardt °22" Wi15 Maldanidse 23 wo Marphysa sp 24 wig Terebellinae 1 24 W22 Lysilla apheles 226, cl4 Crangon sovialiy 27 co Philyra laevis 28 B2 Laternula recta 29 B26 Asleracniea sp 1 50) Bll Batilluria estuarina 3H Bi2 Bedeva paivae 32 X5 Anthothoe albacineta o33% W33 Eupolymnia nebulosa °33 W20: Streblosoma sp 35 BI4 Katelysia peronii aA* W38 Golfingia m udelaidensis °37* B35 Electroma georgiana 33” B20 “Austroliotia densilineata 3R* Wa Schistomeringes sp aut Wwi2 Leitoscoloplos normatis 10.2 73.5 7 95,1 36,8 65,9 3.0 73.0 22 67.6 23 62.2 39 47.0 2.0 56.7 4,3 33.0 1.7 49.2 hg 45.9 0,95 34.0) 0.8 53.5 3.7 23.8 1.95 44,0 46 20.5 0.9 28.1 0.9 23.8 0,35 34.6 0.55 24.3 0.5 28.1 OS 26.5 0,37 28.1 0.6 17.8 0.5 22.2 0.28 23.8 0,26 23.8 0.21 23.8 0.3 25 0.7 9.7 0,17 17.8 0.2 15.) 0.4 97 0.24 13.0 0.14 15.7 O14 13,5 0,2 10.8 0.5 7,6 0.12 15,1 ONS 10,8 * Most common species in the low intertidal zone Group BI Fig. 6. 80 R, C, AINSLIE, D. A. TOHNSTON & E. W. OFFLER WINTER SIMMER WINTER SUMMER WINTER SUMMER WINTER SUMMER WINTER Wu2 1989 1863 1964 1964 1985 1985 1966 1086 GQ Simi, aarry ate % LOW INTERTIDAL STAT INTERTIDAL Fig. 5, Faunal homogeneity dendrograms with respect to tidal zones for all surveys, 1982-1986. wi eo WZ a7 mt s2 we c7 B19 W22 812 xs 82 WIO Ca wih Ge cl4 w3e e20 as wit #4 wa cz we 0811 814 CS s26 wi? c32 W7 W30 W344 82 S35 W339 W20 wi? 10 30 40 50 60- % S!MILARITY 70 80 90 Al SPECIES Bt Fig. 6. Species homogeneity dendrogram determined on non transformed data for the 40 mast common species far the period 1982-1986 as determined by average rank scores assigned as in Hailstone (1976). INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF BL —— Species ——— Individuals 7oa 25 600 SY ly 20 i 500 1 it I th 4000 & a 15 { a E | bd o 3005 bas t a 10 I > a 1 Qa * \ 2002 a | = 9 2 6 z a 1004 = { = { 1 Stations 142 Station 3 Fig, 7. Comparison of numbers of species/m® and numbers of individuals/m? for mid intertidal stations 1 and 2, and for law intertidal stations 3's, Vincent, Thomas et a/ (1986) describe responses of intertidal communities to progressive increase of thermal discharge from Torrens Island Power Station. These include suppression or elimination of populations of bivalve mollusc and worm species characteristic of undisturbed regions, and increases in populations of opportunistic worm species. OF the 120 species recorded from the Port River estuary about 50 species (40%) also contribute to the 160 species of the present study, These include several bivalve and worm species which had a negative response to thermal discharges, and which played a significant part in defining thermal perturbation in the Port River, namely the bivalves Te/lina del- toidalis, Katelysia scalarina and K, peronu, and the worns Néphtys australiénsis, Neanthes: vaulii and Scoloplos cvlindrifer. All of these species are common in. the communities of the mudflats of northern Spencer Gulf, particularly in the mid intertidal zone (Table 2), The presence of these common indicator species provides the basis of a monitoring programme to detect potential thermal perturbation in northern Spencer Gulf. Any changes in abundance of these species in areas near thermal discharges will be reflected in recognisa- ble changes to the stable faunal homogeneity rela- (a) % SIMILARITY SURVEY % SIMILARITY SURVEY Fig. 8. Intersurvey faunal homogeneity dendrograms. (a) stations | and 2, (b) stations 3. Survey number 1 —winter 1982, 2—summer 1983, 3 — winter 1983, 4 - summer 1984, 5— winter 1984, 6 — summer 1985, 7 — winter 1985, # — summer 1986, 9 — winter 1986. tionships described in this study over the period 1982-1986. Such changes in faunal homogeneity dendrogram patierns have been clearly demonstrat- ed by Thomas ef a/ (1986) in studies of changes in the fauna of the Port River estuary with progres- sive increase in thermal discharge to that region. The cirratulid worm Cirriformia filigera is common in the low intertidal.zone (Tables 1-2), Although the life-history of this particular species is not well known, other members of this genus have been shown to have typical opportunistic characteristics, capable of rapid colonisation of disturbed regions 82 KOO. AINSLIE, DAL IGHNSTON AE W OFFLER (George 1964 a,b; Thomas e7 a/ L986}. ‘Nhe ar- cherypal opporiunistic worm species Capirella cupttata (Grassle & Grasse 1974; Gray 1976; Tsut- sumi 1987) has not deen recorded fn the present study. (Cas, however, present in inferlidal commu- nities 30 kin further south (kimhill 19874), and it is nol unusual for opportunistic species in disturbed regions to be imnmugrants from other regions (Tho. mas ef «! 1986), As with changes in the indicator species discussed above, any changes in populations of opportunistic worm species near the thermal dis- charge can be expected to noticeably alrer che pat- terns Of the faunal homogeneity dendragrams, warning of potentially significant environmental perturbation, ‘To 1986 foe impact of the gecusion- | incursions of warmed water [rom Thomas Play- ford Power Siitiun to the intertidal zone, and the initial discharge from the Nerchern Power Station (from summer 1985) can be discerned from com- munity relationships described in the present study. Avknowledpments This work was funded and supported by the Electricity. Trust of South Australia as part of its Ongoing programme of covironmental research. We are grateful to those who identified organisms, espectally Jr P. Hutchings (polychaetes) and hdr Wy, Zeidler (amphipods) and to the South Australian Museum for access to their relerence collections, We also thank Dr M, Geddes for his suggestions and comments, Referéaces AMBER, R. N. & Senne, 1. & (1984) The benthos of a coastal power slafian thermal discharge canal. J Mar tiol, Ass. UK. 64, 603-623, HHCHANAN, JB d& Wanwiek Ro M. 11974) Att eothonate of benthic macrolaunal production in the offshore mud of the Northumberland coast. Ibid. 34, 197-222. ____ ,. SHFADFR ML, & Kinusion, BF (1978) Satioes of variability in the benthic macrotauna otf the South Narthumberland voust, 1971-1976, Mid. 191-209, Cain. Sy AL & Casinu, G. M, be O. (1959) “Manual of Vegetation Analysis” (Horper, New York) Courerr, L, C. Hoterncs, P. A, Giegs, Bob & Coins, A. 3. (19841 A camparative study of the macrphenrhic fauna bof Pardonur australis seagrass meadows Iq New. South Wales, 4qvutic Bar, 18, (11-134, Coun, ec. ©, & Taimace,S. S, (1978) Therma) effects, J Water Boll. Control Fed, 47(6), 1657-1711, CRAWeNS, J. BL HARRELSON, M. EB. & TALMAGE,S. 3. (1984) Therinal effects. (Aid. 55(6), 787-800. Chema, R.& PAGLIAL A.M, B, (LO8Q) ‘The structure of henthig coimunities ia anurea of thermal discharge from u coastal power slatiun. Mar Poller, Bull Vt, 221-224, Fowowns, 8. (1980) A revision of the systematics. of Australian sipunculuns (Sipunculay Rec. S. Aus Mas. 1Bi1), 1-74. Pip SoC) ARK, KOR. d& Warwick, ROM, (62) A practical salegy for analysing multispecies distribulian patterns, Mor Eeal. Pras, Sen ®, 37-452. Pork, Koo. CMON) “Perrology of Sedimentary Rocks” (Hemphill, texas), Civ 1. 1974) Shaflow-water Zonatien of Sea-Lach benthos und its relation to hydrographic and other physical fentures, J. Mar biol Ags, UK, 54, 223-249, Gporoe, J. D. (19648) The life history of the cirratulid warm Cireifanmia lentaculate on an intertidal mudflar. fbd, 47-65, (19646) Qu seme environmental factors affecting the distribution of Cirrifiemia tentaeulute (Polychacta) at Hamble tig 44, 373-388 Grass 6,7, F & Grassre d. P (1974) Opportunistic life histunes and genetic systems in marine henrhic polychaetes, A Afar. Res. 3242), 263-284, Grav i 3. d1974) Animat-sediment relationsiiyss, Ceeunography antl Marine Biol Ann, Rev, LQ. 223-261. _____ (1976) The fauna of the pollited River tes Estuary. Esenar. Coast. Man Seo 1, 203=223, ItaitsTone, T, S. (1976) Deliniitution of subtistal miacroberithas associations at the mouth of the Brisbane Rivet, dust fo Mae Fresh Kes. 27, 217-38, Hurcrians BoA. & MikRAY, A. (1984). Taxonomy of polychaetes from the Hawkesbury River and the sodthean estrarics at New Sourh Wales, Ausrralia, kev, Auyl. Mus, 36, (suppl, 35, T-118, Jones ALR. Watson Rossen, ©. 0, & Munkey Ay (194) Spatial patterns in the macrmbenthic communities ‘of the Hawkesbury Estuars, New South Wales. dust 4. Mar. Prestiy, Res. 37, S21-43. Lanotoku, T. E. 41989) “Elecivicity Generation and the Fealeay of Natural Waters” (Liverpool Liniversity Press, London) Lupakook, NOH, (1984) “Quacernary Moluses of South Austalia" Department of Mines ond Enerey Sourh Austutia Handbook No. 9 Du (Woulman Govt Printer, Adelaide). Muvorcsxn, J. Co & Sieriensen, W. (11977) Unvironmental impact and vaseline studies of the soft bolton marine enyironment un the vicinity Of ocewn sewer Guilally oir the NSW central coast. Third Aust. Conference of Coastal and Occan Engineering, 657-162 (UInsnrution ol Engingers. Austratia, Melbourne), Newrir, RoC. 0979) “Biology of Intertidal Animuls”. Grd ed.) (Faversham, U.K). Nuss RAL & Lisa. G. Wo (L986r Pliysical property distribunons and seasonal trends in Spencer Gulf, Sauil Anistralin: an inverse estuary. use Mar Fresh Rev. I7(\), 39-53. Panken, J Go (980) Elfects of pollution pon phe benthus of Delfast Lough, Mae Poll, Bull U1, 80-83 RAINUR, S.A FITHAROUNC, BR. oO. (1981) Benthic COMMITS bin esluaey with periocie deoxygenarion, Aast + Mor Freshw. Res. 32, 227-43, RKorinsay, K. 1. M. (1985) Temporal changes in the benthic macrofauny of Lake Macquaric, a New South Wales coasial lagoon. Werlands 5(2), 78-90, Yan Dew Veron FT. de Ginus, Pood (983) A survey al the estuarine beathic fauna of Homebnsh Hay, Parramatta River, NSM. (hid, 3(2), 81-84, SAT abe, B, STOMANSON, WL & Moverty, J. (980) The esluatine amacfabenthos of the Calhope River and aaa Crevk, Queensland, Adem Old, Mus, 20(1), 13-161, a YS) Maccrobori has of the coollng Water discharge canal af the Gladstone Pawer INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF 83 Station, Queensland. Aust, J Mar. Freshw. Res. 33, 1083-95. SHEPHERD, S. A., & Spricc, R. C. (1976) Substrate, sediments and subtidal ecology af Gulf St Vincent and Investigator Strait, pp. 161-174. In C.R. Twidale, M_J. Tyler and B.P. Webb (Eds), “Natural History of the Adelaide Region” (Royal Society of South Australia, Adelaide). ____ (1983) Benthic communities of upper Spencer Gulf, South Australia. Trans. R. Soc. S. Aust. 107(2) 69-85. STEFANSON, R. (1977) Spencer Gulf - a review of the oceanography, marine biology and the implications for development. Third Aust. Conference on Coastal and Ocean Engineering, 13-16. (Institution. of Engineers, Australia, Melbourne), Swartz, R. C. (1978) Techniques for Sampling and Analysing the Marine Macrobenthos. (Environmental Protection Agency. Corvallis Environmental Research Laboratory. US Department of Commerce, National Technical Information Service). TALMAGE, S. S. & CouTant, C. C. (1980) Thermal effects. Jour. Water Poll. Fed. 52(6) 1575-1615. THomas, I. M., Atnsie, R. C., JOHNSTON, D, A., OFFLER, E. W., & ZFD, P. A. (1986) The effects of cooling water discharge on the intertidal fauna in the Port River Estuary, South Australia. Trans. R. Soc. S. Aust. 110(4), 159-172. Tsutsumi, H. (1987) Population dynamics of Capitella capitata (Polychaeta; Capilellidae) in an organically polluted cove. Mar. Ecol. Prog. Ser. 36, 139-149. Waro, J. J, & Younc, R. C. (1982) Effects of sediments, trace metals and particle size on community structure and epibenthic seagrass fauna near a lead smelter, South Australia. /bid. 9, 137-146, ,& (1983) The depauperation of epifauna on Pinna bicolor near a lead smelter, Spencer Gulf, South Australia. Environ, Poll. (Ser, A) 30, 293-308, WEINBERG, S. (1978) The minimal area problem in invertebrate communities of Mediterranean rocky substrata. Mar, Biol. 49, 33-40. Womerstk&y, H. B, S. & THOMAS, 1. M_ (1976) Intertidal Ecology, pp. 175-185 In C. R. Twidale, M. J. Tyler, and B. P, Webb (Eds), “Natural History of the Adelaide Region”, (Royal Society of South Australia, Adelaide). ROTIFERA FROM AUSTRALIAN INLAND WATERS. HI. EUCHLANIDAE, MYTILINIDAE AND TRICHOTRIIDAE (ROTIFERA:MONOGONONTA) BY W. KOSTE* & R. J. SHIEL} Summary Diagnostic keys are given to the Australian representatives of the Rotifera: Monogononta in the families Euchlanidae (Manfredium, Diplois, Dipleuchlanis, Tripleuchlanis, Euchlanis), Mytilinidae (Mytilina, Lophocharis) and Trichotriidae (Wolga, Macrochaetus, Trichotria). All species known from Australian inland waters are described and figured, as are some widely distributed taxa not yet recorded from the continent. Distribution data and ecological information also are given. KEY WORDS: Rotifera, Euchlanidae, Mytilinidae, Trichotriidae, Australia, taxonomic revision. ROTIFERA FROM AUSTRALIAN INLAND WATERS. lil, EUCHLANIDAE, MYTLLINIDAE AND TRICHOTRIIDAE (ROTIFERA:MONDGONONTA) by W, Koste* & R. J. SHIELi Summary Kostr, W. & Sxies, R. J. (1989) Rotifera from Australian Inland Waters, If}. Euchlanidae, Mytilinidae and Trichotrijdae (Rotifera:Monogononta). Trans, R. Soc. Aust, 113,85 14, 31 May, 1989. Diagnostic keys are given to the Australian representatives of the Rotifera:Monogenonta in the families Euchlanidae. (Mun/redium, Diplois, Dipleuchlanis, Tripleuchlanis, Euchlanis), Mytilinidae (Mytilina, Lophocharis) and Trichotriidac (Wolga, Macrochaetus, Trickarria) &\| species known from Australian inland waters are described and figured, as are some widely distributed laas not yet recorded from the continent. Distribution dala and ecological information also are given. Key Worobs: Rotiféra, Euchlanidae, Mytilinidae, Trichotridas, Australia, laxonomic revision, Introduction The families of Rotifera:Monogononta considered in this part of our review of the Australian species (see Kosle & Shiel 1984b, 1987h) commonly are encountered in the shallow littoral zones of billabongs or vegetated lake margins {cf. Koste 1981, Tait et af. 1984, Koste & Shiel 19862, 1987a, Koste e¢ a/, 1988). Some species, ee. Euchtanis dilatata and related fortns, are known ta become planktonic in eutrophic waters during cyanobacterial btooms (Ruttner-Kolisko 1974). ‘I'his species Occurs seasonally in the lower R. Murry, S. Aust, (Shiel ef a/. 1982), Otherwise, species of the Euchlanidae (Manfredium, Diptuis, Dipleuchlanis, Tripleuchlanis, Euchilanis), Mytilinidae (Mysilina, Lophocharis) and Trichotriidae (Moleag, Macrochaetts, Trchotrin) generally are benthic or epiphytic in habit, and o¢eur in open water of lakes, reservoirs or rivers only as incursion species during seasonal flushing flaws, Mast of the 3S taxa recorded in the three families were collected as such incidental incursions into the limnetic region of billabongs. It is likely that more taxa remain to be described, particularly endemic species, because our initial sampling program was directed at the planktonic communiry and the sampling bias has “undersampled” the prelerred habitar of these Jittoral rotifers, Only four endemic species are known in the three families, one mytilinid (Lopheecharis curvata) and three trichotriids (Tricholria pseudocurta and T, buchneri from Tasmania, and Meerochaetus darineeli, from ihe N-T.), In contrast, about 25% of the taxa in the Epiphanidae and Brachionidae (Koste & Shiel * Rontgenstr. 1, 2903 Bad, Zwischenatin, Fed. Republic of Cermany tMurray-Darling Freshwater Research Centre, POL Bow 821, Albury, NSW, 2640 1987b), true limmetic species (and therefore more intensively sanipled), apparently are endemic, All known Australian representatives of the three families ave described and figured, and diagnostic keys are provided to enable identification to species. Widely-distributed taxa moc yet recorded from Australia, but which are likely to be found here, are included in some genera. The format follows thar of carlier papers; dichotomous keys are followed by individual species’ descriptions and known Australian records. Relative abundance from > 2000 collections is indicated by: “rare” (fewer than ten tecords, “uncommon” (10-30 localities) and “comnban" Gnore than 30 widely dispersed localities), Brief ecological data are included where available, generally in the sequence: temperature °C), pH, dissolved oxygen (DO, mg 1°‘), conductivity (Kya, pS om ') and turbidity {nephelometric turbidity units {NTU)). To minimise the citation of previous references to the rotifer families reviewed here, early Australian references which are included in Shiel & Koste (1979) are not repeated, nor are the majority of European references, which are included, for example, in the reviews of Hudson & Gosse (1886, 1889), Weber (1358), Voigt (1957), Bartos (1959), Rudescu (1960), Kutikova (£970) and Koste (1978), A more detailed treatment of the Rotifera outside Australia is given by these juthors, Family Buchlanidae Bartos The following genera Were integrated hy Remane (1929-1933) in the subfamily Brachionidae. A special family was erected by. Bartos (1959), later accepled by Kutikova (1970) and Koste (1/978). With the exception of Manfredium the genera are characterized by a lorica with plates which are connected with sulci, a segmented foot, more or less elongated toes, a corona of the same type (Euchlanis-type, Fig3d) and malieate trophi (Fig, 2:2b,3). Bh W. KOSTE & Ro J. SIIEL Key tu genera of the Family Kuchlanidae 1 Lariea thin, without lateral sulci, long toes -Maafrediumn Gallagher Lorica tigiel, With dorsal and sometimes yentral plates v2 ee ee ee W1), Lorica composed of three plates, dorsal divided, separated by sulci. . b ..Diplois Gosse Dorsal plave not divided. 2... 3 3(2).. Dorsal plate arched_ concave, narrower than arched Ventral plate......-- _Dipleuchlanis Reauchainp Dorsal plate arched, convex o7 triangular... .. 4 4(3). Dorsal and ventral plates nearly of same. size, conneeled by «lateral double longitudinal membrane within 4 stiff projection. ... Trpleuchlanis Myers With and without lateral longitudinal sulei vate alia ae Exchlanis Ehreribere Genus Mantredium Gallagher piciaeite Manfredi, 1927, p. 58 (non Walsingham. 1911). Boauchanipiella Remane, 1929, p, 107, Eudactylota Manfredi, 1927 is a junior hornonym of Exdactylota Walsingham, 1911 (Lepidoptera). Recuchampiella Remane, 1929 is a nomen audem under articles 12 and 16 of the International Code of Zoological Nomenclajure (name published without description, definition). Manfredium was Proposed by Gallagher (1957) to replace Eudactylora Manfredi, There is a single species, Man/reditern eudactplotunt. Manfredium eudactylotum (Gosse) FIG, | Sedridiun exdacivlotunt Gosse In Hudsan & Guese 1886, p. 74, Fig, 2h. Eudae -rylota ecigersiont (Gosse) Manlredi, L927, p. 58. Heauchamsiella euductylota (Gosse) Remane, (929, D, Marfrediuim eudactylotmm (Ciosse) alter Gallagher, 1957, p. 182. Djagnosis: Body pear-shaped with soft, transparent cuticle; head stall; foot with two longer segments; toes. long, rod-shaped; striated muscles visible in caudal part of body and foot: masrax rei- lobed, trophi malleate (Mig. 1:1); oesophagus. with small paired glands; large ted cerebral eve; resting egg with fine hairs (Fig. le. Lenguh; 420-760 um, four S0-110 ym, toes 275-320 pm, trophi 25 am, longest uncus tooth 12 wo, manubrium 18am. Oustribuzion: Cosmopolitan, panconitinental in Australia, rare. Warntstenotherm, 11,5-17,0%, pH 4.8-6.5, conductivity to 1600 «S crim. The rotifer may swim) betwee walter plants, also jumps by Vicking the toes. Literature: Koste 1978, 198): Ridder LYBI; Kose & Roberison 1983; Koste & Shiel 1987a Genus Diplo Gosse EXplois Gosse Jv Hudson & Gosse, 1886, Vol. 2, a. 86. Diplois daviesiae Gasse FIG, 2:1 Diplors davesiae Goxse In Hudson and Cinsse, (886, Vol, 2, j. 87. Pig. 24:3. D, phlerrava Vroso, 1910, p. 301, D. seulpturara Yaday, 1897, p. 135 (from New Guines). Diagnosis: Lorica with three plates; arched dorsal plate divided longitudinally by sulcus; lareral sulci small; foot Lhree-segmented; toes thin; lorica surface smooth, flexible; head shart, broad; eye with lens: oesophagus short; stomach compact; gastric lands large. Length: (6 500 pin, width 225 ym, toes 100 jum. Ostribation; May be cosmopolitan in the benthic of freshwater Sphagnum pools. Not yet recorded from the Neotropics. Not recorded in this srudy, bul there is an early record from Old (Thorpe 1887). Genus Diplenchlanis Beauchamp Diplenchlaniy Beauchamp, 1910, p, 122, Dorsal plate narrower than ventral plate; foot three-segemen(ed; tocs more or less long, rod- shaped with tapering points occasionally swollen terminally; rami points with minute comb (Fig, 2:2b); unci with 7-10-teeth; gastric glands long in younger individuals, broader and lobed in adult; red cerebral eye. Two variants of # single species 12 propatula) are known trom Australia; individuals can be discriminated by toe structure. DB. propatula has toes without a swelling belore the points, whereas, D. propaiula macrodactyla has toes with a swelling, Dipleuchlanis prapatula propatuta (Gaosse} FIG. 2:2a—f Diplois Ppropatula Goss /n Hudson & Gasse, ENMG, fi ig Dinlewctanis propaiuia (Gosse) alter Beauchamp. P Diagnosis; Caudal part of dorsal plate sometimes stretched to. rounded point znd reaches nearly te end of ventral plate; tees tapering to start paint. Length; 338-508 ~m, lorica length to 500 pm; width to 200; toes 70-110 pm. Distribution: Cosmapolitan in littoral, sometimes occurs in plankton.of ponds or billabongs, Rare, Vic, NT. Warm stenutherm, pH 4,5-6.4, also in O= poor waters on the suriace af mud. Literature: Koste 1978, (981. Dipleuchlanis propatula macredactyla (Hauer) FIG, 2:22 Dipleuchlarus. maervdacipla Hauer, 1963, p. 351, Dipleuchlanis Prange f, mucrodactyte ( Haner) alter Koste, 1978, Diagnosis: Bava part of dorsal lorica with shore paired cuticular projections; tocs long. Length; 124-244 pm, widil) 69-212 pm, toes 112-168 am, ROTIFERA FROM AUSTRALIAN INLAND WATERS f Fig 1: Manfredium eudactylotum (Gosse Scales lines, top (e) resting egg; (f) trophi; Wulfert (1940); e after Koste (1978). ‘0 pm (e); lower left LO wm (f-i). ): (a) lateral; (b) ventral; (c) dorsal; (d) lateral; Fig. la-d, f-i after (g) opened rami; (h) uncus: (i) fulerum. 100 pm (a-d); lower right 2 zi 88 W. KOSTE & R. J. SHIEL 3b 5a 5b 5c 4h Fig, 2: 1, Diplois daviesiae Gosse, dorsal. 2, Dipleuchlanis propatula propatula (Gosse): (a) dorsal; (b) trophi; (c) yentral, swimming; (d) toe; (e) unci; (f) cross-section; (g) D. propatula macrodactyla (Hauer), dorsal. 3, D. elegans (Wierzejski): (a) dorsal; (b) lorica cross-section (cl, D. propatula). 4, Tripleuchianis plicata plicata (Levander): (a) lateral; (b) traphi; (c) dorsal, swimming; (d) cross-section; (e) dorsal; (f) ventral; (¢) Uophi, apical; (h) lateral. 5, T. plicata razelni Rodewald: (a) lateral, (b) dorsal, swimming; (c) ventral. Fig. 2:1, after Weber (1898); 2a, b, f, 3a-d, alter Myers (1930); 2c, d, 3, after Fadcew (1924); 2g, 3e-g, after Hauet (1965); 5a—c, after Radewald-Rudeseu (1960). Scale lines Fig. 2:1-4, 100um (adult lorica each number group); 5,50 zm; 2b, 4b, g, 10 pm. ROTIFERA FROM AUSTRALIAN INLAND WATERS 89 Distritution: Panjropical, N7., Vie. Water ta 32°C, pH 5.4-65. ‘iferatures Myets 1930; Haver 1965; Koste 1974, 1981; Berzins 1982. Doubtiul or insufficiently described species: Dipleuchtanis elegans (Wierzejski, 1893) = Ewchlanis orapatula elegans {Fig. 2:3) according to Beauchamp, 1910, p. 122, Bipleuchlams canradi Evens, 1947, p, 179, Fie 5. Dipleuchtanis paludosa Haver, 1936, p. 139, Fiz. 1:12. Cienus Tripleuchlanis Myers Tripleucklants Myers, 1930 p. 379- Lorica ovoid in shape, truncate in front, smoothly-rounded caudally; dorsal and ventral plate hearly of same size, connected by pair of lateral longitudinal sulci. Longitudinal flange of stiffened cuticle extending for entire length between each, giving bellows-like appearance to cross-section. Foot three-jolated, guarded by shield-like process extending downwards from median longitudinal Nange; tocs short ending in abrupt paints; mastax of modified malleate type (Fig. 2:4h); six club- shaped teeth on each uncus; rami triangular with minute denticulate combs at inside of tip; double cerebral eye, A single species, 7. pifeate, with a doubtful variant (7 plicata f. rezelmé (Big. 2:5)) frou Romania distinguished on the basis of au elongated dorsal plate. caudally with a rounded notch (Rodewald 1940;88). Tripleuchlanis plicata (Levander) FIG, 2:4 Euchlanis plicata Levander, [RM p. 48, spe neural plicata (Levander) after Myers, 1930, p. Diagnosis: Lorica stout; foot glands and reservoirs elongated; retrocerebral organ small, Length: 250-270 pm, dorsal plare 90~I20 ~m, ventral plate 100-115 pm, toes 25-37 ym. Disiribution: Cosmopolitan; rare in fresh-, more in brackish and marine waters, also (11 warm springs. Qld, Vic literature: Hauer 1925, Koste 1978. Genus &ywohlanis Ehrenberg evenianis Blirenberg, 1832, p. 13). Loriva transparent with oval or ovate outline; dorsal plate arched, convex, sometimes with median keel or lateral wings, larger than ventral plate, variable in height and shape; ventral plate caudally somewhal indented, nearly flai. Two plates connected by thin, Flexible cuticle forming longitudinal sulci. Foot with two or three joints; toes long or short, sword-shaped oF parallel-sided, with sham tips, Paired long selae situated on dorso-distal portion of foot-segments (Fig. 3a:ss). Corona of family type (Fig. 3d); long sensory bristles and cilia in tufts atrd solitary on apical field, Trophi (Fig. 3e, !) moditied malleate; fulcrum (Fig, 3ffr} broad, pointed rami (ra) with and withoul minute comb; uncus with main and variable number of accessory teeth, Brain with large cerebral eye (Fig. 3b ey) behind large mastax, Large retrocerebral organ present (Fig, 3b:re), with subcerebral glands. Dorsal antenna large; lateral antenna (Fig. 3b:la) with sensillae tufts on small tubules. Stomach sometimes with sacculi but not constant between species. Excretory organ with large contractile bladder (Fig. 3a,b:b1), Vitellarium with eight large nuclei, Male thinly loricate (Fig 3i:m) with reduced alimentary tract; protonephridia present. Foot with twa to three joints; two long sensory bristles on terminal joint. Toes short. Dark resting eggs may be atlached to plants (Fig. 3h,j), éuchlanis species ave littoral rotifers living among aquatic plants, byt in the pelagial they occur in eutrophic lakes and ponds at Cyanophyta maxima, especially of Gloeutrichia, the main food of Evchlants dilatata (Ruttner-Kolisko 1974), Other foods consists of diatoms, desmids, other alyae and Getricus. The taxonomy of the cifferent species is difficult: even within the same population lorica shape and cross-section is variable: The shape of the anterior margin of both plates is of little value in jhe determination. Most useful is trophi structure, which is species-specific. Fucklanis should pot be pressed by a coverslip. See for cxample & meretr cross-sections and the different forms of £. diletera Key ta Species of the Genus Fuchlanis 1, Cross section of lorica more or less arched to 2 High keel... .- Teer wie wy wee a 2(1). Posterior edge of dorsal lorica with disiinet nolch 6 Venter membranous ar rudimentary ...,,-.-..5 413), Ventral plate ca. 44 dorsal plate width; wing-like bic} .€. alata Voronkay Ventral plate ca. 44 dorsal plate width, constricted AL posterior end; fiw expanded margins (Fig. 6:3) res -E. tera Hudson 5(3). Lateral constrictions in medial dorsal luriea; flanged Jateral margins (Pip, 8:3. ..£. pyrifarmis Gusse Dorszl forica not constricted, lateral margin not flanged (Figs 4:3, d:d).... .-£, deflexa Gasse 6l2), Cuticular shield-like process just below caudal part of dorsal plate (Fig. 7:1). --.. 8 mene Myers Shield-like process lacking... .....-2.... 2. 7 AG) Ventral plate rodimentary; lungitudinal sulci abserit; toes very Jong (>100 jan), slender (Fig, 4:25 vee ttiiineet vere eee. elpidia Myers Ventrml plate well developed... ocr 4 i ia eras 0 W. KOSTE & R. J. SHIEL Fig, 3: Auehianis dilatate dilatatq Ehrenberg: (a) ventral; (b) lateral; (c) cross-section; (d) apical field; (e) traphi; (1) tYophi, lateral; (z) &. triquetra, amictic egg; (h) mictic epg; (i) EB. dilalafa, mictic egg: (j) subitaneous egg) (k) snerm; (1) male; (m) £. deffeva, male. (Key to a,b: bl = bladder; dpl = dorsal plate of lorica; ey = cye; gg = gastric glands; la = Jaleral-antenna; iv = mouth; me = membrane; ov = ovary; re = retrocerchral sag; sl = stomach; 55 = sensorybristles; vi = vitcllarium; ypl = ventral plate of lorica. Key to f: ma = manubnium; fr = fulcrum; ta = ramus; u = uneus). Fig. 3a, ¢, j, after Koste (1978); b, after Beauchamp (1965); d, afrer Remane (1933); e, alter Hauer (1930); fafter Voigr (1957); g, h, afler Wesenberg-Lund (1929); i, alter Kavar (1914); k, after Ruttner-Kolisko (1972); 1, alter Wullert (1956); 1n, after Leissling (1924). Scales lines, top, a-d, g-m, SQum; ¢, f 10 nm. 8(7). Dorsal loriga with winglike lateral extensions toes dilated distally for 44 length and constrict to deflected ventrally al tips (Fig. 8!2)2. phryne Myers acute points (Fig, 8:1)......,,.8. orapha Gosse No lateral extensions of dorsal lorica.....----. 9 10(1)Mentral plate completely developed, lateral sulci 1 present (Pip. @:1)..... sow stete's etotaa E. tacisa Carlin 9(8). Foat and tocs slender; toes 4 length of dorsal plate, Ventral plate reduced to thin hyaline membrane; wide blade-like, fusiform (Fiz. SabjE. dilatata Ehrenberg dorsal plate flanges (Fig. 8:le-e..... E. trigitetra Foot and toes stout; toes. 14 length of dorsal plate Lhrenberg ROPIFERA FROM AUSTRALIAN INLAND WATERS ut Euchlanis alata Voronkov HG. 41 fuehlanis alerd Voronkoy, 1912, p, 210, Figs 2,3. Diegrasis: Shape of body ovoid: dorsal plate without terminal totch or embayment, but may have lateral wing-like extensions (see Fig, 4:tb, d)- Wingless individuals casily mistaken for & tyre or £. deflexa. Intermediate forms common, Ventral plate elongate, relatively marrow, but without constricuon before end as in E. fvre. Foot twoe segmented and robust; toes stout and fusiform. Tips of tami incurved with pair of finely denticulate combs; four stout teeth in each uncus with several accessory teeth. Length of dorsal plate 260-360 um: ventral plate width 102-156 xm; toes 70-98 ym; toes width 12-20 am, Distribution: Nearctic and Palearctic, in acid boreal waters, littoral and psammal, Not vet recorded from Australia. Liferarnre; Pejler 1962; Kaste 1978; Chengalarh & Koste 1983. Euchlanis calpidia Myers FIG, 4:2 Fuchlanis calpidia Myers, 1930, p. 371, Pl. 20, Figs La Diagnosis, Dorsal plate round, or highly domed (“obscure triradiate", ic. bhelmet-like) in cross section. Middle portions of lateral edges straight or pinched, extending downward below venter: ventral plate rudimentary {cl. #, defleva and E: porYormis), radiments connected wah dorsal plate by flexible membrane; lateral sulci absent; foot iwo- jointed, slender; toes very long, parallel-sided with abrupt point. Trophi characterized by five slender patred tecth in unci with Iwo ur three accessory teeth beside smallest, Tips of rami withour minute denticulate comb. Trophi similar to those of é& deflexa and &. pyriformis, E, calpidia distinguished by presence of distinct posterior netch at end of dorsal plate, and by relatively long toes. Length of dorsal plate 280-400 ym: lorica width 220-397 pmy.toes 12U-155 pm; toe width to 18 xm. Distribution: Nearctic Palearctic (Europe to bar Basi). Single record from a Goulburi R. billabong near Seymour, Vie. 22°C, pH 7,3, DO 8,0 mg 1-3, Literature: Hauer 1936; Wang 196{; Kutikova 1970): Kasie 1978. Evechlanis deflexa Gosse FIG, 4:3 Fuchlanis deflexa Gosse, 1851, p. 200. en deflexa (Gosselin Myers 1940, p. 3689, PL. 21, "Danidit = Euchlents afier Carlen, W354, p, 16) Dirgnosis: Rady resembles are of circle i) cross- section, dorsal plate oveld, without distince Posterior notch, sometimes wilh shallow emargination, Lateral edges of dorsal plate connected by flexible membrane somewhat thickened in position occupied by ventral plate in other Zuch/anis. Posterior portion of this arca more hardened as rudimentary ventral plate No longitudinal sulci present. Foot stow! and awo- jointed, long setae on dorsal end of first joint: trophi with five long teeth on each uncus, clubbed at tips, with 2—3 accessory tveth; rami drawn oul ta long tips, minute inside combs lacking (Fiz. 4:31). Length of dorsal plate 190-350 wm, width 140-240 wm, toes 55-100 um, toes ta 15 wm wide, subltanzous egg !90x89 pm. Disiribution: Cosmopolitan in (he littoral, ovcasionally in the pelagial. Rare; N.SW., Tas., Vic, W.A. There may be local yanants, eg. the Tasmanian form closely resembles & deflexa lore (Fig, 4:4), described from Lake Balkasch, Siberia (Kutikova 1959), [4.0-22PC, pH §,0-7,7, DO 8.0-8.6 me I-1, <70 wS cmb Citerature: Evans 1951; Koste 1978; Shicl & Koste 1979. Euchlaniy dilatata Ehrenberg FIGS 3, S:1 Evrhlenis iifatata Ehrenberg, 1832, p 131, Fig, 4:3. Diagnosis General body shape ovoid, rruncate anteriorly, rounded posteriorly; dorsal! plate variable in height and shape of cross-section, varying from low ate of circle to high triangle: ventral plate almost as large as dorsal; longitudinal sulci nor deep and narrow (Fig. 5;lc), Posterior dorsa! plate divided by deep elongate natch of variable form, ef. f. lucksiana (Fig, 5:4), f. unisetata (Fig, 5:3) or fF, farga (Fig, 5:5), Foot slender and two-jointed: paired setae on distal margin of first segment: toes mostly parallel-sided, tapering terminally to sharp tip, Unci of trophi with four opposing teeth: vetitral large teeth each have rudimentary touth; tips of rami with inner minute denticulate comb: retrocerebral sac large (Vig, 3b:re, cf. also Fig. S:4ik: brain with red eye on dorsal side Intraspecific forms are described exclusively according to differences in the shape of the lorica and cross sections, however research by Parise (1963) suggests that different morpliological types are correlated with ecological differences, This alsa may apply to some distinguished species; & parva Rousselet, tor example, resembles &. difatate and is distinguished from it by smaller size and long slender toes. All known measurements fall within the range of variation of &. dilutata; it is seen as un ecotype of £. dilatuta, Lorica leneth 140-320 ym, dorsal plate width 100-253 am, ventral plate 95-172 am, toes 50-100 am, posterior potch 12-61 win Weep; male 115-148 W. KOSTE & R, J. SHIEL ROTIFERA FROM AUSTRALIAN INLAND WATERS 93 4k Fig 5: 1, Euchianis dilatata dilaratad Ehrenberg: (a) dorsal; (b) toe; (c) cross-section. , E, dilatata macrura (Ehrenberg): ventral. 3, F: dilatata t, unisetata (Leydig): (a) dorsal; (b) cross-section; (c) toe, 4, E. dilatata f. lucksiana (Hauer): (a) cross-section; (b) dorsal; (c) ventral; (d) trophi, apical; (e) swimming; ({-g, k) cross-sections; (h, j) different lorica forms; (i) @. difatata, lateral, swimming. 5, £. dilatata larga Kutikova: (a) dorsal; (b) toe. Fig: lta—c, 2, 3a-c, 5a, b, after Kutikova (1970); 4a-d, j, k, afler Hauer (1930); 4e—1, after Myers (1930). Scale lines 100 zm. Fig. 4: 1, Euchlanis alata Voronkov: (a) without wings, ventral; (b) with wings, ventral; (¢) toes (d) ventral: (c) habit, lateral; ({) cross-section. 2, &. calpidia (Myers): (a) dursal, swimming: (b) trophi, apical; (c-f) crdss-sections; (2) lorica, ventral. 3, E. deflexa Gosse; (a) dorsal; (b) lorica, ventral; (c, 2, j), cross-sections; (d) trophi, apical; (e) tog, lateral; (Ff) other trophi; (h) dorsal; (i) resting egg; (k) lorica, broad form, ventral; (!) toe. 4, E. deflexa larga (Kutikova): (a) lorica, ventral; (b) toe. Fig. 4:1a, d-f, afier Yamamoto (1953); 1b, ¢, 3j, |, 4a, b, after Kutikova (1970); 2, Ja-e after Myers (1930); 3f-h, after Donner (1964), Scales lines 100um (adults lorica in each number group), ud W. KOSTE & R, J, SHIEL jem, toes 34-240 ym. [E, parva Rousselet, 1892; dorsal plate length 140 pm, width 100 «m, Yentral plate length 125 yin, width 75 zr, tes 70-yin, depth al posterior notch 50 pr). Pistriburion: Cosmopolitan in fresh and brackish waler! pH 4-10, Cl 140-4 1-1 (Koste 1978). The Most common and widely tolerant euchlanid in intaid waters; pancominental, including Tasmania, 8.0-25.9°C, pH 35.44-7,9, DO 3.6-13.8 mg I-|, 23-1600 4S cm-1, 2-235 NTU. “iteratures Myers 1930; Mauer 1935; Kutikova 1970; Ridder 1972; Kaste 197%: Shiel & Koste 1979 Koste & Shiel 1980. Evettants tnctse Carlin FIG. 6:1 kuchlenis tncisa Carlin, 1999, p. 17. Euchlanis inguetra Hudson & Gasse, 1389, PT. 23, Fig. 4 Diagnosis Body ovoid in shape, truncate in front with variable anterior notch; dorsal plate triradiate in cross-section with median keel extending from neck to angle of posterior y-shaped notch. Ventral plate completely developed, Longitudinal deep sulci conmect bath plates; foot rwo-yointed, setae on first foot joint; toes slender, fusiform; trophi (Fig, 6:le, hy) with five teeth on cach uncus, First and last (Fig. 6:1h) have accessory shorter tooth; minute combs on rami tips. Male (Fig. 6:11) with normal anatomy for genus (cf. Fig. 3:3), Length of dorsal plate 210-270 ym, ventral plate 180-240 pm, lorica width to 240 wm, toes 70-85 wm; male 145 pm, toes 26 pum. Listribation: Cosmopolitan in the littoral, occasionally in the pelagial of shallow ponds and billubongs Rare, N.T., Tas., Vie: 13.5-29.4°C, pH 5.3-7.4, DO §.45-8.2 mg 171, 23,0-80.2 pS cem-1, 2.0 NTU. Comment A variant, & triguetree mucronata (= & incisa mtucronate) (Vig. 6:2a, b} described by Ahlstrom (1934) from the Neotropics may be an ecotype; it resembles & ineiva and may co-occur. The dorsul plate has an elongated keel. Length of dorsal plate-to 320 am, loriva width ty 252 pm, height to 120 amy; toes lo 120 pn. Not yet recorded trem Australia. Literature; Mycrs 1930; Wulfert 1976; Kessie 1974, 197h, 1981; Shicl & Koste 1979; Berzins 1982. Buenlanis lyra Hudson FIG, 6;3a~j fMuchlarus lyre Hudson In Hudson & Gasse, 1886, ao Fig, 23:1 £ myersi Kutikova, 1959, ys, 224, Oieunosis: Body clongate and ovoid: cross-sevtion resembles are of circle; lateral sulci present; dorsal plate without posterior notch; ventral plate totally developed, somewhat constricted caudally; foot slender, two-jointed; paired long sctae on first Foor joint; ophi with five paired unequal tceth, with {wo or three accessories; rami tips with minute inside comb, Length of toes variable. Length of dorsal plate to 335 jm, ventral plate 302 um, lorica width 110-180 pm, toes 76-90 xm, toe width 14-16 pm. Distribution: Cosmopolitan in inundation areas, also in periphyton, Single record from L. Mulbwala, Vie. 12.0°C, pH 7.5, DO 10.7 nig I-71, 46 pS emn, 17.5 NTU, Variants ure known in che northern hemisphere, ez. 2 dyravryersi (Kutikova 1959; 223) (Fig. 6:4) a Palearctic subspecies with long slender toes (86-107 pani long, 7-8 pm wide) and Lf. lyr larga (Fig. 6:3k, 1) (Kutikova 1959) from N. Siberia. Variants aré Hol recorded from Australia. Literature; Myers 1930; Pejler 1962; Koste 1978; Shiel & Koste 1979. Fuchlanis meneta Mycrs FIGS 7:1,2 Euchlants menera Myers, 1930, p, 378, Pl19, Fizs 5-8 buchianls evophe Lucks, 1912, p. JOS, Fig. 31 Euchlanis proxima Myers, $930, p 377, Pl. 19, Figs bo. Diagnasis> Shape nearly ovoid; well-developed ventral plate almost as large as dorsal plate; cross- section seiiicircular arched bul alsi) roughly tritradiate; dorso-ventral width of Iateral sulet (relatively) uniformly wider than wm any other congener; cuticular shield-like process bencath posterior notch above first foot joint; posteriar notch of dorsal plale very deep and often large (Fig, Tk), foot rwo-joinled; toes very lang and slender, sometimes with minute claw at lps. Maye colourless, with short toes; dorsal and ventral plate still conspicuous; brain long with red eye; two Sensory hairs on Uny Lubules in apical lield, as in female. Trophi (Fig. Tle) with four main teeth, aecessorics not Vistble; minute denticulate combs on inside of rami tips (Figs Tle, 2d), Leneth of dorsal plate 10S-473-hm, width 80-140 yim, depth of posterior notel 35-45 am, width of ventral plale 50-90 pm, toes 60-75 pm, width of toos 3.5 xm, Male total Jengrhy 170 um, toes 25 um Distribucow: Cosmopolitan, mostly in aeidie walers; rare in submerged Spaagiuwm,; N, America, W. Germany, NSW. NLT, Vie WA. 10,0-29,99C pH 5,.3-7.4, DO $.45-10.1 mel 3, 20-523 aS cm | 22 NTU, alk, 2.6-2,7 mg L | Literature: Myers 1930; Hauer 1935 (yrraxinia); Wullert 1991; Pejler 1962; Berzins 1963, 1982; Koste 1978, 1981; Koste & Shicl 1980, Exenlanis oropka Gosse rig. Bl huchlanls oropid Liosse, TSR7, p. 3, Pig. 2:16, non Luchlanis vravhy Lucks, 1912, p. 10S, Fig. 31. ROTIFERA FRQM AUSTRALIAN INLAND WATERS 95 Diagnosis; Resembles EF. ditatata but usually smaller, with stouter foot and more robust, differently-shaped fusiform toes; lateral antennae suluated in cavities beside somewhat keeled end of dorsal plate; (rophi construction as &. dilatare (et. Figs 3e, 5:4d). Comb-like processes on inside of each ramus tip; four apposed club-shaped functional teeth, Length of dorsal plate 164-266 pm, width 127-172 xm, ventral plale length 123-225 wm, toes 49-78 pm, width 1-14 pm, posterior notch 24-29 Distribution, Cosmeopalian. NT, Qld, Vie. 8,0-18.5°C, pH 48-84, Dt) B4-10,0 me b-), 67-400 pS em |, 45-160 NTU. Luerature: Myers 1930, Kutikova 1959; Koste 1978; Shiel & Koste 1979; Berzins 1982, Euchlanis pocameneia Berzins FIG. 7:3 Euchlanis perameneie Werzins, 1973, p, 126, Figs. 4, 5, Diagnosis; Resembles E, meneta (see above). ‘Toes with pseudoctaws; unterier margin af dorsal plate with two short keels. Length of dorsal Jorica 95-104 pm; ventral! plate 70-72 am; width, dorsal 72-75 pm, ventral 50-55 ym; anterior margin width 35 pm, caudal opening 26-30 xm long x 18-25 pm wide; toes 45-50 p.m. Distribution: Recorded from New Zealand, not yet known trom Australia, Luchlanis phryne Myers FIG. 8:2 Euchlanis pkryne Myers, 1930, p. 372, Pl, 14, Fig. 4s Pl. 15, Pies |-4 Diagnosis: Dorsal lorica oval, with slight constriction of median lateral margin; deep inverted L-shaped caudal notch almost a$ long as toes: sciftened yentral plate connected to dorsal plate by longitudinal sulel; toes short, fusiform, ca, '4 length of dorsal plate; trophi with four club-shaped teeth on each ramus; rami with fan-shaped denticulate combs on inner distal margins. A superficial resemblance to E, pyrifermis and &. ealpidia was noted by Myers (1930), and Koste (1978) considered E. phryne 4 possible variant of £, calpidia or E. dilutate. Comparison of lorica morphology of the species above, particularly cros¢- sections, and differences in trophi structure, indigates that E, phryne is a distinct species. Length of dorsal plate 225 wim, ventral plate 210 am, dorsal width 180 pm, ventral plate width 150 pm, toes. 65 am, Distribution: North Americ# (Maine), single lueality, a billabong of the Goulburn R. near Sevmour, Vic, in spring 1976 and again in summer 1973, 17,0-22.0°C, pH 7,1-7.4, DO 8.7-9,8 me b 1, Comnrent: The above anomalous distribution is noteworthy; £. phryne is one of sevetal rotifer and microcrustacean species recorded from Goulburn River billabongs with similar disjunct distributions (Shicl 1976 and unpublished dara; Koste 1979). The importation of heavy dam-building machinery by the Utah Construction Company in the mid-1950"s (during the consituction of Eildon Dam) ts implicated a3 a possible source of transport uf resting eggs/ephippia. Populations have become established ity sheltered billabongs downstream of the dam. site, Euchlanis pyriformis Gosse HIG) 8:3 Evchlaris pyriformis Gasse, 1851, p. 201 Dapidia pyrrifermis Myers, 1930, p. 370, Fig. 15:5-7. Didenosix Shape nearly circular, may have shelit emaremations instead of caudal noteh; lareral margins turned downwards, occasionally pinched in the middle; ventral plate rudimentary, only developed posteriorly; no lateral sulci; foot twe- jointed, pwo long pairs of setaé on dorsal side of the first Foot segment; toes relatively short; trophi resembles that of &. deflexa (cf. Fig 4:3F). Length of dorsal plate 285-320 pm, width 275=<315 zm, tocs 80-85 pm, toe width 10-13 um. Distribuiion> Cosmopolitan in the littoral, occasionally pelagial. Vic, Tas. Literature: Kaste 1978; Berzins 1982; Stel & Kose LORS. Euchlanis triquetra Ehrenbere FiG. 9) Euchilanis triquetre Ehtenberp, 1833, 0, 461, Pig. $7:8. E, pellucida Harring, 1921, p. 6, Fis. 2, E, langubordica Manfredi, 1927, p, 24, Pin. 7h Panidia carinata Cartin-Nilssun, 1934) pW. 6, hig. 2. 