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VOL. 108, 
12 JUNE, 


PARTS 1 & 2 
1984 


Transactions of the 


Royal Society of South 


Australia 


Contents 


Incorporated 


Hutchings, P. A. & Turvey, 8. P. The Spionidae of South Australia (Annelida: 
Polychaeta) - - - - - - - - - 
Austin, A. D. Species of Ceratobaeus Ashmead (Hymenoptera: Scelionidae) 
from south-eastern Australia - - - - - - 
Lansbury, I. Some Nepomorpha (Corixidae, Notonectidae and Nepidae) 
(Hemiptera-Heteroptera) of north-west Australia - - - 
Geddes, M. C. & Butler, A. J. Physiochemical and biological studies on the 
Coorong Lagoons, South Australia, and the effect of salinity 
on the distribution of the macrobenthos - - - - 
Alley, N. F, & Bourman, R. P. Sedimentology and origin of Late Palaeozoic 
glacigene deposits at Cape Jervis, South Australia - ~ - 
Joseph, L & Hope, R. Aspects of genetic relationships and variation in parrots 
of the Crimson Rosella Platycerus elegans complex (Aves: 
Psittacidae) - - - - - - - - - 
May, R. I. & Bourman, R. P. Coastal landslumping in Pleistocene sediments at 
Sellicks Beach, South Australia - : = = 3 - 


Twidale, C. R. The enigma of the Tindal Plain, Northern Territory  - - 


Suter, P. J. A redescription of the genus Tasmanocoenis Lestage (Ephe- 
meroptera: Caenidae) from Australia - - - - 2 


King, M. A new species of Gehyra (Reptilia: Gekkonidae) from 
northern Western Australia - - - zi 2 z , 


Dulhunty, J. A., Flannery, T. F. & Mahoney, J. A.Fossil marsupial remains at 
the southeastern corner of Lake Eyre North, South Australia 


Tyler, M. J. & Davies, M.Uperoleia Gray (Anura: Leptodactylidae) in New 
Guinea - - = < s - s & x : 


Brief Communications; 


Bone, Y. Wardang Island—A refuge for Marginopora vertebralis? - 
Greenslade, P, The identity of Orchezelandia rubra (Collembola: Entomo- 
bryidae)  - - = 4 " = = ¢ 2s 2 


Fisher, R. H. Life history of the Sciron Skipper Trapezites sciron eremicola 
Burns (Lepidoptera: Hesperiidae) - - - - - 


Glover, C. J. M.Further additions to the marine fish fauna of South Australia 


Beveridge, I. A re-examination of Taenia mastersti Krefft, 1871 and Taenia 
fimbriata Krefft, 1871 - - = 2 ~ 2 a. i 


Schwaner, T. D.The identity of red-bellied black snakes on Kangaroo Island 


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STATE LIBRARY BUILDING, NORTH TERRACE, ADELAIDE, S.A. 5000 


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1I3Z 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 108, PART 1 


TRANSACTIONS OF THE 


ROYAL SOCIETY OF SOUTH AUSTRALIA INC. 


CONTENTS, VOL. 108, 1984 


PARTS | & 2, 12 June 


Hutchings, P. A. & Turvey, S. P. The Spionidae of South Australia (Annelida: 


Polychaeta - - - - - - - - - - : 
Austin, A. D. Species of Ceratobaeus Ashmead (Hymenoptera: Scelionidae) from 
south-eastern Australia - - - - - - - - 7 
Lansbury, I. Some Nepomorpha (Corixidae, Notonectidae and Nepidae) 


(Hemiptera-Heteroptera) of north-west Australia - - - 
Geddes, M. C. & Butler, A. J. Physiochemical and biological studies on the Coorong 
Lagoons, South Australia, and the effect of salinity on the distri- 
bution of the macrobenthos - - - - - - “ = 
Alley, N. EF. & Bourman, R. P. Sedimentology and origin of Late Palaeozoic glacigene 
deposits at Cape Jervis, South Australia - - - - - - 
Joseph, L. & Hope, R. Aspects of genetic relationships and variation in parrots of the 
Crimson Rosella Platycerus elegans comples (Aves: Psittacidae) 
May, R. I. & Bourman, R. P. Coastal landslumping in Pleistocene sediments at Sellicks 
Beach, South Australia - - - - - - - > 


Twidale, C. R. The enigma of the Tindal Plain, Northern Territory - m 5 + 
Suter, P. J. A redescription of the genus 7asmanocoenis Lestage (Ephemeroptera: 
Caenidae) from Australia - - - = ys a 3 e 


King, M. A new species of Gehyra (Reptilia: Gekkonidae) from northern 
Western Australia - - - - - - - - - 


Dulhunty, J. A., Flannery, T. F. & Mahoney, J. A. Fossil marsupial remains at the 
southeastern corner of Lake Eyre North, South Australia - - - 


Tyler, M. J. & Davies, M. Upero/eia Gray (Anura: Leptodactylidae) in New Guinea - - 


Brief Communications: 


Bone, Y. Wardang Island—A refuge for Maerginopora vertebralis?- - - 
Greenslade, P. The identity of Orchezelandia rubra (Collembola: Entomobryidae) - 
Fisher, R. H. Life history of the Sciron Skipper 7rapezites sciron eremicola Burns 


(Lepidoptera: Hesperiidae) - - - - : J 4 = 
Glover, C. J. M. Further additions to the marine fish fauna of South Australia - - 


Beveridge, I. A re-examination of 7aenia mastersii Krefft, 1871 and Taenia fim- 
briata Krefft, 1871 - - - - - - - - - 
Schwaner, T. D. The identity of red-bellied black snakes on Kangaroo Island - - 


63 


77 


85 
95 


105 


113 


119 


123 


127 


129 


131 
133 


136 
137 


PARTS 3 & 4, 13 December 


Kangas, M. I. & Geddes, M. C. The effects of salinity on the distribution of amphipods 
in the Coorong, South Australia, in relation to their salinity 
tolerance - - - - - - - - - - 

Bayly, I. A. E. A new species of Calamoecia (Copepoda: Calanoida) from South 
Australia, and comments on three congeners - - - 

Mahony, M., Tyler, M. J. & Davies, M. A new species of the genus Rheobatrachus 
(Anura: Leptodactylidae) from Queensland - - - - - 


Callen, R. A. Quaternary climatic cycles, Lake Millyera region, southern Strzelecki 
Desert - - - - - - - - - - - 
Twidale, C. R. & Campbell, E. M. Murphy Haystacks, Eyre Peninsula, South 
Australia - - - - - - - - - - 2 
Beveridge, I. Dasyurotaenia robusta Beddard, 1912, and D dasyuri sp. nov., from 


carnivorous Australian marsupials - - - - rs 
Beveridge, I., Speare, R. & Johnson, P. M. New records of Globocephaloidinae 
(Nematoda: Trichostrongyloidea) from peop pyeas in north 
Queensland - - - - - - - 
Greenslade, P. & Wise, K. A. J. Additions to the collembolan fauna of the Antarctic 
Jago, J. B., Daily, B. D., Von Der Borch, C. C., Cernoyskis, A., Saunders, N. First 
reported trilobites from the lower Cambrian Normanville Group, 
Fleurieu Peninsula, South Australia - - - s . - 


Brief Communications: 
Lange, R. T. Leaf marking in rangeland grazing studies - - - = 
Schwaner, T. D. & Miller, B. Range extensions of reptiles in South Australia - - 
Schwaner, T. D. & Miller, B. Reptiles new to the fauna of South Australia - - 
Fisher, R. H. Life history of the Narrow-Winged Pearl White Elodina 
padusa (Hewitson) (Lepidoptera: Papilionoidea) - - - - 
Delean, S. & Harvey, C. Notes on the reproduction of Nephrurus deleani (Reptilia: 
Gekkonidae) - - - - - - - - - - 
Preiss, W. V. Correlation of the uppermost late Precambrian succession across the 
Torrens Hinge Zone in the Port Aarmasta region of South Australia: 
a discussion = “ s b. : 
Plummer, P. 8S. Correlation of the uppermost late Precambrian succession across the 
Torrens Hinge Zone in the Port Augusta region of South Australia: 
a reply - “ - - - is “ “ # . _ 
McDonald, K. R. & Tyler, M. J. Evidence of gastric brooding in the Australian 
leptodactylid frog Rheobatrachus vitellinus. - - 
Belperio, A. P. & Murray-Wallace, C. V. Comment: Wardang Island—a nite for 
Marginopora vertebralis?  - - - - - - - 


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203 


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215 
217 


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221 


223 


226 


227 


THE SPIONIDAE OF SOUTH AUSTRALIA (ANNELIDA: POLYCHAETA) 
BY P. A. HUTCHINGS & P. S. TURVEY 


Summary 


Five new species of Spionidae, Aquilaspio pyramidalis, Scolelepis (S.) bifida, Scolelepis (N.) 
edmondsi, Spio tridentata and Boccardia fleckera are described. Descriptions are given of fourteen 
previously described species of Spionidae occurring intertidally in South Australia, together with 
their known geographical range. Two species of Pseudopolydora are described only to genus. A 
Key to all species is provided. 


THE SPIONIDAE OF SOUTH AUSTRALIA (ANNELIDA: POLYCHAETA) 


by P. A. AHurcimcs & §. PL Turvey™ 


‘Summary 


Hurenios, P. A. & Turvey, S. P. (198d) The Spionidue of South Australia (Annelida: 
Polychaeta). Trans, R. Soc, 8. Aust, WR(1), 1-20, 12 June, 1984, 


Five new species of Spionidac, Agdilaspia pyramidalis, Scolelepis (S.) hifida, Seroalelepis 
(N,) etnionds?, Spice tridentata and Boccaritia feekrra are described. Descriptions are given ol 
fourteen previously described species of Spionidae occurring intertidally in South Adstratia, 
together with their known geographical range. Two species of Psendopolydora ure described 


only to genus. A Key to all spegies is provided, 


Key Worbs; Taxonomy, Polychaeta, Spionidae, South Australia, Key, 


Tnatroduction 


Ta 1979 one of us (PH) made extensive 
collections of South Australian polychaetes, 
concentrating on estuarine and intertidal areas. 
Althongh Blake & Kudenoy (1978) have 
Tecently undertaken’ a major review of the 
spionids from SE Australia, we have found an 
additional five vnew species. This probably 
indicates the diversity of the spionids in 
southern atid south eastern Australia, and we 
suspect that many more species remain to be 
described, 

In addition to describing five new species, 
we have included a shori diagnostic account of 
cach genus and of previously described species 
occurring in South Australia. Species identi- 
fied from the key should he checked carefully 
aguinst the descriptions, in particular the scti- 
gers on Which setal changes occur and the 
deinil-of the setal structure, This is particularly 
important for non-South Australian material. 
where other references such as Blake & 
Kudenov (1978) and Hartmann-Sehrdder 
(1979, 1980, 1981) should be consulted. 


Materials and Methods 


Station data have been coded and tabulated 
(Table |) and the codes used in the Material 
examined section of each species description. 
Regsstration numbers of Australian Muscun 
material has been abbreviated to W. plus num- 
her. Paratypes have been deposited wherever 
possible at the Allan Hancock Foundation, 
Loy Angeles (ARLE), British Museum (Natural 
History), London (BMNH) and The National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. (USNM). Other 
abbreviations usexl are us follows: HZM, Zoo- 


“The Australian Muscum, G8 College Street, 
Sydney, N.S.W, 2000, Australia. 


logisches Institut und Zoologisches Museum 
der Universitit, Hamburg; KM, Zoologisk 
Museum, Copenhagen; NMY. National 
Museum of Victoria, Melbourne; SAM, South 
Australian Museum, Adelaide. 

The Australian distribution of each species 
has been given using Day & Hutchings (1979) 
checklist and Blake & Kudenoy (1978). The 
localities are arranged geographically from 
west to cast, and along the east coast of Aus- 
tralia from south to north. Additional locality 
data from subsequent publications wre marked 
with an asterisk. 

In general we have only cited major Aus- 
tralian references. Full synonymies are given 
by Blake & Kudenoy (1978) and Day & 
Hutchings (1979). 

We have followed Foster (1971) in accept- 
ing the various genera within the Prianaxpio 
complex which she recognised based mainly 
on the type of branchiae present, We helieve 
this is a useful division. 


Rey ra the Santh Australian Spientdae 
(after Blake & Kudenov, 1978) 


1, Setiger 5 modified, with specialised 
setae P 2 
Setiger 5S not modified. withoul spe 
cialised setae 11 

2.¢1) Branchige beetining on setiger 2 


Wiig weg roa! Boccard iat 3 
Branchiae beginning on setigers 6.12, 5 


3.(2) Proastomium enthe ..,.. B. prabaseldea 
Prostomium deeply incised 4 
4.43) Neuroselul hooded bhaoks tram 


scliger 7: setiger 5 with simple 
talent spines and spines with con- 
caye cup containing bluntly conical 


toouly B. élilleityiw 
Neurosetul hooded hooks from 
setiver Vd; svtiper 4 with curved 


spines and brush 
wt. fleekera wasp, 


falcute smooth 
lipped setiec: 


rm 


5. (2) 


6.5) 


7, (6) 


8. (6) 


9, (5) 


10, (9) 


11, (1) 


12, (11) 


13. (12) 


I4, (13) 


15. (11) 


A, HUTCHINGS & 8. P. TURVEY 


Setiger 3 slightly modified, with 
prominent parupodia, major spines 
of 2 types arranged in V or J shaped 
row .,.- . Pseudapalvdora 6 
Setiger 5 preatly” “modified with 
reduced parapodia, and 1 type of 
spine arranged in curved row 

Palydora 9 
Prastoniart eiitiré 5 erm 7 
Prostomium deeply incised 8 


Neurosetal hooded hooks, bidentate 
from setiger 3 P. paucihranelitata 
Neurosetal hooded hooks, multi- 
dentate from setiger 7 ; 

Pseqdopely dina sp. 2 


Modified selae on setiger 5S pen- 
noned and simple spines |. P. antenneata 
Modified setae on setiger 5 falcate 
pennoned spines  Pyewdopolydearq sp. | 
Hooded hooks withont constriction 
on shaft; setiger 5 with major spines 
with subterminal boss, compiunion 
setae bilimbate f. PY saciulis 
Hooded hooks with constrictiun on 
shaft: seliger 5 with setae otherwise 
taint ae - 10 
Prostomium weakly incised; setiger 
5 with Falcate spines with large sub- 
terminal flange; companion setae 
bilimbate . PB. hoptera 
Prosiomium melsed, with 2 widely 
separated lobes; setiger 5 with 
curved spines with prominent subter- 
minal tooth and feathered com- 
panion setae BP tani 


Prostomium distally pointed with or 
without subdistal lateral horns... {2 
Prosiomium not distally pointed, 
with or without distal lateral or 
frontal horns, broadly rounded o1 
incised on anterior margin, i5 
Branchine fused to dorsal lamellae 
at lenst basally. contiouing to end of 
body - ed Sealelepis V4 
Branchiae completely free from 
dorsal lamellae, present on variuble 
number of anterior setigers, absent 


posteriorly _ Aatides axveepliali 
Noiosetae all capillaries, at least 
until setiger 93 _ - i4 


Notosetue initially capillaries, biden- 
tate hooded hooks from setiger 38 
S46 — OS. Caracal 
Neurosetal hooded hooks bidentate 
from sctiger 36 S. biftta nap. 
Neurosety! hooded hooks widentate 
from setiger 43 S. edronedst asp. 
Branchise concentrated in 1-22 
anterior setigers, absent posteriorly 

16 


Stewart Istaned, 


Byinchiac present over most of hody 
Tenathe 5 A ee ete 
Branchiae ull cirriform, 10 pairs 
nt Minuspia cirrifere 
Branchiae not all cirriform, 3-4 
pairs 17 
Branchiau all pinnate Aquilaspio 18 
Branchiae pinnate ond cirriform 
Prianaspio niiltlerisiata 
Three paits of branchine 
Aquilaxpia, anehlandive 
Four pairs of branche WwW 
Rounded neuropodial lamella on 
selizer 1. | Aquilaspio multipinnalave 
Neurvpodium of setiger 1 inflated, 
pyramidal in shape Aquilaspioa 
pyramidalix np. 
Branchine beginning on seliger 1 
anteriorly fused to  notopodiul 
Jamellic. neuropodial hooks from 
seliger 28 Spia tridentata nap. 
Branchiac beginning on seliger 2 
stout cirriform completely free from 
notopodin; seuropodial hooks from 
seliger 9 Microspio erentalata 


20 
16. (15) 


17. (16) 
18. (17) 


19. (18) 


20, (15) 


Scolelepis (Blainyille (emended Pettibone) ) 
Prostomium pointed anteriorly and posteriorly, 
Peristomium forming hood about prostamium. 
Branchiae from setiger 2 to near end of body, 
more or Jess completely fused to notopodial 
lamellae at least anteriorly. Neutopadial lamel- 
ise uni- or bilobed. Neuropodial hooks present 
in far posterior or absent. Hooks hooded, 
entire, bi- to quadridentate, 


Scolelepis (Scolclepis) bifida n.sp. 
FIG, Ja-g. 
Sculelepis famellicineta Blake & Kudenov, 1978: 
176-178, fig, la-k Cin part). 
TMatorype: S.A, 09C (W.19283). 
Paratypes: 08C, 2 (W.19284), 1 (USNM 074899), 
1 (BMNH ZB 1982.76), 32C, 8 (W.192R5), 1 
(AHF POLY 1383), 
Other ntaterial examined: Holotype of Seelelopis 
lamellicivera Blake & Kudenoy (NMY Ci102) and 
Paraiype (NMV 62990) Westernport Bay. Vie, 
(SAM &1577) Ellistan, SuA, Holotype of Pxeido 
nevine antipede Augener (IRM) Pegasus Buy. 
New Zealand, 
Description; Holotype posteriorly incomplete, 
partially broken between setigers 63-64; 59 
mm long and 4.2 mm wide at about setiger 
40, for 4 total of 93 setigers, Paratype material 
all incomplete posteriorly, with following 
ranges of dimensions: 40 mm long, 2,5 mm 
wide for 87 svtivers, 25 mm fong, 2:0 mm 
wide for 70 setigers; 18 mm long, 1.5 mom 


SPIONIDAE OF SOUTH AUSTRALIA 


TABLE 1, Collectian data 


Locality, collector and date 


Latitude/ 
longitude 


(Deg. Min.) 


Habitat 


Port Augusta. Hutchings, 14,iii.1979 


Streaky Bay, near carayan park. 
Hutchings, 13.iij,1979 


Streaky Bay, litle island on 
Ouler margin of inner bay. 
Hutchings, 13.11.1979 


Speeds Point, Streaky Buy. 
Hutchinus, 14.11.1979 

Port Kenny, Veniis Ray, 
Hutchioys, 12.17.1979 
Venus Bay, village. 
Hutchings, 12.11.1979 


Elliston, reef al southern end of 
town. Hutchings, 12,iii,1979 


Elliston, reef just past post-office. 
Hutchings, 12.i1i1,1979 

Elliston, jetty. 

Hutchings, 12.17.1979 


Kellidie Bay, Hutchings, 11.11.1979 


Porter Bay, Port Lincoln, near boat 


ramp. Hutchings, 10.11.1979 
Torrens Island, Adelaide Power 
Station. Hutchings, 7.iii,1979 


Flinders Cairn, Port Lincoln. 
Hutchings, 10.i1.1979 


Sleaford Mere, Hutchings, 10.iii.1979 


Sleaford Bay. Hutchings, 10.11.1979 


Sellicks. Bench, reef to north. 
Hutchings, 16,71,1979 


Rapid Bay, jetly between Normanyville, 
and Sceond Valley. Hutchings, 811.1979 


Vicior Harbor, just behind bluff. 
Hutchings, 16.17.1979 


Emu Bay, Kangaroo Island, adjacent 


io old jetty. Hutchings. 1.iii.1979 


Stokes Buy, Kangaroo Island. 
Hutchings & Buller. 5.iii.1979 
Stokes Buy, Kanguroo Island, 
Handley, 4.i11,1976 

Bay of Shoals, Kangaroo Island. 
Hutchings & Edmonds, 1.i1i1,1979 


32-30/137-46 
32~48/134-13 


32~48/ 134-13 


32-48/ 134-13 
33-10/134-41 
33-14/134-40) 


33-39/154-53 


33-39/134-53 
33-39/ 134-53 


34-36/135-29 
34-44 / 135-53 
3447/138-32 


34-49/135-47 
34-50/ 135-45 
34-54/ 135-47 


35-20 / 138-27 


35-37/ 137-12 
35-37/137-12 
35-36/137-37 


Sand on mudflats tn front of 
mangroves. under bridge 
Mussell clumps at mid tide 
level on mud flats 

Mud flats, Posidonia 

Mud sievings, Pesidonia 
Pasidonia and Zostera sievings 
Fauna associated with Zostera 
Sands sicvings 

Sand sievinys, Posidonia 
Under boulders 

Posidonia and Zostera sievings 
Sand sievings among Posidonin 
and Zastera 

Algal washings 

Zostera sievings 

Mussel chumps at mid-tide level 


Algal mat ou reef, south of 

village 

Sand sievings 

Faunu on jetty piles 

putes rocks on low. tide reef 
al 

Algae from low tide reef flat 

Sand sieyings at low tide 

Algal washings 


Amongsl Galeolaria on jetty piles 
Nearby rocks, encrusting algae 
Sand sievings 

Mussel clumps at mid-tide level 
Zostera and sand sievings 
Zastera sievings 


Mudilats in front. of thermal 
effluent (up to 42°C) 

Mud ilats in front of mangroves 
Mud flats in front of mangroves 
with pulchy Zosrera 

Sand at low tide level 

Mussell clumps at mid-tide level 
Mud, salinity 20%, 

Algae on ocean side of bay 
Algal washings 

Sievings in Amphibolis 

Sand sievings 

Sand sicvings near 4renicola 
Fauna. attached to jetty piles 


Crevice fuuna 

Algal washings 
Coralline algae washings 
Crevice fauna 

Algal washings 

Under rocks beside jetty 
Posidonia sievines 
Algal Washings 

Sund sievines 

Algae at low-tide Jevel 
Under rocks ut low tide level 
Zostera sievings 


4 P. 4. HUTCHINGS & §, P, TURVEY 


3 km SW of Cape Rouge. 
Handley, 7.iii.1978 


Hay of Shoals, low-tide. Hoese, iii.1979 
Snellings Beach, mouth of Middle 
River, Kangaroo Island. 

Hutchings & Butler, 5.iii. 1979 
Penneshaw jelly. Kangaroo Island. 
Mandley, 9,if.1978 

Western River Cove, Kangaroo 
Island, Handley, 3.111.1978 
Redbanks, Nepean River, Kangaroo 
Island. Lock and Yoo, 4.iiL 1978 
Muston Point, American River, 
Kangaroo Island, old wharf, 
Hutehings, 2.i.1979 


American River, Kangaroo Island, 

top of river just below turn-off 

to Pentington Bay. Hutchings, 3.1.1978 
Pelican Lagoon, south side, 

Kangaroo Island. Handley, 8.ti.1978 


Cape du Couedic, Kangaroo Island. 
Hutchings & Butler, 4.11.1979 


Harriet River estuary, Vivonne Bay, 
Kangaroo Island. 

Yoo and Handley, 2.i11.1978 
Hanson Bay, Kangaroo Island, 
Hutchings & Butler, 4.11i,1978 


Cape Domby, near obelisk. 
Yoo, 72.11.1978 


Cape Northumberland, on west side. 
Yoo, Loch and Handley, 27,iii,1978 


35-42, 137-06 


35-43/ 137-56 
35-43/136-56 
35-44/ 137-43 
35-47/137-46 


35-47/137-46 


35-40/137-45 


36-03/146—-41 


35-S8,/137-09 


36-02/ 136-51 


37-10/139-44 


38-04/140-40 


Sand flats verging into 
Posidoniaiand Tlormosira 


Pasidenia, Zostera, Wid and sand = 22C 


Algal holdfasts and crevice fauna = 23A 
Sand sievings 236 
In sponges on boom piles at 244A 
5 m, and under rocks 

Sheltered rock pool, under 25A 
rocks and algae 

Sheltered shallow hay at low level 26 
Clumps of sponge at 5m in fast 27A 
flowing channel with many Pinna 

Sund) sponges, and sandy conglo. 27B 
merate rack at 5 m in fast-flowing 
channel 

Zuslera sievings 27C 
Surface detritus and algae 28A 
Under rocks and Hormoasira in 290A 
front of salt marsh, at myld-tide 

level 

Exposed beach, algal holdfasis JOA 
Exposed beach, coralline algac 30B 
and algal holdfasts 

Exposed beach, coralline algac 30C 
washings 

Exposed reef, algal holdfasts 30n 
Exposed reef, coralline algae 30K 
Sievings at loyw-tide level 31A 
Algal holdfasts on reef flat FIA 
Closed mouth of South West 32B 
River,, salinity 30%. 

Exposed beach, sand sievings 320 
Algae from pool on exposed rock = 33.4 
platform, 

Sievings in low Zostera patches 334 
at low-tide 

Sheltered pools behind exposed 344 


rock platform at low tide 


—_— ee —.-— O OOD 


wide for 70 setigers; 15 mm long, 2.0 mm 
wide for 70 setigers; 10 mm long, 1.5 mm 
wide for 37 setigers; 6 tnm long, 2.0 mm wide 
for 20 setigers; 20 mm Jong, 2.0 mm wide for 
66 setigers; 30 mm long, 2.5 mm wide for 
79 setigers and 45 mm long, 2.0 mm wide for 
110 segments. Body broadly rectangular in 
cross-section, broadest in mid-section, tapering 
anteriorly and posteriorly, Colour pinkish 
brown in aleohol.  Prostormurn bulbous 
anteriorly tapering to acute point; posteriorly 
forming small, high, attached keel-like car- 
uncle extending to middle of setiger 1 (Fig, 
la); two pairs of eyes arranged in oblique row 
on cach side of base of caruncle, obscured 
by peristomial hood; occipital tentacle absent. 
Peristomium forming high lateral hood about 
posterior part of prostomium, becoming lower 


anteriorly; pulps thick, smooth, tapering pro- 
sressively with conspicuous basal palpophore 
or sheath extending to seliger 11 (left)-13 
(right). Setiger 1 reduced, with notopodial 
Jamcllace small, thick, blontly triangular, 
neuropodial lamellae smaller than notopodial, 
rounded, cup-shaped. but noto- and newrosetae 
present. Branchiae from setiger 2, thick, cirri 
form, elongating to reach approximately twice 
initial length by about setiger 10 then decreas- 
ing slightly towards end of fragment, each 
branchial pair connected across dorsum by nat- 
raW ciliated ridge: anterior postsctal notopodial 
lamellac extend dorsally as membranous 
borders along Jateral margins of branchiae, 
becoming separate only at far distal extremity 
(Fig. tb); branchiae only slightly longer than 
lamellae giving combined lamellae-branchiac 


SPIONIDAE OF SOUTH AUSTRALIA $s 


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Fig. 1, Scolelepis (S.) bifida nsp, a. anterior end, 
dorsal yiew, b. anterior view of Sth parapodiuny, 
c, anterior view of 20th parapodium, a. anterior 
view of 40th parapodium. c-f, sabre seta at “25 
and “100 magnilication. g. hooded hook. Scales 
mmm, 


uppearance of being distally bifid, with small, 
flattened points (Fig. 1c), From about setiger 
MH40 free tips of branchixe become more 
elongate, digitiform, initially curving around 
medio-distal margins of lamellae before con, 
ltinuing as free processes (Fig. 1d); lamellac 
becoming broader and more rounded distally, 
strongly folded in all but far anterior setigers. 
Presetal notopodial lamellae low, rounded 
ridges in anterior setigers, becoming incon- 
spicuous posteriorly, while body wall support- 
ing hotosetae simultaneously becoming raised 
ridge, Postsetal neuropodial lamellae anteriorly 
semicircular in outline, becoming hi-lobed by 
setiger 30; ventral neuropodial lobe small, semi- 
circular in profile becoming displaced ven- 
trally behind ventral extremity of neurosetal 
fascicle; dorsal neuropodial lobe rapidly torm: 
ing a low, clongate interamal lamella, over- 
lapping with notupodial lamella after a few 
setigers then becoming progressively separated 
posteriorly, Presetal neuropodial  larnella 
similar te notopodial, Notosetae all capillaries 
at least to setiger 93. Anterior notosetac 
arranged in two broad raws, with those in 
anterior row stout, broad, generally bilimbate; 


those in posterior rows longer. narrower bul 
still stout, each seta unilimbate or appearing 
to be without sheath or wings; both types with 
shafts with distal fine granulatians and trans- 
parent limbate processes having faint oblique 
striations; notosetae reducing to single row al 
about setiger 30 with broad vertical group of 
shorter capillaries situated ventrally and 
farrow horizontal group of long, capillaries 
located dorsally, these two groups becoming 
variably separated by a narrow space which 
may include several very short, fine, 
unsheathed capillaries; capillarics) becoming 
less robust with granulations barely noticeable, 
in far posterior setigers, Neurosetae anteriorly 
similar to notosetae except most ventral capil- 
aries developing into a partially separate 
fascicle of 3-7 sabre setac over first 3—4 setigers 
(Fig. le-f£); sabre setae similar to unilimbate 
capillaries except shorter with shafts coarsely 
granular distally in posterfor setigers; capil- 
laries in posterior meuropodial fascicles 
gtadually replaced by hooded hooks from 
setiger 36; initially with only 2 hooks, then 
hecoming more numerous forming broad tas- 
cicle of 7-10 hooks and 3-5 small bundles 
of capillaries; capillaries located between 
hooks in dorsal part of fascicle; hooded hooks 
worn, bidentate, with shafts greatly thickened 
in basal region after emergence from hody 
wall (Fig. 1g). Pygidium and other posterior 
structures missing. 

The paralype material exhibits some varia- 
tion from the holotype including eyes not 
visible, and palps extending to setiger 11-2]; 
neuropodial lamellae bi-lobed from setiger 
23-34; notopodial lamellae variahly but 
noticeably folded at least posteriorly, fre- 
quently in all setigers, Hooded hooks in neuro- 
podia from setiger 32-36, 5-10 in number. 


Comments; Scolelepis bifida usp, belongs 
to the sub-genus Scelelepis as defined by 
Pettibone (1963). Scolelepis bifida n sp, is 
similar to §. s@idamata (Miller, 1806) and 8. 
blakef Harimann-Schréder, 1980 in thal setiger 
1 has notesetae, postsetal neuropodial lamellac 
are divided posteriorly, and hooded hooks are 
bidentate, Scolelepixs squamata ditters im that 
the postsetal notopodial lamellae cxtend only 
slightly along the branchiae, the dorsal lobes of 
the neuropodial lamellae do not form long, low 
interamal lamellae and the hooded hooks are 
not basally swollen, Sealelepis blakei differs im 
2 similar manner and in addition has a trifid 
prostomium, Two other species af Sevifelepis 


6 fh A. MUTCHINGS & S$. P. TURVEY 


fecently deseribec! (rom Western Australia, §, 
(S.) bulihaiensis Hatimann-Sechréder, 1979 
and S$. 03.) Aidenovi Hartmaon-Shroder, 1981 
can be easily distinguished from §. bifida nusp, 
by the absence of notosetae on setiger | in 
these two species. 

Scolelepis Jameilicinta Blake & Kudenov 
1978 was described from SE Australia, in- 
cluding SAL as having unidentate hooded 
hooks, The types of this species have been re- 
examined aid the honded hooks are not 
tnidentate but haye | large tooth plus 2 
smaller teeth, almost forming a cusp shaped 
arrangement. Also in S. Jamelieineta the post- 
setal ettropndial lobe becomes a separate 
entity Whereas in S. bifida nsp., this lobe 
remains allached. One of the paratypes (SAM 
1577) was collected frony Elliston Jetty, S.A. 
and has hiftd hooded hooks and parapodial 
structures similar fo 8. bifida msp. and is 
referred to this species. Aartmann-Schrader 
(1980) deseribed S. laiellicineta from Onslow, 
WA. und also figures unidentate hooded 
hooks, and may represent an undescribed 
species, The type of Secolelepls antipoda 
(Augener) has been examined, however the 
type consists of numerous small fragments, 
but the anterior fragment clestly differs from 
S. hifida n. sp. in the shape of the prostomium 
and the anterior gall struetare, 

The prostomium of S. antipoda is rounded 
in comparison to 8, bifida nap. in which the 
prostomium is bulbous aad anteriorly tapering 
to un acute point, The anterior branchiae of 
§. cttipada ire cyelindrical with a small dorsal 
terminal lanvellae Whereas in S. défidea insp, the 
branehiae are simple ancl cylindrical, 

Eninology: The specific name bifida refers 
to the bi-lobed nature of the postsetal neuro- 
podial lamella trom middle seligers onwards. 
Australian dixtribittion: S.A. (Elliston), 
Habitat; Sandy substrates. 


Scolelepis (Scolelepis) carunculata Bhike & 
Kudenoy 
Sealelepiy varnnentatra Blake & Kuen 
178 180, fig. 2iy i. 
Mutertal evanineds SAN 


197% 


WOR, + (WT9STRS, UTE, 


2 (W.19308), L6D, 3 A W.19R00), 19 A 
(W,19412) 208, 3 (WYANT), 2aBL 4 
(W.193 10), 


Description, Size range of entire specimens of 
66°75 selivers; 16-20 mm long, 10-14 mm 
wide; posteriorly incomplete specimens up. to 
3.0 mn wide, Prostomium slightly fusiform 
pointed anteriorly gd posteriorly; posterior 


part of prostomium frec of dorsum, forming 
earuncle frequently elevated, extending to 
posterior Margin of setiger 2. Setiger 1 with 
postseial notopodial lamellac and notosetae 
present. Notopodial lamellae fused to 
branchiae except distally in anterior setigers, 
becoming more separated posteriorly. Neuro- 
podial pasisetal lamellae single — lohed 
anteriorly, becoming bilobed at setiger 24-37. 
Anterior notosetac all capillaries, bidentate 
heoled hooks. and capillaries trom seliger 3$— 
56. Anterior freurosetae all capillaries with in- 
consmichous sabre setae from setiger 3, not 
noticeably thicker than typical capillaries; 
hidentate hooded haoks from sctiger 31-46, 
with earlier occurrence in smaller specimens. 
Pygidium with ventral cushion and low 
rounded dorsal lobe with simple low tateral 
lobe on each side, 

Comarwnts: Our material agrees closely with 
the original description of Blake & Kudenov 
(1978). Variations in distribution of hooded 
hooks and bilobed neuropodial Jamellac are 
greater than previously recorded. ‘This is the 
first record of the gpecics from South Aus- 
tralia, 

Australian distribution: W.A. (Safety Bay*). 
S.A. (Venus Bay*, Elliston®, Sellicks Beach*, 
Kangaroo. Isl), Vie, (Port Phillip Bay, Wes- 
ternport Bay), N.S.W. fBelmont Beach), Qld 
(Moreton Bay! 


Habsrat- Mud and sand flats, 


Scolelepis (Nerinides) edmondsi n.sp. 
FIG, 2a-e. 

Hela pe Sauth Austialia, OC 6W.19994), 

Paruvpes: O90, 2 (AHP POLY (484) noc, 2 
(USNM 074900), 09C_. 2 (BMNEL ZB ISR2) 77- 
TH), U9e, 6 (W.19395), 24B 1 CWLIN396), 

Deseriptians Holotype, 25 mim tows, 1 inn 
wide tor 9S selugers. Paratypes range in size 
from J2=l4 mm long, O.8-f mm wile lor 
60-65 setigers. All type material posteriorly 
ivcomplele, Prostomiuin acutely pointed, with 
7 pars small eyes: inner pair eliptical hidden 
hy raised clevated earuncle attached to 
dorsum; catuncle with pronounced dorsal 
owelling) occipital tentacle absent. Pecistomium 
forming Ventral rulfle around prostomiimy, 
palps with swollen palpostyle, extending pos- 
terierly Lo setigers 6-7. Setiver 1 with digit 
form notopodial lobe and small globular neuro- 
podial lobe: note: and ieurosetae present 
(Fig, 2a) Branchiae present from) setiger 
2, utluched to notupodiul lamellae basally, 
with free portion of bronchia same length 


SPIONIDAE OF SOUTH AUSTRALIA t 


ay limelian, branchiae with very pro- 
minent blood vessel running along ‘anterior 
margin: branchniw increasing im size pos- 
teriorly, rapidly becoming much longer than 
the notopodial lamellac. Notopodial lamellae 
clongaling aver sequential anterior sctigers 
(Vig. 2b) forming arrow rectangular lobe, 
with development of interamal cirri (Fig. 2c); 
in middle and posterior seligers notopodial 
lamellae reduced to form clongated triangular 
lobe and by setiger 55 (Pig. 2d), becoming 
hilobed in far posterior setigers. Interamal 
citri from setiger 31, becoming triangular in 
shape and greater in size than neuropadial 
lamellae, continuing on all subsequent seligers, 
Neuropodial lamellac initially semi-circular, 
gradually becoming more clongate; then divid- 
ing by setiger 29 ta form interamal cirrus; 
ventral fobe and interamal cirrus initially equal 
triangular lobes, interamal cirrus subsequently 
hecoming Jarger; in far posterior segments 
neuropodial lobe displaced ventrally but 
remaining undivided. Large intersegmental 
oval glandular creamy white patches present 
between neuropodia and interamal cirri. Well 
developed dorsal ridges present from setiger 
2 to end of fragment, low in height, 
Notosetae all capillaries, with most elongate 
setae from setiger 55; as none of the material 
examined is complete, the apparent lack of 
notopodial hooded hooks cannot be confirmed, 
if they occur it is Jater than setiger 98. Neura- 
setae initially capillaries; tridentate hooded 
hooks [rom setiger 43 mostly replacing capil- 
hiries neurosetae dominated by hooks and 1/2 
capillaries. Hooded hooks tridentate with pair 
of stout denticles surmounting main fang 
(Pig, 2e) 
Conunenty: Seolelepis edniendsit asp, belongs 
to the sub-genus Nerinides according to Petti- 
bone (1963). Pettibone described all the 
species which she placed in the sub genus and 
S. edmondyi nap. can be distinguished fram 
all these species by the presence of notosctac 
on seliger 1 and the commencement of iriden- 
late neurosetal hooks on setiger 36-44, Since 
that revision occurred several additional specics 
have been deseribed from Australia, Scolelepis 
(N.) vexillatas (Hutchings & Rainer, 1979) 
which is characterised by posterior segments 
with a lumellar extension of the branchiae. 
Blake & Kudenov (1978) deseribed §. (N.) 
fawra, 8. (N.) precirriveta and 8. CN.) vie- 
toriensis Tvom $B. Australia, two af these 
lack notosetuc on seliger 1, and all have 


Fig, 2, Scolelepis (N,) edmonidsi asp. a. anterior 
end, dorsal view. b, anterior view of the para- 
podium. ¢. anterior view of 40th parapadium, 
d. posterior view of 60th piarapodium. ¢. neuro- 
podinl hooded hook, Scales: in mm. 


hooks beginning yery much earlier than in S. 
(N.) edmondsi asp. 
Etymalagy: This species is named after Dr 
Stan Edmonds who helped and largely made 
possible the field trip undertaken by one of 
the authors (PAH) during which the material 
forming the basis of this paper was collected. 
Australian distribution: $A. (Elliston, Suel- 
lings Beach). 
Habitat: Encrusting algae and algal holdfasts. 
Aanides Clapartde (after Pettibone) 
Prostomium acutely conical; peristomium 
more or less fused with prostomium, Branchiac 
from setiger 2, confined to anterior region of 
body, not fuscd to dorsal lamellae. Hooded 
hooks bi- or tridentate, in both notoe- and 
neuropodia. Pygidium with anal cirri. 


Type species Nerine oxycephala Sars 
Aonides oxycephala (Sars) 

Nerine oxycephala Sats, 1862: 64, 

Aonides. oxyveephala, Poore ef al., 1975: 30, 
— Ramos, 1976: 11-20, text-fles 1-2 (for 
synonymy). — Bluke & Kudenov, 97k TRo- 
191, 


8 P. A, HUTCHINGS & S. P. TURVEY 


Material examined: S.A, 19B, 1 (W.19314). 
Desey!ption; A single specimen incomplete with 
66 seligers, measuring 11.0 mm long, 0.7 mm 
wide. Prostomium conical, with occipital ten- 
tacle present, caruncle absent. Eyes not visible. 
Seliger 1 with nofo- and neuropodial lamellac 
reduced, noto- and meurosetac present. 
Branchiac stout, cirriform, on setigers 2-18. 
Postsetal nolopodial lamellae dorsally elevated 
and pointed im anterior setigers, becoming 
rounded posteriorly, postsctal nevropodial 
lamellae small, roughly triangular, All) anterior 
setae capillaries; bidentate hooded hooks in 
notopodia from setiger 22-24; in nevropodia 
from seliger 22; posterior neuropodia with 
ventrally reflexed capillaries which gradually 
become stouter posteriorly and resemble sabre 
setae, 


Comments: The nutnber of branchiae and 
setigers on which note- and neuropodial hooded 
hooks appear in our specimen are well within 
the wide ranges given by Ramos (1976) for 
this species, and other characteristics are in 
close ayreement. First recon! from South 
Australia. 


Australian distribution: S.A. (Emu Bay, Kan- 
garno JIsland™), Vic, (Port Phillip Bay), 
NUS-W. (Merimbula, Jervis, Bay’, Port 
Hacking*, Botany Bay*). 

Habhitar- Posxidonia seagrass beds. 


Agnilaspia Foster 


subteiangular: with anterior 
border rounded or sometimes extending 
slightly laterally, continuing postenofly as 
more or less developed posterior keel or car- 
uncle, Peristomium surrounding prostomimum 
as hood, developed to Varying degrees. 
Branchiac, two to four pairs all pinnate, from 
sctiger 2. Anterior setae all capillaries; tri- 
dentate or multidentate hooded hacks present 
in posterior svtigers of neuro- and notopedia. 
Pygidium with anal cirri 


Progtomiuns 


Type species Prionesme sexeculara Augener 


Aquilaspio aucklandica (Augencr) 


Prionespio aucklandica Augener, t924: 69-70, 


text-fig, 24, 1926; 158-154, fig. 1. 


Agniluspia aucklandica. Foster, 1971: 105-106, 
Hutchings and Rainer, 1979: 763- 
Prenospio (Aquilaspie) aucklundica. Blake & 


Kinlenoy, 1978; 221-222, real-fig. 25b-g, 
Material examined: S.A OLA. 2 (W.ESSLB). 120, 
5, (W.ANS1E). 1C. 61 (W982). IAL 1 


(W.19317), YE, 3 (W.19316), 33B, 21 
(W,19319), 

Description: Size range: 63-26 mm long. 0.4- 
0.8 mm wide for S113 setigers; anterior 
fragments of lurger specimens present up te 
1.1 min width. Prostomiun anteriorly rounded 
with minor irregularities; caruncle high, keel- 
like, eXtending to posterior margin of sctiger 
1, Peristomium dorsally fused to setiger I, 
forming low lateral wings about prostomium 
at base of caruncle; pulps thick, ercnulate, 
extending to setiger 9-15. Branchiae 3. pairs, 
on setigers 2-4; cach pair densely pinnate, 
similar in length in Jarge specimens, but subse- 
quent pairs decreasing in Jength in small 
specimens with pinnules becoming sparse and 
fewer, occasionally disappearing by third 
pair. Setiger | with reduced rounded noto- and 
neuropodial Inmeliae; notosetae lacking. Noto- 
podial lamellae becoming larger, clongated 
dorsally pointed and medially curved over 
setipers 2-4, then becoming rounded and de- 
ereasing in size posteriorly. Neuropodial 
lamellae similar throughout in size to noto- 
podials generally rounded in shape except sharp 
triangular ventrally directed projection in 
setiger 2, Anterior setae in both noto- and 
neuropodial all capillaries, sheathed, distally 
granular, becoming finer posteriorly; hooded 
hooks from setiger 25-33 in motopodia, 15-18 
in neuropodia with about 5 tiers of apical 
teeth above main fang, primary hood inflated, 
secondary hood distinct; ventral sabre setae in 
neuropodia from setiger 10-11, each stout, 
sheathed, distally granular, tapering abruptly 
to filamentous tip; smyller individuals with 
sabre selae from setiger 10 and hooded hooks 
fu the notopodia from setiger 25 and in the 
Turopodia from setiger 15, Pygidium with 
long cirrus dorsomedially and 2 stout papillac. 
Comment: Size-dependent variations in 
branchiae and setal patterns have not heen 
noted previously. Blake & Kudenov (1978, p. 
222), Slate that a low dorsal crest is present 
ou setiger 7. This was not indicated by Augener 
(1924) and was not observed on our material, 
although the anterior margins of post-branchial 
setigers were slightly raised ta form Jow dorsal 
midges, This is the first record of this species 
from South Australia. 


Australian distribution: SA, (Port Augusta*, 
Torrens Island, Sellicks Beach", Emu Bay*, 
Cape Domby*), Vie. (Port Philp Bay, 
Westernport Bay), NSW. (Merimbula, 
Hotany Hay, Careel Bay’, Wallis Lake), 


SPIONIDAE OF SOUTH AUSIRALIA ] 


Habitat: Intertidal and sub-tidal sediments in- 
cluding seagrass beds, among coralline algae. 
Aquilaspio muttipinnulata (Blake & Kudenoy) 
hew comb, 
Prionespia CAquilaspio) multipinnulata Blake & 
Kudenoy, 1978) 219-221, text-fig, 24a-f, 
Mujlerial examited: S.A, O4B, 2 (W,19324). O7B 
1 (W.19321). 11A, 6 (W.19329). 12B, 1 CW. 
19326). 13A, 4 (W.19330). LOD, 5 (W.19327). 
19, 1 (W.19323). 21B, L (W.19328), 22A, 2 
(W.19322). 270, 12 (W.19325). Onkaparinga 
Estuary, 1 (W.6071) coll. Shepherd. N.S-W., 
Merimbult (W.11736). identified by Blake & 
Kudenov, 


Description: A -single entire specimen (W. 
19326) measures 54 mm long, 1.6 mm wide 
for 137 setigers; posteriorly incomplete speci- 
mens of 0.9-1.8 mm width. Prostomium 
broadly rounded anteriorly with high, keel-like 
earunele extending to posterior margin of 
sctiger 1; two-three pairs of eyes present, Peri- 
stomlum dorsally fused to setiger 1, together 
with notopodial lamella forming low bul dis- 
tinct lateral wings surrounding prostomium. 
Four pairs of densely pinnate branchiae from 
scliger 2. Setiger 1 with notosetae reduced to 
small bundle at base of notopodial lamella, 
neurosetae normal in size, Notopodial lamellae 
becoming more elongate dorsally, pointed and 
medially hooked over setigers 14+ then becom- 
ing rounded, laterally directed, decreasing in 
size posteriorly; in some anterior setigers 
notopodial lamellac extending across dorsum 
to form very low, rounded, hbarely-raised 
dorsal ridges from about setiger 10, occa- 
sionally absent. Neuropodial lamellae showing 
similar to notopodial size variations, generally 
rounded throughout except for sharp ventrally 
directed triangular projection in setiger 2 and 
laterally pointed lamellae in setiger 3. Anterior 
nolo- and neurosetae all capillaries, sheathec, 
distally granular, becoming, finer with less 
distinct sheaths posteriorly; hooded hooks from 
scliger 26-39 in notopodia, 20-24 in neuro- 
podia, hooks with 4—5 tiers of apical teeth 
above main fang, secondary hood distinct; onc, 
or rarely 2 ventral sabre setae in neuropodia 
from setiger 10-11, each sheathed in anterior 
sefigers, distally granular, tapering rapidly to 
filamentous tip. Pygidium with long dorso- 
medial cirrus and a pair of stout lateral 
papillae. 


Comments; Our material agrees closely with 
the description of Blake & Kudenov (1978) 
except for fewer pairs of eyes and slightly more 


posterior appearance of neuropodial hooded 
hooks. Variability in the setiger at which types 
of setac first appear was not recorded by 
Blake & Kudenoy. The pygidium and the occa- 
sional, variable presence of low dorsal crests 
have not been described previously. This is 
the first record of the species from South 
Australia. 

Australian distribution; S.A, (widespread*), 
Vic. (Port Phillip Bay), N.S.W. (Merimbula, 
Wagonga R.*), 


Habitat; Among seagrasses and algae, in nud, 
under rocks, 


Aquilaspio pyramidalis n.sp. 
FIG, 3a-¢, 


Holotype; South Australia, 20A (W.194024). 


Paratypes: OFA, 16 (W.194026). O4A, 1 (CW. 
194025). O7A, 9 (W.194029), 07B, 31 (W.194030). 
G8A, 14 (USNM 074898), 1BA, 12 (AHF POLY 
1382). [8B, 10 (RMNH 28. 1982.66-75). 19A, 
23 (W.194031). 20A, 2 (W.194027), 2LA, 28 
(W,194028). 27B, 4 (W.194032). 33B, 2 (W. 
194033). 


= 1 a 5 ‘ 

t \ 
= Ty 4 
a O57 ‘ 3 
a : é 

= 7 
a { 
\ “f b 
\\ \ 
\ ads 
\ f I 
/ 
) O05 / 
0:14 I 
bes. | ) | | 
Iwi) | i a 
oc ° | 


Big. 3, Aquilaspio pyramidalis tsp. a. anterior 
end, dorsal view (Paratype W.194026). b. 
anterior view of 4th parapodium. ec. anterior 
view of 52nd purapodium. d. notopodial hooded 
hook from 52nd parapodium. ¢. sabre seta. 
Scoles in mm, 


1) P. A. HUTCHINGS & S. P. TURVEY 


Description: Holotype, 10 mm long, 6.7 mm 
wide for 71 setigers, Paralypes range in size 
from 24 setizers, 2.0 mm length, 0.25 tm 
width to SL setigers. 18.5 mm length, 0.85 mm 
width, Prostuntium broadly rounded anteriorly: 
with low, thick caruncle extending to posterior 
margin of sctiger [; two pairs of eyes arranged 
in oblique line of cither side, Peristomium 
fused with nolopodial lamellie of setiger 1 to 
form high Jateral wings about posterior 
margins of prostomioum and base af caruncle, 
extending anteriovly as low lateral wings close 
to sides of prostamium (Fig. 3a): palps lost. 
Pour pairs of sparsely pinnate branchiae on 
sctigers 2-5, becoming shorter posteriorly with 
few pionules. Postsetal notopodial fanielhic 
well-developed and rounded ii setiger 1, in- 
creasing in size and becoming more dorsally 
elongate, pointed and medially hooked ip to 
sctiger + then becoming rounded, laterally 
directed, decreasing in size posteriorly. ter- 
minating laterally on dorsum and not extendine 
to form dorsal crests (Fig, 3b). Neuropoliam 
of setiger | inflated, pyramidal in form with 
postsetal lamellae barely developed as small 
ridge on aper; subsequent neuropodia normal: 
anterior setigers with well-developed postsetal 
lamellae showing similar variation in size to 
nolapadial lamellae; all rounded except 
bluntly triangular dorsally in setigers 2 and 3 
Postsetal lamellae of both nota, and neuro- 
podia reduced to low. thiek ridges, in those 
setigers beuring hooded hooks, with hooks 
often partly surrounded by folds im body wall 
(Fig. 3¢). Presetal lamellae present anteriorly 
in both noto- und netiropeslia as low ridges, 
Anterior noto- und neurosetae all capillaries 
arranged in narrow bundles in setiger |. there- 
utter capillaries in 2 broad, disareanised, par- 
lially sepuratec groups in hoth noto- and 
neuropodia reduced to single fasyciele hy middle 
setigers and | or 2 setae posteriorly; capillaries 
of anteriov setigers slowl, sheathed, distally 


granular. Frequently appearing  unilimbute, 
becoming slender posteriorly with ipcon- 


spicuous sheaths; O-3 hooded hooks froni 
setiger 26 (left)—27 Ceght) in motupadin, O-5 
from setiger 11 in neuropodia, with apical 
tecth in 5—-G tiers aboye main lang, two teeth 
per fier, primiury hood broadly inflated, 
secondary hood not Visihle Chis. 3d); single 
ventral sabre seta present in each neure- 
podium, from setiger 12, each stout, dyasely 
granular distally, tapering rapidly to fila 
méntous tip, sheath well developed anteriorly 


but diminishing posteriorly (Pig, 3e)  Pypir 
dium. with a smle. long dorsomedial cirrus 
and 2 stout lateral papilluc, Coclom loosely 
packed with caus of about 70° jam diameter. 
The paratype material exhibits some variation 
from the holotype. Some have 4 pairs of eyes; 
palps stout extending to setiger 4-10, Number 
of branchiae becoming reduced in small speci. 
mens with a corresponding reduction in 
number of pinnules frequently to otly 1, 2 or 
0 in more posterior branchiac. Smallest speci- 
men with only o single pair of branchiae on 
setiger 2 and completely lacking pinnules, sug- 
gesting thal number of gills and pinnules 
increase with increasing size and presumably 
age. Notopodia with 0-2 hooded hooks fram 
selizer 25-30 in most specimens increasing to 
as many as 5 posteriorly, Neuropodia with 
0-5 hooded hooks generally from setiger 11, 
rarely from seliger 12, One or rarely 2 sabre 
setac generally from Seliger 13, o¢easionally 
from setiger }J/—16, Very small specimens with 
holo- and neuropodial hooded hooks and 
neuropodial sabre setae from as early as 
setiger 18, 8 and 1 respectively. The smallest 
Specimens can Only he assigned to a@— pyre- 
mnidalis nsp. because of the wide range of 
sized miuterial available and this permits the 
sequential development of features tu be fal- 
lowed with increysing: size. 

Discussions Agquilaspio pyramidalis nsp. 1s 
similar to A. sauliipinetlata (Blake & 
Kudenov, 1978), 4. pervaded (Hartmann 
Sehréder, 1962), 4. temuiv (Verrill, 1880). AL 
tevelensis (Gibbs, 1971) and A. rreeedwelll 
(Harton, (951) in possessing four pairs of 
pindate branchiag on seligees 25, Ht may be 
distinguished from all of ihese species by the 
seligers on which neuropadial hooded hooks 
and sabre setae first appear and by whe form 
of the neuropodium of setiger 1. 

Kivrrefogy; The speciic name refers in the 
form of the oeuropodium of setiver 1. 
Anstralian distribution: South Australia (wide- 
spread), 

Habital; Interiidally among algae, seayrasses 
and under rocks, subtidally ameng racks anu 
sponges. 


Mituspie Foster 


Prostomiuim subtriangular, aneeriarts 
rounded, blunt or inflated, catendlig pus- 
teriorly as gx more or less well-developed car- 
uncle, Perislomium forming a hood surrouid- 
ing proastomium, variously develuped. 


SPIONIDAE OF SOUTH AUSTRALIA \! 


Branchiae all ciiform, Irum seliger 2, vary- 
ing from 440 pairs. Anterior setae all capil- 
lavies, Hooded hooks in posterior noto- and 
neuropodia, bidentile to mullidentite. Pygi- 
dim with anal cirri. 
Type species Prienaspie cirrifera Wiréo 
Mintuspio cirrifera Wiren 
Prionaspia (2) cirrifera Wirén, 1883: 409, 
Minnspio cirrifera, Boster, l97L: 1OB-1 12, figs 
262275 (for synonymy). 
Prionospie (Minuspiv) cirrifera. Blake & Kudenoy, 
978: Q2I-224, text-liy, 25a (fot synonymy). 
Matertal examined: S\, 02B, 1 (W,19302), 


Description: Posteriorly incomplete specinien 
of 61 setivers, mensuring 15 mm Jong, Ob 
mm wide. Prostamiun bluntly rounded, ear- 
uncle extending to posterioy margin of setiger 
1. Persiomium torming low Jateral wines 
partly cnelosing prostomium; palps slender, 
extending to setiger & Ter pairs of branchiae 
from seliger 2, all long, cirriform. Setiger 1 
reduced with postsetal notopodial lamella 
larger thin neuropodial but both small, noto- 
and neurosetae present, Posisetal netopodial 
lamellae increasing in size aud beeaming more 
dorsally pointed to setiger 8 then gradually, 
becoming smaller, rounder, more laterally 
directed, Forming low dorsal crests from setiver 
12, decreasing posteriorly to setiger 20 then 
absent, Postsetal neuropodial lamellae small. 
rounded lateral Maps, with those of setipers 
2-3 having slightly dorssl point, Presctal 
famellie smaller, rounded. Atterior setae all 
sheathed distally granular capillaries; setae 
hecoming finer posteriarly. Hooded hooks 
from setiger 49 in notopodia, 19 in. qeuro- 
podia, with apical teeth arranged in 3-4 tiers 
above main fang; secondary hood distinct, A 
single sabre setae in neuropodium from 
setiger 16, 


Comments; Foster (1971, p. 110) states that 
if the hooded hooks of M. cirrifera have n 
secondary hood, then “it is extremely closely 
applied to the hook and is barely distin- 
suishable (fig. 273)". The hook is Mustrated 
as having a secondary hood which is quite 
distinct below the main fang. a condition 
identical to that in our specimen. Notopodial 
hooded hooks appear slightly later in our 
specimen than indicated by Faster (1971) and 
Blake & Kudenov (1978) and the carunele is 
slighlly shorter than described hy the latter 
authors, None of these authors jndicate the 
presence of sabre setae In their texts, although 


they are illustrated by Foster (1971, fig. 269), 
Otherwise our specimen is in close agreement 
with bowl descriptions, This is the first record 
of the species from South Australia, 
Australian distribution: S.A, (Streaky Bay"), 
Vie. (Port Phillip Eny, Gippsland Lakes), 
N,S.W. (widespread), Ok (Deceptiun Bay) 
Habitat: Seagrass beds, mud. sand. 
Prionaspio Malmgren 

Prostamium, With anterior margin incised 
or rounded, without Frontal horns, caruncle 
vatiously ueveloped. Peristomiuim fused ih 
varying amounts with sctiger J often forming 
low lateral wings, Seliger | with reduced 
Pparapodia, notopadia on hranchiferaus seg- 
ments enlarged, post-branehia) notopodia 
becoming smaller, invonspicuous: dorsal folds 
or crests present or absent on posthranchial 
seyments rarcly on branchiferous seyients, 
Branchiae cirritorm and pinnate, limited to 
anicrior sctigers. Anterior setae al! capillaries, 
hooded hooks in posterior noto- and neuro- 
podia; hooks, bi, tri or miultidentate, inferior 
stbre setae present, Pyvidium with 1 long 
medial cirrus and 2 short ventrolateral cirri 
or thickened lobes. 

Type species Prionaspio steeasirapi Malm- 
gren. 

Prigmospio multicristata Hutchings & Rainer 
Prlonasplo multicrisiata Mutchings & Rainer, 1979: 

7THR-771, text, fig, Sa-i, 
Material examined: S.A, 02B, 1 (W.194023). 
TIA, 1 (W194022). IRA, 2 (W.194021), NSW, 
Careel Bay, Posidenia (Holotype W.8286). 
Descriptions A single entire specimen (W. 
19402!) of 77 setigers measures 12.5 mm 
long, 0,75 mm wide: posteriorly incomplete 
specimens. of 0,5-L,6 mm wide. Progstomium 
broadly rounded anteriorly, tapering rapidly 
lo narrow caruncle extending ta posterior 
margin of setiser 4, Two pairs of cys, antenur 
pair small lateral; posterior pair larger, camma 
shaped, Peristomium forming low-taleral wings 
close about prostomium. Four pairs of 
branchiae, on sttigers 2-5, First and fourth 
pairs long, thick, densely pinnate over basal 
2/3, distally bare; second and third pairs 
short, stout. cirriform. Setiger — with noto- 
podial and neurmpodial lamellae slightly 
reduced, both note- and beuroselae present. 
Notopodial lamellae becoming larger and niore 
dorsally pointed on sctigers 2-6, then rounded 
and decreasing gradually in size posteriorly; 
vach poir joined serass dorstim to form high 


12 P. A, FEUTCHINGS & S. P, TURVEY 


crest from setiger 7, decreasing posteriorly ta 
became medially separated at sctiger 24-31, 
then absent. Neuropodial lamellae showing 
similar variation in size ta wotopadial, rounded 
except with sharp triangular downwards pro- 
jection in Seliger 2, Anterior noto- and neuro- 
sctac all sheathed; capillaries, densely granular 
distally, in one specimen (W.194022) sheaths 
of many anterior capillaries also densely 
granular, intensely gold in colour capillaries 
becoming more slender with sheaths reduced 
posteriorly. Hooded hooks from setiger 27-31 
in notopodia, 14-18 in neuropodia with 4—S 
tiers of apical teeth aboye main fang, 
secondary hoad distinct. Qne or rarely 2 
ventral sabre setae in neuropodia from setiger 
Ti. cach stout, sheathed, distally granular. 
tapering abruptly to filamentous tip. Pygidtum 
with long, filiform cirrus dorsomedially and 
two stout lateral papillae, 


Comments; The only substantial differetice 
betwee our materia! and that of Hutchings & 
Rainer (197%) is the earlier appearance of 
nolapodial hooks, This is probably due to the 
smaller size of our specimens, ‘The pyeidium 
of one specimen was intact und there was 
some Vartabilily in the setiger at which hooded 
hooks first appeared. Neither of these features 
have been described previously, This is the first 
record of the species fram South Australia, 


Australian distviburtion: WA. (Cervantes*), 
S.A. (Streaky Bay*, Port Lincofn*}, N.S.W. 
(Merimbula*, Pott Hacking’. Careel Bay), 
Qld (Calliope R.*). 


Afabitat: Sand, seagrass beds, 


Spio Fabricius 


Prostomium anteriorly rounded or incised, 
frantal herns lacking: eyes present or absent, 
Hranchiae from setiger | continuing through- 
out body, sometimes partially fused to dorsal 
lamellae in anterior setigers, [ree posteriorly, 
Notosetae all capillaries, neurosetae including 
capillaries, hooded hooks and sabre setac, 
Pygidium with anal cirri, 

Type species Nerety filicornis Miiller. 


Spio tridentata n.sp. 
FIG. 4a-d 


Holotype: South Australia, 2B, (W.194079). 


Vitter Malerial exuuiiied> Spia parifica NSW. 
Towra Point, Botany Bay, St 329 Mulophila, 13 
Pararypes (WAMO29) coll, SSW, State Pisherics 
Wh, Alike & Kodeniy, 


Descriptions Body robust, broadly fectanenlar 
in cross-section. Colour pink. Posteriorly in- 
complete fragment ol 57 seligers, LS am Jony 
and 1.9 mm wide at setiger 25, Prostomium 
broad, blunt, anterior margin almost truncate 
with faint media) indentation; without lateral 
Wines; eyes not visible, curuncle broad, pos- 
tertorly rounded, extending, to posterior margin 
ol setiger | (Fig, 4a). Nuchal organs not 
visible, but tissue damaged in that region. 
Peristomium broad, not forming lateral wings 
about prostomium. Branchiae thick, cirriform, 
distally rounded, well developed on setiger J, 
increasing gradually in sive over first low 
sctigers to attain a twice initial length by 
setiger 6-8 (Fig. 4b) then decreasing slightly 
over remaining seligers (Fig. 4c), Setiger 1 
with noto- and neuropodial setae and lametiac. 
Subsequent parapodial lamellae all thick, in- 
Hated. Postsetal notopodial lamellae rounded, 
extending dorsally and fused, except for small 
dorsal extremity, to lateral margins of 
branchiac, increasing in size over firsi few 
sclivers: presetal notopodial lamellae low, 
rounded, much smaller than postsetal lamellae 
anteriorly, becoming larger postetiorly but nol 
extenuling more than half way to cdge of 
postsetal lamellae. Postsetal neuropodial 
lamellac semicircular in profile and initially 
smaller than notopodial, attaining similar size 
by setiger 15-16 then increasing further to 
become somewhat larger posteriorly; presetal 
peuropodial lamellac low, rounded, much 
smaller than postsetal in anterior their enlarg- 
ing laterally to reach almost as far as postsetal 
in posterior sctigers. Notas¢tac all capillaries, 
anteriorly arranged in two broad, parallel rows 
with a smaller third group dorso-posteriorly. 
The two major rows then coalescing in middle 
seligers and remaining as single row pos- 
teriorly, Anterior netiresetae all capillaries in 
two broad rows, posterior row replaced from 
seliger 28 (leltj)--29 (right) by a single, simi-: 
larly broad row af 6-11 hooded haoks with an 
addilional, Ventrally-reflexed givup of 4-5 
sabre setae. All capillaries sheathed, those tn 
the anterior of the two major rows of both 
parapodial rami with shafts distally yranu- 
lar, those in’ postetior rows non-wranulary 
cupillary sheaths never graouwar; with dorsal 
granular capillaries in both purapodial rani 
frequently appearing unilimbate. Neuropodial 
hooded hooks tridentate wath large. pointed 
main fang surmounted by 2 stout apical teeth 
decrepsing successively in sive (Mp. dd). Pri- 


SPIONIDAE OF SOUTH AUSTRALIA x) 


mary hood granular, completely cnclosing 
teeth fine but clearly visible secondary hood, 
Sabre setae unilimbate with shafts finely 


granular distally, tapering gradually to a fine 
tip. Pygidium and posterior setigers lost. 


Fig. 4, Spio tridentata n.sp, a. anterior end, dorsal 
view, b, anterior view of 6th parapodium, c. 
unierior view of 30th parapodium, d. hooded 
hook. 


Comments: Spio tridentata n.sp. is similar to 
§. cirrifera (Banse & Hobson, 1968), 8. limi- 
cola Verrill, 1880 (after Holmquist, 1967), 
S, pacifica Blake & Kudenov, 1978 and S, 
pettiboneae Foster, 1971 in possessing triden- 
tate hooded hooks. Tt differs from all of these 
species in the much later appearance of the 
hooks and the almost complete fusion of 
branchiae and notopodial lamellae in at Teast 
anterior and middle setigers. Further dif- 
ferenees include the conspicuous, triangular 
presetal notopodial lamellae of S$, pettibonege, 
the presetal notopodial cirrus in anterior 
seligers and the dorsally bilobed prostomium 
of §. cirrifera and the bilobed caruncle, par- 
tially hooded sabre setae and = granutar- 
sheathed capillaries of S. pacifice, 


Eryinoloyy; the specific mame refers to the 
tridentate hooded hooks. 

Australian distribution: S.A. (Stokes Bay, Kan- 
varoo Island). 


Habitat; Under rocks. at low tide level. 


Microspio Mesnil (following Blake & 
Kudenov) 


Prostomium anteriorly rounded to bilobed, 
without frontal horns; eyes present or absent: 
occipital tentacle present or absent. Branchiaec 
from setiger 2, notosetae only capillary; neuro- 
setae including capillaries, hooded hooks and 
sabre setae. Pygidium with anal cirri, 

Type species Spio 
Claparéde. 


mecznikowianus 


Microspio granulata Blake & Kudenov 
Milorospio pranulata Blake & Kudenov, 1978; 232, 
figs 30-31, 

Material examined: S.A, 02A, 1 (W.19303). 03B, 
13 (W.19306). 03C, 1 (W.19304). O5E, 8 CW, 
19305). 03F, 2 (W.19307). 

Description: Size range of entire specimens of 
27-40 setigers, 3.2-11 mm long, 0.5-1.1 mm 
wide: posteriorly incomplete specimens up to 
1.6 mm wide. Prostomium bilobed, deeply in- 
cised; caruncle extending to setiger 2 with 
prominent pointed or rounded occipital papilla; 
high transverse ciliated ridge behind caruncle 
enclosed laterally and posteriorly by curved 
nuchal grooves; similar ridge on each suc- 
ceeding setiger; two pairs of eyes in oblique 
series; palps stout, basally inflated, extending 
lo setiger 10-14. Branchiae stout, cirriform, 
from setiger 2 to all but last few setigers. 
Setiger 1, reduced without notosetae, noto- 
and neuropodial lamellae small. Postsetal noto- 
podial lamettae of most setigers small, rounded, 
dorsally directed; bluntly pointed dorsally in 
far anterior setigers; becoming elongate and 
tongue-like in far posterior setiger. Postsetal 
neuropodial lamellae small, rounded, decreas- 
ing posteriorly, Presetal Iamellae in both rami 
smaller, low, rounded. Notosetae all capil- 
laries, Neurosetae with capillaries anteriorly 
tridentate. hooded hooks from setiger 9; a 
single sabre setae ventrally from setiger [4—17. 
Pygidium with 4 short, stout anal cirri dorsal 
pair slightly longer and more pointed than 
ventral pair. 

Comments: Our material agrees closely with 
the description ol Blake & Kudenov (1978). 
The pygidium and parapodial lamelae of far 
posterior setigers have not heen previously 


l4 Pr. 


described. This is the first record outside the 
ivpe locality, 


Ansrralian distribution: S.A. (Streaky Bay"), 
N.S.W. (Botany Bay). 


Mabini: Among mussels, seagrasses. in sand, 


Boecardla Carazzi, emended Blake & 
Kudenoy. 


Prostomiun rounded or divided, extending 
posteriorly as carunele. Setiger 1 with or with- 
out notosetac. Setiger 5 modilied with 2 lypes 
of major spines, Companion setae absent. 
Bidentate hooded hooks from setigers. 7-11. 
Posterior notopodial spines present or absent. 
Branchiae from setiger 2, absemt setizer 4, 
present on following variable number of seti- 
gers, Pygidium disk tike with or without 
separate lobes or reduced ta small lobes or 
cults, 


Comment: Boccardia fleckera tsp. has hooded 
hooks trom setiger Lj. The generic definition 
is revised here ty ucconimodate that species, 
Type species Polvdera pelvbranchia Haswell. 
Boecardia ehilensiv Blake & Woodwick 
Roccardia chilensis Blake & Woodwick LOT: 36, 
Blake & Rudenov, 1978; 238-240, fly. 33d-c. 
Material examined: $A, O6A, many (W.19295), 
Coorong (W.19208) coll. M. Geddes. 
Deseriprion: Prostomium deeply divided on 
anterior margin, Seliger 1 with long notoselae, 
Setiger 5 with spines of 2 types, simple faleate 
spines and spines with expanded concave cup 
containing bluntly conical tooth; bidentate 
hooded hooks from setiger 7. Branchiae from 
setiger 2. Pygidium a fleshy pad. 
Comments; Our material agrees well with pre- 
vious descriptions exeept that in the South 
Austtalian material, the oecipital tentacle as 
absent, First record from South Austratia, 


Australian distribution: S.A o (Venus Bay’. 
Coorong"), WA. (Bunbury. Lesehenaule 
Inler#), Vic. (Port Phillip Bay). NSW. 


(widespread) and Macquarie Island, 
Habitat: In amongst algal matt, 


Boccardia feckers o.sp. 
FIG. Sa-f, 
Haletype: South Australia 30B, 1 (W.194020), 
Description Posteriorly incomplete, 23° seti 
gers measuniug + mm Jong and 0.5 mm wide. 
Robust body, speckled with brown flecks of 
pighient, concentrated posteriorly on both 
ventral ancl dlorsal surfaees, Prostomium ceeply 


A, HUTCHINGS & S. PL TURYEY 


imeised, with 2 pairs of spherical eyes; ear- 
unele present, extending to middle of setiger 2 
with short occipital present. Palps with swollen 
bases, extending to setiger [0 (Fig. 5a). Setiger 
1 with large prominent notopodial lamellae 
and notosetae; subsequent parapodia with 
blunt triangular notopodial lobe, and larger 
truncate triangular newropadial lobe (Fig, 
Sheu). Branchiae, stumpy, stout from setiges 
2-4 and 6 onwards, attached to base of nota- 
podial lobe, but Jonger than parapodial lobes, 

Selwer 5 heavily  nrodified, notopodial 

lamellae absent, small globular ocuropodial 
lobe, with 5 worn brush tipped setae (Fig, Se) 
and 3 curved fulcate snioath spines OFig. Sf), 
neurosetal capillaries present, Neurosctae 
anteriorly long thin narrow bladed capillaries, 
from setiger 17, one to two hooded strongly 
bidentate hooks (Fig. 5d) present and by 
setiger 13, hooks predominate. Notosctuc all 
capillaries at least to setiger 23, 
Conunenrm: Boecardia flechera msp. has been 
placed within the genus Boecardia even thourh 
it does not strictly agree with Blake & 
Kudenoy’s emended gencrig description in that 
ihe veuropodial hooded hooks bexin on setiger 
tl and not on setiger 7-8. Rainer (1973) 
erceled the genus Paraboecardia tor species 
with hooks commencing on seiger & which 
Read (1975) reduced to a subgenus, This was 
accepted by Blake & Kudenoy (1978). Wood- 
wick (1964) erected another gens m_ this 
complex Tripolydora, for species With hooks 
commencing on seliger 9. Blake & Woodwick 
(19381) have recently sugzested that this gems 
is more closcly related to the Polydora complex 
than to Bececardia, As we have only iw single 
specimen we have decided to deseribe it ay a 
new species within the venus Boeccardia, as it 
clearly belongs to thes complex from the modi 
fication of sciger 5S and the type of setae 
present. 

Etymology: the specific name fleekera refers 
lo the pigmentation pattern on the bay, 
dusrralian disrributlon: S.A, (Cape du 
Couedi¢, Kangaroo Island). 

Habitat; Exposed algal holdfasis. 


Baceurdia probaseldea Hartman 
Boceurdia probeseidea Horton Wt) 382 
Make & Kudenov, 1978: 238; fig. Vac 
Material exiitined SA, OO OW Ta?) 
(W:19296), mony individiuils at bath siues, 
Description: Prostomium rounded on anterior 
nyatgin; carunele extending to end of setiger 3, 


tial 


SPIONIDAE QF SOUTH AUSTRALIA 15 


ei bey le oo: 
(og bol” 
A Pa ee: 

Bil "4 } Q i 
ae oe 
, 5 

, P o2 i ' 


oo? 


Fig, % Boccardia fleckera nasp, iu. anterior end. 
dorsal view. 6. woterior of view of 4th para- 
podium. ¢. anterior view of Sih parapodium. 
d, neurosetal hook. 2-f. modified setae of sctiger 
5, Scales in mm. 


with two pairs of eyes. Branchiae from setiger 
2, absent from posterior third of body. Setiger 
5 with two types of setac: one type simple, 
faleale, weakly hooked with blunt tips, second 
type with broad asymetrical flattened head, 
slightly domed, densely bristled distally. 
Bidentaute hooded hooks from setiger 7. 


Cominents: Our material agrees with the 
description of Blake & Kudenov (1978) who 
first reported this species from Australia in Port 
Phillip Bay, Victoria, the first record from the 
southern hemisphere, First record fram South 
Australia. 

Australian distribution: WA, (Premanile® ), 
S.A, (Blliston®), Vie. (Port Phillip Bay). 
Habiiat: Ta amongst encrusting algae or Galea- 
laria worm, tubes. 


Polydora Bosc, emended Blake & Kudenov 

Prostomium entire or divided, extending 
posteriorly as caruncle; cyces present or absent. 
Setiger 1 with or without notoselue. Setiger 5 
greatly modified with major spines of one kind 
usually with slender companion setae, spines 
arranged in a singled curved row. Posterior 
notopodial spines sometimes present. Neuro- 


podial hooded hooks bidentate beginning on 
seligers 7-17. Branchiae commencing pos- 
teriorly to seliger 5, Pygidium variable, 
reduced or enlarged, culf-like, satucer-like or 
lobate. 

Type species Palydora cornuta Bosc. 


Palydora haplura Claparéde 

Palydora haplura Claparede, 1870; 58, Read, 

1975; 411. Blake & Kudenov, 1978: 264, fix. 

47. 
Material examined: S.A. V7 (WA9298), 
Description: Latge species up to 40) mm in 
length for over 160 segments. Prostomium 
weakly incised. with caruncle extending to end 
of setiger 3, bearing low occipital tentacle. 
Setiger 1 With neurosclac; notosetac absent. 
Modified setae, sctiger 5 blunt to pointed with 
subterminally lateral flange present, frequently 
resembling a tooth; companion setae bilimbate. 
Hooded hooks. [rom setiger 7 with constricted 
shaft. Far posterior segments with large 
recurved dorsal spines directed toward mid- 
line of body. Branchiae from setiger 7, con- 
tinuing along body until spine bearing region. 
Pygidium broad, flat, with deep ventral notch. 
Comments: This species has been previously 
reported as forming mud blisicrs on oysters, 
In South Australia the species occurred 
amongst encrusting fauna on jetiy piles. This is 
the first record from South Australia. 
Australian distribution: S.A. (Rapid Bay*), 
Tas. (Sirmmons. Beach), Vic. (Port Henry 
Pier, Corio Bay, Port Phillip Bay). 
Habitat: In amongst sessile organisms on 
jetty piles. 


Polydora livni Webster 
Palydora lieni Webster, 1B79: 119, 
1971 und Foster, 1971 for snyonymy, 
Material examined: S.A, 12B, 5 (W.19299), 
Description: Large specimen up to 32 mm 
length for 80 setigers. Prostomium bluntly 
bilobed with occipital tentacle. Two pairs. of 
vyes. Setiger 1 without notosetae, digitiform 
notopodial lobe, Setiger 5, simple falcate major 
spines with blunt subdistal tooth; companion 
setae delicate, feathery; dorsal and ventral 
capillaries absent. Hooded hooks with constric- 
tion on shaft from seliger 7. Posterior modified 
sétac absent. Branchiac from setiger 7. 
Camments; First record from South Australia, 
Australian distribution: S.A (Torrens 
Island*), Vic, (Port Phillip Bay) 
Habitat: Totertidal mudflats. 


See lake. 


16 P. A. HUTCHINGS & S$. P. TURVEY 


Polydora socialis (Schmard;) 

Polydora socialis. Blake, 1971: 20-23, figs 13-14, 

1979: 607-609 (synonymy): Blake & Kudenoy, 

1978: 248-250, fig. 38d-e. 
Material exantined: S.A. 02A, 5 (W. 19300), 02C, 
2 (W.19301). 
Description: Moderately sized individual up to 
9 mm long and 0.75 mm wide for 554 setigers. 
Prostomium deeply incised, caruncle extending 
to xetiger 4—5; occipital tentacle absent; with 
two pairs of eyes. Setiger 1 with notosetae. 
Major spines of setiger 5 simple, falcate with 
subterminal swelling. WNeuropodial hooded 
hooks from setiger 7, without constriction on 
shaft. Modified posterior setae absent, 
Branchiae Jrom setiger 8. Gizzard externally 
shown by dorsal swelling on setigers 18-19. 
Comments: First record from South Australia. 
Australian distribution: S.A. (Streaky Bay*), 
Vic. (Port Phillip Bay). N.S.W. (Botany Bay. 
Sydney Harbour). 
Habital; On mud flats, associated with clumps 
of mussels or Posidonia seagrass, 


Pseudopolydora Czerniaysky emended 
Blake & Kudenov 

Prostomium entire or divided, extending 
posteriorly as caruncle, occipital tentacle 
present or absent, Eyes present or absent. 
Setiger 1 usually reduced, with or without 
notosctae (and in some species without neuro- 
selae if animals are reproducing asexually or 
regenerating). Setiger 5 not greatly modified 
with noto- and neuropodia often well deve- 
loped bearing postsetal lobes, and spreading 
fascicles of capillaries, with curved row of 
heavy modified spines of 2 types or single type 
with companion setae; modified setae often 
arranged in J or U shaped setal group. Pos- 
terior notopodial spines sometimes present. 
Neuropodial hooded bidentate hooks from 
setiger 8. Branchiae present posteriorly 10 
sctiger 5, Pygidium variable, enlarged or 
reduced, collar like or divided inta lobes or 
small Jappets. 

Type species 
(Claparéde), 


Psendopolydora antennata Clapuréde 
FIG. 6a-c. 
Polydora (Psendopolydera) antennata. Hartmann- 
Schréder, 1981: 50, figs 115-118. 
Material examined; S.A. 16A, 1 (W.19986). IBA. 
3 (W.19385). 20A, 6 (W.19387}, 2A, 
3 (W.19388), 27A, 1 (W.19384), NS.W., Merim- 
lula (Wt1703). Old. Mereton Bay, Jackson 


Pyeudopalydora antennata 


Creek (W,6042), Serpentine: Creck (W.6043), 
Brishane R, (W.7474). 

Description; Colourless. Length up to 25 mm 
for 45 setigers, some complete individuals, 
Prostomium deeply incised, with lobes widely 
flaring. Occipital tentacle pointed erect, car- 
uncle extending to middle of setiger 6. Two 
pairs of distinct eyes, Setiger 1 with small 
rounded notopodial lobe, notosetae absent 
(Fig. 6a). Setiger 5 poorly modified with 2 
types of setae, arranged in J shaped row, outer 
row consisting of pennoned setae (Fig. 6b), 
(short spoon-like tips to blades) and inner 
row of simple spines, slightly curved apically 
and tips finely hirsute (Fig. 6c). Hooded 
bidentate neuropodial hooks from setiger 8, 
although in one specimen 2 hooks present on 
sctiger 7, hooks immediately replace neurosetal 
capillaries.. Branchiae present from sctigers 7— 
22. Pygidium 2 semicircular valves. 
Comments; This is. the first record of this 
species from southern Australia, although 
Hartmann-Schréder has recently described il 
from Geraldton in Western Australia. 


0-1 ) 
| 
| 


c b 


tig. 6. Pseudopolydora antennata a, anterior view, 
dorsal view (W.19385), b-c, modified setae of 
Setiger §. Scales in mm. 


SPIONIDAE OF SOUTH AUSTRALIA 1? 


Australian distribution: W.A. (Geraldton*), 
S.A. (Sellick’s Beach, Victor Harbor, Kan- 
garoo Island), 


Habitat: Crevice fauna often associated with 
tufted algae. 


Pseudopolydora paucibranehiata (Okuda) 
FIG, 7a-e. 


Polydora  (Carazzia)  paucibranchiata 
1937; 231-233, fles 11-12. 

Pseudopalyora paucibranchiata. Blake & Kude- 
nov; 1978: 268. 


Pyeudopolydora kempi. Hutchings & Rainer, 1979: 
773-774. Not Southern. 

Material examined: S.A. 11A, 1 (W,19393). 12A, 
14 (W,19389, 19390), 12B, 1 (W.19391), 12C, 
2 (W.19392). N.S.W. Botany Bay, Towra Beach 
(W,13045), Kurnell (W.17427).. Jervis Bay 
(W.5223). Vic. Port Phillip Bay (NMV G3177, 
3178, 3180), Hobsons Bay, Yarra River (NMV 
C3183), identified Blake & Kudenov. 


Okuda, 


0:07 
\ | 
1 a b 
f 
ae / 
0:07| ww) \ | 
|| | | 
\ | }} 4 
\ \ | | \ \ c¢ 
e d 


Fig. 7. Pseudopolydora paucibranchiata a—b. modi- 
ficd setae of setiger 5 (W-.1727), S, Australian 
mitterial. ce, modified setue of setiger 5 
(NMV G3183, NMY G3180) ~~ Victorian 
material, Scales in mm, 


Description: Small individuals 3-5 mm in 
length, 0.5-1 mm in width. Prostomium entire, 
caruncle to posterior margin of setiger 3, occi- 
pital tentacle short, Palps to setiger 13-18, 
extending approximately quarter of length of 
body, Branchiae from setiger 7 extending to 
setiger 20-22. Notosetae absent on setiger 1. 
Setiger 5 barely modified, with simple falcate 
spines (Fig. 7b) and curved pennoned spines, 
arranged in U shaped line (Fig. 7c). Hooded 
neuropodial hooks from setiger 8, bidentate, 
completely replacing neuropodial capillaries. 
Pygidium a small flaring cup. 
Comments: The South Australian material 
differs from the description of Blake & 
Kudenoy in the number of branchiferous 
segments and relative lengths of the palps. 
Examination of this material, much of which 
is in poor condition indicates that the gills 
extend only to setiger 21—23, and not to setiger 
35 as quoted by Blake & Kudenov, and this 
even for gravid females (NMV G3183), 
Blake & Kudenoy'’s material exhibits far 
greater variation in the shape of modified 
selac on setiger 5 (Fig. 7c-e) than exhibited 
by the South Australian material (Fig. 7a—b). 
The type of Pseudopolydora paucibranchiata 
was destroyed during the Second World War- 
However material from the type locality 
should be examined to check the apparent 
wide distribution throughout the Pacific: 
Australian distribution: S.A, (Porter Bay", 
Torrens [sland*) Vic. (Port Phillip Bay, 
Westernport Bay) N.S.W. (lervis Bay, Botany 
Bay). 
Habitat; Mudflats and seagrass beds. 


Pseudopolydora sp, } 
FIG, 8a-c. 


Material examined: $.A., 12A, 2 (W.19397). 12B, 
many (W.19398). 12C, 2 (W.19399). All pos- 
teriorly incomplete specimens, 

Description: Colourless. Prostomium deeply 
incised, lobes widely flaring; small caruncle 
extends to posterior margin of setiger 3-4. 
Two pairs of eyes. Occipital tentacle absent. 
Setiger 1 without notosetae, Setiger 5 mode- 
rately modified, modified setae arranged in 
tight U, of 2 types, simple fulcate spines (Fig. 
82) about 6, and pennoned spines (§), with 
2 longitudinal ridges about concavity, dorsally 
finely hirsute (Fig, 8b-c). Hooded bidentate 
neurosctal hooks from setiger 8, in groups of 
about 13. Branchiae from setiger 7 to ahout 
setiger 25- 


18 P. A. HUTCHINGS & S. P, TURVEY 


0:07 


b 


c 


Fig. 8 Pseudupolydora sp. | a-c modified setauc 
of setiger 5 (W.19398), Scales in mm. 


Coniments: This species differs from both 
Pseudopolydara antennata and P, paucibran- 
chiata which oceur in South Australia, and alsa 
differs from other described species from SE 
Australia, Pseudopolydora sp. 1 differs from 
these other described from SE Australia, P. 
kempi (Southern, 1921), P. glandulosa Blake 
& Kudenov, 1978, P. stolonifera Blake & 
Kudenoy, 1978 and P. prolifera (Augener, 
1914) in the type and ornamentation of the 
modified setae on setiger 5. It probably repre- 
sents an undescribed species, but we have only 
incomplete specimens and as the genus we 
believe is in need of a re-evaluation we have 
decided not to describe another new species. 
Habitat: This species occurs around Torrens 
Island Power Station, both in the thermally 
polluted arcas where temperatures may cxceed 
40°C and in the non thermally polluted areas. 


Pseudopolydora sp. 2 
FIG. 9a-b, 

Materjal examined: S.A. 12C, 1 (W.194034). 

Descriptions Colourless. Entire specimen, 57 
sctigers,, Prostamium entire, rounded. Eye 
spots present, caruncle and occipital tentacle 
present but damaged; setiger 1 small, lacking 
notosctae, Setiger 5 barely modified not en- 
larged, modified setae arranged in small U 
shaped group, consisting of numeraus fine 
pennoned (Fig, Yb) and fulcate spines (Fig. 
91). Hooded multidentate meurosetal hooks 


from setiger 7 and unidentate notosetal hooks 
from setiger I4, immediately completely 
replacing capillaries. Multidentate hooks with 
a group of small denticles closely above main 
fang, Hhumbers of tecth cannot be determined. 
Branchiac from setigers 7-24. Pygidium. an 
anal collar, notched dorsally. 


0:07 


b 


Fig, 9. Pserdopulydora sp. 2 a-b modified setae of 
setiger 5. Scales in mm. 


a 


Comments; This species clearly differs from 
P, anternata or P. pseudopolydora as identified 
by us. As there appears to be some confusion 
within this genus and we have only a single 
specimen, We have decided to just refer it to 
genus, although it clearly differs from all 
species of this genus currently described 
from Australia, We are also reluctant to 
decribe a new species on a poorly preserved 
single specimen. 

Habitat: Mud flats. 


Acknowledgements 


We should like to thank ABRS who funded 
this. study and provided the salary of one of 
us (SPP). Chris Glasby prepared the majority 
of the figures. PH would like to thank Dr 
Stan Edmonds for his hospitality and great 
ussistance in the field, collecting the material 
which forms the basis of this paper. Dr Alan 
Butler ulso assisted in the collection of the 
material. Dr Lu of the National Museum, 


SPIONINAP OF SOUTH AUSTRALIA 19 


Victoria, Mr Zeidler of the South Australian 
Museum, Dr Kirkegaard, Zoulogisk Muscuni, 
Copenhagen Kindly lent ts jaterial, Constric- 


tive criticise af the manuscript was kindly 
eiven by Dr J, Blake and a anonymous 
reviewer. 


Heferenves 


AuceNtR, H. (1924) Papers [rom By 1. A, Mor- 
tensen's Pacific Expedition 1914-1916. No . 14 
Polychaeta 1. Polychuela von den Auckland 
und Campbell-Inseln, Vidersk  Adedde, danish 
#arirh, Foren. 75, 1115. 

—— (1926) Pupers front Dr To H. Mortensen’s 
Pacific Expedition W914 16. Polychaeta Il. 
Polychuelen von Nevseelind, 11) Sedentaria. 
fhid $1 157-294, ; 

Eanse, K, & Lhosson, K. D. (1968) Benthic poly- 
thuctes from Puget Sound, Washington with 
femarks on four other species. Prac, US. Nat, 
Mfrex. 125, 1-53, 

Brake, J. A. (1971) Revision of the Genus Poly. 
dara fram the east coast of Narth America 
Polychacta: Spionidac).. Sediiy. Contribs, Zoal. 
75, 1°32, 

(1979) Revision of some pulydorids (Paly- 

thavta: Spionidve) described and recordel From 

Uritish Columbia by Edith and Cynl Berkeley. 

Proc. Biol. Soc. Wash. 92 (3), 606-617. 

(1981) The morphology of = 7'ripalyearn 
spinosa Woodwick (Polychaeta: Spionidael. An 
application of the scanning electron micnmscope 
to polychacte-systematios, Proc, Biol, Soc Wash, 
94 (2), 352-362. 

—— & Kupenoy, J. BD. (1978) The Spronidae 
(Polychieti) from southenstern Ausiralia and 
adjacent areas with a revision of the genera. 
Mem. natn. Mus, Fiet, 39, 171-280, 

& Woodwick, K. H. (1971) A review of dhe 
eens Bex waredta Carazzi (Polychactas Spionidae) 
with description at two new species. Bull. Sth. 
Calif. dead. Sci, TMCV), 34-42, 

Craparfor, B. (1870) Les Annélides Chétopodes 
du Golfe de Naples, Méin, Soe. Phys. Hist 
Nat. Geneve 20, 365-542, l4 pls. 

Day, J. H, & Hurerinas, P, A. C1979) An anno- 
tated checklist of Australian and New Zealand 
Polychueta, Archiannelida and Myzostomidac. 
Ree, Aust, Mus, 32, 80-161. 

Fosirk, N. M, (1971) Spionidae (Polychaeta) 
of the Gulf of Mexico and the Caribbean Sea. 
Studies on the Faw af Curagaa and other 
Caribbean tstands, 36, 1-183, 

Guns, &. B CL971) ‘Che polychaeta fauna of the 
Solomon Istunds. Bulf, Bro Mus. net. Hest. 21, 
1OT-211. 

Haniman, ©, (1940) Boceardia proboseiden, a 
uew species of sptonid worm from Californin. 
J. Wash. Acad. Set, 30, 982-387. 

— (1951) ‘Phe littoral marine annelids of the 
Gull of Mexico. Publs, Inat, mar. Sei, Univ. 
Tex. 2, 7-124, 

~ (1969) “Atlas of the Sedentariate Paly- 
chactous Annelids from California’. (Athin 
Munecock Foundation: Los Angeles}. 

Hanivann Seuronne, G, (1962) “weiter Hejtrap 
fut Polyohacten-fauie Von Petu. Kieler Meerer 
forsh, 1B, LO9L47, 


— (1979) Jn Hartmann-Schrader, G. and 
Harimunn, G. (1979) Zur kenntnis des Puli 
torals der australischen Kilsten umer besander 
Rerucksichtigune der Polychaeten und Ostra- 
coden. (Teil 2 und Teil 3). Die Palychoeten 
der tropischen — Nordweatkiiste — Australiens 
(zwischen Derby im Norden und Port Hedtarnu 
im Sudden). Mite Ranh. zool. Muy. Inve. 77. 
75-218, 

—— (1980), Die Polychacten der Wopisehen 
Nordwestkiiste -Australiens (zwischen Port 
Samson im Norden und Exmouth im Siiden). 
thid. 77. 4l-110. 

—— (1981) Die Polychacten der (ropisehen- 
subtropischen Westkiiste Austriliens (zwischen 
Exmouth im Norden und Cervantes im Suen), 
thid, 78, 19-96, 

—— (1982) Die polychaeten der subtropischanii. 
borcalen Westktiste Austealiens 9 (zwischen 
Cervantes im Norden und Cape Naturatiste in 
Stiden), bid, 79; 51-118, 

Hotmauist, GC. von (1967) Marenzatlatia wireni 
Atigener —1 pabvelete found in fresh Waters 
of northern Alaska—with taxonomies) coushlers- 
lions On some reluled Spionid worms. Zeal, Syat 
Evalfersel, 5, 298-313, 

Houtcanas, P, A. & Ratner, S, F ¢1979) The 
polychacté fauna of Carcel Bay, Pittwater, New 
soul Wales, Australia. J. mat. Flist. 13, 745> 

hy 

Muiier. O. FB. (1808) “Zoologica Banicu seu 
Animalium Diniae_ct Norvegiae raiyorum, ac 
Minus notoyym Descriplipnes el Historian”, 
Havniae, 

Oxups, 8. (1937) Spionilovm polyehactes Fron 
Japan. fowr Frc, Sei Hokkatda Oniv, §, 217 
254, 

Pernsonc, M. HH, (1963) Revision of some 
genera of polychaete worms of the family 
Spionidac, including the description of a new 
species Of Seolilepis, Prag, Rial, Sov, Wash, 76, 
89-704. 

Poorn, G, C, By, Ramer, S, Seis. R. B. & 
Warp, B. (1975) The zoobenthos proyram i 
Port Phillip Ray, 1969-73, Fish & Wildlife Pap, 
Viet, 7, 1-78, 

Rawer, S. F. (1973) Polydora und reluled genera 
(Polychieti: Spronidae) from Otago waters. 
JiR. Sac, NZ. 3, 545-564. 

Rastos. J. M, (1976) Aenides axveephela (Sars, 
1862) remarques taxonomiques, Vie, Milten 26 
(1A), 11-20, 

Reap, Gi, B. (1975) Systematics und Biolowy of 
Polydovids species (Polychaeta: Spienidie) from 
Wellington Harbour, Jnl Rey. See, NZ, 5 {4), 
395-419, 

Sars, M. (1862) On unnelidslaevien Nerine og 
duns norske arter. Forh. Midensk. Nelsk. Krist. 
TRGT: 59-67, 

Semruere, R. (1921) Fauna of Chilka Lake. 
Polychaeta of the Chilkha Cake and iso of fresh 
and brackish waters in olher parts of Tndin, 
Mem. ladian Mis, 3, 363-659, 


20 P. A. HUTCHINGS & S. P. TURVEY 


VERILL, A. E. (1880) Notice of recent additions 
to the marine Invertebrata of the northeastern 
coast of America, with descriptions of new 
genera and species and critical remarks on 
cthers. Proc. U.S. Nat. Mus, 2, 165-226. 

WEBSTER, H. E. (1879) The Annelida Chaetopoda 
of New Jersey. Ann. Rep. N.Y. St. Mus. nat. 
Hist. 32, 101-128. 


WIrREN, A. (1883) Chaetopoder fran Sibiriska 
Ishafvet och Berings Haf insamlade under 
Vega-Expeditionen 1878-79. Vega-Exped. 
Vetenskapliga lakttagelser 2, 383-428. 

Woopwick, K. H. (1964) Polydora and related 
genera (Annelida, Polychaeta) from Eniwetok, 
Majuiro and Bikini Atolls, Marshall Islands. 
Pacif. Sci. 18, 146-159. 


SPECIES OF CERATOBAEUS ASHMEAD (HYMENOPTERA: 
SCELIONIDAE) FROM SOUTH-EASTERN AUSTRALIA 


BY A. D. AUSTIN 


Summary 


Five species of Ceratobaeus Ashmead that were previously ill-defined are redescribed to facilitate 
their accurate identification. Types are designated for Ceratobaeus clubionus Austin, C. 
cuspicornutus Austin and C. masneri Austin. Ceratobaeus intrudae sp. nov., C. platycornutus sp. 
nov. and C. rieki sp. nov. are described and the male of C. setosus Dodd is recorded for the first 
time. Notes on the diagnosis, biology and distribution of the above species, and some general 
comments on the genus in Australia, are also provided. 


SPECIES OF CERATORARUS ASHMEAD (HYMENOPTERA: SCELIONIDAE) 
VYROM SOUTH-EASTERN AUSTRALIA 


by A. D. AusTIN® 


Summary 


Austin, A, D, (1984) Species of Cerqabacus Ashmead (Hymenoptera: Scelionidae) from 
South Eastern Australia, Trans, R. Soe, S, Aust. 108(1), 21-34, 12 June, 1984, 


Five species of Ceratobaens Ashmead that were previously ill-defined are redescribed to 
facilitate their accurate identification, Types are designated for Ceratobacus clubionus Austin, 
C. cuspicornutuy Austin and QC. masneri Austin, Ceratobaens intrudae sp.nov., C. platycornutus 
sptov, and C. ¢leki sp.nov. are described and the male of Cl. setosus Dodd is recorded for the 
first time. Notes on the diagnosis, biology and distribution of the nbove species, and some 
general comments on the genus in Australia, are also provided, 


Kuy Worps; Ceratobacus, Scelionidac, Araneae, ovipositor, host specificity. 


Introduction 


Ceratobaeus Ashmead is a large genus con- 
taining at Icast 40 described species, They 
occur worldwide, but are most diverse in the 
tropics and subtropics. Available host records! 
suggest thal members of Ceratobaeus are ex- 
clusively parasites of spidet eggs, They display 
a high degree of host specificity and cause 
significant levels of mortality for some spiders 
(Austin, in press). 

The majority of Australian Ceératobaeuy 
were deserihed in the carly part of this cen- 
lury by A, P, Dodd (1913, 19l4a, 1904b, 
1915, 1919), mostly from material collected 
by himself and A. A, Girault along the 
Queensland coast, Subsequently,  Giraull 
(1926) described «a further species from 
Queensland, and WV. V, Hickman (1967) 
deseribed three species from Tasmania, which 
he reared from known spider hosts. Since 
then Austin (1981) has listed and placed 
the types of Australian species according to 
modern concepts, and Galloway & Austin (in 
press) have provided a diagnosis of the genus, 
along wilh a discussion of its biology, «istri- 
bution and relationships with other penera. 

The aim of the present paper is to describe 
three new species from south-easterh Austra- 
lia, and to redefine a further five species whose 
descriptions are presently inadequate, Three 


* Department of Entomology, Waite Agricultural 
Research Institute, Glen Osmond, S. Aust. 5064, 
Prexent address; Commonwealth Institule of En- 
jomology, British Moseum (Naturyl History), 
Cromwell Road, London SW7 SBD, Englund, 

tAusuo, A, D, (1982), The biology and ecolopy 
of Clahiona species (Araneae: Clubionidne) and 
their seelionid parasitoids (Hymenoptera), Ph.D, 
thesis, University of Adelaide.. Unpubl. 


of the latter species (C. elubionas, C. cuspicer- 
metus. and C, masneri) were referred to hy 
Austin (1983) in a study of ovipositor 
mechanics of Ceratobaeus and related genera. 
Although he states that the names adapted are 
manuscript names only, the information pre- 
sented in that paper constitutes their valid 
description under the International Code for 
Zoological Nomenclature (ICZN Artrcles 11, 
13). However, the description of these species, 
along with those of C. lamponae (Hickman) 
and C. sefasus Dadd, are inadequate in that 
they do not separate the species concerned 
from other species of Ceratobacys, The status 
of these eight species is clarified here so that 
information from concurrent studies on their 
ecolory (Austin mm press; in prep.) can be pub- 
lished free of any taxonomle problems, Notes 
are provided on their distribution, hosts and 
relationships with other species. Additional 
diagnostic characters and general comments 
for the genus are also provided, which supple- 
ment those in Galloway & Austin (in press). 


