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VOL. 105, PARTS i & 2 
12 JUNE, 1981 

Transactions of the 

Royal Society of South 



McQuillan, P. B. A review of the Australian moth genus Thalaina (Lepidoptera: 
Geometridae: Ennominae) - - 

Hutchings, P., De Deckker, P. & Geddes, M. C. A new species of Manayunkia 
(Polychaeta) from ephemeral lakes near the Coorong, South 
Australia --- 25 

Butler, A. J. & Keough, M. J. Distribution of Pinna bicolor Gmelin (Mollusca: 

Bivalvia) in South Australia, with observations on recruitment 29 

Pledge, N. S. The Giant Rat-Kangaroo Propleopus oscillans (De Vis), (Poto- 

roidae: Marsupialia) in South Australia ----- 41 

Crook, G, A. & Tyler, M. J. Structure and function of the tibial gland of the 

Australian frog Limnodynastes dumerili Peters ... 49 

De Deckker, P. & McKenzie, K. G. Bennelongia, a new cyprididid ostracod 

genus from Australasia 53 

Bye, J. A. T. Exchange processes for upper Spencer Gulf, South Australia - 59 

Jenkins, R. J. F., Plummer, P. S. & Moriarry, K. C. Late Precambrian pseudo- 
fossils from the Flinders Ranges, South Australia 67 

Womersley, H. B. S- & Sinkora, D. Sonderophycus and the type specimen of 

Peyssonnelia australis (Cryptonemiales, Rhodophyta) 85 

Brief communication: 

Tyler, M. J. & Da vies, M. Partial acquisition of pigmentation in an adult, albino, 

Australian Leptodactylid frog (Limnodynastes dumerili Peters) 89 




CONTENTS, VOL. 105, 1981 

PARTS 1 & 2, 12 JUNE 

McQuillan, P. B. A review of the Australian moth genus Thalaina (Lepidoptera: 

Geometridae: Ennominae) ------- 1 

Hutchings, P., De Deckker, P. & Geddes, M. C. A new species of Manayunkia 

(Polychaeta) from ephemeral lakes near the Coorong, South 
Australia ----------25 

Butler, A. J. & Keough, M. J. Distribution of Pinna bicolor Gmelin (Mollusca: 

Bivalvia) in South Australia, with observations on recruitment 29 

Pledge, N. S. The Giant Rat-Kangaroo Propleopus oscillans (De Vis), (Poto- 

roidae: Marsupialia) in South Australia - - - - 41 

Crook, G. A. & Tyler, M. J. Structure and function of the tibial gland of the 

Australian frog Limnodynastes dumerili Peters 49 

De Deckker, P. & McKenzie, K. G. Bennelongia, a new cyprididid ostracod 

genus from Australasia _-__-. .53 

Bye, J. A. T. Exchange processes for upper Spencer Gulf, South Australia - 59 

Jenkins, R. J. F., Plummer, P. S. & Moriarty, K. C. Late Precambrian pseudo- 
fossils from the Flinders Ranges, South Australia 67 

Womersley, H. B. S. & Sinkora, D. Sonderophycus and the type specimen of 

Peyssonnelia australis (Cryptonemiales, Rhodophyta) - - 85 

Brief communication: 

Tyler, M. J. & Davies, M. Partial acquisition of pigmentation in an adult, albino, 

Australian Leptodactylid frog (Limnodynastes dumerili Peters) 89 


De Deckker, P. Taxonomy and ecological notes of some ostracods from Aus- 
tralian inland waters - - - - - - - - 91 

Beveridge, I. Three new species of Calost auras (Cestoda: Davaineidae) from 

the New Guinea wallaby Dorcopsis veterum - - - - 139 

Tyler, M. J., Davies, M. & Martin, A. A. Frog fauna of the Northern Territory: 

new distributional records and the description of a new species 149 

Baker, A. N. & Devaney, D. M, New records of Ophiuroidea (Echinodermata) 
from southern Australia, including new species of Ophiacantha 
and Ophionereis - - - - - - - - 155 

Jenkins, R. J. F. The concept of an 'Ediacaran Period' and its stratigraphic sig- 
nificance in Australia -------- 179 

Foster, C. B. & Harris, W. K. Azolla capricornica sp. nov. First Tertiary record 

of Azolla Lamarck (Salviniaceae) in Australia - - - 195 

Bull, C. M. & King, D. R. A parapatric boundary between two species of reptile 

ticks in the Albany area, Western Australia - 205 

Brief communications: 

Von der Borch, C. C. Recent non-marine dolomite from the coastal plain, south- 
eastern South Australia ------- 209 

Kailola, P. J. & Jones, G. K. First record of Promicrops lanceolatus (Bloch) 

(Pisces: Serranidae) in South Australian waters - - 211 

Lange, R. T. & Reynolds, T. Halo-effects in native vegetation - - - - 213 

Smith, J. & Schwaner, T. D. Notes on reproduction by captive Amphiboliirus 

nullarbor (Sauria: Agamidae) - - - - - -215 

Smith, M. J. & Rogers, P. A. W. Skulls of Bettongia lesueur (Mammalia: Macro- 

podidae) from a cave in the Flinders Ranges, South Australia - 217 

Zeidler, W. A giant deep-sea squid, Taningia sp., from South Australian 

waters - - - - - - - - - -218 

Addendum - - - - - - - - - - - - -218 

Insert to Transactions of the Royal Society of South Australia, Vol. 105, Parts 3 & 4, 11 December, 1981 


by P. B. McQuillan 


The moth genera Thalaina Walker, Thalainodes Lower and Macqueenia Turner are reviewed. New 
evidence has resulted in an expansion of the concept of Thalaina to include the other genera in its 
synonymy. Nine species are reviewed and T. kimba sp. nov., is described from the mallee areas of 
South Australia and New South Wales. Immature stages, foodplants, flight activity and distribution 
of the various species are recorded where known. The genus has adapted successfully to most major 
habitats in the southern half of Australia wherever their food plants (Acacia and Cassia) are 


In P. B, McQuillan* 

McQuuiAN. P. B. (1981) A review ol ihc Australian moth genus Thalaiua (Lepidoptera: 
Geornctridac: Ennotmnuef. Tn\ns R. Sol. S. Autt,, 105(1), 1-23, 12 June. Hffit. 
The moth genera Thalaiua Walker, Thahiinodes Lower and Macitat-* aia Turner are 
reviewed New evidence has resulted in an expansion of the concept of Thalaina to include 
the other genera in its synonymy. Nine species ate reviewed and 7*. kimba sp. now, is described 
from the malice areas of South Australia and New South Wales. Immature stages, foodplants, 
flight activity and distribution of the various species are iccorded where known. The genus has 
adapted successfully lo most major habitats in the southern hall of Australia wherever their 
food plants (Acacia and tfotfftO M* established. 


This is the firs! of a proposed scries of taxo- 
nomic studies on the ennornine gcometrids of 
southern Australia. It reviews the taxononuc 
status arid known biology of a distinctive 
group of autumn-Hying moths previously re- 
I erred to Thalaina Walker, Tfntlairiot1e\ Lower 
and Afacqncftda Turner 

All the gpflCiCfi arc large, sirtkuiely patterned, 
nociurnal moths with distinctive larvae N«H 
surprisingly, all four species found in coastal 
eastern Australia Were described by 1865. Re- 
siding at Broken Hill O. B. Lower discovered 
and described three inland species between 
1900 and 1502, and his series of two of 'hem 
constitute most of the specimens available lor 
study, An apparently localised species was dis- 
covered in the 1930s at Millerran, southern 
Queensland hy J. Mncquccn and is slill very 
pooily known. Jn the 1960s and early 1970's 
a few specimens of a widespread ecnttal 
Australian species were collected and subse- 
quently described in 1972. Similarly, concen 
(ration of collecting in the semi-arid areas of 
soul hem Australia during late autumn by 
CSIRO collectors over the last ten yeans has 
made available a good series of another new 
species described here. 


The following abbreviations arc used for 
collections: AM = Australian Museum. 
Svdney; ANJC ■= Australian National Insect 
Collection, CSIRO, Canberra; BMNH - 
British Museum (Nalural History). London; 
NMV National Museum of Vicloria, 

* Division nf Entomology. Dcpl of Agriculture 
laboratories, St John's Ave, New Town, Tas. 


Melbourne; PBMcQC P B. McQuillan eoL 
lection, Adelaide; QM — Queensland Museum, 
Brisbane; SAM South Australian Museum, 
Adelaide; TDA Tasmania n Department oi' 
Agriculture, I lobar*; TMAG Tavmanian 

Museum and Art Gallery. Hobart; VQ 
University of Qucenslaud. Brisbane; WAM 
Western Australian Museum, Perth. 

Abbreviations ol the names of collectors 
are PA P Aitken, FMA - F M. Angel, 
KA K Ashby. WBB W. B. Barnard 

RB R. Beresfoid, TJi T. Blackburn. 

TGC I. G. Campbell, DHC - D. H. 

Col less, tPBC I- R B. Common, JRC - 
J. R Cunningham, IWD J, W ( Davies, 

CJD L. J Dumiean, EDE - E. D. Inwards, 
BE B. Evans, RHF = K. H. Fisher.. CWF 
C. W. } -nizier. NG - N. Geary, GCLG 
G. C. L. Goodine, Gtttt G, H. Haidv. 
RJH R J. Hardy. JH - J, Har&lcll. CCI 

C C Ives, Wk W. Klecxaj LEK 
1.. E. Koch. LRK - L. R. fslurtze. AML - 
A M. Lea. RL — R, Lewis, OBL - O. B. 
lower, Gl. « G. LvelJ, NMcF N. McFar- 
land, KJMcK K, J. McKic. JM J. 

Vlacqueen. PBMcQ P. B. McQuillan, 

WLM W. L. May, VHM = V. U Mmcfun. 
BM B. Mollisou, GBM G. B, Montcith 
K.M ), G. Morris, TN T Newberry, 
UN I . J. Newman, KRN K. R. Norris. 
HP H. Pel/. RJP «= R J. PricsL PR 
P. Ranford, LMR - L. Mossc-Robinson. VJR 
V, J. Robinson, ALR A, L. Rogers, RS 
R. Straatman, JJHS - J. J. H, Szeut-lvany. 
JGOT - J. G. O. Temper. HST =*= If- 5. 
Thirkell, LBT - L, B. Thorn, NBT N. B. 
Tindule. H LI H Udell. MSU M. S 

Upton, MMHW — M. M. H. Wallace, RW 
- R, Went, JOW " J O. Wilson. RGW 
R. G. Winks. FWJ - F. Wood-Jones. 

p. b. Mcquillan 

Numbers prefixed by "G" accompanying 
the descriptions of immature stages relate to 
code numbers of specimens in the McFarland 
larval collection housed at the S.A. Museum, 
Adelaide; sec McFarland (1979) for lull 

Genus THALAINA Walker 

Thalaina Walker 1855, p. 659; Type species 
Thalaina klenaea Walker (—Thalaina stivnacu 
(Doublcday)), by subsequent designation bv 
Fletcher 1979. p. 202. 

Ahsyrtes Guenee, 1857, p. 226; Type species 
Absyrtcs magnificaria Guenee, by subsequent 
designation by Fletcher 1979, p. 1. 

Thalainodes Lower 1902, p. 231; Type species 
Arnelora tetraclada Lower, by subsequent 
designation by Turner 1919 p. 386; syn. now 

Macquecnia Turner 1947, p. 101; Type specie^ 
Murqueenia chionoptila Turner, by monotypy; 
syn. nov. 

Adult: Medium sized, moderately robust, noc- 
turnal moths; wing expanse 34-54 mm, Colour 

pattern basically white with or without darker 
geometrical markings. 

Head (Figs 1, 2) with vestiture of crown 
thick, hair-like slightly roughened; Irons well 
rounded (fairly convex), clothed in scales, or 
nearly naked and bearing series of shallow 
transverse ridges beneath large sclerotizcd 
median projection; labial palp short, slightly 
porrect, second segment with projecting scales 
below, terminal segment very short; haus- 
tellum fully developed and functional; antenna 
in male laminate or strongly bipectinate. in 
female filiform; compound eye large, globular, 
without hairs; ocelli absent; chaetoscmata 

Thorax clothed with mixed hair-like and 
spatulate vestiture; forewing triangular, apex 
often subfalcate, in male without fovea, pat- 
tern usually white with ochreous geometrical 
pattern or rarely suffused darker, Sc and R, 
anastomosed, R., anastomosed with lU-i_j to 

Figs 1-2. Heads of Thalaina spp. 1. T. tetraclada; 2. T. macfarlandi. 

Fig. 3. Fore and hind leg of Thalaina sp. E — length of epiphysis; F— length of femur; ISB — distance 

between spur bases; T— length of tibia; TE — distance from top of tibia to top of epiphysis; TSB — 

distance from top of tibia to top of spur base. 


Table 1. Comparison oj some mam ratios ftom 

leg measurement of Thafaina specie* ivt^ fig. J 

for abbreviations). 

-pre lt£ 

Kind les 








wli'ttaw ,{ 








QrtytAoMH j 








elatu J 








Lnscrxpta ( f 








atittt'hroti J 








paronychc ■.■ 







kimhn f J> 








Cflitftlt-I'lltil r$ 








trtwtfiJti g 








ttun lurUunli ,■* 








farm an areoic, R 3 stalked with R 3 , p tVfy 
often arising nearer to M-j than to M : , ; hind- 
wing white, sometimes with darker markings, 
cell rather long; femora smooth-scaled, pos- 
terior tibiae in male slightly dilated, fore tibiae 
with epiphysis, sometimes with apical hook, 
mid and hind tibiae with spurs well developed 
(tot leg measurements, sec Table 1 ). 
Abdomen with eomb on segment 3 in male 
weakly to moderately developed. 
AfdU- ^em/alia with uncus simple, slender 
apically acute; soeii well developed, gnaihos 
simple* slender, with small median recurved 
spine or group of" small spines at apex; juxta 
broadly rectangular or shield -shaped; i'urca 
usually long, biiid; cristate hairs well de- 
veloped; valva elongate, slightly spatulate, 
aedeagus tubular, slightly curved with pointed 
apex and oblique distal orifice, cornuli usually 
a scries of stout, often compound spines. 
ietnnte xenttah'a with papillae anales densely 
hairy, bases Of hairs sel in conical projections; 
apophyses postcriorcs about 1.4 times as long 
as apophyses anterioves, sterigma broadly cup- 
shaped, ductus bursae long, with parallel sides. 
weakly sclcroti/ed, in dtameter about [ that 
of corpus bursae. its posterior part usually with 
ring of longitudinal scleroli/cd slriations; 
corpus bursae ovoid, without signa. 
£(,'(? broadly ovate, with or without darker 

f'n\t fnxtar lar\(\ wilh head capsule brown. 
without pattern; body pale yellow; setae ex- 
tremely long and quite stout; setae XD and D 
on prothorax on a lightly stiorobzed plate; an- 
terior humps on prothorax lacking. Abbre- 
viated prologs present on A4 and A5; 
crotchets arranged in complete circle on A4 
;-nci A5 (numbering about 121 and in half 
circles on A6 and clampers. 

f'inuf instar larva wilh head capsule green to 
yellow-green with numerous small brown 
blotches across- vertex; labrum deeply emar- 
ginate; body fairly robust, pale green to dull 
green with at least a whitish lateral stripe; 
cuticle smooth; setae short and fine; prothorax 
somewhat swollen and produced into two an- 
terior dorsolateral fleshy projections; ab- 
breviated prologs present on A4 and A5; 
crotchets in 3M circle on A4, in 5/6 circle on 
A5 and 1 circle on A6 and claspers, unior- 
dmal on A4-A6 but tending to be. hiordinal 
on claspers of some species: numbers of 
crotchets range: 13-21 on A4, 14-29 on A5. 
25-37 on AG and 28-4S on claspers; length 
23^*1 mm, width 4.5-5.5 mm; head capsule 
width 2.3—3.5 mm. The larvae are foliage 
mimics, being basically green in colouration 
with pale lateral and ventral stripes. Those 
species feeding on bipinnate Acacia usually 
have pale dorsal stripes and more mottled 

Papa dark brown to orange-brown, smooth, 
anterior margin of AID dorsally with 2-4 
widely rounded teeth: cremasler hooks 2 or 
6-8; length 11-17 mm width 5-6 mm. Aesti 
votes in loose cocoon incorporating detritus, 
beneath litter or in the soil. 
Flight Period: Mostly late summet to late 
autumn; flight periods for individual species 
range from less than four to more than twelve 
weeks. Figure 18 shows flight periods for 
species represented in collections by ovet 20 
specimens with full data. 

Distribution: Australia, south of about latitude 
24" 5 see figs 49-51. 

Key tu adults: 

1. Fron* more nr less naked, wilh seterotized 
median projection . 2 

— f ; rons covered with hair-scales, without 
median projection .... . ,., 6 

2. Fort* wing sntiny-whitc with sharply defined 
orange-brown or ochreous* hrown markings 3 

— Forewing mostly pale ochreouvgrey or 
grey ,....,, 5 

3. Forewing wilh stripe along middle one-third 
of cosla ...... A 

— Forewing without stripe along middle one- 
third of costa . .. - .. maefarlandi (Wilson) 

4. Forewing with oblique stripe from near mid 
termen to coital streak at three-quarters 

paronycfui (Lower) 

— Forcwmg without stripe connecting termen 
and costa as above , . ,. telrticfatfa (Lower) 

5. Forewing completely ochreous-grev, minutely 
speckled with black aUochrou (Lower) 

p. b. Mcquillan 

— Porewing while but heavily marked with 
broad. sulTused grey markings kimba sp. n. 

6. Forewillg without markings in disc 7 

— Farswing with markings in disc .. 8 

7, Hindwing with a large black spot near tornus 

....... chionopiilu (Turner) 

— Hindvving without a large black Spot near 
tornus selcnavti (Doubleday) 

(form punetUineu Walker) 

K Horewing with short, ochreoiib brown streaks 

on M|. M-, CuAi and CuA^ near termen 

insert pta Walker 

— Forewiijg without markings at extremities of 
Mi. Usi CuAi and CuAq _ 9 

*J. Porewing with a relatively straight stripe 
ttOTfl tornus to near mid costa or a Jittte 

below mid costa ...._ _._,_,,.,. JO 

Fore wing with stripe from tornus not 

reaching mid costa . angutosu Walker 

10. Forewing with diagonal stripe from mid 

termen to three-quarters of costa 

clara Walker 

— Forewing without stripe from mid termen 
lo costa as above selenacu (Douhleday) 

ThuUufiu selenaea (Doubleday) 
FIGS 4, 14, 18, 19 ; 29, 39, 49. 52. 53. 
CaUtmorphu wlenuea Doubledav 1845, p. 437. 
pi. 5, Rg-. 3, 

Thalaina sek-tHwa, Meyrick 1892, p. 653; Turner 

1919. p. 387. 
Thahthht kletwen Walker 1 855, p. MSO. 
Pompeja ate.tralittria Herrich-Schad'er 1855. pi. <»0. 

fig. 333, 
Ahwrtcs attsimlUiria H.Sch. (magnificuria Gn.) htitftiutru [hierry-Mteg ihs>9, p, 21. 

/tbxyrti'x magniflcaria Gucnee 1857, p, 226. 
I'lidlfu-na mdgnifica Desmarcst 1 858 fyn. mn\ 
(junior objective synonym of mugmfworiu Gn.). 
Tlmlaina punctilinea Walker 1865, p. 228. 

Type o( selenaea: not found in BMNH 
(D, S. Fletcher, pers. coram.) or any Aus- 
tralian collections. 

Types of Meiuuu; Uetotype 2 labelled l 'Cu1ii~ 
ntorphu selenaea H. T. [H abort Town) Aus- 
tralia 44-105" in BMNH, hereby designated: 
1 .; pauila totype labelled "46-46" in BMNH. 
hereby designated. 

Type of uttxfraliuria: not found in Zoological 
Institute, Martin Luther University, Witten- 
berg, GDR (N. Grosser, pers. comni. ). 
HolOtype a of autraliaria var. fortttnata 
labelled TypiCUITI specimen : Ex Musaco 
Aeh. Guencc : Magn. var A Gn. : Ex Ober- 
thur Coll. Brit. Mus. 1927-3" in BMNH. 
Type of imgntficarta. this is the earliest 
Scientific name and formal description ot text- 
fig. 17 p. 5 of Lucas (1857). colloquially 

named by him as "Phalene magnitique" 
according to Guenee (1S57) 4 3 » 1 9 were 
in his collection at the time of description; 
none of these syntypes has been examined. 
Type of magnified: Desmarest applied this 
name to Lucas' unnamed figure; the specimen 


1A + 2A 

Figs 4-5. Venation of Thalainct spp. 4. T. sele- 
naea'., 5. T. chionoptita. 


on which the figure is based bus not been 


Tvpc of pant Hi nea. fectolxpc 6 labelled 
"I i.miuiii 5,S-6U" in BMNH, hereby desig- 
nated; I $ pumU\a>t\pe labelled "V.D.I., 
WHS. |cr WW.SI 43-58" in BMNH, hereby 


Adult (Jig.s 52. 53); Head With lions rounded, 
smoothly hair-scaled, whiiish with fuscous 
band helow vertex; vertex, with ercet hffjr- 
seales, bright red-brown; labial palp wiih 
terminal segmeni and apex o\' second segment 
ImeouN remainder white; antenna of male 
laminate. Thorax aK-vc greyish white on an- 
tcrior margftf, remainder white, base of wines 
orange beneath white; legs white with exterior 
of fore and mid lemur and libia and all tt|T$l 
iiifuseated; forewing (fig. 4) with eosla nearly 
straight, apex pointed, termen rounded and 
somewhat sinuate beneath apex, K t anasto- 
mosed with Se, tLj often anastomosed with 
R., Lp form an ateolc. ground colour above 
shilling while, red-brown streak along eosta 
from base Ua one-haif thence angled inwards 
as a narrowly black-margined streak to lomus: 
similar streak from tonuis along inner margin 
lo base, termen narrowly red-brown* cilia 
red-brown posteriorly fuscous; hindwings shin- 
ing while, moderate fuscous suhapical blotch, 
cilia while; forewjngs' beneath shining while 
infuscatcd near upe\ and with costal streak 
fuscous; hindwings beneath shining while, 
subapical bloich much enlarged, sharply red- 
btown above M, fuscous below; wine 

■x-54 mm, g 52-56 mm 
A comui'Mi variety of this species (fig. 33] 
has a educed costal streak, tacks the diagonal 
fotewing streak and has the streak along ihe 
inner margin poorly developed; the suhapleai 
huidwing blolch may be reduced or even 
flhscrfl but is fully developed beneath 
\t<th' : .ynitalia ilig. L9J with apex of gnaihos 
With small spine, furca with two prongs oS 
equal leoetli; acdeagns < fig. JSft) stout, eornuti 
of two compound spines subeqnul in length. 

Female g&tintfta ("tig. S9.J wilh corpus bursae 

Final insnn tana: Head capsule 3.3 3.5 nun 
wide, pale green with small EuspttuS blotches 
iiiiov, vertex; body green with fine fuscous 
speck I i mi «m anal pi; v claspers, fleshy 
"whitish lateral stripe usually present, 'wo whiir 
sulweiitral stripes present as blotches adjacent 
10 'horacic legs and continuous from nicta- 

IhoruA 10 A9. incomplete White mid ventral 
stripe on A I to A5; crochets 19-21 on A4, 
22-29 on A5, 31 37 on A6 and $5-4$ on 
anal clampers, crochets on anal claspers unior- 
dinal. length 38 — 4 1 mm, width 5.5-6 mm. 
Material: GI19, 

Pupa (fig. 14) mid to dark brown in colour; 
cremastcr hooks 6-.S: dorsal anterior margin 
of A 10 with 4 lobes. Material: CI 19. 

!■><-■<! plant \ Aratia aielaao.xyion R. Br.. A 
reiinndes Schiechl. 

Specimen* t-unninetr 8? <J 45 SL QUE! NSI AND: 
Brisbane, v. ROW I V UO; I amineton Nil Park* 
it., v, WK 2 $ l.'Q: Stanthotpe. iv. I rf I U- 
foowoomba. iii-. iv. HID 3 o UCfc Wybcrha, 
BfOWII Mountain, i. DHC 1 J AMC; « km N of 
Runewahl, iii. |FBC & MSU I rf ANIC". C hureb 
Point, v. II RC i $ ANIC: Clvde Mm 800 m. iii. 
IFBC & MSU I 3 AMC; 8 Km W of Coirs 
Harbour, v. MSU 1 o* ANIC" Depol Beach, iv. 
IPBC I g ANIC; Kangaroo Valley, iii. DIU I 
J ANTC: Kaloomba. iii. I $ SAM; Narara. iv. 
IMR | r - ANIC 5 Ion SSP of Poit Maccmurie 
v MSU I r? ANIC: 2-7 Ui Nl of Oueanbt\san 
fi70 m, iv. fFBC I J ANIC 40 km S of -Sio.e'c- 
ton. iii. IKK I \S ANIC; Tooloom Seruh, iii. I 
,v UQ; « km S of WaucSonc. |IT, II UC ^ MSU I 
,( I ' ANIC. Wolloncone, iii. V.IR 1 rf ANIC. 
NK nf lee's Spring POO m. i II RC I .; I y 
ANIC VICTORIA: Boronia. iv. I d NMV; 
Castlcmaine, iii. I ? S \m S of Gellibrand 2M\ m. 
ii. IFBC I 2 ANIC; Gishorne, iit., iv. 4 £ 3 ? 
AMC, iv f>8 specimens NMV. i-. iii. iv GL 
1 4 fj A 9 SAM; Mcciiian iv. I ? NMV: M.I- 
,_!> am. iv. ]^ 1 4S NMV; Moe. iv. I ? NMV, 
iii GCI tt I 5 SAM: Mordiiilloc, v. CCt I ff 
ANIC: Ml. bifflOtill RUh B- IFBC 1 ^ ANIC: 
SptiflfflValC v. I 5 VO: Turtou'v Pass. iii. NBT 

5 ■ I s sam wiindfn, iv. i 9 nmv Tas- 
mania: Ctmdoniinion Creek, ii. 1 S 2 9 NMV 
Cr.urofi Cios-iirip, if, l V nmv- ftevciapnrt, I £ 
IDA Frcveincl Nil Park, ii. IFBC & MSU I o* 
ANIC Kdso. I # SAM; Kmaston, iii. iv. IRC 2 
cf A"NIC, W. .IRC 2 ? NMV; Lafefl I.euke $£t\ m. 
ii. II-BC & VISU 1 ^ ANIC; I .mmcL-ston. iv. I n 
SAM: In km W of Maydena. ii. 1 i 4 5 NMV; Mt 
Nelson 200 m, iU i( '- P&McQ 4 o TDA: Ml Wd- 
tlYlfiWffi 2>t0 m, i- i'.. iii. BM 7 o 4 9 A1SIC; Ouse, 
ii.. iii. iv 3 3 I ? '^A- Parenna. King tsi.mtl. ii. 
PUMeO 1 $ I'DA; C,)ntrenstown. v. 1 V NMV: 
Ritlge^uv, iv. IRC I ^ 2 y NMV: Rosehei ry ; \t, 
1 V ANIC; Saudford. VVLM I rf ANIC, 5 km V 
of Waratah bfiP rrt. ii CP&t & MSU 2 rf t ? 
ANIC: 13 km SW of Waratah 600 ni. ii IPBC 

6 MSU 2 ti I V ANIC : /eeban. r. CHH 1 i 
r-Hrner. V. 3 J SAM; Naiueoorte, I V SAM. 
YorXe Peninsula, iii.. iv. .IGOT l d 1 ? SAM. 

p. b. Mcquillan 

Figs 6-8. Venation of Thalaina spp. 6. 7\ angulosa\ ?• T. clara\ 8. I. insrrtpta. 

Diurihurkvi: see fig. 49. Flight Period: see 
f?g. IS, 

Comments: T, sehnaca is a dimorphic species 
= one form (var. punctiiinea Walker) without 
markings on the forewing and the other with 
u diagonal bar across the forewing; occa- 
sionally a streak along M s may also be present 
(lig. 52). It occurs in habitats in south eastern 
Australia from open woodland to closed forest 
in areas or greaier lhan 75 cm annual rain- 
fall and at elevations of 0-800 m. Preferred 
hosts arc large phyllodinous wattles such as 
Acacia rctinodes and A. mefanoxylon. An in- 
teresting dwarf population occurs in remnant 
native forest near Parenna on King Island. 
This species is the earliest of the genus to 
appear on the Wing, flying from mid January 
\o April. 

rhahtitta an^ulosa Walker 

FIGS 6, 13, 15, IS, 20, V), 40, 50, 54. 

Thalaina ansttlnsa Walker 1865, p. 2*9; Meyrick 

1892, p. fifi5; Turner 1919, p. 388. 
Holotype 9 without abdomen, labelled 
"S. Aust. 61-104" (F. Waterhouse) in 

Adidt (flfc. 54); Head with Irons rounded, 
smoothly hair scaled, whitish-bufT; vertex with 
rough hair-scales, orange: labial palpi with 
termini! segment and apex of second segment 
fit-.cous. remainder of second segment white: 
antenna o( male shortly bipeclinatc, Thorax 
above greyish-white becoming whiter pos- 
teriorly and on legulae. orange at base of 

wings, white beneath; legs with fore and mid 
femora and all tarsi mfusealed, hind femora 
and tibiae whitish; forewing (fig. 6) with 
cosla nearly straight, apes produced, termen 
strongly arched and sinuate below apex, R 3 
anastomosed with Sc, ground colour above 
shining white, a narrowly black-margined rust 
coloured streak along costa from base to one- 
eighth costa then extending to mid-discal area 
where il divides into an upper arm reaching 
mid termen thence to four-fifths costa and a 
lower arm extending to tornus thence along 
Ihc posterior margin to near base, termen 
narrowly rust coloured, cilia rust posteriorly 
fuscous; hindwing shining white, a large fus- 
cous suhterminal blotch extending from above 
M ; > to near apex with a narrower extension 
to anal angle, cilia fuscous on termen from 
anal angle lo live-sixths with remainder white; 
forewing beneath shining white with upper 
markings visible, a subapical blotch extending 
from four-fifths costa to mid termen, orafltge- 
rust near costa, remainder fuscous becoming 
paler apically; hindwing beneath shining white, 
subtcrmmal blotch as above but less extensive 
and sharply orange-rust above M-,; wing ex- 
panse .* 40-48 mm, g 42-54 mm. Abdomen 

Male genitalia I fig. 20) with apex of gnathos 
bearing a small spine; aedeagus (fig. 30^1 
rather stout, cornuti of two compound spines. 

Female genitalia (fig. 40) with ductus bursae 
swollen prnximally. 


Eg*. Ovoid, greenish, wiihout markings; 

stimulated to hatch by rainfall. See McFjixl&nd 
1971, p. 242 Un' full details. Material: GlOO. 
Final iihstui larva: Head capsule 3.3 mm wide, 
greenish with numerous small brown blotches 
some of which form suffused hand across 
vertex; body bright green with dnrk speell'mg 
on dotsum of prothorax. on anal plate and on 
claspcrs; yellowish-whac lateral stripe often 
heavily edged with fuscous *:^ its lower muT^in, 
-..imeumes four faint parallel whitish IlllCE 
dor/sally, two whitish suhvcntrul stripes preseni 
and incomplete mid ventral Btnpe W Al to 
Af», bases or setae inconspicuous, crotchets 
16-IM on A4 ( 20-22 On AS, 29-30 on A6 and 
33V38 on aiiaJ clampers; length 36-40 mm, 
width 4,5-5.5 mm. Material; G100, 
Papa dig. 15) mid brown ill colour: eremastcr 
hooks 2; dorsal anterior margin of A10 with 
2 lobes. Material: GlOO. 
Food plants: Predominantly Acacia pyenantha 
B-jnth.; also recorded from A brathyho"^ 
Bentb. and Cassia ?mvnophita A. Cunn. 
(McF-irland 1979), 

Specimens rwminetl: 113 o* 77 ?. QUTiENS 
I AND; Stunthnrpe. iv JH 1 J SAM: Talwood, 
h WtfBfi i specimen QM. NfiW SOUTH WALES: 
I km NNW of Goolgowi, v. TFBC & MSU 1 J 
ANIC; 10 km SB of Cm I Go!, v IFBC 1*1 V 
ANIC' lea Tree Creek ni Armidale. v. CWF 
I J fjMG VICTORIA; Biahip. iv. 8 specimens 
NMV iv. I & SAM; Gippshmd, 1 ? NMV; 
Oisborne. iv. I ? NMV, iv. GL 1 ? SAM; 
Hamihon, 1 9 NMV; Itncbkirch. iv LRK I spec- 
men QM; Melbourne, I 5 SAM: Spnnsvale, v. I 
V NMV SOUTH AUSTRALIA Adelaide, iv., V- 
.(GOT r 4 1 ? SAM, v I (f SAM. v. FA 3 9 
SAM; Aldp:itc. V, ( 9 SAM. Aihelstone, IV . V, 
IRIS 7 \$ 2 9 SAM; Pelair, iv. FMA 1 8 SAM: 
Warkwnnd. v. NB'l I rf I S ANIC, in.. W.. v. 
NMel 2 fl 3 9 ANIC. HI ■ « v -i * NBT 20 J 11 
<? SAM, n. VHM I S SAM. iv FA I 9 SAM. v. 
FA I S S\M, v. OBI 1 ,-T 5 ? SAM; BOwhOK [v. 
FMA I 2 SAM; Bunr-iiie, i 7 SAM: IuO km E 
Di QwXim, \ rFBc fi MSU 2 & AMC; Gk-n 
Osmond, v FMA I 2 SAM. Hiehenle, iv. I 3 
SAM. Kadina. v. 1 ? PBMcQC; l.ynton, iv., v. 
KHFl 4 AMC. iv RTIF 4 ,? 4 ? SAM. Mam- 
hr-jv' Creek Nat. Pk., v, IFBC 1 $ AKIC Momma 
'So. .in. v JWD 1 ,-T SAM, 4$ km FSE of Morgan, 
v. IFBC & MSU 1 V AMC. Mt. larfty, AMU 
I ? SAM. Parksidc. OBI I S 1 SAM, iv FMA 1 
<T SAM; Port Lincoln, 1 9 SAM; Renma.V. v. 
PBMeQ t ? PBMcQC: Stonytell, v. FMA I V 
SAM: Waikcnc, iv. 1 ? SAM; Warradale, iv.- v. 
PBMeQ 'I rf 3 9 PBMcQC; 10 km S\V pf Wil- 
mington, v IFBC & MSU 1 *? ANIL; Whyalln. v 
1 / NMV. 4S km S or WhvaU.i, v tt-RC 3 $ "^ 

Jnlct, 1 J WAM; Buragup, 1 2 WAM; Cape 

NatUral.Mt. -v. IFBC & MSU J $ AMC Lar- 
Jinjaup, v. i 9 WAM; CUucmont. 1 $ WAM; 
26 km \ of Collie, iv. IHlC & MSU 1 <$ ANICi 
.1) km WSW of Collie, iv. IFBC & MSU : rf 
AMC; 27 km St ot Coo|i>urJie. iv. IFBC A MSU 
2 r' ANIC, Corrigin, \ $ WAM; Crawley, vi. 
KRN J o AMC: Denuiurk- iii.. iv. WBB 2 
spcCirneiifi QM| Orummoml Cove II km N of 
Geraldton, v„ vi. NAIeP 5 J ANIC: Liumhtcyung 

i ! wam, Dmuias i 2 SAM; Kararf, v I ,-' 

NMV; Kat.innini;. v, KRN 1 $ AMC; I ' '; 
WAM, Kojoaup, iv.. v., vi. ALR &£ 2 2 AMC, 
tv.. v. RJP 9 rf ANIC. iv., v MMHW 6 £ ANIC. 

1 I SAM: Lake Grace, iv. I g ANIC, iv. 3 speei- 
rnens C^M, 45 Km W ot MaJura, iv IFBC & MSl_ 

2 £ ANIC; 2 l J km W of MoRUmber. iv. IFBC &l 
MSU 4 c? ANIC: 98 km E of Norseman, iv. 