0, lata Carlin, 1939, p. 14, aa eae iriquerta plerigaides Grese, 1455, nh &\, cigs non-F. (riquerr after Ruttier-Koliska, (972, p, 182, Fig. b. Diggnesis: Body in dorsal view circular, sometimes somewhat elongated (Fig, 9:1); triangular with a high keel in cross-section; median keel extends fron neck 10 caudal region; dorsal plate laterally with wide Flanges (Fig. 9:te-e); no ventral plate, na posterior notch, no lateral longitudinal sulci; some specimens With semicircular ventral line over base of toes; foot obscurely two-jainted; toes long, slender, nearly straight, ending in abrupt points most loricas very transparent with coloured inner organs (retrocerebral sac dark, mastax yellow, stomach olive green, yellow or brownish), Male (Fig. 33n0) with cuticular plates and dorsal keel, Trephi ef. Figs 9:lj—n) tps ef rami with minute verbs (Fig. S1j-k). %6 W, KOSTE & R. J. SUTEL ROTIFERA FROM AUSTRALIAN INLAND WATERS 97 Lorica length 300-710 am, width to 540 pam, toes 90-15 wm, lorica height! to 300 pm, trophi: rami 40 am, folerum 32 ym, marubria 30 pm. Disirihution: Cosmopolitan in the littoral, sametimes pelagic in shallow waters, NSW, Qid, NG, Vie. Eiteralure: Pejler 1962; Kosice 1978, 1981; Shick & Kaste 1979: Berzins |982, The following species are not considered here: £. arenoya Myers, 1936, known. only fram N. America and ?Volga; £. bioculita Issel, 190} (nosmen nude: name published without description, and apparently tater synonymized with &. plicata Levander by Issel (1906). Incorrectly cited in Voigt (1957) and Koste (1978), & cullysta Myers, 1930 (Fig. 9:4) is possibly a juvenile form of & triquetra, Only known from the Nearctic. Length of dorsal plate 170 xm, lorica width 80 pm, toes 40 pm. (Myers, 1930; Kosre, 1978); £. callimorpha Berzins, 1957 (Fig. 9:2) resembles, £, fncisa, only known fronv Gambia, W, Africas £. contorta Wolter, 1939. known only from central Germany and the Volga estary; £ dapidula Parise, 1966, see Koste 1978 (misquared as Parise, 1963). Doubtful species; E_ hyphidactyla Pavise, 1963, see Koste (9738, Doubtful species. £. Hieulata Kutikova & Vassileva, 1982, endemic in Lake Baikal; &. mamorokuensis Berzins, 1973 (Fig. 9:3), recorded from Madagascar; E. mikropous, Koch-Althaus, 1962 ceniral Germany; £, pannonica Bartsch, 1877, known only from Hungary, single record; £&. perpusilla Ridder Caribbean; E turfoysa (Rodewald, Romania. 5 recorded only fron 1977, known only from the 1940), knawn only from Family Mytilinidae Bartos Loricate rotifers; cross-sections of lorica mostly triangular or nearly rhombic; ventral plate and dorso-lateral plates firmly fused: long dorsum with or without sulcus, latter common with double keel; three or less foot sections; toes pointed, straight or shehtly curved ventralwards; malleate trophi, AU species littoral and benthic, occasionally (but rarely) in the plankton. Two genera (see Bartos 1959, Kutikova 1970, Koste 1978), Key jo penera of Mytilinidae 1. oriea thin or rigiet with dorsal sulcus and double keel; lofida Wnornamented; tocs fong Myltilina Bory de St Viticent Lorica without dorsal sulcus, one strong keel; lorica ornamented wilh dislinet pattern ard cavities; toes short J uphtcharis Ehrenberg Genus Afytitina Bory de St Vingent Mytilina Bory de Si Vincent, 1826, p. 87 ¢=Selpina Ehrenberg, 1831), p, 46 = Diplux Gosse, 1851, 9, 201 = Diplacidium Lauterborn, 1913, p, 483). Two recognized form-series (“Formenkreise"), One series strongly loricate, usually with variable stiff anterior and posterior spines; anterior margin of lorica generally granulated; toes stiff and sword. shaped, Second group thinly loricate without anterior spines (except ML acanthophera, (Fig. 10.1, 2) anterios matgin not granulate, toes mostly long and slightly curved, flexible. Key Species of the Genus Mytilina L. Lorica stout, anterior margin granulated, wes sword-shaped sy...) oc eee ee ee eee Lorica thin, anterior margin not Hrartulated, toes long and curved. q 21), Antetior lorica margin with two short dorsal and two short ventral spines, Mf. rnvcronata Muller Anterior lorica margin with only two ventral spines -.. M. ventralis (Ehrenberg) 31). Ventral margin af ‘lovica with variable projections M, acunthophora Hauer Ventral anterior margin without projections. ..4 AM), Toes with claws...., 202002. Toes without claws, ...-- re 6 3(4), Head part lpricate; 3 foot segments .M. crassipes (Lucks} ! foot scaments -M. ungripes (Lucks) A(4). Toes relatively shorl; ‘anterior fel in of lorica with Lf. CS eee eee WsWleata Lucks) Tocs long, straight, anterior Pack without falds (iesembles Miurcuathophara).M. trigona (Gass) phage Head part iNarivaie; 2 Mytilina ocanthephora Hauer FIGS 10:1,2 Moetifivg acunthophora Hauer, 1948, p, $50, Figs 73a-c. Diagnosis: Dorsal lorica strongly arched, granulated; ventral lociva fattened, with depression beneath head opening; triangular lorica projections flank medioventral head aperture (Fig, 101b); shield-like projection over two-segmented foot; toes Tong, pointed, Length: 195-253 jum (total); to ISO ym (lorica): height to 92 wm; width to 64 um; toes HO-94 yr, An unusually large specimen from Lago Grande, Amuzonia had the following measurements (sequence as above); 342, 198, 125, 137, 140-144 win (Koste unpubl). Distribution: Probably pantropical in polysaprobic waters. Not yel recorded from Africa, Occasiarially Pie. fi: 1, Euchlanss incisa Carttin: (a, 6) lorica cross-sections; (c) yeniral, (d) Tateral, swimming; fe} trophi, apical; (1) dorsal; (uw) toe; (fh) uncus; (i) male. 2. & incisa mucronata (Ahlstrom); {a) dorsal: (b) Jateral, 3, &. dene Hudson: (a) dorsat; (b) veniral; (c) trophi, apical; (d, c) cross-sections; (f) dorsal, swimming; (g, h) Jorica, ventral; (i) toe; (j) tae, (kK) E dvrw darga (Kutikova), vencral; (1) toe 4, E. Gira myersi (Kutikova): (a) yeniralt (bh) another form, ventral: (c) trapihi, Spical, (d)} cross-section; (¢) toe. Fig. 6: la-t, 3a, b, da-d, Myers (1930); if, uy Joel, de, after Kutikava (1970); 2a, b, , alter Koste (1974), Svales lines 100 am (acule jortea in each number group). 98 W. KOSTE & R. J. SHIES, Fig. 7; 1, Buchlanis meneta Myers: (a) lateral; (b) dorsal; (c) ventral; (d) lorica cross-section; (e) trophi apical; (ft) lorica ventral; (g) cross-section; (h) lateral, swimming; (i) lorica dorsal; (j} toe; (k) dorsal, swimming; (1) dorsal; (m) cross-section; (n) male; (0) male, lateral. 2, E. proxima Myers; (a) lateral, swimming; (b) cross-section; (c) ventral; (d} trophi, apical; (e) dorsal; (f) ventral: (g) dorsal. 3, & purameneta Berzins: (a) dorsal; (b) ventral. Fig. 7:la-e, 2e-g, after Hauer (1935); Lf, g, i, j, after Kutikava (1970); Ik, 0, after Wulfert (1960); 3a, b, after Berzins (1973); 2a-d after Myers (1930). Scale lines, top, 100 xm (Fig 7:1, 2), battam, 50 wm (3a, b). ROTIFERA FROM ALSTRALIAN INLAND WATERS 99 in plankton sansples. Qld (Townsville, Mr Isa). Literature: Koste 1978; Shiel & Kuste 1985, Murlling bisyleata (Lucks) FIG, 10:3 phx bisuleara \acks, 1912p. 95, Figs. 18a-c. M. trigune alter Harring, 1913 p. 75. Diagnesis: Lorica with folds on anterior margin; three weak dorsal keels in cross-section; sulcus indistinct, toes shorter than in M. tertguipes: with Jong tips (cf M. wnguipes (Fig, 1232). Length t30-180 jam, toes 60-70 pm, tips 8-14 wm. Distribution: In peat bogs, areas of decomposition. ACE, Vic. Literatures Hauer 1936; Koste 1978: Koste & Shiel 1980; Berzins 1982; Tait ev al 1984. Mynilina crassipes (Lucks) FIG. 10:4 Diplax crassipes Lucks, 1912, p, 96, Fig. 30, Mytilina orassipes after Carlin-Nilsson, 1934, p. IL. Piagnosis: Lorica laterally compressed; head with two lateral rounded plates; trophi small, uncus with six teeth; mouth opening with palpar organs; stomach and gastric glands large; two black spots at “forehead” region (eyes), Length 220-280 am (total); lorica 138 pum (oes 57-75 pln; claw 9 jam; (rophi 21-30 wm, Aistribulion: Cosmopoatiian in polysaprobic waters, occasionally in shallow-water plankton. N.T. (Magela Ck). literature; Kuste 1978, |98l; Koste & Shicl 1980; Tait er al, 1984. Mytitina mucronata (Muller) FIG. 11 Grachionus mucronatig Miiller, 1773, p, 194. Maitilina otucronata alter Holsten, 1908, p. 54. See Koste (1978) for extensive synonymy. Diagnosis; Shape of lorica variable. of Little taxonomic valuc (eg. MM. mucronata var. Spirigera (Ehrenberg, 1832)); corona as in Evchlanis; lorica granulated anteriorly: dorsally curved spines visible in lateral view beside foot opening; keels eud in short hook; foot three-segmented; large cerebral eye present (Pig, Wila eve lateral anrennae indisriner: dorsal antenna (Fig Lh:ta da} visible; trophi of malleate type, unci with five teeth; mastax (Fig. Ula mx) with salivary glands; stomach cellular with small gastric glands (Fig. Il:fa gy), Male approximately half female size, also loticate with cerebral eye (ct. (Fig. 3:1, m)). Length: 170-250 am, height 96-100 um, toes 33-60 pm, subitanedus egg 80 xm, male to 135 zm. Oistribution; Cosmopolitan in eutrophic waters, also on the surface of decomposition areas between witler plants and in brackish waters. Occasionally in the plankton of shallow ponds and billabongs. Vic, 13.5-22,0°C, pH 7.1-7.4, DO 44-102 mg [-1, 240 pS om-), § NTL. Literature: Evans 1951; Koste 1978, Myciling trizena (Gosse) FIG, {2:1 Diplax trigama Gosse, 1851, p. 201. Mytiling trigona after Harring, 1913, p, 75. Diagnosis: Surface of lorica slightly datted; antcrior margin with rounded edge ventrally (ef M. deanthophora Vig, 101, 2; cress section almost triangular; narrow double keel; toes with long tips. Tolal length: 245-245 an, lorica length 150-160 aM, Toes 68-83 pum. Distribution: Cosmopolitan in decomposi- tion/inundation areas, Qld. Literature: Koste 1978; Shiel & Koste 1979; Berzins 1982, Mytilina unguipes (Lucks) FIG. 12:2 Diplay wnguipes lacks, 112 p. 96. Movtiting bisuleara > unguipes (Lucks) after Carlin Nilsson, 1934, p. UL ae ie unguipes (Lucks) afler Wiszniewski, 1983, p, Diagnosis: Anterior margin of lorica without folds, laterally compressed; foot with two joints; toes long, slightly curved with distinct, inflexible claw, Doubtful species, possibly identical with MW. hisulcuta, Length 193 ym, height 94 wm, toes 58 xm, claw 1} pm, Distridition: European peat bog, nor yet recortled from Australiz. LEiferarure: Kutikava 1970; Koste 1978 Mytitina vertiralis (Ehrenberg) PIGS 1:3, 4,5 Salpinia verrratis Ehrenberg, 1832, p. (33, Pl.4, Fig. 7, yf ana veatralis (Ehrenberg) after Harring, 1913. p. "For extensive synonymy sce Harring oe. cit.) and Kunrikova (1970:519-520), Diagnosis: Lorica margin granulated; shape and size of lorica and spings very variable; only ventral anterior spines, Short-spined form (var, brevispina, 176-215 pm) shown in Tig, (:3a-c; long-spined form (var, macracantha, 250-350 um) shown in Fig, 11:4 Pig. 15 shows rare form described by Wulfert (1965) (M. ventrutiy f, longidactyla) with short anterior but long median-and shorter caudal venual hooks, Lorica (90-115 pm) without spines, median caudal spine 62 zm, ventral caudal spines 48 ani, tues 72 wm. These forms may be ecorypes, Distribution: Cosmopolitan. Probably pancontinentul, not yet recorded from S, Aust. Cf. typ. from N.SW., N.T., Qld, Tas., View brevispina from N.T., Qld, Vic; maecrecantha from NT, Gld, W. KOSTE & R, J, SHIEL ROTIFERW "ROM AUSTRALIAN INLAND WATERS tal W.A. 135,.5-28,5°C, pH $.44-7.9, DO 5.8-11.6 me 1-1, 23-1500 pS om>!, 4,8+67 NTU, alkalinity 27 me th Not yet recorded from Australia: M. bicarinata (Perty, 1850) M, macrocéra (Jennings, (894) M. mutica (Perty, 1849) MM. videns (Levander, 1894) faee Kosre 1978 for details), Genus Lophocharis Ehrenberg Lophockaris Ehrenberg 1838, p. 458. Strong lorica, surface soft, granulated, but may have honeycomb structure; cross section nearly rhomboid but specimens with flexible integument nearly oval or compressed; dorsum always arched; more or Jess distinct keel with and without transverse folds; ventral plate with longitudinal and transverse ribs; apical margin with and without servation; ventral aperture rounded, dorsal more triangular; foot-opening ventral; foot with three juris: toes with sharp points; trophi malleate (Fig. L3:Id); cerebral eyes only visible in living animals. Littoral and benthic in habi(, seldom found in the plinkion, Seven species are Known, four have been recorded from Australia, Key to Speeies of the Genus Lophocharis L Lonca >120 um long with distinct dorial heel producing, Wianaular cross-section. a" Lorica 100 pm lone, kel indistinct, crass- -Keehion more elliptical, . --.. 4) curvaia Berzins 20) Anterior margin ‘at lonce sironely serrated, dorsal keel with transverse folusL. selpina (1s hrenberw) Anterior margin smooth or lizhtly serrated, dorsal keel without falds.......-...--..-.--.-2-- 3 (2). Lorica sinooth, dorsal noteh of head aperture V- shaped, ventral a brouder L-shape; fine striae on both sides of dorsal keel..... “, euias Wulferi Lorica sculpiured as 1. sulpina; head aperture notches V-shaped: in striae beside keel... 2. oxpsiernan (Gossey Lophocharis curvata Berzins FIG, (4:3 Lophocharis curvata Berzins, 1&2, p. 12, Fig. '5a—c. Diagnosis: Small species, lorica clongated U-shape without typical dorsal keel, cross section more dorso-ventrally flattened than in congeners; head aperture margin with dorsal und ventral sinuses, ventral slightly broader; foot aperture broadest posteriorly; tacs thin, curved outwards. Lorica length 95 ym, width 55 pm, toes 18 pm, foot apetture width 32 xm, Distributions Endemic; single record from King Parrot Ck, Kinglake, Vic. 18.x,53, Comment: Although the original description and figures of this rotifer do not meet the requirements of the Code, we feel that the distinctive dorsum, foot.aperture and small size warrant retention of spectfic status until material can be examined. Lophocharis naias Wulfert FIG, t4 Laphochoris naias Wulfert, 1942, p, 188, Fig. tak. Diagnosis: Dorsal lorica with shallow keel and lateral longitudinal striations, surface morphology Otherwisesmooth, unornamented; apical margin of lorica finely serrated; uncus with 7 teeth) inner margin of rami with Fine denticles. Lorica length to 170 wm, widith to 105 wm, toes ta 21 um. Distribution; Europe, Single record, from Coongie Lakes, S. Aust.,, coll, Jane Roberts, Botany Dept, Univ. of Adelaide, 06.21.86. Literature: Koste 1978, Lephocharis oxysternan (Gasse) FIG, 13:1 Metopidia oxysternon Gosse, 1851, p, 201, Lophocharis oxysternon (Ciosse) after Harring, 1916, ). 564, Fig, 976-13, Diagnosis: Surface of integument soft or weakly pusiulated; ventral plate with deep depression over foot-opening. Many intermediate forms to L. salpina, e.g. with slightly serrated apical margin, Trophi (Fig. 13:1d-c) with six uncus teeth. Lonica length 120-200 pm, toes 24 pm. Distribution: Cosmopolitan, benthic in fresh and brackish waters, occasionally in plankton. Vic. Literature: Koste 1978, Berzins 1982. Lophochuaris salpina (Ehrenberg) FIG, 13:2 Lepadetla saliping Ehrenberg, 1834, p. 205 Metypidia salpina after Hudson & Gosse, 1889, p\ 46, Fig, 34:4; bs 7 el saline (Ehrenberg) afler Marring, 1916, For extensive synonymy see Koste (1978:153) Diagnosis: Axterior margin always serrated; keel with transverse folds mostly aver caudal part of loriva; variable, intermediate forms to /, oxysternon, traphi also with six uncus teeth: foor with three joints; tocs sharply pointed, ventrally curved. Length 175-224 pm, borica 120-135 wm; wiklth Fig 8: |, Euchlants aropha Gosse: {a) dorsal; (4b) dorsal; (c} cross-section; {d) alternate ane loricas; {e) Lroplii, apical, 2, £, phryne Myers; (a) dorsal; (b) ventral; (¢) trophi, apical; (d) crass-seerion, 3 »& pyelformis Gosre; (ay VeEriyal) (by) larica cross-sections (¢) rrophi. 4, &. ewtlyste Myers: (a) lateral; (b) eross-stctian: (c) trophi; (d) ventral Fig. S:la, after Kosiv (1978); led, after Donner (1964); le, 2a-d, 3a-c, 4a-d, after Myers (1930). Seale lines 100 um (Pig. 81-3); 50 pm (44, b, d), 102 W. KOSTE & R. J. SHIEL 3b 3c Fig. 9: 1, Euchlanis triquetra Ehrenberg, (a) lorica, ventral; (b) lateral; (c—c) different cross-sections; (f) dorsal, swimming; (g) toe; (h) male, contracted; (i) male, lateral, swimming; (j) trophi (fulcrum and rami); (k) ramus; (1) manubrium; (m) fulcrum; (n) uncus. 2, &. callimorpha Berzins, dorsal. 3, &. mamorokaensis Berzins: (a) dorsal; (b) cross-section; (c) ventral, 4, &. perpusilfa Ridder, ventral. Fig, 9-la, after Kutikova (1970); lb-e, after Grese (1955); Lf—n, after Wulfert (1956); 2, 3a—c, after Berzins (1973); 4, after Ridder (1977). Scale lines 100 zm (adult lorica in each number group). ROTIFERA FROM AUSTRALIAN INLAND WATERS 103 3d Af 4g 4h Fig. 10: 1, Mytilina acanthophora Hauer from Magela Ck, N.T.: (a) ventral; (b) lateral; (c) cross-section; (d) lateral (cf. 3a, b). 4, M. crassipes (Lucks): (a) lateral; (b) dorsal; (c) lateral; (d) dorsal; (e) lorica, contracted; (f) trophi; (g) trophi, lateral; (h) uncus; (i) mouth with palpar organs (m = mouth). Fig. 10:3, after Hauer (1936); 4, after Wulfert (1939). Scale lines [00 wm (adult lorica in cach number group). 104 W. KOSTE & R. J. SHIEL 3c 3b Fig, 11: 1, Mytiling mucronata (QO, F. Miller): (a) swimming, laleral [b/ = bladder, br = brain, de = dorsal antenna, eve = cerebral eye, f= foot, 2g = gastric gland, / = intestine, mx = mastax, st = stomach, ¢= toe, vi = vitellarium); (b) male, swimming, lateral; (c) lorica, lateral; (d) lorica, dorsal; 2, Myzilina sp. cross-section. 3, M. ventralis (Ehrenberg): (a) lorica, lateral; (b) ventral; (c) male, dorsal. 4, M4. ventralis macracantha (Gosse) lorica, dorsal. 5, M. ventralis longidactyla (Wulfert), lateral, from E. Africa. Fig. 11:la, 2, 3a, b, 4, after Koste (1978); 1b, c, after Feiler te jd, 3c, after Weber (1898); 5, after Wulfert (1965), Scale lines 100 um. Top Fig. 1l:lb-d; left, 3, 4; ottom, 5. ROTIFERA FROM AUSTRALIAN INLAND WATERS 105 Fig, 12: 1, Mytilina trigona (Gosse): (a) lateral; (b) ventral; (c) lorica, dorsal; (d) cross-section; (e) swimming, lateral; (f) ventral. 2..M. zguipes (Lucks): (a) lateral: (b) cross- section. Fig. 12:1, after Donner (1954); 2, after Hauer (1936). Scale lines 100 zm, 80-95 um, toes 25-40 pm. Distribution; Cosmopolitan, benthic in fresh and brackish waters, also in polysaprobic pools and ponds. Pancontinental, 10,5-30.4°C, pH 5.6-8.4, DO 3,7-11.0 mg 1-1, 28-3330 «S cm~-1, 5~92 NTU, alkalinity 4.0 mg 1 1 Literature: Evans 1951; Shiel & Koste 1979; Koste 1981; Berzins 1982. Not yet recorded from Austratia; Lopkocharis hutchinsoni Edmondson,, 1935 L, naias f. ambidenta De Ridder, 1960 L. parva Rudescu, 1960 L, rubens Wulfert, 1939 (see Koste 1978 for details). Family Trichotriidae Bartos 1959 Three genera with loricate head and body; surface with facets, mostly granulated and with spicules or spines, sec. on dorsum; foot freely movable or with stiff joints; trophi malleate. Generally occur between aquatic macrophytes and in periphyton. In plankton only as migrants. Vig. 13; 1, Lophocharis oxysternon (Gosse): (a) dorsal; (b) ventral; (v) lateral; (d) trophi, apical; (e) trophi, lateral. 2, L. sulpina (Ehrenberg), (a) swimming, lateral; (b) lorica, lateral: (c) lateral; (d) another lorica, lateral; (e) cross- section. 3, L. curvata Berzins, (a) lorica, dorsal; (b) cross section; (c) lorica, ventral. 4, L. naias Wulfert, (a) torica, dorsal, (b) lorica ventral. Fig, 13:1 a-e, 2b, after Harring (1916); 2a, aller Wulfert (1956); Id, 2c, after Haver (1925); 2d, after Klement (1960); 3 after Berzins (1982); Scale lines, centre, 100 pm (la—c, 2-4); lop right, 10 pm (d, e). 106 W KOSTF & R. 1. SHIFI Fiv. 14) Wolea spinifera (Western): (a) dorsal, swiinining; (b) lateral; (c) ventral; (d) cross-sectian a-c. aller Western (1894); d, after Vranovsky (1257), Seale line, $0 yarn, Key tu penera of Trichotriidae 1. Lorica with anal segment... ....-.. a , Lorica without anal segment, ,., Wo/gu Skorik 211), Lorica with distinct elongated spines an dorsum ya sonn fie toe 90rd Macrachaelus Perry peeesuy Erichelria Bory de St Vincem =p ib eee Genus Holga Skorikuv Wolgd Skorikay, 1903,, p. 37, Monospecific, Holem spinifera. Holga spinifera (Western) FIG. 14 Distvlu spinifera Western, [894, p. 427, Fig. 241-4. Cuthypna spinifera after Murray, 1913, p, 545-54. Lecane spinifera afier Harring, 1913, p. 62. Mala spintferu (Wentern) after Skorikov, 1903, p, 37, Diagnosis: Dorsal Jorica divided by ribs into panels or facets; foot not loricate, freely movable; ventral plalé with oval foot-opening undsy transverse distinct line; toes shart, slender, painted. Cross- section of body somewhat compressed (Fig. 14d). Male unknown. Total length 20)-255 um, lorica 95-145 jum, width 70-90. ym, spicules 5-6 pm. Distribution: Cosmopolitan in fresh and brackish water, Tately in the pelagic. Twa localities, probably more widespread, but tare. N,SW., Tas, 10.5-13.0°C, pH 6.7-8.4, DO 10.0 mg! 7. Literature: Koste 1978, Kaste & Shiel [980, Genus Trichotria Bory de St Vincent Trrehtointa Bory de St vincenl, 1827, p, 752. Head, body and three foot segments heavily loricate; cross-section hexagonal (Fig. 15:1f) (see comment with 7) buchneri for exception); Most lorica borders with spicules, grearer on “shoulders” if head contracted (Fig. 15:3, 4a); lorica with facets, granulated; head lorica composed of several plates; second foot segment with relatively long, spines (Fig. \S:la Jsp); toes long; corona of Euchlanis type? trophi (Fig. 15;ld-e) maleate; red cerebral eye (Fig. 15:]4-eye). Male loricate (Fig. 15:1b-c), Food algae and detritus, diatoms preferred. Trichotria accurs tarely in the plankton (as migrants). Seven species recognized, with many variants; five known from Australia. Key to Species of the Genus Trichotria |. Cross-sectian hexagonal, three foot segments. .2 projecting beyond posterior lorica margin; lorica clongated U-shape...) pseudecuria Koste er al, All three, of at least lwo, foal segments and toes projecting beyond lorica margin.........--. - i 3(2), Last foot joint with dorsal minute spine between the toe bases, 6.0.4, 6% poeiiun (Miller) No minute spine between the tye bitses.. os... 2 4(3). Lateral part of dorsal Jorica conspicuously large; marginal spicules of lorica directed forward wey s ot TIM, dates Ob T. truncata Whitelegee Lateral lorica not very expanded, spicules if present not directed forward, ,.. 7) fetractis (Ehrenberg) Trichetria buchneri Koste, Shiel & Tan FIG. 14:1 Trichotria buchneri Koste, Shicl & fan, 1988, pp. 120-12, Figs 3-5. Diagnosis: Lorica of triangular cross section with median keel ending in elongated caudal spine; dorsal lorica with longitudinal rows of denticles to keel (appears granulated by light microscopy); anterior dorsal margin deep U-shaped aperture between two lateral cusps; ventral margin with shallower U-shaped aperture projects beyond dorsal margin to level of lateral cusps (or slightly beyond lips); rectangular plales cover head, when contracted a pyramidal projecrian extends to or past anterior lorica margins; foor two-segmenred, without spines; Loes long, ngid, with acute paints, Length 160-182 pm; width to 125 wm; height to 84 ym; proximal foor segment 24 wm, distal 15 ym; toes 50 um. Distribution: Known only from marginal reedbeds of acid dune lakes north af Strahan, on West coast of Tasmania (1, Garela, L, Ashwood and surrounding dune pools), 17,.0-20.0°C, pH 3.1-5.15, 80.6-188.4 nS em 1, 0.9 NTU. Comment: The mutphological differences between this species und congeners were noted by Koste e/ a, (1988), particularly the domed and keeled cross- section, caudal spine and two-segmented foor, The ROTIFERA FROM AUSTRALIAN INLAND WATERS 107 Fig, 14: 1, Trichotria buchneri Koste et al. a. dorsal, b. lateral, ¢. ventral; 2. Trichotria pseudocurta Koste et al. a. dorsal; b. ventral. After Koste et af. (1988). Scale line 50 yam. armoured head is peculiarly trichotriid amongst the type locality in Jan, 1989, and additional sites for Rotifera, but the distinctive features of T buchneri the rotifer recorded. A detailed SEM study of this suggest that generic status may be appropriate. species will appear at a later date (Shicl & Koste, Accordingly, additional collections were taken atthe in prep.). 108 W. KOSTE & R. J. SHIEL Re ¢ os ey ey it {\ ea —* ROTIFERA FROM AUSTRALIAN INLAND WATERS 1c) Trichoiria pocillum (Miller) FIG, 16:2 Trichoda pocilhum Miller, 1776, po 281. Dinocharis pocillum aller Ehrenberg, 1830, p. 47. Trichotria pocitlum Bory de St Vinecnt, 1827, p. 752. yen extensive synonymy see Flarring 1919) and Koste Diggnesis: Length of font segments and spine on seoond foot segment variable; lorica spindle-shaped, surface divided into granulated facets by ridges, corona simple, can be retracted in neck and lateral head plates; male Ioricate. Rate in plankton. Length 200-300 pm (total), lorica length 110-140 pm, foot spine 55-66 xm, median minute spine on last foot segment 20-28 pm, toes $3-145 um. Distribution; Cosmopolitan in the littoral of fresh and brackish waters. N.S.W., Old_ Literature: Shiel & Koste 1979, Trichotria pséeudocurta Koste, Shiel & Tan PIG, 15:2 Trichotria pseudocurta Koste et al, 1988, pp, 121-3, Fig. 7 Diagnosis: Lorica small, U-shaped in outline with both surfaces granulated; triangular cusps ar external angles; two raised ribs on ventral surface terminate anterior to raiset| tim of foot opening: similar ribs form twin keel on dorsal surface, commence either side of median notch on dorsal anterior margin, fuse to a single keel on posterior lorica; foot 3-segniented, without spines; toes with short claws, Length to 125 pm; width to 74 pm; ioes to 30 pm, Distribution: Known only trom humic waters of Lake Pedder (White Spur Inlet), Tasmania. 21°C, PH 4.2, 32.6 nS em-1, 0.5 NTU, Resembles 7. curta (Skorikow) from the Volga R., however the latter is smaller (110 gm tong) with nore angular morphology, and lacks the frontal cusps. Wriehotria tetractis (Elwenherg) FIG. l6é:1, 3 Dinacherty tetractis Ghrenberg, 1830, p. 47, Trichatria rerractis (Fhvenberg) after Carlin. 193, p, 40. For extensive synonymy see Koste (19781157). Diagnosis: All morphological characteristics variable; loricu spinulated or not; measurements of lorica, foot segments, dorsal and ventral lorica ornamentation difler between individuals, with morphological series between T. ruacate and 7, fetractis similis (latter has. very lang toes; 120-190 ui) (Fig. 17))) caudal part of lorica with one or patred hooks (7) tetractis cvudate; spinules on second foot segment may be very short (7 sepractis Paupera). Length typical form; 218-380 um, toes to 160 win; T. tetractis similis (Stenroos, 1898): 330-40) jum, second foot segment 42-48 pm, loes 120-190 ym; T. letractis caudatar 230-260 pm, caudal hook of lorica 22-28 «im, toes 50-72 um. Distribution: Cosmopolitan in macrophytes, littoral, Commonly in shallow waters of billabongs in N.SW,, Qld, Vie. (typical form) and N.T. (similis), 13.5-24,5°C, pH 6.3-8.0, DO 5.8-9.2 mg 1°), 59-400 «2S cm> 1. 2-160 NTU. Literature: Koste 1978, 1981, 1984; Shiel & Kaste 1979; Green IY]; Rerzins 1982. Trichotria truncata Whitelegge FIG. |6:4a—b Dinacharis truncarunr Whrelogee, TBP px. 315. Dinacharts intermedia Bergendal, 1892, p. 107, Vig. 6:33. Trickotria cornuta Myers, 1938, } tH. Trichotria tetractis var. fruncata (Whitelegge) in Koste, 1978, p. 157. Diagnosis: Margin of loricu with spicules, also on foot segments, all directed forward; most with shorter toes as in 7 tefractis, but many intermediate forms; variable facetration on lorica. Length 95-146 jan, greatest width 28 ym, toes HOMME em, Distribution: Acidophil, ii SpAcgnve, mainly between plants, seldom in open water. N.SW., NOT, Vie, 25,0-28.5°C, pH $.44-7.6, DO 6.0-4.2 mg Ir), 23-145 wS emi-). Literature: Koste 1978, 198), 1984, Comments Although morphologically variable, 77 Jrencafum is readily distinguished from T. retracrés in material we have examined from both Austrahian and Canadian collections (cf, Chengulath & Koste 1983), and we consider it a distinet species, Not yer recorded from Australia perracits caudata (Lacks, 2); cf. Koste 1984, p. L16, i fetractis puupera (Ehrenberg, 1830); T. curta (Skorikov, 1914); T, eukesmrrra Myers, 1934, (See Koste, (978, pp. 187-8 for details). Pig. 16:l, Trickutria tetractis (Ehrenberg); (a) temale, lareral, swimming; [a = anus, Al = bladder, br = brain, al = dorsal lorica, da = dorsafantertna, eve = cerebral eye, fe = toot gland, fj = foot. scement, fsp = foot spines, 8a = aasiric gland, fa = lateral antenna, oe = ocsophagus, sz = sultvary gland, s¢ = stomach, vi = vitellarium, vf = ventral lorical; (b) male [dd = dorsal antenna, eve = cerebral eve, f = foot, p = penis, pe = prostate gland, ss = (estis, vy = vestigial gut], (c) male, Jateral [ef = cihared fields, corona, eve = cerebral eve, p = peiis, ss = pestis, vg = vestigial put); (d) teophi [ra = rainus,# = uncus ma = wanubrium); (¢) uncus, lateral; (f) lorica, wross-section, 2, T pocidian (Miler), fermate, dorsal, swimming; 3, 7) ¢etractis, contracted dorsal, 4, 7) tranceta (Whitelegwe): (a) dorsal, from Canada; (b) from Romania TT, trvacara f, langisping after Rodewald 1935] Fw. 15:la, 1, 2. 3, 4a dher Keste (6984): 4b, after Rodewaldl (1935). Seale Lines, leh 50 jut (Pig. 15da-c); lower cighe VOX) pret (2-4), 110 W, KOSTE & R. J. SHIEL er &), wt % ‘A Ta ROTIFERA FROM AUSTRALIAN INLAND WATERS 11 Genus Macrochaetus Perty Lorica flat, hexagonal cross-section, head part retractile; lorica granulated with minute spinules; more or less developed anal section; two loricate foot segments; toes generally spindle-shaped; paired symmetrically inserted spines (Fig. 18:la); lorica margin with spicules, shoulder hooks larger (Fig. 18:la, 7a—b); integument opaque; corona simple, with central buccal field and longer lateral sensory hairs (Fig. 18:3b); dorsal and lateral antennae on visible papillae; one cerebral eye; mastax with malleate trophi; eggs and males unknown. All species live between or on water plants and are rare in open water. Key to Species of the Genus Macrochaetus [refer to FIG. 18:1a] 1. Only one pair of rudimentary anteriosubmedian spines present (2a, b).M. danneeli Koste & Shiel More spines present..............ccc cece ees 2 2(1). Two anterolateral spines, two anteriomedian spines, two posteriomedian spines present, central dorsal spine pair absent (4a, b)....M. collinsi (Gosse) Central dorsal spine pair present............. 3 3(2). Two anteriolateral- , two anteriomedian- , two central median- and two anteriosubmedian spines present ay ace caennns M. altamirai (Arevalo) Two anteriolateral-, two anteriomedian, two anteriosubmedian- and two posteriomedian spines PIESENL. 5 ji rae Seles eed M. subquadratus (Perty) Macrochaetus altamirai (Arévalo) FIG. 18:3a, b Polychaetus altamirai Arévalo, 1918, p. 1-47. Macrochaetus esthonicus Riikoja, 1925, p. 7, Fig. 2). Macrochaetus altamirai (Arévalo) after Selga, 1952. M. australiensis Berzins, 1982, p. 18, Figs. 17, 38. Diagnosis: Shape, size and spicules at lorica margin variable; distinguished from M. collinsi only by paired central dorsal spines (see key to species). Lorica length 80-115 xm, width 76-108 ym, toes 20-28 pm. Distribution: Cosmopolitan warm stenotherm, rare. Probably more widely distributed in Australia than a single record from Victoria would indicate (as M. australiensis by Berzins 1982). Literature: Koste 1978; Shiel & Koste 1985. Macrochaetus collinsi (Gosse) FIG, 18:2 Dinocharis collinsi Gosse, 1867, p. 269, Figs. 1-4. Macrochaetus collinsi (Gosse) after Harring, 1913, p. 67. For extensive synonymy see Koste, 1978, p. 161. Fig. 17: Trichotria tetractis similis, lateral (toes 190 um). After Koste (1984). Diagnosis; Shape, size, length and number of spinules on margin variable; number of dorsal spines constant; lorica brownish, Total length 94-250 ym, lorica 53-112 wm, width 62-112 ym, second foot segment 20-25 pm, toes 16-20 ym, dorsal spines 48-66 um, posteromedian spines 24-48 wm, anal segment spines 32-35 ym. Distribution: Pantropical and subtropical, warm stenotherm, benthic. N.T., Qld, Tas. 28.5-29.9°C, PH 5.44-S5.47, DO 5.45-6.2mg 1-1, 23-44 pS cm~', alkalinity 2.6-2.7 mg 1-1, Literature: Koste 1978; Shiel & Koste 1979; Tait er al. 1984, Macrochaetus danneeli Koste & Shiel FIG. 18:5 Macrochaetus danneeli Koste & Shiel, 1983, p. 116, Figs. Ja-d. Diagnosis: Lorica granulated/pustulated; dorsal lorica with terraced sides to blunt keel; margin of keel base with longer spinules; uppermost keel dorsum covered with large cuticular ledges and beading; anal segment spineless; shoulder spines (Fig. 18:la, 7a), posterolateral- (Sa, b), TT - —————— Sn— —— SSFSSSSSSSSSSSSSSSSSSSSSFFMsesF Fig. 18: 1 (a) points of insertion of spines and setules in Macrochaetus Perty spp., dorsal lorica and foot segments [/a, 6 = anteriolateral spines, 24, b = anteriosubmedian spines, 3a, b = anteriomedian spines, 4a, b = central dorsal spine pair, 5a, b = posteriolateral spines, 6a, b = posteriomedian spines, 7a, b = shoulder hooks, 8a, b = anal segment spines, 9a,b = foot segment]; (b) Macrochaetus lorica, cross section. 2, M. collinsi (Gosse): (a) dorsal, swimming; (b) lorica, ventral. 3, M. altamirai (Arevalo): (a) dorsal; (b) ventral. 4, M. subquadratus Perty: (a) dorsal; (b) dorsal; (c) ventral; (d) lateral. 5, M. danneeli Koste & Shiel: (a) dorsal; (b) ventral. Fig. 18:la, b, 2a, b, 4a, 5a, b, after Koste & Shiel (1983); 3a, b, after Rodewald (1940); 4a-d, after Wulfert (1964). Scale lines 50 zm each number group. 112 W. KOSTE & R. J, SHIEL posteromedian- (6a, b) and only one pair of rudimentary anterosubmedian spines present. Lorica length 125-130 wm, width 130-137 pm, foot segment 20 pm, toes 16-18 «pm. Distribution: Single locality known: Buffalo Billabong, Magela Creek, N-T. 30.9°C, pH 5.38, DO 4.15 mg 1-1, 29 pS cm-!. Macrochaetus subquadratus Perty FIG, 18:4a-c Macrochaetus subquadratus Perty, 1850, p. 22. Polychaetus subquadratus Perty, 1852, p, 45, Fig. 1:6. Diagnosis: Lorica generally hexagonal (Fig. 18:4a); ten dorsal spines of variable size and shape; shoulder spines most conspicuously large. Lorica length 80-135 ym, width 73-140 um, second foot segment 8-16 zm, toes 15-22 pn. Distribution: Cosmopolitan in periphyton of aquatic macrophytes. Rare migrant in plankton, more common in billabongs. Thermophile. Literature: Koste 1978; Shiel & Koste 1979; Green 1981. Not yet recorded from Australia M. hauerianus Wulfert 1964 (=M. collinsi var. Hauer 1938), p. 534 (Indonesia); M. longipes Myers 1934, p. 13; see Koste 1978 p. 161; M. multispinosus Myers 1934, p. 12; recorded from the neotropics and S. Nearctic; M. philopax Wulfert 1961, p. 87; single find in central Germany; M. sericus (Thorpe) 1893, p. 182; see Koste 1972, Koste & Shiel 1983 p. 117 Fig. 8. Acknowledgments The support of the Australian Biological Resources Study in grants to RJS, the use of the facilities of the Botany Department, University of Adelaide in manuscript preparation and long-term loan of microscope facilities to WK by the Deutschen Forschungsgemeinschaft are gratefully acknowledged. The Department of Lands, Parks and Wildlife, Hobart, supported fieldwork in the World Heritage Area in Jan, 1989, Particular thanks to the staff of the Peter Till Laboratory, Albury- Wodonga Development Corporation (now Murray- Darling Freshwater Research Centre) for providing collections from billabongs in their study area, to Professor D. G. Frey, Biology Department, Indiana University for the donation of an extensive series of rotifer publications from his Limnology Library, and to an anonymous referee for critical comments on an earlier draft. Our thanks also to Charles Hussey (BM(NH)) for checking obscure references, Anne Grant, CSIRO Publications Service, for editorial expertise, and Lor Wai Tan for final corrections. References AHLSTROM, E. H. (1934) Rotatoria of Florida. Trans. Amer. Microse. Soc. 53, 251-6. AREVALO, C. (1918) Algunos Rotiferos planktonicos de la Albufera de Valencia. Ann. Inst. Gen. Tech. Valencia 8, 1-47. Bartos, E. (1959) Virnici. Rotatoria. 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Rotifers ard general character of the planktin.| Aan. Mus. Aad. Sci, St Petershure Wa, 180-214. [Russian]. Vrasovsry, M, (1957) Neikdko virnikay (Raratoriat ¢ ryzovych pali na Sloveusku. Biofagsa Bratiskava 12, 661-67), Want CuiCiu. (1961) [Treshwater Rotifera of China”, Inst, Frestwater Hydrobiol,, Poking, 288 pp, [Chinese]. Wenen, F (t898) Faune rotatorionne du bassin de Leman, Rev, Susce. zool. 5, 263=785. WeSENRERG-LUNPD, ©, (1924) Roratarla. Handbuch a, Zavl. 2, 1-120, Western, G. (1894) On Dustvia spinifera, J. Queckest Micn ©), Sev, 2.5, 427-428. WHITELEGOE. T, (1889) List of the marine ane freshwater invertebrate fauna of Port Jackson and the neighbourhood, Proc, R. Soc. N.S, 23, 163-323, WIERZEISKL, A. (1893) Rutatoria (wrorki) Galivy!, Bell Juternat. Acad. Sci. Cracovie (1892), 402-407 Wisesiewski. J. (1953) Fauna wrotkow Polski j prayleglych, Pol. Areh, Ayadradbioi 1, 417-490 Wuteekl, K. (1939) Beilrdéve sur Kenninis der Radertierfauna Devischlands, 1V. Die Radertiowe der Saale-Blster Niederuny bei Mersebury in okulogiscl:- faunistischer Dezeihung. Arch. Myirobiol. 38, So4-624. (1840) Retatarien einiger oscdeutscher Torimoare. Arch. Aydrotiol. 36, 552-587, (942) Neae Rotutorienarlen aus deulachen Mineralquellen. Zon! Avs. 137, 187-200, —_____ (1951) Das Narurschiuzgebrer auf dem Glarzer Sthuveberp, Die Kaderticie des Natursulitegebictes Arch. Mydrohiat. da, 441-471. Raderticre des teutelssees her Fricdsichshagen, Arch, Avdeobial, 51, 457-495, eee (1960) Die Radertiere saurer Gewsseer ler Duhener Pleide. [. Die Rotatornien des Zadlitzmoprs und des Wildenhainer Britchs. ¢oid, St. 31-359 (1961) Die Radertiore sawer Gowasser dor Dilbuner Heide. IIE. Die Rotatorien des Presseler vind des Winkelmuhler Teiches. Sha. S&, 72-102, (1964) Unsere sepenwarlige Renolois der Rotoriengatlung Waerechootus. Limneaowicu Berlin 2 281-3049. 41965) Raderticre ane cinigen alrikanischert Gewussern. [tid 3, 547-3 YAMAMOTO, K- 41953) Preliminary studies an the Ratatonian fauna of Korea Pei! Sey. 7, (51-104 BRIEF COMMUNICATION 15 REDISCOVERY OF UPEROLEIA ORIENTALIS (PARKER) (ANURA: MYOBATRACHINAE) Glauertia orientalis was described from three male frogs collected by W. H. Stalker in 1905 on Alexandria Station (19°08" , 136° 43" vide!) on the Barkly Tableland, Northern Territory, and a female frog collected by G, H. Wilkins in 1925 from Groote Eylandt'. The species was referred to G/auvertia Loveridge because of its moderately extensive webbing on the toes (Fig. 1) and its widely exposed frontoparietal fontanelle. Fig. 1. Plantar view of the foot of Uperoleia orientalis (Holotype, Alexandria Station), Glauertia orientalis has been reported over a wide range in the north of Australia2,3,4. The genus was referred to the synonymy of Uperoleia® when it was shown that the generic characters represented extremes in continua expressed within Uperoleia. U. orientalis was redescribed from the holotype and topotypic paratypes® but the Groote Eylandt paratype was not examined at that time. Its conspecificity was called into question because of distribution and other data provided in the generic revision®. No other referrable material was available to these authors. Later collecting ® failed to produce further specimens of U orientalis at the type locality, and further studies on Groote Eylandt resulted in the female paratype being referred to U. inundata Tyler, Davies & Martin.’ Uperoleia orientalis thus remained known only from the type locality and from the holotype and two topotypic paratypes. Whilst examining a collection of Uperoleia from the Northern Territory Museum (NTM), I found a specimen referrable to U. orientalis. The frog (NTM RA88) was collected by Mike Gillam during the Cox River Survey on 2.7.1977 adjacent to the Arnold River (15° 43’, 134° 37°), at night in paperbark woodland (Melaleuca spp) in a spring-fed swamp. The species was abundant at the locality9. The specimen is small (probably sub-adult) with a snout- vent length of 17.6mm. Toe webbing is moderately extensive (characteristic of the species) and there are very prominent subarticular and supernumerary palmar tubercles on the hands. The head is small in proportion to the body and the parotoid glands are prominent. Inguinal and coccygeal glands are extensive and coalesce. No mid-vertebral stripe is apparent. The site of collection represents a range extension of about 500 km to the north north-west. Parker, H. W. (1940) Novit. Zool. 42, 1-106. Barker, J. & Grigg, G. C (1977) “A field guide to Australian frogs” (Rigby, Adelaide). Cogger, H. G. (1975) “Reptiles and amphibians of Australia” (Reed, Sydney). Tyler, M. J. (1976) “Frogs” (Collins, Sydney & London), Tyler, M. J., Davies, M. & Martin, A. A. (1981) Aust. J, Zool. suppl. Ser. 79, 1-64. Tyler, M. J., Davies M. & Martin, A.A, (1983) Trans. R, Soc, S. Aust, 107, 237-242. Tyler, M.J., Davies M. & Watson, G. F. (1986) Zool. J. Linn. Soc. 88, 91-101. Tyler, M, J. & Davies M. (1986) “Frogs of the Northern Territory” (Conservation Commission of the Northern Territory, Alice Springs). Gillam, M. Jn litt. MARGARET DAVIES, Department of Zoology, University of Adelaide, G.P.O.Box 498, Adelaide, S, Aust, 5001, THE OCCURANCE OF PLANT PARASITIC NEMATODES IN THE ARID REGION OF SOUTH AUSTRALIA BY JACQUELINE M. NOBBS Summary BRIEF COMMUNICATION ul? ‘THE OCCURRENCE OF PLANT PARASITIC NEMATODES IN THE ARID REGION OF SOUTH AUSTRALIA There is Jittle information on the occurrence of plant parasitic nematodes in arid South Australig. Twenty-four genera of plant nematodes from the area encompassing the Northern Flinders Ranges to Cape Jervis have been identified tovether with a réelanonship between the ihistribuuon of plant species and the distribution of plant nematodes 7. The distribution of plant species was related to Yainifall. Following on from this work, a survey of the arid region (mean annual rainfall less than 250mm) was undertaken over a four year period (1983-1986). The sorl was sampled from under native vegetation, the nematodes extracted using a modified Bacrmann’s funnel technique 2 and where possible the nematodes identified to species level. At many sites few mature female specimens were extracted, so identification to genus only was. possible (Table 1). There were several genera of fungal feeding tylenchids identified which included Aphelenchus avenae, Aphelenchotdes sp, Coslenchus c.f, costatus, Neotylenchus s.1. Dlenchus sl, and Dinvlenchus sp.. These are not included with the plant nematodes in Table 1. ‘Jwelve genera of plant nematodés were identified from the arid region ay well as juveniles and migles of the Heteroderidae which were not identifiable because of the absence of females or cysts. The most widely distributed plant nematode was Tilencharhynchus tebari (Sauer & Aanells, 1981), Of the 374 sites sampled, 63% contained 7 tobari. All other species occurred at less than 8% of the sites. More than Ohe taxon was present in 26% of the sites and a similar percent of sites had no plant nematodes present. The arid regions are an adverse environment for moxt organisiny. To survive, organisms musr be drought resistant and also respond rapidly ta the stimulus of rainfall. Some nematodes are ideally suited to the arid region as they car form an anhydrobiatic state in which metabolic respiration is greatly reduced bur are quick to reyive with addilign of water 3 and begin to reproduce rapidly after revival, T, tobari (Sauer & Annells, 1981) is a widely distrituited nematode within the region and must be well adapted to the environment, The species was first described from a site near Wentworth, N.SW, * The species was. cultured by planting seeds of certain species of Chenopodiacexe (a common family in arid Australia) into soil known to contain Tf tohari. Studies of host/parasite relationships aad covironmental effects on nematode population growth will be presented ia detail in another paper, | meee & Wallace, H. R. (198!) Nematologica 27, 329, 2 Schindler, A. KF. (1961) Plant Discase Reporter 45, 747-748. > Freckman, D, W., Whitford, W. G. & Steinberger, ¥. (1987) Biology and Fertility of Soils 3, 3-10. 