Methods 


Speeimern collection and preparation. Live 
wasps were obtained by rearing them (rom eggs 
of host spiders collected from the ficld. This 
provided valuahle supportive information on 
host relationships and general biology. Other 
material was. collected using yellow pan-traps 
and sweep-netting. 

Specimens examined under SEM were pre- 
pared as Follows: fresh material was. killed, 
washed in 70% ethanol (5 min.). transferred 
to 100% ethanol (2 min.), and placed on a 
glass slide in a desiccator to dry (24 hr), Dry 
material (museum specimens} was softened in 


22 A. D. AUSTIN 


10% ethanol (30 min.) and treated as above. 
Specimens were then mounted on holders with 
electroconductive glue (DAG 915 silver paint) 
or double-sided adhesive tape, couted with 10 
nm cucbo and 30 om gold-palladium, and 
viewed under an ETEC Autoscan SEM 
operated at 5-20 keV. Some specimens were 
examined uncosted using the SEM in the 


Environmental Chamber Mode (Robinson 
l9RO), 
Material examined: The twpes af species 


described here, with the exception of C. ricki, 
were teared Troi a single host exe sae to 
ensure that the scxes of cach species could be 
assochited accurately (they are likely to be the 
offspring of one female), Males of some Cera- 
tobaens species are morphologically very 
similar and ore difficult to separate. Therefore, 
the material listed in “Non-type specimens 
examined” has nat been included in the type 
series in case males of other species have 
been wrongly associated. However, the chanecs 
of this having happened ore sight, as there 
appears to be only @ fow species that attack 
hosts inhabiting the bark of cucalypt trees, 
the habitat from where most of the present 
species were collected. In most cases only one 
paratype female and male were coated for 
SEM: holotypes and allotypes were mounted 
on card-points so as to retain their colour. 

Terminology:  Morphotugical terms ised 
throughout this paper are defined im Masner 
(1980) and Galloway & Austin (in press)- 


Abbreviations: ANIC = Australiin National 
Insect Collection, CSTRO, Canberra; BMNH 
— British Museum (Natural History), London; 
CNC © Canadian National Collection of In- 
sects, Arachnids and Nematodes, Ottawa; 
ODP! — Queensland Department of Primary 
Industries, Brisbane; SAM = South Austr 
litn Museam, Adelaide; WAITE Waite 
Agricultnral Research Institute, Adelaide: 
A.D.A. — specimens collected by A. D. 
Austin; Caringbah — a suburb of Sydney; 
Mylor = study site, 3 km south of Mylor, 
South Australia; H > height: 1 — leneth, W 
= width; S — metasomal sternite: T — meta- 
somal tergite. 


Genus CERATOBAEUS Ashmead 
Ceratebaens Ashmead 1895, pp. 167, 175) Kielier 
1826, p. 139; Masner 1976, p. 65; Huggert 
1979, p. 7; Austin (981, p, 83; Galloway & 
Austin Cin press). 
Type-spesies, Cerarobaeus cornitie Astinead 


Diagnosis: The genus has been adequately 
diagnosed in Masner (1976) and Galloway & 
Austin {in press); however, same additional 
useful characters are given below. 

Head viewed anteriorly usually iriengulsr 
ar subtriangular in shape, sometinies with 
genae prolonged so as to be conical in shape 
towards the mandibles (similar to the condi- 
tion in Odonracalus Kielfer). 

Forewings. Venation clearly delineated though 
sometimes only lightly coloured, with at Teast 


submirginal, marginal and stignmal veins 
present. 
Metasoma., T7 external in) fernale, usually 


Luanpular in shape, always visible when viewed 
from above; ovipasitor held internally, extend- 
ing to the apical end of horn on TI (there- 
fore as long as metasoma). 


Comments: Of the gencra comprising, the tribe 
Idrini (see Austin 1981). Ceratahaens is most 
closely related to J/dris. Hugeert (1979) has 
proposed that Ceratohaeus should be included 
under /dris os a subgenus, due lo the apparent 
existence of intermediate forms. However, 
Austin (1981) maintains that they should be 
kept separate due to differences in the mor- 
phology of the metasoma and posterior meso- 
soma. fyris differs from Ceratebacus in that 
TI i always flat, the metasoma is never 
elongated, the propodcum is net excavated, 
and the propodesl dise is mot divided inte 
laminae, Also the length of the ovipositor 
differs between these two penera. Ceratobueus 
species have very Inng ovipositors that are 
held internally within the octasoraa and 
extend into the anteriorly projecting horn of 
TI. ddérix species. however, have relatively 
short internal ovipositers that do. not reach 
past the posterior margin of T2) (Austin 
1983), 

The length of the metasoma and harn vary 
Substantially between species of Ceratobaeny, 
but remain constant within a species. In some 
species the horn is represented by a small 
hunip, while in others it reaches dorsally above 
the mesosoma. The various stages in horn and 
metasonial lengih hetween these two extremes 
results in there heing a corresponding series in 
the length of the ovipositor. Presumably such 
differences have evolved in response to vary- 
ing accessibility of bost eggs. Ovipositor length 
is well recognised as a factor mvolved in deter~ 
mining host specificity and subsequent specia- 
tion within Che parasitic Ffymenoptera (ec. 
Askew 1971; Gibbons 1979; Henthcote & 


CERATOBAEUS ASHMEAD FROM 8.B. AUSTRALIA 23 


Davis 1976; Price 1972), and undoubtedly a 
similar mechanism has operated in Cerato- 
baens. 

The morphology of the posterior mesosoma 
is Closely tied to the size of the horn: species 
with large horns have the propodeum and 
sculellum more excavated than in species with 
short horns, Although males of Ceratobyeus 
have at most only a slight dorsal expansion 
on T1, many species still display some exeava- 
tion of the posterior mesosoma. This excava- 
tion is always more pronounced in males of 
specics where the female has a long horn 
(eg. C. cuspicornutns, fig. 12)- 


Figs 1-6. Antennue. 1. Ceratobaeus cuspicornutus, 
?, 2, C. cuspicernutus, 3. 3, C. maseerl, 4 
€, masnert, & 5, C. platveornutas, &. 6. C. 
patycornutus, J. Pigs 7, 8. Forewings. 7. ©. cus- 
picornutds, 2. 8, C. masnert, 2. Scale lings. = 
100 @m, N.B, hairs and cilia not drawn on 
anlennae and wings. 


Ceratobaeus chthionus Austin 
FIGS 9-11, 15, 17, 18 


Cerdiohaens clubionus Austin 1983, p. 151 {no 
types designated), 


Types (by present designation): holotype 9, ANIC, 
South Ausiralia: Mylor, 29.317.1979, A.D.A., ex 
ege Clubiony sp. (Araneac). Paratypes reared 
from same egg sac as holotype—Allotype 3d, 
ANIC; Paratypes 2¢, 29, gold coated on SEM 
holders, antennae and wings of 14, 19 on slides, 
Id, 49, ANIC; Id, 49, CNC, 13, 49, QDPT; 
id, 42. SAM; 1d, 49 WATTE. 


Female 


Length 1.3-1.4 mm, Colour, Head and 
mesosoma shiny black; antennae and legs light 
brown, almost yellow; antennal clubs and 
femora slightly darker; metasoma dark brown 
with lighter margin; posterior Tl and anterior 
T2 light brown. 


Head, L:W:H (6.5:19:14), with granulate 
sculpturing and fine scattered hairs; dorsally. 
wider than mesosama and arehed around 
pronotum; occipital carina sharp; eyes large 
and hairy; lateral ocelli touching inner 
margins of eyes; frons slightly curved; 
anteriorly, head ovoid; occiput arched; cyes 
separated by 0.5 ™ width of head: frons 
smooth; frontal carina weakly developed; 
laterally, gena with sides not quite parallel; 
antenna (fig. 17), club with 3 faint incom- 
plete sutures. 


Mesosoma. Dorsally, with coriaceous sculptur- 
ing and sparse short hairs (fig. 9); pronotum 
nat visible; scutum wider than long, L:W (11: 
14); notauli absent; scutellum almost semi- 
circular, L:W (5.5:11), posterior margin with 
wide flange extending over metanotum, slightly 
inflected medially; metanotum narrow and 
crenulated} propodeum vertical and smooth, 
Jaminae diverging ventrally, extended into 2 
small teeth dorsally Cig. 10). 


Forewings not quite reaching to posterior 
margin of metasoma, not particularly broad. 
L:W (36:13); venation distinct, marginal and 
postmarginal veins short; stigmal vein long, 
basal vein present but lighter than other 
veins; lightly infuscated around apex of stigmal 
vein; marginal fringe of hairs moderately long 
(fiz. 15). 


Metasoma wider than mesosoma, L:W (30: 
17), sparsely covered with hairs, pointed pos- 
teriorly; anterior Tl expanded into large 
hump, not reaching above propodeum (fig. 
10); Tl and T2 with coarse Jongitudinal 
striations, T3 with lighter striations; lateral 
margins of T2-T3 and all T4-T6 with granu- 
late sculpturing. 


A, D. AUSTIN 


Figs 9-11. Ceratobaeus clubionus. 9. 2, lateral view 
of mesosoma and TI. 11. 3, dorsoposterior view 
nutus, 12. 3, dorsal view of mesosoma and TI. 
C. intrudae, 3, lateral view of whole body. Wings 


of head and mesosoma. 10. 9, dorsolateral view 
of mesosoma and Tl. Figs 12, 13. C. euspicor- 
13. 9, lateral view of mesosoma and TI. Fig. 14. 
removed on all specimens. Scale lines 100 «m. 


CERATORAEUS ASHMEAD FROM $8.6. AUSTRALIA 25 


Male 

Differing From female in the following: 
length 1.2-1.3 mm; antenia (fig. 18); pro- 
podeal laminar diverging stightly more than 
in female; wings reaching well past posterior 
margin of metasoma; forewing L:W (46-19), 
marginal fringe of hairs long, venation dark, 
basal vein darker than in female; metasoma 
rounded posteriorly, wider than mesosama, 
LiW (26:18); anterior TI inflected dorsally 
into hump, but not as pronounced asin female 
(fix 11), 


Comnients 


C. clubionus parasilises the eggs of € Jubinna 
eyveladate Simon and an undescribed species 
of Cluhiona (Clubtonidae). These spiders 
inhabit the bark of eucalypt trees in the 
Mount Lofty Ranges, South Australia. This 
Species can be distinguished from all other 
Ceratohaeus by its colour, the presence of a 
postmarginal vein, and the hor on TI repre- 
sented by only an anterior dorsal inflection of 
that tergite, jc, not rounded dorsally, 


Non-type specimens examined: SOUTH AUSTRA- 
LIA: 28, 69, ISL IIFS, 2d. 39, 18.41,1979, 
Mylor, A.D.A. Id) 79, Myponea, 441.1979, 
A.DLA. SL m09, S4i1979 Tey 489, ATi ae79, 
Strathalbyn, A.D.A, ANIC: 4c, 279, Mylor, 1979, 
ADA: 1d) 1% Myponga, 447.1979, ADA, 
BMNH; 89, (Saii-1979, Fay 149, 13.v.1979. 
Mylot, A.D.A, CNC, Id. 4%, PRL 1979, 59, 
Dii.1979, 1h, SP, 25.xL 1979, 1d) 59, Vxliay79, 
Mylor, A.D.A., ODPL; 10%, 59, 13.iv.1979, 24, 123, 
207,0980, Mylor, A,D.Ay SAM, 112) 234 L979. 
2, BS. 130.1979, Mylor. A,D.A., WATER, 


Ceratobeeus cuspicerntutus Austits 
FIGS 1, 2, 7, 12, 13 
Ceratobaens euspicornatis Austiq 1983, p, 141 

(not types <lesignated), 

Types (by present designation): holotype 2, 
ANIC. South Australia: Mytor, 18,4.1979, AAA, 
ea gee Olubiana sp, (Aranvae). Paratypes feared 
from same egg sac as haletype—Allelype 4. 
ANIC; Paratypes 13, 29, gold coated an SEM 
tioldeys, antennae and wines of 4 am) 1? on 
slides. 22 dissected on slides, 24, 29, AWIC> If. 
22, CNC; Id, 29, ODPI: If 29. SAM: It. 
29, WAITE, 


Female 


Length 1.50-1.75 mm. Colour. Head and 
mesosoma black; lets and antennae brown ta 
dark brown; metasomal horn black and 
shiny: posterior T] light hrown; T2—T7 brown, 
Head wider than mesosoma, L1W:H £6219: 


15), with granulate sculpturing and short 
scattered hairs, dorsally, occipital curina 
sharp; eyes large, with very short hairs; lateral 
ocelli touching inner margins of eyes; frons 
slightly curved; anteriorly, head subtriangular 
in shape; occiput slightly depressed medially; 
eves separated by slightly more than 0.5 
width of head; lower frons smooth and shiny; 
frontal carina weakly developed; Jaterally, 
ena large, margins parallel; antenna (fig. 1), 
club with 3 faint incomplcte sutures, 
Mesosoma strongly compressed anteriopos- 
teriorly; dorsally, with granulate sculpturing 
and scattered hairs; pronotum not visible; 
scutum much wider than lang, L:W (9:14); 
notauli absent; scutellum transverse, L:W 
{t:10), with posterior fringe of long hairs: 
seutellum, metancium and propodeum 
strongly excavated to receive horn, sloping 
away posteriorly towanls metasoma: propadeal 
laminae parallel, not extended dorsally into 
small tecth (fig; 13); laterally mesosoma 
smooth and shiny, 

Forewings not reaching past posterior margin 
of ‘4, fairly narrow, L:W (44:13); marginal 
vein short, stigmal vein long, postmarginal 
vein less than 0.5 ™ Iength of stigmal vein; 
marginal fringe of hairs moderately long (fig, 
7) 

Metasoma clongated, L:W including horn 
(35:17), slightly wider than mesosoma, with 
scattered hairs; horn long. angled forward into 
mMesosqmal cavity, reaching above level of 
scutellum; apical horn moderately pointed 
and smooth, striate laterally and basally (fig. 
13) base of T1-T4 with longitudinal striations; 
lateral margins of T2-T3 and all T4-T? with 
granulate sculpturing: T7 slightly clongated. 


Male 


Differing from female in the followjne: 
length 1.35-1.50 mm; antenna (fig. 2): dorsal 
mesosoma arched more than in female; 
scutellum more than 2 % wider than long, 
L;W (5:12), posterior margin rounded, in- 
flected medially into small smooth triangular 
patch; posterior mesosoma not excavated but 
flai, sloping posteriorly towards metasoma; 
metanotum narrow and crenulated, visible 
fram above; propodeum smooth; propodeal 
laminae diverging ventrally (fig. 12); wings 
reaching well past posterior metasoma; fore- 
wings moderately broad, L:W (55:20), basal 
vein darker than in female, but still faint eom- 
pared to submarginal vei, marginal fringe of 


2H AOD 


hairs long; metusama moderately clonpiuted, 
rounded posteriorly, LzW (29:16); anterior 
Tl inflected dorsally, striations reaching to 
auierior margin of TI. 


Comments 

This species parasilises the eggs of Cluhiona 
eveladata and an undescribed species of Cli- 
biena (Clubionidae). These spiders inhabit 
the bark of ecuealypt trees in the Mount 
Lofty Ranges, South Australia. C, cuspicor- 
auius 1s related to a group of species that all 
have a long slightly curved horn and an elon- 
gate metasoma. It comes closest to C Jongi- 
cornutis Dodd, but differs in having a darkly 
piuinented basal vein, a brown metasoma (not 
yellow) and a more elongate aatennal pedicel, 
Noa-tipe specimens examined: SOUTH AUSTRA- 
LIA: 34, 9P, Di7.1979, 102, 13.x1.1979, Mylor, 
ADA, ANIC: 2d: 69, Mylor, 27.xii.1980, 
A.D.A. BMNH: 24, 49. Mylor, 15.16.1979, 
A.D.A,, CNC: 1d. 69, Mylor, 13.x1.1979, A.DIA. 
QMPl 24, 59. Mylor, 9.ix.1979, A.D.A. SAM; 
29, 294i.1979, 39, 26.41.1978, Mylor, A.D.A,. 
WALTE. 


Ceratobaeus intrudae sp, nov. 
FIGS 14, 16, 19-22 

Types: holotype & ANIC, South Australias Mt 
Campass, 441979, A.D.A,. ex egg fatrndia sp 
(Araneae). Pavatypes reared fram same eg suc 
as holotype—Allotype 3, ANIC; Paratyjes dat, 12, 
gold coated on SEM holders, antennae and wings 
on slides. 24. 79. dissected on slides. ANIC: |. 
12, CNC. 

Female 


Length 1.30-1.45 mm. Colour. Head and 
dorsal mesosoma very dark brown to black: 
legs, antennae and lateral mesosoma dark 
brown: metasama with a light brown band 
behind horn and a medial brown patch 
becupying approximately two-thirds pF Ta, 
rest of metasama dark brown to black. 

Head wider than mesosoma, not strongly 
curved around pronotum, with granulate 
sculpturing and sparsé short hairs, L:W:H 
(7.5cE8c13): dorsally. occipital carina shurp: 
eyes large, covered with short hairs; lateral 
ocelli touching inner margins of eyes: frons 
slightly curved; anteriorly, head subtriangular 
in shape: occiput flat; eyes separated by more 
than 0.5 & width of head: frons lightly sculp- 
ured: frontal carina well developed, reaching 
hall way to medtan ocellus; laterally, gena 
large, margins almost parallel; antenna (fig. 
19), club with 3 faint ineamplete sutures, 


AUSLIN 


Fig. 15. Ceratebacus clabionns, 3, dorsal Surface 
of whole body. Fig. 16. C. inrrauddiac. 9, dorsil 
surfiice of whole body. Figs 17-20, Antennae. 
17, C. clubionus, 3, 18. C. clubionus, 3.19, €. 
tatrndde, Y. 20. C. intrutdae, & Scale lines = 
100 @m. N.B. hairs and cilia not drawn on 
antennae and wings, 


Mesosoma. Dorsally, fairly flat, with granu- 
late sculpluring and scattered hairs; pronotum 
visible at anterior corners; scutum wider than 
long, L:W (10:14.5); notault absent; seu- 
tellum almost semi-circular, L:W (4:11), 
Posterior border crenulated, with a narrow 
flunge (fig, 22): metanotum narrow and 
crenulatcd; propodeum yertical and smvoth; 
laminae curving dorsally. extended into 2 
small teeth (fig. 21); laterally, mesosoma 
smooth and shiny. 


Forewings narrow, not quite reaching 10 
posterior margin of metasoma, L:W (38: 
12.5); marginal vein short, stigma! vein long, 
postmarginal vein as long as stigmal yein, basal 
vein present but very faint; infuscated around 


CERATOSALUS ASHMEAD FROM $.E. AUSTRALIA 


apex of stigmal vein; marginal fringe of hairs 
short (fig, 16). 

Metusoma slightly wider than  mesosoma 
aid 2 longer than wide, L:W (32:16), with 
scatiered hairs, pomted posteriorly: horn 
ulmost vertical, just reaching above level of 
posterior scutellum, with circular striated 
sculpluring: apically (figs, 21 and 22); TL 
behind horn and ‘T2-T3 with longitudinal 
striatianss lateral T2-T3 and all T4-T6 with 
pranubite sculpturing. 


Mietle 


Dillering from the female in the following: 
dorsally, head slightly more curved around 
pronotum, only slightly wider than mesosoma, 
L:W:tl (7,5:17:13); antenna (fig, 20); pro- 
podeum not quite vertical, sloping away 
slightly towards metasomays, propodeal laminae 
slightly wider than in female; wings reaching 
well past posterior metasoma: forewing L:W 
(42:16), marginal Fringe of hairs Jong; meta- 
soma rounded posteriorly, L:W (25:16); 
anterior TL strongly inflected dorsally, not 
reaching above propadeal Jaminae: without 
striate or punctate sculpturing, but with longi. 
tudinal striations extending to anterior margin 
(fig. 14), 


Conments 


C. inirudae parasitises the eggs of an une 
known species of Intruda (Gnaphosidac), the 
falter being found under the bark of eucalypt 
trees ju the Mount Lofty Ranges, South Aus- 
tralia, This species is related to a large yroup, 
all with moderately short horns, However, C. 
intrudae can be distinguished from all other 
species by the following combination of 
characters: apical horn with circular. striac, 
Inglasoma twice as long as wide, colour as in 
deseription. 


Nielype specimens examined SOASTIL AUSTRA. 
LIA: [e, 39, Bridpewaer 26.47.1979, ADA, 
ANIC) Te, 49. Bridgewuter. 2671-197, ALDLAL, 
SAM. 


Crratohdcns famponae (Hickman 
PIGS 23-25 
Odantacalus lanjponee Hickman 1967, py 1s. 
Ceralobacus lampanar (Hicktnan); Musner 1976, 
p. 46; Austin 1981, p. Bd. 
Lypes holotype 9 on slide, ANIC: Tasmania, 
Domain, Hobart, 29.x,1966, Vo VY, Hickman, ex 
cag Lampona cylindrara (lL. Koch) fArsnene); 
Parutypes 24, 19, on sunie slide as holotype 


m4 
a 


Female 


Length 1.60-1,85. Colour, Head, mesosoma 
and horn shiny black; antennae and metasoma 
dark brown to black; legs brown, 


Read, LiW:t (9:22.5:16), with moderately 
coarse punctate-reticulate sculpturing, covered 
with Jog hairs: dorsally. wider than meso- 
soma, arched around pronotum: occiput well 
exposed; occipital carma sharp, moderately 
angled at corners: eyes large, with long hairs; 
Jateral ocelli touching inner margins of eyes; 
frons straight; anteriorly, head subtriangular 
in shape; occiput straight; eyes separated by 
slightly less than 0.5 > width of head: rons 
flat, with borizontal striac: frontal earina very 
small; laterally, gena with margins parallel, 
rounded ventrally; antennal scape L:W 
(31:6), pedicel (1325.45). elub (27:12) with 
3 faint inconsplete sutures, 

Mesosoma dorsally, with moderately coarse 
punctaly-reficulate seulpturing. sparsely 
covered with long hairs (fig. 24); pronotum 
visible at anterior lateral corners: scatum 
wider than long; L;W (£3.5:17.5); notauli 
absent; scutcllum almost 3% wider than long, 
L:W (4:12), posterior margin straight, fringe 
of long hairs projecting over horn often 
present; metanotuin and propodeum = flat; 
mefanotum narrow and erenulated; propodeal 
laminae diverging ventrally, extended dorsally 
inlo blunt teeth; lateral propodeum and meso- 
pleura margined by single rows of deep pits 
(fig, 23), 

Forewings not quite reaching to posterior 
margin of metasama, moderately broad, L.:W 
(73:27), infuscated medially; venation elear 
and dark; marginal vein short, postmarginal 
vein as long as stigmal vein, basal vein light; 
marginal fringe of hairs moderately long. 
Metausoma wider than mesosoma, L:W_ in- 
cluding horn (42:22), covered with long hairs; 
horn just Teaehing to devel of dorsal scutellum, 
apical two-thirds with coriaceous sculpturing 
winost scaly in appearance (fig, 23); TI in- 
cluding basal one-third of horn and 'T2 with 
lungitudinal striations, 13 strigose with back- 
ground granulate seulpturing, T4-To with 
granulate sculpturing, 


Male 


Differing from female in the followiny: 
length §=1.55-1.70 mm; dorsally, seutellum 
more rounded, though slightly fattened pos- 
teriorly, 2 ™ wider than Jong, b:W (6.5: 


AUSTIN 


A. D. 


CERATORAEUS ASAMEAD FRUM SB. AUSTRALIA 29 


12.5); propodeum flat, almost vertical, propo- 
deal laminae wide, with coarse striate sculp- 
tiring, strongly diverging ventrally, extended 
Wty 2 blunt teeth dorsally which almost 
touch medially (lig. 25); wings reaching well 
past posterior Metasoma; forewing L:W (83; 
32), well infuseated, marginal frinve of hairs 
Jong; metasomu subpedunculate, wider than 
mesosoma, with scattered long hairs, L.sWw 
(35:25), anterior TI inflected dorsally: T1-T2 
and anterior ‘T3 with longitudinal striations, 
Jateral T2 and rest of metasoma with wranu- 
late sculpturing, 


Camanimnts 


C. lampenae (Hickman) parasitises the eggs 

of Lampana eylinedreta (1. Koch) (Gnapho- 
sidae); the latter being widespread through: 
out Australia, C. lamponae, previously only 
known from Tasmania, is recorded from main- 
land Australia (Mount Lofty Ranges, South 
Australia) for the first time, There uppears ta 
be some minor geographic variation in this 
species, Some specimens from Tasmania have 
ise posterior fringe of hairs on the scutellum 
very short or absent, while mainland specimens 
have a very long Fringe. C. lamponqe can be 
distinguished from other species by its colour, 
presence of a postmarginal vein and charac- 
teristic sculpturing on the horn and dorsal 
mesosoma, 
Non-rype speednens examined? SOULH AUSTRAS 
LIA: Ic, 19, gokl eouted on SEM holifers, 
dntennue und wings on slides, 24, 11°, Mylor- 
29.4,1979, ADA, ANIC; 63, 109, Mtytor. 
20.180, Mylor, A.D.A, BMNH; 43, 89, Mylar, 
20,7.1980, A.D.A, CNC; Id, 119, Mylor, 29.7. 
1979, A.D,A, QDPI; 42, 69. Mylor, 20..2980, 
A,D.AG SAM; Ie, 6? Mylor, 14311979, ADLAL 
WAITB. TASMANIA; 14, 115, Domain, Hobart, 
Q11.1967, Vo WV, Hickman, ANICS 3a, 19°. 
Domain, Hobart, 29.57, 1967. Vo V, Hiekrran, 
ONC, 


Ceratataens anaynert Austin 
FIGS 3, 4, 8, 26, 27 
Ceratobhaces mesxeteri Austin 1983, p. Ma te 
types designated). 
Types (by present designation): holovype Y ANIC. 
South Australias Mytor, 18411979, A.D.A. ex 
egg Clubiona sp. (Araneae). Paratypes reared 


Figs 21, 22. Ceratobuens intrudae, 9. 21. Tateral view. of mesosoma und TI. 22. 
mesosuma and Tl Figs 23-25. CL funponae. 23. 2, 


from same ege sac us hololype. -Allotype 
ANIC: Parutypes 24) 3, gold couted on SEM 
holders, untennize and wings of td, 12 ow slides, 


4d, 49, ANIC: Ie, 49. CNC; Tay dy, ODPE: 
13, 49, SAM: Id. 49, WATTE, 

Female 

Length 1,25-1,40 mm. Colour. Head and 


mesosomma black; antennae and legs brown; 
metasoma dark brown; TI light brown, but 
With apex of horn brown to dark brown. 
Head wider than mesosomu. L:W:H (7: 
18:13), urched ground pronotum, with granu- 
late sculpturing and seattered short hairs: 
dorsally, accipital carina sharp; eyes large and 
hairy; lateral ocelli touching inner margins of 
eyes; frons curved; anteriorly, head subtrian- 
gular in shape; occiput curved: eyes separated 
by slightly more than 0.5 % width of head; 
frons smooth, frontal carina weakly developed, 
reaching 0,5 3 distance to median ocellus: 
laterally, gena wide, sides not parallel: antenna 
(fig. 3), 


Mesosoma, Dorsally, with granulate seulp- 
tuniy and scattered hairs; pronotum not 
visible; scutum wider that long, LW (9:12); 
notauli absent; scutellum Lrw (4.5110), 
rounded posteriorly, with crenulated border, 
slightly inflected medially; metanotum narrow 
and ¢renulated; propodeum vertical and 
smooth; laminae diverging ventrally and 
curved dorsally into 2 small teeth, Jaterally 
with coarse striations: (fig. 26). 


Forewings just teaching to posterior margin 
of metasoma, fairly narrow, L:W (40:13); 
marginal vein short, stigmal veiti long, post 
marginal vein approximately 0,75 ™ length 
of stigmal vein, basal vein present but faint: 
marginal fringe of hairs short (fig. §), 


Metasoma wider than mesosoma, oearly 2 
* longer than wide, LeW (35:19), pointed 
posteriorly, sparsely covered with hairs; 'TI 
expanded into a small dorsal horn, not reach- 
ing to level of scutellum (fig, 26); apex of 
horn with faint punciale sculpturings most uf 
hort, posterior TL and V2-1'3 with lonwi- 
tudinal striations; lateral margins of T2, biack- 
ground of T3 and all T4-T6 with granulale 
sculpturing. 


; Dorant view of 
lateral view of mesosoma and Tl 24, ©, 


dorsal view of mesosoma, 25 cl, lateral view of mesosoma and Tl. Figs. 26, 27. ©, puisncis, 26, 
9, dorsolateral yiew of mesosamu and T} 27. J, dorsolateral view of mesoxoma and 7). Big. 28. 
C. platycernutns, 9. dorsoposierion view of mesosanm and Th Wings removed on all specimens, 


Scale lines — 100 am, 


M) A. DB. AUSTIN 


Male 


Dillering (rom female in’ the following: 
Jength 1.20-1.38 mm, antenna Cig. 4); bares 
on dorsal surface of mesosoma slightly longer 
than in female; sculellum oslightly arched 
dorsally; lateral propodcum with fine sparse 
striations; wings teaching well past posterior 
margin of metasoma; forewing L:W (49:18), 
venation same as female, hut with basal vein 
more obvious, almost as dark as submarginal 
vein, marginal fringe of hairs long: metasoma 
tounded posteriorly, L:W (27:16,5): anterior 
TI inflected dorsally, only reaching to 0.5 8 
height of propodeum, striations reaching to 
anterior margin af T) (fig, 27), 


Conunents 


C’. imasneri parasitises the eggs of Cluhjand 

robusta L. Koch, Cl. cycladata, an undescribed 
species of Clubiona (Clubionidae) and Mesmi- 
cloea sp. (Gnaphosidae): all inhabit the bark 
of cticalypt trees throughout south-eastern 
Australia, So far C. mastert has been collected 
from locations in South Auystrutia, Victoria 
and the Australian Capital Territory, “This 
species belongs to a laree group that all have 
short horns. However, ©. orasner? can be 
recognised from all other speetes by the 
presence of a postmarginal vein. a darkly 
pigmented basal vein, and tts colour (see 
description), 
Nan-ipe specinwns examined: AUSTRALIAN 
CAPITAL TERRITORY: 69. Cunberra, 14.1. 
1980, A.\D.A, ANIC. SOUTH AUSTRALLA: 139. 
241979, ex eges Clahbiond sp. (Araneae). fel, 
142, 13.x0, 1979 eX engs Memicloca sp, (Araneae). 
Mylor. A.D.A, ANTC: We. 179. Mylor, 197k 
80, A.D.A.. 19, Myponga, 4.161979, ADA, 74 
339, Mylor, LOW1980, A.D es cans Heri. 
cloen 5p, CNC: do, 69, LN1979, 2, 6. 20.4, 
1980. Mylar, A.D.A. QDPT; Sa. 69, Mylor. 
ITAi 1980, ADA, SAM: 72 bSiw 979, 110, 
12.40i.1979, Mylor, A.DLA., WATTE YICTORIA; 
14, 99, Woordon, 26,ix.1979, ATA, SAM, 


Ceratobaeus platycormutus sp. nov. 
FIGS 5, 6, 28. 29 
Types: holoiype YY ANIC. Australian Cypital 
Temilory: University campus, Canberra, 144-1980, 
A.DLA.. ex eae Cluhiana sp. CAraneae). Paratypes 


reared from sume veg suc as bololype—Allotype 
oy ANIC: Paratypes Iq, 29. gold coated on SEM 
holders, antennae dnd wings of let, LE on slides, 
24 79, ANICS Id. 49. CNC, Te, 42, QDPI: 
1a, 49, WATTR. 


Female 

Length 1.75-1,85 mm. Colour, Head, meso- 
some and apical horn black; antenmac, legs ain 
metasoma dark browits) anterior metasama 
behind horn, with a light brown band. 

Head wider than mesosoma, L:W:tt (7: 
20:15), arched around pronotum, with granu- 
late sculpturing, sparsely covered with hairs; 
dorsally, occipital carina sharp; eyes large. with 
fine hairs; lateral Ocelli touching inner margins 
of eyes: frons curved: anteriorly, head sub- 
Irmngular in shape; oeciput straight: eyes 
separated hy OS & wirlth of head; frons lightly 
punctate; frontal carina very small, laterally. 
gena wide, sides almost parallel; antenna (is. 
5), 

Mesosoma strongly compressed — anterio- 
posteriorly; dorsally, with granulate sculpturing 
und scattered hairs; pronotum not visible: 
scutumy wider than long, L:W (10:15); nataul 
ubsent; scutellum transverse, Pew (1:t0). 
seutellunt, metanotunt and = propodeum all 
strongly indented posteriorly, sloping towards 
metasoma; propadeal laminae vertical, curved 
outwards ventrally, without dorsal teeth (fig. 
28). 

Forewings teaching to pesterior margin of 
T4, fairly narrow, LiW (71:23); lightly in- 
fuscated medially; marginal vein short, styymal 
vein long, postmarginoal vein less than 0.35 
length of stigmal vein, basal vein light; mar- 
ginal fringe of heirs moderately long, 
Metasoma clongated, L:sW= including horn 
(56:18). wider than mesosoma; horn long, 
slightly arched, projecting anteriorly, closely 
fitting to mesosoma, reaching above level of 
scutcllum. flattened aud dise-shaped apically 
(fig, 28); TL except for apical portion of horn 
and T2-T4 with fonvitudinal striations; lateral 
T2-T4 and all TS-T7 with tight granulate 
sculpturing. 


Afale 
Differing from female in the following: 
length 1.4-1.5 mm; antennae and legs yellow; 


” 


Fio, 29. Ceratebacny platyearintus. 


dorsolateral view of mesosoma and ‘Tl Pig. 30, C. 


rieki, ‘?. 


dorsoliteral view of howd. mesosoma sind Th Bigs 31-34. Cerarabacny setasus. 31. 2. umterion 
view of head, iaennue missing. 32. 44, dorsolaternl! view of mesosoma. 33. 9, lateral view of 
mesosoma und Th 34, of, daterl Siew Of miesusoma and Th Wings removed oy all specinens 


excepl on one side of specimens in Pips, 32 and 33, Seale lines 


100 fam 


AUSTRALI 


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ad A. D. AUSTIN 


metasoma dark brown to black, with light 
brown band anteriorly, antenna (fig. 6); head 
not as high, LsW:H (7!20:13.59)) scutellum 
almost semicircular, slightly indented pas- 
teriorly to form smooth triangular patch; 
metanotum and propodeum flattened and 
smaoth; propodeal laminae diverging ventrally; 
wings reaching well past metasoma; forewings 
broad, L:W (80:31), postmarginal vein not as 
long as in female, basal vein slightly darker, 
marginal fringe of hairs long; metasoma not 
as clongated, L:W (30:20); anterior TI in- 
Nected dorsally (fig. 29); Tt-T3 with longi- 
tudinal striations; rest of metasoma with light 
punctate sculpturing. 


Cenunents 

The specific name platycornutis (platys in 
Latin, broad or flat: cornutits, horn) refers to 
the tattened, disc-shaped apical end of the 
metasomal horn. ‘This species has been 
recorded as parasitising the eggs of an unde- 
scribed species of Clubionw (Clubionidae) 
inhabiting bark of cucalypt trees in the Aus- 
tralian Capital Territory. C. platycarnitus is 
very similar to cuspieornutuy and /ongicor- 
nuts, but differs in the shape of the meta- 
soma hom (see comments under cuypiear- 
AUIS), 
Nomrvpe specimens examined: AUSTRALIAN 
CAPITAL TERRITORY: 43, 119. University 
campus, Canberra, 10.7.1980, A.D.A,, ANIC. 


Ceratobacus rieki sp. nov. 
FIGS 30, 35, 39 

Types: holotype 9, ANIC. New South Wales: 
10 mb EB. Trangie, 20.x.1999, FB. F, Riek. Para- 
types 19, gold coated on SEM holdet (mesosoma 
missing), wines on slide, 12 dissected on slide, 
135, ANIC; same duta as holotype. 
Memale 
Length 21-23 mm, Colour. Head and 
mesosoma black; antennae dark brown; legs 
brown; metasoma dark brawn to black. 
Head. L:W:H (7:20:14.5), with granulate 
sculpturing covered with short hairs; dorsiilly, 
slightly wider than mesosoma: occiput exca- 
vated and arched around pronotum; occipital 
earina sharp, not angled at corners; cyes large 
and hairless: lateral ocelli touching inner 
margins of eyes; frons slightly arched; ante- 
riotly, head subtriangular; occiput slightly 
depressed medially; eyes separated by slighily 
more thin 0.5 % width of head; trons smooth: 
frontal carina not developed: laterally, gena 
with margins parallel; antenna (fig. 39), club 
large with 3 faint incomplete sutures. 


Mesosoma strongly compressed anteriopos- 
teriorly, posterior surface sloping towards 
metasoma; dorsally, with punctate-granulate 
sculpluring, covered with short hairs; pronotum 
not visible; scutum 2 ™* wider than Jong, L:W 


Fig. 35. Ceratohacus ricki, 9, 


whole hadyv, Big, 36..C. setests, 2, dorsal surface 
of whole body, Figs 37-39 Antenmaic., 37. C. 


dorsal surface of 


setosus, ¢. 38. CL setosux, 2. 39. C0. ricki, P. 
Scale lines -- LOO som, N.B. hairs and cilia not 
drawn on antennae und wings. 


(8:16); notauli absent: scutellum transverse. 
L:W (1:10) with posterior Fringe of long 
hairs; scutellum, metanotum and propodeum 
strongly excavated posteriorly for reception of 
horn; propodeal laminae vertical, without 
teeth dorsally (figs. 30 and 35). 

Forewings fairly marrow, L:W (47:16), not 
reaching past T4, not infuscated; venation 
distinct, marginal and postmarginal veins 


OERATOBAELS ASHMEAD FROM 5.6, AUSTRALIA i 


short, stizma) veitt long, basal vein present but 
faint; marginal fringe of hairs moderately long 
(hig. 35) 

Metasoma extremely clongated, approximately 
4 “ longer than head and mesosoma com- 
bined, 6 x longer than wide, L:W (90:15), 
covered with short hairs; horn long, strongly 
arched anteriorly, cylindrical but slightly flat- 
tencd apically (fg. 30); TI-VS including basal 
hall of horn with longitudinal striations; lateral 
Margins of T2-13, and all T4-T7 with granu 
late sculpturing. 


Male unknown, 

Host unknown, 

Noretype ypeciniens examined: QUEENSLAND: 
Brisbane, Indooroopilly, 14x76, Boutek, 
BMNH. 


Comments 

[his species is named after ils collecter, Dr 
EB. BF. Riek. ©. niedé is the largest species of 
Crratobaeus so far recorded in the world; its 
clongate inetusoma being unlike that of any 
other species. 


Ceratahdens. setovsas Dodd 
FIGS 31-34, 36-38 
Ceraiobaens setosus Dodd WY l4a, p. 65: Kieflet 
1926, p. 142; Austin 1981, p. 85, 
Tsipes: holotype Sy bead vod wings on slide. No, 
1966, SAM: Queensinnd. Gordonvate (Nelson), 
29.4.1913, A. P. Dodd. 


Female 


Length 1.3-t.5 mm. Colour. Head and meso- 
soma black: antennae and legs brown to dark 
brown; dorsal metasoma dark brown to black, 
ventral surface dark brown, 

Head, LiWrH (8.5:19:15). with granulate 
sculpluring, covered with dense mat of short 
hairs; dorsally squarish, slightly wider than 
mysosoma, not strongly excavated posteriorly; 
occipital carina sharp, not angled at corners; 
eyes Jarge, covered with hajrs; lateral ovelli 
touching inner margehs of eyes; frons almost 
straight; anteriorly, head subtriangular in 
shape; occiput arched; eyes separated by more 
than 0,5 & width of head; frons smooth and 
shiny; frontal carina very short; laterally, gena 
with margins converging, rounded ventrally 
(fig 31); antenna (fiz. 38), club with 3 faint 
Incomplete sutures, 

Mesosoma, Dorsally slightly arched, with 
granolale sculpluring, covered with dense mat 
of short hairs (fig. 32); pronotum not visible 
wf anterior lajeral corners: scutum not much 


Wider than jong, Lew (11iT4.5). noth 
absent; seutellum semicircular, LiW (5:11), 
with erenulated posterior border (fiz, 36); 
roctanotum narrow and crenulated: postertor 
surface of propodeum vertical: laminae diverg- 
ing slightly, extended into 2 small sharply 
pointed teeth dorsally (fig, 33); laterally, 
mesosoma smooth and shiny. 

Forewinys just reaching posterior margin of 
metasoma, L:w (38:15.5); marginal and 
postmarginal veins short. stigmal vein long, 
basal vein darks marginal fringe of hairs short 
(fig. 36), 

Mautasoma broad and at, wider than meso- 
soma, pointed posteriorly, L:W (30:19) (fig, 
36), covered with dense mat of short hairs; 
horn vertical, just reaching to level of scu- 
tcllum., with reneculate-rugose  sculpturing 
apically (fig. 33); base of TI-T3 with longi- 
tudinal striations; lateral margins of T2-T3 
and all T4-T6 with granulaic sculpturing, 


Male 


Differing from female in the following: 
leneth 1.2-1.3 mim; antennae and legs light 
brown, metasoma dark brown; antenna (fig, 
37); posterior mesoasoma almost indentical, 
except propodedl laminae diverging slightly 
more than in female; forewings long, reaching 
well past posterior metasoma, L:W (42:16): 
metasoma broad, slightly wider than meso- 
soma, rounded posteriorly, L:W (24:18); 
anterior T] expanded dorsally info hunip, not 
reaching above propodeum, lonwvitudinal stria- 
tions reaching to anterior margin (fig, 34); 
T2-T6 with long scatlered hairs. 


Comments 


C, setosus bss been recorded along the 
coast of Queensland and New Sauth Wales. 
In New South Wales this species has been 
reared from the eges of Jxenticus rebustuy CL. 
Koch) and /, nartius (Simon) (Armauro- 
biidae), collected From arovnad buildings anid 
under bark of cucalypl trees. Another sce. 
fionid, fdriy txeutrce/ (Hickman), has also been 
recorded as parasitising the cyes of these two 
spiders in Tasmania, Victoria, intand New 
South Wales and South Australia (Hickman 
1967; Austin unpubl.), C. sefoxys can be easily 
recognised from all other species hy its dense 
pilosity aod-sculpturing om the metasoma horn. 
Novweivpe specimens examined: SEA SOUTH 
WALBS: 1d, 49, gold coated of SEM holders, 
winks and antennae of (at. 1Y oo elldes, PS dis- 
secled on slide, te, FPP, ANIC 28 179. ONE. 


34 A. D. AUSTIN 


Pearl Beach, Jan. 1976, A.D.A., ex cays Ineniicus 
robusius (1. Koch) (Araneae): 16, 59, SAM; 
id, 59, WATTE, Caringbah, 12.ii1.1976, A.D.A.: 
39, Peurl Beach, Jan. 1978, A.D... BMNH: 2, 
49, Curingbah, 22.41.1976, ADA, QODPI, 
QUEENSLAND: 19, Brisbane, Sept. 1928, A. D. 
Dodd; 19, Gogingo, Dec. 1928, A, P, Dodd; 
19, Gordonvale, Nov, 1920 (no collector), ANIC: 
2% 29, Maleny, 14.vi.1973, M. D. (D.AL.); 12. 
600-700 m Sunday Creek nr Limna, 28-29.ix, 
1974, I. Naumann, QDPI, 


Acknowledgements 


[ thank Dr G. Gross, South Australian 
Museum, Dr I. D. Naumann and Ms J, C, 


Cardale, CSIRO, and Mr E. C. Dahms and 
Dr G. B. Monieith, Queensland Museum, for 
their hospitality while visiting the above insti- 
tutions and for loan of type specimens. I am 
grateful to Dr L. Masner and Dr 1. D. Galle- 
way for their helpful suggestions, and to Dr 
K, Bartusek for assistance with SEM tcch- 
niques. Early drafts of the manuscript were read 
and substantially improved by Prof. T. OQ, 
Browning, Dr P. W. Miles and Sally Austin. 
Financial support for this study was provided 
by a URG Scholarship from the University 
of Adelaide. 


References 


AsHmMeap, W. H. (1893) A monograph of the 
North American Proctotrypidae. Ball, U.S. natr. 
Mus. 45, 1-472. 

ASKEW, R. R. (1971) “Parasitic Insects”, (Heine- 
mann Educational Books, London). 

Austin, A. D. (1981) The types of Australian 
species in the Tribes [drini, Bacini and Embido- 
biint (Hymenoptera; Scelionidae: Scelioninie) - 
Gen, Appl. Ent. 13, 81-92. 

—— (1983) Morphology and mechanics of the 
ovipositor system of Ceralahbaeuxy Ashmead 
(Hymenoptera: Scelionidae) and related genera. 
Int. J, Inseet Morphol, Enrbryel, 12, 139-155, 

— (in press) The fecundity, development and 
host relationships of Ceratobaens (Hymenop- 
tera: Seclionidae). Ecol, Ent. 

Dopp, A. P. (1913) Further additions to the 
Australian Proctotrypoidea, Areh. f. Nature 
79 Abt. A, Heft, 8 77-91, 

—~ (1914a) Australian Hymenoptera Procto- 
trynoidea No. 2. Trans. R. Soc. 8. Aust. 38, 58- 
131. 


(1914b) Further new genera and species 
of Australian Proctotrypoidea. Proc. R, Soe, Oi 
26, 91-140, 

—— (1915) Australian Hymenoptera Proctotry- 
poidea No 3. Trans. R, Soc. §, Aust, 39, 384- 
454, 

(1919) Notes on exotic Proctotrypoidea in 
ihe Brilish and Oxford University Museums, 
with descriptions of new genera and_ species. 
Trans. R. ent. Sac. Soe. Land. 67, 321-82, 

Garroway, I, D. & Austin, A. D. (in press) A 
revision of the Scelioninae (Hymenoptera: 
Scelionidaé) of Australia. Aust. J. Zoel. Suppl. 
Ser, No. 99, 


Ginvons. J R. H. (1979) A model for sympaine 
specinion in Megarlvesa (Uymenoptera: che 
neumonidae): competitive speciation. 4a Nut 
114, 719-41. 

Grraucr, A. A, (1926) New pests from Australiy, 
V. Private Publication, Brisbane, 20 December 
1926. 2 pp. 

Hratwore, H. & Davis, D. M. (1965) Ecology 
of three species of parasitic insects of the 
genus Megarhyssa (Hymenoptera: Ichneumoni- 
due). Ecolopy 46, 140-50, 

Hickman, V. VV. (1967) New  Scclionidae: 
(Hymenoptera) which lay their eges in those of 
spiders. J. ent, Soc. Aust. (NSW) 4, 15-39, 

Huacerr, L. (1979) Revision of the Palaearctic 
species of the genus /dris Forster sl. (Hymenop- 
tera. Proctotropoidea: Scelionidue), Ent. Scand, 
Suppl. 12, 1-60. 

INTERNATIONAL COMMISSION ON ZOOLOGICAL 
Nomenciature (1964) International Cade for 
Zoalogical Namenelatire, Lendon. xx, 176 pp, 

Kuerrrr, J. J, (1926) Hymenoptera, Proctotry- 
poidea, Seclionidue, Das Tierreich 48, 1-885 

Masner, L, (1976) Revisionary notes und keys to 
world genera of Scelionidae (Hymenoptera; 
Proctotrupoidea). Mem. ent. Soe. Can, 97, 
1-47, 

—— (1980) Keys to genera of Scelionidae of the 
Holarctic region, with descriptions of new 
genera and species (Hymenoptera: Proctorrn- 
poiden). Shid. 13, 1-54. 

Price, P. W. (1972) Parasitoids nitilizing the sane 
fost: adaptive nature of differences in size and 
form, Bealugy $3, 190-95, 

Rowson, B, W. (1980) The backscattered-elec- 
tron Iow-vacuum $.B.M. technique: a users 
evilidion, Mieren 11, 333-4. 


SOME NEPOMORPHA (CORIXIDAE, NOTONECTIDAE AND NEPIDAE) 
(HEMIPTERA-HETEROPTERA) OF NORTH-WEST AUSTRALIA 


BY I. LANSBURY 


Summary 


New taxa of Notonectidae, viz. Walambianisops wandjina n.gen. and sp., Enithares gwini n.sp., 
Anisops douglasi n.sp., and Corixidae, viz. Agraptocorix gambrei n.sp. are described from the 
coastal region of the Kimberley, north-west Australia. Brief notes are given on the water-bugs of 
the Kimberley vis-a-vis those of the coastal zone, the Arnhemland Escarpment, and northern 
Queensland. A key to the Australian genera of Notonectidae, including new genera, is given, as 
well as a supplementary key to the Anisopinae using a combination of secondary sexual characters. 


SOME NEPOMORPHA (CORTIXIDAE, NOTONECTIDAE AND NEPIDAE) 
(HEMIPTERA-HETEROPTERA) OF NORTH-WEST AUSTRALIA 


by 1. LANstury® 


Summary 


Lansuuty, —F, (1984) Nepomorpha (Corixidac, Notonectidac and Nepidue) (Hemiptera- 
Heteroptera) of North-West Australia. Tram. R. Sac, 8 Aust 181), 35-49, 12 June, 1984, 


New taxa of Notonectidae, viz. Walaniblanivops wandjing neon. und sp... Enithares pwint 
Hap, Avivaps denvlast psp, and Corixidac, viz. Agraptocoriva santbre’ nap, are described 
from the coustal region of the Kimberley. north-west Australia, Brief notes are givens on’ the 
witer-bugs of the Kimberley vis-u-vis those of the coastal zone, the Arnhemland Escarpment, 
and northern Queensland. A key to (he Australinn genera of Notonectidue, including new 
genern, is given. as well as a supplementary key lo ihe Anisopinae using a combination of 


secondiiry sexcul characters. 
Ki'ty Worns: Corixidae, Apraptocerixe, 
Walamnbianisops, Zougcogruphy. 


Introduction 


Williams (1979) gives a concise description 
of north-western Australian freshwater fauna 
and a list of the water bugs recorded, Mosi 
of the species he listed are characteristically 
Australian, although some, according to the 
dala given. were collected from habitats east 
of the Kimberley in the Northern Territory. 
The Kimberley data available to Williams sup- 
ported Ais view that north-western Australia 
ineluding the Kimberley is not a distinet fluvi- 
faunular province, bul some more recent data 
derived from: collections in the coastal revion 
shed additional light on the biogeographical 
relationships of water-bugs in this area, They 
show that the water-bug fauna of the coastal 
region is unusual in having at Ieast one link 
with ‘Timor and Lombok, and is clearly not 
allied to the characteristic Australian fauna 
which vatends eastwards to New Caledonia 
god the New Hebrides—as typified by the 
Lnithares woodwardi Lansbury complex of 
sibling species (bergrothi Kirkaldy and hebyi- 
diensis Lansbury). The ew data suggest that 
species groups common to north-western Aus- 
tralia and Timor-Lombok have not in general 
became widespread over Australia, 

There is some evidence [hut a Tew species 
of water-bugs are distributed along the tropical 
fringe from north-west Australia, through the 
Northern Territory to north Queensland, The 
lack of data precludes a realistic discussion 
of the zoogeography of the north-west coastal 
regiow compared with areas of the Northern 
University 


‘Hope Enlomologicnl Collections, 


Museum, Oxford, (de, 


Notonectidae, Anisopy, Euithyres, Paranisaps, 


Territory and Queensland; so many groups of 
water-bugs, ic, Veliidae, Gerridae, Notonec- 
ude (Anisepy), Corixidae (Mieronéeta) and 
Hydrometridae, are insufficiently known from 
the north west coastal region, A few general 
comments are giver ort the relationships of the 
taxa of the north-west with other areas of 
“northern” Australia and elsewhere under 
species descriptions. 


Family: CORIXIDAE 


Agraptocorixa gambrei sp, nov, 
FIGS. 1-12 

Halawspe: Male, WAM 79-294-317. W.A., Port 
Warvender. ix.1976, A. M. Douglas In the Western 
Australian Museum, Perth. 
lMmension of helotype: Length, 7.75 mm, 
Width across the eyes, 2.8 mm, 
Colour; Vertex pale yellow, posterior margin 
Jatk brown. Pronotum dark brown. Clayvus 
yellowish brown with inner lateral anterior und 
posterior angles dark brown, Corium yellowish 
brown with a Jarge dark brown area along 
posterior half of the claval suture, the infus- 
cated area almost reaching the embolium. 
Membrane heavily infuscated. Embolium dark 
reddish brown hecoming paler distally. Thorax 
ventrally pate yellow, abdomen slightly darker. 
Front legs yellow, area of pala claws dark 
brown, Middle femur pale yellow, distal third 
and temaimder of leg dark brown. Hind femur 
pale yellow, tibia and tarsi rather darker, 
Sirveture; Vertex rounded and slightly pro- 
duced between the eyes. Interocular space 
(synthlipsis or natrowest part between the 
eyes) Jess than an eye’s width 1.3714, Mead 
Jengihh to pronotal length 75.100, Facial im- 


36 1. LANSBURY 


XY 


EUR EEeRENe Tey 


Figs 1-5: Agraptocorixa gambret sp. nov. holotype male. (1) head from the front; (2) jhid.. side 


view: (3) osteole of scent gland: (4) lateral 


pression conspicuous (Pigs 1. 2), clothed with 
fine silvery hairs, impression extending from 
lower margin of eyes almost to transverse 
suleations, Pronotal width twice median 
length, covered with fine dark brown ad- 
pressed spines arising from black bases. Clavus 
with similar spines about twice as long as 
those on the pronotum. Corium und right 
membrane clothed in longer dark brown hairs. 
Left membrane with sparse scattered short 
hairs. Pruiinose areca of the claval suture one 
third length of corial pruinose area 34:100. 
Post-nodal pruinose area length about half 
width 4:10. Lateral lobe of the prothorax us 
in Fig, 4. Metaxyphus as in Fig. 5. Sixth- 
eighth tergites as in Pigs 6-8, Sixth tergite with 
an extension, no “strigil’ visible, Front leg 
(Fig. 9) femur with several stout spines, pala 
with fifteen pegs, claws conspicuous. Middle 
femur curved, almost quadrate in section, 
inner margin densely clothed with fine silvery 
hairs. Hind femur (Fig. 10) aitteriorly pubes- 
eent. The relative lengths of legs are shown in 


lobe of prothorax; (5) metaxyphus. 


Table 1, Male genitalia (Figs 11, 12); right 
clusper clongate wedge-shaped, left clasper 
long and thin, 


Comparative Notes 

It is uot possible to key out A. gambrei in 
Knowles (1974) as the species does not have 4 
strigil. The sixth tergite has a projection rather 
like that of 4, paryvipunciata (Hale), A halei 
Hungerford, and A. Ayalinipennis (Babricius), 
the hitter does not occur in Australia, but there 
is no ecomb-like structure present, The number 
of pala pegs climinates A. parvipunetata and 
A. halei, as both have at most twelve pala 
pegs, the right claspers of both are quite 
distinctive, and the Avalinipennis males have 


Relative fengtlis af leys for Agrapto- 
coritn gambret yp. 10V. 


Pala 


Taste od. 


Femur Tibia Claw 


Front leg 100 44 30 25 
Middle leg 100 46 29 29 
Hind leg 100 103 113 an 


NEPOMORPHA OF NORTH-WESUt AUSTRALIA 27 


Tne 


Figs 6-8; Agraprocerixa gambrei sp. nov, holu- 
Iype mule, (6-8) tergites 6-8. 


the face flattened but not depressed and the 
peg row is much longer (22++), extending twa 
thirds the length of the pala palm. The facial 
depression of A. gambrei is clearly concave 
and well-defined, the peg row short nor reach- 
ing half way across the palm. 

There are now six species of Agratucoricn 
known from Australia. Of these, 4. hele? A. 
parvipunctata, A. firtifrony (Hale) and A. 
vambrei sp. n. appear to be endemic, while 
A. euryneme Kirkaldy and A. muacrops Hun- 
gerford are alsa known from New Guinea 
(Jansson 1982). A. parvipunerata and A. 
euryhome are both widespread species occur- 
ring over most of Australia and Tasmania: 
A. hirtifrony occurs sporadically, and A. salted 
seems to be a rather more tropical clement, 
occurring in north-west Australia, the N.T, 
and nerthern Qld. Kaowles (1974) provides 
records for N.S.W. and Alice Spritgs. A, 
macrops is known from New Guinea and 
Bunhury, W.A, A. gamhrei is known only from 
the coastal region of the Kimberley district. 


Avruplocorixa healet Hungerfard 
FIGS 13-19 
Agraprocarixa Nalei Hungerford, 1953, pp. 42-44. 
Agrapracorixa halet: Knowles. 1974, pp. 181-183, 

figs 25-32. 

Hungerford (1935) and Knowles (1974) 

have deseribed and figured 4. /ilei extensively. 
However, the form from Mt Trafalgar differs 
in some minor details from the typical form. 
The claw of the front leg is large but almost 
transparent in the “prepared” state (Fig. 13). 
The strigil is rather larger, almost horseshoe- 
shaped with five combs (Figs |S. 16) rather 
than three as described previously, Rarher 
figures of the 7th tergite show what appears to 
be » “free” triangular lobe dextrally; however, 
the posterior margin of the 7th tergite is con- 
tinuous, and there is a narrow. lightly sclero- 
tised strip) which superficially resembles a 
complete break in the tergite (shown as a 
pair of dotted lines on Fig. 14). Knowles’ 
figure of the left. clasper is drawn from an 
unusual angle; the clasper is broad with a 
promipent finger-like projection at its base 
(Fig, 17). The right clasper is much like 
previous illustrations (Fig. 18). The tip of the 
yedeagus is as shown (Fig. 19). 
Material examined: W.A. Prince Regent River 
Reserve, Ml. Veafalgar, (5° 75'S. 125°04'B, 
26.vil.1974, W. J. Bailey & KR, TL Richards, one 
mille and one femuile (Department of Agriculture, 
Perth. W.A.). 


Family: NOTONECTIDAE 
Anisops douglasi sp. nov. 
FIGS 20-22 

Holotype male: WAM 79-294-317, W.A,, Kim- 
herley region, Port Warrender, ix.1976. A, M. 
Douglas: in the Western Australian Museum. 
Perth. Dimensions of holotype: Length 6.9 mm. 
Colour, ethanal specimen; Eyes grey with 
faucets black. Pronotum anteriorly hyaline grey, 
posteriorly faintly tinged with orange. Seu- 
tellurm creamy yellow. Elytra hyaline, appear- 
ing grey with dorsal coloration showing 
through. Abdemen brown. ventrally rather 
darker with intersegmental membranes pale 
yellow. Connexivum pale yellow tinged with 
orange. 

Structure. Viewed dorsally the head is 
hroudly rounded with the anterior inner lateral 
margins of the eyes convergent. Greatest width 
of head fractionally Jess than pronotal humeral 
width, about ten times. anterior width of the 
vertex, and almost three times the median 
head length. Synthlipsis wide, over half but 


38 I. LANSBURY 


- 


WLS 


| Sty / fi // 


wy WNL 
SW Oh 


\ 
\ ny 
Sui At NN 


imm 


{ 


Figs 11-12: Agraplocorixa pambrer sp, nov. holo- 
type male. (11) left clasper: (12) right clasper, 


less than two-thirds the anterior width of the 
vertex. Median pronotal length about one third 
longer than the head. Pronotal humeral width 
just over twice median length, lateral margins 
straight, slightly divergent over half median 
length. Facial tubercle enlarged with small 
sul hairs (Fig, 21). Rostral prong large, 


OS 


Fig. 13: Agraptecorixa hale’ Wungerford mate, 
(13) front leg. 


the tip almost reaching the anniferous tubercle. 
Labrum slightly broader than long, apex acu- 
minate. Stridulatory comb with twelve pegs, 
outer seven rather longer (Fig. 22). Chacto- 
laxsy of the front leg as shown (Pig, 20). 
Comparative nates 

This species keys out to A. doriy Kirkaldy in 
Lansbury (1969), However, the large rostral 
prong, enlarged facial tubercle, and small 


NEPOMORPHA OF NORTH-WEST AUSTRALIA 39 


Figs 14-19: Agrapiocorixa Aalci Hungerford male. (14) 6th tergite; (15) 7th tergite; (16) strigil; 
(17) left claspers (18) right clasper: (19) tip of uedeugus. 


stridulatory comb distinguishes A. douglasi 
from A. deris, In Brooks’ (1951) key, A. 
douglasi appears to be closely allied to A. 
avsimiliy White, a New Zealand species, but 
the latler does not have an enlarged facial 
tubercle, and the stidulatory comb has about 
25-28 peys (4, douglas’ has rather fewer). 
Young (1962) redeseribed A. assiiilis and 
showed that Brooks’ descriplion of the stridu- 
latory comb was misleading, for ihe pegs 
decrease in size gradually towards the inner 
margin of the front tibia; the “steps” figured 
by Brooks is duc to some of the pegs having 
been broken. 


Anjsopy oceipitalix Breddin 
FIGS 23-25 

aAnisops occipitalis Breddin, 1905, p. £52. 

Anisops occipitaliss Brooks, 1951, pp. 344-5, PI. 
39, fig, 22. 

Anisops accipitalis: Lansbury, 1969, pp, 438-440, 
fies, 18-22. (Anivepy oculariy Pale, 1923 syno- 
nymnised With occipitalis). 

The occurrence of A, occipitalis in main- 
land Australia is rather patchy. Under the 


name “ocularis’, Hale (1923) first described 
and recorded the species from Australia 
(Darwin). Brooks (1951) gives data for Sir 
Graham Moore Island, und the Barron River, 
Qld. I haye collected material from Petersen 
Creek, Yungaburra near Atherton, Qld, which 
has been provisionally assigned to A. occipi- 
talix. This species. hears some resemblance to 
A, deanei Brooks. Both have prorninent rostral 
prongs, and the apex of the third rostral seg- 
ment is clearly wider than the base of ihe 
fourth (Fig, 23); both species have five spines 
on the inner surface of male front tibia (Pig. 
24), and the number of stridulatory pegs is 
roughly the same (20-24). The front femur of 
A. occipitalis is most distinctive, being broad 
proximally with “steps” distally; 4. deani has 
the upper and lower margins of the femur 
more or less evenly curved. According to 
Brooks (1951) .4. deanei males are at most 6 
mm long, but I have found that this species. is 
usually more than 6 mm Jong, with a maxi- 
mum length of 6.75 mm (Lansbury 1969). 
Large collections of A. deanet from “southern” 
Australia made in 1979 show that A. deanei 


40) 1, LANSBURY 


a25mm 
Anisops douglasi sp. nov. holotype 
male. (20) front leg; (21) side view of rostrum: 
(22) stridulatory comb. 


Figs 20-22: 


varies rather more in size than originally 
thought (Lansbury 1969), Extensive series 
from habitats around Alice Springs vary from 
6.5-7.5 mm long. A, oceipitaliy varies suli- 
ciently for smaller males of A. eecipitalis to 
overlap with the A. deanei miles, However, 
the little data there are sugzests that A. veci- 
pitaliy is a “coastal” species in Australiz. 
Majerial examined: Western Australia: WAM 
75—165-9, Wotjulum Mission via Derby, October 
1955, A, M. Douglas, lwo males and two females 
in the Western Australian Museum, Perth. N.W. 
Australia, Prince Regent River Reserve, Charoley 
River, 14.viii.l974, W. J. Bailey & K. Richards. 
one mile in Department of Agriculture, Perth, 
W.A. 


Enithares gwini sp. nov: 
PIGS 26-30, 33-36 

Holotype males WAM 79-294-317, two 
male and seven female paratypes, Western 
Australia, Port Warrender, Kimberley, Octo- 
ber, 1976, A. M, Douglas. One male and 
female piratypes, WAM. 79-218-27, Kalum- 
buru Mission, Kimberley, Western Australia, 
June, 1960, A, M, Douglas & G. Mees in 

Western Australian Museum, Perth. 
Shape: Broad robust species, lateral margins 
of the body converging slightly about midway 


Pigs 23-25; Anisops occipitulis Breddin male. (2: 
side view of rostrum: (24) front leg, (2 
stridulatory comb, 


) 
) 


body length, Median head Icngth appearing 
to be rather short compared with body length 
(1:10). 

Colour, Eyes brown. Vertex and anterior 
half of the pronotum straw yellow, viewed 
from the front, vertex with a brown bar 
between the eyes (not visible from above). 
Posterior half of the pronotum hyaline. Seu- 
tellum bluish-black, lateral margins. yellow. 
Anterior angles of the eclavus and corium 
yellow, pale coloration reaching the nodal 
furrow, remainder of clavus and corium rich 
brown to black in dry mounted specimens, 
colour in cthauol appearing rather darker with 
yellow areas rather more red than yellow, 
Membrane bluish-black. Embolium greyish 
yellow. Sternites black with central keel 
ereyish yellow. 

Structure; Head rounded, anterior width of 
the vertex. more or less continuous with the 


NEPOMORPHA OF NORTH-WEST AUSTRALIA 41 


imm 


\ ™ 


2 30 


025mm 


Figs 26-32: Enithures sp. males, E. gwini sp. nov. Figs 26-30: (26) head and pronotum dorsum, , 
(27) genital capsule: (28) paramere enlarged; (29) labrum; (30) metaxyphus; EL. lombokentsis 
Lansbury. Figs 31-32: (31) labrum: (32) head and pronotum dorsum. 


eyes. Greatest width of head about two thirds 
pronotal humeral width, just over two and a 
half times the unterior width of the vertex, 
and three times median head length, Synth- 
lipsis wide, just over half the anterior width 
of the vertex. Head length slightly less than 
the anterior width of vertex. Pronotal humeral 
width almost three times median length, lateral 
margins strongly divergent, appearing straight 
although they are slightly convex. Dorsal 
margin of pronotal fovea directed obliquely 
laterad behind the eyes (Fig. 16). Nodal 
furrow obliquely turned towards the head and 
less than its own length removed from the 
membranal suture. Labrum acutely triangular 
(Fig. 29). Mesotrochanter rounded densely 
cloihed in Jong curly hairs, inner Jower margin 
of middle femora densely hairy (Fig. 33). Ol 
the middle tibia and tarsi (Fig, 34), the tibia 
is prolonged at outer distal margin, and the 
inner proximal margin has a less prominent 
projection. Hind femur of male produced 
distad-venirally (Fig. 36). First tarsal segment 
of the hind leg with a ridge along the inner 


margin fringed with long hairs along both 
margins (Tig. 35). Metaxyphus triangular 


(Fig, 30). Male genital capsule as illustrated 
(Fig. 27); parameres (claspers) symmetrical 
(Fig. 28), 


Comparative notes 


Similar to Enithares lomhokensis Lansbury 
(Fig. 32) and &. Auhleri Brooks, E. gwini is a 
more robust species. The front tibia of EF. 
gwint is parallel) sided, that of E. buhleri (Fig. 
37) is distally conate, the front tibia of E£, 
lambokensis (Fig. 39) deeply concave with a 
blunt projection distally. The middle tibia of 
E. gwint has a blunt projection apically, and 
18 sharply produced distally whereas E. byhleri 
(Fig, 38) and 2. lambokensis (Fig. 40) both 
have more or Jess parallel-sided mid-tibiae 
which are sharply produced distally, 

Three other species of Enithares are known 
from Australia. E: woodwardi is widespread 
over much of “southern” Australia. 2. huckeri 
Hungerford occurs sporadically in northern 
N.S.W. and Qld. The third species E. loria 


42 1. LANSBURY 


Pigs 33-34; Enithares gwini sp, nov. holotype male. Middle leg. 


Brooks, originally described from New Guinea, 
is folind fairly frequently along the tropical 
fringe of the N.T., Qld, snd in the Salomon 
Islands (Lansbury 1968), 


Walambianisops wandjina gen, and sp. noy. 
FIGS 41-57 


Generic cingnosis; Vertex. extending beyond 
the eyes. Antennae two-segmented. Labrum 
and rostrum conspicuously hairy. All legs of 
both sexes with two-segmented tarsi, Front 
coxa and femur long. Coxal plates (third epi- 
sternum and infracoxal plates of Aucts.) bare, 
cistally fringed with short black hairs. Fifth 
and sixth sternites carinate, narrowly bare. 
fringed with short black hairs, 
Type species: Walambianisops wandjina sp, n. 
In the discussion following the description 
of this new genus and species, a section is 
included on Paranisops which is a superficially 
similar genus, The merits of the latter as a 
generic entity are commented upon. 


Walambianisops wandjina sp. nov. 
FIGS 41-57 
Type series: Holotype male, seven male and six 
femile paratypes WAM 79-287-93 anit WAM 
779-294-317 W.A,, Port Warrender, Kimberley, 


in.1976, AS M. Douglas: iwo male and three 
fomiule paratypes WAM 79-218-27 W.A,, Kalum- 
bury Mission, Kimberley. vi,1960. A, M. Douglas 
& G, Mees; one female paratype WAM 79-1659 
W.A,, Wotjalum Mission via Derby, Kimberley, 
ix,1955, A. M, Douglas in the Western Austeslian 
Museum. Perth. One male and one female para- 
types, N.W. Australia, Prince Regent River 
Reserve, 17°07'S, 125°33°E ex ravine pond, 
17.wili. 1974, W. J. Bailey & K. T. Richards, in 
the Department of Agriculture, Perth, W.A. One 
female paratype, W-A., Koolan Island, permanent 
pool in dry ereck bed, 13.11.1978, W. D. Williams 
in the collections of Adelaide University. 


Lengri: males 9-9.6 mm, females 9-9.9 mm. 


Coloyr: Eyes cither greyish with pale hori- 
zontal bands or dark reddish brown. Head 
yellow with a small brown spot between the 
eyes on the frons. Pronotum anteriorly 
greyish brown, posteriorly more hyaline, 
appearing darker due to dorsal coloration 
showing through. Scutellum dark brown to 
black, shining, apex ond part of the lateral 
margins orange yellow, Clavius and corium 
apically suffused dark brawn to black, occa- 
sionally the dark suffusion extending over most 
of the clavus and along the outer lateral! 
margin of the corium, both the clavus and 


NEPOMORPHA OF NORTH-WEST AUSTRALIA 43 


ae 
‘o 


doin 


37 

Figs 35-40: Enithares sp. males, E. gwini sp, nov. 
Figs 35-36: (35) first tarsus of hind leg; (36) 
hind femur distally; E, buhleri Brooks. Figs 
37-38, LE. lambokensis. Figs 39-40; (37 & 39) 
front tibia; (38, 40) distal end of middle tibia 
and 1st tarsal segment. 


corium faintly iridescent green. Remainder 
of the elytra and membrane hyaline, the red- 
dish brawn tergites showing through. Legs 
dark-reddish brown with margins narrowly 
yellow, particularly conspicuous on the front 
and middle legs. Thorax reddish-brown with 
yellow patches. Sternites dark brown to bluish 
black. Lateral margins of the connexivum nar- 
rowly pale yellow. 


Structure: Eyes large and protuberant, vertex 
extending beyond the eyes (Fig. 42), dorsally 
between the eyes shallowly depressed, Facial 
tubercle enlarged and rounded, frons above 
facial tubercle deeply concave. Labrum and 
rostrum hairy, labrum slightly broader than 
long. Anterior lateral margin of the third 
rostral segment finely serrate (Fig. 41). 
Greatest width of head five times anterior 
width of vertex, and about six times width of 
the synthlipsis which is almost as wide as the 
anterior width of the vertex. Head width more 
than twice median head length. Anterior 
margin of pronotum raised and produced 
between the eyes. Pronotum almost one and a 
half times longer than the head, humeral width 
greater than head width and almost twice the 
median pronotal length, Jateral margins 
strongly divergent. The female is much like 
the male, head width three times head length, 


Figs 41-43: MWalambianisops wandjina gen. and sp. nov. male. (41) head and pronotum Irom the 


side; (42) ibid., dorsum; (43) antennac. 


44 I, LANSBURY 


ES 
SSS Se Se 


SAAS 


~~ 


0-25mm— 


Figs 44-49; Hulambianisaps wandiina gen. and sp. nov, mate. (44) front leg: (45) enlarged deta 


of front leg; (46) enlarged detail of front tibia: 


Parameres. 


pronotum almost twice as long as the head. 
Scutellum large, strongly convex, about as 
broad as long, 

Front coxa and femur longer than tibia, 
front tarsi two segmented, claws short and 
blunt apically, those of the female spinose, 
Femur with four groups of elongate setae 
arranged like inverted cones (Figs 44, 45). 
Front tibia moderately spinose with a row of 
“pegs” separate from each other forming a 
“stridulatory-comb” (Fig. 46). Middle femur 
elongate and spinose, chaetotaxy not differing 
greatly from that of front leg, but without the 
peg-row. Very little difference between chacto- 
taxy of males and females. Hind leg elongate, 
fringed with moderately long swimming hairs, 
tarsi of middle and hind legs two-segmented, 

Coxal plates bare and shining with coarse 
irregular oblique striations, distally fringed with 
black hairs (Fig. 51). Third sternite just visible 
latero-distad of coxal plate. Fourth sternite 
with a trichome, fifth-seventh sternites variably 
carinate, apically narrowly bare, fringed with 


(47) genital capsule; (48, 49) right and Te 


shart black hairs. Female seventh sternite not 
as acutely carinate as preceding sternites, 
males rather more sharply convex than eari- 
nate. Outer lateral margins of connexival 
segments two-four with small half-moon 
shaped projections arising from depressions in 
the integument (Fig. 51). Arising from the 
lateral margins of the seventh and eighth 
tergites pale yellowish hairs which are much 
longer than those along the inner margins of 
the connexivum. Male seventh abdominal 
tergite with a prominent sclerotised spine on 
the caudo-sinistral margin (Figs. 53, 54), 
Male operculum (subgenital plate) with 
fringes of hairs distally (Fig. 57). Female 
operculum much larger, deeply convex and 
fringed with several rows of hairs and spines 
(Figs 55, 56), 


Antennae two-segmented, both segments 
with modified spatulate hairs (Fig. 43). 


Male genitalia (Fig. 47): capsule partially 
cleft posteriorly and heavily sclerotised, para- 


NEPOMORPHA OF NORTLLEWES! AUSTRALIA as 


tex —_- 

imin 

Vips 50-52: Walambianisops wandjina gen. und 
sp, nov, (SQ) mule aedeagus; (51) coxal plates 
und connexivum of male; (52) female genitalia; 
terminology 8-9 paratergites 8 & 9, 1 & 2 gpo 
Ist und 2nd gonuphysis, 1 & 2 ex Ist und 2nd 
fonocoxa, ga gonangulum, gpl gonoplac. 


meres (claspers) large and  asymmictrical 
(Figs 48, 49), aedeagus membranous (Fig, 
52). 


Female genitalia (Pig, S2y: first: gonocoxa 
elongate, anterior margin infolded forming a 
partial ramus. First gonapophysis large, sclero- 
lised With stout spines distally, ventrally fringed 
with short hairs, Gonanguluin clongate, almost 
reniform, fused to paratergite nine. Second 
gonocoxa membranous and acutely triangular. 
Second #onapophysis membranous, distally 
lightly selerotised. Gonoplacs stylus-like with 
scattered hiirs. 


Discussion 


Superficially, Walambianisops resembles 
Paranisaps endymion (Kirkaldy). However, 
despite resemblance in size and coloration, 
only one primary character is common to both 
genera: all legs of both sexes have two-seg- 
mented tarsi. Walambianisops has many fea- 
tures in common with Aniseps, but the aper- 
culum is not sharply carinate as it is in Ani- 
sops; it is variably convex, ie. it is structurally 
midway between Aniseps and Paranisops. The 
males of Walambianisops have a large sclero- 
tised spine on the seventh tergite (Figs 53, 
54): Anivaps and Buenoa Kirkaldy (New 
World counterpart of Anisops) have a 
variously shaped projection on the same ter- 
gite. This projection is absent from Puranisops 
species (Bigs 58-60). The seventh abdominal 
tergite of Walambianisaps is asymmetrical, that 
of Paranivops symmetrical with a prominent 
sclérotised projection extending caudad dor- 
sally (Figs 38-61), The cighth abdominal seg- 
ment is much the same in both genera. The 
male genitalia of Paranisopy (Fig. 62) are 
robust, well-sclerotised structures, cleft pos- 
teriorly, The parameres of P. endymion are 
symmetrical, of P.. inconstans Hale, asym- 
metrical; those of Wealanihianisops arc much 
like the generalised type found in Anisops and 
Buenoa, The first gonapaphysis of Anisaps, 
Buenoa and Walambianisaps ave all heavily 
sclerotised and spinosc, whereas Paranisops 
has a feebly sclerotised first gonapophysis 
which are plate-like with a few spines distally. 


Features unique to Walambianisaps males 
include the four groups of setae om the front 
femur and the row of pegs across the front 
tibia. (The latter are homologous. with the 
more complex stridulatory peg row of Anisops 
and Bueneoa, The row of serrations along the 
third rostral segment of Walambianisops can 
be equated with the rostral prong of related 
vénera.) Finally, there is a series of half-moon 
shaped projections on the outer lateral margins 
of the connexivum. Female Malambianisops 
can be distinguished from Paranisopy by the 
smooth coxal plates; in Paranivopy these plates 
are covered in black hairs, Females of Walam- 
biunixops may be distinguished from Anisopys 
by their two-segmented untennae (those of 
Anixsops are always three-segmented). 
Although the antennae of Walambianisops are 
two-sexrnented, the general chactotaxy closely 
resembles thal of Anisopy CLundblad 1933). 


46 T. LANSBURY 


SSS 
SO 


sot 
at 
Ta a 


ZZ 
ern 


EN: 


Figs 33-59: Walambianisops waning gen. aod sp. nov. Figs 53-57: (53) male 7th tergite side 


view: (54) ibtd.. dorsum; (55) 


side view of 


female operculiim; (56) ibid. ventrally: (57) 


male operculum ventrally; Paradisaps imeonstans Hale mile. Fies 38-59: side and docsul aspects 


of 7th and Sih tergiles, 


MWulambianisops shares a number of features 
with Antsops and Parayisops. However, the 
combination of characters found in Walam- 
Aranixsops supports the hypothesis that Anisops- 
Buenouw and Waliunbianisops are more closely 
related to cach other than they are to Parani- 


soppy. The key to the Australian genera of 


Notonectidae refleets the close relationship 
of Waluaibiantisops and Anixopy. However. it 
secondary sexual characters are used, as in the 
alternative key to the Anisopinae, Helam- 
Miunisaps is shown to be quite distinct fram 
other genera of the subfamily. 

Anisops, itself is a large genus with ca. 34) 
species Known from Austraha (well over 100 
in the Old World). One species, 4. agalin 
Hinehinsen from South Africa, was placed 
i a sepiirate subgenus Aniyepoides as the 
mates lack a rostral prong, the stridulatory 
pegs are arranged in the same way as in 
Wokvahianisops, the front tibia is enlarged 
apically (thase of Halunthienisops are nat), 


Aniseps avalia males, ike all other species in 
the genus, have one-segmented front tarsus, 
No informution is available on the presence of 
discrete auditory specialisations on the ros- 
trum, 

Buenow males have two-segmiented front 
tarsi, a rostral prong, and most, in addition 
to the stridulatary comb on. the [ront tibia, 
have a stridulatory area on the front femur. 
The secondary stridulatory zone is absent from 
all Old World Anisopinae. Possible secondary 
stridulatory features have been deseribed on 
Anivops nullati Poisson from Madagascar 
(Lansbury. 1966). 

Unusual if not unique in the Anisopinae, the 
female of Paraniseps endymion hears several 
secondary scaual characters (extensions to 
hind margins of eyes, and projections on the 
lateral margins of the pronotum). They are 
more fully deseribed in Lansbury (1964), 
Females of the related species P. incunstans 
do not exhibit this reversal of secondury 


NEPOMORPHA OF NORTH-WEST AUSTRALIA VW 


Figs 60-63: Puranisops endyvmien (Kirkaldy) 


sexual characters. ‘The data available for 
Paranivops show that P. endymian is restricted 
to south-west Australia, and the distributional 
gap between P. endymtion and Walanibianisops 
is extremely wide. Lansbury (1964) observa- 
tions on P, incanyteany were based on limited 
numbers. of specimens from various localities. 
There was no data on the relative abundance 
of the “macropterous” or dark form P. ineon- 
stains inconstins compared with the Jeuco- 
chroie form P_ incenstany vat lutea Hale in a 
population. During 1979 at Cedar Falls near 
Brishane P. inconstans was found in relatively 
large numbers in deep, shaded water under a 
hridge. The “macropterqus” form P, inconstans 
incenstany uncommon; one male and two 
females in a sample of 34 males and 56 
females, remainder being P. ineonstans var 
lutea, P. ineenstans is an eastern coastal 
species recorded from N.S,W, and Queensland. 

The two species of Paranivops are charac- 
teriscd by the flat operculum (Figs 60, 63), 
the posteriorly cleft genital capsule, the hairy 
coxal plates, the carinate frons and the thin 
plate-like first gonapophysis. P.  endyinion 
males have symmetrical parameres, females 


mide. 


(61) 


(60) side view of 7th teryite, 
dorsum of 7th tereite: (62) genital capsule, after Lansbury, 1964; (63) operculum, 


ihid,, 


have secondary sexual characters. The chacto- 
taxy of the male and female front legs is 
similar except in the claws which are 
dimorphic, P. incenstaris males have asym- 
metrical purameres, the chaetotaxy of the 
male front lee differs markedly from that of 
the female, the claws are not sexually dimor- 
phic, and the female does not have any 
obyious secondary sexual characters (as 
exhibited by P. eadymion). The similarities 
and quite striking differences between these 
twa species suggests that generic concepts 
within the Anisopinae are fairly flexible. 


Key to Australian genera ef Noioneetidac 


1. Hemelytral commissure cominous, without 
a hair-lined pit close to the apex of the 
seutellum (Notonectinge) ‘y 

— Hemelytral commissure with a prominent 
hair-lined pit close to the apex of the 

scutelhim (Anisopinach _ 4 

Mid-femur with a large spine ante-apically 
(Fig. 33). Eyes dorsally widely separated 
(Fig. 26) , 3 
— Mid-femur with a smull spine unte-api- 
cally, Eyes dorsully contiguous forming an 
ocblir commissure, Le, appearing to be 


i) 


N 


AR I. LANSBURY 


joined or ovetlapping, no syn{blipsis 
Nyehia 
3. Antera- Jateral margins of the pronotin 
not foveate . Netortecta 
Antero-literal margins of ‘the pronouin 
fovyeate (Figs 26, 32) Enithares 
4+, Coxal plates shining, sometimies coursely 
striate, distally occasionally fringed with 


black hits at ete 5 


black hairs 
Parnnixaps 
s Antennae tavo- segmented (Fig. 43) , 
Walambianilsaps gen, nov 
— Antennae three-segmented Anisaps 
Alternative key te Australlan penera of Anisopinae 
(. Prominent rostral prong, Front tibia en 
Jarged aploally with a cow of strtdulatory 
pegs. Front tarsus one-segmented in male, 


Coxal plates covered with, 


wwo-sepmiented in female. Opercolum 
carimebe Anisaps 
Rostral prong athsent. Front tibia not 


enlarged apically ta accommodate sipidu- 
latory pegs (if present). Front tarsi two: 
seamented in beth sexes. Opeteulum 
either fat or convex o 
Coxal plates bare Front ferme with four 
kroups of setue. Pegs of sividulatory row 
ull clearly separate, Antennae two-sep- 
mented. Frons’ and vertex bathaus 
Walambianisags 
— Coxal plates covered in black hairs, Front 
femora without groups of setae. Stridu 
latory pegs absent. Antennae three-seg- 
menied. Frons and verlex curinate 
Puranisops 


ts 


Family: NEPIDAE 
Aistronepa anevsra CHale) 

Crnfer angusta Hale, 1924, pp. 308-809 
Austronepa ancuyta: Menke .& Stange, 

67-72 (new senus for aayusia), 
Austrenipa anpnsht: Lansbury, 1967, pp. 641-644 

Auspronepa is widely distributed through 
the tropical fringe of the Northern Tertitery 
and in Queensland down to Stradbroke sland. 
Breeding sites vary, They include toad-side 
pools and ditches with or without dense 
growths of macrophytes (as in) Queensland 
near Mareeba), small shallow grassy pools 
(az. by the Mary River along the Arohem 
Highway, W.T.), and large billabong (e.p. 
near Nourkingee Rock. Northern Territory, 
where Anstrenepa was found in deep water 
amongst Pinidanus roots). ft is fairly common 
aut Foge Den near Darwin which has clenr 
water, and abundant at MeMinns Lagoon also 
near Darwin, where the water is extremely 
turbid, Althaugh found in a wit variety of 
hobiruty, all are lentic or slow totic. Little 


1964. pp, 


variation i$ apparent in size and external mor- 
phology of the species, and the male genitalia 
are uniform throughout the range. 

Marrial wxvamined: WAS WAM 79 149-545. 
Kulumburu Mission, Kimberley, 7.1960, A. M. 
Douglas & G. Mecs, two mules, two females and 
three immatures (4th-Sth instar), in the Western 
Australian Museum, Perth, 


Ranatra diminuta Montandon 


Ranatra leorpipes. var diminuta Montandon, 1907. 
p57 
Ranatra diininuta Montandon; Lansbury, 1972, 


pp. 323-326 (var dinrinita clevated to species), 

In Lansbury (1972). 4pecimens from north- 
west Australia of R, diminuta key out as R, 
langipes Stil, because the eyes are eleauly 
wider than the interocular space. However, 
the male genitalia of the two species are dis: 
tinctive; the parameres of R. diminuta have a 
prominent Tooth distally, this projection is 
missing in R. lonulpes, The status of several 
Species Of Ranatra from the Australasian 
region is Open to question. A possible synonym 
of R. diminuta is R, birei Lundblad (1933) 
described from New Guinea (Astrolabe Bay), 
Comparison of the types of R. birot with 
those of R. diminuta seemed to indicate that 
the status of hoth species was valid (Lansbury, 
1972), However, more recent studics of addi- 
tional material from Australia, New Ciuinea 
and the Philippines has shown that characters 
previously used, such as eve width-interocular 
space, distance helween middle coxac com: 
pared with hind coxwe, and elevation of vertex 
above eyese-are all taxonomically unretiable- 
Another species described from Millstream. 
Western Australia CR. occidentalis \ansbury) 
is also part of the R. diminuta complex. As 
R. oceldenttalty appears to be geographically 
isolated from mainland Australian Re dorinute, 
it is sail tentatively regarded as 4 distines 
species. Typical R. diminiuta is fairly common 
in the Philippines, but there is a high level of 
trivial variation iit the shape of the parameres 
(see Lansbury, 1972), A Torge series of R. 
dininuta trom Queensland has: 49% have the 
eye width equalling interocular space, 37% 
have the eye width less than interocular space, 
14% have the eye width greater than inter- 
ocular space. ‘The simplistic shape and Jack of 
“ornamentation” throughout tnest-oaf the sniall 
oriental Menara renders the species limits 
obscure. 
Mrrerial 
Wotjuluin 


exanivieds WyA.s WAM 79-126. 
Mission via) Derby, Kimberley, 


NEPOMORPHA OF NORTH-WEST AUSTRALIA A) 


xii.1955, A. M. Douglas, two males and two 
females in the Western Australian Muscum, 
Perth. 


Acknowledgments 


J wish to thank Dr T. Houston, W.A. 
Museum, and the Director, Department of 
Agriculture, Perth, W.A. for making available 
to me via the Division of Entomology, 
C.S.L.R.0,, Canberra, the remarkable aquatic 
Heteroptera described in these notes. I am 


deeply indebted to Professor W. D. Williams, 
Adeluide University, for the loan of material 
and cditorial assistance, to Dr Houston for 
information on the Kimberley region and to 
Miss Sandra Lawson, Secretary, Department 
of Zoology, University of Adelaide, for the 
preparation of the final manuscript. This work 
was commenced during the tenure of a grant 
from the Leverhulme ‘Trust (London) and 
grants from A.B.R.S. and C.S.1.R.0., Can- 
berra. 


References 


Brepoin, G. (1905) Rhynchola Heteroptera ats 
Java, gesammelt von Prof. K. Kraepelin 1904, 
Mitt, Naturhist,. Mus, Hanthurg. 22, 152. 

Brooks, G. T. (1951) A revision of the getus 
Anisops, Kars, Univ. Sci. Bull. 34, 301-5 19. 

Have, H. M. (1923) Studies in Australian Aquatic 
Hemiptera. Ree, §. Aust. Muy. 2, 379-424. 

—— (1924) Studies in Australian Aquatic Hemip- 
tera No. 3, (hid, 2, 503-520. 

Huncervorn, H. B, (1953) Notes on some Cori- 
xidae from New Guinea and New Caledonia, 
Pacific Insects 24, 95-103, 

Know es, J. N. (1974) A revision of Australian 
species of Agraplecupiva Kirkaldy and Dia- 
prepocoris’ Kirkaldy. Anst. J. Mar. Presiiwater 
Rey. 25, 173-191. 

Lanspury, |. (1964) A revision of the genus 
Paranisops. Proc. R. ent. Soc. (B) 33, 181-188. 

(1966) Notes on Aniveps niilloti Poisson, 
1948, Ann. Mag, nat. Hist. (13) 9, 375-376. 

—— (1967) Comparative morphology of the male 
Australian Nepidae. Aiat. Jc Zool, 15, 641-649. 

(1968) Enithares (Notonecidae) of the 

Oriental region. Pacific Insects, WW, 353-442. 


(1969) The genus ¢lnivepy in Australia. J. 
nat, dist, 3, 433-458, 

—— (1972) A review of the Orientil species 
of Runatra, Trans. Ry ent. Soe. Loanil. 124, 
287-341. 

LunNpeLap, O, (1933) Zur Kenntinis der aquatilen 
und semi-aquatilen) Hempiteren you Sumiutra, 
Java und Bah. Archiv, f, Hyvdrobiolagu Suppl. 
12, |-489. 

Menkr, A, L. & Srasce, b, A. (1964) A new 
venus of Nepidae from Australia with notes on 
the higher classification of the family. Prac, 
roy, Sec. Od, 75, 67-72. 

Munranpon, A. L. (1907) Quelques especes du 
genre Rauatra. Ann, Sec. ent. Fr. 77, 49-66, 

Trouxat. B.S, (1952) The comparative mor- 
phology of the male genitilia of Ihe Notonec- 
lidag. J. Kaas. Entumal, Soc, 25, 30-38, 

WituiAms, W. D, (1979) Notes on the freshwater 
fauna of north western Australia, especially the 
Kimberleys. Rec. West. Aust. Mus, 7, 213-227. 

Younac. BE. (1962) The Corixidae and Notonec- 
tidae of New Zealand. Ree. Cunterbury Mus. 7, 
327-374. 


PHYSICOCHEMICAL AND BIOLOGICAL STUDIES ON THE COORONG 
LAGOONS, SOUTH AUSTRALIA, AND THE EFFECT OF SALINITY ON 
THE DISTRIBUTION OF THE MACROBENTHOS 


BY M. C. GEDDES 


Summary 


During 1982 there was no outflow from the barrages at the mouth of the River Murray and the 
Coorong was a hypermarine system. In the North Lagoon spatial and seasonal patterns of salinity 
were apparent with salinities highest (to 80%c) at the more southerly stations in the summer months. 
Salinities were between 90 and 100%c in the South Lagoon. The macrobenthos in the North Lagoon 
was dominated by the amphipods Melita zeylanica, Paracorophium sp. and Megamphopus sp., the 
polychaetes Ceratonereis pseudoerythraensis, Nephtys australiensis, Capitella capitata and 
Ficopomatus enigmaticus, the bivalves Notospisula trigonella and Arthritica semen and the 
gastropods Hydrobia buccinoides and Salinator fragilis. The species number was low, possibly 
reflecting the extreme salinity fluctuations in the Coorong. High salinity in the southern end of the 
North Lagoon restricted the distribution of most species. The fauna of the South Lagoon was a salt 
lake assemblage including dipterans, especially the chironomid Tanytarsus barbitarsus and the 
ephydrid Ephydrella sp., the ostracod Diacypris compacta and the isopod Haloniscus searlei; the 
only fish present was the hardyhead Atherinosoma microstoma. The salinity regime in 1982 
represented an extremely hypermarine phase in the long term salinity fluctuations of the Coorong. 
In periods of sustained high River Murray outflow in the mid 1970s, salinities were estuarine in the 
North Lagoon and from 30-70%o in the South Lagoon. 


PHYSICOCHEMICAL AND BIOLOGICAL STUDIES ON THE COORONG 
LAGOONS, SOUTH AUSTRALIA, AND THE EFFECT OF SALINITY ON 
‘SHE DISTRIBUTION OF THE MACROBENTHOS 


by M, C. Geppes & A. J, BUTLER*® 


Summary 


Gepors, M,C, & Baviun, Ao (1984) Physicochemical and biologival studies on the Courony 
Lagoons, Soutl) Austialia, and the effect of salinity on the distribution of the macrobenthos, 
Trans, R. See. §. Aust. W801), 51-62, 12 June, 1984, 


During (982 there was no outflow from the barrages at the mouth of the River Murray 
ind the Coorotg wos a bypermarine system, In the North Ligoor spatial and seasonal patterns 
of suinity were apparent with sulinilies highest (ta BOS) at the more southerly stations in 
the suminer months, Salinities were between 90 und LOOS, in the South Lagoon. The magro- 
benthos in the North Lagoon wis dominated by the amphipods Melita ceylanive, Paracore- 
phivn sp. and Megamphopas sp.. the polychuetes Ceratonere’s preudoersiiraensis, Neplitys 
austrulionsis, Capitella capliata and Ficopamatas euusidticns, the bivalves Natoaspisala tiga. 
nella wnd Ariieiieca semey, and the gastropods Mydrohia buceinwides and Salinator fragilis. 
Che species number was low. possibly reflecting the extreme salinity fluctuations in the 
Coorong. High salinity in the southern end of the North Lagoon restricted the distrifvution 
of most species, The fauna of the South Lagoon was a salt lake assemblage inchiding 
dipterons, especially the chironomid Tiuvtytersas barhitaryes and the ephydeid Ephydrella sp. 
the ostracod Diveypriy compacta and the isopod Haleniseus scorlet; the only fish present was 
the hardyhead Aiherinaxonim micrastoma, The salinity regime in 1982 represented an 
exiremely hypermarine phase in the long term sulinity fluctuations of the Coorong. In periods 
of sustained high River Marray outflow in the mid 1970s, salinities were estuarine io the 


North Lagoon und from 35-707. in the South Lagoon. 
Kry Worns: Physiochemistry, Coorong Lavoons, salunty, macrobenthos, bypermarine 


system. 


Introduction 


The Coorong is a coastal lagoon system 
which extends trom the mouth of the River 
Murray some [00 km southeast along the 
South Australian coast (Fig. 1). (t is divided 
into North and South lagoons by a constricted 
area which limits waler exchange. The North 
Lagoon has an area of approximately 73 km* 
and a volume of approximalcly 73.000 ML in 
summer; corresponding Values for the South 
Luyoon are 8O km* and 93.000 Ml Both 
lagoons are shallow with meun depths of 1.0 
mand 1,1 m for the North and South lagoons 
respectively, The Coorong was. formed when 
rising sea level Hooded an old interdunal 
corridor about 6,000 years BLP.) (yon cer 
Borch 1975). A barrier dime, the Youugtius- 
band Peninsula. was built up and seawater 
access was gradually resirieted, Deposits 
Within the Coorong suggest that the system 
assumed its present lagoonal character perhaps 
3.000 years BP. (Browne L965)1. The present 
Coorong has only a restricted entrayice via the 
narrow chinnel at the Murray mouth, The 


 Zoolory Department, University of Adelude, 
Box 498, G.P.O,, Adelaide, & Aust. 5001, 


only freshwater inflow is fram the River 
Murray which iscat the extreme northern end 

of the lagoon system, There ix at present na 

significant run-off elsewhere along the Jagoons, 
although historical revords indicate that flows 
may haye entered via Salt Creek in the first 
half of the century (Noye 1975). Under the 
present regime much of the environment of 
the Coorong is likely to he hyperhaline (sense 

Barnes TYSO), 

Most af the work on the Coorong lagoons 
has been of a geologicul (Browne 1965!: von 
der Borch 1965, 1975) or hydrological (Noye 
19702; Noye & Walsh 1976) nature, although 
some consideration of the plants, fish and 
aquatic birds is given in Noye (1975). ‘The 
Coorong is acknowledged as a major habitat 
for aquatic birds and some work has been 
done to estimate bird numbers and to study 
the major food chains in the Coorong leading 
1 Browne, R. G. (1965) Sedimentation in the 

Coorong, South Australia, Ph.D. Yhesis. Uni- 
versity of Adelaide (unpublished). 

“Noye, BJ. (1970) On the physical Jininolagy 
of shallow takes aiid the theory of tide wells, 
Ph.D, Thesis. University of Adekvide  (unptity 
lished), 


M. C. GEDDES & A. J, BUTLER 


Touwitenere Is, 


Ceaorong 
Channels 
Murray Mouth 


WOE 
TH LAGI 


South 
Australia 


SOUTHERN OCEAN 


Stony Weil Basin 


Cattle Is, Basin 


VQ G@{ 
|S Policeman's P 


¥Sc EK: 


Salt, Ck. Pr. 
= S 


Seagull Is 


Saft Ck. Basin Flax Pt 


Tea Tree Pt. 


Policeman's Pt Basin Bul Bul Basin 


Fig. |. The Coorong lagoons showing stations (1-9) sampled in_ this study. Bathymetric contours 


are based on Noye (1975) and are approximate. 


to aquatic birds (Delroy et al. 1965%; Paton 
1982'). The Coorong also supports a major 
fishery and over the six years to 1981-82 the 
annual catch of yellow-eye mullet (Aldrichettu 
forsteri) has been 105-235 tonnes, that for 
mulloway (Sceiaena antarctica) 24-115 tonnes 
and for black bream (Acanthopagrus butcheri) 
from 10-72 tonnes®, Some studies have been 
made on the biology of the yellow eye mullet 
(Harris 1968) and the black bream (Weng 
1970"), 

The present study constitutes a limnological 
survey of the Coorong during 1982, with par- 
ticular emphasis on the effect of salinity on the 
distribution of macroinvertebrates. In addition 
some analysis is made of the longer term 
“Delroy, L. B., Macrow, P. M. & Sorrell, J. B. 

(1965) The food of waterfowl (Anadidae) in 

salt water habitats of South Australia. Unpwuhb- 


lished report of Fisheries and Fauna Conserya- 
tion Department of South Australia. 

4 Paton, P. (1982) Biota of the Coorong. South 
Australia Department of Environment and Plan- 
ning, Nov. 1982, S.A.D.B.P. 55 (unpublished ). 

“SAFIC February (982, April 1983  Cunpub- 
lished). 