FBC k MSU 2 J AMC. 24 km N of Nor 
rhumpion. iv. IFBC Sc MSU 1 rf ANIC; II km 

5 of Pemberton, iii, IFBC k MSU t <jf l2 ANIC; 
Perlh, v.. viii. I J 1 f NMV; Piihjra, iv IFBC & 
MSU 1 rf ANtC, v. t ? WAM. Ravens woo d 
vi. f-M A 3 # I 5 SAM: Slockyard Gully, .lurien 
Bay, vi. BH 1 ? WAM: Swan River, vii. I rf OM. 
Uiinmin, v.. vi. 9 specimens NMV, Y; ( nchep Nail 
Pk, iv, IFBC & MSU : £ ANIC; Vtma. iv. IFBC 

6 MSU 2 i ANU:. 

Distribution, see fig, 50. F/&fc Period: see 

Comments: T. ungnloui is widespread in dry 
sclciophyll forest and malice areas in South 
and Western Australia extending to wet sclero- 
phyll forest in southwestern Australia. How- 
ever it is rart in this habitat further east It 
is syniputric with I. lettathula over a large 
part of iu range A favoured hahitai is The 
open woodland ot the Ml Lofty Ranges where 
Aauia pytnaaiha is commonly a dominant or 
Lo-dominant shrtih. AdtiHs enierge after a 
sticeesbiun of eold nights in mid autumn and 
persist until late May. 

Jluilama tiara Walker 

rn.s 7, 16. IS. 21, 3I ; 4U 50, 55. 

ihxihtinn •t""* W.ilkcr lft5S t p. 660: Meynck 

Mkyritifc L8SO, p. 654: 'turner lr?|9. p. 38S. 
Holoivpr- • labelled "Australia 52-39' 1 m 

Adah <hg. 55). Head with frons rounded, 
smoothly ^ hair-sealed, whitish-huff; vertex 
roughly hair-sealed, nisi colour: labial palpi 
With terminal segment and apex of second 
segment fuscous, remainder white; antenna 
ot male laminate. Fhorax white, anteriorly 
i:re\ -white, orange at base i}\' wings; legs with 
fore and mid femoia and tibiae and all tarsi 
infuscated, hind femora and tibiae white; Tore- 

P. B. WcQUlll AN 

wing (tig. 7) wilh costa stra^hl, gently ar«-heU 
apically, termen arched and stieluly sjiviuio 

heneath apex, R, anastomosed whh Sc and 

again wilh R 2 » ground colour shinine while, 
narrowly Mack-margined red-hrown streak 
tit'in hase l" CrtW ihiid costa then angled to 
mid-discjil hhm where it bifurcates emitting 
iiiu: siieyk lo iornus i hence alone, posterior 
mareiii to near and another stieak tracing 
IVT ;{ Lo mid termen then back to fonr-Oflhs 
cosl;i. termen narrowly light fuscous 1 , cdla 
orange posteriorly fuscous; hindwiny shinm.i 
white with a rounded fiwous siihapicai blotch 
never exiemling to anal anjile, cilia white; 
lorewinu beneath white, cosla iind <i (nangnCi 
subnpical blotch fuscous, costal edec oi lhi> 
blotch oranac. hmdwing beneath white, stiha- 
pical Notch usually larger than an ripperslchj 
and orange above M T ; witig expanse I 
H 48 mm. 9 40-4S mm. Abdofteti WhiU- 

Male xcfiitttiia (lie. 2\ ) with apex of enalhos 
with small spine; aedeagus \ hg. 31 ) with 
eornuti of two compound spines one nohceaMy 
longer than other 

Female xeniiaha { Qg, 41) wilh sclerotised sec- 
tion ot dUCtUfi bursac longer than COlllculuTf?. 
i'itnd ifistar lai\;t. Head capsule 3.2 rum wide 
blue-green with small fuscous htdttthes 
upper half; body bright olive ereen with ).-,, 
Wavy whilish lines dorsally; enclosed green 
areas irregularly mottled wilh bluc-e-.iecn, cv- 
trcme posterior margin of each segment uhjiisn. 
fleshy yellowish lateral line irregularly edged 
with pinkish and fuscous below, lateral Sreaa 
mottled With while and finely speckled MflJ&k. 
two whitish subventral siripes on most Mo- 
ments and diagonally extended to thoracic legs 
and prologs, ventral areas pidc green with 
several whitish parallel lines; cmichcts 1 3-1*3 
on A-l. 15-19 on AS, 25-28 on A6 and "_"-' 
On .mil claspers: lem-th J3-37 mm. width 
1 5 13 mm. Material GI2X 

Pupa fiig- lb' daik brown in colour; crcni> '. ' 
hooks f« dorsal qmcrior margin of A 10 With 

: lobes. Material Gt2& 

Fitotlptant: Acacia tlc< arrcti* ( 1. Wendl, ) 

Spa tnti m - < arnau\i 117 4 '' ''Ml \ S 

lAND; Hlackbnit. 1 ? I.:0; Millmei ran. v. 1M 
1 ? AMC. v. : ? NMV iv IM I ,-f I in 'i»m,- 
woomba, iv 2 1 i 5 nmv, ll)., Iv l( v., vin. EJD 
9 g 4 V U£j, v |OM 1 S UQ: NF-W SOU •• 
WAI I s AutUev. v. HST I eJ ANK rCrn.nuion 
iiotise via Salisbury, v GP.M I V VQ, Panvr ! 
iv. HST t S ANIC; 6 km V of BilttftWaM iii 

IHbC & MSU I ,-; AMC; Church Poidt, V JFBC 
i 1 \N<C; 7 km S\V of Goslord, iii. IFBC & 
MSU 4 J ANK: llornsbv. iv. 2 o 2 + ! NMV. 
KJIUrw, .v . v 3 ' I ; N\tV. Marulan. iv. 1 ? 
SAM: Millag-m-, iv. I g 2 r t NMV; Mttlgoa, LMR 

i , : r am<\ Naiata. Iv, t.MR 4 .; n S ank 
Njtioo.iJ park, [v, I ' NViv. Onuige, \i, fisi 1 
.V I .' ANU ! Pine ficck via loi)\ Harlun, v 
GBM I rf UQ; Ko^cvilie. n IMS 5 o ANIC; 
40 km S of Singleton, in, UHC I rf ANK . 
Sydney, iv l J \\1V, Scrub, Iii. 6JD 
2 J ANIC: i... iii. I JO 7 o ' V UQl '» ubrahue. .. 
< ork, H.iMtnviot. Tops. i. RS I J ANIC, i. I . ? 
NMV; M Km S of Wuiichnpc. iii. IFBC <^ MSU 
I $ AMC. I? km NL- of Win<lsor. iii UHi A 
IhRKITfWsV. Black Mounlain, ii., iii. II IK 5 i 

ANir. in, it-Kc t ,(• uo; Canberra, N II '■' 

1 C ANIC: Condor Creek 8UU ni, ii. 1MK' I . 
ANIC. 3 km N of Lee's Spring 1201) in. ii. IPB< 

A IDE I cP ANIC. VICTORIA. Uulwvn, \i. I d" 
NMV. bi-tka Ris'cr. Maltaeoola. jit. I V MV1V. 
I ...n'tTbury, iv 10 ^pccimenv NMV, V, 1 J SAM; 
CastlemaniL'. iii. I " NMV; ChcUcnham, iv. 20 

ftfwehrrcitf NMV; Cilh lVmu. [v. CC1 l r 2 5 

AM' ; OaiKlentmv- iv r > ipecunen> NAIV; Ois- 

t>ornti IUh iv. cl :n $^ccUnc»s nmv. Lit. jv. 

CI. 5 di ' 9 SAM. tv vo. I J LTQ; H,/,lv.ooa 
iv JHC 1 c* ANK. .VT.ieedon, \i.. ii.. 2 § NMV; OW 2 rf I 9 SAM: M.ieh.m l\ 
5 f 1 ';' NMV: Moe, ni, iv. UCLG 1 r t 1 ? 
ANK 1 , .v RAIV 2 (? ANIC. o QCJJ3 2 .' SAM. 
IV, I d CO. Ml Donna Huang, i. 1 ,, NMV, Ml 
l.i ica. fi I " N'VIV. Ml \V;ivcilcv, i -r NMV, 
MynJclonl. iv. I £ NMV; Oakleieh | 9 \\IV: 
Otway K..nyex. II, PRMcQ I o J V PRMeQC '.: S:.k, 
IVi 2 _? NMV; Sprinyvale, iv.. v. 6 speeimcnN 
NMV. v I , : : SAM, iv- I HI I J UQ; 'C.. 
Valley Nad l>aik, ii. PBVlcO 2 ? PBMeQC* 
rhiin.i Rivei in. Cape HvcrarJ, iii. I ,t NMV. 
li.ualcon. iv. 2 d" 2 2 NMV; Wakroanya. .v., 1 $ 
SAM; Wan.lin, ; ^f NMV. Woori Yalloek, iic. 
I J NMV- TASMANIA: Mole Creek, tv, IKi 
OBI J f SAM; Ni.nKoo.te, iii., iv. IOW I r 

\nic. Wo0dvilk i\. IB ] 5 sam 

Oisirilmtuar ^cc fig. 5U. fUtfta PcikkI: 'a:e 
% L8 

( i-miiunt< 1 ditw fanyes over mucli of the 

vanie 'vi'c «■! hahiCat a* V \>'i,u<n'n liowever 
ils presence in Tasmania and near A.U.ndc 
requires eonfiimunori- 

IhaJtiina inscnptii Walker 

FIGS % W ::. \2 K Sfff, 56. 

I fintiuttti 'tsu'rifhi Walker »K55 ( p, f»6|, M<-\i - k 
|; '. p Sy3 I QWCI IX-TV p 2vi> ihuvj.): 

Turner W9 p, 3W. 

rh/.\</n:\rHi firiticintriit Morn'ch-Sehallei |^55, pi. 

78, lit- 446. 



Ah.writ'y print i par ia Iferrich -Selidller. Guenee 

1*57. p. 227. 
TlWtfim htmyHfphltn lower IS'H, p. 2h0, Cn|J- 

Jinch 1944, p, 191. 
Topics ol in&rripia: krtotvpe 6 labelled 
"V.D.L 51-153" in BMNH. hereby cfcutg- 
natcd, \ S I 9 (>antl<ct<>rypc\\ 1 g labelled 
"V.D.L. 54-9" in BMNH, hereby designated, 
I , labelled as Tor leciotype, in BMNH, 
hereby designated, 

type o( prnutpana not in Zoological Insti- 
tute, Martin Luther University Wittenberg, 
GDR (N. Gosser, pers. cormu.), 
ffaiotypc % of httro^typhha labelled 'Wpril 
"93, SG6 Blackwood y0§ V. hietoxh'ptiim 
Lower" in SAM. 

Adult (Fig, 56»: Head with irons rounded, 
smoothly hair-sealed, whitish-buff; vertex with 
rough hair-scales, pale fuscous; labial palp with 
terminal segment and most of second segment 
pale lu.scous, second segment beneath with 
long white scales, antenna of male laminate, 
t boras .grey-while becoming whiter posteriorly, 
pale fuvcous at base oi wings; lorewing (fig, 
8) with costa nearly straight, apex produced; 
(ermen strongly arched and sinuate beneatb 
ape\: K, anastomosed with Sc and again wilh 
R., ; shilling white markings oehreoir-. io pale 
fuscous, narrowly margined darker: streak 
from hJWW to one-third ctifltS 'hen exlendiug to 
m!d disc where it bifurcates mlu an upper orm 
icaching mid termen rhence to eostn at three- 
quarters, and a lower arm reaching Io tornus\ 
tbeiiee thieklv along posterior margin to base, 
termen with small semi-eiruilar marking ai 
the extremities of M.., CuA, n and CuA , and a 
larger triangulnr blotch below apev. cilia pale 
fuscous; hind wing shining while with a 
moderate fuscous subapicrd blotch sometimes 
extending to termen and costa; lorewing be- 
neath white with upper-side markings visible; 
costal half o\' these hcing faintly outlined wiLb 
pale Inset mis below; hindwing beneath while, 
stibapical blotch reproduced and usually larger; 
wing expanse fl 3-S 44 mm c 40-42 nun 
Ahdomen white with pale luscoits shadings 

Malt' HeMftiiUl (6g- 22} With tegumen rather 
narrow: aedeaeus (tig 32) wiih eornuti of two 
subci|iia1 compound spines, 
PcmaU* x'uijuliti (fig. 42") wiih papillae anales 
rather la fee: ducui-. hnrsae relatively short. 
/ dial iir.fnr larva: \ modified from Lower 
IS')?). Leijgtb 26 mm Head capsule 2.3 mm 
wide, pale green wilh small fuscus blotches 

across upper frons: body green, lightly decked 
darker, a mid dorsal snipe o\ dense blackish 
speckling; two fate dorsolateral \*, hilish stripes; 
fleshy whiiish lateral line irregularly blotched 
with fuscous above arid Ivlow; a thin whiiish 
subventrol stupe; vent rally pale green Willi u 
white mid vcntial stripe; crotchets 13-14 on 
A4, 12 on A\ 25 on A6 and 2K -29 on anal 
claspers Material: One specimen "Tas., 15 
km NNW o( Auckland. 1^ viii. 1**80 On 
t.wv/a mwtrwft De Wild P. fe McQ'\ in 

Piijuf: mid brown in colour; 6 eremaster hooks, 
dorsal anterior margin of A 10 with 2 lobes. 
Material; One specimen labelled as above, but 
"pupa a i\. 19K0'\, in IDA. 

Fooilplanit; Acacia da nm/rs (L Wend I. ) 
Willd.. A mcanisa' De Wild. 

SpfitfmtWt cxuntim'tl: 53 3 21 ?. QUI ENSLAND: 
Veppoon, i. HBC I *? ANIC; NLVV SOUTH 
WALES Km S of Ml Timidly 1600 m, ii. 
II tu" & MSU I ,' \N1C; x km SF of Pilot Hill, 
BtgO Ffll&t, Hallow, iti. IOC I o ANIC. AUS- 
Creek Rd. 1000 m iii. If BC I rf \MC; VIC- 
TORIA: Hendigo, iv. I S NMV. Faalemaine. iv. 
4 4 I V NMV; Gippsland, iii. iv. 30 specimens. 
NMV: G.shoinc. iii.. iV, 4 £ 2 ? SAM. iii. iv. 
30 sptrcinneus NMV" Macedon. iii. 2 S NMV; 
Melbourne, iv. I n NMV; Tuol.tnyi, ie. 3 rf I ? 
NMV, Warntin, iv. 5 spccimi*ns NMV. T \S 
MAMA: Bellenve. iii BM 1 .' ANIC, Cressy. 
IV, 1 ,1 IDA: Fern Tree in. RJH 1 ( MDA; Hcll- 
yer c.oiec \l 1PBC & MSU I 9 ANIC, Kingston 
iv. JRC 2 ,' tJQ iv. JRC 2 rf ANIC. iv. IRC 3 g 
4?TMU<. iv, JRC t cJ NMV; L. l.eake 1500 m. 
ii, JFHC .v Msu i v ANIC; Iti km W tff Way- 
dena. ii. I J NMV; Mi Hanuw 800 m. iii. IFBC 
& MSU I & AM<- Ml Field NtttalMl Pflr* 1*0 
m, ii. ITBC & MSU 1 4 'VN1C: Ml Nelson 700 
iti, iii.. Iv. PBMcQ S cf 3 <j lOA: Ml M'elbneion 
271) m. ... ii . iii. iv . BM LJ g 5 ? AMC. Qtiie, 
iii.. Lv- 3 cJ TIM: Pyermana 310 it., iii. lt*HC $ 
MSL' 2 ? I ? ANK Riilg0W*V, lj- » "I NMV 1 . 
Iravv.lbn. iii. RJlf i r? TDA; 21 Vm S of WVm 

bury iii. "THC $ MSU I -* anic. 
DfsirihHtion; see tiu. 50. Fffefo PfWori: see fig. 

Comments f- inscriput is an interesting species 
wilh larvae adapted in colour pattern lo living 
on bipinnate Acacia food phints. It is the 
most cotd-tolcranl of the germs, being the 
commonest species over much of Tasmania 
where it occurs in open woodland up to 
1000 m; it also occurs in the Australian Alp9 
up to 1500 ni. I he single record from central 
Queensland is noteworthy in view of the ab- 


p. b. Mcquillan 

Figs 9-11. Venation of Thalaina spp. 9. 1\ paronycha; 10. T. allochroa; 11.7". kimba. 

pro meso A1 



* Dl 



) ^ ® di 







9 SD 







6 SD! 

in * L3 

S ® 

© Sit 



SV •« 




• 5VI 






- e 


" VI I 



A5 A6 A7 A8 A9 A10 





















Vt ■ \ 

Fig. 12. Sctal map of final instar Thalaina clam. 


sence Of records from northern N.S.W. and 
southern Queensland. 

Adult moths were abundant in an open 
eucalypt woodland in which Acacia meanmi 
was the dominant tali shrub about 8 km south 
of Steppes, central Tasmania, on 6 iii. 1980. 
In the mid-afternoon sunshine months were rest- 
ing on the undersides of Acacia leaves some- 
times two or three per tree, and were readily 
put to (light by walking near them. Their flight 
was extremely erratic and sustained; and 
generally less than 5 m above the ground. Al- 
though conspicuous in flight, they were not 
easy to see at rest on the trees in the dappled 
light when looking against the sky. 

Thalaina chionoptilu (Turner) comb. nov. 
FIGS 5, 18, 23, 33, 43, 49, 57. 

Mttcqitccnia chio/taptila Turner 1947, p. 102. 
Types: leciotype & labelled "Milmerran, Q. 
25 APR. 1936 J. Macqueen : Macquecniu 
chionoptila Turn. TYPE" in ANIC, hereby 
designated: 4^4? parak'crotypes, 1 ,5 . 
3 9 "Milmerran, Q. 14-5-31 J. Macqueen", 
1 3 "Milmerran, Q. 27-4-31 J. Macqueen", 
1 3 Milmerran, Qld. 27 APR. 1935 J. 
Macqueen", 1 ;. Milmerran. Old. S May 1935 
J. Macqueen". 1 9 "Milmerran, Q. 20-4-31 
J. Macqueen : Genitalia MS IS P.B. McQ. 
197S" in ANIC, hereby designated. 

Adah (hg. 57): Head with frons rounded, 
smoothly scaled, whitish buff; vertex with 
erect hair-scales, bright red-brown: labial palp 
buff, becoming whitish towards base of second 
segment; antenna of male strongly bipectinate. 



1 MM 

Eig. 1 3. Head capsule of final 

inslar Thalaina 

Thorax greyish-white on anterior third, re- 
mainder white; legs white with exterior of fore 
and mid legs infuscated; forewing (fig. 5) with 
eosta straight, apex pointed, tcrmen slightly 
sinuate beneath apex and hardly arched, R x 
often connected with Sc by short bar and some- 
times anastomosed with R 2 ; ground colour 
above shining while, orange-brown spot at base 
of Cflsta, extreme costal, termen and posterior 
margins orange-brown, cilia orange-brown; 
hindwing shining white; subapical and sub- 
tornal fuscous spot present, cilia while; fore- 
wing beneath furtively infuscated; orange cos- 
tal streak from near base to five-sixths, inner 
margin streak absent; hindwing beneath white, 
fuscous spots as above but larger; wing ex- 
panse 3 4S-52 mm, 9 52-54 mm, 
Male, genitalia (fig; 23) with socii rather pen- 
dulous, apex of gnathos enlarged with 
numerous small spines surmounted by a larger 
single spine, furea relatively short with prongs 
of unequal length; aedeagus (fig. 33) with 
cornuti of two groups of small spines. 

Figs 14-17. Pupa and cremaster of Thalaina spp. 14. T. selenaca; 15. T.; 16. T. ciara; 17. T. 


p. b. Mcquillan 










-* selenaea 

-♦ clara 

"■ angulosa 

■* tetraclada 







Fig. 18. Flight period of adulls of some Thalaina spp. 

Female genitalia (fig. 43) with colliculum 
rather broad. 

Immature stages not recorded except that larva 
pupates underground. 

Foodplant: Acacia harpophylla Bcnth. 
Specimens examined: QUEENSLAND: Millmer- 
ran. iv., v. JM 5 J 4 ?, ANIC: Millmerrun, iii., 
iv., v. JM 28 specimens, NMV: Millrnerran, iv., 
v. JM X specimens, QM-, Miltmerrun. iii.. iv., v. 
JM 4 d" 8 5, SAM: Millrnerran. iii., iv., v. JM 
5 rf 4 ?, UQ. NEW SOUTH WALES; Trangie, 
IV. RL 1 <J. ANIC. 

Distribution; see fiy. 49. Plight period: see 

fig- is. 

Comments; T. chionoptila is mainly known 
from a long series of adults taken at Mill- 
merran in the 1930's and 40s by J. Macqueen. 
Land clearing since then has much reduced 
the availability of its foodplant in southern 
Queensland. On present evidence it is the most 
localised member of the genus. 

Thalaina tetraclada (Lower) comb. nov. 
FIGS 1, 18, 24, 34, 44, 51, 58. 
Arncloru tetraclada Lower 1900. p. 406. 

Thalainodcs tetraclada I .ower 1902, p. 23 1 : 
Turner 1919, p. 386, Wilson 1972, p. 123. 

Thalainodcs nessostoma Turner 1919, p. 386; 
Wilson 1972, p. 123. sytu nov. 

Holotype £ of tetraclada labelled "3459 

TYPE Broken Hill 6.5.1899 : tetraclada Low. 

3459", in Lower's hand, in SAM. 

Holotype 9 of nessostoma labelled "'probably 
Bourke coll. Helms : Thalainodes nessostoma 
Turn. TYPE : G. M. Goldfinch Collection" 
in AM. 

Adult (fig. 58): Head with frons naked and 
bearing rounded projection (fig. I); vertex 
orange or tan; labial palp pale brown above, 
white below; antenna of male strongly bipec- 
tinate. Thorax orange or tan above with inner 
margin of tcgulae and sides of thorax white: 
legs white, exterior of fore and mid femur 
and tibia and all tarsi infuscated: forewing 
with costs nearly straight, apex produced, tcr- 
men gently arched and oblique, ground colour 
above shining white, orange or tan costal 
streak from apex to one-half costa attenuated 
anteriorly then continued as fine streak to base: 
faintlv black-marained orange or tan streak 




i'.i & li / 





Figs 1 9-21. Vlnic genitalia. 19. T. selenaea; 20. T. an^utosa; 21. X. c/ffrw. Scale lines I mm. 

from base to just above mid lermen tracing 
M ;{ and emitting similar streak from about 
one-fifth to one-third tracing CiiA., to lermen; 
streak along posterior margin from near base 
to tornus, lermen narrowly orange or tun, 
Cilia orange or yellow; hindwing shining while; 
large fuscous subapical blotch, projection of 
which often touches termen or is continued 
siihtcrminally to anal angle, cilia white; fore- 
wing below white, lightly infuscaled basally 
and below CtfAn, costa and lermen narrowly 
orange Of tan; fuscous subapical blotch pos- 
teriorly orange or tan; hindwing below shining 
white; maeulation as above; wing expanse c? 
38 48 mm. S 38-46 mm. 

Mate genitalia (fig. 24) with uncus rather 
long, unathos with a large apical spine hearing 
numerous smaller one-, f urea rather linear and 
both forks of equal length; aedcagus (fig. 34) 
curved basally, cornuti of two groups of about 
3 spines, 

Female genitalia (fig. 44) with apophyses an- 
teriorcs nil her short, colliculum longer than 
sctcrotiscd hind of ductus bursae. 
Specimens c.\«tni»rd: 37 4 33 ? NEW SOUTH 
WALES: broken Hill. v. OBI. 2 r ? SAM; Bourke, 
1 $ AM: SOU IN AUSTRALIA: 10 km SW of 
Iron Knob. v. IFBC 4 ,V 10 9 AN1C; 64 km F of 
Nnllahor, iii. IFBC & MSU 1 tf AN1C: 89 km E 
of Nullabor. v. IFBC & MSU 1 d. 1 5 ANIC; 

46 km SW of Whyalla. v. IFBC 4 2 ANIC; 
Whvalla, v. 2 ? NMV; WESTERN AUS I "RALIA: 
27 km SE of Coolganlie. iv. IFBC & MSU 3 o* 
2 r ANIC: Dahvallinu. iv. LEK I specimen 
VVAM; Dumbleyung, IV, HU - specimens WAM; 
Kaftarri Nil Park, iv. IFBC & MSU 2 rf ANIC; 
Kojonun. iii, RJP 1 J, iii. KB 1 A iv. AIR 5 <J, 
iv. RJP 6 rf 2 $, iv. M.MIIW 3 J 1 i\ v. RJP 
I ,;\ v. AI.R I ,:. v. MMHW I , ;'. all ANIC; 
45 km w of Madura, iv. IFBC & MSU 2 ? 
ANK : Merredin, LJN 1 specimen QM; 46 km 
W pi Mcrrcum. iv. IFBC 8t MSU 1 rf ANIC; 
29 km W of Mogumber, iv. IFBC & MSU 1 S 
ANIC; 97 km E of Norseman, iv. IFBC & MSU 
t J ANIC; 24 km N of Northampton, iv. IFBC 
& MSU 3 J I S ANIC: Pilhara, iv. IFBC & MSU 
I & 4 8 AMC; Tammin. v. 2 J NMV. 

Distribution: see fig. 51. Flight period: see 
tig IS. 

Comments: This species ranges widely across 
the subinlcrior of southern Australia, reaching 
the coast along the Nullabor Plain and south- 
we&t Australia. East of W.A. its range lies be- 
tween the 20 cm and 35 cm isoh>cls whereas 
in southwestern Australia it extends to the 
100 cm isohyct, apparently occupying the 
niche filled by the three forest species in south- 
eastern Australia. Some clinal variation exists; 
specimens from the central and eastern parts 
Of its range are usually smaller (mean wing 
expanse 40 mm) and the forewing markings 


p. b. Mcquillan 

Figs 22-24. Male genitalia. 22. 7\ inscripta; 23. T. chio/toplUa; 24. T. tctrartada, 

arc tan or ochreous-orange, whereas in S,W. 
Australia Ihey are usually larger (mean wing 
expanse 46 mm) and with bright orange mark 
ings. There are no genital differences between 
extremes of Ihc dine. 

Ihalai/m tnavfarlamli (Wilson) eomb. nov. 

FIGS 2. 17. 25, 35, 45, 51, 59. 
riuilnnioth's awvJarlanM Wilson 1972, p. 12."*. 
Types: Iwlotype l ? labelled "NTHN. TERR.. 
148 km S of Alice Springs 26 April 1%6 N. 
Mefarland u( uv light" in SAM; allotype and 
painty pes. see Wilson (1972). 

Adult ( figs 2, 59) adequately described by 
Wilson (1972); wing expanse J 3S-44 mm, 
9 38-50 mm. 

Male genitalia (fig. 25) with tCgUfTKtfi '"id 
valva elongate, apex of gnathos with large 
recurved spine bearing some smaller ones. 
Furca goblet-shaped, juxta elongate; acdeagus 
(fig. 35 ) with apex produced, cornuliis a 
9Clcrotised plate hearing small marginal spines. 
Female genitalia (fig. 45) with very long duc- 
tus bursae and without sclerotisation. 
Egg ovoid, pale green with an elongated brown 
blotch, 1.0 mm long \ 0.S mm wide. Material: 

Pinal instar larva; Head capsule 3.3-3,4 mm 
wide, pale green with white band across vertex 
bearing many small fuscous blotches; body dull 
green, white lateral stripe strongly developed 
on pro- and mesothorax and again on A6 to 

A9 but usually poorly developed or absent on 
intermediate segments, spiracles black and 
placed above this line. L setae on A2 to A 7 
arising in black blotches on lower margin of 
lateral stripe, anal claspers with two black an- 
terior vertical stripes, ventral areas pale green 
with two whitish subventral stripes on A7 and 
AS only, crotchets 13-14 on A4, 14-15 on 
A5, 28-30 on A6 and 45-48 on anal claspeis, 
those on hind claspers tending to be biordmal: 
length 27 "U mm, width 5.0 mm. Material: 
G 1 80. 

f'u/Hi (fig 17) pale orange-brown; crcmaster 

hooks 2; dorsal anterior margin of AH) with 

2 lobes; length 1 I mm. width 4.5 mm. 
Material: G180. 

Food plant \: Unknown, but larvae thrive on 
A cat ia pyrnaathn as a surrouale host 
(McLatland 1979.). 

Sprrimcns examined: 14 g 16 ?. QUEENSLAND: 
Curmamulla. v. NG I g AM; 24 km S of Mill 
men an. iv. IFHC 1 ? ANIC NHW SOUTH 
WALKS: S0.S0S I4G.33B 23 km SSh of Bvrock, 
v. EDE .v. MSU 1 o* 2? ANIC-. Cobar. iv,. 
v V.IR 2 ? ANIC; 29 km li of Vaughan 
Springs HS. vi. KJMcK t ? ANIC. SOUTH A US 
TRA1.IA: 27.I8S 133.251- Ammuroodinna Creek. 
v. PBMcQ | cf PRMcQC; 160 km NNVV 0f 
Coobet Pedy nr Winlirma, v. NMcF & TN 1 g 
SAM; 61 km S of Kulgero, v NMcp A TN I ■•* 
Alice Spnnzs. v. NMcF & IN I ' ANIC I / 
I ? AM, I 9 BMNH, 1 tf I ? NMV 45 km 
VVSW of Alice Spruits, v, NMef* & 'IN 1 V 
SAM: [48 km S 1st Alice Springs, iv. NMcV 



Figs 25-28. Male genitalia. 25. T. macfarhtndi; 26. T. kimbu: 27. T. paronycha; 28. T. aUochroa. Scale 
lines 1 mm. 


p. B. Mcquillan 




. ; 



Figs 29-38, Aedeagi of Thalaina spp. 29. T. sele- 
naea; 30. T. angulosa\ 31. T. clara; 32, T. in- 
scripta; 33, 7. chionrtptila\ 34. T. t tlrac I ad a; 35. 
T niacfarlandt; 36. T. kimbtr, 37. T. paronycha\ 
38- T. allochroa. 

I 5 SAM; 24.20S 13I.35E Amadeus Basin nr 
Reedv Rockhole. vi. PR 4 5 ANIC: 23.48S 
1 32.2 1 F 5 km NE of Go&se's Bluff, v. HP 4 o* 
I ? ANIC. 

Distribution; sec flg. 51. Flight Period; laic 
April lo early June. 

Continents: The dispersed nature of the few 
locality records for T. macjarlandi suggests 
a wide distribution in central Australia. 

Thalaina kimba sp. nov, 
FIGS II, 26, 36, 46, 5K 62, 63 
Types: holotxpe <? labelled "'32.5 IS 14I.37E 100 
kin S by E of Broken Hill, N.S.W. 3 May 1976 
I.F.B. Common E. D. Edwards: genitalia slide 
M846 PBMcQ 1978" in ANIC; 10 cf 8 ? pom- 
types, 2 J same data as holotyne, in ANIC; I d* 
labelled "6 miles S.W. of Iron Knob, S.A. 7 May 
1970 I.F.B. Common; genitalia slide M842 
PBMcQ 1978" in ANIC; I <J labelled "6 miles 
SE of Got Gol. N.S.W. 5 May 1970 T.F.B. Com- 
mon" in ANIC; I e? labelled "100 fcm SE of 
Broke* Hill. N.S.W. 29 April LA76 1 .F.B. Com- 
mon E. D. Edwards" in ANIC; 1 o labelled " 1 
km NNW of Goolgowi, N.S.W. 5 May 1976 
J, F.B. Common E. D. Edwards" in NMV; I o* 
same data as for previous specimen in SAM; 2 & 
same data as for previous specimen in ANIC; 
1 tf I 9 labelled "33.23S I4E40E 82 km NW of 
Wentworth, N.S.W, 28 April 1976. I.PA Common 
E. D. Edwards" in BMNH; 1 ? labelled "31.49S 
141. I2F. Umbenimbcrfcn Reserve, 9 km NNW of 
Silvcrton, N.S.W. I May 1976 I.F.B. Common 
E, D. Edwards" in ANIC; ] ? labelled "30.50S 
146. 33E 23 km SSE o\' Byrock, N.S.W. 8 May 
1973 E. D, Edwards & M. S. Upton: genitalia 
slide M845 PBMcQ 1978" in ANIC; I 5 labelled 
"Mambray Creek Nat. Park. S-A. 11 May 1970 
I.F.B. Common: genitalia slide M843 PBMcQ 
197K" in ANIC; 1 s? labelled "30 miles SW of 
Whvalla, S.A. 9 May 1970 I.F.B. Common: geni- 
talia slide M822" in ANIC: 1 ? labelled "Kimba 
12.5,1963 R. E. Harris" in SAM; I ? labelled 
"Minnipa, S.A. May 1970" in PBMcQ; 1 g 
labelled "Moorunde Wombat Reserve, nr. 
Blanchetown, S.A. 12 Mav 1974 P. B. McQuillan" 
in PBMcQC. 

Adult (Figs 62, 63): Head with frons naked, 
bearing long rectangular projection emarginate 
at apex; vertex of head rough-scaled, greyish 
with suggestion of fuscous transverse bar be- 
hind antennae; labial palpi with terminal seg- 
ment white; antenna in male shortly bipec- 
linate. Thorax with anterior and posterior 
thirds fuscous-grey, mid-third and tcgulae 
pale grey; legs infuscated, except hind femur 
ind tibia; forcwing (fig, 1 1 ) with costa straight 
in male, slighlly recurved in female, termen 
sinuate beneath apex and strongly arched, R t 



tin I | 

Figs 39-43. Female genitalia. 39. T. selenaea; 40. 7*. anguhsa; 41. 7\ dura; 42. T. inscripta; 43. J. 
chionoptila. Scale lines 1 mm. 


p. b. Mcquillan 

Figs 44-48. Female genitalia. 44. T. tetraclada; 45. T. macfarlandi; 46. T. kimba; 47. T. paronycha; 
48. T. allochroa. Scale lines 1 mm. 



allochroa O 

paronycha A 

macfarlandi A 

tetraclada • 

Fig. 49. Distribution of Thaiaina selcnaea and T. chionoptila. 

Fig. 50. Distribution of Thaiaina angulosa, T. clara and T. inscripta. 

Fig. 51, Distribution of Thaiaina allochroa, T. paronycha, T. macfarlandi, T. tetraclada and 3". kimba. 


v. b. Mcquillan 

anastomosed with Sc. ground colour white, 
niai kings suffused grey nij sprinkled with 
bluish-white scales, base of cost a hlackisb; 
veiy broad black-margined streak from one- 
third costa dilated posteriorly td mid disc 
where it bifurcates, one arm lo mid Icrrncn 
thence angled to four-h\fths eost.i and extend- 
ing to half eo.sta, other arm to lornus theme 
alooju posterior margin to near base where it 
is mixed wilh black; broad JudefUed streak 
along termen. eilin brown is h-orey becoming 
crcv below M ;i ; Itindwing translucent whin* 
Willi several irregular terminal and subtermmnl 
fuscous blolches; lorewing beneath white with 
streak along posterior margin discal 
streaks narrower and more sharply delincd 
with fuscous, icrrncn streak 08 RbcFW hind 
winp beneath while, a subapieal and n stih- 
lomal blotch only; wine expanse fi 38 -14 
mm, ? 40-44 mm. 

Mah QCttftidta ( lie. 2b ) with e.dva ralbcr 
broad, apex ol gnathos with 3 or 4 small 
Bpia&r, futca long and sinuate with one brant h 
poorly developed; aedcagus I he- 36) with 
cornult of single stoul spine and group ol 3 
or 4 spines. 

CWTt&te f&fttttllia (tig. 46) wiih eolliculum 
stpiare, shfligma well developed, ductus bur- 
sac '.rry lonu its proximal thud with thick 
fold^. remainder thinly membranous. 
Divrti'tttton: see rig. 51- Probably widespread 
m mallee habitats from Eyre Peninsula 
through the Murray Mallee to western New 
South Wales. 

Flight period; late April 10 early Mav. 

Ihnlmm pamnytha i Lower) comb r<..\ 
FIGS 9, 27, 37.47. 51. 61. 

timlfra rjun-jnyrha Lower i^OO, p. 407. 
Thalointnlrs i>un>n\chu lower ojli.' p 2TI . 
Tomer l r J/^, p. .1S6; Wilson 1972, p. J 23. 

Hoforyp* y lahelied "3 4fi0 TYPE broken Hill 

24.5.n.S" in lower's hand, in SAM. 

Adiill (Gg. 61): Head with from, naked, 
bearing moderate truncate projection wilh 
longiiudmal rib beneath, vertex wilh ochreous 
hair-scdes which extend nlmosL to extremity of 
frontal proieetion; labial palpi whitish: antenna 
Ol male strongly bipeetinate. Thorax ochreous 
above, whitish below; legs tinned oChr&OUS, 
fore-nbia very short, bearing apical spine; h-rc- 
wme. dig, 9} wilh costa ^traighl, apex pointed, 
le'rueo moderately arched. R, anastomosed 
with Sc, ground colour shining while, marking 

dark ochrooic: or tan finely edged wilh brown, 
extreme costal edge while slightly broader 
trom one-sixth to one-hall costa, costal 
sireak trom base to apex narrowly continued 
along termen and slightly extended hut 
broader along posterior margin: streak from 
one-quarter costfi lo termen above middle trac- 
ing M ; , thence angled on termen to costa .a 
three-quarters, diagonal streak from half dbtciil 
>treak to just above torhus. cilia ochrcoxis: 
hindwiuy shining while tinged ochreous, pale 
fuscous subapieal spot, cilia white; fttfWing 
beneath white, tinged ochreous on basal half 
and on margins, pale fuscous diagonal suha 
pieal spot; hmdwing beneath white, subapieal 
spot slightly larger and darker than ahovc. 
small fainl fuscous spot near icimcn between 
CuA.i and A,; wing espanse , 56-40 mm. 
v 42-44 mm Adbomen ochreous. 