4 rhe i R. & Aunells, CM, (198) Nematologics 27, JACQUELINE M. NOBBS, CABI Institute of Parasitology, St Albans, Herts,, UK. AL4 OXU. TABLE |. Nuytiber und percent of sites Jrom whieh plant nematodes (PP) were extracted in the years 1983 t0 1986 rarciusive. Genus and Species Trlenchorhpncdus tobari Saver & Annelis, 1981 Pratyleachus sp. Morulaimus sp. Helicotylenchus sp. Seutellonenia sp, Radopholus sp. tulenchorhynchus velatus Sauer & Annells, 1981 TWienchorhynchiss sf. Reivlenchys sp, Heaplolatinus sp. Morulaimus geniculatus Sauer, 1966 Puralongidorus sp- Heteroderidae males aud juvenites Séutellanema laeviflexum Phillips, (971 Trlenchorhynchus unmiulgies (Cassidy, 1930) Golden, 1974 Scutellonema minutum Sher, 1964 florlenchus hastulaius (Golden, 1960) Siddiqi, 1963 Melicotylenchus yariebilis Phillips, 1971 Radopholus crenatus Colbran, {971 Paratylenchus sp. Hemicycliophora sp. foral number of sites sampled Sites with >1 genera af PP Sites with PP Sirés without PP Sites Percent of Sampled Total Sites 237 63.3 24 6.4 16 43 16 4.3 ity 4.3 14 37 12 3.2 12 3.2 ? 1.9 7 1 § 13 4 tl 3 0.8 3 0.8 4 0.8 2 05 1 0.3 1 03 1 03 “4 0.3 i 0.3 374 100.0) 07 25.9 276 Ti8 98 26.2 VOL. 113, PARTS 3 & 4 30 NOVEMBER, 1989 Transactions of the Royal Society of South Australia Incorporated Contents. Koste, W. & Shiel, R. J. Rotifera from Australian inland waters IV. Colurellidae (Rotifera: Monogononta) - - 2 : : = : = : Short, A. D., Buckley, R. C,, & Fotheringham, D. G. Preliminary investigations of beach ridge progradation on Eyre Peninsula and Kangaroo Island - Pledge, N. S. The occurrence of a diprotodontid marsupial referable to Euowenia grata De Vis in western New South Wales - - = = = Austin, A. D. & Allen, G. R. Parasitoids of Uraba lugens Walker (Lepidoptera:Noctuidae) in South Australia, with description of two new species of Braconidae - = = a * = p ey Barker, S. Contributions to the taxonomy of Australian Buprestidae (Coleoptera): New species of Astraeus and Stigmodera (Castiarina) and a key to Astraeus CS ee Jenkins, J. F. & Hasenohr, P. Trilobites and their trails in a Black Shale: Early Cambrian of The Fleurieu Peninsula, South Australia - - - = Gowlett-Holmes, K. L. & Holmes, N. J. C. Rediscovery of Primovula (P) heleneae Cate and description of a new species of Crenavolva from South Australia (Mollusca: Gastropoda: Ovulidae) - - = = = Davies, M. Developmental biology of the Australopapuan hylid frog Litoria eucnemis (Anura: Hylidae) - - - = Ee a 4 z Brief Communication: Beardsell, G. R. Hybridisation of Litoria chloris and L. xanthomera, (Anura: Hylidae) - - - © : J , = : ¥ 119 145 163 169 185 195 205 215 221 PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 113, PART 3 ROTIFERA FROM AUSTRALIAN INLAND WATERS IV. COLURELLIDAE (ROTIFERA:MONOGONONTA) BY W. KOSTE*, & R. J. SHIELT Summary Diagnostic keys are given to genera and species of the rotifer family Colurellidae (Colurella, Squatinella, Lepadella, Heterolepadella) recorded from Australian inland waters. All species are figured and available distribution data and ecological information given. Some widely-distributed taxa not yet recorded from the continent are included. KEY WORDS: Rotifera, taxonomy, Australia, known species, Colurellidae Transeenions OF he Ravel Suctety af §. Atyt, (S989), 10303) 19-145 ROTIFERA FROM ALSTRALIAN INLAND WATERS [V, COLURELLIDAE {ROTIFERA:MONOGONONTA) by W, Koste* & R. J. SHIELT Summary Kosta, W., & SHlel, R. J, (1989) Ronfera fram Australian inland waters, CY. Colurellidae (Kocifera: Menogononta), Trans, R. Sov, 8, Alust, 113(3), 119-143, 30 Noveinber, 1989. Diagnostic keys are given Lo genera and species of the rotifer family Colurellidae (Coluredia, Squatinalla, Lepadella, Heteralepadella} recorded from Australian infand waters. All species are figured and available distribution data and ecological informavion given, Sonte widely-distesbuted taxa nor yet fcoordéed trom the continent are included. Key Worps: Rotifers, iaxdnomy, Austraba, known species, Colurellidae Intraduction The monogonont rotiter families considered lu ate in our revision of the Australian Rotitera (Koste & Shiel 1986b, 1987b, 1989) have included the mosr common planktonic taxa in Australian inland waters. Here we review the Australian represen- (atives of the family Colurellidae, several genera of which: have widely-distribured species in plankton and littoral collections in Australian waters and are considered heleoplankters (semsw Hutchinson 1967) rather than accidental ineursions, i.e. pseudoplankton (= fychoplankton af Ruttner- Kolisko 1974) (eg, Lepadella patella). In general, however, the Australian representatives of the Colurellidue (Colurella, Squutinella, Lepudella, Hererolepadella) are littoral (epiphytic or epibenthic) in habit. They are encountered cammonly in the shallow vegetated areas of billabongs and other werlands, where diverse mImgermphyte associations effectively partition the habitat, providing abundant microniches. Diagnostic keys, brief descriptions and figures of all known Australian representatives of the family are provided to enable identification to species, Some widely distributed taxa not yet recorded from Australia are included, The formar follows that of eurher parts; dichotomous keys are followed by individual species” descriphions and koown Australian records, Relative abundance is indicated bys “conmmoan” (more than 30 widely dispensed records), “uncommoan" (10-30 localities), “rare” {<10 records). Brief ecological dats are included Where available. Early Australian records cited in Shiel & Koste {1979} are nol repeated here, nor are the majority of northern hemisphere citations given by Koste (1978), which contains a detailed treatment * 5, Ludwip-Brill-Strasse, Quakenbrtick 9-4570, Fecera) Repubtic of Germany } Murray-Darbng frestiwwier Research Contec, PO. Box 92), Albury, 6 SW 2640) of the Rortifera outside Australia, Keys to families are included in Kostc & Shicl (19876). Family Colurelidae Bartos Loricate rolifers, the vorena in all genera with 4 membraneous cap or semicircular shield (retractible in all except Squatinel/a); no lateral sulct; Colurella and Paraceluretia with ventral sulcus; (rophi malleate; ventral plate with caudal foat- Opening or enlarged Jor basal toot segment; toes slender, sharply pointed, Four genera; Colurella, Lepadella, Squatinella, Paravoluretla. Key to genera of the Family Colurcllidae }. Lortea laterally compressed, wilt venwal or sori and ventral apertures (Fig. 2),....-...-...... 2 Lorica dorso-yentrally flattened, without such apertures -.--...-.--- ----- - 3 2(l), Lorica a single structure, with ventral aperture (Pig. U.....---..Calurella Bory de St Vincent Lorica two plates, with. dorsal and ventral apertures (Fig, 2:8)....... Puracolaretia Myers (not known from Australia). 3(1). Hood or head-sineld large, non-retractible (Figg) oo... Squatinelia Bory de St Vincent Hood small, retractible (Fig. 5) poems Lepadella Bory dv St Vincent Gentis Calyrela Bory de St Vincent Colunmila Bory de St Vincent, 1824, 17.203, One piece loriva, in lateral view oval, egg- or hatchet-shaped (outline of lorica differs according to degree of contraction, particularly ventral margin); anterior lorica margin more or Jess (rancated or rounded: posterior margin rounded, gradually merging into blunt, straight or curved proj-tions, in some cases sharply pointed, offset win ike extensions of posterior margin (Fig. Ic}: lan W, KOSTE & RK. J, SHITE) ventral cuticle between mussel-like lorica borders delicate, membrancous, may be distoricd in preservative; corona of the Euchlunis type (cf, Koste & Shiel 19876); loot with three or four segments, basal segment mostly pliable and often of indistinct structure; distal foot segment (before toe) with dorsal sensory pore; toes of variable length, species with long toes more compressed laterally; toes separate, however may appear fused as frequently are stuck together; gastric glands mononucleate; trophi malleate; eyeless or with two. eyespots, which may or may not be red-pigmented; lateral antennae very conspicuous; male little known and insufficiently described. Resting eggs retained in lorica of parent an deuth (cl. Fig. 2:3). Variability in the described species requires investigation, particulatly whether the morphology of different races (“var”) or forms represents thes Pig. tl, ecotypic variation [“Millienfaktoren” (Hauer 1924)], e.g. the variability in the lorica ends in C uncinata or © adriatica, Eeotypic variation is common in the Australian brachionids (Koste & Shicl 1987b), and variants are known in the five species of Colurella recorded from Australia. It is likely that other species occur here; in view of their small size (most <100 ym) colurellids are easily overlooked in detnitus-rich samples. Key to species of the Genus Coturefla {, Lorica surface smooth. 2.00. 00.2-- 02.00.0008, 2 Lorica surface with longitudinal furrows or facets icht it pew. tr CG. fesselala (Glascolt) 2(1). Loriva "Vale (lateral) slender, posteriorly with Watiable apices,,.. 2.0.0... e ee ee ees Lorica valve rounded, Pliiied Faocek here ses a Oe 0 uN us Colurella adriativa (Ebrepberg): (a) taterat (an = anus; du — eyes er > subilaneous eg, ax = mustilx; vi = vitellarivint); (b) lateral, contracted; (e) ventral: (dd) posterior lorica showing foot and we (DS. = dorsal sensory mit}; (e-1) lateral views showing Variation in lonca morphology, 2 lateral: (dg) male, lateral. 7, ¢* Corda (1834); 1h, 3d, alter Carlin (1939): le, (1953): Va-c, atter Hauer (2924). Scale lines 20 pm, WC. colurus (Ehrenberg): (a) dorsal: ¢b) ventral: (c) vitusy (Coasse), (2) dorsal: (b) ventral: (a Vater, (d) anorher morph, lateral. la, afrer d, wtier Waulfert (1980); fest, Zuse, aller Hauer (1928); 20) aller Wiszmewsky ROTIFERA FROM AUSTRALIAN INLAND WATERS Pal 3(2).. Posterior end of lorica with raised lines terminating in varnible tupering, downward-curviny (oveasion- ally lightly upward-curving) spines; lorica high (eneth/height rario 1.8); toes long Sein Lorica Valve postenarly rounded, or at most extended tes blint apex (Fig. 1;2c)...,, 128906 -C ecalurus (Ehrenberg) Loriva valve posteriorly ‘with curved spines (Fig. Hc) Baalebne'a''s g & adriatica Bhrenhere ColureHa adrartica Ehtenberg FIG, V4 Coluretla adctatiea Ehrenberg, 1831, Fig. 3:3. Piapnosis: Extremely variable lorica size and phenotype, particularly points at posterior end of lorica. These are the only features separating C adriatica from C calurus, Pejler (1962) noted transitional forms, Different ecotypes were des- eribed by Hauer (1924), with increased loricu size and Woe length correlated with increasing salinity, The measurements below encompass the range of ecolypes (Koste $978), Length 85-113 ~m; width 29-48 pm: height 44—66 pam; toe length 28-48 xm. Disiribution: Probably cosmopolitan, Euryhaline. Rare: fresh and slightly saline waters of southeast S. Aust, Tas. and western Vic. Berzins (1982) recorded C udriatica asx widely distributed in eastern Australia. 5.0-30.0°C, pH 5.5-10.5, dissolved oxygen (DO) 6.0-11.2 mg | |, conductivity (K,,) §7-13,940 uS cm‘ turbidity 21-67 nephelometric turbidity units (NTU). Literature: Ridder 1972; Kusle 1978 Colurella colurus (Etrenberg) FIG. b:2 Meno colurus Ehrenberg, 1430, p, 44, Colurella colurus after Harriny, 1913, p, 29, Diagnosis: Distinguished trom C adriatica only by the rounded or blunt-spined posterior lorica, Resting eve and male known (Fig. 12d). Length 71-110 um; width 25-37 wm; heieht 39-54 xm; toe length 25-40 ym. Distribution: Cosmopolitan, widely tolerant, oecuc- ting in fresh, arhallasic saline and coastal waters. Huropean temperature 1,0-28,0°C and pti 4.9-10,0, Ridder (1972) records ©. colurus from 15,035 2 Cl Ll . Rare: Tas, Vie 1.8-16.0°C, pH 3.7-6.4, 495-2130 eS cm. Lilerature: Koxte |978, Colurella vhtusw (Gosse) FIG, t.3 Colurus obsusus Cisse fr Hudsen & Gidsse, 1886, p. 103, Fig. 26:3, Colurella abtusa after Holsien, 1909, (x. 84. Diegnasis; Lonca outline plump; margin of bead- uperture directed obliquely from ventral lorica; ventral lines variable; head width approx. halt Jorica length, or smaller; eyespots with or withour pigment. Length 60 pm; width 28 pm; toes 17 am, Distribution: Cosmopolitan between aquatic mac- rophytes; pancontinental, uncommon, 8,.0-29,0°C, pH 3.4-7.5, DO 11.2. mg 1", 36.4-1600.p5 cm !, {5 NTU. Literature: Eyans 1951; Koyte 1978, 1981; Shicl & Koste 1979; Green 1981: Koste ef of, 1988. Colurella lesselata (Glascat) FIG, 2:1 Calurus tesselatus Glascott, 1893, p, 74, Colurella tessélata atier Hofsten, 1909, p. 84. Diagnosis, Loriea with dorsal keel; facet-like pattern with ribs. on lorica surface; toes relatively short, Length 55-64 am; width 24-31 pin; height 32-41 puny, toes 18-27 wm. Distribution: IC -osmopohtan (nat recorded from N.Z,), sporadic in oligotrophic waters, mostly in moss, also in psammion, Single record from Coghills Creek, Tourello, Vie. Literature: Koste 1978; Berzins. 1982, Colureila uncinata uncinala (Miller) FIG, 2:2 Brachionus uncinatiy Muller, 1773, pp, 134, ColureHa uncinata after Having, 1913, p, 30, Diwgnasis: Lorica posterior end gradually merges into downward-curving short spines; head-opening smaller than greatest lorica height. Length 77-106 jem; width 30 pm; height to 60am: toes 18-23 wm; male 50-85 «wm, Distribution: Cosmopolitan, eurvhaline, Common- pancontinental. &.5-24.5°C, pH 4.8-8.0, DO 7.8-11,2 mg | ', 20-1600 ws cm~', 3-160 NTU, ‘jtérature; Koste 1978, Comments; Two. morphologically distinguishable taxa described originally as distinct species, but regarded by Koste (1978) as “forin” yariants of C uncinata also até known from Australia. Colurella uncinata bicuspidara (Ehrenberg) FIG, 2:3 Coluras bicuspdaias Ehrenberg, 1832, pm 123, Cohtrella necinatd C. bieuspidata alter Mewche, 1939, px W. KOSTE & R. J. SHIEL ROTITERA FROM AUSTRALIAN INL ANI? WATERS 121 Diegnosis: Loriva width up to twice that of noininate species; highest extension of torica al head-opening; lorica ends short, most cume downwards; toes relatively short; resting exe with punctiform cavities, Lengih 60-108 «mj; width 53-62 pm; height to A um; toes 13-27 pm. Distribution: Cosmopolitan in littoral of fresh and athallasic saline waters, alsa marine Littoral. Uncomnion: N.SW,, Qld, 8. Ause, Tas. Vic. 10,0-23.5°C, pH 4.5-7.5, 93-927 pS am | Literature: Koste 1978, Stuel & Koste 1979; Green IYRI; Berzins 1982, Calurella unciaala deflecd (Ehrenberg) FIG. 2:4 Cotarus defleris Ehrenberg, 1834, p. 203. Celurelia uncinata \, deflexa alter Meuche, (939, p, 400}, Diagnosis: Lorica end robust, clongated, turned outward, most directed ventrally. Very variable. Length 60-100 am; width to 55 yin; height 35-55 um; toes. 17-25 pny, Distriburion: Cosmopolitan, sporadic, euryhaling. Rare. Qld, Tas, 9.5-20,0°C, pH 6.5. Hiterature: Shiel & Koste (979; Koste & Shiel 1987a. Not recorded trom Australias CO. denticauda Carlin, 1939. Scandinavia dicentra (Gosse, (887), N. Eurype, tance | hindenburai (Steinecke, 1917)(Fig. 2:6), Probably cosmopolitan ©. geophila Donner, 1951. Burape ©. mariqavi Althaus, 1957, Black Sea, mesopsammal Aan i © mondodacivlos Althaus, 1957, Black Sea C. mucronulatea Atitstrom, (938, U.S.A, © ob/onga Doaner, 1943. Europe C. oxycauda Carlin, 1939, Scandinavia C. paliidasa Carlin, 1939, USSR, C satiaa Althaus, $957 (Fig. 2:7). Europe, N.4. C. sinisira Carlin, 1939, Furope C. suleata (Sienroos, 198K Fyr. 2:5), Probably cosmopolitan, warm 2000 collections, however it Was recorded by Whirelegge (1889) from N.SW,, also by Evans (1951) front Victorian waters. 1 iterature; Pejler 1962; Kaste 1978. Fig. 2, Coluretfa tesselata ({Glaseott); fa) dorsal; (bu) lateral. 2, C vnemnara uncinare (Miter): (a) lateral: (b) dorsal te) male, ventral; (d) male, lateral. 3, C wactnala bicwspidaty (Fhcenbers): (a) tatecals db) another moeph, lateral: fe) ventral (d) ventral, with resting egy. 4, CL uavinare defieve (shrenberg): (a) literal; (b) hubiv, dorsal. 5, C. stefeutes (Stcnrnos), lateral 6, C. hiadendured Steineekes (a) dorsal; (b) literal. 7, CL valine Alnaus: (a) laterals (b) ventral. 4, Paravalirefie lagna Myers: (a) lateral; (b) ventral, Ja, 6, 2b, 46, afer Haier (1924)5 lo, 2a, Ra-c, da. 3, alter Curlin (1940); Ze, U, alter Weber (1898); 3d, Sat. b, aller Kasic (1978); 6a, by alter Steineeke (YLT), Fa, by after Alulmuy (19571 Seale lives SO pan. 124 W. KOSTE & R. J. SHIEL KOTIFERA FROM AUSTRALIAN INLAND WATERS 124 Squarinella devdlell lougiseta (Pourriot) FIG, 34 Squadinetla teveigiiT, fangisera Pouiriot, [Y71, pp, 99-100, ripe 4s Lnagnosis: Robust body; dorsal spine longer than body; two foot segments; no spite on distal end of terminal segment (at base of toes) Much larger animal than §. /ongispinate. Body length 210-235 am; dorsal spine 270-378 pm; ratio spinethody length 1.6; tocs to 37 pm. Distribution: Rare ino submerged Sphagnum, Myriophytium, etc, Single record trom @ pool near L. Gareia, western Tas, 17.0°C, pH 3.1, 80.648 cm 7, Comment: The nominate S. feydigil is not recorded from. Australia, [t is distinguished from S. longispinata by foot morphology and from the ssp, longiseta by its markedly smaller dimensions (body length 10-130 um, dorsal spine 90-110 jm: spine:body ratio 0.8-0.95; toes Lym). Literature: Koste 1978, Squatinella mutice (Ehtenbere) FIG. 3:2 Stephanopyomuticus Ehvenberg, 1832. p. 138. Squaetinel/a mutica after Harring, (YI, ps. 97, Diagnosis: Caudal mann uf dorsal lorica semi- cirenlay, tongue-shaped, three-lobed, with three short spines or truncate (Fig, 3:2d-g), All forms may’ occur in same popwation, Foot three-seemenied, Body length 100-226 pm; headshield width 60 ym; toes 10 30 pun. DEiribyhon: Europe, N.Z. Rare, from billahongs, nonds or dams: N.SMW,, NW, Tas., Vic. 16.5-24,5°C, pH 3.8-5.3, 45.5-46.2 pS um |. Literature: Russell 1961; Koste 1978; Berzins 1Y82: Koste & Shiel 1987. Squatinella Mmuricn tridertteata (Fresenius) PCH, 3:5 Srepharnans tricdendius Fresenius, 1858, p. 28, PL 1d, Fig, 1 Sauatineda maticu tridentata afler Wisaniewsh), 1953, p. 401. Diagnosis: Posterior lorica oval, terminating in three pointed, clonguted spines of variable length. Resembles §. rostrum, but lacks spine at base ul toes, Headshield semicircular. Body tength 134-190 pm; toes 20-30 jatn, Distribution: Ponds and pools, often abundant in Sphagnum. pH 4.8-6.2, Rare, Qld, N.SW., Vie. Literature: Evans 1951; Koste J978. Squatinella rosirum (Schmarda) FIG, 3:1 Lisirion rostran Schimarda, 1646, p 20, Pl 2. Pig. t. Syuatinella rostrum after Carlin-Nilson, 1949, p. 3 Pragnosis; Caudal lorica with three long or short pointed processes which arise from narrow bases, Lateral antennae sunken in dorsal lorica margins. Length 150-217 pm; toes 24-28 pm; headshield to 76 pm; caudal processes fo 36 fm; spine at base of tes ta 23 pin. Distribution; In Aulwuchs (i.¢, periphytun attached to submergent stems), Pwo records only, Brisbane, Qld and from Creswick Ck. Clunes, Vie. Literature: Thorpe 1887; Koste 1978; Berzins 1982. Not recorded from Australia: S. biferca (Bolion, 1884) in Hudson & Gosse (L886) Kurope, lecland. S. mivredactyla (Miicray, 1906), Europe, Newtoundland, in SpAagerum, Genus Lepadella Bory de St Vincent Lepadella Bory de St Vittcerit 1R26, p &6, Outling-oval, pear-shaped or circular, more or less dorso-ventrally Hattened; dorsal and ventral lorica plates firmly joined: deep notch for head and wor; ocvipital margin of lorica often with granulated collar; head with cap-like plate, Dorsal lori¢a domed, smooth, with or without keel development, also with several ledges or fillers; lorica margins ih some species drawn out into winglike extensions. Head and foot apertures of different Shape; differences useful in taxonomy of group, Foor of three flexible segments, one inflexible basal segment, Toes short and pointed or long and nvedle- like, similar or dissimilar in length, inserted laterally or dorsal-ventrally, in some species completely or partly fused. Corona single ciliary wreath with lateral ciliary tufts and buccal field. Two lateral eve spots, occasionally with separate pigment granules Fig. Mb Squutnedts rostram (Sehmarda): (a) headshield, dorsal; (6) headshicld, ventral; (c) dersal td) lateral amenna: (ep ventral: (1) lateral, 2; So inuticg (Ehrenberg): (a) ventral; (by lateral: (ce) dorsal (d-e) variations in vaudal morpholowy, 4.8. ompispinata (Tatem): (a) taveral; (b) dorsal; (c) foor, lateral. 4, 5. /evdieit tonetsera Pourriol. SS. nitien tridentate (Fresenius): (a) dersals (b) lateral, lH, by afler Webou (1898), Je-t, 2a, fy, da, alter Wultert (1999); 2c-z, ather Koste (1978); 3a, after Pejler (1962); 3b, atler Slenroos (1898), Ju alter Mycrs (942). dh, ater Gillard (1948), 8, after Pourtie) G971). Serle lines $0 yam, 126 nearby. Dorsal and lateral antennae present, lattct generally dorsally situated at level of upper border of fuot-opening, Last foot segment usually with caudal antenna. Trophi maleate, uncus with five teeth. Gut with large, sometimes lobed gastric glands. Vitellarium large with eight nucleii. Proionephridia with distinct terminal ces and large bladder, Litthe known of reproductive cyele, Some males (90-120 pin) wre Known, Resting ees, will prickly outer shell, retained in lorica of parent on death. JTepadella species. are benthic, commen in billabongs and ponds, but occasionally collected as incursion species in the plankton of reservoirs and rivers, og. L. patelhy (Shiel ef al, 1982) Further details wre given by Koste (1978), who notes thar all Spevies can be identified trom preserved nvaterial, although caution must be takeb with preseruition artelucts, particulady uo the case of formalin- preserved specimens, Several of the 45 tasa ot Lepetelia identified from Australia probably are misidentified for this reason. These are considered in the texc. Nevertheless, about half of the known species upper to be indigenous; Lhe genus is thus second only wo Brachionys (Kost & Shiel 1987b) in ats vonsplerment of endermiv taxa. Three subgenera are recognized on the basis of toc morphology; i) fepedella (Woes of equal length, entirely separated); (ii) Hererolepadella (toes af unequal lenethy; (iii) Nenolepadella {toes complelely or partly Tused). Por convenience, the Australian species of the three subgenera are considered separately in the following keys. Key to species of the genus Lepadelia (s. str) 1, Dersa) lorica without keelor ribs (may be lightly suieiate, Hur otherwise wnomamented)....2 Doroal Jones with keet declan longuudinal ribs or Other ornamentation... . ccs peeee esl? 20). Lorea tength > 3s lorica width, wlmast cylinitrival (rig, 4:4) ody anensra Berzins Loviea length <3adocica with, ovoid, circulay, trapezoidal Or rhormboidal,... 2.2.2... 26. 3 Dorsallonica cross-section 1/3 of eirele ta highly- domed hemisphere (Pig. $:2d)......----.-- 6 Dorsal lonica a stiatlow dome, or darsoventrally Wattenud -- é a! Loriva oval to circular; head aperture notehed both Roof und ventrally (Fig, t:3a)....... 4. evalis (Muller) 312). Lori iva 2x ‘wide in posterior third Man anterior, dorsal murgin of head aperture straielit, ventral mitt only weakly y-shaped (fig. @:9b) 4 5 eee oe - W. KOSTE & R. 3(4). (3). 7(6). YS), L(A), HIG), 1211). 13(12). 14(13). 15¢14) 1612), (7d). 107) W177. JUSHIEI Ventral lorica with twer lateral spines directed forward: (ues striuglue (Pig, O82). 000.00 ee, penta corey be tana Koste & Shiel Ventral Jorica Apineles a4 ae curved, ee bee 13 netmlsst Bervins Devipit al maint ol head ape ee protruding (Fig, SOMEED Heenan on a): : _.. Qecipiial margin of head Aperture concuvely noiched (Fin. S4a)......, pecininiciibic! Lopes = 7 Lorica outline elongate oval to cireutay (Fig. 5;7a) Lutica outline rhombaidal (Pig, 3: da}, se 1. vhensalathi Kosic Posterion margin ot Joriva rounded oF trungaie, never pointed (Pig, SFad. vere ee Posrerion maraih of loriga tapers lo a pointed projection extending wes bitse OF roes (hig. 9:7) wilhatis) Kose & Shiel Head! aperture vential sinus with parallel cuticular ribs; dorsally with a koe prantlared collar ellipned Willert Head apeitiise- Wittiout rs or dorsal collar , patella (M*iler) Ventral forica with long pleaty....... trifa Myers Ventral tories “umpleared sda ome ue heath Cross-section with Sie bead- like foteral margink. sc... voceyedy daerplisera (Stencons) Crossesection lateral margins produced lo aqgute en ere it BIC Wee ete sc veatalle ai leletene tn 12 Toot-opening excised dorsally. .-...., 5 ao TES Foot-opening nat excised dorsally... 0.0, iy Extremely wide foot-opening, tonger than wide; corners With ourwardly-cHived points... ._. i4 Foot openine wider dhan long, no points on 57 tt TR a L. henxamow Warring Head ajentuce ventrally with lateral biftreare pointed extensions extending beyond dorsal Margins... 0. -L. cornuti (Koste) Nor balecal poly oo head aperture, ,,, 15 Head aperture with convex dorsal margin last foot seument 1.5; toes 20-25 yrn.. Mistribution, In Tresh and brackish water pools, often sympatric with the nominate species and L, patella similis. Bidrklund (1972) demonstrated blometrically distinguishable populations, however infergrades with 4. patella typ. occur, Only Australian record from Sheepwash Billabong, near Yeu, Vic, 17iv.76(Shiel unpubl.). 19.9°C, pH 7.2, DO 8S mye) , 85 pS em |, Secchi trans- parency RO cm, Giterature: Koste 197i Lepadelta patella similis (ducks) FIG. 74 Metopidia similiv Lucks, 1912. p. WY, Pie, 34, Lenadelta similix (Lucks) afler Hauer, 1925, p. 17 t patella vay, similis atten Remuine, 1924, 9, Fig. VAA-E. “. budngensis Bereiny, 182, p, 1 Me. 9. Diagnosis, Lorca outline ellipucal: taay-openiig with teinforced uppec arch (Fig. 7:4 f, 2}, mustly hexagonal with angled edges and base straight to indentud. Not reliable taxonomic character because oF vanability in foot-opening. Distinguished fram L. guitella (s,.sir,) by lorica lengih:width ratio (> (2) and overall smatler habit. Length 68=85 pm; width 49-60 xm; wes 17-21 pin, Dixiribution: Bewween macrophytes, detrivus, in psamnmon and rock pools; fresh, brackish and salt; pH to 10 (Koste 1978). Single record from Australia (as £. buangensis Berzins), but recognizable as a nreservation artefact of L, patella similis, Mt Donna Buang, Vie Lepadella plilota Berzins FIG. 834 C. pliléta Berzins, 1960, pp. 83-85, Figs 4-6. Diagnosis; Body ovoid in outlines venira) surface slightly medially convex, dorsal surface very strongly 50; dorsal median line with sharp, uneven longitudinal ridge; frontal margin dorsally convex, with blunt projection in the middle; venually with rounded sinus; laterally, lorica enlarged with three pairs of thin, transparent tamellary projections; lorica prolonged posteriorly (to rounded Jobe: crass section somewhat triangular, Length 72~75 wm width 60-67 pm; ventral sinus depth 8 xm; length of foot groove (2-15 ym; toes 12 yarn, Distribution. Endemic. Only known locality Creswick Creek near Clunes, Victoria, Ecology unknown, Comment Resembles J. ¢riprera (Ehrenberg) but differs in (he form of the trantal margin of the Torica, the sharp dorsal ridge, and the presence of lateral cuticular projections (termed "flappers” mn the original description althaugh they have ne muscular contact], Lepadella quadricurinata (Stenroos) FIG, 8:2 Melopidia quadricarinaia Stenroos, 1R98, 5. 165, big. 3-2, Diagnosis: Dorsal lorica with four short ribs over foot-opening; posterior end tapering to short tongue; ventral lorica flat; foot opening variable. Lorica length 81-92 ym; width 61-72 wm; height 1o 34 um; toe length 20-26 jam. Distribution: Littoral, between macrophytes, pr 6.2-10.2 (Kaste 1978), Only record that of Berzins (1982) trom Coghill’s Creek neat Tourello, Vic. /, q. Sexcarinata (Klement), recorded fram the same localily, muy be an ecotype, Literature: Koste 1978, Lepadella yuinquecostata (Lucks) FIG, 8:3 Metopidia quinquecestata Tucks, 1912. p. 126, Pig. 47. Diagnosis: Lorica pyriform, narrowing towards head; dorsal lorica with median low keel which hegins as witely separated ridges behind callar.of head aperture; two or three pairs of side ribs: distinctive cross-seclion (Fig. &:3c): Jorica end variable, sometimes with short ridges foot-opening variahle, Lorica length 92-112 ym; width 60-70 wm; tne lenyth 22-30 yin, Distributions Cosmopolitan in tirural, efter W. KOSTE & R, J. SHIEL 136 ROTIFERA PROM AUSTRALIAN INLAND WATERS ity collected in Sphagnuen. pH 4.$-6.7, temperature to PC (Koste 1978). Single record: dam near Chillagoe, Qld. Lepudelta rhomboides (Gosse} FIG. 84 Metopidivrhamboides Giosse in Hudson & Classe, 1886, p. 18, big. 25:70, Lepudela rhamboides alter Harring, 1913, p. 65. Diagnosis; Dorsal lorica. with moderately high, mare or less arched median keel, bounded by lateral grooves; anterior margin Width 4 lorica length; dorsal sinus broadly U-shaped; ventral sinus ¥- shaped; dorsal aperture of head-opening with collar, whieh can be bordered by cuticular thickening; foot opening narrow, U-shaped with parallel sides: distal foot seament long (cl. L, elfipticah; toes short (<4 body length) slightly decurved, tapering to points. Lorica length }10-120 pm; width 35-88 yam; wwe length 20-28 pm, Disicibution: Cosmopelitan im periphyton, pH Tolerant, Rate: N.S.W., NLT, Tas,, Vie. 10.5-30,7°C, pR 4.7-7,.9, DO 4,6-6.0emg 1 ', 23-750 pS om | to 67 NTU, alkal, 2.7~2.9 me L-'. Comunent: Two variants are kitown From Australia, described elsewhere as ssp, form or var: df. rhamboides hauer{ (Wulfert) (Fig-8:5) from a stack vam south of Burnie, Tas. (168°C, pl 4.7, 42 uS em and L. /homboides caringta (Danner) (Fig. 8:6} from the Uroken R. at Benalla, Vie. (27,0°C, mil 7.6, DO 7.0 mp t-', $2 pS em '), Literature: Kutikova 1970; Koste 1978: Koste er al. IDS; Koste & Shiel 198ha, Lepadella rarrenhursi (Lucks) FIG. Yt Metapidia redtentyied Lacks, (912, p. 127, Pip. 48ay 0, (.rovenburei atler Harring 1913, p, 65: Digenosis: Side margins of lorieu cross-section earend to pointed tips below level of median ventral bulge; head aperture dorso-ventrally deep, only: ventral sinus; dorsal margin straight, without sinus: fool-opening variable. Resembles £. dactylisera, Can be-contused with Colitrella spp. in lateral view, Lorica length 75 am; width 50 pm; toes 25-28 pm, Oistribution: Isolated occurrences in Sphagnum and. peat swamps. Only record Riddel’s Creek, Sunbury Vic. Liferature: Koste 1978, Berzins |82. Lepadella tana Koste & Shiel FIG. 9:2 Lepadedle tara Koste & Shiel, 1987, p. 102, Fig. 19. Diagnosis: Loviva hraadly ovoid, ventral loriea Mat, dorsal micdially conver tn median cross-section: dorsal margin of head-opening almost straight. ventral margin with weak V-shaped sinus: fold behind middle of bedy on ventral surface; with pointed, slightly curved spine on either side, directed apically; foot-opening broad; toes relatively long, straight, sharply pointed, Lorica length 79 xm; width (medially) 61 yam; anterior width 29 am; foot-opening 18xl8 pm; toes 29 wm} fateral spine length 18 am. Distribution; Eudemic. only known lucality L, Pedder, Tasmania. 165°C. pH $.3, 46,2 aS emo. Comment: Resembles L. nedoissi, a Victorian endemie (Fig. 6:31, however |he latter has ventral folds (no spines). and differs in foot and toe morphology, Lepadella triba Myers FIG, 9:3 Lepadella triba Myets, 1934, pp. 4S. Figs 10-12. Diagnosis: Lorica elongated; crostseytion shallow, evenly arched dorsally; dorsal margin of head aperture almost straight, ventrally broadly V- shaped; no stippled collar; ventral lorica with distinctive Euch/anis-like slightly clevated mid- section (Fig. 9:3c), lateral margins run to head Aperture or diverge io side apices; foot opening Maring: distal foot segment approximately 4 toe- length; toes long, tapering to slender drawn-out tips. Lorica length 71-80 am; width 42-45 jim; ventral sinus depth 12 jan; foot opening depth 20-any; foot length 26 wm}; distal segment 14 jam; toes 30-33 pn. Distribution: Noth Ameriga, Single record from Old (Berzins 1982) needs verifieution. Lepadella Sriptera (Ehrenberg) FIG. &:4 Metopitia triptera Ehrenberg, 1830, pp. 74-83, Lepadelta triptcra Ehrenberg, 1832, p: 72, Fig. Rl, Lepedelia ptilota Berzins: (a) dorsal; (h) ventral; (ec) crass section. 2) 2. quadricarinate (Stenroos):; (a) dorsal: (h) ventral: (¢) cross-seetion: (0) dorsal: (ey ventral; (f) dorsals (8) cros#section. 3, 2, guinguecostate (Lacks): (a) veniral; (b) laccrak; (ev) crass-seetion; (d) dorsal. 4, 2. rhermdbeices (Gosse): (a) ventral: (b) lateral: (c) dorsal: (a) cross secuion. $5, 2. rhamboides hauert (Wulterl): (a) SCT. form, dorsal; (b) Luropean form, ventral: (c) utass-secrion. 6, { rhomboides Corina (Dunner): (a) ventrals (b) averal (c) venieal: GH cross section Spee. dh CO} Choss-secuiOn spec, 11) loriea stinfaee morphology. L, alter Berzins (1960); Jaw, 1 Shu, alter Bartos (1959): 2d, fy after Pejler (1982); Te alter Biciblugd (972). 4, 4, ater Harring (916) 3, aller Koso 98th after Donen (943). Seale lings 30.j0i, 138 W. KOSTE & R, J, SHIEL ROTIFERA PROM AUSTRAI LAN INLANIY WATERS {39 Diagrosis; Lorica outline vatiable: circular ta chombic according to Koste (1978). Dorsal lorica vaulted, with high, somewhat protruding median keel; head aperture with shallow U-shaped sinus ventrally, usually slight median notch dorsally: forica surface may be Vinely striped (Fig, 9:41) wirh small founded proirnsions, mast, however, are smueth, Lorica length 58-65 ym: width 30-57 pm, toes 13-22 wm, Oistelution: Cosmopolitan in fresh and brackish Waters, and on muddy litrorals. Rares NSW. NT, Old, Tas, Vio 12.4-20.0°C, H 48-77, )7-3330) Soom '. Literature: Shriel & Koate 1979; Koste (981+ Berzins 1982, Lepodella tyleri Koste & Shil FIG, 9:5 Lepudella tyler] Koste & Shiel, W8Fa, p. 37, Pigs 3, 4. Diogaosis: Rigid loricu of trapesoidal outline; cross- section nearly triangular; dorsal plate with median keel} dorsal and yentral plates finely pranulated; head aperture lightly convex dorsally, deep V-shaped sinus ventrally, with lateral lamella; caudal Jorica with slight constriction aboye blunt lateral corners; fool-opening broadly flared; three foot segments, distal with caudal antenna: toes very thin, elongated, pointed. Total length 123-125 pm; lorica length 78-SD pm; greatest width 58 vm) height 36 pm; head aperture width 21 wm, depth 22 pm; foot opening 18x18 um: distal foot segment 12 4m; toes 32 pm, Distribution: Endemic; only known locality a stock dam south of Swansea, east voast of Tasmania, 134°C, pH 6.0, 390 nS cm ', 82 NTU, Lepudetla vitrea (Shephard) FIG. 9:6 Metopidieavatis Shephard in Anderson & Shephard, TRYD, p. 78, Fig. 66a, Won ML valis (Muller, 1786) M, vitrea Shephard, Wil. p, 53 Lepadella wired alter Harring, L913, p. BS. Diaynosis: Ovoid, tansparent Jorica, taperine to cach end from widest part posterior to midline: head aperture slightly convex dorsally, without sinus, ventrally with deep, broadly elliptical sinus: dorsal margin of foot opening convex between lateral points of slightly outcurved caudal Jprica projections; ventral foot apenine deep s-ellipse; distal foot segment almost twice length of proximal sépments, more than half length of toes, toes short, Tapering to pointed fips. Length 123 pam; width 36 wm, [Na other measurements given in ihe original description, however from Fig. 6 in Anderson & Shephard (1852), on the basis of cheir lengrh/width measurements, the following wre estimated: head aperture 23 jun wide x 20 pm deep; foot openiny 20 am wide x 22 ym deep; distal tue joint 12 pm; tors 17 ym] Distributions Subsequent to the ormeinal Cine LBrizghton, Victoria), L. vitree was recorded from South Westland, N.Z. (Russell 1954), Dimensions were slightly different (14 «m Jong, toes 20 ym), with “a series of dots round the lateral edge of the dorsal plate”. L. virrea is possibly more widespread- Comment 1. vitrea was synonymised with LZ. fu(usinus by later ceviewers (ee. Koste 1978), however can be distinguished from che latter by more elongated lorica (lengthiwidth ratio 2.2 vs <15 in L. fitusinus), absence of cowl-like head aperture, elongated distal foot-segment and markedly shorier 10¢s, Lepadella williams Koste & Shicl FIG, 9:7 fepadetla williarns! Koste & Shiel, 1980, pp. 138-139, Fix 4 Olasnusiss Smoothly oval, highly vaulted lorica, with ventrally-directed head-opening; dorsal lorica tapers caudally to variable-length laterally curved point, foot-opening narrow, ending.at base of dorsal lorica projection; three foot segments af similar length; long, dorso-laterally curved sharp toes, Lorica tength 112-6 pm; width 60-64 pm; height to 56 xm; head uperture 33-36x28-30 pm; caudul lorica projections 19-28 p~m; foot opening 17-20x19-20 jam, distal fant segment 10 jm; toe length 36-40 um, Distributions Kuala Lumpur, Malaysia, and Jabj Iuka, NT. only known localities, 25.5°C, pH 6.2, DO 29 mel |, 62 pS ecm ') 5 NTU, Fie. 4, Lepedella rotienburgi (Lucks): (a) ventnats (b) fatural: (¢) verttrals (el) cross-section 2, £. ane Aoste & Shivl, venifl, 3, 2. frida Myers (a) dorsal; (b) ventral; (c) cross-section. 4, 2. edprera (Etirenberu): (a) dorsal: (b) ventral: {(c) eross-Keurion, (d) cross-sectiom: (¢) lareral: ((-h) different Joriea forms, ventral; Gi) Tas. farm, ventral; (j) 2d ‘Tas, form, wential 5) 4, Wert Koste & Shiel: (a) venenaly (4) dorsal (c) lateral QD) cross-section: 6. L. Witrea (Shephard }: (a) dorsal; (b) ventral. 7, 2. williams? Koste & Shiels (a, b) lwo morphs, ventral: (c) fateral: (d) cross-section. 1, b, afler Caling (939); bo, d, after Bartos (1959), 2, 4), 5, ier Koste & Shret (ORG): J, alter Myers (1934): tit hy, ates Having (1916), 5, ater Rosle A Shiel (9874); Git. orig, 6h, aber Shephard (90s 7, aller Koste Moshe (4880) Seale lines 40 pm, 140 W. KOSTER & R. 1. SHEEL Key to species of the genus Lepudelliz (Heterolepadelia) 1 PR Pec (Gs Le 2) | nnn DO re eat Leariea thombotdal.. 00.00... 2 eee ee Arar) Ze). Head aperture with Veshapead dorsal ancl ventral KIMUSOS, oo sere bh. HY, Aeteredavivia Tadeey Head aperture with only a sential sinus, dorsal Margin stright. ....... L. 7H.) upsicara Myers AA), Larice wilh posteriori pore spurs, loot puoove Wilh lateral pointed piujections... ,, - ; Cod. CAD @hrenberai (Peri) Lateral wings of loriva.arid postenar margins af foot groave rounded... .. L. UY) heteroxtvle Murray poets Lepitdella (Heteralepadella) apsicora Myers ric. We Fepadeia upsicare Myers, 1934, pp. 5, 7, Migs t6-18. Diaenosis Loyica oval; crass section shallow, evenly arched dorsally; dorsal anterior margin nearly straight, ventral margin with deep V-shaped sinus; stippled collar present; foot groove wide, flared posteriorly; discal foot joint 2x length of combined first and second joints; foot twisted, left toe lies beneath rights (oes asymimetrie. right 24 fenuth of left, Lorca lengrh 80 jams width 65 am; ventral sinus depth 18 prs anterior points 28 pm. foot groove Zt pms foor 30 pny distal segment 20 yr; right foe 30 pms lett yam. Oistethution: North America, Two records from Attstrahia: L. Boort, central Victoria, and Winmurra Billabong, Magela Creek, NUT. 23.5°C, pH 3.4-7.9,, DO 60-6.2 me | 1, 23-750 pS cm alkal, 2.7 mel '. hiterufure: Koste & Shiel 1980; Koste 1981. » 67 NTU, LL. ¢H). ehrenhergi (Perty) PVG, 10:2 Niutawania ehrenhersi Perty, 1850, p. 20. Moerapidia clhrenbergi otter Jennings, 1894, py 26. Lepudelia ehrenberge ater Harring, (913, p. 63 Lvagnosis: Lotica rhombic, with posterolateral margins produced to triangular, dorsully ridged spurs reflexed upwards in cross-section (Fig. 10:2¢); smaller painted spurs on each side of foat granve; stippled collar on ventral plate, with less obvious Jornal stippling; foo. groove Vi lenegrh of lorica, rounded anteriorly, flaring slightly posteriorly; foot stout, 3 length of lorica; distal foot joint longest; toes long, asymmetric, laper evenly, Loriva tength 70-95 pin; with 70-90 pn; toe length 27-32 wm and |9-27 yon. Divtribution: Cosmopolitan in periphyton, Orie laria and moss. Rare; in billabongs and vegetated lake margins in N.SW,, NTL, Qld, Vic. 24.0-29.2°C, pH 6.3-7.5, DO 5.1-8.0 mg 1-', 42-85 pS em |, 6 NTU. Literature: Shiel & Koste 1979; Koste & Shiel 1980; Greeiy 1981; Koste 1981. L. (H.) heterodactvia Fadeev FIG, 10;3 L. (F.) heterodacivla Fadeey, 1925, p. 73, Phot, big. 8, Diagnosis; Lorica oval; dorsal and ventral mareins of head aperture with deep, almost triangular sinuses; last foot segment longer than preceding segments. Lorica length 106-113 pm; width 70-75 pro; distal foot segment 22~23 pm; toes 33-30 and 20-25 jum. Distribution: Rurope, S. Ameriva (Amazonia), Single record, L. Boort, central Vie. 20.0 °C, pH 7.4, DO 8.0 ing Io! Literature: Kutikova 1970; Kaste 1978: Koste & Shicl 1980, 1, (fh) heterustvia (Minrray) IG, 10.4 Metopidia heterostvla Murray, (913, p. 489, Ph19, Pig. 6a-c. Diagnosis: Lorica rhombic; lateral wings rounded rather than pointed (cf. L, ekrenbergi), with tips more or less deflected dorsally; ventral lorica wider than dorsal in region of head sperture; stippled collar present. Distribution: Probably cosmopolitan in littoral zones, stagnant waters, Rare: N.SW., Tas., Vic. 10.0-23.5°C, pH 5.8-7.9, DO 40-98 mg 1°, 80-750 pS cm |, 2-67 NTU. Literature: Koste 1978; Shiel & Koste 1979: Koste & Shiel Lgo. Not revordéd from Australia: L..