“Weng, H. T, C. (1970) The black bream A cai- 
thopagrus butcheri (Munro): its life history and 
its fishery in South Australia, M.Sc. Thesis, Zoo- 
logy Department, University of Adelaide (un- 
published). 


salinity variations in the Coorong and their 
likely biological consequences, Comparisons 
are made with studies on other Australian 
coastal lagoon systems, including the Pecl— 
Harvey system in Western Australia (Hodgkin 
et al, 1980"), the Gippsland Lakes in Victoria 
(Poore 1982) and Lake Macquarie 
(MacIntyre 1959) and the Tuggerah Lakes 
(Powis & Robinson 1980; Collett et al. 1981), 
New South Wales. The Pecl—Harvey system 
is most similar to the Coorong as it also has a 
hyperhaline zone, 


Methods 

Phy sico-chemical 

Monthly visits were made to the North 
Lagoon from December 1981—March 1983, 
and samples were taken from 9 stations (Fig. 
1). Samples were collected from the South 
Lagoon in April (7 sites) and November (10 
sites) 1982, At each site conductivity (K.5) 
and temperature profiles were measured with 
a Martek Mark V Water Quality Meter. Light 
penetration was estimated with a Secchi disc. 
Surface and bottom (10 em above sediments) 
water samples were taken from stations 1, 3, 
5. 7 and 9 in the North Lagoon and from the 
sites sampled in the South Lagoon. In the 
laboratory the conductivity (Radiometer CDM 


PILYSICOCHEMISTY AND BIOLOGY OF THE COORONG 


% meter) and turhidity (Hach 2100A Tur- 
bidimeter) were measured, chloraphyll a con- 
centration was calculated by the SCOR 
UNESCO irichromatic method (Anon 1976) 
and total P determined by digestion with 
perchloric acid and measurement via the 
Stannous Chloride Method (Anan 1976). In 
addition to the samples collected in 1981-83, 
conductivity measurements were made on 
came earlier collections. 

Conductivity values were generally hyper- 
marine and so practical salinity could not be 
calculated from: the International Oceang- 
graphic Tables (UNESCO 1981). As an estt- 
mate of salinity, conductivities were converted 
to values for total dissolved solids (TDS) 
using the regression equation of Williams 
(1966). This equation was developed for Aus- 
tralian saline lakes but comparison of mea- 
sured TDS (dry weight of residue after drying 
at 102°C) for 30 sumples from the Coorong 
over the K. range 18 to 130 mS cme! with 
calculated TDS values showed close agree- 
ment (Fig. 2). This is to be expeeted consider- 
ing the similar nature of ionic dominance in 
Coorong water and Australian salt lakes 
(Williams & Buckney 1976). 


20 


{00 we 


Je] bid 


Poe) 
4 
% 


60 Po 


TDS. 


40 $ 


oO 


~ 


20 40 60 80 10020 
Conducrivity [K35) 
Fig. 2. Relationship between conductivity (Kas) 
and ‘TDS (%.). The line represents the regres- 
sion equation of Williams (1966), The points 


are values of Ke; and TDS by weight for 


samples from the Coorong lagoons. 


Biolagical 

Biological samples were taken from stations 
1, 3, 5, 7 and ¥ in the North Lagoon each 
month, avd from the siles in the South 
Lagoon, Mucroinvertebrates were collected 


ws 
es 


in three ways. Ekman Birge grab samples 
were taken from the mud in mid-lagoon 
(1.5-2.5 m depth) while in the littoral region 
a handnet was used among the muerophyte 
beds and an epthenthic trawl was pulled 
through the shallows (both nets with mesh 
size 0.5 mm). On return to the laboratory, 
collections were washed through a 0.5 mm 
sieve and animals picked out und preserved. 
Later each sampie was inspected under a 
sterodissecting microscope, specimens identi- 
fied and the relative abundance of species 
noted. Animals on hard substrates were col- 
lected occasionally but eryptic or small 
organisms on or under rocks were not cal- 
lected. No samples were taken from sand 
substrates on the Younghusband Peninsula 
shore of the lagoons. 

Samples of aquatic macrophytes were taken 
for identification and on same occasions fila- 
mentous algae were collected. Water samples 
were examined lo determine which algae were 
the dominant phytoplankters. 


Station Number 


FMA M J 
1962 1983 


JAS GON Du FM 


Fig. 3, Surface salinity (TDS ‘eo) at 9 stations in 
the Coorong, North Lagoon, over 15 months. 


Results 
North Lagoon 


Surface salinity at the 9 stations over the 
15 month period is shown in Fig. 3, In 
December 1981 salinities at stations 1-5 were 
below that of sea water, but all other salinities 
were hypermarine, A gradient persisted with 
salinities always lowest at station 1, closest to 
the Murray mouth, and highest at station 9; 


54 


salinities along the lagoon ranged 20-50%. in 
December 1981 and 40-80%. in January 1983. 
A seasonal pattern was apparent with salinitics 
high in summer 1981-82, falling during May, 
June and July and rising again in October to 
a peak value in January 1983. However the 
changes in salinity were not purely gradual 
seasonal ones; the abrupt rise in salinities in 
October was followed by a fall in November 
before a rise again in December and a rather 
abrupt fall in February-March 1983. There 
were also some occasions when a_ vertical 
salinity gradient existed with salinities at the 
bottom up to 5‘ above those at the surface. 
These variations probably reflect wind-induced 
currents, with either seawater entcring from 
the Murray mouth flowing southwards over 
saltier water, or highly saline water coming 


northwards along the bottom from the South 
Lagoon. 
eT i) Flax Point 7 50 
(2) Son Creek 
1901 | (3) Policeman's Point 40 
(4) Magrath Flat 
2c ® Needle Island (20 
(6) Long Point 
oo} | (2) Mark Pom oO 


TDS. Yo 


(6) 


i) T T 7 T 
1975 1976. 1977 1978 1981 i982 (983 


Fig, 4, Surface salinities (TDS “,) measured at 
various sites in the Coorong lagoons. Broken 
lines are used where samples were taken at 
long time intervals apart. Note the break and 
change of scale of the X axis. The values for 
Salt Creek, Magrath Flat, Long Point and Mark 
Point from May—October 1981 are calculated 
from field conductivity measurements made by 
the Engineering and Water Supply Department. 


Some information on longer term fluctua- 
tions in salinity is presented in Fig. 4, indicat- 
ing that salinities during 1982 were particularly 
high. In the mid 1970’s the North Lagoon was 
estuarine and from 1975-1977 the southern 
end of the South Lagoon (Flax Pt) showed 
fluctuations from around 40%-—70%-. In early 
1981 salinities were high in both lagoons but 
they fell to low values, particularly in the 
North Lagoon, prior to the commencement 
of the present sampling program. 


M. C. GEDDES & A. J. BUTLER 


(°C) 


257 
204 


Temperature 


m) 


( 


Secchi 


Turbidity (NTU) 


Total P_ (mg/m3) 


Station 1 
Station 3 
Station 5 
Station 7 
+ Station 9 


Chlorophyll a (mg/m) 


1983 


Fig. 5. Physicochemical measurements at 5 sta- 
tions in the Coorong, North Lagoon over a 15 
month period, Surface temperature did not 
differ significantly between stations. 


Water temperatures varied 11-25°C and 
there was never more than 0.5°C difference 
between surface and bottom temperature (Fig. 
5). Water was always clearest, as indicated 
by high secchi transparency and low turbidity, 
at station 1 and clarity decreased southwards 
(Fig. 5). There was no apparent. seasonal 
pattern to water clarity and it is likely that 
variations reflect local weather conditions. 
Total P showed no apparent pattern seasonally 
or along the lagoon (Fig, 5), The mean value 
was 79 mg m=". 

Chlorophyll « values were generally below 
10 mg m™“ (Fig. 5), highest values were at the 
southerly stations (7 and 9). At most times 
diatoms were the dominant algal group in the 
phytoplankton. In January, February and 
March 1982 and 1983 blooms of flagellates 
occurred at station 5, 7 and 9 and chlorophyll 
a tose to high tevels. 


PHYSICOCHEMISTY AND BIOLOGY OF THE COORONG 


Although no systematic collections were 
taken of macrophytes or filamentous algae, 
some comments can be made on their distri- 
bution and abundance. The macrophyte beds 
occurred on the shallow landward shelf where 
water depth was less than 1 m; occasionally 
patches occurred in deeper (to 2 m) water 
mid-lagoon, Ruppia megacarpa was the domi- 
nant macrophyte, occurring at all sites 
throughout the study. Lepilaena cylindrocarpa 
was common from stations 3-7. Zostera mutel- 
leri was common from stations 1-5 and a 
few plants were collected from station 7. 
From October to February mats of filamen- 
tous algae, including Cladophora, Entero- 
morpha and Oscillatoria, occurred among the 
macrophyte beds and covered much of the 
bottom of the lagoon, 

The macroinvertebrates collected in the 
North Lagoon are listed in Table 1. At stations 
1, 3 and 5 there were no seasonal patterns of 


{>< 
Station 1) 
Melita zeylanica 


+ ——__ Paracorophium sp. 


+ + Megamphopus sp. 
nn Ceratonereis pseudoerythraensis 
+ Nephtys australiensis 
Capitella capitata 
+ Ficopomatus enigmaticus 
———— Capitellides sp. 
Fabriciinae 
Hydrobia buccinoides 
Salinator fragilis 
Notospisula trigonella 
Arthritica semen 
Atherinosoma microstoma 
* Pseudogobius olorum 
+ Arenigobius — bifrenatus 
—— - Macrobrachium intermedium 
Tanytarsus pbarbitarsus 
Ephydrella sp. 


TDS. (%e) 
pf oO 
oo. 95 

ofp sts 


e4 
a4 


++ 
+ 
+ 


| 


< 
7 
z 
> 


TDS. (%s) 
a oo 
o. 6 

apres, 


: 


- — + 


Paracorophium sp. 
Capitella capitata 
Capitellides sp. 
Ceratonereis pseudoerythraensis 
+ Hydrobia buccinoides 
+ + Ficopomatus enigmaticus 
> ——— + Salinator fragilis 
Atherinosoma microstoma 
Tonytarsus barbitarsus 
Ephydrella sp. 


Fig. 6. Salinities (TDS %-) at 5 stations in the 
Coorong, North lagoon, over 15 months (sim- 
plified from Fig. 3), and presence of animal 
species at the two most saline stations, 7 and 9. 


a 
WW 


occurrence with most species collected on 
most occasions. Mean summer - salinities 
(October 1982—March 1983) at these stations 
were 38.8, 44.0 and 50.6% with peak values 
of 42.6, 50.5 and 57.6% (Fig. 6). At stations 
7 and 9 mean summer salinities were 61.5 
and 72.3%- with peak values 68.5 and 81.6%. 
Although many species were collected during 
the winter months, especially from station 7, 
as salinities rose most species became rarer 
and only Capitella capitata, Salinator fragilis, 
Atherinosoma microstoma and the dipteran 
larvae were common in the summer of 1982— 
83. 


Pseudogobius olorum 
Arenigobius bifrenatus 
therinosoma microstoma 
Amarinus lacustris 
Macrobrachium intermedium 
Haloniscus searlei 

Melita zeylanica 
Paracorophium spp. ies 
Megamphopus sp. SS 
Osticythere reticulata ——_—_—_—_—_——_ - 
Diacypris compacta 
Ceratonereis pseudoerythraensis —————————--_ -__ - 
Nephtys australiensis ————_ -- 
Ficopomatus enigmaticus 
Capitelia capitata 
Hydrobia buccinoides ————— ee ot 
Salinator fragilis _——_—___ 
Notospisula trigonella 
Arthritica semen 
Tanytarsus borbitarsus 
Ephydrella sp 


- T + T 
30 40 50 60 


T.D.S. 


an 
70 80 90 
(%s) 


Fig. 7. Distribution of animal species in the 
Coorong lagoons related to salinity (TDS ‘,). 
Broken lines indicate isolated collections of 
only one or a few specimens. 


The distribution of the fauna with respect 
to salinity is shown in Fig. 7. All species were 
collected up to 50% but above 55%. many 
species were absent. At above 65%- several 
species were represented by only a few 
individuals; only five species were collected 
above 70%. 


The distribution and abundance of each of 
the groups of animals collected will now be 
considered more closely. 


Fish 

Fish were collected in the trawl and the 
handnet in the littoral regions. Many fry were 
collected but not identified. The small adult 
fish comprised three species. The blue spot 
goby (Pseudogobius olorum) was most 
common from stations 1-7 over the salinity 
range up to 68%. The bridled goby (Areni- 
gobius bifrenatus) was collected less fre- 
quently, but over a similar salinity range (to 
65%.). Both gobies occur in estuaries and 


56 M. C. GEDDES & A. J. BUTLER 


TABLE 1. Macroinvertebrates and fish collected from the North Lagoon of the Coorong during 1982. 


Fish 


Arenogobius bifrenatus (Kner) 


Pseudogobius olorum (Sauvage) 
Atherinosoma microstoma (Giinther) 


Crustaceans Decapods 


Amarinus lacustris (Chilton) 


Macrobrachium intermedium (Stimpson) 


Amphipods 


Melita zeylanica Stebbing 


Paracorophium spp. 
Megamphopus sp. 


Ostracods 


Osticythere reticulata Hartmann 


Diacypris compacta (Herbst) 


Polychaetes 


Ceratonereis pseudoerythraensis Hutchings 


Nephtys australiensis (Fauchald) 
Australonereis ehlersi (Augener) 
Ficopomatus enigmaticus (Fauvel) 
Boccardia chilensis Blake and Woodwick 
Capitella capitata (Fabricius) 


Capitellides sp. 
Fabriciinae 
Gastropods 


Hydrobia buccinoides (Quoy and Gaimard ) 


Salinator fragilis (Lamarck) 


Bivalves 


Notospisula trigonella (Lamarck) 


Arthritica semen (Menke) 
Tellina mariae (Tenison-Woods) 
Soletellina donacioides Reeve 


Chironomid 
Ephydrid 


Insects 


Tanytarsus barbitarsus Freeman 
Ephydrella sp. 


——- — eer 


coastal lakes from Queensland to Western 
Australia (Hoese & Larson, 1980). The small 
mouthed hardyhead (Atherinosoma  micro- 
toma) was collected from all stations and was 
particularly common at 7 and 9. This species 
also occurred in the South Lagoon at salinities 
up to 100%, A. microstoma is common in 
shallow lagoons and brackish water lakes 
from the Tuggerah Lakes in N.S.W. to the 
Coorong (Ivanstoff 1980), 


Crustaceans 


The small spider crab Amarinus lacustris 
was collected on two occasions from station 3 
and once from station 5 at salinities of 38.2, 
43.3 and 53.2%. This crab is normally 
restricted to fresh or slightly brackish water 
with other species of Hymenosomatidae 
usually found in estuarine and marine situa- 
tions (Walker 1969; Lucas 1980). The 
shrimp Macrobrachium intermedium was col- 
lected in handnet samples from among macro- 
phyte beds on ten occasions from stations 3, 
5S and 7 in salinities ranging up to 63%. The 
species is common in weedbeds in the S.A. 
gulfs (Hale 1927) and in estuaries in south- 
eastern and southwestern Australia (Poore 
1982; Hodkin pers. comm.). 

Amphipods were very abundant on the 
plants and mud jn the littoral regions. Three 
species occurred in high abundance, Melita 


zeylanica, Paracorophium sp. 1 and Megam- 
phopus sp. at stations 1-5. A few specimens 
were collected from station 7 during winter 
and spring but none were collected in January 
or March 1983. The distribution and salinity 
tolerance of the amphipods is discussed in 
Kangas & Geddes (in press). 

The ostracod Osticythere reticulata was 
common in many collections at salinitics up to 
60%. and a few specimens were collected at 
72.4%, This is an estuarine species described 
from the Hopkins River and the Gippsland 
Lakes, Vic. (Hartmann 1980). Diacypris com- 
pacta, a salt lake ostracod recorded from 
ephemeral salt lakes along the Coorong (De 
Deckker & Geddes 1980), was collected on a 
few occasions. 


Polychaetes 

Of the several species of polychaetes listed 
in Table 1 only four were common. Cera- 
tonereis pseudoerythraensis was abundant in 
all benthic samples from stations 1, 3 and 5 
and Nephtys australiensis in benthic mud 
samples from those stations. The former is 
known from estuaries and embayments from 
S.A., W.A. and N.S.W. (Hutchings & Turvey 
1982) while the latter is found in sand and 
mud substrates in bays, lagoons and estuaries 
from southern Qld to Spencer Gulf (Rainer & 
Hutchings 1977; Hutchings 1982). Most 


PHYSICOCHEMISTY AND BIOLOGY OF THE COORONG $7 


records of CL psendovrythraensis and Ne ats- 
tralicasis were from below 65 and 57% respec- 
lively, alfhough some speeimens of the tormer 
species were collected at salinities us high as 
72) and same of the latter species at 64%. 
Capifella capituta was collected occasionally 
from stations 3 and 5 but was mest abundant 
at stations 7 and 9 at salinities Up to 824 
it was the only polyehucte at stations 7 and 
9 in the summer of 1982-83, Lt ix recorded 
tram W.A, along southern shores to N.S.W,, 
in rnuddy sediments in estuarine and shallow 
protect areas (Hutchings 1982). ‘The cosmo- 
politan brackish water serpulid Fieopomtarus 
eiivinaticuy was common in the littoral areas 
ot stations 1, 3 and § and large mounds of 
live tube worms occurred, Mounds of enypty 
tubes were common at stations 7 and 9 “The 
highest salinity at which large populitions of 
aclive warms were scen was 67%. but some 
newly setthed solitary individuals were found 
at higher salinities. 


Mollases 


Of the gastropods, Hydrabia buecinoidey was 
extremely abundant in the littoral macrophyte 
beds at stations |, 3 and 35 throughout the 
study, at station 7 until December 1982, and 
ul station 9 during the middle of 1982, The 
highest sulinily at which large populations 
occurred was 66!5 although a few individuals 
were collected up to 74, Salinater fragilix 
was present in lower numbers but it persisted 
wt station 7 throughout the study and was 
often collected from station 9 at salimties up 
to 74%, Species of Aydrobia are characteristic 
of estuarine and lagoonal environments in 
Europe (Barnes 1980, p. 59) and A. bucet- 
noides Was common in the Tuggerah Lakes 
(Collett e¢ al, 1981). Salinapor frayilis is 
common in estuaries and sheltered embay- 
ments if southeastern Australia (Poore 1982, 
Collett et af. OSL, Butler ef al. 1977), 


The most common bivalve was Nofospivida 
winonela which was collected from. stations 
1, 3.and 5 throughout the study and station 7 
until December 1982; highest salinity for the 
species wis 66%), At station 9 there were 
extensive deposits of empty shells, The small 
hivalve Artiritica senten generally was col- 
lected along with Norosplyla. The other 
species of bivalves. were collected only ocea- 
sionally, Notospivale tigonelta is recoynized 
as ou widespread estuarine species in southe 
enstemm Austealin estuaries (Poore 1982) and 


itand Arhvitica semen are among the domir 
nunt bivalves in the Peel-Warvey system 
(Hodgkin es al. L980)", 


Dipreraiy 

The chironomid Tanyrarsuy — barbitarsus 
oveurred occasionally all along the lagoon but 
very large numbers were collected at stations 
7 and 9 from October 1982 to Mareh [983 
when they were casily the most numerous 
invertebrate, At stations 7 and, especially, 9 
Kplivdrella sp. and a few specimens of Cera- 
fopovonidae were collected, These dipterans 
are characteristic of inland saline lakes 
(Williams 1981). 


ts ee a ee, a ES EE I 
gt ee gg ge ut ey * 
s cp ge es a eat ag 
Ca gs a & ae 
“ ov a ge! ot 


Fig. 8. Salinities (TDS ‘i, ) measured on Iwo ocea- 
sions at various sites in the Cooropgz, South 
jugoon, Open circles: 16 and 17, April 1982. 
Closed circles; 9 and 10, November 1982, 


Sauth Lagoon 


The South Lagoon was sampled on two 
oceasions and the salinities are shown in Fig. 
8, On both oceasions the salinity was rather 
similae from Stony Well Island southwards 
24 km to Salt Creek, at values around 97 ir 
in April-and 93%. in November. This indicates 
that the lagoon was well-mixed and/or that 
it had little exchange with the North Lagoon 
wid reecived no significant inputs of fresh 
water, 

Mean total phosphate was (10.5 mg nr 
(SD 15) in April and $7.9 mg m~ (SD 23.6) 
in November, Mean scechi depths were 45 cm 
and 30 em and mean chlorophyll a values 
21.6 me me! (6.3) and 29.2 mg m* (5.0). 
There were no elear trends in any of these 
parameters along the lagoon on either sampling 
Uitte. 


7 Hodgkin, FE, P., Birch, P. Bu Black, R. E., Hum- 
phries, R, B, (1980) The Peel-Harvey Fstuarine 
System Study (1976-1980) ReporL_ No, 9, De- 
partment of Conservation and Environment, 
Western Australia, 72 pp. Cunpublished), 


58 M. C. GEDDES & A. J. BUTLER 


8) Murray Mouth 


sa aia ioe fi 10 20 30 40 
B. = 15m4 we 26-11-75 
¢ 
ah at = 
80 


eat be Mouth 


36! ‘ 40 
Cc. 15 23-11-81 
! 
L 


8 Murray Mouth 


s 70 180 90 100 
D. 15m 10-11-82 


8 Murray Mouth 


(MI x 1078) 


Monthly Barrage Flow 


150 


100 


50 


Monthly Rainfall (mm) 


1973 1974 1975’ «#1976 1977 1978 1979 1980 1981 " 1982 


PHYSICOCHEMISLY AND BIGLOGY OF THE HOROSG Su 


No live Reppia wos collected on either 
trip although small yuantities of dead Ruppia 
lowves, presumably KR, mberoxa, were collucted 
in the littoral samples, The only fish collected 
from the South Lagoon was Adherineselna 
mierustariay it occurred in very large numbers 
and was collected ai nost stations. The erustite 
evans comprised the isopod Halonivens scarlet 
and the osttacod Dizevpris compacta, a Tew 
specimens of Cypridus quxtraliensis and Reti- 
cyprly sp, were also collected, Dipterans wete 
the Most HuUMeroUs animals and included cers- 
topogonids,  tabanids, — strattamyids and 
especially the chironormad Turyrarsiy uth 
fersuy and the ephydrid Ephydrelly sp. On 
mud flat areas, partigularly south of Salt 
Creek, the gastropod Codielfa sp, and the 
beetles €lavinia sp. (Carabidae) and Blecius 
sp. (Staphylinidse) were abundant, 


Discussion 


‘The Coorony lagoons ure characterized by 
the variety and degree of their salinity Muc- 
inatiins, Salinity varies alone the length of 
the Coorong, On ogeasions there gre vertical 
swinity gracdtents, and there are seasonal and 
long-term patlerns, A fange of salinity pat- 
terns is represenicd in Fig 9 which shows 
horizontal and vertical gradients in salinity at 
different occasions sinee 1973. Gn cach ocen- 
sian there is a gradient tawards hypermarine 
sultities in the South Lagoon. This suggests 
that the dominawat freshwater influence on the 
Coorong is trom the River Murray. barrages. 
Salinity values in the South Lagoon vary 
widely Hom 400) to more than 100%,. When 
the barrages are open or only recently closed 
(Fig, 94,B,C), a vertical salinity profile exists 
al the northern end of the Nerth Lageon. 

‘Yhe salinity pullers can be interpreted with 
respect to outflow from the River Murray 
barrages add to rainfall itv the southern region 
of the Coorong (Fig, 9). The high River 
Murray flows tp 1973-75 resulted in the 


Cie, 9. Horizontal and vertical patterns of solinity 


and the relationship between Courong sulinity 


bariages being open almost cormlintiously pro- 
viding fresh water al the Murray Mouth, The 
fresh walee was ayvuilable for mixing to the 
south, maintaining estuarine conditions over 
most of the Coorong in 1975 (Pig. 9B). These 
Jow sulinitics occurred even though rainfall 
in the southern area of the Coorong was holow 
average in 1974 and 1975. For must months 
in 198O-and the first part of 198) the barrages 
were elosed, salinities af the mouth would 
have beet at about that of sca wile and 
southward the Coorong would have beconre 
progressively more hypermarine, “The brict 
period of high Now trom the Murray in July 
to October L981 provided some dilulian. of 
the North Lugoon, but seems to have had 
little effect on the South Lagoon (Fig. 9C) 
The barrages were then closed for the 12 
months up to November 1982. sea water 
would have re-entered the mouth afea, and 
galinities alone the lagoons became progres 
sively more hypermarine (Fig. YD) The 
above-average rainfall in the southern 
Coorong area im TYSt did not reduee salinilies 
significantly. The important role of freshwater 
from the River Murray in controlling salinities 
in the Coorong is also apparent in earlier 
periods. The high salinities in the late 1960's 
and carly 1970's which pronypted much 
environmental concern (Noye 1975), followed 
low flaws from the River Musray and long 
periods of barrage closure from 1965 to 1969, 

Although there is a relationship between 
River Murray flow and salinity in The Coorong 
lagoons, it may be that the period of lime lor 
which lreshwater is avatlahle at the Murray 
nrouth 1s mere important than the flow per se 
Fiows were hish in mid-1981, providing an 
abave-jverawe flow for the year 1981-82, but 
(he barrages were open for only a few months 
and the availability of fresh water for this 
short periad did not bring about « geweral fall 
in salinities in the southern parts of the 
Coorong, When there js Fresh water at the 


wong the Coorong kiwoons ut four diferent periods 
and River Murray outflow vin the barrages and 


raintall in the southern Coorong region, The four salinity profiles are taken from the Murray mouth 
io Bul Bul Lagoon and correspond to the mip on the top of the Fivurc, Broken lines ure poes- 


sible values. Burrage outflow valves are estintated by the 
when salinity profiles were taken. 

water overlies saline water in the North Lagoon, 
urragey the North Laguow is esiugrine and the Sourh 


ment, Arrows ou the flow diagno show Himes 
A. Following opening of the barrages fresh 


ft. After lwo years of goad flow from the b 


Lagoon is al about seawater salinities, 
©. Following closing of the barrages. 
Lignan % estuarine. 


Engineering und Water Supply Depart- 


sed witter intetdes through the mouth; most of the North 


D, Afier a lung period af birrage closire the Coorome. bevomes & hypernuume sysigm with 
salinities up to three times seawaier in’ the South Liapwun, 


60 M. C. GEDDES & A. J. BUTLER 


Murray mouth it can be moved southeast and 
mixed by wind action (Noye & Walsh 1976), 
thus 


gradually reducing salinity in more 
southerly areas of the Coorong. Lengthy 
periods of continuous outflow from the 


Murray may be necessary to. significantly 
reduce salinity in the South Lagoon. The 
complex relationship between River Murray 
flow and salinity in the Coorong needs further 
investigation. 

The extensive beds of macrophytes, 
generally dominated by Ruppia, are important 
in the ecology of the Coorong. The macro- 
phytes are covered with epiphytic growth 
which probably is grazed by many of the 
invertebrates and they provide shelter for the 
invertebrates and the small fish. In the North 
Lagoon Ruppia megacapa was abundant from 
station 3 southwards, This species is common 
in estuaries and lagoons in southeastern and 
southwestern Australian (Brock 1982a), The 
plants continued to grow in the North Lagoon 
while salinities were above 60%- and as high 
as 80%, These values are above the salinity 
range of 12-50%» found by Brock (1979)8. A 
second species, R. tuberosa, has been recorded 
from the South Lagoon. This is an annual 
species found in shallow and ephemeral saline 
water in 8.A., W.A. and Vic. (Brock 1982a). 
It regenerates from sexual perennating organs, 
the turions, and it also produces large amounts 
of seed (Brock 1982b). This species appears 
to be particularly important to the bird life 
in the Coorong as ducks and swans graze the 
plants, especially the starch rich turions 
(Delroy et al. 1965)8, and the seeds and 
turions form a major food source for many 
of the waders (Paton 1982)*. R. tuberosa 
occurred in many salt lakes in the southern 
Coorong area and at Flax Point in the South 
Lagoon from 1975 to 1978 (Brock 1979)8. 
The salinity range was 13-230; however 
there is little information on the range of 
salinitics at which germination or regeneration 
occurs. No growing R. tuberosa was collected 
on either of the trips to the South Lagoon jin 
1982, and it appears that because of the high 
salinities this very important plant was rare 
or absent, 

The fauna of the North Lagoon was similar 
to that recorded in other coastal lagoon 
systems including the Tuggerah Lakes, N.S.W., 


S Brock. M. A. (1979) The Ecology of Salt Lake 
Hydrophytes. Ph.D. Thesis, University of Ade- 
laide (unpublished). 


the Gippsland Lakes, Vic., and the Peel— 
Harvey system, W.A. The dominant macro- 
invertebrates in the Corong were Notospisula 
trigonella, Hydrobia buccinoides, Melita zey- 
lanica, Paracorophium spp., Megamphopus sp., 
Ceratonereis pseudoerythraensis, Nephtys 
australiensis and Macrobrachium intermedium 
and all are recorded in one or more of these 
other lagoons. The amphipods seem to be par- 
ticularly important in all of these coastal 
lagoon systems. Ficopomatus enigmaticus was 
abundant in the Coorong but not collected in 
the other studies, possibly because collecting 
concentrated only on the infauna. Only 21 
species were collected in the Coorong com- 
pared to numbers in excess of 100 in many 
large estuaries in southeastern Australia 
(Saenger et al, 1980; Rainer & Fitzhardinge 
1981). In the Gippsland Lakes 90 species 
were recorded (Poore 1982). Although col- 
lecting effort was probably lower in the present 
study, the fauna does seem to be considerably 
Testricted and many species common in 
estuaries were absent. This low diversity may 
reflect the extreme fluctuations in salinity in 
the Coorong. Species number is also low in the 
Peel-Harvey System, and especially in the 
Harvey estuary section where salinities vary 
most widely (Hodgkin et al. 1980)7. The 
species assemblage in the Harvey estuary is 
similar to that of the Coorong, including 
Arthritica semen, Notospisula — trigonella, 
Salinator fragilis, Capitella sp. Ceratonereis 
pseudoerythraensis, Melita sp. Paracorophium 
sp. and chironomids (E.P. Hodgkin pers. 
comm.). 

The species in the Coorong perhaps repre- 
sent that group from within the broader 
estuarine fauna that is most euryhaline. All of 
the species showed especially wide salinity 
tolerance and all persisted throughout the 
year thus displaying wide temperature tole- 
rance as well. Our data suggest that for Ama- 
rinus lacustris, Melita zeylanica, Paracoro- 
phium sp., Megamphopus sp., Osticythere 
reticulata, Hydrobia buccinoides, Notospisula 
trigonella, Arthritica semen and Ruppia mega- 
carpa, the salinities tolerated by the popula- 
tions in the Coorong may be higher than those 
recorded elsewhere. Although all species 
recorded were present at salinities up to 55%, 
in the hypermarine conditions in the Coorong 
in [982 increasing salinity along the lagoons 
did restrict the distribution of the fauna. When 
salinities in the North Lagoon reached levels 


NHYSICOCHEMISTY AND BIOLOGY OF THE CUOROMNG fl 


above 70% the character of the fauna changed, 
with the estuarine assemblage /eplaced by one 
dominated by haluphilie dipterans and salt- 
lake crustaceans. 

The salinity regime i [982 represents an 
extremely hypermarine phase ia the long-term 
salinity fluctuations of the Coorong lagoons, 
and this is reflected in the restricted distribu- 
tion of the fauna. Mast members of the fauna 
have good dispersal powers and at times of 
lower salinity) their distribution would’ pre- 
sumably be expanded. In the South Lagoon 
carlier oceurrence of an estuarine-lagoonal 
fauna was evidenced by mounds of tubes ol 
Ficapomatiy eritgmerieus aud shells ol Noto- 
spisule miganella, Purther study is needed 10 
clucidate the conditions under which recoloni- 
vation of the South Lagoon might proceed, 


Acknowledgments 


We thank Julie Francis, Helen Vander- 
woude and Mervi Kangas for help in the field 
and the laboratory and the following people 
fer identification of specimens, Pat Mutchinys: 
(polychaetes), Margaret Drummond (amphi- 
pods), Patrick De Deckker (ostracods), Lance: 
Lloyd (fish) and Margaret Brock and Enid 
Robertson (planix). The Engineering and 
Water Supply Department provided unpub- 
lished data on outflow from the Murray bar- 
rages and on salinity in the Coorong. Thanks 
to Saundra Lawson and Heather Kimber for 
typing the manuscript and to Ruth Altmann 
for the artwork. This study was supported by 
an MST grant which is gratefully acknow- 
ledped. 


References: 


ANON L1976) Standard methods for the examinu- 
lion of water ond wastewater. [4th Edition, 
(American Public Health Associauion: Washing 
tot.) 

Barnns, R. 8. KB, 
natural history of a neglected habitat 
bridge University Press: Cambridge, 

Brock, M. A, (19824) Biology of the salinity 
tolerant genus Ruppia L. in saline takes in South 
Australia, | Morphological variation within ane 
between species und ecophysivlogy. Aqdur Mer 
13, 210-48, 

(1982b) Biology of the salinity tolerant 
cenus Reppia L. iq saline lakes in South Aus- 
tralia, IL Population ecology and reproductive 
hidlogy. Lbjd. 13, 249-68. 

BeTLER, A, -, Derrers, AWM. Mckaaur, S.C. 
& Thomas, D, PL 41977) A survey of mangrove 
forests in South Australia, 8) dast Nat Sits), 
34-49. 

Comm. LC. Ceruins, A, J. Cipes, Pod, & 
Wesr, R. J. (1981) Shallow dredging as is 
strategy foe the control of sublittoral mucro- 
phytes: a case study in ‘Tupgerah Likes, New 
see Wales. Ansr. J. Mar. Freshwater Res. 32. 

(4-7 t- 

Dr Drerxve, P. & Gepprs, M, C. 119801 
Seusunal fuuna of ephemerul saline lukes near 
che > tabi Lagoon, South Australia. fbi. 31, 
ay7-89, 

Hace, H) M. (1927) The Crustaceans of South 
Australia. (Government Printer: Adelaide.) 


Hare, J, A, C1968) The yellow-eye mullel [rom 
the Coorong Lagoon, South Australiu. Travis, 
R, Sor, S, dust, 92, 37-50, 

Hagraann, G, Cl980) Die ostracoden der Ord- 
nun Podocopra G. W. Mueller, 1894, der 
warm temperticrien und  subtropikisch-tropi- 
Kyschen kusten-ubschmitte de Sucl-und sudasi- 
kusic Austrahens (Zwischen Ceduna in Western 
une lakes Enttanee in Osten), Mitr Mah. 
fool Mus, dost, TT. 111-204, 

Horse, DF) & Lassen, TK. F198) Rarily 
CGiobiidae Ciabres, Chupler 32. fa RM. 


(1980) Coastal laggans A 
(Cam- 


McDowell (Ed) “Freshwater Fishes of South- 


eustern Australia.” pp, 186-92. (Reed: Syd- 
ney.) 
Hurcuines, Po (1982) Aristlewormy (Phylam 


Annelida). fi S, A. Shepherd & 1 M. Thomas 
(Rds) "Maring Invertebrates of South Anstralia 
Part 1.” (Government Printer; South Australia, 
pp. 228-98, 

—— & Tunverx, S. PB. (1982) The Neteldidae of 
South Australia. ‘Trevi, R. Soc, 8. Awst 106031, 
4.1.44, 

Lvansrorr, W.. (1980) Family Atherinids. Silver- 
sides or Hardyheads. Chapter 20. da RM, 
McDowall (Ed.) “Freshwater Fishes of South- 
eastern Australia.” pp. 132-37, (Reed) Syd- 
ney.) 

Kanaas, M, & Gepprs, M, C. (in press) The 
effects of salinity on the distribution of smphi- 
pods in the Coorong, South Australia, in relas 
lion to thelr salinity tolerance. Trans. R. Soc, 
SN, Aust. 108. 

Lucas. |. S, (1980) Spider crabs of the fumity 
Hymenosomatidae (Crustacea; Broachyora) with 
particular reference to Australian species Kee. 
Aust, Mus. 33, 143-247, 

MacInryre, R. J, (1959) Some aspects of the 
ecology of Lake Macquaric, N.S.W., with 
regard to an alleged depletion of fish. VIT The 
benthic macrofauna. Aust. Jo Mar, Freshwater 
Res, 10, 341-53, 

Novy, B. J. (1975) The Coorong, (Department 
ol Adult Education: University of Adeliyide.) 
Publ, No. 39, [31 pp, 

—— & Warsn, Put. (1976) Wind-induced water 
level ogcillations in shallow. lagoons, das, J. 
Mar. Freshwater Res, 27, 417-30, 

Poort, G, C. B. (1982) Benthic communities of 
the Gippsland Lakes, Victoria, Pid, 33, 90115, 

Powrs. M. J. & Rosson, KOT. M. (1980) Beit- 
thie macrofaunal communities if the Tuggersh 
Lakes, New South Wales. Jbid- 31, 803-15 

Rainer, 8. & Frrzearoinar, RoC, C1981) Benthic 
communities in an usiuary with) perinidic de 
oxygenation. Uhid. 32, 127-43. 


62 M. C. GEDDES & A. J. BUTLER 


—— & Hurcnines, P. (1977) The Australian 
Nephtyidae. Rec. Aust. Mus. 31, 307-47. 


SAENGER, P., STEPHENSON, W. & MOover_ey, J. 
(1980) The estuarine macrobenthos of the Cal- 
liope River and Auckland Creek, Queensland. 
Mem. Qld Mus. 20, 143-61. 


UNESCO (1981) International Oceanographic 
Tables. UNESCO Technical Papers in Marine 
Science. No. 39, Vol. 3. United Nations Edu- 
cational, Scientific and Cultural Organization. 


VON DER Borcu, C, C. (1965) The distribution 
and preliminary geochemistry of modern car- 
bonate sediments of the Coorong area, South 
Australia. Geochem. et. Cosmochim. Acta 29, 
781-99, 


—— (1975) The geological history of the 
Coorong Lagoon. Chapter 2. In J. Noye (Ed.) 
“The Coorong” (Department of Adult Educa- 
tion: University of Adelaide.) 

WALKER, K. F. (1969) The ecology and distribu- 
tion of Halicarcinus lacustris (Brachyura: 
Hymenosomatidae) in Australian inland waters. 
Aust. J. Mar. Freshwater Res. 20, 163-73. 

WILLIAMS, W. D. (1966) Conductivity and the 
concentration of total dissolved solids in Aus- 
tralian lakes. Ibid. 17, 167-76. 

—— (1981) The limnology of saline lakes in 
Western Victoria. Hydrobiologia 82, 233-56. 

& BucKNEYy, R. T. (1976) Chemical com- 

position of some inland surface waters in south, 

western and northern Australia. Aust. J. Mar. 

Freshwater Res. 27, 379-97. 


SEDIMENTOLOGY AND ORIGIN OF LATE PALAEOZOIC GLACIGENE 
DEPOSITS AT CAPE JERVIS, SOUTH AUSTRALIA 


BY N, F. ALLEY & R. P. BOURMAN 


Summary 


The most extensive exposure of the Permian glacigene Cape Jervis beds was investigated in the 
type area, Cape Jervis. The beds are divisible into five units which are interpreted, from the base 
upwards, as fluvioglacial and glaciolacustrine sediments, lodgement till, fluviolacustrine deposits, a 
flow till complex and a glaciomarine unit. The basal beds are considered to be the proglacial 
deposits of an ice-mass advancing northwards along the present Backstairs Passage trough. At its 
maximum extent the ice over-ran Cape Jervis and plastered lodgement till over a partly frost- 
shattered bedrock surface. Deglaciation was marked by the development of a kame terrace on which 
fluviolacustrine sediments and a flow till complex were deposited. Glaciomarine silts and clays 
were laid down during a late deglacial marine transgression into an isostatically depressed coastal 
lowland. 


SEDIMENTOLOGY AND ORIGIN OF LATE PALAEOZOIC GLACIGENE 
DEPOSITS AT CAPE JERVIS, SOUTIT AUSTRALIA 


by N. FP, Aniey® & R, P. BouRMAN? 


Sunmiary 


Airy, N. PB, & BougmMAn, RB. P. (1984) Sedimentology and ofgin of Late Palaeozoic glaci- 
pene deposils at Cape Jervis, South Australia. Trans. R. Sec. S. Anst. UBC), 63-75, 12 


June, 1984, 


The most extensive exposure Of ihe Permign glacigene Cupe Jervis Beds was. investi 
muled in the type urea, Cupe Jervis, The beds are divisible into five units whith are inter 
preted, from the base upwards, ws Huvioglacial and glaciolacustrine sediments, lodgement till, 
fluvidlacusiine deposits, w Now till complex and w glaciomarine unit, The basal beds are 
considered to be the proglacinl deposits of an ige-mass advancing northwards along the present 
Backstairs Passage trough. AL its masimiin) extent the ice over-ran Cape Jervis aad plastered 
lodgement till over a partly frost-shattered bedrock surface. Deglaciation was marked by the 
development of a Kame terrace on which fluyiolacustrine sedimenis and a flow till complex 
were deposited, Glaciomuarine silis and clays. were laid down during a late deglacial murine 
Lhanspression into an isostaticully depressed coastul lowland, 


Key Worbs: Cape Jervis. Late Pulavozoic glaciation, Jithostrarigraphic units, lodgement 
WL Now till Huviolacusirine sediments. glactomarine deposits. 


Introduction 

The results presenied in this paper form part 
ol a larger investigation of Late Palacozoic 
tlacigene deposits in southern South Aus- 
tralia, The paper reports om the results of a 
reexamination of the type area of the Cape 
Jervis Beds at Cape Jervis (Ludbrook 1967) 
and a sedimentary model is proposed to 
uccount far the facies ohserved, 

We have retained the original designation of 
“Cape Jervis Bedy" from the holostratotype 
section (Ludbrook 1967) rather than assign 
ing them “Formation” status. The latter may 
be considered when the extent of the beds Js 
better known following an examination of 
other facies exposed at parastratolype sections 
or intersections in boreholes. 


Previous Investigations 

The Cape Jervis Beds in the type area 
extend north afd east of the Cape Tervis 
lighthouse (Fig, 1) see Fig. 10 for general 
location), oceupying N-S trending bedrock 
depressions, AL te coust the beds form part 
of a cliff that has been extensively croded 
(Pig. 2), thus leading to the concentrations 


* Geological Survey, South Australian Department 
of Mines & Energy, Box 151, Easiwood P,O., 
South Australia 5063. 

7 Dept of Soil Science, Waite Institute, Univer- 
sitv of Adelwide, Urrbrae. 

Published with the permission of the Director- 

Gener, Depurtineot of Mines & Energy. South 

Ausiralia, 


of lurge erratics hoth in the more recent 
alluvial cover and in the near shove zone. 

Exposure of the Late Palaeozoic glacigenc 
sediments at Cape Jervis has long heen known, 
but so far studies of them have failed to recog- 
nise the complexity of the sediments and the 
association of (heir characteristics with distinc» 
tive glacial depositional environments. 

The significance of these beds wis. first 
established Jate last century (David & Howchin 
1897). A report on the Cape Jervis area 
(Glacial Research Committee (898)  con- 
sidered the glacigene sediments there to 
represent the most extensive occurrence of a 
genuine till “yet discovered in South Aus- 
tralia”, The till was described as an unstratified 
deposit, varying in colour from grey to almost 
black, and full of boulders of all sizes. Tb wis 
estimated to exceed 30 in in thickness and was 
cupped by “variegated (Miocene) chiys™, now 
known (oO be glacio-nurine sediments (Lud- 
brook 1967), The committee invoked the 
influence of shore-ice to explain some of the 
characteristics of the deposits, particularly the 
number of walerworn but striated pebbles 
found i the drift. 

Based on these observations and other evi- 
dence Benson (1911) proposed a glacial 
origin for Buckstaits Passage, a proposal sup- 
ported by many subsequent workers (Madigan 
1925; Campana & Wilson 1955; Bauer 19591; 


'Bauer, FM. C1959) Regional eeogriphy of 
Kangaroo Island. Ph.D. thesis, ALN.U. Cunpubl,). 


64 N. FP. ALLEY & R. P. BOURMAN 


LEGEND 
QUATERNARY Calcrete 
PERMIAN Cope Jervis Beds 


CAMBRIAN Kanmantoo Group 


A Type section of Ludbrook (1967) 
B Measured section (this paper) 

Topographic. contours 
(50m interval) 


Fabries on 


Lodgement Till SITE’ 


CAPE JERVIS 
Lighthouse 


83-399 DEPARTMENT GF MINES. AND ENERGY —SOUTH qusTRALIA 


Fig. |. Geologieal sketch of Cape Jervis urea (modified from Ludbrook 1980) shawing position of 1ype 


section and new meitsured section. 


Bourman 1973"; Daily et al. 1979), During 
the course of regional geological investigations 
on Fleurieu Peninsttla Campana er al. (1953) 
noted the exposure at Cape Jervis and 
deseribed the beds as moruaines’ consisting 
“mainly of unsorted, unstratified boulder clay 
and sundy clay, containing erratics of all sizes 
and degree of angularity” (p. 5). They also 
interpreted the rhythmically bedded materials 
containing dropstones as varves. However, 
from these local and more extensive studies on 
Fleuricu Peninsula they did not establish any 


*Bourman, R. P. (1973) Geomorphi¢c evolution 
of southeastern Fleuriew Peninsula. M.A. thesis, 
Univ, Adel, (unpubl.). 

“The term “moraine’ is normally restricted to 
topographic forms rather than to deposits. 


clear lithostratigraphic succession in the beds 
apart fram recognising the abundance of 
boulders and pebbles near their base and sands 
in their upper sections, 

Deposits. of till some 20 m thick with inter- 
bedded thinly laminated silt and clay beds 
were described at Cape Jervis by Brock 
(1964'). However, in his map and fizure 
descriptions, bedded sediments were labelled as 
tills (Brock 19644, fig. 6 and plate 12), 

The most definitive work is that of Lud- 
brook (1947) who established the site as the 
holostratotype section of the Permian Cape 


‘Brock, E. J. (1964) Denundation chronology of 
Fleurieu) Peninsula, M.A. thesis, Univ. Adel. 
(unpubl, }. 


CAPE JERVIS BEDS, SOUTH AUSTRALIA OF 


j ae 

Fis, 2, Coastal slope at type urea. Steeply dipping 
Kanmanlog Group mictasediments form shore 
plattoem and steep clits in distance. Positions of 
features shown by symbols arc; G, gully of mei- 
sured section; BD. earthen dunt; BE, line of large 
erratics paralleling Wow till complex unit. View 
looking. northeast, 


Jervis Beds of the St Vincent basin and auljoin- 
ing -areas. From the base, the seetion is 
deseribed as: (1) bouldery, sandy till (6.7 m 
in thickness) unconformably overlying Cam- 
brian Kanmantoo Ciroup bedrock, (2) dark 
prey boulder till (1.5 m); (3) gritty send- 
stone with boulders and interstratified clay 
(3.9 m); (4) crosshedded sandstone (1.5 m); 
(5) clay shales with grits and boulders (9.0 
a}: (61 and obscured sedintents, but pro- 
bably a continuation ef the underlying clays 
(7.2 m). Using the presence of arenaceous 
foraminifera in the clay shale unit 1.adbrook 
determined a Lower Permian (Sakmarian) 
age and a murine influence in the environment 
of deposition for at least the upper part of the 
sequence. Harris (1971)9 curried out a further 
stratigraphic study of Ludbrook’s type section. 
It is apparent from his descriptions that the 
“fills” do not appear to have the characteristics 
of true lodgement till particularly in’ his 
second unit. From the base upwards, Harris 
(1971) interprets the sediments us: (11 dia- 
mictite: clay matrix with large erratics (6-7 
m); (2) slumped till with slump rolls of 
sandstones (1,5 mm); (3) boulder beds in 
sunds, claystones and cross-bedded sands (3.9 
m); (4) claystones and sind beds with pro- 
minent yellow Fontainbleau sandstone, pebbly 
in places (1.5 m); (5) well laminated grey 
clay containing arenaccous foraminifera and 


hiferix, Ro P. (1971) The geology of Permian 
secliments and erotics, Proubridge Basin, South 
Austowy, @&Se. Hons, thesis, Giiv Adet, 
Cunpubl, |, 


dispersed ertatics (9.0 m); and (6) poorly 
exposed but probable boulder clay at surface 
{exceeds 7 m), 

Methods 


Despite their great antiquity, the general 
un-lithilied nature of the Late Palagozoie sedi- 
ments in South Australia permits the use of 
techniques normally employed on Plefstocene 
ilacial deposits. Pebble counts of clasts greater 
than 5 mm diameter have beech undertaken to 
characterise individual tills and le indicate the 
possible provenance of the clasts. Although 
some doubt exists about the statistically accept- 
able number of pebbles required to adequately 
characterise sediment fithologics for each 
lithostraligraphic unit, we have used the 
usuully ueceptable minimum population of 300 
(Dryden 1931: Krumbein & Pettijohn 1938; 
Hubert 1971). Till fabric analysis was utilised, 
in the absence of small scale crosional fea- 
tures, to determine the direction of ice move- 
ment. This technique has also served as an 
aid in establishing the origin of certain dia- 
inictons and at Cape Jervis helped to distin- 
guish lodgement and flow tills, 

Till fabric analysis rests on the assumption 
that the long axes of certain pebbles reflect 
the dircetion of ice flow at the base of the 
glacier during their deposition (Holmes 1941). 
The orientation of a minimum of 50 pebbles 
represents & statistically acceptable population 
(Harris 1969; Lindsay 1970), Pebbles selected 
for analysis were larger than 7 ent abd were 
characterised by the ratia between a and b 
axes being 3:2 or greater, Results were plotted 
on Tost diagrams in. 20° classes since this range 
hest reflects the level of accuracy of both the 
measurement and the relationship of the Inng 
axis to the direction of ice movement, 


Stratigraphy of the Type Section 

A measured section was examinacd in a guily 
which heady in an earth dam immediately 
north of a fence line in the northern part of 
the type arca (Figs 1, 2). This gully presents 
the best continuous exposure of the Cape 
Jervis Beds, and reveals a sequence of older 
glacigenc sediments not present in the original 
measured section which lies several hundred 
meles to the south of our site (Ludbrook 
1967, 1980 p. 76). All other exposures in the 
outcrop were also examined, 

The Cape Jervis Beds have been divided 
into five unils on the basis of sedimentological 
and wenetic differences. We Interpret the 


66 N. F. ALLEY & R. P. BOURMAN 
LEGEND 
y Soil and vegetation SCALE 
> ie) 50 100 
S ~- Quaternary alluvium 
5-4 
S|lece|] Calcrete METRES 
oa 
“8-.-|| Pleistocene sands with marine fossils V2 SE 
aa HOT — 
-©_=| Glacio-marine fossiliferous Permian B 
| ry 
A —..y 65 
rgo,0") Flow till complex Trateenle 
r 60 
P= Sandstone beds ? a 
e 
a cay eal bss = 
8 ~~~] Glaciolacustrine beds with dropstones a 
3] f 50 w 
Ox A Isolated erratics wl 
2 = bas th 
et 2.8, Lag concentrate = 
-—— fr 40 
A « Lodgement till 
Ff 35 
& Shattered bedrock (gelifracts) z 
el z 
=f } Cambrian Kanmantoo t 30 
INN metasedimentary rocks ee sy 
Ss (SoS b25 2 
NM. UNIT SoS SS 20 
4 2 ans St bis W 
Unit | occurs south CO a wa Sees 
of measured o* 4 4 = Lag 
section \ a 
NG 
5 
: KG 
Be KS 


83-400 DEPARTMENT OF MINES AND ENERGY — SOUTH AUSTRALIA 


Note 


For location of measured section see FIG! 


Fig. 3. Stratigraphy of Cape Jervis Beds at measured section. Unit numbers equivalent to those in 


text. 


sequence to comprise, from the base up, fluvio- 
glacial and glaciolacustrine sediments, lodge- 
ment till, fluviolacustrine deposits, flow. till 
complex and a glaciomarine unit (Fig. 3). 

These sediments are in part plastered un- 
conformably against a steep, irregular bedrock 
slope, and elsewhere occupy north-trending 
bedrock depressions. These depressions are 
strongly influenced by the northerly strike and 
45°E dip of the bedrock of Cambrian Kan- 
mantoo Group metasediments. 


Unit I—Basal fluvioglacial and glacio- 
lacustrine sediments 

The base of the measured section comprises 
approximately 50 cm of angular rubble derived 
from fracturing of the underlying bedrock. 
Since excavation revealed that the rubble con- 
tinues under Unit 2, it is interpreted as frost 
shattered rock debris (gelifracts) which pro- 
bably formed in a_ periglacial environment 
prior to the advent of ice in the area. 


At some other places, Unit 2 directly over- 
lies non-shattered bedrock but attempts to 
locate buried striated bedrock surfaces by 
excavating till along the contact were unsuc- 
cessful. 

At a few other exposures south of the 
measured section, Unit 2 unconformably over- 
lics a few metres of cross-bedded medium 
sands at the base, overlain by level bedded, 
indurated medium to fine sandstone and finely 
bedded clays. The upper two intervals are 
characterised by striated dropstones and thin 
lenses (30-40 cm) of diamicton. Since there is 
a pronounced erosional surface at the contact 
with the overlying till unit, the basal beds were 
probably more extensive than their present 
meagre outcrops. 


The basal beds are interpreted as a pro- 
glacial facies of an advancing ice-mass. The 
cross-bedded sands are probably outwash, 
whereas the finely bedded clays were possibly 


CAPE JERVIS BEDS, SOUTH AUSTRALIA 67 


deposited in a small lake dammed between the 
ice and the steep bedrock slope. Dropstones 
are thought to have been derived from float- 
ing ice, and the diamicton from icebergs or 
the adjacent glacier. 

Unit 2—Lodgement till 

Directly overlying the rock rubble at the 
measured section are 3 m of compact, non- 
stratified diamicton containing pebbles and 
boulders of various shapes, sizes and litho- 
logies (Fig. 4a), The larger clasts are com- 
monly polished, faceted and striated. These 
are sect in a matrix of sandy silt. The diamicton 
is essentially unbedded but at a few sites 
incorporates small lenses of coarse sand, 
which may be remnants of deposits from sub- 
glacial melt-water channels or the remains of 
blocks of frozen pre-till sediments eroded by 
the glacier. Approximately 50 cm of pebble 
and cobble gravel forms an irregular layer 
on the upper surface of the diamicton (Fig. 
4b). 

The diamicton is interpreted as a lodgement 
or basal till (an interpretation further sup- 
ported by the till fabric analyses discussed 
below) and the thin uppermost gravel layer 
as a lag concentrate formed by erosion of the 
fines from the till during deglaciation. 

The till varies in thickness across the expo- 
sure, reaching a maximum of 5 m at one site. 
On the southern part of the exposure the till 
is plastered on a steep, channelled bedrock 
slope: it is evident that the sub-till topography 
is very irregular (up to 40 m of relief) and 
that the till locally occurs at a topographi- 
cally higher level than stratigraphically 
younger sediments. 

Till fabric analyses undertaken at the mea- 
sured section (site 1) and at a site on the 
southern part of the type exposure (site 2) 
reveal a strong NNW-SSE orientation (Fig. 
S). 

Generally, the till at our measured section 
contains a low frequency of pebbles (Fig. 4c). 
Pebble lithologies (Table 1) indicate that the 
larger clasts are dominantly metasandstones 
and were probably derived from local Kan- 
mantoo Group bedrock. Clasts of Encounter 
Bay type granites are present in low fre- 
quencies and form some of the larger erratics 
scattered across the exposure. The Encounter 
Bay Granites have a_ broad distribution 
between Port Elliot (Fleuricu Peninsula) and 
Cape Willoughby, Kangaroo Island (Daily 
et al. 1979) and were probably more widely 


exposed prior to the Late Palaeozoic glacia- 
tion. Together with the strong NNW-SSE 
orientation of the till fabrics, the presence of 
erratics of the Encounter Bay Granites, implies 
a local ice movement from the SSE. Pebbles 
of unknown provenance (not apparently 
derived from the bedrock on Fleurieu Penin- 
sula or adjacent islands) are represented by 


Fig. 4. Units at base of Cape Jervis Beds: 


a. Frost shattered bedrock (B) overlain by 
lodgement till (T). View looking southeast, 

b. Lodgement till (T) overlain by irregular 
layer of lag concentrate (LG) and fluviolacus- 
trine beds (1). Small spade 75 cm long; look- 
ing south. 

c. Close-up of lodgement till showing Jow fre- 
quency of pebbles and cobbles. View looking 
south. 


68 N. F. ALLEY & R. P. BOURMAN 


LODGEMENT TILL FABRICS 
SITE | 


FLOW TILL FABRICS 
UPPER FLOW TILL 


83-40) DEPARTMENT OF MINES AND ENERGY - SOUTH AUSTRALIA 


Fig. 5, Fabrics of lodgement till and flow tills. 


low percentages of granitoid types, porphyritic 
volcanics and pink, micaceous quartzite. The 
porphyritic volcanics were probably derived 
from the Mount Monster porphyry in the 
South East of the State. 


Unit 3—Fluviolacustrine beds 

Overlying the lodgement till are 15-20 m of 
interstratified sands, silts and clays contain- 
ing lenses of gravel and isolated pebbles and 


LOWER FLOW TILL N 


boulders (Figs 6a, b, c). Above these beds 
are 10-12 m of sediments that are largely 
obscured with debris and soil in the measured 
section. However, exposures in adjacent gullies 
indicate that the unit probably extends up to 
the base of the flow till complex (Unit 4). 


Although the beds are variable in lithology, 
cross-bedded sands dominate in the lower part 
of the unit and finely-bedded silt and clay 


CAPE JERVIS BEDS, SOUTH AUSTRALIA 69 


TasBce 1. Pebble lithologies of lodgement till (LT) 

and flow (FT) at Cape Jervis. Figures are per- (ay 

centages of total for each sample of 300; less than 
1 percent shown by x 


—— 


LT samples FT samples 
2 


Pebble lithologies 1 2 1 
Meta-sandstone 61 74 75 62 
Arkosic sandstone — 2 1 Xx 
Quartzite 

* Undifferentiated 20 5 4 14 
* Pink 3 2 2 4 
Siltstone 1 — x x 
Clay nodule X x — x 
Chert x x x x 
Schist x 2 1 1 
Phyllite — 4 2 2 
Gneiss — 1 x —_ 
Quartz 1 z 3 X 
Grinitoid types 7 7 8 13 
Volcanics — x x x 


beds are more common nearer the top. Pro- 
minent beds of calcareous sandstone contain- 
ing pebbles crop out at several intervals; these 
vary from coarse, well consolidated sandstone 
to fine friable sand. Rounded to angular intra- 
clasts of lacustrine clay occur throughout the 
unit and occasionally form thin lenses. 
Although such clasts are found in a variety of 
sedimentary environments, here they were 
probably eroded from frozen sediments and 
transported rapidly in a frozen condition to 
be re-incorporated into adjacent sediments, 

Near their base the beds dip eastwards at 
15—20° into the underlying bedrock palaeo- 
slope. Since there is no evidence of similar 
tilting of units above and below these beds, the 
tilting could be the result of sub-aqueous 
slumping, which may have occurred shortly 
after deposition. 

Polished, faceted and striated pebbles are 
common; stones in the finest beds that pene- 
trate to subjacent layers are regarded as drop- 
stones. Pebble lithologies are dominated by 
rocks of local origin although = granitoid 
erratics (including Encounter Bay type) are 
not infrequent. 

Because of the highly variable character of 
the beds and the presence of glacial erratics 
and dropstones, the unit is interpreted as of 
fluviolacustrine origin deposited in close proxi- 
mity to ice. The beds are characteristic of ice- 


Fig. 6, Fluviolascustrine beds: 
a, Interbedded sands, silts and clays showing 
eastward dip, 
b, Dropstone in bedded silts and clays. Hand 
lens approximately 135 mm long. 
c. Finely bedded clays interbedded with & ‘tee uit 
medium sands. as ‘ ( ba 


ita 
2 eee wy > 


70 N. F. ALLEY & R. P. BOURMAN 


contact environments where disintegrating 
ice produces temporary lakes and meltwater 
streams, the positions of which are constantly 
changing (Flint 1971 p. 184). Possibly the ice 
which deposited the lodgement till lay disinte- 
grating against the bedrock ‘slope thereby 
damming meltwaters and forming a kame 
terrace, 


ars Leo " 
wee. 78. 


Fig. 7. Flow till complex: 

a. Pebbly and bouldery lower flow till bed. 
Small lenses of bedded sands (S) near hammer 
handle. View looking east. 

b. Thin lens of flow till (FTL) interbedded 
with medium/coarse sands (S) overlying the 
upper flow till bed (FT). View looking east. 

c. Three flow till beds (FT) intercalated with 
pebbly medium/coarse sands (S). Lower flow 
ull beds pinch out to right. View looking south. 


Unit 4—Flow till complex 


Overlying Unit 3 are 8-10 m of sediment 
consisting of two very pebbly and crudely 
stratified diamictons intercalated with hard, 
superficially calcareous, bedded sandstones and 
other minor bedded silts and clays (Figs 7a, 
b). The sandstones are the Fontainbleau sand- 
stones of Harris (1971)°, although our work 
reveals that in places the calcareous cement is 
found only in the outer 50 cm of the exposed 
rock and is not present at depth. This sug- 
gests that the calcification may be the result 
of relatively recent pedogenic processes. 

The matrix of the diamicton beds is a silty 
sand; it is thus coarser than the matrix of the 
lodgement till. Gravel and sandy lenses occur 
in the diamictons, and the larger clasts range in 
shape from angular to rounded and are com- 
monly striated, polished and faceted. This unit 
is characterised by a concentration of very 
large boulders, one measuring 1.5 m diameter. 
This concentration of boulders is a most strik- 
ing feature of the type area and is readily 
traceable across the exposure (Fig. 2). Fabric 
analyses of the lower and upper diamicton 
beds reveals chaotic pebble alignments, when 
compared with the analyses of the basal tills. 
However, the lower diamicton displays a poor 
east-west alignment and the upper diamicton, 
a crude NW-SE alignment (Fig. 5). Pebble 
lithologies are dominantly local bedrock types 
but with a slightly higher frequency of grani- 
toid erratics than the lodgement till (Table 1). 
Harris (1971)" made a detailed lithological 
study of erratics from till at Cape Jervis, but 
from his descriptions of the stratigraphy of the 
site we conclude that the crratics were obtained 
from the flow till complex (see Fig. 9). The 
study, however, provides valuable information 
on the lithology of those distantly derived 
erratics found in the fiow tills. 

The sandstones interstratified with the dia- 
mictons contain numerous pebbles and thin 
lenses of fine gravel and grit. The sandstones 
are both flat- and cross-bedded, and the bed 
immediately below the lower diamicton is con- 
torted. The shape, surface features and litho- 
logics of the larger clasts within the beds are 
similar to those in the diamictons. 

Exposures in adjacent gullies reveal that 
similar suites of diamicton beds and associated 
sediments occur. In a gully immediately south 
of our measured section a diamicton bed form- 
ing part of such a suite was observed to pinch 
out in a coarse sand bed (Fig. 7c). Elsewhere, 


CAPE JERVIS BEDS, SOUTH AUSTRALIA 71 


fewer sandy diamictons were found, thus sug- 
gesting that the diamicton unit consists of a 
number of individual tabular lenses. 

The diamictons are interpreted as flow tills 
on the basis of the above evidence and the 
association with underlying ice-contact sedi- 
ments, These tills are believed to form in a 
variety of ways. Most commonly they develop 
by movement of supraglacial debris from the 
glacier onto adjacent proglacial sediments or 
by underwater flows in lacustrine or marine 
conditions (Dreimanis 1976), Flow tills are 
generally regarded as indicative of deglacial or 
ice-stagnation conditions. 

The characteristics of the flow tills described 
above are similar to those of Pleistocene age 
documented at a number of sites in the 
Northern Hemisphere. Where successive mud- 
flows have moved from glaciers out over pro- 
glacial scdiments, flow tills are often inter- 
bedded with stratified drift and resemble 
multiple lodgement tills produced by successive 
glacial advances (Boulton 1968, 1972). Since 
flow tills are derived from supraglacial debris, 
much of the fine clay and silt material is 
washed out so that the matrix is commonly 
sandier than that of lodgement tills (Dreimanis 
1976, Evenson ef al. 1977, Hicock ef al. 
1981). Pebble fabrics are found to be random, 
or unrelated to the direction of glacier move- 
ment (since pebble orientation is destroyed 
during flow from the glacier), or a preferred 
orientation may be developed in the direction 
of the mudflow (Marcussen 1975, Dreimanis 
1976; Evenson et al. 1977, Haldorsen & Shaw 
1982). Mud flows may also produce intra- and 
interformational fold structures and = shear 
planes (Dreimanis 1976, Evenson et al. 1977, 
Hicock et al. 1981). Some researchers also find 
that flow till, because it is derived from supra- 
glacial debris, contains more distal to interme- 
diate clast lithologies, whereas lodgement tills 
contain mastly local stones (Marcussen 1975, 
Hicock eft al. 1981). 

The fiow till complex in the Cape Jervis sec- 
tion is thought to have formed by successive 
supraglacial debris flows from ice stagnating in 
the present Backstairs Passage area. These 
flows spread out over an adjacent kame terrace 
thereby blocking streams to form shallow, 
ephemeral lakes. Fabrics suggest that the 
lower flow till may have formed by debris 
flows that moved eastward from the ice and 
the upper flow till by a northwestward move- 
ment of debris. 


Unit 5—Glaciomarine sediments 


Overlying the flow till unit are at Ieast 14 m 
of fine silts and clays which were measured 
up to the small dam at the top of the gully. 
Sporadic exposures above this level elsewhere 
indicate that the clays probably extend for 


Be LX a 
SOS 


oes 


till complex and ; 
marine silts and clays. Contact characterized by 


overlying massive glacia- 
low amplitude flame structures (FS). Drop- 
stone (DS) also shown. View looking south- 
east. 

b. Horizontally bedded clays of upper part of 
glaciomarine unit. . 

c. Plan view of bedding plane showing develop- 
ment of biscuit-shaped joint blocks characteristic 
of upper glaciomarine clays. 


72 N. F. ALLEY & R. P, BOURMAN 


another 30 m. The clay unit is capped by 
massive calcreted Point Ellen Formation of 
Ludbrook (1983). Ludbrook (1967) deter- 
mined a Lower Permian (Sakmarian) age 
for the clays and the underlying sediments 
from the presence of marine foraminifera in 
the clay unit. 


At the base of the unit the silty clays are 
massive and display conchoidal fracturing 
(Fig. 8a). This part of the unit also incor- 
porates thin beds of sand exhibiting slump 
structures, and thin lenses and blebs of sand 
and gravel composed of a variety of litho- 
logies, including some granitic types. A few of 
the pebbles in the gravel and isolated pebbles 
in the clay are polished, faceted and striated 
and are clearly of glacial origin. 


The silty clays are conformably overlain 
by 3-4 m of bedded clays containing pebbles, 
nodules of sand and numerous very thin lenses 
of sand or grit measuring at most 10 cm in 
length. Deformation of the clays under the 
pebbles and gravel lenses indicate that the 
coarser clasts in the lower parts of the clayey 
unit are of ice-rafted origin. 


The remaining part of the unit consists of 
horizontally bedded clays containing drop- 
stones and occasional thin lenses of grit 
(Figs 8b, c). Generally, the bedding becomes 
better developed higher in the sequence but 
there is no evidence that the beds are rhyth- 
mites. Numerous pebbles and boulders are 
strewn across the surface near the upper part 
of the beds and make the clays there appear 
till-like. The resemblance is only superficial, 
however, because the larger clasts are a lag 
derived from the clays by constant erosion of 
the slope. 

The sedimentary characteristics of this unit 
and the presence of arenaceous foraminifera 
suggests that deposition at first occurred in 
shallow marine conditions in close proximity 
to ice and meltwater streams. This was fol- 
lowed by deposition in progressively deeper 
water further away from the influence of the 
glacier and icebergs. Marine foraminifera 
recovered from the clays are indicative of low 
temperatures or low salinity (Ludbrook 1967) 
which would be consistent with expected in- 
fluxes of meltwater into a transgressing sea 
around a stagnating ice-mass. 


Relationship to the type section 


Although the stratigraphy we present above 
appears to differ significantly from that of 


Ludbrook (1967) and Harris (1971)5, it is 
only because the gully we examined contains 
a more extensive record of glacigene sediments. 
In the case of the type section, the base is 
located on portion of a bedrock palaeoslope 
that stands some 20 m above the base of the 
section we describe. We belicve that the bed- 
rock topography influenced the deposition and 
preservation of the lower beds, and that the 
base of the type section and the section 
described by Harris (1971)* corresponds with 
the flow till complex (Unit 4) of this paper 
(Fig. 9). 


NE 
This Paper 
f+ 300 mstres ——»} 


Point Ellen Formation| ~~ ~~ ~~ ~~ ~~ 


Unit 5 


SW 
Ludbrook (1967) 


Kunkar 


Units 5-6 


Units 1-4 


Frost shattered 
rock 


Fig. 9. Diagram showing probable stratigraphic 
relationships between units in this paper and 
those of Ludbrook (1967). 


Discussion and Conclusions 
Ice movements 


Evidence presented from areas adjacent to 
Cape Jervis indicates that movement of Late 
Palaeozoic ice was generally westward over 
Fleurieu Peninsula (Fig. 10). Till fabric data 
at Cape Jervis, however, show that local ice 
movement was NNW, and it is likely that this 
was related to the orientation of the bedrock 
controlled palaeoslope. Similar controls over 
local ice moyement have also been suggested 
at Hallett Cove to the north of the study area 
(Sprigg 1945; Milnes & Bourman 1972). Local 
deviations from the general direction of move- 
ment of large masses of ice are commonplace 
for the Pleistocene ice-sheets in the Northern 
Hemisphere. 

Striae on the western side of Backstairs 
Passage at Smith Bay, Kangaroo Island, are 
used to infer a northward direction of ice 
movement (Daily et al. 1979). The fabric of 


CAPE JERVIS BEDS, 


Mallell Gaye 


‘Direction of ine Movement 
from gtrige 


STR faoric 


Vieror Harbor 
GROMITE IS. 
WRIGHT 5 
ROSETTA, 4Eeth 


“west 15 


Encounter Boy 


Fig. 10, Map of ice movement across Fleurieu 
Peninsula as derived from striac, Source: Sprigg 
(1945), Milnes & Bourman (1972), Bourman 
& Milnes (1976) and Bourman ef al. (1976). 


the lodgement till overlying this striated bed- 
rock surface showed a north-south alignment 


Ww 
OF 


x fc x E 


tee hicknese unknown 
x 


x 
Fypataciar aebris 
=< 


Actve acial KE + peewee 
a 


~ : ae 

Sheot plone RS ww 

= — Ansal debris SI 
‘ in jamment RS 

— INRA 


% x. 


% , 
Konmantoc Group melasediments . 


© 


Supraglaciol debris 


a4 Fabtestiont gill) 


Fluvial onal 
focustrine 
sediments 


. 6 NN ON 
Kanmantoy Group metasediments \. 


* 


RA-40E NEFAATWEHT OY MINES amily EMRIOT —smuy + aus ALA 


SOUTH AUSTRALIA 13 
af pebbles which is consistent with such move- 
ment, 

We conclude from the above evidence that 
movement of ice across Fleurieu Peninsula was 
essentially westward but locally in the Cape 
Jervis area was guided by a north-south trend- 
ing depression. It is possible that Cape Jervis 
lay in close proximity to the confluence of ice 
flowing westward across Fleurieu Peninsula 
and ice moving northward along the eastern 
side of the modern Gulf St Vincent. 


Model of deposition 


The conclusions drawn concerning the 
genesis of the major facies comprising the 
Cape Jervis Beds are summarised in Figure 
11, This depositional model is characteristic 
of large ice-miasses stagnating in coastal low- 
lands (Alley & Chatwin 1979). During degla- 
ciation, retreat of the ice margins is accom- 
panied by general down-wasting of the ice 
surface. Uplands emerge through the ice first 
as it wastes down into the major valleys and 
coastal troughs where it may remain covered 
and protected by supraglacial debris for a con- 
siderable time. In these situations the stagnant 


Phivial and 
lecusirine 
sediments 


Flowriits 


| ai ¥ at é 
alles 
are , 
 Stognant ice N 
“ F uv x m& . 
s ; onmontod Group mete 
of ss 
a . - a % ‘ SA * 
A - Xs >< 
Ww E 
Ire hers, 
® = Bow Seo lyel 
‘d cS 
=“ 7 f= 


lee rotled debris 


GZ Ldagamany ts 
YOM x %. 
a, 


Slaghdnr ice Mey te ~ Ss 
rate fi — Can 
~ = eee “3 SNM SAXXALK 
ae ste Group ageOENS, 
= oS y N . 
Lee ee KK NSS SER SOAR 


Fig. 11. Model of deposition for Cape Jervis Beds at Cape Jervis. 


74 N, F, ALLEY & R. P. BOURMAN 


ice dams meltwaters and streams flowing from 
the uplands against the adjacent slopes. The 
ice marginal zones then become elongate 
basins in which complex suites of ice contact 
and proglacial sediments are deposited. 


Deposition in coastal areas is’ usually com- 
plicated by late deglacial transgréssion of the 
sea over and around the stagnating ice. For a 
short period there is an apparent rapid glacio- 
eustatic rise in sea level since this outstrips 
the rate of isostatic recovery in the lowland 
areas where rebound is slowed by the presence 
of thick masses of stagnating ice. With large 
volumes of meltwater and glacial debris avail- 
able, a considerable thickness of sediment may 
be deposited in relatively deep, brackish-water 
conditions. 


In summary, our model for deposition of 
the Cape Jervis Beds comprises the following: 


(1) Lodgement till was laid down over rem- 
nants of proglacial sediments or plastered 
on a partly frost-shattered, irregular west- 
facing slope by glacier ice moving 
roughly from south to north (Fig. 11a). 

(2) Deglaciation led to ice stagnation during 
which ice down-wasted into the coastal 
trough, exposing first the uplands and 
then the lodgement till on the palaeo- 
slope, A kame terrace formed between 
the ice and adjacent slope. The terrace 
initially consisted of a complex suite of 
interfingering alluvial and lacustrine sedi- 
ments (Fig. 11b). 
Further down-wasting of the ice led to the 
development of a thick cover of supra- 
glacial debris which, from time to tirhe, 
became unstable enough to flow from 
the ice out across the adjacent kame 
terrace, thus forming a series of flow till 
beds (Fig. lic). 


(4) Eustatic rise in sea level led to a rapid 
marine transgression that submerged the 
slope and probably part of the stagnant 
ice in the trough. Subsequent sedimen- 
tation occurred in brackish water turbid 
with fines brought to the sea by abundant 
meltwater, while Ienses of coarser clasts 
and isolated = pebbles/boulders were 
dropped from icebergs into the finer 
sediment below (Fig, 11d). 


Conclusions 


Our cxamination of the thickest and most 
continuous exposure of the glacigene Jervis 
Beds suggests that the sediments are related 
to only one glacial advance. All of the 
characteristics of the sediments can be 
accounted for in terms of deteriorating climatic 
conditions preceding glaciation, the passage of 
a wet-based (temperate) glacier over the site, 
and various proglacial environments associated 
with the stagnation and decay of an ice-mass. 
Although at least four till-like deposits occur 
in the type area, only one can be identified 
as a lodgement till. Hence, this site presents 
no evidence, at least locally, for multiple 
glaciation in the Late Palaeozoic. 

Finally, our studies reveal that the sedi- 
ments from which the Permian age of the 
glaciation is determined represent the final 
deglacial episode and are the youngest part 
of the Cape Jervis Beds. Whether the under- 
lying beds are earliest Permian in age or older 
however, could not be determined. 


Acknowledgments 
We are grateful to Drs W. V. Preiss, G. E. 
Williams, A. R, Milnes, N. H. Ludbrook and 
V. Gostin for their critical reviews of earlier 
drafts of the paper. Line diagrams were 
drafted by the Drafting Section, Department 
of Mines & Energy, South Australia. 


References 


ALLEY, N, F. & CHaAtwin, S. C. (1979) Late 
Pleistocene history and geomorphology, south- 
western Vancouver Island, British Columbia. 
Can, J. Earth Sci., 16, 1645-1657. 

BENSON, W, N. (1911) Note descriptive of a 
stereogram of the Mt Lofty Ranges, South Aus- 
tralia. Trans. R, Soc. S. Aust. 35, 108-111. 

BOULTON, G. S. (1968) Flow tills and related 
deposits on some Vestspitsbergen glaciers. J. 
Glaciol, 7, 391-412. 

(1972) Modern Arctic glaciers as deposi- 

tional models for former ice sheets. J. Geol. Soc. 

Lond. 128, 361-393, 


BOURMAN, R. P. & Miutnrs, A. R. (1976) 
Exhumed Roche Moutonnée. Aust. Geogr. 13, 
214-216. 

——, Maup, R. R. & MiLnes, A, R. (1976) Late 
Palaeozoic glacial features near Mt Compass, 
South Australia. Search 7, 488-490, 

CAMPANA, B., WiLtson, R. B., & WHITTLE, 
A, W. G. (1953) The geology of the Jervis 
and Yankalilla Military Sheets. Geol, Surv. S, 
Aust. Rept. Invest. No. 3. 

— & WILSON, R. B. (1955) Tillites and related 
Glacial Topography of South Australia. Eclog. 
Geol, Helv. 48, 1-30. 


CAPE JERVIS BEDS, SOUTH AUSTRALIA 1S 


Day, B. Minis, A. R, Twipare, C. R, & 
Bournr, J. A. (1979) Geology and Geomor- 
phology. Ja M. J. Tyler, C. R. Twidale & 
JK. Ling, (Eds) “Natural History of Kanguroo 
Island” (R. Soe. S. Aust.: Adelaide), pp. 1-38, 

Davip, T. W. BE. & Howcnin. W. (1897) Notes on 
the glacial features of the Inman Valley, Yan- 
kallla and Cape Jervis district. Trams, R. Soe. 8, 
Aust. 21, 61-67. 

Dremanis, A, (1976) ‘Tills: Their origin and 
properties. Ja R. F. Leggett (ed.) Glacial Tih 
Spec. Publ. No, 12, R. Soe. Canada, pp. 11-49. 

Drypen, L., (1931) Accuracy in the percentage 
representation of heavy mineral frequencizs. 
Proc. Nat, lead. Se. 17, 233-238. 

Por, R, FB. (1971) Glacial and Quaternary 
geoloyy. (Wiley & Sons: New York). 

Evinson, FE. B., Dremanis, A. & Newsome, J. W. 
(1977) Subaquatic flow tills: a new interpre- 
tation for the genesis of some luminated till 
deposits. Borcay 6, 115-133, 

GLACIAL RESEARCH COM™MTiTEE (1898) On the 
evidence of glacial action in the Port Victor 
and Taman Valley Districts. Aust. sles. Adi. 
Sef. 87, 114-127. 

Harporsen, S, & Suaw, J. (1982) The problem 
of recognizing mel-oul till, Borews 11. 261-277. 

Harris, S. A. (1969) The meaning of till fabrics. 
Mn J. G. Nelson & M. J. Chambers (eds.). 
Geomorphology. Selected Readings. (Metheun: 
Toronto). pp. 143-164. 

Hhicock, &, R. Dreimasis, A. & Broster, B. EB. 
(1981) Submarine flow lls at Victoria, British 
Columbia. Can, J. Barth Sci. 18, 71-80. 


Houmrs, C.D, (1941) Vill fabric, Ball geol. Sec. 
Aim, 52, 1299-[354. 

Hubert. J. BF. (1971) Analysis of heavy-mineral 
assemblages. Jn R, BE, Carver (ed.) Procedures 
in Sedimentary Petrology (Wiley-Interscience: 
New York), pp, 453-478, 

KRUMBEIN, W. C. & Prettizonn, PF. J, (1938) 
Manual of sedimentary petrography. (Appleton- 
Ceniury-Crofts: New York). 

Linpsay, J. F. (1970) Clast fabric of till and its 
development. J. Sedim. Petrol. 40, 629-640, 

LuUpnroox, N, H, (1967) Permian deposits of 
South Australia und their fauna. Trans. R. Soe. 
S. Aust. 91, 65-92, 

(1980) A guide to the geology and mineral 
resources Of South Australia. (Department of 
Mines & Energy: Adelaide). 

—— (1983) Molluscan faunas of the Early Pleis- 
tocene Point Ellen Formation and Burnham 
Limestone, South Atistralia, Frans. R. See. S. 
Ansi, 107, 37-49. 

Manican, C, T. (1925) The geology of Fleurieu 
Peninsula Part t. The coast from Sellicks Hill 
to Victor Hurbour. /hid. 49, 198-212. 

Marcussps, 1. B. (1975) Distinguishing between 
lodgement till and flow (ill in’ Weichselian 
deposits. Boreay 4, 113-123. 

Muanrs, A. R. & BourRMan. R. P. (1972) A 
Lite Palaeozoic glaciated granite surface ut 
Port Elliot, South Australia. Trans. R. Sac. S. 
lust. 96. 149-155. 

Serica, R, C, (1945) Some uspects of the geo- 
morphology of portion of the Mount Lofty 
Ranges. lhid. 69, 277-302. 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 108, PART 2 


ASPECTS OF GENETIC RELATIONSHIPS AND VARIATION IN PARROTS 
OF THE CRIMSON ROSELLA PLATYCERCUS ELEGANS COMPLEX 
(AVES: PSITTACIDAE) 


BY LEO JOSEPH & RORY HOPE 


Summary 


Enzyme electrophoresis was used to examine genetic variation within and between three of the four 
principal parrot taxa of the Crimson Rosella taxa of the Crimson Rosella Platycercus elegans 
complex of eastern Australia. Comparisons were made with the results of recent studies of other 
birds, mainly Northern Hemisphere members of the Order Passeriformes in which it has been 
shown that although populations of birds show levels of variation in isozymes comparable with 
those in other vertebrates, levels of isozymic differentiation and thus structural gene differentiation 
between bird taxa are unusually low. The data from rosellas, which belong to the Order 
Psittaciformes, although necessarily limited and therefore somewhat equivocal with respect to these 
generalizations, appear to be consistent with them, especially the former. No significant or 
consistent geographical patterns of gene frequency differences were detected among the rosellas 
studied. 


ASPECTS OF GENETIC RELATIONSHIPS AND VARIATION IN PARROTS 
OF THE CRIMSON ROSELLA PLATYCERCUS ELEGANS COMPLEX 
(AVES: PSITTACIDAE) 


by Leo Joseru & Roky Hope* 


Summary 


Joycen, f. & Hore. R, (1984) Aspects of genetic relationships and yaralion i) the parhots 
of the Crimson Rosella Platyeercus elegans Complex (Aves: Psittacidae) Trany. RE See. 
4. Aust, VOR(2), 77-84 12 June, 1984. 


Hingvine cluctruphoresly was used lo examine genetic varialion within and between three 
af the four prineipal parret taya of the Crimson Rosella Plaiycereus elegais complex of 
eastern Australian. Comparisons were made with (he resul(s of recent studics of other birds, 
mainly Northern Hemiscphere members of the Order Passeriformes in which it has been shown 
(aut although papilations of birds show levels of variation in isozymes comparable with those 
in Other vertebrates, levels of isozymie differentiation and thus structural gene differentiation 
between bird taxa are unusually low, The data from rosellas, which belong to the Order 
Pyiltvertormes, although necessarily limited and therefore somewhat equivocal with respect 
to these eeneralizations, appear to be consistent with them, especialy the farmer. No signilieant 
or consistent geographical patterns of gene frequeney diflecences were detected among the 


rosellas studied. 


Key Worns: Birds, electrophoresis, poptilation genetics, Plumweerewy clear. 


Introduction 


Four principal parrot taxa comprise the 
Crimson Rosella Plaryeercus elegans complex 
jv eastern Australia (see Pig. la and Forshaw 
1981, Tor details of habitat and distribution), 
One, the Crimson Rosella P. elegans, ranges 
south from the Atherton Tablelind, Queens- 
lan, along Australia’s eastern coast to south- 
eastern South Australia with an isolated popu- 
lation on Kangaroo Island. Adults are pre- 
dominantly ¢rivson, immatures green, A 
second, the Yellow Rosella P, flaveelus, is con- 
fined ta the Murray-Darling river system, In 
veneral, the crimson in the plumage of P. 
elesany 18 replaced by yellow in this form, 
In the Mt Lufly Ranges and southern Plin- 
ders Ranges there occurs a third form, a series 
of populations all of which are variable in 
eolour but intermediate helween P. elegans 
and #. flaveolus, These are collectively termed 
here the Adelaide Rosella P. adelaidae. There 
is clipal variation in plumage in FP. adelaidac, 
particularly on the ventral surface (Torshuw 
J981). In the southern Mi Lofly Ranges adults 
are, wenerally, cieh scarlet. Northwards, they 
hecome progressively lighter and more orange. 
Populations in the Plinders Ranges wre pre- 
dominantly yellow yertrally but usually have a 
strang wash of orange, The fourth member of 
the complex, the Green Rosella P. cale- 


* Department. ol Genetics, Universily of Adetaicte, 
GV... Box $98, Adelunic, S. Aust. S001, 


donicuy of Tasmania and the larger Bass 
Strail islands, was not included in this study, 

There has been considerable debate over the 
relitionships and taxonomic stalas of these 
parrots. Condon (1954) preaposed that. P. 
adelaidae evolved = through hybridization 
between P. elecuny and PL flaveolus, This is 
suggested by the intermediate colouration of 
Po adeluidae (see alsa Martindale 1974", 
Forshaw 1981). Also, Ihe southenorth cline int 
colouration of FP, adelaidae 1% most simply 
explained perhaps a4 having arisen through 
introgression of P. flyveo/ny genes lrom the 
north and FP, elegans genes fram the south. 
Ifoso, this would afford turther support for 
some previous hybridization, Cain (1955) and 
Keast (1961), on the other hand, suggested 
that the members of the complex evulved in 
Miu in response to climatic changes, 

Two generalizations have emerged from 
recent electrophoretic studies of genic variation 
in birds. Firstly, populations of birds do not 
(differ significantly from those of other verte- 
brates in levels of Within-population varialian. 
Sceondly, there seems to be considerably Jess 
genic differentiation between bird taxa thah 


' Martindale, J. (1974). Some ceegeretic relu- 
fionships and their origins in platycereine species 
complex (Order Psittaciformes, Cliss Aves) of 
south-eastern Australia, Honours Thesis, Depart- 
ment of Genetics and Homan Wariation, das 
Trobe University, Victoria. Unpubl 


78 L. JOSEPH & R. HOPE 


between taxa of other vertebrate classes at 
equivalent levels of the taxonomic hierarchy 
(see, for example, Avise & Aquadro 1982 and 
references therein and an alternative view 
offered by Sibley & Ahlquist 1982). 

These generalizations have grown largely 
out of studies of passerine birds. Concerning 
Australian species, data are at present available 
only for the Grey-crowned Babbler Pomatos- 
tomus temporalis, a communally breeding 
passerine, and from some species of the pas- 
serine family Hirundinidae (Johnson & Brown, 
1980; Manwell & Baker, 1975). Thus, the 
present study of non-passerine birds aimed (1) 
to measure levels of electrophoretically detec- 
table genic variation between and within some 
members of the Crimson Rosella complex, 
and (2) to compare them with such levels 
measured in other birds and vertebrates in 
general. 


P flaveolus 
P adeladae 


Materials and Methods 


Collecting Procedures: Specimens of rosellas 
were collected under a permit from the South 
Australian National Parks and Wildlife Service. 
Within an hour of death, samples of liver, 
heart muscle and breast muscle were extracted 
for electrophoresis and transported in dry ice 
to the laboratory where they were stored at 
—20° C. Specimens were sexed by dissection 
and aged as adult or immature from plumage 
(Lendon 1973). Stomach contents have been 
preserved and the birds have been prepared 
as voucher study skins and lodged in the 
South Australian Museum, Adelaide. The col- 
lecting localities are shown in Fig. 1b. 

Adelaide Rosella P. adelaidae: 59 specimens 
were collected to sample P. adelaidae as evenly 
as feasible throughout its range. They were 
collected in six geographical sectors: S, 
extreme southern Mt Lofty Ranges = 


Fig. 1. (a) Approximate distribution of members of the P. elegans complex in eastern Australia. (b) 
Localities and sample sizes collected for this study; letters are the abbreviated names of each 


sample. 


GENFIICS OF THE CRIMSON ROSELLA COMPLEX 74 


Fleuricu Peninsula (10 specimens); RB, 
southern central Mt Lofty Ranges between 
Aslibourne and Meadows (9); Ma, lower 
Marne River and environs (9); M, northern 
central Mt Lofty Ranges between Kapunda 
and Saddleworth (10); C, northern Mt Lofty 
Ranges between Auburn and Clare (11); and 
F, southern Flinders Ranges between James- 
lawn and the Dutchman’s Stern (10), No 
morc than three specimens were taken from 
one locality within these sectors with the 
exceptions of 15 km NB of Saddleworth and 
the Dutehman’s Stern. Sample F was taken 
aver a larger urea than the other samples 
hecalise of the Jogistic difficulties involved in 
sampling the Flinders Ranges birds and 
hecause it was intended to treat these birds 
as a distinet sub-population of P. adelajdae in 
subsequent dala analyses, 

Crimson Rosella PL elegans: six specimens 
were taken from four localities at the western 
end of Kangaroo Island, In addition, the livers 
of four fromm the South-East of S.A, and of six 
from the Atherton ‘Tableland, Qld were 
examined, Youcher specimens of the latter 
iwo samples are held in the Australian 
National Wildlife Collection, Canberra. 

Yellow Rosella P. flaveolus: five specimens 
fram near the Murray River between Barmera 
and Waikerie were examined. 

Other species: organ extracts were obtained 
from the parrot genera Barnardius and 
Psephotuy, both of which are closely related 
to FPlatyeercus (Cain 1955) and of a more 
distantly related bitd, & pigeon, Details of 
these ure as follows: the exolic Spotted Turtle- 
Dove Streptepelia chinensis (one iadividual, 
liver only. collected in grounds of Liniversity 


of Adelaide), Mallee Ringneck Parrot Barnar- 
dius barnardi (two, lier and heart miusele, 
near Swan Reach), Mulga Parrot Psephorus 
varius (one, liver and heart musele, near Swan 
Reach), Red-rumped Parrat Psepharus haenta- 
tonabiy (one, liver only, near Mannum).. The 
availability of these samples tnade possible 
genetic comparisons between the members of 
the P. elevans complex and other species. 

Electrophoresis: Electrophoresis was carried 
out on cellulose acetate gels (Meera Khan 
1971) following proeedures of Baverslock cf 
al, (1977), When electrophoresis revealed iwo 
forms of an enzyme (manifest as distinct sets 
of bands), the presumptive locus encoding 
the most anodal form was designated 1 and 
the other 2 e.g. Jedhi-l, /dh-2 for isocitrate 
dehydrogenase, The presumptive allele encod- 
ing the most anodally migrating produet of a 
locus was designated @, the second most anodal 
hand so on. In one case, Peed. fb indicates an 
allelic form of the enzyme intermediate in 
mobility between Pydh and Ped 

Table | shows the enzyines assayed and 
the organ and electrophoresis buffer used for 
each enzyme, In all, nine enzymes were elec- 
trophoresed and choice of these was not 
entirely random, being determined partly by 
which enzymes were under investigation in the 
laboralory for other projects. Aconitase was 
examined to test a hypothesis of sex-linkage 
in birds (see Baverstock er al. 1982). 

The symbols used to represent ihe locus or 
loci corresponding with cach enzyme are: 
6-phosphogluconate dehydrogenase, Ped) 
purine nucleoside phosphorylase, NP: adeny- 
late Kinase, ak—-7, Ak-2) phosphoglucomutase, 
Pam: isocitrate dehydrogenase, fdh-]. Bdh-2; 


TABLE 1. Enzvntes assayed and buffers and organs bsed for clectrapharesiy. 


Ensyme 


é-Phosphogliconate dehydrogenase 
Purine nucleoside phosphorylase 
Pyruvate kinase 

Adenylate kinase 
Phosphoglucomutase 

Isovitrate dehydrogenase 

Agonitaxe 

Glucose phosphate isoinerase 
Glilamate oxaloacttate Lransaminase 


Buffer! Orgare 


B -+- NADP L 
A L 
C. L 
8 L 
C L 
i 
L 
L 


__ 
B+ NADP 
B 


B 
B L 


jai nea te heh eae Lala cen acne EE 
Buffer code: A—OAN M cilrate-phasphate, pH 6.4; R—0.02 M phosphate, pH 70. C—(.5 
M trisemuleate, pil 7.8; D—0.12 M tris-glycine, pH 95 


"Organ code, b—liver; H heart. 


} NADP indicules that 600 xl of NADP (10 mg/oal) were added te the buffer placed in (he 
cathudal compartment and in which the gel was souked_ 


80 L. JOSEPH & R. HOPE 


aconitase, Acon—1, Acon—2; glucose phosphate 
isomerase, Gpi, glutamate oxaloacetate trans- 
aminase, Got and pyruvate kinase, Pk. 

Analysis of data: The proportion of poly- 
morphic loci, P, was calculated directly using 
two definitions of polymorphism: one where 
the most common allele had a frequency of 
less than 0.99 and the other where this fre- 
quency was less than 0.95. 

The average heterozygosity per locus, H 
(see Nei 1978), was calculated as: 

y= 2h 
r 
where r is the number of loci scored, and h 
is the expected heterozygosity at each locus, 
calculated as 1 — x, where x; is the frequency 
of the ith allele at each locus. 

Standard errors of H estimates have been 
discussed by Nei (1978) and Nei & Roychoud- 
hury (1974), who concluded that it is more 
important when estimating H to screen many 
loci in few individuals rather than the con- 
verse, Gene frequencies were compared with 


t-tests and Fisher’s exact method. So that 
comparisons could be made between P. elegans 
or P. flaveolus on one hand and P. adelaidae 
on the other, the P. adelaidae data for each 
locus were pooled, their homogeneity first being 
assessed by the method of Hancock (1975). 
Heterogeneity was detected only for the data 
from pyruvate kinase, which were accordingly 
not used in such data analyses. 


Results 


Electrophoretic Typings and Isozymic Varia- 
tion within P. adelaidae 

Tables 2 and 3 present the results of elec- 
trophoresis and Table 4 presents measures of 
genic variation within the rosellas and in ver- 
tebrates in general. The standard errors of the 
rosella H estimates are relatively large, e.g. 
approximately 0.05 for the H estimate of 
0.104 in P. adelaidae (see Nei 1978). The 
estimates are, therefore, only coarse approxi- 
mations. The data for Pgd, Np, and Idh-2, 
the most variable loci, showed P. adelaidae to 


TABLE 2, Numbers of individuals of indicated genotypes in population samples of the P. 
elegans complex. In samples of P. elegans itself, KI = Kangaroo Island, SE = South-East of 
South Australia, and A = Atherton Tableland, Qld. For brevity, genotypes are represented 
thus: at an indicated locus e.g. Pgd, a/a designates Pgda/Pgd8. Wholly invariant loci are 


omitted. Heart samples for typing of Acon—2 


and Idh-2 not available for samples SE and A. 


P. elegans 
SE 


Genotype KI A P. adelaide P. flaveolus 
Ped a/a 2 
a/b 1 10 2 
b/b 4 2 5 39 2 
b’/c 1 
b/c ] 8 1 
c/¢e 1 
Np a/a 5 1 - 29 2 
a/b 3 23 2 
b/b 5 1 
a/c 2 
Ak-1 a/b 1 * 
b/b 6 4 6 58 5 
Ak-2 a/b 1 1 
b/b 5 3 6 59 5 
Pgm a/b 1 
b/b 6 4 6 58 5 
Tdh-1 a/a 1 
a/b 2 
h/b 6 4 0 59 5 
Idh—2 b/b 5 
a/c 2 
a/b 1 1 
a/a 6 48 4 
Acon-—I a/a 1 
b/b 5 57 5 


GENETICS OF THE CRIMSON ROSELLA COMPLEX 81 


Tare 3, Numbers of individuals of indicated genotypes in population samples of P. adelaidae 
designated 8, R, Ma, M, C and F (see text), Genotypic symbolism as for Table 2, Whally 
invariant loci are omitted, 


¢ Sample 
Genotype 5 R Ma Cc F 
Ped a/h 3 4 2 i 
hifh 8 5 6 5 6 9 
b/c 1 
hfe 2 3 | 2 
cfc 1 
Np asa 6 6 3 6 1 & 
afb 2 3 5 3 8 2 
hfh 2 | ] 
are 1 I 
Ak-I a/b | 
h/h 10 g 9 9 im 10 
Prem a/b ! 
bib 10 9 9 ine) 11 9 
Tuh-2 e/a 2 
hfe I 
c/e 7 7 6 10 8 10 


Tabin +. Values of P, the praportion af polymorphic loci, and H, the average heterozygasity 
per locus, ebtained in the P. elegans complex. Walues far Wertehrata and Aves are fram 
Neva (1978). 


P H 
Loci Freq. of most common allele 
Taxon scored <0.99 <0.95 
P. adelaidae 10 0.3 03 0.104 
P elegans 
Kangaroo Ishind 11 0.18 O18 O07 
South-East 8 0,375 0.375 ().137 
Atherton Tableland 8 Q.125 0.125 0.05 
P, flaveolus il 0.27 0.27 0.102 
Vertebrata a 173 + O19 0.0494 
+0.0365 
Aves — O150£0,111 0.0473 
+ 0.0360 
be in Hardy-Weinberg equilibrium (Ped, P. elegans samples only in gene frequencies at 


xz = 0.74, 0.7>P>0.5; Np, Xj — 0.025, 0.9> 
P>0,8; Idh—-2, x3 — 2.16, 0.2>P>0.1). 

Two detected alleles were unique to P, 
elegans and six ta P. adelaidae (Table 2). 
Nohe occurred at frequencies significantly 
higher than zero except Pgin® in the com- 
parison of the Flinders Ranges P. adelaidae 
sample with all other samples of P. adelaidae 
pooled together, 

Comparisons of Gene Frequencies 

Even allowing for the small sizes of the 
samples, significant differences in gene fre- 
quency were few, especially between P. elegans 
and P, flaveolus. Atherton Tableland P. elegans 
despite its geographical isolation, differed 
significantly from P, flaveolus and the other 


the /dh—/ locus, South-East P. elegans showed 
no significant differences from P. flaveolus. 
One significant difference between South-East 
and Kangaroo Island P. elegans was detected 
at Np. The six samples from Kangaroo Island 
all lacked the Np® and, as well, the Pgd* genes 
found in P. flaveolus and South-East P. elegans 
and also the Jdh-2> gene found in P. flaveolus 
and P. adelaidae. Only for Np” in the Kan- 
garoo Island P. elegans-P. flayealus compari- 
son was such a difference significant (P = 
0.043). 

Ped® occurred at a significantly higher fre- 
quency in Kangaroo Island P. elegans than in 
P. adelaidae samples, 8S, Ma and F and all P. 
adelaidae samples pooled, and was greater 
also in P. flaveolus and sample M than sample 


82 L, JOSEPH & R. HOPE 


S, Np®* occurred at a significantly greater fre- 
quency in Kangaroo Island P. elegans than in 
P. adelaidae samples Ma and C and all P. ade- 
laidae. 

Within P. adelaidae, significant differences in 
gene frequency were few and showed no con- 
sistent geographical pattern. The Flinders 
Ranges sample of P. adelaidae differed signifi- 
cantly from other P. adelaidae samples pooled 
only for Pgm (0.05>P>0.02). 

Enzyme Expression 

The enzyme products of Jdh-1 and Acon-I 
were strongest in liver extracts, while those of 
Idh-2 and Acon-2 were strongest in heart 
extracts. Avise et al. (1980) noted the same 
difference for Idh loci in North American 
thrushes and their allies. Wholly or largely 
invariant enzymes were both glucose- and non- 
glucose metabolizing. 