Mate genttmlhi [fig. 27) with teguraeti broad, 
npc\ of gnulhos elongate and hearing lonei 
nidinal row oi' about G stout spines, valva 
relatively long, forca reduced to small lobe 
aedeagus (ftg. 37) strongly curved, eomuti 
ol single spine and another group oi' fused 

Ft'm<tlv gtlilttilte (fig. 47) wilh apophyses an 
leriorcs nifher short, a band of sclerotisatiou 
ai top of ducius bursae similar in lenglh lo 
eolliculum, ductus bursae very long. 

s>t7>//ou txaminviL II rf 26 V NEW SOUTH 
WAI KS: Broken Hill, iv.. v. OBI. HI rf Ifi 
LIA; Ammaroodinna Creek 27.18S I31&3E \ 
PBMcO | £ I'BM.'.h 

Oistrihttfion; sec fig- 5!. flight periud: mid 
April to late May 

Commrntx: This and the next species are 
rather isolated from I he rest of the genus bv 
structural features iuetl as evmiaha. presence 
of a spine on the Fore tibia and the form ol 
ihc frontal process. 

Thofaina utltnhrou \\ ower) comb, nov 
TfGS 10 2S, 3.X, 4X. 51. 60 

TJh<t f <it,<nlt>\- ulfavhrOa i o*er ino:, p, 23.?; 
Turnt'i jyiy, p SX7; Wilson 1972, p. 12^. 

tottitfp* labelled "3 5 02 Broken Hill : 
3??9 ; Amelora alloebroa Lower TYPr" in 
Lowefs hand, in SAM 

Adult (flg 60) with head as for J. paroni'dia: 
vcrica With oehrct)us-bulf hair-scales which ex- 
tend onto frontal proteclion; iabiaJ palpi white; 





f A 



^^ '^»m^.._ . 


J. *W 




Figs 52-63 Adults of Thakdna spp. 52. T. selenaea eft 53. T. A^e/mea var. punctilinea fc 54. J. angK- 
losa tf 55 T c/tfra J 1 ; 56. T. inscripta rf; 57. 7\ chionoptila $\ 58. 7\ tetraclada c?; 59. i\ mac- 
farlandi d; 60. T. allochroa <?; 61. T. paronycha g\ 62. 7". *«w&a holotype d; 63. T. fcim&i para- 
type ?. 

antenna of male strongly bipectinate. Thorax 
ochreous-bulT above, whitish below; legs tinged 
ochreous; fore tibia very short, bearing apical 
spine; forewing (fig. 10) with costa straight, 
apex round-pointed, termen moderately arched, 
R-, anastomosed with Sc, uniformly ochreous- 
buff, often sparsely flecked with black scales, 
extreme costal edge white especially between 
one-sixth costa and one-half, cilia ochreous; 
hindwing white, slightly ochreous tinged, pale 
fuscous subapical blotch emitting faint sub- 
terminal line to tornus, cilia white; forewing 
beneath whitish tinged with pale fuscous to- 
wards apex and costa; pale fuscous diagonal 
subapical spot; hindwing beneath white, suba- 
pical blotch slightly darker than above; wing 
expanse $ 36-44 mm. Abdomen ochreous. 

Male genitalia (figs 28, 38) similar to 
T. paronycha, but apex of gnathos usually with 
4 spines. 

Female genitalia (fig. 48) almost indistinguish- 
able from T. paronycha. 

Specimens examined: 5 <J 2 $. NEW SOUTH 
WALES: Broken Hill, v. OBL 1 3 1 ? NMV, v. 
OBL 6 2 SAM; Mootwingee Historical Site 
31.14S 142.18E, v. IFBC & EDE 2 ? ANIC. 
SOUTH AUSTRALIA: Ammaroodinna Creek 
27.18S 133.25E, v. PBMcQ 1 <3 PBMcQC; 
McDouall Park, v. FWJ 1 $ SAM; Tallaringa 
Well. v. PA 1 $ SAM; 107 km S of Coober Pedy, 
Giles, v. 2 c? PBMcQC; Skirmish Hill, 1 rf SAM. 
Distribution: see fig. 51. Flight Period: May. 
Comments: T. allochroa exhibits a radical de- 
parture from the basic colouration of the 


p. u. Mcquillan 

genus Hut structurally it is virtually indistm- 

.uuisluhlc trom paronychti II is just possible 
that ulltHhtoa is only a MtMtdeltan segregate 
of punviyifui but breeding c\pennicni> arc 
needed la confirm this. Though infrequently 
collected, both species appear to be widely 
distributed within the 15-25 cm isohyets south 
Of IS S latitude. 


lliere arc no consistent venalioual, genua lie 
ov other structural differences which juMdy 
I lie .separation of I hutuinot{<'\ I owcr and 
Mat -itHi-aiui Turner from T/mktinu Walkct. 

Thfl presence o( an areolc in the t'orcw ine i^ 
not correlated with any other features Q\ 
diagnostic value beyond species leve! An 
yrcoJc K present in one form in aWe/mcw, in 
another iorm in <hiono(>tHu, clam and far- 
senpio and is ab>eni in the rest. Tower's im- 
pression i)t a longer cell in the hindwing of 
his Thohiittotlvs is erroneous as measurement 
will show. 

Nome diversity cxisN in the male ;iiucnimc. 
they are laminate with vcnfrally produced, 
cihaicd scL-mcnts in seknaca, ikserfptfl and 
tiara, shortly bipeclinafe in nuqnh>^a. mat fell 
tatidi and kimha. and strongly bipcettnatc in 
uliiH it n ><s, initnttyclut, thionoptila and fr/ra- 
clmia Earlier nuthnrs have previously over- 
looked the alight hm definite bipcetinntc nature 
of the antennae of uiixulo\a. 

The corneous frontal process appears to 
have ;irisen independently several times in the 
genus, as it has in many other arid /one genera, 
This structure, in conjunction with the forfr- 
tibial spines in athnhroa. puroitvt'/m and 
kimha. prohahlv assists the imajju to find its 
wav to the surface v{ the soil after emerging 
fioni I he buried pupa. 

A number o^ biolopieal features :m- 
shared, All are late summer to tolc autumn 

fliCPS with an annual life-cycle, li i- very 
likely that eggs of all species are stimulated to 
hatch by ram as angnloxa is (MeFailand 
1973). Larvae are associated with Anvia (or 
less oltcn CjUSftt) and where known, complelc 
their feeding in early spring and pupate just 
helow the surface of the ground where tliev 
i'vcr-siimmcr. Pupal aestivation is a phenome- 
non shared by many autumn-flying eunomines 
in southern Australia, such as the species \4 
Chlmioy {Madden & Bashlord 1977) Tho- 
ittttnt has successfully exploited a very wide 
range of habitats in the. southern half of Aus- 
tralia, u few species ha sin- adapted to each 
major ecological zone within the overall ranee 
of the genus. 

Previous attempts to fragment the genus 
have been based on either incorrect evidence 
or are unnecessaiy, since several other genera 
(e.g. one to contain allochroa and paronvcha) 
could be erected oty similar evidence. There- 
fore. I feel it is better to slightly expand the 
onemal definition c>\ Thala'uui to contain ill 
of the ahove species, thus reflecting their close 


1 am indebted to I3r L P. B. Common, 
C.SI.K.O. for helpful discussions and com- 
ments on the manuscript ;tnd to Mr D. S. 
Fletcher, BM(NK), for generous assistance 
with information on type specimens and titera 
Hire A generous grant from the Royal Societv 
Qj South Australia Research Fund enabled me 
io visit the Australian National Insect Collec- 
tion, Canberra, for which [ am grateful 
Thanks arc also due to the curators who 
loaned material or sent information, to Mr M. 
rotgeitcr for phou-grnphv and to Dr J. J H 
Szent Ivany and my wife Kathtyn who helped 
in various ways. 


Df.smarest. M. E. f185fti Tn Chemi. I C. Encv- 
tfjop&lie J'llistoire OOtUtelle —Tabic nlpfube- 

liqiji.: lepitlopteres. (Rons I 
Oman h>.\y, E. CI645) Descriptions of some new 
Aaslralian I epictopterons mseeis. In f. I. Kvr.:. 
Jt.\ fixfuhts' Pttc. MM AttSt. T Apnendiv pp. 
437-439, pi. 5. 

Flint in ft. D S. f I97M ) The eencric names of 
moths of the world. Vol. L Cieometrnide H 
1. W. B- Nye fed.. I. (British Museum, London, 'I 

CtomriNMi. (., M, (10441 Notes on (he Auslra- 
h.m Hoarmiidac and Oenochromnlnlat; (I cpi- 
dnplet'ii) with tlesciinnons of new species Or,,, 
Utm, Sor. A' SW. 69, 189-192. 

Gut m r V I 1857) Hisloire natiirelle des insectcS; 
Species general dfifl Irpidoplijivv. Tiftlte 10. Ufa 
niilcs et Phalenift.-s ft, pp. 1-S&4 ( Rorcl. Paris-. 

UtRurrii-SmArrLK. Q, A. W. (1855) Siimmln.'v 
neuei ouei wenie beknnntet .nisscrenropaiseh^r 
Schmetterhnge Volume ' Scries 1. 2n Heft. 96 
pi., fiyv 1-551. 

LpWER. O. H. f I89^» Ocscriptions oi uov ,\ns(i;«- 
Han Heterocera. 7^//;v. /?. .Vr.r V. Au*t. 17. 287 

(P-HUM Dcseriplions of new Australian Hcic- 

roccni Pror. Um Sor. NSW, i3(3i, 403 -A2 I 
— i I9Q0] DcNCriptions of new genera and spr 
cits of AostrMi<in Lepidoptern Txinw /.' S,n 
-V. Auk> 26. 212 247. 


Lucas, H. (1857) In Chenu, J. C, Encyclopedic 
d'Histoire naturelle — Papillons diurnes, 310 pp., 
40 pi., text illustr. (Paris.) 

McFarland, N. (1971) Egg photographs depict- 
ing 40 species of southern Australian moths. /. 
Res. Lepid. 10(3), 215-247. 

(1973) Some observations on the eggs of 

moths and certain aspects of first instar larval 
behaviour. Ibid. 12(4), 199-208. 

(1979) Annotated list of larval foodplant 

records for 280 species of Australian moths. J '. 
Lepid. Soc. 33(3), Suppl. 72 pp. 

Madden, J. L. & Basheord, R. (1977) The life 
history and behaviour of Chlenias sp., a geo- 
metrid defoliator of radiata pine in Tasmania. 
J. Aust. ent. Soc. 16(4). 371-378. 

Meyrick, E. (1892) Revision of Australian Lepi- 
doptera V. Proc. Linn. Soc. N.S.W. 6(2), 581- 

Thierry-Mieg, P. (1899) Descriptions de lepidop- 

teres nocturnes. Ann. Soc. ent. Belg. 43, 20-21. 
Turner, A. J. ( 1919) Revision of Australian Lepi- 

doptera VI. (Third instalment.) Proc. Linn. Soc. 

N.S.W. 44(2), 258-310 and 383-413. 
(1947) New Australian species of Boarmia- 

dae (Lepidoptera). Proc. R. Soc. Qld 58(6), 71- 

Wai ker, F. (1855) List of the specimens of lepi- 

dopterous insects in the collection of the British 

Museum. Part 3, pp. 582-775. (British Museum, 

(1865) List of the specimens of lepidop- 

terous insects in the collection of the British 

Museum. Part 31. Suppl. pp. 1-321. (British 

Museum. London.) 
Wilson, J. O. (1972) A new species of Tkalai* 

nodes (Lepidoptera: Geometridae: Ennominae) 

from central Australia. Mem. Natl Mus. Vict. 

33, 123-124, pi. 11. 


by Pat Hutchings, Patrick DeDeckker & Michael C. Geddes 


The polychaete Manayunkia athalassia n.sp. is described from ephemeral lakes adjacent to the 
Coorong Lagoon, South Australia. This is the first record of this genus from Australia. Manayunkia 
athalassia is active over a wide range of salinities (27-95%c) and persists in dry lake beds during the 
summer months. 


hy Em Mm. .him;$t\ Patrick Dr. Dr-CKMiKt 1 & Michael C. 0EBBCT1 


IJiit'iiiNos. P., Or Drr^KTR, P. & (fi:DDis, M. C (1981) A new species of Mumn/tnkw 
< Polychaetn) from ephemeral lakes near the Cooron?> South Australia. Trans. H, SvjL; S 
4un »*5(1 L S5-2& iZJune. Iffl-L 
The polychaete Manaynnkia nfhatassiu n.^. \$ described from ephemeral lakes adiuccnt 
to Ihe Coorong Lagoon, South Australia This is ihe first record of this genus from Aus- 
tralia. Xf<<rt<iYntik!« utlntfassiti is active over u wide range of salinities (ll-i*5'A) and persists 
in dry lake beds during the summer months. 


Ihe l.mna of athatassie (non marine) saline 
lakes tn south-eastern Australia has been listed 
in several studies (Bayly & Williams 19*6, 
Bayly 1970, Geddes 1976. Williams I97S, 
Do DecUcr & Gcddes I960). Hie only toomvI 
of polyehacte worms is from a small lake near 
Lake F.Iiza, South Australia where Ci-ratonc- 
reis eryfJiraeenxix Fauvel a lid Cupitclla c^pi- 
fata (Fabricius) were collected (Bayly 1970). 
These species are known to tolerate a wide 
range of salinities but cannot resist desicca- 
tion During a teceut study of 23 ephemeral 
lakes near the Coorong Lagoon. South Austra- 
lia (Dc DecMer & C.eddes |y80) another 
polychacte. a new species of Mannyunkia. 
was common in several localities. 

Munavuitkia athalassta n sp. 

Type material 

Holotype (AUsti Mus. W 17671) from ephe- 
meral lake at 36M9'46"S, ]39 a 44'48"E, adja- 
cent to Coortme, S.A., coll. P. De Dccfcker & 
M. C. Gcddcs 17.vii.1978. 

Para types from Various ephemeral lakes 
ad/acent to Coorong (20 specimens, AM 
VVI 7672), United States National Museum 
(1 specimen USNM f>337<»> British Museum 
(Natural History ) ( 1 specimen ZB l c >R0: 
265). Further material in Australian Museum 
im specimens AM WI7677). 

Ten tHCU tot crown colourless, antcrioi body 
up to scligcr 3-5 darkly pigmented, rest of 
body colourless, Thorax o\' eight 

Australian Museum, P.O. Box A2S5. Sydney 

Smith N-S.w 2000_ 

I Department of /oology. Lhiiversiiy of Adelaide. 

S A. 
1 Present address: Department of BjOgCOgrapfry & 

CiiDiiuirphology, Australian National Umver- 
stly, Canherra. A.C.T. 

abdomen of three setters, All thoracic setiecrs 
similar in size, abdominal setigers equal in 
lengths slightly shorter lhan thoracic seugers. 
Tentacular crown of paired symmetrical halves 
with semicircular bases, each hall" with two 
short compact radioles. Outer radioic with 
four short stumpy pinnules and inner radiolc 
with three, originating close to the base of the 
radioles. Radioles ciliated and not connected 
by web-like membrane. Ventral paired palps 
vascularized, smooth and with skeleton: palps, 
thicker than radioles but similar in length. 
Prostomium bluntly rounded with pair of 
pigment eye spots, visible only after removal 
of tentacular crown. Perisiornial collar well 
developed ventrally, triangular with rounded 
apex: collar developed laterally as narrow rim 
and present dorsally as two small lappet,. 
Setigcr 1 with notosctac only, subsequent 
sctigers with noto- and neuruselat*. Notosetac 
Of two kinds, basically broad bladcd capillaries 
and hastate setae: thoracic ncurosetae long 
handled hooks, abdominal ncurosetae long 
handled tincini wilh six or seven horizontal 
rows of teeth, each row with four or five 
teeth, teeth not perfect lv aligned in rows, 
fewer teeth per row towards apex. Number 
of noto- and ncurosetae per sciiffcr shown in 
Table 1 Triangular pygidium with no pygidinl 
eye spots. 

Length of holotype 5 mm. paratypes 3-4 
mm. Width of holotype 0.5 mm, paratvpes 
0.4-0.5 mm. 


\ftvhivunkia athafassia occurs in ephemeral, 
atbalassic. saline lakes adiaeeni to Coorong 
Lagoon. South Atismdia. The lakes, In which 
M itthtihiwia occurs, are not connected to the 
sea, and are characterised hy fluctuating 
salinities and varying water levels, due to 
the raising and lowering of the saline water 




Fig. la. Manayunkia athalassia n.sp. entire animal, dorsal view, b. capillary seta, c. hastate thoracic 
seta, d. thoracic neuroseta, e. abdominal neuroseta, lateral and head on view. 



Tablc I. Nitmbri of riotoxrttic and tKumsviw 
per xeiigei. 






(all uncini) 



























Abdominal \ 









tabic and to sensorial precipitation. A descrip- 
tion of the physical and biological features is 
given by Dc Deckkcr & Cieddes fl980>. The 
polyehactes collccled by Bayly (1970) were 
not found in this study 

Manayttnkia atholassia lives in translucent 
gelatinous tubes, in soft clayey carbonate sedi- 
ments which some times contain shell debris 
(ostracods. gastropods) Occasionally M. otha- 
hmia was found living in a colony of up to 
20 individuals Empty gelatinous tubes also 
were found, 

This species certainly persists in the lake 
beds while lakes are dry over the summer 
monihs. Even when the lakes arc dry some 
moisture may be trapped by hygroscopic sail 
crystals, and an occasional cover of dead 
aquatic plants such as Leptlaena sp. and 
Rupf'in sp. and. less frequently, the alea 
Lamprnthanwinn papillosum on the surface 
of the mud. However, the summer air tem- 
peratures may exceed 40°C Tn February 1979 
distilled water was added la samples of mud 
collected from dry lakes where M. athaluma 
occurred the previous season. Within one day 
active adults were seen at the wide range oI : 
salinities of 27 -05"/.. Further laboratory 
studies showed thai ir could be maintained in 
an aquarium of 82& for several months 


Manayunkia ntluthsshi differs from con- 
geners in being found in empheuieral saline 
environments isolated from the sea; the other 
species are found in marine or brackish water 
conditions. M. A/r/wV/V/u/v Bansc, 1956 col- 
lected in mangroves In Canuea in Brazil may 
experience some hyper-marine salinities in the 

interstitial walcr during low tide, but no 
salinity measurements are given. 

Mimay ttrtkict ulltahssia also differs morpho- 
logically from the other species. M, caxpfcu 
Aunenkova, l l >28 has six or seven pinnules 
per radiole and Af. spttftisa l.cidy, 1859 has 
six. radioles with nhout 60 pinnules per radiole. 
Southern (1921) iUggCSte that The arrange- 
ment of the filaments, which appear to be 
attached in groups to a short common stem, 
is an artifact and probably due to ihc contrac- 
tion of the basal membrane. However, we 
suggesL that the filaments arise from a short 
common stem as in conveners, but we have 
not examined the type. M poinds Zcnkewitsch. 
1935 has equal numbers of pinnules on both 
radioles, whereas Af. athulnsxia has three pin- 
nules on (he inner and lour on the outer 
radiole. A/, acsluarina (Bourne I S 8 3 > has 
four pinnules per radiole with the paired palps 
considerably longer than the pinnules, whereas 
in A/, uilwlussiu the palps and the pinnules are 
similar in length. 

Delailed setai counts are given only for Af. 
bravilivnsis and M. polarh. Af. hru\ilicnsis has 
more setae of all kinds on both abdominal and 
thoracic seligers than M atluilussia. In addi- 
tion Af. hnisiiiensis- has capillary setae of two 
different lengths whereas M. athnJnssui has all 
capillary setae of similar length. Af poralis 
has considerably more abdominal uncini than 
M, athalaxxut. For these reasons Af. athahiwm 
is described as a new species. The specific 
name refers to the type of saline lakes in 
which this species lives, 

Only three geuera of the subfamily Fabij- 
cinae have been recorded from Australia: 
Desthiuonu. Fabn'cia and Oriopsls (Day & 
Hurcbimis 197V). Hartmann-Schroeder & 
Harunann (1979) have described an Oriopsis 
sp. and Fabrtcmac gen. and sp. indct. from 
Port Hedlatid. Western Australia Manayunkia 
ar/inhnshi is ihe first record of this genus 
from Australia, but several other species of 
the subfamily Fahrieinae are present In Aus- 
tralia and shortly will be described by JJui- 
chines. Earlier, general marine collections 
overlooked the Fabrieinac because they are 
small and often have specialised habitat 


We thank Anna Murray tor the artwork, 
and Karl Ranse for his comments on the 
eencric status of the specimens. 




ANNtNKOVA, N. (1928) Oeber die pontokaspis- 
chen Polychaeten. 2. Die Gattungen Hypaniola, 
Parky pania, Fabricia and Manayunkia. Ann. 
Mus. Zoo}. Leningrad 30, 13-20. 

Banse, K. (1956) Beitrage zur Kenntnis der 
Gattungen Fabricia, Manayunkia und Fubriciolu 
(Sabeliidae, Polvchaeta). Zool. Jb, 84, 

Bayly, L A. E. (1970) Further studies on some 
saline lakes in south-east Australia. Aust. J 
Mar. Freshwat. Res. 21, 117-29. 

& Williams, W. D. (1966) Chemical and 

biological studies on some saline lakes of south- 
east Australia. Ibid. 17, 177-223 

Bourne, A. G. (1883) On Haplobranchus, a new 
genus of Capitobranchiatc Annelids. Q. J. 
microsc. Sci. 23, 168-76. 

Day, J. H. & Hutchings, P. A. (1979) An anno- 
tated check-list of Australian and New Zealand 
Polvchaeta, Archiannelida and Myzostomi'da. 
Rec. Aust. Mus. 32, 80-161. 

De Deckkkr, P. & Geddes. M. C. (1980) The 
seasonal fauna of ephemeral saline lakes near 
the Coorong Lagoon. South Australia. Aust. J. 
Mar. Freshwat. Res. 31, 677-99. 

Geddes, M. C. (1976) Seasonal fauna of some 
ephemeral saline waters in western Victoria with 
particular reference to Parartemia zietziana 
Sayce (Crustaceat: Anostraca). Ibid. 27, 1-22. 

Hartmann-Schroeder, G. &. Hartmann, G, 
(1979) Zur Kenntnis des Eulilorals der aus- 
tralischen Kiisten unter besonderer Benicksich- 
tigung der Polychaeten und Ostracoden. (Teil 
2 und Teil 3). Mitt Hamb. Zool. Mus. Inst. 
76. 75-218. 

Leidy, J. (1 859 ) Manayunkia speeiosa. Proc. 
Acad. nat. Sci. Pit Had. 10, 90. 

Southern, R. (1921) Polychaeta of the Chilka 
Lake and also of fresh and brackish water in 
other parts of India. Mem. Indian Mus. 6, 

Williams, W. D. (1978) Limnology of Victorian 
salt lakes, Australia. Verb. Internal Verein 
LimnoL 20, 1165-74. 

Zi nkewitsch. L. (1935) Ober das Vorkommen 
der Brackwasserpolychaele Manayunkia [M. 
Polaris n.sp.) an der Murmankusle". Zool. An:,. 
109, 195-203. 


by A. 7. Butler & M. 7. Keough 


A diving survey was conducted in January 1980 at 43 sites from Port Broughton in Spencer Gulf to 
Ceduna in the Great Australian Bight, to observe the distribution of the bivalve Pinna bicolor 
Gmelin, its density, habitat-types and associated species. Earlier records from Investigator Strait, 
Gulf St Vincent and Spencer Gulf are also reported. At 11 sites samples were taken to determine 
distributions of shell length, counts of growth checks and gonad states. 


by A. J. Butler* & M. J. Kuouuu* 


Buuifl, A. .1. ft KfcOUGii. TVI . f. (1081) DiMpbulion of Pinna bnalor Gmelin (MoIIuaca' 

Hivulviu) in South Anstuilu, wnh observations on recruitment. Tratis\ R Soc. S. Alltf. 

rifft), 29*}©, t2 June, IS*L 

A Jiving survey was conducted in January 1980 at 43 sites from Port Broughton in 

Spencer Gulf to Ceduna in the Great AltoiMHSin Bight, to observe the dnrtribUtioq of the 

bivalve Pinna bicnlia Gmelin, Its density, habilal-types and associated species, fc&rher rC£0t<fc 

from IimaJigBter StraiL Gull* Si Vincent and Spencer Gulf Are atso reported. At ll sites 

samples were taken 10 determine distributions- of shell length, counts of growth checks and 

gonad -(talcs. 

Although P. bicofor is widespread in suitable habitats Throughout South Australia, lis 
distribution is patchy 1 on Lirge and small nodes. Recruitment is shown to vary m space and 
lime and the significance of Jus in the ecology of the specie* is discussed. 


I lie ecology of Pinna biColor Gmelin is of 
intrinsic and practical interest ( Buller & 
Krvwsicr 1979) and wc have been studying 
both the population ecology of the bivalve 
(Butlw & Brewster 1979) and the cpibiota on 
its shells (Kay&Koough 1981,Keougb 1981') 
at a few sites in Gulf St Vincent. However, 
there is no systematically collected informa- 
tion about the distribution and habitat-types of 
this species throughout the rest of the State. 

It is well known lhat the 'recruitment' of 
many marine organisms, especially those with 
pelagic larvae, is variable in both space and 
lime. By 'recruit menf we mean entry to the 
population a! a size such that they can be de- 
tected ot captured — in this case, seen by a 
diver. This is not the same as •settlement' from 
the plankton, because newly-settled larvae may 
die before they are detectable. Variability it) 
recruitment may be extremely important in 
the ecology of such species (e.g. Bowman & 
Lewis 1977, Keough 1 , Sutherland 1974, 
Sutherland & Karlson 1977). Although varia- 
bility in recruitment is reasonably well docu- 
mented for certain commercially important 
species (e.g. LoosanofT 1966. Andrews 1979) 
there is a paucity of pubished data about its 
occurrence in a wide variety of organisms, and 
a paucity of detail about the spatial and tem- 
poral scales of "paichincss* in recruitment. 

: Oqi.niMient ot" Zoology, University of Adelaide. 
|iu\ 498 G.KO.. Adelaide. S. Aust. 5001. 

1 Keough. M. J. [1981) Dynamics o( the epifanna 
of the bivalve Pinna blcolor Gmelin. T*h t> 
thesis. University of Adelaide (submitted). 

Such data are needed for the development of 
methods for investigating patchy recruitment, 
and for the development of models, and 
management policies, for species which have 
large random variation in certain components 
of their environments 

Pinna bicolnr is such a species, and in South 
Australia it is at the southern edge of its 
tropical and subtropical range i" Rose-water 
1961 ). so it is especially interesting to know 
how its recruitment varies between places and 
times in South Australia. 

Finally, the spatial distributions of sessile, 
henthic animals may be 'patchy' in the sense 
that their density appears to be non-uniform 
and to vary nou-randomly over areas that ap- 
pear to an observer to be uniformly suitable 
(e.g. various papers in Coull 1977). Tt Is, of 
course, possible that the area is not in fact uni- 
formly suitable, but also possible that the 
animals are absent from some habitable sites, 
perhaps as a result of 'patchy' recruitment. 
Again, this phenomenon requires documenta- 
tion as a first step in its study. II is important 
to produce distribution maps showing eon- 
firmed absences as well as records of a species. 

This paper reports a survey designed to pro- 
vide jicneral observations on the distribution 
of P, bicolor in South Australia, the habitat - 
types in which U occurs, the organisms as- 
sociated with it and the regularity of its re- 
cruitment. Certain conclusions can be drawn 
from these general observations made at one 
time: detailed explanations must depend upon 
long-term observations and experimental tests 
of hypotheses. 




Pirwa hits been recorded from depths .a 
gieai as 30 m on the lloor of Gulf St Vincent 
(Shepherd & Sprieg l l >76| but ihts survey was 
Confined to mens within 2 km o( the shore flnj 
depths no more than IS m Our object was to 
visit js m:my Efc$ possible oi those sites where 
Pintiu had been reported or where ii might 
have been expected to occur from the Lype 
of bottom and degree of exposure. Ouj as- 
sumption was simply thai Pbma requires a %a£\ 
bolrom and no more than moderate wave-ex- 
posure, That part of the vSoulh Australian 
coast from the Victorian herder to Backstairs 
Passage is not discussed here. Before- this sur- 
vey we had many records from Gulf Si 
Vincent and some from .Spencer Gulf and 
Investigator Strait; those records are sum- 
marised here, and in partieuat wc discuss dau 
from cF^hr sues scored within two months ol 
the main survey iv^\ using I he same pro- 
cedures (sites t-N in Tabic 2\ The survey 
itself covered 43 sites from Port brouchton 
in Spencer Gulf to Ccduna in the Great Aus- 
tralian Bight in January, i9fto 

Subtidal sites were surveyed using SCUBA 
from an inflatable dinghy, iuterlidal sites on 
fool, At every site, position was determined by 
landmarks nnd bearings, and a mop reference 
recorded- Depth was measured fa m hy shot 
line, surface and botiom water temperatures in 
C by mercury thcrmometei, and Ihe state erf 
tide and current noted. An estimate was noted 
Qf prevailing wave and current conditions, 
indued i'rom aspeet, present conditions and 
weather, botiom topography and sunoundings 
The diver swam over :i variable (hut always 
recorded.) distance, making notes on a pre- 
paied data-state, He recorded a qualitative 
statement of bottom type and dominant matro 
biora :uni estimated percentage cover for each 
of the seagrass genera Zostera, Pnsidon'nt, 
Ainphlholis and Httfophtla, The density of P: 
hin,ior was estimated separately for each of 
two size-classes (dorso-ventral shell height 
' : -\ 1 cm, and height > 7 em in No m--'). The 
observer carried an aluminium rod I m long 
lo remind him o\' scale, and had extensive 
prior experience of measuring density using a 
I m quadrat, and so Ihe estimates can he 
taken as sufTicrenily reliable for use as an 
index of density. Ihe smaller size-class ]& 
likely to have settled within the last year 
(Outlet & Brewster I97V >; il was scored 
\cpaialcly to give an index of recruitment M 
sues where samples were not taken, He aj$0 

noted qualitatively the si/.c distribution, spatial 
distribution, morphology and epibioLi ot 
P, hinilor. Densities of the animal \\t 
groups listed below were scored on the fol- 
lowing qualitative scale- none seen, wc. com- 
mon, abundant. Theft categories were bases! 
on previous experience of Typical 1 densities for 
mese $pefcies, and have dilYcrent meanines i\.r 
each group The grgtW -.cored were ttie bi- 
valves Malleus merit fitwu\. f_ hhtmvx asper- 
rimuw and C. htlronx, gastropods of (he genu* 
Po/intces and family Muncidae. hotolhunans. 
cchir.oids. asteroids, fish and ecphnlopodv 
Under the headings Muricids, Urchins and 
Asteroids, and commonly under cithers, it was 
pMSMble to identify the parlicuar specu-s 

At certain sites, which are marked with 
asterisks in Table I, in adidtion to a|t the 
above observations, random samples oi 
/'. bfCaUfi were collected by clearing a I nv 
wide transect in a randomly-chosen direction, 
and examined in the dinghy or ashore, For 
each animal, anteroposterior shell length and 
dorsii-vcntral shell height were measured AS 
described by Butler & Brewster (1979). Scars 
left by the posterior adductor muscle ui the 
narreoii= layer Of the shell were eounted: 
these are counted with error, but the number 
of 'major* scars appears lo be an index of ggc 
(Butler & ttrewster 197*5) and in this studv 
they were always counted by the same Ob 
server. A crude index of age is also avail;;!-'!.- 
from the cpibiota of the shell, given a know 
ledge Of the bioloey ot the epibroiic species 
(KcoughM. which were recorded in this study 
on the qualitative scale k\^^\ by Hutler & 
Brewster f \ L Jl9) with notes on species-compo 
silion, The reproductive tissue in Pinna 
i Is diffusely n\M\>iv the mantle anterior lo 
Ihe posterior adductor muscle and is not al- 
ways detectable macroscopicfllly. Its develop 
meni is as yet poorly understood. In Ihis study 
i' was scored qualitatively on ihe following 
M.alc; 0, none visible; P, poorly developed, a 
Ihin layer of what appears to be gonad visible; 
M, moderately developed, Undoubtedly gonad 
tissue present obscuring underlying organs, VV 
well developed, massive gonad concealing; 
large, area of underlying organs. 

■ inilty, for *ach animal we noted sbcM 
damage due to bieakage or sponge boring, and 
the presence of suhfuhular spiucs on the ex- of the shell. Table I shows the sites 
investigated In addition to visiting widely 
spaced loeaUons. we commonly sampled 


Table 1. Sitrs inspected and estimated densities of P. bieolor, summer, 1979-80. ^Random samples 
of \\ bicotor were collected for measurements, etc. at these sites. ^Densities at these sites were mea- 
sured Using a f m- twutar. Density columns arc left blank for sires where no Pinna BW found. 
S: P. bicolor of 7/ < 7 cm. L: P. bicolor of H > 7 cm. 


Lat r S/ 





Long. °E 

No. m- 

No. m - 


1 ocation [ I - 



1-2 km w of Semaphore jetty 


34.83, 138.45 




3 km NW of Si Kilda 





2 km E of Ardrossan 






Aiiliossiin: beacon N of bulk loading jetty 





Stansbury: on mlcrlidal sand spit 


34.92 137.83 



6* - 

ndithbuiRh: site of Butler & Brewster (1979) 






Troubridgc Island: intertidally on SW side 






Wallaroo: to ISO m W from site of old jetty 






Port Broughton: o\xr 4 km travelled in channel 

and around mangrove island 




Chinaman Cicek 2 km WSW of shacks 



1 I 

Chinaman Creek 




1.5-2 km WSW of Chinaman Creek 




4(10 m WSW of Chinaman Creek 






Chinaman Creek 


32.65/137 S2 


Porl Auyuata: Playford Power Station ielty 






Port Augusta: first normal Channel marker 

S from Power Station 

7 5 





franklin Harbour: en. 400 m S of Cowcll jetty 


33.70 136.94 


Franklin Harbour 




Franklin Mainour; 500-50Q m E of jetty 



<0 0l 


Franklin Harbour: Cowcll jetty 




Franklin Harbour: (locally called 

Or Thompson's Reef) 


33.71 <136.94 





Tumbv Bay; 600 m ESE of jetty 
Tumbv Bay; jciiy 






Tumbv B;iv: 200 m off end OT jetty 


34.39. 136.12 



Tumbv Bav: 100 m off entrance to caravan park 


34.38/136.1 I 



Port Lincoln: 3(10 m E of caravan park jetty 


34.73/1 35. X9 


Port Lincoln: Kcrton Point jetty 





Port 1 incoln. 150 m off caravan park 




Port 1 incoln: 3<W ffl WSW of 1st pOU 




channel murker 
Port Lincoln n 
Coffin Bay: between Jetty and point to NW 
Coffin Bay: in channel leaving Coffin Bay 
Coffin Bav 

15 IS 



34.70 135,88 
34.63 135.-1/ 





Coffin Bay: between Goat Is. and other 

side Of fc*y _, _ _ 
Coffin Bay: point Ht entrance to Coffin Bay 
Kellidie Bay 
Kellidie Bav 




34.62 /I 35.46 



Coffin Bay "jetty 
Klliston: near jetty 
Venus Bay: near jetty 
Venus Bay 





33.04/ 134.89 
3.3.23 '134 72 


Venus Bay: downstream from 2nd upstream 
channel marker 




Veons Bay: side channel on way back to jelly 




Venus Bay: 1st upstream channel marker 
from jclty ^ , , 
Venus Bay: channel SW of dermem Island 
Venus Bay 


3V22/1 34.67 




33,22/ 134.66 



Venus Bay: 1st downstream channel marker 

from jetty 
Streak v Hay 
Streaky Bay: lOL) m inshore from 48 






50 : 

Streaky Bav: 200 m S of 1st outgoing 



channel marker, near Crawford Landing 
£ eduna jetty 





A.J. Dirrt.BR ftM. J. KtOtJGH 

several sites separated hy short distances 
within one v irca or cmbayment Nine? one ob- 
ject, of the survey was to investigate the isrnal!- 
scalc patchincss' of P. bicofot, and since on? 
object of this paper is to allow future worker 
to investigate changes over time, (he locations 
qf our sites are given in its much detail is pos- 

&(bl« in Table I. 