(H.) eyrtopus (Harring), Eastern Europe, N. and Central America. A single free-living species of the subgenus Lepadella (XNenalepadella) is tecorded from Australia. In view of the preferred habitat ol most oi the other known species of the genus, ie. the branchial chambers of a range of crustaceans, it is likely chat orher species oceur here but have heen overlooked. Lepudelle (Nenolepadella) mondactyla Berzins FIG. 10:5 Lepadolla monodactyla Reveins, YU, pp. 5,6, Fizs B=, ROTIPERA FROM AUSTRALIAN INLAND WATERS 4] Fig. 10:1, Heterolepadella upsicura Myers: (a) dorsal: (b) ventral; (e) cross-section; 2, 4. chrenberei (Perty): (a) dorsal: (b) ventral; (c) cross-section; (d) lateral, 3, Af. heterodactyla Fadeev: (a) ventral; (b) toe detail. 4, /1. heterost via Murray: (a) ventral; (b) dorsal; (¢) lateral; (d, e) different cross-seclions: 3, Xenvlepidella rronodaciyla Berzins: (a) dorsal; (b) ventral; (¢) cross-section, |, 2, aller Harring (1916); 3, after Kudkova (1970); 4, after Harring (1916); 5, after Berzins (1960), Scale lines 50 pm, Diagnosis: Lorica outline elliptical; dorsal lorica arched, punctate or lined; ventral plate with light bulge medially, otherwise unstructured; head aperture with V-shaped dorsal and ventral sinuses. Total length 170 um; lorica length 127 wm; width 77 wm; foot-opening 31x10 um; toe 57 pm. Distribution: Madagascar, variants known from Brazil. Single record, billabong, Jabiluka, NT. 25.5°C, pH 6.2, DO 2.9 mg 1-!, 62 zS cm !. Literature: Koste 1978; Koste & Shiel 1980, Not recorded from Australia: L. (X.) astacicola Harring, L. (X.) borealis Harring, L, (X.) branchicola Haucr, L. (X.) haueri Rodewald, (.. (X.) lata Wisniewski, L. (X.) parasitica Hauer, £. (X.) pygmaea (Gosse). See Koste 1978:198-200 for detuils). Acknowledgments Collectors acknowledged in our earlier Papers also contributed further material to this work, Their assistance is appreciated. The Deutshcen For- schungsgemeinschaft, Bonn-Bad Godesburg, provided microscope and photographic facilities to WK. i2 WoRKOSTR & RoI, SHIP References AvbetROM, 1H (HY S8) Alankrom Rotutona tron Nerrlt Cawlmoa, J tlishe Murchell Ser Soe, 54, KR-LD Anoiaes, FL (957) bauuistisch-okdogmch Studien tn Romiorien saltzinitiiwe Gewusser Mitteldeutschlands, His. 2. Martin-furhertiniys Halle Witheabers erath neturwiss, 6, L738. Aworrsem, Ho fP & Stipiagn. 2 (1892) Notes an Viclorian routers, Prac kh. Soe. Mit. 4d, 9-80. Ragios, ©. (955) Viner rode hepartediiow Wick gest li unwovuni, Crea Naren, as, Ode, pitrnd. th. f2-44_ (1959) Vermeil — Roraroria, aura O SR. 45, 1-464. Bekvins, KR (4960) Neue Rotirorien aus Madwwaskar. Men [insh Sei. Madan. sé A, 14, 1-6. — (982) Contribution lo lhe kiowlgede¢ of Rorararia of Austaila, (Universiny oF Lund, Launscdl) Hiorntuwo, BG, (1973) Phe rorifer taunt of the rock Mooly inthe Tvarminne Archipelago, sourhern Makan, Aetna Zool bern, 135, 1-A0. Bowy pe Sy Vincest, IB oG. MLM. (IR24) ney. Merb, 2. Oazer "hssar dune classifigulion sles aonmanys Mivroscapiyucs’ (Pars) Cyrevik, (19399) Ober Rotutorien voter Seen. bei Aueboda, Weld. Fiemeal. instr. Littds baty 2, 3-68, Canine sos, B. (1999) her die diniden fur Sehweden nene Rotutorien, irk, Zec/, 264, I-14, Corus, A. Pb (1838) Nouvelles Observations imiroserpiques. Aloweduch de Corlsdad A. 179-198. 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Soe, Old. 4, 28-30. Weber, FP. (1898) Faune rotatorienne du bassin de Leman. Rev. Suisse zoo! 5, 263-785, WiiteLecce, T. (1889) List of the marine and fresh- walor invertebrate [auna of Port Jackson and the neighbourhood, Proc, R. Soc. N.S.H2 23, 163-323. WisaNitwski, (1983) Fauna wrotkow Polski i reyonow pravieglych, Pol. Arch. Hydrobial 4, 317-490, WULFERT, K, (1939) Beitrage zur Kenninis der Rader- Uerfauna Deutschlands. 1V. Die Radertiere der Saale- Elster Niederung bei Mersburg in Okologisch- faunisiischer Reziehung. Arch. Hydroabial. 35, 563-624, —_—— (1950) Radertiere. Naiurschutzgehiet auf dem Glatzer Schneeberg. /hid 44, 441-471, (1960) Die Raderticre saurer Gewasser der Diibener Heide. [, Die Rotatorien des Zadlitzmoors und des Wildenhainer Bruchs. 7hbid 56, 261-258, PRELIMINARY INVESTIGATIONS OF BEACH RIDGE PROGRADATION ON EYRE PENINSULA AND KANGAROO ISLAND BY A. D. SHORT*, R. C. BUCKLEY**, & D. G. FOTHERINGHAMT Summary Field investigations along 2150 kin of the South Australian coast including the Great Australian Bight, Eyre Peninsula and Kangaroo Island revealed numerous sites with low wave energy beach ridge and occasional chenier plains. Examination of 14 mainland sites and six sites on Kangaroo Island indicate that all ridges are wash deposited during episodic wave events, probably over a period of decades to centuries. The ridges overly intertidal shelly sandflats. The oldest innermost tidal flats date at 6.3 ka and 5.9 ka BP, but the oldest ridge dated at 4 ka BP suggests a period of tidal flat formation and progradation is required before a source of sediment and plat form for ridge formation is available. All ridges and ridge sequences dated between 4 and 2 ka BP with no ridges material younger than 1.8 ka BP. While this confirms a delayed start for ridge development it does not necessarily imply no present ridge building as reworked, older shells may be incorporated in the modern ridges. The age and elevation of the sites lend support to a published sea-level curve for northern Spencer Gulf. The limited evidence available suggests that sea-level reached its present level in the Bight by 6 ka BP with no conclusive evidence for changes since. KEY WORDS: Beach ridges, sea-level, Eyre Peninsula, Kangaroo Island. Hansuctions af the Raval Suctety of 5. odast, (1989), 1304) 144-16). PRELIMINARY INVESTIGATIONS OF BEACH RIDGE PROGRADATION ON FYRE PENINSULA AND KANGAROO ISLAND bv A, D. SHOR’, RoC. Buckiey**, & D, G. FOTHERINGHAM/ Sutnmary SHORE. A. DL, Boehtbh, ROCL & PorirkingHam, DOC. (1989) Preliminary investigations of heaeh ridge mogradauion on Eyre Peninsula and Kangaroo Island. 7rans. R. Soc. §. Aust. 113(3), 145-161, 30 Novernber, 1989, Meld nywstwutons dong 2150 km al the South Austrilian coast indudiiy the Great inant Breht, Eyre Peninsiute apt Kangitrow [stand revealed mumeratts sites with los wave encres beach ridge and pecasiioal ehemer pliitis EXAM Aron Ol 14 mainland sites aad sis sites on Kangaroo tykind indiguhe hat all mages Ones wiih deposited during episadic wave events, probably over a period of devades Wy contunies Chie rides overly inferiehil shelly samdflives, the oldest innermost sida) (has dare at 6.4 ku ane 29 ba WPS bur the midest Hidhve daledal ka BP stiepests a pertod of tidal tid fornition and progradariny 1 tequired before weenie OF scdiment and phiitform for ridge formarion is available, AN rilges ain ride scquences died belween 4 and 2 ke BP with nev ridves manernal younger clan 8 ka BP, While this confirms a delayed stint Ton fidve development itdoss, notnevessarily triply no present ridge buildin as reworked, older shells Iidy he ineorpardied ithe modern ridges, The age and clevarion ob ute sites tend support toa published sda-hvel burve lor dortherd Speaver Gull) The limited evideriee availible sugeests chat sea-level roagheal is present level in the Bishi by 6 ka BP with no conelusive evidenee tor changes since. Kru Words: Beach ritlyes, sea-level, Byre Peotnoula, Kangaroo [ylud Introduction The South Australian coast, west of Whyalla and including Kangaroo Island and the 7& islands of the Eyre Peninsula, represents 60%; of the state's coasting. Phe cous! contains a variety of Flalacene Shoreline types ranging from. tide dominated. saudflats fo, Will iereasing eave energy, cheniers, beach ridges ind regressive (lorechine ridges) iid transgressive barrier-dune svsiems (Short f988a). The Jatter include some of Australian's ogost extensive voustal dune systems, including more than S00 kin of clifftogy dunes (Shore t988b). Whilst the 3700 kis, Aust. coast possesses a wide range of Holovene coastal depositional systenis, The nutune and evolidon of rhese systents received Only cursory attention unit Chis decade. Revional studies und detailed invesdauion have only just begun to address (ie status of Lhe couse, In the cull regions Hologene sedimentation and Pluistovcne and Holocene sea-level history wxsocraicd wath beach tides plains huve been investivaled by Burne (1982), Belperio et at (1983), Hails & Ciostin (1984) and Belperioa er af (988). fails & Ciostin (984) and colleagues also presented aseries of papers on the evolution af northern Speneer Gull and its implications for modern sedimentation. In St Vincent Gulf, Bowman & Harvey (1986) conducred extensive coring and = (ous) Studies Unit, Dental Cremgniphy, University af Sydney, Sydiwwy, SSW. 200A. sOAMPEL. 162% Miller St, O'Conmmen, ACT 2600, 1 Coast Managenrtent Brareh. DA. Oo Box 667, Adtvlnide, So Aust S00), cating of the Le Fevre Peninsula to reconstract the evolution of Wiis imipertant beacheridge cand plex. On the open coast three revenl regional stuaies of the Seuth East, Kangaroo Island and Eye Peninsula (Short & Hesp 1984; Shan & Borhenmyehainn 1986; Short er al, 1846) provide the first comprehensive assessment of the morpho dynamtivs and Holocene evolution of these seerions of coasiline. The aii of this paper is Co assess We evojution of the low wave energy beach ridge and chenier sites on Kangaroo Island and Eyre end their implications for mid to late Holovene sen- fevel in the region, Beueh vidges are tere defined its Shore linear, low sundysstelly swash deposits, separated by wider iitet-and suprasGdal sane Mats. They are a product of periodic wave acon and showld nor be contused with aeolian foredune ridves, Wineh have also been called beach ridges (see Hesp 1954). Cheniers have similar ridees but are vader lain aid separated by finer sediments such as minds, silts or fine muddy srids, Reach ridges occupy only 4°) (66-km) of Evre Peninsula and (44.1 30-kmn of Kangacoe stand. This is Tt sucprising given the generally high wave enerey together with high percentage Of racky east (66 and 4") respectively), On Fyre Penttisulie beach ridges ate common in thy ventral gull and oeeur it several protveted bays on the Bivhe coast, while on Kangaroo Island they ure restricied do the proiccted north eastern caves (see Short et al JORG and Short & Fotheringham 1986 for locwtion of all stiles). These sttes are, however, very importint for the mierpretation of hor Hotovene shoreline evolution and sea-level history. parnenierly gevern M46 A, D. SIIORT, R. C. BUCKLEY & 1 1G, FOTAHERINGHAM the noise-or ubsence (due io erosion) of the more duminant higher energy beach and dune deposits, Bach Burne (1982) and Belperio eral. (1983, 8k) use such deposits (0 interpret Holocene sea-level history, particularly invorthern Spencer Gull, while Burne (1982) and Bowman & Harvey (1986) identified episodes of ridge building in northern Spencer and Sc Vincent pulls. respectively, This paper provides a preliminary assessment of the initiation and timing of ridge building at 2) sites together with some implications for mid to late Holocene sea-level in these regions, Study Area All beach cridge-chenier sites were identified by mapping the entire coastal geomorpholoey using 1:10,000 calour aerial photography. Twenty representative sites were selected for field investigations (Pig. 1). They represent about 300 al both regions beach ridge systems. Table 1 lists environmental parameters of each site, Field investigations involved surveying ul cross ridge tratscety using a theodolite, and bath surface and shallow (<1 m) subsurface sampling, the latter using both shovel and hand auger. All samples were collected within 10-cny Stratigreplic unin usually from in situ tidal flat deposits, All shells were collected within cach layer, For dating, only the mosi intact valve or bivalve shells were collected assuming that the more intact and fresh the appearance, the less distance they have been transported and therefore (he younger they ruight be, thereby providing a more accurate Mininium age of the deposit. ‘Lonless otherwise indicated, dated shells were obtained trom the base of the anger-shovel hole The locdhion of each sample is Tllustrated in the accompanying figures. All sites are chacavterised hy low breaker wave corditions, with essentially zero modal occan wave effects, Wave energy is delivered bath by greatly attenvated (<1 im) extreme icean waves and, more lreyuently, by local wind waves ( cssia Tidal Flats 2120'80 ——S= Site 3: The knob Erasion Scarp Po 5. Tulka Wells a WKS37 2900760 b.CS631 3510+ 80 cWKS36 2420160 100 200 DISTANCE (meters) 100 200 300 400 Fig. 3, Spencer Gulf field sites, Chronological ridge numbers indicated above some ridges. Mount Dutton Bay. On the north-western side of Mount Dutton Bay (site 9, Fig. 1) is a number of very low energy low beach ridge sites, One of these was surveyed and sectioned (Fig. 4e). The innermost sample (WK 833) dated at >40,000 yr BP indicating Pleistocene material. A sample obtained from tidal flat deposits below the base of the innermost ridge yielded a date of 6330 4 90 yr BP (CS 530), the oldest Holocene date recorded in this study. This date confirms the arrival of the Holocene sea-level stillstand by at least 6.3 ka BP, in agreement with its arrival in Spencer Gulf (Belperio e/ a/. 1984) and Australia in general (Hop!ey & Thom 1983). This date also is supported by the age of the first swale which dated at 4590 + 80 yr BP, the third oldest date in this study. The elevation of the samphire — beach ridge, and tidal flat — samphire contact between ridge | and 2 (Fig. 4e) suggests a slight (~50 cm) fall in relative sea- level. The timing of this apparent fall however cannot be determinded from the existing data. Streaky Bay Perlubie Beach (south) Perlubie Beach (site 10, Fig. 1) is located in south- east Streaky Bay and consists of a 1 km wide plain of Holocene beach and foredune ridges (Fig. 5a), A single date was obtained from the shell-rich swash zone immediately behind the innermost Holocene ridge (CS 499). It dated at 3290 + 90 yr BP, again Suggesting a time gap between the stillstand and the onset of low energy beach ridge progradation. Acraman Creek The Acraman Creek (site 11, Fig. 1) region in northern Streaky Bay consists of a 10 km section of shoreline dominated by multiple episodes of recurved spit development and bayward beach ridge progradation (Fig. Sb). The shoreline at its maximum has prograded up to 3 km seaward. One date was obtained from the inner portion of the back barrier depression. Whole shell samples (CS BEACH RIDGE PROGRADATION 1st TABLE 2.. Radiocarbon Dates Sample Lab. Location CSIRO Environment Material Elevation Age BP Corrected Age No, No. km (m MSLW) Yrs BP = la Yrs BP i da a. Eyre Peninsula See Spencer Guil j-2 cS 50 Glensea 157¥ Back Swale shell 3 3020 = 280 zsali = 280 3-6 CS dt Gilensea 1579 Beach Ridge shell 23 3580 = 80 220 + 40 4-| CS 324 Glensea 1579 Beach Ridge shell 3 250 + BO 2070 4 80 4-4 CS 525 Glensca 1579 Beach Ridge shell | 2470 + 80 2010 = 80 4-5 WK 826 Glensea 1579 Swale shell 2 mos 30 B30 4 50 a7 CS $26 Point Germein 1628 Beach Ridge shell 3 Fasu - 80 JI 2 BO 4-50 CS 531 Tulka Well 18560 Tidal Flats shell | 3970) + RO 3510 4 a0 AY. WK 837 ~~ Tulka Well 1856 = Tidal Flats shell 16 M60 4 AO 20) ~ 60 4-48 WK 836 = Tulka Well le56 Tidal Flats shell 12 ZAR) + 60 2420 + 60 Coffin Way 4-25 WE R28 Gate Waterhole 2089) inter Barrier shell ~| 4150 + 190 4690 4 190 4-24 Wh 827) Morgans Landing 2089 Back Swamp shell = 3000 = 60 2540 + 40 4-23 Cs 427 Morguns Landing 2089 BReavh Calearenite — shell inter lidal 3140 4 Bt MR + 80 4-26 WK §29 0 Salt Warerhale- 214 Back Swamp shell ~1 3000 + Al) W404 60 4-27 WR #300 (Collin Bay) 214 0 Swast Deposit shell ~2 4390 + 60 3930 + €D 4-28 Ch 528 (Coltin Bay) 214 -Swash Deposir shell -~2 3650 + SU Win + BO 4-37 CS §29 Yargzie Island 2193 Inier Tidal shell 07 3240 4 80 2780 = 80) 4-38 WK 832 -Yangic Island 2133 Inter Tidal shell ~08 2540) + 60 TORO = AN) 4-34 WE 8330 Mt Dutiog Hay 272 Unter Tidal shell 17 40,000 > 40,000 4-40 CS 330 Mr Dutton Bay 2172 Inter Tidal shell 15 679 - 90 6330 + 90 4-41 WK 834 ML Dutton Bay 2172 Inter Tidal shell 13 05 + 80 459) 4 ORO Gteat Austratian Bight 3-103 CS 499 Perlutic (south) 2544 Inet Ueach Ridps shell tt 3750 4 0 324) a 3-54 CS us Avantman Cresk 2543) Innermast Swale shell ? 610 4 9G S630 + 96) re ee ail f : ; _ ayy + 3000 yy yoy + F000 3-88 CS 469 Point Collinson 2611 Shell Rock shell 1-45 42,400 arin 42,300” sn00 ne athe : » AF0G + 1700 —* ¢ By 5 , . =) < a dV 4 4 3-4) CS 469 Point Collinson 2611 Shell Rock shell 1-1.4 17,500 1400 ARAM) © i400 3-10) CS 509 Evie Istarid 2033 Inter Tidal shell ~1 T8604 80 140) b RD 3-71 CS 508 Eyre Island 2033 Inter Vidal shell ~1 3300 = 90 2kd0 4 YO 3-71 CS 506 Fyre Lslanel 2635) Inter Vidal shell ~1 3680 + 4 3220) + 9 3-73 CS 308 Fyre Island 2635 Inter Tidal shell -) 3770 = 160 VO + 160 4-24 WK SIS Laura Bay 2683 Beach Ridge shell -l 6l4@ - 90 S6k + 90 4-45 CS S04 Latira Bay 2683 Beach Ridec shell 2 740 4 90 2280 4 00 3-46 WR A24 Laura Bay 2683 Beach Ridge shell 23 2340 1 70 WW» 3-07 C§ 302 laura Bay 2683 Beach Ridge shetl 2 3520 4 91) 260 + 90 3-98 CS 50) Laura Bay 2683 Reach Ridge shell 3.5 2760 + 90 2 + 90 3-99 Wh s25 Laura Bay 2683) Beach Ridge shell 24 3480 = 60 3020 + a 3100 CS SQ) Laura Biv 2681 Swale shell zs 4 - WA, YO 3-3) Wh 816 fourville Bay 2739 Bench Ridge shell Z 2050 | il ZL) + A) 4-34 WR RIT Davenport Creek 277h Chenier Ridge shell =2 2470 + © 90) 201 = Yd 4-35 WK RIB Davenport Creek 277 ~=Chenier Base mangrove ~I 2860 4 SO 2860 4 50 3-36 WK 819 Davenport Creek 2776 Chenier Base mangrove ~1 2370 4 Sal 2571 + 50 3-07 WK AIO | Davenport Creek 2776 Chenier Base nud ~1 2540. 5h 2340 + 30 3-38 Wk 821 Davenpori Creek 2776 = Intertidal mangrove =~) 269) » 50 260). FD b, Kangaroo Island nn een ee kl 13 WK 839) Mutrion Beach 42 Reach Ridge Suale — shell mn S40 — 70 ask = Th KL Su Wh K42 Cyenet River a She!l Ridge shell 25-3 30. AU 2A + AD KL SI Wk 84] Cyenet River dM Shell Ridge shell 254 2750 + 90 D2 4 MI a 1 ~ unditates uearveved, exiiete Only 2 Colrecvon Paton ef 466 years (sed, sue Henerera a Hlarvee (1983), 152 A DOSHORT, B,C. BUCKLEY & BD. G TOTTERINGHANM 503) gave uw dale ol! $850 ~ 90 vr BP the second vldest a! the Holocene dates, This alsa supports fheampirom age ot the Holocene stillstand in ibis ienion a >6 ke BP. Fyre dstunil Fyre Island (site 12. Fig, 1). whieh partially blocks Smoky Biv isthe only unconsolidated inland inthe Right. teconsists of wamoltiple series Of onlapping recurved spits. beagh ridges and low foredunes whieh surround a core of samphire = mioigrove Hats (Pig. 4). Ao transect was made across. [he westernmost and oldest of the existing ridges (Fig, Se). Phe date obtamed trom the three swiles and behind the present beach indicate that Chis scetion of the ishind bexan building around 3.8 ka BPoand progracded to around L4 ku BP. This sechion is presently crogding as sedimerms are reworked eastward alone both sides ol the land fo progride Iherecupved spit complexes, As the island may have been reworked uastward prot to tka BP the dates can only be used as a pide (o (he ape of the oldest section of the existing istand, and donot nevessarily indigate the commencement of iskted formation laura Bay Laura Bay (site 13, Tie. Dis a 2 kin wide bay within Deeres Bay, Locontains a 1.2 km wide beach ridge plain frontal by several huadred metre wide intertidal sand flats, The bay receives easentially no Dcean wave energy us attested by the growth of mangroves on irs northern sand(lats. A senies of Seven dates obtained from two swales and the base of five ridges (Fig. Sd) suggests that progradation commenced alter 4.0) ka BP (CS 500). A period of slay progradation (100 m between 4-3 ka BP, ridges | and 2), was foliowed by rapid progradation (S00 m) between 3-2 ka BP (Fig, 5d), All ridges ure within the reach of present storm swash elevations und do nol suggest any apparent change in sca-level. Like the Glensea ridges, the 2.3 ka age for the outer active ridge (6) may indicate age contamination by older shells, and hence i younger ape may be appropriate, loweville Bas Taurville Buy (Site M4, Big. )) isthe westernmat enclosed bay in 8. Ausi. and contains the wesleromiost mayigreve woodland am South and southern Australia. A tora) of five dates were obtained in the bay, lour lor Dayenport-Creck andl ove from the northern side of ihe bay (Fie. Se). Davenport Creck consists of a 10 kim’ memgrove woodlind described by Butler eral C977) within whiel He rhree disvaminuoas cheniens (Eig, 7). The second of the three cheniers was seetiondd and provided j sequence ob Three dates (WOR 820, 818, 819, Fig, 30), These dates confirm the enrplavenient of this Second ehemer at between 2.5 and 2.0) ka BP, and the development ial the underlying Heliatl flats by 2.5 ka BP. Mangrove stumps on the south side ol! Davenport Creck, presently being Huried by dune transgression datecl af 2690) 4) 30 (Wh R21), conteniporanous with the chenier hase mangroves, The date on the nerthetn shore of Tourville Bay was obtained fram the intertidal hase of the imnermest beachoaidge (Wik $16). Its dare of 2190 © OO Ve BP coincides with the development of the secund chenier. these dates are confirmed by the rysulls oF Belpeno eat (988) who present the results af 13 vibrocores across Tourvillé Bay, They lound the basal Posidome seagrass facies bevan accutane about 6800 vr BP, with the embayment taracty infilled by 5000 vr BP Bench ridge formation followed the mantling of (he seagrass banks by intertidal and mangrove factes. Phere uinerniost arid single teach ridge date of 1840 yr BP is in aurcement with our furdings. Kangaroo Island Mostof the Kangaroo Island coast is (Wypitied by high bedrock aod calcarendte city exposed to tush wave and wind conditions, However the north east leeward pare of che island contains rhree embayments — the Buy of Shoals, Western and Eastern Coves which offer varying degrees of low cuerty Holocene sforeline progeddation, Westert Cove bas experienced che most sobstanital progradation with sediment beinw delivered Alonyshore from ityeasiern und western arm, as well as from the Cvener River and (4 sity depasits, Four aves were Caamined around the shoreline (rigs 1, &). Brownlow A series of live. low, shelly, swash-deposiled bench ridges was surveyed at Browlaw (site 15, Lig, 1; fig. Ba). Their uniform elevation suggests no aupparcnl chive to seadevel Na dithes aye been oblained, srownlow wolf course Sever low shelly ridges were Survevedt immediately south of the Brownlow golfoourse (silt 10. Fig. 1 and avain their relatively uniform elevation gives no incication af sea-level change (Pig. Sb), Cyenen River Aseres of low discouttueus beach ridge and possibly eheriers were surveyed | km south ol phe Cyonet Riverimourh (sive (7, Fiz. (i. The two ridges A, D. SHORT, R. C. BUCKLFY & 1, G. FOTHERINGHAM 154 Site 7: Salt Waterhole Site 6:Margans Landing Holocene Beach 7 \. Coffin Bay. | Ridges ge hae dress wae? i .< . Coffin Bay rier i Sand Flats ATS ; Samsnire, Bieta a.WK828 3690-190 0 tkm b.WK827 2540-60 c.CSS27 2680+ 80 d.WK829 3440 “60 &.WK830 3930-60 1.CS528 3190-80 Site 8: Yangie Island (m) ee Tidal \eo 8 ye Channehy 2 transect g.CS529 2780:80 h.WKB32 2080°60 pe ia lly Sand 7 5 ae DISTANCE (m) | WK833 -40,000 |. CS530 6330-30 k WK 834 4550 ° BO = jis Or L] o> F ee, ae [Calesrentte pe te ge ELEVATION (ISLW) nN ! . Ae 20 40 ~ 60 80 100 DISTANGE (m) Fig. 4. Coffin Bay field sites, Sites 6, 7 and 8 were not surveved. Sketches based on 1:10,000 aerial photographs. 154 A. D. SHORT, R. C, BUCKLEY & D, G. FOTHFRINGHAM Site 10: Perlubie Beach Streaky Bay Pigfuce Is- Holocene Ridges a G3499 3290:'90 Site T2: Eyre Island ©. 68505 3310160 d.CS506 3220°90 8.68508 2840-90 E5509 }400180 4 Z 200 400 * DISTANCE (m) \\ Mangroves 6 pay ( = ~~ oO > z g = < > [3 am wW Southern PiJaines he Ocean ikm| Site Td: Acraman Creek b CH50393 5650: 90 Mangroves Se x) Acraman Ck =, Beach Ridges “Su )/ a PL. Lindsay Streaky Bay Site 13: Laura Bay 060-80 770'70 OSS00SN Oe mutA 0 Shelly Sand £ Calcarenite 4 L L z00 400 1000 1200 1400 400 300 DISTANCE (m) Site 14: Davenport Creek Chenier Section Organic Shelly Sand Manygrave Facies nWKE16 2190 "60 owWK B17 2010'90 bp WKET8 28860.°50 qg-WK81S 2570 °50 1WKS820 2540'50 B.WRKB21 2690'50 Fig. & Western Eyre Peninsula field sites. Sites 10, {1 ard 14 not surveyed. Sketches based on 1:10,000 aerial photographs. ~ Site 12 survey elevations suspect, distances correct. Chronological heach ridve numbers given for sites 12 and 13. yielded dates of 2650 . G0 and 2290) + 50 yr RP (WK 840. WK 841). The ridges are now frotited by 300 m of densely vegetated samphire flats.(Fig, 8c). Their distance from the present shorefine and the elevation of the samphire flats could have resulted Irom a slight fall in sea-level subsequent ta their formation. However this cannot be confirmed from the present data. Morrison Beach Two sites were examined at Morrison Beach (sites Tk and 19, Fig. , Ina waterhole al the eastern end of the heach (Site 18) the elevation of intertidal shingle exposed in the waterhole is ~0.5 or higher than the present beach shingle (Fig. 8d), and could be attcibuted to a slighr fall in sea-level, However the clevation of the inner shingle lies within the reach of storm swash activity. The beach-foredune ridge plain at Morrison beach comsists of a 2-3 km wide plain that hag prograded northward and westward into the bay (Fig. 1; Mocris 1976). Shells abtained from a swale hehind one of the innermost ridges (site 19) (WK 839) dated at 4480 + 70 yr BP, indivating (hat BEACH RIDGE PROGRADATION 155 Fig. 6. A view looking eastward along the Smoky Bay side of Eyre Island. The subtidal sand-flats are backed by a low energy reflective beach, and multiple recurved spits/beach ridges with samphire flats and mangroves composing the core of the island. Fig. 7. A view southeast across Tourville Bay shows the Davenport Creek tidal channel and sandflats backed by mangrove woodlands and the discontinuous chenier ridges (C). Active dune transgression (D) in the south is burying the mangroves. 156 A. D. SHORT, R. C. BUCKLEY & D. G. FOTHERINGHAM TABLE 3. Beach ridge molluscs in approximate order of abundance a, Intertidal mollusc species commonly found in Eyre Peninsula Holocene beach ridge and swale deposits. All these species are associated with low energy mud/sand flat situations, generally with associated Zostera beds or silted rocks etc... Coxiella sp. is usually associated with salt pans or salty lakes. Bembicium melanostoma and Salinator fragilis are found on mud flats at the upper intertidal zone in sheltered estuarine areas. Anadara trapezia is usually associated with Zostera beds; it is extinct in S. Aust. b, Rocky shoreline species (also found in Eyre Pensinula beach ridges). c, Kangaroo Island beach ridge molluscs, primarily in Western Cove region. a Katelysia peroni (Lamarack) Katelysia scalarina (Lamarck) Amesodesma erycinaea (Lamarck) Amesodema cuneata (Lamarck) Spisula trigonella (Lamarck) Anadara trapezia (Deshayes) Brachidontes erosus (Lamarck) Batillariella estuarina (Tate) Zeacumantus diemenensis (Quoy Gaimard) Salinator fragilis (Lamarck) b Tellina (Macomona) deltoidalis (Lamarck) Trichomya hirsuta (Lamarck) Cominella eburnea (Reeve) Austrococholea constricta (Lamarck) Phasianotrochus rutilus (A. Adams) Sanguinolaria biradiata (Wood) Mactra Pura (Deshayes) Fulvia tenuicostata (Lamarck) Ostrea angasi (Sowerby) Bembicium cf, auratum (Quoy & Gaimard) c Katelysia scalarina (Lamarck) Katelysia peroni (Lamarck) Katelysia rhytophora (Lamarck) Zeacumanties diemenensis (Q & G) Mesodesma cuneata (Lamarck) Fulvia tenuicostata (Lamarck) Austrocochlea concamerata (Wood) Nassariu pauperata (Lamarck) Pisulatrigonella (Lamarck) Batillaria estuarina (Tate) Thalotia conica (Gray) Prothalotia sp. Bembicium meeanostoma (Gmelin) Coxiella sp. Phasianella australia (Gmelin) “Bulla botanica” Hedley Diala sp. initiation of this substantial and continuing shoreline progradation commenced by 4.5 ka BP. Shore Platform Evidence Shore platforms cut into Pleistocene beach and dune calcarenite are widely distributed around the western Eyre Peninsula and Kangaroo Island open coast. Whilst shore platforms are highly questionable indicators of sea-level (Twidale e¢ al, 1977; Hopley & Thom 1983) they do warrant brief attention in regard to possible changes in Holocene sea-level in the Bight and Kangaroo Island. Three platforms in the Bight were examined for evidence of Pleistocene sea-level (Buckley et al. 1987). Two of these platforms contained a well defined upper, inactive Pleistocene platform, fronted by a wide active intertidal Holocene platform (Merdayerrah and Clare Bay; Fig. 1). Subsequent low altitude aerial reconnaissance of the entire Eyre Peninsula and Bight by ADS and DGF revealed the existence of relict elevated platforms, presumably of Pleistocene age along much of the calcarenite coast. Active horizontal shore platforms occupy 440 km of the Eyre Peninsula coast (Short et al. 1986) and are usually located intertidally close to mean sea-level. All western Eyre Peninsula platforms examined possessed a single active Nerita atramentose (Reeve) Brachydontes erosus (Lamarck) Tellina (Macomona) deltoidalis (Lamarck) Cominella lineolata (Lamarck) Cominella eburnea (Reeve) Austrocochlea concamerata (Wood) Salinator sp. Diala sp. Bittium sp. platform surface. On Kangaroo Island, however, Short & Fotheringham (1986) describe a Pleistocene platform at Pennington Bay (site 20, Fig. 1 fronted by two Holocene platforms at different elevations (Fig. 8e). The upper platform (No. 1, Fig. 8e) lies 50 cm above the lower platform (No. 2). The difference in elevation and abrupt drop from platform 1 to 2 may be a result of a slight fall in sea-level (~ 50 cm) producing a lowering of the level of saturation and hence platform erosion surface. Discussion These data have implications both for the evolutionary history of the low energy beach ridge sequences and for mid-late Holocene sea-level history in the Bight and on Kangaroo Island. Beach Ridge Progradation The initiation of beach ridge progradation at nine of the 12 sites dated did not commence until 4 ka BP and in some areas as late as 2 ka BP (Fig. 9). Most sites indicate an hiatus between the Holocene sea-level stillstand and inner ridge building of up to 4 ka. This is in general agreement with Burne (1982) who found that in northern Spencer Gulf the BEACH RIDGE PROGRADATION I Site 15: Brownlow (m) ' “FF Aeolian Sand — 3 oe ; E ate ete NN PEN ee 2b Shotly Sand : + 100 200 is) site 17: Cygnet River 4b = 1 2 Fine Sand 7 \ Salt Fat Ae = at 2 o3kh y z ° - 2 < > wi a 1 200 i] at} aite 18: Marrison Beach 5 5 i 4 3 2 1 eh | | i 20 40 60 Lay ~ bb | Site 16: Brownlow (Golf Course) i = Shelly Sand * "34" #3 aWK840 2650: GO b.WK841 2290' 50 i Tidal Crock 400 600 {m) Pleistocene Shore Platfarm Platfarn 1 Galcarenite Platform 10 20 30 DISTANCE (meters) Fig, 8. Kangaroo Island field sites. Beach ridges are numbered chronologically, development of low energy beach ridge pluins uwaited sufficient subtidal carbonate production and as a result progradation did not commence until 4-3 ka BP. Belperio e¢ a/, (1988) also concluded that carbonate secreting organisms uggrade and prograde the subtidal sand banks in association with the subtidal seagrass facies. Once the banks accumulate to low water level, subsequent intertidal und beach tidge progradation can oceur. They found this sequence al Tourville Bay on Nyre Peninsula as well as several sites in Spencer and St Vincent Gulfs, In Gulf St Vincent, Bowman & Harvey (1986) found that the extensive Le Fevre Peninsula beach ridges have been prograding continuously over the past 7000 years with pluses in ridge formation apparently linked to longshore (northward) sediment supply, This higher energy west facing As A, DO SHORT, ROC. BUCKLE RY AD, G, POTEIERINGH AM system Gnodal Hb = 0.5 mi) represents a transiion belween the low ederey beach ridges discussed in this paper and the high eneray¥ open coast regressive depowts which experenced rapid progrudation following the salistand. Their mode and sequetice of esolution of the Peninsula is therefore oe direcily applivuble to the lower cheney systems of all which experienced a substantial tig in ridge development, On the (wo sites where a sequence of beach ridges was dated (excluding Eyre Island) the soungest Tides dated 2010 vy BP (Glensea> and 1770 yr BP {laura Bay), suggesting that ridges are either nat currently prograding or older material is being deposited during ridge building. Whilst ridge cessaliod 4 i aercentent With sedimentation models ivhigher energy sites do south-east Australia (Thom etal. )984), itis surprising in ad environment where most sediments are derived front jt sida carbonale produetion from fronting intertidal and subtidal sand flais, the so cullud ‘cay bonate factary' {Burne 1982). In northern Spencer Gulf, Burne found periods of shoreline (ic. sundflat) progradation, aliernacng with beach-ridge progradation. The most notable shorcline progradation was between Jand 2 ka BP, On the Eyre Peninsula, however, that was the period of maximum beach ridge construction. A similar contrast applied te (he penod from 2 ka BP to presents beach ridges in the north-east Spencer Gulf prograded, while those in ihe Eyre appear (o have rervained static. Evidently nore information iy required before genctal models of S Aust. beach ridge-sand flat progradaliqn can be clarified, Eyre Island liss a slightly younger outer ridge «400 yr AP) and the ishind is vontinuing to develup new recurved spits, However this may be accounted for by erosion and dowadrift reworking of istand sediments father that external sund supply. Hulucene Sea-Level Yhe exdet timing of the Holocene sea-level Maxi eatanot he resolved from these dites However the date of 6330 - 90vr BP from Little Douglas as Well as the 3450 | 90 yr date trem Acranmin Creek lend support to the general curve adupted by BelperiG ef af, (1984) for Spencer Gull aid | opley & Thorn (1483) for Australia as a whole, Phat is it dppears thae the sea-level reachyeel its present position by 63 ka BP in the Greal Australian Right, Warren, dK, (982) Nate Quarernare evaporite cambodatecdepourional covianmenisad Saul Amaral, PHD esis, Plaideds Gil. Adeluide (uopulht Inevaluating evidence lor relative sea-level history during the mid to late Holocene, tectome differences between (he three main regions, the Bight, Spencer Gulf and Kangaroo Island must be taken inro aecoult. The most stable resion appears to be western Lyre Peninsult (the Bight), Teetonic stabiliry i§ Suggested by the widespread occurrence of the + 2-3 Pleistocene shore pluctorms (Buckley ef a/. 1986), the continuity of the presently. active shure platforms. the single intertidal solurion notches in limestone clifts in Coffin Bay and the evidence from dated beach ridge and tidal sites (Figs 4, 5), In the Bight the s¢a-level maximum was reached at least by S650 yrs BP AFig. 5b) and has been stable since. In northern Spencer Gulf there ip, However, evidence for Holucene reeronism. Belperio ef af (1984) proposed a sea-level cucve for vorthern Spencer Gulf which has a srillstand being reached ar ~ 6.5 ka followed by an abrupe fall ta present sea-level at 1.2=l.6ka BP. They auribure (his relative fall te local tectanism, The present Speucer Cull dala are not sufficient lo.add to the Belperia ey a/. curve und may well be tectonically or isastatically distinct, The Gilensea bec rides aod swales slupe seaward Fig, Ja); however (his 1s not conclusive evidence for a slight fall iv sea-level without subsurface morphostratigraphy. I could also represent iv decrease in waveencrey as the T Am wide intertidal and subtidal sand Mats accunmulated, Likewise, while other Spencer Gull’ sites logerher with Yanwe Isiand and Mount Dutton Ray sites nay suggest a mudesi fall in sea-level (<1 m), errors inherent in surveying, facies identification and elevation, dating and establishing reliable davums, place all of (he possible changes within the sampling error bands, Conse quently (here iy no conelusive evidence ura small relative (Hl) insca-tevel, Wi any has oceprred it would be only on the order ofa few eleciinelers. Together with the Bight data this suggests a relatively stable sea-level site the sea-level maaima ar or prior tu 6300 yr BP Summary aed Canchisiuans The following sequence oF events appeiins to have Ooeeld red ulong Me lay eferey Bight, Evre Penisulit and Kangaroo Ihind coyists sites Hie Holoeene sed level srillstana, 7-6 ka BP. The stillsiand was achieved by at leasi 6330 + 9041s, BP and most likely by 6,6-6,4 ka AP based on evidence from northern Spencer Gulf (Belperiés ef af L984), 6—1 ka BP Pollawing the stillsiand mast law energy secliuns of caast experienecd minimal shoreline proanmidition, Exceplons were areas where shingle Was immediaigly available, particularly ib Spencer 159 BEACH RIDGE PROGRADATION “puels] OOIesULY puke e[NsutuAag aAq Uo Sars aEpi4 Yea AB19U9 MO] IOJ WOTIBADTS SNSI9A Sa]Bp SUSIO[OH JO JO} 6 “A dd SUVSA (ll of Holocene sea fevel and enastal progradati vi, northeastern Spencer Gull, South Nustcalia. BMH fr Aust, Geol. Geophys, 7, 35-45 Burterk, A... Dreers, A. ML, McKitiup, S.C, & THowss, DP. (1977) Distribution and sediments of mangriowy elds in Soulli Australia. Tras. R, Soe. S. lust, WOH, 38-44, CHtiTON, Dh, Hussey, K. J & Parke, M. E. (1981) Global satelliile measursiicnis of water vapour, wind speed vil wave height, Satire 294, 529-532. Dany, B. Mirsss, ALR, Twtbare, C. RB, & BOURNE, JA. (1979) Geology aad geamorphology pp 1-38. /7 M, J. Tyler C. R. Twidile, & dK, Ling, (eds.) “Natural History of Kiingaroo Island" (R. Soc. S. Aust, Adelaide) DE DECKKIR, P., BOLLD, J. & BURNE, R. V. (1982) Pillie Lake, Eyre Peninsula, South Australia: Modern environment and biota, dolomite sedimentation, and Holocene history. Trani. R. Soc. 5. Aust, 106, 169-181. Galioway, R, W., Story. R.. Cooper, R. & YApp, G. A. (1984) Coastal lands ot Ausiralia, CSIRO Diy. water and Land Resources. * ‘Natural Resources Series 1. Haris, J. R. & Gostin, VN. (Eds.) 1984 The Spencer Gulf region, Marine Geol, 6L, 111-424, Hesp, P, A. 1984 The fo qutton of sand ‘beach ridges’ and foredunes. Search 13. 289-291. Hopcey, D. & THom, B. G. (1983) Australia sea levels in the last 18,000 years: a review. Jn Hopley, D. (Ed,) Australian sea levels in the last 15,000 years: a review pp 3-36, Qcvas. Pap. 3, Monog. Ser. Dept Geography. James Cook Uniy,, Townsville. Morris, Bo 4 (1976) Investigation of heavs imineral sands, Morrisua Beach, Kangaroo Island, Adineri/ Resources Mev, 145, 34-46, SHorRi, A. 2. +i98Sa) Response of a high iraugh low energy Sandy coast to sea level transgression and stillstand, South Australia, Geogruphical Review 78, 119-136, — (98KS! Modes of Holocene coastal dine transgression in southern Australia, Sedimentary Geol. 55, i21-l4e - FGI edeam, DL G, (1986) Coastal mMmorpho- dynamics and Holocene evoludion of the Kangaroo Island. voasi, South Australia, Coastal Studies Unit Tech, Rept, 86/1, +s uastal Studies Unit Univ. Sydiey, Sydney). - > ——_. » & Brentey, R. C_ (1986) Caasial marphodynamics and Holocene evolution of the Lyre Peninsula coust, South Australia, Coastal Studies Unit ‘Tech. Rept, $62, (Coastal Studies Unit, Univ. Syditey, Sydney). & Hew, PA (1984) Beach and dune morpho dynamics o! the South East coast of South Australia, Coastal Studies Lnit Tech. Rept 84/1 (Coastal Studies Unit, University of Sydney, Sydney). THom, BL G.. Rowman, G. M,, GILLispre. Ra, Pesan, R, & Baker iii, M. (1981) Radiocarbon dating of Holacene bevch ridge sequences in southeast Australia, Monog. LH, fue Geography, R.M.C., Duntroon, ACT. TWiDALE, C. R., Bourne. AL & Twroatn, N. (1977) Shore platfort witd sea-level changes in the vulls region of South Awenha Trans, R, Sac. 8. Aust. 1, 63-74. THE OCCURRENCE OF A DIPROTODONTID MARSUPIAL REFERABLE TO EUOWENIA GRATA DE VIS IN WESTERN NEW SOUTH WALES BY N. §. PLEDGE* Summary A complete last upper molar (M?) and fragments of several other teeth found in a bore on the Talyawalka Anabranch, east of Menindee, N.S.W., are identified as the Pliocene species Euowenia grata De Vis. Euowenia grata is redescribed and refigured, and the new M? is compared with the equivalent teeth of Diprotodon optatum, Zygomaturuwss trilobus, Meniscolophus mawsoni, Nototherium mitchelli and Palorchestes azael. The Talyawalka specimen extends the range of Euowenia grata by almost 1000 km and suggests a Pliocene age for the deposit. KEY WORDS: diprotodontid marsupial, Euowenia, Diprotodon, Zygomaturus, Meniscolophus. Nototherium, Palorchestes, Talyawalka, Menindee, New South Wales, Pliocene. Transactions of the Royal Society of S. Aust., (1989), 113(3) 163-168. THE OCCURRENCE OF A DIPROTODONTID MARSUPIAL REFERABLE TO EUOWENIA GRATA DE VIS IN WESTERN NEW SOUTH WALES by N. S. PLEDGE* Summary PLebGe, N. S. (1989) The occurrence of a diprotodontid marsupial referable to Euowenia grata De Vis in western New South Wales. Trans. R. Soc. S.Aust. 113(3), 163-168, 30 November, 1989. A complete last upper molar (M°) and fragments of several other teeth found in a bore on the Talyawalka Anabranch, east of Menindee, N.SW., are identified as the Pliocene species Euowenia grata De Vis. Euowenia grata is redescribed and refigured, and the new M’ is compared with the equivalent teeth of Diprotodon optatum, Zygomaturuwss trilobus, Meniscolophus mawsoni, Nototherium mitchelli and Palorchestes azael. The Talyawalka specimen extends the range of Euowenia grata by almost 1000 km and suggests a Pliocene age for the deposit. Key Worps: diprotodontid marsupial, Euowenia, Diprotodon, Zvgomaturus, Meniscolophus, Nototherium, Palorchestes, Yalyawalka, Menindee, New South Wales, Pliocene. Introduction De Vis established the species Owenia grata (De Vis 1887), for a skull and jaws (Fig. 3, QM F519) found at Chinchilla, Queensland. Later he renamed it Euowenia (De Vis 1891) after finding the earlier name to be preoccupied by an annelid worm. At the same time, he named a second species, Euowenia robusta, for a set of dentaries found at Freestone Creek, near Warwick, Qld, in Pleistocene alluvium, Woods (1968) suggested that Euowenia robusta is a junior synonym of Nototherium inerme which could effectively restrict Euowenia to specimens from the Chinchilla Sand, and to a Pliocene age (Woods 1960; Bartholomai & Woods 1976). Euowenia grata occurs with certainty only at Chinchilla (Stirton ef a/, 1968). M. Archer has informed me (pers. comm. 3.xi.1975): “In many cases, specimens should more honestly be referred to merely as undetermined small nototherines”, and there is no evidence at present for any Pleistocene occurrence. Recently, Ewowenia has been listed (Tedford, Williams & Wells 1986) in the Plio-Pleisto- cene Kanunka local fauna at Lake Kanunka, South Australia, but this record needs confirmation. The purpose of this note is to record a new occurrence of the species well beyond its previously known locality. Material and Methods In 1969, Mr D. Walker of Talyawalka Station east of Menindee, New South Wales, presented to the South Australian Museum some unusual tooth fragments found in the sludge from a newly-drilled water bore in “White Water Lake” (32°25’S, * South Australian Museum, North Terrace, Adelaide, S. Aust. 5000. | | Broken Hill 3 | a al 2 z= | < | ne i L.Menindee | 2 | L-Tandou | | Peet | NEW SOUTH SOUTH WALES AUSTRALIA | { | | | 0 Km 100 Fig. 1. Locality map. White Water Lake, Talyawalka Anabranch, New South Wales. 