Discussion 


Although breeding studies in rosellas have 
not been conducted to test the mode of inheri- 
tance of the proteins examined in this study, 
Mendelian inheritance has been assumed 
because: 

(i) the proteins examined display Mendelian 
inheritance wherever studied in other organisms 
(see, for examples, Harris & Hopkinson 1976); 
and 

(ii) population data satisfy Hardy-Weinberg 
expectations such as to render unlikely any 
other mode of inheritance approximating 
Mendelian expectations. 

Thus, the isozymic data obtained in this 
study may be used to discuss genetic relation- 
ships and variation, 

The sample sizes used in this study, par- 
ticularly those of P. elegans and P. flaveolus 
were rather small. Baverstock ef al. (1977) 
have shown nonetheless that such samples can 
be adequate to indicate relationships in elec- 
trophoretic studies, although they did not 
negate the desirability of having larger 
samples, especially for studies of within-popu- 
lation variation. 

The genic variation measured in the rosellas, 
genically limited though it is, is consistent with 
evidence that levels of genic variation in birds 
are much the same as those in other verte- 
brates. Similarly, the paucity of genic differen- 
tiation between the rosellas may support the 
concept that in birds morphologically quite 
different taxa exhibit relatively less structural 
gene differentiation within an Order than do 


other animal groups. Consistent with this were 
the observations of alleles shared by the parrot 
genera Barnardius, Psephotus and Platycercus. 
Nevertheless one should note that although P. 
elegans and P. flaveolus appear quite different, 
there could be a simple genetic basis to their 
plumage differences. A full discussion of the 
concept of weak structural gene differentiation 
in birds is not intended here; the reader is 
referred to the reviews and alternative opinions 
presented by Sibley & Ahlquist (1982) and 
Avise & Aquadro (1982). 

If sampling in P. adelaidae has been com- 
prehensive, the evolutionary and genetic signi- 
ficance, if any, of ‘unique’ alleles in P. ade- 
laidae shall remain obscure until more exten- 
sive samples of P. elegans and P. flaveolus are 
collected. Bearing in mind the possibility that 
P, adelaidae evolved by hybridization, we 
would note that the phenomenon of ‘unique’ 
alleles in hybrid populations has been observed 
previously and discussed by Golding and 
Strobeck (1983), 

Some of the differences in gene frequency 
between Kangaroo Island P. elegans and other 
populations may be due to stochastic effects, 
This population has been isolated for some 
10000 years (Lampert 1979). 

Four presumptive pyruvate kinase hetero- 
zygotes (double-banded) were detected in 
liver extracts of P. adelaidae. Heart extracts of 
three of these individuals were found to be 
single-banded. Possibly, pyruvate kinase is 
modified in the liver, producing a secondary 
band on gels, thereby creating a false impres- 
sion of heterozygosity in homozygotes. Alter- 
natively, a gene for pyruvate kinase may be 
‘switched-off in the hearts of rosellas hetero- 
zygous for it. Extracts from other organs and 
analysis of data from parents and their off- 
spring would resolve this anomaly. Almost 
certainly, this difference explains the statistical 
heterogeneity in the P. adelaidae data for 
pyruvate kinase. 

Relationships in the P. elegans species-group 

As noted above, Cain (1955) and Keast 
(1961) suggested that the members of the P. 
elegans complex evolved in situ in response to 
climatic changes. Alternatively, Ford (1977) 
proposed that P. flaveolus originated in the 
Mt Lofty Ranges while isolated from P. 
elegans by arid country in the region of the 
Coorong during an arid part of the Pleistocene. 
According to this interpretation, P. flaveolus 
moved northwards into the Flinders Ranges 


GENETICS OF THE CRIMSON ROSELLA COMPLEX a 


and castwacds to the Murray-Darling system 
when this barrier broke down, while 2. elegans 
moved westwards and freely hybridized with 
P. flaveoluy in the Mt Lolty Ranges, producing 
PF adeldidae, Although Ford (1977) was able 
to Suggest dates for these propased events 
from geological data, one of his postulates, 
namely that of Plinders Ranges PL adeluidae 
being isolated by seventy kilometres From 
other populations of P, adelaidae to the south, 
appeurs lo be exaggerated. Any ‘isolation’ of 
this population can be by no mere than thirty 
to forty kilometres, Moreover, it is unlikely 
to be permanent owing to the vagility of 
rosellas combined with the existence of habitat 
corridors, and has undoubtedly been promoted 
by, iP dt is not entirely consequent upon, agri- 
culttiral clearing of vegetation in the northern 
Mi Lofty Ranges within the last 100) years. 
Thus, given the likelihond of eene flow 
through all populations of Po adelutdae. the 
mechanism of the maintenance of  elinal 
plumage variadon m P, addeldidae remains 
problematic. Neither Hardy-Weinberg 
equilibria nor regression and eorrelation 
analyses suggested that strong selection dif- 
ferentials act ow electrophoretic chatacters in 
P. adelaldae, The existenee of some measure 
of sclection is suggested by the clinal varia- 
lion in plumage itself and hy the Pem and 
Alli-2 gene frequency differences between 
the Flinders Range and other P, adelaidae 
samples. 


Concerning the relitionship between P. 
flavealus and PL adelaidae, Short (1969) 
argied that it should be determined whether 
the two overlap with only some hybridization 
or wiih extensive hybridization: the former 
situation would indicate effective reproduc- 
tive isolation. the latter conspecificity. Unfor- 
tunately, the observations and claims of 
Lendun (1973) and Condon ($969, 1975) 
concerning overlap between the two lack 
adequate supporting evidence. Present-day 
hybridization between then of, more specifi- 


cally. introgression of P, flaveoluy genes inte 
P. adelaidae 1 suggested hy the fdlh data. 
fdh-2> was not deteetcd in FP. elevans hot it 
was in P, flaveolus and several samples of 1, 
edelaidae, eluding that taken from the Mame 
River region where FP. adelaidae and P. 
Haveoluy are perhaps closest (unpubl, data), 
More fyaterial would be useful in evaluating 
this und any olher interpretations, 

Overall, the genic data Indicate a close rela- 
tionship between the three rosellas studied but 
ul present they are insufficient to determine 
with certaincy whether the birds constitute a 
single biological species, ‘There 1s scope for 
much further work on the group, The 
mechanism und mainienance of the clinal 
variation in, and the siznificance of ‘unique’ 
cones in FP, adelaldae could be clarified with 
more extensive field and Jaboratory analyses. 
Also, the relationship of 2. ealendonicus to 
the other members of the complex warrants 
invesligalion ax does a much closer examina- 
fion of The genetic relationships between all 
the members, A karyotypic analysis may 
prove useful, specially to test further the 
hypothesix of a hybrid origin of P, adelaldae. 


Acknowledgments 


We thank Prof, J. H, Bennett and the 
referees of drafts for their encouragement and 
constructive criticisms. Specimens collected for 
this study were taken under the conditions of 
a permit from the South Australian National 
Parks aol Wildlife Service. For his generous 
and enthusiastic belp in collecting we should 
particularly like to thank Dr J. D. Dwyer. C 
Baxter, R, Beckwith, J, Cox, R. Drummond, 
R. Ellis. T. Fraser, 7, Aney-Williants, D, 
Jones, L, Parker and L. Pedler provided in- 
Valuable help with collecting. M. Adams, P. 
Baverstock, (, Chesson. T, Hancock, C. R. 
Leach, D. Potirell and Ry Schodde all helped 
in Various technival aspects of laboratory work 
and data analysis; ta them we arte also grate- 
ful. We Jastly thank Miss G, Psaltis for typing 
and Ruth Altmann ter preparing Vig, 1- 


References 


Avist. J.C) & AOVanno, CoB. (1982) A com- 
parative summary of genetic distances in the 
Vertetirites. Eval. Bial WS, 131-185. 


——, Parron, $0, & Agusona, C.F. (1980) 
Evolutionary eeuctics of birds. [. Relationships 
aimone North Ameri¢an (heushes and allies. 
The Anak WI, (45-147, 


favrestoce. POOR, Waris, ©, 2, 
S. Ro (19771 Bleetrepharetic 


S. & Core, 
COMPHTSANns 


hetween allopatric populations Of five Austratian 
pseudomvine rodents (Muridae), Aas! J. frat, 
Sel, 30, 471-485, 

——, Anpams, M., PorkincHorne, R, W, & 
Ghiber, M, (1982) A sex-linked enzyme In 
hirds-Z-chromosome conservation bur hu dosage 
conipensation, Nature 296, 763-760. 

Gain, AL J, (1955) A revision of brie fomlossius 
huematntus and the Australian platycercive 
parrots. Shir 97, 442-479 


84 L. JOSEPH & R. HOPE 


Conpon, H. T. (1954) Remarks on the evolution 
of Australian birds. §. Aust. Orn. 21, 17-27. 
(1969) “A Handlist of the Birds of South 
Australia.” (South Australian Ornithological 

Association: Adelaide. ) 

(1975) “Checklist of the Birds of Australia.” 
Part I. Non-passerines. (Royal Australasian 
Ornithologists Union: Melbourne.) 

Forp, J. (1977) Area of origin of the Yellow 
Rosella. Emu 77, 229-230. 

ForsHaw, J. M. (1981) “Australian Parrots.” 2nd 
edition. (Landsdowne: Melbourne.) 

GOLDING, G. B. & Stropeck, C. (1983) Increased 
number of alleles found in hybrid populations 
due to intragenic recombination. Evolution 37, 
17-29. 

Hancock, T. W. (1975) Remark on Alogrithm 
434 (G2). Exact probabilities for R xX C 
contingency tables. Comm. Assoc. Computing 
Machinery 18, 117-119. 

Harris, H., & Hopkinson, D. A. (1976) “Hand- 
book of Enzyme Electrophoresis in Human 
Gentics.” (North Holland Publishing Company: 
Oxford. ) 

Jounson, M. S. & Brown, J. L. (1980) Genetic 
variation among trait groups and apparent 
absence of close inbreeding in Grey-crowned 
Babblers. Behav. Ecol. Sociobiol. 7, 93-98. 

Keast, A. J. (1961) Bird speciation on the Aus- 
tralian continent. Bull. Mus. Comp. Zool., 
Harvard 123, 307-495. 


LampPerRT, R. J. (1979) Aborigines. In M. J. 
Tyler, C. R. Twidale & J. K. Ling (eds.): “The 
Natural History of Kangaroo Island.” (Royal 
Society of South Australia: Adelaide. ) 

Lenpon, A. H. (1973) “Australian Parrots in 
Field and Aviary.” (Angus and Robertson: 
Sydney.) 

MANWELL, C. & Baker, C. M. A. (1975) Mole- 
cular genetics of avian proteins. XIII. Protein 
polymorphism in three species of Australian 
passerines. Aust. J. Biol. Sci. 28, 545-557. 

MEERA KHAN, P. (1971) Enzyme electrophoresis 
on cellulose acetate gel-zymogram patterns in 
man-mouse and man-Chinese hamster somatic 
cell hybrids. Arch, Biochem. Biophys. 145, 470- 
483. 

NEI, M. (1978) Estimates of average heterozygo- 
sity and genetic distance from a small number 
of individuals. Genetics 89, 583-590. 

, & Roycnoupuury, A. K. (1974) Sampling 
variances of heterozygosity and genetic dis- 
tance. Genetics 76, 379-390. 

Nevo, E. (1978) Genetic variation in natural 
populations: patterns and theory. Theoret. Pop. 
Biol. 13, 121-177. 

SHorT, L. (1969) Taxonomic aspects of avian 
hybridization. The Auk 86, 84-105. 

Siptey, C. G. & Antauist, J. E. (1982) The 
relationship of the Yellow-breasted Chat 
(Icteria virens) and the alleged slowdown in the 
rate of macromolecular evolution in birds. 
Postilla 187, 1-18. 


COASTAL LANDSLUMPING IN PLEISTOCENE SEDIMENTS AT 
SELLICKS BEACH, SOUTH AUSTRALIA 


BYR. I. MAY & R. P. BOURMAN 


Summary 


A pronounced bench occurs at approximately 25 m asl in a short section of the 50 m high coastal 
cliffs eroded in Quaternary sediments at Sellicks Beach, South Australia. Below the bench the 
sediments are disturbed and the stratigraphy is confused. As a result of studies of the regional 
stratigraphic succession we have identified the presence of a large rotational landslump as well as 
evidence of earlier phases of slump activity. Here we describe the Late Cainozoic stratigraphic 
record near Sellicks Trig., the morphology of the bench and the character of the underlying 
sediments, the probable age of the slumping responsible for the bench and factors involved in its 
formation, and implications for planning, development and management of this coastal region. 


COASTAL LANDSLUMPING IN PLEISTOCENE SEDIMENTS 
AT SELLICKS BEACH, SOUTH AUSTRALIA 


by R, I, May & R. P. BoURMAN* 


Sunumary 


May, Rv 1 & Hourman, Ro T (198d) Coastal landslimping in Pleistocene sediments at 
Sellicks Beech, South Australia. Trams. A. Seo, S. tust. 10802), 85-94, 12 bane, 1984, 


A pronounced bench aceues at approximately 25 m asl tn a short section of the SO my 
high constal clits eroded in Quaternary sedimenis at Sellicks Beach, South Austriis. Below 
ihe bench the sediments are disturbed und the stratigraphy is confused. As a result of studies 
of the regional steatigraphic succession we have identilied the presence of o large rotational 
lundslump as well as evidence of earlier phases of slimp activity. Here we describe the Late 
Camozoie stratigraphic record near Sellicks Trig. the morphology of the bench and the 
character of (he undechiow sediments, the probuble age of the slumping: responsible for the 
bench und faetors involved in its formation, and implications for planning, development and 


Management of this coustal region. 


Kiy Worps: Pleistocene stratigraphy, slumping. Sellicks Beach, munagement, 


Introduction 

Investigations of the Lule Cainozoie strati 
graphic suceession exposed if the coastal xtitls 
south of Adelaide ave currently being Ueader- 
tuken ji the light of previous work it the 
region (eg, Howehin 1923, Campana & 
Wilson 1953, Ward 1966, Cooper 1979) to 
provide information on the mineralogy, 
chemistry and environments of deposition of 
the sediments. About 200 m north of Sellicks 
Tig. within Section 673, which is designated 
a Recreation Reserve (Big. 1), is a bench thot 
has previously eseaped notice and commenti 
apart from a possible passing reference by 
Howchin (1923, p. 310). An understanding 
of the stratigraphic suecession of the Late 
Cainozoie sediments described by Ward 
(196) provides the basis far the interpreta 
tion of the structure an! its possible age and 
origin. 


Stratigraphy 

The oldest rocks ot the sie of the beneh in 
the coastal eliffy are Miocene limestones of 
the Port Willunga Pormation (Cooper 1979). 
‘These reeks, which oceur al levels up to Jom 
above sea level, have been warped into a slight 
anticlinal structure under the boneh. A karst- 
like and calereted surface is preserved an these 
limestones with a Vertical relief in execss of 
4m. The presenee of sificried plant roots on 
the calereied surfaee judicates. the former 
existence of a soi and prolonged exposure 


* Department of Soil Science, Waite Agricultural 
Research Institute, Che University of Adelatle, 
Glen Osmond. SA. 5064, 


prior to the deposition of the overlying sedi- 
ments, 


The major part of the cliffs al Sellicks Trig. 
consists of uUncorsalidated Late Cajnozoie 
mottled sandy clays and thick gravel layers 
and lenses, correlated by Ward (1966, fig. 8) 
with the Late Pliocene Seaford Formation and 
the Barly Pleistocene Ochre Cove Formation. 
A stratigraphic column measured by Ward 
(1966) at Sellicks Trig. is shown in Figure 2. 
The base of (he column, which lies 7.3 m ams, 
is marked by a 7 om thick unil of boulder con- 
wlomerate Which was regarded as being equiva- 
lent in age to the Pliocene Hallett Cove Sand- 
stone, Above this layer 22 m of red, yellow 
and grey clays and gravels wete correlated 
with the Seaford Formation from their type 
section at Ochre Point, Ward (1966) tegarded 
the basal beds of the Seatord Formation as of 
Lower to Middle Pliocene age because they 
allegedly interfinger with Hallett Cove Sand- 
stone at Maslin Bay, und the youngest deposits 
of the Seaford Formation to be oo younger 
than the Late Pliovene, The Seaford Forma- 
tion js overlain by 16,5 m of Ochre Cove For- 
Toation sediments, consisting of red and grey 
angular gravels and conglomerates with some 
lavers of stronyly mottled clays, and inter- 
preted as of Early Pleistocene age (Ward 
1966). Sediments of the proposed Late Pleista- 
cene Taringa Formation lie above those of the 
Ochre Cove Formation, forming a 7 m thick 
unit of grey clay and gravels, grey [rifting 
sandy clay ond a thick gravel layer. The 
sequence al Sellicks Trig. is capped by 6.7 m 
of brown to reddish-brown alluvial gravels 


86 


Coastal Land Slump — Sellicks Beach 


Location Map 


50. «100 


R. I. MAY & R. P, BOURMAN 


Source 1949 Aerial Photograph Svy 42 Photo No 20 
12,500 Topographic Sheets 6527 -35-k and 
SA Dept of Lands 


150 


N 


fo 
-ONEDIN AVE 
DUNE Bl a] 


200 Metres 


fe 
~“SELLICKS_ t 
ATRIG Soe view RE 


Fig. 1, Coastal landslump at Sellicks Beach, Extent of slump is shown by (a), backwall by (b), bench 


by (c) and locality of older slumping by (d). 


and moderately calcareous clays which Ward 
(1966) assigned to the Christies Beach Forma- 
tion of Late Pleistocene age. 

Though Ward’s (1966) interpretation is 
generally acceptable, the discovery and identi- 
fication of the Early Pleistocene marine Burn- 
ham Limestone at the base of the sequence 
requires a re-evaluation of the ages of the oldest 
formations in particular. 

Directly seawards of Sellicks Trig. near a 
small archway cut in the Miocene Port Wil- 
lunga Formation, 1-2 m of sandy clays and 
gravel beds varying in colour from brown, 
grey-black and light olive green to grey with 
weak orange mottles rest on the calcreted sur- 
face of the Miocene limestones. The sandy 
clays in places resemble the Seaford Forma- 
tion. Interbedded with the basal sandy clays 
are several layers of clasts, dominantly cobble- 
sized, which may be the boulder conglomerate 
referred to by Ward (1966). Overlying this is 
a | m thick layer of relatively soft, friable and 
marly carbonate rock which contains a rich 
marine fauna. The limestone has been iden- 


tified as the Early Pleistocene Burnham Lime- 
stone because of its lithology and the presence 
of the distinctive Early Pleistocene gastropod 
Hartungia dennanti chavani (kindly identified 
for us by Dr Ludbrook) described from the 
Point Ellen Formation of Kangaroo Island 
(Ludbrook 1983) and the Roe Calcarenite of 
the Eucla Basin (Ludbrook 1978). The Burn- 
ham Limestone (Firman 1976, Ludbrook 
1983) has not previously been reported from 
this locality, nor has Hartungia previously 
been reported from the Burnham Limestone, 
Between 0.5 to 2 m above the Burnham Lime- 
stone a 10 cm layer of soft, white dolomite 
rests on grey clays. The sandy clays imme- 
diately above the Burnham Limestone appear 
similar to some of those below it; thus the 
limestone may represent a marine interval dur- 
ing the deposition of the sediments of the Sea- 
ford Formation, which in this locality must 
be regarded as having an Early Pleistocene age 
rather than the Mid- to Late Pliocene age 
preferred by Ward (1966). Moreover, we con- 
sider that the Hallett Cove Sandstone at 


COASTAL LANDSLUMPING IN PLEISTOCENE SEDIMENTS, SELLICKS BEACH 


59: 7masl 


CHRISTIES BEACH 
FORMATION 


LATE 53-0m asl 


PLEISTOCENE 


TARINGA 
FORMATION 


45:75m asl 


OCHRE COVE 
FORMATION 


EARLY 
PLEISTOCENE 


29-3masl 


SEAFORD 


PLIOCENE FORMATION 


7-3masl 


PORT WILLUNGA 


MIOCENE FORMATION 


87 


SOIL 


RED BROWN CLAY & GRAVEL 


GREY CLAY & GRAVEL 


GRAVEL LAYER 


GREY SANDY CLAY 
(FRITTING OUTCROP) 


RED & GREY GRAVEL & CLAY 
(FRITTING OUTCROP) 


RED & GREY COARSE GRAVEL 
(CAVERNOUS OUTCROP) 


YELLOW & GREY CLAY 
& GRAVEL 


GREY CLAY & GRAVEL 


RED, YELLOW & GREY CLAY 
& GRAVEL 


HALLETT COVE SANDSTONE 
-BOULDER CONGLOMERATE 


FIG 2 PLEISTOCENE STRATIGRAPHY AT SELLICKS TRIG 
ACCORDING TO WARD (1966) 


Fig. 2. Pleistocene stratigraphy at Sellicks Trig according to Ward (1966). 


Maslin Bay near Tortachilla Trig. does not 
interfinger with the Seaford Formation as sug- 
gested by Ward (1966) but that the Seaford 
Formation overlies the Hallett Cove Sandstone. 
We consider the thin (approximately 1 m 
thick) layer of limestone that occurs within 


the Seaford Formation to be the Early Pleis- 
tocene Burnham Limestone. Consequently, the 
Ochre Cove Formation may be younger than 
the Early Pleistocene age ascribed to it by 
Ward (1966), An Early Pleistocene age for 
the Ochre Cove Formation is essential to the 


88 R, 1. MAY & R. P. BOURMAN 


sea level scheme proposed by Ward (1965, 
1966) and defended by him (Ward 1967) 
against the criticisms of Twidale et al. (1967), 
although subsequently Ward (1975) conceded 
that there are no localities where there is 
certain proof of coastal stability. from which 
reference sea levels can be calcylated. New 
information presented in this paper demon- 
strates that the Ochre Cove Formation is not 
of earliest Pleistocene age, so that a re-evalua- 
tion of both the ages of the sediments and the 
eustatic and climatic history of the area as 
proposed by Ward (1966) is required. 

The precise characteristics of the Seaford 
and Ochre Cove Formations at this locality are 
sometimes difficult to determine because of the 
colluvial mantle of coarse gravels masking the 
underlying sediments. However, occasional 
heavy rains in the winter of 1983 tended to 
strip the colluvial mantle and expose the 
underlying sediments, especially in several 
steep-sided gullies through the section. 


Ward (1966, p. 43) considered that part of 
an inaccessible outcrop at Sellicks Trig. might 
be Ngaltinga Clay although he mapped it as 
part of the Taringa Formation. We believe 
that this outcrop is, in fact, Ngaltinga Clay. 
The inclusion of clasts within these sediments 
may have influenced Ward’s (1966) decision 
to map them as Taringa Formation. However, 
we consider that the location near the base of 
the Willunga escarpment would lead to the 
occurrence of clasts within the generally clast- 
free Ngaltinga Clay. No other characteristics 
of this unit suggest that it is the Taringa For- 
mation, which is typically a grey, calcareous 
mudflow deposit with a pronounced columnar 
structure. Conversely the deposits at Sellicks 
Trig. are olive-green grey in colour with a 
friable structure that is a distinctive feature 
of the basal part of the Ngaltinga Clay. 

A distinctive calcareous breccia containing 
fragments of purple and green metasiltstones 
originally derived from the Cambrian and Pre- 


cambrian basement rocks forms a capping over 
sections of the Christies Beach Formation near 
the modern ground surface. 


Morphology of the bench 


The bench, which lies between 20 m and 
26 m amsl, has a maximum central width of 
50 m and is backed by an arcuate cliff that rises 
a further 30 m to the surface of the piedmont 
plains fronting the Willunga escarpment (Fig. 
1). In detail the bench surface is stepped with 
four major levels being apparent; the highest is 
at the northern end and each is separated by 
a drop of one to two metres to the south, 


Gullies occur in the lower cliff and they have 
eroded headward about 20 m into the bench 
surface, following the bases of steps on the 
bench. Towards the seaward edge of the bench 
crenulated and hummocky features rise up to 
2 m above the general bench level. Exposures 
in these micro-forms reveal disordered gravels 
and clays. Vegetation consisting of grasses and 
stunted bushes is well established on both the 
bench and the northern section of the steep 
backing cliff. Towards the central section of 
the bench, however, the cliffs are bare indicat- 
ing that more recent spalling or settling may 
have occurred here. 


Description of the slump 


Both the gross and detailed morphology of 
the slump are consistent with a large section of 
the cliff having subsided and tilted. The rota- 
tional movement is indicated by the disposi- 
tion of gravel layers exposed in the most 
northerly gully across the bench. The bedding 
of the Pleistocene sediments at Sellicks Beach, 
as a result of their deposition on aggrading 
alluvial fans at the base of the Willunga 
escarpment, dips at approximately 5° to the 
north. On the other hand, the beds in the gully 
dip at up to 45° to the east demonstrating 
pronounced dislocation. At this site recent 


Fig. 3a: Back-tilted Late Pleistocene Christies Beach Formation sediments (CBF) and an inclusion of 
Taringa Formation deposits (TF), overlain by seaward dipping recent colluvium (c). View in 
northern most gully looking south. 3b; Hammer rests on inclusion of mottled Ochre Cove Forma- 
lion sediments within much disturbed deposits of Christies Beach Formation. 3c: Erosiona! hollow 
in Miocene Port Willunga Formation (PWB) limestones through which Pleistocene sediments (P) 
were funnelled. 3d: Vertically tilted Middle Pleistocene Ochre Cove Formation sediments (OCF) 
overlain by horizontally disposed Late Pleistocene cobbles and boulders. 3e: Stranded boulder beach 
(bb) rests on Port Willunga Formation limestone several metres above the modern shore platform. 
3f; Small alluvial fan (f) developed on slump surface by gullying of undisturbed sediments exposed 
in backing cliff. 3g: View of slump surface looking north. Note stepped nature of bench and hum- 
mocky rolls of material in left foreground. 3h: Back-tilted Pleistocene sediments related to older 


phase of slumping near Sellicks Trig. 


COASTAL LANDSLUMPING IN PLEISTOCENE SEDIMENTS, SELLICKS BEACH 89 


90 R. I. MAY & R. P, BOURMAN 


colluvium with a gentle seaward dip overlies 
the steeply dipping gravel beds (Fig. 3a). In 
some other gullies below the bench level the 
sediments have been totally masticated, with 
all original bedding having been destroyed. 
Both the disturbed nature of the sediments and 
the undulating topography near the seaward 
margin of the bench can be attributed to sea- 
ward flow of material following rotational 
slumping of the former cliff-top. 


Sediments of the Ochre Cove Formation 
and younger formations, as described above, 
are exposed in the cliff backing the bench. In 
exposures below the bench, however, sediments 
of the youngest formation at this site, the 
Christies Beach Formation, rest directly on 
the Miocene limestones of the Port Willunga 
Formation or on the Early Pleistocene Burn- 
ham Limestone. The beds of the Seaford For- 
mation have been totally removed by the 
slumping and only occasional patches of dis- 
located Ochre Cove Formation sediments 
occur near the seaward edge of the slump. 
Sediments of the Christies Beach Formation 
have been greatly disturbed with the original 
bedding having been destroyed. Small isolated 
inclusions of mottled Ochre Cove Formation 
sediments up to one metre in diameter occur 
sporadically within the disrupted Christies 
Beach Formation (Fig. 3b). In the northern- 
most gully a two metre diameter slab of a 
grey calcareous sediment that resembles the 
Taringa Formation occurs as an_ inclusion 
within the sediments of the Christies Beach 
Formation even though no Taringa Formation 
appears to be exposed in the cliffs immediately 
backing the bench. It may have derived from 
a small, isolated channel fill. 


Approximately 300,000 m® of material has 
been removed from the cliff face as a result 
of the slump and the total volume of material 
involved in the mass movement must have 
exceeded twice this amount. The volume of 
material involved was such that a frontal lobe 
is likely to have flowed across the beach into 
the sea. The stepped nature of the bench. sur- 
face, the arcuate and steep backwall, the hum- 
mocky nature of the seaward edge of the 
bench and the flowage of a frontal lobe into 
the sea together with backward tilting of the 
sediments is typical of coastal rotational land- 
slumps such as those of Axmouth and Folk- 
stone Warren (Ward 1945), and = smaller 
slumps on the Christchurch Bay coastal cliffs 
(Barton et al. 1983). 


Heath (1963) recognised major slump struc- 
tures in bleached Cretaceous shales near Oodna- 
datta, several of which are of comparable size 
to the slump described here. These structures, 
which occurred on the margins of a duricrust 
capped plateau were established as slumps on 
the basis of excessively steep local dips in 
comparison with the regional disposition of 
bedding. The occurrence of these slumps along 
the weathering front was attributed to under- 
cutting of less resistant shales at the cliff base, 
thereby initiating gravity flows along water 
saturated joints and partings in the kaolinitic 
shales. 


Possible causes of the slump 


Major dislocation of the stratigraphic suc- 
cession is apparent in the structure we have 
identified, and because of the disrupted nature 
of the Christies Beach Formation sediments 
and the occurrence of inclusions of blocks of 
material from older formations within it, 
slumping and saturated flow are the likely 
mechanisms involved. Sediments underlying 
the bench have been tilted but are otherwise 
not disrupted, while at the seaward edge sedi- 
ments of both the Christies Beach Formation 
and the Ochre Cove Formation have flowed 
together in an uncohesive mass. Thorough 
wetting of the sediments facilitating flowage 
in a near saturated condition appears to be one 
prerequisite to explain the evidence observed. 
This would have required intense and/or pro- 
longed rainfall. An impermeable layer at the 
base of the structure would have assisted the 
saturation of the overlying sediments and in- 
creased the possibility of mass failure. The 
Pleistocene sediments at Sellicks Beach are 
relatively permeable because of the coarse 
gravel layers. Similarly the underlying lime- 
stone of the Port Willunga Formation is also 
permeable, but the clays incorporating the 
Burnham Limestone interval may have pro- 
vided a sufficiently impermeable layer to assist 
saturation, and the slip appears to have 
occurred essentially along this layer. In the 
<2 pm fraction of these clays, kaolinite and 
illite are the sub-dominant clay minerals 
present, whilst smectite, which has expansive 
characteristics, is the dominant clay mineral 
and could have acted as a catalyst for the 
slump. 

Several large channels 5 m to 30 m wide 
form part of the karst surface developed on 
the upper surface of the Port Willunga Forma- 
tion and extended to or below beach level 


COASTAL LANDSLUMPING IN PLEISTOCENE SEDIMENTS, SELLICKS BEACH 91 


(Fig. 3c). In these hollows the Barly Pleistn. 
cone Burnham Limestone has been removed, 
whereas on the witervening higher sections, the 
maring sediments usually remain. These 
hollows may have acted like funnels, channel- 
ling the slumped sediment seaward in long 
tongues. 


The proximity of the slump to the Willunga 
Fault sugeests the possibility Of tectonic trig- 


perma, (here om evidence of considerable 
tectonic = uUisplacement aeross the — fault 


zone, Lidhrook (1983) points out that the 
Burnham Limesteng oecurs as thin discrete 
remnants alang the coast south of Adelaide 
ram Kigyston Park to Port Willunmia, docreas- 
ine i Glevation from 30 m at Hallett Cove 
to 20 mat Maslin Bay The distribution of the 
Burnhun Linestone ean aw be extended to 
Selheks Beach where it lies at approximately 
8m ams! thereby corrohoruling Lhe soultrward 
dip of the formation, The die equivalent Point 
Ellen Formation occurs at 50 m ams! at Cape 
Jervis, Variations in elevation between this site 
atid the lower oecurrenees on Kungaroo Island 
Were interpreted by Ludbrank (1983) lo be 
due to gentle warping of les than 1. How- 
ever, the proximity of the Burnham Limestone 
at Sellicks Beach to the marked disturbances 
of Migvene und Middle Pleistocene sediments 
suggests considerable jvclamic dislocation of 
the Early Plestocene Uepnsits across the site 
of ihe Willunga bault at least, 


Campana & Wilson (1953) iusteate Barly 
Pleisiocene gravels mm a clearcut tectonlie Con- 
act with, Cambrian rocks in the Mt Ternble 
Gully 500 m south of Selliecks ‘lrig. The 
oruvels are lilled and ae overlain by younger 
horizontal heds (Campana & Wilson 1953, fig. 
2). These authers also ligure tilled upper 
Pleistovene eravels near the mouth of Mt 
Yerrible Gully (Plate V, fig, 2) to illustrate 
post-Pleistoeenc teetonic uisturbanees. We 
were not uble to locate this last-meniioned 
site. but soully of the mouth of Met ‘Terrible 
Gully where Miocene limestones of the Port 
Willunga Formation have been tilled steeply by 
tectonic movements, Ochre Oove sediments, 
which we regard to be of Middle Pleistocene 
awe, huve been Hhted into a verhcal position 
(Fig, 3d), These beds wid the adjoining Mio- 
vene limestones have been iruneated by the 
sea witen (he shoreline stood shout 4 m higher 
than present, during whieh time a boulder beach 
(Fig, 3¢) was produced and incorporated shell 
fragments and aecasianal Intack mofluses that 


have been superficially abraded. Behind the 
boulders ate back-beach or duae sands con- 
taining calearcous rhizomorphs. This former 
shoreline is horizontally disposed, and extends 
along the eoustline for hundreds of metres, 
although in places there are erosional breaks, 
A fadiocarbon date on the shells incorporated 
within ihe boulder beuch yielded un age in 
excess of 30 000 years B.P, (GaK-6095). Thus 
the heach is probably related to the |ate Pleis- 
tocene Glanville Formation deposited during 
the last interghicial, This shoreline has not 
been abviously tilled. 

The above evidence suggests that tectonie 
cdislacation of Miqvene, Early Pleistocene and 
probable Middle Pleistocene sediments has 
oecurced, Moreover, seismic activity bas con- 
tinued throughoue the St Vineent Basin ta 
moderna times (Sutton & White 1968, Steel 
1962). so that there has been aniple potential 
lor semmic triggeriug of saturated eoastal 
sediments, 

Many coastal cliffs fail due to undercutting 
of the base of the cliff by marine erosion (see, 
for example, Ward (945, Rozier & Reeves 
1979), At present the limestone of the Port 
Willunga Formation provides a bullress to 
erosion of the soficr Pleistocene sedimenis and 
would have done so in the past, which might 
sugeest that manne erosion ig less important 
(haw other factors in generating the slump. 
However, there is considerable evidenee for 
active marine erosion along this section of 
coastline (Mowehin 1923, p. 313, Bourman 
1976). The possibility of a world-wide 1 m 
highee stand of the sea during the Holocene 
lias been suggested (Chappell er af, L982, 
Chappell er al. 1983) and there is some evi- 
dence for a higher shoreline along SL Vineent 
Gulf and Spencer Gulf during the Holocene 
(Ward 1966, Hourman 1972, Gastin et a, 
1981), However, Belperio e¢ al. (1983) main- 
jain thal the evidence is equivocal. If tis pas- 
tulated higher sea leyel did occur, although its 
wecurrenee is not vilul to the initiation af the 
slump, then the processes of marine erosian 
would have latensified, The higher shoreline of 
probable Late Pleistoceng awe that truncates 
the tilted Ochre Cove Formation sediments bs 
not relevant lo this discussion beeause it pres 
dates the Slumping by many thousands of 
years, 

It may scem anomalous that the slump 
uclually occurs behind the buttress of Port 
Willunga Formation limestone, However, the 


92 R. I, MAY & R, P. BOURMAN 


presence of the limestone would have allowed 
the development of a steeper cliff face here 
than elsewhere, whereas the slopes on uncon- 
solidated sediments would have re-adjusted 
rapidly to basal steepening by marine erosion. 
Moreover, where the Port Willunga Formation 
crops out the clay-rich layer including the 
Burnham Limestone is also exposed above sea 
level to provide a slip-plane layer. 

The present steepness of the coastal cliffs 
backing Sellicks Beach is a consequence of 
marine erosion. Moreover, as active rotational 
slipping is essentially confined to slopes under- 
going basal erosion (Brunsden & Jones 1976, 
Goudie 1981, p. 158), the role of marine 
erosion in developing the rotational slump at 
Sellicks Beach is probably far more significant 
than other factors. Immediately south of the 
slump an extensive shore platform occurs in 
the intertidal zone, which has the effect of 
refracting waves to concentrate their attack 
on the cliff fronting the site of the slump; 
this factor may help to explain the precise 
location of the slump. 

Basal steepening by wave attack and satura- 
tion favoured by the presence of an imper- 
meable layer probably established the ideal 
circumstances for rotational slumping to 
occur, while seismic activity or even vibrations 
generated by storm waves may have acted as 
the trigger for the mass failure. 


Possible age of the slumping 


No indications of the slump having taken 
place since European settlement have been 
found. Consequently the failure occurred more 
than 150 years ago. Debris on the beach and 
in St Vincent Gulf would be expected to 
result from outward flowing of the toe of the 
slump. However, no remnants of the flow 
remain seaward of the cliff. Arber (1973) 
described a land slump at Dowlands, Devon, 
England, which produced a 1200 m long reef 
in the sea, but which survived only a few 
months. Thus lack of debris on the beach may 
not be crucial in assessing the age of the 
slumping. Similarly, the vegetation on the 
bench could have become established in a 
relatively short period especially as much of it 
is exotic, Gullies which have been eroded into 
the bench down to beach level and into the 
backing cliff have involved the removal of 
considerable amounts of sediment and the con- 
struction of a small alluvial fan on the bench 
surface (Fig. 3f). Erosion of the bench and the 


backing cliff suggests that more time has lapsed 
since the slumping occurred than indicated by 
either the vegetation present or the lack of 
flow debris on the beach. 

The preservation of detailed hummocky 
topography on the seaward edge of the bench 
surface (Fig. 3g) is suggestive of a relatively 
recent age for the slumping. The bench surface 
topography appears to be more subdued in the 
northern section of the slump, which may sug- 
gest that the southerly section is of a more 
recent age. However, there is no corroboratory 
evidence to confirm that this is the case. An 
S-shaped feature occurs at the base of the cliff, 
immediately landward of the central gully that 
dissects the bench, on the 1949 aerial photo- 
graph (Fig. 1). This structure is now obscured 
by colluvium, derived from the backing cliff. 
This process of colluviation on the bench sur- 
face continues to the present. 

The local stratigraphy is most useful in 
determining the age of the earth movement. 
Sediments of the Christies Beach Formation 
are clearly disrupted by the slumping as is the 
youngest sediment in the backing cliffs, the 
calcareous breccia that overlies the Christies 
Beach Formation. Blocks of the calcareous 
breccia occur on the seaward margin of the 
bench. Slumping therefore postdates the Late 
Pleistocene. A lens of dark sandy clay which 
resembles the Waldeila Formation that occurs 
in nearby Sellicks Creek (Ward 1966) is 
present in a small gully below the bench sug- 
gesting that slumping occurred after or during 
the period of time when the Waldeila Forma- 
tion was being deposited. Shells incorporated 
within the Waldeila Formation in the lower 
Onkaparinga River Valley have been dated at 
4580 += 160 B.P. (Bourman 1979) (GaK- 
6094). Hence a Late Holocene age is postu- 
lated for the rotational slump. The colluvium 
which overlies the back-tilted Christies Beach 
Formation sediments at the northern end of 
the slump supports a Late Holocene age for 
the slump and may offer opportunities for 
colluviation dating, a technique applied suc- 
cessfully on an abandoned London Clay sea 
cliff at Hadleigh in Essex by Hutchinson & 
Gostelow (1976). 

Implications 

The identification of a large land slump at 
Sellicks Beach indicates potential instability 
along the cliff line. Not only is there evidence 
of a large Holocene coastal slump north of 
Sellicks Trig. but immediately south of the 


COASTAL LANDSLUMPING IN PLEISTOCENE SEDIMENTS, SELLICKS BEACH 93 


Fig. 4. View of sediments involved in older slump which occurred through a depression in the Port 
Willunga Formation (PWB). Upthrusting of Burnham Limestone (BL), dolomite layer (d) and 
other light-coloured sediments (1) is especially notable on right side of photograph. Minor bench 
level about half way up cliff at (b) represents remnant of a once more extensive slump surface, 


trig, older slumping has also occurred. There is 
no clear topographic expression of this earlier 
phase of slumping, but sediments have been 
back-tilted and disturbed by mass movement 
(Fig. 3h). This older slump appears to have 
been funnelled through a broad depression in 
the underlying Port Willunga Formation. The 
originally horizontal Burnham Limestone and 
the thin dolomitic layer above it have been 
disrupted and thrust to higher levels by rota- 
tional back-slumping (Fig. 4). Both of the 
major slumps discussed above occurred behind 
buttresses of Miocene limestone, so that a 
natural buttress to erosion does not ensure 
protection from cliff failure. 

Smaller slumps have also occurred on the 
northern side of Mt Terrible Gully in similar 
sediments where fluvial undercutting has 
steepened slopes. This suggests that further 
slumps could be expected along the coastal 
cliffs at Sellicks Beach. During the winter of 
1983 a small scale slump occurred at the 
head of the gully near Sellicks Trig. Thus 
slumping appears to be an important process 
in both the headward erosion of the gullies 
in particular and in the retreat of the coastal 
cliffs generally. 

Small scale slumps pose little immediate 
threat to the houses built on the cliff top, but 
a major slump of the size of the one reported 
here would place a number of dwellings at 


risk. Indications of potential failure might be 
the development of tension cracks or structural 
weaknesses in buildings. Investigations along 
these lines could be used in developing a 
coastal management plan for the area, and 
could be extended to other similar areas such 
as at Kingston Park, where houses have been 
built on steep cliffs underlain by Pleistocene 
sediments close to a fault zone, and at Witton 
Bluff where coastal erosion is very pronounced 
in Cainozoic sediments. 

Man’s effect on the potential for further 
earth movements also requires investigation, At 
Sellicks Beach a pathway has been built down 
to beach level from the cliff top at the northern 
end of the land slump, Large gutters are 
being eroded along the path and these may 
encourage infiltration of water into the cliffs 
and create the potential for further move- 
ment. Rates of infiltration may also be altered 
by runoff from roads, by heavy watering of 
domestic gardens and by extensive use of 
septic waste disposal systems. Any of these 
factors which increase the infiltration of water 
into the sediments has the potential to increase 
cliff instability. 


Acknowledgments 
We are grateful to Professor J. M. Oades 
and Dr A. R. Milnes for comments on the 
paper. Dr N. H. Ludbrook kindly identified 


94 R. I, MAY & R. P. BOURMAN 


Hartungia in samples of the Burnham Lime- 
stone. The illustrations were prepared by 
Michael Tscharke and John Coppi prepared 
the photographs. The aerial photograph was 


reproduced with permission of the S.A. Lands 
Department. Funding for the radiocarbon 
dating was provided by the former Adelaide 
C.A.E. 


References 


ARBER, M, A, (1973) Landslips near Lyme Regis. 
Proc. Geol. Ass, 84, 121-133. 

BeELpPerio, A. P., Hairs, J. R. & Gostin, V. A. 
(1983) A review of Holocene sea levels in 
South Australia. James Cook Univ. Monograph 
Series Occasional Paper No. 3, 37-47. 

BARTON, M. E., Cores, B. J. & Titier, G. R. 
(1983) A _ statistical study of the cliff top 
slumps in part of the Christchurch Bay coastal 
cliffs. Earth Surface Processes and Landforms 
8, 409-422, 

BourMaNn, R. P. (1972) Geomorphic aspects of 
the Onkaparinga Drainage System. Taminga 9, 
117-143. 

(1976) Environmental Geomorphology. 
Examples from the area south of Adelaide. 
Proc, R. Geog. Soc. Aust. (S.A. Branch) 75, 
1-23. 

— (1979) Geomorphological contributions to 
coastal management. Proc. Focus on Our 
Southern Heritage. Con, Council of S.A. & 
Dept. Cont. Ed., Univ. of Adel., pp. 80-88. 

BRUNSDEN, D, & JonEs, D. K. C. (1976) The evo- 
lution of landslide slopes in Dorset. Phil. Trans. 
R. Soc. Lond. A283, 605-631. 

CAMPANA, B, & WILSON, R. B. (1953) The Geo- 
logy of the Jervis and Yankalilla Military 
Sheets. Geol. Surv. S. Aust, Rept. Invest. No. 3, 
26 pp. 

CHAPPELL, J., RHopEs, E. G., THom, B. G. & 
WALLENSKY, E. (1982) ,Hydro-isostasy and the 
sea-level isobase of 5500 B.P. in North Queens- 
land, Australia. Marine Geology 49, 81-90. 

, CHivas, A., WALLENSKY, E., PoLacu, H. A. 
& AHARON, P. (1983) Holocene Palaeo-environ- 
ment changes, central to north Great Barrier 
Reef Inner Zone. B.M.R, Jl Aust. Geol. & Geo- 
phys. 8, 223-235, 

Cooper, B. J, (1979) Eocene to Miocene Strati- 
graphy of the Willunga Embayment. Geol. Surv. 
S. Aust. Rept. Invest. No, 50, 101 pp. 

FirMan, J. B. (1976) Limestone at the base of 
the Pleistocene sequence in South Australia. 
Quart. Geol. Notes Geol, Surv. 8. Aust, 58, 
1-5. 

GosTIn, V. A., Hats, J. R. & Potacn, H, A. 
(1981) Quaternary sea levels in the Upper 
Spencer Gulf, South Australia. Search 12, 43- 
45. 


GouptE, A. (Ed.) (1981) Geomorphological 
Techniques. Geo. Allen & Unwin, London 395 


pp. 

Heatu, G. R. (1963) Large scale slump struc- 
tures southeast of Oodnadatta. Quart. Geol. 
Notes Geol. Surv. S. Aust. 7. 

Howcuin, W. (1923) A geological sketch-section 
of the sea cliffs on the eastern side of Gulf 
St Vincent from Brighton to Sellicks Hill with 
descriptions. Trans. R. Soc. S. Aust. 47, 279-315. 

HutcuHInson, J, N. & GosteLow, T. P. (1976) 
The development of an abandoned cliff in 
London Clay at Hadleigh, Essex. Phil. Trans. R. 
Soc. Lond, A283, 557-604. 

Lupprook, N. H. (1978) Quaternary molluscs of 
the western part of the Eucla Basin. Geol. Surv. 
W. Aust, Bull. 125. 

(1983) Molluscan faunas of the Early Pleis- 
tocene Point Ellen Formation and Burnham 
Limestone, South Australia. Trans, R. Soc. S. 
Aust, 107, 37-50. 

Rozier, 1, T. & Reeves, M. J. (1979) Ground 
movement at Runswick Bay, North Yorkshire. 
Earth Surface Processes 4, 275-280. 

STEEL, T. M. (1962) Subsurface stratigraphy in 
the western suburbs of Adelaide. Quart. Geol. 
Notes Geol. Surv. S. Aust. 2. 

Sutton, D. J. & Wuire, R. E, (1968) The seis- 
micity of South Australia, J. geol. Soc. Aust. 
15, 25-32. 

THom, B. G. & CHAPPELL, J. (1975) Holocene 
sea levels relative to Australia. Search 6, 90- 
93, 

TWIDALE, C. R., DAiLy, B., FIRMAN, J. B. (1967) 
Eustatic and climatic history of the Adelaide 
area, South Australia: a discussion. J. Geol. 75, 
237-242. 

Warp, W. H. (1945) The stability of natural 
slopes. Geogrl J, 105, 170-197. 

Warp, W. T. (1965) Eustatic and climatic history 
of the Adelaide area, South Australia. J. Geol. 
73, 592-602. 

(1966) Geology, geomorphology and soils of 
the southwestern part of County Adelaide, South 
Australia. C.S..R.O. Soil Pub. 23. 

— (1967) The Adelaide area; a reply. J. Geol. 
75, 352-357. 

— (1975) Geology of coral terraces, Huon 
Peninsula, New Guinea: A study of Quaternary 
tectonic movements and sea level changes: 
Discussion. Bull. Geol, Soc. Am, 86, 1482-1484. 


THE ENIGMA OF THE TINDAL PLAIN, NORTHERN TERRITORY 


BY C. R. TWIDALE 


Summary 


The Tindal Plain, near Katherine, N.T., is a karst plain notable for its pinnacles, for prongs 
developed beneath the natural land surface, and for its survival. The Plain is of exhaumed type, and 
is the surface upon which the Lower Cretaceous Mullaman beds were deposited. It is suggested that 
the bedrock has survived marine transgression because the sea was essentially staurated with lime, 
and has resisted subsequent epigene attack beacause the bare bedrock is essentially dry, though 
doline and cave development are active where there is a (moist) soil cover. 


THE ENIGMA OF THE TINDAL PLAIN, NORTHERN TERRITORY 


by C. R. TwiDALe* 


Summary 
TwiDALe, C. R. (1984) The enigma of the Tindal Plain, Northern Territory. Trans. R. Soc. S. 


Aust, 108(2), 95-103, 12 June, 1984, 


The Tindal Plain, near Katherine, N.T., is a karst plain notable for its pinnacles, for prongs 
developed beneath the natural land surface, and for its survival. The Plain is of exhumed 
type, and is the surface upon which the Lower Cretaceous Mullaman beds were deposited. It 
is suggested that the bedrock has survived marine transgression because the sea was essentially 
saturated with lime, and has resisted subsequent epigene attack because the bare bedrock is 
essentially dry, though doline and cave development are active where there is a (moist) soil 


cover. 


Key Worpbs: Karst, exhumed surface, dolines, Tindal Plain, Northern Territory. 


Introduction 


Between Venn and Katherine (Fig. 1) the 
Stuart Highway, and the now disused North 
Australian Railway, both pass over a virtually 
flat limestone surface, underlain by and deve- 
loped upon the Tindall Limestone, of Middle 
Cambrian age (Randal 1963) and here called 
the Tindal Plain after the air base centrally 
located upon it. (The discrepancy in spelling 
is due to error by the geological nomenclature 
authority.) This karst plain occupies an area 
of roughly 45 km* extending several km on 
either side of the S.E.-N.W. transport corridor. 


if 
SAE ATHERINE 


Fig. 1. Locality of Tindal Plain in Northern Ter- 
ritory, Australia, 


* University of Adelaide, Box 498, G.P.O., Ade- 
laide, S, Aust. 5001, 


It is a broadly rolling feature situated between 
140 m and 200 m above sea level. Though 
there is a relief amplitude of some 60 m 
within the area occupied by the plain, slopes 
are gentle save in the vicinity of the few 
widely-spaced incised river channels, some of 
which drain eastwards to the Gulf of Carpen- 
taria and others westwards to the Joseph 
Bonaparte Gulf, and around the quite 
numerous but areally minor low hills that 
stand above the level of the plain. Many of 
these residuals can be regarded as properly 
belonging to other, adjacent physiographic 
regions, but they are of particular interest in 
that they provide vital evidence concerning 
the essential character of the Tindal Plain. 
There is an anomaly in that though all earlier 
writers, whether considering the plain in 
general terms (Wright 1963, Randal 1963) or 
as a karst form (Jennings & Sweeting 1963), 
regarded it as a youthful feature, the field 
evidence strongly suggests that it is an ancient 
feature of complex derivation, and in some 
respects difficult to explain. The limestone 
plain crossed by the Stuart Highway appears 
to be at least 135 Ma old, and the question 
arises as to how a feature eroded in such a 
weak material as limestone has so successfully 
survived attack by weathering and erosion. 
This is the enigma of the Tindal Plain. 


Background 


The Tindal Plain is located within the 
Daly Basin, a broad, shallow intracratonic 
structural basin underlain by folded Protero- 
zoic strata and associated igneous emplace- 
ments, and occupied by a suite of Cambrian 
rocks that includes the Antrim Plateau Vol- 
canics and various limestones prominent 


96 Cc. R. TWIDALE 


amongst which are the Montejinni, Oolloo 
and Tindall formations, and by the Lower 
Cretaceous Mullaman beds.'. The basin struc- 
ture predates Cretaceous sedimentation and 
has suffered recurrent reactivation, the Lower 
Cretaceous quartzitic sequence, for instance, 
appearing from beneath the succeeding beds 
on the divide between the Daly Basin and the 
Wiso Basin to the south. 


The Tindall Limestone is a massive crystal- 
line rock with dolomite bands and lenses and 
thin beds or attenuated lenses of chert that 
are early diagenetic features. In the Tindal 
area the limestone is well bedded and jointed 
and essentially flat-lying, but the fractures are 
widely spaced so that outcrops are dominated 
by large angular blocks. The limestone is 
inherently white in clolor, but where exposed 
in the subsurface typically appears to be of a 
creamy hue, this impression being due partly 
to a skin of calcite and partly to the con- 
trasted red colour of the adjacent soil. Where 
exposed to the air, however, the limestone is 
consistently dark grey or black, this colour 
being due to an algal encrustation that forms 
a thin surficial patina, Soot from the periodic 
burnings to which this district is subject may 
also have contributed to the surface discolora- 
tion. The Tindall Limestone retains these 
characteristics on both sides of the Stuart 
Highway, but the Oolloo Limestone exposed 
in the Fitzmah Quarry some fifty km south- 
west of Katherine is finely bedded and grey in 
colour, and, unlike the Tindall, has no pin- 
nacles or other residuals associated with it. 
Again, the karst plain developed on the well 
bedded, almost flaggy Montejinni Limestone 
around Top Springs, is featureless and lacks 
the blocky residuals characteristic of the 
Tindall Limestone outcrops. 

The Cambrian rocks are unconformably 
overlain by the Mullaman beds which consist 
of a basal quartzite exposed on_ structural 
highs. According to Skwarko (1966) the basal 
member of the Mullaman beds, the Lees Sand- 
stone, is a massive, poorly sorted saccha- 
roidal quartz sandstone. The lower members 
are characterised by cross-bedding and plant 
remains suggestive of a lacustrine environment, 
though the higher part of the sequence is 
regarded as marine, An overlying argillaceous 
member is well-bedded and jointed and con- 
sists of siltstone and mudstone, but is com- 


'Mullaman beds, lower case “b’, is the official 


stratigraphic term. 


monly weathered to a lateritic profile consist- 
ing of a silty A-horizon, a pisolitic ferruginous 
B-horizon and a C-horizon that comprises 
mottled and pallid porcellanic material. Where 
exposed, the sandstone too is silicified and 
slightly ferruginised. In places a pisolitic ferru- 
ginous layer is preserved either as a surface 
cover or as an infilling to minor pipes and 
basins developed on the quartzite. 


Lateritisation took place in this region in 
the later Cretaceous and Early and Middle 
Cainozoic, ceasing in the Miocene, for various 
later Cainozoic sediments and lavas remain 
unaffected by lateritic weathering (Stewart 
1954, Hays 1967, Twidale 1956, 1983). Judg- 
ing from the conditions in which lateritic 
soils are forming at present, the climate 
during this extended period must basically 
have been warm and humid (Prescott & 
Pendleton 1952, Sivarajasingham et al. 1960, 
Maignien 1966) a conclusion confirmed by 
palaeobotanical investigations (see e.g. Kemp 
1978). The present climate is warm and sub- 
humid, Katherine for instance averaging 966 
mm of rain per annum, 95% of it falling in 
the summer seven months October-April, but 
laterite is not evidently forming at present, and 
though contrary views are expressed from 
time to time, it is generally agreed that the 
laterite of northern Australia is a relic feature. 


Lateritisation ceased sometime during the 
Miocene, partly as a result of regional warp- 
ing that caused the rivers that had reduced 
the land surface to one of low relief to be 
rejuvenated and resume valley incision, partly 
as a consequence of major world wide climatic 
changes which both directly affected the 
environment and also, through the concomi- 
tant lowerings of sea level, added to the ten- 
dency of exorcic rivers such as those that drain 
the Daly Basin to incise their beds, According 
to Wright (1963) these streams exploited 
variations in the laterite profiles to produce 
several plains and benches, and in toto a 
stepped relief, The master surface capped by 
laterite and named the Bradshaw Surface is 
represented by various scattered but prominent 
mesa and plateau remnants, with the Maran- 
boy and Tipperary surfaces at lower levels. 
Hay (1967) mentions a _ sub-Cretaceous 
exhumed surface that is lateritised in the area 
west of Mataranka, but does not discuss the 
Tindal Plain, which is not lateritised and 
which lies northwest of Mataranka. 


TINDAL PLAIN, N.T. oT 


In the course of a brief reconnaissance, 
Jennings & Sweeting (1963) noted the bare 
karst developments and also some recently 
formed dolines sguth of (Katherine. They 
related the soil cover to the present climatic 
conditions and accepted Litchfield’s (1952) 
interpretation of the sandstone that forms 
scattered outcrops on the plain as part of the 
same stratigraphic formation as the limestone 
that underlies it, a view which, in turn, 
reflected the prevailing geological opinion of 
the time. Thus they were led to consider the 
karst plain as a youthful feature, despite the 
presence of limestone pinnacles which are 
prominently developed on the Plain, and 
which are generally considered as representa- 
tive of mature karst landscapes; though the 
term “youthful” may have been used in the 
general sense of “little-dissected”, which the 
Tindal Plain certainly is. 


Evidence 

The Plain Proper: Two types of surface can 
be distinguished within the Plain, First, there 
are the many areas of exposed limestone. They 
have a well developed system of N—-S and 
E-W trending joints disposed — essentially 
normal to one another (Fig, 2) and with the 
prominent horizontal bedding, forming an 
orthogonal set that subdivides the limestone 
into cubic and quadrangular blocks. In detail, 
these rock outcrops are rough due to the 
development of innumerable clints, tables and 
blocks separated by wide grikes or widened 
and weathered vertical joint planes (Fig. 3). 
The limestone blocks are typically fluted, 
though the furrows are vertically zoned, the 
lower parts of the blocks being not only 
smoother but also steeper and paler in colour, 
suggesting a recent lowering of the soil surface 
between the blocks of some 20-30 cm, In 
places the presumed former soil level is 
mirked by a rim of calcite, Also the thin 
interbeds and attenuated lenses of chert stand 
out as minor ledges and rims. There are also 
miuany small, shallow dolines which lead down 
to quile extensive cave systems such as that 
at Cutta Cutta, These vertical shafts provide 
good exposures of the honeycombed nature of 
the rock immediately beneath the blocky out- 
crops (Fig. 4). 

The second type of surface (Fig. 2a) is 
underlain by a greater or lesser thickness of 
red or light brown soil. The presence ol 
quarries in this soil immediately adjacent to 
rock outcrops (as, for instance, near the 


Fig. 2. (a2) Rock and soil plains, Tindal area, N.T. 
(b) Detail of karst plain, near Tindal. 


Fig. 3. Limestone tables with fluted sidewalls, and, 
near hammer head, Cretaceous sandstone pre- 
served in bedding plane. 


southern extremity of the Tindal runway) sug- 
gests that the contacts between fresh rock and 
the soil are steep. According to Litchfield 
(1952) the soils consist predominantly of fine 
sand, though with an increasing proportion of 
silt and clay with depth. They are only mode- 
rately alkaline (pH 6.2—7.2) throughout, and 
their depth varies between a few em to a least 


98 C. R. TWIDALE 


m wl 


Fig. 4. Honeycombed limestone exposed near 
mouth of doline near Tindal. 


9 metres. These soils are unlikely to have been 
derived entirely from the weathering of the 
Tindall Limestone, and are more likely a mix- 
ture of ancient red earth and basal Mullaman 
sandstone. 

Dolines developed in the late ‘seventies 
near the old Venn airstrip provide excellent 
exposures of the soil-rock contact. A group of 
six dolines is developed in a broad, shallow 
topographic depression between the Stuart 
Highway and the old North Australia Railway 
embankment about 3.5 km northwest of the 
Cutta Cutta Park turnoff (Fig. 5). Two of 


eA es ie 

3 ie = = Jat ik. ~ is ee 

Fig. 5. Dolines formed within the last decade near 
Venn, N.T. 


the dolines are of quite large diameter, the 
others small. The largest doline has a diameter 
of about 13.5 m and though it can fairly be 
described as circular in plan it is in detail 
irregular and scalloped in outline. It is about 
9 m deep. The others are similar in depth 
but are of lesser diameter, The dolines are 
interesting not only because of their recent 
development and their having formed in an 
ill-drained depression that is partly of anthro- 
pogenic origin, but also because the form of 


the contact between the red soil and the 
cream-coloured limestone is well exposed, 
After the development of the dolines, soil 
from the adjacent areas was washed into the 
depressions, exposing the irregular basined 
bedrock surface (Fig. 6). The limestone basins 
are separated by sharp ridges or arétes that 
form a polygonal pattern in plan. The area of 
such basins and arétes extends beyond the 
confines of the topographic depression and to 
the west of the railway embankment, suggest- 
ing that the fine soil has been washed through 
the matrix of the ballast that forms the 
embankment and into the dolines. 


Fig. 6, Basined limestone surface exposed as result 
of original soil cover being washed into recently 
formed dolines near Venn. 


Isolated blocks (Karrenblocke, Karren- 
steine) can. be seen in various stages of expo- 
sure from the natural soil cover, and the 
naturally buried rock surfaces display a dif- 
ferential etching of the calcite crystals to pro- 
duce a rough surface comparable to the pitting 
described from granitic terrains (Twidale & 
Bourne 1976) and seen also developed on 
limestone, though in the zone of seasonally 
Galong in central N.S.W. The presence of 
corestones set in a matrix of red soil shows 
beyond doubt that some, at least, of the free 
boulders exposed at the surface owe their 
origin to differential subsurface moisture 
attack, 


The upper surface of the limestone mass is 
highly irregular with tall prongs or fingers 
(one of them with flared sidewalls) up to 3 m 
high and separated by deep soil-filled pipes 
comparable to the gulls, puits naturely or 
orgues géologiques of the European chalk 
country (Fig. 7). 


TINDAL PLAIN, N.T. 99 


. a y 
+ te ot 
Fig, 7. Sidewall of recent doline near Venn, with 
limestone prongs and pipes filled with red soil 
between. 


A similar bedrock morphology, with lime- 
stone prongs, about 6 m high, projecting into 
the soil cover was noted in 1959 by M. M. 
Sweeting and J. N. Jennings in a_ freshly 
developed doline adjacent to, and indeed 
encroaching upon, the Stuart Highway, some 
27 km southeast of Katherine (Jennings, pers. 
comm., December 1983), It is interesting to 
speculate that the small pinnacles that are 
prominent in parts of the area (see below and 
Fig. 8) may be, in part, such prongs that 
have been subjected to sculpture and super- 
ficial modification after exposure. 

About | km to the southeast (between the 
main doline site and the Cutta Cutta turnoff), 
but also in a broad natural topographic 
depression and located between the railway 
embankment and the road, there were 
(August, 1983) two shallow (less than 1 m) 
depressions in the red soil plain, They are 
incipient dolines, and are defined by dirt 
scarps; associated with them are a few arétes 
of limestone, indicating that even at this early 
stage of development soil has already been 


washed into the depression from the sur- 


rounding areas. 


Fig. 8. Pinnacles with fluted sidewalls, northwest 
of Katherine, N.T-. 


Residual Hills: The plain is broken by residual 
hills of several types. Though quite numerous, 
their total area is small; nevertheless they 
provide essential evidence as to the age and 
character of the Tindal Plain. Small groups of 
limestone boulders together form low rises 
that might be termed low blocky nubbins, The 
limestone blocks, tables, pinnacles (or minor 
towers) and perched blocks are black and 
intricately fluted (Fig. 8). Some honeycomb 
weathering is present. The open bedding 
planes are prominent, and rock basins are also 
developed. In some areas pinnacles up to 4m 
tall dominate the groups, as for instance on 
the western side of the Stuart Highway, some 
13 km northwest of Katherine. 

Also present are several low hills, standing 
up to 7-8 m above the plain level, that are 
blocky, but nevertheless plateau-like. Again, 
every exposed surface is black and fluted, and 
bedding planes are prominent. In several areas, 
and especially toward the margins of the 
Plain, there are mesas of lateritised Lees 
Sandstone that are preserved by virtue of the 
ferruginous capping, or lower mesas of porcel- 
lanite, some of which lack any caprock. 


Perhaps of greatest interest, however, are 
the many residuals, high and low, that consist 
of Lees Sandstone boulders, most of them in 
situ. In some instances there is a scatter of 
blocks and boulders, elsewhere merely a “skin” 
of sandstone on the underlying limestone (as, 
for instance, near the display centre in the 
Cutta Cutta Park), but at some sites there are 
blocky sandstone hills standing up to 10 m 
above the plain. Weathering rinds are well 


100 C. R. TWIDALE 


developed on some of the boulders. Other 
features include mushroom or hoodoo rocks, 
polygonal cracking, flared boulders and slopes, 
grooves or Rille, and several more or less 
bizarre conformations. In these areas the soil 
is locally and superficially sandy due to wash 
from the residuals, but the most significant 
feature of these sandstone hills, a characteristic 
they share with the lateritised and porcellanic 
remnants, is that the unconformity between 
the Cretaceous rocks and the underlying 
Cambrian limestones is consistently contiguous 
with the level of the adjacent plain. 


exhumed type of pre-Cretaceous age is surely 
the only explanation that accommodates the 
observed data. It cannot be argued that the 
plain is a Late Cainozoic, much less a recent 
feature. What are construed as basal Mulla- 
man beds are intimately associated with the 
karst forms, major and minor, It is concluded 
that the Plain and many of the karstic forms 
associated with it were already in existence 
when the Mullaman beds were laid down. 
The karst forms cannot be attributed to 
differential weathering beneath a cover of 
Mullaman beds, for the basal Mullaman is 


laterite 

capped (K) (K) 

plateau sandstone sandstone limestone sandstone 
nubbin block mesa pinnacle nubbin 


' 


- oF - A A unconformity 
5 a Sete SS 


limestone 
table 


Fig. 9. Diagrammatic section through Tindal Plain, showing relationship between laterite, Mullaman 


beds and Tindall Limestone. 


Nature of the Tindal Plain 


There are so many residuals in which the 
unconformity is continuous with plain level 
(Fig. 9) that it cannot be fortuitous. To argue 
that the limestone between the surviving 
remants has everywhere been reduced to the 
level of the unconformity is asking overmuch 
of coincidence. The Tindal Plain is essentially 
the surface on which the Lower Cretaceous 
Lees Sandstone was deposited. It is therefore 
an exhumed plain of pre-Cretaceous age, This 
interpretation is confirmed by the common 
occurrence of quartzite boulders, some of them 
with honeycomb weathering, both on _ the 
rocky limestone plain and scattered amongst 
the limestone blocks of the plain and of low 
rises. The presence of small blocks of Lees 
Sandstone (as, for instance, at an outcrop 
about 4.5 km northwest of the Cutta Cutta 
turnoff) in widened bedding planes, on joint 
planes and within the flutings and tubes that 
comprise honeycomb weathering lends support 
to this suggestion. Also, and more commonly, 
a skin of red gritty sandstone with a thin skin 
of silica, which is interpreted as a mixture of 
basal Lees Sandstone and the red soil and 
developed on the Tindall Limestone in pre- 
Cretaceous times and incorporated in the basal 
Mullaman beds (Fig. 3), is preserved on the 
limestone blocks. That the Tindal Plain is of 


essentially unaltered and rests on fresh Tindall 
Limestone. The basal Mullaman beds are 
slightly more ferruginous than the rest of the 
section (and this may be associated with the 
later lateritisation) but nowhere is there a red 
soil preserved between fresh sandstone and 
fresh limestone. For the same reason, the 
karst plain and associated low limestone hills 
cannot be interpreted as a simple etch surface 
due to stripping of an earlier developed 
regolith, for if such were the case a regolith 
would surely be preserved between sandstone 
and limestone? On the other hand there is 
clear evidence of active dissolution beneath 
the land surface, beneath the soil cover; con- 
temporary solution has at least maintained the 
karstic nature of the rocky plains. 


Sequence of Events 


The most likely sequence of events to 
which the present landscape may reasonably 
be attributed involves the Cambrian strata 
being weathered and reduced to a surface 
of low relief by the Late Jurassic, prior to the 
deposition of the Early Cretaceous Mullaman 
beds. As mentioned previously, these earliest 
Cretaceous strata are supposed by Skwarko 
(1966) to have been laid down in shallow 
lakes, and, if so, it may be that wave action 
stripped the regolith veneer and incorporated 


PENDAL PLAEN, NLT, Wt 


iL infu the newly deposited basal sediments. 
Certainly, where wave achon is known to have 
eroded } regulith (as, for instance, on north- 
weste¢n Eyre Peninsula during the Late 
Conovoic (Twithile ef af, 1977)) packets af 
soil Have been preserved beneath the incoming 
rock cover, and these may iclide the red 
soils exposed in the Venn dolines and ctse- 
where, despite Litchfield’s (1952) assertion 
that the red soils are im equilibrium with the 
present covitonnient, The present conditions 
uy not be very different from those that 
obnine! 1301-140 Ma ago, am the regolith 
could be a polyeendtig Feature that bas altered 
in respunse to changing environment. What is 
cern is that siream rejuvenation during the 
fate Cainoroig hus caused the stripping of 
mast of the Cretaceous cover, resulting in. the 
reesposure of the unconformity between the 
Mesozoic find Palacavuie beds. The Tindal 
Plain, and the agsodiiied limestone pinnacles 
and low plateaus. melding several of the 
minor karst features, ace exhumed limdforms 
ol preCretaceous age 


Preservation 

Apart from ay antiquity and complex origin, 
the preservation of the Tindal Plain presents 
several difieulies. The Mullaman beds that 
formerly blaiketed the reyian were subyected 
1) deep. intense and long-eantinued weathering 
during the Late Cretavcous and through much 
of the vacher Caiezoie, This weathering 
resulted in widespread lateritisation wherever 
the wrujllaccaus sequences were affected. ‘The 
sandstone was superficially silivified and lerru- 
viniserl, wilh coheomitant piping and hasin 
development indicative of silica solution. dn 
these conditions TL iy difficult to understand 
how (he underlying limestone was nol alfered 
la such un extent (hat all original structure 
was last. ‘Ihe limestone is certainly well 
bedded and jointed, and presumably allowed 
free pisage of the mereorie or vadose walers 
infiimiing inte the subsurface. 

Vhere is some suggestion that a silieitied 
lmmpermeable horizow evolved within the 
niin Taterite progile (Wright (9463) but it is 
unlikely fa have been completely watertight 
and, indeed, the depth of weathering beneath 
ilus horizon within the Mullaran beds argues 
weit sich it caplanation, Likewise, the 
porevllanite that is developed on the arwilhit 
ceeous Mullan beds is silitied. but is too 
well (raetured to be impervious, The busal ced 
beds af the Mullaman. those that are found 


plastered on joint faces, etc,, are silieifivd, but 
there is no evidence that the silicified material 
is any more than au patina: permeable Lees 
Sandstone occurs within widened Cractures, 
and this would not prevent the pitssage of 
groundwuters. 


It may be suggested that the unconformity 
und the limestone were allected by stagnant 
saturated groundwaters within a closed system, 
but, in view of the known perviousness of the 
Palueozoie basin sediments and of the Pro- 
terozoic basement rocks, this is unlikely. 
Similarly, any argument (hat the water table 
was deep below the uncontormily and that 
groundwaters merely passed through the rele- 
vant vones does nok bear eXamination, for it 
is preeisely such throughflow that 1s conducive 
to karst development, especially vertical 
shafts, vMlso, the develapment of 4 lateritic 
profile on the Mullaman beds surcly argues a 
fluctuating water table well above the uncon- 
formity, and the latter located within the zone 
of permanent saturation. It can be argued 
thar the groundwaters, moving — laterally 
through the system, were of low pH, but this 
is unlikely in view of their provenance and 
the evidence of silica solution at the time of 
lateritisation, 


What appeurs at first Sight to be a com- 
parable situation obtains in the lower Mt 
Arden Valley, in the southern Flinders Ranges, 
S.A. There, considerable elements of a planate 
surface cut io Cambrian liniestene are pre- 
served #8 a high plain, now dissected, but with 
ycatlered remoants of a cross-bedded econgla- 
merate and sandstone persisting in luw, bloeky 
ourcrups. The arenaceous strata which have in 
detail been sculptured by alkaline sar} meis- 
jure, producing rocks. busins, minor flared 
slopes, ete, were laid down in early ‘Pertiary 
(Foeene) Gmes in a lake that eectipied the 
northern Willovhrsa Basin (Twulale 1966, 
1980), Agaip, the question arises as fo how 
the limestone survived solution No pre- 
Tertiary regolith is preserved at the wicon- 
formity und the limestone surface, though 
weathered Lo produce minor karst features 
Clingerprints, mindr Karen, basins), has not 
so tar yielded any dolines or cave systetis. 
But ihe Cumbrian strata are folded and the 
fractures to the synelinal stricture are argue 
ably so tight as not readily to permit the 
pussuge of water. Ne such tentative explana 
tion, can, however, be applied to the ‘Tindal 


102 C. R. TWIDALE 


region, where the limestone is flat-lying with 
open joints and bedding planes. 


The problem of the preservation of the 
Tindal karst can be eased, though not resolved, 
if a basic assumption made by an earlier 
worker is rejected. The greater part of the 
present karst, including the differential com- 
partmental weathering, was developed by the 
end of the Jurassic. The old landscape was 
then buried by the Mullaman beds, with 
regolith material being stripped and redistri- 
buted by wave action. Skwarko (1966) sug- 
gested that the earliest Cretaceous beds were 
deposited in lakes, largely on the basis of the 
contained plant (fern) remains. But this 
interpretation is open to question. The Lees 
Sandstone is so widespread that the series of 
discrete lakes envisaged must have been of 
enormous extent (cf. Twidale & Milnes 1983 
with respect to Miocene lake basin deposits 
in central Australia). Also the sub-Cretacecous 
surface is consistently of low relief and the 
question arises as to what rivers carried the 
sands and spread them over the landscape? 
There is no large scale cross-bedding and rapid 
step-up of beds, such as is commonly found in 
fluvial or lacustrine sandstones, It is much 
more in keeping with the distribution and 
character of the Lees Sandstone to suggest that 
it is marine and associated with an epicon- 
tinental sea. The plant remains could have 
derived from islands, or simply have been 
washed into such a sea. In such conditions, 
leaving aside the associated relatively narrow 
coastal zones, where various types of erosion 
would likely have been active, the limestone 
floor may have been only superficially affected, 
for the sea water, if comparable with that of 
today, would have been saturated with lime 
and therefore incapable of carbonate dissolu- 
tion. Even so, it is difficult to understand 
why marine erosion (physical, chemical, 
biotic) did not eliminate such comparatively 
fragile features as the pinnacles. It may be 
that the small limestone plateaux and _ hills 
stood as islands in a shallow sea and survived 
in a low energy environment. 


Finally, why has the limestone surface not 
been weathered to a far greater extent than 
it has since exposure in Late Cainozoic times? 
After all, the region receives a considerable 
rainfall, albeit on a markedly seasonal basis, 
and the area is covered by a woodland that 
produces a considerable litter which decays to 
produce various organic acids (see e.g. 


Hingston 1962). Rates of limestone solution 
vary considerably (see e.g. Sweeting 1973 pp. 
41-42) but are almost everywhere, and cer- 
tainly under the prevailing conditions, enough 
to have caused a deep soil to develop through- 
out the plain under consideration. Un- 
doubtedly there has been some recent solution 
and, indeed, recent doline development, but 
large areas of the plain have been preserved. 
Moreover there is evidence that the rocky 
nature of the plain is being enhanced as soil 
is washed away, possibly as a result of anthro- 
pogenically introduced environmental im- 
balance. Even in areas of well-bedded and 
jointed limestone, such as that exposed in the 
Fitzmah Quarry, little subsurface weathering 
is in evidence. 

One possible explanation is that, as has 
been suggested by several authors (de Saussure 
1796 in general terms, Monroe 1964 p. 108, 
Panos 1964 p. 12, Panos & Stelcl 1968, Gams 
1979, 1981 with respect to limestone, Barton 
1916, Bain 1923 with regard to granitic 
rocks), rocks that are dry are weathered only 
very slowly, whereas those that are wet, and 
especially those that are covered by a moisture- 
retaining regolith, are altered very rapidly. 
And the Tindal Plain may well be effectively 
arid, There is a long dry season; in the 
summer wet, water rapidly disappears beneath 
the surface into the cavernous riddled rocks; 
burning ensures that there is little in the way 
of ground cover to retain moisture; and human 
interference added to the natural systems has 
recently caused accelerated soil erosion, caus- 
ing the moisture retaining soil to be stripped 
away to a depth of 20-30 cm and leaving the 
bare rock that much in relief. Moreover, there 
is a reinforcement effect (Twidale er al. 1974), 
for the more an outcrop stands in relief, the 
greater is the tendency for water to run from 
it to the adjacent plains. 

Beneath the soil-covered plains, on the other 
hand, the exposures revealed in the Venn 
dolines suggests that solution not only has 
been, but probably still is, active and has 
produced a differentiated relief at the weather- 
ing front. Thus Monroe (1964 p. 108) had 
good reason to state that “The solution of 
limestone is most active under soil cover and 
almost ceases when soil is removed”. 


Conclusion 
The Tindal Plain, which heretofore has been 
accepted, when it has received any attention 
at all, as a simple, youthful landform, is in 


TINDAL PLAIN, N.T. 103 


reality a feature of considerable antiquity and 
complexity, the ortgin and persistence of which 
stil] pose considerable problems, 


Acknowledgments 


The writer thanks Australian National 
and the Northern Territory Department of 


Mines & Energy for facilitating some of the 
field work on which this paper is based; 
Dr J. N. Jennings (Institute of Research 
Studies, A\N.U.) and an anonymous referee 
for consiructive comments on a draft of this 
paper; Mrs E. M, Campbell and Ms Sabina 
Douglas-Hill for assistance in the field in 
1979 and 19834 respectively. 


References 


Bain, A. DB. N. (1923) The formation of insel- 
berge, Geol. Mag, 60, 97-107. 

Barron, D. C. (1916) Notes on the disintegration 
of granite in Egypt. J. Geol. 24, 382-393 , 

Gams, I, (1979) International comparative study 
of limestone solution by means of standard tab- 
lets. Actes Symp. Imer, sur Verasion Karstique. 
UALS, 1979, pp. 71-73. 

——. (1981) Comparative research of limestone 
solution by means of standard tablets. Proc. 8h 
Tnier. Cong. Speleol, l, 273-5. 

Hays, §. (1967) Lund surfaces and Jaterites in 
the north of the Northern Territory. pp. 182- 
210, Ja J. N. Jennings & J. A. Babbutt (eds) 
Landform Studies for Australia and New 
Guinea. (Aust Natl Univ. Press: Canberra,) 

Hinasron, PF, J, (1962) Activity of polyphenolic 
constituents of leaves of Eucalyptiy and other 
species in complexing and dissolving iron oxide, 
Aust. J. Soil Res, 1, 63-73, 

JENNINGS, J. N. & Swoetinc, M, M. (1963) The 
Limestone Ranges of the Fitzroy Basin, Western 
Australin, Bonn, Geogr. Abh, 32, 7-60. 

Keme, E. M, (1978) Tertiary climatic evolution 
and Vegetation history in the southwest Indian 
Ocean region. Palaeogeog,  Palaeoclimatol, 
Palacwecol, 24, 169-208. 

Lircacinnp, W. (1952) Soil surveys in the 
Katherine district, Northern Territory, 
C.S.1.R.0. Div. Soils, Div, Rept, 7/52. 

Mauinten, R. (1966) Review of research on 
laterites. Nat, Resour, Res, IV (UNESCO; 
Paris.) 

Morro, W, H. (1964) Lithological control and 
iropicul karst topography, Proc. 20th Int. 
Geogr! Congr., London, 107-108. 

Panos. V. (1964) Der Urkarst im Ostflugel der 
Bohmischen Masse. 4, Geamearph. 8, 105-162, 

& Srrreu, O. (1968) Physiographic and geo- 

logic control in development of Cuban mogotes. 

thid, 12, V1L7-165, 


Prvscorr, J. A. & Penpieron, R, L, (1952) 
Latente and lateritic soils, Comm. Bur. Seil Sci. 
Tech. Conun, 47. 

RawpaL, M. A. (1963) Katherine N.T., B.M.R. 
Explanatory Notes, 1:250 000 Geological Series, 
Sheet SD/53-9. 

Ssussure, H, B.. de (1796) 
Alpes. (Fauche: Neuchatel.) 

SAVARATASINGHAM, §,, ALEXANDER, L. T,, Capy, 
J. G. & Crm, M. G. (1962) Laterite. Ady. 
Agran, 14. 

Skwarko, S. K. (1966) Cretaceous stratigraphy 
and pulavontology of the Northern ‘Verritory. 
BM.R, Bull. 73. 

Srewarr, Gi, A. (1954) eden kynalcry of the 
Barkly region. Jv Survey of the Barkly region 
1947-8. Land Res, Series 3, 42-58, 

Sweerna, M, M. (1973) Karst 
(Columbia U.P. New York.) 

Twoare, €. R. (1966) Chronology of denudation 
in the southern Flinders Ranges, South Austra- 
lin. Tras, Ry Sec, S. Aust. 90, 3-28, 

— (1980) Landforms, pp. 13-41, Ja D. W. P, 
Corbett (ed.) A Field Guide to the Flinders 
Ranges, (Rigby: Adelaide.) 

(1983) Australian laterites and. sileretes: 
ages und significance, Rev, Geol, Dynan, & 
Giagr, Phys. 24, 35-45, 

—_, Bourne, J, A, & Smrn, D. M. (1974) Ree 
inforcement and stabilisation mechanisms in 
lnndform development. Rev. Géamorph, Dynam. 
23, 115-125. 

. Bourne, J. A. & Twipace, N, (1977) Shore 

platforms and sealevel changes in the Gulfs 

region of South Australia, Trans. RK, See. 8. 

Aust. 101, 63-74. 

& Minnes, A. BR. (1983) Aspects of the dis- 
tribution and disintegration of siliceous duri- 
sates in arid Australia. Geol, Mijn. 62, 373- 
383, 

Wriour, R. 1. (1963) Deep weathering and ero- 
sion surfaces in the Daly River Basin, Northern 
Territory, J. geal. Soe. Aust. 10, 151-164, 


Voyave dans les 


Landforms, 


A REDESCRIPTION OF THE GENUS TASMANOCOENIS LESTAGE 
(EPHEMEROPTERA: CAENIDAE) FROM AUSTRALIA 


BY PHILLIP J. SUTER 


Summary 


The genus Tasmanocoenis Lestage is redescribed and nymphs directly associated with adults of T. 
tonnoiri, the type species, are described. Nymphal characterization of Tasmanocoenis is given. The 
recently established genus Pseudocaenis Soldan is shown to be congeric with Tasmanocoenis and is 


suppressed. 


A REDESCRIPTION OF THE GENUS TASMANOCOLNIS LESTAGE 
(EPHEMEROPTERA;: CAENIDAE) FROM AUSTRALIA 


by Putte J. Surer* 


Sumimury 


Sutre PL J, 


(1984) A redeseriplion of the genus Tusaiecoenis Lestage (Ephemeroptera: 


Coenidve) from Australia, Trans, RL Soe, 8. Aust, 108(2), LOS5-111, 12 June, 1984, 
The genus Pasatoeoents Lestage is redescribed and nymphs directly associnted with adults 
of PF romoiri, the type species, are deseribed, Nymphal characterization of Tasmanocoenis 1s 
given. The recently established genus MMsewdecaenis Soldin is shown lo be congeneric with 


Tiusmanevoeniy and is suppressed, 


Key Worbs! Tasmuanoceensis, Ephemevoptera, Cacnidue, Msevdacaents, nymphal charac 


terization, 


luiraduction 

Prior to 1978 the Cacnidac was considered 
to be represented in Australia by only one 
genus, — Tavatiacoens Lestuge (1930) 
(Demoulin 1955; “Chew 1960; Williams 1968; 
Riek M970. Suter 1979; auch Williams 1980). 
However, Soldin (1978) established a new 
genus, Preudocaeniy based only on hymphal 
material From one locality in Queensland. and 
one in New South Wales. The description of 
this Hew genus was based On & comparison 
with deseriptions given by Harker (1950, 
1957) of nymphs of Tayirenoeoents, 

Inv recent shidy of South Australian may- 
flies, Souler! ceared imagos from nymphs 
referrable to Prendocaenix. The udults were 
typival Taynanocoeniy. These observations sug- 
ested that the genera were synonymous. 
Examination of mptertal of all) deseribed 
species of Tusmanoceveniy Cwith the exeeption 
of Vo fillenyi Warker) and Pseudacarnis 
demonstfated that the generic diagnosis given 
for Prudtncaeniy (Soldin, 1978) — clearly 
characterized Tasmanacnenis. Therefore 
Prerdocanis is suppressed and beeomes a 
synonym of Taaierocoenty, 


Materials and Methods 


Nymphs and adults were associated in the 
Juaboratory, and the reared udults were pre- 
served with their individual cust skins. Dis- 
yected appendages were mounted in polyvinyl 
lacto-phenol mounting medium, 


and 


“Stare Water Laboratories, — Hngineering 
Hag, 


Water Supply Department, Private Mail 
Salisbury Post Offiee, S. Aust. S108, 


Suter, PJ. (1980) ‘The taxonomy and ecology 
ot the Ephemeroptera. (Mayllies) of South Aus 
tralia. PhD, thesis, Dept, OF Zoolopy, University 
of Adelaide. Unpubl. 


All measurements ave given i mm, Each 
segment of the fare. middle and hind legs of 
the nymph is compared to the length of the 
femur. as a rao, The absolute length of the 
femur is given in parentheses, Comparative 
measiirements of the scyments of the tabial 
and maailary palpi are also expressed as 
ratios, compared with the proximal segment 
length, given in parentheses, 

fo figures of the labium, the methad of 
Peters & Edmumils (1964, 1970, 1972) is fol- 
lowed, wilh the ventral surface shown on the 
left. 

Material examined was made available from 
linstitut Royal Des Seiences Naturelles de 
Belgique (I.R.SeN,B.) and the National 
Museunt of Vietoria (NMYV), 


Genus TASMANOCOENITS Lestage 
Tasinuneoents Lestage, 1930, p, 53, Type species 
Tasmanococnis waned! Lesiage, original designa- 

' 
oe (930- 53-54; Tillyard, 1936: 56 (Part in 
Cavnivis Harker, 1950: 24-26, 29 (referred 10 
as Cuenis): 1954 266 (referred to us Tasmano- 
cdeniy sie, path ty Cauenis)s Demoulin, (a5: 1-7: 
Harker, 1957" 76; vin Brugeen, L957: 32-33: 
Thew, L960; 202: Riek, 1970; 238; Puthz, LOTAs 
412, Soldan, 1978s 124-128; Huler, LV79> &2, 

The genus Tasmanecoeniy was established 
by Lestage (1930) when 7. fenneiri was 
vescribed fram a single adult specimen col- 
lected at Geeveston, Tasmania by Tonnoir in 
1923, The deseription was nol Hustrated and 
no nymphs of this species were ceeorded, Th 
1936, Tillyard deseribed u further caenid 
species from Tasmania (Caenis seertiy but 
Lestage (1938) noted that the specific nime 
was pre-oveupied, and renamed this species 
Coeris tillvardi (sic), 

Harker (1950), wpparently unaware of 
Leslige’s paper. maintained Caenis scot Tor 
the speeies described by Tillyard, and 


106 


described a nymph and female imago which 
she assigned to this species. These nymphs 
were not directly associated with a male 
imago, and there remains some doubt about 
the identification of the nymph described. 


Demoulin (1955) reviewed the Australian 
Brachycercidae (Caenidae) and re-described, 
with illustrations, Tasmanocoenis tonnoiri, the 
type species of the genus. He recognised that 
Caenis tillyardi belonged to the same genus 
as T. tonnoiri. Demoulin also presented a key 
to the genera of adult and nymphal caenids. 
He based the generic separation of adults on 
the length and width of the prosternum, length 
of the antennal pedicel, and length of leg seg- 
ments, especially the comparative lengths of 
the fore tibiae and tarsi, The nymphs of Tas- 
manocoenis were distinguished from other 
genera by having a three or four segmented 
first gill. This, and the following characters 
were mentioned by Demoulin as interpreted 
from Harker’s (1950) description and illustra- 
tions: gill II not joining at the mid dorsal line 
of the abdomen, lamellate gills II-VI fringed 
with tracheal filaments generally simple, rarely 
bifid; anterior margin of labrum with a median 
concavity, and denticles on each side of the 
concavity; second segment of maxillary palp 
almost as long as third segment; glossae and 
paraglossae of labium widely separated. The 
nymphal characteristics were therefore estab- 
lished not from actual specimens, but from 
Harker’s illustrations and descriptions of T. 
tillyardi. 

Thew (1960) revised the genera of the 
Caenidae and listed the following nymphal 
characteristics as distinguishing Tasmanocoenis 
from other genera: mandibles lacking mar- 
ginal fringes on both sides; maxillae with only 
a few hairs, no thick spines; labrum lacking 
marginal fringe of hair; gill covers without 
triangular ridge and without marginal fringe 
of hair; first gill three or four segmented and 
lamellate gills with fringe of single or bifid 
tracheal filaments. Soldan (1978) also noted 
these characters, all of which are consistent 
with Harker’s (1950) illustrations. 

Suter! questioned the status of Pseudocaenis 
when adults of Tasmanocoenis tillyardi were 
reared from nymphs which displayed charac- 
teristics used by Soldan (1978) to define 
Pseudocaenis. To establish the status of 
Pseudocaenis it was necessary to examine the 
type species of each genus in either adult or 
nymphal stage. Nymphs were collected and 


P. J. SUTER 


reared from the type locality of T. tonnoiri 
(Geeveston, Tas.) but all were T. tillyardi. 
However, material from the La Trobe River 
and the Tyers River, Vic., enabled association 
of nymphs and adults of T. ronnoiri. Examina- 
tion of this material, the holotype of T. ton- 
noiri, and paratypes of P. queenslandica and 
P. rieki Soldan, show that the generic criteria 
of Demoulin (1955), Thew (1960), and 
Soldan (1978) for Tasmanocoenis are erro- 
neous, and that the characterization given for 
Pseudocaenis (Soldan, 1978) is actually that 
of Tasmanocoenis. The genus Pseudocaenis 
thus becomes a synonym of Tasmanocoenis. 
Tasmanocoenis is redefined based on examina- 
tion of the holotype, and nymphs associated 
with male imagos of the type species, and of 
associated adult and nymphal material of T. 
tillyardi. 

Imago Characteristics 

Male: body length 3.1-4.2 mm, forewing 
length 2.9—4.0 mm. 

Female: body length 5.0-6.5 mm, forewing 
length 4.5-5.2 mm, Head: eyes separate, lateral. 
Dorso-lateral ocelli raised, black; median 
ocellus small, black, Antennal pedicel twice 
length of scape. Thorax: robust, dark black/ 
brown. Pronotum narrower than head. Pros- 
ternum triangular, apex truncate, lateral mar- 
gins separated, slightly longer than broad, 
Mesonotum strongly humped, slightly broader 
than head, median notal suture divided just 
anterior to wings to form pale area. Legs: 
pale grey, slender and delicate; forelegs longer 
than middle and hind legs. Fore, middle and 
hind legs of male with five tarsal segments; 
female with all tarsi four segmented. Tarsal 
claws of male imago similar in foreleg, both 
blunt and club shaped, dissimilar in middle 
and hind legs, one blunt, club shaped, one 
slender, curved and sharp. Female with each 
pair of tarsal claws dissimilar: one blunt, one 
curved and sharp. 

Wings: forewing length 1.7—2.0 * width; 
hyaline with milky-opaque pterostigma. Vena- 
tion reduced, almost lacking cross veins, pos- 
terior margins lined with very fine setae. 


Mature Nymph Characteristics 

Head smooth, lacking protuberances. Pedi- 
cel of antenna 2-3 length of scape, covered 
with long setae. Tentorial body rectangular, 
length 0.75 X* width. Gills, six pairs on abdo- 
minal segments 1-6: first abdominal gill fila- 
mentous, two-segmented with long setae; 


REDESCRIPTION OF TASMANOQCOENIS 17 


second gill operculate with triangular dorsal 
ridge, mesal fork with bifid setae, posterior 
ridge not extending to posterior margin of gill 
cover, margin lined with setae; gills 3-6 
triangular, pigmented belween trachea (pig- 
ment may be lost after long preservation), 
tracheal filaments single to multifid, and with 
longitudinal band of short bifid bristles on 
dorsal surface. Seeond abdominal segment 
with a small blunt dorsal median spine, Abdo- 
minal segments 3—9 with postero-lateral pro- 


jections. 

Mouthparts: labrum rectangular, 2-3 
broader than long, anterior margin with 
shallow median concavity, lateral and 


anterior margins with spine setae. Mandibles 
stout, With marginal setae, outer incisors with 
3-4 teeth, inner with 2-3. Glossae of hypo- 
pharynx oot produced, anterior margin: con- 
cave, paragnaths lined with setae. Muxillac 
slender, with wroup of teeth at apices, palpi 
three-segmented. Labium with 3-segmented 
palpi. Lee murgins lined with spine setae, 
femur of fore leg with transverse row of setac 
on ouler Jateral edge, tarsal claws short, 
curved with blunt teeth near base, and smaller 
distal teeth. Male and female nymphs similar, 
hut females more robust. 

Tusmenocoenis closely resembles. Cacnis in 
both nymphal and muginal characters, but the 
following combination of characters distim- 
guishes Tasmanoeeeniy from all other genera 
in the Cuenidae, tr the nymph: (1) pedicel 
of antenna with setae, (ii) mesal fork of gill 
cover with bilid setue, (ii) posterior ridge 
not eatending to posterior margin of gill 
cover, (iv) tarsal claws with blunt teeth near 
biuse and smaller distal denticles, (v) sub- 
murninal row of seales on pill cover. Male 
imago: (i) forceps strongly bowed (ii) lobes 
of penes fused, with apieal indentation, 


Tasnuntocoenis tonnot Lestage 
PIGS. 1-13 
Leslage, 1930; 53-54; ‘Villyard, 1936: 56: Harker, 

1954! 266; Demoulin, 1955; 2 3: Thew, 1960; 

202, 

This Species wus described by Lestage 
(1930), from a dry specimen glued to a card; 
no ilusteationy were given, Demoulin (1955) 
redeseribed the holotype, aml mounted the 
genitalia, [ees and wings onto slides. and 
placed the body in spirits. In the present stucly 
the holotype was examined, and although 
Demoulin’s description Qvith the exception of 
the penitalia) is adequate, Iresh material has 


cere St | ? 


Vigs. 1-2. Tashwinocoenis tonnajry > 1, Genjlatia 
of holotype male, showing the distorted penes 
and forceps; 2, Genilalia of a male imago from 


the La Trobe River. Vic. The small busal 
tubercles are indicated by the arrows, Scale 
lines = OF mm, 


heen used Lo add to this description, Thorax: 
fees slender, fore femur equal in length to 
middle femur, but shorter than hind femur, 
Ratios of lez segments: fore ley 1.00 ; 1.85 5 
O13 : 0.61 2: 0.28 : 0.32 2 0.19 (0.54 mm), 
middle leg 1.00 : 0.68 : O11 : 0.09 : 0.08 : 
0.04 : 0.23 (0.53 mm); hind leg 1.00 : 0.72 : 
0.12 : O09 : 0,07 : 0.05 + 0.21 (0.58 mm), 
Genitalia: the genitalia of the holotype, now 
on a slide, are badly distorted, in parts torn, 
with the penes folded back within themselves, 
giving a broad curved posterior murgin as 
illustrated by Demoulin. and in Fig. 1, Geni- 
talia from specimens from the La Trobe 
River, Victoria, were dissected, and the actual 
shape of the penes is shown fo Fig. 2 

The forceps of the holotype are also twisted 
and the resultant structures cannat he viewed 


108 P. J. SUTER 


Figs. 3-7. Tasmanocoenis tonnoiri mature nymph: 3, Basal part of antenna, scape, pedicel, and basal 
segment of flagellum; 4, Fore leg; 5, First abdominal gill; 6, Second abdominal gill; 6a, Enlarge- 
ment of the bifid setae of mesal fork of the dorsal triangular ridge; 6b, Enlargement of the sub- 
marginal scales; 7, Third abdominal gill. Scale lines; Figs 3, 4, 5 and 7 = 0.1 mm, Figs 6, 6a 
and 6b = 0.05 mm. 


REDESCRIPTION OF TASMANOCOENIS 


NN 
\ 


\\\ 
aK 


he a ‘ z= 
Figs 8-13. | 


Yasmanocoenis tonneiri mature nymph: 8, Dorsal view of labrum with the enlarged antero- 

median emargination; 9, Left mandible. ventral view; 9a, Left incisors and prostheca enlarged; 

10, Right mandible, ventral view; 10a, Right incisors and prostheca, enlarged; 11, Hypopharynx: 

12, Right mansilla, ventral view; 13, Labrum, dorsal (left) and ventral (right) view. Scale lines 
0.1 mm. 


é: 


109 


110 P. J. SUTER 


in ventral orientation. However, along the 
mesal margin of the forceps are 3 small 
tubercles. These are also present on the forceps 
of the holotype, but appear as ventral tubercles 
(Fig. 1). 

The narrow, strongly bowed forceps with 3 
mesal tubercles and the shape of the penes are 
diagnostic characteristics of T. tonnoiri. 


Mature Male Nymph 


The following description is based on one 
individual, but the range of variation observed 
in the examined material is given in paren- 
theses. 

Head width 0.70 mm (0.70—-0.94 mm); body 
length 2.62 mm (2.62—4.15 mm); cerci length 
1.99 mm (1.99-—2.35 mm); terminal filament 
2.64 mm (2.64—-2.88 mm). 

General body colour brown. 

Head: brown with darker regions between 
eyes, lateral margins smoothly convex. Eyes 
black, ocelli brown. Antennae yellow-brown, 
pedicel 2.60 X length of scape (Fig. 3), fla- 
gellum 1 mm long. 

Thorax: pronotum brown, antero-lateral 
margins with few spine setae, width equals 
head width. Mesonotum dark brown, width 
1.5 X head width. Legs light brown; femora 
without dark markings (Fig. 4). 

Ratios of leg segments: fore leg 1.00 : 0.72 
: 0.56 (0.57 mm); middle leg 1.00 : 0.71 
0.50 (0.56 mm); hind leg 1.00 : 0.78 : 0.49 
(0.62 mm). Femur length to width ratios; fore 
leg 2.41 (2.30-2.63), middle leg 2.48 (2.33— 
2.63), hind leg 2.63 (2.52-2.75). 

Abdomen: brown. Operculate gills extend- 
ing over segments 3-7. Cerci and terminal 
filament dark brown. Gills; first gill (Fig. 5) 
apical segment length 3.7 X basal segment 
length, lined with setae. Second gill operculate, 
length 1.16 X width (Fig. 6), mesal fork 
of triangular ridge with 8 bifid setae (range 
7-10) (Fig. 6a); submarginal row of scales 
each with 8-10 bristles present (Fig. 6b). Gills 
3-6 triangular with multifid tracheal branches 
(Fig. 7). 

Mouthparts: labrum (Fig. 8) 2.03 X 
broader than long. Left mandible (Fig. 9) 
outer incisors with 4 teeth with short setae on 
ventral tooth (Fig. 9a), inner incisors with 3 
apical teeth with short setae on ventral tooth, 
prostheca robust, bifid with bifid or trifid 
setae apically. Right mandible (Fig. 10), 
outer incisors with 3 apical teeth, inner incisors 
with 2 teeth (Fig. 10a), prostheca simple 
apically with bifid and trifid setae; margin 


between prostheca and molar region with large 
tubercle. Hypopharynx (Fig. 11). Maxillae 
(Fig. 12), galeo-lacinia with 4 apical teeth, 
palpi longer than galeo-lacinia, segment ratios 
1.00 : 0.70 : 1.00 (0.14 mm), 

Labium (Fig. 13) proximal segment of 
palpi 1.64 X longer than broad; second seg- 
ment convex; distal segment short, triangular; 
segment ratios 1.00 : 0.84 : 0.49 (0.13 mm); 
glossae rectangular, rounded apically, para- 
glossae curved. 