Distribution in Smith Atlstrulu 

fiuna bhaiiH has been recorded by us at the 
locations, for which positive densities are gwe(J 
in lahlc I and from Fishery Reach (Flcuncn 
Peninsula). American Kiver Inlet near Mustoti 
(Kangaroo Island), Rapid Buv Aldtnga Red 
and Price (all Golf St Vincent), Goose and 
Wardane, islands (Spencer Gulf). Shepherd & 
Sprigg (1976) recorded it at many sites on she 
floor of Gulf Si Vincent, and Coth.n (1961 ) 
recorded it from "Beaehport to Fremantle' 
Thus, the species is widespread o^ sheltered 
shores or in deeper water throughout the State. 
However, nolo that P. hicolor was not found 
at all of the locations in Tabic I and that ak- 
<I|?SC tO one another often t\i\J\tr (e.e. China- 
man Cre-eK, sites 10-14; Franlhn Harbour. 
sites 17-11; Fnmhy Bay. sites 22-15: Port 
Lincoln, sites 26-30). Note also that m some 
embay menis which appeared suitable fat 
Pinna, we (ound none ( keilidi;* Bay, sites 
31-3K. Venus bay. sites 40-47; Fllistnn, site 
39) or very few (Franklin H.uboiu, mIc.s 
17-21; IV,n Lincoln, sites 26-30). Tims, the 
distribution of /' hicolor appears 'pab_hv' 


\ few shell-* found at various sites were 
similar to the species which Cotton (1961) 
identified as Snbitopiniw »ir%dta. but almost 
all were typical of his Pinna <ial«hrahi. Both 
of thoe were referred by Kovewaler <196lt 
to the variable species P hicolor. I he r datum- 
.ship between shell lenglh and shell heiebt will 
be discussed elsewhere, but on preliminary 
analysis it appears not to differ si-n.hcjnllv 
amongst all the locutions sampled. At any 
lot aiion some shells bore more submhuKr 
Spines than others; these were more prominent 
m. young individuals; the typical form at all 
locations is fairly smooth-shelled <Coli..n 
IS&I, Figs 68 & 69; Rosewuter 1961, Fl 147. 
151 & 152). 

Density in different habitats 

This survey did not provide data suited fol 
powerful tests of null hypotheses about the re- 
latmaship between P. hicolor density and such 

variables as bottom type, depth, current and 

the presence of other organisms Nevertheless, 
some evlreme posibiliiies can be eliminated 
from The available tiara. Tabic I shows esti- 
mated densities al those sites where P. hicolor 
was found. P. hicolor occurred in hottom 
sediments ranemi' from very fine sand lo vcty 
coarse sand, we could detect no rebtiotiship 
beiwccn our qualitative notes on sediment iype 
and ihe presence, or density, of P hicolor, the 
prevailing* or 'average" conditions of tempera 
fur., and cut tent could only be estimated 
roughly from olir measurements and nolcs on 
a single dive, but again wc could not sec a 
possible explaniion foi the presence, density, 
OF estimated age-dislnhulion (see below) of 
P hicolor in either of these variables. 

Thero is no significant correlation between 
P. hicolor density and depth (data in Fable I 
for positive p. hicolor densities; zero densities 
included for all other sites; r — - 0.17 n *= 
51, P > ff05), nor between P. hicolor densiiy 
and the percentage cover of seagrasses (Hie 
luller transformed to angles. Rohlf & Sokal 
1969, p 129, r - 0.02, n 51. P > 05). 
Since we already had reason to suspect a nega- 
tive correlation hetwecn P, hicolor and sea- 
trasses f. unpublished datal, this was rechecked 
hj excluding data pertaining to embuymenis 
where P hicolor was rare or absent, and where 
one might argue la/vac have, for some reason, 
failed lo afflve (namely Pt Lincoln, Kellidic 
Hjv Venus Bay and Flhston) The correlation 
between P hicolor density and seagrass covet 
remained non-siumlicant (r — 0.17. n = 29, 
P > 0s . 

There Was no significant Correlation between 
P hicolor density and latitude- (For sites with 
positive densities, r 0.37, n — 22. P > 05" 
for njj si'es. r 0-17. n - 51, P > H 05 > 

The densities of other species which miglil 
COHCCI v ably i n fl uence P. hicolor, or h a \ t 
siimlai requirements, were scored on qnaltta 
live scales, The reasons for scoring dies, 
species were as follows. The bjvah\-> 
M I'lrritltdhu*. C . asfttrrinnts and C*. bilroiw 
arc ecologically similar to Pinna. Certain 
usltroids prey on P hicolor. Ihe gastropods 
l s olimt c\ spp. and probably some murictdr. ah 
thought to tin so Some fish and cephalopods 
may do so, especially on small Pinna. Hole 
thurians and eehiooids may influence the sur- 
vival ot recently-settled post -larvae. We were 
interested in any hint of association; (positive 
or negative I bei\ve.:n Tt<c presence of P. hi- 
color, particularly of recent (vermis, and Ihe 



"1 \hi * 2 AwiHhiiuu, h,t\vri_/t qualitative xeures for tin- densities of P. bfcolOT ami thtcc Sptfftafl ul 
epihmifae bivalves. Each figure in the Tttbk h the taonhrr of sites at which that combination of senrex 


Density of 

low g I mr* 
high > 1 m-- 

V-Uor2 X2 
1 contingency table 

Malleus invrttlianns 
low hi^h 


1 .38 


Chhi/nys asperrimus 
Jow high 





Jow high 

i 30 

abuiidanec of any of these species, Inspection 
of a table of ihcse scores showed no obvious 
relationships Willi P m hicoior density. Mosi of 
the data do not warrant statistical analysis, but 
the association between the bivalves P. bicahr> 
M, mericliantix, C aypern'mus and C. hi from 
was examined further. Scores for each species 
were grouped into two categories — 'low* (= 
-f rare) and 'high 1 (common + abundant) 
— and the scores lor P hicoior were tested 
for independence of each of the other species 
in three 2 >C 2 contingency tables; none was 
significant at the 5% level (Table 2). This 
is not a powerful test; it merely indicates that 
the other species are not strongly associated 
with P. htvolor. 

Gonad condition at different sites 
The Scoring of gonad development as None, 
Poorly. Moderately or Well Developed is a very 
crude index, not only because it is somewhat 
subjective, but also because the histology of 
gonad development in P, hicoior has not yet 
hcen established and related to ihcse scores. 
(I his work is in progress) Nevertheless, if 
the populations at ditlerent sites were pre- 
dominantly in ditlerent stages of the repro- 
ductive cycle, this might he expected to be 
reflected in the scores, whatever their detailed 
histological meaning, To test this, we first 
determined (or each of the 12 sites at which 
gonads were examined within the lime-period 
December 1 979-Januury 1980. the minimum 
number of adductor muscle scars -it which any 
animal was scored 'moderately' or 'well de- 
veloped' (M or W). Next, wc considered only 
animals with that number of scars or more, 
zm\ calculated the proportion of Uiem scored 
M or VV. This was done because at some sites 
the proportion of l he whole sample with de- 
veloped gonads would be depressed by the 
presence of a large number of very small pre- 
reproductive animals. Scar counts are D$ed 
here as an index of age (see below) but 
similar results arc obtained if shell length, in- 

stead of scars, is used to determine which 
animals me potential breeders. 

The results are shown in Tabic 3. The pro- 
portions scored M and W show highly signifi- 
cant heterogeneity between &rtG$ when the 
whole set is tested us a 2 X 12 contingency 
table. However, this may possibly be due to 
the length of time (more than one month) 
between sampling the first and last sites. There- 
fore, consider only the SCVCn sites sampled 
over 10 days (5-I5.LNO) and sampled sequen- 
tially from Franklin Harbour to Ceduna so 
that latitude rose and then fell during the 
period, This set is also highly significantly 

There are three pairs of sites sampled close 
together in both time and space; sites 13 & 16 
in Upper Spencer Gulf, sites 23 & 25 in 
Tumby Bay and sites 48 & 50 in Streaky Bay. 
Tested by a 2 < 2 contingency table, each of 
these pairs is homogeneous for (he proportion 
M or W. The proportions scored M and W 
( tranformed to angles t arc not significantly 
correlated with latitude (i — 03, n - 12. P 
0.05). In summary, despite our crude 
method of scoring gonad condition, it is clear 
that sites spatially far apfttti even if sampled 
at about the .same time, dilTer in the propor 
tion of animals in breeding condition. Sites 
close together in space And time receive similar 

Counts Of adductor muscle scars 

Adductor muscle scars are counted with 
error, but Buller 4 Brewster (1979) argued, 
for sile 6, Table 1. that major scars probably 
represent winter gtowth checks. This awaits 
cMiitirmation from current work on tagged 
animals, and it is also important to note that 
variables which cause a slowing of growth, 
such as temperature, food supply, breeding or 
various kinds of stress (Clark 1974), may be 
distributed differently at different sites. At one 
site, scar COUfltS are probably an index of age. 



Taw d 3. Gonad development in Pinna bieolor 
siimpled at 12 sites. Site numbers correspond to 
ItihJe I Ganads were i'isitully scored as '0'— not 
apparent; '/" — poorly developed; 'A/' -moileiaicly 
ileiilofH'd and 'W — well developed; these lnt\t> 
heen pooled into two categories here. Only animals 
ifi reproductive "age^-class, as determined by ad- 
ductor muscle scars, are included {see text). 

Site Date 





M + W 

3 lS..vii.79 


5 20.xii.79 



6 I3.xii.79 



13 24.1.80 



16 23,i.80 



21 5.i.80 


23 7.i 80 



25 6JX0 



26 9.i.80 



48 13.L80 



50 13.i.80 



51 15.i.80 



x'- : tests for homogeneity: 

Whole 2 X 12 table: 

3 1 


(P< 0.001) 

Sites 21,23,25.26,48,50,51 

x- - 


54, n (P.;oooi) 

Sites 13,16 


0.10 (P>0.05) 

Sites 23.25. 




Sues 48.51): 

K 2 




but they do nol necessarily estimate chrono- 
logical age in the same way at all sites. We 
know that the relationship between shell length 
and scar count differs between sites. For 
example, the average length of animals with 
five scars from site 3 is 31.5 cm; that from 
site 5 (which is intertidal) is 20.0 cm. Wc 
have at present no way to test whether inter- 
tidal animals produce more sears per unit time, 
or simply grow more slowly. However, from 
Ihe data available to Butler & Brewsler (1979) 
and various oservations obtained subsequently 
(Butler unpublished) it seems likely that scars 
do provide an estimate of age in years, in seve- 
ral different habitats (sites 3. 5, 6, 7 in Table 
1). We shall therefore base our interpretation 
of scar counts on this assumption. 

If the number of major adductor scars is an 
estimate of age in years, then even ihough 
scars are counted with error a comparison of 
the distributions of scars counts from two sites 
should test whether the age distribution is the 
same at the two sites. The distributions of scar 
counts at 13 sites are shown in Table 4, They 
are highly significantly heterogeneous when the 
whole set is tested, or when only the set 
sampled in January 1 980 is tested (see G-valucs 
in Table 4). Comparing pairs of sites close 
together in space and sampling date, wc find 

Taiuh 4. Frequencies of counts of adductor-muscle scats in samples of P. bicolor. 


Number of Scars 
4 5 6 7 

No. of animals 
10 11 5=12 in 


















9 8 
13 1 

13 7 

















































13 30 

3 1 







Log-likelihood ratio tests for heteunrraeitv 

Whole 13 X 13 table: G 1156 

Sites 13-51: G = 499 9 

Sites 13 & 16: G = 24 | 5 

Sites 2^ £25: G = 28.57 

Sites 48 & 50: G 14.87 
Scar classes 0-5 (0& 1 pooled) 

All sites except 26 G = 463.8 

Sites 13-51 (except 26) G - Mi.* 

Sires 13 & 16 G 18.42 

Siles 23 & 25 G - 16.73 

Siles 48 & 50 G = 7,45 















P < 0.001 
P <r oool 

0.001 < P 
0.001 < P 
P > 05 

.. 0.005 
< 0.005 

nrsTRimuioN and ritrikmiiM 01 r/v.v.i tttCOLOR 

i - 

TAIUL 5. tnji tetters altalit density of recent ny 

caiittnrnt and rcffimrUy a) revrti\tnicri\ tiv& mr- 

VlOliS 5-(t yew \. htMt •/ on L'C&lPtIS oj adductor 
munch sans \)tible <1) >>n ii\.su.mptions that stars 
represent wtHW growth checks, and \Jftf* /■'<'■<'- 
nirmttnent mortality rates and their \car-lo-xc,t> 
\uriutiotw ate mou. at all sites. See ten lor 
method*. O, no recrudtnent; ta. aamv recrudment; 
A/, major rcernttmeta. I, irrcsulat: R> rcu'd<<> 
tuhh- aha shows in/eretiees uhottt den.uiy ol 
rcceni recruitment hasetl only on tiller's cxtttnatc 
ol density of l J inna of It ^ / i ta at sites when' 
I'inn.i thnsitv VMS > 0, 1 1 ahie 1 ). 0. denstt* 
0, S, y{Htt\L, Q < density ^ 0.! m~'-: />. <htiu>, 

drn\H\ > 0.1 ttr'-. 



Rcgulaiii', Ol 


reel Liilment 



from scat 

J'lom SCUt 

from density 



of small 
































































that one is homogeneous (sites 48 & 50), the 
other two heterogeneous (sites 23 & 25. sites 

ta & is). 

The sear distributions were examined further 
f0 make inferences about recruitment. First, we 
considered 'he density of recent reeruilnmnt 
Because of the difficulty in scoring the first, 
faint scar the categories and one scar Were 
pooled, and assumed to represent 1978- 9 
recruits. The size of this class relative to the 
rest of the sample was scored for each site 
into one of three categories; 0, no animals 
with zero or one scar; minor, < p 20 
where p percentage of the sample having 
zero or one scar; major, 20 p 100. The 
results are shown in Table 5, As for most 
animals with planktouie tarvne. the density of 
recruitment is not expected to be const ont from 
year to year, cvoi it some recruitment always 

occurs; l his appears to be borne out by Tabic 
4. Fttfthet, the fluctuations in density of ie- 
eruitment do not seem m be in phase ..t all 
sites- If fluctuations in recruitment Were in 
phase, and if subsequent aye specific mortality 
rates were also the same, the conspicuous 
ovules should be in the same scar-classes at all 
TbCJ flrC not. as ts shown by 1 able 4. To 
test this, we considered only the first six xcai- 
elasscs (0-5), because for older animals wc 
have less confidence in the assumption ol a 
constant schedule of age-specific mortalities. 
The results of tests for homogeneity are shown 
m fable 4, The whole set is highly signilicantly 
heterogeneous, as is the set of nine sites sam- 
pled in January I "80. More imporlantly, two 
of the pairs of nearby sites sampled close 
together in lime are highly significantly 
heterogeneous (sites 13 & 16 and sites 23 & 

Next, wc jskcU whether recruitment appear:-, 
to be 'regular' at each site, that is, whether 
some recruits appear each year, albeit at 
varying densities. To do this, high 
were ignored because mortality may have 
reduced their numbers so much that sampling 
error becomes important. The tirst sis. sear- 
classes (0-5) were examined; recruitment nl a 
site was called irregular' if there were anv vero 
frequencies fn Lhe lirsl six classes, otherwise 
it was 'regular'. Table 5 shows the result. It 
also shows a ranking of the diver's estimate 
of density of small rintm. Note that this is 
an 'absolute' index based on the numher of 
small animals per m- of bottom whereas the 
above method is based «m the proportion of 
rhc sample which was young. Also, an animal 
7 cm in shell height may, al some sites, he 
several years old. The index based on density 
of small animals is thus of limited value, hut 
is included because it is available where satu 
pies were not collected 

the maemtude 0t the 197ft-'} Tccru.tment. 
on cither index, appear-, to have ditfered be- 
tween sites. Many have irregular rccriiinticnl 
In jane ease a pair of sites which ditfered in 
sxar~f reqtteney distributions above (sites- 23 S* 
2M also differ in their L97&-9 lecuulmenl and 
in their regularity of recruitment 

The data collected concurrently with 1ne 
samples give no suggestion ol explanations for 
these variations. Neither depth nor percentage 
cover of scagrass was significantly associated 
with regularity of recruitment usine eithei the 
Fisher CKSCJl test (assiening scagrass or depth 
values to two categories) or a two-sample runs 


\ I. BUTI IK & l\f. J KKOL'L.H 

test (ordcrifig the depth Or scagrass values ant! 
then counting runs of regularity scores); to 
both cases, P > 0.05. Similarly if the scores 
fot 1978-9 recruitment were grouped into two 
categories t 0-Hn, M ) then they were not sig- 
nificantly associated at the 5% level with either 
depth or seagrass cover mine c if her test. 

Regularity of recruitment was not signifi- 
cantly associated with latitude. This was rested 
by grouping those sites scored I and those 
scored R and eomparine, their mean latitudes 
(r n - 1.07, P > 0.2). 

The magnitude of 197S-9 recruitment 
(grouped into two categories) was not signi- 
ficantly associated with regularity of Temirr.- 
ment ( Tisher exact probability test: P = 

When the notes on associated species were 
grouped into two categories (0 - 1 rare; 
common -f abundant) and tabulated against 
the scores for 1978-9 recruitment or for 
regularity, no positive or negative associations 
were apparent On inspection, and certainly 
none was statistically significant at the 
level using Fisher exact tests. 

The density ot small animals (H £T 7 cm) 
is positively correlated with that of larger ones 
(both estimated in xitu hy the diver). For ftitCS 
where any P. Iricolor were found. Pearson's 
r - 0.4'A P < 0.0V Spearman s n 0.76, 
P < 0.001. This test was repealed, excluding 
sites 5. 7. 13. 21, 23 25. 48. 49 because the.r 
leneth-scars relalionship showed that animal of 
K — 7 cm may have more than two scars, 
and thus the density of small animals may nOI 
be an estimate of the density of recent recruit- 
ment. The conclusion remained the same 
(Pearson's r - 0.54. P < 0.05; Spearman's 
P - 0.75, P < 0.005) 

However, regularity of recruitment was nor 
significantly associated with total density either 
by a 2 x 2 contingency table with density 
classified as ^ 2m- or > 2 m- (Fjshcr exact 
test P 0.085) or by a runs- test as used 
above for depths (P > 0.05) , 


This survey has provided a dtstrihulion map 
for /' tricolor In South Australia. It is based 
on visits to many .sites apparently .suitable in 
having low wave-action with sof» bottoms 
There is a temptation to assume that when ;■ 
species has been recorded ,.r two points, it may 
Ik . \pcctcd to occur in suitable habilal.s in be 
twfen (thus, distribution maps are often 

hatched), but that seems not to be so in this 
ease. The distribution is patchy; P. tricolor 
ts absent from some apparently suitable sites. 
The patchiness occurs on a local scale; /'- 
hiculor may he found on some but not other 
dives on apparently similar bottoms wilhm 
1 km or so — eg., sites 10-14, 17-21, 22-2 \ 
26-30. Hut n is ,i|no evident on a Jailer 
scale; the specieb seems to be absent from cer- 
tain lar^e and appatcntly habitahle embay- 
ments (Kclhdtc Bay. Venus Bay, lilliston). 
thoueh present in others north and south of 
them. Note that these arc well-enclosed cm- 
hayments; perhaps the current pal terns an? 
such thai the arrival ol planklooie larvae there 
from outside is a rare event. Jf so, then by 
chance a recruitment mi$hl occur from time 
to tune and establish a temporary 'population'. 
This seems to have happened al Port Lincoln 
rsite 2b t The reason for giving the details in 
Table I is to document this patchiness: later 
workers miehi want to check the same loca- 

Organisms ate rarely if ever distributed 
evenly. Some of the unevenriess in their distn- 
hutions can be explained by i\n understanding 
of their ecology two can sny why the Un- 
occupied sites are unsuitable or have not been 
colonised) bur there may remain a compo- 
nent which cannot be explained, even reuta 
tively. with existing knowledge. The possibility 
remains that the vacant sites are unsuitable 
or inaccessible, but the reasons are not at 
present known. We shall call this 'iiiioplaincd 
pale-biuess 1 ; / 3 . hicolor provides an example. 

There is no detectable relationship between 
density of P. hicoior and sediment type, cur- 
rem regime, water depth (Table I), cover ol 
se.i^rass, or associated animal specie- 
l p.- iuIiy other ecologically similar bivalves 
(Table 2) t One might not have expected a 
competitive interaction of any importance 
bciween these bivalves (Stanley 1977), but 
perhaps their ecological similarity, or even the 
fuel that Malleus and CI usperrimnx ftsc 
P. btcolor for attachment, might have led to 
a positive association None is evident. We 
note Ed passing thai the other three specus 
of bivalves were, like P, b'h-ntnr. more oftei'i 
scored low than high in density even though 
many sites appeared suitable, and any diver 
knows that they can be abundant. These 
species, too, appear *patehy\ 

Species which are either predators '►. 
'malcni tides* (Andrewanhu 1970) might be 



expected a fvJyrv to have mosl of their in- 
fluence on younger stages of P. hkofor: slill, 
\vc note lhat they showed no association with 
the density of P, jWcefttf large enough to be 
seen by a dJvci. 

I he above is based on imprecise data 
(mostly subjective rankings) and so there is a 
possibility lhat real associations exist but were 
not delected. However, one might have ex- 
pected such associations to be at least 
noticeable in the Kind ot data we collected, 
even if they were not statistically signilicam. 
no trends, however slight, were apparent. Thus 
we conclude that with respect to the presence 
and density ot P. bu"l'>r we ore observing 
unexplained patchiness. 

The proportion of the population with 
developed gonads appears less 'patchy*. 1* 
differs between sites even considering only 
those sampled close together in time, hut 
spatially-close sites (pairs ^ sites in the some 
embaymcnt'l do not differ significantly t Table 
3J< We cannot infer that these populations are 
in ihe same phase (because we do not know, 
for example* whether a gonad scored T' is 
developing or spent) but it seems likely. 

Counts of adductor-muscle scars were 
heterogeneous between sites, including some 
nearby pairs. These scars probably represent 
check-*, in fhc growth of the animal, oilt the 
reasons for the checks, and their periods, are 
noi known with certainty. There is reason to 
assume that they represent winter growth 
checks find our interpretation of the counts 
was hased on lhat assumption On that assump- 
tion, the at!e-diMribolions of the standing 
populations of P. hhoior at different sites (in- 
cluding some nearby pairs') differ. We 
.■•aimned those distributions in more detail 
and found that the proportion of the popu- 
lation with low scar counts (recent recruits) 
.lilTers between swte including nearby OHM 
(Tabic 4h In other words, the density of 
recruit relative to thai of adults varies Fur- 
ther, the presence or absence ol* whole classes 
(interpreted as •regularity"' of recruitment) 
diflers between sites. The absence- of an entire 
class is a stringent criterion of irregularity', 
given the error in counting rings, It seems clear 
thai recruitment fluctuates from year to year, 
and the fluctuations nre not in phase at ail 
sites, nor necessarily even M nearby sites. 
T his contrasts with the proportions with deve- 
loped gonads, which were similar at nearby 

Recent recruitment and the regularity of 
recruitment wen: not correlated with dcpih, 
cover of sea grass, nor with each other. 'Regu- 
larity was not significantly associated with 
total density of P. bicotur as estimated in Nm, 

No relationships could be detected between 
recruitment and the densities of associated 
species. This is not to s..y that the associates 
haw no effects Firstly, il is possible that '.heir 
abundance is correlated with the recruitment 
of P. hiroior hut our data are too imprecise to 
denct it Secondly, they may move about. 
so lhat i heir ahundance at a particular place 
and time hears little relationship to their effects 
on P. bicofar (here at some earlier time 
Thirdly, their effects may be masked by other 
variables, especially the density of settlement 
of P hiroior. The lack of correlations in our 
data does eliminate the grossest hypotheses, 
e.g. that dense holothurians will, by killing 
newly-settled larvae, lead to sporadic recruit- 

The above discussion concerns recruits ;is .i 
proportion of the population. Actual density 
ol reeruits would be of the only 
relevant daia we have are the divers hi tfrri 
estimates of the densities of two s^e-classcs. 
The density of small animal* i( positively 
correlated with thai of large ones, considering 
all sites where P hicolor were found. How- 
ever, this mav be an artefact, because although 
an animal ol H < 7 cm at site 3 Would very 
hkclv be tinder two years old and probably 
under one fBuller £ Brewster 19731, this will 
not necessarily he true at all site* (above). 
However, if we eliminate sires where animals 
have a large scar count lor a given lengih. we 
siill find the same conclusion; density of small 
P hicolor is positively correlaled with that of 
large ones. This sCcms to be tather in con- 
trast to the Lonclusions drawn above from the 
s^ar counts. However, it is consistent with 
them if the events leading to recruitment are 
viewed as follows. 

Larvae of Pinnidae can travel lone distances 
in the plankton (R, S and A. Schehema. /nrv 
fonttn.). Thus, the fact that animals hreed at 
all sites does not guarantee lhat settlement 
(still less, recruitment) will occur at all sites. 
and those larvae which settle ai a site may not 
have been spawned there. Larvae move about 
with the currents and may well be distributed 
patchily within the water I sec reviews in 
Steele 1978) Thus, their probability of stic- 
cesslul recruitment at a given henthic bite 


V i Rll I Efi <ic M, J. K POUCH 

depends firstly on their being carried there on 
a current &J suitable strength, etc. lor settle- 
ment, jnd secondly on subtle properties of the 
bottom (which may vaty from time to time), 
the presence or absence of mobile or ephe- 
meral predators, the availability ol food fY.r 
newly settled post-larvae (which itself may 
depend upon planktonie patehiness and on the 
vagaries of the currents), .umA >o on We stress 
the term 'probability', The mere fact that 
recruitment is partly dependent on currents, 
on the shapes of land masses and channels JUrf 
on the topography of the bottom, will mean 
that sites differ consistently in the probability 
thai tarvae WiH settle. This can account For 
some very well-enclosed cmbayments 
apparently containing few or no P. hivolo^ 
and for a correlation between the densities i.t 
adults and young, but it leaves recruitment rs 
a random variable with a targe variance which, 
ow the present state o( our knowledge, WC 
cannot explain Most sites receive VanaNc, 
and some even irregular, recruitment and we 
cannot explain or predict this using depth, 
associated species, sediment type or latitude. 

Recruitment is n mnior "mystery stage" 
(Spioht 1975) in the ecology ol many species 
With pelagic larvae (e.e Andrews 1079, Milci 
kovskv 1971, Sastry 1979, Undciwood 1979). 
It is important hecause it vanes so widely 

Wc can hope to utidcrskmd the ecol.t.ey of 
the species after successful recruitment, but 
recruitment itself is ihe mam event that deter- 
mines the density of such a species at n given 
site. Whilst it may be possible in some case* 
in predict rceiuilrnent from independent 
variables which influence larval survival, our 
data (especially the very Miuifl-scalc 'patehi- 
ness ) L-ive no encouragement thai it will be 
possible in this case For purposes o( a general 
understanding oi the ecology oi the system for 
for long-term planning, if the species were a 
commercially important one) we may make 
use of a probability distribution Tor recruit- 
ment. t~ov purposes of short icrin prediction, 
the only course is 1o monitor recruitment 
directly, as done for commercial species h 
Brand el a/. 1 980, Phillips 1972. Ph. Hips A 
Hall 1978). 

The above should not be taken as an asser- 
tion that nothing can he known about uV 
recruitmcni of Pinna. Knowledge of a proba- 
bility di-trihufion can he powerful, Thov 
species which interact with Ptnnu — feed on 
its y&ung, live on its shells, etc— must be 
adapted to that probability distribution, ll ts 
a challenge to ccoloev to produce useful 
models foi systems m which many of the 
important events have probability distributors 
with laree variances. 


Anuki w.Mtriu. H. G. (1970) 'Introduction to the 
studv of animal populations ' (Mclhucn. Ion- 
don ) . 

Anuiows, j. D. (IW) Pelecypoda: Osircidae. 
in A. O. Onree & J S. Pcafse i h (fe | Uepro- 
ttnerlon of marine invertebrate^ Vol. V. Mn|- 
luscs- Pelccvpods and Le$SCr Hav-.c pp. 293- 
"Ui iAcji<krnic Prrsv London), 

Bowm*R R S. & lewis. I. R fl977J Annual 
llueluutioas jn the leeminncnt of Patella vuU 
tyttn I. 7 r?>u>. btot Kv C.K. 57, 795-815. 

Krano. A, R.. P\(.|. J, D. & Hooc.tsrr.riiK. J.' N. 
( l')Mi;. Spa settlement of Hie scallops Cfi/nntw 
opcit uhtrts (I.) arid P<Th'n nuninms ft ) q^ 
nrtifieiHl collectors. Ibid 60, 379-90V 

HmirM. A. J. & Bwivvsirk, K. I. fIV7?l Sfec 
distributions and crowd, ol I he f.m-shell Pinna 
hhnlov Gmolin I MotlMca! r.ntamelhhranchki ') 
in South AiiNlnili'a, Ausf. J. Mar (n^nuu 
AVv 30, 25-39, 

Clark, G. K. (1974) drouth lines in invertebrate 
skeletons. Ann. AVi Enrih am/ Phtnsuvv Aw 
2, 77- 99. 

Cotton, B. C. (\9(A) -South Australian Mol- 
Insca bslfcEyperfft. 1 (SA. Govt Printer. Ade- 
laide'. 1*. C M977) (EJ.) 'Ecology .if Ma. me 
Benthos.' (Univ. of Soulh r.imli.,;* i J i,*-.v 
( :oliiMihiit. s.i i 

KAY, A. M. & Krorrui. M. J. H9S1* Gte^MpDtWi 
hi i utch^ in i Ik- cpifaunal cOmniUnilte^ on 
pier piling :.n,l ihr biVuJVe Pinrm biwlor a1 
Itliltihtiryli. Ot'coloxin, Btrl In pits-.. 

I 0O5ANOM -, V. L, ( 1 0r,6 1 Him and intercity of 
settling of Ihe ovsUt Cn\.s\o\trt'o ^i^nii./i. ir. 
Lonji Mural Sound- Btoi Butt. Wi>wix Nob 

I3H, 2it it, 

MH rn.mSM. ';, ,\ i |97l » Tvpc^ ol larval 
J^vrlopnunt in murine hoilom inVerttl'Q >•'■.- 
Ihen itisiiihntidn and CCOlflgicJll siei.'tiLiM;; 
.i ic-evatu;nton. Mm\ Mai. 10. 19;.2]>. 

t'tniLii-s. H, I- it)?. 1 ) A scmi-quunuutive ..ol- 
lector ot the piiertilus Iarv;iL- ..if I be Upitlffrl 
rock iohstt.1 p,wt<tir/i\' fonafftofA cytinu) Ofcor^c 
I Oeeapocla. Palinuriaae) CUMta/funn 22, 147 

A II -ai. V C T H'^H) C&tCheS of piicruUc 
Urrvac t>n c«»t!c ;ior s n-, .i measure of nalilKill 
settlement oT Ihe wesrern iocK U>haer ftirw 
lirm <xyntt\ tk-orgc. C.S.I.K.O. Uoasion of 
i i--herifs and Oewnography, Report n.». ys 
pp. |-lj 

Koiur, h .1, & gQKkl, R R, 11969) 'Siajhrical 
Tables' (Freeman, S,in Kr,aii.iscoL 

Roffl ft Mi.*, i flftsjl t/k- rWft Pinnuluc in ihe 
in l l.v^. t .tic iH-ln-hu. Mnlluscv I, 175-217. 



Sastry, A. N. (1979) Pelecypoda (excluding 
Ostreidac). //; A. C. Giese & J. S. Pearse (Eds) 
'Reproduction of marine invertebrates. Vol. V. 
Molluscs: Pelecypods and lesser classes' pp. 
113-292 (Academic Press, London). 

Shepherd, S. A. & Sprigg, R. C. (1976) Sub- 
strate, sediments and subtidal ecology of Gulf 
St. Vincent and Investigator Strait. In 'Natural 
History of the Adelaide Region' (Eds C. R. 
Twidale, M. J. Tyler & B. P. Webb) pp. 161-74 
(R. Soc. S. Aust., Adelaide). 

Siegel, S. (1956) 'Non-parametric statistics for 
the behavioural sciences'. (McGraw Hill: New 
York ) . 

Spight, T. M. (1975) Factors extending gastro- 
pod embryonic development and their selective 
cost. Occologia, Bcrl. 21, 1-16. 

Stanley, S. M. (1977) Trends, rates, and pat- 
terns of evolution in the Bivalvia. In A. Hallam 
(Ed.) 'Patterns of Evolution as illustrated by 
the fossil record' pp. 209-50 (Elsevier, Amster- 

Steele, J. H. (1978) (Ed.) Spatial Pattern in 
Plankton Communities. (NATO Conference 
Series IV Marine Sciences 3) (Plenum, New 

Sutherland, J. P. (1974) Multiple stable points 
in natural communities. Am. Nat. 108, 859-73. 

& Karlson, R. H. (1977) Development and 

stability of the fouling community at Beaufort, 
North Carolina. Ecol. Monogr. 47, 425-46. 

Underwood, A. J. (1979) The ecology of inter- 
tidal gastropods. Adv. Mar. Biol. 16, 111-20. 


byN. S. Pledge 


Isolated teeth referable to one juvenile Propleopus oscillans have been found scattered in a cave 
deposit at Naracoorte. They have been interpreted as a lower premolar (rP 3 ); four upper molars : 
rM (described for the first time), rM , two of rM or ; a lower incisor (Hi) and four lower molars ; 
IM2, rM 3 or 4 , 1M 5 , rM5. The 'deciduous" molar M 1 is rectangular and quadritubercular, showing 
greater similarity to the M 1 of Bettongia spp. than to Hypsiprymnodon. The form of the incisor 
indicates that this tooth had a period of continued 'rootless' growth until maturity. A large humerus 
is referred tentatively to P. oscillans and body proportions are calculated. P. oscillans was an animal 
of about the same bulk as an Eastern Grey Kangaroo (Macropus giganteus), but much stockier and 
with longer legs. Apparently it inhabited dense scrub, living on a diet of herbaceous vegetation, 
occasional carrion and small animals. 


by N. S. PUMft* 

Pinter. N. S. < 1 *>S 1 > The Gfwit Rtit-Karmuioo PfaplMpUi nsrilluns < Dt- Vtf>, (Poloroidje 
Marsupmlial in South Austialia. 7><,m, R 5&£ V- -f/'sf. WIS H >, 41-47, 12 June, 1981. 
Isolated teeth referable to one juvenile Proplroptts osailans have been found scattered 
in a uave deposit at Naracnorte. Ihev have been interpreted as a lower premolar MM; 
tour npprr molars : rM* (described for the first time). rM~, two of rM 3 or l , g lower 
incisor (II, i and four iowei malar, ; IM,. r\|. ; or j. IM ,, rM. v Tfe "deciduous" molfll W 
is reetanmilin and tjuudritubercuktr. vhnui.u: &rC$|£l similarity to the M 1 of Bmbngfa ^TP 
than to l/vn^rn'mntuhv). fhe form of ihe incisor indicates that this toolti hud a period 
of continued "rootless 1 growth until maturity A large humerus is referred tentatively to 
P. t'Htllans and body proportions arc calculated. /'. Otirffkfnti *ftS Ml animal of about the 
same bulk as Ml l.asicrn Grey Kangaroo {Xhh.iopus ^^tnfrus), but much stockier Mid With 
loneer tegs. Apparently it inhabited dense scr lb. living on a diet of herbaceous \cgetalkm, 
occasional carrion and small animals. 