143°18’E) on the Talyawalka Anabranch of the Darling River (Fig. 1), The new specimen is very fragmentary, comprising the complete left M’, two enamel fragments with differing degrees of wear, believed to be anterobuccal corners of the protolophs of M' and M' of the same tooth row, the anterobuccal corner of the metaloph of M', the anterior half of M,, an unidentifiable molar fragment and a fragment of incisor. Because of the poor quality of De Vis’ figure (De Vis 1887), the new specimen was for several years considered by the author to be a possible very large new species of Palorchestes, because it had a fairly distinct midlink between the lophs. However, direct comparison with a cast of the upper dentition of Evowenia grata (holotype QM F519) by courtesy of Dr M. Archer, has now proven its identity. The specimen also has been 164 N.S. PLEDGE THE OCCURRENCE OF A DIPROTODONTID MARSUPIAL compared with teeth or casts ot Dipratodon optatum, Zveormaturus trilabus, Meniscalophus mawsoni, Nototherium mitchell and Palorchestes azuel, Material is from the South Australian Museum, Adelaide (SAM), Queensland Museum, Brisbane (OM) und British Musetum (Natural History), London (BMNH). The status of the genus Evoweni¢ is thought to be in doubt and in need of mayor revision (Archer & Wade M976), but wai this done itis felt that Aadcdesery ton of fhe species is desirable, based partly on this new material. Tooth notation follows Archer (1978), SYSTEMATICS Euowenia grata De Vis FIGS 2-4, Owenia grata De Vis, 1887, p. 100, pl. I-IV Upper premolar two-rooted, about half the length of M* and triangular (De Vis 1887, 1888). Protocone slightly smaller than parametacone. Protocone and anterior part of parametacone joined by narrow transverse loph to form short side of a right triangle. Buccal side formed by buccal crest from parametacone to anterior end of the 165 tooth, Third side formed by anterolingual cingulum. Small deep central basin with rugose enamel, Upper molars large (Tables 1, 2) and bilophodont, with lophs convex forwards and somewhat tumid with incipient midlink development — postproto- crista posteromedially from protocone and pre metacrista(?) extending anteromedially from meta- cone. Midlink development virtually absent in M¢ but increases slowly to its greatest extent in Mé where pre- and post-cristae are contluent fora few millimetres vertically, M°~“ subrectangulary MM? trapezoidal, with metaloph abour three-quarters width of protoloph. In M! and M>, metaloph forms distinct dog-leg bend where midlink joins it. Anterior cingulum extends full width of tooth, but is longer at its lingual end. Posterior cingulum narrow, particularly on M’ where small pocket may be formed by weak postmetacrista, Only trace of a lingual cingulum may be seen, at bottom of transverse valley. Buccal cingulum better developed, across mouth of transverse valley. Lower molars more rectangularly elongate than uppers. Both protolophid and hypolophid are oblique and concave anteriorly. Pratoconid end of protolophid enlarged by development of preproto- cristid extending anteromedially. Similarly, hypo- conid end of hypolophid is enlarged by cristid obliqua which forms posterior half of weak mid- TABLE 1. Cheek Teeth Dimensions, Euowenia grata Except for (he premolar, measurements are made in planes parallel io and normal to the lingual edge of the tooth. SAM P14506 QM F519 (epoxy cast) SAM P14506 LM. LP* LM? LM" LM! LM’ LM" A-p Length ~ 15.0 26.5 32.6 36.2 35.7 34.8 Ant, Diam. 19.3 — 20) 26.2 30.3 28.7 27.7 Post. Diam. — 15.7 24.2 25.9 28.4 23.2 23.1 TABLE 2. Euowenia grata Dimensions as given hy De Vis (1887) These are not directly comparable with those to Table 1, as M” for example is apparently measured along the outer edge. PL M. Pp! M’ Length 14,5 34.5 1540 38.0 Max. Breadth 85 22.0 14,5 29.5 Vip. 2. Euowenia grata De Vis. Lett ipper molar M’ of SAM P14506. Stereopair photographs: a) Occlusal; b) buccal; ¢) (itigual view, Seale = | em. 166 N. S, PLEDGE FIG. 3. Euowenia grata De Vis. Holotype palate and dentaries, QM F519. Scale bar = 10 cm. link. In the holotype, lowest part of midlink is intact in last molar (De Vis 1887). Anterior cingulum almost full width of tooth. Postcingulum divided unequally into two parts at its highest point by fine submedian posthypocristid. Comparison with other species The Talyawalka specimen may be compared with this (Table 1) and other diprotodontoids. The upper molar Mis the same size as that of the holotype of Euowenia grata, and same order of size as Meniscolophus mawsoni. It is smaller than Zygomaturus trilobus and slightly more than half the length of Diprotodon optatum. The Talyawalka tooth differs from Diprotodon (SAM P23406, Lake Callabonna) in: its much smaller size, relatively higher crown, more convex protoloph, slightly narrower precingulum with greater development at the lingual end, a buccal cingulum across the end of the transverse valley, a narrow but strongly developed postcingulum, the absence of cementum, and particularly the presence of a Y-shaped hypoloph formed by a distinct premetacrista. It differs from Zygomaturus (SAM P29889, Naracoorte) in: its smaller size (about three-quarters the length), shorter precingulum, absolutely higher lophs, thicker lophs, presence of a distinct buccal cingulum across the transverse valley, deeper and more acute transverse valley, distinct premetacrista, more distinct postmetaloph-pocket, and presence of rugose/punctate enamel. It differs from Nototherium mitchelli (cast of BMNH M5002) in: its smaller size, its longer and better developed precingulum, presence of a buccal cingulum across the transverse valley, relatively higher lophs, presence of a premetacrista, and better developed postcingulum. It differs from Meniscolophus (SAM P13647, of which M? is unknown) in slightly smaller size, higher lophs, presence of distinct premetacrista, and narrow postcingulum. THE OCCURKENCE OF A DIPROTODONTID MARSUPIAL. th’ Pi. 4, Euowenla grata De Vis, Cast of left upper maxillary tooth row of holorype, OM FSi9 Ni differs trom rhe holotype of Auowenia grata (cast of QM F519) im having a distinct buceal cingulum with two small (one twice as big as the other) cusps across the transverse valley, and a more distinel posimelaloph-pocket. I differs from Palorchestes azael (QM F773, fide De Vis 1895) in: its larger size, relatively greater length, lack of a forelink, presence of a strong buccal cingulum across the transverse valley, strong postcingulum, and absence of a complete midlink. The new specimens (SAM P14506) agree closely with the holotype, considering the difference in degree of wear and preservation, In M1’, the enamel is just breached so that a good indication of the height and torm of the tooth ean be gained, The lophs are high (eqnal to the width of the tooth) aid stout so that the transverse valley is particularly deep and narrow, a harrowness accentuated by the bend caused by (he premetacrista, ‘Together, the metaloph and premetacrista have a distinct Y-form, Centra De Vis (1887), 1 found no indication of a forelink on the upper molars of the holotype cast, bul there is a slight protuberance i (his position on the new specimen of M®°. The molar cnamel is notably rugose and punctate, similar to that of Diprotodon and Euryzyzoma (Archer 1977). In the worn M,, frag- Ment, the anterior cingulum is heavily abraded by interdental appression and has a small cusp de- veloped at the anterolingual corner; The “midlink" is discontinuous and does not cross the transverse valley as an integral structure: The incisor is too fragmentary to describe, save that it would be roughly circular in section with a diameter of at least 22 mm, Only a portion of the rool is preserved and, from De Vis’ deseription (1887), it could be either upper or lower II, Discussion The teeth came trom a depth of about 28 m, in a 1.5 m thick bed of coarse, yellow-brown dirly quartz sand, below sandy clay, The age is uncertain, but an early Pleistocene, or even Pliocene, age 1s possible, based on the absence of the species rom lale Pleistocene surficial deposits nearby — hinette sands dated at 25,300 years BP at Lake Menindee (Tedford 1967) and 32,000 years old at Lake Mungo (Barbetti & Allen 1972), Although speculative, this determination is supported by the Pliocene age of the holotype (Woods 1960). The specimen extends the known range of the taxon by almost 1000 km, It seems that the bore drill struck a more-or-less complete skull and jaws of Euowenia because all tragments appear to relate to a single individual, there being no duplication of elements. Conclusions These fossil teeth, recovered from a water bore on the Talyawalka Anabranch, eas! of Menindee, N.SW., match those of Evowenia grata (De Vis) and, because (hey are Jess worn than the holotype, add fo (he knowledye of the species. The occurrence extends the known range of the species. Acknowledgments | thank Mr D, Walker for donating the specimen to the South Australian Museum, Mrs Debbie Brunker and Mrs Debbie Lowery for typing the manuscript and Mrs Linda Blesing for drafting the map. References ARCHER, M. (1977) Origing and sublanilial relationships of Diprotodon (Diprovodontidae, Marsuplatia), Mem. Qld. Mus, V8, 37-39. —_—— (1978) The nature of the molar-premolar boundary mi marsupials anda reinterpretation of the homology of marsupial checkteeth, Jbid 18, 187-164. & Wabe, M. (1976) Results of the Ray EB. Lemley Expeditions, Part 1, The Allingham Formation and a new Pliocene vertebrate fauna from northern Queensland. /hid 17, 379-397. Baruetii, M. & ALLEN, H, (1972) Prehistoric mar at 168 N. S. PLEDGE Lake Mungo. Australia, by 32,000 years BP. Nature 240, 46-48. BARTHOLOMAIL, A. & Woops, J. T. (1976) Notes on the vertebrate fauna of the Chinchilla Sand. Appendix 2 in Bur. Min. Resour. Bull. 166, 151-152. De Vis, C. W. (1887) On an extinct Mammal of genus apparently new. Proc. R. Soc. Qld. 4, 99-106, pl. 1-4. (1888) Notes on the genera Zygomaturus and Nototherium. Ibid 5, 111-116, plate. _ (1891) In confirmation of the genus Owenta so- called. Proc. Linn, Soc, N.S.W. (Ser. 2) 6, 159-165, pi 13. (1895) A review of the macropodidae in the Queensiant Mossum 75-134, pls. 14-18. STIRTON, R, A., TEDFORD, R. H. & WoopBuRNE, M. O. (1968) Australian Tertiary deposits containing terrestrial fossil vs af the lind 1, mammals. Univ. Calif. Publs Geol. Sci. 77, 30 pp. TeDFoRD, R. H. (1967) The fossil Macropodidae from Lake Menindee, New South Wales. Univ. Calif. Publs Geol. Sci. 64, 156 pp, 4 pls » WILLIAMS, D. & WELLS, R. (1986) Late Cainozoic sediments and fossil vertebrates pp 42-72. In Wells, R. T. & Callen, R. A. (Eds) “The Lake Eyre Basin — Cainozoic sediments, fossil vertebrates and plants, landforms, silcretes and climatic implications”. Australian Sedimentalogists Group Field Guide Series No. 4+. (Geol. Soc Aust... Sydney) Woops, $ T. (1960) Posstliterous thiviatle and cave d (Eds) The yaits. Un Hill, O A& Denmead, ALK. wy of Queensiand. £ Geol Soe. Aust. 7, 393-403. — (1968) The idenuty of ihe caisnci marsupial genus Nototherium Owen. Mem. Qld. Mus. 15, 111-116, pl. 13-14. TRANSACTIONS OF THE ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED VOL. 113, PART 4 PARASITOIDS OF URABA LUGENS WALKER (LEPIDOPTERA: NOCTUIDAE) IN SOUTH AUSTRALIA, WITH DESCRIPTION OF TWO NEW SPECIES OF BRACONIDAE BY A. D, AUSTIN & G. R. ALLEN* Summary Information is presented on the large complex of hymenopteran and dipteran parasitoids associated with Uraba lugens Walker (the gumleaf skeletonizer) in South Australia. A key to the 22 species involved is presented, along with notes on identification and relationships with their host. Two species of microgastrine braconids are described, Cotesia urabae sp. nov. and Dolichogenidea eucalypti sp. nov.; both are parasitoids of the larval stages of U. lugens. KEY WORDS: Uraba_ lugens, Noctuidae, parasitoids, hyperparasitoids, Braconidae, Ichneumonidae, Aphelinidae, Chalcididae, Elasmidae, Eulophidae, Eurytomidae, Eupelmidae, Trichogrammatidae, Tachinidae. Transwelions of the Koval Sociey of & Aust, (1989), LIM4) 169% 184. PARASITOIDS OF URABA LUGENS WALKER (LEPIDOPTERA; NOCTUIDAE) IN SOUTH AUSTRALIA, WITH DESCRIPTION OF TWO NEW SPECIES OF BRACONIDAE by A. D. AUSTIN & G. R. ALLEN* Summary Ausrin, A. D. & ALLEN, G R. (1989) Parasitoids of Urabe (ugens Walker (Lepidoptera: Noctuidae), with descripuian of rwo new species of Braconidac. Trons, R. Sov. S, Aust. 113(4), 169-184, 1D Noveniber, 1989. Information is presented on the large complex of hymenopteran and dipteran parasitoids assucialed with Uraha luzens Walker (the gumleaf skeletonizer) in South Australia. A key to the 22 species involved is presented, along wilh nates an identificanon and relationships with their host, Tao species of micengasirine bravonids are described, Caresio nndlive se mo, and Nolichogenidey eucelypli sp, now.; both are parasitoids of the larval stages of Of lugens. Key Worps Graha dueens, Noctuidas, parasitoids, Hyperparasitoids, Bracomdar, Ichneumonidae, Aphelinidae, Chalcididae, Elasmidae, Eulophidae, Kuryiomidae, Eupelmnidae, Trichogrammatidac, Tachiniduc. fntroductiva Uraba fugeas Walker, he gumleal skeletonizer, is 3 native noctuld moth and has been. collected from all Scares of Australia (Turner 1944), It has been recorded as damaging stands of cucalypt species in eastern Australia (Brimblecombe L962; Campbell 1962; Harris 1972!, 1974; Harris er ai. 1977°y, the Adelaide region and south-western Western Austialia (pers, comm. F. D. Morgan; Strelein 1988). Occasionally outbreaks of this species can defoliate large arcas‘of native forests. Several such ourbreaks have occurred in stands of Eucalpytus camaldulensis Dehnh. along the Murray Valley region of N.SW. and Victoria, where, on at least four Occasions, More than 30,X) fia have been affected (e.g, Campbell 1962; Harris 1974; Harris ef al. 1977"). Apart from the widespread damage associated with this spevies, it is also responsible for the partial defoliation of individual eucalypt trees planted as ornamentals in parks and gardens. Other than the work of the above authors and those of Morgan & Cobbinah (1977) and Cobbinah (1983), very little has been published on the biology and ecology of L! tugens, while even less has been reported on its parasitoids, Brimblecambe (1962) reported five species of primary purasitoids as "Harris, 1A. (1972) Che effect of flooding on papulation density of the gum leaf skeletonizer morh, Urabe lugeny Walk. in Barmah State Forest, Fores) Commission, Victoria, Research Branch Report, No. 25 (unpubl) “Elarris, J. AJ Neumann, FG. & Ward, B. (1977) An outbreak of the gum leaf skeletonizen Urabe lugens Walker, in river red gam Torest near Barynah. Fores! Commission, Vicioria, Research Branch Report, No. 87 sonpubl) * Department of Entomology, Waire Agricultural Research Tnstinite, Universiny at Adelaide, Cilen Osmond, S. Aust, S064. altucking LU dugens and Campbell (1962) ten species However, in most cases these parasitoids were no! identified Jurther than family level and, if so, their identification was not reliable. One of us (G.R.A.) has recently completed a major sludy on the interaction of this insect and its parasitoids in the Adelaide region. This work shows thal the immature stages of U, lugens support a diverse complex of hymenopteran and dipteran parasitoids (22 specics — Table 1), which includes both primary parasitoids and hyperparasitoids, In this paper we provide 4 taxonomic framework for the information on the behaviour and ecology of’ this: parasitoid complex ard its wteraction with &, fugens, which will be published elsewhere by ChR.A. Here we present a key to identify all the parasitoid species involved, and provide notes on their taxonomic position and biology, including information on the stage attacked and place of pupation, Two of the more common species reared from U /ugens larvac. which are members of che braconid subfamily Microgastrinae, and are the subject of detailed behavioural studies by G.R_A., are described here as new. Materials aod Methods All lite bistery siages of U dugens were collected froin eucalypts (mostly FE, cumaldulensis, & feuvoxylon F. Mucll. and &. pricrocanpa (Maiden) Maiden) between 1985 und 1988 at several sites within 7 km of Adelaide G. P.O. They were brought to the laboratory, held gt 20°C in rearing cages, and larvae provided with new foliage, so that any parasitoids present could develop and emerge normally, Parasitoid cocoons found with ibeir dead host in the field were also collected and reared in the laboratory at 20°C until adults emerged, Vi) 4.1, ALIST IN A Ci R, ALLIN TABLE 1 Sueterant af relationships berween UU, lugens and its parasitoids and. hyperparasitaids a Parsi Primary (P).or Hi perparasnaid (H) Li. levers suive Stage viteryis pels Family Solitary (8) or Gresuniaus (C1 netacked trom A a Treitiourantind Sp, Trichegrammatidge Pis ene 22 Cola yrahae Braconidae BS larva larva Doalivhogenilea eucalypt Braconidae PS larva larva fuplecveus ap. Eulophidae BN larva larva Casihora mer {ehneumoniclae BS lorva larva Exorista Navivepy Tachinidae WS larva larva Priborus sp. IchneumoniWae RS hanya pupil Yenthopunpla rhopalaceroy lehneumociudee PS pupa? pupit Anirocephatis ay Craleididae PS nope pupa Brachemeria sp. | Chaleididae RS pupa pupa Winthemia larenalis Tachinidae Iss i pupa? Eearktorna sp Eulophidae PGS pupa pupa /purasitoid cocoon Centrudora sy. Aphelinidae TG — parasitoid coenon Urachymerid 4p. 2 Chaleididae Hs ~ Parasitoid covoon Klasius aastrulienses Fla\midae HG — parusitoid cogaon qpeviys ines, Ewlbophidae IG _— parasitoid eacoon Podtobns sp. Fillapaidac HG = psrasitad cacoon species indet, Eupelmidae HS parasitoid carood Anasiatuy sp. Eupelmidac HG — parasitoid, cocoon Mesachorus sp. Iulneumonidae HS — paHasditmd cocoon Paraphylax sp. (chneumomdae His - parasitoid cocaon Pleririalus sp. Precomilidae Hib - parasitoid cocoon Parusitoids were stared in a freezer or in 70% 3, ethunol prior to mounting on pins or card points, Material for S.E.M. study was washed in half strength concentrated liquid soap, rinsed in distilled waiter, dehydrated in an alcohol seriés and critical point dried using an Emscope CPD 750, before being cxamined under a Cambridge Stercoscan 250 (Mk 3B) electron microscope. Terminology for morphology follows Boutek (1988), Eady (1968), Gauld (1984), Harris (1979), Mason (1986) and van Achterberg (1979), The term ‘alitrunk’ is used for the tharax plus propodeum, and ‘gaster‘ is used for the post-ptepadeal segments, The abbreviation ‘T’ refers to the gastral tergiles. Abbreviahons for collections are; ANIC, Austratian National Insect Collection, CSIRO, Canberra, WARL, Waite Agricultural Research Institute, Adelaide. Voucher specimens of all species are lodwed in the Waite Institute collection. Key to the parasitoids of L'raba lugens in South Australia 1, Two pairs of wings developed; dorsal surface: very rarcly with slout bristles: wasp- lke i appearanee (ovi posiror always developed in fepygle and wsuatly clearty visible (Migs 5, 7, 29)) (Tymenoptera), , 2 Only one paar od wins developed (fore wine 28); ‘dorsal surface with nunierods stout bristles; blow fly like in appearance (Figs 48. 49) (scutum with several black lensiniding] bands) (Tachinidacy --21 Pow wing with relatively complete venation (eg. Digs 1,2, 6, 19), 3 hore wing wilh pigrnemed venation Feabestt lo dmlenor Maren (ee Paes 2, 23, 27, FB eee ee ed hy 6. More wing, with venation distal to prerosiigma wanting (Fies J, 2), sem 2 absent (Rraconidaey..-. 4 Fore wing with) distat veins peesent and well- pizmented; vein 2ancu a te 4, “Hy. {etc monde), ee a i ett rréncideum with lonwivaglisal nvedial carina, coarsely sculptured at least anteriorly (Pig. 12); legs red to red: yellow . . Cotesta urubue ap. nay, Propodeum with large arinate areola and horizontal carinae extending to lateral margins of propodeum (Fig. 13): leas dark brown fo black with distal parts reddish... 2... Doalichogenidea eucalypt sp. nov. fore wing wih an dredlet (Pigs 4, 6).... 0... -- fi Fore wihg without dn dreolet (Figs 19, 20)......7 Scatum and prapodelin coursely punctate or ruiwulase: ovipositor very shorn (Pig, 95 oO Benimalny Wifthoutlone rods protruding posteriorly (body dark Hrewir to black, fews reddish), 0... er een Custraria Uvera tooman & Giauid Seucuim and propodenm generally unsculptired (oxcepr for propaded) eurinae); OVIpOXiton extending well past posteriar easter (Fig. Ty, ot pentratia wirk pair ol lon fads (poresquama) probity poster ian Lye (body vellow-browin with uitrher Mmarkinsh oc. Mesochorus sp, Body brush) yellow will block markings: EE shorn ancl breistel hasatlly (Vig, 16)... Nunthapinphr rhapaline eres ‘Krieger Body nut so coluureds TH narrow Iasi (bigs 17, 18) fe ee a) Fore wing with radial cell} short and browd (Fig. 19): ‘TT tlat, broadening distally (Hig. 17) (small species, lenwth 23-3.2 mm wot including 2 ovipgsilor, body black except for T2 which is yellow-brown Paraphylax sp. kore wing with radial cell lane and narrow (Fig. 20). MT tubular in basal ball, balbouws in distal half (Fig 18) (large species. lenath 6.5-8.7 mim not including ovipesitor tor » head and alitrunk black, gasrer reddish-brown - Frihorus ay) eae 10. a = 4 if PARASIVOIDS OF WAAR FUGENS V7 Femur of hind leg greatly expanded, juolaed ou sepnited aloe lower margin (Fig. 24) (hal white) ee pit eco se ey Le Tymar of hind leg normal, smgott along lower tiangin (Pipe 25; 35) oy ecaarpeeesse 4 ee wae ee 12 fore wing willt marginal ver much longer than pastmarginal vein (Fla. 22); apex of hind tibia tapering Imo strong spine (Brachpmneriay. 00.000, Pere wing, with marginal vein about same length as pasiparginal vein (Fig, 2 apex of hind sihia perpendicularly (runeale (Fig. 24) (large spevics, 4.8 mm in teneth; body black, bine lee dark red-brawn marked witth black) Antrecephalas sp, a Boy black with reel lined lenmurand tities 42-44 myn in Jength, dag ey eet vee Brachymeria sp | Kody black with Whikkyellow markings on rowalau yd leps; L8-23 mim in Jength,.... Brachynerie sp. 2 - Hibd voxa developed as large [at dise; hind tibiy waph Uistinel criss-cross patterd al setae (Fig. 25) (fore wing with stigmal vein very short (Fig. 23). body dark. tezula and legs excepr for hind coxa pale; body length ot 18-27 mm, ¢ 13-19 mm) (Elasenidac) vevtreees cs 2 Blass australensis Girault Hind coxa, hind tibia and stiemal vein nat as above ane, tri teet th Mee. te 13 . Pronotum (seen derselly) targe and qiradeangutiar; dorsal surtave of abtrunk coarsely sculptured (Lig. 2h), fole Wing Venation asin Fig, 27 (hody black aod Hon-metilhe, legs with seme pale “tnarkings) (Buryronmdaey, oc .. eee) Burytone sp. Pronoturny not large and quadrangular and ahrrunk withoul such sculpraniies: body offer meruic in OTE usttetn. ee, 345 ath Body length greater than about 1.5 ome: waster sepiraicd from alinnink by narrow waist (ges 28, 29) 1S Ai ote, & Het - oo 20 Tarsi S-segmented (ull Jigs 24, 257... , 16 Farsi 4-segmiented (Pig. 359 ito LB Mesopleurion hot cotired and shield-lke (Ei. 28); body rather robust with large hoad ancl alitrunk (2 aaster i laigral view Sharply angled: cr paster snvall and (Tallened; body length of & 23-33 mm, o [7-24 mince inetallic green ia colour with yellow-hrawi legs and antennae) (Pieromatidae) » Preramalus sp. body sumewhett 17 - Fore wing with stigmal and postmaruinal veins very Shor (Fiy. 34); anterior sewlellum longitudinally strivte (Pig. 43) (hady dark with merallic ureen tinge, 11-15 mm in engi), 00. Pediohnes sy. Vivre Wiig wilh stigmal and postinarginal veins long (Fig 7)> scutellum sinmorh or with fine rencntate UD UUTINE 5) ; resold 19. Dorsal head and alitrunk with scartered long bristle like hairs (hig. 34) (body mostly dark and non- metallic; eyes red; antenna Jegs.and broad pareh on dorsal gaster yellow-brown), 2... Euplecius sph Dorsal head und alitrunk with shorter liner fairs (dorsal alitvunk with distinenve avetaltic green ane yellow markings (Fig. 4%)), 280. & species inder 20. farsi 3-scumented (big 40); antennae S-senmented (Figs 4), 42); fore wing very browl (Fig, 39) (Trichograntiniatidue). .... Trichogrammd sy farsi S-seemented (Fig, 47); antennag 6-sepmenied (Higs 45, 46); fore wing narrower (Fiz, 43) (Aphel- inidac). -. GUnTRodara sy 21) Hues cayering occiput (pasterion purt of head )silver. grey, abdomet in dorsal view with anterior, medial and posterior parts black, latlurs! areas brown (Fis. FA) eee Winthemia lateralis (Macquart) Hairs covering ogviput salden brown, Ist seument ot wbdomen black, other segments blagk with parclics of silva (Pin. 49)... Beoriste Hayiveps Muaciuart ‘Treatment af species HYMENOPTERA bamily Beaconigies Cotesia urahae sp, nov, FIGS 2, 8, 0-12, (4 Malotwpe: F, ANIC, 8. Aust., Adelaide (Mitcham), reaied ex Uraha lugenson Kucalvpiys microdarpa, coll, 14...J985, emerged 3,xi.)985, G. R. Alten, Paratypes: 8 O, 20 o, same dala as holotype except some with different suburbs of Adelaide and dilferont dates, 179 9, orc, S. Aust, Adelaide (Waite Institue campus), various collecting and emergence dates during 1964, |. Hope, §¢ >,10, 5. Aust., National Park, Belair, coll. 20,x7,1964, emerged 25.41.1964, RDM. (59 9, Jar, ANIC: rest of material including 1% gold coated for SEM (Wings slide-mounted) in WAR ). Femate Length. 2.9 mm (2.53.2 nm, r= oVipositor. Colour, Body black; palps yellow: lugs cxcepr far coxa yellow-brown, hind leg infus cute distally, distal chd of hind tibia with dark patch; anterior pleural- sternal region of gaster dark red, wings liyaline, stigma cvenly and darkly pivsmented, as is rest of fore Woe venation, Heal. Mn anicrior view vertex arched su that head is somewhat circular; face and gena with lopewh White hairs and associated ting collieulaie sculp- turing; ther margins of eves siriually parallel: in dorsal view ovelli in wide triangle, posterior uangem of median ocellus coincident with imaginary line across {he anterior margins af laicral ocelli: frons and medial occiput siiooth and talrless; lemples with white hairs avid Yssociated collivulate aculp 10) including 172 A, D, AUSTIN & G. Ro ALLEN — Oe Figs \-7, 1-2, Fore and hind wings: |, Dotichogeniaea excalypul sp, Nov,, 222, Culesia urabae sp, nov., 3; 4, Lateral view of vaster uf Dolichoxenidea eucalypt spinov., 95 4, Distal tore wing of Mesockorus sp, 9) 3-6, Cusinaria atiera Jerman & Gaull, 9: 5, Lateral view of easier; 6, Fore wing: 7, Lateral view of gaster of Mesochors ap. 9, Scales: Figs 1, 2, 4.and 6 = 0.5 mm: Fie. 3 = 200 um; Figs Sand 7 = 1.5 mm, Abbreviations: a = areolet; pt = pterostigma, turing which ig slightly coarser than on face; vertex with few scattered short hairs otherwise smaoth; antennae slightly longer than body, distal 4-5 segments only slightly Jonger thar wide. Alitrunk. Seutum punctate with covering of shortish hairs, punctation denser along courses of notauli and along lateral margins (Fig. 8), punctation along posterior margin becoming slightly longitudinally elongate; courses of notauli faintly depressed, these faint depressions broadening posteriorly; scutellum faintly punctate; phragma of scutellum exposed along posterolateral margins (Fig. 8); propodeum PARASITOIDS OF URABA LUGENS 173 Figs 8-11, 8-9, Dorsal view of alitrunk; 8, Cotesia urabae sp. nov., 9; 9, Dolichogenidea eucalypti sp. nov., 9; 10-11, Cotesia urabae sp. nov., 9; 10, Dorsal view of gaster; 11, Lateral view of gaster, Seales: = 200 am, 174 ©, 1. AUSTIN & 6, BR ALLEN coarsely rugose to rugose-striate in anterior half, generally smoath with faint ragose punctation in posterior balf, medial longitudinal carina well developed with associated short horizontal and oblique carinae (Fig, 12); mesoplewron finely punctate in anterior half with associated short hairs, smooth and hairless in poslerior hall except for compact group af 5-6 foveae medially; metapleuron smooth antenorly, rugase-punctate posteriorly (Fig. 14); hind coxa faintly punctate on dorsal surface with associated short hairs, this sculpturing becoming coarser on ventral surface. Wings, Fore wing with veins c and 2-SR sharply angled, r slightly Jonger than 2-SRy cu-a almost striate; distal part of basal cell and anterior part of sub-basal cell devoid of hairs or almost 50; discal cell sparsely covered with hairs; hirid wing with yein r (spectral) present (Fig. 2), Gasier, T1 as wide as long, broadening posteriorly, surface in posterior two-thirds coarsely punctate to rugose-punctate, becoming coarsely striate jn posterior one-quarter (Fig. 10); sclerotized part of T2 rectangular, slightly wider than T1, coarsely Tugase-punctare With a few longitudinal striations, sculpturing fading lo nearly smooth in pastero- Jateral corners, longitudinal midline smooth; T3 usually rugose-punctate in anterior ane-quarter to two-chirds, with seattered hairs, posterior part smooth, in some specimens T3 virtually smooth throughout but always with al least anterior margin with band of punctation; rest of tergites smooth wilh scattered hairs; in lateral view hypopygiurt pointed, extending past posterior easter, with scat- tered fine Nairs, ventroapical margin not indented (Fig, 11); ovipositor sheaths with few apical hairs, Male As for female except lor length, 2.8 mm (2.6-3,1 mm, nm = 10) and sexual differences (genitalia and development of hypopygium), Qlther material exantined: S. Aust., suburbs of Adelaide. various dates and vollectars, 109 9, 16c 4 -WARI (excluded from type series because material is damaged or inadequately labelled), Camments: The sculptunng on the propodeum and TI-f3, the shape of these sclerites, the form of the hypopygium and ovipositer, clearly place (his species in Coresia Cameron. Colesia has previously been referred to as (he g/omeratis species-eroup of Apanteles s./. (see Mason 1981; Nixon 1965), and is the fatgest generic level taxon in the subfamily Microvastrinae, the latter comprising same 1300- 1500 described species world-wide (Mason 1981), Ju Australia Colesig ts common and diverse, but other than several species intfeduced trom Europe and North America as bjo-control agents tor certain lepidepteran pests (viz, G /lavipes Cameron, glomerala (L,) © kuzak (Telenga), C marginiyentris (Cresson), C plure/lae (Kurdjumay), C. rubecula (Marshall) and C. rufierus (Haliday)), che Australian Fauna rentains unstudied, A few poorly characterized species, which presently remain under the name Aparteles 3,1, May turn oul to belong in Cotesia, These species are unlikely to be conspecific with the present species as their type localijies are outside rhe known distribution of C urabge sp. nov. or they are asgociated with other hosts, There are no workable keys fo Indo-Australian species of Cotesia. However, the key in Nixon (L974) to the north-western European fauna can be used io separate C, wrubae sp, nov. fram four af the seven introduced species mentioned above. Of the other three species, © flavipes, is very different to C, urabae sp, noy. in that its body is strongly flatrened dorsoventrally (see Austin 1989), while C. kuzak and (~ rnarginiventris can only be identified reliably in association with their hosts, Helicoverpa spp. and Myrhitina coavecta (Walker), respectively, Biology; C. urabae sp, nov, is a solitary, primary eridoparasitoid and oviposits into early to inter- mediate larval instars of its host, emerging from intermediate to Jate instars before pupating. The pupal cocoon is alongside the host and is sulphur yellow-green with a surrounding silk matrix, This species has only been reared from t/ dugens. Dolichogenidea eacaly pti sp, noy. FIGS 1, 3, 9, 13, U5 Holarype: 9, ANIC, S. Aust, Adelaide (Highgate), reared ex Uraba lugens on Excalyptus camal- dulensis, collected 1,x7.1985, emerged 17.x7.1985, G, R, Allen, Paratypes; 18° 9, 20o°o same data as holotype except some with different suburbs of Adelaide and different dares; 139 9, Soo, S, Aust., Adelaide (Wajle Institute campus), various collecting and emergence dares during 1964, L- Hope (59 9, Sorcr, ANIC; rest of material including 19 gold coated for SEM (wines slide- mounted) in WARE). Female Length 2.9 mm (2.53.1 mm, 1 = 10) meluding OVIPASITOT, Cu/our, Body black, wings hyaline woh darkly pigmented venation, palps brown, legs black with distal Fore legs, tarsi of mid and hind legs and ubio- femoral joint yellow-brown. Head, Mostly smooth except for tine colliculate sculpluring associated with dense covering of short PARASITOIDS OF URABA LUGENS 175 Figs 12-15. 12-13, Propodeum and first and second tergites of gaster: 12, Cotesia urabae sp. nov., 9; 13, Dolichogenidea eucalypti sp. nov., Q; 14-15, Lateral view of alitrunk: 14, Cofesia urabae sp. nov., 9; 15, Dolichogenidea eucalypti sp. nov., 9. Scales: = 200 pm. 176 A.D AUSTIN & GR, ALLEN hairs; surface with Characteristic dull lustre; in antefior view inner margins af eyes virtually parallel; in dorsal view ovelli forming wide triangle, posterior rangent af median ocellus coincident with imaginary Line across the anterior margins of lateral weelli, fine colliculate sculpturing and assacialed pilosity slightly denser across vertex and occiput, excenl for smooth narrow band around posterior margins oF eyes; antennae reaching to posterior gaster or shghtly beyond, uistal three segments slightly longer than wide and sometirnes shghtly compressed. Alftrunk. Scutum coarsely punctate with punctures mastly closer to each other than their own diameter, except along posterior border and along. courses of notaull, which are thus faintly indicated (Fig. 9); scutellum smooth; scutum and sculellum densely cavered With short hairs; metanoium rather broad, an(erolateral margins emarginate so that phragma of scutellum well exposed; caringe forming propodeal arevla raised well above surface, carinae extending laterally belaw lyorlzontal midting, these carinac with dorsal and ventral extensions forming cristulae, but nat enclosing spiracles (Fig. 13); anterior part of propodeutn mastly smooth and setose, posterior part with faint rugose-puinctate sculpturing which becomes more obvious laterally; mesopleuron sefose in anterior half, smooth posteriorly; metapleuron mostly smooth, except for ventrapostertor one-third which is rugose-puncvrate (Fiz, 15); distal fore rarsus without spine opposed to tarsal claw: Wings. Fore wing venation as in Fig. t; costal and basal cells bare posteriorly; Kind wing broad; vein LSC+R deeply bowed; r present but faint; cubilellan cell moderately broad: submediellan cell minded posteriorly. Gaster. 1) as wide as long, slightly widened In posterior half, jateral margins slightly emarginale, surface mastly punctate, striale-punclate along lateral mains and stale in posterolateral corners and along posterior margin (Fig. 13); sclerotized part of T2 slightly wider than TL, 25 x wider than long, mostly smooth wih faint scattered punctures: TS slightly longer than T2 (14:11); T4-16 shorter than T2 (R11), T7 very short, about one-quarter length oF T2; 13-16 all smooth; T2-17 each with single transverse row of hairs; ovipositer and sheaths shorl, not extending far past posterior gasier: OVipositar with strong distal attenuation (Fig. 4}; hypopygium lacking obylous lateral creases though weakened normally im ventral midline: Male As for female except as follows: Length 2.7 mm (A528 mm, n = LW); alitrunk very slightly flanened dorsovenirally; fore wing stigma unpie- mented in medial area so that it is transparent: rest of wing venation generally with less prementation than female; TI sometimes with dense rugose- Pulictate sculpturing merging with posterior striations, otherwise same as female except for male genitalia and Lacking hypopygium, Comments: The sculptunng of the sculunmt and propodeum, shape of Tl and T2, form of the hypopygium, and shape and fringe of the vannal lobe of the hind wing clearly place this species in Dolichogenidea Viereck. Previously Delichogenidea was considered as three related species-graups in the genus Apanteles s./,, vic. he ultor, laevigains and lodpipalpis species-groups (see Mason 1981; Nixon 1965, 1967). This species falls into the witor group Which was revised by Nixon (1967) for the [ndo- Australian region, Jn this work 2 euculypit sp. nov. keys out to _D, cleo (Nixon) (couplel 23), a species Known only from India and associated with a nymphalid host Eriboae arja Felder, or with some difficulty it keys as D. cuniae (Wilkinson) (couplet 3b), which is known only from Java and associated with a limacodid, Cania bandura Moore (Austin 1987). Apart from having different hosts, these species differ from 2 evcalypti sp. nov. in that 2, voniae has an unusual striate seulpturing pattern on T) and T2, and D, cleo has the sub-basal cell evenly and darkly setose, the hind femur yellow in colour, the proximal half af the ovipositar very broad, and the mesopleuron coarsely rugose- punctate anteriorly. This is the fourth species of Dalichozenideu tecorded from Australia, tte others being DO Jipsis (Nixon) comb. noy., DB niris (Nixon) comb, noy. and Q tasmanica (Cameron) comb. navy. Examination of the holotypes of these species shows that they differ from & eucalypt sp, nov, in a number of important characters, All three have the oviposior much Jonger, being at least as long as the hind tibia, and, if addition, D, Jipsis and DB, tasmanice have a while spot ou the cheek and a much reduced propodeal areola. These species can be readily separated from 2. eucalypt sp, poy, using the key in Nixon (1967), Biology: D. eucalypli sp, nov, isa solitary, primary endoparasitoid which oviposits into early to intermediate larval instars of its host, emerging From inlermediate to late instars before pupating, The pupal cocoon is spun alongside the host and is white in colour and lacks a surrounding silk matrix, This species has only been reared from UO lugens. Family tehneumovidae Xanthopinpla rhopalaceras Krieger FIG. (fi PARASITOIDS OF URSHA LUGENS 37 This species (s easily identified by its distinctive colour pattern and fore wing venation. In South Australia it is Knows from two specimens (WARL),. one collected 4s an adult in the Adelaide region, the other reared from U. /vgeas at Keith, It is also known from Queenstind and Tasmania where it has been recorded from Ll /ugens, ard (he Lortricids Epiphyas postvittane (Walker) and MeropAves divulscina (Walker) (Brimblecombe 1962; Jumble. ton 1940; Gauld L984) also see Townes & Chui 1970), It is a solitary, primary endoparasitoid and emerges from dhe pupal stage of its frost. See Gauld (1984) fur additional taxonormie information and hst of synonyins. Parapivlax sp. FIGS t7, 19 Peraph ay is a large Old World genus with more than 50 recognized species from Australia, the majority of which are undeseribed (Guuld 1984), Where their biology is known Porgpiviax spp. have been recorded as primary and hyperparasitoids, mostly of lepidopteran hests, Apart from the charagters given in the key this species 14 notable in comparison to other parasitoids assactated willt UL /ugens for its relatively smooth unseulptured body {except for propodeal carinae) and lateral teeth on the propodeum, The species here belongs to the cormvax species-group (see Gauld 1984) and is only khown from the Adelaide region, where it has been reared as an obligare, solitary hyperparasiloid through ©. urabae and D. euculypti. Erthorus sp. FIGS 1%, 20 This is a distinctive species when compared to ihe other ichneumonids associated with UL, frees: In addition to the characters given in the key this species has distinctive reticulatc-puncrate sculp- (uring on the scutum, seutellum and propodeunt. Friborus sp. is 4 solitary primary parasitoid of L ingens, ovipositing into the larval Siages and emerging from the pupa. dels poly known from the Adelaide region and has been reared from its host On various occasions since 1965 (WARL, unpub- lished recorels), Casinaria micra Jerman & Gauld FIGS 5, 4 This species is a solitary, primary enduparasitoid eisily recognized by its lore wing venation, shor ovipasitor and colour, ft has been recarded trom ull states in Australia aod, although it has been most commonly associated with U fiers, C mucru has been reared [rom species belonging ro three other distantly related lepidopteran families — Geomet tidac, Qecuphoridae, Notodontid&e (see Gauld 184; Serman & Gauld 1988), The pupal cocoon is constructed near to (lerman & Gauld J98X) or underneath (observalions in this srudy) its dead larval hoscand is attached firmly to the jeaf surface Tt iS grey-browi and marked with characteristic black spo. See Jerman & Gauld (1988) for add- ional taxonomic information and list of synonyrnos, Mesochorus sp. FIGS 4,7 This is a large cosmopolitan venus of printary and hyperparasitoids of lepidopteran and colooys teran hosts; mos} Indo-Australian species are tindes- enibed (Gauld (984). The species recorded here Is a solitary obligate livperparasitoid of &. fugens through C micra, C. urahge and DB. enculyprti, and is known only from the Adelaide region. It is a pale coloured delicate species. with fine thread-like antennae, The male is distinetive in having the sxonosquama Of the genitalia extending from the posterior gaster as 4 pair of tony rods, Family Chaledidar Brochvinerta spp, PG, 22 This ts a large genus ui Australia with nearly 70 described species, the majority of whieh are primary. pupal parasitoids of Lepidoptera (Boutek 1988), The two specics meorded here (both unidentified) can be separated easily by the characters in the key. Spevies | is a solitary, primary parasitoid of GO. /daens and emerges. from the host pupa. Species 2 16 solitary and hyperparasitie through ©) urabee and 2. enealypri, thowah it ts noi known whether this relationship is facultative or obligatory. In eastern Australia Iwo species of Grachymeria have been reared from LU, /uvens, vie. 8. Jroggutti (Cameron) (Rrimblecambe 1962) and A. rubripes Girwalt (Campbell 1962) (B rbripes is considered a jubior synonym ot B fear (Watkerk: sec Boutck 1988). However, the matertal from these records Would have been identified ar atime when the Australian species in the gsenus were confused by most authors and hence might be misidentified, Phese numes should thus be used with some care, especially since Routek (1988) did not see any matetial in Australian collections or elsewhere revred trom Of fugens that he could assign. ro either 8: froggattt oc Bo teuta, Autravephalus sp. PGS 2b, 24 This spevies is fepecsetited here hy a site 178 A. D. AUSTIN & G, R, AIT EN 16 PAKASITOIDS OF URA#A LUGENS \7y Specimen reared as a primary parasitoid trom the pupa of UL lugers in the Adelaide region, Ti is easily distinguished by the characters in the key, There are more than 60 described Australian congeners which are discussed by Bouck (1988). Family Eurytomidae Fueyroma sp, FIGS 24, 27 (his is 4 large cosmopolitan genus with more thin 60 described Australian species (Boucek 1988). Biologically the group is very diverse including phytophagous species, primary parasitoids (mostly of tepidopteran hosts) and typerparasitoids, The species recorded here develops as either a primary Bregarious parasitoid of Ui /uvens, or as asolitary hyperparasitoid through (2 wrehee. Family Pleramalidac Pferonialus sp. FIG. 28 This specigs ts-relatively easily separated from other Chalcidoidea associated with U, lugens by its robust body and metallic green colour, The genus ig Gixonomically very complex and the Australian species are in need of revision (Bougek 1988). The species recorded here is an obligate, solitary hyperparasitoid reared from U, Jugens through Cc micra, C. wrafae and D, eucalypii in the Adelaide Tegion. Family Flasmidae Elasmus australfensis Girault FIGS 23, 25 A distinctive species recarded here as an obligate, solitary hyperparasitoid of UL fugens through C. auera, C. urabae and D: excalypti. Previously it was known to be hyperparasitic and occasionally gregatious through an unknown ichneumonid assuciated with CC /ugens in the ACT, and also fias been collected from N.S.W., and northern and southern Qid (type locality: Gordonyale, QId). This ty the first record of E. australiensis from §, Aust., indicating that it is probably distributed throughout south-eastern Australia. See Riek (1967) for addifional taxonomic information and list of synonyms. Figs tqy-28 Aeros sp oe Vetravephaliy ap. 25 22, Brachyieria sp. 2, Anravephalys sp. IG-15, Virst tergite of gaster: 16, Aanthapinpla rhapaloceroy Krieger, (97 1-20, Distal tore wings: 19, Paraphylie sp. Family Eulophidae Euplectrus sp. FIGS 36, 37 Eviplectrus is a cosmopolitan genus of preparious ectoparasiloids of lepidopteran larvae, Which 1s represented in Australia by 13 described species (Boutek 1988). The spevies recorded here is solitary and only known from the Adelaide region. altacks the early to intermediate larval stages, killing them before they pupate while pupating itself Undernesth the dead host. Pediobius sp. FIGS 33-33 This is a large, cosmopolitan genus of primary and hyperpurasitoid species that attack a wide range of insect groups (Boutek 1988). There are more than 30 described Australian species, mast of which do not have associated host information (Boutek 1988), The unidentified species recorded here is easily distinguished from other Hymenoptera associated with Uf lugens by its distinerive venation and sculpturing on the scutellum. Jt is represented by four specintens (WARI) reared in the Adelaide region from L. dusens and developed as a solitary hyperparasitoid through Evplecteus sp. and-as a gregarious hyperparasifoid through 1. excalypri. Fulophidae (genus & species indet.) FIG. 38 The two specimens reared as hyperparasitoids of O. lugenrs through D. eucalppti in the Adelaide région could not be identified to genus due to phe poor condition of ihe material, They are different from the other eulophids recorded here and can be distinguished by the dorsoventrally flattened body and distinctive colour pattern, Family Eupelmidac Anastalus sp, FIGS 29-3) This species is easily separated front the other chaleidoids associated with UL /ugens by its distinctive body shape and banded wings. The genus is cosmopolitan ard Known mainly as parasitoids of the eggs of Lepidoptera and Heteroptera, 17 Peraphvlis sp. 95 UR, V7 20, Erifarus sp. G. 21-23 Fore wires: 21, B28, Elasmus dustralieusis Giraull, GO: 24-25, Hine levs: 14, 25, Elaxemues australensis Gira, &. Scales: Tigs 16-19 = 250 pins bigs 20 aud 24 = 0.5 mm Pye 2) = ON maw Pig, 22 = 250 pans big, 23 = 200 yin; Fip, 28 = Loimm. Abbreviaiionss uy = cova t= hide, Ob = mureinal ver: pm = postMarginal veins R = radial val, st = Sstigmal vem; is = aisal seMINCAS, 180 “4D. AUSTIN & GR, ALLEN Pigs 26-35 26-27. Aurvieme yp, 4 26, Dorsal wew af atiorunk; 27, Pore wing; 28-29, Lateral view of body: 28, Pieramulies sp, 29, -bauwarus sp., 2) 30. Forge wing of Avfesfafis sp,, 27 31-33, Dorsal view of alitvunk: 31, Anasweiy sp. 7732. Eupelmidae, genus & species indet., oy 33, Pediobius sp., 9; 34, fore wing of Pediobivs yp., 2, 38, Hind feo of Peeiabins sp. %, Seales, Figs 26, 27, 29, 34 and 35 = 250 wm; Fig. 28 = 0.8 mm; Pin. 30. = 200 wan; Pigs 31-33 = 150 am. Abbreviations: mp = mesopleurony pm = postmurginal vem; ke = sculums: xl = scurellumy st = stigma) veing is = tarsal segments. PARASITOIDS OF URABA LUGENS RI a felt { rN | | bigs 36-42. 