Material examined: holotype male, Geeves- 
ton, Tas., 7 Dec. 1922, A. L. Tonnoir, in 
I.R.Sc.N.B, 

La Trobe River, Rosedale, Vic. 10.ix.1980, 
R. H. Norris and P. Mitchell (nymphs and 
adults) in NMV; 27.11.1974, J. Blyth 
(nymphs) in NMV; Tyers River, west of 
Tyers, Vic., 24.11.1974, J. Blyth (nymphs) in 
NMV. 


Discussion 


With this redefinition of the genus Tasmano- 
coenis the following species are now recog- 
nised in Australia; T. tonnoiri Lestage, T. till- 
yardi (Lestage), T. jillongi Harker, T. queens- 
landica (Soldan), and T. rieki (Soldin). The 
latter two species were described in the 
nymphal form only, but on examination of 
paratype material presented to the National 
Museum of Victoria, the two nymphal para- 
types are indistinguishable. The characters 
listed by Soldén (1978) to distinguish the two 
species do not separate the paratypes. How- 
ever, on the limited material available, and 
in the absence of reared adults, a synonymy 
of T. queenslandica and T. rieki would be 
premature, but some doubt must remain as 
to the validity of these species. 


Acknowledgments 
I would like to thank Dr G. Demoulin of 
Vinstitut Royal Des Sciences Naturelles de 
Belgique for making available the holotype of 
Tasmanocoenis tonnoiri, and Dr T. Sold&n 
for depositing paratype material of Pseudo- 
caenis queenslandica and P, rieki in the 
National Museum of Victoria. I would also 
like to thank Dr R. Norris and Messrs J. 
Blyth, P. Mitchell and L. Metzeling for col- 
lecting the material from the La Trobe River 
and for assistance in rearing the adults of 
T. tonnoiri. I am also grateful to Dr A. 
Neboiss, Dr A. Calder and Dr D. Towns for 
their critical discussions and suggestions in the 

preparation of this manuscript. 


REDESCRIPTION OF TASMANOCOENIS WW 


References 


Demoucin, G. (1955) Les Brachycercidae Aus- 
traliens. Le genre Tusmanocoeniy Lestage. Bull. 
Mus. R, Hist, Nat. Bele. 31, 1-7, 

Harker, J. E, (1950) Australian Ephemeroptera. 
Part 1. Taxonomy of New South Wales species 
and evaluation of taxonomic features. Proc. 
Linn, Soc. N.S.W. 75, 1-34, 

(1954) The Ephemeroplera of Australia, 
Trans, R. Entomal. Soc. London. 105, 241-268. 

—— (1957) Some new Australian Ephemeroptera, 


Proc. R. Entomol, Soc. Landon. Ser. B. 26, 
63-78. 
Lesrace, J. A. (1930) Notes sur le premier 


Brachycercidien decouvert dans la fuune aus- 

tralienne Tasmanocoenis fonnoiri sp. nov. 

(Ephemeroptera) et remarques sur la famille 

des Brachycercidae Lest. Mem. Soc. R. Belge 

Entamal. 23, 49-60. 

(1938) Contribution a Vétude des Ephéme- 
ropteres XX. Notes synonymique: Coenis scotti 
Till. (1935) nec Ulmer (1930) Coenis 
tillyardi nom. noy. Bull. Ann, Soc. R. Belge 
Entomol, 78, 320, 

Peters, W. L. & EpMunps, G. F, Jr. (1964) A 
revision of the generic classification of the 
Ethiopian Leptophlebiidae (Ephemeroptera). 
Trans, R, Entomol. Soe. London, 116, 225-253. 

& (1970) Revision of the generic 

classification of the eastern hemisphere Lepto- 

phlebiidae (Ephemeroptera). Pac. Insects 12, 

157-240, 


- (1972) A revision of the generic 
classification of certain Leptophelbiidae from 
South America (Ephemeroptera). Ann, Ento- 
nal, Soe. Am. 65. 1398-1414. 


Puruz, V. (1975) Eine neue Caenidengattung aus 
dem Amazonasgebiet (Insecta: Ephemeroptera: 
Caenidac). Amazoniang 3, 411-415. 

Riek, E. F. (1970) Ephemeroptera (Mayflies). 
In The Insects of Australia, sponsor C.S.LR.O, 
anon of Melbourne Press: Melbourne). 

SOLDAN, (1978) New genera and species of 
aa anc (Ephemeropicra) from Iran, India 
and Australia. Acta Entomol. Bohemosloy. 75, 
119-129, 

Suter, P. J. (1979) A revised key to the Aus- 
tralian genera of mature mayfly (Ephemerop- 
tera) nymphs. Trans. R, Soc. S. Aust. 103, 
79-83. 

Tuew, T. B. (1960) Revision of the genera of the 
Family Caenidae. (Ephemeroptera). Trav. 
Am. Entomol. Sac. (Philadelphia) 86, 187-205. 


Tittyarp, R. J, (1936) The trout food insects of 
Tasmania. Part 17. A monograph of the mayflies 
of Tasmania. Pap. Proc. R. See. Tasmania 
1935, 23-59. 


VAN Bruacen, A. C. (1957) On two new species 
of mayflies from the Wissel Lakes Central New 
Guinea (Ephemeroptera), Neva Guinea ns. 8, 
31-39. 


WictramMs, W. D. (1968) Australian Freshwater 
Life: The Invertebrates of Australian Inland 
Waters. (Sun Books: Melbourne). 


(1980) Australian Freshwater Life: The 
Invertebrates of Australian Inland Waters. 2nd 
Edition. (The Macmillan Co, of Australia Pty. 
Ltd.; Melbourne). 


A NEW SPECIES OF GEHYRA (REPTILIA: GEKKONIDAE) FROM 
NORTHERN WESTERN AUSTRALIA 


BY MAX KING 


Summary 


Specimens of an undescribed form of Gehyra are compared with populations of G. australis found 
in adjacent areas of the Kimberley in Western Australia and are described as a new species of the G. 
australis species group. 


A NEW SPECIES OF GEHYRA (REPTILIA: GEKKONIDAE) FROM 
NORTHERN WESTERN AUSTRALIA 


by Max KiING* 


Summary 


kinc, M. (1984) A new species of Gelivra (Reptilia: Gekkonidae) from northern Weslern 
Australia, Trans, R, Soe, 8, Aust, 108(2), 113-117, 12 June, 1984. 

Specimens of an undescribed form of Gehyra are compared with populations of G, 

australis found in adjacent areas of the Kimberley in Western Australia and are deseribed as 


anew species of the G, australis species group. 


Kry Woros: New species, Gelyra, Gekkonidac, Kimberley. 


Introduction 

A karyotypic analysis of population of the 
widely distributed, and morphologically diverse 
Australian gekko Gehyra australis, revealed 
considerable chromosomal helerogeneily, 
Seve chromosome races occur in northern 
Australia; 2n — 44, 2n «= 42A, 2n -= 428, 
2n 420, 2n 40A, 2n —- 40B and 2n — 
38 (King 1982, King 1983a). Each chromo- 
some race is allopatrically distributed, cither 
geographically or because of habitat pre- 
ferences. Vhese isolated forms are chramoso- 
mally monomorphic for a series of fixed dif- 
ferences. In areas of possible contact between 
chromosome riuces, there is no evidence of 
hybridization, 

A subsequent morphometric analysis of the 
2n — 427A, 2n = 42B, 2n = 42C, 2n ANA 
and 2n 38% chromosame races (King 1982, 
King, 19836) has greatly modified our concept 
of Gehyra australls. This species was redefined 
and its new distribution was shown to approaxi- 
mate that of the 2n — 40A chromosome race 
(King 1983b). It is therefore now restricted 
lo the northern sector of the Northern Terri- 
tory. and a small area of northern Western 
Australia, G. dubia Macleay was resurrected 
to accommodate the 2n 42C chromosome 
ruee and the following new species were 
described: G, parnela (2n 472A): G. robusta 
(2n 42B) and G. harrelvela (20 38) 
(King 1982, 1983b). Too few specimens of 
the 2n —— 40B and 2n — 44 races were avail- 
able fora taxonomic reappraisal of these forms 
to be made, 

‘The present paper deseribes the results uf oa 
morphometric analysis of specimens of the 2n 

44 chromosome race of Gehyra, and of 


* Department of Population Biology, Research 
School of Biological Sciences, The Australian 
National University, P\O, Box 475, Canberra 
City, A.C.T. 2601. 


populutions of G. anstraliy sensu sifieto, from 
nortbern Western Australia. A new species 1s 
described, 


Materials and Methods 

Three specimens karyotyped by King 
({983a) were measured, as were an additional 
seven museum specimens which were mor- 
phologically identifiable as belonging to the 2n 

44 chromosome race. These animals were 
compared with 27 individuals of G. australis 
from adjacent areas of the Kimberley. The 
distribution of the animals examined js shown 
in Fig. 1, 

All specimens were measured with micro- 
meter-adjusted calipers and a steel rule. 
Dimensions tuken in this study are those of 
King (1983b), 


Results 
The specimens analysed fell into two unam- 
biguously distinguishable morphological 
groups. One of these groups comprised typical 
G. australis, although certain minor variations 


122-F [24 130 
. | 
LR cg | 
Vs 
| 7 
| 1a" ° ’ 
ad — ‘ 
* e ef “ 
“ L x 
i «° 
pws 
ca.) e 
‘ae 
\ 
ako 
18° 


4 


Fix, 1, Distribution of Gehyra aceidentaliy (die 
monds) and G, anstralis Chlack spots )- 


114 MAX KING 


(see later), were detected. The second group 
of ten specimens represented an undescribed 
form, some of which had been karyotyped by 
King (1983a) and shown to have 2n = 44, 
This form is described here. 


Gehyra occidentalis sp. nov. 
FIGS 1-5 


Gehyra australis part: King 1983a p. 723. 
Gehyra australis part.: King 1983b in press. 


Diagnosis: Gehyra occidentalis is distinguished 
from other membets of the G. australis species 
group by the following combination of 
characteristics. It differs from G. baliola in 
lacking a ‘U’ shaped rostral scale, and by the 
absence of skin folds on the back of each hind 
limb. It is distinguished from G. xenopus by 
the absence of basal granules dividing the 
fourth toe subdigital lamellae. G. occidentalis 
is distinguished from G. australis, G. robusta 
and G. dubia by having longer postmental 
scales. Males of these species and of G. borro- 
loola have fewer than 19 preanal pores, 
whereas, G. occidentalis has 23-49 pores, 
G. occidentalis is most similar to G. pamela 
from which it is distinguished by the following 
characteristics: the rostral scale is deep and 
its dorsal surface strongly gabled, in G. occi- 
dentalis, whereas it is oblong and_ slightly 
gabled in G, pamela; when viewed from below 
the rostral scale projects forward of the snout 
line in G. pamela, but not in G. occidentalis; 
the background colouration of the back pat- 
tern is chocolate brown in G. occidentalis and 
grey in G, pamela. In those specimens of G. 
occidentalis with pronounced patterning, bands 
of black spots predominate in size over the 
interbands of lighter spots. In G. pamela the 
bands of off-white spots form the predominant 
coloured bands, the darker spots being reduced 
In size. 


Fig. 2, Diagrammatic representation of the snout 
of G. occidentalis showing the steeply gabled 
rostral scale, and two small internasals lying 
between the larger internasals. 


Description: 

Holotype: Male W.A.M. R83711. Collected 
on a rock face at night in Manning Gorge, 
W.A. (16°44'S, 125°57’E) by D. R. King on 
3.xii. 1980. 

Head: Width 11.0 mm, depth 6.1 mm, length 
14.0 mm. Snout 6.1 mm long from tip of 
rostral scale to anterior margin of orbit. Face 
and head covered by small rounded scales, 
those on face larger than on top of head. 33 
interorbital scales. Nostril surrounded by 
rostral, internasal, two posterior nasal and first 
supralabial scales. Rostral scale oblong and 
relatively deep. Dorsal surface of rostral 
steeply gabled to midline apex (Fig. 2). 
Median groove on rostral extending for 1/3 
of scale depth from middle of dorsal surface. 
Nostrils separated by two large internasal 
scales. Two very small internasal scales located 
at apex of rostral separating large internasal 
scales (Fig. 2). Nine supralabial and eight 
infralabial scales on each side of jaw. Mental 
scale triangular. Postmental scales long (3.3 
mm) and not in contact with second infralabial 
scale (Fig. 4b). 

Body: Depressed, slender build (Fig. 3). Snout 
vent length 59.4 mm, tail length 65.00 mm. 
Tail round in section tapering into a point. 
Larger scales on ventral surface. Dorsal sur- 
face of body covered by small rounded scales. 
Scales on ventral surface larger and flatter 
than those on dorsal surface. 126 scales around 
circumference of abdomen in midbody. Nine 
subdigital lamellae on dilated section of 
fourth toe. Subdigital lamellae divided along 
midline (Fig. 4c). 29 preanal pores in chevron 
formation in front of cloaca (Fig. 4d). Two 
postnatal tubercles in cluster at base of tail on 
each side, 

Colouration: Background dorsal colouration 
chocolate brown in life. Head and faces with 
alternate very dark brown and off-white spots, 
separated by background colour, Two parallel 
eyestripes extending from snout and finishing 
above ear. Back pattern consisting of bands 
of black spots (which have coalesced to form 


Fig. 3. Holotype of G. occidentalis in life. Bar 
scale = 10 mm, 


NEW SPECIES OF GEHYRA 115 


bars) interspaced by bands of off-white spots. 
Bands of spots separated by chocolate brown 
background. Alternate coloured bands like 
those on the dorsal surface extending along 
length of tail. Limbs spotted with black and 
off white (Fig. 3). 

Paratypes; There are 9 paratypes: W.A.M. 
R83712 14°53’S, 125°45’E. W.A. 27.vii.82, col- 
lected by J. Dell, W.A.M R83713 32 km E of 
turnoff to Napier Downs on Gibb River road, 
W.A, 27.viii.80 collected by D. King, W.A.M. 
R45009 Napier Range 170°18’S, 124°S0’E W.A. 
1.xi.73 collected by W. H. Butler, W.A.M. 
R70587 11.5 km S.E. of Mt Percy, W.A. 17.v.80 
collected by G. Harold, P. Griffin and G. Barron, 
W.A.M. R70553 8.6 km S.E. of Mt Amy (Napier 
Downs) W.A, 18.v.80 collected by G. Harold, P. 
Griflin and G. Barron, W.A.M. R70664 8.6 km 
S.E. of Mt Amy (Napier Downs) W.A. 18.v.80 
collected by G. Harold, P. Griffin and G. Barron, 
W.A.M. R58757-9 Wombarella Creek, Napier 
Range, W.A. 9.xi.76 collected by R. E, Johnstone. 
Distribution: The known distribution of G, 
occidentalis is restricted to the western section 
of the Kimberley division of W.A. Populations 
extend from the Mitchell Plateau in the north 
to the Napier Range in the south, G. occiden- 
talis is an exclusively rock dwelling form, 
most specimens having been collected on rock 
faces at night. 

Variation: The range of variation in a series 
of morphometric and meristic characteristics of 


j-. 


1. 
pire 
ot ae 
AO 


Fe hg 


‘ tae 
Coat # Magar’ 

, Pare os 

i J 


G. occidentalis and G. australis from the 
Kimberley, are shown in Table 1. Specimens 
of G. occidentalis always have longer post- 
mental scales than those of G. australis of 
comparable snout-vent length (Fig. 5). Males 
of G. occidentalis are also readily distinguished 
from G. australis by the higher number of 
preanal pores (23-49 compared to 11-19). 
Moreover, G. occidentalis has fewer fourth toe 
subdigital lamellae (9-10) than G. australis 
(10-12). The lamellae are completely divided 
in G. occidentalis but only depressed in the 
midline in G. australis (Fig. 4c). 


The 27 specimens of G. australis examined 
here show certain differences from the 49 N.T. 
animals analysed by King (1983b). Kimberley 
specimens tend to be larger (x S.V.L. 68.7 mm 
compared with x 60.6 mm) although this may 
be due to a sampling bias. Specimens from 
the Kimberley are slimmer in appearance and 
often have eyestripes, a feature absent from the 
N.T. populations. They also have more preanal 
pores in males (K = 16.5 compared to X = 
13.5), and a larger number of subdigital 
lamellae on the dilated section of the fourth 
toe (K = 11.4, range 10-12 compared with 
xX = 10, range 9-12). G. australis were found 
on human habitation or on trees in the N.T. 
whereas those in the Kimberley were also 
found on rock outcrops. These differences may 


Fig. 4a. Side view of the head of the holotype of Gehyra occidentalis. 
b. A ventral view of the snout of the holotype showing large postmental scales. 
c. Ventral view of the foot of the holotype showing divided subdigital lamellae. 
d. Ventral view of the holotype with 29 preanal pores indicated. 
e. Ventral view of a specimen of G. occidentalis with 49 preanal pores indicated. 


116 MAX KING 


TasBLe 1. Morphometric and meristic characteristics of G. occidentalis and G. australis. Means with ranges in 


parentheses. 
N_ snout vent taillength snoutlength postmental head width head depth head length 
length 
mm mm mm mm mm m mm 
SS eS” TO Cn 
G. occidentalis 10 58.9 — 6.8 3.5 12,2 7.0 15.2 
(48-67.3) (up to64) 5.6~-7.5) (2.9-3.9) (10.1-14.2) (5.9-8.3)  (12.9-16.5) 
midbody interorbital preanal postanal subdigital supra- infra- 
scales scales pores tubercles lamellae labials labials 
G. occidentalis 112.2 32 30(63) 2.5(629) 9.7 9.2 8.4 
(96-126) (26-36) (23-49) (2-4) (9-10) (8-11) (7-10) 
N_ snout vent taillength snoutlength postmental head width head depth head length 
length 
mm mm mm mm mm mm mm 
G. australis 27 68.7 ee 7.3 3.0 13.0 7.7 16.6 
(58.6-75.0) (upto 84) (5.7-8.1) (2.1-3.5) (11.2-14.6) (6.7-8.7) (14.1-18.0) 
midbody _interorbital preanal postanal subdigital supra- infra- 
scales scales pores tubercles lamellae labials labials 
G. australis 117.6 32.4 16.5(188) 2.71184) 11.4 10.3 8.7 
(102-131) (27-37) (11-19) (1-3) (10-12) (9-12) (7-10) 
4e). A further difference between these species 
is seen in the morphology of the rostral scale 
which is deep with a strongly gabled dorsal 
surface in G. occidentalis, and oblong and only 
slightly gabled in G. pamela. When viewed 
4.0) ; from below, the rostral area projects anteriorly 
< * so n§ to the snout line in G. pamela but not in G. 
E ’ _ occidentalis. In G. occidentalis a series of 
= i weft 0(2), 1(4) or 2(4) small internasal scales 
4 3.0) . eh ates occur between the large internasals, at the apex 
| oe of the rostral (Fig. 2). Sometimes one small 
3 : internasal is seen in G. pamela. 
£20 [ G. occidentalis is also distinguished from 
2 G. pamela by its chocolate brown rather than 
| grey colouration. Some specimens of G. occi- 
| dentalis lack a pronounced back pattern; others 
i a Ye eee an are strongly marked, with bands of black spots 
Snout — vent length [mm] predominating. In G. pamela the dark colours 
Fig, 5. Postmental scale length plotted against are much less pronounced and the bands of 


snout-vent length for specimens of G. occiden- 
talis (triangles) and G. australis (spots). 


simply reflect clinical variation within G. 
australis; they are much less profound than 
the differences which separate G. occidentalis 
from G. australis. 

G, occidentalis is morphologically most 
similar to G. pamela (King 1982). G. pamela 
is found on the Arnhemland escarpment and 
probably throughout Arnhemland in the N.T. 
Both G. occidentalis and G. pamela have long 
postmental scales, and a higher number of 
preanal pores than G, australis (up to 28 in 
G. pamela, up to 49 in G, occidentalis) (Fig. 


light spots predominate (see Fig. 4a, King 
1982). The back pattern of the holotype (Fig. 
3) is similar to that of some specimens of 
G. borroloola, but distinction from that species 
can be readily made by the rostral shape, and 
greater number of preanal pores in males. The 
only other species of Gehyra which has a back 
pattern similar to G. occidentalis is G. cog- 
natus (Rudiger-Borner & Schuttler, 1982). 
This species, based on a_ single specimen 
shares many characteristics with G. pilbara, 
including the presence of 8 subdigital lamellae 
(although they are grooved, not divided), 8 
supralabials, 6 infralabials and a deepset, blunt- 
snouted head. The animal was apparently 


NEW SPECIES OF GEHYRA V7 


captured with specimens of G. pilbara and on 
the basis of the published information is 
probably an unusual G. pilbara. In any case, 
itis not a member of the G. australis species 
complex, 

Lrymology: The specific name G. occidentalis 
is derived from the Latin “occidentalis” mean- 
ing western, and refers to the distribution of 
this. species. 

Other material examined: W.A.M. R70153-4, 
R70156, R70146—-8 28 km S.E. Kununurra, W.A.. 
21.iv. 1980, W.A.M, R60345 35 km S.W. Kunu- 
nurra (Saw Ra.). W.A,, 2.vi.1978. W.A.M. R70688 
3.5 kim N.W, New Lissadell H.S., W.A., 6.v. 1980, 
W.A.M, R44037 Sir Graham Moore Is.. Bona- 
purfte Arch, W.A., 2.vii.1973. W.A.M. R27571-4 
Purry Ck, W-.A,, 20.viil.1965. W.A.M. R42788-90 
Old Lissadell HLS., W.A. 16.%.1971 W.AM. 
RS50776-7 Drysdale R. Nat, Pk. 14°40°S, 127 00° 
EB. 9-12.viii.1975, W-A.M, R50794 Drysdale R. Nat. 
Pk, 14°40'S. 127°00"F,  12.viin.1975.  W.ALM, 
R70451 10.7 km S.W. New Lissadell H.S., W.A,, 
26.iv.1980. W.A.M. R64919 Drysdale R, Nat. 
Park 15°O8'S, 126°55°E, 3.viii.1975, W.A.M. 
R64920 Drysdale R. Nat. Pk. 15°03'S, 126°44° 
B, 18.vili, 1975, W.A.M. RS0807 Drysdale R. Nat. 
Pk, 14°40°S, 127°00°F, 1 4.viii. 975. W.ALM. 
R50869 Drysdale R. Nat. Pk. 15°02'S, 126°49°R, 
18.viil, 1975. WAM. RS50595 Drysdale’ R. Nat. 
Pk. 14°46'S, 127°05°R, 14.viii.1975. W.AM. 
R50960 Drysdale R. Nat. Pk. 15°02'S, 125°49'E, 


14.vii.1975. W.A.M. Ri3582 Kalunibutu, W.A., 
26.vi L980. 


The Gehyra australis species group 


The G. australis species group now com- 
prises G. baliola, G. xenopus. G. berroloola, 
G. qustralis, G. robusta, G. dubia, G. pamela 
and G. vecidentalis. 

Karyotypic analysis has shown that these 
large Northern Australian forms, while being 
in the same lineage as the other Australian 
Geliyra, have evolved as an independent group 
(King 1982, 1983a). Their general morpho- 
logical similarity is accentuated by the fact that 
six of the species (G. australis, G. borroloola, 
G. robusta, G. dubia, G. pamela and G. occti- 
dentalis) were, until recently, all included as 
G. australis. Adult specimens of this species 
complex have a snout-vent length of 50-9] 
mm and at least 9 subdigital lamellae on the 
dilated area of the fourth toe. 


Acknowledgments 


The author is indebted to Dennis King, John 
Dell and Richard How for collecting live 
specimens of G. occidentalis for analysis. I 
thank Glen Storr and Laurie Smith of the 
Western Australian Museum (WAM) for 
providing a series of Gehyra for examination. 


References 


Kinc, M. (1982) Karyotypic evolution in Gehyra 
(Gvukkonidse; Repulia)y. 1. A new species from 
the Alligator Rivers Region in northern Aus- 
lraba. Aust, J. Zool. 30, 93-101. 

—— (1983a). Karyotypic evolution in Ge/ryra 
(Gekkonidae: Reptilia), UL The Gehyra aus- 
tralis complex. (hid. 31, 723-741. 


(1983b). The Gehyra avstraliy species com- 
plex (Sauria: Reptilia), Amphibia-Keprilig in 
press. 

RupiceR-Borner, A. & Scuurrier, B, (1982), 
Notes on the Australian Lizard Genera Ge/ryra, 
Hemidaen|us and Hetcronatia (Geckonidae). 
Mise, Art. in Saurolagy X, 1-17. 


FOSSIL MARSUPIAL REMAINS AT THE SOUTHEASTERN CORNER OF 
LAKE EYRE NORTH, SOUTH AUSTRALIA 


BY J. A. DULHINTY, T. F. FLANNERY & J. A. MAHONEY 


Summary 


Marsupial fossil remains, interpreted as being Pleistocene in age, occur in a restricted deposit lying 
disconformably on Tertiary dolomite along the eastern side of Price Peninsula at the southeast 
corner of Lake Eyre North. Fossil remains include Sthenurus andersoni Marcus, 1962, Diprotodon 
sp. and Macropodinae indet. 


FOSSIL, MARSUPIAL REMAINS AT THK SOUTHEASTERN CORNER 
OF LAKE EYRE NORTH, SOUTH AUSTRALIA 


by LA. DuLnunry, T.F. Prawwery? & J. A. MATIONEY® 


Summary 


Dunwtniy, J. Ay FLANNERY, T. F. & MaAniowey, J. A, (1984) Fossil marsupidl remains 
wl the southeastern corer of Lake Eyre North, Sauth Australia. Trany, R. Sac. 8. Aust 


HO8(2), (19-122, 12 June, 1984. 


Marsupial fossil remains, interpreted as being Pleistocene in age, oceie in it restrieied 
deposit lying disconformahly on Tertiary dolomite along the eastern side of Price Peninsula iat 
the sourheast corner of Lake Eyre North. Fossil remains inelude Srhenurus andersent Marcus, 


1962, Dipretodon sp. and Mactopodinag tet, 


key Woross Muarstipial fossils, Lake Fyre, South Australia, Srheuweny, Dipreveton. 
Macropadinac, aeeutrence, taxonomy, chronology, 


Introduction 


In 1980 J. A. and R. Dulhunty found an 
oecurrenes of fossil marsupial remains at the 
sOulheasier corner of Lake Vyre North, The 
geological sctting of the oewurrence was studied 
by them and hones were exatvinod and iden. 
tified by Tob, Plannery and J. A, Mahoney 
during J98l-1982. 

The purpose of this paper is to record the 
location ahd geological selling of the occur 
renee, und the taxouamy aod ehrotology of 
the fossil remains: 


Location and peologicyl setting 

‘The fossil remains eceur along the eastern 
shore of Price Peninsula which separates Luke 
Clayton from Lake Eyre, south of the Prome- 
Clayton Pstuary near Level Post Bay (Fig. 1A), 
The sediments containing the fossils vary spora- 
dically from lightly compacted sands to semi- 
consolidated uryitlaceous, gypseous and cal- 
carvous sundy mudstone. They crop out as u 
continuous bed, 2-3 m thiek, at and a little 
above shoreline level along the eastern side of 
Price Peninsula (Pig. 18). The bed is referred 
ta itorwially for field purposes uy the “hare 
hed", Th lies disconformuably upon the eraded 
surface of dolomite beds of the Etadunna For- 
mation which crop oul along Ihe western side of 
Lake Clayton and dip goocrally morthwest to- 
wards und beneath the bed af Madigan Gull. 
Section A-B (Fig. 1C) is a tentative illustration 
of a stricture in Price Peninsula indicated by 
preloyinuary field studies. The bone bed is over- 
Iain conformably for the most part, but with 
' Department of Geology & Geophysics. Univer 

sity Sydeey, NLS Wo UNG 


i School of 4oolopy, University ao NSW. Ken- 
singionm, NUS.W, 2033. 


minor disconfarmities, by semi-consolidated 
clays and argillaceods sands up to at least 13 im 
thick which crop out werass the southern shores 
of Madigan Gulf above the Etaduona Forma- 
lion dolomites (Pulhunty 1982, 1983), The 
geolugy of Level Post Bay. Price Peninsula 
and the southern shores of Madigan Gull was 
described first by King (1956), and later by 


Johns (1963) and Williams (1976). ‘To the 


cast of Lake Eyre, the geology of the Tirari 
Desert was described by Stirtow er al. (1961), 
The beds overlying the bone bed are now 
helieved to be Pleistocene age, and the under- 
lying dolomite of the Etadunna Formation is 
revarded as late Tertrary (Wopfner & Twidale 
1967, Wopfner 1974, Callen & Tedfore 1976. 
Callen 1977). 

Bores by King (1956) and the South Aus- 
tralian Department of Mines (Johns 1963) on 
the shore at Level Post Bay revealed the upper 
auirface of Etadutina dolomite at about 4 m 
below the shoreline level of the bay, but sur- 
lee outcrops of dolomite along the western 
and southern shores of Lake Clayton were not 
reeorded, Recent fieldwork by Dulhunty 
hetween the western shore of Lake Clayton 
and Frome Creek (Fig. 1A) found the upper 
surface of dolomite cropping out ut uboul 
4 (0 6 m higher than in Level Post Bay, This 
indicated a faull, warp or erosion between 
Level Post Bay and Lake Clayton (Fiz. IC), 
A Sparker seismic traverse from Madigan Gulf 
into the Frome-Clayton Estuary, by C. V, G. 
Phipps (pers, cow.) during the major filling 
of the lake in 1974, yielded evidence of fauit- 
ing as indicaled in Figure 1A, This, in Tine 
with the Price Peninsula structure, could well 
he part of o lineation running north-north-west 
which postdated the bone bed and ¢ontributed 


120 J. A, DULHUNTY, T. F. FLANNERY & J. A. MAHONEY 


FIG.IA 
L. EYRE NTH. 


pn GULF L Fos 
FRESGOTT 1 


LOC. PLAN 


EIS. EVID. OF FAULT 


‘ SEE FIG.1B 
MADIG % ENLARGEMENT 


RY LAKE CLAYTON 


10 15km 


YOUNGER 
QUATERNARY SEDIMENTS 


CLAYTON 
——> 
ESTUARY 


ETADUNNA 
FORMATION 
(DOLOMITE BEDS) 


m below sea level 
(Aust. Helght Datum) 


Fig. 1. Occurrence of fossil marsupial remains at the southeastern corner of Lake Eyre North. 


to the original position of the shoreline on 
the eastern side of Lake Eyre. 

The bone bed crops out along the western 
shore of Lake Clayton on the eastern side of 
Price Peninsula (Fig. 1B). At its limits of out- 
crop the bed is overlapped by younger sedi- 
ments onto the eroded surface of dolomite. To 
the east of its outcrop the bed has been 
removed by erosion during excavation of 
Lake Clayton. No bone bed has been found 
above dolomite surfaces which crop out along 
the southern shores of Lake Clayton. To the 
west of its outcrop the extent of the bone bed 
is obscured by younger sediments and the 
westerly dip takes the dolomite surface and 
overlying horizon of the bone bed below the 
shoreline of Lake Eyre (Fig. 1C). The dolo- 


mite emerges again 20 km further west, 
between Shelly and Bluebird Islands (Fig. 


1A), but no fossil marsupial remains have yet 
been found immediately above its outcrop, In 
1982 R. A. Callen found a large vertebrate 
bone (Callen, pers. comm.) in a section above 


the surface of the lake bed at Prescott Point, 
some 8 km west of the bone bed outcrop in 
Lake Clayton. This bone may have occurred 
on a horizon above that of the bone bed in 
Lake Clayton. 

From limited field evidence available it 
would seem that the bone bed was very 
restricted in original area of occurrence, and 
that it probably terminated by overlap beneath 
Price Peninsula 1-2 km west of its present out- 
crop, as shown with query in Figure 1C. 


Taxonomy and chronology of fossil remains 

The remains of three marsupial taxa have 
been identified among a small collection of 
fossil bones and teeth from the bone bed (AM 
F65475-9). These are Sthenurus andersoni 
Marcus, 1962, Diprotodon sp. and Macropo- 
dinae indet. The specimens are in the Aus- 
tralian Museum, Sydney. 

Sthenurus andersoni is represented by a 
broken mandible with teeth in an advanced 
stage of wear (Fig. 2) (AM F65476). S. 


FOSSIL MARSUPIAL, REMAINS, LAKE EYRE 


. mandible of Sthevuras anderson’ Marcus, 1962 from the 
corner of Luke Eyre North, South Australia. Occlusal view (slereopair of right ramus and anterior 
part of t eft ramus). 1.1, 


122 J, A, DULHUNTY, T. F. FLANNERY & J. A. MAHONEY 


andersoni can be distinguished from near rela- 
tives in the following ways: it differs from the 
species of Simosthenurus Tedford, 1966 in 
having more gracile dentaries, and in lacking 
the conspicuous crenulations on the molars. 
S. andersoni is smaller than the other species 
of Sthenurus (S, tindalei Tedford, 1966, S. 
notabilis Bartholomai, 1963 and S, atlas Owen, 
1838). While closest in size to S. atlas, S. 
andersoni differs from that species in possess- 
ing lower crowned molars with less well- 
developed linking. 

Sthenurus andersoni is known only from 
sediments of Pleistocene age. It has been 
recorded from several undated sites in South 
Australia including, tentatively (as S. cf. 
andersoni), localities along Cooper Creck, 
northeast of Lake Eyre (Williams 1980). The 
most recent date for S. andersoni is 20000 
years B.P. at Spring Creek, southwestern Vic- 
toria (Flannery 1980, Flannery & Gott in 
press). 

Diprotodon Owen, 1838 is a poorly under- 
stood genus and it is not possible at present to 
determine the specific name for most speci- 
mens. The Diprotodon remains from the bone 
bed are incomplete, consisting of molar (AM 


F65477) and postcranial fragments (AM 
F65478). As far as it is known, Diprotodon is 
restricted to the Pleistocene. Rich et al. (1982) 
record the genus from the Upper Pliocene, but 
this is an error (T. Rich pers. comm.). Ter- 
minal dates for Diprotodon are around 25 000 
years B.P. at Lancefield, Victoria (Gillespie 
et al. 1978) and 20000 years B.P. at Spring 
Creek, Victoria (Flannery 1980). 


The macropodine is represented by an astra- 
galus (AM F65479). It is similar in mor- 
phology and size to the astragalus of a large 
species of Macropus Shaw, 1790. 


The association of species of Srhenurus and 
Diprotodon is a characteristic feature of many 
Pleistocene fossil localities in southeastern Aus- 
tralia. In South Australia such an association 
has been recorded from several localities along 
the Warburton River and Cooper Creek near 
the northeast corner of Lake Eyre as well as 
from many other parts of the State (Williams 
1980). The latest date for sites containing 
these taxa is about 20 000 years B.P., but some 
such sites may be up to 2 million years old. 
On this basis, the Price Peninsula bone bed is 
interpreted as being Pleistocene in age, but 
probably older than 20 000 years. 


References 


CaLLeN, R. A. (1977) Late Cainozoic environ- 
ments of part of northeastern South Australia. 
J. geol. Soc. Aust. 24, 151-169, 

& Teprorp, R. H. (1976) New late Caino- 

zoic rock units and depositional environments, 

Lake Frome area, South Australia. Trans. R. 

Soc. §. Aust. 100, 125-167. 


Dutuunty, J. A. (1982) Holocene sedimentary 
environments in Lake Eyre, South Australia. J. 
geol. Soc. Aust, 29, 437-442. 

(1983) Lake Dieri and its Pleistocene 
environment of sedimentation, South Australia. 
J. Proc. R. Soc. N.S.W. 116, 11-15. 

FLANNERY, T. F. (1980) The Late Pleistocene 
Spring Creek site. Ree. Bur. Miner. Resour. 
Geol, Geophys. Aust. 1980/67, 27. 

& Gott, B. (in press) Spring Creek, a late 
Pleistocene megafaunal site in southwestern 
Victoria. Aust. Zool. 

Gittespiz, R., Horton, D. R., Lapp, P., 
MAcuUMBER, P. G., RIcH, T. H., THORNE, R. & 
WricHT, R. V. S. (1978) Lancefield Swamp 
and the extinction of the Australian megafauna. 
Science 200, 1044-1048. 

Jouns, R. K. (1963) Investigation of Lake Eyre, 
Part 1, The sediments of Lake Eyre Basin, 
evaporites and brines, their constitution and age. 
Rep. invest, Dep. Mines S. Aust. 24, 1-69, 


Kinc, D. (1956) The Quaternary stratigraphic 
record at Lake Eyre North and the evolution of 
existing topographic forms. Trans. R. Soc. 8. 
Aust. 79, 93-103. 

Ricw, P., Ricw, T. & BEARLIN, R. (1982) Mam- 
malia. Jn P. V. Rich & E, M. Thompson (eds), 
“The fossil vertebrate record of Australasia’, 
pp. 709-712. (Monash University Press: 
Clayton). 

STIRTON, R. A., TepForD, R. H. & MILLER, A. H. 
(1961) Cenozoic stratigraphy and vertebrate 
paleontology of the Tirari Desert, South Aus- 
tralia, Rec. S. Aust. Mus. 14, 19-61. 

WILLiAMs, A. F. (1976) Lake Eyre map sheet 
and explanatory notes, Geological Atlas of 
South Australia, 1:250 000 series (Geol. Surv. 
S. Aust.: Adelaide), 

WILLIAMS, D, L, G. (1980) Catalogue of Pleisto- 
cene vertebrate fossils and sites in South Aus- 
tralia, Trans, R. Soe, S. Aust. 104, 101-115. 

Woprner, H. (1974) Post-Eocene history and 
stratigraphy of northeastern South Australia. 
Thid, 98, 1-12. 

& TWIipace, C. R. (1967) Geomorphological 

history of the Lake Eyre Basin. In J. N. Jen- 

nings & J. A. Mabbutt (eds), “Landform studies 

from Australia and New Guinea”, pp. 118-143. 

(Australian National University Press: Can- 

berra). 


UPEROLEIA GRAY (ANURA: LEPTODACTYLIDAE) IN NEW GUINEA 


BY MICHAEL J. TYLER & MARGARET DAVIES 


Summary 


Uperoleia lithomoda is reported from New Guinea for the first time. The occurrence of the species 
across the north of Western Australia, the Northern Territory and Queensland is recorded. 


UPEROLEIA GRAY (ANURA; LEPTODACTYLIDAE) IN NEW GUINEA 


by Mictiath J. TybeR & MaArcAretT Davins' 


Summary 
Tycer. M1, & Dayers, M. (1984) Uperoleia Gray (Anuar: Leptodactylidac) in New Guinea, 
Trans. R. Soc. S. Aust. W8(2), 123-125, 12 June, 1984, 
Uperaleta lithomeda is reported from. New Goinen for the first time, The occurrence uf 
the speeies seross the north of Western Australia, the Northern Territory and Qucenslaud is 


recorded, 


Key Worns: Uperaleia, New Guinea, Cupe York Peninsuki, Osteology, Distribution.. 


Introduction 


The Jeptodactylid (imyobatrachid of some 
wuthors) genus C/peroleia Gray is one of the 
most poorly krown Australopapuan frog 
venera, Tyler, Davies & Martin (t9Sta) 
revised the Australian members of the venus, 
described fine new Species, resurrected twa 
others and raised to 16 the number of species 
recognised, Subsequently, ‘Tyler ec ab. 
(J98tb.c) ceseribed two further hew species 


from Western Australia and the Northern 
‘Lerrilory respectively, 
Tyler (1972) reported an Unidentified 


species Of L/peroleia collected by Fred Parker 
in the southern lowlands of New Ciuinea, 
representing the first record of the genus from 
Papua New Crome, Phese specimens wre the 
bisis on which Menzies (1975) and Zweifel & 
Tyler (1982) included the genus in the New 
Guinea fauna, 

Recently we have vompared the New 
Ciuinca speeimens with material from Horn 
Ishund Coll the lip of Cape York) and from 
various sites on the Anstralian mainland, 
These data demonstrate that the specimens 
represent U. fithontoda Tyler, Davies & 
Martin, Here we report on the New Ciunen 
specimens amd review the distribution af UL 
lithamoda 


Materials and Methods 


‘The specimens reported fure ae lodged in 
institutions abbreviated us follows: MCZ 
Muscum of Comparative Zoology, Ularvard 
University; NIM Northern Territory Museum 
and Art Giallery, Darwiny ONPWS Queenslind 
National Parks and Wildlife Service, Towns- 
ville; SAM South Australian Museum, Ade- 
laide, 


* Department of Zoology, University of Adelside, 
CPO. Box 498, Adehude, S. Aust, SOOT, 


Methods of measurement follow Tyler 
(1968). Skeletons were cleared and stained for 
bone after the method of Davis & Gore 
(1947). Recordings of male calls were 
analysed on a Kay Digital Sonagraph 7800 
and compared with sonagrams of U. litho- 
vue published by Tyler et al, (1981), 


Uperaleia lithomoda Tyler, Davies & Martin 
1981 

Uperoleia sp: Tyler, 1972, p. 237. 

Material examined: Papua New Guinent MCZ. 
Al06605, Y37616 (cleared and stained), SAM 
R29425, Moorehead, Western Districts; Australia, 
Queensland: MCZ Al06606, ABNI34, SAM 
R25423-24 Horn Island, Torres Strnit; QNPWS 
N28871, Lakefield, N.P., Cape York Peninsula; 
N32319, Coen, Cape York Peninsula: SAM 
K4941 Bentinck Island. Northern ‘Territory! NEM 
ROS, Fore Dam; RORSO-81, 2020-24 Shoul 
Bay, Darwin; R2885. Berry Springs; &2912, 25 km 
NE Noooamah; RS&868, Mary River, Annaburta 
Sin, SAM R24011-14, R2510-11, tl km NE 
Katherine, R24015-16, Suddle Creek. Victoria 
Hwy; R2d009, 25.6 km E NU/WA border, Vic- 
lorke Hwy: R25/08-9 17 Km Roper River Rd, 
Stuart Hwy Junctions R25106—-7, B enc Angurige 
Airport, Groote Eylindt; R24010 (Desmonds Pas- 
sae), Victoria Hwy, 409 Km W Katherine. 


Distributional date fram call pecards: 

W.AL 206. 5.3. 7.6, 91, 100, 10.8, 16.9, 19.3. 
OPA, 2b.4, 26.1, 26.7, 28.5. 32.4. 33.8, 37.1. 421, 
HO & RE km FE oof Wyndham township on 
Wyndhom/KRunniuea Rd) 93 & 43.7 km E 
Wyndhium aippartk tanoll, Wyndham Rundnurra 
Kal. 

NP) 1h2 & 13.3 km & Victoria River on Vie- 
foria Hwy: 2.0, 7.6, 92, 106. thd, 12.4, 13.5, 
I3.8 & I4.4 km EF Saddle Ch un Victorian Hwy: 
100, 295, 403.9, 406, 406.8, ANOS. Stl, 413.2 & 
415.1 km W Katherine on Victoria Hwy; 18.4 & 
20 km W Daly River /Stuart Hwy Jen, Duly River 
Rd 7 km W Mare River, Arnhem Tlwy; Mary 
River Bridye, Arnhem Hwy; Unibakumba, Groote 
Liylandl. ; ’ 

The two New Cininea specimens are adult 
females wilh snout to vent leneths of 19.7 and 
73.0 mim respectively. The type series did not 
include w single female, and the 16 males 


124 M. J. TYLER & M. DAVIES 


ranged 20.9-25.2 mm (Tyler ef al. 198la). tion in populations in Queensland and the 
Because female frogs tend to be larger than Northern Territory. 

conspecific males, the size of the smallest Osteologically the single New Guinea speci- 
female from New Guinea might be considered men cleared and stained (MCZ Y37616), 
sufficient to question its identity. However, the conforms closely with representatives of U. 
specimen falls within the range of size varia- lithomoda from the Kimberley Division of 


Fig. 1. A Dorsal and B ventral views of the skull of Uperoleia lithomoda from New Guinea (MCZ 
Y37616). Scale bar = 5 mm. 


iN 
a 
thijtad ayn ey 
YK ; ~ \ 
cial ue 
as a NEA JZ eS 
3 ae ih NA eK 
ite 
cae ce { 
ane, J a5 / “at, ° \ 
be LE ag e e ) C | aa 


> aw & 
WA 
3 ‘ 
BOT NS. 
=) e 
34> \ 
j ph ad 
| 
| 
| 
| 
| 
i 


Fig. 2. Distribution of Uperoleia lithomoda in Australia and New Guinea. 


UPEROLEIA IN NEW GUINEA 125 


W.A. Exposure of the frontoparictal fontancelle 
is greater than that of N.T. specimens illus- 
trated by Tyler ef al. (1981a) but is similar 
to the condition exhibited by specimens from 
the vicinity of the type locality, The medial 
edves and anterior extremities of the tronto- 
parictals are characteristically crenulated 
(Fig. JA). The nasals are triangular and 
widely separated posteromedially, The zygo- 
matic rami of the squamosals are minute and 
the medial arm of the pterygoid is charac- 
teristically broad and not acuminate, The pala- 
tines are slender and reduced laterally, angled 
to the midline, and the premovers are absent. 
The maxillary arch is edentate (Fig. 1B). 

Call: Parker did not record the call of the 
New Guinea or Horn Island specimens, but 
described it as a single, very loud note on 
Horn Island. This description fits our field 
observations, The nearest locality where calls 
have been recorded is Coen on the Cape York 


Peninsula (specimen QNPWS N32319; K. R. 
McDonald), That call is within the parameters 
of WU. lithomoda defined in the type descrip- 
tion, 


Distribution: Uperoleia lithomoeda ranges 
from the Kimberley Division of W.A. through 
the N.T, and Qld to New Guinea (Fig. 2). 
The wide gaps between some localities pro- 
bably reflects Jack of collecting, or our lack of 
access ta specimens from these areas, We 
suspect that the distribution will prove to be 
continuous on the Australian mainland, 


Acknowledgments 


This work was supported by an Australian 
Research Grants Scheme award. We thank 
Fred Parker, Keith McDonald, David Carter, 
Terry Schwaner and Pere Alberch for access 
to material. Fred Parker also is thanked for 
ficld notes and Keith McDonald for call data. 


References 


Davis, D. D. & Gore, V. R. (1947) Clearing and 
staining entire skelelons of small vertebrates. 
Fieldiana: Techniques (4), 1-16. 

Menzies, J. 1. (1975) Handbook of common New 
Guinea frogs. Wau Ecology Institute Handbook 
No, 1; (Wau, New Guinea). 

Tyuer, M. J. (1968) Papuan hylid frogs of the 
genus Myla, Zool. Verhand. Leiden (96), 1-203. 

, Davins, M. & Martin, A. A. (1981a) Aus- 

tralian frogs of the leptodactylid genus Upero- 

leia Gray. Aust. J. Zoal. Suppl. Ser. 79, 1-64. 


,— & (1981b) New and rediscovered 
species of frogs from the Derby-Broome area 
of Western Australia. Rec, W. Aust Mus. 9, 


147-172. 

& (198lc) Frog fauna of the 
Northern Territory: new distributional records 
and the description of a new species. Trans. JR. 
Soc. 8S. Aust. 105(3), 149-154. 

Zwrirer, R. G. & Tyter, M. J. (1982) Amphibia 
of New Guinea. In J. L. Gressitt (Ed.), Bio- 
geogruphy and Ecology of New Guinea, 
Monogr, Biol. 42, 759-801. 


WARDANG ISLAND - A REFUGE FOR MARGINOPORA VERTEBRALIS? 


BY Y. BONE 


Summary 


Marginbopora vertebralis Blainville is reputed to have become extinct in South Australian waters 
after deposition of the Glanville Formation although it still flourishes today at Shark Bay, and along 
the Great Barrier Reef where water temperatures are much higher than in the Southern Ocean. 
Samples have been retrieved from contemporary bottom sands at Quindalup, Geographe Bay (M. 
Lindsay pers. comm.) whilst the species has been reported as “rare” at Oyster Harbour near Albany 
and as “isolated to frequent” in bottom samples from the continental shelf south of the Eucla Basin. 
On Wardang Island it occurs in the Pliocene Hallett Cove Sandstone with examples up to 2.5 
centimetres in diameter, and in lightly indurated beach rock on the 2 metre high stranded beaches 
(Fig. 1) where it is of the order of <0.5 centimetres in diameter. This latter material may belong to 
the Glanville Formation, although extensive searching failed to locate any specimens of species 
commonly associated with M. vertebralis at this time, e.g. Anadra trapezia (Deshayes) or Pinctada 
carchariarum Jameson. 


BRIEF COMMUNICATION 


WARDANG ISLAND—A REFUGE FOR MARGINOPORA VERTEBRALIS? 


Marginopora vertebralis Blainville is reputed to 
have become extinct in South Australian waters 
after deposition of the Glanville Formation!.?, 
although it still flourishes today at Shark Bay®, 
and along the Great Barrier Reef where water 
temperatures are much higher than in the Southern 
Ocean. Samples have been retrieved from con- 
temporary bottom sands at Quindalup, Geographe 
Bay (M. Lindsay pers. comm.) whilst the species 
has been reported as “rare” at Oyster Harbour 
near Albany? and as “isolated to frequent” in 
bottom samples from the continental shelf south 
of the western side of the Eucla Basin®, On 
Wardang Island it occurs in the Pliocene Hallett 
Cove Sandstone with examples up to 2.5 centi- 
metres in diameter, and in lightly indurated beach 
rock on the 2 metre high stranded beaches (Fig. 
1) where it is of the order of <0.5 centimetres in 
diameter. This latter material may belong to the 
Glanville Formation’, although extensive searching 
failed to locate any specimens of species com- 
monly associated with M. vertebralis at this time, 


(\coose Island 


. 
x 
‘, 


. 
+ Reef 
\ ‘ 


Pliocene 


Permian 
(not to scale) 


\ 
Stranded | |! 
beaches / / 


t/ 
/} 


Cliff Point 


Fig. 1. Locality map. 


® 
Ali e= 
Beus, 


Fig. 2. M. from tidally deposited 


vertebralis 
detriths on northern beach of Wardang Island. 
Scale: 1 cm = 1 mm. SEM image. 


e.g. Anadara trapezia (Deshayes) or Pinctada 
carchariarum Jameson. 


However, it is also found as complete, <0.5 
centimetre diameter, discrete individuals amongst 
the daily tidally deposited detritus at the edge of 
the reef between Wardang Is. and Goose Is. (Fig. 
1). Numbers seen in this detritus are of the order 
of approximately 10 samples/5 metres/30 minutes 
searching time. Microscopic observation of M. 
vertebralis from this site, the beach rock and the 
Pliocene from Wardang Is., Glanville Fm, material 
from St Kilda beach and recently living specimens 
from the Great Barrier Reef leads one to suspect 
that this foraminifer may still be extant on the 
modern Wardang Is. reef. The tests at the latter 
locality show either none or little evidence of 
cementation (Fig. 2), and overall, surprisingly 
little aggradation, suggesting a nearby source. A 
determined effort to locate the source met with 
failure. so that it is likely that neither the rocks 
of the adjacent coast lines of Wardang and Goose 
Islands nor the reef rock are the source. This leads 
back to the possibility of the reef being a favour- 
able habitat for the continuing survival of the 
species after Glanville Formation time. 

To further test this hypothesis, a comparative 
analysis of the Mg content of the tests has been 
made (Table 1). For each location, 2 specimens 
were sclected, with care paid that they did not 
contain or have adhering any particles of matrix. 
These were crushed, taken up into solution in acid 
and then analysed for Mg by atomic absorption 
spectrophotometry. The living foraminifer secretes 
a high-Mg calcite test’, which after death and 
incorporation into the sediment, rapidly inverts to 
low-Mg calcite by diagenetic processes. 

Although the analyses do not prove the hypo- 
thesis, they certainly support the suggestion that 
the reef between Wardang and Goose Islands may 


bo 
ow 


TABLE 1. Chemical analysis of MgCOx content of 
Marginopera vertebralis 


Location Age MgC0O3% 
Wardang Island __ possibly present 8.99 
(shore-line 
detritus) 
Great Barrier present 9.75 
Reef 
Wardang Island Younger than Glanville 10.31 
(beach rock) Formation 
St Kilda Beach Glanville Formation 6.27 
Wardang Island Pliocene— 3.35 


(S.E. coast) Hallett Cove Sandstone 


be a refuge for M. vertebralis. The high MgCOs 
figure for the Wardang Island beach rock is inex- 
plicable, although suggestions such as groundwater 
influence (1) from high-Mg halophytic vegetation 
decomposition, (2) from underlying bedrock (R. 
Oliver pers. comm.) or (3) leading to the for- 


1Howchin, W. (1923). Australas. Ass. 
Advmt. Sc., 16, 94-101. 

2Howchin, W. (1935). Trans. R. Soc. S. Aust., 59, 
68-102. 

‘Logan, B. W., Brown, R. G. & Quilty, R. G. 
(1976). 25th Inter. Geol. Cong. Excursion Guide 
No. 37A. 

IMcKenzie, K. G. (1962). J. Proc. R. Soc. West 
Aust., 45, 117-132. 

5Chapman, F, & Parr, W. J. (1935). J. Proc. R. 


Soc. West. Aust., 21, 1-7. 


Rep. 


mation of microscopic dolomite rhombs, as 
commonly occurs in this type of environment (V. 
Gostin pers. comm.), are all possibilities. Although 
this high figure downgrades the validity of the 
hypothesis, the other analyses are nevertheless 
sufficient to warrant questioning the age of the 
reef samples, especially when compared to the 
analyses of 8.8%, 9.55%, 10.50% and 12.52% 
MgCOs3 content quoted for contemporary M. verte- 
bralis specimens from Fiji®. It is desirable that a 
thorough search be made for living specimens in 
this area. Care will be needed. Hasty decisions 
based on protoplasm-staining techniques could be 
misleading as various algae and bacteria rapidly 
invade vacated tests. 

Due to changing policies regarding the use of 
Wardang Island, the author has been unable to 
return and follow up this work. It is, however, 
hoped that this note will stimulate further investi- 
gations into the occurrence of M. vertebralis in 
South Australia’s past—and present? 

SBone, Y. (1978). B.Sc. (Hons.) thesis Univ. 
Adel. (unpubl.). 

7Firman, J. B. (1966). Q. geol. Notes, geol. Surv. 
S. Aust., 17, 6-8. 

‘Bathurst, R. G. C. (1975). Carbonate sediments 
and their diagenesis. (Elsevier: New York). 
‘Loeblich, A. R. & Tappan, H. (1964). Jn: 

Moore, R. C. (Ed.), Treatise on Invertebrate 
Paleontology. Part C, Protista 2, Vols. 1, 2. 

(Geol. Soc. America & Univ. Kansas Press). 


Y. BONE, Department of Economic Geology, University of Adelaide, G.P.O. Box 498, Adelaide, S. 


Aust. 5001. 


THE IDENTITY OF ORCHEZELANDIA RUBRA (COLLEMBOLA: 
ENTOMBRYIDAE) 


BY PENELOPE GREENSLADE 


Summary 


In the course of a study of the scaleless Orchesellini of Australia, a New Zealand species, 
Orchezelandia rubra Salmon, 1937, which had been placed in the tribe, was examined and found to 
belong to the genus Entomobrya in the Entomobryini of the same family. 


BRIEF COMMUNICATION 


129 


THE IDENTITY OF ORCHEZELANDIA RUBRA 
(COLLEMBOLA: ENTOMOBRYIDAE) 


In the course of a study of the scaleless 
Orchesellint of Australias! a New Zealand species, 
Orchezelandia ribra Salmon, 1937, which had 
been phiced in the trihe, Was examined and found 
to belong to the genus Entomeabrya in the Ento- 
mobryini of the same family, 

With the transfer of this species ram the 
Orchesellint to the Entomobryini, there remain 
no Sealeless Orchesellini species recorded from 
New Zealand and only one scaled species, Hetero- 
wineus tides (Templeton)? Like New Zealand 
(he tribe is poorly represented in Australia also 
compared wilh Europe. A single tropical sculed 
species, Feterannurus (Alloscopus) tetracanthuys 
Borner. is Known from north Queenslund', and i 
few species of ihe genus Ausiruloramnnuy Stach 
from the southeast! of Australia. The latter genus 

as originally deseribed in the Isolomidae but is 
now known to be 4 member of the Orchesellini, 


Entomohrya rubra (Salmon) nov, comb, (Pig, 1) 
Orchezelandia rubra Salmon, 1937, Trans, Rov. 
Soc. N.Z, 67(2). 356, plate Slu-e. 
Orchesellides rubrum Bonel, 1942, 
3(2)> 46 
Orehesellidey rubra Sulroon, 1944, Ree Dom. 
Mus. 1/2): 165, 


Ciencia 


ry i j 
\ L i 

i ‘ 4 

7 

j \ 

. "i 
ho aN 
a 


Big. t, Gtehezelandia sebra dolotype. a. lott side 
of tend afd antennal segment fob. right side 
ol tend anh antennal segment 1, ¢. dorsal view 
thurys Howbuumen IVI, a. claw EL e, dorsal 
view abdomen Tt right side showing miacro- 
vhueliue. 


Material examined. Holotype. Orehenellides 
(reherelandia) rabia, dew IT. Salman, ‘Type 
(Mrtd euparil) NMNZ 3/197, Newberry, Palm 
Nth, in soil, (933 (hatrelled by TIES), 

Salmon erected a mew genus of Scaleless Orehe- 
sellin, Qrefeselundia i 137 on the basis of a 


specics with five-segmented antennae, tridentate 
mucrones und ratio of abdominal segments IIT: 
IV of 1:3-4, These churucters were fizured, He 
therefore clearly separated Orchezelandia from 
Orchesella Templeton on the basis of antennal, 
mucronal and abdominal morphology. Although 
Hot definitely stated, i appears Salmon only had 
oné specimen of his species and in 1941 he 
recorded the registration number when he pub- 
lished a pew deseription of O. rabra with redrawn 
figures’; he also corrected the description of the 
mucro to “bidentate with basal spine”. By so doing 
he removed one of the characters by which 
Orchezelandia dillered from Orchesellides, ww genus 
described eurlier by Bonet also with five-seg- 
mented antennae, leaving only a differing ratio of 
ubdominal segments WL wnd LV to distinguish the 
Iwo gene»ra, 

In 1942 Bonet synonymised Oreherelandia with 
Orchesellides stating that Salmon's deseripion 
concurred ahsoliutely with his own. He mentioned 
the differences in the proportions of abdomen TT 
und (Y (ie, from 1:3-4 in O. rubrion lo 3:4 in 
Orehesellides horael) and stated they were “clearly 
of specific value’. Salmon later ugread with 
Bonet’y synonymy und pubhshed it as Orchesel- 
lidey rubra nov. comb, in 1944, 

A recent study tus shown thal the (ype and 
sole specimen has four-segmented  antennue, 
althooeh the insertion of (he first antennal segment 
on the right side is unusually prominent on the 
mounted type but it is clearly without setae. Ta 
orher respects the specimen sgrees with Salmon's 
description although some ol the churucters cannot 
be seen clearly Le, the muoecrones and tip of ant. 
IV, because of the alignment of the specimen and 
wl darkening of the mounting medium, ‘The ratio 
ul abdomen TITY is as Salmon stated (123.5) 
aml the body is covered with short ciliated setoc 
wilh some macrochuelae wilh flexed tips which are 
depse on Thorax UP and less dense on other 
seamente and on the head. In all respects the 
specimen agrees (in those characters which vat 
be seen) with Evfomehrya BRondani and itis 
hereby transferred to (his genus. Bonet conunents 
thal Orechesellides is closely related to Enip- 
molrye cupart from the scementation of the 
antennae dnd ratio of abdominal segments UP and 
IV. A complete description of the species should 
uwwuil recollection from the type localily and revi 
sion of genus Lrromohliva from New Zealand. 

Enfomebrya rubra seems wearest to 8, hand 
selin’ Stach in colour pattern ratio of ubdomen 
Hl to TV, antennal length and ratio of antennal 
segments and claw, The trochanter organ, Lip 


130 


of ant. IV, bothriotricha, and labrum cannot be 
seen clearly on the specimen. However I have 
compared E. rubra with various species of Ento- 
mobrya, Australotomurus Stach and Orchesella 
and it is more similar to species of Entomobrya 
than to those of the other two genera both in 


IMari Mutt, J. A. & Greenslade, Penelope, In 
press. 


2Wise, K. A. J. (1977). Bull. Auckland Inst. Mus. 
11, 1-176. 


general appearance and in other morphological 
details. 


This work was carried out while the author was 
in receipt of a grant from the Science and Indus- 
try Fund, and a French Government Scholarship. 


3Mari Mutt, J. A. (1982). Pacif. Insects 24, 84— 


PENELOPE GREENSLADE, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000. 


ERRATA 


BLACKBURN, G., ALLISON, G. B. and LEANEy, F. W. J. (1982). Further evidence on the age 
of tuff at Mt. Gambier, South Australia. Trans. R. Soc. S. Aust. 106(4) 163-167. 


The isotope data reported were slightly in error. The correct data are given in the Table. 


TABLE 1. 


Details of charcoal samples and isotope measurements. 


Depth of 


Depth of 


sample Thickness sample below 14C Age 
Distance from below of tuff vole. affected Acid pre- Alkali pre- 
Volcanic Blue Lake surface  layer(s) material 513C treatment treatment 
Sample No. CSIRO No.(s) Ash Zone (km) (m) (m) (m) (%- Tel to PDB) (+ 1 sd.) 
1 CS44 — Outer (B) SE 0.6 470 + 210 n.d. 
2 CS35 CS76 Inner (A) 4SSE 0.6 0.03 0.3 _ —24.3 7240 + 250 7300 + 250 
3 CS34. -CS74 Outer SNNE 0.45 = 0.15 _— —25.5 7670 + 240 7710 + 150 
4 CS45 — Outer 8NNE 0.20 860 + 210 n.d. 
5 CS46 _— Outer 4N 0.6 3600 + 220 n.d. 
6 — CS101 Outer 4N 0.5 —_ 0.02 — —24.2 n.d. 470 + 200 
7 CSs43 CS75 Outer 5SSW 0.5 _— 0.04 — —24.4 5680 + 230 5550 + 245 
8 _— CS102 Inner 2.5N 1.1 0.45 0.10 _ —25.5 n.d. 4670 + 240 
9 CS40 _ Inner 2.5N 1,1 0.3 0.15 — —_ 4060 + 285 n.d. 
10 C842. -CS77 Inner 2SE 1.6 0.6 0.10 _ —24.8 4450 + 230 3380 + 200 
11 CS41 CS78 Outer 4NNE 0.9 0.04 0.23 _ —25.1 8190 + 260 6700 + 240 


LIFE HISTORY OF THE SCIRON SKIPPER TRAPEZITES SCIRON 
EREMICOLA BURNS (LEPIDOPTERA: HESPERIIDAE) 


BY R. H. FISHER 


Summary 


The skipper butterfly Trapezites sciron eremicola inhabits open heathland in mallee areas from Eyre 
Peninsula to western Victoria. The nominate subspecies, T. s. sciron Waterhouse & Lyell, was 
described from southwestern Australia. The life history and early stages of ssp eremicola are 
described here from material collected in Ngarkat Conservation Park (35°40°S, 140°30°E), South 
Australia. 


BRIEF COMMUNICATION 


LIFE HISTORY OF THE SCIRON SKIPPER TRAPEZITES SCIRON 
EREMICOLA BURNS (LEPIDOPTERA: HESPERIIDAE) 


The skipper bullerlly frapezites sciron eremi- 

coalu inhabits open heathlind in mallee ureas from 
Eyre Peninsula to western Victoria, ‘The nominate 
subspecies. 7, 9. sefron Waterhouse & Lyell. was 
dexcribed from southwestern Australia. The life 
history and carly stages of ssp erenticala are 
described here from material collected in Nearkat 
Conservation Park (35°40, 140°30'B), Sonth 
Australia, 
Larval food plant: Lemandra glauea (R. Bri) 
Ewart (Liliaceae), In the butterfly’s habitat the 
plant occurs predominantly on the northern aspect 
of the crests of ridges and sandhills (Fig, 1), 


Billiatt 
Larval 


Fig. 1. Habitat of T. seiran ecrenucola, 
Conservation Park, South Australia, 
food plants in left foreground, 


Deseription of immature stages, Egg (Pig. 2A): 
diameter | tim; almost hemispherical with 19-22 
distinet verticnl ridges intersected by numerous 
obscure literal lines; pale cream when newly tit 
but developing a broad brown lateral band after 
several days. 

Kirst instar larva (Fig, 2B): length 3 mm: body 
white with a few long posterior setae, prothoracic 


!Atking, A. FL & Miller, C. G. (1977), Aust, ent. 


R. H. FISHER, 21 Seaview Rd., Lynton, 5, Aust. 


plate a dark brown transverse band; head shining 
black with a few short setae. 

Mature larva (Fig. 2C, D): length 20 mm; body 
white with obscure grey markings and a distinct 
grey dorsal line, spiracles black, prothoracic plate 
with a black posterior margin, anal plate with 
numerous swollen white setae arising from black 
bases and with four short, black-tipped posterior 
setae; head capsule rugose and with short swollen 
setae, dark brown with scattered paler markings: 
frons with paired longitudinal pale brown bands 
diverging ventrally. 

Popa (Fig. 2E, F); length 17 mm; cylindrical, 
abdomen tapering sharply and terminating in a 
red-hrown cremaster with a cluster of hooked 
setae; pale brown with darker markings, particu- 
larly on the head and thorax, body surface except 
wing cases with groups of branched white setuc. 
The setae resemble those described on the pupae 
of Trapezites heteromacula Meynck & Lower. 
Biology: Eggs are Jaid singly on the leaves of the 
food plant. The young larva emerges froni the egg 
alter about five days and makes a simple shelter 
by joining the bases of a few fresh leaves of the 
food plant with silk. When more mature it con- 
structs a silken shelter incorporating debris, dried 
lenves and sand which is attached to residual 
leaves where they arise from the basal sheaths of 
the food plant (Fig. 2G, H). Pupation occurs in 
late August within the shelter, which is open al 
the top and well-concealed. Adults appear from 
spring to early summer and both sexes exhibit 
hill-topping behaviour in open areas on {he crests 
of sandhills. 

I thank the Wildlife Conseryajon Fund for 
financial assistance in field work, the National 
Parks & Wildlife Service for permission to collect 
material in Ngarkat Conservation Park, D. F. 
Crosby and A. E, Mitchell for the use of vehicles 
and J.S. Womersley for botanical identification, 


Mag, 3, 104-106, 


5062, 


Fig. 2. T. sciron eremicola. A egg. B first instar Jarva, C mature Jarva. D head of mature larva, E 
pupa with larval head cast at left. F branched setae on pupa, G, H Jarval or pupal shelters, H 
partly Opened to show pupa. Bar scales A, B, D, F=1 mm; C. E. G, H=1 cm, 


FURTHER ADDITIONS TO THE MARINE FISH FAUNA OF SOUTH 
AUSTRALIA 


BY C. J. M. GLOVER 


Summary 


Six species of fish are newly recorded for South Australia, most on the basis of single specimens. 
One species, Idiacanthus niger, represents the first record of the family Idiacanthidae in the region. 
A seventh species (Metavelifer multiradiatus) and family (Veliferidae), are reported from South 
Australian waters for the first time. Four of the species were included in a checklist for the south 
east of the State but without further details. All species reported here have been recorded elsewhere 
in temperate waters around Australia; some also in tropical waters. All occur beyond Australia. 


BRIEF COMMUNICATION 


133 


FURTHER ADDITIONS TO THE MARINE FISH FAUNA OF 
SOUTH AUSTRALIA 


Six species of fish are newly recorded for South 
Australia, most on the basis of single specimens. 
One species, /diacanthus niger, represents the first 
record of the family Idiacanthidae in the region, 
A seventh species (Metavelifer multiradiatuy) and 
family (Veliferidae), are reported from South 
Australian waters for the first time. Four of the 
species were included in a checklist for the south 
east of the State! but without further details. All 
species reported here have been recorded else- 
where in temperate waters around Australia; some 
also in tropical waters. All accur beyond Austra- 
lia. 


Some of the species are acknowledged wide- 
ranging oceanic forms (Heyanchus griseus, Gas- 
terochisma melampus, Makaira indica and Lago- 
cephalus lagocephalus, but lack of supplementary 
records does not indicate permanent extensions of 
distribution into the South Australian region, As 
with some earlier records?-7 these latter records 
probably represent vagrants. 


The discovery of H. griseus, I, niger and Cyttus 
novdezelandiae reflect recent increased deepwater 
commercial fishing off South East South Australia. 


Family Hexanchidae. Wetanchas griseus (Bon- 
naterre, 1788). 


A Specimen (Fig. 1) was captured on a long- 
linc, set at 329 m depth, on a flat “muddy” sea 
bed, about 46 km southwest of Beachport. S. Aust. 
(approx. 37°43'S, 139°35°E), on 17.vii,1982, by 
R, Morgan. 

I. griseus has been recorded in Australia off 
Tasmania, Victoria and New South Wales, be- 
tween Port Fairy (Vic,) und Norah Head 


Fig. 1, Hexgnehus griseus, SAM FA740, TL. 1450 
mm, 


(N.S.W.)§, unpubl. rees.. 1 is found in Lropical and 
temperate waters worldwide." 


Family Idiacanthidae. Jdiacanthuy niger Regan, 

1914, 

One specimen (Fig, 2) was trawled, at 549 m 
depth, about 47 km southwest of Cape Buffon, 5. 
Aust. (approx. 37°52'S, 139°43’B), in iv. 1981, by 
J. Sealey. 

I, niger has been recorded in Ansiralia in deep 
water off western Vic. and N.S.W.(unpubl. recs), 
and is also known off New Zealand, Chile and 
South Africal.11, 

Family Veliferidae, Metavelifer multiradiatus 

(Regan, 1907). 

A specimen (Fig. 3) was trawled, at 40 m 
depth, 4 km southeast of Evans Island, near 
Ceduna, S. Aust. (approx. 32°24’S, 133°31'B), 
on 3,xi.1983, by A. Olsen. 

M. multiradiatus has been reported previously 
from W.A., N,S.W.!2, and “the Great Australian 
Bight", It has been recorded also (but not re- 
ported) in the Great Australian Bight specifically 
off the 8. Aust. coastumpubl. rees.. Beyond Australia, 
the nominal Felifer mualtispinosus Smith, 1951, 
from southeast Africa, is a synonym of M, piglit- 
radiatus™*, 


Fig. 2. Idiacanthuy nirer, SAM F4757, TL 400 mm. 


134 


Fig. 3. Metavelifer multiradiatus. SAM F4746. TL 
109 mm. 


Family Zeidae. Cyttus novaezelandiae (Arthur, 

1885). 

Five specimens (Fig. 4) were trawled, at 549 
m depth, about 47 km southwest of Cape Buffon, 
S. Aust. (approx. 37°52’S, 139°43’E), in iv.1981, 
by J. Sealey. 

C. novaezelandiae has been recorded in Austra- 
lia from Tasmania, Vic. and N.S.W.S8 It is also 
known from New Zealand!°, 

Family Scombridae. Gasterochisma melampus 

Richardson, 1845, 

A specimen (Fig. 5) was trolled, about 23 km 
west-northwest of Port MacDonnell, S. Aust. (ap- 
prox. 38°03’S, 140°26’E), on 7.vi.1982, by I. J. 
Carrison. 

G. melampus has been recorded in Australia 
from off Tas., Vic., N.S.W. and Qld.§ It is also 
known from other temperate waters of the 
southern hemisphere, e.g. New Zealand, Argen- 
tina and South Africa19.11, 


: venga * 
am, + 


ee 


kh ‘ee 
ee 


Fig. 6. Makaira indica. TL 3820 mm. 


ie = 

si a —. 2h = 

— Meee ee } ‘ : 
‘4 


Fig. 4. Cyttus novaezelandiae. SAM F4745. TL 
(largest specimen) 195 mm. 


Family Istiophoridae. Makaira indica (Cuvier, 
1831). 


‘A dead specimen (Fig. 6) was found floating 
at the surface near the grain wharf at Thevenard, 
S. Aust. (pprox. 32°09’S, 133°39’E), on 15.iv. 
1983, by A. E. & D. J. Holder. Fisheries officer 
J. N. R. Smith took the photograph (Fig. 6) and 
measurements. A. E. Holder of Thevenard kept 
only the head. 


Fig. 5. Gasterochisma melampus. SAM F4759, TL 
970 mm. 


ee 


2 . 


fee)? * 


> ire* 
esr 
> bas 


_ 


_ ; eho ; 
ee 3 4 
ae fi 
rsaled 


M. indica has been recorded in Australia from 
W.A. (Shark Bay and off Albany) and N.S.W. 
(Port Stephens) (unpubl. recs.), Tt ranges widely in 
circumglobal temperate and tropical waters!. 


Family Tetraodontidae. lago- 


cephalus (Linnaeus, 1758). 

A dead specimen (Fig. 7) was found on the 
beach near the jetty at Port MacDonnell, S. Aust. 
(approx. 38°03’S, 140°42’E), on 9.v.1983, by P. 
Cawthorne. 


Lagocephalus 


Po Set iF 
ee * ie 
* “4 


» 


——. ~ 
bg rs 
> Sen e-3te oa 


135 


L. lagocephalus has been recorded in Australia 
from Tasmania’. This species is well known in the 
Atlantic, Indian and Pacific Oceans!®. 


Those persons mentioned (all professional 


fishermen) are thanked for collecting and donating 
the specimens to the South Australian Museum. 
G. Bond (Arena Sports Store, Mt Gambier) and 
J. N. R. Smith (S.A. Dept. of Fisheries) are 
thanked for assistance with the L. lagocephalus 
and M. indica specimens respectively. 


Fig. 7. Lagocephalus lagocephalus. SAM F4733. TL 520 mm. 


1Glover, C. J. M. (1983) Freshwater and marine 
fishes. In Tyler, M. J., Twidale, C. R., Ling, 
J. K. & Holmes, J. W. (Eds) “Natural History 
of the South East”, pp. 157-167. (Royal Society 
of South Australia: Adelaide.) 

2Glover, C. J. M. (1966), Rec. S. Aust. Mus. 15 
(2), 353-355. 

8Glover, C. J. M. (1974). South Aust. Nat. 49 
(1), 12-13. 

4Glover, C, J. M. (1976). Ibid. 50(4), 69-72. 
5Glover, C. J. M. & Branden, K, L. (1978). Ibid. 
52(4), 55-60. 

6Glover, C. J. M. & Branden, K. L. (1983). 
Trans. R. Soc. S. Aust. 107(2), 134-136. 
TKailola, P. J, & Jones, G. K. (1981). Ibid. 105 
(4), 211-212. 

SLast, P. R., Scott, E. O. G. & Talbot, F. H. 
(1983). Fishes of Tasmania, (Tasmanian 
Fisheries Development Authority: Hobart.) 


°Bass, A. J., D.’Aubrey, J. D. & Kistnasamy, N. 
(1975). Invest. Rept. No. 43, Oceanographic 
Resch. Inst., South African Assoc. Mar. Biol. 


Resch. 
10Whitley, G. P. (1968). Aust. Zool. 15(1), 
1-102. 
Usmith, J. L. B. (1950). The Sea Fishes of 


Southern Africa. 
South Africa.) 

12Munro, I. §, R. (1957). Fisheries Newsl. 16(9), 
15-18. 

13Whitley, G. P. (1962). Marine Fishes of Aus- 
tralia, Volume 1. (Jacaranda Press: Brisbane.) 

14Walters, V. (1960). Copeia, 3: 245-247. 

Nakamura, I, (1975). NOAA Tech. Rept. 
NMEBFS SSRF-675, part 3, 17-27. 

16Fowler, H. W. (1928). Memoirs Bernice P. 
Bishop Mus., X, i-iii, 1-540, pls. I-XLIX. 


(Central News Agency Ltd: 


C.J. M. GLOVER, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000. 


A RE-EXAMINATION OF TAENI MASTERSII KREEFT, 1871 AND 
TAENIA FIMBRIATA KREFFT, 1871 


BY I. BEVERIDGE 


Summary 


In 1871, G. Krefft published brief and inadequate accounts of a number of species of cestode 
collected from Australian mammals and birds. None of the species is recognisable by contemporary 
standards. Johnston redescribed and rehabilitated, from Krefft’s type specimens, most of the 
cestodes from birds. He listed Taenia mastersii Krefft, 1871 and T. fimbriata Krefft, 1871, both 
from wallabies, but did not redescribe them or comment on their affinities. Beveridge, in revising 
the Anoplocephalidae of Australian marsupials, pointed out that Krefft’s names, Taenia mastersii 
and T. fimbriata, could take priority over newer names currently in use if the species could be 
rehabilitated. 


BRIEF COMMUNICATION 


A RE-EXAMINATION OF TAENIA MASTERSII KREFFT, 1871 AND 
TAENIA FIMBRIATA KREFFT, 1871 


{1 1871, G. Krefft published brief and inade- 
quate accounts of a number of species of cestode 
collected from Australian mammals and_ birds. 
None of the species is recognisable by contem- 
porary standards. Johnston? redescribed and 
rehabilitated, from Krefft’s type specimens, most 
of the cestodes from birds. He listed Taenia 
mastersii Krefft, 1871 and T. fimbriata Krefft, 
1871, both from wallabies, but did not redescribe 
them or comment on their affinities, Beveridge®, 
in revising the Anoplocephalidae of Australian 
marsupials, pointed out that Krefft’s names, 
Taenia mastersii and T. fimbriata, could take 
priority over newer names currently in use if 
the species could be rehabilitated. At the time, 
serial sections of mature proglottides of the 
types of 7. mastersii in the Australia Helmin- 
thological Collection (AHC) housed in the South 
Australian Museum (S70) were examined?, and 
were considered to be identical with Calostaurtus 
thylogale Beveridge, 1975 a davaineid cestode 
recently described from the pademelon, Thylo- 
gale billardierii in Tasmaniat. However, because 
the features of the scolex are essential for specific 
identification in this genus and because no scoleces 
were present in the type material examined, no 
change could be made to the status of T. mastersii. 
No type specimens of T. fimbriata were found in 
Johnston’s collection (AHC), but specimens 
identified in Johnston’s handwriting as TT. fim- 
briata, collected from Thylogale stigmatica in 
Queensland, were located, and after examination 
were re-identified as Progamotaenia zschokkei 
(Janicki, 1906)°8. 

The writer has recently located type material 
of both T. mastersii and T. fimbriata in the Aus- 
tralian Museum (G11160 and G11156). The 
type specimens of T. mastersii (G11160) consist 
of 4 complete specimens including the scoleces, 
In an intact scolex examined, the rostellar hooks 
were arranged in a_ six-lobed circle with hook 
lengths between 30-40 um long. These two 
features, namely the shape of the ring of rostellar 
hooks and their lengths, are characteristic of C. 
thylogale and clearly identify T. mastersii and C. 


'Krefft, G. (1871). Trans. Entomol. Soc. N.S.W. 
2, 206-232. 

2Johnston, T. H. (1912). Rec. Aus. Mus. 9, 1-35. 

“Beveridge, I. (1976). Aust. J. Zool. Suppl. Ser. 


No. 44, pp. 110. 


thylogale, supporting earlier observations on the 
morphology of the mature proglottides?. C. thylo- 
gale therefore becomes a synonym of T. mastersii 
as Calostaurus mastersii (Krefft, 1871) comb. 
nov. C. mastersii was originally collected from an 
unknown species of wallaby in Queensland. All 
recent collections are from T. billardierii in 
Tasmania‘. 


The type specimen of 7. fimbriata in the Aus- 
tralian Museum (G11156) has external features 
similar to P. zschokkei with a broad, prominently 
fimbriated velum and paired genital pores. Serial 
histological sections of a fragment of the type 
specimen also reveal internal features similar to 
P. zschokkei, namely paired genitalia, a single 
uterus, heavily armed and coiled cirri and large 
seminal receptacles. However, even the youngest 
proglottides of the type specimen are nearly gravid, 
with the uteri obliterating most of the sexual 
organs and consequently the distinguishing charac- 
ters of the mature proglottis (distribution and 
number of testes) cannot be determined. P. 
zschokkei and P. villosa (Lewis, 1914) are the only 
known species of the genus with fimbriated vela 
and a single uterus. They can be distinguished 
from one another by the number of testes per 
proglottis, by the length of the velum and by the 
pattern of development of the genitalia®. T. fim- 
briata has a velum similar to that of P. zschokkei, 
but assessment of this character is somewhat sub- 
jective and neither the number of testes per pro- 
glottis, nor the pattern of development of the 
genitalia can be determined from the type frag- 
ment of 7. fimbriata, Therefore, although T. fim- 
briata and P. zschokkei are possibly the same 
species, their identity cannot be demonstrated 
beyond reasonable doubt. Since the host and col- 
lection locality of the type of T. fimbriata are not 
known, and thus further collections of the parasite 
cannot be made, T. fimbriata must continue to be 
regarded as a nomen nudum, 


Thanks are due to P. Hutchings for the loan of 
type specimens from the Australian Museum, 
Sydney. 


+Beveridge, I. (1975). J. Helminthol. 49, 129-136. 
"Beveridge, I. (1981). Trans. R. Soc. S. Aust. 105, 
139-147, 


I. BEVERIDGE, Veterinary Sciences Division, South Australian Department of Agriculture, Institute 


of Medical and Veterinary Science, Frome Road, Adelaide, S. Aust. 


5000. 


THE IDENTITY OF RED-BELLIED BLACK SNAKES ON KANGAROO 
ISLAND 


BY TERRY D. SCHWANER 


Summary 


The red-bellied black snake, Pseudechis porphyriacus, although not reported previously from 
Kangaroo Island, was listed as occurring on the island based on specimens in the South Australian 
Museum. The authors stated, “This snake, like the Black Tiger, is jet black above and grows to 
about 1.5 m, but is distinguishable by its whitish belly which is usually tinged along its lateral 
margins with pink or crimson ... On the island it has been seen in coastal heath on the south side, 
but it is not common and its usual habitat there is in doubt.” Two recent observations have led me to 
conclude that Pseudechis porphyriacus is not a member of the herpetofauna of Kangaroo Island. 


BRIEF COMMUNICATION 


137 


THE IDENTITY OF RED-BELLIED BLACK SNAKES 
ON KANGAROO ISLAND 


The red-bellied black snake, Pseudechis porphy- 
riacus, although not reported previously from 
Kangaroo Island,!.2,% was listed as occurring on 
the island based on specimens in the South Aus- 
tralian Museum.t+ The authors stated, “This snake, 
like the Black Tiger, is jet black above and grows 
to about 1.5 m, but is distinguishable by its whitish 
belly which is usually tinged along its lateral 
margins with pink or crimson ... On the island it 
has been seen in coastal heath on the south side, 
but it is not common and its usual habitat there is 
in doubt.4 Two recent observations have Jed me to 
conclude that Pseudechis porphyriacus is not a 
member of the herpetofauna of Kangaroo Island. 

The only specimen identified as Pseudechis por- 
phyriacus in the South Australian Museum Col- 
lection (SAM R4366, Kelly Hill Caves, K.1.) is 
reidentified as a melanistic tiger snake of the 
Notechis scutatus/ater complex, No specimens of 
Pseudechis porphyriacus from K.I. have been 
located in the collections of the Australian 
Museum, Western Australian Museum, National 
Museum of Victoria, Museums & Art Galleries 
of the Northern Territory, Australian National 
Wildlife Collection or the Queensland Museum 
(R. Sadlier, G. Storr, J. Coventry, P. Horner. J. 
Wombey and J. Covacevich pers, comm.). 


Recent studies on variation in colour pattern of 
tiger snakes (Notechis  scutatus/ater complex) 
of Kangaroo Island involved the collection of 
fresh, roadkilled specimens by rangers of the 
South Australian National Parks and Wildlife 
Service and local residents. Among these speci- 
mens were tiger snakes with reddish bellies (Fig, 
1). Red-bellied tiger snakes are colour forms of 
the highly variable Kangaroo Island populations 
(Schwaner, in prep.), However, these are easily 
distinguished from Pseudechis porphyriacus by the 
shape of their subcaudal scales, in which most 
scales are entire, undivided plates (divided and 


1Waite, FE. R. (1925), Rec. S. Aust. Mus. 3, 17-32. 

“Waite, E. R. (1927). Trans, R. Soc. S. Aust, 51, 
326-329. 

*’Waite, E. R. (1929) “The Reptiles and Amphi- 
bians of South Australia.” (Government Printer: 
Adelaide. ) 


paired in P. porphyriacus). Localities for two 
specimens with reddish bellies are both in the 
Hundreds of Haines district in South Central KJ1., 
although similarly coloured individuals could 
be found elsewhere on the island. 

Thus red-bellied black snakes on Kangaroo 
Island are not Pseudechis porphyriacus but red- 
bellied, melanistic tiger snakes (Notechis scutatus/ 
ater complex). 

I thank T. Dennis, M. McKelvey, H. Stitchell 
and J. Watkins for collecting the specimens. 
Roman Ruehle took the photograph in Fig. 1, and 
Lila Schwaner typed the manuscript. 


Fig. 1. Ventral pattern of a red-bellied, melanistic 
tiger snake (Notechis scutatus/ater complex) 
from Kangaroo Island. The specimen (SAM 
R24987) is an adult female (SVL = 1115 mm) 
collected on a road at Ayliffe Hill (Hund. of 
Haines) by M. McKelvey on 3.xi.83. Light 
areas of the belly are Spinel Pink (Colour 
108C).5 Note the undivided subcaudal scales. 


4+Houston, T. F. & Tyler, M. J. (1979) In M. J. 
Tyler, C. R. Twidale, & J. K. Ling, Eds. “Natural 
History of Kangaroo Island”. (Roy. Soc. S. 

Aust.: Adelaide). 

5Smithe, F. B. (1981). Naturalist’s color guide. 

Part IIL (Amer, Mus, Nat, Hits.: New York.) 


TERRY D. SCHWANER, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000. 


VOL. 108, PARTS 3 & 4 
13 DECEMBER, 1984 


Transactions of the 


. 


Royal Society of South 
Australia 


Incorporated 2 
a pa ery Saeate  e h, l S S el e 


Kangas, M. I. & Geddes, M. C. The effects of salinity on the distribution of amphipods 
in the Coorong, South Australia, in relation to their salinity 


tolerance - - - - - - - - - - 139 
Bayly, 1. A. E. A new species of Ca/amoecia (Copepoda: Calanoida) from South 

Australia, and comments on three congeners - : = = 147 
Mahony, M., Tyler, M. J. & Davies, M. A new species of the genus Rheobatrachus 

(Anura: Leptodactylidae) from Queensland - - - - - 155 
Callen, R. A. Quaternary climatic cycles, Lake Millyera region, southern Strzelecki 

Desert - - - - - - - - - - - 163 
Twidale, C. R. & Campbell, E. M. Murphy Haystacks, Eyre Peninsula, South 

Australia = - - - - - - - - - - - 175 
Beveridge, I. Dasyurotaenia robusta Beddard, 1912, and D. @asyuri sp. nov., from 

carnivorous Australian marsupials - - - - - - 185 


Beveridge, I., Speare, R. & Johnson, P. M. New records of Globocephaloidinae 
(Nematcda: Trichostrongyloidea) from Macropodidae in north 
Queensland - - - - - - 2 - - - 197 

Greenslade, P. & Wise, K. A. J. Additions to the collembolan fauna of the Antarctic 203 

Jago, J.B., Daily, B. D., Von Der Borch, C. C, Cernovskis, A., Saunders, N. First 
reported trilobites from the lower Cambrian Normanville Group, 
Fleurieu Peninsula, South Australia - - - - - - 207 


Brief Communications: 


Lange, R. T. Leaf marking in rangeland grazing studies - - - - - 213 
Schwaner, T. D. & Miller, B. Range extensions of reptiles in South Australia - - 215 
Schwaner, T. D. & Miller, B. Reptiles new to the fauna.of South Australia - - 217 
Fisher, R. H. Life history of the Narrow-Winged Pearl White El/odina 
padusa (Hewitsun) (Lepidoptera: Papilionoidea) - - - - B48 
Delean, S. & Harvey, C. Notes on the reproduction of Nephrurus deleani (Reptilia: 
Gekkonidae) - - - = - - - - ~ - 221 
Freiss, W. Y. Correlation of the uppermost late Precambrian succession across the 
Torrens Hinge Zone in the Port Augusta region of South Australia: 
a discussion - - - - - - - - - - 223 


Plummer, P. S. — Correlation of the uppermost late Precambrian succession across the 
Torrens Hinge Zone in the Port Augusta region of South Australia: 


a reply - - - - - - - - - - - 225 
McDonald, K. R. & Tyler, M. J. Evidence of gastric brooding in the Australian 

Jeptodactylid frog Rheobatrachus vitellinus. - - - - 226 
Belperio, A. P. & Murray-Wallace, C. V. Comment: Wardang Island—a refuge for 

Marginopora vertebralis?  - - - - - - - - 227 


PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS 
STATE LIBRARY BUILDING, NORTH TERRACE, ADELAIDE, S.A. 5000 


TRANSACTIONS OF THE 


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OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 108, PART 3 


THE EFFECTS OF SALINITY ON THE DISTRIBUTION OF AMPHIPODS 
IN THE COORONG, SOUTH AUSTRALIA, IN RELATION TO THEIR 
SALINITY TOLERANCE 


BY M. I. KANGAS & M. C. GEDDES 


Summary 


The tolerance of Melia zeylanica and Paracorophium sp. to a range of salinity and temperature 
combinations was investigated by LDsp analysis and response surface analysis. At the optimum 
temperature of 18-20°C, the salinity tolerance of both species was from 1 to 62%c; at high and low 
temperatures, tolerance to high salinity was reduced. Melita zeylancia, Paracorophium sp. and 
Megamphopus sp. were common in the Coorong, but all species generally were restricted to 
salinities below 53%c. No deleterious effects of salinity on the reproductive condition of populations 
were identified. 


THE EFFECTS OF SALINITY ON THE DISTRIBUTION OF AMPHIPODS 
IN THE COORONG, SOUTH AUSTRALIA, IN RELATION TO THEIR 
SALINITY TOLERANCE 


by M. I. KANGAS & M. C. GEDDES* 


Summary 


KANGAS, M. I. & Geppes, M. C. (1984) The effects of salinity on the distribution of amphipods in the 
Coorong, South Australia, in relation to their salinity tolerance. Trans. R. Soc. S. Aust, 108(3), 139-145, 13 


December, 1984. 


The tolerance of Melita zeylanica and Paracorophium sp. to a range of salinity and temperature com- 
binations was investigated by LD.) analysis and response surface analysis. At the optimum temperature 
of 18-20°C, the salinity tolerance of both species was from 1 to 62%o; at high and low temperatures, tolerance 
to high salinity was reduced. Melita zeylanica, Paracorophium sp. and Megamphopus sp. were common 
in the Coorong, but all species generally were restricted to salinities below 53%. No deleterious effects 
of salinity on the reproductive condition of populations were identified. 


Key Worps: Melita zeylanicu, Paracorophium, Megamphopus, salinity tolerance, amphipods, 


Coorong, South Australia. 


Introduction 


Most investigations of the salinity tolerance of 
estuarine and coastal marine amphipods have con- 
sidered tolerance to dilute conditions (Vlasblom & 
Bolier 1971; Dorgelo 1974, 1976; Ritz 1980). Few 
studies have been made on tolerance of amphipods 
to concentrations greater than seawater (McLusky 
1967; Marsden 1980), although amphipods are often 
important in hypermarine systems (Hedgpeth 1967). 
In the present study the salinity tolerance of M. 
zeylanica and Paracorophium at various 
temperatures were examined in the laboratory and 
the results related to the field distribution of the 
amphipod species in the Coorong lagoons. In the 
field study the relative abundance and reproduc- 
tive status of the amphipod species are investigated 
to look for possible sublethal effects of salinity and 
temperature on amphipod populations. 

The Coorong is a coastal lagoon system situated 
in the south east of South Australia (Fig. 1). The 
Coorong waters show a marked longitudinal sali- 
nity gradient which varies in direction and 
intensity seasonally and from year to year (Noye 
1975), In 1982, the Coorong lagoons were hyper- 
saline (Geddes & Butler, 1984), and this provided 
the opportunity to investigate the distribution of 
organisms along a hypermarine salinity gradient. 
Amphipods form a major part of the macroben- 
thic fauna and this study investigates the distribu- 
tion and salinity tolerance of three common 
species Melita zevlanica Stebbing (Melitidae), 
Paracorophium sp. (Corophiidae) and Megaim- 
phopus sp. (lsaedidae). Melita zeylanica is a 
cosmpolitan species, commonly found in 


* Zoology Department, University of Adelaide, Box 498 


G.P.O., Adelaide, S. Aust., 5001, 


estuarine systems (Croker 1971: Barnard 1972; 
Griffiths 1973; Krishnan & John 1974, 1975; Boltt 
1975) and has been recorded in Australia from the 
Peel-Inlet (Potter e¢ a/, 1981) and Lucky Bay, 


Western Australia (Barnard 1972), from the Tug- 
gerah Lakes, New South Wales (Collett ef al. 
1981), and from the Gippsland Lakes, Victoria 
(Poore 


LD 


1982), The other two amphipods are 


jg ae’; | 


é SOUTH AUSTRALIA. 


Fig. 1. The Coorong lagoons showing sampling localities 
(*) and sites where populations of M. zevlanica and 
Paracorophium sp. were collected for salinity tolerance 
experiments (*). 


140 M. I. KANGAS & M. C, GEDDES 


undescribed. The genus Paracorophium is 
endemic to the Southern hemisphere and found in 
Australia, New Zealand and South America 
(Barnard & Karaman 1983). Paracorophium sp. 
differs from P excavatum Chilton, the only 
described species known from Australia, in the 
structure of the third uropod and the number of 
setae on gnathopods 1 and 2 of the male. Very 
small numbers of a second undescribed species of 
Paracorophium were also collected. Neither 
species is similar to known Paracorophium species 
(Karaman 1979). No species of Megamphopus 
have been described from Australia (M. Drum- 
mond pers. comm.) but species have been found in 
abundance in the Tuggerah Lakes, N.S.W. (Collett 
et al. 1981) and the Gippsland Lakes in Victoria 
(M. Drummond, pers. comm.), Reference material 
of the species considered in this paper has been 
placed in the South Australian Museum, Adelaide 
(SAM C€3924-C3927). 


Materials and Methods 


(i) Salinity Tolerance 

Amphipods were collected in March, April and 
June 1982 from two localities (Fig, 1) at salinities 
of 51.3, 41.9 and 44%. respectively. 

Salinity tolerance was determined at five tem- 
peratures for hypersaline conditions (5.5, 14.4, 18.5, 
26, 32.5°C for M. zeylanica and 6, 14.4, 19.5, 26, 
30.5°C, for Paracorophium sp.) and three tem- 
peratures for dilute conditions (14, 18, 25.4°C for 
M. zeylanica 14.4, 19.5, 27.5°C for Paracorophium 
sp.). Amphipods were acclimated to test tem- 
peratures in 35% (seawater) for two days prior to 
experimentation. 

9-12 amphipods were directly transferred to five 
salinity dilutions (0,1-10.5% ) and ten hypersaline 
media (38.5-73.9%.). Dilute media were prepared 
by mixing seawater and distilled water; hypermarine 
media were mixtures of seawater and Coorong 
water. Conductivities (K2;) were measured with a 
Radiometer CDM2e Conductivity Meter and total 
dissolved solids (TDS) calculated by a regression 
provided in Williams (1966). This regression was 
developed for saline lake waters but comparison of 
dried TDS for samples from the Coorong with 
values calculated from conductivity via the Williams 
equation showed very close agreement (Geddes & 
Butler, 1984). This is to be expected considering the 
similar nature of ionic dominance in Coorong water 
and that of Australian salt lakes (Williams & 
Buckney 1976). The TDS values were used as a 
measure of salinity. 


At all salinity-temperature combinations, adult 
individuals were used without regard to sex. 


Gentle aeration and a light/dark regime of 12 
hour: 12 hour was maintained but no food was 
added. Fine debris and filamentous algae was sup- 
plied to Paracorophium sp. to enable it to con- 
struct tubes (considerable mortality was ex- 
perienced when this tube dwelling species was kept 
in clear water). The number of animals surviving 
were counted at 6 and 24 hours and every 24 hours 
thereafter for 96 hours. Following Ritz (1980), 
death was taken as a cessation of pleopodal 
rhythmic beating motor response to tactile 
stimulation. 

Data were analysed in two ways: determining 
LD,» values and fitting response surfaces to sur- 
vival data. For LD;) determination, the dose and 
response (% survival) values were transformed to 
log dose and logits (Hewlett & Plackett 1979) and 
regression equations calculated with the form Y 
=a + bX where Y is logit + 10. This form allows 
symmetrical confidence limits to be placed on LD.y 
values. 

The response surfaces were fitted according to 
a BMDP program Stepwise Logistic Regression 
(PLR) accessed via the Cyber 173 computer. The 
PLR estimates the vector of parameters (ji) for 
the linear logistic model E(s/n) = e8X/1 + ePX 
where s is the sum of the binary dependent 
variable (dead, alive) and X represents the in- 
dependent variables (salinity, temperature), The 
parameter fi may be expanded to a quadratic # y 
+ BX, + B aX, + BsX?, + BsX*, + AsXiX2 
where X, is the temperature, X; is the salinity, 6 o 
is aconstant, 8, is the linear effect of temperature, 
8; is the linear effect of salinity, @; is the 
quadratic effect of temperature, f; is the 
quadratic effect of salinity and A, is the interac- 
tion effect between temperature and salinity. Con- 
tour lines for specified percent survival were then 
plotted. 


(ii) Field Observations 

A series of offshore stations (Fig. 1) were sampled 
in the North Lagoon of the Coorong at approx- 
imately monthly intervals from January 
1982-March 1983, The northern end of the South 
Lagoon was sampled in August 1982 (Fig. 1). At 
each station, surface temperatures and water 
samples were taken. Amphipods were collected by 
towing a trawl net (160m) through algal beds for 
5-10 minutes. The samples were returned to the 
laboratory, sorted, preserved in 10% formalin and 
identified. One hundred randomly selected in- 
dividuals were identified to record species composi- 
tion and relative abundance; within each species, 
sex ratio, female reproductive condition, and egg 
number were noted. 


SALINITY EFFECTS ON COORONG AMPHIPODS ial 


Results 

(i) Salinity Tolerance 

Tables 1 and 2 summarise results found through 
logil analysis for Af, zevlanica and Puracorphium 
sp. respectively. The lower LD, value for AZ 
zevlunica and Paracorophium sp. is 1.0% and the 
upper LD. value for M. zevlanica is 62% and for 
Paracorophium sp. is 60.5%, indicating a wide 
tolerance range for each species. Due to the wide 
confidence limits, the respective values of LD, at 
each temperature trial overlap wilh the preceding 
and followilg values. General trends are that highest 
LD. values occur et medial temperatures while they 
decrease at both lower anc mgher temperatures. For 
M. zevlanica at 5,5°C the LD, value is 52%. and 
at 325°C is reduced to 49%. For Paracorophium 
sp. the LD,y value is 59% at 6.0°C and 48%6 al 
30.5 °C 

The contour patterns Jor salinity-temperature 
combinations are shown in Fig. 2. Both species 


show wide temperature and salinity tolerance with 
greater than 90" survival over most of the experi- 
mental range. The central region in (he contour 
pattern provides an estimate of optimum condi- 
tions (Alderice 1972). The close spacing of the 
contours indicates relatively low variability of 
response in the experimental animals. 

Temperature and salinity values and the relative 
abundance of species at stations |, 2, 3, 4 and 5 
(Stauions 1, 3, 5, 7 and 9 in Geddes & Butler, 1984) 
during the period December 1981-March 1983 are 
shown in Fig, 3. Low salinities in Dec, 1981, 
presumably the result of freshwater influx from 
Lake Alexandrina, were followed by an increase in 
salinity during the summer months, a lowering 
during April-June and an increase the following 
summer, At stations 4 and 5 highest recorded 
salinities were 68% and 82%o in January 1983. 
Surface temperature reached 27°C in summer and 
the minimum was 1"C in June. 