The kangaroos, wallabies and their allies 
(the Macropodoidca), contains many species, 
living, and losstL soma are quite common and 
others exceedingly rare Modern tuxonomtsts 
(e.g. Archer & Barlholomai 1978) split this 
large group into two families: the kangaroos 
and wallabies in the family Maeropodidae, 
and the rat kangaroos (poloroos, bcltongs, 
etc.) in the family Potoroidac. The lallcr 
family is divided into Ihe subfamily Potoroinac, 
containing potoroos and bettongs, and the 
subfamily Hypsiprymnodontinae. containing 
(be living Musky Rat-kangaToo Hypsiprym- 
tunlon niosrhtftnx Ramsay and the extinct 
PraplcvpUK specie - 

Modem // itwschatux is a small, rat-like 
animal living in restricted areas of rain forest 
in northern Queensland, li tS distinguished 
I font other kangaroos by a combination ol 
several primitive characters (Ride ]9M. 
1964). such as a simple alimentary canal, the 
presence of the hallux on the inner side of the 
foot, the presence of an upper enninc and a 
second lower incisor, and relatively simple 
hunodont molars. These features are combined 
voili a number of specializations. The most 
sinking of these are the large 'plagiaulacukP 
premolars, i.e. secant (bladed) premolars, 
which arc larger than the adjacent molars, 
with n curved and serrated cutting edge, and 
laces strengthened with a number of parallel. 
vertical ridges corresponding to the points o\' 

'■ Souih Australian MtJgettPI NoiO. Terrace. Ade- 
laide. S, AitM. 5000. 

the serrations. Such premolars arc, in fact, 
most distinctive teeth (Woods I960' 

The fossil species of Propleopwi are believed 
to be the closest known relatives of Nvp.u'p- 
rvin/trnhm. All have large secant premolars, 
although m Proplvopus these are more than 
three limes the size of the modern teeth The 
molars arc also similar. The Mountain Pigmy 
Possum, Bunamys parvus has similar premo- 
lars but a different molar structure and a jaw 
structure that precludes macropodoid affinities 
(Ride 1950. 1964) Despite the relatively 
large size and robustness of the fossil teeth 
and jaws, however. P. osciUam is known from 
only a handful of specimens. By comparison. 
kancaroos ol similar size front the same depo- 
sits often arc represented by hundreds of speci- 
mens, Nevertheless, occurrences of Preplcopux 
are widespread. Woods (I960) listed two 
specimens, a fragmentary' incisor and a near- 
complete lower jaw, and Bartholomai (1972) 
described a partial maxilla, from the Eastern 
Darling Downs. 

A further specimen has been reported from 
Wellington Caves. N.SW. (Woods I960) and 
.mother from L. Menindee (Tcdford 1955. 
1967). A second Pleistocene species P. chllla- 
pocnslv has been found in cave deposits at 
Chillagoe. north Queensland (Archer iff <tl 
1978)' Archer & Barthoiomai (1978) mention 
a specimen from a Pliocene deposit in northern 
N.S.W. This dentarv mav be conspeeific with 
a single isolated molar (Gill 1953, 1957, Rtde 
1 9M ) from a sub-basaltic Pliocene deposit 
umt Hamilion, Vic dated at 4.3 million years. 



Williams (1980) lists two additional speci- 
mens til P. QscUlam^ found recently in South 
Au-aralia ai Hookino Creek (P22425) and 
near Ml GumbiLT ( Green Waterholt-. P20XI \ ). 
Th«sa have been inspected, bin are undet 
Mtuly elsewhere so only the cheek-tooth inoa 
sure mcriW will be given in this paper. 

Tooth nomenclature follows the system ol 
Archer (l<>7Xr the total dental formula foi 
PrOpJe&pus would probably be dl^, EL dC A » 
C J ?M Mf|. The first molut Ml and pre- 
molar P2 are replaced by the erupting 1M 

New South Australian occurrence 

Rich Pleistocene bone deposits arc currently 
being excavated in caves at Naracoorle. These 
deposits have produced large and varied los- 
&il faunas ol mammals, together With associated 
amphibians, reptiles and birds (Williams 
1980), One ol these deposits partially filled 
and sealed a small cave, the Henschkc Fossil 
Gave, that was discovered by quarry opera- 
lions on the edge of Naraeoortc township in 
1969. Fxcnvations ot the deposit wctc under- 
taken hy me and continue to yield an abun- 
dance of Fossil bones. Associated charcoal has 
been collected, and preliminary radiocarbon 
results give the deposit an age of aromul 
15 000 years for the upper metre or SO Of 
sediment, (SUA- 1 40, depth 105*120 cm, 
15 000 years BP. *C 1 ' 997.7-+ 4.0; SUA 243, 


- IhOti 

years BP. flC 1 

30-75 cm. 33 800 

SW5.1 ± 3.5), 

Maeropodoids constitute more than half of 
the llcnsehke Fossil Cave assemblage, and 
potoroids are well represented. Amongst 

these specimens are several isolated tcctb. 
found over a period ot eight years, thai may 
be relerred confidently to P. oscHkms-, This 
Sfnall sample comprises om: lower premolar 
(rP.s) and seven molais. one ol which I con- 
sider to be a deciduous molar (m 1 ). An un- 
usual lower incisor, by elimination from all 
other ta\a in the assemblage, also appears to 
be r. i>\(il/nn\. Most of the specimens consist 
of enamel crowns only. They show little or no 
wear, indicating a juvenile age for t he m- 
dividual(s) represented. There is no evidence 
in ihe lorm of duplication to suggest more 
than o\k Individual, but the preservation and 
spatial distribution of the teeth might indicate 
otherwise (Fig. I), The teeth with their fo- 
iVrred serial position, area of occurrence, and 
dimensions tire listed in Tabic I 

1 \bi_i I. Mt'iiMtremtiUs of P. nwiUans teeth. 
Henschkt Fossil Cavr, 





SPCciittoi N 

O, area level 




i Mi 










i\r- to 

1 F22K15 

Aft/ 1 1 




1 M HT 

1 P22S2fi 

All X 14-^-; 






All s/!4±i- 


II. R 








PI 7692 

X3/j i 






A.?,, t r 






-\7 M 

1 (.2 





Alt) 12 




Measurements in mm. * : approximate, j Specimen 
round during cleanup of slumped sediment from n 

Knee area centred on All, previously excavated to 
level 17. Levels excavated Mere 15 un'thiek except in 
i his instance. 

rj ^J^' X g i ^ 

'. ■■ 'L I I > 


I I 

! | I 

A "3 A*. |Ae|A^Aio; All I A12 l AH 

B I <"rov-..cclion Of deposit. HfflfldChice IosmI Cave, Naraeorlc. showing distribution of fossil 
'rvpteoptis leeih - (Tabic 2 ) <_-J4in Al indicaies projected position of dated charcoal sample 
SUA- 1 4u f>330M years b.p. *C" — «7, £ 4.0). 


The check-teeth of Pr&pi&Hn& have been 
described adequately by Woods (1960), Bar- 
Iholomuj (1972) and Archer vi «L M l >7SK 
and will he discussed here only in general 
terms. One of the notable features about ibe 
leeth is the very slight dilTcrenee in cJlatflCfWfl 
that distinguish upper from lower, and indeed 
left from right, molars. They have roughly 
the saute proportions in both upper and lower 
molars, and differ only in minute features 
such as a small lingua! ridge coming 
forwards from the hypocone and a broad an- 
terior cingtilum on upper teeth, both absent on 
lower molars. These teeth bear a superficial 
resemblance to the deciduous molar M* Of 
some short-faced kangaroos, Sthetutrn\ spp, 
(sensu lato), in which the lopbs are not as 
well-developed as in M- r \ However, they are 
distinguished by different development of the 
midlink (mainly on the protoloph in Sth?nn- 
rns) and of the eresl joining the paracone and 
metaeone (straight and more vertical in P. 
owillanx). The M a o\' SttfWUKKS k also less 
rectangular than are the molars Bfl P. osdflans. 

One toolh (P22730) is considered here to 
represent an M' of Propleapuw This tooth is 
quadnlubcreular, almost rectangular and 
slightly longer than wide It resembles the other 
molars in general form, but is smaller and 
relatively shorter, and cannot bo matched with 
any described toolh. To some extent u also 
resembles uppeT molars (e.g. VI- and M :; ) of 
the Koala. Ph(t\cohtrrti^ rmem<v t which dif- 
fer ffl being selenodout and less rectangular. 

While the molar teeth of P. vstithfH'9 are all 
similar (Fie,. has been possible to iden- 
i if y iwo of them with some certainty as last 
lower molars (VI:.) by the reduced size of 
Ihe talonid, the posterior half o( the lower 
molar (Woods [9601. This is a feature Of 
mpnv marsupials. It WAS Ihus possible to cheek 
Whether this tooth [PAWS6) was the last 
upper molar (M*l of P. osrilhinv by testing 
the occlusion rviwcen it and the lower molars 
Tliere wa>, m fact. no possible match, because 
of the great size discrepancy, 1 therefore rule 
out the possibility that the toolh P227.V» fe 
a barely erupted M". which tooth is yet un- 

The specimen consists only of the enamel 
crown of the tooth, and is slightly worn on 
the cusps and crests. In the Heusehkc Fbs$il 
Cave this son of preservation is typical of 
deciduous teeth, and those barely erupted 
(eclh of Juvenile individuals where the roots 

ami dentine apparently have not been fully 
calcified, allowing 'hern tn roL away. 

1 conclude that the Looth is an \\\ despite 
the fad fhir it differs so greatly from the W 
of HypyprymnotioH (Ride 1?61), which is a 
rather irregular three-cussed tooth with a 
poorly developed hypocone This may imply a 
gtfcaUir systematic separation from ttVptipryftl' 
ntnion than is currently accepted (e.e Bar- 
tholomai l l »72). The condition of the tooth 
is much more like that of M 1 in Btttott.L'ia, 
where it is small and somewhat triangular but 
definitely quadri-tubercular. TM* agrees with 
the observations of [tartholomai (1*^721 on 
the permanent molars of Propleypus. 

One of the major problems of "cave 
palaeontology" is ihe ever present risk of re- 
working of the fossifs as they are moved 
piecemeal from the entrance to their final 
resting place. Some ol : the clangers are de- 
scribed by Archer 0^74). in the present c.^e. 
despite trie considerable lateral and vertical 
distribution of the specimens, there is no real 
evidence for more than one individual and, 
if P22736 is an M\ the animal was a juvenile. 
This favours the interpretation that P22S1o is 
a lower incisor o\' a juvenile P, o\ci!l<mx and 
explains the difference from the only figured 
specimen (in OM F3302* Woods <|9(S0i and 
Bartbolornai 1 1972 t have remarked upon the 
unique pattern of wear of h This is not 
readily evident in P22816 due to its youth, 
but a similar wear profile (especially at the 
tip) may be seen and the enamel pattern 
corresponds in its ventrolateral distribution to 
that in P. oxcijfans (WooA i960) The enamel 
is broken oil short with the tooth, and stain- 
ing indicates I hat only half the specimen was 
es posed i(i the jaw. These features and the 
lapcriug form ol the tooth (which is smaller 
in both diameters than F3302) Suggest that it 
undergoes considerable open-tooted growth 
during ontogeny. 

Dimensions of known specimens of Prop- 
/raptis teeth arc given in Table 2. While the 
Naracoortc teeth (Table 2) are shghtlv longer 
and narrower where comparable, they clearly 
lit the proportions atuJ description of P. wctl- 
/rt/K belter than P, eliltlaffptfttstj 

The natural history of Fropkupus 

The reeih. pw fragments. :ind associated 
fossils in |bc same deposits provide circum- 
stantial evidence for interpretation of aspects 
of the habitat, as well as the ecological role 
of the animal within the habitat 






Fig. 2. Isolated teeth of P. osciltatis, Hcnschke Fossil Cave. Naracoone. 



Table 2. Proplcopus spp. tooth measurements (in mm), length x width (anterior/ posterior) 

QM K6675 

NMV H159I7 


P. oscillans 

P. chillagoensis 


15.2 x 10.8 

21.] \ I i .0 


10.5 x 9.7 


M l 

ll.l x 10.3 

9.7x11.0/ 9.6 

M 1 


10.2 x 9.5/ 7.6 

M 3 


9.3 x 7.5 

QM F3302 

UCMP 51697 

NMV P15919 

UCMP 45171 

SAM P20815 

SAM P22425 

P. oscillans 

P, oscitlans 

P. chillagoensis 

Propieopus sp. 



P. oscillans 


22.9 x 6.9 


21.1 --. 1 M 





13.9 x 9.7 

*14x 10 


13.8 x 9.8 


14.7 x 7.7 


9.5 x 8.7 

* 9x 9.5 

9.6 x 9.7 

10.0 x 8.9/ 9.1 

9.8 x 9.1/ 9.2 

tlO.Ox — ,8.8 


10.8 X 9.8 

*llx 9.5 



11.3x10 /10.3 

t — x9.1/— 


ll.2x 10.3 


10.6 X 10.7 

12.2 x 11.2/10.4 


til. 5x9.2/8.3 


ll.Ox 9.6 

10. 1 x 9.0 

1 1 .2 x 10.1/ 8.5 

11.6 x 9.9/ 8.6 

f _ x 9.7 — 

* approximate, measured from Tedford (1967, Fig, 5.). 

t approximate, damaged or in alveola. (D. L. G. Williams pers. comm. 198U.) 



0.5cm 2cm 

Comparison of macropodoid right humeri in anterior aspect. A: Bettongia penicittata. 


Uy'psiprymnodon moxchatus, C: cf. Propleopus, D: Simoxthenttrux maddocki. 

1. Size: The molar teeth are as large as 
those of the Eastern Grey and Red Kangaroos, 
Macropus giganteus and Megaleia ruja. The 
only described lower jaw (the holotype) is 
also as large. J assume that P. oscillans had 
roughly the same bulk as the large kangaroos 

(Woods 1960). In body proportions and 
build, however, it differed. The jaw is robust 
and similar in shape to those of H. moschatus 
and Bettongia (Woods 1960) rather than 
Potorous. In view of the apparent close rela- 
tionship of Propleopus with Hypsiprymnodon, 


N S PI | OOl: 

we rvuv aSSUrnfl they tfid hud similar hotly 
proportions. // moschcttitx differ; from the 
potoroines in having relatively long torelimbs 
(Woods I960), apparently n primitive feature 

retained because of some advantage m its 
dense brush habitat. This in fl 
fmni the other rat kangaroos is well shown 
\n Troughlon (1173, pi. XT ♦. I calculated the 
approximate limb-bone ratios tor H tuowhtt- 
U8* using the specimen QM JM27s>9. ,k fol- 

humerus • radius : femur j tihiu — I ; 1.17 
I 1.57 . 1. 11. This compares with I ■ 1.19 : 
2.3 2.78 for Betro/igia pivMHitiA Gray, 
IR37: I : E.33 : L9 ! 3.07 for (he Western 
Grey Kangaroo Mtirroptf.y fultghwfut (Oes- 
marest): and 1 : 1.24 : 1-48 ; 2.0 l > foi ll "' 
cxtith.r shortfaccd kangaroo. Simoxrtutwnr, 
nuuhtnckt Wells $ Murray. 1 979, (S A M 
P17471-S2) all of which atv, or were, inhahi 
lauis of thick scrub. Only M fn/ightoxus is a 
grazer, corninc out into eras<y clearings 1o 

In (ho Henschkc Fossil Cave material then. 
i$ t largo humerus, ft is straighfer, more slen- 
der and more cylindrical than that of the 
kangaroos and potoroincs, and it has maiXcdry 
reduced deltoid and pectoral ridges and a 
shorter supinator crest. Of the marsupials 
compared with it (including specie* of Bit 
tnngitTt tfvptfpi | mnodon, Met topus, Sthemt 
fits, ihyfacmus. T/iydtcolfo, Pimu OfyrctQd I 
the fossil hone most closely resembles ih;_ 
hutMiru? ol H, mowhatHx, hut is even 
straiuhtcr and more cylindrical (Ffc 3>, 'Hie 
total length of the fogwl humeri* is estinr.iied 
at 195 mm. Assuming lhat it represents P. 
(>\<ifl,in\. and applying the Hypsiprynw,. ./.^ 
ratios, the radius length is T2H nun. femur *CW) 
nun. and libra 335 mm: measurements null 
eating an animal as bulky as a grey kangaroo 
hi|r \vn\i sfiMjter hind legs and much longer 

2. Pacnl\ 'II. c icih art relatively simple in 
fnrn., gu...lptubcrcu!ar and bunodont. Tt] i 
features are also typical of mammals such as 
Man. pies .md bears, all q( which have an 
omnivorous pi browsing vegetarian diet, a.s in 
fact have living pntoroids. Hume (r'*7S> wort- 
Kfdcfcd //, mcscha/iiK to be omnivorous, and 
Ramsay (187h) recorded that it ears 
insects, worms and tuberous tools . " a.'-J 
palm berries {Aechanfopfm t 'w'\ (Pfy, 
\f>crmo\ ,rWW/f?* i. flie retention of rela- 
tively J>mi- forelegs may reflect its methnd of 

food gathering, namely turning over scrub 
debus, and digging likv a bandicoot (Trough- 
ton 197?) The large seeanl premolars also 
seem connected with an omnivorous diet, 
possibly being used to cut Hcsh .is well js 
vegetable matter. P <><:rittans probably lived 
on sob herbaceous vegetation, carrion, invct- 
b'hr..(.;-v, and meat 

3. Jfahiiat; Hvp,\iprymn(>dnn and some of 
Ihc other potoioids hvc m dense wet scrnb 
where ihey can obtain protection ayainst pie 
dators. Although so much larger than the 
living rat-kangaroos, Pn*pl t _utpn\ may have bid 
the same need. Its presumed stocky build 
•as -uld be an advantage in thick hiush, just as 
it seems to he to die stocky Kangaroo Jsland 

i 53r00 (Af, /. tultyino\H\). The associated 
fauna adds some support to the idea. Although 
(here are a few lossd;. species present (such as 
f.T.i.vhifUiv sp. and Pntcopiodon BJ>.) that 
might suggest b more open environment, the 
abundance of brush -dwelling animals, such 
as poi, rfoines, small wallabies (mainly 
Mavwpus ritfogn'wu.s) and bandicoots, indi- 
cates the presence of moderately shrubby. 
<«p-.n forest in the Naracooiic area during the 
late Pleistocene. The browsing short -faced 
kangaroos iSitmnthctutnn- spp.) arc common 
A\u\ the cow-sized diprotodontid ZyxomaturtiS 
triln&M also is present: these are believed to 
have inhabited moderately Ihick scrub. Dt;v< 
todOrt opafum, which I consider i.o be an open 
scrub or plains unuvt.d, is rare: only fragments 
of three teeth have been recognized. Tor- 
toises (Chclodhta cf. toKgicoW 1 *) were abun- 
dant In a nc;obv swamp. The overall pftftift 
given by the lossd fauna is of an area at 
Naracoorfc more thickly forested and with 
heavier scrub or docker undcrstory. and a 
higher rainfall than today. 

thus Proplr,sp,t\ mav be seen a-, a la.i'C, 
bulky, relatively diorl-leggcd animal, living in 
dense th/cKcts and scrub, and eating sueeulem 
heibaceous vtgerarion, insects and other srn.d! 
anim-ds . 4 no* possibly carrion, tls rarity in the 
fossil record may rcllcct its strong preference 
Uw thick scnifl where remains seldom become 
fossili^eiJ Ii may be signilicant that the to ,JK 
rcpoilcd tiete represent a juvenile individual, 
one thin was perhaps less cautions than an 

I Itwnfc Off M. Archer for helpful initi.l 
diseicsion ol the teeth, the Queensland 
Mn-r.iro for providing reference casts of the 



type and figured specimens, Dr R. T. Wells 
for allowing me to measure the Mt Gambier 
(Green Waterhole) specimens, and Mr D. L. G. 
Williams for measurements of the Hookina 

Creek specimen. Dr J. K. Ling and Mr P. F. 
Aitken gave constructive criticism of the manu- 
script, which was typed by Mrs Joan Murphy. 
Jenni Thurmer drew the figures. 


Archer, M. (1974) Apparent association of bone 
and charcoal of different origin and age in cave 
deposits. Mem. Qd Mus. 17, 37-48. 

(1978) The nature of he molar-premolar 

boundary in marsupials and a ^interpretation 
of the homology of marsupial cheek leeth. Ibid. 
18, 157-164. 

Arc hlk, M. & Bartholomai, A. (197X) Tertiary 
mammals of Australia: a synoptic review, 
Alchfringa 2, 1-19. 

, & Marshall, L, G. (1978) Propleopus chil- 

lagoensis* a new Norlh Queensland species of 
extinct giant rat-kangaroo (Macronodiduc:Poio- 
roinae). Mm. Nat. Mus, Vict, 39, 55-60. 

Bartholomai, A. (1972) Some upper cheek teeth 
in Propleopus oscilUms (De Vis). Mem. Qd 
Mus. 16, 211-213, pi. 8. 

On L, K. D. (1953) Australian Tertiary marsu- 
pials. Ami. J. Set. 16, 106-108. 

(1957) The stratigraphieal occurence and 

palacoecology of some Australian Tertiary 
marsupials. Mem. Nat. Mu.s. Vict. 21, 135-203. 

HUME, 1. D. (1978) Evolution of the Macropo- 
didae digestive system. Aust. Mammalogy, 2, 


Ramsay, E. P. (1876) Description of a new- 
genus and species of Rat Kangaroo, allied to 
the genus Hypsiprymnos, proposed to be called 
H\psipr\mn<nion moschatus. Proe. I inn. Soc. 
N'.S.W. 1, 33-35. 

Rtt)L, W. I). U < 1956) The affinities of ttunamys 
parvus Broom, a fossil phalangeroid marsupial. 
Proe, Soul. Soc. Land. 127, 413-429. 

(1961) The check-teeth of llypsiprymnodon 

moschatus Ramsay 1876 <Macropodidae:Mar- 
supialia). J. R. Soc. W . Aust. 44, 53-60. 

- (1964) A review of Australian fossil mar- 
supials, ibid. 47. 97-131, 

Tedioro. R. H. (1955) Report on the extinct 
mammalian remains at Lake Menindee, New 
South Wales. Rev. S. Aust. Mus, II. 299-305. 
(1967) The fossil Macropodidac from Lake 

Menindee. New South Wales. Univ. Calif. Puhl. 
Geo}. Sci. 64, 1-156. 

TKOucmoN, E. (1973) "Furred animals of Aus- 
tralia." (Revised and abridged.) Angus & 
Roherlson: Sydney.) 

Wimiams, D. L. G. (19801 Catalogue of Pleis- 
tocene vertebrate fossils and sites in South Aus- 
tralia. Tton$. R- Soc. S. Aust. 104, 101-115. 

Woods, J. T. (1960) The genera Propleopus and 
Hypsiprxmnodon and their position in the 
MaerapOdidae. Mem. Qd Mus. 13, 199-221. 


by G. A. Crook & M. 7. Tyler 


Limnodynastes dumerili has a large, oval, dermal gland on the tibia; it is perforated with ducts at a 
density of 6/mm 2 . In horizontal section the gland appears as a series of large horizontal chambers 
filled with secretions. The ontogeny of glands in tadpoles is described. 


by G. A. Crook & M T Tyler* 


G. A. Crook & rV£, f, Trim I 1981) Structure and function of Ihc libud ftlafld tiff d«s frog LinwtulynttMes dimwrili Peters. T/vw.v. R, tffltt S*. ,-1f/vr. 105(2 1, 49-52. 12 

June, 1981. 

!.hrtnoti\rwMvs' tlunarili has a large, oval, dermal gland on the IWftfli H is performed With 
ducts at a density of 6 mm- In horizontal section ihe gtfcfld appears as a serrOl of I I j 
horizontal chambers filled with secretions. The ontogeny of glands in tadpoles is described. 

When ihc gWl secretion is expressed manually to the Surface, rats avoid the frug. In the 
:( bsL-ncc of expression the ffog is eaten hut the glands- are left untouched. Predators of the 
frog and closely leered species with similar glands am listed, 


In the Anura there arc diverse kinds of dis- 
crete dermal ghnds. Examples are the pare- 
toid glands o( many species of Bufo (Blair 
1972); the dorsal lipid glands of South Ameri- 
can phyllomedusine hylids (Blayluck ct ui 
1976); Ihe supralabial glands of Rana (Smith 
1954) and the tibial glands of some species 
of BUfo (Blair 1972) and some Australian 
froes of the genus Lfmtwdvnasfi's (Martin 
1972; Tyler 1976). 

Much attention has been devoted to pavotoicl 
glands and their secretions (Low 1972): the 
structure and possible functions of the tibial 
gland* have not been examined. Here we 
describe the gross structure and ontogeny of 
these structures in ihe southern Australian 
species Limnoi/ynastcs tlumeiili\ and investi- 
gate their possible role. 

Lifimofivnavh'S' dttmerHi is a robust species 
measuring up lo 75 mm snout to vent length 
when fully grown. It is widely distributed in 
southeastern Australia and is 3 member of the 
/ dorsal's- group of species, whose represen- 
tatives extend over the eastern and south- 
western portions of the continent (Martin 
1972). Illustration-, of the frog appear in 
Barker & Grieg (1977), Tyler (1977, ISTfrl 
and Coggpr (1979). 

Material and method* 

The specimens used in this study were col- 
lected at various localities within an SO km 
radius of Adelaide and were maintained for 
varying periods in vivaria in the University of 

J Department of Zoolotrv, University of Adefaulc. 
Box 498, G.P.O.. Adelaide, S. Aust. 5001 

Glands were obtained from freshly decere- 
brated and hpinalised animals. They were 
dissected free from the ttbia and lor histo- 
logical preparations were fixed in 10% 
hultercd formalin. Sections were cut at 7 //m 
and stained with haemaloxylin and eosin. For 
studies where glands were exposed to potential 
predators, minima! lime elapsed between the 
death of the donor and such exposuie. In six 
trials, groups o\' four rats were fasted for 4S 
hours and the entire dead frog offered, the 
entire frog minus the glands, or gland secre- 
tions wiped upon the surface of rat pellets or 
fresh meat 

Studies of ontogeny were based on tadpoles 
collected al stages 25-32 of Limbaugh & Volpe 
(1957) and reared TO a diet of boiled lc'tuce 
leaves. Samples were taken from Ibis stock at 
interval* determined by hmdlimb development. 

Gross and histological structure 

The tibial glands of Unworiynustrs .hamuli 
arc single, raised, ovoid, dermal structures 
On the dorsal surface of each tibia and dCCUpy 
mg more than 50% o( the dorsal aspect of that 
limb segment (Fig i). Dimensions of the 
gland in a 75 mm frog are 15 x II mm. Ill a 
series of nine specimens the length of the 
gland was 51-699& of the length of the tibia. 
The gland is entirely dermal and does not 
adhere to Underlying muscles- It is palpahle 
and resilient. 1 he dorsal surface ot the gland 
is perforated by Ihe apertures of mum-roiis 
d'Kts at a frequency of approximately (i/mni- 
ln some frogs the gland is pigmented with a 
metallic sheen contrasting won the remainder 
of the dorsal skin, however in the majority 
there was no such colour dilleicnliutiou. 

In transverse section (Fiji. 3) 1he gland has 
.i regular, columnar arrangement of cavities 



Fig. 1. Dorsal aspect of right hindleg of Limno- 
dynastes dumeriii showing large, protuberant 
tibial gland. 

filled with vast quantities of secretion, and 
communicating to the exterior via short, nar- 
row ducts. The stratum corneum is of 
moderate thickness, and mucus glands are 
comparable in size with those of other frog 
species. In horizontal section (Fig. 2) the 
regular shape and repetitive form of the secre- 
tory cavities is revealed. In the illustration 
the secretory contents have contracted from 
the very narrow matrix of connective tissue. 
We attribute this to the dehydration process 
during histological preparation. At higher 
magnifications a few small, circular vesicles 
could be detected within the secretory 

Fig. 2. Horizontal section of tibial gland. Dense ar 
secretions have contracted slightly. Scale bar = 
tibial gland. Note columnar arrangement of the 
secretions in the gland is fractured as a result of 
4. Transverse section of skin of tibial region of 
Note occurrence of cell nuclei in most superficia 
= 50 Mm. Fig. 5. Transverse section of skin of 
Volpe (1957). Granular glands (G) and mucus 
with the surface via open ducts (arrowed). Scale 

eas are the internal secretions of the gland. These 
100 Mm. Fig. 3. Transverse section of portion of 
internal matrix, and duct (arrowed). The smear of 
histological preparation. Scale bar = 1 mm. Fig. 
tadpole at stage 39 of Limbaugh & Volpe (1957) 
portion of stratum corneum (arrowed). Scale bar 
tibial region of tadpole at stage 43 of Limbaugh & 
glands (M) are well developed and communicate 
bar = 100 Mm. 



The gland is innervated by a branch of the 
sciatic nerve, and there is a network of blood 
vessels upon the ventral surface of the gland. 

The tibial glands first become evident in 
the tadpole at stage 42 and become more 
prominent until, at stage 46, they assume the 
oval form characteristic of those of the adult. 
At stage 39, the general adult structure of the 
skin is recognizable (Fig. 4). but the second 
layer of the stratum corneum contains cells 
with nuclei whereas in the adult no nuclei 
occur there. The stratum spongiosum is very 
thin and non-glandular. Glands are represented 
in the epidermis by aggregations of cells. 

At stage 40 the ultimate layering of skin is 
distinguishable. Glands are present in the 
stratum spongiosum and some have formed 
lumina. The evidence suggests that the glands 
develop downwards into the stratum spongio- 
sum, and continue to enlarge there. By stage 41 
the lumina of the granular glands are fully 
formed and secretion by the now comparatively 
thin inner layer of epithelial cells has begun, 
The secretion is in the form of spherical glo- 
bules. Ducts connect the glands to the surface. 
The epidermis has changed and assumed the 
adult form with enucleate stratum corneum cell 
layers. Aggregations of epidermal cells repre- 
sent incipient mucus glands. By stage 43 the 
mucus glands have developed lumina contain- 
ing mucus. The granular glands have expanded 
to occupy nearly the entire stratum spongio- 
sum (Fig. 5). 

At stage 44 granular glands in the tibial 
gland are in various developmental states. The 
glands at the centre have expanded to occupy 
the entire stratum spongiosum and are very 
tightly packed together. However, the peri- 
pheral ones are small and less dense. By 
stage 45 the mucus glands are well developed 
and the granular glands occupy the entire 
stratum spongiosum. At stage 46 the granular 
glands have increased further in area and the 
stratum spongiosum has expanded to accom- 
modate them. The glands are tightly packed 
and thus comparable with the form of the 
adult tibial gland. 

Function of tibial glands 

Because the tibial gland appears during late 
larval life, it is clearly of functional significance 
only to the adult. The potential range of func- 
tions is extensive, but protection from preda- 
tors appeared the most fruitful direction of 
investigation. Examination of the literature 
established that L. dumerili and its close rela- 
tives sharing tibial glands are normally eaten 
by a wide variety of vertebrates (Table 1). 

We fed freshly killed frogs to several verte- 
brate predators: Chelodina longicollis, Fresh- 
water Tortoise, ate tadpoles and juvenile frogs 
with no adverse effects; Notechis seutatus, 
Tiger Snake (no effect upon the snake); Anas 
platyrhynchos, Mallard (juveniles up to 40 
mm in length eaten readily) and also to 
Rattus norvegkus, rat. Rats ate every portion 

TABLE 1. Predation upon frogs of the Limnodynastes dorsalis species group 

Species of 

L. dorsalis* 

L. dumerili^ 
L. dumeriliX 

L. dumerili 

L. dumerili 



Vidpes vulpes (Red Fox) 

V. vulpes (Red Fox) 

Noteehis seutatus 

(Tiger Snake) 
Pseudeehis porphxriacus 

(Black Snake) 
Litoria raniformis 



In \% of stomachs 
investigated. 1958 \ 

In 7.7% of stomachs f 
investigated, 1959 I 

1 in stomach 

Fed to captive 

Observed swallowing 
L, dumerili on two 
occasions. Predator 
Bones of numerous 
frogs found in 
regurgitated owl 



J. D. Croft 
(pers. comm.) 

E. Worrell 

(pers. comm.) 



M. J. Tyler 

* Frog species identified as "£. dorsalis" were reclassified as a complex of species 
f Identity uncertain: captured in areas of sympatry between L, dumerili and L. te 
t Identity uncertain: locality data unknown. 

by Martin (1972). 


of the frog except the tibial glands and a small 
strip of skin joining the glands together. The 
experiment was performed six times with the 
same results: namely that rats fasted for 48 
hours would eat the entire L. dumerill with the 
sole exception of the tibial glands. However 
in the experiments in which exudate from the 
glands was expressed onto the outer surface 
of the skin, the fasted rats would not eat any 
part of the frog. Having appeared to taste the 
gland's contents they exhibited great distress, 
running around the cage, and making wiping 
motions with their paws. However, when the 
tibial glands were removed from the frog, the 
rats ate the hind limbs with the rest of the 

When food objects were coated with the 
secretion and offered to fasted rats, the items 
were rejected after initial investigation (usually 
by licking, and this activity was followed by 
extensive cleaning of the mouths and paws). 


We have not performed pharmacological or 
biochemical investigations to determine the 
identity of the glandular secretion, but we note 

that in their survey of pharmacologically 
active substances from the skin of Australian 
frogs, Roscghini et al. (1976) reported that in 
the L. dorsalis group a variety of substances 
widely distributed in other species was absent, 
with the exception of 5-Hydroxytryptamine 
(5-7 Mg/g skin). Although we are unable to 
identify the substance involved, our observa- 
tions indicate that the secretions of the tibial 
glands afford the animal a degree of potential 
protection from at least some of the animals 
likely to predate upon it if the secretions are 
released from the glands. However the efficacy 
of the mechanism is uncertain. The secretions 
can be expressed by applying lateral pressure 
to the glands, but we have failed to observe 
release under other forms of stress. 


We are indebted to Professor I. S. D. de la 
Lande for advice in the course of this investi- 
gation. We are also grateful to Marjorie Quin 
for help with histological matters. The study 
was funded by a research allocation from the 
University of Adelaide. 

Barker, J. & Grigg, G. (1977) "A Field Guide 
to Australian Frogs". (Rigby: Adelaide). 

Blair. W. F. (1972) "Evolution in the Genus 
Bujo". (Univ. of Texas Press: Austin). 

Blaycock, L. A., Ruibal, R. & Platt-Aloia, K. 
(1976) Skin structure and wiping behaviour in 
Phyllomedusine frogs. Copeia 1976, 283-295. 

Cogger, H. G. (1979) "Reptiles and Amphibians 
of Australia". (2nd Edtn) (Reed: Sydney). 

Limbaugh, B. A. & Volpe, E. P. (1957) Early de- 
velopment of the Gulf Coast toad, Bujo valli- 
ceps Wiegmann. Amer. Mus. Novit. (1842), 

Low, B. (1972) Evidence from Parotoid-GIand 
secretions. Ch. 13. In W. F. Blair (Ed.) "Evo- 
lution in the Genus Bujo". (Univ. of Texas 
Press: Austin). 


Martin, A. A. (1972) Studies in Australian Am- 
phibia: III The Limnodvnastes dorsalis com- 
plex. Ausu J. Zool, 20, 165-211. 

McIntosh, D. L. (1963) Food of the fox in the 
Canberra District. CSIRO Wild, Res. 8, 1-20. 

Roseghini, M., Erspamer, V. & Endean, R. 
(1976) Indole-, imidazole- and phenylalky- 
lamides in the skin of one hundred amphibian 
species from Australia and New Guinea. Comp. 
Biochcm. Physiol 54C, 31-43. 

Smith, M. (1954) "The British Amphibians and 
Reptiles". (Collins: London). 

Tyler, M. J. (1976) "Frogs". (Collins: Sydney). 

(1977) "Frogs of South Australia". (2nd 

Edtn) (South Australian Museum: Adelaide). 

(1978) "Amphibians of South Australia". 

Handbook of the Flora and Fauna of South 
Australia (Govt Printer: Adelaide). 



BY P. De Deckker & K. G. McKenzie 


A new Australian ostracod genus and species Bennelongia harpago (Family: Cyprididae Baird, 
1843) is described from Queensland. 


by P. Dk Dgckkf.r* & K. G. MtKi\/iP.r 


Dp DrxkKUK, p. & \ltki\yii:. K. C. (1081) Benneloiuuu, a new cyprididid ostrcicott genus 
from \nstralasii. I'xws: ft. Sov. S. Mxt> 1U&J, 53-58. 12 June, U'Sl. 

A new Australian nstracod gfefllt* and species ihtmclon^ta liffrpcigo t Family; Cypriuidae 
Btlirdj 1543] IS deseiibed from Queensland. 


The ostraeod Chliiinytlothcen titisi ralis 13i~ud\, 
[RSG was described from empty shells col 
lecled at Pcnola* South Australia. Sars l 1 S'>4 ) 
synonymi/.ed C. ttitMralis with t ypris mine 
tQtTg King, 1 SS5 aTrer examination of New 
Zealand specimens but Mullcr ( 1912) queried 
iheir congeneric status. Henry (1923) follow- 
ed Stirs' (1894) concept of the synonymy and 
Chapman (1967) laler transferred the species 
to < ypntwttis. (1971) in hi-, review of the 
palaeozooycography of freswater Osiracoda, 
referred to the ahovc species as the ^Chlanlv- 
th>t/i('cti tiu\tratis species croup", and pointed 
otlt anatomical differences between this group 
a I'd Cxpn\ and Chlamvilothectt s.s. This 
therefore indirectly suggested the need to erect 
a new yenus to include the Australian and 
New Zealand species. 