36-37, Euplectrus sp., 97 36, Dorsal view af head and atitrunk; 37, Pore wing: 38; Dorsal view af alitrunk of Eulophidae, genus & species indel., 2 (stippling indicating colour pattern); 39-42, Trivhogrammnia sp. 39, Pore and hind wings, a; 40, Hind leg, 9; 41, o antenna; 42, 9 antenna. Scales: Figs 36 and 37 = 250) yim; Pigs 38 and 39 = 150 pin, Fig. 40 = 100 um; Pigs 41 and 42 = 50 #m (same seale line). Abbreviations: pm = postmarsinal vehi, st = atigmal yein; ts = larsal seoments, although a few are recorded as hyperparasitoids through bravonids, Anestatus is represented in Australia by 40 described species (Bouéek 1988), The species recorded here is represented by three specimens reared as a gregarious hyperparasitoid through © wrabae and D. eucalypti in the Adelaide region. Eupelmidae (genus & species indet,) FIG. 32 A single male specimen reared as a hyperpara- sitoid of Uo lugens through CG urabue in the Adelaide region could not be identified 1a genus, but it is not Anastatus sp., from which it van be separated by the characters in the key. Family Trichogrammatidae Trichogramma sp. FIGS 39-42 The members of this cosmopolitan genus are 1k? A.D, AUSTIN & GR. ALT EN Figs 43-49. 48-47, Centradora spc 43, Fore wing, 7-4, Dorsal view af whole body, Fs 44) 2 antenna, 46, cr antenna: 47, Hind lee, 9:48, Abdumnest of Mlthentiatoveraly. 9) (Miaxcquartk: 49, Dorsal siew of Peoresta flaviceps Macquart, 2 (stippima showing colour patiern in Figuees46 and 47). Seales: Fig. 43 = 200 jum: Fig. 44 = 300 yan, Fieg 45-47 = 50 aim (same scate tine for Figs 43 and 16); Figs 48 and 44 = Lijmm, Abbreviation: 1s = tarsal segments, obligate, primary parasitoids of insec! eggs, most frequently those of Lepidoptera. [t is the anly egg, parasitvid of U. lugens so far recorded, and it can be identified by the characters in the key, as well ay its minute size, dislinctive lore wing venation, fore wing setal pallern, and very narrow. hind wing. This species is a solitary parasitoid known only front the Adelaide region, We. have not seen material of the Jrichograrmima sp, reared [rom U. lugeris in the Murray Valley in N.SW. (Campbell 1962), which may be the same species to that recorded here, Tamily Aphelinidae Centrodort sp. FIGS 43-47 This is a cosmopolitan genus of about 40 deseribed species (Llayar 1983), most of which are primary parasitoids of the eggs af Orthoptera and Homoptera, although at least one Species is reported to be hyperparasitic (Gordh 1979; also see Viggiani 1984), The species recorded here is an obligate, gregarious hyperparasitoid of U fugens PARASITOIDS OF URAHA LUGENS 1s3 through C. urahae or 2 eucalvpti. Apart from the characters in the key and the life history stage attacked, Cenfrodora sp. can be separated From the other parasitoids associated with UL lugens by is minute size, wing venation, and ovipasiter which 1% more than half the length of the guster {the oviposiior being significantly less than half the gastral length in Trichogramma sp.). DIPTERA Family Tachinidae Winthenvia lateralis (Macquart) FIG. 48 Previously this species has heen collected at various localities in all states of Australia. It has been reared from host species belonging to seven lepidopteran: families, viz. Arctlidas, Noctuidae, Pieridae, Notodontidae, Nymphalidag, Saturnidac and Geometridae (Crosskey 1973; Cantrell 1986, 1989), The only record trom U fugens is from specimens if this: study (3 specimens, Waite Institute campus, J. Cobbinah, 1975, WARI). 4 tuteralis oviposits onto the external surface of a hast larva. Alter hatching the fly larva penetrates the host larva and usually emerges from the host pupa, alchough we were not able to confirm its’ biology in this study, Set Crosskey (1973) and Cantrell (1986, F989) For additional taxonomic information and list of synonyins. Exortsla flaviceps Macquart FIG. 49 This species has been recorded from all states of Australia and the N.T, (Cantrell 1985), and has been reared from members Of nine lepiduptcran families, Vaz. Lymeanimidas, Anithelidac, Pieridae, Aguris- lidac, Sphingidac, Genmetridae, Notodontidae, Lasiocampidae and Noctuidae (Crosskey 1973; Cantrell 1986), Lt ovipeastts unto the surface of U. fugens laryac. After hatching the fly larva burrows into che lost ro feed and develop internally, finally emerging from late larval instars fo pupate outside the dead host. The colour pattern on the abdomen and occiput of the head ts the easiest way (0 clis- tinguish (his species trom Ff laferalis. See Caitcell (1985) for additional taxonomic information and list_of synonyms Other Parasitnids trom LU. /ugens in Queensland Brimblecombe (1962) reared (wo ichneumonord species Which have not been recorded in S. Aust, viz. Jrebuthe sp. (Ichneumonidae) and Camprloneure sp. (Bracon- tac). Also Gauld (1984) reports the following ichneumonids as having been reared trom U dugens: Stiromesostenus spp., Campoplex sp. and Pristomerus sp,, but again, they were not reared during this study and may: not be found in S, Aust. Acknowledgments We thank Dr lan Naumann (Australian Natiunal Insect Collection), Mr Geatf Hollowsy (Australian Museum) ail Dr Bryan Cantrell (Qucenstand Deparunent of Primary Industries) for assistance with identifications: Mr Pan! Dangerfield for the line drawings and scanning micrographs, and Dr Mike Keller for reading a draft of the manuscript. This: work was supporled by a Commonwealth Postgraduate Research Award to GRA and a grant from the Australian Biological Resources Study Participatory program to ADA, Refercnees AvsTIS. A, DO ISB7) A review ef the Braconidae (Hymenoptera) thal parasitize Limacodidae in South- tam Asia, particularly those associated with coca and ail palm. pp. 139-164, fn Covk, M-. 1. W., Godfray, H.C. J, & Holloway, J. D, (Eds), “Slug and Nettle Caterpillars. The Biology, Taxonomy and Control of the Limucodidac of Feonomic Importance on Palas in South-east Asia”. (CAB International, Wallingford. (1989) The taxonomy of New World inigragastring diavonids (Hymenoptera) parasitic on Ditereaed spp. (Lepidoptera; Pyralidae), Bull. ent. Res. 7, 131-144, Brimaceeomar, ALR. (1962) Ourhreaks of the eucalypt leah skelotoniver. Qf J. aurte. Sef, 19, 209-217, Bourtk, 4. (1988) “Ausinuasitn Chatcidoides ¢Hymen- optera}, A Dosystematic revision of venera af fourteen lurmilics, with @ teclussificatlon of species”. (CARB International, Wallinwfard). CAMPUELL. K. CG. (1962) The biology of Raesetia lugens {Walk.), the guntleat skeletonizer moth, with particular reference to the Eucalyptus cumakdulensis Dehn, (river red guin) forests of the Murray Valley Region, Mroc, Tainn, Boe. NS 87, W6-338, CANTRELL, B, K. (1985) A revision of rhe Australian species of Exorivia Meigen, with notes on the other acnera Of Austratian Rxoristim (Diplent: Tactinidaey, Aust, J. Zval. 33, 547-576, (1986) An updated host catalogue for the Australian Tachinidae (Diptera). /, Aust, ent, Soc, 25, 255-265. — —_ (1989) The Australasian species at Rartemia Robinew-Desvoidy (Diptera: Tachinidae) with neites on Oceanian and Ociental specivy. Met 28, 93 10. CORBINAU, cL R- (1983) Sintability of various euvalypts for frowth of (he larvae ot Crater dasiens (Walk), bor Leal, Manag, 6, 361\-366, Crossxey, Re W. (1973) A comspectas of the Tachninidag (Diptera) of Australia, imeliding keys to The Supraspecific (AXA ahd TaxonOmiv and host catulewucs, Bull, Br Mus. far, Hist, (Ent.) 24, 1-221 Domeleron, 1. 1. (L940) Tarirex pasrvitrana Walk. and 184 A.D, AUSTIN & GR, ALLEN its parasites in Australia. N.Z. J. Set. Technol. (A). 21, 322-327. Eapy. R. D. (1968) Some illustrations of microsculpture in the Hymenoptera. Proc. R. ent, Sac. Lond., Ser. A. 43, 66-72. Gacip, 1, D. (1984) “An introduction to che Ichneumonidae of Australia”. (British Museum (Natural History), London), Gorpy, G. (1979) Superfamily Chalcidoidea. pp. 743-1043, In Krombein, K. V., Hurd Jr, P. D., Smith, D. R. & Burks, B. D. (Eds), “Catalog of Hymenoptera in America North of Mexico, I°. (Smithsonian Institution Press, Washington, D.C.). Harris, J. A. (1974) The gum leaf skeletonizer Uraba fugens in Victoria. For, Tech, Paper, For. Common. Vie. 21, 12-18. Harris, R. A. (1979) A glossary of surface sculpturing, Calif. Dep. Food Agric, Bur, Ent. Occas. Pap. 28, 1-31, Hayat, M- (1983) The genera of Aphelinidae (Hymenoptera) of the world. Syst. Ent, 8, 63-102, Jerman, E. J. & GauLp, [. (1988) Casinaria, a paraphyletic ichneumonid genus (Hymenoptera), and a revision of the Australian species. J. Naf. Hist. 22, $89-609. Mason, W. R. M. (1981) The polyphyletic nature of Apanteles Foerster (Hymenoptera: Braconidae); a phytogeny and reclassification uf Microgastrinae. Mem. ent. Soc. Can. 15, 1-147. (1986) Standard drawing conventions and definitions for venational and other features of wings of Hymenoptera. Proc. ent. Soc, Wash, 88, I-7. Moraan, F. D, & Caueinan, £ R, (1977) Oviposition and establishment of Uraba lugens (Walker), the gum leaf skeletoniser. Aust. For, 40, 44-55. Nixon, G, E. J. (1965) A reclassification of the Tribe Microgasterini (Hymenoptera: Braconidae). Aull. Ar Mus. nat. Hist. (Eint.) 2, 1-284. (1967) The Indo-Australian species of the ultor- group of Apanteles Forster (Hymenoptera: Braconidac). thiad. 21, 1-34. (1974) A revision of the north-western European species of the glomerarus-group of Apanteles Férster (Hymenoptera, Braconidae). Bull. ent. Res. 64, 453-524. Riek, E. F. (1967) Australian Hymenoptera Chalcidoidea Family Eulophidae, Subfamily Elasminae. Aust, 2 Zool. 15, 145-199, STRELEIN, G. J. (1988) Gum leaf skeletoniser moth. Uraba tugens, in the forests of Western Australia, Aust. For. 31, 197-204, Townes, H, & Cut, $-C. (970) The Indo-Australian species of Xanthopimpla (Ichneumonidae). Mem. Amer, ent. Inst. 14, 1-372. Turner, A. J. (1944) A revision of the Australian Nolidac (Lepidoptera). Proc. R. Soc. Old 55, 13-50. Van ACHTERBERG, ©. (1979) A revision of the Subfamily Zclinae auct. (Hym., Braconidae). Tijdschr Ent. 122, 241-479, VigGiAnt, G. (1984) Bionomics of the Aphelinidae. Ann, Rey, Ent, 29, 257-276. CONTRIBUTIONS TO THE TAXONOMY OF AUSTRALIAN BUPRESTIDAE (COLEOPTERA): NEW SPECIES OF ASTRAEUS AND STIGMODERA (CASTIARINA) AND A KEY TO ASTRAEUS (S.S.) BY S. BARKER* Summary Astraeus meyricki Blackburn is resurrected from synonymy, redescribed and illustrated. Astraeus badeni van de Poll is redescribed and illustrated and the following five new species of Astraeus (sensu stricto) are described: A. aridus sp. nov., A. occidentalis sp. nov., A. princeps sp. nov., A. williamsi sp. nov., A. yarrattensis sp. nov. A revised key to Astraeus (s.s.) is provided. The egg of A. pygmaeus van de Poll is illustrated and some biological information given. Two new species of Stigmodera (Castiarina) are described: S. garnettensis sp. nov., S. wellsae sp. nov.; the first has sexually dimorphic colour pattern, rare in the sub-genus. KEY WORDS: New species, Astraeus, Stigmodera (Castiarina), Coleoptera, Buprestidae. Transactions of (he Raval Society af S, Ause, (989), TL3(4) 185-184, CONTRIBUTIONS TO THE TAXONOMY OF AUSTRALIAN BUPRESTIDAE (COLEOPTERA): NEW SPECIES OF ASTRAEUS AND STIGMODERA (CASTIARINA) AND A KEY TO ASTRAEUS (s.s,) by &. BARKER* Summary Barker, 5. (1989) Contribulions to |he taxonomy of Australian Buprestidue (Coleoptera): New species of Axrraeus and Sigmodera (Castiarina) and a key to Asrraeus (s.8.), Trans. R, Nae, 8, Aus 134), TRS-194, 30 November, 1989 Asiraéus meyvrick? Blackburn is resurrected from synonymy, redescribed and illustrated. Astraeus budent van dé Poll is redeseribect and illustrated and the following five new species of srraeus (sens sfricto) are described; 4. an/dus sp. nov., A. eeeldentalis sp. nov, As prineeps sp. nov. AL williams’ sp. Way, A. Jarrattensis sp, noy, A revised key lo Asirueus (5.4) is provided, The egg of 1, pugmeeus van de Poll is illustrated and some biological information given. Two new species of Sriginadera (Castiarinia) are described, J S. gornettensis sp, nov,, 5. wellsue sp. nov; the finest bas sexually dimorphic colour pattern, rare in the sub-gerius, Koy Worps: New species, Aatraeus, Sigmodera (Custiarina) Coleoptera, Buprestidae. Introduetion The buprestid genus sfraevs was revised by Barker (1975) and additional species described by Barker (1977). No further information has been published. Fresh material has now come to hand and five new species are described herein including one species from the Flinders Ra., S. Aust., which was misidentified in the revision of Barker (1975). The separate Stalus of A. baden*yan de Poll and A, meyricki Blackburn has been in doubt since they were deseribed, A re-examination of the specimens available has convinced me that they are separate species. The key to Asyraeus (s,s,) given by Barker (1975) requires Updating to include three new species published by Barker (1977), five new species described herein and one species resurrected [rom synonymy. A new key is presented and new lovality records of two species recorded. The egg of A. pygmaeus van de Poll is illustrated, the first such observalion of an Asirdeus species, together with biological information on A. pygmaeus. Two new species. of Srigmodera (Castiarina) are described and illustrated; one being unusual in showing sexual dimorphism in colour pattern, a rare occurrence in this sub-genus, Materials and Methods Male genitalia were prepared and displayed by the method deseribed by Barker (1987). Specimens for S.E.M, were dehydrated prior to being coated with gold-palladium, Abbreviations used in (he text for museum and private collections following Wall * Department of Zoology Liniversity of Adelaide GPO. Box 498 Adeliuide, S. Aust. S001 (1979) are: BMNH_ British Museum (Natural History), London; JHQA Mr J. Hasenpusch. Innisfail; MNHN Museum National d'Histoire Naturelle, Paris) NMVA Museums of Victoria, Melbourne; SAMA South Australian Museuni, Adelaides WAMA Western Australian Museum, Perth GWNA Mr G. Williams, Landsdowne, GWOA Mr G. Wood, Atherton, Resurrection of Astracus nreyricks, Van de Poll (1892) considered A. mepricki Blackburn a synonym of 4, hadeni van de Poll but Blackburn (1895) disagreed, Barker (1975) followed van de Poll, | have re-examined the two male syntypes of A. meyricki held respectively in the BMNH and SAMA collections and compared them with specimens in the SAMA identified by comparison with the female holotype of A, badent held by the MNHN, The pronotum of 4, badeni is Widest medially and dilters trom A. meyrickl which is widest basally, Male genitalia of A, meyricki are broader than those of 4. baderi and are rounded more abruptly al the apices (Figs 1, JE). | conclude that 4, mevricki{Pig. 2B) isa good species and hereby resurrect it from the synonymy of A, badeni (Pig, 2A), Both species are redescribed here. Astnreus (Astraeus) williaims/ sp. n0y. FIGS 1A, 2D Holotype: &, Peninsular Rd, Lansdowne, N.SW,, 10.45.1987, G, Williams, SAMA 1 28219. Allutype 9, same data as holotype, SAMA 121220. Purarypes) NSW. S oc, Peninsular Re, 186 S. BARKER G 1mm Fig. 1, Photomicrographs of male genitalia of the following Astraeus (Astraeus) species: A, Astraeus williams sp. nov., B. A, aridus sp. nov., C. A. meyricki Blackburn, D, A. occidentalis sp. nov, E, A. badeni van de Poll, F. A. yarrattensis sp. nov,, and the following Stigmodera (Castiarina) species: G. Stigmodera garnettensis sp. nov., H. S. vallisi Deuquet, I. S. wellsae sp. nov. Lansdowne, 17.xi.1980, 23.x.1980, 30.x.1980, 22.x.1982, 28.x.1988, G. Williams, GWNA. Colour: Head blue-green or blue. Antennae, segments: 1-3 coppery; 4-11 blue, Pronotum blue laterally, purple medially. Elytra dark blue, each elytron with following yellow markings: basal spot; pre-medial fascia represented by mark over humeral fold and spot near suture; post-medial fascia touching margin but not reaching suture; small pre- apical spot. Ventral surface dark blue. Legs: femora dark blue; tibiae mostly dark blue, distal tip testaceous; Ist tarsomere mostly testaceous, distal tip blue, 2nd tarsomere mostly testaceous, distal tip blue, 3-4 tarsomeres blue. Hairs silver. Shape and sculpture. Head closely punctured, without median keel, hairy. Pronotum closely punctured; laterally parallel-sided at base, rounded posteromedially to apex, hairy. Elytra costate, intervals flat and smooth each with basal row of punctures; laterally parallel-sided basally, rounded posteromedially and tapered to sharp marginal spine; sutural spine sharp, rounded inner margin, humeral fold moderately developed, angled (vide Barker 1975 Fig. 1B). Ventral surface shallowly punctured, moderately hairy, hairs short, Size. Males, 7.5 + O18 x 2.9 + 0.08 mm (6). Females, 7.9 x 3.2 mm (2). Male genitalia: (Fig. 1A). Parameres parallel-sided from basal piece, widened medially, tapered, rounded and tapered to apex, Apophysis of basal piece broad, rounded apically. TAXONOMY OF AUSTRALIAN BUPRESTIDAE 187 Fig. 2. Habitus illustrations of the following species: A. Astraeus badeni van de Poll, B. A. meyricki Blackburn, C. A, aridus sp. nov., D. A. williamsi sp. nov., E. A. occidentalis sp. nov., FA, yarrattensis sp. nov., G. A. princeps sp, nov., H. Stigmodera wellsae sp. nov. {88 S BARKER Remarks; Named after Mr Go Williams, Latis- downe. Asiraeus (Astraetis) yarrattensis sp, nov, FICIS IF, 2 Holomwpe o, VYarratt State Forest, NE Wingham, N.SAW., TEXLI98L, G. Williams, SAMA 1 2122), Allolype: @, 10 km SSW Laurieton, N.SW,, 6.xi, 1980, G. Willian, SAMA 1 21222, Paratvpes: NSW. | oct, same data as holotype, GWNA3 oo, Yarratt State Forest, NE Wingham, ZU.xPYBL, 16.01.1982, GC) Willais, GHNAS 1 cr, Bom Bor State Foresi, SE Grattan, 14,%0,1984, G. Wilhains, GWNAs TO. Mitwhells* Island, (Factory Rd), & Taree, 9.xi,1988, G. Willians, GWNA. Colour, Male, Head blue-green, dark blue basally with purple reflections. Antennae, segments: I-3 coppery; 4-11 blue, Pronotum dark blue with purple reflections, blue at lateral and basal margins. Female. Head and pronotum dark blue, rest as in male, Elytra dark blue with following yellow markings On each elytron: large basal spot) pre- medial fascia covering tumeral fold bul not reaching suture; post-medial fascia covering margin but not reaching suture. Ventral surface blue. Legs: femora bluc; tibiae blue at proximal end, rest (eslaceous; Ist larsofiere lestaceouss 2-4 tarsomeres blue, Hairs silver Shape and xculplure: Wead closely punctured, glabrous median keel, hairy, Pronoetum closely punctured; laterally parallel-sided basally, rounded posteromedially, tapered to apex; median impressed line from (iiddle to apical margin, laterally hairy. Elyira costal, intervals flat, basally wrinkled, each with row of punctations; laterally tapered from base, rounded posteromedially, tapered to sharp marginal spine; sharp surural spine, rounded inner margins humeral fold well developed and angled. Ventral surface closely punetured, moderately hairy, hairs medium length, Sizes Males, 6.9 x 2.9 om (6). Females, 74 x 31 mim (2), Male genituliar (Fig. We) Parameres angled outwards fron basal piece, more or less parallel- sided, rounded to apes Apophysis of basal piece broad, rounded apically. Remarks: Named after che wpe localicy. Astraeus (Astraeus) aridus sp, noy, FIGS. 1B, 2C Astrueus badeni: Barker 1975, p 131, Holowpe o, Puttapa Gap, 21 km 8 Copley, 8. Aust., Flinders Ra., 21.x.1971, §. Burker, SAMA I 21223, Allotype: 9, same data as holotype, SAMA | 21224. Paratypes: §, Aust; 6 co, 8 9 Q, same data as holotype SAMA; 8 vo, 6 99, Derna Pass, 25.x.1969, N. MeFurlund, SAMA, Colour Head, antennae, pronotam black with blue reflections. Elytra black with blue reflections and following yellow markings: basal spot; fascia covering humeral fold, not reaching sulure, concave anteriorly, often represented by two spots; short post-medial fascia touching margin but not suture, coneave posteriorly, Ventral surface and legs black with purple reflections, Hairs silver. Shape and sculpture’ Head closely punctured, na median keel, hairy. Pronotum closely punctured: laterally rounded and narrowed from base to apex; hairy. Elytra costate, intervals flat, each with row of punctures; larerally parallel-sided basally, rounded ‘posteromedially (o marginal spine; both spines sharp, sutural spine with rounded inner margin, humeral fold poorly developed, slightly angled, Ventral surface shallowly punctured, moderately hairy, hairs medium length. Size: Males, 8.9 + 0.10 x 3.5 + 0.07 mm (15). Fenvales, 9.1 4 0.14% 3.6 + 0.07 mm (15). Male genitalia: (Fig, 1B), Parameres angled oulwards Irom basal piece, rounded anteromedially to apex. Apophysis of basal piece medium width, rounded apically, Remarks: All specimens were collected on the leaves and stems of Melaleuca glomerata F, Muell. from the same locality, Derna Pass is a synonym ol Pullapa Gap, The specific name is derived from aridus L.,. dry. Astraeus (Astracus) badeni yan de Poll FIGS 16, 2A Asiraeus badeni van de Poll 1889: RA, 93, 94, pl, 2, Figs 7, 7a, Blackburn 1891; 496; Kerremans 1892: 101; van de Poll 1892; 67; Blackburn I895: 45, 46; Kervemans 1903: a8; Carter 1929 282; Obenberger 1930: 365; Barker 1975; 13h. Asirueus budeni dixjunctus Qbenberger 1928: 204; 1930: 465; Barker 1975: 130, Astraeus meyricki Blackburn 1890; 1256, 1257; van de Pall 1892; 67; Kerremans 1892; 101; Kerremans 1903: 148. Carter 1929; 282; Obenberger 1930; 365; Barker 1975: (31, IAXONOMY OF AUSTRALIA. BUPRESTDAL 189 Tiper Holotype 9, Gawler, MNHN, Colour: Head, antennae, pronotum black with blue and purple reflections. Elytra black wih purple reflections and following yellow markings on each elytran: basal spot; pre-tnedial fascia from marain over humeral callus, not reaching suture, concave anteriorly; short post-medial fascia from margin not reaching suture, concave posteriorly: small pre- apical spot. Ventral! surface black with bronze and purple reflections. Legs black with blue and purple reflections. Hairs silver. Shape and sculpture: Head closely punctured, no median keel, hairy. Pronotum closely punctured; laterally parullel-sided basally, rounded, indented to apex, hairy, Elytra costate, intervals flat each with row Of punctures; laterally parallel-sided basally, rounded posteromedially, abruptly tapered to sharp nrarginal sping; sharp surural sping, rounded inner nrargin, huniensl Sold moderately developed, angled. Vential surface shallowly punctured, mauderately hairy, pairs long, Size! Males, 7.9 * 3.4 mm (6). Females, 8.4 x 3.6 mim (8). Mate genitalia: (Fig, 1E). Parameres angled outwards from basal piece for most of length, rounded to apex, Apophysis of basal piece medium Widtl, rounded apically. Distribution: §. Aust. Gawlet, Morgan. Murray Bridge, Nuriootpa, Vics Inglewood. W,A,; Marvel Loch, Southern Cross, Remarks: A, baeleni differs from congeners in the bulbous appearance of the pronotum and the more abruptly tapering apices of the elytra. Ip is a short broad species. A specimen in the SAMA calleeted by Tepper at Nuriootpa, S. Aust. has ‘ou Culfitris” oo the label, the only specimen scen with w recorded Hust pln, Astroens (Astracun) mevricki Mackburn FIGS 1c, 2B ANeraous Meveiek: Blackburn, 1890: 1256, 1297) 1K95) 45, ah Tipe Syntype o, no data, BMNIT syriype er, W, Auswalia, SAMA, Colour Head, aitennae, pronutim black with purple rellections, Blyrra black with following yellow marks on each elytron: stall basalspot; pre- medial fascia covering humeral fold, not reachivig suture; post-medial Mscua not reaching mann ur suture, Ventral surface black with purple reflections. Leys dark brown with purple reflections. Halrs silver. Shape gnd sculpture; Head closcly punctured, shallow basal median sulcus, without median keel, hairy, Pronotum closely punctured, with prominent basal fovea, median glabrous line from base Lo middle; laterally rounded from buse in apes, Elytra costate, intervals flat, each with row of punctures; Jatcrally tapered from base, rounded pusteromedi- ally then tapered to sharp marginal spine; sharp sulural spine, rounded mner margin; humeral fold very poorly developed, very slightly angled. Ventral surface shallowly punctured, moderately haicy, Hairs medium length. Size; Male, 10.6% 4.5 mim (2). Male genitalia: (Vig, IC). Paramens angled outwards from basal piece, rounded te apex. Apophysis of basal piece medium width, rounded apically, Remarks: There is a temale spectinen lodged in the SAMA possibly belonging to this species which ! identified as A, badeni (Barker 1975 p. 133). It was collected 18 km SW Three Springs, W.A,, on Dryandra cirisoides, 8.41.1968, NO Me Farland, Astraeus (Asiraeus.) princveps so. nov PIG. 2G Rololype: 9. Prince of Wales Island, Qid, Feh 1939, SAMA | 21225. Paratype: Qld: |, same data as holotype SAMA. Colour; Head black with blue reflections and yellow frontal spot. Antennae black with blue reflections. Pronotum black with blue reflections and lnree lateral yellow murks, Flytra black with blue teflections and following yellow. marks on each elytron: broad pre-medial fascia covering margin but not reaching suture, obliquely angled anteriorly: broad post-medial fascia covering margin bur nol teaching suture, Veutral surtace dark blue, Legs red brown, Hairs silver, Shape and sculpture: Head shallowly punctured, no median keel, few shart hairs. Pronotum shallowly punctured, laterally rounded from base to. apex, widest medially, few short hairs. Elytra costate, Intervals Mat, each with row of punctures, surface wrinkled basally; laterally tapered trom base, rounded postcromediatly, tapered to sharp marginal Tt & BARKER spine; Sharp sutural spine, inner margin straight and cured upward, with extended marginal im(erval between spines; humeral fold absent. Ventral surface shallawly punctured, moderately hairy, hairs medium length. Size: Females, 7.9 x 3.4 mm (2). Remarks: Roth specimens were given jo Mr §. Watkins by Robert G. Wind, insect supply dealer and tesident-of Monterey, California. His Australian wite had been stationed on Prince of Wales [sland priocto World War Tl and presumably she was the collector. They are unique to the sub-genus due to the coloration on the head and pronotum. The dorsal coloration may have been pink or red in fife and now faded, The name is derived from princeps. L., leader. Astracus (Astracus) occidentalis. sp. wow FIGS TD, 2b Hotaiype: cr, 99 km S Overland Roadhouse, Carnarvon Hway, WiA,, 25.1x,1980, on Casuarina dielsiany, 5, Barker, SAMA, | 21226 Allotvpe: 24227 G. same dala as holotype, SAMA, | Paratypes: W.A., 2 ors, 1 G, WAMA, SAMA. Culow: Head. amennue, pronutum voppery-brornze. Elytra blaek with blue and purple reflections and allowing yellow markings on each elytron: basal spot; pre-medial fascia covering humeral fold, Louching basal margin bur not reaching suture, concave anteriorly; medial spot on margin and one near suture fi some specimens; large post-medial Spot near margin; pre-upical spot neat suture. Hairs silver. Shape and seulptures Head closely punctured, thin medial imipressec line from base, continued as.short glabrous median keel af apex. hairy. Propotum closely punctured; prominent basal fovea projecting anteriorly for short distance as impressed ling, anterior murgin projecting medially; laerully rounded and narrowed trom base to apex, hairy. klyira costate, intervals fat, each with row af punctures; laterally parallel-sided basally, rounded medially and narrowed to small marginal spine; slurp sulural spine. rounded inner margin; humeral old poorly developed, slightly angled. Ventrul surface shallowly puncrured, moderately hairy, hairy lone. Size: Males, 11,2 x-4.4 mm (3), Females. 11,8 ¥ 4.6 mm (2). Male genitalia: (Fig, 1D), Parameres. angled ounvards from basal piece, shghtly rounded medially, rounded abruptly to apex. Apophysis of basal picee broad, rounded apically, Remarks: The name is derived from occidentalis U.., western. Key to the species of Astraeus (sensu stricta) 1. Fleud with median keel... eeeeee CITT Head without median keel, ...,,----------- 22 2) Hats silver... ee ee eee eee eee 3 Hairs tyellow. 22. 00. eee ee bet }\. Part or all of anterior ventral surtace Ted-brown None of anterior ‘Ventral surface red-brown... - 7 4. Gulat prosternum, mesa- und metasternum, coxae 2-3, ubdominal segment 1 red-brown... 1.0.0... a. A. bekeri Barker Less of vertical’ surface red browat oe iter od 5, Prosternum, coxae red-brown, , hoster ys (eect et On Bett tetie 6 6. Shorter than 7.5 ni, humeral told well developed, ucutely angled . sda! «IN deur A. fraseriensis Barker longer than 7.5 mm, humeril fold moderately developed, aneled..,,......, 24, vbscurus Barket 7. Most or part of lew jestaccous - HEE ela None of ley testaceous - 54 tt 14 R.. Legs 1-2 festaceous except for onirer fiarin. af femora, lew 3 lestaceous except for femur, ....-, erdQesd eae -- cee i eg A. dilufipes vau re Pol} 1 ess of leg than above testacequys.,---.. ----.-- vf] 9. Tibiac, tarsomeres 1-2 restaceauy.. 2. ee ee OSE aie Al sme Barker Less of lez than above testaceous. ptt _- Mt 10. Tibiae testaceous, distal tips blue, tarsomere | testuveaus . AL J artittensis sp. Noy, Less of Jeg than above lestaceous..-- == il Il, Distal tips of tibiae tesraceous; tarsomeres l-2 (estaceous, distal tips blue... A. willie sp. nov. Less Of Teg thin above [ebtaceous eer | J 12. Hips of tibiae and jarsomere { tesiueeouy,..... 13 Tursomere |} testa, gents ett lp AAAS olm AL notraneeeas/]s Barked 13, Usually shortee than Fo MM, ., 0. ecs sce eens saat AL pereeus van de Poll Usually fohget ‘Than 7mm. ......-5, id, Eluieral fold well developed, aculely angled... AL enasterst. Macleay Humeral fold ‘well developed, umyled......-- pee Pict oh samonelli Saunders - Fumera! fold moderately developed, angled... 16 Tumeral fold poorly developed, slightly anvled- W tn TAXONOMY O)} AUSTRALIAN BUPRESTIDAE 191 16. Head green and coppery purple or blue-green. . 17 Head black......... da elon t t29Lanucraty 18 17. Head, pronotum green and eoppery purple... , peeeesy tet, _. A. intricatus Carter Head blue- -green, pronotum with medial, cordiform purple mark, anteriorly green, laterally blue... -. adi nt tqmpape's -AL blackdownsensis Barker 18, Browd, rounded gpecies Aa bio A, vlobosus Barker Elongute species............,, 4. watsoni Barker 19. Body blue; elyira with 2 “yellow fasc1d. ii. eee ejetleg @49 A. “Fruteresihus van de Poll Body black or coppery-bronze....., 22... --20 20, Body black; elytra with numerous yellow spots. . axe : oA. crassus van de Poll Body coppery- bronze: each elytron with yellow fascia and 4-5 yellow spots. ... .4. occidentalis sp. nov. 21. Elytra with 3 yellow fuscia, red areas. ___- CCOOP Mme eeisc em Ets Ea eae A, major Blackburn Elytra with 2 1 yellow Taisen: no red, i) De Aan od. Mavate his (Tv homEony 22. Body elongate and cylindrical... ... dae Rela apd Body teardrop shaped... ...... cep eens 24 23, Pronotum conically elevated medially. 600199 mene tea JA. prothoracicus van de Poll Pronotiltt convex medially, .... 6.04 ‘ we Medes cots et le oe Be A. elongatus van de Poll 24. Sutiral spine with rounded inner margin —~.. 25 Sulural spine with straight inner margin.......37 25. Legs red-brown colour.-.,........, 116 pnorefetete 26 Legs other than red-brown colour... ,, 222.27 26, Elytra marked with spots and faseia ..._, Sao Rede A, macmillani Barker Each elytron with 2 > vittae A. vitiatus van de Poll 27. Head, pronotum, legs metallic brown or bronze. r= voce. Jlavopictus C & G Head, pronotum, ‘legs other than brown of Brorize ee ne ...28 28. Fiuimeral told well developed, ‘angled, tiewec dd Humeral fold moderately or poorly developed. 30 29. Head black or coppery purple; ventral surface coppery-purple, oo. 6.0645 .-AL adanisi: Barker Head blue of green; ventral surface blue-green Tee | bd, geet = ae A. simulator van de Poll 30, Humeral fold moderately developed, angled, ,..3 Humeral fold poorly developed, slightly angled... A asTL SRA TOS Teen Comes Hd dote ole de 31. Pronoruin laterally rounded from base to apex, , , ee eet ey se eee A. crockerae Barker Pronotum parallel-sided ap base, rounded, indented 10 AON LL. a A. badeni yan de Poll 32. Basal spot touching basal margin of elytron, Did wreetewa € aD aD ve ety ed A. carnabyt Barker Basal spot not touching basal marvin of elytron Aecosm oP ARE T ON oho s eee O60. 33 33, Blytron with 2 spots und 2 fascia or 4 spots and a fascia’ ....5 54 reat 34 Elytron with 6 gnats ‘and. a fascia, 7 T spots ors spors 35 34. Head pmnotum ‘and “ctytra black, ' fL3900R.3.emm wo AL meprick! Blackburn Head pronotum and elyira plac with Blue and purple reflections... .... -A. eridus sp. nov. 35. Each elytron with 6 spots and a fascia or & spors Vit ratte c vol fansoni yon de Poll Each elytron with 7 ‘spots 1 Peri te DOCer 36 36, Pronotum parallel-sided ram base to ‘middle strongly rounded and jiarrowed to apex; dorsally conver (i lateral profile. ..22.....4. oherthuri vay de Poll Pronulur gradually rounded laterally, narrowed from base ta apex; dorsally flattened in lateral profite se Ese on oc ul oa -AL carter( Barker 37, Head with frontal spor... .A. princeps sp. nov, Head withoul frontal spot,,,....... 00222) 38 38. Head with shallow median suleus.-.,..,,,,,.... elaete sop at ‘ ect guerlingi Barker Head with deep median SUIEVS no setew essen, oo Head with basal median sulcus; provotum laterally inflared, medial oval patch of hexagonal cells. b dete oe oA, evaneus Kerremans Head with apical median sulcus; pronotum laterally rounded, no medial hexagonal cells. . b caledonteus Fauvel 39 New locality records A single female specimen of 4. irregularis van de Poll was collected by Mr E, E, Adams and myself at Blackdown Tableland, Qld, oo 17.1.1979 on the foliage of Casuarina inephioia, This isa common W.A. species recorded from 5. Aust. by Barker (1977), In the revision of Astraeus (Barker 1977) t did not record 4, jumsoni van de Poll outside of S, Aust, except for a doubtful Queensland record. Carter (1929) listed the species as occurrilig in the Blue Mts. | have examined Carter specimens in the SAMA collection from the Blue Mts and more recent material collected by Mr S. Watkins on Mg. 3, SDM. microwraphoof che elarion af Asrraeus pygroens Van de Poll 192 5S. BARKER Callitris muelleri at Wentworth Falls and Kings Tableland in the Blue Mts and confirm them to be A, jansoni. Egg of A. pygmaeus van de Poll A female specimen of Astraeus pygmaeus captured by me 24 km east of Armidale, N.SW., on 19.xii.1989, laid 13 eggs onto the cardboard label in the vial in which it was placed. These hatched approximately 28 days later. The hard sculptured chorion of an egg from which a larvae hatched is illustrated (Fig. 3). Crowson (1981) indicated that most beetles have smooth eggs with a soft surface. A micropyle cannot be distinuished, but a row of tubercules, each pierced by an aeropyle, is clearly visible around the margin (Fig. 4). The eggs are oval with a flat ventral surface. The larvae emerged through the ventral surface and burrowed directly into the label. The chorions adhered to the label and covered each individual gallery. Presumably the same happens in the field situation. These are the first eggs of Australian buprestids that I have seen. Fig. 4. S.E.M. micrograph of aeropyles (arrowed) on chorion of Asfraeus pygmaeus, Stigmodera (Castiarina) garnettensis sp. nov. FIGS 1G,5 Holotype: o, Mt Garnett, Qld, 24.ii.1989, G. Wood, I 21228 SAMA. Allotype; 9, Mt Garnett, Qld, 24.11.1989, J. Hasenpusch, | 21229 SAMA. Paratypes: Q\d: 1 9, Mt Molloy, 9.ii.1987, G. Wood, GWQA; 2 99, Kuranda, 6/8.i.1989, J. Hasenpusch, JHQA; 1 9, Mt Garnett, 20.11.1989, G. Wood, GWQA; 1 o, Mt Garnett, 21.11.1989, J. Hasenpusch, JHQA;2 oc, 3 9 9, 22.11.1989, J. Hasenpusch, JHQA; 5 oo", 2 9 9, same data as allotype, JHQA; 5 oo, 6 99, Mt Garnett, 21/22.11.1989, G. Wood, GWQA; 10 oo”, same data as holotype, GWQA; 1 9, Mt Garnett, 26.11.1989, G. Wood, GWQA. Colour: Male (Fig. 5A). Head, antennae and pronotum coppery-purple. Scutellum green with coppery-purple reflections. Elytra yellow with following markings either black with coppery- purple reflections, or dark blue with green reflections: variable basal mark, continuous with basal margin, not reaching lateral margins, reduced to basal margin and extension along suture in one specimen; post-medial fascia reaching margin; apical mark. Ventral surface coppery purple. Legs green with coppery-purple reflections. Hairs silver. Female (Fig. 5B). Head, antennae and pronotum green with yellow reflections, occasionally head and pronotum coppery-bronze. Scutellum green. Elytra yellow with following dark blue markings: broad basal fascia; broad post-medial fascia; basal mark. Ventral surface green with yellow reflections. Legs blue-green. Hairs silver. Shape and Sculpture: Head closely punctured, median sulcus, short muzzle. Antennae, segments: 1-3 obconic; 4-11 toothed. Pronotum closely punctured, narrow basal fovea extending forwards to middle as glabrous line, basal notches on each side more marginal than medial; apical margin projecting medially, basal margin barely bisinuate; laterally angled outwards from base, rounded one third of distance from base, bulbous, tapered to apex. Scutellum scutiform, punctured, flat. Elytra punctate-striate, intervals convex, punctured; laterally angled out from base, rounded at humeral callus, concave, rounded posteromedially, abruptly rounded to spineless apex; apices diverging, apical margin subserrate. Ventral surface with shallow punctures, edges of abdominal segments glabrous, moderately hairy, hairs medium length. S, rounded in both sexes. Size: Males, 14.2 + 0.15 x 5.8 + 0.07 mm (24). Females, 15.4 + 0.27 x 6.6 + 0.12 mm (17). Male genitalia: (Fig. 1G). Short and_ broad. Parameres angled outwards from basal piece, rounded posteromedially then parallel-sided, rounded to apex. Apophysis of basal piece medium TAXONOMY OF AUSTRALIAN BUPRESTIDAE 193 mm § Fig, 5, Habitus illustrations of the following species: A. Stizinodera gurnettensis sp. nov, male, B. S. garnetlensis sp. nov. female. width, tapered to rounded apex. Medium lobe pointed, sides obtusely angled away. Remarks: This is a member of the 8. cruenta Saunders species group as well as S. flava Saunders, S. ovala Barker, S. pallas Blackburn and S. vallisi Deuquet. Like S. va//isi the sexes are dimorphic and can be separated on colour and pattern differences. In the male genitalia of S. yal/lisi (Fig. 1H) the apophysis of the basal piece is narrower. The specific name is derived from the type locality, Stigmodera (Castiarina) wellsae sp. nov. FIGS 11, 2H Holotype: &, Portland, Vic., 21.xi.1944, FE. Smith, NMVA. Allotype; 9, 20 km N Moe, Vic., 8.xi.1979, A. Wells, SAMA | 21230, Paratypes: Vic 1 co, 1 9, Emerald, 3.xi1.1907, Jarvis, NMVA, SAMA; 3 oo’, Grampians, Nov. 1950, B Given, NMVA; 4 oo’, Portland, 6.xii.1918, NMVA; | ov, Ferntree Gully, 30.x.1919, NMVA; | oO, Portland, NMVA;1 0, | 9, ‘Taruna’, Croydon, S. HW. Fulton, NMVA; 1 9, Belgrave, 19.xi.1946, F £. Wilson, NMVA; 1 9, Healsville, Nov. 1916, NMVA; 19, Monbulk, 12.xi.1902, Jarvis, NMVA; 1 9, Boulora, 4.xii.1954, 4. L. Brown, NMYA; 1 ®, locality indecipherable, 14.xi.1903, NMVA; 1 o, 49 9,nodata, NMVA; 4 o'o",5 9 9, Mt Rosea, Grampians, 15.xi.1950, G, F Gross, SAMA; 1 co, Hall’s Gap, Grampians, 14.xi.1950, LeSoue/, SAMA. Colour; Head and antennae blue with bronze reflections. Pronotum blue with bronze reflections medially, yellow laterally. Scutellum blue with bronze reflections, Elytra yellow with following blue markings with blue-green reflections: very broad basal mark only touching margin at humeral callus; broad post-medial fascia; pre-apical spade-shaped mark, reduced in holotype and allotype to short horizontal bar, all marks connected along suture to apex. Hairs silver. 