PABLE |. Relationship between logit + 10 of % mortality (x) and In K;. (v) and the calculated LD. values Jor M. 
zeylanica for high and low salinities and at various femperatures 


LD,, + 95% LD,, + 95% 
Temp. Regression Equation r confidence limits confidence limits 
(Ka) (Salimty: TDS) 
5.5e Iny = 3.896 + 0.0364 0.744 70.18 + 14,92 s1.Y 4+ 9.0 
I4.de ly = 4.382 4 0.006 x 0.010 71.6% + 75.60 53.3 + 56.9 
18.50 Iny = 4.111 * 0.025 x 0.731 78.49 > 16,29 61.8 4 ¥9 
26,0 Iny = 4.168 + 0.017 © 0.194 76.72 + 35,19 58.0 + 23,0 
32,5¢ lyy © 3.8L + 0.04 x 0.682 67.47 = 18.73 49.4 4 11.5 
14.0 lyy = 4.57 — 0.376% 0.947 2.25 + 6.03 134 35 
18.0d [yy 3,83 -— 0,329 ¥ 0.925 1.72 + 6,83 10+ 4,0 
260d Iny = 4.771 — 0.356% 0.834 3.36 * 28.90 1.9 + 184 


c—intermediate and concentrated media. 
d—dilure media 


Tabrt 2 


2, Relationship berween logit +10 of % mortality (x) and Inky, ()') and the caleulared LD. values for 


Parucorophium sp, for Wigh and low salinities and at various temperatures 


os ————————— ee ————— 


LD. + 95% LD,, + 95% 
Temp. Regression Equation re epntidence limits confidence limits 
(Ky) (Salinity: TDS) 
6,0¢ Iny = 2.867 + 0,049 x 0.518 78.33 = 25.81 59.5 + 16.3 
l4.de Iny 4.092 + 0.028 + 0597 79.44 + 24.6 60.5 — 15.4 
19,5¢ Iyy > 4,002 + 0.037 x 0.610 79.21 + 23.45 60.3 + 14.6 
26.0 Iny = 3.794 -+ 0.049 4 0.614 72.61 + 18.26 34} 4 11.2 
30.5 In 3.749 + 0.04d x O78 66.07 + 15.39 4AK2 +4 OF 
15.00 Iny = 4.401 O.476 x 0,536 0.82 + 33.17 O.3 + 21,8 
18.0d Inv = 4.136 UAT xX 0576 0.54 + 30,56 0.3 + 19,6 
275d Iny = 4.961 ~ 0,442 4 0.640 1,72 + 26.82 1.0 = 17.0 


c—inlernediate and concentrated media. 
d—dilvle media. 


142 M. 1. KANGAS & M. C. GEDDES 


0 
30 
30 
S) 
2 20 
wv 
2 
2 
a 
Ee 
we 10 
. 10 20 30 40 50 60 70 80 90 
Kp (mScm7') 
ie) 10 20 30 40 50 60 70 


TDS (%0) 


36 S50 


20 
10 
10 20 30 40 50 60 70 80 90 100 
-1 
Ky, (mScm™) 
oO 10 20 30 40 50 60 70 


TDS (%o) 


Fig. 2. Estimation of percent survival based on a fitted response surface to observed mortality at 96 hr. under 65 


conditions of temperature and salinity. 
(a) M. zevlanica (e'/1 + e' = —: 
(b) Paracorophium sp. (e'/1 + e! = 


M. zeylanica, Paracorophium and Megamphopus 
sp. were found at stations 1, 2 and 3 throughout 
the study period. Paracorophium sp, and Megam- 
phopus sp. occurred at station 4 in Jan. 
1982 at a salinity of 60%-, but seem to have suc- 
cumbed to the increasing salinities in the follow- 
ing months. In Nov. and Dec. 1982 
Paracorophium sp. reappeared after a period of 
lowered salinities. Melita zeylanica was collected 
from station 4 in May when salinity dropped to 
51%. Paracorophium sp. was the only species 
found at station 5 with 5 individuals collected in 
June. In the South Lagoon salinities were above 
80%. and no amphipods were found. 

All three species maintained large populations 
at stations Il, 2 and 3 throughout the study period. 
Paracorophium sp. generally had the greatest 
relative abundance and seasonal fluctuations in 
the abundance of Paracorophium sp. were small, 
M. zeyvlanica occurred in higher numbers from 
March-June with a lowering of numbers from 
Feb-April 1982 but their abundance was high 
from June-Nov. 

Table 3 documents the effects of salinity and 
season on the reproductive condition of females; 
it compares the mean percent ovigerous females 
and the mean egg number per ovigerous female 
for the different stations in “summer” (Jan.-April 
1982 and Nov. 1982-March 1983) and “winter” 
(June-Oct, 1982). 

All three species breed throughout the year with 


—4,.695 + .237s + .55It — .003s* — .019t* — .39Its). 
—2.183 + .18s — .002s° — 


OL6t*). 


similar numbers of ovigerous females being pre- 
sent during the summer and winter months. The 
percent ovigerous females does not show a consis- 
tent change between stations, although a substan- 
tial decrease occurs at station 4 for 
Paracorophium sp. The mean number of eggs per 
ovigerous female for M. zeylanica was similar bet- 
ween stations and seasons except for a low egg 
number at station 3 in summer. For 
Paracorophium sp. and Megamphopus sp. there 
were often significant differences in egg number 
between stations, with highest egg number 
generally recorded at station 2. 


Discussion 

A longitudinal gradient of increasing salinity 
persisted in the Coorong throughout 1982 with 
hypersaline conditions being maintained over 
most regions. In years of high River Murray flow 
the North Lagoon of the Coorong experiences 
marked lowering of salinity levels (Noye 1975) and 
so to persist in this region the fauna must be able 
to tolerate both estuarine and hypermarine condi- 
tions. This may limit species richness. In the event, 
amphipods in the Coorong form a_ simple 
assemblage with only three common species. In 
comparison, many estuaries have a much larger 
assemblage of amphipods (Gable & Croker 1978; 
Collet e¢ a/. 1981). Although Melita zeylanica, 
Paracorophium and Megamphopus are found in 


SALINITY EFFECTS ON COORONG AMPHIPODS 


Se 
0) a 
5p, <<< 50 


309 9" ~~ Salinity 30 
20| Temperature rae) 
1 ; 10 
OvF MAMS J SOND FM 
STATION 2 
V2, —— $a 


ie — a oe 


ee Pens 50 


= Irth . Laat 40 
304 a tai 30 
> | es 20 
fr 10 t—t—rot tI 8 
ro od FMAM YY AS OND UE M = 
o 2 
& [STATION 3} c 
2 = 
M , —_—e 2 
ra a 
ee 
‘ sp =r 
ie one, pO 
30) Set 40 
20 oo et 30 
yor + *20 
w 


(: <i .... 


ee oe 
Msp — 

Pspd 4 wg 80 

ne st OS ba 

att =n wee * 4 +60 

30 4an 39 

20 40 

190 30 


DvVFMAMJ I ASONDUEW 

hi, 3. Seasonal Muctuations in lemperature and saliniry 
and the relative abundance (%)) of MM) zeydaniew (M2) 
Pearaceroiun sp. (Paap) and Aves garmphops (Map) 
from Dee I98) March 1983. " indicates times when less 
than 20 individuals were found in samples. Psp.) fram 
stitivn 5 is represented by au open block, 


estuaries, they appear to be particularly well 
adapted to the extreme conditions met in coastal 
lagoon systems. Some or all ob these Uiree ame 
phipods are present in other Australian coastal 
lagoon systems as previously indicated, 
Laborarory studies on AY cevlanied and 
Paracorophian sp. show that they are euryther- 
Mal anceuryhaline with asalirtity toleranee range 


133 


al | 62%» ut the optimal temperature of 18°C, 
folerance al temperature extremes (5°C ancl 
2.5°C) was somewhat restricted bul salinities of 
So could be tolerated at all temperatures in- 
vestigated, A wide tolerance range is charalernstic 
ol estuarine or coastal lagoon species which ex- 
perience fluctuations in environmental conditious 
(MeLusky 1967, 1968; Jones 1972; Dorgelo 1976), 
and studies on the tolerance of two estuarine am- 
phipods, Orehestia chiliensis (Marsden (980) and 
Corophium volutatar (McLusky 1967) showed a 
tolerance of 3-3) %e ancl 2-S0%. respectively, The 
present study, and the field records of amphipods 
from 50-80% from the Laguna Madre in North 
America (Medgpeth 1967), sugwest that aeclima- 
tion in hypermarine environments produced 
hieher salinity tolerance than is normal for 
estuarine species, 


Maoasl work using response surface analysis has 
involved fish and larae decupods. None has invelyed 
umphipods. In the present study, response contour 
lines were more closely spaced than in studies on 
decapods and fish (Costlow er ad, 1962; Kinne 197); 
Aldetice 1972) indicating little individual variability 
in response. This may relate to the osmotic 
behaviour of the decapods and fish studied which 
are osmoregulators, while the amphipods in the pre- 
sent study are probably conformers in hypermurine 
water. There may be more individual variability in 
the response of regulators to salinity stress than for 
conformers. 

In 1982, amphipod distribution was not lunited 
by low salinity as is the case in most estuarine 
svstems (Meadows 1964; McLusky L968; Mills & 
Fish 1980), but the field distribution of all species 
wets limited by high salinities in the stations furs 
ther from the mouth of the lagoon, Under the 
rather stable salinity. pattern) which persisted 
throughout 1982, ne extensive changes in am- 
phipod distribution were seen. Cienerally aiii- 
phipods were restricted to salinities less (han 53% 
although sporadic records of a few individuals 
were made to sulinitics Up to 63%. These valucs 
are sumewlhiat lower thaw found in the laboratory 
tolerance studies. Field studies in conjunction 
with laboratory investigations provide informa- 
lion on other factors affecting distribution, One 
factor whieh may explain differences between 
laboratory results and field distributions is the lag 
Tune in reeraiiment of individuals into an area 
Which has only recently become favourable. It is 
possible that amphipods were absent trom 
southern stations in the winter months when 
salinities were apparently suitable because 
previous high summer salinities there had exceed- 
ed (olerunce limits. The young have direct develop- 


144 M. I. KANGAS & M. C, GEDDES 


TABLE 3. Comparison of percent females ovigerous and mean egg number per ovigerous female 

between different stations and different “seasons”: Figures for percent females ovigerous bas- 

ed on 10-100 females and for egg number on mean of 5-75 brood pouches. The * represents 
significant differences between mean egg number (Student’s t-test, P<0.05). 


M. zeylanica 


tig % Ovigerous Mean # Eggs 
Station “Summer”’ “Winter”? ““Summer”’ ““Winter’’ 
1 — 33.6 — 9.2 
2 79.7 §3,3 »{l4-6 12.3 
3 72.3 65.2 7.4 9.8 
Paracorophium sp. 
1 24.7 20. { 1.9 2.7 \, 
2 50.4 25.8 4.8 5.897 * 
3 32.8 35.9 { aah 4.9 
4 6.0 — 2.0 = 
Megamphopus sp. 
1 44.5 22.7 + 43 4.3 
2 27.5 28.7 +{ 7.9 5.5 
3 28.0 31.0 4.2 3.9 


ment and so there is no planktonic dispersal 
phase. Thus, in a system with seasonal and long- 
term fluctuations in salinity and with animals hav- 
ing poor dispersal abilities, it is possible that there 
is a time lag between the advent of suitable 
physico-chemical conditions in an area and the 
establishment of a viable population. 

There were no clear effects of salinity on the 
reproductive condition of populations. The propor- 
tion of ovigerous females and the mean egg number 
per female showed no seasonal change although 
summer salinities were considerably higher than 
those in winter. At stations | to 3 there was no major 
difference in percent ovigerous females but 
Paracorophium sp. from station 4 showed a mark- 
ed reduction in the percent ovigerous females. For 
all species there were some differences in mean egg 
number between stations, with stations 3 and 4 hav- 
ing lower egg number than station 2. This may in- 
dicate some lowering of reproductive capacity at 
higher salinities, but the evidence is not conclusive. 


The amphipods form a major part of the 
macrobenthic fauna of the Coorong, and are pro- 
bably important in food chains leading to fish and 
birds. High salinities in the Coorong results in nar 
rowing of the range of distribution and lowering 
of abundance of amphipods and this may have 
significant effects on animals further up the food 
chain. 


Acknowledgments 

We thank Julie Francis and Helen Vanderwoude 
for help with collecting and sorting amphipods, Dr 
Keith Walker for advice on data analysis and Phil 
Leppard for advice on computer analysis and in the 
design of programs. We are grateful to Prof. W. D. 
Williams and Dr Margaret Drummond for discus- 
sions on amphipod taxonomy and distribution. 
Thanks to Heather Kimber and Sandra Lawson for 
typing the manuscript and to Ruth Altmann for the 
art work. This study was supported by an M.S.T. 
grant which is gratefully acknowledged. 


References 


ALDERICE, D. F. (1972) Factor Combinations. Jn: O. 
Kinne (Ed.), ‘Marine Ecology”, Vol. I, Pt. 3, pp. 
1659-1772. (Wiley-Interscience: New York.) 


BarNarD, J. L. (1972) Gammaridean Amphipoda of 
Australia, Part I. Smithsonian Contributions to 
Zoology 103, 1-327. 

& KARAMAN, G. S. (1983) Australia as a major 

evolutionary centre for Amphipoda (Crustacea). Jn: 

J. L. Lowry (Ed.) Papers from the Conference on the 

Biology and Evolution of Crustacea. Australian 

Museum Memoir 18, 45-61. 


Co.uett, L. C., CoLuins, A. J., Gipps, P. J. & West, 
R. J. (1981) Shallow dredging as a strategy for the con- 
trol of sublittoral macrophytes: a case study in Tuggerah 
Lakes, New South Wales. Aust. J. Mar. Freshwater Res. 
32, 563-571. 

Bottt, R. E, (1975) The benthos of some southern 
African lakes. Pt. [V: The benthos of Lagoa Poelela. 
Trans. R. Soc. 8. Afr: 41, 273-282. 


CostLow, J. D., Bookout, C. G. & MONROE, R. (1962) 
Salinity-temperature effects on the larval development 
of the crab Panopeus herbstii Milne-Edwards, reared 
in the laboratory. Physiol. Zool. 35, 79-93. 

CrOKeER, R. A. (1971) A new species of Melita from the 
Marshall Islands, Micronesia. Pac. Sci. 25, 100-108. 

Day, J. H. (1979) The Estuarine Fauna. Jn: J. H. Day 
(Ed.), “Estuarine ecology with particular reference to 
Southern Africa”. (A. A. Balkema: Cape Town.) 

DorceE1o, J. (1974) Comparative ecophysiology of gam- 
marids from marine, brackish and freshwater habitats, 
exposed to the influence of salinity-temperature com- 
binations. 1. Effect on survival. Hydrobiol. Bull. 8, 
90-108. 

—(1976) Salt tolerance in Crustacea and the influence 
of temperature upon it, Biol. Rey, 51, 255-290. 

GABLE, M. F. & CRoKER, R. A. (1978) The saltmarsh 
amphipod Gammarus palustris Bousfield at the nor- 
thern limit of its distribution. Estuar. cst/. mar. Sci. 6, 
225-230. 


SALINITY EFFECTS ON COORONG AMPHIPODS 145 


Geppes, M. C. & Burter, A. J. (1984) Physicochemical 
and biological studies on the Coorong Lagoons, South 
Australia, and the effect of salinity on the distribution 
of the macrobenthos. Trans. R, Soe 8, Aust. 108(1), 
51-62. 

Gruriuiis, C. L. (1973) The amphipods of Sourhern 
Africa. Part 1. The Gammaridae and Caprellidae of 
southern Mocambique. 47a. S. Afr, Mus. 60, 265-306 

HeporetH, J, W. (1967) Ecological aspects of the Laguna 
Madre, a hypersaline estuary, Ja: G, H. Lauff (Ed.), 
“Estuaries", pp. 408-419, Am. Assoc. Advan, Sei, Publ. 
No. 83, 

Hewrerr, PS. & PLAcKeIT, R. L. (1979) The Interpreta- 
tion of Quantal Responses in Biology, Ch. 3 (Edward 
Arnold: London.) 


Jowes, M. B. (1972) Effects of salinity on the survival of 
the Jaeru alhifons Leach group of species (Crustacea; 
Isopoda). 2 exp. mar. Biol. Beal. 9, 231-237, 


KARAMAN, G, S. (1979) Revision of the genus Paracoro- 
phium Stebb with description of # chelatum, n. sp, and 
genus Chaelocorophium, no, gen. (Fam. Corophiidae) 
(contribution to the knowledge of the amphipoda 100), 
Prir, Glas. Republ. Zavoda Zast. Pri. Muz. Titogradu, 
12, 87-100, 


Kune, ©. (1971) Salinity: Animals—-Invertebrates. J; O. 
Kine (Ed.), “Marine Ecology”. Vol. 1, Environmental 
Factors, Pt, 1, pp. 821-995, (John Riley: London.) 


KRISHNAN, L. & Joun, P. A. (1974) Observations on the 
breeding biology of melita zevlanica Stebbing, a 
brackish water amphipod. Hydrabiologia 44, 413-430, 

— & (1975) Breeding biology of (he amphipod 
Melita zeylanica in a tropical monsoonal lake. Zool, 
Anz. 194, 328-334. 


Magspen, L. D. (1980) Effects Of constant and cyclic 
temperances on the salinity tolerance of the estuarine 
sandhopper Orchestia chiliensis, Mar, Biol, 59, 211-216, 


McLusky, D. S. (1967) Some effects on the survival, 
moulting and growth of Corophiumn volutator, - Mar. 
biol. Ass, UK. 47, 607-617, 

——(1968) Some effects of salinity on the distribution and 
abundance of Corophium volutatar in the Ythan 
Estuary. /bid, 47, 607-617 

Meapows, R. S. (1964) Experiments on substrate selec- 
tion by Coruphium species: films and bactena on sand 
particles. J. exp. Biol, 41, 499-511. 

Mitts, A. & Fisu, J. D. (1980) Effects of salinity and 
temperature On Corophium volutator and C. arenarium 
with particular reference to distribution. Marine Biol. 
58, 153-161. 

Nove, B. J. (1975) “The Coorong—past, present, future”. 
Publ. No. 39.—1974 revised. (Department of Adult 
Education, University of Adelaide: Adelaide.) 


Poore, G, C, (1982) Benthic communities of the Gipps- 


land Lakes, Victoria. Aust. J. Mar Freshw. Res, 33, 
901-915, 

Porver, |. C., Lenanton, R. C. J., LONERGAN, N., 
CrHerystat, B., Caputi, N. & GRANT, C. (1981) “The 
Fish and Crab Fauna of the Peel-Harvey Estuarine 
System in Relation to the Presence of Cladophora", 
Bulletin No. 88 (Department of Conservation and En- 
vironment: Western Australia). 

Rrry, D. A. (1980) Tolerance of intertidal amphipods to 
Fluctuating conditions of salinity, O; and copper in 
Gammarus duebeni and Marinogammarus marinus. J. 
Mar. Biol. Ass. UK. 60, 489-498. 

Viaseiom, A. G. & BoLicr, G. (1971) Tolerance of em- 
bryos of Marinegammarus marinus and Orchestia 
gamarella to lower salinities. Neth, J. Sea. Res. 5, 
334-341, 

Witiiams, W. D. (1966) Conductivity and (he concentra- 
tion of total dissolved solids in Australian lakes. Aust. 
J. Mar. Freshwater Res. 17, 169-176. 

— & Bucknty, R. T. (1976) Chemical composition of 
some inland surface Waters in south, western and 
northern Australia, (bid, 27, 379-397. 


A NEW SPECIES OF CALAMOECIA (COPEPODA: CALANOIDA) FROM 
SOUTH AUSTRALIA, AND COMMENTS ON THREE CONGENERS 


BY TI, A. E. BAYLY 


Summary 


Calamoecia zeidleri sp. nov., a comparatively large species of Calamoecia, is described from fresh 
waters near Lake Eyre and Oodnadatta. 


A NEW SPECIES OF CALAMOECIA (COPEPODA; CALANOIDA) FROM SOUTH 
AUSTRALIA, AND COMMENTS ON THREE CONGENERS 


by L 


N\, EB, BayLy* 


Summary 


BAY LY |, ASB, WOR4) A yiew species of Co/amoeeta (Copepoda: Calanoida) from South Australia, and com- 
mens on three congeners. Trams, Ro Soe 3. Anse. WOK(9), 147-154, 13 December, 1984. 


Caluinoecia cedlert spenoy., a comparatively large species of Calanecia, ts described (Om Mesh waters 


nea Lake Eyre and Qudnadiasta. 


Two Western Australian populations of ©. /vcasi, which have diverged markedly boul structurally and 
ccologically from populations inthe eastern hall of Australiaaind in New Zealand, are described in detail, 
Both populations have an abnormally large body size for (his species, and (he cluleh size of the females 
of one is unusually high, A palaeocliniatological explanation for the subspecifie divergence of Western 
Ausvaliah populations of C. dueayivand C. g/fboxe trom those in the east is presented, 

New information is presented on the disiibucion of ©. canberru, 


Kis Wokbs! Copepoda, Calahoida, Colwnmoccia, fresh water. 


Jntroduetion 


The genus Culameecia, which contains small 
non-marine calanoids, was revised by Bayly (1961, 
1962). A. further species was added (Bayly 1979) to 
bring the total number of deseribed species to 13, 

During 1Y8t and 1982 | examined a series of 80 
collections of zooplankion made by Mr Wolfgang 
Zeidler of the South Australian Museum (SAM) 
rom inland waters of South Ausiraha and the 
Northern Territory, Included amongst this material 
were five collections from the northern part of S.A, 
(to the north of Oodnadalla and west of Lake Eyre) 
which contained a bighly distinctive undescribed 
species of Culamoecia, This is deseribed below, 

Additionally, two isolated and peculiar popula- 
lions al © lucas? Brady sampled during the field 
work associated with the paper of Geddes er al, 
(1981) on saline lakes in Western Australia (bul not 
recorded in that work because of their oveurrenee 
in fresh waters) and passed on to me are deseribed. 

Possible reasons tor the east-west divergence hin 
the morphology of C, dveasiund ©) eibbosa are 
discussed, 

Kinally, new information is presented on the 
distribution Gl C. canberra Bayly hitherio known 
from few localities but Which occurred in LS of rhe 
Zeidler collections. 

Although (wo species of Culunteecia occur in 
saline walers, und saline waters are comiNon in those 
eeneral regions of Australia referred to ii) this paper, 
wll Culumoeciv material discussed below came from 
fresh waters, 


+ Zoolowy Department, Monash University, Claynon, 
Vin 3168 


Calamoccia zeidleri sp.nov. 
FIGS |-2 

Type Material: Holotype a, allotype?, paratypes 30a, 40 
g (trom swamp 29°57'S., 136°14'E) of Billa Kalina Hostd; 
holutype and allorype stained with Chlorazol Black, 
dissected and mounted in balsam on mucroslides, puralypes 
preserved in formalin, inmounted in vial; SAM C. 3961-7, 
’uralypes from dam rir William Creek (28°55'S,, 
136°20'E.) 304, 308, unmowited in formalinin vial, SAM 
C. W969-70), 


Descripiion of Male: 

Size. (a) Swamp tir Billa Kalina Hota mean (n 
10) length to end of uropods (formerly Furcal paint) 
110mm, (b) Dam 16 km N, Willian Creeks mean 
length as above 0.99 mm, 


Lith legs (Figs 1A-1B). Right exopod with com 
paratively short proximal segment, middle segment 
with tooth on inner edge slightly proximal of mic 
polit and second tooth on outer distal cde near 
point of insertion of seta on posterior face (My, 
LA), distal claw strongly bent inwaids trough 
approximate right-angle (as inC, g/bbosa) then cur- 
Ving outwards towards distal extremity, lackinw 
sccondary Spur (present in seven other species oF 
Cilamoecia) on inner proximal edge of claw; right 
cndopod 2-segmented, proximal segment only 
aboul \4 length distal seement, distal segment with 
highly distinctive thumb-like spur arising ot outer 
distal corner and orientated at right-angles (o long 
axis ol sexment, with two long spines al distil ex- 
tremity, that next to “thumb” (» “indes finger) 
strongly curved neur base, minute spine oceasionally 
present at inner distal corner near base of inner 
distal spine (Fig, 1A); left exopod 2-sezmented on 
anterior face (Fig. 1B) but line of segmentanon 
jurvely Obseuréd of posterior face (Vig. LA), distal 


148 I. A. E, BAYLY 


Fig. 1. Calamoecia zeidleri sp.nov, A and B, ¢' fifth legs, 
showing posterior and anterior aspects, respectively; C, 
2 fifth leg; D and E, lateral aspect of 2 urosome show- 
ing, respectively, the ventral bulge, or genital operculum, 
closed and open; F, ? urosome, ventral aspect. 


segment with conspicuous seta inserted short of ex- 
tremity on anterior face and with elongate concavity 
on posterior face near inner edge; left endopod 
l-segmented expanded distally and typically with 
five spines—two (1 long, 1 short) terminal, two sub- 
terminal, and one on outer edge % of total length 
of segment from distal extremity. 


Description of Female: 
Size. (a) Swamp nr Billa Kalina Hstd: mean (n = 
10) length to end of uropods 1.42 mm. (b) Dam 16 


km N. William Creek: mean length as above 1.26 
mm. 


Fifth legs (Fig. 1C). Terminal exopod segment with 
five spines, largest or terminal spine only slightly 
longer (ca 1.3x) than segment itself (compare with 
most species of Calamoecia in which terminal spine 
> 2x length segment); endopod 1I-segmented bear- 
ing eight (or occasionally seven) setae, seta 
immediately to inside of terminal seta very short 
and spine-like. 


Genital segment (Figs 1D-1F). No lateral 
outgrowths (Fig. 1F) as in C. gibbosa, C. clitellata 
and W.A, forms of C. /ucasi, genital operculum with 
distinct posterior ‘nipple’ as in C. /ucasi, C. 
australica and C. canberra. 


Remarks: This species is easily recognised by the 
large outer distal spur on the right endopod of the 
fifth legs in the male, and by the relatively short 
terminal spine on the terminal exopod segment of 
the fifth legs in the female. In the latter feature on- 
ly C. salina, in which the terminal spine is about 
1.6x the length of the segment bearing it, ap- 
proaches C. zeidleri. In C. salina, however, the ter- 
minal exopod segment of the female fifth legs bears 
only two spines (cf five in C. zeidleri). 

The body size of the female of this species is 
relatively large for Calamoecia and may be com- 
pared with that of the female of C. attenuata. 

C. zeidleri coexisted with C. canberra Bayly at 
all five localities and also with Boeckella triar- 
ticulata (Thomson) at three of the five localities. 
The size relationships existing for one situation in 
which C. zeidleri was one of three coexisting 
calanoids, and another in which it was one of two, 
are shown in Table 1. There was no overlap in the 
mean lengths of the adults of different species. 

Congeneric occurrences are not common for 
Calamoecia in Australasia as a whole (cf. Bayly & 


TABLE 1. Size relationships of coexisting calanoids. 


Swamp near Dam 16 km N. 
: Billa Kalina H.S. William Creek 
SRR and ® (n=10) xX @ length X (n=10) x ? length 
length (mm) x dlength length (mm) x ¢ length 
Boeckella triarticulata 
(Thomson) 1.17 
female 1.81 
male 
Calamoecia zeidleri sp.nov. 1.29 1.27 
female 1.42 1,26 
male 1.10 0.99 
C. canberra Bayly 1.16 1.13 
female 0.88 0.77 
male 0.76 0.68 


A NEW SPECIES OF CALAMOECIA 149 


Willams 1973, table 6:3), However, they are not un- 
common in the far south-west of W.A. where C 
atfenuala may coexist with a smaller Calamoecia 
such as CL fasmanica or C. elongata. 

Table | shows that the ratio (mean female 
length):(mean male length) tor C zeidleri is relative- 
ly high (1.27-1.29) for Calamoecia (cf. Bayly 1978, 
table |, group C), 


Material Examined: S.A.: Swamp (Devils Playground) 
6km 5.6. of Billa Kalina Hstd (29°55'S., 196°H’E.), 450, 
409, 5.xi,1974; dam 16 km N. of William Creek (28°55'S., 
136°20'E.),45¢, 409, May 1976; dam 35 km N. of 
William Creek, If, May 1976; Alberga Creek road cross- 
ing 47 km NNW. of Oodnadatta, Le, 3.v.1976; waterhole 
5 km N. of Mt Sarah (26°55'S:, 135°20'E.), 2¢, 4.1976: 
all five coll, W, Zeidler. The distriburion is shown in Fix. 2. 


T 
\, \4H°E 


aI 


South 4usvala H 


‘kQodnadatta 


Ch eee 
sols ‘ <n an 
} vw | \" aie 
le 
f Lake = [3 
—~~—s«&yre \ 
(4 
\ . { t 
_ ‘\ ho AW 
“MS ® \ Le 
bind N te! \ 5 \ \, 7 


“‘, J L 
J ott 
y am wy 
n sa { Dong 
Kill 
A, 
fee's et iaT°E 30°s 


Fig. 2. Distribution of Culameecia zeidlert sp.nov. 


Isolated Western Australian populations of 
Calamoecia ana their marked morphological 
divergence 
Cu/atmeoecia fucust Brady 
As shown by Bayly & Williams (1973, Fig, 6:3), 
and as indicaled in Fig. 3, most Australian popula- 


~ oh . $ 
; @ } 


Fig. 3. The main, eastern areas of Australia occupied by 
Calamoecia lucasi and C, gibbosa and the isolated W.A. 
populations of these species. The arrows indicate ex- 
tensions to previously known distributions—not direc- 
tions of dispersal. 


tions of C. /ucasi are restricted to the eastern half 
of the continent (the species also occurs in the 
North Island of New Zealand), However, the ex- 
istence of some isolated populations in what are 
almost certainly temporary waters in arid regions 
of W.A, is now known. These W.A. populations 
have diverged remarkably, both morphologically 
and ecologically, from those in the eastern half of 
Australia and N.Z. The morphological divergence 
is evident with respect to both body size, which is 
much larger, and the details of secondary sexual 
characteristics. If one of these W.A. populations 
Was transported to N.Z., | doubt if it would be im- 
mediately recognised as C. /ucasi when first en- 
countered there. The possibility exists that breeding 
experiments would justify the W.A. form being 
treated as a separate species. However, 1 consider 
the aberrant W.A. populations are properly 
referable lo C /ucasi. 


(a) The Cue Population 
FIGS 4A-D 
Material Examined: W.A.: 20%, 104 pond close to Nallau 


(27°16'S, 117°59'E) 21 km N.N,E. of Cue, coll. M,C. 
Geddes et al. viii 1978, 


Body Size (mean prosome length). Female, 0.96 mm 
(n = 10); male 0.86 mm (n = 10). 


Male Fifth Legs (Pigs. 4A and 4B). These differ 
from those of eastern populations as follows; 
(1) the proximal segment of the right exopod has 
no projection at the inner distal corner (compare 
Figs. 6A and 6B) 


150 I. A. E. BAYLY 


Fig. 4. Calamoecia lucasi Brady from population near 
Cue, W.A. A and B, ¢ fifth legs, showing posterior and 
anterior aspects, respectively; C, ? fifth leg; D, ? genital 
segment (and extensions of last prosomal segment), dor- 
sal aspect. 


(2) there is a strong projection on the inner edge 
of the middle segment of the right exopod which 
is not seen in eastern populations 

(3) the distal segment, or terminal hook, of the 
right exopod is more strongly bent 

(4) the middle segment of the right endopod is 
enlarged so as to present a semicircular outer edge 
(5) the left endopod invariably has an armature of 
5 spines (2 terminal, 3 sub-terminal) instead of the 
usual four spines; however, variation in spine 
number from two-five has already been 
documented (Bayly 1961) 

(6) there are quite strongly developed denticles at 
or near the distal extremity of the left exopod 


Female Fifth Legs (Fig. 4C). The distal exopod seg- 
ment differs from that of eastern populations in 
bearing six spines instead of the usual five. 


Female Genital Segment (Fig. 4D). This differs from 
that of eastern populations in having a more pro- 
nounced lateral bulge on the left side (compare Figs 
6E and 6F). 


Clutch Size. The mean number of eggs was an 
unusually (for this species) high 44 (Table 2). 


Remarks: C. lucasi was the sole calanoid present 
in the zooplankton collection from this site which 
had a maximum depth of about a metre, a very high 
turbidity, and a T.D.S. value of 41 mg/l. The tem- 
porary nature of the pond was emphasised by the 
presence in the collection of an abundance of con- 
chostracans. Also present were ostracods, 
cyclopoids, Chydorus and Keratella. 


(b) The Population Near Lake Grace 
FIGS 5A-5F 


Material Examined: W.A.: 10 ¢, 102, roadside pool on 
northern side of road, 3.5 km W. of Lake Grace township, 
coll. M. C. Geddes ef al, viii.1978. 

Body Size (mean prosome length). Female, 1.00 mm 
10); male, 0.93 mm (n = 10). 


(n = 


Male Fifth Legs (Figs 5A-5C). These differ from 
those of eastern populations as follows: 

(1) the proximal segment of the right exopod has 
a more strongly developed projection at the inner 
distal corner 

(2) the distal segment, or terminal claw, of the right 
exopod is more strongly bent, as for the Cue 
population 

(3) the terminal segment of the right endopod 
typically (Figs 5A and 5B) has one or two greatly 
reduced, or only vestigial, setae, but occasionally 
(Fig. 5C) a longer seta is present 

(4) the left leg has the same peculiarities as describ- 
ed above for the Cue population 


Female Fifth Legs (Fig. 5D). These have the same 
peculiarity as detailed above for the Cue 
population. 


Female Genital Segment (Figs 5E and SF), This is 
distinctive in being essentially similar to that 
described above for the Cue population although 
the left lateral outgrowth is even more pronounced. 


Remarks: Two other calanoid species, Boeckella 
opaqua Fairbridge and B. robusta maxima Sars, 
were also present in the collection examined. Both 
of these species are characteristic of shallow, tem- 
porary waters. A T.D.S. value of 980 mg/1 was ob- 
tained for a water sample taken from the pool. 


(c) C. lucasi from New Zealand 
FIGS 6A-6F 


Drawings of material collected by the author 
from Lake Alice (40°08 'S, 175°20‘E) near Marton, 


A NEW SPECIES OF CAL AMOECTA 


151 


TAtib 2. Length aad clutch size of Catanioecia lucasi Jemales. 


X prosome 


Nature and location of population length Nu: Peinales Slaveth Hee Cost var. 
(mm) examined 90. BBS (a) 

(A) Hoa. seasonal leniporury-wyler 

populations 

Poot near Lake Grace? VW) 

Pond near Cue 0.96 x 44.1 12 
(Hh) NA. perennial lacustrine populations 

Lake Ololou O.57¢ 18-19 

Lake Rotorua 0.65* 3.4 Alig 

Lake Roti 0.645 2.0€ 27 

Nowell’s Laguwon! ~ 25 13,8 2) 


1 Length data froty 1U individuals measured along a mid-dorsal line and omitting the well developed, 
posteriorly praycetihe “wings” On the fast segment of [e'prasome, 


* No ovigerous females present 


© From Green (1976, table 5), The data represent oud) wieans obtained from (he measuremens of a laroe 
number of individuals rom vavh of a substantial series oF samples. 


“rom Bayly (1961, table 2) 


© From Chapman (1973, table 3). Mean dara front a large number of individuals collected over a bwo- bu 


Threeevedr peru 


N.Z., are included for comparison with the W.A,. 
populations. 


Discussion 


As shown in Table 2, individuals from these two 
desert populations of C. /weusi are 50% or more 
(up to 75%) larger than those belonging to N,Z, 
populations, This probably underestimates the size 
discrepancy because the prosome measurements of 
the N.Z, specimens apparently include the postero- 
lateral “wings” of the last prosomal segment. The 
type of measurement specified in Table 2 for the 
W.A. specimens although slower is preferable 
because Of intraspecific variation in the relative 
degree of development of these wings. 

Gigantism in calanoids in Australian desert pools 
ix noted by Mitchell (1984) who referred to 
BoeckKella triarticulata reaching a length of up to 
3.2 mm in a temporary pool near Lake Eyre. 
However, Mitchell’s explanation, “Organisms in 
these localities often attain very large sizes due to 
rapid growth rates [my emphaisis]” seems invalid: 
in planktonic crustaceans large adult body size ts 
associated with long development ume (slow 
growth) aod both of these correlate with low 
temperature alone if food is sufficiently abundant 
(MeLaren 1963), 

The large clutch size found for the Cue popula- 
lion (Table 2) is in accordance with the principle 
(Belk & Cole 1975) that where a calanoid species 


occurs in both permanent and temporary waters, 
populations in temporary waters typically have a 
larger Clutch sizé thah those from permanent waters. 
A larger clutch size also would be expected in (his 
instance because a positive correlation between clutch 
size and female body size generally applies within 
the Copepoda (McLaren 1963). It may be noted, 
however, that in Boeckella symmetrica an increased 
clutch size in temporary waters (Bayly 1979) does not 
appear to be accompanied by the striking gigantism 
reported here for C /ucasi, 


Typically, freshwater species of Culwroecta occur 
in permanent waters (Bayly 1978). The chief excep- 
lions are the W.A. species, C attenuata and C 
elongata, W.A, populations of C ampulla, and © 
canberra, all of which occur net uncomnionly in teni- 
porary waters even if they also occur in permanent 
ones. Maly (1984) confirms that, considering the 
genus Calamoecia as a whole, it is much less 
common than Boeckella in temporary pools. Timms 
(1970, table 12) assessed C. /ucasi as having poorer 
powers of dispersal in north-eastern N.S\W, than 
three species of Boeckella that occurred in the same 
area. Additionally, C /ucasi seems not to have been 
recorded from temporary waters in N.%. Despite 
these generalizations concerning the genus Cala- 
moecia as a whole, and C /ucasi in particular, at least 
two W.A, populations of this species undoubledly 
are adapted for habitat ephemerality. 


152 I. A. E. BAYLY 


Fig. 5. Calamoecia lucasi Brady from population near 
Lake Grace, W.A. A and B, @ fifth legs, showing 
posterior and anterior aspects, respectively; C, portion 
of ¢ right fifth leg, showing endopod with (for this 
population) unusually long terminal seta; D, 9 fifth leg; 
E and F, ventral and dorsal aspects, respectively, of % 
eerital segment, showing pronounced outgrowth on left 
side. 

Should the W.A. populations be regarded as relic- 
tual in character or relatively recent derivatives from 
the east? Structural evidence favours the former view; 
the W.A. populations may be regarded as being more 
primitive in having a less reduced armature on the 
fifth legs of both sexes (the armature of the male 
right fifth endopod of the Lake Grace population 
excepted). The relatively poor dispersal ability of 
Calamoecia (Maly 1984), combined with the fact that 
westerly or south-westerly winds predominate 
throughout much of the southern half of Australia, 
would tend also to favour transport from west to east 
over the reverse. 


Calamoecia gibbosa Brehm 


A parallel situation exists for this species as for 
C. lucasi (Fig. 3). For many years C. gibbosa was 


known only from south-eastern Australia. It was first 
described in 1950 from Lake Dulverton in Tasmania. 
Two further Tasmanian records and one from 
Flinders Island were added by Bayly (1964), and three 
mainland records (all lakes at or near Mt Gambier) 
were added by Bayly & Williams (1964). Two further 
unpublished records (a fourth Tasmanian locality and 
a second one on Flinders Island), making nine in 
all, were known at the time of preparation of the 
map presented by Bayly & Williams (1973) for C 
gibbosa showing it restricted to south-eastern 
Australia. However, in 1977 an isolated population 
was found at Newmann’s Rocks in W.A. (Fig. 3) and 
described by Bayly (1979) as a new subspecies, C. 
gibbosa newmannensis. 

A previously unpublished record of C. gibbosa gib- 
bosa (incorporated into Fig. 3) is that from Fresh 
Dip Lake between Beachport and Robe at 37°16'S., 
139°49’E, (collection 1,xi.1979). 


General Discussion of Western Australian Forms of 
C. lucasi and C. gibbosa 


The situation described above for C. /ucasi and 
C. gibbosa is not unlike that recognized by Bayly 
(1961) for C. tasmanica (Smith), with C. fasmanica 
tasmanica in the east, and C. tasmanica subattenuata 
in the west [the position with C ftasmanica is, 


Fig. 6. C. ducasi Brady from Lake Alice near Marton, New 
Zealand. A and B, ¢ fifth legs, showing posterior and 
anterior aspects, respectively; C, ? fifth leg; D, terminal 
exopod segment of fifth leg enlarged; E and F, @ 
genital segment, dorsal aspect (different individuals and 
orientations). 


A NEW SPECIES OF Coll oh Went has (s3 


however, thore complex than orjwinally supposed 
(Bayly (979)), What explanation can be alfered for 
the subspecific divergence af WA. populunous of 
© lucayeand Co gibbase (and C2 tasmenice) (rom 
those in the caster hall’ of Australia? 


In the carly Miocene, 20 inhon years azu, the 
environment yn ie southern coast of Australia was 
subjeet to high humidity that penetrated far ince 
the continent (Bowler 1982), There were extensive 
freshwater lakes jn (he interior where mun salt lakes 
doininale, Despite a summer mesinum i the rave 
fall, Bowler (1982) considered that even in winter 
surplus moisture prevailed right across the conti- 
Hen and inland foges conditlons were common, 
With such a cline i might he supposed thal 
populations of freshwater ealanvids suel us € 
lucasiund C, gibhosa exvended freely across the con- 
tinent from east to west (except (hal marine |run- 
seressions into the Bucta and Murray basis would 
have interrupted the continuum along the southern 
border). Subsequently, however, the development af 
wl intense zone of aridity in the Nullarbor region 
and |ts northward extension seems likely to have 
splil The east-west ontinuunr lito two segments, the 
caster being somewhat larger than the western one 
In lute Miocene times, six million years aga, there 
was intense seasonal aridity (winters were now ery) 
across southern Ausiraia reaching a maximuin in 
the Nullarbor region. In the late Pliocene, 2,5 
million years agu, (he present climatic zonation of 
Australia developed for the frst time, and by one 
Million years wee central Australia was already dry 
without nevessarily being as arid as subsequently 
(Bowlet 1982).. Huwever, there was 4 major phase 
during the late Pleistocene frotty 30-50 QO) years 
BP. the Mungo lacustrine phase, of lake expan- 
sion and (allowing tor a reversal of seasonality in 
precipilation) uw return alriost 1 the conditions 
described for the carly Miocene, 

The W.A, populations of C. diteasi and & gih- 
bosa may be regarded os reliclual, and a product 
of geosraphical isolation by arid nortti-south ¢lissee- 
Hoy Of a previgus east-wes! conlinuum, Bulowtich 
Ol (he arid dissections was the operative ore’? be 
the absence of w fossil record we wae prescnely say 
Vinle coneerning paces of evelution in. caljureid 
eopepods. However, the face (hat we are dealing 
With Only subspecific levels of differentiation would 
tend to suppest (hal an iiterruplion to gene Mow 
Oveurred i the late Plemtocene rather than al some 
earlier link Tis reasonable lo suggest, theretone, 
(hal (he relevant lisseclion post-dared the 30-30 G00 
years BP Mingo lucusoine phase referred to hy 
Bowler (1982), bur mut the pened of maxi, 
ariuiry 1 OOO years BAL 


The question still remains ts to why populations 
of, (ueasi and C. sibbosa are ot wow found in 
(he wet Far soullrwes! corner oF WoA, (say to the 
south-west of a straight tine frou Buseelron to 
Albany). One can only suppose that, alihough these 
species hud almost continuous and extensive cast 
west distributions prior lo dissection by an and cor- 
ridor through the Nullarbor region, they did oot 
extend ro the extreme south-west of W.A,, ane have 
been Unable to achieve dispersal there since. 

I may be jolted thal the population of ©. lacasi 
near Cue inhabited a body of water that way pro- 
hably at least parcially ol wuny-madeé origins field 
notes stated that the depression was “likely to have 
been artificially deepened’. 7 he man-made nature 
Of the povd Occupied by OC. 2xibbosa al Newmann's 
Rocks was emphasised by Bayly (N74). Poputa- 
tions of C fueass and C. elfbose i the desert 
roviors.of WA, must have been very sparse if recent 
tijies before the adven!| of European man, and il 
is possible ihat man-made excavations have allow- 
ed significanr expansion of populations this century. 

No alleriative iiferpretation to that presented 
above is rhiatiie WA, populations of CG fceusi amd 
C, sibbosa represent recent penetrations fram ite 
east, such movement perhaps being favoured by 
anthroapogeue moditicauon of desert habitats, This, 
hawever, apparencly ruins counter to the mor- 
Pholopiwal evidence wi the case of Co lerasih 


Burhan af, canberra Bayly 


The triangular dist bution shows for €. curberra 
by Bayly & Williams (1973, liz. (4) was based on- 
ly on five records; the tap left apex was for two 
dams elase to.Alice Springs, the top right apex was 
lor two lakes (Barcoorah and Dujol near Araynae, 
and the bollom apes was lor the Lype locality, Like 
George, near Canberra. New teeords, summarised 
wd combincd with the alder ones i Pigs 7, areas 
follawse 

Sede Kilely dim oor Paring (Od 'S 28°17), 
TT U7 wunterlolenr Dalkuninn (9001'S. Ss 281.) 
Boslsville Track, PsiilO74: Cooper Crock erdssing i 
braduimia HS, (28 48'S.. 1B8°S8'E.) Birdsville Troek. 
Lei l874o din aot lL Plribbs (29°32'S.. a7 -ny bo) an 
romd ia Siuart Creek Sutin Ti i974: swamp (oeuts 
Playyroundy 6 key SE of Billa Kalina HS (29°55), 
{3h V1 'E.), Sx t974; Berestord Dani & of Willigng Creek 
(28°55 5 IH 2b), 6 x AYTS Paradise Dam 19 km. 
ab Williaa Crovk, Txii 19745 dan 1 ke Not Willie 
(Seok, y 976; dim 3S ke Not Witham Creek, «197: 
Albertus Creek road crossine 44 bin AN W oul 
Mocwadaita, 2V197A: waterhole § kim Nooat Mb Surah 
(20°55 'S,, 195720 b i dv t8F6: all dh coll, Wo Aeidlen 
Herestord railway dean (2974'S (aae8o by, 1978, coll. 
B19. MitelicIL Dain tr Carrietan (32°26'S., (AX 32'E 
lO.xii 197, voll, ML. Geddes. Old: TL, Kuelivow 
14555, 5995S TGS kis of Bedourie, [Ki 1897; 
Loughe) watertole {22°46/S, 12K SPE) betwecon 
Colohorinanol and Roxhonvuell Downs sudions, 20.ie 1477, 


154 I. A. E. BAYLY 


both coll. W. Zeidler. N.SW.: Dam 2 km from Wanaar- 
ing (29°42'S., 144°09'E.), 1.1969, coll. W. D. Williams er 
al. Pond 16 km SM. of Narrandera (34°45'S., 146°33'E.), 
10.v.1982, coll. E. J. Maly. N‘7.; Waterhole under McGrath 
Creek bridge 47 km N. of Alice Springs (23°19'S., 
133°47’E.), 20.iv.1979; roadside ditch 7 km N. of Stirling 
(21°44'S., 133°46’E.), 20.iv.1979; both coll. D. Black. 

These records show that C. canberra is widely 
distributed in the central arid portions of Australia 
to the east of the eastern border of W.A. Most of 
the water bodies from which it has been recorded 
are specifically described as being shallow and 
highly turbid. 


Acknowledgments 


I wish to thank W. Zeidler and other collectors 
mentioned above for providing me with the material 
on which this account is based. 


Fig. 7. Distribution of Calamoecia canberra Bayly. 


References 


Bayty, I. A. E. (1961) A revision of the inland water 
genus Calamoecia (Copepoda: Calanoida). Aust. J. 
Mar. Freshwater Res, 12, 54-91. 

___ (1962) Additions to the inland water genus 
Calamoecia (Copepoda: Calanoida). Ibid 13, 252-264. 

____ (1964) A revision of the Australasian species of the 
freshwater genera Boeckella and Hemiboeckella 
(Copepoda: Calanoida). Ibid 15, 180-238. 

____ (1978) Variation in sexual dimorphism in nonmarine 
calanoid copepods and its ecological significance. 
Limnol. Oceanogr. 23, 1224-1228. 

_____ (1979) Further contributions to a knowledge of the 
centropagid genera Boeckella, Hemiboeckella and 
Calamoecia (athalassic calanoid copepods). Aust, J. 
Mar. Freshwater Res. 30, 103-127. 

—__ & WILLIAMS, W. D. (1964) Chemical and biological 
observations on some volcanic lakes in the south-east 
of South Australia. /bid 15, 123-132. 

__ & ____ (1973) “Inland Waters and Their Ecology”. 
(Longman: Melbourne.) 

BeLk, D. & Core, G, A. (1975) Adaptational biology of 
desert temporary-pond inhabitants. /n N. F. Hadley 
(Ed.) “Environmental Physiology of Desert Organisms”. 
(Dowden, Hutchinson and Ross: Stroudsbury.) 

Bow er, J. M. (1982) Aridity in the late Tertiary and 
Quaternary of Australia. Paper 4 /n W. R, Barker & 
P. J. M. Greenslade (Eds) “Evolution of the Flora and 
Fauna of Arid Australia’. (Peacock Publication: 
Adelaide.) 


CHAPMAN, A. (1973) Calamoecia lucasi (Copepoda, 
Calanoida) and other zooplankters in two Rotorua, New 
Zealand, lakes. /nt. Rev. Ges. Hydrobiol. 58, 79-104. 


Geppes, M. C., De DeckkER, P., WILLIAMS, W. D., 
Morton, D. W. & ToppiNc, M. (1981) On the 
chemistry and biota of some saline lakes in Western 
Australia. Hydrobiologia 82, 201-222. 


GREEN, J. D. (1976) Population dynamics and production 
of the calanoid copepod Calamoecia lucasi in a 
northern New Zealand lake. Arch. Hydrobiol. Suppl. 
50, 313-400. 


McLaren, I. A. (1963) Effects of temperature on growth 
of zooplankton and the adaptive value of vertical migra- 
tion. J. Fish. Res. Bd Can. 20, 685-727. 


Maty, E. J. (1984) Dispersal ability and relative abun- 
dance of Boeckella and Calamoecia (Copepoda: 
Calanoida) in Australian and New Zealand waters. 
Oecologia (in press). 


MITCHELL, B. D. (1984) Limnology of mound springs and 
temporary pools south and west of Lake Eyre. /n 
P. J. M. Greenslade (Ed.) “South Australia’s Mound 
Springs”. (Nature Conservation Society of South 
Australia: Adelaide.) 


Timms, B. V. (1970) Chemical and zooplankton studies 
of lentic habitats in north-eastern New South Wales. 
Aust. J. Mar. Freshwater Res. 21, 11-33. 


A NEW SPECIES OF THE GENUS RHEOBATRACHUS (ANURA: 
LEPTODACTYLIDAE) FROM QUEENSLAND 


BY MICHAEL MAHONY, MICHAEL J. TYLER & MARGARET DAVIES 


Summary 


A leptodactylid frog Rheobatrachus vitellinus sp. nov. is described from the Clarke Range near 
Mackay, Queensland. The new species is larger than the gastric brooding frog R. silus, and is 
distinguished from it by a suite of external and internal characters. The two species are separated by 
a distance of approximately 800 km. 


A NEW SPECIES OF THE GENUS RHEOBATRACHUS (ANURA: 
LEPTODACTYLIDAE) FROM QUEENSLAND 


by MICHALL MAHONY*, MICHAEL J. TYLER & MARGARET DAVIEST 


Summary 


MEAWONY, ML, PYlbR, M4. & Dravins, M- (1984) A jew spucies Of the genus Rheubelrachus (Anura 
Leptodactylidie) fron Queenstand, Trens. R. Soe, 8. Aust, 1083), 195-162, 4 December. 1984 
A leprodactylid rae Rieaburnichus vires sp, voy. is desenbed fromthe Clarke Ranwe near Mackay, 
Queensland. The new snoeies is catger than the wastrie braoding frag R. sida, and is distinguished from 
i) by a suite of external aod internal characters. ‘The rwo species are separated by a distance of apprey- 


itarely BOO hi 


KEY Wokos Aju, beplodactylidae, chromosoines, RAeobarrachys vi/ellinus sp. nov. 


Iniradwelion 


The trog Rheebatrachus silus Laem (1973) was 
demonstrated by Corben e/ al, (1974) to be unique 
in the Animal Kingdom in brooding its young in 
its stomach, and eventually giving birth through the 
mouth, Che listological and physiological 
modifigations (hat accompany the conversion ofa 
stomach to a brood sac have been the subject of 
extetisive investigations based in Adelaide, and sum- 
miarised by contributors to the yolume edited by 
Tyler (1983), More recently ofher aspects of the 
cytology, pharmacology and physiology of gastric 
brooding have been dowumented by Gibbins & Tyler 
(1983), Tyler ef af (1983), Laidler er af (1984), de 
la Lande ef ad (1984) and Shearnian eal, (1984). 

In the lighr of the considerable interest in 2. silis, 
the sudden demise of the population in 1979 
assumed special sigmificance, Despite extensive sear- 
ches, particularly | 1982-83, nota single represen- 
lalive al RK si/us has been found, bul if is nol passi- 
bie to make-a definite statement tbat the species is 
exuinel. 

In January (9844 new species of RAeebatrachus 
was discovered in the Clarke Range in the vicinity 
ol Rungella National Park, west of Mackay, 
Queensland, approximately BOO km north of the 
known geographic range of Ro ails Tere we 
describe the new species. 


Materials and Methods 


The specimens reported here ure deposited in 
miuscum collections abbreviated as follows: AM 
Australian Museum, Sydney; OM Queensland 
Musecurn, Brisbune: SAM South Australian 
Museum, Adelaide 


‘School of Hrological Sciences, Macquarie University, 
Noth Ryde, NSW J1h3 
Y Departuient of Zootowy, 

Adelaide, SA, S00) 


Universiiy of Adelaide, 


Methods of measurement and abbreviations 
employed in the text follow Tyler (1968), Clearing 
and staining techniques lor bone and cartilage are 
\hose of Dingerkus & Uhler (1977). 

Milolic chromosomes from one individual (sub- 
adult female QM J42145) were obtained fram short 
term lymphocyte cultures using standard rechniques 
applied to mammalian cultures (see Sharman e/ wl. 
1970), the only modification being incubation at 
25°C. Whole blood (0,2-0,5 ml) was obtained by 
icant puncture using a sterile heparinized syringe. 


Rheobatrachus vitellinus sp, nov, 
FIGS 1-11 
Hololwpe: QM J42529, an adult female collected 
at Eungella National Park, 148°38 ‘00°E.; 
2°01'30"S., Queensland on 271,84 by Rk, R, 
MeDonald and V. R, J. Hansen. 


Desvription of holotype: Head flattened, approx- 
imately as long as broad, Snout not prominent, 
evenly rounded when viewed from above, rounded 
and projecting shghtly in profile. Nares dorsal and 
inclined superiorly; surrounded by loose, fleshy 
margin and with small papilla at posterior border, 
Distance between naris and tip of snout approx 
imately equal (o eye to paris distance. Internarial 
span ereater (hin clistance separating eye from haris 
(E-N/EN 0.80), Canthus rostralis hot defined, Eye 
prominent (Fig. 1). 

Tympanim not visible externally. No yomerine 
teeth. Choanae large, widely separated and sur- 
rounded by rim of soft Ussue, Tongue large and 
adherent to floor of mourh. Lower jaw with 
superior symphysial prominence inserting into deep 
ciastenia between premaxillae. Upper jaw with 
medial imimargination. 

Mingers cylindrical with lateral fringe on medial 
surface of digit Il. In order of length 3>4>2>1. 
Subarticulaur tubercles very poorly defined, uo 
palmar (ubercles, Sllehtly developed terminal discs 


156 M. MAHONY, M. J. TYLER & M. DAVIES 


Fig. 1. Upper: Female Rheobatrachus vitellinus in life. Lower left: Ventral aspect of R. si/us. Lower right: Ventral 
aspect of R. vitellinus. 


NEW XMEOBATRACHUS FROM QUEENSLAND 7 


Vig. &. Aheabarrachis vitelinus, A. Palmar surface of 
hand. B, Plantar surface of foot, 


lacking cirevimmarginal grooves. No interdigital 
Webbing (Fig, 2A). 

Hindlegs short (TL./S-V 0.47). Toes long, sub- 
articular tubercles moderately developed but lack- 
ing on metatarso-phalangeal joint of toe LV (Fig. 
2B). 

Expanded dises on tips of toes larver than those 
on fingers. Webbing reaching discs of all toes; broad 
medial flange on toe [, In order of length 
4>3-5>2>1, Large flattened inner but no outer 
metatarsal (ubercle. No supernumerary tubercles on 
fool: 

Dorsal skin irregular and foveolate. Large, 
irregular, bullate projections on upper eyelid. 
Distinet fold in customary supralympanic position 
extending from posterior corner of eye to insertion 
of forearm. Narrow, dermal fold superior to 
antenor i of supratympanic fold, No tarsal fold} 
few small, prominent Cubercles on posterior surface 
of tarsus, No tubercles on flanks, Anus with fim- 
brialed border. Ventral surface of body and limbs 
smooth, 

Alter three months in preservative, dorsum 
mottled irrealarly with dark brown and with small 
patches of cream upon a light brown background, 
Ventral surface cream With dense stippling of dark 
brown upon the throat and chest, but becoming 
more sparse posteriorly, Back of thighs heavily 
pigmented with dark brown. Palmar and plantar 
surfaces dark brown. Remainder of ventral surface 
pale cream. 


Osteology: 


Cranial Features 

Skull poorly ossified; sphenethmoid poorly 
ossified, modified anterolaterally to orm articu- 
lating surfaces, normally overlain by cartilaginous 


ey Ps 
ep 


—— 


\" 
\ \e 


Fig 9) Dorsal view of sphenethmoid of RAeubarachies 
vitedlinus. 


cap (Fig. 3); nor in bony contact with nasals, 
extending ‘4 length of orbit in ventral view, 
Elongate small bone located medially above 
sphenethmoid (Fig. 3), Prootic and exoceipital com- 
pletely fused dorsally, slizht reduction of ossifica- 
tion in plectral region, Crista parotica short and 
stocky, not articulating laterally with otic plate, 
Dorsally prootic extended posteromedially to form 
(wo flanges dorsolaterally to exoccipiral. 
Frontoparictal fontanelle ovoid except for indented 
anterior extremity. Frontoparietals well ossified, 
anterior extremities slender, asymmetrical, not 
reaching anterior extremities of sphenethmoid. Or- 
bital edges of frontoparietals straight, angled slighr- 
ly posterolaterally. Anterior margins of 
frontoparietal fontanelle formed by sphenethmoid 
at level about anteriorly along length of orbit. 
Posterior margin aboul 44 posteriorly along leugeh 
of orbit (Fig. 4A), 

Nasals small, slender, widely separated, expanded 
anteromedially, horizontal. Nasals not in bony con- 
lact with any roofing bones. Palatines broad, curv- 
ed, unvidged, overlying sphenethmoid ventrally to 
level of anterior extremities of frontoparierals (Fig. 
4B), Parasphenoid tnoderately robust. Culiritorm 
process short, tapering anteriorly, not reaching ar- 
ticulation Of anterior ramus of pterygoid. Alary 
Processes arising from ventral flanges in 
posteromedial region of cultriform provess, short, 
moderately slender and crenate laterally. 

Pterygoid extremely robust. Anterior rami in love 
contact with palatal shelf of maxillary. Medial rami 
extremely broad, blunt. Posteromedial {lange at 
junction of three rami and ventromedial (lange at 
posterior extremily of anterior ramus. Posterior 
rami moderately broad, long, acuminate. Quadrato- 
jugal robust and entire; squamosals robust with 
long acuminate zygomatic ramus and shovrtey 
expanded oOlic ramus, 


158 


M. MAHONY, M. J. TYLER & M. DAVIES 


Fig. 4. Cranium of Rheobatrachus vitellinus. A. Dorsal aspect. B. Ventral aspect. Scale bar=10 mm. 


Maxillary and premaxillary dentate. Teeth fang- 
like. Pars facialis of maxillary shallow with well- 
developed preorbital process, not in bony contact 
with nasals. Alary processes of premaxillaries short, 
broad, perpendicular to pars dentalis of premax- 
illaries, but inclined medially. Palatal shelf well 
developed with poorly developed palatine processes 
of premaxillaries. Lateral extremities of palatal shelf 
of premaxillaries elongated to lie medially to 
anterior portion of palatal shelf of maxillaries (Fig. 
4B). Pterygoid process not developed. 


\ Cc ‘ aw 
\ A va en 
| / Lp iN 
| F f fy ~ 
ff “. 


Fig. 5. Rheobatrachus vitellinus. A, Left ventral aspect 
of pelvic girdle, B. Humerus, C. Dorsal view of 
mandible. 


Vomers considerably reduced. Remnant denti- 
gerous processes present, but edentate. Alae poorly 
developed. Bony columella extremely long. 

Ligaments joining mentomeckelian bones on 
lower jaw directed dorsally to form cartilaginous 
protuberance fitting into notch between palatine 
processes of premaxillaries. Meckel’s cartilages 
poorly differentiated (Fig. 5C). 

Hyoid plate broader than long. Alary processes 
with broad base, variable shape (Fig. 6). No aper- 
ture present on alary process for passage of 
hypoglossal nerve. Cricoid ring complete. Branchial 
processes simple, moderately long; oesophageal pro- 
cesses short. Posteromedial processes ossified; car- 
tilaginous tips confined to lateral and medial 
extremities (Fig. 6). 


Post Cranial Features 

Pectoral girdle arciferal and robust (Fig. 7). 
Omosternum and ziphisternum present; xiphi- 
sternum % calcified. Clavicles robust, moderately 
separated medially. Coracoids robust, widely 
separated medially. Scapula bicapitate, very broad 
and stocky. Suprascapula about % ossified, 
proximal and posterolateral edges crenate (Fig. 7). 

Well developed anteroproximal crest on humerus 
(Fig. 5B). Phalangeal formula of hand 2,2,3,3. 
Distal tips of phalanges knobbed. Six carpal 
elements present: radiale, ulnare, preaxiale centrale, 
postaxiale centrale, carpales 2 and 3. Bony prepollex 
(Fig. 8A). 

Eight procoelous presacral vertebrae. Vertebra I] 
imbricate, others non-imbricate (Fig. 9). Relative 
width of transverse processes II] >1V>SD>II> 
V>VI>VII>VILI. Sacral diapopyses widely 


NEW RHEOBATRACHUS FROM QUEENSLAND 


Fig. 6. Ventral view of hyoid plate and larynx of 
Rheobatruchus vitellinus. 


SSS 


= 


LAS 


Se 
=S. 
SS 


Fig. 7. Dorsal view of pectoral girdle of Rheabatruchus 
vitellinus. 


expanded, Bicondylar sacrococeygeal articulation, 
(Fig. 9). Transyerse processes present on urostyle. 
Urostyle long with poorly developed crest extending 
about Vs its length. Moderately developed dorsal 
prominence on ilium; tiny dorsal protuberance 
(Fig. SA). Pubis cartilaginous. 

Phalangeal formula of foot 2,2,3,4,3. Three distal 
tarsal elements present; O. fibulare and O. tibiare 
fused. Small bony prehallux (Pig, 8B). 

No sesamoids present on appendages; free 
epiphysial joints not apparent. 
variation: There are four paratypes: AM RIII733 
an eviscerated adult male collected at Eungella 
National Park by K. R. McDonald and VY. R. J. 


\ 
| 
A jr 
a 7 
& VI (4 
a | a 
m4 i 


Foot. 


A a B ao 
f _ ah} Psat 2 4 
ee ee eee ea 
ee i, CoQ 
case oN oA 
SS SOs 
= a ay 
aan a ee 
rene Ag 
( es bY [ as 
ES aw AOS 
\ ra ao Broa } iw JA & \ | 
q | [ ~ vy 


Fig. % Rheobatrachus vitellinus, Vertebral column. A. 
Dorsal aspect. B. Ventral aspect. 


Hansen on 28.1.84; QM J42145 a sub-adult female 
collected at Tree Fern Creek, Clarke Range by M, 
Mahony on 2.i,84; SAM R25447 a cleared and stain- 
ed aclult female collected at Eungella National Park 


160 M. MAHONY, M. J. TYLER & M. DAVIES 


Fig. 10. Rheobatrachus vitellinus, Scanning electron 
micrograph of nuptial pad. Bar=10 » m. 


by K. R. McDonald and G, Chester on 12.i.84; SAM 
R25446 an eviscerated adult female collected at 
Eungella National Park by K. R. McDonald and 
G. Chester on 10.i.84. 

The adult females have S-V lengths of 68.9 mm 
and 62.2 mm respectively, the sub-adult female 
measures 41.5 mm and the adult male 55.7 mm. 

In their habitus the paratypes do not differ 
significantly from the holotype. The head propor- 
tions are uniform; the eye is protuberant but the 
E-N/IN ratio is slightly higher (0.86-0.98) com- 
pared with 0.80 in the holotype. 

The adult male has an unpigmented nuptial pad 
on the medial and dorsal surfaces of the first digit. 
With a magnification of 50x it is possible to see that 


1 


SE RR ae 
7 


Fig. 11. Karyotype of Rheobatrachus vitellinus. 


it is covered with numerous small spines (an SEM 
illustration is shown in Fig. 10). The male has a 
vocal sac with paired apertures on the floor of the 
mouth that are surrounded by fleshy margins, 


Colour in life: All specimens examined by us have 
a pale brown dorsum with obscure darker patches 
on both the body and limbs (Fig. 1). The ventral 
surface of adults bears an extensive area of vivid 
yellowish-orange (Spectrum Orange of Smithe, 
1975) covering the limbs and extending for varying 
distances up the abdomen. There also are patches 
of similar colour on the undersurface of the arms. 
The specimen illustrated (Fig. 1) has the remainder 
of the ventral surface unpigmented but in some in- 
dividuals it is dark brown. 


Karyotype: The karyotype of R. vitellinus is shown 
in Fig. 11; the diploid number is 2n=24, The 
chromosomes are arranged in two groups on the 
basis of size. Pairs 1-6 are large with relative lengths 
(R.L.) ranging 15%-10%; pairs 7-12 are small with 
R.L. ranging 6%-3%. Chromosome pairs 1, 5, 7 
and 8 are metacentric; pairs 3, 4 and 6 are submeta- 
centric; pair 2 is subacrocentric; pairs 9-12 are 
acrocentric. A prominent secondary constriction 
occurs procentrically on the short arm of pair 6. 


Comparison with other species: In appearance R. 
vitellinus differs from R. silus principally in its 
larger size and more spectacular ventral coloura- 
tion. The three adult female paratypes of the new 
species have an S-V range of 62.2-68.9 mm (and the 
largest live specimen now in captivity is approx- 
imately 83 mm long). This size range compares with 
44.5-53.9 mm S-V for 19 female R. silus examined 
by Tyler & Davies (1983). Similarly the sole adult 
male S-V of 55.7 mm compares with the cited range 
of 32.9-40.6 mm S-V for R. silus. 


TET 


ae hh hh 
12 


SEW RFAEORATRACHUS FROM QUEENSLAND {Al 


The stuiking difference in ventral colouration is 
aluwn in Pigs b ARheoharruechussilus has the ventral 
surface of the hindlimbs very pale yellow compared 
with the britht and more extensive yeltaw mark- 
ings of RK, vrlelenss, No individual of &, silus 
exhibit (he brown colouranon seen i some, bul ol 
all, Ry vite/linus. 

Ovivologically R vitellinus differs from &. sillus 
ina number of feattires. Cranially the presence of 
alticulaling facels anlerolaterally on the 
sphenethmotd and uf posterouivdial lanes on the 
dorsal surface of the prvotic, and the small 
supernumerary bone dorsal to the sphenethmoid 
are Lnique tof, irellinasy and in Vact are nol shared 
by any other Ausiralian leptodaetylid. The position 
ofthe anterior extremities of the frontoparietals and 
posterolateral anule of the orbital edges of the 
fromoparictals differ between the (wo species, Thy 
skwlot A. Virellinus is more extensively ossifed in 
the crista parotiea region and the frontopariveal tisn- 
tanelle is less extensively exposed than in @ siluy 
(Davies 1983). 

Absence of the apertures for (he hypoglossal 
nerve on the alary processes of the hyaid and 
minimal development of the menromeckelian ear- 
Hlages of the lower jaw are foarures unique to &, 
viellinny in ihe genus, 

Posteranially, the arciferal pectoral girdle of Ay 
Vitel/mus (Modified arcileral in A, silus) separates 
the [we species, ws does the orenate edves on the 
suprascapula and the felanve widtlts of the 
tramsveTse processes Of the presacral vertebrae. 

The karyotype of R, vite/linus is sinlar te those 
of the majority OF Ausiralian leptodacylid frogs, 
includiny A. si/us, iu diploid number and relayive 
chromosome lengths. When compared with the 
karyotype of &, sifas (Moresealehi & Ingram, 1974) 
ifferences are apparent i the cemromere positions 
OF several corresponding vhromosome pairs and in 
the location of the secondary constriction. The 
most Obvious dillerences in centromere position 
gecur in pair 6 Which is submetacentne im R: 
Vitellinus did acrocenee in A sins: pairs 9 and 
[Dare werocentric in A. viteliras and oaeracenerte 
(We situs A prominent secondary comstrietioat we- 
Cus procentnically on the shor aem af pair 6 in 


R, vitellinus, Moresealehi and Ingram (1974) did not 
identify any secondary constrictions in the 
kar yolype of R. silus, however the acrocentric mor- 
phology of pair 6 means that a secondary constrie- 
Lion cannot possibly be in the sare position as in 
R. vitellinus. These differences in chromosome mor- 
phology indicate thal structural chromosomal rear- 
rangements have occurred since the two species had 
a common ancestor and verify the specific identity 
of &. vitellintus. 


Habitat: Rheobatrachus vitellinus is an aquatic 
species inhabiting shallow sections of fast lowing 
erceks in rain forest, Preliminary observations 
suugest that the species is confined fo areas above 
approximarely 300 m a.s.l, where the crecks flow. 
a¢ross granitic rocks. K. R. MeDonald currently is 
exanining habilat preferences and distribution of 
lhe species, 

In January 1984 the only other species of frogs 
observed at the creeks with R, vite//inus were 
Tavidactvins eungellensis which was active on 
emergent rocks in (he creek beds, and 7 /iemi which 
was calling from crevices in the creek bank. 


Etytiplosy The specific name is derived from the 
Lalin viledinus “of the yolk of an ege” and refers 
to the ventral colouration. 


Acknowledgments 


We record our erati(ude to the Peter Rankin Trust 
bund and the Australian Biological Resources Study 
for funding the visic by one of us (M.M.) which 
resulted in the discovery of this species, We are also 
deeply grateful to the Queensland National Parks 
and Wildlite Service for granting collecting permits, 
and particularly to Mr Keith MeDonald for his 
enthusiastic cooperation, advice, and the provision 
of specimens, M.M, thanks §. Bergin, G. Johnston 
und T Maidens tor tield assistance. A visit to the 
type locality by M.JT was made possible by the 
Australian Broadcasting Corporation, Laboratory 
studies were lunded by a grant to M.T. and M.D, 
hy the Australian Research Grants Scheme, The 
ticlusian of calour plates was made possible by a 
grant fram the Mark Mitchell Foundation, 


References 


Corberx, ©. 1, deGaaM, Cio |e Tybee, Mod, (1974) 
Gastric branding: Wigue foe ot parental cane in we 
AUstiahyu Vrog, Severo 8G, Ode. 7 


Davies, M. (1983) Skeleton, da M.S. Teler (Ed) the 
Gastric Brooding Trog, pp, Sh68 (Croom Metin 
London and Cunberra.) 


eta Lanpr, |S, O'BRiex, P, SHEARMAN, DL. C., 
Taypor, P und Tyuer, M. J. (1984) On the possible 
role of prostiglandin £, in intestinal stasis in the gastric 
brooding lrog Rheatatrachus silus. Aust, J. Exp, Biol. 
Mee Sei, 62, 417-323. 

Dinutanos, G. & LHF eR, L. D, (1977) Pozyme clearing 
Of alcian blue stained whole small vertebrttes for 
déeinonstration of cartilage: Stein Technol, $2,229 23), 


162 M. MAHONY, M. J. TYLER & M. DAVIES 


GipBins, I. L. & TyLer, M. J. (1983) Changes in the 
organization and ultrastructure of smooth muscle cells 
in the stomach of the gastric brooding frog, 
Rheobatrachus silus, during brooding. Cell Tissue Res. 
231, 451-6. 

LAIDLER, P., TYLER, M. J. & SHEARMAN, D. J. C. (1984) 
The intestine of the gastric brooding frog 
Rheobatrachus silus during and after brooding: a mor- 
phological study. /bid. (in press). 

Liem, D. S. (1973) A new genus of frog of the family Lep- 
todactylidae from SE Queensland, Australia. Mem. Qld 
Mus. 16, 459-70. 

MORESCALCHI, A. & INGRAM, G. J. (1974) New 
chromosome numbers in Australian Leptodactylidae 
(Amphibia, Salientia). Experientia 30, 1134-5. 

SHARMAN, G. B., ROBINSON, E. S., WALTON, S. M. & 
BERGER, P. J. (1970) Sex chromosomes and reproduc- 
tive anatomy of some intersexual marsupials. J. Reprod. 
Fertil. 21, 57-68. 


SHEARMAN, D. J. C., TAYLOR, P., TYLER, M. J., O'BRIEN, 
P., LAIDLER, P. & SEAMARK, R. F. (1984) An update on 
the role of prostaglandins in the stomach and intestine 
of the gastric brooding frog Rheobatrachus silus. In 
A. Allen, G. Flemstrém, A. Garner & W. Silen (Eds). 
“Mechanisms of mucosal protection in the upper 
gastrointestinal tract?’ (Raven Press: New York.) 

SMITHE, F. B. (1975) Naturalist’s color guide. (American 
Museum of Natural History: New York.) 

TyLer, M. J. (1968) Papuan hylid frogs of the genus 
Hyla. Zool. Verhand. 96, 1-203. 

—— (1983) (Ed.). “The Gastric Brooding Frog?’ (Croom 
Helm: London and Canberra.) 

——., SHEARMAN, D. J. C., FRANCO, R., O’BRIEN, P., 
SEAMARK, R. F. & KELLy, R. (1983) Inhibition of 
gastric secretion in the gastric brooding frog, 
Rheobatrachus silus. Science 220, 609-10. 

— & Davies, M. (1983) Superficial features. Jn M. J. 
Tyler (Ed.). “The Gastric Brooding Frog’, pp. 5-15. 
(Croom Helm: London and Canberra.) 


QUATERNARY CLIMATIC CYCLES, LAKE MILLYERA REGION, 
SOUTHERN STRZELECKI DESERT 


BY R. A. CULLEN 


Summary 


Following the drying of the very extensive lakes which existed during the Tertiaty, there were three 
complex Pleistocene climatic cycles, or portions thereof, recorded in sedimentary sequences at Lake 
Millyera near the southeastern margin of Lake Frome. It is suggested these reflect changes in world 
climate related to glacial/interglacial episodes, though at present dating is only sufficient to identify 
the younger phase with certainty (16-21 000 years B.P.). The earliest phase records mainly high 
lake levels, culminating in complete drying of the lake bed (the top of the beach deposits was +16 + 
1.5 m a.h.d., the bed of Lake Millyera is + 4 m, and that of Lake Frome is close to sea level). This 
was followed by semi-arid fluvial activity and aeolian deposition, during which time Lake Frome 
regressed to its present shoreline. Watertables were lower than before, but higher than at present. 
The uppermost sequence represents part of the longitudinal dunes which built the Strzelecki 
Dunefield, again indicating a drop in watertable, but still periodically higher than present. A new 
rock unit, the Coombe Springs Formation is defined, and the Millyera Formation redefined. 


QUATERNARY CLIMATIC CYCLES, LAKE MILLYERA REGION, SOUTHERN 
STRZELECKI DESERT 


hy R.A, CALLEN* 


Summary 


Cary, R.A. (984) Quaternary climatic cveles, Lake Millvera region, southern Strzelecki Deseri, 
fens R. Sav 8S. Aust, (0803), 163-173, 13 Devember, 1984, 


Nollowing the drying of the very extensive lakes whieh existed during (he fertiary, (here were three 
cumples Pleistocene climatic evcles, or portions thergal, recorded in sedimentary sequences at Lake Millyera 
near the southeastern margin ol Lake Frome. 1 1s siigeesred these reflect changes in world climate related 
to placial/inerglacial episodes, though at present dating is only sufficient to identify the younger phase 
with certainty (16-21 O00 yrs BLP), The earliesr phase records mainly high lake levels, culminating in eom- 
plere drying of rhe lake bed (the lopof the beach deposits was 416 1.5 abd, the bed of Lake Millyera 
is 4 41), and thatol Lake Frome is close to wea level). This was followed by semiarid tlhivial activily 
and aeolian depustiion, during whieh dime Lake Frome regressed to its presemt shoreline, Waterlables were 
Jower than before, bur higher than at present, The uppermost sequence represents part of (he longinidinal 
dunes which buill (he Sirzelecki Dunefield, again indicating a» drop im watertable, but still periodically higher 
than presen. A new rack unit, the Coombe Springs Formation is defined, and the Millyera Formation 


redefined, 


Key Worps Quuternary climate, Strveleck! Desert, Lake Frome, Lake Millyera, Coombe ‘Springs 


Formanoan, Millyer Formation, 


Introduction 


Around Loke Millyera in the Strzelecki Desert 
(Fig. 1) dre a number of well-exposed Quaternary 
sections, «woot whieh have been used to define rock 
Stratigraphie unity ia the region (Callea & ‘led ford 
1976), These seetions contain an excellent revord al 
former climatic Huctuacions, relating to the prior 
expansion and contraction ol Lake Frome curing 
Ihe Quaternary. Both these lakes are now playas, 
which {ill occasionally under conditions of excep- 
tional local raiis (Callen 1983), 

Lake Frome has a critical size in relation to its 
culchment, such that it should respond to major 
changes in runolf bui not local Muctuations (Bowler 
1981). Thus variations in shoreline should reflect 
significant climatic events. Possibly superimposed 
ov this is the effect of Quaternary uplift of the 
Flinders Ranges (Callen & Tedford 1976) which 
night have deereased runoff in the western catch~ 
ment of Lake Frome by the rainshadow effect. 

Past shoreline positians can be identilied by 
appropriate facies changes in thie sediments. Degree 
of aridity can be micasured by identifying aeolian 
sequences, evaporites, and palacosol horizons, sup- 
plemented by fossil evidence, Some limitations are 
imposed by the lack Of accurate height data, lack 
of knowledge about movements on faulty if the 
vicinity, and problems with radiocarbon dating of 
calcareous palaeosals (Callen ef af, 1983), 

Mis short paper describes four sections in detail 
and thelr iuterpretation, and redefines the Villyera 


* CGeologwal Survey, Dept ot Mines de Energy, 19) (Green 
hill Road Parksice, S4,, 3063. 


Formation, introducing a new unit, the Coombes 
Springs Formation (Callen ef @/, 1983). Their 
significance in relation to world climatic change is 
assessed, 


Results of the Investigation 


Lake Millyera is located adjacent (0 the southeast 
ol Lake Frome, cutting across a series of palaeo- 
shoreline features. Good exposures of Late 
Cuinozoie sediments are to be found along the nor- 
thern shore, and in gulhes to the southeast (Fig, 1). 
The northern shore is dominated by low cliffs of 
the Tertiary Namba Formation dolomite and clay 
(Callen & Tedford 1976), blanketed hy red aeolian 
and lluyial sands of the Quaternary Period, The 
best Qualernary sequences are found along the 
soulheastern shore (Fig. 2), where (te Namba Por- 
ination is not exposed, and in a depression cut in- 
(o the Namba Formation in the northeastern part 
of the lake adjacent to the old track crossing (Fig. 
1). These Quaternary sequences range from 3 m to 
25 m thick, and reveal green lacustrine clays and 
fine sand, red fluvial sands, arid an overlay of two 
aticient aeolian sequences, Shelly beach deposits 
form prominent white benches, Caleareous 
palaeosols and algal limestones are useful markers. 
The Nainba Pormation forms a base ro the entre 
sequence, and can be found just beneath the lake 
floor, 

Descriptions of the sections, with interpretations, 
are presented in igs 3-6. A summary of correla- 
tion and environments appears in Fig. 7, 

Thin sections, X-ray diffraction and radiocarbon 
daling were used to study rhe sequence, 


164 


R. A CALLEN 


4 Y Site iit yy 
3 FP 1 CALLABONNA 
8 MARREE J, 
2 BLANCHE ER SS SE te 2 yg 
SCALE IN KILOMETRES 
190 20 30 ao so 
| S 
we 


30° 00 ES 


\ ‘N). 


Se 


f 


Frome Downs 7 
ji 


2 * Godnadorte 


nv 


Lake Eyre 


Morree™ 


Lake Torrens 


dekh 


MI ARROWSMITH 


Fig, 1. Location of Lake Millyera. 


LOCALITY MAP /\ 


peed 


nT | 


ALICE SPRINGS w 


LEGEND 


CAINOZOIC & MESOZOIC. — L_] 
Pre-Creinceous \N 
(Combrian and Precambrian sediments N 


and Crystalline bosement) — — 


WA a i an 1 Dunp frends —— eee eT 
QODNADATIAg “] 
Les BRISBANE Edge of high level plains — mad 
ae * 
. NSW Section and Ret number____ FR 
ADELAIDE 
[~-\, svoNey Trg Pom _A 
0 sO TRS 00 MELBOURNE pVIC. Aduitithod ee i 
CF ete sce eg eg _—— 


QUATERNARY CLIMATIC CYCLES, LAKE MILLYERA 165 


To _\ 
Lake Frome 
) he 


AUGER HOLE @ PIT 
* 
SECTION 4 


SCALE 


100 is) toa rane] 300 490, Bhs) 


METRES 


REFERENCE 


Recent loxe sediments = 


Recent slope wash and small fans 


Coonarbine Formation with 
Chara Limestone 


Eurinilla Formation’ Clay junette and probable alluvial 
deposits to ne south May include some red beds of CZes. 


Mitlyera Formotion with 


Lofgituding! dunes, usually with cores of Ophe 


Suytical sand. sprend of interdune oreos and 
Holocene valley fil! (par! asohan, part slope wash) 


oO | 
Ls ) 
r Ons: } Car J 
Copse dunes 
Qha, ] 
[ Ore | 


platy Chara Limestone 


Gecs | Coombes Springs Formation 
SS Reu sands 
—— Shell layer 


i 


Namba Formation (Carly to ?Lore Miocene}, 


SAOME 8 /)644I 


Fig. 2, Western Lake Millyera—geological map and location of sections, 


The sections around Lake Millyera have useful 
height data and do not have their relationships com- 
plicated by faulting. However, similar units to the 
east may possibly have been affected by northerly- 
trending structures. Nevertheless, the eastern lakes 
region offers the potential of resolving older 
shorelines through detailed drill traverses coupled 
with accurate height data, 


Correlation and Nomenclature 

Detailed mapping, and practical aspects of depic- 
ting shoreline deposits of different ages on 
1:250 000 scale geological maps of the South 
Australian Department of Mines and Energy (con- 
sider the implications of the section, Fig. &), have 
lead to redefinition of the Millyera Formation and 
introduction of a new unit, the Coombes Springs 


166 R. A CALLEN 


10 SECTION 4 
rs 
2 
° 
E = : : 
COOMBES SPRINGS “4 Fg ee RE SU as Te YM Z 
2 Fi beset tcaeteoes Cee © SS ee x [7] 
re FORMATION é es 
a5) 8 (6) 
Sih hoo 7 6 ee 
[3] 
om 1& 3 
F er 5 O— [3) 
[e} FORMER — aan 
a hss ae | MILLYERA ~ ze 
a ere, «= FoRMaTION ¥ = 89) a 
pe | ix 
og (8b) 
WwW 
“ | { 7 tt 
NAMBA FMN @) 


REFERENCE FOR SECTIONS 


FOSSILS 
[od Gastropods, bivalves ® Land snails 
¢ ..Vertebrates (mostly fish) a Aboriginal artifacts 
8 Chara oogonia € Egg shells 
v Ostracods 
LITHOLOGY STRUCTURES 
7 5 clayey 
Sands 4 Bea gypsiferous eae Small scale cross lamination 
Clays UZ Aeolian cross-beds 
ooo 
go], Pebbles — ...Horizontal lamination 
Erstad Chara Limestone Raid 
=e ai Erosive contacts 
a .Lacustrine gypsum 
—— Sharp contact 
| —=|) Clayey lens, clay clasts @ 
SS a Beds of this paper 
Coquinite 
2] Formal units of Callen & Telford (1976) 
er Disc- shaped gypsum crystals - simple 
a or rosettes 
any NOTE: 
v Gypsum patches in paleosols The level of Lake Frome is close to Mean 
Th Sea Level. 
| Pedological jointing Levels are tied to Lands Dept. bench 
marks by barometric survey. 
§ $5 Calcareous paleosols 


SADME S$ 16938 


Iron mottling 


Fig. 3. Lake Millyera section 4, includes. Millyera Formation type section. 

1. Orange brown loose sand, fine grained. 0.3 m. Modern longitudinal dune sand. 2. Coarse sand with gypsum 
grains and angular milky quartz pebbles, hummocky “cow pat” gypsum crust. 0.7 m. Interdune deposits of gyp- 
sum dunes, gypsum tepees. EURINILLA FORMATION: 3. Clayey very fine greyish-orange sand with numerous 
Chara oogonia. Multi-coloured grains. Capped by gypsum crust 1.10 m. Overbank deposits with old saline groundwater 
horizon (represented by gypsum). COOMBES SPRINGS FORMATION: 4. Soft clay with sharp upper contact, 
dark yellowish brown. Oxidised and crumbly, grades down by alternation to fine medium sand with Chara oogonia. 
This sediment overlies, with sharp contact, well sorted clayey sand of greenish yellow colour. 1.40 m.?Lacustrine 
deposits and ?beach sands, soil profile at top. MILLYERA FORMATION TYPE SECTION: 5. Interbedded clay 
and clayey very fine sand in very thin millimetric laminae. Sand very well sorted and rounded, grades down to 
6. 0.70 m. Lacustrine. 6. Laminated yellowish grey to green clay with silt laminae. 0.30 m. Lacustrine. 7. Thin, 
platy, charophyte limestone, consisting of Chara tubules, rare ?Coxielladda gastropods, interbedded with clay as 
above and below. 0.40 m. Saline lake, probably ephemeral. 8a. Brittle, soft, waxy clay, distinctly laminated and 
thin-bedded, each lamina grades up to fine silt with Chara oogonia and ostracods. As for 6. 0.10 m. Lacustrine, 
possibly seasonal. 8b. As above, but oxidised brown, lamination not clear, grades abruptly to red sandy clay with 
reworked dolomite granules at base. 1.30 m. Lacustrine, lag preserved at base. NAMBA FORMATION: 9. White 
dolomite, thin bedded, hard, micritic, clayey. Very sharp erosional contact at top. 0.10 m. Lacustrine. 


QUATERNARY CLIMATIC CYCLES, LAKE MILLYERA In 


SECTION 5 


A NeuNie 
ete 


Tenn 


Familet 


lary 


Hiewhy 


tiv, 4 Lake Millyerg, sewuion S, mieludes Coumbes Springs Formation type seciion. 


Ta, Brown sand, 0.30 m. Modern ephemeral beach und fan deposits. 1b. Red brown loose fine to medium sand. 
3,50 m, Modern Jongitudinal dunes, lc, Brown to red fine cross-bedded sand, 1.00 m, Modern copse dunes around 
shrubs, 2. Red- brown sand, fine ta medium, poorly sorted. Blocky ped structure 10 ™ 5 ems, 0,50 mm. Slope wash 
dnd dritt sand. COONARBINE FORMATION SUPPLEMENTARY SECTION; 3, Light brown sand as Jor 4, 
weak diffuse carbonate crust. Blocky polygonal joint pattern, 0.40 m, Longitudinal dunes with weuk pulacosols. 
4, Red-brown well sorted, well rounded orange-brown sind, massive. Soft white carbonate mottles ar top, 0,60 im. 
Longitudinal dunes with semi-arid calearcous palacasol, §, Porous Chara oogonia and algal platelet limestone with 
rare voarse sand and gastropods, 0.20 m, Probably windblown trom nearby beach— suggests ephemeral lake in 
vicinily. KURINILLA FORMATION SUPPLEMENTARY SECTION AND PINPA PALAEOSOL: fa, b, Fine 
to medium sand, poorly sorted, massive light brown sand grades down to inedium brown sand with conver laree- 
scale false-bedding dipping towards Lake Millyera. Pinkish irregilar carbonate nodules in upper parl, and capped 
by several well-developed calcareous rhizonodule and gypsum mottle horizons (Pinpa Palaeosol), Some carbonate 
granules. Sund lunettes with some clay pellet layers, capped by calearcous palacosols (semi-arid), Shell and Charu 
oogonia lenses iy centre. Windblown shell from nearby beach, Basal part of section contains clay pellet layers inter- 
bedded with small-scale cross laminated coarse sand rich in Chura oogonia (tangential bottomsets, foresers face 
away trom Luke Millyera) and rare gypsum sand grains, impregnated with gypsum. S04) 1. Gypsum and clay lunetle, 
Cross lamination probably represents upward migrating megaripples on aeolian dane front. Seasonally flooded 
pan, COOMBES SPRINGS FORMATION TYPE SECTION 7s, Hard pale olive clay with irregular shiny ped 
surfaces, mangans and ferrans well developed, Reticulate gypsum cylindroids duncated at surface, Orange brown 
patches. Upper contact sharp, flat, eroded, Grides down Lo light green solt clay interbedded with fine white sand 
nich in Chare ooponit. 2.50 m, Lacustrine deposits capped by lake mud gypsum of lowered water table, Broded 
lake bed. 7b. Yellowish grey silt to fine sand with numerous Whit clay lamellae cich in Chara tubules like those 
in platy limestones of Millyera Formation, Some very coarse lenses, Good sorting and well rounded, Clay pellet 
layers (course), 10 cm thiek smiil scale cross-bed sets, Mish vertebrae, rare egg-shell ar base. Coxie//adde and similar 
gastropods, ostracods. 5,00 m, Beach sands of saline lake with permanent water lacustrine phases. 7c, Bright ced 
brown very fine sand, grading to above unit, Very coarse sand to granules at base. Contact with underlying unit 
inrepulir, Miatssive gypsum rosettes with dise-shaped crystals. 0,93 m, Aeolian source, but fluvial overbank deposits, 
or playa margin fans? (lacking lamination, mud drapes and mud eniecks), Saline proundwater horizon, MIT LYERA 
FORMATION SUPPLEMENTARY SECTION; 8&1. Rippled to nodular gypsum laminae in red sand as above. Some 
interbedded charaphiyte limestone Lo southeast of main section, Chara oogonia and single course sand grains svar 
tered through gypsum. Playa lake with some aeolian content. Ephemeral. Sb. Yellowish orange to ureenish while 
sand, fine grained, with Chara oogonia, very coarse laminae, Reworked distorted clay fragments al hase on hart 
cracked surface. 0.70 im. Lake or beach sands. NAMBA FORMATION: 9. Hard black clay with greasy skew plane 
surfaces, Disturbed upper contact. 3.28 m. Bioturbated lake and swamp deposits, Palaeosol, 


168 R. A CALLEN 


20 SECTION 16 


? COONARBINE FORMATION 7? 


3 15 

@ COOMBES 
a SPRINGS 

a 

wn 

= FORMATION 

WwW 

3 

m 10 

aq 

— 

= > MILLYERA FMN. ? 5 ® 
wu 

x 


LAKE 
5 . MILLYERA 


= 
YFG ASILAY ER FI SE I NIA SS 


SADME S16942 


Fig. 5. Lake Millyera—Section 16. (Section “100 m NW section 5” of Callen ef a/. 1983). 

2? COONARBINE FORMATION: 1. Coarse, orange-brown sand with egg shell, landsnails, algal and root tubules, 
aboriginal artifacts. 1.10 m. ?Aeolian dune. 2. Limestone of large Chara oogonia and algal platelets. 0.25 m, 
?Aeolian—blown from nearby beach—implies seasonal lakes. COOMBES SPRINGS FORMATION SUPPLE- 
MENTARY SECTION: 3. Well sorted medium grained beach sand with no shell over alternating green and brownish 
green clay and clean white sand with much shell (Corbicula, Coxielladda, etc). Persistent thin layer of ornamented 
spiral gastropods. Bedding diffuse and disturbed. Lenses of bioturbated beach sand. Greenish yellow colour. Mottled 
with white soft carbonate of calcareous palaeosol. 8.4 m. Beach deposits, with some ?aeolian clay pellet layers. 
Saline to fresh lake. Semi-arid soil-carbonate cap. MILLYERA FORMATION: 4. Laminated clay very similar to 
Millyera Formation. 1.20 m. Lacustrine, 5. Hard platy Chara limestone, possibly Millyera Formation. As float. 
Ephemeral lake. HOLOCENE: 6. Copse dunes. 1.0 m. Recent aeolian sand accumulated around shrubs. 


Formation. The beach deposits are mappable as a 
lithostratigraphic entity, though consisting of a 
number of separate phases. These phases are 
grouped into the Coombes Springs Formation, 
which now includes the upper part of the Millyera 
formation in some of its former supplementary 
sections. 

The Millyera Formation type section (section 4, 
Figs 2, 3) has been correlated with a supplemen- 
tary section (Section 5, Figs 2, 4, 7) using platy 
charophyte limestone as a marker unit. A persis- 
tent bed of laminated, rippled powdery gypsum is 
found interbedded near the base of the section (Fig. 
4) and in a gully to the south; laminated Chara 
limestone is interbedded with this. Between these 
two sections, the limestone is found at several loca- 
tions as float (Fig. 2). In addition, laminated green 
clay like that of the Millyera Formation was found 
beneath beach deposits at section 16 (Fig. 5), and 
interbedded with the limestone in section 5 (Fig. 4). 

The beach deposits above the limestone and green 
clay can be traced between sections 5 and 16, but 
the relationship with section 4 is not seen. Nor is 


it known whether they are composite beaches or 
represent single episodes of stable lake levels. 

In Section 17 (Fig. 6), not previously published, 
beach deposits of similar aspect are found resting 
directly on the Millyera Formation, which is iden- 
tified here by the interbedded charophyte limestone 
and clay. The fine red sand beneath the Millyera 
Formation in section 5 (bed 7c, Fig. 4) has been 
found in the vicinity of section 17 (Fig. 6), but 
definite relationships were not observed. Clasts of 
this material are in bed 5, and the soil developed 
on it closely resembles the Pinpa Palaeosol (Callen 
et al. 1983) found on the Eurinilla Formation (bed 
6a) at section 5, all of which suggests it could be 
equivalent to or older than the beach deposits at 
section 17 (Figs 6, 7). This red sand is 
topographically higher than the beach deposits at 
this site, and also crops out along the northern edge 
of Lake Millyera, where it rests on the Namba For- 
mation. There remains the possibility that the beach 
deposits of section 17 (Figs 6, 7) are cut into this 
unit. Thus, although section 17 bed 7 is confident- 
ly equated with the Millyera Formation at section 


QUATERNARY CLIMATIC OVYCLES, LAKE MILLYERA ay) 


1 SECTION 17 3) 

r (2) 

: 

" TOO NARBINE 
a FORMATION 
z 

> We i a pe ee AR en SE re 

mn CAKE COOMBES SPRINGS 
=z MILLY ERA FORMATION 

E 

E = ee a 
wo MILLYERA. FORMATION 

w 

zr 


SEUMe = eae 


Pig. 6. Lake Millyera—Seetion 17, 

1, Brown fine sand. 0.30 m, Modern lake, marginal small-scale fans. 2. Yellowish fine to medinm loose sand 7.) 
m. Modern longitudinal and transverse dunes. COONARBINE FORMATION: 3, Light brown quatiy sand with 
Polyzonal joloe pallerns and weak calcareous patches ay top. 1.0m, ‘Transverse dune and interdune deposits with 
semi-arid palueosot 4, Fine brown quarta sand to brownish ercy pypsacolianile, lurge scale cross-heddineg at low 
angle (10-15), Polygonal joints 40-50 ems across, small gypsum stringers. Numerous Chure Oogopin, Sut lave lil- 
tered with lanl snails, aboriginal urtifacts. Very irregular lower contact, 7,0 1n. Transverse sand und gausum dimes 
of Jeeside mounds along cast side of L, Frome, 5, Coarse sand with large clasts of underlying beds and bed § 

Much disrupted by rabbit burrows, Calcarcous patches at top. 0.90 m. Probably interdune tat deposits. COOMBES 
SPRINGS FORMATION SUPPLEMENTARY SECTION: 6. Light greenish brown to light brown meédiun) ta vers 
coarse sand, becoming clayey and laminated in lower half. Lenses of gastropod and shell in upper part, Foagryents 
vPexg shell, scattered fish bones, numerous Chara oogonia and ostracods. Some lenses of green clay, Small seale 
cross lamination Upper surface very irregular. 1.50 m. Lacustrine beach and nearshore deposits. MILLY RRA FOR- 
MATION: 7. Laminated light green soft clay, sub-conchoidal fracture, with laminae of CAura tubules, Pitted upper 
Contact, Ostracods present, 0.40 m, Lacustring, offshore, becoming ephemeral at top. 7COOMBES SPRINIS HOR: 
MATION: 8, Bolin brownish red fine sand with strongly developed white carbonate and gypsum parehes Rela 


Honship to @ uncertain. 2.0m, Overhank deposits or lukesedge fans with semi-arid soil horizon at top, 


5, il is notcertain whether the beach deposits above 
fare equivalent or tepresent a younger and distirict 
beach. Added to this is the problem (hat extensive 
pedogenesis has taken place on bed 10 of section 
8, far In excess of anything encountered on the 
Millyera Formation elsewhere (though it may have 
been eroded), 

For these reasons and practical mapping cvon- 
siderauious, the Millyera Formation has been 
redelined to exclude the beach deposits, which are 
now grouped into the Coombes Springs Formation, 
Within these unils, considerable voniplexity is to 
be expected, along the lines of the model (Fig. 8), 
Which demonstrates a repetition of facies of differ 
ing. ages, The new use and first publication of the 
new nane arc jy Callen ef a/, (1983), 

The Millyera Formation ts redefined (o exclude 
Units (3-5) of section § (Fig. 4), Units 5-7 in the 
lype section 4(Fig. 3) and all the Millyera Forma- 
lion of sections 6 and 7 in Callen & Tedtord (1976), 
Which are vow included in the Coombes Springs 
Formation, Unit | of the type section (Fig, 3) has 
been extended downward by means of ai auger 
hole, Which intersected Namba Formation dolomite. 


Age of Uniis 

The radioearbon age of calcareous palaewsoals has 
been diseussed iu Callen et af (1983) lor Section 
S (bigs 2. 4). Results suggested a “last glacial” age 


for beds 3 and 4 of the longitudinal duries (Fig. 4), 
and that the Eurinilla Formation lunetbe is al les! 
95 000 yrs old, 

Section 4 (Fig. 3) was sampled by J. M. Bowler 
(pers, comm. 1981) for magheue reversal stfati- 
graphy, and has been determined as maenetically 
normal throughoul, However ir is not known 
whether this is the Gauss or Bruntes episede. The 
degree of induration suggests the younwer age is 
vorrect, Thus the Millyera Formation is probably 
between 95 000 and 2.2 million years old, and the 
Coombes Springs Formation at the type section is 
probably also within this time episode bur younyer 

Correlauon based on gross lithological ane 
fatinal similarities suggest (he Coombes Springs 
Formation overlies (he Millyera Formation, thoweh 
itis possible the relationships are more complex 
For (he present, the simpler explanation is nevepted 
Height dilferences bel ween sections are within ihe 
limits of the barometric method used (= 1.5 m). 