Since a collection of ostraeocls from Queens- 
land, made available to both authors by Mr 
p. Bcntlew yielded a new species belonging 
to the new genus, it was decided to describe 
it iointlv, and «t is here referred to as Benne- 
lonuiii hiirptiw* n.eeu.. n,sp. 

Systematic description 

Family ( vrnmmAt Baud, 1845 

Sublamily CYPKIDINAE Baird. 1S45 

Bi'tnul'invia n.ucn. 

f\l>c spa its: Rcrmelonw'n H&rpegQ n.sp. 

Rcnnelongia n.cen. 

n/ f i : .;no.\i*: Adult with strongly asymmetrical 

valves anteriorly: left valve beak-like an- 

terovent rally: selvage displaced inwards with 

* Department of /ouloj-'y , I'niversity Of Adelaide. 
Present Address Department of Bioe.eoLruipliv 
;.nd Geomorpholoyv. Australian National Uni- 
vctsily. P.O. Box 4. Canberra, ACM. 2600. 

r School <ii \pplieii Science. Rivcnna College o\' 
Advanced Education Wngga Wagga. N.S.W. 

inner list forming broad but short lip-like 
structure which does noi extend anlerodorsally. 
Anterovcntral area of right valve usually 
smooth-curved bul occasionally with beak-like 
tlanye. Juvenile with symmetrical valves and 
no beak-like structure; external surface cither 
deeply pitted or reticulated, often with many 
wan-like tubercles. 

Asymmetrical male maxillary palps; thora- 
copoda I 1st segment with two unequal setae 
and penultimate segment weakly divided; 
lateral lobe of hemipenis broadly boot-shaped. 

Derivation of name; From Bennclong, the 
first aboriginal to have a long association with 
the early European settlers of Australia. 

Discussion: McKciV.ic ( 1 971 ) noted ana- 
tomical differences belween the new genus 
and the genera Cypriv O. F. Mullcr, 1776. 
Cftlamvtfothrcti Saussure, 1K5S, Mocypns 
Khe, 1935 and Glohocypris Klic. 1939. A 
unique feature of the shell distinguishing 
Bvnni 3 loti[tni from the otherwise similar South 
African Cypridinae, is the broad hut shorl 
lip-like inner lisl which does not extend 

Bvtmclonviu anstmlis (Brady) and conve- 
ners are described in De Oecktier (1981). 

Bcmti-ion.vitf horpa^o u.sp. 
FIGS 1-19 
Desrwptionx: Carapace (external) oval to sub- 
rectaneular in lateral view; oval in dorsal view. 
Greatest height at about 1/3 from anterior 
where dorsal thickening of shell forms overlap 
in larger left valve. Valves asymmetrical: 
antcroventrally, left valve ll beak M -shaped and 
extends much further than right valve which 
is broadly rounded in that area. Deep con- 
cavity just posterior to *'bcak"-1ike feature of 
left valve whereas mouth region only slightly 
inflextcd in both valves. Left valve overlaps 
other in anterior region of hinge, posteriorly. 
and vcntrally where overlap is broadest. Shell 



Figs 1-6. Bennelongia harpago n.gen., n.sp. 1. Lateral view of carapace showing RV of paratype, 
P31613. 2. Dorsal view of carapace of paratype P31614. 3. LV internal of holotype, P31612 
4. RV internal of holotype, P31612. Fig. 5 a-b— Detail of anterior area of Fig. 1 (stereo pair). 
Fig. 6 a-b Detail of anterior area of Fig. 3 (stereo pair). Scale 1 — 150," for Figs 1-4, 2 = 50n 
for Figs 5 a-b, 3 = 100^ for Figs 6 a-b. LV, RV = left valve, right valve 



-_ - i • -■..-- 

500 U 

I.l's 7. Htiuulo/iGia huriniuo n.gen., n.*p. Internal 
view ol l.V paratype, P3I6I5. 

pseudopunetate with simple rimmed type nor- 
mal pore canals and with number of broad 
pustules anteriorly and posteriorly, some of 
which arc perforated by normal pores. Peri- 
phery of right valve tubcreulate along ventral 

( Internal) Inner lamella broader anteriorly 
and broadest in left valve. Inner list develops 
into broad lip antcrovcntrally in left valve in 
"beak" region and absent antcrodorsally; in 
ff0jtt ol hp is a deep gTOOVc; selvage nearly 
peripheral in posterior region. In right valve, 
inner list narrow and nearly peripheral all 
touud except anteruventrally where it is broad- 
er; anteriorly groove runs parallel to it. Sel- 
vage tuberculaie vcnirally Anteriorly, radial 
poie canals long, numerous and straight. 
Adductor scars consist of two rows of three 
sears, anterior sears being slightly broader; 
an additional smaJI scar behind and below 
bottom scar oi front row, and anterior to 
and below posterior row. The two mandibular 
scars long and uarrow, 

Anatomy: Anlenuula: (Tig, S) 7-segmcn- 
ted: length width ration of last six segments: 

Id M. A. J*- 1L1 N 1 Natatory setae 

2 • 3 ' r 1.4 1 ' 3 

slightly longer (ban all segments together* 
Penultimate segment with one small hrisile 
and lour natatory setae and dtstal set-merit 
with three natatory setae. For placement of 
other setae and bristles sec Fig. 8. 

Antenna; (Fig. 13) with lour claws: three 
r M n,.l ones all -stijjjltfy pectinate on penulti- 
mate segment ptus additional claw on last 
segment reaching tip of other three. Distal 
segment with additional short, slim pectinate 
claw. Nalatory setae reaching tips of claws. 

Mandible: (Tig, 12) epipod with five long 
and plumose Strahlen and a smaller one on 
side of plate; coxalc with seven teeth; endopod 
Willi a bristle long, narrow and smooth, (3 
bristle short, stout and tutted all over, y 
bristle almost twice length of distal scgrnenl, 
and pilose in distal half. 

Labrum: For detail sec Fig. 9. 

Rake-likc organ: (Fig, 17) five teeth and 
additional bifid one on inner side of each 

Maxillulu: (Fig. 151 epipod with 17 
Strahlen and four downwards pointing setae; 
3rd lobe with two smooth Zahnborstcm length 

ratio of palp segments ~^\ distal segment 

of palp rectangular. 

Maxilla: (Fig. 11) endopod with three 
terminal bristles, one short and two others of 
almost equal length and slightly more than 
twice length of short om\ epipod with five 
plumose Strahlen. For chactotaxy of proto- 
poditc, see Fig. 1 J. 

Thoraeopoda I: (Fig. 14) protopod coxa 
with one lone dorsodistal bristle, geniculate 
basis with one long bristle; 2nd segment with 
one long inner distal bristle; penullimale seg- 
ment divided and bearing long inner bristle at 
mid-length where it is divided and two un- 
equal inner bristles distally; distal segment 
with two short distal hristels, one on inner 
side and other on outer side. Claw long and 

Thoraeopoda II; (Fig. 18) terminal segment 
with two uncc|iial bristles; short one hook- 
shaped and half length of other. 

Furca: (Fig. 16) claws unequal; anterior 
one 1.4 x length o( other; anterior bristle hall 
length of posterior one which is 2/3 length 
of small claw. 

Furcal attachment (Fig 1$) median branch 
slightly curved: ventral and dorsal branches 
forming right angle iu\^\ hooUshaped at Up. 

Genitalia: weakly chitinous. For outline 
refer to Fig. 10. 

Colour of shell" green with beige strips in 
dorsal area. 

lenght. height 

Size : holotvpo : LV 1 02O 60* 

KV 920" 580m 

Holotync; AM P3 1 6 1 2; Paratypcs: AM 
P3lfiji'— P34615 (Australian Museum) 




Ffgs 8-12. BenncUmxta harpaga n.gen.. n..sp., holotvpe P3I6I2. 8. Anlennula; 9. Labrum; 10 Geni- 
talia; 11. Maxilla; 12. Mandible; all x 150. 



Figs. 13-19. lir/mclongia harpago n.gen.. n.sp., holotypc. P 31612. 13. Antenna, 14. Thoracopoda I, 
15. Maxillula — palps and lobes, 16. Furca, 17. Rake-likc organs, IS. Thoracopoda II, 19. F ureal 
attachment. All x 150. 



Type locality: Dam (19°16'18"S, 144°36' 
22"E) near Mt Teddy, beside McKinnons 
Creek, Lyndhurst Station, on the Kennedy 
Highway near Einasleigh, Queensland. 
Distribution: B. harpago also has been col- 
lected at the following localities in Queensland: 
L. Powlathanga, near Charters Towers; 
Cauckingburra Swamp at L. Buchanan, near 
Charters Towers; roadside swamp at Calen; 
Farm Dam, Mingela. 

Ecology: Found in fresh waters. At the type 
locality, it has been collected from the margin 
of the dam among aquatic plants at a depth 

of about 30-45 cm. No males have been 


We are grateful to Mr C. Bentley for the 
type material and Dr B. V. Timms for the 
other Queensland collections. We are thankful 
to Mrs R. Hughes who inked the original 
drawings, Mr P. G. Kempster for his help 
with the photography, and Ms C. Twang and 
Mrs J. Harden for the typing. 

The research of one of us (K.G.M.) was 
supported by A.R.G.C. Grant E7615127. 


Brady, G. S. (1886) Notes on freshwater Ento- 
mostraca from South Australia. Proc. zool. Soc. 
London, 54, 82-93. 

Chapman, M. A. (1967) Ostracoda. Jn I. A. E. 
Bayly, J. Bishop and I. D. Hiscock (Eds.) 
'An Illustrated Key to the Genera of the Crus- 
tacea of Australian Inland Waters". Aust. Soc. 
Limnol. Spec. Iss. 1, 2-7. 

De Deckker, P. (1981) Taxonomy and ecological 
notes for some Australian ostracods from inland 
waters. Trans. R. Soc. S. Aust. 

Henry, M. (1923) A monograph of the fresh- 
water Entomostraca of New South Wales. Part 
III. Ostracoda. Proc. Linn. Soc. N.S.W. 48, 

McKenzie, K. G. (1971) Palaeozoogeography of 
freshwater Ostracoda. Bull. Centre Rech. Pau 
—SNPA 5 suppl., 179-190 

Muller, G. W. (1912) Crustacea. Ostracoda. 
Das Tierreich 31, I-XXXIU, 1-434. 

Sars, G. O. (1894) Contributions to the know- 
ledge of the freshwater Entomostraca of New 
Zealand as shown by artificial hatching from 
dried mud. Fork. Vidensk. Selsk. Krist. 1894- 
5, 1-62. 



BY J. A. T.Bye 


Upper Spencer Gulf, which is defined as lying north of Lowly Point in Spencer Gulf, South 
Australia, is characterised by a salinity regime in which there is an approximate equilibrium 
between a northward advection of salt by a small mean current of about 0.1 mm/s that is necessary 
to maintain the water level constant in the presence of the annual net evaporation, and a southward 
diffusion of salt, probably by the shear effect induced by strong tidal currents. These processes have 
been studied theoretically using diffusion coefficients determined from the available salinity and net 
evaporation data. Approximate agreement with the observed mean and annual variation of salinity 
is obtained for the reported diffusion coefficients if the actual evaporation rates for upper Spencer 
Gulf are about 0.5 that of the regional pan evaporation rates. 


by J. A. T, Bye* 


bVEp J. A. T. (iMl) Exchange processes for upper Spencer Gull, South Australia Trans. 
K. Soc. S. Aitst. 105(2), 5M-6fi, 12 June. 1981. 

Upper Spencer Gulf, which is defined as lying north or Lowly Point in Spencer Gulf, 
South Australia, is characterised by a salinity regime in which ihere is an approximate equi- 
iihfium between a norlhwatd adveetion of salt by a small mean current of about 0.1 mm 's 
that is necessary lo maintain the water level constant in the presence of the annual net evapom 
lion, and a southward diffusion of salt, probably by the shear ellecl induced by strong tidal 
currents. These processes have been studied theoretically using diffusion coefficients determined 
from the available salinily and net evaporation data. Approximate agreement with the observed 
mean and annual variation of salinity is obtained for the reported diffusion coellicicnts it 
the actual evaporation rates lor upper Spencer Gulf are about 0.5 thai of the regional pan 
evaporation rates. 

The response inic scale for a dissolved substance is a function of the departure from 
equilibrium of its concentration, and for the annual salinity cycle, the time scale is about 
308 days. Substances introduced into upper Spencer Gulf at approximately its midpoint would 
have a residence time of about 180 days, and transient injections would initially disperse 
on a lintL- scale about 120 days, but this lime scale would increase as the remnant concen- 
tration of substance declines. 

A hiyher pan factor with proportionately higher diffusion coefficients would also approxi- 
mately reproduce the observed salinity patterns, and for a pan factor of 0.fi7 (as reported for 
lake Eyre) the dispersion limes would be reduced by about 16%. This ngute is the suggested 
overall accuracy for the predictions. 


Upper Spencer Gulf may be defined as 
that part oi Spencer Gulf to the north of 
Lowly Point. It contains a water volume of 
approximately 4 km 1 , and ha* a surface water 
area of about 500 km-\ and a length of about 
65 km (Fig. 1). Below Lowly Point, Spencer 
Gulf increases markedly in width, and this 
region appears as a transition region between 
a basically longitudinal hydrological regime, 
and the more general circulation patterns of 
lower Spencer Gulf (Bullock 1975). Indeed 
the longitudinal nature of upper Spencer Gulf 
allows methods of occanoyruphical analysis 
which are much simpler than in the general 
situation to be applied, and results on the 
exchange rates between upper and lower 
Spencer Gulf to be obtained. 

Physical Principles 

The dominant currents in upper Spencer 
Gulf arc tidal cut rents ( Radok l c >78), which 
normally generate sufficient turbulent mixing 
in the vertical to cause dissolved substances 
to become almost uniformly distributed in the 

; FHml£nS Institute for Atmospheric and Marine 
Sciences, Flinders University of South Australia. 
Bedford Pork, S. Anst. 5042. 

water column. Thus the concentration patterns 
can be described basically using vertically 
averaged values. The observed salinity patterns 
now suggest that a further approximate 


Fig. I. Upper Spencer Gulf, x indicates location 
of sources. 


J. A. T. BYE 

averaging may be taken for each cross-section 
of the gulf to yield an essentially longitudinal 
property distribution. An understanding of this 
simplified situation yielding information on 
exchange rates is readily obtained, and is 
presented in later sections. It should be noted 
however that there may be significant cross- 
sectional distributions associated with the in- 
flow regions of substances introduced within 
upper Spencer Gulf, as distinct from sea salt 
which has no significant internal sources. 

The turbulent motions induced by the tidal 
currents give rise to a longitudinal diffusion 
probably by a process known as the shear 
effect (Bowden 1965). The resulting co- 
efficients o( longitudinal diffusion are large, 
and tend to increase with water depth. Thus 
there is a mechanism that transports substan- 
ces in the opposite direction to the mean con- 
centration gradients within the gulf, i.e. south- 
wards; this process is present independently of 
any mean water transport. In fact a steady- 
state concentration distribution occurs when 
the southward turbulent diffusion of substance 
is balanced by the northward advection in- 
duced by the flow into upper Spencer Gulf 
necessary to compensate for the water lost by 
the net evaporation at the surface. This balance 
is the key to the understanding of the mean 
salinity distribution, or indeed the average 
concentration pattern for any substance, in- 
cluding that with a source in upper Spencer 

Thus, measurements of the salinity dis- 
tribution in upper Spencer Gulf together with 
the net evaporation rates enable calculations 
to be made of the diffusion coefficients, which 
in turn allow predictions to be made for the 
behaviour of any introduced substance. The 
theoretical formulation of these processes is 
given in the next section. 


Consider a one-dimensional (longitudinal) 
distribution of substance along Ox, then its 
conservation equation (Nihoul 1975) has the 

**» + &<*-£ **& 


in which t is time, Q(x,t) is the volume 
transport along Ox, K(x) is the (local) 
coefficient of eddy diffusion, A(x,t) is the 
cross-sectional area, P(x,t) is the net rate of 
production of substance/ unit length, c(x,t) is 

the concentration of substance, and P is a 
reference water density. The volume transport, 
Q, is given by the equation, 

§ - - b cjf + E - *> 

where b(x) is the breadth of the channel, 
E(x,t) is the evaporation rate, R(x,t) is the 
rainfall rate, and ffl is the rate of change of 

water level. This pair of equations, together 
with the boundary conditions that the con- 
centration is specified at the open end of the 
gulf (Xj) i.e. 


c (t) 

and that the flux of substance, and the volume 
transport through the top end of the gulf (x ) 
are zero, i.e. 



- Qc) x = 0, QK 

is sufficient to determine solutions for c(x,t). 
It is assumed for simplicity that K is inde- 

pendent of time, and that 



is indepen- 

dent of distance. This latter condition is ap- 
proximately valid for the progression of long 
period (.> 1 week) water level changes, 
mainly of meteorological origin, which are 
observed to propagate into the South Austra- 
lian Sea. Changes in cross-sectional area with 
time for these waves are assumed to be given 
by the relation. 




The net rate of production, P, consists of two 
parts; a production rate, p, associated with the 
introduction of substance, and a decay rate 
which is due to (a) breakdown of the sub- 
stance, d, and (b) exchange with the atmos- 
phere or sediments, s. Thus, 
p = p _ d _ s 

where d = pAxc and s = pb,uc 

in which a is a breakdown time constant, and 
fi is an exchange velocity with the atmosphere 
or sediments. 

The method of solution oi the equations 
uses a finite-difference representation with a 
resolution of 2 km, and extends over 180 km. 
approximately from the top of Spencer Gulf 
above Ft Augusta to Wallaroo. Solutions of 
the steady-state concentration equation for the 
same region have been discussed previously 
in Bye (1976). 



Determination of the diffusion coefficients 

FOi a steady-state distrihuhon with P 
the pair of governing; equations may he 
hincd !o yield fhfl relation. 



Tabi t ( EwporuUvn K/id Mtotfoll rat?$ for rV 
Aukihtn anil Waltowi) 


where S(jt) 


A £S 

bdv is the surface area ol 

the guff to the norlh of x. On substituting for 
the quantities ow the rfgW hand side of the 
equation from observed salinity data, and 
estimated net evaporation rates, estimates of 
Klx) can he obtained for at! \. Details of the 
method, logclher with cross-sectional area, 
breadA, and surface area data for upper 
Spencer Gulf are giYGJi iil Hofloway (1974), 
The results of his calculations Indicate that the 
dilliisioii coefficients increase from small values 
(— 5 m-/s) at Pt Augusta lo an approxi- 
mately constant vaiue (- 80 m-/s) soulh 
oi Lowly Point. In this study, the deduced 
variation pt lv has been approximated by the 
I unction, 

aunty i 

SOm- .-s 

-: 3.3 km 

:: 3.3 km 

The Annual Salinity Cycle 

I he purple of this section is to estimate 
the \-\c\ evaporation rates applying over upper 
Spencer Gulf such that the observed mean 
salinity distribution and annual salinity cycle 
are predicted theoretically using the diffusion 
coefficients, it is well known that pan evapora- 
tion rates obtained from shore station-- cue 
grealei than the evaporation rates from neigh- 
bouring waier bodies. For example, In a recent 
study ol (he water balance of flooded I . Eyre 
(Tet/lalT & bye 1976), »• was found thai the 
evaporation rate over the lake was approxi- 
mately 0.67 of the mean evaporation tales 
(or Woomcrii Oodnadalta and Moomhu and 
thai the proportionality factor was almost in- 
dependent of the lime oi year. No similar 
cumparisons between the evaporation rate for 
Spencer Gulf and that of neighbouring sta- 
tions are known, and hence theoretical predic- 
tions o( the salinity cycle have been made 
USJng a linear interpolation between pan 
evaporation rates (each adjusted by identical 
factors) M Pt Augusta and Koseworthy 
( Hounaul 19b I ), and rainfall rates (Anon 
1975| at Pt Augusta multiplied by a factor 
of IJ3 to allow for run-off, :\nd at WallBtTOo; 
as end points for the section (Table I). The 

PT U_ 







lion 1 





tiniii M 














I 1 ) 




1 12 


VI uv 

























Oc toper 










l_>. .ember 











1 Pt Avigastn pan evaporation 
- Koseworthy pan evaporation 

net evaporation rates were obtained by sub- 
traction, and reasonable agreement wilh the 
annual mean snlinhy profile, and the ampli- 
tude of the annual salinity cycles along the 
gulf was obtained using a proportionality fac- 
tor for the evaporation of 0.48 

The annual mean salinity profile obtained 
from these data, and the annual cycle of 
salinity at Pt Augusta and Lowly Pom!, for a 
salinity at Wallaroo o( 3K\'<, are shown in 
figures 2 and 3. They are representative of 
salinity data taken at various limes and loca- 
tions and reported in Thomas & Edmonds 
(1956), Holloway (J.974) and Anon (1980). 

[I is interesting that the volume transports 
(due io the evaporation in the north) which 
maintain 1 lie salinity distributions are ex- 
tremely small* for example the northward 
annual mean volume transport at Lowly Point 

Fig. 2. Predicted profile of annual mean salinity. 


J. A. I. RYE 


T — i 1 r 

~T i 1 n 1 t 

.G'At . t ■ 


Fig, 3. Predicted seasonal variation of salinity 
at Port Augusta and Lowly Point. 

is 8 m ; /s which corresponds lo a mean current 
of 05 mm. s. Furthermore, on dividing the 
volume transport into the voluir.e of tipper 
Spencer GuJf, wc find that a period of ap- 
proximately 15 years is requited to renew the 
water vulume completely, This time however 
has Jitttle relevance to the adjustment of the 
concentration profiles, since this adjustment 
is controlled by advection and diffusion. II 
would only he the time Scale 5 lor exchange 
of substance instanlancously at a uniform 
concentration within the gulf, The actual time 
scales for exchange of substances would be 
quite short while high transient uonequilihriiim 
concentration profiles persist, and thai 
lengthen steadily as the diffusive and advec- 
Uvc fluxes tend to their equilibrium values. 

These factors are reflected in a salinity 
variation which responds to (he seasonal 
variations in net evaporation only partially, 
with a response factor* R *- O.IK, where K 
is the ratio of the amplitude Of the observed 
salinity cycle to that of a hypothetical cycle 
In complete adjustment with the annual riel 
evaporation cycle. The corresponding lag (L) 
of the observed salinity cycle relative to the 
annttal net evaporation cycle varies between 
60 days at Vt Augusta aud 80 days at Lowly 

On approximating these results by a simple 
harmonic cycle, in which the response factor 

(R), and the lag (L) arc related to the fre- 
quency (a) oi' the cycle by the expressions, 

L J Ian 1 err and U - 1/VT+ (ffr)-: 

one finds that the time scale tor sail exchange. 
r — 300 days. 

The above results indicate that longer 
period climatic variability in net evaporation 
also may be significant in controlling the ob- 
served salinity variation. Kor example, a net 
evaporation cycle of period 5 years would 
have Ihe parameters. R —0.7 and L <•*■ 200 
days, and consequently would be attenuated 
much less than the annual cycle 

Dispersion from within upper Spencer Gull 

I heorelica! predictions for the dispersion of 
substances introduced within upper Spencer 
Gulf can be readily made for the two instances 
of a continuous constant production rate and 
an instantaneous injection of a fixed quantity 
of substance. For deliniteness the substance 
will he introduced at a location 34 km from 
Ihe head o\' Spencer Gulf (Fig. I), and 24 
km south pj Pi Augusta. 

(I) A continuous production run- of 6 kfifx 
for a conservative tracer 
The concentration pattern for a coiiset valive 
tracer (d s 0) is shown in Figure 4. 
Ihe concentration falls sharply soulh of the 
source, and is approximately constant north of 
the source with a value of 48 ppm; the Lowly 
Point coiiccnlrulion being 15 ppm. The small 
rise north i2i' 34 km mirrors the salinity dis- 
tribution since the fields are hoth sotiTCc free 
in this region. The total mass of substance in 
the steady-state equilibrium above Lowly Point 

' Ihe lime scale fr) is defined as the non-equili- 
bcinm mass of substance in upper Spencer GUil 
divided by the southward thi* of substance past 
Lowly Point Tjme scale is used rather than nun- 
constant ftmCe r may vary appreciably With t. 
The mass and tlu\ ate positive quantities fot loss. 
and negative quantifies for yain of substance. 








I , 


Fig, 4 Predicted concentration profile tor con- 
servative tracer for continuous source of h 

kg/s located 34 km from head at Spcneei 



hit> 5, Predicted concentration profiles 40 *md 
20U days &ftCf instantaneous release of 10 I ol 
conservative tracer a( location 34 kin fiom 

bead ol Spencer Gulf. 

is 90 kl, and hence its residence time in upper 
Spencer Gulf is ISO days. This residence lime 
is less than the exchange time scale for salt 
of 300 days, due to the concentration gradients 
being relatively greater than for the distribu- 
tion o\' an introduced substance than for salt, 
and leading lo a more rapid dispersion, in 
fact the residence time is controlled almost 
exclusively by lateral diffusion (ef. Table 3). 

(2) An instantaneous \ource of 10 t of a con- 
servative tracer 
the distribution pattern initially shows a 
lateral dispersion approximately symmetrically 
centred on the source. After about 150 days 
however the northward dispersion impinging 
on the head of the gulf has redistributed the 
substance in a similar manner to the steady- 
stale concentration pattern (Fig, 5). Thus at 
Pt Augusta a maximum in concentration of 
5 pph occurs after about 60 days. 


r » t , i 1 | 




i ip, h Predicted percentage ol* ins|;inhmeous re- 
lease located 34 Km from head of Spencer 
Gl4f. north of Lowly Point, as function or time 
: .ikr release, f ) indicates loss time scale 
in days. 

Fig. 7. Predicted concentration profile for frflCCt 
which exchanges with atmosphcie lor COfl 
linuous source of 6 kg/3 located 34 km from 
head of Spencer Gull 

The dispersion past Lowly Point is charac- 
terised by an initial interval of small loss prior 
to the arrival of substance by diffusion, fol- 
lowed by a loss which decreases in rapidity 
as the concentration decreases (Fig. 6). Tn 
fact, initially the loss time scale is about 120 
days, but this increases to ahout 250 days 
when only 25% of the substance remains. 

For the reasons discussed previously, the 
characterisation of the loss process by a simple 
exponential law is clearly not adequate, An 
approximate analysis indicates that the loss 
time scale (>) In fact increases exponentially 
with time, i.e. 

r — r ( C W 

in which r n and r are constants, that for the 
location of the instantaneous source have the 
approximate values, r : 110 days, and f 
350 days. Thus although the initial stage of the 
dispersion is relatively rapid, the final stage is 
highly protracted. 

(3) A continuous production rate of 6 fcgt* 
with surface exchange 
The distribution of a substance with a decay 
time constant of less than ~ 100 days would 
he expected to be changed significantly from 
that for the conservative substance, In particu- 
lar, for exchange with the atmosphere a typical 
exchange velocity, P> 1 0-"' m/S (1 laney 

1^72). thus at 34 Km the local decay time 

constant (-%) would be about 6 days. This 

short decay time constant causes a 1 arge reduc- 
tion in concentration in comparison with the 
conservative tracer (Fig. 4); the maximum 
value o\ 6 ppm now occurring at the source 



\ A T ftvi 

The above result can also be interpreted in 
terms or the temperature Held arising from n 
hsal Source with subsequent atmospheric ox- 
ehjiogc Pot exaiTlprte, Cor a ffisohage (q) of 
10 ,: mvday ;i! an excess temperature (AT) 
of 5'C. The apparent source. 


pdx ,,qAT 

where x. and V are the bounds of the source, 
and the concentrations In Figure 7 correspond 
approximately to cucss water lemperatiircs in 
I ''1 00th of L C. e.g. the maximum excess on 
the scale of averaging of the calculation (2 
km) is 0.06T, and is aligned with the source. 
In the near field however much higher tem- 
perature anomalies would occur, and the dit- 
Flisive heat flux divergence would be negli- 
gible compared with the surface exchange flu*, 
so thaf for a semi-circular temperature distri- 
bution appropriate to a source on the coast, 
one obtains. 

Tin ATe-' r 'r, ' ' r <: b ( x, 

where i is the radial co-ordinate, and r., 

(-M ■'*■'»:• h the dceuy length. The travel lime 
■ i„ " h a. where h is the thickness of 
the 'aufaec heated layer, and thus approxi- 
mate^ the near field temperature structure has 
a lerttilb scale. r ( , = t km. and OH assuming 
that h - 0.1 m, a time scale \ u = 3 hrs. 

f4) T/ Ir e()e< j &f lotti? period crWKgfS in wafer 


It is known that long period changes in sea 
level distinct from the astronomical tides occur 
along the south coast of Australia (Provis & 
Radok [§79 Js These fluctuations m sea level, 
which t\pieallv have a magnitude of 0.2 rn, 
piopngak- into Speneer Gulf, and those of long 
period (> 1 week) cause the sea levels in 
upper Spencer Gulf to change approximately 

The effect of the level changes is 1o oscillate 
the water longitudinally with a range of 2a 
* V where a is the vertical amplitude of the 
I"i.g pentad motion. The ratio - A is almost 
congou with distance, varying from about 
' 111 ft Augusta to 4000 at Wallaroo, Thwi 
for a 0/2 m. the range o\' the water particles 
i- i 2 km. and this leads to cyclic changes of 
salinity of amplitude 0.03'-. at Lowly Point 
and 0.4 m Pt Augusta. Otherwise no impor- 
tant effects on the propctty distributions are 


Several matters which follow from the dU- 
cussion of the previous sections, can now be 

( I ) The accuracy of the estimates 

The main question concerning the estimates 
i$ oi course their accuracy. The principle of 
the analysis has been to reproduce reasonably 
well the tnCUl salinity profile, and the annual 
salinity cycle, and then to predict the disper- 
sion of other substances. No data sets are 
known to exist for the distribution of other 
substances however which can provide inde- 
pendent checks on the deduced mixing para- 

In view of this situation recourse must be 
made lo a theoretical error analysis, table 2 
shows the Variation in mean annual salinity 
levels al Pt Augusta and Lowly Point that 
would occur for dilTusion coefficients and net 
evaporation rales varying from those of the 
analysis by factors of 1.50 and 0,75. The most 
important conclusion is that the mean salinity 
levels are determined by the ratio (Er-R.) . k, 
(cf. Determination of the dilTusion eoeiii- 
eieutsL thus the vigour o\' the exchange pro- 
cess is not monitored by the mean salinity 
profile. TJlC vigour however is monitored by 
the observed salinity cycle, such that for a 

Iabil 2 PrctlieteJ mean annual salaafirs inui 
tlh nitweal salinity naive for varit>a\ (F-lt) 


(I. Kl 

I -\( I OR. 75 1.00 1.50 

0.7S 45.8 4S.7 53.1 

12.0 4L4 $6.4 

K I .(Ml 43.7 45,8 50*3 

41.0 42L0 44.2 

J. 50 41 7 -Mo 45.8 

■10.(1 40.0 42.0 


Mcttiw Imviv 


r 50 1.50 48.4 43.4 43.0 41.1 

1-00 1,00 47.8 44iJ 42.7 41.4 

0,7s ujf. |7 44.4 42.6 41,4 

l The two values are respectively the mean anntiaJ 

salinity at Pt Au^iM;i and lowly Point, 
" 4 tic (actor I 00 COl I esporid--. Lo that for the main 

calculations, the other factors are multipliers Foi 

>I-R) or K. 
•* The two values are respectively the maximum 

and minimum salinities during the annual cycles. 



I'ADil- 3, P't'dUte,! rcshhUi r titttf in days for a 
iitntinwns conservative atiurCe located 34 km 
from the head of Spencer Gulf for various* 
tL-Ki an. J K 

(I Ri 

\ Arrow 








K 1.00 







l r> 

1 rftu Factor I.oo eoirL-spnruU [o dun lot ibe rmtin 
cjlcuiuuuns. i he other ruCtOrb ttfi multfptfett 
For ILK) ur K 

constant * ER > lv rhe amplitude of the sali- 
nity cycle is reduced as B-R (or K) is in- 
creased. This irenci nevertheless is not well 
marked f Tabic 1\, and hence the true solution 
for E-R and K cannoi he denned precisely 
from the available data. In the event that Ibe 
proportionality factor for the pan evaporation 
& 100 small, a reduction in the time scales for 
dispersion would occur, e.g. for a pan factor 
of 0.67 (cf, L. Eyre) I he residence lime for a 
continuous conservative source would be re- 
duced rrom ISO days to 130 days (Tahle 3). 
Similar reductions would apply for other time 
scales, and the diffusion coefficients south of 
LowJv Point would have the value I 10 m : 7s 
instead of K0 m- s. Thus it is probable thai 
the dispersion estimates have an accuracy of 
about 25 c £. with the likely bias (from con- 
siderations o( the net evaporation) being fct- 
Wfirdfi lime scales somewhat less than those 
round yjrfrvS llollowav'x estimates of the dilYu 
siou coefficients 
(2 ) Additional factors 

Wind CffCCtS have not been considered ex- 
plicitly since it is probable that the main 
agCttl of the turbulence is the tidal current; 
however a proportion of the mixing may he 
wind induced, and thus one would expect some 
seasonal variation in the diffusion coefficient** 
It is not anticipated however that individual 
storms would have a LargQ etfect on the dis- 
persion processes. Periods of calm associated 
with minima in the tidal current amplitudes 
such as occur around 'dodge tides* also max 
have a transient effect un mixing by allowing 
a stratified How regime to exist 

All these effects have their place, howeve/ 
an average corresponding io the saliniLy dis- 

tribution is perhaps that which is required 
initially In a study of i racer distributions, and 
Sato surest that the salinity distribution is a 
reasonably well deliued regime within the 
recent 25 years during which measurements 
are available (Thomas & Hdmonds 1965; 
Holloway 1974; Anon 1980) 

The special behaviour o\' tracers other than 
well vertically mixed conservative and atmos- 
pherically exchanged quantities also has not 
been considered, for example, the behaviour 
of surface and bottom seeking substances 
would be significantly different and require 
individual treatment. A comprehensive treat- 
ment of course must extend to an evaluation 
of the results of exchange of sirbstancv for 
marine life and for the atmosphere. 
{}) Proposed programmes of observation 

A fuller understanding of the dispersion pro 
cess would of cuttrsc rely on extended pro- 
grammes of observation. In particular, obser- 
vations of the aclual water balance of upper 
Spencer Ciult, including evaporation, rainfall 
and run-oll* are required as time series extend- 
ing over several years. These data could then 
be used together with observed profiles of 
salinity, and other introduced tracers to specify 
in detail the dispersion process-. In p.oticukir, 
attention should be given to the conditions 
just south of Lowly Point which control the 
response of upper Spencer Gulf to the north. 

In summary, the property distributions in 
upper Spencer Gulf are derived from an in- 
teresting balance between the net evaporation 
and lateral diffusion, and arc a eood example 
of what has been called a 'reverse estuary* 
Several similar inlets exist around the Austra- 
lian coastline (notably in Western Australia) 
for which this kind of analysts would also 
appear to be applicable 


The ideas presented in this paper have been 
stimulated by the necessity for obtaining >ome 
understanding of the dispersion regime in 
upper Spencer Gulf in the event of rhe pro- 
posed industrial development at Rcdclifl', 
which is situated 34 Km from the head of 
Spencer CJulf approximately at the source 
location used in the dispersion studies 


J. A. T. BYE 


Anon (1975) Climatic averages. S. Aust. and 
N. Terr. Dept. of Sci. & Cons. Aff. Bur. Met., 

(1980) Redcliff Project Draft Environmental 

Effects Statement Dow Chemical Co., 178 pp. 

Bowden, K. F. (1965) Horizontal mixing in the 
sea due to a shearing current. /. Fl. Mech. 21, 

Bullock, D. A. (1975) The general circulation 
of Spencer Gulf, South Australia in the period 
February to May. Trans R. Soc. S. Aust. 99, 


Bye, J. A. T. (1976) Pollution dispersal in water. 
Proc. S. Aust. Inst. Tech. Water Pollution 
Workshop, 1-13. 

Haney, R. L. (1972) Surface thermal boundary 
condition for ocean circulation models. /. Phys. 
Oc. 1, 241-248. 

Holloway, P. (1974) Determination of eddy dif- 
fusion coefficients for Northern Spencer Gulf. 
Cruise Rep. 4, Flinders Institute for Atmo- 
spheric and Marine Sciences. The Flinders Uni- 
versity of South Australia. 

Hounam, C. E. (1961) Evaporation in Australia. 
Com/n. Aust. Bur. Met. Bull. 44. 

Nihoul, J. C. J. (1975) Modelling of marine 
systems. Elsevier 272 pp. 

Provis, D. G. & Radok, R. (1979) Sea level 
oscillations along the Australian coast. Aust. J. 
Mar. Freshw. Res. 30, 295-301. 