194 5. BARKER Shape and Sculpture: Head closely punctured, median sulcus, short muzzle, Antennae, segments: 1-3 oabconic; 4-11 toothed. Pronotum closely punctured, narrow basal fovea extending anteriorly to apical margin as impressed line, shallow fovea at each basal angle; apical margin projecting medially, basal margin barely bisinuate; laterally rounded from base to apex, widest medially, Sculellum scutiform, few punctures, flat. Elycra punctate-striate, intervals convex, punctured; laterally angled out from base, rounded at humeral callus, Concave, rounded posteromedially and narrowed to spineless apex; apices hardly diverging, apical margin rough. Ventral surface with shallow punctures, edges of abdominal segments glabrous, moderately hairy, hairs short, S.; male truncate; female rounded. Size: Males, 10,1 + 0,18 x» 4.0 4 0.09 mm (19), Females, 10.5 + O19 x 4.1 4+ 010 mm (16). Male genitalia: (Fig. (1) Short, Parameres parallel- sided basally, rounded posteromedially, parallel- sided, rounded apically. Median lobe sharp, sides acutely angled away, Apophysis of basal piece medium-width, tapered, flattened apically. Remarks; Member of the S. distinguenda Saunders species group together with S. arida Barker, S. blackdownensis Barker, S. garrawillae Carter, S- grata Saunders, S$. militaris Carter, 8. subgrata Blackburn, 5, supergrata Barker, S, watkinsi Barker, It is closest to S. warkinsi but differs in having: blue markings on the dorsal surface whereas those in S, watkinsi are green, a different distribution; different male genitalia (Barker 1988; Fig. 1B). Specimens in the NMVA had been misidentified as 8, grate Saunders. Named after Dr Alice Wells. Acknowledgments I thank the following for their assistance: Miss Cc. M. H. von Hayek, BMNH; Dr A. Neboiss, NMVA; Mr G. G. Burns, Mornington; Mr J. Hasenpusch, Innisfail; Mr 8. Watkins, Caparra; Mr G, Williams, Lansdowne; Mr G. Wood, Atherton; Mr P. G. Kempster, Mr C. Miller, Ms H. Vanderwoude, Dr A, Wells, Mr D, J, Williams, Department of Zoology, University of Adelaide. References Barker, S. (1975) Revision of the Genus Asiraeus LaPorte & Gory (Coleoptera: Buprestidae). Trans. R. Soc, 8, Aust. 99, 105-141. (1977) Asiraeus (Coleoplera: Buprestiducy; A description of three new species and new lacality records. (hid, Wl, 11-14, (1988) Contributions to the taxonomy of Stigmadera (Castiarina) (Coleoptera: Buprestidae). Thiel. 112, 133-142. Brackuurn, T. (1890) Notes on Australian Coleoptera, with descriptions of new species, part v, Proe, Lina, Soc. NSW. 4, 1247-1276, (1891) Notes on Australian Coleoptera, with descriptions ol new species, Joi, 5, 479-550, (1895) Further notes on Australian Coleoptera, with descriptions of new genera and species. Trans, R. Soc, §, Austr. 19, 27-60. Carrer, H. J, (1929) A check list of (he Australian Buprestidae. Aust, Zool. 5, 265-304. Crowson, R. A. (1981) The biology of the Coleoptera. (Academic Press, London.) KeRREMANS, C, (1892) Catalozue synonymigue des pupeete decrits de 1758 & 1890. Mer. Soc. ent. Belg. __. (1903) Coleoptera; Servicornia Fam, Buprestidae. Genera Inseetorum 12, 1-338. (P. Wytsman, Bruxelles.) OBENBERGER, J. (1928) Opuscula Buprestologica 1. Beitrage zur Kenntnis der Buprestiden (Col.), Arch, Narurgesch, 92, 113-224, (1930) Buprestidae I], Coleoptorum Catalogus NIT: 213-568. (W, Junk Berlin, Den Haag.) VAS DF Pont, J. R, He N. (1889) Monographical essay on the Australian Buprestid genus Astraeus C, et G. Tjschr. v. Ent. 32, 79-110. ___ (1892) Note sur quelques especies d'Asrraeus, bid, 36, 67-68, Watt, J. C, (1979) Abbreviations for entomological collections. N. 2. Zool. 6, 519-520, TRILOBITES AND THEIR TRAILS IN A BLACK SHALE: EARLY CAMBRIAN OF THE FLEURIEU PENINSULA, SOUTH AUSTRALIA BY RICHARD J. F. JENKINS* & PAM HASENOHR Summary An Early Cambrian trilobite from the Normanville Group in the Sellicks Hill area, Fleurieu Peninsula, South Australia is here identified as [vshiniellus briandailyi sp. nov. The trilobites occur in ‘black shale’ in close stratigraphic proximity to a bedding-plane surface showing trails made by large trilobites. The environment was below storm-wave base and the traces indicate that the black shale deposited in water sufficiently well oxygenated to support large metazoans with a presumed oxidative metabolism. Black coloration in shales may not be taken as a necessary indication of anoxic bottom waters. KEY WORDS: new species, conocoryphid trilobite, trace fossils, black shale, Cambrian, South Australia. Transactions af the Roval Saetety af 8. Aust, (1989), 113(4) 195-203, TRILOBITES AND THEIR TRAILS IN A BLACK SHALE: EARLY CAMBRIAN OF THE FLEURIEU PENINSULA, SOUTH AUSTRALIA by RICHARD J. F. JENKINS* & PAM HASENOHR Summary Jenkins, R. J. b. & HaAsenorr, P, (1989) Trilobites and their trails in a black shale: Early Cambrian of the Fleuricu Peninsula, South Australia. Trans. R, Soc S. Aust. 113(4), 195-203, 30 November, 1089, An Early Cambrian rrilobire from the Normanville Graup in the Sellicks Hill area, Fleuricu Peninsula, South Australia is here identified as /vshiniellus briandailyi sp. nov. The trilobites occur in “black shale’ in close stratigraphic proximity to a beddiig-plane surface showing Lrails made by large rrilobires, ‘Lhe environment was below storm-wave base and the traces indicate that the black shale depasiled in water sufficiently well oxywenated to support large metazoans with a presumed oxidative metabolism. Black coloration in shales may not be taken as a necessary indication of anoxic bottom waters. Key Worns: new species, conocoryphid trilobite, trace fossils, black shale, Cambrian, South Australia. Introduction Jago et al, (1984) reported the first discovery of trilobite remains in the Early Cambrian Norman- ville Group on Fleurieu Peninsula, South Australia. They described two specimens of a conocoryphid trilobite from the Heatherdale Shale in the Sellicks Hill area (Pig, 1), bul considered the material so poorly preserved as to preclude formal description. While exploring the same site during late 1984, one * Department of Geology and Geophysics, University of Adelaide, G.P.O, Box 498, S. Aust, S001, of us (P.H.) discovered a spectacular trace fossil made by an arthropod, Further searching revealed two more body fossil remains of trilobites, This paper Teports these new discoveries. Of the new trilobite remains, one shows the cephalon and anterior thorax and the other is a fragmentary thorax. They were discovered on surfaces that had split open to the weather, and the very incomplete specimen is preserved essentially as a ferruginised film, The more complete individ- ual, an external mould, resembles specimen SAM P24321 described by Jago ef al. (1984) and provides 2/0 Granilas Hectiierenitalnel Kangaroo (stand lreup a Kanwiantow Grown = P| Adelman (Matern a = Dey alalliay Maw irriertt 4 CAMBRIAN —rT Theat Fauits ——"_ Faults TAN MLOMb IES io 20 40 Ww FRE AMI Fig. 1. Locality thap showing find of trilobites ahd trace fossils at Sellicks Hill and the placement of this discovery in relation to the framework of the Adelaide Pold Bele and adjacent foreland basin. 196 R, J. &. JENKINS & P. HASENOHR Fig. 2A-C. /vshiniellus briandailyi nov. sp: A, holotype SAM P24321, with anterior glabella relatively well preserved and composite mould of hypostome present, x3; B, C, paratype SAM P24844, B, external mould and C, silicon tubber cast of individual made from mould, both x1.5. D, conocoryphid gen, et sp. indet, SAM P24845, fragment photographed under a film of water, 1.5. TRILOBITES OF THE FLEURIFU PENINSULA 17 sufficient! additional morphological information ta justily formal deseripoion of the taxon. The arthropod traces in near association with the body fossil specimens of the trilobites are of especial interes! as the host rock vepresenis a ‘hlack stale" facies’ Thus the traces provide an unusual palaenhiological perspective on this interesting find, The studied speciens are registered in rhe Palneontolugy Collection of the South Ausualian Muscum, Adelaide (SAM). SYSTEMATICS Soperfainily CONOCORVYPHACEA Angelin, 1854 Family CONOCORYPHIBAE Angelin, [854 Remarks: Divisions amongst the various genera of vonocoryphids tend to be ill-defined due to description ol cranidia alone or because many remamas of these thin-shelled trilobites are murkedly distorted. Furthermore, workers have tended to stress thal the group is palyphiyletic (Westereard F950, Sdzuy 1961; Hutchinson 1962; Rasetti 1967). On the basis of the evident positioning of the facial suture on the brim, the present form is a ‘thc’ conocoryphid (Figs 2 & 4), The family Conocoryphidae is rare in the Early Cambrian and much more common globally in the Middle Cambrian (Koroboy. 1973). Lochman-Balk & Wilson (1958) considered thac the blind Conocorypludac and Eodiscidae were characteristic of eXtracratonic, enxinic biofacies in the Cambrian uf eastern Nerth America and suggested that the blind condition wis favoured by some imporcant ecological factor, Genus Justinrel/lus Korabov, 1986 Remarks: The better preserved of (he orivinal specuens (SAM) P2432}) is described as having a rearwardly directed furrow on the side of rhe anterior part of the glabella (lage ef af, 1984): medially this par of the glabella seems 1o have been entires und undivided. Though the front of the elabella of the new specimen is crushed, there appear to be tour pairs of glabellar furrows (Fig. 3); the elabella tapers anteriorly and is somewhat constricted at abour a (third its length. These charactenstics arn! other aspects of the cranidium conform to the diagnosis of the Early Cambrian venus /vsftinieliay Korobov, 1966, Most Cambrian conocoryphid genera show three pairs of glabella furrows {in advance of the occipital groove). The distance the glabella reaches forwards towatds the frontal border, the possible presence of a raised pre- glabella lobe, and the width of the froneal fleld, if it is developed, are same of (he significant character- istics important lor dislinguishing conocoryphids. While Korobov (1966) compared his new genus with the Middle Cambrian taxon Builiella Mathew, 1885. ir is similar ta the Carly Cambrian Pseudatops Lake, 1940. Species of Psevdatops may show Tour pairs of glabella furrows, but the alabella is only weakly lapered forwards and differs from thar in Mvshiniellus in encroaching on the frontal border, Pig. 3. Representation of the elabella wid cence! pan ol theeranidiany of SAM) 24844 shaw tag posiian ot ihe four parrs.at ehabetlar Jurpaws, t=. ii udvaitee of phe ovoipilal furrow. Gee. The riore urterion glubetlar lobes aic strongly campressed ih an atlerforepasteriu threanan, Consperv wid) Peni 2B; Ivshiniotlus brianduilyi sp, sv. FIGS 2-4 Conovoryphid sp, indet, daee er af, 1984, pp. 208-210, ‘fies, 2o-c, Etymology: Lonouring the lite DY Brian Raily (1931-1986) for his contribution to Australian feology and (caching. Diagnosis: Brim weakly inflexecd medially, crani- dium encireted by border furrow and rather evenly vovered by tine to coarse, pointed granules {spin- ules) in immature specimens, glabella almese amooth in more mature exumnples; pleura with acicular, weakly deflexed terminal spines; eleventh segment macroplenrous. Material: Holotype SAM P2432], lacking reariust parts Jago er af. 1984, liv. 2), by; two paratypes, SAM 224322, rear thoracic segments and Tragmentary pyeidtur (lage er wh M984 fie, Bey SAM P24844, cranidium and parts of 14 choracie LeAOenNts, Ayseripiivn Tt is unecessary 16 repear te desenpions ul (he holotype and second fragmeni 198 kK oIF hip. 4. Reconsrruetion af eephalon cand miajol part a Hark of fevdinelus brigndailvi sp. tus, abo 2.9 Himes dimensians OF halorype. provided in Jago-er af (1984, pp. 209-210). The new specimen confirms the arcuate shape of the cephalon before, and slight median inflexion of the border, as well as indicating that rhe facial suture is cunlined to the brim. In the holotype, the granulose brim is poorly preserved and is wider than might seem obvious in the previously published Fipures; its posterion is well defined and che wide border furrow more fincly and sparsely eranulose than the ceptral parts of the cranidium. The new remains show a nearly smooth, narraw glabells tapering forwards. with four pairs of glabells furrows directed rearwards, The (riagmentary base of an occipital spine is present in the holurype. The additional individual indicates at least 14 thoracic segments and confirms that the eleventh is Tnucropleurous. The thin pleural ridges om the holotype show a line of 10-1 spinules or pornted eranules on the an(erior segments and 8-9 such iramules on the posterior ones; other Cine granules accur between. Meusurements: The hulotype, SAM P2434), indicates an animal caceeding 25 mnvin length. ‘The distorted cranidiun of SAM P24844 is 50 mm wide and the maximum width oy ihe glabella Is 8.5 jm: fourth iboracie segment wboul /45 ain wide, JENKINS & T HASENOHEK excluding the spines and with the axis about 70.2) the width of the segment; animal in excess of 55 mm long, Comparison: The genus includes two previously described tormns, /vshiniellus nikolii- ard Iyshiniellus pululus Korobov, 1966, both from the Aldanian Stage in Tuva, southeastern Siberia, within the Ezim Suite on the Ezim River. Both are represented by distorted cranidia and so any comparison with the present distorted remains must neecssarily lack precision, 7, hbrigndai/yi resembles f. ackold in the shape of the cephalon, save for the distinctive medial inflexion vf the trontal border: rhe glabella also seems narrower. 7. pa/w/as differs from both of the former in us pointed glabella and the somewhat quadrate shape of the fixed cheeks. Mr J. G. Gelling has kindly provided photographs of cranidia and loaned several nearly complete dorsal exoskeletons of conocoryphid trilobites collected from the Parara Limestone at Bunyeroo Gorge in the Flinders Ranges. This material is of a taxon different ta £ briandailyi as the cighteenth thoracic segment is inacropleurous Hot the eleventh, and the medial thoracic segments have remarkably long and strong pleural spines. The tather long glabella, which reaches a depressed horder furrow behind the medially widened border, has three pairs of incised, backWardly directed furrows and a weak pair of depressions on the exireme anterior portion} laterally, the facial sutures cut off the major part of the brim. En its cephalic characteristics this form from the Parara Limestone shows similarities (oO the EBurly Cambrian genus Arops Emmons, 1844. Conocoryphid gen. et sp, indet. Remarks: Vhe additional fragmentary thorax (SAM P24845) dues not show the eleventh segment (counted from the most anterior segment preserved) macropleurous, and the pleural spines are more deflexed and evidently shorter than in the preceding form. However, the pleura show the characterise wide-flat pleural grooves and narrow pleural ridges of conocoryphids, ‘Plic thoracic segments are about 11.3 times as wide as long, excluding the spines, with the axis about 0.24 the width of the segment, The poor preservation tias largely obliterated any surface detail, Disarticulated thoracic segments associated with cranidia of Clendcephius Hawle & Corda, 1947, in material from the Middle Cambrian of France, show the pleural terminations narrowed abruptly and thence produced into a somewhat deflexed, slender spine, The shape of rhe pleural terminations in SAM P24845 is markedly similar; This specimen coukl be a dimorphic individual of 5 briandaily TRILOBITES OF THE FLEURIEU PENINSULA 19 or even represent a separate conocoryphid taxon, but basically is too fragmentary for determination, Age of trilobite finds The occurrence of Jyshiniellus on the Fleurieu Peninsula is of potential significance with respect to the timing of a major stratotectonic development within the Adelaide Fold Belt, Cambrian rocks of the Normanville Group below the Heatherdale Shale are of both basinal and shelf aspect (Daily 1963, 1976; Jago et af. 1986), The trilobites occur in the upper metnber of the Heatherdale Shale high in the section of the formation exposed (Jago er al. 1984), At Carrickalinga and more southerly parts of (he Fleurieu Peninsula the Heatherdale Shale is ubruptly succeeded by up to ca. 1300.m of curbidites of the Carrickalinga Mead Formation (sertsw late of Daily & Milnes 1971, 1972; Milnes 1986), the basal interval of the thick psammific and/or pelitic sediments of the Kanmantoo Complex, Sandstones of subtidal or lidal origin (Backslairs Passage Formation) succeeding the Carrickalinga Head Formation record a phase of basin Silhing. Jenkins (1989) identifies this overall basinal deepening and shallowing as the ‘Ramindjeri/Jadliaura strarotect- onic cycle’ and (he organic-rich sediments (Foster et al 1988) of the Heatherdale Shale may be considered as The eusiule’ phase Marking the scary of rapid subsidence associated with renewed litha- spheri¢ extension, The likely age of /vefuiel/as lo Siberia is fate Aldanian (Korobov 1973; Jago ef af 1984) or medial Early Cambrian, and the comparable genus Pseudafags also oecurs in the Imid Early Cambrian of eastern North America syd the British (sles. The present Finding that 4 brivadaity; and ia conocoryphid from Bunyeroo Gorge are differed taxa Weakens che claim of Japo er vf (1986) that conoeeryphids suggest a votretation between che Heatherdale Shale and higher parts of the Parant Litiestone in te Flinders Ranges, though on a tecloric basis the Onset of deposition of the dark, nodular offshore facies of the latter formation mity well have corresponded wilh (he deepening signalled by the Mearliendale Shale (see Jenks 1989), Trace Fossils Ab upper-surface beddina plane at che trilebite silt shows a relatively well preserved arthnopod (rail extending over a distance of BO vin, parts of two other traile probubly made by large trilobites and & poorly preserved section al! the back-flled burrow produced by a worn dike creatine (Figs #6), This surface has heen preserved due lo an unusual CUCULTES EAN The arthropods evidently moved over a relatively fine-grained and soft, muddy substrate with the ends of their limbs making deep impressions. Had this surface been subsequently buried by further silr, the cleavage now present in the rock due to deformation (Cambro-Ordovician Delamerian Orogeny) would probably preclude splitting along the particular bedding surface, However, the traces were buried by a 3.5 cm thick bed now comprising intergrown phyllosilicates and uinor tine (— 0.1 mim diameter), angular quartz grains, which are commonly amalgamate and show undulose extinetion, The quartz is probably detrital. XRD analyses of the rock confirm visual observation that the bed is inhomogeneous with a (less weathered?) fraction comprising predominantly illite 2M), minor muscovite M , and kaolinite-smectite, and a second (weathered?) fraction including illite arid Ca montmoriilonite. Iron rich opaques are concentrated in parts, Mite 2M is consistent wih a paragenesis involving deep burial and alteration of possible older mineral phases, The bed is continuous along strike over an exposure distance of 60m and thickens slightly lowards the south (up to 4.5 cm), tn places, obvious eranular components that are apparently ‘reliets’ of the original sedimentary parucles grade in sive from ~ 0.5 - 12 mm near the base of the bed to ~ 0.1 mm al the top, The granular aggregates of phyllosilicates posubly represent the tnghly altered components ol an airtall cuff that wixed with a small amount of detrital material and formed a slurry which buried and moulded the trilobite traces. During: modern Weathering 4 ling of parting, developed subjacent to the Underlying silistone such that avery thin veneer of granular material still adheres jo and completely covers the traces, The trail traced over 80 em consists of sub parallel rows of rather closely spaced iniprints of ambulatory limbs and averages 75 fo 80 aim in width. The Individwal imprints wend ro be ser obliquely or en-echelon and the best preserved are about 1 mm tony. Pils urace is af the form itlusteated by Seilacher (1955, fig, la, b) and was evidently made by a perambulating trilobite with Irs body axis alned in che direction of travel. In one area il is overlapped by a ericely segmented trace nade hy & Wolfielike anual, Nearby exposures iy the Hesiherdale Shale show much hetier preserved examples of whit is probably this Same race OCCUTTINg Lh assomation wilh nuiterous burtows of a relatively large furm of Planolites Nicholson 1872. The segmented trace is probably Of endogenic ongin aid well preserved examples show @ reeqlar back-filling. of large ovate faecal pellets arranged Irninsversely 200 RK. J.B. JENKINS & P. HASENOHR Pig. 5. Bedding plane below tull horizon, showing trace lossils made by trilobites, A-C, and structure, Dy artibuted fo ain unknown worn-like animal. A-A’, trace made by perambulating trilobite with body aligned in direction of travel B-B', macks made by large trilobile moving, obliquely; C-C', indication of a trilobire travelling relatively quickly, Seale bar equivalent to 20 em. pil f / f LIL SE ff Lip fe v PLILEEEL SPELL E /) f a — ia ao aed - Cc 9 p 20 cm % i Fi ; ae ao® d es LLL SPITS LLLLLLL LLL Tee 2 © Fiv, 6. Representation of bedding surface portrayed in Fig. 5 indivaling (he several sets of markings made by trilobites (A-A’, B-B’, C-C") and another unknown animal (D). TRELOBITES OF THE FLEUKIFL: PESENSULA alt] The two other traces show relatively widely spaced series of imprints interpreted ay the inipressions Of che tips of the telopodites of trilobites In the larger trace. the matvidual imprints ae up to 20 mm Jong and on one side of the complete trail dre arranged in ratbec Wregular rows positioned in an eeechelun manner Phis sore of marking is well expluined by Seilacher (1945S, fig. le V) and indicates a trilobite moving obliquely erab-wise, perhaps partly in response lo a eentle side current. Che tins of the limbs which formed the racks on elher side al the itace spanned approximately 23 cm, indicating a quite large animal, The imprints of the remaining tre are widely spaced and suggest a smaller animal travelling relatively quickly with the body aligned in dhe diteetion of travel. These marks. are tol especially disanenve (aml wete only seen in cera Hating) but dre comparible with sont ol the lane HHobile (rails Occurring in the late Eariy te Middle Cambri Kangaroo ishind Group of Daily (1956), waethe berch couse of Kangarog Intima, Che body fossil rennains of the tilobites desenhed Jictern occurred tespectively. 1.6m siratigraphically below (he surface with trilobite trails (SAM P24844) and #8 cm above (his suclaee (SAM P2484), The taces suvges! (hat (rilobites were live jnhabitaats of the horton environment Patacobjological signifivance The thinly laniinated argillites of the upper member ( ~240 nm thick) of the Hewtherdale Strate are almost cutirely far bedded and certainly jocomulated at depts below storm wave base (cf, daeo er df, (986). A single rippled surfidoe observed riray reflect localised bottom currents. Considering the thickness of the succeeding Larbidites, che depth of water in the deposibonal envirgiment was prabably of the arder of hundreds of retres. Jago et al. (1984) cite and illustrate apparent tiuderacks in the Heutherdale Shale Oheir fig. 4) ws possibly indivating a shallow environment of depusition bul (hey concede that the eracks may have formed due to syiaeresis, DAiasiiski & Walton (1965) illustraie their tigs 12 & 113) compurdble eracks in turbidites. Trilotile trails oveur abundantly in parts of the late Barly (& Midulle Citrbrinn sequence on the north coasr of Kangaroo Island (Sprisuz 1985; Daily eral \979). Lxarmples of such rails up bo LU-32 em in Width gre especially dumerqus in iitervals of (ese forks deposited in tidal and subtidal facies. This assoeiahion of t(rdobite wails ii shallow-water sediments is common world wile duriuig the older Vulacozpie and constinites the Cruzer facies of Seilueher (46d). Mis u is apparenc that crilobires had anordinary oxidative kind of metabolism such as is characteristic of modern, free living arthronads and metazoans in general, The wtthropod tras in the Beylherdl: Stale suggest relatively larce animals moving over the bottom in seme numbers. Lhe implication is thar the bocly fossil remidins of urilabites were part ul un indigenous papulation and not isolated individuals somehow transported inte the enviror- ment by chance. Presuming sn oxidative meta- bolism for (he (rilobites, and [he worm-like crealures responsible for various burrowed horizons, the holtom waters of the Heatherdale Shale enviroment were clearly not uniformly anoxic and must have been at lease moderately well oxygenated al the fines Gf broturbation, Chis comtradiers the commonly treld opinion that black shales characterize oxygen starsed basins; fluctuating oxvgen tensions may be more likely (Marris 1980); Kuufiiman 1982; Kuispert 1982; Wetzel 1982; Conway Morris 1985). By measunney the displacement of a eonveniently sived plaster model of a trilobite, the soft tissue mass of the anjmal which made the 23 cm wide trail tay be estimated at ~ 500 9, With yeferehee to fie. #6 of Alexander (1979), the oxygen consumption of wnanimal of this size may have been about 0.045 ml ein hie’ Jor for the whale organism, 23 al hr . Runnevar (1982) demonstrated that the surface area or rhe respiratory Organs (eg. gills) of aqueous animals iy approximately proportonal to the body mass; this sugeesis a cespiratory exchange surface of ~S500cm) for the trilobite. Apptivation of equation 13.5 of Alexander (1979, p. 279) indicates that the partial pressure between the sea water and blacd aver the respiratory surface needed to supply the required OX Veen consupipsiarn is -0.06 atm. Assuming that the respiratory pigment in the animal's blood was saturated at 0.02 aim. the (OUI Oxy Ben COnventtation: i the seu Waree Hecessury far (he tnlobile was - (LOR aim or ~ 38% PAL (~60 mm He of 0). This vahte lies within the medial part of the speerrum of oxygen conech- Hations in the oxygen midtimuto layer of modern oceans, effectively 2.8% PAT. 10.60% PAL (Barnes & Hughes (982, fie, 1,9), There is corvently a renewed wave of speculicion concerning the possible icrense ol atmospheric oxyeen during the late Prevambliriad atd ity relationstip to Me cinergerice Of Ne metacoa ane higher plant life (Runnegar 1982a, 6 aind spers comm; Glaessner 1983, 1984), The present exanple Sugeest that Siniple observation of dark coloration in parts of Precambrian sequenees (eu, arey anc blick phyllites in the “Tarrensian’ Series Ot the Adelaidean) is unlikely 1 give relluble inlocmation OW ambient wsyeen tenons, The coloration of 402 R. J.P JENKINS & P. HASENOHR shales Will also reflect diagenetic processes and the éffects of inapient metamorphisre, for exaniple fécrystallizatian of Naematite to specularite may cause beds to become almos! black (Clemimey & Badham 1982). ! briendaily! is remarkable for the tong genal spines, the attenuated spines on the pleurites and iS Macropleurous eleventh segment. Similar long Spines in adontopleurid trilobites have been considered to have functioned in helping to support the animals on ihe sea bottom, either during ‘resting’ ur while they were feeding with the aid of turrenls passing, het ween che substrare and the lower side of the thorax (Wittington 19S6a, b: Clarkson 1969, for a different opinion see Bergstrém 4973). Such an adaptation is consistent with L brianedilyi inhabiting a soft muddy bottom, Because of the lack of streamlining brought about by the spines the trilobite was probably not ati ace swimmer and this avcards. with the finding of irackways in vlase proximity to its temains, However, the crackways indicate larger animals than the body fissil Specimens and there is no way of telling: whether the iruees were made by more mature indivaduals of conaderyphids or orher trilobites kitewn bo reach large sive locally, such as Kedlichia Cossmann. 1902 (sve Conway Morris & Jenkins 1985), A host of modern day créatures have adapted to inliaht caves and other dark places and beeume secondarily blind, and loss of sight in conoguryphids would have posed no disadvantage if these primal denizens of the deep were bottom dwellers who lived below the effective perelration of light, sustained primarily on settled detrjpus, References ALExanubk, Re MeNriit, (979) “The Tivertebrates.” (Cambridge University Press, Cambridge.) Wyanrs, Ro S KO & Plucis, BR. ON, (198?) an Introduccion ta Marine Reology” (Blackwell, Ox ford, Tondon.) Burs ROM, J, (1923) Ci ganization, life and systenians Of (ilobites, Fossils wit Surat 2, |=69. Ofannsos, Fo N. K. (1969) A Tunetional study af the Silurian odentopleuril crilobite Leonasns deflexa (lake), herkeia 2, 319-344, frpveey, H. & Bauliaw, No 1982) Gkypen in the Precambrian ajmosphere: #9 cyalualion of the zeological evideline, Geufogy IM, [41-146, Conway Morais, S, (1985) Cambmran lagerstatcen: iteir distribution and significance, Pil Thans, R. Soc. Lane, BSil, 99-A5 & Jenkins, KL K (1985) Healed injiirtes any Farly Cambrian trilobites tram South Australia. Aleheringa %, ta7-177- Dairy, & (1956) he Cumbrian in South Australia, der “EI sisteina Cambrich, su palaeoerafia ye el protiloma de su bise”. 20h /nt, geol Curige. Aferioa 2, 91-142. __ — (1963) The tossiliferous Cambrian succession on Flenrict Peninsula, South Australia Hee 4, As, Afus. 14. £79-601. —____ {1976) The Canibrei of the Flinders Ranges, {7 K.P Thomson, BR, Daily, R. P. Cours, & B, PB, Forbes, “Late Precambrian und Cambrian eealegy of the Adelaide “Geosyneline” and Stuart Shelf South Austraha’. 257 Ine, geal. Cungr Excursion Guide Na 335A, 1-56. & Minnes, Ry (971) Stratgraphic notes on Lower Cambrian fossiliferqus inelasediments berween Campbell Creek and Tunkalilla Beach in the type section ofthe Kaamantoo Group, Fleurieu Peniisula, Soutlt Australia. Trans. 2. Soe. S Anse ¥5, 199-214, & — (1972) Revision of the stratigraphic nomenclature Of ihe Cambrian Kanmantoa Cronp, South Australia, J Geal, Soc. Ause, 19 197-202, ——~—. Pwanarm. C.K & Boukne, £ A. (1979) 1: Geology and geombrphology. pp 1-38 fa MF Tyler, © _R.Twidale & J. R. Ling, 1Fds) “Natural History al Kangaroo Island" (Royal Sucicty of South Australia, Adelaide.) Divgyachy Sy & Watton by Ny 11965) "Sedimentary Keatures or Flysch and Cirevway kes" Deyelapinents in Sedimentology 7. (Fisevier Netherlands| Posen, C. G., Crenovsnis, A. & O'BRINN, G. Wo U1985) Organic-walled nnerafossils from the Faw Cambrian at South Australia. 4lcheringa 9, 25V-268, CILAESSYER, M_ fF. (1983) The emergence of Metaena (n the early history of tite, Precamb, Kes 2. 427-441. US) “Pho Dawn of Ammat (ite A Biobistorical Stidy", (Cambridge University Press Canibridge.1 HUTCHINSON. KD, (1962) Cumbrian stratigraphy and trilobue faunas of southeastern Newtourlland, dell, Geol Suu Cun. 88, I-56. Jaco. LB, Dany, B, Von Bek Bowen, ¢. ©, CEENOWSEES, A, de Sabi Obs, N. (1984) First reporied trilobires trons the Lower Cambrian Normanvyille Group, Heurigu Peninsula, South) Australia. Trans. &. Sad S Aust, 108, 207-211 4 GNDING, Jt. & Dairy, B (TRG) Canbrian sediments of the Sellick Hill-Carnickelinea Head area, Fleurieu Peninyula, South Australia. dA. | Parker (Ed.) “One Day Geolovival Excursion ot we Adelaide Region" pp, 67-81, (Geological Society al’ Australia, SA. Div, Adelaide) Jiwinins, Ry KOP988) The Adelaide Fall Belle leer feanpiaisal J}. B, Jago, & POS. More, /Puts) "Aspeeis of che Adclaidean and Cumbrian Gealoes wl ihe Adelaide Geasynctine” (heal Soc. Aus. Spec Publ in press.) KoarirrmaAt, E.G. (0982) The communry airnetnee of ‘shell islands' on oxygen deplered subst rales in Mesozoic dark shales and larninated carbonates, JG. Einaele de A, Seilacher, (dst “Cyclic and event sirarification” pp, 502-503 (Springer Verlag, Berlin Heldetbere ) Koxovov, MN. (1966) New vilobites of (he amily Conowtryphidas fom the Cambrian of the Siberian Platform and Tuva. Paleant 4h. 186d, Nu. 4, 92-97 (In Russian). —— - (75) The iilotite family Conocaryplndar and its significance (o the stratigraphy of the Canibriat denosily, Trvdygeel, inst, Moscow, 201, (ly Russian}. KispcetT, W. (1982) Environmental changes during att shale deposition ag deduced from stable serine raring fa G, Einsele & AL Seilacher, (Ed) “Cyclic and event strauilication” pp, 482-501, (Springer-Verlau. Berlin, Heidelberg.) LOCUMASEBALK, CL & Wilson, 10, (1958) Gumbrian Wiostraligcuphy in Neri Amencu, 2 Galena: a. 312-350, TRILOBITES OF THE FLEURIEU PENINSULA 203 Minnis, A. R. (1986) The Encounter Bay Granites and their relationship to the Cambrian Kanmantoo Group pp. 43-52. In A. J. Parker (Ed_) “One Day Geological Excursions of the Adelaide Regian”. (Geological Society of Austraha, S.A. Div., Adelaide.) Morris, K. A. (1980) Comparison of major sequences of organic-rich mud deposition in the British Jurassic, J, Geol, Soc, London 137, 157-170. Rasserrt, F (1967) Lower and Middle Cambrian trilobite faunas from the Taconic sequence of New York. Smithson, misc, Colins, 152, (ll. RUNNEGAR, B. (1982a) Oxygen requirements, biology and phylogenetic significance of the Jate Precambrian worm Dickinsonia, and thé burrowing habit. Alcheringa 6, 223-239. _ __., (1982b) The Cambrian explosion: animals or fossils? J. geol, Soe. Aust. 29, 398-411. Spzuy, K. (1961) Das Kambrium Spaniens/Teil 11: Trilobiten. AbA, math-naturw. K. Akad, Wiss. Mainz 1961, 597-693. SFILACHER, A. (1955) Spuren und Lebensweise der Trilobiten. /n O, H. Schindewolf & A. Seilacher, Beitrage sur Kenntnis des Kambriums in der Salt Range (Pakistan), Ab. math-naturw, Kl, Akad, Wiss. Maing 1955, 342-372. (1964) Biogenic sedimentary structures pp. 296-311. Jn J. Imbrie & N. D, Newell (Eds) “Approaches to palaeoecology” (John Wiley, New York). Sprica, R. C. (1955) The Point Marsden Cambrian beds, Kangaroo Island, South Australia. Trans. R. Soc. S. Aust. 78, 165-168. WESTERGARD, A. H, (1950) Non-agnostidean trilobites of the Middle Cambrian of Sweden. Sver geol. Unders.. ser, C, No. Sli, Arsb, 43(9), 1-56. Wetzer, A. (1982) Cyclic and dyscyclic black shale formation pp. 431-456. Jn G. Einsele & A. Seilacher (Eds) “Cyclic and event stratification” (Springer-Verlag, Berlin Heidelberg.) Wittincron, H. B. (1956a) Type and other species of Odontopleuridae (Trilobita). J. Paleont. 30, 504-520. (1956 b) Silicified Middle Ordovician trilobites: the Odontopleuridae. Bull. Mus. Comp, Zoot. Hary. 137, $57-170. REDISCOVERY OF PRIMOVULA (P.) HELENEAE CATE AND DESCRIPTION OF A NEW SPECIES OF CRENAVOLVA FROM SOUTH AUSTRALIA (MOLLUSCA: GASTROPODA: OVULIDAE) BY K. L. GOWLETT-HOLMES* & N. J. C. HOLMES* Summary The first live collected specimens of the South Australian allied cowrie Primovula (P.) heleneae Cate, 1973 are recorded. This species, previously known from a single shell collected prior to 1908, is redescribed, and details of the animal and its habitat given. Crenavolva cruenta sp. nov., 1s described from Spencer Gulf, $.A.; it most closely resembles C. striatula and C verconis but is distinguished by its more elongate form, more produced ends and colour pattern. KEY WORDS: Gastropoda, Ovulidae, South Australia, Primovula heleneae, Crenavolva, new species, habitat. Transactions af the Reyal Saciery af S. Aust, (1989), L304) 205-214. REDISCOVERY OF PRIMOVULA (P) HELENEAE CATE AND DESCRIPTION OF A NEW SPECIES OF CRENAVOL¥A FROM SOUTH AUSTRALIA {MOLLUSCA; GASTROPODA: OVULIDAE) by K. L, GoWLETT-HOLMES*® & N. J.C, HOLMEST Summary COW LETT-HOL bs, K, b. & Horns, N. J.C. (1989) Rediscovery of Prinvrovele (Pd helenewe Cate and description ala new species af Crenuvalva (rom South Australia (Mollisca: Gastropoda: Ovulidae). Trans. R. Soc. 8, Aust. 1134), 205-214, 30 November, 1989, The first five collected Specimens of the South Australian ullicd cowrie Prinrcvala (27 feleneae Cave, 1973 wre recorded, This species, previously known from a single shell collected prior to 1908, is redeseribed, and details of the wnimal and its habitat given, Cren@volve crvenra sp, Nov, is deseribed from Spencer Gulf, SA. i most closely resembles C. siriatuda und © yerconis but is distinguished by its mare elongate form, more produced ends and colour pattern, Key Wonns; Gastropoda, Ovyulidac, South Australia, Primovuly heleneae, Crenavolva, new species, hubiiar, Introduction Three species of the allied cowrie family Ovulidac are recorded from South Australia: Creravolva verconis (Colton & Goudlrey, 1932), Primovula (P) heleneae Caie, 1973, and an Ac/yvelva sp. (Coleman W9R1). All are regarded as very rare, The first two species are known from single specimens dredged dead prior to 1908 (Vereo 1908, as Oviilea formosa). Ol Douglas Point in northern Spencer Gulf, S. Aust, one of us (NICH) found a number of allied cowries living on fans of the gorgonian corals Muplexaura sp. and Mehinugergia sp. in 10-15 m depth, These specimens were identified as Primovula (P) helerieae Cate, 1973, (living on Euplexaura sp), and an undescribed species of Crenavolya, (living on Echinegergia sp,), Here we describe the new species of Crenavolva, redescribe F (PF) heleneae, and include details of the animal and its habitat. Materials and Methods The material reported here is deposited in the South Australian Museum, Adelaide (SAM), Australian Museum, Sydney (AM), Museum of Victoria, Melbourne (NMYV), Northert) Territory Muscum of Arts & Sciences, Darwin (NTM), Queensland Museum, Brisbane (QM), Western Australian Museum, Perth (WAM), Tasmanian Museum and Art Gallery, Hobart (TM) and National Museum of New Zealand, Wellington (NMNZ). Living material was collected by “SCUBA” and “Hookah” diving and using a Smith- Melntyre grab dredge. Underwater photographs of the living animals were taken using Nikonnos IL and Nikonnos V cameras with macro extension * South Australian Museum, North Terrace, Adelaide, S, Ausr. 5000 ) Kinhill Mating Scenes, 200 East Terrace, Adelaide, S. Aust. S000, tubes and clectronie flash units, Wet material was preserved in 75% methylated ethanol saludo oe 2% formalin/propylene glycol solution, Radulae Were prepared for examination under scanning electron microscope (SEM) afler (he method of Bandel (1984), Colour descriptions follow Kornerup & Wansehe: (1978). Primoyvula (Primovula) helerete Cate, 1973 FIGS I, 3, 4, Ovule farmoasa; Verws, 1908, p. 342 (non Actans & Reere, RAB), Prosuiunta verconis: Couion & Godley, 1932; po a6, pl. 1, fig, TA (in part). Pellusimriia verconis: Tredale, 1935, f, 105; Coren, TYS9, p. 368, (in part), Primovula (Primovula) vercanis: Schilder, 1941, p. 107, (in part). Neaspinia (Pellasininia) verconis: Allan, 1956, p. 130, pl. 14, Ne. 40 (in part), Primovula (Primovula) helencue Cale, |873a, po 4a—4, li, B9, Type: Holotype (SAM 1915943), part of the type series examined by Cotton & Godfrey (1932); a paratype of Prosiminia vercenis. Type locality: Holotype dredged in Gill St Vincent or Spencer Gull, S. Aust., by J. C. Verco; the holotype of PB yerconis was dredged of f St Francis Island, S. Aust, by J C. Vereo. Other material: SAM D17973 (4 + eggs, i spiril, SAM DI7975 (2, dry) Douglas Poilit, northeri Spencer Gulf, S. Aust., N.C, Holmes, 171.1987; SAM D18434 (13, dry), SAM D18435 (8, in spirit) Douglas Point, northern Spencer Gull, §. Aust, N, J.C, Holmes, 5.v, 1988, Species description: Small allied cowrie, adall length 9.15-13.80 mm, mean 10.77 mm + J.004 Elongare; both ends slightly produced, posterior end mare so; widest part of dorsum in posterior 4 (Fig. 3B), - GOWLETT-HOLMES & N. J. C. HOLMES Fig. 1. Primovula (P) heleneae, two adults, one with partially extended mantle, on the gorgonian coral Euplexaura sp. Fig. 2. Crenavolva cruenta sp. n , two adults with partially extended mantles on the gorgonian coral Echinogorgia sp. REDISCOVERY OF PRIMORULA (PR) HELENEAE CATE 2U7 Dersim (Fig. 3A) sculptured with numerous cransverse Wavy striae; funiculurn prominent, with three weak transverse ridges, Base (Fig. 3B) convave, outer lip and patietal-columeflar lip Nat and inward sloping; aperture narrow with a strong fossula, both sides strongly calloused. Parietal-columellar base smooth but wih body whorl striae visible beneath the callus. Outer lip with weak transverse ridges posteriorly, inner margin slightly dentate Posteriorly, Outer margin with three to ten slight fo prominent nodules at posterior end, mare prominent in smaller shells; usually with one to three slight nodules at anterior end, Posterior canal somewhat produced, with a basally projecting terminal knob dividing canal inte distinet Jeft and right channels (Pigs 3C, 3D). Auiterior canal shallow “U-shaped, sliglitly Narng and jurned to the right, bordered on both sides by distinct and slightly projecting terminal ridves. Shell pastel red to pale orange re ercyish yellow, anterior and posterior tips usually orange; paler band across widest part of dorsun\s colameliar vallus, outer Jip and labial callus all paler than dorsum, Animal mantle lobes (Fig. Lb with oo papillae; translucent pastel red to dark orange, with irregular, reddish brown patches varving in size, area immediately around each pateti not patterned, remainder of mantle patterned wih slightly raised, while dots, Poot pinkish white; anterior dorsal foot with a series GF prontinent, vertical lamellae; posterior dorsal foot with a longitudinal, violel brown (6 brown stripe, Tentacles with while up for 1/6 length, violet brawn for almost half length, remainder pinkish white with basal 1/5 from eve swallen, with violet brawn stripe from eve to base. Siphon and preboscis violet brown to brown, Radula (Fig. 3F) with central teeity narrow at base, will large, broad, flal heads, outer edge with many perpendicular, sharp cusps, central cusp very large, mairow at base, Widening for about half length, where there is usually a lateral denticle on ar least one side, then natrowlnie apically fo sharp point) with eight or qine small