All units are younger than the Namba lorma- 
tion (Callen & Tedford 1976) which may extend to 
Late Miocene age oy evel Phioeene iy the upper 
part. 


Discussion and Interpretativn (Pip. 7h 


Although the detailed relationship benween these 
sequences has not been fully resolved, the follow- 


170 R. A CALLEN 


CLIMATIC SUMMARY 
(Based mainly on Section 5) 


WET DRY REFERENCE 
LAST ENVIRONMENTS 
GLACIAL 
au ob itding’s isod Pr ent (? alkaline) lak 
june- building episodes. ‘erman © Glkall e 
>95000 (Callen ef al 983) 
Drop i ter table, , 
channeling of He bed. Permanent shallow lake (? brackish) 
> 
S Ephemeral playa 
= 
Ephemeral ( Chara limestone ) 
16 CF Fluvial 
Yili, 
aoe Beach 


Aeolian (sand, longitudinal dunes ) 


Aeolian (gypsum , clay, sand 
lunettes or leeside mounds ) 


? Aeolian (Chara oogonia) 


Paleosols 


L2| RN COONARBINE FORMATION 
EF. .EURINILLA FORMATION 
CSF..............COOMBES SPRINGS FORMATION 
ME:.,...-csuats MILLYERA FORMATION 
NF .............NAMBA FORMATION (TERTIARY) 
CONTACTS 
raz 
> 
< ~~. ....... Erosional 
E ——_....... Disconformable 
wv ---—-........Gradational 
= 
< 
a VERTICAL SCALE 
fa) ° 5 
= ———E————EEE 
| METRES 


Mean Sea Leve/_+ 1-0 metre SADME S$ 16939 


Fig. 7. Lake Millyera. Correlation of sections and environmental/climatic interpretation. 


=| Chara Limestone — ephemeral lake 


Beach deposits 


Clays — permanent lake 


Fig. 8. Diagram of beach and lacustrine facies for a shallow lake of fluctuating depth. 1-4 are successive lake levels. 


QUATERNARY CLIMATIC CYCLES, LAKE MILLYERA 71 


ing comments can be made, and are summarized 
in the Figure captions (Figs 3-6). They supplement 
and modify the outline given in Callen (1977, pp. 
164-5 and Fig. 6) and Wasson (1983a, b). 

Deposition in the Lake Millyera region in post- 
Namba Formation times (Late Tertiary or after) 
began with the lacustrine Millyera Formation. The 
deposits have features indicative of a standing body 
of saline water. They are finely laminated green 
clays, with increasingly abundant calcareous algal 
laminae near the top (stem moulds and fruiting 
bodies of charophytes). The clays contain ostracods, 
including Reticyprus kurdimurka (pers. comm. P. 
De Deckker 1983), and rare spiral gastropods. 
Charophytes can reproduce in salinities up to twice 
that of seawater, but may require lower salinities 
for germination of zygotes (Burne et al. 1980), The 
algal limestone beds grade into rippled gypsum, a 
relationship observed in modern environments 
nearby (Callen & Tedford 1976), and showing the 
water was highly saline in the final depositional 
phase and at least some of the time during clay 
deposition. 


These sequences suggest a permanent lake as 
there is no evidence of drying and palaeosol for- 
mation during clay deposition. The proportion of 
carbonate laminae increases upwards over about 
50 cm and terminate abruptly, suggesting increas- 
ing salinity variations and drying of the lake. There 
followed a brief return to a permanent lake before 
the watertable dropped and the shoreline of this 
precursor of Lake Frome retreated west of Lake 
Millyera. 

As the lake dried, coarse aeolian sand grains were 
blown onto the gypsum deposits. This was follow- 
ed by deposition of horizontally laminated red- 
coloured fine sand as the lake regressed, probably 
representing an ingress of waterborne aeolian sedi- 
ment to the lake margin. At this time the lake 
shrunk to near its present shoreline. The red sands 
are an early phase of the Coombes Springs Forma- 
tion. They are cemented with large disc-shaped 
gypsum crystal rosettes, deposited from saline 
groundwater in subsurface. Although indicating a 
drop in watertable, this was still much higher than 
present. 

The red sands were followed by fine white to 
greenish sands rich in fossils. They contain an abun- 
dant shelly fauna, dark-coloured smooth eggshell, 
fish remains, and calcareous Chara algal remains. 
The gastropods and bivalves include Coxiella, 
Coxielladda, Potamopyrgos, Corbicula and 
Pisidium species, all of which are salinity tolerent, 
though not of high salinity like that in present day 
desert salt lakes of this area (Buonaiuto 1982!). 
The beach deposits resemble those of modern Lake 


Eyre formed in recent times, though with a more 
diverse shelly fauna. The comparison suggests a 
great potential for bird fossils. 

Higher in this sequence are massive alternating 
green clays and sands. The clays, though massive 
and rather crumbly, do not exhibit aeloian clay 
pellets, though containing rounded clay-flakes 
indicating exposure of mudflats. Some of this struc- 
ture is probably the result of soil processes. Slicken- 
sided skew planes with clay and iron oxide coatings 
are common. These sediments are interpreted as 
lake deposits affected by later soil processes. The 
soil is best developed at the top, where a reticulate 
mass of gypsum penetrates the clay bed, and has 
been truncated at the surface. Thus pedogenesis 
took place on a former lake bed, implying the water 
table dropped beneath the lake floor. This lake bed 
forms the foundation to the extensive sand-covered 
flats in the vicinity. 

Elsewhere, along Lake Tarkarooloo, the Coombes 
Springs Formation can be traced laterally into 
brown fluvial cross-bedded sands and gravels 
(Callen 1977), cemented with carbonate. These 
deposits are in channels cutting into the Namba For- 
mation and contain abundant burrows, possibly of 
insects such as ants (Callen & Tedford 1974, Callen 
1977). 

Lunette dunes of the Eurinilla Formation were 
built upon the lake bed near Lake Millyera. These 
are essentially sandy, but have clayey layers with up 
to 40% aeolian clay pellets at the base, alternating 
with cross-bedded sands. This small scale cross- 
bedding dips away from the lake, whereas the domi- 
nant low angle large scale crossbeds dip towards it. 
The small scale crossbeds were probably formed by 
lee eddy deposits. The base of this section is im- 
pregnated with secondary gypsum, and occasional 
rounded gypsum grains are present, suggesting there 
may have been more abundant aeolian gypsum 
previously. The sands are rich in charophyte 
oogonia and contain occasional shelly layers, sug- 
gesting periodic flooding of the surrounding lake 
floor. The sequence is capped by calcareous 
palaeosols, demonstrating stabilization of the 
dunes, and the presence of rhizomorphs indicates 
growth of plants. Rare Diproftodon tooth enamel 
fragments and Genyornis and emu eggshell are pre- 
sent. Similar deposits are present along Billeroo 
Creek and at Lake Moko, but no clay pellets were 
identified. 

The origin of clay pellet dunes and building 
lunettes has been adequately discussed by Bowler 


'Buonaiuto, J. M. (1982) Late Cainozoic non-marine 
mollusca of the Lake Frome area and other selected 
localities from northeastern South Australia. S. Aust. 
Dept Mines & Energy Rept, 81/68 (unpublished). 


172 R. A CALLEN 


(1983), and for Lake Frome, by Callen (1983). Clay 
is broken down on mudflats where groundwaters 
periodically reach to the surface, permitting 
crystallization of salts and “fluffing” of clay, which 
is then blown off the dry lake bed. Gypsum is 
broken down into cleavage flakes and blown into 
dunes; rounded edges and sorting are evidence of 
wind transport. 

Elsewhere the Eurinilla Formation is essentially 
fluvial, with shallow ephemeral streamflow (Callen 
& Tedford 1976, Callen et a/. 1983). Some redbeds 
mapped as this unit on the FROME geological sheet 
are likely to be aeolian, or fluvial equivalents of the 
Coombes Springs Formation. The temporal rela- 
tionship between the aeolian and fluvial facies in 
the Eurinilla Formation is uncertain. The streams 
cut down into the older lake deposits, and the 
shoreline of Lake Frome regressed to its present 
location. The channels contain locally abundant 
vertebrates similar to those at Lake Callabonna,. 

Thus there was a profound drop in water table, 
suggesting a significant decrease in rainfall. The 
absence of extensive aeolian deposits suggests lack 
of windiness, though deposition in many desert ter- 
rains tends to be dominantly fluvial, and other 
aeolian facies may yet be recognized. 

Longitudinal dunes were built upon all of these 
sequences, recording a change to aeolian conditions, 
and increased windiness. However, the abundance 
of clay pellets in these dunes indicates periodic 
flooding of the interdune corridors, so the water 
table was still higher than present (Callen er a/. 1983, 
Wasson 1983a, Ash & Wasson 1983). The pure algal 
oogonia limestone beds suggest Lake Millyera was 
flooded quite regularly. 


Conclusions 

The catena of landforms described above has 
been controlled by a fluctuating watertable and 
ultimately by climatic change. The cycles begin with 
a permanent brackish lake extending east of pre- 
sent day Lake Frome. This lake became ephemeral, 
decreased in size, and semi-arid conditions prevail- 
ed, though watertables were much higher than pre- 
sent (10 m or more). Processes were dominantly 
fluvial and lacustrine, though there is some evidence 
for wind-blown material. This lake expanded again, 
and a shoreline was established in the vicinity of 
Lake Millyera. Following subsequent development 
of full lacustrine conditions, the lake finally dried, 
and the watertable dropped well below the lake 
floor. 


Lunettes developed on this old lake bed, recor- 
ding the demise of permanent long term lakes in 
the region, The Billeroo Creek found its way 
through the barrier of aeolian and beach deposits, 
and cut down into the lake floor. Although runoff 
and sediment load appear greater than at present, 
aeolian deposition became a prominent feature. 
These deposits record a major drop in watertable 
indicating a significantly drier climate was establish- 
ed between about 0.1 and 2 million years ago. 

The last cycle of events records the building of 
the longitudinal dunes and origin of the Strzelecki 
Dunefield, though this is better recorded elsewhere 
(Callen ef al. 1983) in the region, only the later 
phase being present at Lake Millyera. This 
represents the effects of the last major glaciation, 
an event abundantly recorded throughout southern 
Australia (Bowler et al. 1982, Wasson 1983b). These 
dunes indicate strong directionally variable winds 
from a narrow westerly sector of the compass, a 
higher watertable than present, rather limited sand 
supply and more arid climate (Wasson 1983a, 
Wasson & Hyde 1984). 

At present, dune building is still active, though 
flooding of interdune corridors and formation of 
clay pellets is rare. Quartz sand dominates the 
dunes. Fluvial sedimentation on this side of Lake 
Frome is restricted to clays in suspension and locally 
reworked aeolian sand. Downcutting has been 
limited by failing streamflow and a hard sub-surface 
in the form of the dolomite of the Namba Forma- 
tion. Lake Frome very rarely fills to its present 
shoreline. 

Thus is recorded the demise of permanent 
brackish lakes in the northeast of South Australia, 
and development of an increasingly arid climate 
though with periodic fluctuations, at least partly 
due to world glaciations. Aeolian processes and 
ephemeral watercourses are now dominant. 


Acknowledgments 


Dr J. M. Bowler (Research School of Pacific 
Studies, A.N.U,) assisted under the SLEADS pro- 
gramme in trenching on section 16, and provided 
palaeomagnetic data for section 4. 

B. G. Forbes, G. W. Krieg and an unknown 
referee are thanked for constructive criticism of the 
manuscript. 

The Director-General, Department of Mines & 
Energy permitted publication of these data. 


QUATERNARY CLIMATIC CYCLES, LAKE MILLYERA 17 


tod 


References 


ASH, J. E. & Wasson, R. J. (1983) Vegetation and sand 
mobility in the Australian desert dunefield, Z. 
geomorph, 45, 7-25, 

Bower, J. M, (1981) Australian salt lakes, a 
palaeohydrologic approach, /n Williams, W. D. (Ed.), 
Salt Lakes. (Dr W. J. Junk Publishers: The Hague.) 

—— (1982) Aridity in the late Tertiary and Quaternary 
of Australia, /n Barker, W. R. & Greenslade, P. J. M. 
(Eds), Evolution of the Flora and Fauna of arid 
Australia, (Peacock Publens/Australian Systematic 
Botany Society/Anzaas (S.A, Div.); Adelaide.) 

—— (1983) Lunettes as indices of hydrologic change: a 
review of Australian evidence. Proc. R. Soc. Vict, 95, 
147-168. 

Hore, G. S., Jenninas, J. N., SINGH, G. & 
WALKER, D. (1976) Late Quaternary climates of 
Australia and New Guinea. Quat. Res. 6, 359-394. 

Burne, R. V., BAULD, J. & De Deckker, P. (1980) Saline 
lake charophytes and their geological significance, J. 
sedim. Petrol, 50, 281-293, 

CALLEN, R, A, (1976) FROME 1:250 000 scale geological 
map, sheet SH54.10. Explan. Noles geol. surv. §. Aust. 

——~ (1977) Late Cainozoie environments of part of nor- 
theastern South Australia. J. geol Soc. Aust. 24, 
151-169. 


Note added in proot 


—— (1983) The Islands of Lake Frome—an insight to Last 
Glacial climate in northeastern 8, Aust. Q. geal. Nates, 
geol, Surv. S. Aust. 88, 2-8. 

—— & Teprorp, R. H. (1976) New Late Cainozoie rock 
units and depositional environments, Lake Frome area, 
South Australia. Trans. R. Soc. S. Aust. 100, 125-168. 

——, Wasson, R. J. & GILLESPIE, R. (1983) Reliability 
of radiocarbon dating of pedogenic carbonate in the 
Australian arid zone. Sediment. Geal. 35, 1-14, 

Wasson, R. J. (1983a) Dune sediment types, sand colour, 
sediment provenance and hydrology in the Strzelecki- 
Simpson Dunefield, Australia. Jn Brookfield, M. FE, & 
Ahlbrandt, T, 8. (Eds) Folian sediments and PFOCeSSES. 
Developments in Sedimentology 38. (Elsevier: 
Amsterdam.) 

—— (1983b) The Cainozoic history of the Strzelecki and 
Simpson dunefields (Australia), and the origin of the 
desert dunes. Z. Geomarph. 45, 85-115. 

—— & Hype, R. (1984) A test of eranulometric control 
of desert dune geometry. Earth Surface Processes, 
Landforms. 8, 30)-312. 

WILLIAMS, D. L. G. (1982) Multiple episodes. of 
Pleistocene dune-building at the head of Spencer Gulf, 
South Australia. Search 13, 88-90, 


Coombes Springs should read Coomb Spring, based on latest information from S. Aust, Dept 
of Lands. The new rock unit name is, therefore, Coomb Spring Formation. The location of 
section 5 is 31°02'59" 139°56'36" not as given in Callen & Tedford (1976), 


MURPHY HAYSTACKS, EYRE PENINSULA, SOUTH AUSTRALIA 
BY C. R. TWIDALE & ELIZABETH M. CAMPBELL 


Summary 


Murphy Haystacks consist of two groups of large residual granite pillars and boulders located near 
the west coast of Eyre Peninsula, between Port Kenny and Streaky Bay. Many of the pillars and 
boulders have flared sidewalls and several are partly hollowed out through the development of 
tafoni. The gross forms were in existence at least by the later Pleistocene though there has been 
some slight further exposure during recent times. The pillars and boulders could be derived from the 
differential subsurface weathering and exposure of larg cubic or quadrangular blocks defined by 
orthogonal joint sets. Alternatively they could be remnants, again modified by subsurface moisture 
attack, of massive convex-upwards sheets of rock. Which of these possible origins applies to 
Murphy’s Haystacks cannot be unequivocally demonstrated, but on balance the evidence favours 
the second theory. 


MURPHY HAYSTACKS, EYRE PENINSULA, SOUTH AUSTRALIA 


hy CLR. TWibaALe & ELizABeiH M. CAMPBE! 1! 


Summary 


Pwibare, COR, & Campprr rn, LM. (LO8d) Murphy Haysticks, Fyre Peninsula, Soul) Auseratia. Tens, 
R. Suc. So hus MKB), 174-183, 13 December, (¥s4, 


Murphy Maysiueks consist of two groups of large residual granire pillars and boulders located near 
fhe west coust pf Eyre Peninsula, berween Port Kenny and Streaky Bay, Many of the pillars anc boulders 
fave Maree dewalls gid several ape partly hollowed oul (hroueh the development of (afoni.. The gross 
fornis Were in existence at leas! by the later Pleistocene though there has been some slight hither exposure 
clivia revent Times, Phe pillars and boulders could be derived trom the cifferential subsuriaee weathering 
und exposure of large cubie or quadrangular blucks defined by orthogonal joint sets, Alterqatively they 
could be reninants, again moditied by subsurface moisture attack, of Massive eonver-apwards sheels of 
rovk, Whivh of these possible origins applies to Murphy's Wavstucks cunnot be unequivocally demonstrated, 
bul on balance the cyidence favours the second cheory, 


Kiy Wrens) Pillans, granite formes, Byee Peninsula. 


Introduction 


IUis said that when couchmen on the old Streaky 
Bay-Elliston-Port Lincoln putt reached a point a 
few kilometres NW of Port Kenny, they used fo 
point oul to thei’ passengers a group of large resi- 
dual grarure pillars and boulders standing on the 
skyline near the ecest of a hill, and which loak like 
Ihe upper halves of hourglasses. Nol unreasonably 
they likened thers to haystacks or hayricks, and as 
They stood on the Murphy property they became 
known as Murphy's Haystacks, The name has en- 
dured, partly because of its origin in the romantic 
(if uncomfortable) clays of the borse and carriage, 
but also, it is supgested because ibe name is as 
descriptively apposite 4s can be found in the English 
languaue (see below), 

Located in the Calva district some 25 km NW 
of Port Kenny and 35 kum SE. ol Streaky Bay (Pig. 
1), Murphy Haystacks (Mig, 2) are a well knows 
focal landriark. They are becoming internationally 
known in Lhe popular literature because of their ode 
name, large size, iniricace sculpture, and jocinsie 
beuucy. 

The Haystacks offer many points of interest, in. 
cluding the origin of the gross forms. Although 
some of the residuals are Iree-stunding, most are 
apparciily contiguous projections of the granite 
that underlies (he hill on which they stand, and the 
questions arise ay to whether the nature of these 
subsuplice extensions are modified projections of 
quadranvular blocks or exposed portions of 
spheraidally weathered kernels or coresiones 
(Fig. 3), uid whether the forms are derived frum 
a ass of pranite dominated by orthogonal frac. 
ture sets or by ureuate, convex-pward sheet siruc- 


* Gimeraphy Depe, University of Adelaide, GPO. Box 
8, Adelaide, So Aust 


ture, The intricate sculpture of (he Haystacks may 
be due lo subsurface weathering or vo epigene at- 
tack. Agswers to these questions are (undamental 
1o.a debate concerning the origin of large residual 
boulders and related forms (see Linton 1955; Ring 
1958: ‘Twidale 1982), Orthogonal fractures are due 
(o shear Siresses and reflect regional tectonic style 
(Cloos 1936), whereas arcuate convex-upward sheets 
are variously interpreted as due to erosional of f- 
loading (Gilbert 1904), Or lo lateral compression 
thal is genetically related to crustal stress (| Twidale 
1964, 1973, 1982). 


Description 

The Haystacks consist of two areally separate 
though genetically related groups of pillars and 
boulders standing Hear, (hough not on, the crest of 
# convex-upward roughly cireular hill. They are 
about 40 m above the adjacent valley floors and 
100 m asd. (Figs | & 4). The granite from which 
the residuals are shaped is a pink, massive, coarsely 
equigranular rock consisting mainly of quartz and 
orthochase. Similar granites near Baird Bay and on 
the Investigator Group islands near Elliston are of 
Middle Proterozoic age and 1456+ 26 Ma old (Webb 
er ai, 1982'), The crest and flanks of (he hill 
however, though eroded in granite, carry a veneet 
ul calcrete, a pedogenic limestone that in the area 
under discussion is derived from dune calearenite 
ar ucolianite (Crocker 1946) and carbonate dust, 


‘Webb, A.W, Thomson, BP, Hlissert, A. HL, Daly 
So. Plint, Re B & Parker, A, he (1982), Geachronology 
of the Gawler Craton, South Australia, Depry Mines & 
Energy, Sout Australia, Rept. Bk Mo. 82/86 (unpubl), 


176 C. R. TWIDALE & E. M. CAMPRELL 


STREAKY 
ENLARGEMENT 


Streaky Bay 
1 
a Murphy's Haystacks 
S Port Kenny 


EYRE 
PENINSULA 


= 
Adelaide 


SCEALE 
BAY 
Murphy Haystacks, 
‘Oakfront’® 
Cape Blanche 


CALCA : 
a Cash Hill 


Freeman (Malijay) Hill” 


Pe 
Te, 
oy 
he} 


ott —_ 


» ey eel *eeNere ee ee 


= e 
rt 


_ s 
es * 


ae o 
hat iel * 


A ee 
Lae 


‘ 
te 
ais oe 


Fig. 2. General view of Murphy Haystacks (eastern 
group) seen from the east (Port Kenny—Streaky Bay 
road). 


The term ‘boulder’ is suitable for those rounded 
masses that are wholly exposed and detached but 
it cannot be used of the many others that merge 
unbroken with the underground rock mass. They 
seem to be attached to a solid cohesive base but, 


«Mount Hall 


e 
Calca Quarry 


as stated previously, it is not known whether they 
merge with the base of a corestone or with an 
essentially unaltered block (Fig. 3). In this cir- 
cumstance it is preferable to use a descriptive and 
non-genetic term, and it is suggested that the appa- 
rently attached forms be called ‘pillars’, used in the 
sense of upright columns, and additionally to bor- 
row the colloquial term ‘haystack’ for those pillars 
with sidewalls so flared that they widen appreciably 
from the base to the upper shoulder. 

Both groups of pillars and boulders are of King’s 
(1958) “skyline” type for though they are not on 
the crest of the hill, they stand high on the convex- 
upward sideslopes. The eastern group is smaller 
than the western, but the two comprise similar suites 
of forms. Both consist mainly of large pillars up 


MURPHY HAYSTACKS 177 


Heeler shit 


ee ae 
(a) 
etn 
4 ‘ 
a ~s 
\b) 


Tig 1. Alternative explanations at haystacks: (a) boulder 
(1) anid pillar (ii) derived from granite dominated by or- 
(hogunal fracture sets; (b) bowlders anu pillars from 
granite subdivided by arcuate, eouves-upward sheeting 
JONES, 


to 1 om tight aud characteristically with Mared 
sidewalls (Pie 4), Some, and two ui the eastern 
group in particutar, fave well developed lafori, or 
jiollows, One of We two is nereworthy for irs steer 
size (Fig. 6), The other because vertical grooves 
(Rillen or Nurren) are laintly developed on the 
overhaniing inner wall of the hollow, Some of the 
boulders are similarly modified, and one of the 
laiwe boulders in the western group displays burh 
Hares ond a larve tafone with ribs protected by 
hiehen. Less common are blocks that have been only 
slightly modified by weathering, giving rise to 
tabular lorms, 

These pillars, boulders and blocks are the basic 
forms but there are many variations, combinations 
und modifivations, One pillar ia the eastern group 
is tall and nasrow and resembles an hourglass. Some 
(l(devounded blocks. stand squarely ane upon the 
Olher to form cottage loaves. Some blocks have 
sidewalls so weuthicred and flared that quite Wide 
platforms are developed around their bases, and in 
extreme cases the plathorms are so wide and the een- 
tral pillars so reduced in height as well as diameter 
tharthe whole looks like a boss and shield (Pig, 7 
With further weathering the pillar las been wholly 
eliminated to produce a platform flush with the 
vround surface (big. 8) Many of the bloeks and 
pillars retain secondary fractures within their 
masses, and many carey detached shells. 

Thouth apparently haphazard in their distribu. 
LOK, Mappings stows thar marly of lhe boulders are 
mintly defined by fractures (hug. 49, Moreover many 
are in play disposed in orderly fashion with respect 
to imlerseeting fracture sets. In the eastert group 
[he major fractures trend SSE-NNW (13521607) 
and WNE-SSW (20°25"), with an east-west set also 
present. tn che western! group however the form and 
cisposinion uf the residuals are related to two iitter 


secling urcuale sels disposed i tadiating fan-lke 
palterns with axes trending roughly east-west and 
SW-NE (Fig. 4), 


Age of the Forms 

The plains and slopes separating and surroun- 
ding the granitic residuals are covered by a veneer 
Of valerete derived from caleurenite (the agolianite 
oF dune sand of Crocker 1946) and carbonate dust. 
Though in many places wash from the residuals has 
caused the calerete to be dissolved, creating annular 
depressions around the bases of the pillars and 
haystacks (ef, Jennings 1973), (he culeréte essentially 
laps up against the bases of the forms. Moreover 
al one site calcrete with fossil foraminifera 
fagments occurs within a hollow formed along the 
Joint that cuts through the base of the residual, This 
limestone and one from nearby Freeman (Malijay) 
Hill have been dated by the Cl4 method (Gak-5266 
and 5267) as of late Pleistocene age (Twidale er al. 
97H). Thus there is the Suggestion thal the 
Huystacks were essentially in existence before the 
Pleistocene dune calearenites and pedogenic 
calereres. Some ol the miner as well as the grass 
lorms Nave been exhumed from beneath the dune 
wover (of. Jack 1912), 

Many blocks and pillars stand on low plinths, 
whose pilted surfaces indicate recent exposure 
(Twidale & Bourne 1976x). Ina few cases valeium 
carbonate has impregnated the granite exposed at 
the bases of the residuals. This pilling and the steep- 
sided, lowermost slopes of the pillars (Fig. 9) argue 
recent exposure through soil erosion of approxi- 
mately 25 cm (probably following clearance of 
vegetation by Eurapean settlers, follawed by 
ploughing and pastoralism), but the gross forms of 
Murphy Haystacks predate the late Pleistocene 
carbonates and can be regarded as partly exhumed 
fran’ beneath those carbonates, 


Origin of ihe Farms 

The problems: Whether the large residuals of 
Murphy Haystacks are pillars or boulders, the pro- 
blem of their rounding remains the same. [t has long 
been recognised and accepted ryat some boulders 
owe their rounding to abrasion by rlyers and waves 
but, equally, chat Weathering is differential and 
causes the conversion of angular io rounded forms. 
As MacCulloch (1814 p. 74) noted “Nature aiuvar 
yuadrata rotundis*’: granite blocks have been 

“rendered spherical by decomposition” as a result 
of fhe more rapid attack by moisture on corners 
und edges than on plane faces. The problems eon- 
eerning the pillars al Murphy Haystacks are 
Whether The lwo upper corners of a block have been 
st rounded or whether all four have been allected 
(Viv, 3), and whether the fractures exploited are of 
orthogonal or sheeting sets, 


178 C. R. TWIDALE & E. M. CAMPBELL 


© pillar 
height inmetres (approx.) 
flared slope 
pronounced flare 


» tafoni 


= gnamma 


joints (mainly inferred) 
loose boulder 

boss and shield 
whaleback 


platform 
"~~-= crest of hill 
+—- fence 


Ometres 


6 Cte 


ac 
2 Be 2 


Fig. 4. Plans of Murphy Haystacks, and contour plan of the area showing also locations of plans of eastern group 
(area A) and western (area B). Contour plan adopted from S. Aust. Lands Dept. 1:50,000 series Calca 5731-1. 


Fig. 5. Prominent haystacks at eastern margin of western 
group seen from east. Hill crest is to left. Note that 
flared sidewalls are better developed on upslope (left) 
side; also isolated horizontal fracture. 


The question of whether the weathering took 
place beneath the land surface as suggested by 
Hassenfratz (1791) and many others (see Twidale 
1978) through to Linton (1955) or after exposure 
as suggested by King (1958), is readily resolved, for 
most of the pillars and boulders are in some degree 
flared. 

Flared slopes are a particular form of the 
weathering front (Mabbutt 1961) which may in 
general terms be defined as the lower or lateral limit 
of significant weathering. Flared slopes are best 
developed in piedmont or scarp foot situations, or 


MURPHY HAYSTACKS 179 


3 


Fig. 6. Large boulder tafone, eastern group. 


ES a i 


lO ee 


Fig. 7. Boss and shield, eastern group. 


Fig. &. Platform in western group, 


along open joints in massive rocks, and in tec- 
tonically stable regions (Twidale 1962, 1972): 
anywhere there is a long-term concentration of 
water in a rock type that is altered in such a way 
as to produce an abrupt transition from weathered 
to unweathered rock. Granite is an eminently 
suitable medium in most respects. Water running 
off the hill or large boulder permeates into the rocks 
at the base of the residual. There, because of water 
retention, chemical weathering is enhanced. Also, 


Fig. 9. Base of flared haystack with narrow platform and 
steep basal slopes; evidence of recent soil erosion from 
surrounding plain (eastern group). 


because of surface desiccation the weathering front 
advances laterally more rapidly at depth than at the 
surface (Fig. 10). Thus when as a result of lower- 
ing of the plains the weathered detritus of the pied- 
mont is evacuated, the exposed weathering front is 
concave in form. The essential feature of this ex- 
planation is two stage development, the first con- 
sisting of subsurface weathering, the second ex- 
posure by erosion. Incipient flares, in the form of 
concave weathering fronts still beneath the natural 
land surface have been observed in excavations at 
Yarwondutta Rock, Chilpuddie Hill and several 
other sites on Eyre Peninsula (see e.g. Twidale 1962, 
1982, pp. 243-257). 

The common occurrence of flared sidewalls on 
the pillars and boulders at Murphy Haystacks 
shows that the granite blocks on which the forms 
are based were subjected to weathering beneath the 
land surface. The moisture attack was directed along 
vertical and near-vertical joints and the weathering 
was most effective 6-8 metres beneath the land sur- 
face. Some of the tafoni that are evident at the 
Haystacks may have been initiated in the same man- 
ner; they may be flares at which weathering has 
been especially rapid (perhaps by reason of minera- 
logical or structural weakness), though there has 
undoubtedly been development after exposure (see 
Bradley ef al. 1978; Twidale 1982). 


Orthogonal or Sheet Jointing: Evidence and 
Argument 

The various major forms and some of the more 
prominent minor features present at Murphy 
Haystacks have their origin in the shallow subsur- 
face. But what was the structural base: did subsur- 
face moisture attack affect orthogonal or sheeting 
joints? The exposed residuals vary in size, indicating 


180 C. R. TWIDALE & E. M. CAMPBELL 


F 
+ + + + 
tote Ot Ft 
ering front + + 
+ + + + 


= 

9 

= 
++ 


+ + 
+ + + + + eH + + + 
+ + 


Fig. 10. Development of flared slopes. 


either variations in original fracture spacing or in 
the intensity of weathering, or in both factors. No 
pattern is evident. The variations are equally, if in 
rather general terms, explicable in terms either of 
orthogonal or of sheeting sets. Some arguments and 
evidence are however diagnostic. 

Lack of basal separation: Although some pillars 
have horizontal or sub-horizontal fractures at or 
near their bases, most do not: they merge without 
physical break with the underlying granite. This is 
equally explicable in terms either of corestone 
development in the context of orthogonal blocks, 
or in terms of the weathering of radial fractures 
developed within sheet structure. No grus, or 
weathered granite, has been found beneath boulders 
or pillars, but this may be due to a lack of suitable 
exposures. 

Location on hill: That neither group of pillars 
and boulders stands on the crest of the host hill may 
be significant. If the pillars were developed from 
orthogonal joint blocks the residuals would on the 
one hand reasonably be expected to survive longest 
on the crest of the hill, most distant from the erosive 
effects of rills and streams that before the accu- 
mulation of calcrete would have coursed down the 
slope. On the other hand, the larger corestones 
might be expected to occur on the lower hill slopes 
because the deeper zones would not have been sub- 


jected to moisture attack for as long a time as those 
near the surface. Yet in reality the surviving pillars 
and boulders occur just below the crest. It may be 
that the orthogonal fracture sets are nonpervasive 
and are heterogeneously distributed within the 
hillmass. The hill crest may be upstanding by vir- 
tue of such a lack of open fractures. On the other 
hand the pillars and boulders, though not on the 
crest are quite close to it. In any case even if the 
distribution of fractures is uneven, strain patterns, 
being regional, are not; and such strain zones are 
as readily exploited by weathering as are fractures. 
An alternative explanation is that the hill is 
underlain by sheet structure. In these terms the 
crestal zone, being antiformal, would be in tension 
and therefore vulnerable to weathering by water. 
The arcuate fractures on the other hand allow per- 
colation of water into the synformal (valley) zones 
that for that reason are deeply weathered, so much 
so that no corestones or masses of fresh rock have 
survived, In other words the plan distribution of 
the pillars is consistent with the sheet structure 
concept. 


Survival of minor forms on crest of residuals: At 
Caloote, in the eastern piedmont of the Mt Lofty 
Ranges, some 65 km E. of Adelaide, a rounded 
granite hill carries several large sculptured granite 
blocks notable for their roughly flared sidewalls (the 


Fig, 11, Residual block with crestal grooves, Caloote. 


MURPHY HAYSTACKS IMI 


eTanile is coarse seained) and (he odd shapes that 
have resulted from subsurface moisture attack. For 
jostance one bloek is shaped hike awn anvil, and has 
preserved on its gently rounded crest a ghajma ot 
rock basin. The crest of another is scored by several 
parallel tullers (har rum entirely across che crest (Fig. 
1). Such minor forms as basins aud gurters are 
characterisne of the gently anelined slopes of in- 
selhetes cul in massive rock, and remnants of such 
forms a Che crests Of pillars and boulders at ihe 
site under discussion would point to 1beir being 
fragments of a disintegrated dome, A search was 
mude of the crests of the pillars and boulders at 
Murphy Haystacks but only one large squat pillar 
(Xin Pig. 4) has a snamma developed on iis ¢resi, 
and an the same residual there are two putters. Oue 
is clearly Pravture-vaonourelled and van. be discounted 
in the present context, The other is partly controll. 
cd by structure (iL runs along several disconlinuous 
veins) though it does diverge in places and especially 
Toward the edve of the pillar But it-does not rin 
across the residual, as do those at Caloote, and 
thoumh if may have had its origin On rhe mare et 
repisive massive sloping Subtace, i jay plot. Similarly 
jhe rock basin may have evolved on a structurally 
Weak site and does nol eonstiule undeniable prool 
that the Haystacks are denved trom the 
disimegrabion af a domieai inselberg, 


fay 12 Preferred develupinent of flares an opposed 
aspects. 


fractures: UW the etanite mass beneath the bill 
were subdivided by orthogonal sets then some 
Nhonzautal or subhurizontal members ought te be 
ievidenke Such hanzantal fractures as do occur 
are notably localised, frequently restricted to one 
pillar and absent from (hose nearby (see eg Fip. 5), 
The verlical merthers ought to run in parallel. i 
on the other hand the hill underlain by sheet 
structure the secondary jolts i section Ought to 
form o radiating or fan-like pattern, Unfortunately 
most of the (ats are covered hy rubble of calcrete 
anid toc few joints are exposed for a patrerti to be 
plotted, (hough same indirect evidence on This point 
has been deduced (see below), On the alher hand 


the convex-upward form of the hill is suggestive of 
determination by sheet structure, and some of the 
subhorizontal {ractures present. are associated with 
(nangular wedges that are typically associated with 
sheel structure (Twidale 1964, 1973, 1982). Similar 
wedwes ovcur on the crests of some pillars, They 
are related to differential rnovement along the 
sheeting planes and ultimately to the rock musses 
beme in stress (Twidale 1964, 1982; Twidale & Sved 
1978). One sueh horizontal fracture in one of the 
prominent haystacks of the western group has such 
(rianigular wedges on both sides of (he residual, that 
on the northern face being single but those on the 
southern being complex and in detail consisting of 
at luast six distinet wedges (cf. the complex wedge 
at Ucentitehie Hill} Twidale 197], p. 71). Such frac 
ture pallerns are readily explained by a single phase 
of differential movement, as are other wedves, bur 
they could relleet vertical pressure, 


Flares and aspeck Given a vertical (racture that 
contrals water percolation and the development of 
flates, all else being equal flare development ought 
to be similar on both sides of the subsequent 
fracture-controlled cleft. IF however the fractures 
were inclined, and given that water percolates under 
graviry, the upslope facing side of the fracture ought 
(a be more weathered and develop a more pro- 
nounced flare than that associated with the 
overhang facing downslope (Fig, 12). 1f the frac- 
ture system (hat controlled weathering were or- 
thoganal and vertical then there ought to be no 
preferred distribution of flares, Conversely if the 
jointing is orthogonal but inclined then flares will 
be consistently asymmetrically distributed, If om the 
other hand the fractures are part of fan sets 
associated with sheer structure then there ought ro 
be contasted asymmetry on opposed lanks of the 
hall 

In the field there is a tendency for there ta be a 
greater development of Flares on the upslope-facing 
side of pillars, and for that tendency to be in the 
reverse aspect On Opposite sides of the hill (see e.g. 
Fig. 5) and though there are many exceplions, and 
evell great variations of fare development on the 
same aspect on adjacent blocks, (here are many 
assemblages that suggest sheet structure and radial- 
ing Secondary fractures rather than orthogowal sets. 


Lvidence from nearby hills; Since tracture patterns 
in erystalline rocks such as erarite are arguably due 
10 regional stress, Comparisons with adjacent 
uplands are relevant to |he problem of Murphy 
Haystacks.. Some of the nearby hills offer better 
exposures af the granite than can be obtained at 
Murphy Havstacks. Thus at Cash and Freeman 
hills (Fig. 1) ealerere ts preserved on the hill crests 


182 C. R. TWIDALE & E. M. CAMPBELL , 


and lower slopes but there are extensive outcrops 
of granite at midslope. The Calca Quarry also pro- 
vides excellent exposures of another granite rise. 
And it is clear from all of these that the granite mass 
is dominated by arcuate, convex-upward sheet struc- 
tures, that have disintegrated to give orthogonal 
blocks. It is notable that at Freeman Hill there are 
large residual boulders protruding above the late 
Pleistocene calcrete, though they are neither as large, 
nor as numerous, nor as intricately sculptured. But 
these more southerly hills are slightly higher than 
Murphy Haystacks and possibly did not receive 
the same thickness of sand cover, so that subsequent 
erosion has been more effective and allowed exten- 
sive exposures of the granite country rock, 

The suite of forms at Murphy Haystacks is 
similar to that which would develop if a domical 
inselberg or bornhardt like Ucontitchie Hill or Mt 
Wudinna were partially buried by dune sand, and 
particularly calcareous sand. Ground water per- 
colating through such material would become 
alkaline and so more actively attack the granite (e.g. 
Alexander ef a/., 1954; Twidale 1979). The crests of 
Mt Wudinna and Ucontitchie Hill are boulder and 
block-strewn, but the mid slopes are essentially 
smooth, being largely an expression of sheet struc- 
tre. On the other hand, though largely similar, there 
are important differences in detail between this 
assemblage and some of those, such as Richardson 
Rocks, described from the Upper South East 
(Twidale & Bourne 1975b; Twidale ef a/. 1983). It 
may be that Richardson Rocks, standing and devel- 
oped by differential weathering beneath a plain, 
show water table effects in greater degree, in for in- 
stance the common development of low tabular 
forms. 


Conclusion 


The groups of large residual boulders and pillars 
that are Murphy Haystacks stand on the upper 
slope of a convex hill underlain by granite but with 
a veneer of calcrete derived from dune calcarenite. 
Though not conclusive, the weight of evidence and 
argument suggests that the boulders are remnants 
of a sheet structure, The massive sheet is subdivided 
by fan joints. Water penetrated down these, caus- 
ing the widening of the fracture clefts and the for- 
mation of flared slopes on the bounding walls dur- 
ing subsurface chemical weathering. Erosion of the 
weathered granite or grus allowed exposure of the 
sculptured blocks. This took place prior to the 
spread of coastal dunes during the Pleistocene. The 
dune calcarenite spread far inland and buried not 
only many of the hills of the Calca district (though 
it is not known whether the pillars and boulders 
at Murphy Haystacks were totally overwhelmed) 
but also a granitic landscape of considerable relief 
that occupied much of the northwestern Eyre Penin- 
sula. It is clear however, that the calcarenites and 
calcretes at least lapped around the bases of the 
residuals, Following an amelioration of climate and 
rise of sealevel, much of the dune material was 
eroded, though some was precipitated to form 
calcrete, and soil erosion of some 25 cm took place, 
but only the topsoil was stripped, exposing the 
calcareous duricrust that has essentially stabilised 
the local land surface. 


Acknowledgments 


The authors thank Dennis and Mary Cash, of 
‘Oakfront’ for their hospitality and interest in the 
research on which this paper is based. Some of the 
field work was funded by awards from the 
Australian Research Grants Committee, and the 
Australian Research Grants Scheme. 


References 


ALEXANDER, G. B., Heston, W. M. & ILER, R. K. (1954) 
The solubility of amorphous silica in water. J. Phys. 
Chem. 58, 453-455. 

Brapbtey, W. C., Hutton, J. T. & TwipAce, C. R. (1978) 
Role of salts in development of granitic tafoni, South 
Australia. J. Geol. 86, 647-654. 

Coos, H. (1936) Plutone und Ihre Stellung im Rahmen 
der Krustenbewegungen. /6th Int. Geol. Cong. Rept. 
1, 235-253. 

Crocker, R.L. (1946) Post-Miocene climatic and 
geologic history and its significance in relation to the 
genesis of the major soil types of South Australia. Bull. 
Coun. Sci. Indusir. Res. Aust. 193. 

GILBERT, G. K. (1904) Domes and dome structures of the 
High Sierra. Geol. Soc. Amer. Bull. 15, 29-36. 

HaAsseNFRATZ, J-H. (1791) Sur arrangement de plusiers 
gros blocs de differentes pierres quo l’on observe dans 
les montagnes. Ann. Chimie. 11, 95-107. 


Jack, R.L. (1912) The geology of portions of the 
countries of Le Hunte, Robinson and Dufferin, with 
special reference to underground water supplies. Geol, 
Surv. S. Aust. Bull. 1. 

JENNINGS, J. N. (1973) The disregarded karst of the arid 
and semiarid domain. Karstologia 1, 61-73. 

Kina, L. C. (1958) The problem of tors. Geogrl J. 124, 
83-87. 

Linton, D. L. (1955) The problem of tors. /bid. 121, 
470-487. 

Massutt, J. A. (1961) “Basal surface“ or “weathering, 
front”. Proc. geol. Assoc. London 72, 357-8. 


MacCuttocn, J. (1814) On the granite tors of Cornwall. 
Trans. geol. Soc. 2. 66-78. 

TwibaLe, C, R, (1962) Sieepened margins of inselbergs 
from north-western Eyre Peninsula. Z. Geomorph. 6, 
51-69. 


MURPHY HAYSTACKS 183 


(1964) Contribution to the general theory of domed 
inselbergs. Conclusions derived from observations in 
South Australia. Trans. & Papers Inst. Brit. geogr. 34, 
91-113. 

—— (1971) Structural Landforms. (Austr. Natl. Uniy. 
Press: Canberra.) 

—— (1972) Flared slopes, scarp-foot weathering and the 
origin of the piedmont angle. S. Afr geogr. 4, 45-52. 

(1973) On the origin of sheet jointing. Rock 
Mechanics 5, 163-187. 

— (1978) Early explanations of granite boulders, Rev. 
Géomorph. Dynam, 27, 133-142. 

— (1979) The character and interpretation of some pedi- 
ment mantles. Sediment. Geol. 22, 1-20. 

— (1982) Granite Landforms. (Elsevier: Amsterdam). 


& Bourne, J. A. (1976a) Origin and significance of 

pitting on granite rocks. Z. Geomorph. 20, 405-416. 

& —— (1976b) The shaping and interpretation of 
large residual granite boulders. J. geol. Soc. Aust. 23, 
371-381. 

—, & SmiTH, D. M. (1976) Age and origin of 
palaeosurfaces on Eyre Peninsula and in the southern 
Gawler Ranges, South Australia. Z. Geomorph. 20, 
28-55. 

——, CAMPBELL, E. M. & Bourne, J. A. (1983) Granite 
forms, karst and lunettes, pp. 25-37 in M. J. Tyler, C. R. 
Twidale, J. K. Ling & J. R. Holmes (Eds) “Natural 
History of the South East” (R. Soc. S. Aust: Adelaide). 

— & SveD, G. (1978) Minor granite landforms asso- 
ciated with the release of compressive stress. Aust. 
geogrl. Studies 16, 161-174. 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 108, PART 4 


DASYUROTAENIA ROBUSTA BEDDARD, 1912, AND D. DASYURI SP. 
NOV., FROM CARNIVOROUS AUSTRALIAN MARSUPIALS 


BY IAN BEVERIDGE 


Summary 


Dasyurotaenia robusta Beddard, 1912, is redescribed from specimens collected from the type host, 
the Tasmanian devil, Sarcophilus harrisi (Boitard). The rostellar hooks are described for the first 
time. Dasyurotaenia dasyuri sp. nov., from the tiger cat, Dasyurus maculatus (Kerr), in Queensland 
and Tasmania differs from D. robusta in the size and shape of the rostellar hooks, the presence of 
transverse osmoregulatory canals and the number of uterine branches. Specimens from Dasyurus 
maculatus, described in earlier works as D. robusta can now be assigned to either species with 
certainty. Lesions associated with D. dasyuri sp. nov. are described and the taxonic position of the 
genus discussed. 


DASYUROTAENIA ROBUSTA BEDDARD, 1912, AND D. DASYURI SP, NOY,, 
FROM CARNIVOROUS AUSTRALIAN MARSUPIALS 


by LAN BEVERIDGE* 


Summary 


Bevekibok, 1, (984) Dusyarolgenta robustly Beddard, 1912, and 2. dase sp. flow, (Onn CArAibecaents 
Australiaiy marsupials. Trym, R, Soe, 5) Aus? 10804), 185-195, 13 December, 1984, 


Dasvurolgenta robusta Veddard, 112, is redeseribed tran specimens collecred from The (ype Hest, 
the Tasmanian devil, Sureophilus harrisii (Boitard)., The rosiellae hooks are deseribed for the first titre. 
Dasyvurotaenia dasyun sp. poy., (rom the wer eat, Dasvurus maerduis (Ker), 1 Queensland and lasmauie 
differs trom 1. rebusta inthe size and shape of the rostellay hooks, the presence of (ransverse osmoregulatory 
canals and Lhe number Of ulerine branches, Specimens from Dusyarus Hacklatios, described V0 carer works 
a5 DP. robusta cannot now be assigned to either species with certainty. Lesions assaciated with D. dastiri 
gp nov, aire deseribed and the faxenemie posibien of the gents discussed. 


Kiy Words; Cestoda, Taenidae, marsupials, Dasvaromwentu. 


Introduction 


Dasyurovteenia robusta was lirst deseribed by 
Beddard (1912) from specimens found in a Tasma- 
tian devil, Sareophiliis frarrisit (Bojrard) 
( Dasvarus ursinusy which died in London at the 
Gardens of the Zoological Society, Beddard (1912) 
desenbed a number of unusual morphological 
features including suckers armed with hooks, and 
placed the species, with some reservations, 1 the 
Taenioidea. Baer (1925) re-examined Beddared’s 
types and indicated that a number of misinterprera- 
tions of the morphology of the cestode had been 
made, Tncluding the “armed suckers” which proved 
to be an armed rostellum, Baer (1925) concluded 
that the species belonged to an independent! genus 
within the Taenioidea, while Wardle & McLeod 
(1952), with considerable reservation, placed the 
genus within the family Taeniidae, 

Subsequently, Sandars (957) redeseribed the 
apecies bused On cestodes collected from two 
Dasyurus maculatus (sert) trom Tasmania, conti 
ming most of Baer’s (1925) observations and eon- 
eluding thar the genus did belong within the 
Taeniidde, This taxonomic position was accepted by 
Yarnagutr (1959), but at has been questioned by 
Rausch (1981) on phylovenetic grounds, 

Recent collections of cestodes from dasyurids in- 
dicule thal two tidependent spevies of Daxvure- 
faenia have been formerly confused under a single 
gpeertic name, largely because Lhe rostellar hooks 
of the Species have never described (Beddlard 1912, 
1918; Baer 1925; Sundars 1957). In uddition, a re- 
examination of the morphology of the two species 
supports Rauseh’s contention (M981) that this genus 


* Dieision af Velorimary Sefenees, South) Australian 
Department of Avticullure, evo boscitute ob Medical 
and) Veterinary Seienee, brome Road, Acelitide, 
Soule Ausrralia, 


may not belonp to the laeniidae. Lt this pauper, 
Dasvurotaenia rabuvid is redescribed (rom Sea 
cophilus harrisii, We type host, and a be spectes 
is deseribed frank Daxyvurws macudaly 


Materials und Methods 

Cestodes were relaxed in water; fixed in 10% 
neutral bullered formal salting, and stared in FOM 
ethanol, Whole maynts were stained with Celestine 
blue, dehydrated it graded cthanols, cleared im clove 
oil and mounted in balsam. Scoleces were mounted 
in, Berlese's fluid, and digital pressure wits applied 
to the cover slip to enable examination ar the 
rostellar hooks. Serial sections cut at a@ thickness 
ofS om, were stained with haematoxylin any casin, 
Gravid proglouides of D. dasvuni whieh Had heen 
fixed in formalin were diced into sovall cubes, post- 
fixed in osinium tetroxide and embedded in araldin. 
Thin sections. were stained with lead cilrile any 
uranyl acetate and viewed witha Joel Od CX elec. 
(ron microscope. Additional specimens of Dasviyre- 
laenia were Obtained front preserved earcasses ol 
Dusyurus neeniyiis held inthe National Museums 
of Vieloria., Methourne 

Measurements are given in the lexi, in mim, as 
the ranve followed, int pareniheses, by the mean and 
the number of measurenients mude. 

Abbreviations of institutions cited in text 
AHC Auseruliin Helminth Collection, housed in 
the South Australian Museum, Adelaide, BMNH— 
British) Museuiy (Natural History), London. 
MHNCi—Muséum d'Histoire Naturelle, Cieneva, 
SAM—South Atstralian Manseurny, Adelaide. 
WAM—Western Australian Museunr Perth, Wi— 
Commonwealth Suicim Tie and Iudusieiael Research 
Orgunisuvion, Division of Wildlife and Rangelands 
Reseurch, Canberra. 


IRG 


Dasyurotaenia robusta Beddard, 1912 
FIGS 1-11, 26-28 

Description: Cestodes of moderate size, up to 140 
in length, 4 wide with up to 290 proglottides in 
gravid strobila. Scolex large (Fig. 1), 2.36 (n=1) in 
diameter, deeply embedded in intestinal mucosa of 
host. Suckers 0.30-0.38 (0.33, n=3) in diameter; 
rostellum 0.44 x 0.15 (n=1) retracted within scolex, 
extremely muscular, sucker shaped, with 42 (n=1) 
rostellar hooks arranged in 2 rows. Large or anterior 
rostellar hooks (Figs 2-4, 26) 0.046-0.058 (0,054, 
n=10) long, base 0.056-0.062 (0.059, n=10) long; 
blade large, core striated, sometimes vacuolated; 
handle extremely short, relatively wide; guard long, 
wide, single lobe (Figs 4, 27). Small or posterior 
rostellar hooks (Figs 5~7) 0.042-0.052 (0.047, n=10) 
long, base 0.044-0.060 (0.054, n=10) long; blade 
large, core striated; handle extremely short, knob- 
like; guard large, flattened, almost bilobed distally 
(Figs 7, 28). Neck present. 

Musculature of mature proglottides poorly 
developed. Outer longitudinal muscles single or in 
very small bundles; inner longitudinal muscles in 
larger bundles containing 20 or more fibres. 
Transverse muscles in several bands; including bands 
internal to inner longitudinal muscles and 
separating inner and outer longitudinal muscles; 
more poorly defined bands between bundles of in- 
ner longitudinal muscles. Dorso-ventral muscles 
sparse, crossing cortex and medulla at irregular in- 
tervals. Longitudinal osmoregulatory canals paired; 
ventral canal 0.10-0.19 (0.13, n=5) wide in mature 
proglottides, not joined by transverse canals, with 
valve-like flaps protruding into lumen at junction 
of proglottides; in one strobila, ventral canals of 
gravid proglottides with several smaller projections 
of canal wall in addition to major valves. Dorsal 
canal extremely narrow, sinuous, 0.05 (n=1) in 
diameter in mature proglottides, dorsal or external 
to ventral canal. Mature proglottides 0.35-0.60 
(0.48, n=5) * 2.45-3.05 (2.77, n=5), length:width 
ratio 4.8-7.1 (6.0, n=5) (Fig. 8), Gravid proglottides 
1.45-2.20 (1.75, n=5) x 2.60-3.50 (3.18, n=5), 
length:width ratio 1,2-2.3 (1.9, n=5). Genital pores 
almost exclusively unilateral, occasional genital pore 
on alternate side. Genital atrium narrow, situated 
in middle of lateral proglottis margin in mature pro- 
glottides dividing margin in ratio of 1:0.67-1:1.00 
(1:0.88, n=5); in middle or posterior half of margin 
of gravid proglottides, dividing margin in ratio of 
1:1,00-1:1.66 (1:1.35, n=5). Genital ducts pass 
between longitudinal osmoregulatory canals. Cirrus 


I, BEVERIDGE 


sac elongate, thin-walled, invariably extending 
beyond osmoregulatory canals into medulla, 
0.60-0.85 (0.74, n=10) * 0.06-0.09 (0.07, n=10) 
in mature proglottides. Cirrus slender, approxi- 
mately 0.01 in diameter, coiled, armature of 
extremely fine bristles visible on mid-region of cirri 
in section. Internal and external seminal vesicles 
absent. Vas deferens greatly coiled, narrow duct, 
loops medially, then at midline turns posteriorly, 
terminating between lobes of ovary. Vasa efferentia 
not seen. Testes numerous, situated in 1-2 layers in 
dorsal plane. Testes occupy most of medulla 
between osmoregulatory canals, occasionally extend 
over osmoregulatory canals on poral side of pro- 
glottis with small numbers of testes being outside 
canals (Fig. 8); testes confluent anterior to ovaries 
and frequently confluent posterior to vitellarium 
in | or 2 rows; row of testes posterior to vitellarium 
sometimes interrupted: always some testes posterior 
to vitellarium; small numbers of testes overlie 
ovaries. Testes number 170-223 (200, n=10) per pro- 
glottis; diameter 0.05-0.10 (0.08, n=10). Vagina 
0.010-0,020 (0.015, n=5) in diameter, straight, lin- 
ed internally by hairs or bristles, surrounded by 
single layer of glandular cells. Proximal of 0.16 of 
vagina of wider internal diameter, unarmed. 
Seminal receptacle small, 0.06-0.08 (0.07, n=5) x 
0.02-0.04 (0.03, n=5), situated in mid-line between 
lobes of ovary (Fig. 10), Ovary bilobed, poral lobe 
smaller, 0.14-0.32 (0.21, n=10) x 0.21-0.40 (0.29, 
n=10), aporal lobe 0.16-0.30 (0.23, n=10) x 
0.27-0.48 (0.38, n=10), joined by narrow isthmus. 
Vitellarium posterior to ovary, elongate laterally 
0.07-0.14 (0.11, n=10) x 0,47-0.90 (0.70, n=10). 
Mehlis’ gland spherical, 0.08-0.10 (0.09, n=5), in 
diameter, between vitellarium and seminal recep- 
tacle. Uterus arises as tubular structure in midline. 
Uterus in gravid proglottides with 6-9 (7, n=10) 
poral and 7-10 (9, n=10) aporal lateral uterine bran- 
ches; uterine branches frequently subdivided lateral- 
ly (Fig. 11). Eggs approximately spheroidal 
0.033-0.048 (0.042, n=10) x 0.035-0.040 (0.038, 
n=10)(Fig. 9); embryophore thick, homogenous, 
non-striated, oncosphere 0.028-0,033 (0.030, n= 10) 
x 0.023-0.030 (0.027, n=10); oncospheral hooks 
0.008-0.010 (0.009, n=10). 


Development of genital organs in single specimen 
140 long: testes first visible in proglottis 95; first 
mature proglottis approx, 160; uterine filling com- 
mences in proglottis 190; male and female genitalia 
involuted by proglottis 230; fully branched uterus 


Figs I-ll. Dasyurotaenia robusta Beddard. |. Scolex with rostellum retracted; 2-7, rostellar hooks; 2, 3, large or anterior 
rostellar hooks, lateral view; 4, large rostellar hook, radial view; 5, 6, small or posterior rastellar hooks, lateral view: 
7, small rostellar hook, radial view; 8, mature proglottis; 9, egg; 10, female genitalia; 11, gravid proglottis, Scale lines: 


fig. 1, 


1.0 mm; figs 2-8, 11, 10, 0.1 mm; fig. 9, 0.1 mm. g—guard; h 


handle; (—tip. 


or 


DASYUROTAENIA SPB FROM MARSUPIALS 


I. BEVERIDGE 


188 


PASYUROTAENI A SPP PROM MARSUPLALS au 


present by proglotts 250; first gravid proglortys 278; 
total Number proglottides 290, 

Host: Sareophilus harrtsii (Bowward, 1841) (Mare 
supialia; Dasyuridae). 

Sue in hosp Small intestine. 

Types, Collected: London Zoological Saciety 
Gardens; slides of serial seetions MHNG 24/53-61, 
Materiel examined: wpes, 3 specimens, Adelaide 
Zoologldal Caiidens, Too l9k0, colleeted by M. CG. 
O'Cullagtnin, ALC 824) arf 513845 | Specimen (without 


scolie), Chicago /oologival Catrders, collected by V- 
Rowlatl, BMNMA|967,0 28,23. 


Dasyurotaenia dasyuri spjov. 
FIGS 12-23, 24, 25. 29-31 


Deseriplion (from types): Large cestodes, 237-306 
(400, 1 = S) in deneth, 2.4-3.9 (2.8, n =3) wide, with 
280-330 (310) progiottides in gravid strobilae, Scolex 
larec, 2.40 (a=!) it diameter, deeply embedded in 
mucosa of hast. Suckers 0.32-0.38 (0.35, n-9) in 
diameter, rostellum 0.38-0.50 (0.43, n=4) in dia- 
meter with 36-38 (m4) rustellar hooks arranged 
in two pows (Fig. 12), Large or anterior rosteljar 
houks @,105-O.11) (0,107, n-20) long, base 
0.077-0.093 (0.083, u= 20) long (Figs 13, 14, 24): 
blade large, vere frequently vacuolated; handle ex- 
tremely amall; guard clongate, nor enlarged (owards 
extremly (Figs (9,25). Small or pastenor rostellar 
hopks O.080-0,093 (0.087, n=20) long, base 
().060-0.083 (0.075, 1 = 20) ony (Pigs 17, 24); blade 
large, core trequently vacuolated; handle virrually 
absent, gic large, broad, Irequently bilobed 
distally (ius 18, 25) Neck present. Museulature rot 
stronuly developed. Outer longitudinal museles 
single or in siall bundles. of 2-5 fibres, uiner louei- 
Ludivial pmuseles in Jareer bundles, up to 0,025 iy 
diameter, containing 20 or more fibres. Transverse 
niuseles iy several bawds; wo mos: prominent bands 
immediately internal to inner longitudinal muscles 
and separating inner and outer longitudinal 


muscles; poorly delined bands between bundles af 


inner lonvitucdinal muscles. Dorso-vencral muscles 
spirse, Sintle, crassing cortex and medulla al i 
regular intervals. Lonvitucdinal osmorevulatory 
cunts pied; ventral vanals 011-017 (O15, a> 5) 
in odidneter in outdne proglottides, joined at 
posienar murgin al each proglotis by broad 
Iramsverse canal. Ventral canals with valve like Haps 
promruding iro lumen af junction ot provlottides, 
Dorsal cami extremely narrow, sinuous, 01 G7 3) 
di diameter it mathure prowlouides, Mature provlot- 


ddes 14-25 (17, n= MW) 26 25-30 (29, 10), 
leneth:width ratio 1,.02=2.64 (154, 1 = LO) (ir, 20). 
Gravid proglotlides L839 (2,7, n=T) & 1.93,3 
(2.8, n- 10), lengthewidth ratio (271-459 (107 
n=10). Genital pores mainly unilateral ovvasionally 
iternate irregularly, Genital atriym shallew, 
situated i anterior half of lateral proglottis maiuin 
in mature proglotoides, dividiny margin in ratio 
1:1.3-1:2.4 (218. n=10): in middle of marpin in 
wravid provlotiides, dividing marge in fano 
1:0.9-115 (211, n—10), Genial duets pass between 
longitudinal osmoregulatory canals, Cirrus sae 
clongate, thin-walled (Fig. 1%) invariably exiending 
beyond osmoregulatory canals inta medulla, 
0.50-0,71 (0.57, n=10) © O.08-0,12 (0.10, p =O) in 
mature proglottides. Cirrus slender, 0.01-G,02 (0,015, 
o=5)in diameter, voiled, armature of five hairs vist 
ble on distal region of sume cure under high 
magnifiedtion, Internal and external seminal 
Vesigles absent. Vas deferens erearly eoiled. rlarrow 
duct. loops medially and anteriorly, then at mid- 
line turns posteriorly, terminating near seminal 
receptacle Vasa efferentia not seen. Testes 
numerous, situated iit I-2 layers i dorsal planes. 
Testes oeeupy most of medulla beoween 
osmoregulatary canals, eseepl area of female 
genitalia; testes confluent anterior to vis déterenss 
restes usually confluent postertor te virellarium, ot 
with 1-3 tesies posterior ta and overlyire 
vitellarium; occasionally no testes posterior ro 
Vitellariure. Testes number 150-160 (n= 2) per pro. 
vlovtis; diaineter 0.06-0,08 (07, 4 = 10). Vagiia ape 
proximately G,12 in diameter, straiet, lined inter- 
nally by ting hairs, surmonnded externally by single 
layer ot glandular cells. Proximal 0.15 of vagina mir- 
row, surrounded by. thicker muscle layer than re 
tainder of yayinad, Leckie eladdular cell tyest- 
ment. Seminal reveplacie ovoid (hig, 21), (1-014 
(O14, n=10) = 0.07-0.10 (0.9, n= 10) when Milled. 
situated in mid-line betwen lobes of Ovary, Ovary 
hilobed, paral tobe small, 026-039 (O31, =10) 
« ON2-25 (0.20, 7=10), uporal tobe 032-044 
(h37n- 10) & OLS-0.30 (0-21, n=10), joined by 
vurrow isthinus. Vilellarium postertar lo ovary, 
reniform (12-24 (9.20. n= tO)» OTB U31 (0.23, 
ne 10). Mehlis’ gland spherical, G.O7-0.1) (0.08- 
n=10) in diameter, between vielluriuny and seamal 
reveplacle, Uterus atises as tubular sirietire in 
midline. Uterus in gnivid proglorrides (Fig 23) with 
6-17 (12. = 10) poral and 10-20: (15, 8 = 10) aperal 
faleral ulerine branches: wlerine branches frequern| ly 
subdivided lalerally. Egg approximately spheroidal 


Pips 12-23. Daywinaiacni , dasyurd sp.nov. 12. Seales. apie views 13-18, nastclian books, 13, M4, larvee or anterior 
rostellar hooks, Mleral views 15, large rostellar hook, apreal view, 16, 17, saigll or posterior postetkur bouks, literal 
views HS amiall postellar hook, apical views 1Y, citrus sav and distal vaginas 20, mature proglidtis: 21, female geniiatins 
22, eves; 23, eravid progloltis, Seale lines: Pigs 12, (3-19, 21, O.booim; tie,20, 35 ems tie 22, ON iit fie 23, 


Phen gener; handles t-—tip- 


190 I. BEVERIDGE 


Figs 24-25. Rostellar hooks of Dasyurotaenia dasyuri, sp.nov.; 24, apical view of rostellum, hooks in lateral view; 
25, apical view of hooks showing differences in shape of guards of large and small hooks (arrowed). Scale lines 


0.01 mm. 


Figs 26-28. Rostellar hooks of Dasyurotaenia robusta Beddard; 26, large rostellar hooks, lateral view; 27, 28, apical 
view of hooks showing difference in shape of guard of large and small hooks (arrowed). Scale line 0.01 mm. 


Figs 29-31. Histological features of scolex of Dasyurotaenia dasyuri sp.nov. and associated pathology; 29, sagittal 
section through scolex showing partly withdrawn rostellum; 30 scolex(s) lodged in muscularis externa immediately 
below mucosa (m); 31, scolex(s) lodged in muscularis close to serosal margin showing dome shaped projections 
of tissues (d) beyond normal serosal surface intestine and mucosa (m). Scale lines 1 mm. 


DASPUROTAESIA SPP, FROM MARSUPIALS 19 


(hig. 22), 0,035-0.040 (0.038, 9 = 10) 0.030-0.4134 
(0.032, n=l): embryoplrore thick, homogenous. 
nonsrnaied, onvosphere 0.022-0,028 (0.024, n= 10) 
« O16 O.018 (L017, 2 =10); oneaspheral hooks 
0,008-0.010 (0,008, 4 =10). Development af genial 
organs in 5 specimens: anlage first visible in pro- 
glotlides 60-120 (85); lestes Tirst visible in proylot- 
hides TRO. 190 (187): Viest mature proglottis approx- 
imately 205-220 (215); uteline [ilinye comuiences 
In progloliides 225-265 (245); first wravidl prowlot- 
Tis 270. 420 (305); total prowlottides 280-340 (310), 


hanidtior: Specimens from Lis.; wlentieal co (ypes, 
except in the following minor features: rostellar 
ligok number more variable than in 1ypes, 32 a 
(37, n= 3); large rosicllart hooks O.110-0.120 (0.116, 
ne lO), and small postellar hooks 0.093-0.098 
(0.095, 1=10) bath slightly larger (approximately 
0,010) than hooks of type specimens; testes auntber 
169-226 (186, n—10) per praglottis, higher than in 
types; vitellariun O.10-0.15 (0.13, 7=10) © 0.21-0,38 
(0.30, n=10) celatively wider wad shorter thang ta 
Types 

Host? Dasvuruy macilatus (kerr, 1792) (Mar- 
supialia:s Dasyuridae). 

Sue in hosts Small intestines 


Types: Hololype, & parilypes, Mt Windsor 
Tableland, Qld. 16'12'S, 145°045'T, (7 vii, 1982, coll. 
D. M. Spratt. Holorype. 2 slides SAM V3459, 3 
paratypes, SAM V3d60-V 3462; 4 paratypes, AHO 
S2169-S2172, HC 12322; 1 paratype, BMNH 1983. 
6,13, 1-2; | scoles, strobilar tragments, paratypes, 
in colleehion of R. 1. Rausch. 


Hine yiructure af the eee envelapes (Fie. 32). The 
following chyelopes were recognised surrounding 
the oneosphere The outer envelope of the eve is 
bounded by a thin virellbne membrane, ericlosine 


it ouregular cytoplasmic Jayer. Thre outer 
. | 
4 mB 
~ ' fa 
- 
' ’ 4 
yr 
a] wh 
7 
vi 4 
. 
’ 
i . 


Pig, 32. Tritosmission electron mictograph of ep 
envelopes, Oaiveroraenia dasvari Scale line, SP pie 
c—embryophor, g—eramudar layer ie—inher em 
biyopharic tembrine, oe —ouler embryophoric tmem- 
brane: vl—Vacualated layer; vni—vilelling membrane. 


embrvaphori¢ membrane lies inmediately external 
to the thickened embryophore. The embryophore 
is Of uniform thickness and is composed of relative- 
ly homogeneous election dense marerial which is 
nol ordantised info reaiular embryophoric blocks and 
is Without lacunae or internal. ci@ular bodies. Some 
areas within the embryophore are less electron dense 
and sugrest cavities between blocks but are not ar- 
ranged in a regular fashion. A. zoue of electron 
dense granules, the granular layer, lies between the 
embryaphore and a broad sub-embryophone 
vacuolited area which may represent aggregations 
of lipid bodies. The vacuolated zone and granular 
fave if bounded jnternally by the inter em- 
bryophoric membrane Internal to (hs lies the on- 
cosphere bounded by the Oneosphetal membrane. 


Material examined: Olds typesy Was 3 specimens, 
Simithron. eel BOL. Mirra, vii t979; 3 speviniens, 
Sinithtion, coll. DM. Spritt, 26.41.1968, WE C282, Ca3; 
2 specimens, Lilydale, coll BOL. Munday, 2viis976, 
AHC, HOW732; fragments of specimens, Wynard, voll. 
unknown, 224).1922, whole preserved carcass NMV 
CAO cosiodes ANC 9TKS, 


dxsuerled Lexions (Figs 29-31); Seoleces of D 
dasviri lie dveply embedded in the external musele 
layers of thre wall of rhe small ivestine of the best, 
dither superticially, that is immediately below the 
submucosa, or, close to the serosal margin of the 
risculature such thar the position of the cestode 
scoles 15 indicated by a raised dome-shaped projec- 
Hon on the serosal surface. The neck and anterior 
region Of the strobila lic in & narrow tunnel which 
opels ite the ifleshoal luinen, Two specimens of 
D dasvuri were embedded singly, two were 
enibedded together, and a funher three worms were 
embedded at a single site. The superficial layers 
lining the cavities induced by cestode invasion con- 
aist primarily of neeratic cells and of cell debris 
together with viable cells compressed by the cdisten- 
Hon of Surrounding tissues. There are in addition, 
surrounding the scoleces, a few small localised areas 
of tecrosis of the myoeytes and infiltrations by ine 
Manromatory cells. The principal host reaction is a 
chronic inflummatory one with an infiltration of 
macrophaves and lyniphoetyes and a few plasma 
cells ato Tissues Surrounding the cestode, Polymor- 
phlonuclear leukoevies are unvommon, but 
Langhans-type giant cells are ovecasionally present 
at the edees of lesions. Fibroblasts are prominent 
ia low areas on the outer edges of iufilimabed areas. 


Discussion 
In pone oF tbe previous descriptions of Das viuro- 
tventa hy Beddard (1912, 1915), Baer (1925) or San- 
dars (1957) have the rostellar hooks heen adequately 
deseribed, Beddard (912) provided drawings af the 


192 I. BEVERIDGE 


histology of a purported sucker of D. robusta 
showing sections of sclerotized hooks but did not 
describe the size or shape of the hooks. Baer (1925) 
had no scoleces to examine, but concluded from 
Beddard's description that the cestode in question 
possessed a rostellum armed with taeniid-like 
hooks. Sandar’s specimens (1957), here attributed 
to another species, were without rostellar hooks, but 
were re-described under the name D. robusta. She 
estimated that some 96 hooks were normally 
present. 

The material described above indicates that in the 
past two (or more) independent species have been 
confused under the single name D. robusta. The two 
species described in this paper differ markedly in 
the size and shape of the rostellar hooks, but have 
few differences of note in strobilar morphology, and 
this has undoubtedly led to the confusion. 


D. robusta is known only from the type series col- 
lected from Tasmanian devils in the London 
Zoological Gardens, from the three specimens 
described here, collected from the same host species 
in the Adelaide Zoological Gardens and from a 
single specimen without scolex from the Chicago 
Zoo. Beddard (1915) reported the species in four of 
nine devils dying in the Gardens; however, examina- 
tion of 294 devils in Tasmania has not revealed its 
existence (Gregory ef al. 1974). 

The present redescription, although based upon 
a very limited series of specimens is considered 
justified as the species may now be rare, and since 
the material available allows a fairly full descrip- 
tion to be made for the first time, including the 
distinguishing features of the rostellar hooks. 

D. robusta is distinguished from D. dasyuri by 
the form and size of the rostellar hooks (Figs 2-7, 
13-18). The large rostellar hooks of D. robusta are 
only 0.046-0.058 long compared with 0.105-0.120 
in D. dasyuri. In addition, the size of the hook blade 
is relatively smaller in D. robusta so that the length 
of the hook base is larger, on average, than the total 
length of the hook; in D. dasyuri the length of the 
base is less than the total hook length. In both 
species, the guard of the small hook is particularly 
broad, and this is more marked in D. robusta than 
in D. dasyuri. 

The principal strobilar character distinguishing 
D. robusta from D. dasyuri is the transverse 
osmoregulatory canal joining the ventral canals on 
both sides of the strobila in each proglottis, The 
difference is not immediately obvious in mature pro- 
glottides, but the canals are readily visible in most 
gravid and near gravid proglottides and the presence 
or absence of transverse osmoregulatory canals can 
be readily ascertained. Beddard (1912) noted the 
lack of transverse canals in the type specimens of 


D. robusta, and Baer (1925) confirmed that the 
canals were not visible in Beddard’s sections, but 
suggested that their absence might be more ap- 
parent than real owing to the severely contracted 
nature of the type specimens. The new material con- 
firms Beddard’s (1912) observations that transverse 
canals do not exist. 

Beddard (1912) discussed at some length the 
“membranes” stretching across the lumen of the 
osmoregulatory canals, noting that the lumen was 
occluded by “membranes” once in each proglottis. 
Baer (1925) explained Beddard’s observations in 
terms of oblique histological sections passing 
through consecutive coils of the osmoregulatory 
canal, suggesting that the “membranes” were 
essentially artefacts due to the state of contraction 
of the specimens. The new specimens indicate that 
Beddard’s observations were correct. At the pos- 
terior end of the proglottis in D. robusta and D. 
dasyuri, the lumen of the osmoregulatory canal is 
largely occluded by a valve-like extension of the 
canal wall. The structure is in most respects iden- 
tical to valves which occur in comparable positions 
in the osmoregulatory canals of species of 7aenta, 
described in detail by Kohler (1894). In specimens 
of D. robusta, the ventral canals are of extremely 
variable diameter, and in some incompletely relax- 
ed proglottides, there are occasional folds in the 
canal wall similar to the incomplete ‘‘membranes” 
described by Beddard (1912), Baer (1925) was pro- 
bably correct in ascribing these changes to the state 
of relaxation of the specimens. 

The two species also differ in the number of 
Jateral uterine branches, with 6-10 (9) in D. robusta 
and 6-20 (14) in D. dasyuri. Although these dif- 
ferences appear to be consistent in the material ex- 
amined, they should be treated with some caution 
since Verster (1967) in a rescription of Taenia solium 
Linnaeus, 1758 and T. saginata Goze, 1782, two 
species which have frequently been identified from 
gravid proglottides by difference in the number of 
uterine branehes, found that overlap in uterine 
branch number occurred if a sufficient number of 
proglottides was examined, Some overlap obviously 
occurs in uterine branch numbers of Dasyurotaenia 
spp. and more extensive series of specimens are 
required to test the validity of uterine branch 
number as a taxonomic character in this genus. 

A number of minor morphological differences 
noted between D. robusta and D. dasyuri require 
more detailed examination in larger numbers of 
specimens before their reliability can be established. 
(i) The vitellarium was much shorter and wider in 
D. robusta (Figs 10, 20); however, this may have 
been due to the incomplete state of relaxation of 
the specimens of D. robusta. In addition, there was 
variation in the dimensions of the vitellarium be- 


DASYUROTALT NIA SPP TROM MARSUPTALS 193 


tween specimens of 2 dass trom Tas. and Qld 
the laller specimens (fhe types) aving muely jer 
cower vilellana. Gi) In specimens of DL refute. 
testes were occasimnally found overlying the 
osmorogulutory Canals or even entirely laveral ta 
them, lt contrast to 2. desvuri in which the testes 
invariably lie befween the canals. Gi) The seminal 
reveplacle in D. robust was smaller than D. dusweri 
and the cirrus sae slightly longer and more promi- 
nee, All features Mentioned require examination 
in an estensive series Of Specimens belore any con- 
fidence can be plaved upon cheir ability ro 
distinguish the two species. 


Phe data presented above suggest thar O robusta 
is conlined to Sarcaplites Farrisi’ and thal 
dasyurt vecuts Only in Dasyurws acwlarus, San- 
dir’s (1957) specimens described under the name 
D. robusta bul callected trom Desvurns meeilaras 
m Tas cannot deliniely be assigned to cilher 
Species, siuee they hud oo rosrellar hooks. She siawd 
that ne transverse osmoregulatory canals could be 
seen, but prefaced her remarks by saying that dertils 
of the ostnorepulatory canals could nat be deter- 
mined tt would therefore be unwise Lo assume (hal 
het 4pevinens were J. robusta based on her tailuce 
to find transverse Osmoreulatory canals. Her figure 
(Fig. 27) of a weavid proglottis reveals 12-14 lateral 
ulerine branches, suggesting (hal the specimens she 
deseribed may have been in fact @ dasvuri and nor 
D. rebusta. However, Dasyruuiuenia is probably 
represented by several species in dasyurid mar- 
supials. In addition to the new speeies deseribed 
here. a single juvenile cesiode with 96 rostellar 
hooks, probably representing yeranochte: species. 
Was oo-parasitic with the ype specimens of 2 
Hesvurd and has been deposited in AH (2173). 
Sandar’s (1957) specimens could be attributed bo 
rhis species or could bave been a mixture of two 
species, 2, dasvuricand the undeseribed species. 
Another probably new species with 18-22 hooks, 
0100-0108 and 0.092-0.104 lone fram Dasyures 
whoprnenius Schlegel, 1880, in New Guinea is 
represented by two specimens oniv in the collection 
ofthe BMNH (1973.7,0.5-6), More specimens are 
required before the species can be ceseribecl ade- 
quately. binally, juvenile cesiades with 34 hooks 
QA520.15@ and 0,122 0.126 long were present in i 
specimen Of Safanedius hallucatus Gould, (842 from 
W.A. The specimens probably: represent a new 
species ot Desvaratvenw and have been deposited 
in) WAMT (79, SO, BI-1983). 

A metacestade of Daseurotaenia, identilied ast 
robusta was teported lroro the peritancal cavity of 
Patoraus tridactylus (Ket, 1792) in Tas. by Gregory 
(pers, corm. in Beveridge, 1978). This parneular 
specimen hod been identified by comparison wath 


seuvleces front Lasvuruy nievlarus, and is mow con- 
sidered to be a metacestode of D. dasviny, 

The occurence of Ue sealex deeply eniedded 
within the intestinal wall of the toast ty urnestial 
among cesiades, Pavadi/epis sco/ecina (Rudolphi. 
VR1Y9) burrows info the small imesting of Cormorant 
Phalacrocoray carbo Livnaens, the scolex lodging 
in the muscularis externa close to the serosa 
(Karstad ev a. 1982), and a stiytlar lovalisuryoan tis 
been reported far Paradilepis delachwuxi 
(Futhenanin, 1909) in Phalacracoran africans 
(Gmelin) by Baer (1859) In mammiuls, the 
anuplovephalid keraporephaliven aber Rausch & 
Obbayashi, 974, Gooeurs wil is seolex deeply 
buried in the wall.of the saceutis potundus of the 
pikas Ochulona ravler (Geilby) and OL nlacratis 
(Guuther) (Rausch & Ohbayashy, 1974) The 
mecharisins of invasion of Dusvarolaenia were Wot 
clear from the material studied, Dasvurataenia spp, 
de not have prominent postellar glands co seerete 
proteolylic enzymes such as are present in dy abel, 
bur maienal from /2 rmaculacuy (WL C43) does sup- 
zest that the puvenile cestodes of D. dusuri beeanie 
deepty embedded in the small intestine wall befare 
the Injhation af progloilisalion The hestolugresat 
reaetiuin to the seolex of B dasvurt ts similat to that 
desenbed tor PF sealeving and & uber, 

The genus Dasyyrarecviia was allocated re the 
Taeniidae by Baer (1Y25) and (his was contiemed 
subsequently by Sanddars (1957). Rausch (981) 
however has emphasised that the family, iu the form 
recognised by Abuludse (1965) and Yarmtaguti (1959) 
iy obviously polyphyletic, Cladolaeria Cohn, 190! 
as indicated by Freeman (1973), belangs to the 
Dilepididae, based on the morphagenesis of the 
nictacestodes, (hoveh the morphology of matune 
and gravid provlotiides is similay to the taeniids. 
Aneploraenia Beddard, VOU, a parasite at the 
Tasmaniag Devil, likewise has a proglottis 
morphology akin to the Tweniidae while 
melucesiode developinent uidicates affinities with 
the Linstowiidae (Beveridge (982) In asceriaming. 
the (rue relationship ol genera witht che ‘laenidae, 
iy is obvious that a knawledee of ineracescode 
development is a pre-requisite, and these dia eur 
renuy are lacking dor Daseurarenu As a conse- 
quetice, Morphological data canuot be tle basis tor 
a final determination of its tixenamic position, but 
may provide clues. 

Dasyurotdenia \s disungdished from other genera 
ol the Taeniidae (sensi Yaniagull 1959) by the large 
seolex embedded deeply in (he tissues of the host 
and hy the essenoally unilateral genital pores, San- 
Jurs (1957) mentioned rhe vereral body shape. the 
structure al che seolex, the form uf tie cirrus sae 
and the development of the musculature as features 
Jistinsuishing the genus, While the form af the vir 


194 1. BEVERIDGE 


rus sac is markedly different from A. dasyuri, a 
coparasite of the Tasmanian devil, it does not dil- 
fer from most Taenia spp. and therefore cannot be 
considered diagnostic. Of the characters mention- 
ed by Sandars (1958), only the arrangement of the 
musculature seems worthy of consideration as il is 
apparently unique in the family. Unfortunately, the 
musculature of many species of the Taeniidae has 
not been described in detail, and its value as a taxo- 
nomic character for Dasyurofaenia is therefore open 
to some doubt at present. 


The structure of the egg likewise is inconclusive. 
The embroyophore is extremely thick, as in taeniids, 
but is not composed of radially arranged blocks 
with lacunae (see Fairweather & Threadgold 1981) 
nor is it characteristic of dilepidid eggs (Pence 1967). 
In A. dasyuri, the structure of the egg was inter- 
preted as being typically taeniid (Beveridge e/ al. 
1975) yet the morphogenesis of the metacestode of 
this species indicates linstowiid affinities. Hence, 
there is some doubt as to the taxonomic significance 
of egg structure, and little weight can be placed 
upon the presence of a thick embryophore and in- 
significant outer envelope in the egg of D. dasyuri. 

In D. robusta, the rostellum is apparently retract- 
able, and can be retracted fully within the scolex. 
This characteristic, shown in Fig. 1, has been 
overlooked by previous writers, but it is not a 
characteristic of Taenia or Echinococcus (see Wardle 
& McLeod 1952), the only two genera considered 
by Rausch (1981) as belonging to the Taeniidae. A 
retractable rostellum is a feature of the Dilepididae 
and Hymenolepididae (Wardle & McLeod 1952) and 


may indicate an affinity with these groups rather 
than with the Taeniidae. 

In summary, none of the morphological data pro- 
vided allows the definitive allocation of Dasyuro- 
taenia to a family. Superficially it resembles the 
Taeniidae, but the retractable rostellum of the type 
species, the musculature, and structure of the egg, 
cast doubt on such affinities. 

Rausch (1981) suggested that Dasyurotaenta 
could not be allocated to the Taeniidae on 
phylogenetic as well as morphological grounds, 
alluding to the evolution of the Dasyuridae in isola- 
tion from eutherian mammals, and the belief that 
the true taeniids have evolved exclusively within 
recent Carnivora. If this is the case, Dasyurotaenia 
may exhibit a strobilar morphology convergent with 
species of Taenia, yet be derived from alternative 
origins, either the Linstowiidae of dasyurid and 
peramelid marsupials (Beveridge ef al, 1975, 
Beveridge 1982) or Diplepididae from accipitriform 
birds (Beveridge ef a/. 1975). Elucidation of the life 
cycle of the parasite will be required before a final 
answer can be given. 


Acknowledgments 


My sincere thanks are due to D. M. Spratt for 
providing most of the material described in this 
paper and for reading the manuscript, to R. Bray 
and C. Vaucher for lending material in their care, 
to M. G. O’Callaghan and B. L. Munday for col- 
lecting the material from Sarcophilus and Dasyurus 
respectively and to Dr R. L. Rausch for reading the 
manuscript. The electromicrograph was taken by 
K. Smith. 


References 


ABULADSE, K. 1. (1964) Taeniata of animals and man and 
diseases caused by them. /” Essentials of Cestodology, 
Vol. 4. Ed. K. [, Skrjabin. Akademia Nauk SSSR. 
(English translation, Israel Program for Scientitic 
Translations, Jerusalem, 1970). 

Baker, J. G. (1925) Some Cestoda described by Beddard, 
1911-1920. Ann. Trop. Med. Parasit. 19, 1-22. 

— (1959) Helminthes parasites, Exploration des Parcs 
Nationaux du Congo Belge, Mission Baer-Gerber 
(1958). (Institut des Pares Nationaux du Congo Belge 
:Bruxelles.). 

BEDDARD, F, E. (1912) Contributions to the anatomy and 
systematic arrangement of Cestoidea, V. On a new genus 
(Dasyurotaenia) from the Tasmanian devil (Dasvurus 
ursinus), the type of a new family. Proc. Zool. Soc. 
Lond. 1912, 677-695, 

(1915) Contributions to the anatomy and systematic 
arrangement of Cestoidea, XVI. On certain points in 
the anatomy of the genus Aymabilia and Dasyurotaenia. 
Thid. 1915, 175-191. 


Brverivce, I, (978) Helminth parasites of Australian 

marsupials. /m Proceedings of Course No. 36 for 
velerinarians—Fauna. (Post-graduate committee in 
Veterinary Science, University of Sydney, Sydney.) 
— (1982) Specificity and evolution of the 
anoplocephalate cestodes of marsupials. Mes, mus, 
nat, Hist, nat. nouvelle serte, Serie A., Zoologie 123, 
103-109. 

— Rickarp, M. D., Grecory, G. G. & MUNDAY, 
B. L. (1975) Studies on Anoplotuenia dasvuri Beddard, 
IYLL (Cestoda: Taeniidae), a parasite of the Tasmanian 
devil: observations on the egg and metacestode. /nt. J. 
Parasit. 5, 257-267. 

PAIRWEATHER, T. & THREADGOID, L. T. (1981) 
Hymenolepis nana; the fine structure of the embryonic 
envelopes. Parasitology 82, 429-443. 

FREEMAN, R.S. (1973) Ontogeny of cestodes and its bear- 
ing on their phylogeny and systematics. Advances i 
Parasitology WW, 481-557 (Academie Press: London). 


1. BEVERIDGE, R. SPEARE & P. M, JOHNSON 198 


Grecory, G. G., Munpay, B. L., Beveripar, 1. & 
Rickarb, M. D, (1974) Studies on Anoplotaenia 
dasyuri Beddard, 1911 (Cestodae, Taeniidae) a parasite 
of the Tasmanian devil: life cycle and epidemiology. /nv. 
J. Parasit. 4, 187-191, 

KARSTAD, L., Sitbo, L., OkecH, G. & KHALIL, L. FE 
(1982) Pathology of Paradilepis scolecina (Cestoda: 
Dilepididae) in the white-necked cormorant 
(Phalacrocorax carbo.) J. Wildl. Dis. 18, 507-509. 

Koner, E. (1894) Der Klappenapparat in den Excretion- 
sgefassen der Taenien. Z. Wiss Zool. 57, 385-401. 

Pence, D, B. (1967) The fine structure and histochemistry 
of the infective eggs of Dipylidium caninum. J. Parasit. 
53, 1041-1054. 

Rausch, R. L. (1981) Morphological and biological 
characteristics of Twenia rileyi Loewen, 1929 (Cestoda: 
Taentidae). Can. J. Zool. 59, 653-666. 


& OHBAYASHI, M. (1974) On some anoplocephaline 
cestodes from pikas, Ochotona spp. (Lagomorpha) in 
Nepal, with the description of Eefopocephalium abei 
gen. et. sp. n. J. Parasir. 60, 596-604. 

SANDERS, D. F. (1957) Redescription of some cestodes 
from marsupials. I. Taeniidae. Ann. Trop. Med. Parasit. 
51, 317-329. 

VERSTER, A. (1967) Redescription of Taenia solium Lin- 
naeus, 1758 and Taenia saginata Goeze, 1782. Z. Parasit. 
29, 313-328. 

Warb_e, R. A. & McLeop, J. A. (1952) The Zoology of 
Tapeworms, (University of Minnesota Press: 
Minneapolis.) 

YAMAGUTI, S. (1959) Systema Helminthum, II. The 
Cestodes of Vertebrates. (Interscience Publishers: New 
York.) 


NEW RECORDS OF GLOBOCEPHALOIDINAE (NEMATODA: 
TRICHOSTRONGYLOIDEA) FROM MACROPODIDAE IN NORTH 
QUEENSLAND 


BY I, BEVERIDGE, R. SPEARE & P. M. JOHNSON 


Summary 


Globocephaloides affinis Johnston & Mawson, 1939, is resurrected as a valid species and 
redescribed. The male is described for the first time. G. affinis differs fron congeners by the 
possession of a gubernaculum, in the form of the spicules and in the size of the buccal capsule. It 
occurs in Macropus dorsalis and rarely in M. giganteus. G. macropodis Yorke & Maplestone, 1926, 
was found in Macropus agilis, M. dorsalis, M. parryi, Wallabia bicolor, Lagorchestes conspicillatus 
and Aepyprmnus rufescens. M. parryi, W. bicolor, L. conspicillatus and A. rufescens are new host 
records. Amphicephaloides thylogale Beveridge, 1979, was found only in Thylogale stigmatica, and 
has not previously been recorded from this host in Queensland. 


NEW RECORDS OF GLOBOCEPHALOIDINAE (NEMATODA: 


TRICHOSTRONGYLOIDEA) FROM MACROPODIDAE IN NORTH QUEENSLAND 


by I, BEVERIDGE*, R. SPEARET & P.M, JOHNSON{ 


Summury 


BEVERIDGE, |, Spharh, Ry & Jonsson, PM. (1984) New records ot Cilohocephaloidinae (Nematoda: 
Trithostrongyloidea) Hon Macropodidite in north Queensland, fravs, R. Soe, S. Auer W8(4), 197-201, 
13 December, (984. 


Globovephaloides affinis Johnston & Mawson, 1939, is resurreeted as a valid species and redescribed, 
Vhe male is described for the first time ©. a/finds differs trom congeners by [he possession of a gubernaculum, 
inthe form of the spicules and io the size of the buceal capsule. I Occurs in Macropus dorsalts and rely 
in MA gizariteus, G. macrepodis Yorke & Maplestone, 1924, was found in Macrapus agilis, M. dorsalis, 
M. parryi, Wallabia bicolor, Lagorchestes conspicillatus and Aepypeyninus rufescens, M. parryi, W. bievlar, 
1, conspicillaius and 4, ryfescens are new host records, Amphicephaloides thylagale Beveridge, 1979, was 
found only in Tivlogale stigmratica, and has nat previously been recorded from this host in Queensland. 


Kiy Worps Ohihavephaloides, Amphicephaliides, Nematoda, disitibution, hosts, morpholopy, 


Maciapadidae, 


Introduction 


The trichostrongyloid subfamily Glohocepha- 
loidinae Inglis, 1968, consists of three nematode 
species belonging to the genera Globocephaloaldes 
Yorke & Maplestone, 1926 and Awmphiceplialoides 
Beveridge, 1979, occurring in the duodenum ul 
macropodid marsupials and currently placed within 
the family Herpetostrongylidae (Durette-Desset & 
Chabaud 1981). Ina recent review of the subfamily 
(Beveridge 1979), (wo species of Glohucephaloides 
were recognised as valid and a new monotypic 
genus, Amphicepha/loides, restricted to pademelons 
(Thylogale spp.) was described, The distribution 
and prevalence of G. srifidospiculuris, a species 
occurring in south-eastern Australia, was reported. 
The lack of extensive parasite collections rom 
northern areas of the continent restricted any con- 
siderations of the distribution and abundance of 
G. macropodis which is apparently limited to 
tropical Australia, Recent collections of nematodes 
from kangaroos and wallabies in north Queensland 
have provided additional information on the preva- 
lence and distribution of G. macropadis and 


* Veterinary Scienees Division, S.A. Departinent of 
Agriculture, © Trstitute of Medical and Veterinary 
Science, Frome Road, Adelaide, 8. Aust, 5000, 

7 Graduate School of Topical Veterinary Selence, lames 
Cook University of North Queensland, Tawnsville, Qld, 
4810. 

} Queensland National Parks and Wildlife Service 
Northern Regional Centre, Pallarenda, Townsville, Old. 
4810, 


Amphicephaloides thylogale as well as indicating 
that G, affinis Johnston & Mawson, 1939, a species 
described originally from females only and 
suppressed by Beveridge (1979) as a synonym of 
G. macropudis, 1s in fact a valid species. In this 
paper, G. affinis ts resurrected, the male of 
G. a/finis is described for the first tume, and addi- 
fional hosi and geographic revords are given for G. 
macropodis and A. dylogale, 


Methods 


Nematodes were collected Horr the content of the 
duodenum of kangaroos and wallabies either 
immediately following the death of the host or from 
viscera preserved in 10% formol saline. Duodenal 
cortent was washed in a fine sieve and the residue 
examined using & dissecuon microscope, Live nema- 
fodes were !ixed in boiling 70% cthanol and stored 
in 70% ethanol wich 5% glycerine. All specimens 
are deposited in the Australian Helminthological 
Collection (AHC) housed in the Suuth Australian 
Museum, and collection numbers are cited ti the 
text. 

Drawings were made with the aid of a camera 
lucida from specimens cleared in lactophenol. tn 
the descriptions, measurements are wiven in mil. 
litnetres as Lhe range followed by the mean of five 
measurements in parentheses. 


Results 
The prevalence of Glohocephaloides and 
Amphicephaloides in macropodids from ort 
Queensland is shown in Table 1, 


198 


I. BEVERIDGE 


TABLE 1. Prevalence of Globocephaloides and Amphicephaloides in macropodids from northern Queensland. 


G. macropodis 


G. affinis A. thylogale 


No. No. No. 

No. Prevalence worms Prevalence worms Prevalence worms 
Macropodid species examined % (mean) % (mean) %p (mean) 
Macropus agilis (Gould, 1842)* 40 30 1-10(3) 0 — 0 — 
Macropus antilopinus (Gould, 1842) 10 0 — 0 —_— 0 — 
Macropus dorsalis (Gray, 1837) 22 59 2-28(7) 68 1-11(4) 0 — 
Macropus giganteus Shaw, 1790 26 0 — 4 | 0 _ 
Macropus parryi (Bennett, 1835) 15 13 1,32 0 = 0 — 
Macropus robustus Gould, 1841 21 0 — 0 _ 0 — 
Wallabia bicolor (Desmarest, 1804) 9 11 | 0 — 0 — 
Aepyprymnus rufescens (Gray, 1837) 1] ) 4 0 — 0 — 
Thylogale stigmatica Gould, 1860 10 0 — 0 — 40) 1-28(9) 
Largorchestes conspicillatus Gould, 
1842 9 1] 9 0 — 0 — 
Onychogalea unguifera (Gould, 1841) 10 — 0 _— 0 = 
* Data from Speare ef al. (1983). 
Globocephaloides affinis Johnston & Mawson, 1939 Male 


(FIGS 1-18) 


Material examined: From Macropus dorsalis: Qld: 6 ¢, 
2?, Milman, AHC 12741; 1? , Mt Surprise, AHC 
12742; 74, 18 , Warrawee Station via Charters Towers, 
AHC 12321, 12316, 12737; 1¢ , 62, Pallamana Station 
via Charters Towers, AHC 12738; 8d, 16 2, Harvest 
Home Station via Charters Towers, AHC 12739, 

From Macropus giganteus: Qld: 1é , Harvest Home 
Station via Charters Towers, AHC 12740, 
Description: 

Small nematodes, coiled in loose, flat spiral, 
ventral surface on inside of coil. Body without 
longitudinal ridges or alae, covered by numerous 
transverse striations. Striations extremely fine, close 
together, interspersed irregularly with more promi- 
nent striations, 0.024-0.048 apart. Mouth opening 
oval, slit-like when closed, dorsoventrally elongate. 
Lips absent; mouth surrounded by 2 lateral, jaw- 
like structures joined at dorsal and ventral margins. 
Each jaw bears 3 finger-like extensions of pulp- 
tissue which reach mouth opening. Cephalic 
papillae not seen. Jaws supported posteriorly by U- 
shaped buttress-like elevations of cuticle and sub- 
cuticular tissues on dorsal and ventral aspects, 
extending from well posterior to anterior extremity 
of oesophagus, running anteriorly and laterally ter- 
minating at origin of 3 extensions of pulp-tissue. 
Buccal capsule large, urceolate, with thickened rim 
at base, thinning markedly anteriorly; walls also thin 
dorsally and ventrally. Single pointed, elongate 
dorsal tooth arises from oesophagus; subventral 
teeth absent. Dorsal oesophageal gland opens 
through apex of dorsal tooth. Oesophagus elongate, 
clavate, slightly wider at posterior end. Nerve ring 
encircles oesophagus near anterior extremity. 
Excretory pore at level of nerve ring. Deirid small, 
not lobed, at level of excretory pore. 


Length 4.9-5.6 (5.2); maximum width 0,.25-0.32 
(0.28); buccal capsule 0.11-0.13 (0.12) x 0.11-0.13 
(0.12); tooth 0.035-0.044 (0.040) long; buttresses 
arise 0.20-0.23 (0.21) from anterior end; oesophagus 
0.78-0.88 (0.82); nerve ring 0,37-0.40 (0.39) from 
anterior end; excretory pore 0.35-0.38 (0.37) from 
anterior end; deirids 0.35-0.37 (0.36) from anterior 
end; spicules 0.26-0.32 (0.29); gubernaculum 0.11 
long. Lateral lobes of bursa enlarged, fused with 
ventral lobes. Dorsal lobe very small, not separated 
from lateral lobes. Ventroventral and ventrolateral 
rays separated, slender, each reaching margin of 
bursa. Lateral rays broad at origin, externolateral, 
not reaching margin of bursa; other lateral rays 
reaching margin of bursa. Externodorsal ray 
slender, originating from dorsal ray, not reaching 
margin of bursa. Dorsal ray slender, divided into 
2 very short branches at distal extremity, each 
minutely divided at tips. Bullae over origin of lateral 
and ventral rays absent. Genital cone not promi- 
nent. Anterior lip of cone very small, conical, with 
tooth-like appendage ventrally. Posterior lip gives 
rise to rounded, distally bifid projection. Spicules 
short, complex, not heavily sclerotised, yellow. 
Spicules trifid distally. Main branch of spicule with 
serrated lateral margin, and bi-cornuate distal 
extremity, Lateral branch of spicule slender, simple, 
no longer than medial branches, tip simple. Third 
branch simple, elongate, poorly sclerotised. Distal 
tip of spicule with large transparent flange. Guber- 
naculum slender, elongate. 

Female 

Length 5,5-6.2 (5.9); maximum width. 0.42-0.48 
(0.45); buccal capsule 0.12-0.14 (0.13) x 0.13-0.15 
(0.14); tooth 0.035-0.050 (0.044); buttresses arise 
(0.22-0.24 (0.23) from anterior end; oesophagus 
0.87-0.98 (0.94); nerve ring 0.35-0.40 (0,37) from 


NEMATODES FROM QUEENSLAND MARSUPIALS 19¥ 


Figs t-18. Globocephaloides affinis Johnston & Mawson. 