Radok, R. (1978) Oceanography of Northern 
Spencer Gulf. A response to a brief by the 
South Australian Department for the Environ- 
ment. Dept for the Environment, S. Aust. 

Tetzlaff, G. & Bye, J. A. T. (1976) Water 
balance of Lake Eyre for the flooded period 
January 1974-June 1976. Trans R. Soc. S. Aust. 
102, 91-96. 

Thomas, I. M. & Edmonds, S. J. (1956) Chlorini- 
ties of coastal waters in South Australia. Ibid. 
79, 152-166. 



byR. J, K Jenkins, P. S. Plummer & K t C. Moriarty 


Enigmatic, subcylindrical to conical, sediment-infilled structures occurring in the Trezona 
Formation and the lower and upper parts of the Moorillah Formation of the late Precambrian of the 
central Flinders Ranges, South Australia, resemble certain fossil burrows such as Bergaueria Prantl, 
and also late Precambrian body fossil remains including the sack-shaped, soft-bodied form Ernietta 
Pflug from South West Africa (Namibia). The structures occur up to 3,300 m stratigraphically 
below the well known Ediacara assemblage. The lack of disruption or vents in their matrix 
seemingly negate the possibility that they are water escape structures, but studies of their infilling 
and literature research indicate that their true origin is almost certainly inorganic, their resemblance 
of various fossil remains being fortuitous. Their recognition emphasizes the apparent non- 
occurrence of metazoan remains below the Ediacara assemblage in the local rock record, lending 
negative support to recent theories postulating either late evoulution of the Metazoa or rapid 
diversification of animal life in the latest Precambrian. 



by R. J F, Jenkins*, P. S. Plummer* & K. C, Moriarty! 

JENKINS, R. J, F„ Plummik, I*. S. & Moriarty. K. C. < IV8I) Late Prceambrian pseudofossils 
from the Flinders Ranees, South Australia. Trans. H. Soc, S. AUSh 105(2), 67-83, 12 June, 
i ML 

Fnigmalic, subcylinditeal 10 conical, sediment-infilled structures occurring in the Trczona 
I oi malum Unci Ihe |0W*|' ;iiui upper pails of the Moonllah horriiHlion of the late Prccambr'mn 
ul ihe centra! I binders Ranges. South Australia, resemble ccilain fossil burrows such 
;is Hwgautria Prand. and also lute Prceambrian body fossil remains including the sack-shaped, 
nol'l-bodicd form htrijciht Pilug from South West Africa (Namibia). The structures occur 
up Id 3,300 m strattgraphKally below the well known Ediacaia assemblage. The lack of 
disiupliun or vents in their matrix seemingly negate the possibility that they are water escape 
Structure?, but studies ot their infilling and literature research indicate that their true origin 
is almost Certainly inorganic, their resemblance of various fossil remains being fortuitous. 
I hen recognirinn emphasizes ihe apparent non-occurrence of mctazoan remains below the 
biiacara assemblage in the local rock record, lending negative support to recent theories 
postulating cither late evolution of the Metazoa or rapid diversification of animal life in the 
latest Prceambtian. 


In 1075, a discovery a! ilunidant. peculiar 
cylindrical structures was made by Moriarty 
in the low a part ol the Moorillah 
Formation (Plummer rWTKJ in Utunoima 

Creek, within the Bunblnyunmi Range, south- 
west of Wilpena Pound in the Flinders Ranges 
(Fig. I) Ai the time he considered that the 
structures may have been biogenic. During late 
1976, in Ihe course of extensive field mapping 
ot the Ikachiiia Subgroup, Plummcr found n 
comparable structure in float within Bunycmo 
Creek, and later a second specimen only a tew 
kilometres from the original fttWl of Moriarty. 
This latter specimen was shown to Jenkins who 
noticed a radial pattern ol markings on Ihe 
base of the (short) column, and longitudinal 
strialions on its sides As such, the form 
showed a resemblance to the Cambrian to 
Jurassic trace fossil fh'ry.attrritt Pi'aUtl, and also 
to various late Prceambrian mctazoun remains 
Subsequent exploration has widened the 
known distribution of Ihe structure and con- 
firmed its constancy of general form and the 
more or less stable sedimenlologieal charac- 
teristics attending its preservation. 

Two specimens of similar structures in 
stratigraphic collections ol the University of 

r Centre for Prceambrian Rcseatch, The Univer- 
sity of Adelaide, North Terrace, Adelaide, 
S.Aust. S00T. 

I School oi l.<rth Sciences. Flindcfs tjinversue 
S.Aust. 5042. 




Wtovdpi MS 



CREEK r. i 

BLACK GAP--' % ' 



X Itdmu 

HS HoinestMd 


^-- Sealed 

_.— " Unsealed 


Fig. I- locality map of pseudofossi! occurrences 
in central Flinders Ranges, South Australia. 

Adelaide were found by M. A. Reynolds in 
1950 west of M Bunyeroo Hut" (probably Yan- 
yanna Hut on current maps) apparently within 
the Tre/ona Formation of the Urnberatana 


The Trezona Formation reaches a thickness 
of about 240 m in the Oraparinna area and 
consists of mud flake rich pelletal limestones* 














RAWN5LFY <tt Z. 

BONNE* ^-S. 

















cr to 




o Interval with /x^p 

Pseudoto&sils \ 


Interval with N 

J PseudofossilsN 

ii/ , 

- Pseudo*os&lls 


Fife. 2. Charl summan' regional stratigraphy 
of youngest precambrian succession in central 
Flinders Ranges, and showing sfrutigraphk 
intervals in which pscudofossils arc found in 
relation lo Edjaeara assemblage. Only upper 
part of Umberalana Group shown. Pound Sub- 
group after Jenkins IVtfSv). 

cryptalgal-Iaminitcs and associated stromato- 
lites, oobtic limestones and intraformational 
limestone conglomerates intcrbedded in green 
sillstoncs (Thomson ct ul. 1976). Jt is of 
shallow waier origin and comprises the upper- 
mosi unit of the Farina Subgroup, within the 
Umheratanu Group, and is of early Marinoan 
age in terms ol the local late Preeambrtan 
1ime-rock stratigraphy (Thomson 1969). It is 
succeeded by the approximately 60 m thick 
Elatinfl Fdnftation which includes the tillilic 
deposits of the third and youngest glacial phase 
of I he Adelaideun ( Mawson 1949), and is 
part of the WilJochra Subgroup of the 
Umberalana Group. 

The Brachina Subgroup (Plummet 1978) 
of ihc Wilpcna Group is a coarsening upward, 
doimnanlly clastic sequence with an average 
thickness oi 1500 tfL It rests mostly conform- 
ably though locally disconformahly on the 
I ialina Formation, Lying within the Subgioup 
approxnnalely 600 m above this glaciogcmc 
unit is a 460 m (hick sequence consisOue 
predominantly of thinly interhedded red and 
purple shales and sillstoncs. I his sequence is 
defined as the Moonllah Formation and com- 
monly interposed within it are medium to thick 
beds of purple tufTaeeous silistone which often 
display intra tormational conglomeratic tex- 
tures, and lenticular erossbedded sandstones. 
Soft-scdiinent deformation structures are ubi- 

The structures which form the subject ol 
this paper occur in two broad intervals within 
the Moorillah Formation. In Bunhinyunna 
Kmgc (Fig I ) several hundreds of" specimens 
were observed in a hroad zone spanning much 
of the basal half of the Formation. Very few 
specimens were observed near Ihc lop of ttw 
Formation in this area. Specimens were, how- 
ever, moderately numerous neat i he top of the 
Formation at Mount Chambers Creek. 100 km 
to the northeast, 


Representative specimens ate lodged m the 
collections of the University of Adelaide; 
catalogue numbers prefixed "A. I 1 ,". Studies 
were also made of a large collection of late 
Precambrian fossils obtained by Dr Hans 
Pfjlig, G lessen. West Germany, FrOflJ 'be N'iim.i 
Ciroup oi Namibia (South West Africa). 
These specimens, catalogued ' 'PI ." are ulti- 
mately to be relumed by Hr Pllug to the 
appropriate ffULtUtOTy body in Windhoek 


1 he shape of the structures in vertical 
seclion is commonly cylindrical (Fig. 1A & 
M), wilh eilher a rounded in hemispherical 
base (Fig. 4A), or a flat, U-shaped haw 
(Fig. 3F; Fig. 4G), which is often lopsided 
and with one or several ceniraJ indentations 
(Fig. 4C, L), fi & F). A snb-conieal shape is 
also occasionally displayed (Fig. ^B & C> 
The upper part frequently expands outwanK 
like the mouth of a trumpet. In section para- 
llel lo the bedding the structures display ,\ 
circular to ovoid outline I Fig. 3C k D). The 
width of the struct ores (disregarding the 
irumpetTike upper rim) averages 6 B cm. but 



vanes between O.S and 20 em. The expanded 
mouth of the trumpet may reach over 30 em, 
I heir width is frequently less than their depth. 
With some specimens roughly twice as de^p 
,is wide, rita depth may reach about 20 em. 
Often their surface is nearly smooth, but on 
many a series of fine longitudinal ridges 
and grooves c\\eiuK along the cylinder or cow^ 
Hvd iajis out on the Irumpel-Iike upper Tim. 
Occasionally these ridges and grooves con- 
tinue across the base of the structure as radial 
markings (Fig- 4F). The significance of these 
markings is questionable, however, as they are 
often indistinguishable from, and giade into 
slickensides which also occur commonly 
within the matrix. 

Jn the Moorillah Formation the structures 
arc found Within mterbedded .hales, siltstones 
and line sandstones. In the majority of speci- 
mens the baste of Ihe suucU'rc is either 
sunken down into, or in close juxtaposition 
with Ihe top of a siltsronc or iinc sandstone 
bed (Fig. AC). The upper parts of ihe *truc- 
hucs arc often enveloped by laminated shales 
which sometimes include thin heds of siltstone. 
When- KtK ^ihsiouc below the structures js 
thin it is sometimes bowed downwards and 
there is ;i tendeney for the -.hale laminae and 
thin Siltstones subjacent to Ihe sides of tfW 
structures also to be gently curved downwards 
(Fig. 3 A & 6; b'g 4C & Dt. The uppermost 
part ol (he stiuciun-s may Terminate within 
shaU-. or be truncated or capped by another 
hc(l (rf siltstone wh»eh is often bowed Up? 
v\m\K. lite capping shale or siltstone par- 
ticipated m pactiaily'tilling the structures. Rare 
specimens occur r.ior? or less entirely within 
shale it-;'. IF). 

Sections ol the structures show that they 
had a complex history of infilling and sub 
sequent compaction (Fig 4C„ D. F & Gi- 
Otlen the initial infilling consists ol fine to 
coarse sand witb nnrulai or subrounded mud 
and silt clasrs which came to rest in a aib- 
hori/onlal attitude, or sometimes m an edge- 
wise fashion | Fig 4G.I. The sand infilling may 
show rather irregular cTosional surfaces within, 
oi gOldfl upwards into lammalcd siltstone slill 
coniaining occasional clasts. It is apparent that 
cJuiine de-watering of the surrounding matrix 
appreciable compaction occurred. The sandy 
material and stlty Infill of ihe forms seems 
generally to have behaved in a more compe- 
icni manner and compacted less than the 
enveloping matrix, resulting in the observed 

downward deformation of the external sedi- 
ments below and upward deformation above 
(big. 3£). the silt laminae of much of the 
infill are frequently bowed upwards (Fig. 4C. 
D & fiL The specimens from the Tre/ona 
Formation have a slightly calcareous silly infill, 
but ol her wise show similar characteristics \o 
the material from the Moorillah Formation. 



The nature of the infill indicates that the 
structures must have initially been hollows or 
holes wkhin the sedimentation surface. These 
holes thus acted a v s traps for any coarse sedi- 
ment being curried in traclion before being 
buried by the more turbid finer sediment- A 
syngeneic origin is theteforc envisaged for 
which cither an organic origin, or production 
by inorganic precedes, jj$ possible. 

Possible Organic Origin 

Two possible organic origins could produce 
syngeneic structures similar to those described 
here. Firstly, the burial of dwelling or hiding 
burrows of aetinum-like creatures could pro- 
duce trace lossils resembling such forms as 
Hct^itirrut, or Conusuchits Lcsquercux, Alter- 
natively, the preservation of cup- or sack- 
shaped animals could yield body fossils com- 
parable to NuituitUt Germs, the "Trmetto- 
morpha" of Pllug ( IA72fl) or Haikalwu Soko- 

» 'rev < 1^70) describes the borrows of 
several living aclinians as being considerably 
deeper than wide, or sometimes cUremeW 
elongate and occasionally branched. Shinn 
(1968) shows that adjacent to such burrows 
it is common to sec a downwarpiug of the 
sediment lamination. In a study which 
Mangum (1970) made on the burrowing be- 
haviour oi' the anemone Fhyilnciis Milne 
EdW&ttfs & Haime, she slated that rhythmic 
contractions of the column musculature mani- 
pulate the fluids of the column's hydrostatic 
skeleton to push rhc pedal disc into the suh- 
strale; "afler the pedal disc reaches a hard 
substratum and sand surrounds the column 
(burrowing) ceases.'* The bower Cambrian to 
Jurassic trace fossil Berpputrib is normally 
attributed to uet'mians burrowing .'ind resting 
m ihis manner (Crimes a nL 1977). Accord- 
ing (o \lpert (1973) the depth oi' Brr^ancria 
is less than or equal to its diameter, although 
Crimes et nl. have observed deeper specimens 
in the Tower Cambrian of Spam Externally, 




some of our more shallow specimens are essen- 
tially indistinguishable from Bergaueria except 
for being generally greater in size. Sectioned 
specimens of Bergaueria from the early Cam- 
brian of Alberta, Canada (Needier Arai & 
McGugan 1968; Germs 1972a) show a 
coarser, less regular internal lamination than 
our material, and sometimes show slumping 
which is presumed to reflect decomposition of 
the inhabitant of the burrow. 

More conical examples among the present 
structures show a marked resemblance to 
another trace fossil, Conostichus, which is best 
known from the Mississippian-Pennsylvanian 
of North America and is also attributed to 
burrowing actinians (Chamberlain 1971; 
Crimes et al. 1977). Conostichus is a more or 
less conical burrow, tapering downwards, 
reaching about 1 I cm in depth and 8 cm in 
diameter (Branson 1960) and variably marked 
by transverse constrictions and longitudinal 
furrows and ridges. The sides of Conostichus 
consist frequently of several "packed" layers 
apparently formed as the anemone wallowed 
in its burrow. Where it is assumed that the 
actinian abandoned the burrow abruptly, 
"shifting sand and debris filled the void with 
cross laminae" (Chamberlain 1971). Where 
the actinian apparently migrated upward in 
order to keep pace with sedimentation, the 
centre of the burrow may be filled with trans- 
verse sand laminae forming apically-convex 
concentric cones. While the present structures 
are sufficiently similar to Conostichus to sug- 
gest that they could have a broadly similar 
biological origin, that is formed by a "poly- 
poid" organism largely buried in the substrate, 
some important differences are apparent. Our 
structures are never observed to show layers 
of sediment plastered subjacent to their walls, 
and both the even lamination or the small 
erosiona! features within the infill show that 
the origin of this material is entirely sedimen- 
tary and not the packing behind a polyp mov- 
ing upwards in its burrow. 

The second possible organic interpretation 
of the structures is that they are actual body 

fossil remains, with the integument of the 
organism represented by the surface of parting 
between the enclosing matrix and the infilling 
material. They could thus be considered to be 
preserved as either an external mould and 
counterpart cast or as composite moulds in 
the terminology of McAlcstcr ( 1 962) and 
Wade (1968, 1971). As such, they might be 
further interpreted as showing affinities with 
a rather variable group of fossil forms known 
from the late Precambrian. These forms are 
Namalia villiersiensis Germs and the variety 
of specimens which Pflug (1972a) includes in 
the "Erniettomorpha", both from the Kuibis 
Subgroup of the lower part of the Nama 
Group, Namibia, and Baikalina sessilis Soko- 
Iov from the Ayankan Formation of the Upper 
Yudomian, on the Malyi Anai River, near 
Lake Baikal in Siberia. 

Our structures strikingly resemble Namalia 
villiersiensis and the variable fossils Pflug 
(1972a) collectively describes as the "Erniet- 
tomorpha". Within this division Pflug includes 
13 genera with 28 species in five subfamilies, 
four families and two orders (Glacssner 
1979). One of us (Jenkins) has examined 
Pflug's material and considers that all the 
specimens he refers to as the "Ernietto- 
morpha" belong to a single genus and species, 
Ernietta plateauensis Pflug; the other nominate 
taxa seemingly differ only in their complete- 
ness of preservation, their degree of com- 
pression, or the amount by which syndeposi- 
tional erosional processes have truncated the 
upper parts of the buried remains (see also 
Glaessner & Walter 1975). The remains of 
E. plateauensis seem so similar to N. villier- 
siensis in their morphology, preservation and 
provenance that it is very likely that they are 

Ernietta is usually preserved in apparent 
life position with recorded field occurrences 
(Germs 1968, 1972a, 1972b) and specimens 
(Pflug collection) indicating that numerous 
individuals lived in close proximity to one 
another. The organism was essentially sack- 
shaped in form (Fig. 5A, B, C & E; Fig. 6A) 

Fie 3 Field photographs of uncollected pseudofossils; examples from lower part of Moonllah 
Formation except D and F from upper part; scale divisions in cm. A, subcylindncal form with 
beds in adjacent matrix curved downwards at sides and thinned near base: Utanouna Creek. 
B conical specimen: Utanouna Creek. C opening where conical specimen has eroded from matrix; 
upper edge of form apparently extended to line x-y and striations on mould of its surface are 
arrowed- Black Gap. D, external mould of large bowl-shaped form: Mount Chambers Creek. E. 
subcylindrical specimen showing gentlv domed ripple bedded sands above and downward deforma- 
tion of beds near base: Utanouna Creek. F, profile of shallow structure within shale: Mount 
Chambers Creek. 





and composed ol several layers or soil tissue 
which could undergo ready deformation. The 
sack is inferred In have been attached at it? 
closed vrnl, cither to the substrate, or vtty 
frequently to small clay galls or some- other 
projection on the surface ol the substrate 
(fig, 513 4 L 7 ; Fig. oA) The several wall 
membranes o\ the sack were interconnected by 
thin, flexible, radially arranged, longitudinal 
septa wh«»sc position on any particular wall 
layer is marked hy either narrow riders or fine 
longitudinal grooves (Fig. oH, C & D). The 
elongate tubular SpflCC* so formed between the 

walls and the sepi;i are frequently packed with 

silt or sand to form internal muulds. I he 
Upper -onl apparently open end of Lwistui 
terminates in a ciem-Kate margin in the lew 
fare instances whcie it is preserved Mote 1ie- 
quently. however, it is found collapsed and 
de-formed (I'ij?. ?A & E>. This discussion oi 
oiher details concerning Ihc structure of Ihis 
>.\ti (Ordinary organism fall outside the scope 
of" the present work 

One. of the must remarkable charauiii-iK - 
ol hrnh'liii is its preservation, which is one of 
mlilhuL' comparable to the structures from Ihc 
I l.'.iuVrs Ratines, Thus, sections of Lnuettu 
may show a layered and graded sand or silt 
infill, with medium-grained sand at the closed 
end of the sack and vciy fine-grained sand 
near the apparent open end I Kie. 5C>. Bed 
dint* surfaces indicate the original deposhional 
orientation. Blebs o! clay-nch material and 
limonitc psiMidornorphs, presumably ,iHei 
autbigenic pyrite. are frequent in the inftlimcs 
The tew specimens still within their original 
matrix art enveloped by medium-eramed sand 
(now quarl.'itc), The ipfillinc oi Lniic/lu d< BS 
mil. hmvLver, show the fine lamination IH1 
nhserved in our structures. 

Plfufc (1972b 197-1 ,i nriemated the "Crniet- 
lomorpha 11 in the reverse aspect 'o thai SU&^ 

yested here, that is with me closed end of the 
sack upwards. Published data concerning their 
occurrence ut the field (e.g. Germs 1968, 
IV72a) are ambiguous as to their orientation. 
Nevertheless, evidence from their distortion 
and layered infill is considered substantiaiive 
of our interpretation. 

From a slraligraphic viewpoint u is intrigu- 
ing that 'he Kuihis Subgroup, which contains 
ErmetttL is succeeded by a sequence containing 
^lactogenic sediments and scraped and grooved 
beddine planes loans ol the Schwarzrand 
Subgroup ) reflecting several cold phases of late Precamhnun elacjation in southern 
Africa (Germs 1472a* 1974 Kroner 1977; 
Kroner ci a/ JSS6j and that our structures 
occur only relatively short straligraphie dis- 
tances both bclnw and above the last late Pre- 
camhrian ^lactogenic deposits in the Flinders 
Ranges (parts of the Elatina Formation). 
However, the glacial events recorded in ihe 
Schwarzrand Subgroup arc ftjggo&led CO have 
bc«n of local extent, and radiometric data ol 
relevance lor the Nama Group (Kroner 1977; 
preliminary data passed in personal eomnumi- 
eation between Professor Kroner and Dr Pilug; 
Kroner ct uL 1 9801 together with fragmentary 
dating information from Australia reviewed 
try Williams ( 1975) suggests the likelihood 
that the Nama ^lactations arc significantly 
youpger than the Flatma Formation. 

The Siberian form Buikahnt, fe&ilis h 
appatently closely similar to fcrnietta, but is 
too incompletely described for farther com- 

One objection to a biological origin is that 
if our forms did represent soft-bodied fossil 
organisms then the creatures must be inter- 
preted as having maintained ihcir life func- 
tions even during the tirac of their infilling, 
as Iheir presumably soft integument would 
..urcly have collapsed flat if death were earlier. 

pM 4. J 'holographs of infilling sediment and seciions of psetidotossils and field photograph of depres- 
sions .onnecled bv puller- ftottl tower part of Moon II ah Formation at Utanouno Creek, except F, 
from same sirntiorapliic interval at Black Gup and G from loose boulder in Hunycroo Oeek. scides equivalent to 1 cm A, Nidc view of plug of iofill0l£ sediment of pseudofossd eroded free, 
A U No 24731. ft, field photograph ot iirepnUu depressions interconnected by gtllters: end ol 
ruler lo Ffin&c M) cm. C-V, reflected light photographs of sections of psettdOfoftsfltt C, rrosivc base 
(h) of I'-ini CfoaerV juxtaposed with bedding inleiface (\-y) helwccn silicone bed {&} and over- 
tying shale tshh A. I!. No. 2472-9 £>« W# .mill with erosive mi face, arrowed, and cone-shaped mmc- 
(f-f) extending below: shale laimnac in matrix down-iunied at hides ct sirnclure, A.U. No. 
24730 >-. specimen with etn.t.dlv domed bast and domed, finely laminated silt InftHMtfi, A.U. No. 
2472S h undciitealli view of part of base of mfllUnd plug o* larye pseud. ifossil with arrows in- 
dieao'nr- wore or bsfl radially ;itrj.npcd (nnricingf, A.LI. No, 747^3. G broken section through, 
pseiidofossil within fine sandstone: inlillin^ also tint .sandstone, include-; aUtfWUOliS shale Hakes, some 
»>l wind, ara Oltent4t«l h edgc-wiw fashion: nvci1\ing bed of laminated fine sandstone, A.U, No. 





Maintenance ot life during sedimentation 
SCCms manifestly impossible. This argument 
makes i.he c\ plii nii ti on of the preservation ol 
ht.-tnitu difficult", but for this form il might 
he presumed that the omel ot sedimerilation 
was extremely rapid, perhaps in the form of 
a turbid fay current which gave rise to the ob- 
served graded infill. In any event, the Lrnielia 
organisms appear to have ultimately suc- 
cumbed to sedimenlation as their tops are 
variuMv pursed and compressed or ovcrfolded 
(Pig. 5A & h) Our structures show no evi- 
dence of the multiple wall tissues present in 
Ltnictla, nor definite evidence of septa or the 
precise and regular organization so charac- 
terislie Of this fossil lorm. The most convinc- 
ing explanation ot Ihe striations on 'he -.ides 
ot OUT structures arc as sliekensulcs Joe tq 
differential compaction. In a number q1 -pevi- 
mens ihe slickeusided surfaces project into the 
malrix below the actual infill in the form o\ 
a more or less cone-shaped iracttire (Ffe, AD1- 
'llie systematic position of N'tiD'itlin, .In 
"Ernietlomorpha^ and BaikttlfMi rbgfcfliDl with 
sever. J other problematic Prceanibnan fossil 
onanisms, was reviewed by Glaessner & 
Walter (ls»7S), who suggested them lo he re- 
laled lo their (axon Arnmhcna hanku', I rum 
the Arumbcra Sand:- lone in the Amadeus 
Basin, central Australia. Glaessner & WfilfcC* 
considered thai vvhilc ibese forms edufd gene- 
rally he referred In PI1uu> U'/ZOa, WOh. 
1971, Ui72a, 1972b) division of the **Fciib 
lonainac' . tfie tomral application of this name 
was not to be recommended bccatt$c of its 
hypol helical concept and various speculative 
implications applied to it, Thus tffcy rejected 
Ihe ideas of PflUg fWM. 1974 > tnal ihc " Pi ''' 
latiUftaao" represent a peculiar branch if TVe- 
l unbnan evolution intermediate in position 
between the kingdoms of plants and animals. 
Nevertheless Glaessner k Walter .maintained 
that the forms just cfecusssd a»d various Ptv- 
. amhiian frond-like organisms snow a general 

similarity which implies an underlying t&KO* 
nomic unity, and that the tnerubm of this 
grouping ' may be classified provisionally as 
Coclenterata of uncertain systematic position''. 
This Viewpoint IS criticised by Jenkins and 
Grilling (1978) who suggested thai the origi- 
nal concept of the "Petalonamae" may involve 
ihe artificial amalgamation of quite disiincnve 
classes or even of separate phyla. Varying 
further interpretations of the Pctalouainae arc 
giVen by Ford (I979J, Brasicr (19791 Scrui- 
ron (IV79) and Glaessner (1979), but this 
problem falls essentially outside the scope of 
rhe present study. Based on comparison with 
the geological Mudies and experimental work 
oi Diutynski & Walton (l%5), "Artonberur 
is considered by Brasier (1979) io be "prob- 
ably a p^ClldofoSfii] caused by oirbid waTcr 
(lows' New widespread discoveries of "Anon- 
/**riV* made bv Dr li Dady and Jenkins m 
ceulral Australia (>ee ttlfiO Kirschvink 1978), 
where the form is several times found in near 
association with coarse grained sandstones or 
conglomerates, provide a body of additional 
evidence supportive o\ Brasier's conclusion. 

Fr.'lmble tru>n>.c,nic Origin 

111 tin excellent review of inorganic cylmdii- 
cat structures found within sediments. Bailey & 
Newman (I97M indicate thai the usual pos- 
tulated mechanisms for I heir formation involve 
early diagenetic upward-flowing springs or 
I -scaping pore waters (see also QtfftkC 1930; 
Havvley & Han I9.U; Gahelman t955 Cony- 
hcarc & Crook |^4j8>. Such smicluics are fre- 
quently filled with sediment of similar grain 
m./c, sorting and texutre to the enclosing (Silt- 
stone or sandstone) matrix, or show a crude 
pqU|9Cttit>Jl ir > pilB si/e towards the centre of 
the cylinder. Occasionally, BVcn an invcTlvd 
cmic-iii-c-i-inc arrangement of .-all and sand 
lamination can oceui (see Dionrie & LflVcr- 
dierc 107?: C.:mg1o*T 1974). Often the tower 
end of ihe cylinder is irreguinr and shows 

Fifc 5 


Remains of fiftjicto pftfWVntis Pflttg. 1%*. from Kninis Subgroup o( lower pan of N«nHI 
,..* N-imibiir all natural SIX* V sJdfi of neaity eomplrte specimen viewed aiAnjB 'me ul Wlflffl 

VteSwd no.., mI to wider l£n«VWM axis; at ItaM. three separai: wall membrane^ ( W\ W*- .ind\\ • ) 
shown bv interna! moulds of saml. e.MU.mimc "hn.libne ttltdte oceu.s ,.!ony b-b Pf. No, 1^. C, section ol sand .ntillmu envitv showing layers P)l ptpmm finer M'd.tneni 
nut denosUtonal surfaces (-0 ra-sumabiv ttllatune mtBrmllient singes ot ini.ihn- and onenimiun 
ul organism at each stase; ct-iik miu.hv and MM* limon.te after authjfeBftiC pynte b?hi MhluM 
cctu-L ol [he in-line due 10 v:eaUi-,r ( .,:z PI No 5(V8. O. ot l«Mfi ot srecmcB ri|OWl>1* fietormcU 
nrofl [dl ;mO m>plc-Jike prominence (p) wtiid. ruij watt ihe tomi Of aitactimera 01 foe 

or^miM., /,^ai' sniare hW bnc n-v, Pf. No. 09 K, „de v-exv o speumen With some Of 
endosino rnatro.: mHlinfi i'tdvi.1nal cotlap.scd downward*, u.tb Wi$e llauenin- and upper pail 
parsing nnvar.N, Pf No. 287. 




F.g- f> Ernttttto pfalcutunsis Pflug. 1966. A, reconstruction of group of individuals living on sandv 
Prccambnan sea boltom. Several possible alternative, hut noi necessarily mutually exclusive re- 
constructions of individuals shown. Youthful specimens evidently attached to mud galls 01 other 
prominences on sea bottom, more mature specimens may have lived with basal pirn buried in 
substrate. Outermost membrane seems to have been caul like and only faintly showing underlying 
loneiHklmal structure. "Mouth" sometimes preserved btoadJy Oared, or more frequently collapsed 
01 pursed Individuals of all sizes may shown enigmatic transverse suture or possible zona of hud 
• iinj: at ;.bo.H half their leneth. About .7 natural size. HI), sketches of sectional cuts through 

RCtual specrmens; b, traces of bedding in matrix (impure sandstone or quartzile); r adhering thin 
illVn 0\ rock; s septal membranes evidenced by surfaces of parting or thin limonite stained traces 
w similarly evidenced wall membranes; z, known or inferred position of basal zig-zag suture all 
d* C .^ ^Plong.tudinal section almost normal to aig-ettjj axis, three wall membranes evidenced 
H. Mo. iajj. C, section 01 another specimen almost transverse to lonfiilUdlnal surface markings D 
section ot different specimen cut oblique to longitudinal surface markings 



fissure fillings. Jn the present structures from 
the Minders Ranges, aowever, the frequently 
line lamination of ihe more silly phases of the 
inflUmg shows no evidence of disruption olher 
than slight deformation during compact ion, 
and the thin sand beds which often occur at 
the base of the cylinder show no sign of 
Assuring or disruption Mich as miebl he ex- 
pected with the rapid escape ot fluids. 1 'hus 
our structures were evidently not formed by 
the upward escape or llowaec of pore Hinds. 
Also, the syngenesis of these structures elimi- 
nates au epigcuetk coiicrctionat y origin Such 
syngeneic structures could, therefore. Ofity ^ 
formed inorganically as lotoids or pot-casts; 
these names both essentially referring 2D the 

casts of circular $wu* pits. 

In past literature such -o>ur pits are olicn 
suggested to have been formed by whirlpool 
action generated along the boundary between 
currents flowing m opposite direct" ne-. As such 
they might be expected to occur in high energy 
shallow '-water environments in both marine 
and non-marine realms. According Pr Cony- 
he.trc & Crook (1968*1, circular loraicfcs tfft 
usually wider than deep. The external mor- 
phology ol their C3MI ts suggested io be 
chafaetenaieally swirled, like that of a 
"folded bun", generally with a smooth -uiiacc, 
although the formation of roughly conecninc 
patterns by scouring is common. Internally, 
loroid easts may display structural and lex- 
lural homogeneity 

Our structures show a marked resemblance 
to the "rippled toroies' 1 described by Don & 
KaulTman ( 19K3J from the Mississippian 
Napoleon Sandstone of southern MiehL'.m. 
They considered these loroids to have been 
loimed by Ihe action of VorleX currents on 
LiMcansOlidiiled Mid «i shallow wuAct Of the 
inner suMittoral 7o\m:' and suppnitcd (heir 
firuline-S by inducing vortices in a \\ at t:r-fi'lcJ. 
Sand -floored bOX and producing Attiffcial 
loroids- Sonic of the structures they described 
were elongate, oblique cones, a shape occa- 
sionally shown by our material 

This precise explanation seems inapplicable 
wiih respect If) OUT structures for several 
reasons Dorr A; Kaallmann considered that 
Ihe environment of deposition of the Napo- 
leon Sandstone to*$ one of relatively high 
CnciBV and this is consistent With the texture 
of the rock fsand). Our structures o.-.m 
within a variety of titfiotvpes representing mud 
to fine sand grades, reflecting micTocm ^ roti 

menls of varying eneigy levels, and pte-sum- 
ably mostly of taihcr low energy. The \ or lex 
hypothesis invoked by these authors implies 
tidal shear or near-shore pheuomenn which, 
by their wry nature, musi be restricted to a 
relatively narrow beit parallel to '-he pulaeo- 
coasl. We. however, have located our struc- 
tures at geographic separations of 20 km 
parallel to the inferred depostitioilal strike :md 
perhaps fcfl km at right angles to it. Also, our 
forms occur a) various levels over a t0t«f1 strati- 
•jra^bic thickness in excess of 1400 m. 

Aieuer & I uttcrei I J97&1 and Aiger . I97S) 
docuincnt and explain inorganic pot-casts ami 
guUcf-cavts from the Wuschelkalk of south- 
west Germanv Some ot the po*~CfcStf w:hich 
iiir., inmrc (i.e. Aigncr & FbKcrfcr l^7H. fig*. 
JA AV, D. iV and t > au identical m :.bape 
and internal layering to the Flinders Ranees 
specimen*, except that the coarse material Bl 
the base of the '"pois" coomsU of shell dehris 
rather than day galls. The Musehelkalk pot- 
may be packed wdlhn an edgewise WHl 
glomerate of shells similar to the edgewise 
conglomerate of mud flakes present in some 
of our specimens (Fig 4<J). 

On the basis of limine experiments A.gner k 
Futterer attribute the Muschclkalk pot-holes 
|0 the erosive effects of eddy currents 1'ormrng 
in shell-filled depressions due to the influence 
of a steady or increasing unidirectional eur- 
lenf over the general surface of the substtate. 
Ihey note that such pots arc unknown on 
present-day tidal flats and consider that their 
onyin is entirely submarine. The implications 
of 'this are at some variance wiih Plummer's 
(|*)7S| finding that on the basis of such struc- 
tures as flat-topped ripples, interference 
ripples and shallow channels with ebb-orien- 
tated asymmetric ripples, parts of the MOOTII 
Ijjfi Formation in which our structures occur 
frequently accumulated in a low intortidat mud 
Hal environment. However, we did not obscrvr 
anv sedimcntologieal structures in direct » u 
ci;-;'h.ii with the poi-easts that would imr>"ly 

An attractive aspect ot' this explanation for 
the Hinders Ranees structures is thai somc- 
times the pot-castv are peculiar ly coolesced nr 
joined bv une\en Ir^uelvtil-L forms [Fig. ^rO 
which are exactly parallelled in the Musehel- 
kalk material 'Avjuer 5 I*.ilterer [SW. hflS. 
3B. G and F). In rare inslaiices there rruy be evidence of a ripple hollow or other 
depression in which (he clay galls (hat 


250 i 

200 ■ 







Number of Readings = 72 
(Level II) 


Maximum Diameter (mm) 

Number of 
Readings = 230 
{Levels I -IV) 


Level IV 

Level HI 

Level II 

Level I 





Direction of Alignment 

100 150 

Maximum Diameter (mm) 

>^^. . P^^, 







tonncd the cjomvc loots may have originally 
.aiumulatcd (Fig. -3 A). 