cusps on either side of central one Firs} lalyral teeth elongate. strongly curved, heads tricuspidate, cusps Lineven, central cusp largest. Secand Jateral teeth elongate, with broad, Nat heads, heads with aboul five long, curved, sharp-tipped cusps. Eee mass (Fig, 4A) with 57 colourless, transparent, jellylike capsules laid on a single branely of the host sequentially in an elongated spiral and fused together, Fach capsule contains minty White-shelled embryos, shells wider than long, shell lengli 150-230 um, shell round, punctate with thin operculum (Figs 4B, 4). Range: The new material examined here is fram nothern Spencer Gulf, S. Aust. Cate (1973a) pave the type locality of # (PB) heleneae as the “S. end ol St Vincent Gull, South Australia, (34°5S'S 138°10'B)", but the holatype is labelled in Verco’s handwriting "“G: St Vincent or Spencer Gulf, dredved”, so Cate's (1973a) restriction of the type locality is suspect, The range of this species is assumed to be central S. Aust. It is noteworthy thar Cotton (1935) refers to the occurrence of Crenavolva verconis in southern W.A., but we can lind no specimens to substantiate it, Care (1973) refers Cotton's (1935) reference (0 a Queensland species, Mrimovula (P) platvsia Cate, 1973 tas Ovula Jormosa Vereo), but does not mention if le cxamined the specimens, Habitat: On fans of (he gorgonian coral &uplewaure sp, (Caldaria; Octocorallia: Plexauridae), in areas OF strong tidal movernerit. Remurks: The specific identity of the host coral (SAM H355) could pot be determined (PL Alder- slade pers. comm,), The white spicules (Pig. §C) are typical of Euplexeuna, ln northern Spencer Gulf, colonies (Fig. SA) are multibranehed fans In one plane perpendicular to the strong tidal currents, (ey ire attached by & thickened base to a large shell fragment or other piece or pieces of solid debris in the substrate; a poorly sorted, silty, shelly sand. Colonies are relatively large (ca 30 cm |ileh}, and Widely and irregwlarly dispersed at 1-20 m depeh_ Distribution appears to be associated with tidal current velocity and substrate siltiness, Buplevaneg SP. appears Lo be less toleraw of silt than Behine gorgia sp., and is less common. Each Explexmne sp. colony usually hosts two to five individuals of P (PF) heleneae. Polyps are usually withdrawn during daylight on all of the colony regardless wr the presence of PF (FP) helenewe. Orientation of (he shell is always with the longit\dinal axis parallel ro the branches of the host, with po observed preference fora head up of livad down attitude. Na particular position on (he host is favoured, and adults and juveniles are present together, Fags have been observed in Jate autumn and early winter, laid on a single branch of the host and pot altered by the parent, although adults were present on the Lost. The colour pattern of ihe extended mantle lohes Of F(R) heleneae is almost identical to that of Euplexaura sp, The background colour of the coral is pastel ted, and rhe withdrawn polyps are red to reddish brown, The pattern of reddish brown patches an the mantle lobes of PP) heleneae mimics the position of the polyps on the earal, ane! an animal with ihe mantle lobes fully extended is 208 K. L. GOWLETT-HOLMES & N. J. C, HOLMES hi very Well camouflaged. The very close mimicry by P. (P) heleneae of its host suggests that it is an obligate predator and that its distribution will be limited by that of Euplexaura sp. The distances between colonies of Buplexaura sp. (10’s of metres) suggest that adulis of PR (P) heleneae would not move between colonies, so dispersal is probably by a free-swimming veliger stage of short duration. Crenavolva cruenta sp. nov. FIGS 2, 7. Holotype; SAM D18431, dry adult specimen 10.45 x 3.60 x 3.00 mm, collected on the gorgonian coral Echinogorgia sp. at 15 m depth, 1.6 km E. of Douglas Point, northern Spencer Gulf, S. Aust., 32°5)'4R"S, 197°49'12"E, by N. J. C. Holmes, 5.v.1988. v. 3. Primovula (P) heleneae (SAM D17975), A. dorsal view, x 7.5; B. ventral view, x 7.5; C & D. lateral views, x 7; E. size range, x 2.5 (SAM D18434); F. radula, scale bar = 10 pm (SAM DI7973). Paratypes: Same locality data as holotype. Spirit specimens: SAM D18433, 14 juv., 4.65-9.60 mm, 24 ad., 7.35-11.65 mm, Dry specimens: SAM D18432, six juv., 6.30-9.15 mm, 27 ad., 7.15-12.35 mm. SAM D18437, 11 ad., 6.70-12.60 mm. AM 156094, two ad., 9.55 and 10.90 mm respectively. NTM P1384, two ad., 945 and 10.20 mm respectively. NMV F52836, two ad., 9.35 and 9.95 mm respectively. QM MO.19385, two ad., 9.40 and 9.70 mm respectively. TM E17744, two ad., 9.15 and 9.35 mm respectively. WAM 152/88, two ad., 8.30 and 10.30 mm respectively. NMNZ MF. 49494, two ad., 8.65 and 10.60 mm respectively. Other material examined: Two broken specimens in spirit from Douglas Bank northwest, northern Spencer Gulf, S. Aust., 32°47°18"S, 137°49°12"E, E. Oks, Sept. 1987 (SAM D1i7976); Backy Point REDISCOVERY OF PRIMOVULA (P) HELENEAE CATE 209 Fig. 4. Primovula (PR) heleneae (SAM 117973), A. egg mass on branch of Fyplexaura sp., x 4; B. embryonic shell, dorsal view; C, embryoni¢ shell, showing aperture and operculum, scale bar = 10 ym, cast, northern Spencer Gulf, S. Aust., 32°55'00"S, 137°51'24"E, E. Oks, Sept, 1987 (SAM D17977). Diagnosis; Small allied cowrie, adult length 6,70-12.60 mm, mean 9,51 mm + 1.2. Elongate; both ends produced. Dorsum sculptured with numerous transverse wavy striae; funiculum not prominent; aperture narrow with strong fossula; Outer lip outer margin with slight to prominent terminal nodules; both canals bordered by slightly projecting terminal ridges. Shell greyish orange to brownish red; tips red to orange; dorsal bands and patches usually present, white, pale yellow to pale orange; columellar callus, outer lip and labial callus yellow to light orange; fossula pinkish white. Animal mantle lobes translucent pale greyish pink with dark red spots and small and large white papillae; foot ruby, underside with median longitudinal very pale pinkish white Stripe; tentacles with white tip for 4 length, remainder of tentacle, siphon and proboscis ruby. Description of Holotype: (Figs 7A-D) Shell clongate; both ends produced; widest part of dorsum in posterior 2/5, Dorsum sculptured with numerous transverse wavy striae; funiculum not prominent, smooth. Base somewhat concave, outer lip Mat, inward sloping; parietal-columellar lip convex; aperture narrow with a strong fossula, both sides strongly calloused. Parietal-columellar base smooth with faint transverse striae present on central inner part, Outer lip with rounded transverse ridges, becoming weaker anteriorly; inner margin smooth, outer margin with four slight nodules at posterior end, three slight and one prominent nodule at anterior end. Anterior canal shallow “U"-shaped, slightly flaring, bordered on both sides by distinct and slightly projecting terminal ridges. Posterior canal somewhat produced, shallow “U-shaped, strongly flaring, bordered on both sides by distinct and slightly projecting terminal ridges, outer lip ridge terminating in a nodule, parietal-columellar lip ridge with two nodules, one terminal, Shell orange red, anterior and posterior tips bright red; light yellow median band across widest part of dorsum, continuing onto base; light yellow diffuse band midway between median band and posterior end, continuing onto base; light yellow patch almost midway between median band and anterior end. Outer lip and labial callus yellow: columellar callus translucent yellow with dorsal colour pattern visible. Interior of posterior and anterior canals red, becoming orange red at very tips. Fossula pinkish white. 210 K. L. GOWLETT-HOLMES & N. J. C, HOLMES A » Fig. §. A. Euplexaure sp. colony in situ, x 0.8; B. Echinogorgia sp. spicules, scale bar = 100 pm (SAM H356); C. Euplexaura sp. spicules, scale bar = 100 pm (SAM H355). Etymology; “Cruentus” (L.) meaning bloody, with random, dark red spots, rows and groups of pertaining to the main colour of the shell and the — small, simple white papillae and occasional much animal. larger, white, elevated, rounded papillae with wart- like knobs, mantle more opaque white at base of Additional Characters from Paratypes: Animal papillae. Foot uniform ruby, underside of foot with mantle (Fig. 2) lobes translucent pale greyish pink median, longitudinal, very pale pinkish white stripe; REDISCOVERY OF PRIMOVULA (PR) HELENEAE CATE 211 Fig, 6. Echinogorela sp. colony in situ, x 0.75. anterior dorsal foot with series of prominent, vertical lamellae, becoming less distinct posteriorly, disappearing about '4 length of foot from anterior end, Tentacles with white tip for 4 length, remain- der ruby with basal Ys from eye swollen. Siphon and proboscis ruby, Radula (Fig. 7F) with central teeth narrow at base, with large, broad, flat heads, outer edge with many perpendicular, sharp cusps, central cusp very large, tapering apically to sharp point, with sharp, lateral denticle on either side at about half length; with five or six small cusps on either side of central one. First lateral teeth elongate, strongly curved, heads tricuspidate, cusps uneven, central cusp largest. Second lateral teeth elongate with broad, flat heads, heads with eight long, curved, sharp-tipped cusps Fariation: Paratypes differ from the holotype in size, shell colour and sculpture (Fig. 7E), Shell colour is from greyish orange to brownish red: anterior and posterior lips red to orange; dorsal hands and patches white, pale yellow 10 pale orange, and yary from very prominent (often forming three distinct bands) to almost absent, median band being most conspicuous; columellar callus, outer lip and labial callus yellow to light orange. Funiculum varies from smooth to with up to five weak nodules on inner edge. Outer lip outer margin nodules vary from very slight with one prominent nodule at anterior end, to three prominent nodules at posterior and two prominent nodules at anterior end. Posterior end usually much more produced in small specimens, almost rostrate. Posterior canal parietal-columellar lip ridge occasionally with three or four nodules. Range; Northern Spencer Gulf, S. Aust. FHlabitat; On fans of the gorgonian coral Echinegorgiu sp, (Cnidaria: Octocorallia: Paramuriceidae), in areas of strong tidal movement. Comparison with other species: The new species belongs in Crenavalva, which is distinguished from other genera in the Ovulidac by the small, elongate shell with median angulation, narrow, produced but truncate ends, dentale, curved aperture, striate dorsum and ridged outer lip (Cate 1974), C eruenta most closely resembles C. striatula (Sowerby, 1828) and C. verconis, the congeners recorded from Australia, Cate (1973a) records two subspecies from Australia, C. s. traillii (A, Adams, 1855) and Cs, tinctiira (Garrard, 1963), both from Queensland. C cruenta can be distinguished from C. striatula and C. verconis by its narrower, more elongate form, more produced ends, the shape of its more elongate aperture, and shell colour pattern. Remarks: The specilic identity of the host gorgonian coral (Eehinogorgia: P, Alderslade pers. comm. SAM H356) could not be determinded: the red thorn scale spicules (Fig. 5B) are typical for the genus. In northern Spencer Gulf, colonies of this species (Fig. 6) are sparsely branched fans in one plane, attached by a slightly thickened base to shells, fragments or pebbles in the substrate: a poorly sorted, silly, shelly sand. Colonies are relatively small (ca 15 em high), and appear much more tolerant of silt (han Euplexaura sp., being widely 212 K. L. GOWLETT-HOLMES & N. J. C. HOLMES Fig. 7. Crenavolva cruenta sp. nov. holotype (SAM D18431), A. dorsal view, x 6.6; B. ventral view, x 6.6; C & D. lateral views, x 6.6; E. size and colour pattern range, paratypes, x 2.25 (SAM D18437); F, radula, paratype, scale bar = 10 pm (SAM D18433). but patchily distributed in 10-20 m of water, often in quite high densities (up to 20 colonies/m’). When C. cruenta is present, each Echinogorgia sp. colony usually hosts one to three individuals. From observations made in daylight, the host’s polyps are usually withdrawn on all or most of the colony when C. cruenta is present. Orientation of the shell is always with the longitudinal axis parallel to the branches of the host, with no observed preference for a head up or head down attitude. No particular position on the host is favoured, and adults and juveniles are present together. Colour pattern and form of the extended mantle lobes of C cruenta are almost identical to Echinogorgia sp. The background colour of the gorgonian coral is red: the polyps are reddish orange when withdrawn, yellow when extended. The thorn scale spicules in Echinogorgia sp. protrude through the surface of the colony, giving it a rough texture. These protruding spicules help retain a ragged layer REDISCOVERY OF PRIMOVULA (P) HELENE AL CATE 213 of mucus and detritus upon the surface: this nucous layer is white, and patchily masks the red background colour The pattern of reddivh brown patches and white papillae an the mantle lobes ol C. erventa mimics the ragged mucus on the vorgonian coral, and an animal with the mantle lobes fully extended is very well camouflaged on ity hose. The very close mimiery of C. ervente to Its host suggests that it is an obligate predator and that its distribution is limited by thar of Eehinogergia sp. The distances between colonies of Echinogorgia sp, (often <0.5 m) and the suitability of C. cruente§ camoullage suggest that the latter would be able to nove across the substrate between colonies, so dispersal is probably only partly by a lree-swimming velier. Discussion The generic names used in this paper follow the sysiem and modifications proposed by Cate (19734, 1973b, 1974). The distinctions between some of the genera used there are inadequate, The two species of Ovulidve previously named from, Aust: Creravolva verconis and Primovuta (PR) feleneue, are both based an damaged, dead shells. Vereo (1908) had three specimens und recorded them as Ovula formosa A. Adams. & Reeve, 1848, a species described from Borneo, The deseripvion and photographs of the holotype of O. Jormosa [= Delonavolya_forniosa (A, Adams & Reeve, 1848)] in Cate (19734) were compared with the holotype of CG verconis, with the holotype and other specimens of P (P) kelencae, and with the types Of ©. crventa.and D, formosa was lound 10 differ sivnificanily from them, D. Jormoasa is bright purple with vellow tips: the fossula is not prom: ihent; the whole shell is-much narcower overall, with a narrower outer lip than either C. crwenta, C verconiy or F(R) helenewe, and a more pointed posterior Up than either © eryenra or GC. verconis, and which lacks the basally projecting terminal knoh of PR (P) heleriewe Coton & Godfrey (1932) described Verco’s specimens as a new species, Prosimnia vercanis, Their desctiptian obviously was based on all the specimens, although the holotype is clearly defined as the shell trom St Francis Island (SAM DI3476, = DJ0I74). Cate (1973a) re-examined the Lypes of & verconis, which riow consisted of two specimens, (one specimen of the original series is missing, presumed lost). and discovered that they were no! conspecific. Cate avknowledged the designation by Cotton & Godfrey (1932) of (he specimen registered SAM DI3476 (= SAM. DI0I74) (very worn and faded), as the holotype of Crenavelya verconis (Cotton & Godfrey, 1932), and used Cottan & Godfrey’ (1932) paratype (SAM D15943), a worn, faded and broken specimen, us the holotype of Primovule (P) helenege. The live collected material from upper Spencer Gulf was compared with the types of both species and, although both types are damaged, the material callected from Euplexaura sp. was clearly conspecific with P (P) heleneae. and all of the new material was different from the designated holotype of GC. verconiy. According to Coleman (1981), the specimen illustrated by: Coleman (1975, fig, 262) as Neosimuia verconis (Cotton & Godfrey, 1932) is actually a specimen of an unnamed species of Ac/yvolve. Acknowledgments We are grateful ta Dr B. R. Wilson for his considerable assistance and critical comments, and to Mr PB. Alderslade for identification of the gorgonian corals, Mt W, Zeidler, Dr -C. Watts, Dr TE. Wells and un anonymous referee are thanked for critical comments, We thank Mr R. Thomas, Mr D. Evans, Mr M. Moore and Ms J. Young, all of Kinhill Marine Sciences, for assistance in the held. The photograplis (other than Figs 1, 2, 7, SA, 6) were taken by Mrs J, Forrest and Ms A. Rentrey (SEM). The colour plates were fiinded by the Research Fund of die South Australian Museum. 2l4 K. L. GOWLETT-HOLMES & N. J. €, HOLMES References ALLAN, J. (1956) “Cowry shells of world seas.” (Georgian House, Melbourne). BANDE, K. (1984) The radulae of Caribbean and other Mesogastropoda and Neogastropoda. Zool. verhandl. Leiden 214, 1-188, pls 1-22. Cate, C. N. (1973a) A systematic revision of the recent cypraeid family Ovulidae (Mollusca: Gastropoda). Veliger Suppl. 15, 1-116. (1973b) The Ovulidae: Replacement names for four ovulid homonyms. Veliger 16(2), 238-239. ____—s«(1974) The Ovulidae: A key to the genera, and other pertinent notes (Mallusca: Gastropoda). Ibid. 16(3), 307-313. COLEMAN, N. (1975) “What shell is that?” (Hamlyn, Sydney) _ (1981) “What shell (Lansdowne, Sydney). is that? 2nd Edition.” Corton, B. C. (1935) “Combing the southern seas.” (Rigby, Adelaide). (1959) “South Australian Mollusca: Archaeo- gastropoda.” ( S. Aust. Govt Printer, Adelaide). & Goporrey, F. K. (1932) South Australian shells (including descriptions of new genera and species). Part LI. S. Aust. Nat. 13, 35-86. GARRARD, T. A. (1963) New species of Mollusca from eastern Australia. J. Malac. Soc. Aust. 1(7), 42-46. IRLDALE, T. (1935) Australian cowries. Aust. Zoal. 8(2), 96-135. Kornerup, A. & Wanscier, J. H. (1978) “Methuen Handbook of Colour. 3rd Edition.” (Eyre Methuen, London.) ScHILDER, F. A. (1941) Verwandtschaft und Verbreitung der Cypraeacea. Archiv Mollusk. 73(2/3), 57-120. Verco, J. C. (1908) Notes on South Australian marine Mollusca, with descriptions of new species. Part LX. Trans. R. Soc. S. Aust, 32, 338-361. DEVELOPMENTAL BIOLOGY OF THE AUSTRALOPAPUAN HYLID FROG LITORIA EUCNEMIS (ANURA: HYLIDAE) BY MARGARET DAVIES* Summary The egg mass and tadpole stages of Litoria eucnemis are adapted for a stream-dwelling habit. The egg mass is compact and adherent whilst the tadpoles have suctorial mouthparts and shallow fins. The eggs have a large yolk reserve used by the tadpole for nutrition between stages 19 and 26 after hatching. At 16-24°C, L. eucnemis metamorphoses 74 days after egg laying and 69 days after hatching. Trammactions ul the Royal Society af S Aust, (989), TIB4) 2S-20 DEVELOPMENTAL BIOLOGY OF THE AUSTRALOPAPLAN HYLID FROG LITORTA EUCNEMIS (ANURA: HYLIDAF) by MARGARET Davies* Summary Davirs. M. (1989) Develupmental biptogy ul the Anstralopapuat tiylid thoy Litera eecneniy (AnuTa Hylidae), Trans. Rk, Soc, S. Ase, 113(4), 215-220, 30 November, (989, The eve mass and radpole stages. of J oma egenentis are adapted for a stream-dwelling habit. The cog Mass J compact and adherent whilst the tadpoles have suctorial mouthparts ani shallow fins, The caps have a large yolk reserve used by the midpole tor nutrition between stages 19 and 26 wfler hatching, At 16-24°C. 1, ehore ms Metamorphoses 74 days alter eye laying and 69 days afer hareliine. Kiy Wrebs: Liter eucaesns, tadpoles, developmental biology, Joie adayrarons introduttion fuloria encnemis (Lonnberg) is disrribured in rainforest streams act widely-separated localities in New Guines (Tylec 1968; Zweifel 1980) and on the Cape York Peninsyla in northern Queensland (Zweifel 1980, Couger 1986). The species has had un unstable nomenelatural history in Australia (Tyler & Watson 1985) and the names /iteria ewnemis, L. serrata und ©, genimecidata have variously been applied 10 Australian representatives of the species. Life history data are confined to 4 comment that the eggs are dark brown and that nothing appears distinctive about the tadpole (Menzies 1974). A pair of £. eucnemis bred in the laboratories of the Zudlogy Department, University of Adelaide. The tadpoles were reared to metamorphosis and the development of the spevics is reperted here Matevialy and Methods Specimens of 1. ewcnenris were collecred by M. Mahony, J. Read, M. Schmidt and T, Haff at Charmiltin Ck, Qld (17°43 145931‘) in late August 1988 and reached Adelaide on |ix.1988. A pair of frogs, amplectant upon arrival, Was (ransferred to a glass aquarium with a pond of water atone end. The aquarium was muimiained at oem temperature. Spawn was removed from the pond, and placed in dechlotinated, acrated jap water in a glass aquurium 25%25x8 em for 24hr at 30°C. The aquuzium was then maintained at room temp- erature, aerated and the walter chiinged daily. Tadpoles were fed on lightly-boiled lertuve leaves. The culture was divided in two on 26.48.1988. To give the ladpoles roweh surfaves on which to attach, alga-covered stones were introduced to the aquaria when the spawn hatched, * Deparoment of 7Goloyy, Universiry af Adelaide, GPO. Bax 498, Adclaide S. Aust. S001 Samples of embryos and larvae were vollected as shown in Tuble PF and preserved in Tylor’s fixative (Tyler 1962), Measurements of developmental stages were taken usity vernier cullipers reading, fo 0.05 min. Hiustrations were made with the gid-of a Wild Ms stereodissecting microscope and camera lucida. Developmental stages were classified acyardins: to Gosner (196()), Results Aniplevuss Amplexus was axillary (Fig. 1), The pair was amplectant upon arrival in the tbordtory possibly ue response to the reduced aie pressure wi the cargo hold of the aeraplane (M, J ‘Tyler & M. Davies unpubl. obs.), &ggs: Oviposition took place overnight, 2 and 3.ix, 1988 Temperature of the water in which the egps were laid was 18,5°C, The egy tatss of 843 Guys was 67.5. mim at its greatest width und compact Individual jelly capsules were tightly adherent to each other (Fig. 2). The animal pole was dark brown and approximately half of each egg on the vegetal hemisphere was white. Only one membrane was detected in each capstle: Mean diameter of six eggs was 2.25 mm (range 2,10-2.56 mm) and of the capsules, 2,98 nim (range 2.40-3,60 mm). On the day following oviposition, the eges were in lare cleavage (Stave 9), Within 24 hours they hadl progressed to (he neural fold stage (Stage I4) and within a further 24 hours. the embryos were at Stuge 17. The cinbryos were perched on a large chovolate-coloured yolk sae (Fig. 3); larpe crescentic adhesive organs were located anterodorsally, A prominent stomoducal pic was loculed posteradorsally io rhe adhesive organ (lig 3), On Tax. d9RK, rhe embryos remiined in the jelly capsules. The adhesive Organs were beginning: to divide ag head flexion conuneneed, The storbodaent netch was prominent and jhe pronephiros wus apparent laterally; the will plate was vleurty 216 delineated. The tail was differentiated and the embryos were at Stage 18. Larvae; The embryos hatched on 81x.1988 al Stage 19. The head was flexed ventrally and the adhesive organs situated ventrolaterally as two raised pigmented suckers. A stomodaeal depression was dorsomedial to the suckers, The primordia of two pairs of gill filaments were present and eye bulges were apparent (Fig. 4a). The pronephros was M, DAVIFS still apparent. A large yolk sac provided nutrition for the embryo. By 9.1x.1989 the embryos were at Stage 20. The two pairs of gills were now filamentous, the anterior pair with four to five branches, the posterior pair with one to three, Ventral suckers were ventrolateral to an open mouth. The optic bulges were well developed hut opaque. Nutrition was supplied by a large yolk sac (Fig. 4b), TABLE |. Dimensions of developmenral stuges of Litoria eucnemis. Age Slage Body length Total length il (days, date) A, range in A, range in parentheses (mim) parentheses (mim) 6 19 312 5.07 6 (8.1x.1989) §2,.96-3.18) (4.95-5.25) 7 20 3.43 6.6 3 (21x. 1989) (3.3-3.3) (6.4-6.8) 8 20 3.47 8.03 3 (10.1x.1989 (3.25-3.9) (7.8-8.2) y 21/22 3.65 8.68 \2 (LLix A989) (3.4-4.05) (8,0-9,4) 12 a4 10,96 Ms (15.1x.1989) (9.9-12.0) 14 28 12.3 13 (17.18.1989) (11.0-13,2) 23 26 17.07 10 (27.1x. 1989) (16,1-18.1) i at 16.8 3 (5.x.1989) (16.7-17,0) 42 30 42 1 (26.*.1989) 42 4 7,92 24.5 5 (26.x.1989) (7.0-8.4) (24,1-25,2) 42 a OAS ate: 2 (26.x. 1989) (8.8-9,35) (26.0-28,5) 42 33 Ya 27,9 ! (26.x.1989) 42 44 9.1 28.9 3 26,.,.1989) (8.7-9_6) (27,0-30,8) 50 a4 B98 27,5 4 CXL TMD) (8.7-9.2) (26.5-24,9) 50 35 985 2k A 2 (A.NL1989) (97-100) (27,5-28,7) 50 36 10,4 33.7 2 (3.81. 1989) (10.3-10,5) (33.5-33.9) 54 6 9.65 30,3 z (7x1. 1989) (9,3-10.0)) (29,3-31.3) 54 a4 7 29.6 2 (7.x), 1980) (24-101) (2K.()-31.2) $4 4) Wd 34.6 | (Txt A9RY) 57 41/42 9.8 20.4 | (10.81.1989) 38 42 10.6 31.2 | (xi 989) 6] 3 Hk 22.1 | (4.x. 1989) 6l 4S 10,5 14.0) | (14.n1.1989) 46 1.2 - 2 63 (16.x1.1989) DEVELOPMENT IN LITORIA EUCNEMIS 7 Fig. 1. Amplectant pair of Litorta eucnemis. By 10,ix.1988, the larvae remained at Stage 20 bul olfactory pits were now clearly visible. Larvae had reached Stage 21 within the next 24 hours. The cornea was (transparent, pigmentation of the adhesive organs was fading and the gills were more filamentous (Fig. 4c). Ridges could be distinguished on the lateral and posterior margins of the mouth, foreshadowing the labial teeth and papillae. The tail had elongated; nutrition was still supplied by yolk. By 15.ix.1988, the external gills were completely covered on both sides by the operculum. The adhesive organs were degerating, although some pigmentation remained. The horny beak was keratinized, but the tooth rows had not yet formed. Nutrition was still provided by yolk and the embryo was at Stage 24. Some tadpoles were now free swimming. A hig. 2. Dorsal view of a spawn clump of Litoria eucnemis. Greatest diameter ix 67.5 mm. The spiracle had begun formation by 17.ix.1988. It was situated ventrolaterally and the opening directed posteriorly. Two upper and the two lower entire tooth rows were keratinized. The gut was loosely coiled but yolk-filled, and embryos were at Stage 25. Larvae were nol sampled again until 27.ix,1988. The spiracle had elongated and its orifice was now directed posteradorsally. Mouthparts were com- plete, the larval gut had formed and the tadpoles were feeding on vegetable matter. The development of the limb buds indicate that the tadpole was at Stage 26. eS | Fig. 3. A) Dorsal and B) anterior views of embryo of Liraria eucnemis at Stage 17, Scale bar represents 1 mm. 218 M. DAVIES ea Fig. 4. Litoria eucnemis larvae: A) lateral view at hatching, Stage 19; B) lateral view Stage 20; C) lateral view Stage 21. Scale bar represents 1 mm. DEVELOPMENT IN. LATORTA EUCNEMIS 214 | dt (nea Yl } Duy Ae Fig. 5, A) Dorsal and 8) lateral views and €) mouth disc of a lacva of Litarid edcnemis ar Stage F1. Seale bur represents 5S mm (A,B) and 1 mm (C). The next sample of larvae, taken on 5.x.) 988, was at Stage 27. The body of the larva was covered with brown pigment dorsally, and was pale ventrally. A disrupted fine of pigment granules bordered the dorsal myotomes and further pigment granules were scattered laterally and on the dorsal fin. Dimensions of developmental stages of Ly eucnemis are shown in Table |, A larva at Stage 31 preserved on 26.%.1988 is illustrated in Fig. Sa.b, The anus was dextral and the spiracle elongate and sinistral. The dorsal fin way shallow and tapered to a genily-rounded tip, General body colour was brown with faint pigment granules on the tail myotomes and the dorsal fin. The mouth disc of a larva at Stage 31 is shown mn Fig. Sc. There are two upper and three lower rows of labial teeth, Both the second uppec and the first lower row of teeth have a median gap. There are two rows of labial papillae forming a wide band around the lateral and pasterior murgins of the mouth. Metamorphosis: By 10.xi 1988, the lett forearm had protruded in one larva and the first froglet ut Slage 46 was recorded on 16.x7. 1988, a toral of 74 days after the egws were laid and 69 days afler hatching. Water temperatures varied throughout the pertad of development from 16.4 -23.8°C, Larval behaviour: Larvae frequently avereuated, mostly beneath the stones within the aquaria or on the surface of the lettuce. ‘They attached theriselyes by their suctorial mouthparts to the floor of the aquaria during water changing, 70 NM Discrission Nhe develupment of Litoria enonenus is wrusual amongst Australian hylds tutilising a lange yotk reserve and showing relatively lite development ol the larval ut (the speaes bilehes at stage 19 hurt the gut docs not form until stage 26, 9-19 dityn utter hatching), Such development is more representative of dircet-developing species or those (hit spend a considerable period of Iheu development within the cee capsule [as Avartans sphaunecotus (Moure 1961: Anstis }981)]. Amougst Ansiralian species, this mode ot development is restricted fo leprodactytid frogs. £. ewcnens is a New Guinea species that extends its range wito Australia, and irs relationships lie with the new Guinean hylid fauna, in particular the momtune species of Liter and Nictinysies. Such species lay huge unpigmented eves: tadpoles, where known. exhibit love adaptations (Menzies 1974; Ther & Davies 1979; Zweifel 1983). However, complete life bistiory data gre unknown for these species, The exes of L. eucnemis ure Jame in Comparison with other Australian hylids (see Lyler 1989 for review), direetly related to the made af early development. [Montine New Guinea Litata and Nyediniystes species alsa are described as huving hinge eggs (Menzies (974), The single adherent e2 mass is an adapuon in Lirarta eucnenis lo 4 orig lifestyle: (he tooth rows and orher sttwerires are typical of other Australian hylids (Marin & Watson 1974), but dig suctorial moulli disg is an adaptation to fast-flowing waters, ASIES Amongst Australopapuan amuriins, there ire several kinds of lofic adaprationts of (He MOUTH cise. Nhe vstreme claborations exhibited by Aitartd ndjnols, bo rheocala (Liem W874), L. urhek heed (Menzies 1974), and Nvetourstes teed (see Lyler 1989 for Plate) consist of a large Suetormil Arai oveupying the entire ventral preoeular reyian of the tadpole. Juuducev/us spp have umbrella-shaped hips hanging vertramedially (Licm & Hosoter 1973) whilst a2. subvlendulosa (he moutliparts form a funnel in whieh all labial tooth rows and piznentation af the homty beak are absent, Oral dise tubercles have keratinised tips whilst unusual black filaments sre suspended anteriorly i the pharyiny (lyler & Anstis 1975). Lhe less elaborate acaprations occupying & lesser Proportion of the anleroventral surface of the Gtdpote ane exhibjted by f encnemnts are shared by L, copland? and L. theirlana (Tyler ef al, 1983), L. deseenes (Martin & Watson 197), Afeeistofaris fipmariay Clyler eral 1979) and Aiivopives balbus (Watson & Martin J97A9). Orher torrent adapiions include the narrow (ail tins and the ventrolateral spiricular opening, Ack rowledgiients thank Michael Maliony for the provision of the adult frogs, Michael. Tyler assisted with ulpote rearing and critically read the manosedpr “This work was supported by Liniversity of Adelaide Deparment of Zoology funds, References Anolis Vi (O81) Breeding biology and ramige extension Jor he Sew Sail: Walos frog Avee cide spatagarcolis cami Leptadaervlidaed wtiat A Aerperok b. e9. Comoch H.-G. (986) 'Reprles ind Amphibians of Anatralia” 3rd cdi (Reed, ‘Syelney), Cine tt. Ky (M60) A Simplified Galle len staging anus embryos and kavae with oles on idenrifieation, Herpetolamicn WG, TRIAT. Lira, Po oS. 9TH) A review Of the Liferte tn ony spemes BOP and a desenpned Of a few spouts it Citurie Tom serthers Queenshaids Australia (Agora, Hylidae, Mew Od Muy. (71), 181-168. —_ & Hosen, We (M74) Frags of the genus Tundaoss hes vith descriptions of Awe fiw species jranura! beplodactylidae) Aten, Qa Miia. 1603), 444457, Martin, ALA. & Wavptoem, Gy be (ISL) Life history as an aid (oy senerie delineation in Wie lnnly Myhidae, Cuprerer (UY), FR T9, Misges bol. (1974) Handbook Gf eonmen New brute frogs, WAL Leology Tastinite, Handbook not, Nionar DA. UY6L) Fons of eistern New Soh Wiles. Bult Am Mus. dat, Host, W234, 14-3R6 Trice M. 4. (19862) On the preservedon ar anuran tapoles. Anat de Seb 25, 222, —___()908) Papuan hytid frogs of (Ne genus Lyle feed Ferhanid, Leiden Ga, iid, C1989) “Australian NMelhourne), A Aves M. (1975) Tuxanony and biobogy of (rogs of Uy (iota ctrapa comples (ANOKE Mytidae), Reo, So oteert Ms, V7, 41-40). 8, Choke Ch AL & Dawes, M, (1989) Repracuertye Wiolowy ar the fross of the Magehe Creek system, Northern Terriers Mid 18, $15 -44b, — J& Taviks, My, (979%) Kedefiniion und by evolutionary ongio of the Austalopapnian frog genus Nowdionvszes Steyneger, AWM A Zu0f, 27, T38-972, Manin, Ay Ao Davis, M. 11979) Biology ected eesrermaticy of anew Tiniiodyoasting gemus (yaa Leptodacivlidae) from mgr thowestornd Alsiritia. lest A Foul, IT, V3AS-350. & Wasson, Gok. (O84) On ihe nomenchiare of oy hytid tree frog trom Queenston Phan Ae Sec 8. dase WO84), 19s t94, Woavvsos, Ch FA MARIN AL AL (1973) Lite besten, larval morphology and pelavodships of Australian Jepto- davtyhd frogs, Mad, 97, 34-45, ZW), Ro Ge U9880) Results of the Agglitatd EXpeditians no 107, broys and Gaards from the Huo Ponuisilo, Papua New Gritoes Ail da Mis, nat list, LOS(5), 490-434, (983) Two ny hylid tows trany Papow New Hine anu) a discussion af The Nverariisreny: Aaya spewies wimpy, di dite, Novildtes 2754, 12h brops? PViking O'NaTh, HYBRIDISATION OF LITORIA CHLORIS AND L. XANTHOMERA. (ANURA: HYLIDAE) BY GARY R. BEARDSELL Summary Hransactions af the Rewil Society aj S$. Aust, (1989), Disa) 221-224. BRIEF COMMUNICATION HYBRIDISATION OF LITORIA CHLORIS AND L. NANTHOMERA. (ANURA:HYLIDAB) A study of intraspecific variation among the three disjunct populations af Litoria chloris (Boulenger) of Eastern Australia showed that the northern population differed in morphology from the olher two populations!. Iscenzyte electrophoresis demonstrated extensive Benetic divergence between the northern and southern populations resulting if the recognition of the Horthern population as @ tlistinet species, L. xanihomera’. Farly development and larval nerphology of /. ehloris have been described! and (he larvae of 4, xanthomera are identical to those of L. vhloris’. Here I (est genetic compatibility through a fertile cross between a captive male L. chloris and a female L. yonthomera. Two adult male L. ch/oris were acquired from Finch Hatton Gorge, Mackay, Queensland in January 1984, and one adult female of L. xenrhomiera Irom Crystal Falls near Cairns, Old on 10.xi1.1987, The frogs were housed in an indoor terrarium. Amplexus occurred on 28.i1.1988. Embryos were reared in an aquarium with rainwater to a depth of 24cm. Water temperature was 18-29°C, Larvae were ed pond algae, commercial fish food and hoiled lettuce. When they approached metamorphosis they were transferred to a terrarium, The frogléts were fed on house flies and flour moths. Measurements were made using yernier calipers reading (0.0.1 nim and an ocular micrometer to 0.1 mm. The larval mouth dise was drawn under a stereoscopic microscope while the larval and froglet drawings were inked directly Onto enlarged photographs, that were later treated with polassium jodide and a sodium thiosulphate fixing bath to semove the photographic image and iodine stain respectively’. Developmental stages follow Gosner', Calling, commenced on. 17.xi.1987 and continued on Most nights through to mid-January when the female Z. xanthamera became gravid. Amplexus pceurred on Pip. |. Developmental stages of a fertile cross between Litoria chlorts and L, xanihomera, A, Stage 29; B, Stage 34; C, Stage 42. Seale bar = | mim. 28.11.1988 at O800 hr, and the hale continued to call for one minute while the female produced a soft call, The amplectant pair was then placed into a smaller breeding aquarium. Oviposition occurred from 0630 ro 0700 hr of 29,1), 1988. Approximately 600 eggs were laid as a fat, jelly-like mass on the surface of the water; mean ovidiameter = 1.5 mm, capsule diameter = 4.4 mim, (n = 6). The animal pole was dark brown, the yegetal white. The fertile eggs began to sink and adhere to the botiom and sides of the aquarium in the first hour, Dimensions of the embryos and larvae are shown in Table {. Hatching began 1,iii.1988 al stages 20 & 21, The embryos were dark to mid brown; length! 6.5 mm, Each embryo had three pairs of external gills with numerous branches, On 3 iii. L988 the larvae were at stage 23, The operculum developed and melanophores appeared along the mid- dorsal ling, over the tail musculature and as circular patches over the dorsal surface of the abdomen. The larvae were pale brown and the coils of the intestines visible through (he abdominal wall. The larvae reached stage 25 at 1700 hy (4.ii1.1988). AL stage 29 (Fig, 1A) the larvae had a total length of 15.5 mm and a body length of 5 mm, Most larvae developed @ lateral curvature of the (ail of varying degrees of severity between stages 28-30 in weeks 4 & 5. Artis point there was marked variation in growth: some larvac artained a total length 2x that of others, As the larvae approached stages 34-38 between weeks § and 7 the severity of the fail bend j/ic¢reased in(o an S-shape in many individuals (Fig. 2), At stage 34 body colour was light brown wilh a transparent area over the abdominal wall, Pigmentation was apparent on the dorsal tail (in and tail musculature (Fig. 18). On day 49 (he largest larva mMeasurect 52 jam in stage 40 (Vig. tC), The mouth dise (Fig. 3) of a larva in stage 34 has the typical hylid. formuta of hig. 2, Abnocmal tail bend in a hybrid larva al Stage 40 Total length 52 nom. 222 TABLE |, Dimensions of developmental stages of a cross between Litoria chloris and L. xanthomera. Means (mm) with ranges in parentheses AGE STAGE SAMPLE EMBRYO DIAMETER (Hours) CAPSULE DIAMETER BODY LENGTH TOTAL LENGTH 0 2 6 15 (1.5-1.6) 15 3 — — 2.75 4 = = 3.75 8 — — 6 10 —~ — 10.75 Wel — = 11.75 ul = = 12.5 12 = = 15 13 — — (Days) I 18 5 44 (4.2-4.8) 2 21 6 6.5 (5.9-6.9) 3 23 5 1 (7.0-7.3) 4 25 5 19 (7.7-8.0) 8 26 5 — 37 33 4 = 38 38 4 — 45 40 3 = 51 46 4 — In many larvae extra papillae with labial teeth appeared between the upper rows of labial teeth (Fig. 3). Four froglets completed metamorphosis 20.iv.1988 and had snout-vent (S-V) lengths of 14.4, 14.6, 14.8 and 15.1 mm. The remaining 52 surviving larvae completed metamorphosis by 26.iv.1988. The majority of juveniles metamorphosed with pelvic girdle abnormalities (Fig. 4A) to varying degrees of severity. Approximately on third of the juveniles displayed no external sign of skeletal abnormalities (Fig. 4B). The colour of juveniles was mid- - 43 10.3 (4.1-4.4) (9.8-10.6) - 12.1 38.2 (10.2-12.3) (36.7-39.1) = 134 39.3 (13.1-13.7) (38.9-39.9) = 22.4 52.1 (21.6-22.7) (50.1-52.8) = 147 52.1 (14.4-15.1) a brown with a very fine, diffuse speckling of white disappearing after one week. The embryo and larval lifespan extended over a period of 51-56 days at 18-29°C. The froglets were measured at monthly intervals after completion of metamorphosis (Fig. 5). Within a few days they developed a white supralabial stripe disrupted in some individuals and complete in others, The mid-brown colour changed to a light mustard- brown within two weeks of metamorphosis. By 5.v.1988 rare a yn 11 cal yay. ‘diayodinyul mire marvacun tormenta Fig. 3. Hybrid larval mouth dise at Stage 34. Scale bar = | mm. Fig. 4. A, Pelvic girdle abnormalities in a three month old hybrid froglet (S-V 23 mm); B, Normal three month old hybrid froglet (S-V 27.2 mm). 223 the supralabial stripe developed a green tinge while green pigmentation developed on the supraobital skin. The thighs developed the orange of L. xanthomera by May-mid June, the supralabial stripe became indistinct concomitantly, with a green tinge over the brown on the body parts that are green in the adult life. After eight months all 25 surviving froglets continue to grow regardless of the presence or absence of pelvic girdle abnormalities. On 6,xii.1988, the largest fraglet had a snout-vent length of 37 mm and had developed nuptial pads. The life history pattern of the hybrid conforms to the pattern of Australian hylids", The larval life span of 51-56 days is in agreement with the 41 days recorded for L. chloris' and 58 days for L. infrafrenuta’ considered usual among tropical species adapted to life in ephemeral waters”. More importantly, development of a lateral curvature of the tail in larvae at four weeks manifesting as pelvic girdle abnormalities in newly metamorphosed froglets supports the premise of genetic differentiation attained by L. xunthomera and ils recognition as a separate taxon from L. chloris*. The development of some apparently normal froglets suggests that these species are closely related. 35 t Bt] = | ‘ n-s ? 35 n=5 = ns5 5 20 z n=6 ~ 2 n-5 184t n=10 10 0 1 2 a 4 5 6 Time from Metamorphosis (months) Fig. 5. Growth rate of hybrid offspring from a cross between Litoria chloris and L. xanthomera. (Mean and Tange; n = sample size). I thank Mr M. J. Tyler and Dr M. Davies who critically read the manuscript. Special thanks are due to Richard Perkins for his expert skills in photography and introducing me to the technique of bleach-outs and the science department of Ingle Farm High School for the use of various equipment and their general interest in the frogs, Thanks are due to my brother Kym for typing the manuscript. 224 'Davies, M & McDonald, K. R. (1979) Aust. Zool. 20, 347-359. z & Adams, M. (1986) Proc. R. Soc. Vict. 98(2), 63-71. ‘Watson, G. F. & Martin, A. A. (1979) Aust. Zool. 20, 259-268. ‘Paterson (1979) “Book of the Darkroom”. (Paterson Products Ltd. Sawtells of Sherborne Ltd, England.) °Gosner, K. L. (1960) Herpetologica 16, 183-190. °Martin, A. A. & Watson, G. F. (1971) Copeia (1971) 78-79. Banks, C. B.; Birkett, J. R.; Dunn, R. W. & Martin, A. A. (1983) Trans. R. Soc. S. Aust. 107(4), 197-200. ‘Duellman, W. E. & Trueb, L., (1986) Larvae, Chapter 6 in “Biology of Amphibians”. (McGraw-Hill Book Co., New York.) GARY R. BEARDSELL, Ingle Farm High School, Belalie Road, Ingle Farm, S. Aust. 5098. ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED Patron: HIS EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA LIEUTENANT-GENERAL SIR DONALD DUNSITAN, K.B.E., C.B. OFFICERS FOR 1989-90 President: G. M. E. MAYO, B.Ag.Sc., Ph.D., D.Univ. Vice-Presidents: Cc. J. M. GLOVER, J.P., M.Sc. N. A. LOCKET, M.A., B.M., B.Ch., Ph.D., D.O. Secretary: Treasurer: G. A. CROOK, B.Sc. R. H. WATERS, J.P., F.A.I.M. Editor: Assistant Editor: M. DAVIES, M.Sc., Ph.D. V. A. GOSTIN, M.Sc., Ph.D. Librarian: Programme Secretary: M. ANTHONY, A.L.A.A. R. W. INNS, B.Sc., Ph.D. Minute Secretary: Membership Secretary: D. FINLAYSON P. M. THOMAS, M.Sc. Members of Council: R. V. SOUTHCOTT, M.D., D.Sc., DIM&H C. B. WELLS, E.D., M.Ag.Sc. B. J. COOPER, B.Sc., Ph.D. R. H. FISHER, E.D., A.U.A. D. J. WILLIAMS Printed by Graphic Services Pty Ltd, 516-518 Grand Junction Road, Northfield, S.A. 5085