Vig, 1, Cephalic end, lateral view. Fig. 2, Cephalic end, ventral view, Fig, 3, Cephalic end, en fuce view. Fig. 4, 
Anterior end, literal view. Pig. 8, Deirid, ventral view, Fig, 6, Transverse body striations in region of excretory pore. 
Fig, 7, Female tail, lateral view, Fig. 8, Vulva and ovejector, lateral view. Fig. 9, Gubernaculum, lateral view. Fig. 10, 
Gubernaculum, dorsal view, Fig. 11, Bursa, lateral view. Fig. 12, Dorsal lobe of bursa, dorsal view, Fiz. 13, Termination 
of dorsal ray, dorsal view. Fig. 14, Genital cone, lateral view. Fig, 15, Genital cone, yentral view. Fig. 16, Spicule, 
lateral view. Pig. 17, Distal lip of spicule with surrounding cuticular flange, lateral view. Fig. 18, Main branch of 
spicule showing denticulate margin, lateral view. Seale lines: Figs 1-3, 5, 6, 9-12, 16-18 to same scale, 0.1 mm; 
Figs 4, 7, 8 to same scale, 0.1 mm; Figs 13-15 to same scale 0.01 mm. 

Legend: a, anterior lip of genital cone; d, deirid; e, excretory pore; n, nerve ring; p, posterior lip of genital cone. 


200 


anterior end; excretory pore 0.35-0.39 (0.37) from 
anterior end; deirids 0.37-0.41 (0.39) from anterior 
end; tail 0.18-0.20 (0.19); vulva to posterior end 
1.41-2.10 (1.76); egg 0.08-0.11 (1.10) = 0.06, Tail 
short, simple, conical. Vulval lips not prominent; 
lips approximately equal. Ovejectors paired; vaginae 
uterinae short. Uteri large, sac-like opposed. Egg 
thin-shelled, ellipsoidal with two-cells when laid. 


Site in host: duodenum. 


Globocephaloides macropodis Yorke & Maplestone, 
1926 
Syn. Globocephaloides wallabiae Johnston & 
Mawson, 1939 
Globocephaloides thetidis Johnston & Mawson, 
1939 


Material examined: From Macropus agilis: Qld: 1d, 4 
9, Townsville, AHC 7565; 1¢ , 5% , Hervey’s Range 
(Townsville), AHC 7165; 1 2 , Black River (Townsville), 
AHC 7515; 2 2, Inkerman Station via Home Hill, AHC 
7525; 2, Wenlock River, Cape York, AHC 7407, 
From Macropus dorsalis: Qld: 14 ¢, 192 , Milman, AHC 
12023, 12253; 7¢, 12 2, Clements Creek, Marlborough, 
AHC 12199; 24, 3? , Warrawee Station via Charters 
Towers, AHC 12743, 11298; 2 ¢, Mt Surprise, AHC I1141; 
1¢ ,69%, Pallamana Station via Charters Towers, AHC 
12745: 8d, 16% , Harvest Home Station via Charters 
Towers, AHC 12744, 

From Macropus parryi: Qld: 93 , 24% , Inkerman Station 
via Home Hill, AHC 11931, 12364. 

From Wallabia bicolor: Qld: 1% , Harvest Home Station 
via Charters Towers, AHC 12334. 

From Aepyprymnus rufescens: Qld: 2d , 2? , Ayrsville, 
AHC 8841. 

From Lagorchestes conspicillatus: Qld: 63 ,3 2 , Bohle, 
AHC 12837. 


Amphicephaloides thylogale Beveridge, 1979 


Material examined: From Thylogale stigmatica: Qld: 12 , 
3. Peeramon, AHC 11235; 1d, 3¢, Tolga, AHC 735]; 
1d, El Arish, AHC 7383; 17¢ , 112 , Wongabel State 
Forest, AHC 12354. 


Discussion 


Globocephaloides affinis was initially described 
from the small intestine of Macropus dorsalis by 
Johnston & Mawson (1939) on the basis of female 
specimens only. Beveridge (1979) considered that the 
only distinguishing feature of this species, namely 
the larger buccal capsule, was not a valid specific 
character given the variation seen in other species, 
and in the absence of males suppressed G. affinis 
as a synonym of G, macropodis. In recent collec- 
tions from M. dorsalis, male and female nematodes 
with extremely large buccal capsules, similar to the 


I. BEVERIDGE, R. SPEARE & P. M. JOHNSON 


original description of G. affinis, were found in| 
association with G. macropodis. The characters of 
the male copulatory apparatus indicate clearly that 
G. affinis is a valid species and is not synonymous 
with G. macropodis. G. affinis possesses a guber- 
naculum, unlike congeners, lacks bullae overlying 
the lateral lobes of the bursa, has a much longer 
dorsal lobe of the bursa and the spicule bears den- 
ticulate processes on its main branch. The spicules 
are lightly sclerotised and yellow in colour, as in 
G. macropodis, but, like G. trifidospicularis, they 
have three distal branches compared with two 
branches in G, macropodis. The presence of two 
simple subsidiary branches of similar length differs 
markedly from G. trifidospicularis in which the 
minor branches are sinuous and differ in length. 
In addition, the genital cone differs from congeners 
in having a diminutive anterior lip, and a short, 
stout posterior lip. The simple deirid differs strik- 
ingly from the bilobed deirids present in both 
G. macropodis and G. trifidospicularis, The 
presence of a gubernaculum, denticulate spicules, 
the lack of paired bullae on the internal surface of 
the bursa, the features of the genital cone, the lack 
of subventral teeth in the buccal capsule and the 
presence of a thickened ring at the base of the 
capsule indicate similarities with Amphicephaloides 
thylogale rather than with congeners. The buccal 
capsule of G. affinis is considerably larger than 
either G. macropodis or G. trifidospicularis and 
appears to be a reliable distinguishing character. 
Females of G, affinis can be distinguished by the 
size of the buccal capsule, by the absence of promi- 
nent vulval lips, and by the extremely fine body 
striations, interspersed at irregular intervals by 


coarser striations. ; ' 
GC. affinis occurs commonly in M. dorsalis and 


is frequently found in mixed infections with G. mac- 
ropodis (Table 1). In the Charters Towers area of 
Qld, G. affinis was found in 13 of 14 M. dorsalis, 
one of six M. giganteus, none of 21 M. robustus and 
none of five Wallabia bicolor, suggesting that M. 
dorsalis is the normal host species, but that it may 
occasionally infect other sympatric macropodid 
species. 

G. wallabiae Johnston & Mawson, 1939 was 
placed as a synonym of G. macropodis by Beveridge 
(1979). Initially described from Macropus dorsalis, 
the type specimens have been lost, and in relegat- 
ing the species to synonymy, Beveridge (1979) relied 
on new material collected from M. dorsalis which 
consisted of seven specimens only. The frequent 
occurrence of G. macropodis in M. dorsalis and 
careful comparison of this new material with a simi- 
larly extensive series of specimens from the type 
host, M. agilis, confirms the synonymy of G, 
wallabiae with G. macropodis. 


NEMATODES FROM QUEENSLAND MARSUPIALS 20 


G. thetdis Johnston & Mawson, 1939, was also 
placed. as a synonyili ot G, macropodis by Beveridge 
(1979). G.. theudis had been deseribed originally 
from a few specimens found in Phvlovale thesis 
from the New England region of N.SW., yet 
attempts to collect additional specimens of the para- 
site in New England and in southern Qld yielded 
only 4, fhylogale (Beveridge, 1979), Other nema- 
tode species described from the same host animal 
from New England from which Johnston & 
Mawson (1939) described G. thelidis suggest that 
the host has been misidentified. Species of 
Rugopharyax Noenniz, 1927, and Zoniolainus 
Cobh, 1898, originally deseribed from "7 fhelis”’ 
i fact oceur only in M, dorsalis (Beveridge 1982, 
1983). Misidentification of (he host would account 
for the otherwise inexplicable oceurrenee of 
G, macropodis iT. thetis, and he common occur 
tence Ol G. wacropoiliy in M. dorsalis (Table 1) 
(herelore supports the synonymy of G. theridis with 
G. macropodls. 

G. macropodis as previously been reported fromm 
M, agilis, M, dorsalis, M. gigantens and Petrogale 
inarnala in Old (Beveridge 1979; Speare ef a/, 1983), 
Data presented in this paper confirm thal 
G, macropodis is a common parasite of M. itorsalis. 
The farhire to find G, macropadis in M. gigunieuy 
(Table 1), even when AM. giganrous was sympatric 
with other infected macropodid species, suggests 
that it is only dn incidental parasite of this host. 
Beveridge (1979) found only a single female 
Cf, macropodis in Mf. giganteus wear Rockhampton 
G. mucropodis is reported for the first time fron) 
Aepyprymnus rifescens, Macropus parryi, Lager- 
chestes conspicillains and Wallabia bicolor, bur wi 
each instance the prevalenee is relatively low, stug- 


gesting that they are incidental hosts, while AL agilis 


and M. dorsalis are (he principal lrosts of This jienta- 
tode species (Table 1). 

Beveridge (1979) concluded that both G, ¢rifide- 
spicwiaris and G, macrepodis have an extremely 
wide host range, the forier species aecurring in 
southern Australia and being replaved hy 
G macropodis inthe north. The recent callections 
support this view, Af. parryi, M. wigarnleus and 
Wi hicolar are hosts al G trifidaspicularis in 
Victoria and N.SW. (Beveridge 1979), while in 
northern Qld the same host species are parasitised, 
if infrequently, by G, pracropedis. By contrust, Ci. 
affinis is apparently restricted lo MW, dorsalis, with 
only an oveasional infeetion in M. giganreus in areas 
where the Iwo macropodid species are sympatrle, 
A, (hylogale oceurs in Thylogale thedis in SSW 
and southern Qld and in 7 stigmutiew trom 
florthern N.SW,, bai has nor beet reported 
previously io 7) srignaticw from northern Qld where 
it is apparently restricted to this host species, 

To date, only G. trifidespienlarin has been 
implicated in causing disease in kanparous (Arundel 
et al. 1977). One ol us (R.S,) has recently autopsied 
a 12 month old captive AZ, eigenieus which died 
from anaemia and hypoproleinaemia caused by 
2,300 G, mucropodis, The juvenile ML gigueneny 
grazed a confined area together with adult Mf wes/is 
and Md? parryi, indicating that under con! ined con 
ditions, G. macropodis as well as G. brifidaspiculins 
cun be a serious pathogen. 


Acknowledpments 


Our thanks are duc to P, Speare fo) assistance ni 
collceting and examining specimens, 


References 


ARUNDEL, Hy BARBER, b oB.& Beveriooe, L. (1977) 
Diseases uf trarsuyrals, pp. 141-154. JB. Stamehouse 
& BD. Gilmore (eds.) “The Biology of marsupials” (Mac 
Millan: London.) 


Brviginue, 1 (979) A review of the Clobovephatoidinac 
Inglis (Nematoda: Amidostomartidac) trom macrapadid 
toarsupials. lies. A 4aad 27, 181-174. 


(1982) A lasonomic revision of the Phuryngo 
stronglyined Popova (Nematoda: Strongvloidea) from 
macropodid mansupints, Joie Suppl Ser. No, 83, 150 
rp. 


(1983) laxononmic revision Of tie Zonrolaimiyes 
(Popova) (Nemialodur Strongyloided) (nam maernprectia 
marsupials, /hfd. Suppl Sen No. Yl, RB pp. 

Duk cry Dessert, Me. C. & CHsnauu, A Ce LSet) 
Nouvel essai de ehissifennon des nemarodes Tighe 
stroupyloides. vii, Parasin a, 247 312, 

Tous tin, TA & Mawson, P.M. C934) Steeiievte 
HeMmtodes from Queenshind marsupials Tears he, Sac 
S. Aut 03, 12h 148 

SPrARL, BR, BEVERIDGE, 1. Tonssas, POM, & Cons 
L, A. 983) Parasites of the agile walliby, Avorn 
auilis (Marsupiatia) alas Wild Nes Ub S904 


ADDITIONS TO THE COLLEMBOLIAN FAUNA OF THE ANTARCTIC 


BY PENELOPE GREENSLADE & K. A. J. WISE 


Summary 


Two new species records of Collembola from Deception Island are noted. Onychiurus sp., and 
Folsomia candida. A description is given for the former. Neither of the two genera have been 
recorded previously from the Antarctic. A discussion on their possible recent introduction to the 
island is given. 


ADDITIONS ‘TO THE COLLEMBOLAN FAUNA OF THE ANTARCTIC 


by PENFLOPE GREENSLADE* & K, A. J, Wise? 


Suminary 


GaLPSsLADE, P, & Wisk, K. A. J. (1984) Additions to the collembolan fauna of the Antarctic, Trans. R. 


Sou. So Ase DORE) 203-205, 14 December, 1984. 


Iwo mew species records of Collembola from Deveption Island ure noted, Onvesiurus sp., and 
Fulsonia candida, \ deseviplion is given for the former, Neither of the Wwo genera have been recorded 
previously Trane the Antarctic. A discussion on their possible recent introduction to the island is uiven. 

Key Wokos: Collembola, Jaxonomy, Antarelic, Deception Is., introduced. species, 


Introduction 
Wise (1967,197]) recorded 1 species of 
Collembola from Antarctica, (the mainland 


together with nearby offshore islands, including 
the South Shetland islands, which lie at 61-62° §). 
Re-exuminalion of two collections made by 
J. Strong on Deception Island (South Shetlands) 
in 1965, revealed two previously unrecognised 
species representing the genera Onyehiurus and 
Folsotia, wot previously recorded from the An- 
farclic. The species from these collections are 
listed in Table 1 and the new records given in more 
detail below, 


Family Onychiuridae 
Onyehlurus sp. 
MGA 

Material examined 7 2% 13 1 Imm. Umotd) 
SAMA, 1% (mntd) BPBM, |  (mntd) AMNZ, 
40 spee, (ale) BPBM, 1 spec. (alc) SAMLA, South 
Shetlands, Deception Island, Whalers Bay, under 
whale bones on beach, 3.411965, J. Strong; sanie 
locality and date, under whale bones and debris 
on beach, 12 (mntd) AMNZ, 2 spec, (ale) BPBM. 
Deseriplion, Body Lenvuasy) LS 1.6mm, 1.2 
mm. Colour: white (after preservation). 

Posteuntennal organ consisting of many 
simple vesicles i two. parallel rows! pseudovelli 
paliven 33, 022, 333330, with slight variation (ie, 
antenual base pseudocelli 4, 3 in one of the 
specimens examined); seta mon thorax | present: 
subeosue Ll and TE without pseudocelli; claw 
Without tooth, empodial appendazes with macrow 
lamellae; ventral tube with 10+ 10 setae; anal spines 
present and single furcal rermman| with 2 setae; ab- 
omen Vo owith s! ela amerion to S seta. 
maeroehaene about twice anal spines. tn lengths 
abdomen. VE with slightly divergent lateral setae 
wich ad single median seta (Pig: 1) 


‘South Avistralian Museum, Nori Terrace, Adelaide, 
S Aust, 5007 

) Auckiiadl Tostitute ind) Museum, 
Zoning, 


Auckland, New 


Distribution of such characters as the 
pseudocelli and sefal patterns within the popula- 
tion is difficult to analyse becuse of Lhe poor 
slate of preservation of the material and prevents 
identification to species. Using Fjellbere’s (1980) 
key the animals are near fo the Arctic littoral 
species, O. islumedicus Gisin although the »! setit is 
silualed more anteriorly in the Antarctic species, 
We consider the present species to be a member of 
the wrnatus group. Controversy over the specitic 
value of characters has led to both a proliferation 
Of species mames in the genus and a number of 
synonymies (Gisin 1960, Hale & Rowland 1977, 
Pitkin 1980). 

This is a new record for Onvehiurus. The 
specimens Were previously misidentified as 
Tillhergia mixta Wahlgren (Wise 1971), however 
ihe idenury ol other reeords of 7 wityra from 
Deception Island by Gressitt er af (1963) and Wise 
(1967) have been confirmed (K.AGEW. det, 1983). 


Family Isotomidae 
Folsomia candida Willen 
Muterial examined 1 imm. (mmtd) BPBM, South 
Shetland Islands, Deception Island, Whalers Bay, 


I aad ‘ 
‘ia a 
7 
— A _ 
" | ‘ xs — . 
' os ~ ~ 
/ , an 
® / ~ a ~ 
_ = 4 = 
i ae ps. 45 
fal ~S 
2 Ps 
{ ke M = { \ 
~ \ 
—— all ~ “ “ 


Vig. b. Dorgal-yiew of abdomen Vand Viol 2 shows 
peemtocelll aad chaerotasy Mo = maerochacta, s'S sun 
sory acta Seale line 100 jk. 


204 


P. GREENSLADE & K. A. J. WISE 


TABLe 1. Collembola collected from Deception Island 1965. C=Cosmopolitan species, N=Native species 


Collection Data 


Whale Bay, under whale 
bones on beach 3.1i.196S. 


Whale Bay, under whale 
bones and debris on beach 
311.1965, J. Strong 


Species Distribution Published Records of 
Specimens 
Hypogastruridae f r 
Hypogastrura viatica (Tullb.) Cc Wise, 1967: 129 (2 specimens) 
Onychiuridae this work (recorded in error as 
Onychiurus sp. J. Strong LE Tullbergia mixta Wahlgren by 
Wise, 1971: 59) 
Isotomidae ; 
Cryptopygus antarcticusWillem N Wise, 1971: 65 
Cryptopygus caecus Wahlgren N this work 
Hypogastruridae : 
Hypogastrura viatica 16 Wise, 1967: 129 (1 specimen) 
Onychiuridae this work (recorded in error as 
Onychiurus sp. 27C Tullbergia mixta by Wise, 
1971: 59) 
Isotomidae 
Folsomia candida Willem Cc this work 
Archisotoma brucei(Carpenter) N Wise, 1967: 137 
Cryptopygus antarcticus N Wise, 1971: 65 
Cryptopygus caecus N this work (2 imm. specimens) 


Other collections from 
Deception Is. recorded Wise, 
1967: 136 


NN 


under whale bones and debris on beach, 3.11.1965, 
J. Strong. 

Although the specimen is immature, it is of 
adult size, (1.8 mm) and agrees in all respects with 
Stach’s 1947 redescription of the species and with 
other reliably identified specimens. 

Six species were found in the two collections 
made by Strong (Table 1) and this is a rich fauna 
for a species poor environment such as the 
Antarctic. The reason for this could be the high 
level of decaying organic matter which was almost 
certainly present. It is usual for temporary 
habitats of this type to be colonised by widely 
distributed r selected species (Greenslade & 
Greenslade 1983). 


Discussion 


Both Onychiurus and Folsomia are most highly 
diverse in the Northern Hemisphere with many 
species in and near the Arctic Circle. The southern 
hemisphere equivalents of these two genera are 
probably Tiullbergia and Cryptopygus respectively, 
which dominate antarctic faunas both in numbers 
of individuals and species. In Australia 
Onychiurus is found predominately in disturbed 
humid habitats; it is rare in native vegetation. In 
contrast Ti//bergia species of the southern antarc- 
tica Lubbock and mixta groups are restricted to 
soils which have retained their native vegetation 
cover. It is likely that all Onychiurus species in 
Australia have been introduced while species 
belonging to these groups of 7u//bergia are native. 


Onychiurus had not been found previously further 
south than Campbell Island (52° 33’S) where 
O. subantarcticus Salmon 1949 has been recorded 
from a disturbed habitat. Onychiurus species of 
the armatus group until now had been recorded 
on every continent but Antarctica. Folsomia can- 
dida is a species also found on every continent up 
to the present except Antarctica but is not abun- 
dant. Both species are easily cultured in the labora- 
tory with plentiful food and moist conditions and 
can mount dense populations under these condi- 
tions. 

It seems likely from this and other distribution 
records that both the Cryptopygus species and 
Archisotoma brucei aré native to the area while 
the Onychiurus and Folsomia may be relatively re- 
cent introductions. Hypogastrura viatica is a 
widely distributed species in the southern 
hemisphere including other subantarctic islands 
but, because there are no endemic congeners in 
the Australasian Region, (all species which have 
been found so far also occur in Europe), it is 
possible that this species also is not native to the 
region. What is known of the biology of this and 
the other genera would tend to support these con- 
clusions. There is evidence for the recent introduc- 
tion of another Hypogastrura species to a suban- 
tarctic island. Hypogastrura denticulata (Bagnall) 
was collected for the first time on Macquarie 
Island in 1974 despite intensive earlier collecting 
there in the 1960’s. The collecting site was a rub- 
bish dump near the biology hut. Since that year 
H. denticulata has been collected from a number 


ANTARCTIC COLLEMBOLA 205 


of different areas on the island (P. Greenslade & 
kK. A. J. Wise, unpubl.). 

Further collecting on Deception Island is 
needed to determine whether Onychiurus and 
Folsomia have become established. 

Acknowledgments 


This work was carried out as part of a project 
funded by the Trans-Antarctic Research 
Association. Thanks are due to L. Deharveng for 
comments on the manuscript. 


References 


FIELLBERG, A. (1980) Identification Keys to Norwegian 
Collembola. Norwegian Entomological Society. 

GISIN, H. (1960) “Collembolenfauna Europas” (Mus. 
Nat. Hist. Geneva: Geneva). 

GREENSLADE, P. J, M. & GREENSLADE, P. (1983) Ecology 
of soil invertebrates. /n “Soils: an Australian view- 
point’. Division of Soils, CSIRO, pp. 645-669. 
(CSIRO: Melbourne/Academic Press: London), 


Gressit1,J. L., LeecH, R. E. & Wise, K. A. J. (1963) 
Entomological investigations in Antarctica. Pacif.Jn- 
sects 5, 287-304. 

HALE, W. & RowLanp, J. P. C. (1977) Biochemical 
“finger prints” as indicators of taxonomic status 
within the genus Onychiurus. Revue Ecol. Biol Sol. 
14, 535-562. 

PITKIN, B. R. (1980) Variation in some British material 
of the Onychiurus armatus group (Collembola). Syst. 
Ent. 5, 405-426. 


SALMON, J. T. (1949) New Subantarctic Collembola. 
Cape Exped. Ser. Bull. Dpt Sci, & Ind. Res, 4, 1-56, 

—(1964) An index to the Collembola, R. Soc. N.Z. 
Bull. 7, 1-651. 

StacH, J. (1947) The Apterygotan fauna of Poland in 
relation to the world fauna of this group of insects 
Family: Isotomidae. Pol. Acad. Sc. & Lett, Cracow. 
1-488. 

Wise, K. A. J. (1967) Collembola (Springtails). 
Antarctic Res. Ser. Washington. 10, 123-148. 

— (1971) The Collembola of Antarctica. Pacif. Insects 
Monogr. 25, 57-74. 


FIRST REPORTED TRILOBITES FROM THE LOWER CAMBRIAN 
NORMANVILLE GROUP, FLEURIEU PENINSULA, SOUTH AUSTRALIA 


BY J. B. JAGO, B. D, DAILY, C. C. VON DER BORCH, A. CERNOVSKIS & N. SAUNDERS 


Summary 


Two recently discovered fragmentary trilobites are the first found from the Lower Cambrian 
Normanville Group of Fleurieu Peninsula, South Australia. They occur about 340 metres above the 
base of the Heatherdale Shale, the uppermost unit of the Normanville Group. An exact age within 
the Early Cambrian cannot be given on available evidence although a middle Early Cambrian age 
seems likely. The species represents one of the earliest members of the Conocoryphidae. The 
trilobites are associated with probable conchostracans. Trace fossils and possible desiccation 
(?synaeresis) cracks are present within the Heatherdale Shale just below the trilobite-bearing 
horizon. 


FIRST REPORTED TRILOBITES FROM THE LOWER CAMBRIAN NORMANVILLE 
GROUP, FLEURTEU PENINSULA, SOUTH AUSTRALIA 


by J. B. JAGo*, B.D. Day, CC. VON Beek Borer, AL CLRNOVSKIS & N) SAUNDERS 


ah minary 


Jago. J BY DAY, Bo. Vom Ob Boren, € 


» Chiewskis, No, SAUNDERS, 


A, (1984), Pivst Reported 


Iilobties trom the Lower Cumbrian Narvuraille Croup, leurioy Penifisala, South Australia. Truss, 


RK, Sac 


S. Ausi 18 (4), 207-211, 13 Deweniber, 


1984. 


Iwo pecenily discovered fragmentary trilobites ate the first found from (he Lower Cambrian Norman- 


ville Group ol Pleuries Peninsula, South Australia 


Chey oecur about 340 metres above the base ot the 


(leatherdale Shale, the uppermost unit ol (he Normanville Group, An exact age witli rhe Raely Cambrian 


cannot be viven unavailable evidence although a muddle Larly Cambrai ase seems likely. 


The species 


represents one OF (he earliest members of the Conovoryphidae Che trilobites are associated with prababte 
conchostracans, trace fossilsaind possible desiccation (2synaeresis) cracks dec present within the bleather- 


dale Shale just below the trilobite bearing horivon, 


Kiv Worhs: Setlick Hill, Fleurieu Peninsula, Heatherditle Shale, Normanville Group, rrilobites, Con- 


oceryplidue, Barly Cambrian, 


Introduction 


The purpose of this work is to report the discoy- 
ery of trilobites trom the Heatherdale Shale in the 
Sellick Hill area, 55 Kirt south of Adeliide, The two 
specimens figured herein represence the firsr trilo- 
bites known from the Normanville Group (Daily 
& Milnes 1973), The Normanville Group, which 
rests Unconformably on the late Precambrian 
Adelaidean Marino Group, comprises rhe Mt Ter- 
rible Formation (base), Wangkonda Formation, Sel- 
lick Hill Formation, Pork Tree Limestone and 
Heatherdale Shale (top), [tis overtain sharply but 
conformhably by the Carrickalinga Head Formation, 
the basal member of the Kanmantoe Group. Li the 
Sellick Hill area the Normanville Group ts 
comuontly overturned (Pig. 1), with the finer 
grained nits showing a promibent cheavape, 

Early Cambrnan Archaeoevatha were first 
reported trom what is now knownas the Fork Tree 
Limestone at Sellick Hill by Howehin (1897). Abele 
& McGowran (1959) gaye a bricl summary of 
pre-1959 work in the Sellick Hill area. Since 1897 
Humerous fossils have been reported from all mem- 
bers Of the Normanville Group by various workers 
(au. Abele & McGowran 1959: Daily 1963, 1969, 
19760; Daily et a/. 1976, 1982; Wrieht 1969). includ- 
ing archagocyathids, hyolithids, pastropods, 
sponges, brachiopods, tommotiidés and others. 

The Heatherdale Shale (Abele & MeGowran 
1959) comprises a lower calcareous member and an 
upper dark grey lo black shale and siltstone, eencr 
ally lacking in carbonate, Nodules and stringers of 


*Schoolot Applicd Geolowy, South Australian (nerivure 
of Teclinolovy, P.O, Bos 1, logle Farm, 5. Aust, 509% 

) Department of Geology and Mineraloyy, Universiny a 
Adelitide. 

PE School olf Furth Sciences, blinders University 


blavk phosphate oecur, and are particularly con- 
spicuous in the upper member (Daily er wi 1976). 
Laleral and verdical changes in carbonate content 
within the formation are quite marked (Daily 1963; 
Daily e¢ a! 1976), Hyolithids, sponges, brachiopods 
aid gastropods occur sparsely throughout the 
Heatherdale Shale, 


SELL i, a 
BEACH 


CAINGZOIC 


ATWOWS CIPO Loramie tecewe 


‘CAMBRIAN 


HEATHEROALE arlaly 
| =URK THEY IMM Cur 


[T= | 
PST} WAN TRONTA PoMMAr Koh 


| MAT) TEM FARIS ID PPA POT IL the 


PROTEROZOIC 


WRG Pe RUMI 
Wiel Si at Se 


Sree tne ceo 
overl unmet bec 


Pete ring 


Ta tly nandes (ically 


Mam Cie nin 


hie TE Geolaey ot the Setliek BDH ares, 


ih i showing the 
iriubite Loewy. 


208 Jj. B. JAGO, B. DAILY, €. C. VON DER BORCH, A. CERNOVSKIS & N. SAUNDERS 


Pig. 2. a P2432la, latex cast of external mould of concoryphid trilobite, X3. The spine bases on the axial rings of 
the thorax probably supported spines of a length about equal to half the width of the axial ring. b. P24321b, internal 
mould of conocoryphid trilobite, X2. c. P24322a, latex cast of external mould of second trilobite specimen, N3. 


In February 1983, the first trilobite (Figs 2a, b) 
from the Normanville Group, was found by three 
of us (C. C. V. B., A. C., N. S.) in the upper mem- 
ber of the Heatherdale Shale about 340 metres 
above the base of the formation in a road cutting 
on the Main South Road (Fig. 1). Subsequent 
searches by all authors have yielded only one fur- 
ther specimen (Fig. 2c). Abundant specimens of 
what are probably small conchostracan arthropods 
are found at the same stratigraphic level, where they 
occur mainly in small black phosphate-rich nod- 
ules. Organic-walled microfossils have also been 
found in the Heatherdale Shale (Foster ef al. 1985). 
Abundant trace fossils (Fig. 3) are known in several 
horizons one to two metres below the level where 
the trilobites were found. This is the first record of 
trace fossils from the Heatherdale Shale. 


Possible desiccation cracks (Fig. 4) are present 
about 2 metres below the trilobite horizon. If these 
are indeed desiccation cracks, rather than synaere- 
sis cracks, then this suggests a very shallow water 
marine depositional environment for this part of 
the Heatherdale Shale. However, it should be noted 
that the Early and Middle Cambrian conocoryphids 
in eastern North America are characteristic of 
faunas found some distance offshore (Lochman- 
Balk & Wilson, 1958) 


Palaeontology 
Both available trilobites are preserved as external 
and internal moulds. They are described briefly 
below. The specimens figured herein are housed in 
the palaeontology collection of the South Aus- 
tralian Museum, 


LOWER CAMBRIAN TRILOBITES, FILEURIEU PENINSULA 209 


Tig. 3a, b. Trace fossils in an horizen one to two meres below the level of the Lilobites. 


Fig. 4, Desiceation(”) or synaeresis (?) cracks, Heatherdale 
Shale, about 2 metres below the level of the trilobites. 


Specimen I 

Description: The first specimen (P2432la + b, Figs 
2a, b) is distorted and crushed; it comprises a cepha- 
lon, a hypostome and parts of thirteen thoracic seg- 
ments. The surface is coarsely granulose with 
exception of the pleural furrows. Before distortion 
the cranidium was wider than long. The border fur- 
row js maderately deep; the border is narrow. The 
génal spines extend posteriorly to at least opposite 
the fifth thoracic segment. The glabella is narrow; 
but the preservation makes it difficult to determine 
just how far forward it extends. The occipital fur- 
row is moderately deep abaxially, but very shallow 
across the centre of the glabella. Other details of 
the glabella are obscured by poor preservation, 
although near the front of the glabella is a short 
furrow which is directed abaxially and posteriorly. 
Narrow, low eye ridges extend from the glabella 
almost to the border furrow. Although facial sutures 
appear to be absent, it is difficult to be certain of 
this due to the poor preservation, The shape of a 
small hypostome shows up on the right hand side 
of the anterior part of the cephalon where the shell 
of the vephalon has been crushed against the under- 
lying hypostome. 

Parts of 13 thoracic segments are preserved. Each 
Segment is about 11 times as wide as is long. The 
axis has a width about 0.3 that of the segment 
(excluding pleural spines), There is a large centrally 
placed spine base on each axial ring. Wide pleural 
furrows deepen abaxially; the pleural strips are nar- 
row and granulose. The pleural spines are long and 


210 J. B. JAGO, B, DAILY, C. C. VON DER BORCH, A. CERNOYSKIS & N. SAUNDERS 


narrow with the spine on the eleventh thoracic seg- 
ment being extraordinarily long and extending well 
to the posterior of the projected position of the 
pygidium. It is similar to the very long pleural spine 
on the eleventh thoracic segment of the late Middle 
Cambrian conocoryphid, Dasometopus maensis 
Korobov (1973, pl. 5, fig. 1). 


Specimen 2 


Comments: This specimen (P24322a + b, Fig. 2c) 
comprises parts of several posterior thoracic seg- 
ments as well as part of a very poorly preserved 
pygidium. This specimen probably belongs in the 
same species as Specimen 1, but does not warrant 
further description. 


Discussion: The specimens probably belong to a 
new genus of the Conocoryphidae, although they 
are too poorly preserved to make an accurate 
assessment. One of us (B.D.) has collected what may 
be a representative of the same conocoryphid genus 
in association with other polymerids and an eodis- 
cid from the upper part of the Parara Limestone 
in the Chace Range of the Flinders Ranges. When 
this material is studied it may assist in positively 
identifying the specimens from the Heatherdale 
Shale. The specimens described herein belong in the 
Conocoryphidae and add to the rather limited num- 
ber of Conocoryphidae reported from Lower Cam- 
brian sediments. As noted below, the Heatherdale 
Shale is of Early Cambrian age, whereas most 
known Conocoryphidae are of Middle Cambrian 
age. 

However, the concoryphid genera Pseudatops 
and Atops are known from the upper part of the 
Lower Cambrian of eastern North America 
(Lochman-Balk & Wilson 1958, fig. 4). Pseudatops 
is also known from England and Wales where it is 
found in the ‘Protolenid-Strenuellid” Zone of 
Cowie et al. (1972) which is regarded as of late Early 
Cambrian age by Cowie et a/. (1972) and Rushton 
(1974). 

Korobov (1966) described two new genera of 
conocoryphids, Atopina and Ivshiniellus from the 
Lower Cambrian Aldanian Stage of Tuva. Korobov 
(1973, tables I and II) suggested that all known 


of the Aldanian Stage. Fritz (1973) reported a 
conocoryphid from the middle Lower Cambrian ol 
north-western Canada. Further discussion on the 
subject of Early Cambrian conocoryphid correla 
tions is reserved until two of us (B.D. and J.B.J.) 
describe the material mentioned above from the 
Flinders Ranges. 


Age of the Heatherdale Shale 


The two trilobite specimens figured herein are so 
poorly preserved as to preclude formal description 
and adequate comparison with known taxa, and 
hence are of limited biostratigraphic use. However, 
as noted above, a conocoryphid trilobite is known 
from the Parara Limestone in the Flinders Ranges. 
Daily (1972, 1976a) has shown that the Parara Lime- 
stone is of Atdabanian age within the Early 
Cambrian. 

The Fork Tree Limestone, which conformably 
underlies the Heatherdale Shale, comprises two 
members, the lower one of which contains abun- 
dant, but poorly preserved archaeocyathids. D. 
Gravestock (pers. comm.) has identified the follow- 
ing archaeocyathids:—?Ajacicyathus  sellicksi, 
Dokidocyathus and Pycnoidocyathus, but notes 
that because all are long ranging genera an exact 
age within the Early Cambrian cannot be given. 

A correlation chart given by Daily (1976b) sug- 
gests that the Heatherdale Shale is stratigraphically 
well below the late Early Cambrian fossiliterous 
sequences exposed on the north coast of Kangaroo 
Island as described by Daily e¢ a/. (1979). Hence 
at this time on the basis of fossil evidence it is not 
possible to give a more precise date to the Heather- 
dale Shale than ‘somewhere in the middle part of 
the Early Cambrian”. Suggested Early Cambrian 
correlations within southern Australia are given in 
Cooper & Grindley (1982). 


Acknowledgements 
We wish to thank J. Shergold (Bureau of Mineral 
Resources, Canberra), R. Jenkins (University of 
Adelaide), P. Jell (National Museum, Melbourne), 
A. R. Palmer (Geological Society of America, 
Boulder) and C. B. Foster (Western Mining Cor- 


Early Cambrian Conocoryphidae are of approxi- poration, Adelaide) for valuable advice and 
mately the same age, equivalent to the upper part — discussion. 
References 


AneLe, C. & McGowran, B. (1959) The geology of the 
Cambrian south of Adelaide (Sellick Hill to Yankalilla). 
Trans R. Soc. S. Aust. 82, 301-320. 

Cooper, R. A, & GRINDLEY, G. W, (Eds) (1982) Late 
Proterozoic to Devonian sequences of Southeastern 
Australia, Antarctica and New Zealand and_ their 
correlation. Geol, Soc. Aust, Spec. Publ. 9. 


Cowir, J. W., RUSHTON, A. W. A. & STUBBLEFIELD, C, 
J. (1972) A correlation of Cambrian rocks in the British 
Isles. Geol. Soc. Lond., Spec. Rep, 2. 

Dairy, B. (1963) The fossiliferous Cambrian succession 
of Fleurieu Peninsula, South Australia, Ree. S. Aust. 
Mus. 14, 579-601. 

—— (1969) Fossiliferous Cambrian sediments and low 


LOWER CAMBRIAN TRILOBITES, FLEURIEU PENINSULA 2 


grade metamorphics, Fleurieu Peninsula, South 
Australia. /# Daily, B. (Ed.): Geological Excursions 
Handbook, 49-54, ANZAAS, Section 3, 1969, 

—— (1972) The base of the Cambrian and the first 
Cambrian faunas, Univ, Adelaide, 
Precainbrian Research, Spec. Pap. 1, 13-41. 

(1976a) New data on the base of the Cambrian in 
South Australia. /zv. Akad. Nauk. Ser. geal. 3, 45-352 
(In Russian), 

—— (1976b) The Cambrian of the Flinders Ranges. /n 
Thomson, B. P., Daily, B,, Coats, R. P. & Forbes, B, 
G, Late Precambrian and Cambrian Geology of the 
Adelaide ‘Geosyneline and Stuart Shell, South 
Australia, 2514 Int, Conger, Excursion Guide No. 334A, 
1-56, 

——-, FIRMAN, J. B. Forpes, B, G. & LInpsay. J, M, 

(1976) Geology. /n Twidale, C. R., Tyler, M. J. & Webb, 
B. P. (Eds), Natural History of the Adelaide Region, 
§-42. (R. Soe. S. Aust.: Adelaide). 
—, Jaco, J. B. & JAxtes, P. R. (1982) Lower Cambrian 
sediments, Precambrian-Cambrian Boundary and 
Delamerian Tectonics of Southern Fleurieu Peninsula. 
Jn Oliver, ROL. & Gatehouse, C. G. (Eds). Excursion 
Guides B, B. B,, B, Geology of the Adelaide 
Region, Fourth International Symposium on Antarctic 
Earth Setences, Adelaide, 30-4). 

& Mirnrs, A. R. (1973) Stratigraphy, structure and 

metamorphism of the Kanmantoo Group (Cambrian) 

in ils lype section east of Tungkalilla Beach, South 

Australia. Trams R. Soc. S. Aust 97, 213-241. 


Cearre for 


——, MILNes, A. R., Twibane, C. R. & Bourne, J. A. 
(1979) Geology and Geomorphology. /n Tyler, M. J., 
Twidale C. R, & Ling, J. K. (Eds) Natural History of 
Kangaroo Island, 1-38 (R. Soc, S, Aust,: Adelaide), 

Foster, C. B., CeRNovSKIS, A. & O'BRIEN, G. W. (1985) 
Orvanic-walled microfossils from the Early Cambrian 
of South Australia. A/cheringa Y (in press). 

Fritz, W. FH. (1973) Medial Lower Cambrian trilobites 
from the Mackenzie Mountains, northwestern Canada. 
Geol. Surv. Canada, Paper 73-24. 

HowceHin, W. (1897) On the occurrence of Lower 
Cambrian fossils in the Mount Lofty Ranges. 7rans R, 
Soe. S. Aust 21, 74-86. 

Korornov, M. N, (1966) New trilobites of the family 
Conocoryphidae from the Cambrian of the Siberian 
Platform and Tuva. Paleont. Zhurnal. 1966, no. 4, 
92-97. (In Russian). 

(1973) The trilobite family Conocoryphidae and its 
significance 10 the stratigraphy of the Cambrian 

‘deposits. Trudy. geo. Inst, Moscow, 211. (In Russian). 

LOCHMAN-BALK, C. & Witson, J. C. (1958) Cambrian 
biostratigraphy in North America. J Paleont, 32, 
312-350). 

Rusuton, A. W. A, (1974) The Cambrian of Wales and 
England. /n Holland, C. H. (Ed.), Cambrian of the 
British Isles, Norden and Spitsbergen. 43-121. (Wiley: 
London). 

Wricht, R. G. (1969) Geological and Geochemical 
Report on the Sellick Hill Area, Mining Review, S. Aust. 
Dep. Mines 126, 21-32. 


LEAF MARKING IN RANGELAND GRAZING STUDIES 


BY R. T. LANGE 


Summary 


Sheep graze extensive areas of arid zone vegetation in South Australia. A simple leaf-marking 
technique is described that provides a quick and novel method of assessing the grazing pressure. 


BRIEF COMMUNICATION 


M3 


LEAF MARKING IN RANGELAND GRAZING STUDIES 


Sheep grave extensive areas of arid sone Vegetation in 
Sour Australia. % simple leafmarking technique is 
described (hal provides a quick amd novel method of 
assessing the grazing pressure 

On saltbushes (4 /riplex vesicuria) it is quick and simple 
to mark hundreds of the outer leaves each witha paint 
dot abou! 2mm in diameter and to provide for rapid 
reinspection OF the dots bv painting guidemarks on me 
fishes! stems, 

Free grazing sheep are colourblind! and geraze the 
marked leaves along with Che rest, showing no bias for 
Or against marked leaves provided that (he dots are small 
and odorless, That was shown by § trials in-gach of which 
approximately 200 random shoots were marked and 200 
more were photographed before exposure Lo six weeks of 
pastoral grazing. Contingency tables of the results (as in 
‘Table | for example) were all non-significant. Fenced-off 
control marked shoots shawed no. losses. 


TAH 1. Contingency table campuring sheep grazing 
lasses uf physically tagged (painted) and untauched 
(photographed) leaves an salthushes, x} = 0.05 whieh 
iy nonsignificant, Showing that sheep did nol vraze 
proimed leaves. differently to nan-painted leaves. — 
ileal Dante aa sec ath A SOE 

aller @ weeks 


missing temaining totaly 


painted M) l74 24 
intially photegraphest 33 173 206 
Totals 63 347 40 


--—————_——————— 


So this technique establishes, for exch locality where 
alarge number of leal-dots is deployed, the likelihood of 
an Guler shoot of the salitosh being grazed during any 
month or similar period (the percentage chance of being 
grazed), 

To Hustrate these ikelihoods a uniform saltbush plain 
wats studied near Whyalla, South Australia, where 3 sheep- 
socked paddocks shared a fenveline Tjunction. The pad- 
docks fotalled about 5000 ha and had independent locks 
of about 275 sheep each (6 ha sheep '). Ouadrats 60 © 


Leaves 


Percent Lass Of Marked 
5 


rs | 
O65. 1 
Egesta Acclimulation (Ka) 


Fig. |. Percent of marked saltbush leaves mken versus 
sheep evesta accumulated over a six-week period ar each 
of 1) focalines in saltbush vegetation. Further 
explangdcon is io the text, 


4m were marked in each paddock near the Tyunctian and 
200 leaves were marked on the saltbushes on each, as 
follows. 

All bushes on the quadrat were numbered and were 
selected by random numbers. On large bushes 10 leaves 
were marked and on small bushes 5. Each leaf was on a 
separate terminal shoot of the outermost foliage, Terminal 
shoots were selected to scatter the marked leaves over the 
bush. The pant used was yellow quick-drying enamel 

The quadrats were then picked clean of egesta and were 
left for 6 weeks, When the marks Were recounted and rhe 
cgesia weighed that had fallen on each quadral during the 
period, Fig. | shows (he results, 


30 


20 


10 


Percent Loss Of Marked Leaves 


10 20 
Percent Loss Of Total 


Foliage Biomass 


30 


Fiv. 2, Pereen| loss of marked saltbush leaves versus 
percent loss of total foliage biomass in a small-paddack 
praving trial. The samples were 10 » 10m areas within 
a 10 %* 200m paddock which sheep grazed unevenly 
for a week. 


‘As can be seen, the likelihoods were in the surprisingly 
tigh range 828" and much of the variation from quadrat 
to quadral could be accounted for by quadrat egesta 
accumulation, which is known from other siudies* to 
index Nocktime. From this iv should be clear (hat the 
marking technique has much potential for use in rangeland 
studies, As further examples, Fig, 2 shows thar percen- 
tage loss of marked leaves correlates with percent loss of 
total foliage biomass’ in small-paddock experimental 
Work, Where if is also gecounted for in terms of sheep 
egesta accumulation (Pig. 3). 

The following caution must be heeded. Care when 
applying the paint dots is essential Experience with this 
techtique in the hands of students shows that spillages 
on foliage and dots bigger [han prescribed can result in 
sheep avoiding the marked leaves. 

The particular value of this technique is that it allows 
much more sensitive discrimination of the depradations 


N 
= 


ne] 

oO 

¥ 30 

oO 

= 

5 » 20: 
g 

no 

no 

S540 

€ 

oO 

2 

® 

a 


3 4 5 6 
Egesta Accumulation (Kg) 


Fig. 3. Percent loss of marked saltbush leaves versus sheep 
egesta accumulated in a small-paddock grazing trial 
similar to that described in Figure 2. 


of sheep on saltbush than is otherwise possible. The error 
terms associated with plant biomass estimation techniques, 
which are the only alternatives, are in general very great’. 


"Huxley, J. (1972) “Memories I’ (Penguin Books: 
London). 

"Lange, R. T. (1983) Trans. R. Soc. S. Aust. 107, 137. 
‘Andrew, M. H., Noble, I. R. & Lange, R. T. (1979) Aust. 
Rangel. J. 1, 225-231. 


R. T. LANGE, Botany Dept, University of Adelaide, Box 498, G.P.O. Adelaide, S. Aust. 5001. 


RANGE EXTENSIONS OF REPTILES IN SOUTH AUSTRALIA 


T. D. SCHWANER 


Summary 


The ranges of some reptile species already known to occur in South Australia have been greatly 
extended by recent field studies. Here we report these findings and briefly comment on their 
significance. 


BRILE COMMUNICATION 


RANGE EXTENSIONS OF REPTILES IN SOUTH AUSTRALIA 


The ranges of some reptile species already known to 
decurin Sonth Australia have been greatly extended by re- 
cent field studies. Here wereportrhese findings and briefly 
comment on their significance, 

Voucher specimens reported here aie located in the 
following museum collections: National Museums of Vie- 
fora (NMYV); South Australian Museum (SAM). The 
nomenclature follows Comser except lor Gehvru par 
Mirescens (sens Start’). 


katnily: Gekkonidae. Gelivra purnurescens Storr 


Locality regard) S, Aust SAM R2400L Clayton Bore, 
Cordilla Downs Stn (29°17 'S, [38°23 EB). 


Comimicans: Shorr reported the distribution of G. puer- 
Purescens its Hie arid interiar OF Western Australia, cen- 
tral and southern Northen Terrirocy and torthery SA, 
The species is distinc) chrontosemaily from all other 
Gehwra (OC. Moaly, wi litt. 7.81982), Moritz (pers. corn) 
has calleeted another specimen recently front MeDauall 
Peak Sty, S.A, (29 30'S, 194754 bE). These records repre- 
SeHE ANIONS OF Hie raneeal Lhe species well inte cen- 
lal SAL 


Tamily: Scincidve Beenie cummnghanie tray), 

Locality records: S. Aust: SAM RIG9R9, R17355, 
R22293-97, West [sland (35°37'S, 138°96°R) SAM 
RI7352, AM R93474, Pisheries Boaeh (35 40'S, LIN -07"E), 
SAM RIT&AS4, Deep Creek (3536'S, 148 1S°L): SAM 
R17353,, Blowlwle Creck (45°3R'S, [38 10'b), 

Comments: In S, Aust this species wine knowrl einly 
fram The Westen slopes of the Mi Lolly Ranges! * 
Speeimens were volleeted droufid the Coast of Pleurieu 
Perosula trom Cape Jervois to West Esland, Phe West 
Ishundl specimens represent the only known offshore tstartel 
population Gk This species in Australia. Patan &: Pabor 
listed (he spevies as oveurring on West dsland, bul ap- 
parently were ynaware of phe wniqueness al the peapula 
fon, dnd fo voucher specimens were collected. These 
coun, Chillwelling populafions are sawiller iy adull 
boty sive aid mon bnghtly patterned than ilune pepuli- 
Hons SOA. Tn ies respeet they resemblespeenens ram 
the Moonta Kanges i NSW, and might represent a new 
subspecies (G. Sha & Ro Sadlien fi eh, Va TYR4), 


Lerishe ierar’s arentcalir Stuavtr 

hovalily ‘eeards: S. Aust SAM) R296. Nuyis 
Archipelapo, Crour Island (32°.0'S, 132°30'Ly: SAM 
R2S08K, Franklio (shind West (32 27'S, 132 40°F). 

Commicniss Dhese are new records ob 2. a arernticatiy 
trom afshore istinds of S Nast. Previously, bwu specinens 
(SAM RIS99 gid RSRGO) were culldeled from “West C aust. 
SAT’ by "Constable Hollorin, Powers Bay" an 
“VENT LQIOY and from “Head of Riehl, SAS in couscal 
sound hills by “HL, Bowshall” on 187 L964!" respectively. 
The Croat tstaind speciiner wis colleeted from under ce Tay 
limestone roek pear the centro of the island, The branklie 
To specinien was foun in sam under an uprooted shrub. 
Aldiateh the ishtid) specimens contormicad ta previons 
dese riplrone Willy titbats i contact and 23 youle pows at 


Pi b. devise mucratis urenicoly. 


midbaly, the dorsal pariern consisted ar twa bale dar 
solidral stripes, (wo ra rower paravertebral scripes une 
a narrower vertebral stripe (Fig. 1), similar ro We specnnens 
PL, Pe Reerets from Israelite Bay WAS! 


Menena grey Cray 

Locality revords: S. Aust: SMV 12566697], Kanpando 
[shind, Smith Creek valley between Linu Bay ail Cape 
Cassio) § kar WSW bin Bay township (35°38, 
137° 25'b). SAM R23601-05- L kink. Prospect Hill Penne 
shaw Rado nr Salt Lagoon (39 50S, 137 47'b) SAM 
R239635e44, Didley Conservalion Park oe "Sandhurse 
(38°S1'S, 137 SVE SAM R23547, RISZT3IO, Kimescore 
(35°R9'S, 137°99'R), 

Comments: The first specimens of A sre trom 
Kungardd [sland were collechal from under (lat stones 
along Sine Creek (Hutchinson, JW /ife, JUATAON |) Ade 
Witional specimens subsequently were culleercedl inde rub- 
bish and amo vatura liler fron three arher widesprened 
lowahbes an (he island. Give indivdiial was dbserveet (bul 
not capnired) ay the tacthwest ond at ihe istand at 
Vharvey’s Return!’ (35°48 'S, (36°38 1). This, chris supath. 
comment Hit elusive skik mh Nave aun istamd wine 
distribution, Hirelviesen (it dive. 2ON P98) riotedt har 
most A wee! from KE exhibited 24 longinidinsl rows ul 
scales on midhorty, suoikir fo hose reported trom favre 
Peninsula and citferent fran: the modal Wanmie (2) 
reporred for ere: pupulavions. 


amily: Glapidae, Dewmunsi psanieyis (Selileeel) 

Locality, record: 5, Austr SAM R275], Kinser 
Rungarva Isfand (35°39, 197 38 bs 

Cammentt Tis lirst reporled specnmen of Use yellow. 
taced wiipsnuke Prot KE wits colleeted from ander a street 
of galvanized iron in the vard of a Kinescote resulenee The 
snibaclull femahke (smourro-verr lenge 303 Hn) won 
similar in scalation anc coloue parrern ut tevelanel 
specimens of D psammophis.' The lone oecurrenge ot 
this diurnally active snake, and the failive to locare acluli- 
ional speciniens in reeent wilensive searches, susaeals {tal 
Ihis recard may represent d recent, aceidewial minrodiehon 
from rhe mainland, 


We |tiank A, Editrds, bk. Growler IN blalinies, ML. Tut 
whinsant. BL Plutcliinsou, Gt oterar, Wk. Miller, C) Moritz, 
R. Sudlien 1. Sehwanen A Seb waren, Schwarten OG. Shea, 
D. Watkins, 4 Watkins, wid S. Walkin for ther puny ven 
Mibihons to Chis repart. Bo Rushle prepated the 
Pholoerdoh, by Sehwaner rypwel thie eneeciieser tt 


216 


'Cogger, H. G. (1983). “Reptiles and amphibians of Paton, J. B. & Paton, D. C. (1977). Corella 1, 65-76. 


Australia?” 3rd Edtn. (Reed: Sydney.) °Storr, G. M., Smith, L. A. & Johnstone, R. E. (1981). 
°Storr, G. M. (1982). Rec. W. Aust. Mus. 10, 53-59. “Lizards of Western Australia. I. Skinks? (University of 
‘Ehmann, H. F. (1976). Herpetofauna 8(1), 2-5. _W.A. Press and W.A. Museum: Perth.) 

: (1976). Ibid. 8(2), 5-13. Storr, G. M. (1976). Rec. W. Aust. Mus. 4, 189-200. 


T. D. SCHWANER, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000 and B. MILLER, 10 Yarrow 
Crescent, Hope Valley, S. Aust. 5090. 


REPTILES NEW TO THE FAUNA OF SOUTH AUSTRALIA 
BY T. D. SCHWANER 


Summary 


Distributions of many reptiles in South Australia are poorly known, in spite of several attempts to 
document their ranges. Recent fieldwork and re-examination of old specimens lodged in the South 
Australian Museum and elsewhere, have resulted in the discovery of nine species and subspecies 
not previously recorded from the State. Here we report these findings and briefly comment on their 
significance. 


HEIDE COMMUNICATION 


REPTILES NEW TO THE FAUNA OF SOUTH AUSTRALIA 


Distributions of many repriles in Seuth Australia are 
poorly Keown, la spite of several attempts to document 
their ranpes.’’ Reeent fieldWork and re-examination ol 
old speelinens lodged in the South Australian Museum 
and elsewhere, have resulted in (he discovery of nine species 
and subspecies not previously revorded. fran the Stare, 
Here We wport these findings and brietly comment on 
their significance. 

The vougher specimens reported here are located in 
the following museniy calleetians: Australian Museum 
(AM). Sow Austrian Museum (SAM) Westert 
Austiatian Museu (WAM), The nomenelitune follows 
Cogger, except lor Clenaphorty precutalies dials 
(ens Stone sind Store en at ye 


Damilys Gekkomie, 2yaloddentias willanisi Kluge 

Locality reeords: 3. Aust: SAM RISL TAB, R28228, 
Danggali Cons. Park tr Morwanvale HS (3314'S, 
140 °42'h), SAM RISTOA-D, 4 km S Canopus HS 
GQ333'5, M042 by, SAM RISI20A-C, 20 km 5 
Canopus FPS (4940'S, 4042’ 2) SAM RIS042, Ralali 
Sr (43° 99'S. A088"), SAM RIGLOI, te Canopus Sin 
(33°45'S, WO37'E) SAM R21462, 20 kin N Renmark 
(Gas, OS by SAM: RI6097, RITI27, Dangwali 
Cone. Park (3341'S, 140°33'T); SAM R9915, Taldra R,, 
Loaton (34°27 'S, 4034 'LOy SAM R16297, tn, Birthday 
Dam (33) 20'S, 40r54 "by 


Conmmenis: 2 willed was found primarily under 
the bark of Yence posts in the riverlund ane prallee cour- 
iry Near Renmark, SA, d tone esrensin of same KOO 
kin alone (he Rives Mitrray devine tran NSAV! 
Subsequently, specimens of 2. wilt un the S.A. 
Museum from fhe same op nearby louatities were found 
1 he niisidentificd 9s 2% wnreriiedins Do wilhansé is 
disnngnished lam 20 intermedius in S.A. by fewer and 
more irregbirly spiteced tubercles on che dorsi and hy 
dW lick Of enlarged scales hetweer the tows of tubengles 
on the cil (ig, VAG 


bumly: Aparndie Crenaphorus peeiiatis Aili: 
Storr 


Locality records & Ause, SAM R22983, 17 km BE al 
the WIA-S.AL border (QL °35"S, 129 °(O'E). 

Comments: A single specimen was collected in tow 
coos shrabs above (he Nullarbor elffts, ©. pa diutis 
was reporteel Fram Old Buda arty 25 km Woof our 
locality.” Colon: ane pudtern of the specimen wenerally 
comfatnis ta Shore's deseripuan’) but ditters Tr having 
distinct blick edges te keels on all head seales (Pig, 1B). 

anily: Seineidac Learinve coven Start 

Locality reserd; & Avan SAM R271, Leek NS Cape 
Wamks Livlittouse 27 $8 "S. 022 Ey 

Comments: A specinen pt fo wteercee way colleeted 
in dn Dior trap fi open scrub, closed sedgelaniat 
labia Ch Slew (vr (fe, ta 2OS84) Heted War tive 
specimens Uf SE, Tucrtose,” rhe WoS, coutterpart ull & 
Cavern. previously were depoesital si the Brilish 
Museum (Nutt Histors? fram Soul Auseralia’! 


Shen wrote, “The locality is very suspect. The listing 
predates the description of 2. cavenrtey so could be 
either ds. dderiose ord covenievi” We lave nol examined 
the BM specimens and teel that the more recent finding 
iy suPlieienthy important to werranh reporting ws a new 
reeard for the State, The nearest locality previously 
recorded for this species is Portland, Vietoria, [00 kn E 
of the SA tewaliry, Colour and pattern af the specimen 
(Pig. 1) are bulder than reperted in previous deserip- 
trans!) > 


Lvernid cariiata Smith 

Localitv recurds; S. Aust) SAM R2910, Ooldea 
(30°27'S, I31°50-) SAM. R2d78, RYSIO, RIUS3S—41, 
Flinders tsland (44 43'S) 194 °91'R) SAM R3062, Cor 
albjenie VIS (82°37'5, 16°2L'b); SAM R3346, Penong 
(3156'S, (33°0l'E SAM RK 12727, Kounalda Cave 
(37°24'S, 129°50'Ei: SAM RIDR4E, 1 kin NS. Inala 
rovkhole Yuniburra Cans. Park (41°46'S, 133°29'L); 
SAM R22977-78, AM RIO7937, WAM R70803.04, 
15-17 kin E SAC WA border (319355, 129 "10/E), 


Comments: SAM specimens of &, carinala Sram SA 
were misplaced amore the similar species, &. striofara (Cr. 
Shea, vedint, (.x.1983). & carinede (Pig. 1D) differs trom 
& stride in umber of upper caudal scales (44-46 vs 
56-64) and number of subcaudals minus expanded up- 
per caudal seales (16-21 vs 9-7), respeetiwely. The 
nonmenelatural seaitus of &. curinete is in doubt. Cogger 
erat” qotd thar “the holotype (auc only specimen) of 
neha: (Peters) ts identical emost respects to A gerniy 
variety Savith, 1939 from southern WAL andl over which 
its name would have preeedence if the two dre showh to 
be conspecihe, 


Lerisia picluruia bayvnesii Storr 

Lowality reeerds: S. Aust: AM RI07936, T6 kin EB. 
SA/WA border (3142'S, 129°02 1h); SAM K23025-24, | 
kin E SAYWA border (31°38"S, 129°0L'E). 

Comments: The iva SAM specimens were ealleered 
from wider coastal enicalypr teal tien The AM specimen 
was collected from the Nullarbor elifl edge, 02 kin S of 
the Eyre Pighway, in leat liter beneal) Mefeleuce tees. 
Ot the same day, (eee specimens of 1. pierarand edward. 
sue (AM RID7944-40) 42.3 kon Eb. of the “Nullarbor 
Roadhouse ss were colleetod. This locality plaves yy 
havnesiiand Lop. edwardsue within 2X) kin of each arher 
und suyyests Pha the (Wo popiilittiona diay compacd along 
(re cost rather thaimte the Northolt the Nullarbor Plain 
is Suuewested by Storr! 


Lerisia savithire Stor 

Localily reterds: S. Avs SAM RIYO74, 3 kim SW New 
Muluaria Als (0°ld'S, (37°39) SAM RIYOTS. 15 kn E 
Prome Downs HS (3b 10'S, (39°58 lh SAM R20041, 
R20962-H3, Olympic Dam area (Roxby Downs) (0328, 
136° 56'E), 


Comments Cogger foted me distribution of Le veer 
thane as “Ciibsen Desert, WAL te soucliwesterr: NSW" hut 


218 


questioned its range in S.A. Specimens of L. xanthura 
(R19074-75) were taken from sand dunes with low shrubs 
and Casuarina trees, nr Lake Frome. Other specimens 
(R20941, R20962-63) were caught in pitfall traps in 
similar habitats nr Roxby Downs. Scalation and colour 
pattern of these specimens followed Storr!’ and Cogger.* 


Fig. 1. 


A. Diplodactylus williamsi, 
maculatus dualis, C. Egernia coventryi, 
carinata. 


B. Ctenophorus 
D. Egernia 


'Cogger, H. G. (1975), “Reptiles and Amphibians of 
Australia” (Reed: Sydney.) 

°____ (1979). Ibid. 2nd Edtn. 

(1983), /bid. 3rd Edtn. 

*Houston, T. F. (1973). Reptiles of South Australia, a 
brief synopsis. /n- ‘South Australian Year Book, 1973? 
(Govt Printer: Adelaide.) 

(1978). “Dragon lizards and goannas of South 
Australia” S. Aust. Mus. Spec, Ed. Bull, oer 84 pp. 
yler, M. J. (1979). In: Tyler, M. J., Twidale, 
C. R. & Ling, J. K. (eds) ‘Natural History a Kangaroo 
Island? (R. Soc, S. Aust.: Adelaide.) 

‘Thompson, M. B. & Tyler, M. J. (1983). Reptiles and 
Amphibians. /m: Tyler, M. J., Twidale, C. R., Ling, J. K. 
& Holmes, J, W. (Eds) ‘Natural History of the South East? 
(Royal Soc. S. Aust.: Adelaide.) 

“Waite, E. R. (1929). “The reptiles and amphibians of 
South Australia?’ (Govt Printer; Adelaide.) 


T. D. SCHWANER, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000 and B. MILLER, 


Crescent, Hope Valley, S. Aust. 5090. 


Notoscincus ornatus (Broom) 

Locality record: S. Aust: SAM R17324, Mokari, Simp- 
son Desert (26°19'S, 136°27'E). 

Comments: This first specimen was collected on red 
sand dunes among canegrass and scrub nr _ rotten 
Casuarina trees. The range of this species now extends 
well into northern S.A. 


Proablepharus reginae (Glauert) 
Locality record: S. Aust: AM R17385, Mt Davies, 
Tomkinson Range (26°15S'S, 129°16'E). 


Comments: Although used to construct Australia-wide 
distribution maps'*" this first record specimen of P 
reginae from S.A. never was formally reported (G. Shea, 
in litt, 13.11.1984). 


Family: Elapidae. Simoselaps anomalus (Sternfeld) 
Locality record: S. Aust: SAM R17466, Serpentine 
Lakes (28 °52'S, 129°11'E). 


Comments: This specimen was collected during a 
survey of the ‘Unnamed Conservation Park” in extreme 
western S.A, Previously included as a subspecies of S. ber- 
tholdi, Storr!’ elevated anomalus and littoralis to full 
species noting subsequently" that “these ‘subspecies’ are 
in fact parapatric or marginally sympatric species?’ The 
S.A. record extends the range of S. anomalus 350 km S in 
W.A. and the N-T. to deep within the recognized range of 
S. bertholdi’, further confirming Storr’s taxonomic 
assessment. '* 


We thank S. Berry, P. Bird, A. Edwards, J. Fowler, G. 
Harold, C. Harvey, M. Hutchinson, R. Hutchinson, K. 
Miller, B. Taverner, T. Morley, A. C. Robinson, R. Sadlier, 
S, Sarre, G. Shea, M. Thompson, R. Wells and J. White 
for their many contributions to this report. R. Ruehle 
prepared the photographs from colour slides. L. 
Schwaner typed the manuscript. 


°*Storr, G. M. (1982). Rec. W. Aust. Mus. 10, 199-214. 
"___ Smith, L. A. & Johnstone, R. E. (1983). “Lizards 
of Western Australia. Il. Dragons & Monitors?’ (Univ. 
W.A. Press & W.A. Museum: Perth.) 
"Kluge, A. G. (1963). Proc. Linn. Soc. 
230-34. 

"Storr, G. M., Hanlon, T. M. 
Rec. W. Aust. Mus, 9, 23-39. 
3 et J. Proc. R. Soc. W. Aust, 48, 45-54. 
West. Aust. Mus. 6, 147-87. 
“Commer H. G., ‘Cameron, E. E. & Cogger, H. M. 
(1983). “Zoological catalogue of Australia, 1, Amphibia 
and Reptilia?” (Australian Govt. Publishing Service: 
Canberra.) 

"Storr, G. M. (1982), Rec. W. Aust. Mus. 10, 1-9, 

; (1976). Ibid. 4, 241-256, 
'S____ (1967). J. Proc. R. Soc. 
(1971). Ibid. 54, 59-75, 


N.S.W. 88, 
S. & Harold, G, (1981). 


W. Aust. 50, 80-92. 


10 Yarrow 


LIFE HISTORY OF THE NARROW-WINGED PEARL WHITE ELODINA 
PADUSA (HEWITSON) (LEPIDOPTERA: PAPILIONOIDEA) 


BY A. R. MILNES, N. H. LUDBROOK, J. M. LINDSAY & B. J. COOPER 


Summary 


The pierid butterfly Elodina padusa (Fig. 1) occurs widely across northern Australia and south into 
central New South Wales and northern South Australia. It is commonly found close to its larval 
food plants, Capparis spp. A brief description of the early stage is given by Common & 
Waterhouse, but illustrations of these do not appear in any literature on Australian butterflies. A 
detailed description is provided here with illustrations and with notes on the life history, using 
material collected at Gammon Ranges National Park (30°30°S, 139°20°E), S. Aust. 


BRIEF COMMUNICATION 


214 


LIFE HISTORY OF THE NARROW-WINGED PEARL WHITE ELODINA PADUSA 
(HEWITSON) (LEPIDOPTERA: PAPILIONOIDEA) 


The pierid butterfly Elodina padusa (Fig. 1) occurs 
widely across northern Australia and south into central 
New South Wales and northern South Australia. It is 
commonly found close to its larval food plants, Capparis 
spp. A brief description of the early stages is given by 
Common & Waterhouse', but illustrations of these do 
nol appear in any literature on Australian butterflies. A 
detailed description is provided here with illustrations and 
with notes on the life history, using material collected at 
Gammon Ranges National Park (30°30'S, 139°20’B), 
S, Aust. 


Fig. 1. Elodina padusa, recently emerged from its pupa. 
Bar scale 1 em, 


Larval food plant: Cappris mitchellii Lindley (native 
orange, Capparaceae), a shrub or small tree up to 4 m 
high and with ovate or broadly oblanceolate leaves, 
Description of tmmature stages. Egg (Fig. 2A): height 
about | mm, diameter 0.5 mm; tapering uniformly to a 
blunt base and apex, with 16 vertical ridges of which some 
extend slightly beyond the apex and with numerous fine 
lateral lines; white at first, becoming pink or reddish. 

First instar larva (Fig. 2B): length 2 mm; head yellow- 
green with selae projecting anteriorly; body pale yellow- 
green, each segment with raised subdorsal, lateral and sub- 
lateral grey or black spots, each with a long black seta 
curved anteriorly at first then becoming erect. On 
abdominal segments 1, 2 and 3 the spots are enclosed in 
a red area which sometimes extends dorsally across the 
segment, while some segments have a few additional raised 
spots and setae. Second instar larva (Fig. 2C): length 
10 mm; head green with short setae; body pale green with 
a white dorsal line and numerous scattered setae, pro- and 
mesothorax and abdominal segments 2, 3, 4 and 8 with 


dorsal red-brown tubercles, those on segment 2 most 


pronounced, posterior segment bifid. 

Third instar larva (Fig. 2D): length 15 mm; head and 
body as in second instar but with dorsal red-brown 
tubercles usually prominent only on abdominal segments 
2 and 8. 

Fourth instar larva (Fig. 2E): length 20 mm, head and 
body as in third instar but prothorax with white dorsal 
area and several red-brown tubercles; dorsal line edged 
cream, 

Pupa (Fig, 2F): length 18 mm; slender, pale green, 
anterior produced into a long tapering point; thorax with 
a dorsal ridge; abdomen with a faint dorsal and paired 
lateral white lines, tapering uniformly posteriorly, slightly 
flattened, ridged laterally and with three pairs of dorso- 
lateral brown spots. 

Biology: Eggs are laid singly on either surface of the 
leaves of the food plant and occasionally on the stems. 
Young larvae make their first meal by eating a hole in the 
surface of a leaf, but in later instars larvae feed on the 
edges of leaves, their slender pale green bodies conforming 
cryptically with the eaten edge and thus providing some 
protection from predators and possibly parasitic insects. 
Pupation occurs on the leaves and stems of the food plant, 
the pupa being fastened by a cremaster and girdle. When 
attached to leaves the pupa is usually aligned with the 
central rib of the leaf and, as with the larva, the cryptic 
colour and shape make detection difficult. 

Eggs collected in early March were reared in Adelaide 
and reached the adult butterfly stage an average of 29 days 
after eclosion. The larvae passed through four instars. 
Such a short cycle from egg to adult suggests that a number 
of generations of this butterfly may occur in a single year, 
or in a single summer season. 

I thank the Wildlife Conservation Fund for financial 
assistance in field work, the National Parks and Wildlife 
Service for permission to collect material in Gammon 
Ranges National Park and A. E. Mitchell for the use of 
his vehicle, 


‘Common, I. F, B. & Waterhouse, D. F, (1981). “Butter- 
flies of Australia” Revised edition. (Angus and Robertson: 
Melbourne.) 


R. H, FISHER, 21 Seaview Rd., Lynton, S. Aust. 5062. 


270) 


— = 


D third instar larva. E fourth instar (mature) 


Fig. 2. Elodina padusa. A egg. B first instar larva. C second instar larva. 
larva. F pupa. Bar scales A, B = | mm: Cc, D, E, F = lcm 


NOTES ON THE REPRODUCTION OF NEPHRURUS DELEANI 
(REPTILIA: GEKKONIDAE) 


BYSTEVEN DELEAN 


Summary 


Reproductive biology of the knob-tailed geckos (Nephrurus spp.) is largely known. Although the 
ecology of some Nephrurus has been extensively studied in Western Australia, there has been only 
one report of clutch and egg sizes for this genus. Here we report the first successful hatching of 
eggs from a captive Nephrurus. 


BRIEF COMMUNICATION 


221 


NOTES ON THE REPRODUCTION OF NEPHRURUS DELEANI (REPTILIA: 
GEKKONIDAE) 


Reproductive biology of the knob-tailed geckos 
(Nephrurus spp.) is largely unknown. Although the 
ecology of some Nephrurus has been extensively studied 
in Western Australia', there has been only one report of 
clutch and egg sizes for this genus.’ Here we report the 
first successful hatching of eggs froma captive Nephrurus. 

Nephrurus deleani is the only knob-tailed gecko 
endemic to South Australia, where it is restricted to the 
Acacia dominated sand dunes surrounding Pernatty 
Lagoon’. On 24.x,1982 at 2015 hrs. we collected a gravid 
N. deleani (SVL. 78 mm, weight 13.0 g) in sand dunes at 
the type locality of this species (Fig. 1). The specimen 
was retained to determine egg size, incubation time and 
hatchling size under laboratory conditions, 

The specimen was placed in a small vivarium on a 
substrate of moistened Vermiculite and maintained at 
20°C. On 30.x.1982, two eggs were found buried 
approximately 2 cm beneath the surface. These were weigh- 
ed, measured, marked and placed in a sealed container 
in sterilized Vermiculite mixed with distilled water (100 
ml water/1SO ¢ Vermiculite)." The container was placed 
in a thermostatically controlled chamber, where the 
temperature range was 29° -30.5°C. The eggs were light- 
ly sprayed fortnightly to prevent desiccation and 
measurements of egg sizes and mass were made regularly 
(Table 1). Approximately one week prior to emergence the 


Fig. 1, Female Nephrurus deleani (SVL—78 mm, weight 
13.0 g) 2 days before paturation. 


one 


Fig. 2. Hatchling male Nephrurus deleani (SVL 36 mm, 
weight 1.9 g) with unhatched egg. 


eggs appeared desiccated and although daily spraying was 
carried out the appearance of the eggs remained 
unchanged. 

Hatchlings emerged on 25-26,xii.1982 after 55-56 days, 
incubation. Both specimens emerged from the egg imme- 
diately after completing a longitudinal slit in the egg shell. 
No part of the yolk sac was visible on either specimen. 

The hatchlings were much darker than the adult female 
in colour and possessed a pale vertebral stripe which ex- 
tended from the occiput to the tip of the tail (Fig. 2). The 
presence of this vertebral stripe, found only in juvenile 
N. deleani, previously led to some confusion between this 
species and N. vertebralis.’ Both hatchlings has a SVL 
of 4.5 mm less than the smallest SVL we have recorded 
from field collected specimens. 

Unlike many other species of reptiles, no significant 
changes were noted in the size of either of the eggs dur- 
ing the incubation period (Table |). 

We have collected gravid specimens of this gecko in 
January, April, May and October, which indicates that 
N. deleani may reproduce opportunistically, rather than 
seasonally, 

The Peter Rankin Trust Fund for Herpetology funded 
our field work. Michael Delean assisted in the recording 
of data and Winnie Feijen typed the manuscript. Terry 
Schwaner made constructive comments on the manuscript. 


TABLE L, Length (L), width (W) and mass (M) of eggs, hatching dates and snout-vent length (SVL), tail length (TL), 
sex and weight of hatchling Nephrurus deleani. L, W, SVL and TL in mm, M in g. 


Dates Measured 


24.x.1982 22.x1,1982 


Dates 
Hatched 
(Dec. 1982) 


20.xi1, 1982 


Egg No. L Ww M L Ww M 


rt 
bh 
aos 
a 
ee) 
a 
id 
a 
tu 
a 
LA 


222 


'Pianka, E. R. & Pianka, H. D. (1976). Copeia, 1976 (1), “Harvey, C. (1983). Trans. R. Soc. S. Aust. 107, 231-235. 
125-142. ‘Barnett, B. (1982). Vic. Herpetol. Soc. Newsl. 1, 1-10. 
°Gow, G. F. (1979). N-T. Nat. 1, 9-10. 


STEVEN DELEAN, 49 Davenport Terrace, Seaview Downs, S. Aust., 5049 and CHRIS HARVEY, 20 Crozier Ter- 
race, Oaklands Park, S. Aust., 5046. 


CORRELATION OF THE UPPERMOST LATE PRECAMBRIAN 
SUCCESSION ACROSS THE TORRENS HINGE ZONE IN THE PORT 
AUGUSTA REGION OF SOUTH AUSTRALIA: A DISCUSSION 


BY W. V. PREISS 


Summary 


In his recent paper, Plummer proposed new arguments in favour of resurrecting a correlation of the 
Simmens Quartzite and Corraberra Sandstone Members of the Tent Hill Formation (Wilpena 
Group) on the Stuart Shelf, with the Pound Subgroup of the Adelaide Geosyncline. This correlation, 
as well as previous ones proposed by the other authors quoted by Plummer, had been made at an 
early reconnaissance stage of mapping in both regions. Only those correlations published after 1965 
were based on adequate knowledge of the regional stratigraphies of the Stuart Shelf and Flinders 
Ranges, but even so, the details of facies relationships on the Stuart Shelf did not become available 
until the extensive drilling by mineral exploration companies in the late 1970s and early 1980s. 


BRIER COMMUNICATION 


223 


CORRELATION OF THE UPPERMOST LATE PRECAMBRIAN SUCCESSION ACROSS 
THE TORRENS HINGE ZONE IN THE PORT AUGUSTA REGLON OF SOUTH 
AUSTRALIA; A DISCUSSION 


In fis recent paper, Plummer! proposed new 
arguments in favour ob resurrecriny a varrelarion of the 
Simmens Quarizile and Corraberrs Sandstone Members 
of the Tene Hill betination (Wilpena Group) on the Stuart 
Shell, with the Pound Subgratp of the Adelaide 
Geosvncline, Uhis correlarion, as well as previons ones 
Proposed by Me onler authors quoted hy Plummer hau 
been mudesitan early reconnaissance stage oF mapping 
iW both regions. Quilty (hose correlauons published afer 
1965 were bused on adequate knowledge of (he regional 
sturiiigvaphies of the Stuart Shell and blinders Ranges, bul 
ever su, (he details of davies pelartonships on the Stuart 
Shelf did pio become available unril the extensive drill- 
ine bY mined exploration campanes en the fale (970s 
und curly LYROs. 

The crilicisor thot the pirevions earreluions were ull bas- 
ed on lithostrabwraphy is a tittle misplaced when 
Plummer’s own arguments are Oi (he absence of indepen- 
dent evidence such us biostlatizraphy, geochronology: or 
Palveomagoetism), alsa encrely tithastrareraphic. fe is 
therefore necessary 10 examine closely the eriteria amet 
asSuMpions Gn whiell his correlations are based. 


In the enurse of stinlying the regional sedimentolosy 
and palaeogeorraphy af the lower Wilpend Graup in the 
Adelaide Geosvilelnig, Plummer mapped a previousty 
unreenznised loeul crosional contact between the ARC 
Range Quartzite and the overtying Bunyeroe Formation, 
near Buckdiringa Gore in rhe southera Dfinders Ranges. 
Having visited this lowality, Fant happy to accepr bis map- 
ping (Plummer, Pig, 4) a5 aveurate The same map 
indicates. hal just north of the erosional channel cut in- 
tothe ABC Ranwe Ouurtzite, the (Wer Units interlonwue, 
fram this celanonstip alone if therefore seems 
Unreasonable (o useribe a tegiouul siguiicanve to the eno- 
sional break, and ro extend Wis discomlormily. westward 
on le the Stuart Shell 

The only evidence presented by Plunimer! for such an 
erosional break on the Stuart Shelf (the large lacuna int 
his Pig. 2) is based on a srratigraphi section uf the 
Tregoluna Shale, Corraberra Sandstone and Simoarens 
Quartvile Members published hy Thomson' This see- 
hon, based on onitrop, Widicates clearly in Upwatd bran 
sitidn trom shale to saidstone, avd inay be vonlipmed by 
Inspection of punwrous drilleares from the Stuart Shell 
The apparent “sharp boundary separating the nvo shaly 
units al the Tregolina Shale Member" potted oul by 
Plummer'!, os @ caplogrape peculiar. stave all 
ihological boundaries On (His diagram are Shown by a 
similar heavy fine, and it was never intended to tndicate 
a lithologie discontinuiny (BH. PO Theisen, pers. con, 
1984). No new evidence fram either odterop or drifleare 
lias been ofleredk 

The presence wl Bunyervo Pormauon in fhe Wilkatana 
LOU Bore ts irrelevant to the discussion since the bure 
Was NUL SUTTiciently deep fo intersect the BunyeronyA BE 


Range Quarivite contact The existence af a discon lor 
mily here fs therefore purely speculative. 

Plummer dues not state reasons. other than the 
posulated regional disconforminy, why the simplest in 
(erprelavon proposes! hy previous authors aad reeenlly 
stummarised by Preiss’ is incorreet. Chis is parueularty 
Hi icule to uftcterstanel ny rhe Wiipena Cirouj nt the Stuart 
Shell isan upward coarsening sequence (locally avertain 
bu fine-erairied unit, Me Vartoo Sliale}, essentially super 
lo the wpressive sequence he has deseribed> from the 
lower Wilpena Group of the Minders Ranges. The only 
senihounl itference seems to be [he replacemen| of the 
Moorillah and Bayley Range Siltskdne Members of the 
Braching Formation hy the cross-bedded, red Corraberra 
Sandstone on the Stuart Shell. However, med sandstones 
lucully Gectir at the base of the ABC Ranve Quartile, 
Possibly representing tongues of Corraberra Sutidstone, 
Moreover. no facies resembling the Wonoky Formation 
Nas been found on the Stuart Shelf, Plummer’s correla- 
fun table (Fig. 2) implies that the Wonoka Fornation 
passes laterally westwards fito Bunyeroo Furnanen, but 
there js no dievet evidenve of (his. Indeed, the Wonoka 
Formation isd laterdity persistent unit, recently iren weed 
it part as a galcureous flvsch facies’. The presence of 
{imestone iniractists jn dolraformanonal breccias sugpests 
thats shallow-water mareingl laciés were also Galgareous. 

Pluintier has nepleeted toaeeount for the Yarlou Shale. 
(owas lirstimapped by Johns overlying the Sinmmerns tor 
Arcoona) Quartyire aid discanlormably uverlain by the 
Cambrian Andamovuka Limestone on the northeasten 
Stuart Shelf, amd hes been correhued with the Bunyeroo 
Formation’. The Yarloo shale, andjlsgradalional pasvage 
down into fivies that are typical of he ABC Ranee Quart- 
ain may be abserved in the Arigea SCY WIA drillhole 
core Wlrilled qear Yarra Waria cliffarthe northern enct of 
Lake Torrens), This cacellens section penetfales the whole 
Snatigniphy froin Cambrian vedbeds down fo the Sturn 
Maal. Allintervening Adelaidean units can be identified 
conhidenniy with rhe excepliod of the Wonka Formation 
wou Pound Subgroup. Theseareabsent, either having been 
never depasited onthe Stuart Shell (they reflect an overall 
regressive Pllase of deposition, al least (othe top at the 
Bow mey Saudstone} er haying been eroded ineartivss Cam 
heen rine 

In contelision, although there is no proof ot absolute 
synchroneit of he Stuart Shell Wilpena Group with the 
lower Wilpena Group of the Flinders Ranges {up (oundin- 
cluding the lower part of the Bunyeroo Formathen) rhere 
are compelling reasons ip consider them as part of the 
sunic depositional sysiem, unrelaied to the deposition of 
the Wonoku hormution and Pound Subbraup, which 
resulted Hon reyeweu feclonie activity? und may have 
been confined to the Adelaide Geosynecling and parte at 
the Torrens tinge Zone, 

This discussion is publisheel Wi0h che persist at (he 
Direcor-General, Deparment uf Mines & Eucrey 


224 


‘Plummer, P. S. (1983). Trans. R. Soc. S. Aust. 107, 
171-175. 

?Plummer, P. S. (1978). Trans. R. Soc. S. Aust. 102, 25-38. 

3Thomson, B. P. (1965). Q. geol. Notes, geol. Surv, S. 
Aust. 13, 4-5. 

4Preiss, W. V. (Compiler), (1983). Adelaide Geosyncline 
and Stuart Shelf: Precambrian and Palaeozoic Geology 
(with special reference to the Adelaidean). Geological 
map, special series, 1:600 000. Geol. Surv. S. Aust. 


sJenkins, R. J. F. & Gostin, V. A. (1983). Abstracts, 
symposium ‘Adelaide Geosyncline sedimentary en- 
vironments and tectonic settings”, Geol. Soc. Aust. 10, 
39-44. 

°Johns, R. K. (1968). Bull. geol. Surv. S. Aust. 41. 


W. V. PREISS, South Australian Department of Mines & Energy, Box 151, Eastwood, S. Aust. 5063. 


CORRELATION OF THE UPPERMOST LATE PRECAMBRIAN 
SUCCESSION ACROSS THE TORRENS HINGE ZONE IN THE PORT 
AUGUSTA REGION OF SOUTH AUSTRALIA: A REPLY 


BY P. §. PLUMMER 


Summary 


The discussion by Preiss warrants comment on a number of points. Firstly, the accusation that the 
correlations by Plummer are purely lithostratigraphical is erroneous. By _ correlating 
palaeoenvironments into a palaeogeography, a chronological component is injected into the system. 
In sequences such as the Precambrian that are largely devoid of biostratigraphical, palaecomagnetic 
and direct age determinations, such correlations are the only hint available of time equivalence. 


BRILE COMMUNICATION 


225 


CORRELATION OF THE UPPERMOST LATE PRECAMBRIAN SUCCESSION ACROSS 
THE TORRENS HINGE ZONE IN THE PORT AUGUSTA REGION OF SOUTH 
AUSTRALIA: A REPLY 


The discussion by Preiss! warrants comment on a 
number of points, 

Firstly, the accusation that the correlations by 
Plummer are purely lithostratigraphical is erroneous. By 
correlating palaeoenvironments into a palacogeography, 
a chronological component is injected into the system. In 
sequences such as the Precambrian that are largely devoid 
of biostratigraphical, palacomagnetic and direct age 
determinations, such correlations are the only hint 
available of time equivalence. 

The presence of the hitherto unrecognised discontfor- 
mity at the ABC Range Quartvite/Bunyeroo Formation 
level is agreed to. However, Preiss reyerts to a logic of layer- 
cake stratigraphy in an endeavour to lessen the significance 
of this horizon, Such logic is nol applicable to the stable 
shelf/active basin setting under consideration. In such set- 
tings, shelf unconformities can frequently be time 
equivalent to thick basinal sedimentary packages. Such 
is the case here. 

The lateral persistence of the Wonoka Formation is 
ageed to, but only within the context of the geosyncline, 
The Wonoka Formation is a basinal flysch facies, and 
hence not expected in its same form on the adjacent stable 
shelf. Here again, however, Preiss applies a layer-cake logic 
for his expectation of just such an equivalent facies. 

Describing the sharp boundary separating the two shaly 
units of the Tregolana Shale Member as a “cartographic 
peculiarity”, is surely not intended to be taken as a serious 
scientific argument! 

‘Twice in his discussion Preiss refers to the lower Wilpena 
Group as representing a coarsening upward sequence, then 


draws an equivalence with the coarsening upward Tent Hill 
Formation on the Stuart Shelf. it should be pointed out, 
however, that the upper Wilpena Group is also a 
coarsening upward sequence. Both sequences are, in fact, 
regressive, but the lower Wilpena Group sequence is 
regressive to the point that erosion occurred within the 
basin. In such a case, any time equivalence correlation with 
a similar regressive Sequence on the adjacent stable shelf 
must be viewed with Scepticism, 

Finally, the only locally occurring sandy facies within 
the basal portion of the ABC Range Quartzite is a deep 
purple, heavy mineral rich, trough crossbedded, medium 
0 Coarse quartzite Which originated in deltaic distributary 
channels. These are not typical Corraberra Sandstone 
facies, and hence highly unlikely to be the tongues that 
Preiss suggests. 

In conclusion, it is agreed that “there is no proof of 
absolute synchroneity of the Stuart Shelf Wilpena Group” 
with the upper Wilpena Group in the Flinders Ranyes, 
but the chronological component injected into the system 
by palacvenvironmental correlations renders this syn- 
chroneity the most likely, 


'Preiss, W. V. (1984) Trans. R. Soc. S. Aust. 108, 233 
224, 

“Plummer, P. S. (1983) Trans. R. Soe, S. Aust. 107, 
171-175. 


P.S, PLUMMER, v/o Shell Petroleum Development, P.O. 
Box 2032, Dar es Salaam, Tanzania. 


EVIDENCE OF GASTRIC BROODING IN THE LEPTODACTYLID FROG 
RHEOBATRACHUS VITELLINUS 


BY KEITH R. MCDONALD & MICHAEL J. TYLER 


Summary 


The first record of gastric brooding in the Animal Kingdom was reported in the leptodactylid frog 
from Rheobatrachus silus Liem in 1974, and the first photographs of oral birth were published in 
1981. The description of R. vitellinus includes morphological evidence of a close phylogenetic 
relationship to R. silus. Here we report that R. vitellinus also broods its young in its stomach and 
gives birth through its mouth. 


BRIEF COMMUNICATION 


EVIDENCE OF GASTRIC BROODING IN THE AUSTRALIAN LEPTODACTYLID 
FROG RHEOBATRACHUS VITELLINUS 


The first record of gastric brooding in the Animal 
Kingdom was reported in the Australian leptodactylid frog 
from Rheobatrachus silus Liem in 1974', and the first 
photographs of oral birth were published in 1981°. The 
description of R. vitellinus’ includes morphological 
evidence of a close phylogenetic relationship to R. silus, 
Here we report that R. vitellinus also broods its young 
in its stomach and gives birth through its mouth. 

On 12.1.84 a single male and female R. vitellinus were 
collected at approximately 2130 hr in a creek bed of large 
boulders within complex notophyll vineforest in Eungella 
National Park, Queensland. The water temperature at the 
site was 19.4°C. The female was greatly distended, and 
during road transport to Mackay on 13.i1.84 she began to 
give birth to young. 

The first juvenile was born at 0700 hr and, during the 
next 27 minutes, 14 more were born individually or in twos 
or threes on nine occasions. These individuals were born 
underwater and it was noticed that the female opened her 
gape greater than 90°. The subsequent birth of babies was 
spaced out at less frequent intervals, with single indivi- 
duals born as follows: 0811 hr, between 0855 and 0915 hr, 
between 1210 and 1219 hr, between 1630 and 1635 hr and 
between 2245 hr on 13.1.84 and 0600 hr on 14.i.84. The 
mother was despatched by air to Adelaide, and a further 
juvenile was born in transit sometime before 1655 hr. 

At 1900 hr on 14.1.84 the mother was cooled and then 
packed in crushed ice prior to removal of the viscera for 
histological and biochemical studies. On removal of the 
entire alimentary canal a bulge was observed in the 


stomach, and upon compression a further baby frog was 
expressed. The juvenile was placed in warm water and 
recovered consciousness 30 min. later. 

The total number of young brooded by the mother was 
therefore 22, which is within the range for R. silus 
(18-25)*. The larger body size of R. vitellinus’ has not 
been accompanied by an increase in the number of young. 
However the snout-vent length of the young at birth 
(15.1-15.9 mm in two preserved specimens) is larger than 
the known range for R. silus 11.9-12.8 mm)’. 

Rheobatrachus silus gives birth to young at the surface 
of the water. We remain uncertain whether the underwater 
birth of R. vitellinus was a natural phenomenon or a con- 
sequence of the artificial conditions in which the female 
was constrained. 

The female on which these observations are based subse- 
quently was cleared and stained for bone and cartilage 
and is a paratype (South Australian Museum R 25447)’. 

Field assistance was provided by Veron Hansen and Guy 
Chester, and helpful advice offered by Margaret Davies 
and Jeff Miller. 


' Corben, C. J., Ingram, G. J. & Tyler, M. J. (1974). 
Science 186, 946-47, 

"Tyler, M. J. & Carter, D. B. (1981). Anim, Behav. 29, 
280-82. 

‘Mahony, M., Tyler, M. J. & Davies, M. (1984). Trans. R. 
Soc. S. Aust 108, 155-162. 

“Tyler, M. J.(1983). Oral birth and perinatal behaviour. 
Chap. 5, pp. 36-43. Jn M. J. Tyler (Ed.). The Gastric 
Brooding Frog. (Croom Helm: London & Canberra). 


KEITH R. McDONALD, Queensland National Parks & Wildlife Service, Pallarenda, Townsville, Qld, 4810, & 
MICHAEL J. TYLER, Department of Zoology, University of Adelaide. 


COMMENT: WARDANG ISLAND - A REFUGE FOR MARGINOPHORA 
VERTEBRALIS? 


BY A. P, BELPERIO AND C. V. MURRAY-WALLACE 


Summary 


Bone recently suggested that tests of Marginopora vertebralis found on the intertidal flat between 
Wardang Is. And Goose Is. are of recent origin, and that this species may be extant within the 
adjacent reef waters. This hypothesis was based on the fresh and uncemented appearance of 
individual specimens, on the MgCO; content of various samples, and on the apparent absence of 
outcropping (erosional) sources of older material. As part of ongoing studies into the stratigraphy 
and chronology of coastal Quaternary sediments of South Australia, we are examining the wider 
problem of reworking of Pleistocene bioclastic detritus into Holocene coastal sediments. Previous 
studies have demonstrated that reworking of older material is a significant process which must be 
considered, particularly where isotopic dating of the carbonate fraction of sediments is 
contemplated. One verified indicator of reworking is the presence of the bivalve Anadara trapezia in 
Holocene coastal sediments of S. Aust., and a similar conclusion has been assumed in the case of 
M. vertebralis. Our recent but unpublished data on the extent of amino acid racemisation in M. 
vertebralis found in Holocene strata from various areas of the State confirm that in each case, its 
presence also results from reworking of Pleistocene deposits. 


DRIED COMMUNICATION 


WIT 
“ 


</ 


COMMENT: WARDANG ISLAND—A REFUGE FOR MTARGINOPORA VERTEBRALIS? 


Bone recently siivgesred that testis al Mferginepore 
vertehrats ound on the intertidal flat between Wardany 
Is. ane Crouse Is, are of revent origin, and that this spe- 
cies may be extant: within the adjacent reef waters. This 
hypothesis was based on the fresh and uncemented 
appedranee OF individual speeimens, on the MgCO, con- 
tent of various samples, and on the apparent absence of 
OnTCrOpping (erosional) sources of older material, As part 
Gf ongoing studies Hilo The stratigraphy and chronology 
of coastal Qiiaternary sediments of South Australia’? 
We are examining the wider problem of reworking of 
Pleistacene bivelaste detritus into Holocene coastal sedi- 
ments, Previous studies" have demonstrated that re- 
working of alder material is a significant process which 
Must be considered, particularly where isotopic dating of 
the earhonate fraction. of sediments is contemplated. One 
verified indicator wt reworking is (he presence of the 
bivalve eliadare frapeste in Uplocene coastal sediments 
OS. Aust vind a similar conclusion has been assumed 
in (he cise OF AM verretralis’, Our recent bur unpub- 
lished daly on the extent of amino acid ravemisation in 
AT. vertehralis found in Holocene suala [rom various areas 
of (he Stan: contipn that in each case, ips presence alsa 
results Hom reworking of Meisrocene deposits. 

Tu test the validity of Bone’s hypothesis for Wardany 
Tyhind, we have exinined the Wardang Island reef site, 
am! have used radiocarbon!’ and amino avid 
dawemisation dating techniques on the samples collected. 

The results show that the reef connecting Wardany Is. 
and Coose ts rather than being modern', is of Pleisto- 
cone uve (Table 1), Phe siittace of tis reef is exposed al 
low (idecanad is tatled with a thin veneer (mostly <0 
cin of teeent inlertical sand. The ree? rock comprises 


poorly sorted and weakly wemerted bioelastic detritus, in- 
cluding numerous specimens of MZ vertebredis upto 8 mm 
in diameter Detritus from the crumbling seaward reef 
edge, including M. vertebralis, is swept over the reel sur- 
face and is the major souree of the sediment of the inter- 
tidal veneer, Thus the “modern” intertidal veneer is largely 
composed of reworked Pleistocene skeletal detritus 
(lithoskels) and The ave indicated by radiocarbon data 
(Table 1) is the “average age’ of the skeletal constituents 
which make up the sediment. Inicividual fragments of the 
clumbliig reef rock are visibly calureted. However evidence 
of cementation and caleretization enveloping individual 
grains (§ apparently reduced and eventually eliminated by 
continual abrasion in the intertidal environment. in par- 
Hieular specimens of M. verrebralis have secondary car 
honate infillings progressively removed with distance land- 
wards irway from the reef edge, lenee their “fresh” and 
“uncemented” appearance, 

Amine acid raicenisation measurements were also 
undertaken on M, vertebralts extracted from the reef rock 
(lible 2), Orher specimens of known age from northern 
Speneer Gull (Late Pleistocene)! and the Great Barrier 
Reef (Recent) provide a basis for calibration, The 
foraminifera extracted from the Wardany Is, reef rock are 
clearly of Late Pleistocene age (ea, 110-000 yrs by ana- 
logy with the northern Spencer Gulf sample’). Amino 
acid racemisanion measurements could not be undertaken 
onthe foraminifera loose on the tidal flat as bacterial and 
dlgal contamination penetrates the porous skeletal frame- 
work and vannot he satistierarily removed. 

We conclude that the specimens of M. verfebradis found 
on the udal lar between Wardang Is. and Goose ls. are 
devived fram erosion of underlying weakly cemented 


TAME 1, Rediocurbon meusurements on samples fran) vb urduniwe island, 


Fractien 


Satiple Dated 


HAQVRS7S Reet rock Bulk earbonate 


6329RS76. linlertidal 
sand 


Bulk sediment 


Radiocarbon Creologival 
Age Aue 

40 B00 * i yes! Late Pleistocene 
6 24) + 90 yrs > Halocene 


SSSSsSSSSSSSSSSSSSSSSSSSSSSSSSSSFSSSSSSsSsSsSh 


. a aPparen| aye’ resuliing from the mcarporution of seme modern earbor juo Lute Pleistocene (Glanville Foi.) 
sadinernh 


AN “avenue ave" resulunig [rom physical intermiviag of Late Pleistocene irhuskels with modern skeletal detritus, 


Tani 2. kvteet af wining aed racenielian in samples af Margivopera vertebralis ("feral Avdrolvsele" DEL ratios 
of alanine, proline wud aspartic acl). 


No. of Amino acid D/1. ratios 
Lowaliny analwses ALA PRO ASP Civoloyicul Age 
Cen Barrier Reel 3 0.04 0,07 (14 Holocene 
Ujiper Speneer Gulf | 0.25 0,29 0.37 Late 
Pleistocene 
Wardane Istaned x O25 0,27 0.32 


228 


Pleistocene marine sediments. Consequently it is unneces- 
sary, and is most likely incorrect, to invoke an hypothesis 
that M. vertebralis is extant in the adjacent waters. 
This paper is published with the permission of the 
Director-General of Mines and Energy. 


'Bone, Y. (1984). Trans R. Soc. S. Aust. 108, 127-128. 

Belperio, A. P., Hails, J. R. & Gostin, V. A. (1983). 
James Cook University Geog. Dept Monograph Series 
3, 37-47. 

’Belperio, A. P., Smith, B. W., Polach, H. A., Nittrouer, 
C. A., DeMaster, D. J., Prescott, J. R., Hails, J. R. & 
Gostin, V. A. (1984). Marine Geology 61, in press. 


Cy 


‘Belperio, A. P., Hails, J. R., Gostin, V. A. & Polach, 
H. A. (1984). Marine Geology 61, in press. 

SMurray-Wallace, C. V. (1983). B. A. (Hons) thesis, Univ. 
Adelaide (unpublished). 

‘Burne, R. V. (1982). BMR J. Aust. Geol. Geophys., 7, 
35-45. 

"Hails, J. R., Belperio, A. P. & Gostin, V. A. (1983). 
James Cook University Geog. Dept Monograph Series 
3, 48-53. 

8Gostin, V. A., Hails, J. R. & Polach, H. A. (1981). 
Search 12, 43-45. 

°Schroeder, R. A. & Bada, J. L. (1976). Earth-Sci. Rev. 
12, 347-391. 


A. P. BELPERIO, Department of Mines and Energy, P.O. Box 151, Eastwood, S. Aust. 5063. 
~ MURRAY-WALLACE, Department of Geology, University of Adelaide, G.P.O. Box 498, Adelaide, S. Aust. 5001. 


ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED 


Patron: 


HIS EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA 
LIEUTENANT-GENERAL SIR DONALD DUNSTAN, K.B.E., C.B. 


OFFICERS FOR 1984-85 


President: 
J. 5. WOMERSLEY, B.Sc. 


Vice- Presidents: 


D. W. P. CORBETT, B.Sc., Ph.D. M. J. TYLER, M.Sc. 
Secretary: Treasurer: 
P. M. TAYLOR, B.Sc. R. H. FISHER, E.D., A.U.A. 
Editor: Assistant Editor: 
M. DAVIES, M.Sc. K. H. NORTHCOTE, 
: B.Ag. Sc., D.Sc. 
Librarian: Programme Secrétary: 
N.-P. McKELLAR-STEWART, P. HORTON, B.Sc. 
B.Sc., Dip.Lib. 
Minute Secretary: Membership Secretary: 
R. J. SHIEL, B.Sc., Ph.D., T.ST.C. P. M. THOMAS, M.Sc. 
Members of Council: 
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D. C. LEE, M.Sc. C. B. WELLS, E.D., M.Ag.Se. 


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