Measurements pf the diameter of the pot- 
casts in the Fliudcrs Ranges {Fig. 7A & B) 
snow a range of variability quite comparable 
to gutter-casts studied by Aigner & i utlercr 
<uul the depth o\ specimens is similar. A histo- 
gram of diameter againsL frequency (Fig. 7A) 
shows u somewhat skewed distribution with 
the modal diameter ul 6-7 em. A suggestion 
of a polymodal component in this histogram 
probably reftee's the observation lhat spci. i 
mens of the pot-casts on giVWl bedding planes 
tend to be closely similar in sue. a likely in- 
diealtun of then nearly ;>inuibaneous gjRflfifO- 
tioa. Karez ?t uL (1974) describe how a held 
of small pits | individually reaching up 10 
9 mm diaint'K'i I may lonu on freshly deposited 
clay undei the influence of a current, and it 
iinv be possible that such depressions serve 
as a nucleus lor the subsequent formation of 

Ihc study of Aieuer & Fultcrer also de- 
mon*»inies a very marked preferred oricnla- 
tion foi the JVIuschelkalk gutter-casts, parallel 
to the coastline and ba.sin axis. Our poU 
••'iuc'u'vs arc usually ovoid in outline 1 1 j >j 
7B) ami measurements on four bedding-pkmes 
ill ascending straiigraphic order at Utanouna 
Creek also demonstrate a marked pi cf erred 
orientation tor any given bedding plane (Fig, 
7C>i dtough tins direction jhauges tiy 90 
hi twee. i ihc lowest and the (bird surfaces 
studied. I he more or less east-west oiienta- 
ihhi- Kir Levels I find II are approximately 
normal to the palaeodepositional trends for; 
thc Hinders RrinjfiS aica. whilst ihe N.N.W. 
SS,1 diloCHPlM foT Levels til ami IV arc 
mui-hly parallel to the basin axis and (wes- 
tern) shoreline (e.g. stfic Plummer 1 97N ) , 

The individual pot-easts are spaced on 
bedding planes at intervals of several freriS Ql 
centimeters lb seveial meters. In rare instances 
-iwirnens arc grouped on parr o( a bedding 
plane, while none are present on ihc remainder 
of Ihc surface Doff & KautTm-in (1963) 
pointed out die "rippled toroids** which 
they described also show a patchy distribution, 
Similarly Norrman (I°M ) indicated a 

clumped distribution for modern day pot-holes 
eroded into a clay lake bottom, 

Associated pseudnfossils and oldest local 
record of metazoans 

Giaessncr < 1 969 j described the complex 
liace-Jike marking fiunyerichmis dul^urnoi 
from the base of the presently recognised 
Mooriilah Formation at Bunycroo Gorge. He 
considered that this marking may have been 
made by an animal related to a primitive mol- 
lusc without a shell. Large bedding plane 
exposures aic a persistent feature of many 
outcrops oi the Mooriilah Formation ;U this 
locality, but despite extensive searches no new 
finds of B. daignrttoi have been made during 
Ihc decade since its discovery. Alter critical 
examination of b. dalxarnoh Jenkins (1975a) 
suggested thai it is a composite of primary 
impressions occurring on two separate bedding 
planes, and thus unlikely to be ot nicla/oan 
origin. The great frequency with which in- 
organic tool markings occur in this formation 
ted Jenkins to consider it \o be "a Unique 
accidental set of markings made by a tethered 
implement moved by ihe current". The imple- 
ment may have been a small mud clasl caught 
iu n tassel of twisted algal threads. It is sug- 
gested to have moved in a series of small 
jumps in several arcuate swaths to leave over- 
lapping, cujvcd bands of more or less regular, 
transverse imprints marking -.atccessivc bed- 
ding planes. Thalus-bearing. ribbon-like aigac, 
which might have provided the tethei. appear 
in ihc Riphean of the U.S.S.R. and form ac- 
cumulations grading into sapropelic laminae 
and films in the Vcndian (Sokolov 1 977). 

Small circular markings on a bedding; plane 
ol a single loose block of rock found within 
Ihe Mooriilah Formation have been considered 
as structures formed by escaping gas 
(Plummer I9K0). 

I be oldest probable record of metazoans in 
the blinders ranges is. of markings resembling 
small medusoids and fine sinuous trails found 
by Dr Mary Wade al Brachina Gorge wilhin 
the Bouncy Sandstone (previously the "Red 
Pound Quart/ate 1 } which is the lower forma- 
tion of Ihe Pound Subgroup (Wade 1970, p. 

Pip 7. Measurements of pseiutofossils at Uumouna Creek. Bunhinyunna Ranee. A. histogram of 
maximum transverse diameter of specimens plotted againsi frequency; specimens occur on four 
larfce hrrWing planes, levels I IV in ascending slraligraphic order. B. plot oi maximum and minimum 
transverse dMmcieis of individual specimens occurring on level II, C, direction oj orientation of 
loiter transverse axis of ovate tpecimenes plotted for each bedding plane. 


R. K. i ll-NKINS. I\ S. ri IJMMDR & K C MOKIARi Y 

92). Other fiiuN or Precambftafi MetaAaana in 

Ausiudw qrc apparently either approximately 
eqmvalenr ir. age to ihc L.diaam» assemblage 
(wmii ifrir/v or ore likely ol younger age .i> 
is mJcrred foi the Mi Skinner 1;mnj M ■ <f the 
Amadous and (ieorgina Uasins (see Winlc 
1369; Daily 1972; Kirschv'mk 1978). 1i diuS 
becomes timely to consider why ihc s*.v- ■ n 
iIkmil-.huI moires ol Mannoan or youngest pre 
Cambrian sediments which precede ihe Pound 
Subgroup m ihe Flinders Ranges, aiul which 
arc well exposed and otherwise occur exten- 
sively throughout the Adelaide Fold-Belt, are 
apparently butren ol mela/oan fossils, Nume- 
rous stnt (graphical and sedtmcritokigieal 
studies gem-rally imply marine origins foi 
much •_>! tfns succession ( I hornson 1909; 
Thomson el al. 1 976; Plummcr 1978). Argu- 
ments rhat the I acres may noi ba\c been 6Ufc 
able tor the preservation of small soft-bodied 
K'limins i.i Olaessner I972) are not fillty 
i QftVfncfng as fine!} laminated beds preserving 
ihe smallest inorganic tool markings (en- 
(J.J mm) are hequent. and Wade (1968. 1470) 
has found that the conditions needed for the 
preservation uf a soft-hodiod assemblage WCtt 
not particularly stringent, Indeed ihe Fdiaeurj 
assemblage \ru\u lam is now known vo occur 
in quite variable faeics ai diH'erenr World 


Aii extensive literature concerns supposed 
finds Of either body fossils or 1race IosmIs pre- 
dating the Fdiacara assemblage srn.stt Uao, hut 
subsequent studies have led frequently t«. ;i 
questioning of the interpretation or sometimes 
ihe age of these reported occurrences (Cd;it_ ;:- 
ltd 19iS9. |97s»andpers comrn.; Ctoftjd 1973. 
I97K; riouci rt ai 1980; Sokolov 1977; Yakoh- 
m-iii & kiylov 19771. Other of these finds 
continue to be cited as possible evidence sup- 
pt'iiive of ideas ot a gradualist!", evolution flrf 
• ;iily rn-eta/oans (&& Glaessnei 1'*?2. l r ' 7. 
Hem-stun 1977; Rrasier 1979). The studied re- 
cord of the sequences tn The Flinders Ranges 
docs noi provide evidence of an extended 
early histoiv fof the metazoans. 

Rulher negative evidence ami the known 
incurrence of fossil* tn the Flinders Ranees 
lend support in ideas such as those of Stanley 
(1973. iPTfift. WbM a late radio- 
Of meia/oans. predicted on a basis of 
ecological modelling, or to an exponential in- 
crease in animal life from ihe laic Precambrian 
to the Cumbrian, shown by analysis of age 
data and counts of recognized taxa (Sepkoski 

197S. 1 979 1. Stanley considered that the early 
radiation of cuknryoics may have been in- 
hibited by the saturation effect of Precatubnan 
algfci] systems and uicgcsled .hat ihe advent of 
cvM-eating hetcrouophy triggered a "kind u)j 
sell -propagating feedback system of divcrsifi- 
caiiou*' cumulating in the initial major rHoUUl 
(ions of bolh ihe melaphyles and melazoans. 
Recently Chouhcn & Fame Murer <I9KD> 
a^tgncd a middle Ripbcm age lo ihc rock 
containing Precambrian mctazoau fossils on 
the Avalon Peninsula. Newfoundland, and at 
Chat 1 1 wood Forest, England. This finding in 
patently incorreci. and not only overlooks 
lines ot geological evidence which suggest 
that these rocks are late Precambrian (Wil- 
liams & king 1979; l>atshctt ft nL lo.S0). but 
ITCS a 20 year cycle of research documeul 
wg Vcndian or Ediaearan affinities for the 
genera (lunula Ford and CtlCfWfodkcltf Ford 
present in the fossif assemblages ftdaessner 
I 977. I979K 

( -mi. Ins inns 

The present study describes Precambrian 
slructures which have a relatively complex 
L'L-uesis and show resemblances to CCTtSUd 
cylindrical lo conical fossil hurrows. as well as 
to sack-shaped, longitudinally striate body 
fossil remains of comparable age. However. 
these resemblances appear to be fortuitous and 
the true origin of the structures is suggested 
to he as erosive pits formed by vortices induced 
i->v episodic currents and the scouring aetion 
of trapped mud gall*. The hydrodynamic con 
difions leading to their formation \rc not fully 
understood and to our knowledge no ;iueh 
siruciurcs have yet been described from 
present da v. marine, sub-iidal environments 
the degree of resemblance between thes-- 
pscudofossils and several bin row lorms and 
sack-shaoed bodv fossils emphasizes that a 
variety of both inorganic ^nd Organic pro- 
o sses may lead to the formation of closely 
Similar structures which may even show de- 
grees of gradation. The corollary of this 
finding is lhat the description of such kiml- 
of Precambrian structures and the assignment 
of a particular genesis, especially one of bio- 
logical origin, should be approached with 

The finding of these structures and recog- 
nition of other associated nseudofossils sue 
gests lhat true indications of Metazoa are 
restricted tn the highest part of the local 



Precamhrian sequence, ihus providing nega- 
tive evidence supptyltrtg recent theories that 
postulate a late evolulion or radiation ot 
animal lite towards »he close of the Precarn- 


Dr Hans Pllug of Ihe Justus Liebig LJtii 
vesitv. Giesscn, Wo*t Germany, is warmly 
thanked lor allowing examination and study 
o( Ins lurec collection of Precamhrian lossits 
from southern Africa, and for his hospitalny 
and numerous valuable discussions on his 
material. The University of Adelaide is ac- 
knowledged lor rnakinc a Study Leave Grant 

available towards this purpose. 

Dr B. Daily, University of Adelaide, and 
Dr W. V. Preiss, S A. Department of Mines 
and hnergy, are ^ratelullv acknowledged lor 
their constructive criticism of the manuscript, 
and extremely helpful remarks were also made 
by an anonymous reviewer. Dr V. Gostin, 
University of Adelaide, assisted with litera- 
ture, and Professor C. von der Boreh, Flinders 
University, is thanked for discussion. Dr T. D. 
ford, University of Leicester, England, read 
the manuscript ai the "Symposium on Life in 
The Precamhrian. Leicester. April 10-12. 
|y8(T. and passed on the remarks of the 
assembled meeting. 


Animk. '!'. I 1978) 3 Slorm-generalcd eoquinas. 
In 1. AlfiJltr, 'L Hnjjdom & R. Mundloo (Edsi bioslromal and worm generated 
eoquinas in ihe Upper Muschtlkalk. pp 49-5fc 
X'nn-s //'- Cfol. Ptilaont Ahh 157 

$ Ku.hih. E. tLJ7«) Kolk-Toplc und- 

Rinncn Ipol nnd gtiller-easts) im Mu*ctac1k«Ik- 
An/eiijer Fur Wnucnmeer ihkl. 156, 2K5-30u 

Aiimui. S. P. fr£7Sj Bemtn>>ria feantl (C im- 
btian and Ordovieiim'l. a nrohahle aeliniun trace 
Insvil, J Ptltt'OtU. 47, sJ]9-924. 

Hmuv. R. H. & Niwmmn. W. A. M C >7H> Origin 
and significance- of cylindrical sedimentary BtrUG- 
lures from the Boston Bav liroun, M.i^wchu 
sells. -1>»- /. St/'- 72H. 713-7 M 

HiNfisruN. s. (l t >77» "Aspects nf prabtenwiic 

fossils in the early Palaeozoic." lAtfl Lmvcrsi- 
'atis Upsalicnsis 115. Uppsala,) 
Rkanson. C\ ( 1460) Conosiu-futs. Ok ht Ccol. 
flfrjfn 20, H»_v207, 

huasiml m. d. r ]«>7y ) The Gamfcrfeui rajliatfoo 

tvcHf U M R- House (Pd_> "The Oiiuin M 
Major Invertebrate Groups." pp. 103-159. Syst, 
Assoc. Spec Vol. 12. (Academic Press: I ondon 
X New York.) 
( :n4MU Ri A»N. C. K. t 1971 ) Morphology unci 

ethology ot rrucc fossils from the Ouachita 
Mountains. 8»R MMioau. / ftftaftft 45. 212 

: a, 

I iipUUfrfCT. <• (El Iauki -IvUkm, A. (IVSt)) The 
iTeciimbrian in norih pcri-Allantic ami 80Utb 
MraiUTnuitian moh-lc /ones, general result*. 
Etfth&t '*'• »«> ftS 219 

ClCltii). P- CW3J Pseudotossils: A plea for 
cumiuii VtroJaqy 1. 123-127, 

( 107K) World's oldest ammal \XS&4$ \»i«r< 
275. "Wl 

, (Ommson. I H. & W\rsoN. ). A. L, 

1,1'lSO) ) Im- works nf llvlritf BOSJal insects as 
pst nd*ilossiK :nnl [he age Qfl the oleics! Vnown 
MetflZOQi Hi rVnctf 210, I0I5-10M. 

LoNVnrvkt, ( E U. & Ckook. K. A. W. ( l»>fiK ) 
M ( inu;il ttl srOitm-iiTnry Mrnellile^. fl/tfl V«» 
rVfi»tfr, Hi'xotir. <7eoL Gtaptiyt U\l. 327 p. 

C "RiMhs, I. P., Iron, J., Maucos, A. & Arholuya, 
M. (|577) -'l.aic Ptcuimhrnm-low Lower Lam- 
hriun (nice fossils from Spain, in T, P. Crimes 
& J. C. Harper (Eds) "Tiaec Fossils 2', pp S»2- 
138. Lpool Momhr. £tol. J V't't tssiu 9. U. * 1972 ) The base ol" the and 
the Itrst Camhrmn fiiimns. hi j, B. lones iL B. 
McOowr^n lEUs) '•Stratieraphie Proltlems of 
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DzutYNSK], S. & Waiton, E. K. (1965) "SbcIj- 
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huw, R. W. (1970) 'Hie lebensspurcn ol nunc 
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Ioiuj. I. D. (I979) Piec;imbrian fossils and Ihe 
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(loorvrw, I. W. ll»JJ5.| Cylmdiical SlrUClUW rn 
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Ganoioii\ P. (1974) Lcs striicmres .ylindi iqiies 
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COkms. G. .1, B. (I'^M) Diseoveiy of n new 
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- ()'>72ai "The Shtr-aTigriphy and paleontology 
of the liHver Nama GrOUp, ^Ql\th Wesi Africa", 
i' .;/>'. ^rtpr if*H(( Dipt th<>lr<$\ Ciuwrhcr of 
Mines PriCOmh /?r.v, Unit, Hntt \Z, 250 p, 


l VMh l I 'ice tov.ils /-oir! iltc Nama 

( 19741 The Niimu CStqup in South West 

\lnc-l and lis rebtiimsbip If) the- P/m-Atracm 

GeosyncJfne, ) Gt&i *z, an-317. 

Gimnsmk. M. Fi il96*M Tnrce fossils from Die 
Hcecumbrioil and basal Cambrian, tt'ihaia 2. 

It&fcj »'n i.,.mhiian puliteuzoulog>. h\ I. B. 

Junta $ U. McCiow 'ran < Eds i "StrMtigrafihii 
Problem* eri ihe I utei Precambruo and Early 

C tivibm.n , pp 4342 itifcJV. Add. Centre fot 
Pteeamk Res. Spec. Pap. I: Adelaide) 
— (14771 The F-diacara fauna and irs place 
in the evolution ol ihe Mettaeitf. ffl A. V. 
renko (Ed I t orrelaia-n Of Ihr Piecambnan" 
Vol I, pp. 257'258. (Naukj. Moscow j 

i I97VJ Presamtomn. to R. A. Robinson ..nd 

f' Teiclntt ltd--} irc.iise on Invertebrate 
&luj£0fltolo£} Pari A Inlroduclinn FovrilizultOP 
( I aphonomy) Biogeour.iphy and BioMmh 
poiphy", pp A79-AI18. tCcol Soc Am.. Umv 
Kansas Lawrence. Kansas. J 

& Wai im. M. |-\ fW75) New Pre-cumbruin 
fossils from tlit Aturnbcra Sandsiouc, Nonbcin 
Territory, Australia Afckcrttt$a I. 56-69. 

lUwuv. J. E. &. Haki. R. C. (1934) Cvlindncal 
simctutes in sandsrone. *"" v.W y ^ J| ff/l 45, 

^'■mnv R. F. f. <(97_Sa> The JiverMiicMfon of 
carls me'a/oaus Abstr 1st Aust Ceo! r'nnv 
A.IiL.ide I Mr. May, 10??. pp |<J-2«. fGeOl 
Sot Ausl.. Adelaide, t 

(1975b) An environment. j! ntUiiy of (lie 
rtjefa combining Ibe EJincanf ■•isM.-mbl.iye in the 
Flinders Ranuts. th>d, py. 2\ ': : 

-k Gluing. J G . I978)A review of the 

frond-like tossils of 'he F.diac .r.< assren 
iiii s. ANXb A/ '" 17. 347-359. 

KaRCV I l-Nas, P. £ UNGil.Lf. >, MO;.), 

SimctUrw iicnetuted in fluid Hr&fting. of freshly 

deposited days resemble ichnotossiK (;,■<<( 

2. Zm-VJU 
KlfiSCUVINK, J. I M978I The Prccamt a,:, 

i i mhti. mi i-tuun-Jaiy problem: magneto-strati 

eiaphv of ihe Amndcus Bu&in. Ceninil Austra 

I-.. Qcoi Mais 115. 139-150 
M:om «. A. i i^ll \ Non-syiKhroneity of the 

LaiC rVeea-rnhuan gfaci lljooS iji Africa. /. 

CiArf 85. 289-joii. 

. VhWutHMS, VI, O, v,rr«M8. Cj I. H. 

KMU V H \ Sn.MK. K 1£ l_ (1980) Haleo- 
maynetism ot late Ptvcambnan [p early Palpa 
/oil nii>.tiie-brar r ing Jormaiions m Namibia 
( South West \tncMi: The Nam.* GfOiip aod 
Hljiuhckcr I oimaiion 4>ft^ /_ V» . 2K0, *>42- 

MaN(.um. O- C, (1970) bchaviom Dd 
I hi m-.i anemone P/nllartix H\ol ttult khtK 6/of 

Mrwsi.N. O. | |Q49) KlfHina vlaciation: a lhir*| 
urrpficc of plMC'atton evidenced in ijie Ade- 

liiidf System. TfUUS. fi. /Stir, s Auto 73. M7 

Mi Ai.mih. a. L. d962t Mode or preservation 

m early Palsieo^oic pelevvpods and its iviorphu- 

lopit am* cCOlOfijc significance. / Pdteont. 3(i. 


Nuuiik Arai, \T & McGi'.vs, A » I%K) A 
^ifhiemahcal uoelenlerafe (7) ftom the Lower 

Cambrian, near Moraine L^ike, Banlf Arc-. 
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N(*RR\tAs ) O iVW) Lake Vallem, IrtvcMipa 
lions on shore and bottom morphology. CY. 
inn!. 46. 1-3 

PAiitnir, J. P^ Gau, N. H., Goodwin, R. & 
Hi mm, M, i. 11980) Rb-Sr whole rock iso- 
ChTpn Hfifffi of laie Piccambrian to Cambrian 
igneous rocks trom souihern Britain, ;, Ocot 
$0c. fhm\d ) I3t. 649-655. 

Pf'tjUb. H. D. (!970iO 2lU hauna der Namu- 
Schiehtcn in SiidweM-Afrika I. Pteridinia, Km.- 
und svstematische /itgeb«:»riekeit. Palaeanto- 
tyfiphtefi A, 134. 226-2lO 

il970b» /ui r-aiiua Jc Nama-Schichten >n 
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(1971) Neue K'ossilfnnde tm Jung-Prakam- 

hnum und ihrc Aussagen xtir Entstehung tier 
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(P*72a) Svslematik der jitnc-prakambrisehen 
Pttjlonomav f'Mug 1970. Palttant 7.. 46, ^6-67. 

- (I972bi /ur Fauna ijer Pfamfi Bcbicftten in 
StJdwesi Atrikn III. Emiettomorpha. Ban und 
Systenia'ik Pahiconto^taphint A 13'). 134-170. 

- (1973) /in l-auna tier Nama-Schichten in 
Siidu est Afrika IV Miktoskopische Anaiomie 
dcr PeEalo-Orgamsmen. Ibni 144, 166-202. 

( 1974) FcinttntktUl und Onlqgeme dcr 

jiinci-priikajnbiischtfn Petalo Oryanismen. 

Paiiunu /, 4K, 7 7-||)y. 

Pi umvii r ? P. S. ( 1978) Stratigraphy of the lower 

Wilnena Group (late Prcvumbrian ), Flinders 

KaiiL'es .South Australia. / runs. H Sac. A. 

Ana. 102, 25-"iH. 

( I98(M Circular stiuctures in a Lale Prt 
i ii bt tan sandstone" PossH medusoids or r\ . 

tlcnce af fluidization':' (hid 1»»4, 1 3- 16. 
0''i<Kf i t (1930) Spring pits, sedimentation 

phenomena. /. Qjtal. 38. KK-91, 
StHi nov. C T. ( 1M7*J "i Early fossil cnidarians 

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0C-. Spec;. Vol. No. 12. (Academic Presv 

London A New Vork | 
SrpxOSKL I J. (1978) A kirjeriC model of 

Phanert.i/oie la\unomic Oivci^n I \nalvsts ..| 

marine ordeiv P'ifc<'hioloy\> 4, Z23-25I 

(197^1) A kinetic irmdcl Of Phaneto/oic 

i i.-:nr.omie diversity \[. Troly Pfuincrozorc 

UiniUr^-- and mulliplt* equilibria tbttL 5. 223- 

25 i 
Shinn. L, A (I96S) burrowing in Recent (Jmc 

scilimenl?* of Morida and the Baham -v 

P.tteont. 42. 879-894 
SoK.nnv. B« S. (1977 1 Heispeetivts or. Prccam- 

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r;.y./ <-,<-<>})■,_ is, 54-70. 

S'amiiv, S. M. (197?) An ecological theotv for 
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Intc Piecambriaii Prat tsutn Acad Set 
T/mS A. 70. I4B6-WW, 

I J 976a 3 Fossil dftUj and the Precambrijn 
Cambrijii evohu- .„a. v irariMlion. Am. } Sit 
276, 56 76 

- 11976b) Ideas of the filHIBfi of meta/oan 
OivcrsificaLior. PuUfthioloyy 2, 209-219. 



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, Daily, B., Coats, R. P., & Forbes, B. G. 

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— — ( 1969) Medusae from uppermost Pre- 
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— (1970) The stratigraphic distribution of the 
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(1971) Bilateral Precambrian Chondro- 

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Wii liams, G. E. (1975) Late Precambrian glacial 
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Williams, H. & King, A. F. (1979) Trepanessy 
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7, 57-70. 




The type specimen of Peyssonnelia australis Sonder in MEL is a Peyssonnelia and is distinct from 
the taxon known as Sonderophycus australis (Sonder) Denizot. The latter, which is based on P. 
australis Sonder, is distinct generically from Peyssonnelia and is therefore re-described as 
Sonderopelta coriacea gen. et sp. nov. P. australis Sonder is an earlier name for P. gunniana J. 
Argadh but a synonym of P. capensis Montagne. 


by H. B. S. Womersley* & D. SlNKORAT 


Womersley, H. B. S. & Sinkora, D. (1981) Sonderophycus and the type specimen of 

Peyssonnelia australis Sander (Cryptonemiales, Rhodophyta). Trans. R. Soc. S. Aust. 

105(2), 85-87, 12 June, 1981. 

The type specimen of Peyssonnelia australis Sonder in MEL is a Peyssonnelia and is 

distinct from the taxon known as Sonderophycus australis (Sonder) Denizot. The latter, which 

is based on P. australis Sonder, is distinct generic-ally from Peyssonnelia and is therefore 

re-described as Sonderopeita coriacea gen. et sp. nov. P. australis Sonder is an earlier name 

for P. gunniana J. Agardh but a synonym of P. capensis Montagne. 


Peyssonnelia australis Sonder (1953, p. 
685) has been recently referred to as Sondero- 
phycus australis (Sonder) Denizot (1968, pp. 
260, 307). Earlier it had been referred to as 
Ethelia australis (Sonder) Weber van Bosse 
(1921, p. 300), though Weber van Bosse's re- 
cord "Archipel lndierr seems likely to apply to 
a different taxon. The description of Denizot 
was probably based on material such as that 
illustrated by Harvey (1859, pi. 81), and this 
is indeed a distinctive southern Australian 

However, the type specimen of P. australis 
Sonder in MEL (573182) is not the plant now 
known as Sonderophycus australis but is Peys- 
sonnelia gunniana J. Agardh (1876, p. 387), 
which Denizot (1968, p. 123) places as a 
synonym of the South African P. capensis 
Montagne (1847, p. 177). 

The type specimen of P. australis in MEL, 
from Holdfast Bay, South Australia (F. 
Mueller), includes a small sheet with Sonder's 
handwritten notes on both sides; several 
phrases are repeated in the type description. 
This number, with four pieces of thallus in an 
envelope, is regarded as the holotype. There is 
a further specimen in MEL (573183) labelled 
by Ferdinand Mueller and which is probably 
isotypc material. Sonder later also included 
some Sonderophycus specimens under his 
P. australis. 

P. australis Areschoug (1854, p. 352) from 
,v sinu Port Adelaide" (specimens in S) is the 
same as Sonder's type; Arcschoug's name was 
apparently indepedent of Sender's . 

* Department of Botany, University of Adelaide, 
Box 498 G.P.O.. Adelaide, S. Aust. 5001. 

t National Herbarium of Victoria, Royal Botanic 
Gardens, South Yarra, Victoria 3141. 

It seems likely that all references to Son- 
derophycus or its synonyms apply to this genus 
as understood by Denizot, apart from Sonder's 
original description. The misinterpretation of 
Sonder's type probably dales from Harvey's 
1859 description and his Alg. Aust. Exsicc. 
328E from Port Phillip Heads, Vic. (also 
Harvey, Trav. set 434 from the same locality). 

Since the generic name Sonderophycus is 
based on Peyssonnelia australis Sonder, it must 
be relegated to synonymy of the latter. No 
alternative generic name or specific epithet 
exists for Sonderophycus, which dates only 
from 1968 and was imperfectly presented 
(Denizot did not see and failed to cite the full 
date for Sonder's publication, viz. 18. . for 
1853, and the French discussion and citation 
of basionym are given on p. 260 with the latin 
diagnosis on p. 307). 

Accordingly this distinctive taxon is here 
described as a new genus and species. 

Sonderopeita coriacea gen. and sp. nov. 
Peyssonnelia australis sensu Harvey 1859: 
pi. 81. 

NON P. australis Sonder 1853: 685. 
Ethelia australis? (Sonder) W.v. Bosse 

1921: 300. 
Sonderophycus australis (Sonder) Denizot 

1968:' 260, 307. 

Sonderopeita gen. nov. 

Thallus uncalcified, with a short, thick and 
felty stipe of entangled rhizoids and an ex- 
tensive relatively thick and cartilaginous lamina 
extending eccentrically from the stipe position 
with concentric growth zones, variously lobed 
and lacerate; lamina in section composed of 
radiating filaments of cells of similar size, 
spreading both to the underside of the thallus 
with frequent terminal cells of the filaments 



Fig. 1. The holotype specimen of Sonderopelta coriacea. 

producing attachment rhizoids, and to the 
upper (light-facing) surface with the filaments 
becoming erect and short celled to form the 
cortex. Reproduction unknown. 

Thallus non calcareus, stipite rhizoideorum 
brevi et crasso et lamina extensiva crassa 
cartilaginea a positione stipitis zonis concen- 
tricis auctus ecccntrice extensa, lobata vel 
lacera; lamina in sectione e filamentis radiatis 
cellularum amplitudinis similaris composita, 
filamenta utrinque extendentia, ad paginam in- 
feriorem cellulis frequentibus terminalibus 
fllamentorum haptcra efferentibus, ad paginam 
superiorem filamentibus erectescentibus et 
cellularibus brevis cortex formantibus. Repro- 
ductio incognita. 
Type species. S. coriacea sp. nov. 

S. coriacea sp. nov. 

Thallus (Fig. 1 ) eccentrically peltate, 
spreading from a short (to 2 cm long and 

li cm broad), fibrous and often divided stipe r 
usually growing under overhangs in low light 
intensity; lamina cartilaginous, i-i (-H) mm 
thick, radiating eccentrically from the stipe, 
often deeply divided or laciniate with lobes to 
20(-25) cm long and io 15 cm broad, margin 
smooth, convex to rounded, dark red-brown 
above (side to light), grey and fibrous (from 
septate rhizoids) below where in larger plants 
this side is 1-2 cm from the rock substrate. 

Thallus eccentrice peltatus, ab stipe (ad 2 cm 
longo et 1] cm diametro) breve saepe diviso 
patens, plerumque sub petra imminenti in luce 
demissa vivens; lamina cartilaginea, i-1 (-H) 
mm crassa, a stipe eccentrice radians, saepe 
divisa profunda vel laciniata lobis ad 20(-25) 
cm longis et ad 15 cm latis, margo laevis, con- 
vex ad circularis, pagina ad Iucem superior 
sanguinea, pagina inferior cinerea fibrosa (per 
rhizoidea septata) in plantis magnioribus 
i-2 cm a substrato. 



Type locality. Pondalowie Hay, Yorke Penin- 
sula, S. Aust. (2-3 m deep in shade. 14 ILtSBl; 
S AY. Clarke). 

type- ADU, A5Z035 (Fig. 1) [solypcs to be 
distributed as No. 214 in "Munnc Algae of 
Minthern Australia", 

Sotulcropclta is named to commemorate 
Otto Wilhelm Sondcr ( I SI 2- 1 SSI), combined 
wilh the peltate form of the well-developed 

Distribution; From the Isles of St Fran*. is 
S. Aust. to Waratah Bay, Vic. and around Fas- 
mania, mainly on BOU^h-WAttf coasts in depths 
of 1-25 m, usually in heavy shade. 

Sonderopelta differs from Pcyssonnelia in 
thallus structure, having longitudinal filltffgntfl 
which diverge to both upper and lower sur- 
Faces, whereas Peysstmnclia has a distinct basal 
hypolhallial layer producing filaments above. 
and attachment rhizoids below Ltlwtia, in 
which Suf}dvropclta was placed with some 
doubt by Weber van Bossc, differs in having 
upwardly and downwardly directed filaments 
produced from B central, apparently limited, 
layer pi distinct I v larger filaments; it also 
differs in being fully adherent to the subslrale 
but without producing attachment rhizoids. 

Peyssoncliii anstralis Sondcr 

Pevssonnefia rmvtniiix is a common subtidal 
alga Ofl southern Australian coasts. P gffjfr- 
n'tanu I Agardh ( 1 876, p. ?S7h based ou it 
collection oi' Harvey (327J) Horn George- 

town, Tasmania (type in Herb. Agardh, LD, 
Z709S) which had been earlier referred to the 
European /'. rubra Harvey, becomes a syno- 
nym of P, australis, and P, cocctnea J. Agardh 
(1870, p, 385) from Western Australia (prob- 
ably near Bunbury) (type in Herb. Agardh, 
LD, 27650) is probably also synonymous. 

Denizot (1968, p, 123) placed P* gwmiam 
as a synonym of P, capensis Montague. (1847, 
p. 177) Jrom South Africa (type in PC?) but 
without detailed comments on their identity. 
Denizot regarded the presence of internal cal- 
eilied granules as well as hypobasal calcifica- 
tion as characteristic of P, capensis (as well 
as other features such as the septate rhizoids), 
and Australian specimens appear to be speci- 
fically identical with ones studied from South 
Africa (e.u. Isaac 307 from Teruuiet, near 
Mossel Bay, 13.x. 1954; ADU, A40825). 

Hence P. akstmlls Sondcr should be re- 
yarded as a synonym {along with P. gunniana 
J. Agardh) of P. capensis Montagne. 


We are grateful to Dr Paul Silva and Dr 
Jim Ross for comments on nomenelatural 
aspects, and to Mr S. A. Shepherd and Mr 
S. Mi Clarke for comments on the growth 
habit and habitat o\' Soncleropelta. Dr E. M. 
Ooi don Mills kindly supplied the latin diag- 

Appreciation is expressed by the first author 
to A.R.G.C. tor provision of technical assis- 


Af.Aunu. J. G. (1876) "Species, Cienera et Ordines 
AIjmuhm." Vol 3(1); 1-724, lipicrisis systema- 
tic Plnridearuni. (Lund,) 

Aium uiiih,. J, E« ( I S54 > Phyceae novae et minus 
im-iiii-ic in marihus extraeuroruraifl collectae, 
Ada H. Sot. SVv, t'jwfa, Ser. III. 1, 329-372. 

Dr Sii/ot, Mi < 1968 ) Los Aluucs Floridees en- 
croutantes t a lYwclusion des C'oi jillinacees ) . 
310 |>p. ( Museum national d'Histoirc naturdle: 

Harm v. W H. i 1859) Phvcologia Australia. 
Vol. II. plrties 6M20. (Rccvc: London.) 

MoNUt.NK C (1847) Enumcralto Funporum 
cjiios a CI. Dregc in Africa Meridional! col- 
lects. rlflA. Vr/. Nat. Dot. ill, 7, 166-181. 

Sondir, O. W. ( 1 85^ ) Flanlae Muellerianae, 
Algae, finiuna 25. 657-709. 

WtBhR v\n Hossf. A. M92I > Liste de.s Algues du 
Sibogn. fT. Rhodophyeeue Pi 1. Protoflorideue, 
Nema'ionales, Crvptonemiales. Monogr. Stboga- 
Expect. 59b. 187-310, pie 6-8. 



byMichaelJ. Tyler and Margaret Davies 


Although there are sporadic reports of the discovery of albino frogs, the individuals involved 
generally are stable in their lack of pigment. Exceptions are the observations of European Rana 
species in which the tadpoles derived from albino ova acquired normal pigmentation gradually over 
a period of approximately two weeks. We have not located the description of pigmentation 
developing in albino frogs in later stages in their ontogeny. 


flfi I. 

Partial albino l.tmnotlyntixfes Jnr?o-nh 




Although there are sp° rJ ^ c reports of the dis- 
covery pf albino froys. the tmtividuals involved 
generally arc stable in Ihcir lack of pigment. 
|-:,\ccptions are the observations' J of European 
Ram species in which ihe tadpoles derived from 
albino ova acquired normal pigmentation gradually 
over a period of approximately two weeks. We 
have not located the desct iplion of pigmentation 
developing in albino frogs at later singes in (heir 

On 22.8.79 we received a gravid female i\inut> 
Jvttash.s dtrnwrili collected at Fvunston neai 
Cniwlet, South Australia. The body was dull pink 
with darker areas on the flanks where the internal 
or gum. tunld be seen in partial transparency 
through the bodv Mh. The eyes were pigmented 
normally and for this reason, conform lo the 
description of "partial albino" ftoys in the termi- 
nology Of Pavosi\ SmaJleombe- and Dubois' 1 . 

The female was placed with u group of 
normally pigmented /.. iluntcnli from North Ade- 
laide, On 22.9.79 she muled with one o\' the 
normal males and laid approximately 3.100 piu- 
mented eggs of which 1,170 were infertile. The 
tadpoles were pigmented normally, and the 
■ esuhani metamorphosing frogs were perfectly 
normal in exlernal appearance and behaviour. A 
very hfgfc death rate tn ihe offspring, following 
metamorphosis was attributed to overcrowding at 
a lime when competing requirements for Other 
research animals limited the srace available For 
them. Ten individuals remained in October 1980, 
but they were relatively small (snout to vent 
length 35 mm), and they died over ihe following 
three months. 

In November 1980 the female began lo acquire 
a tew small (3 mm diam.)* circular -pots of black 
pigment Initially these were confined to the skin 
rovi'ting Ihe tibial glands, and the pigment aicus 
expanded there until each ylnnd was pigmented 
entirety. Over a period of several months a dark 
mosaic of pigment then developed upon Ihe 

»Sm;*H*o.nbc, W. A. (1*381. Nature 14| (357V), 

(1949). 1. Genet. 49. 21*40. 

dorsum and limbs (Fig. t », so producing a piebald 
appearance, Unfortunately ihe female died before 
pigmental ion had extended further. At death her 
snout to vent length was 66 mm which is within 
the normal range for the species. 

Wc have been unable to trace in the literature 
oilier examples of the delayed acquisition of pig- 
mentation comparahle HO that reported here. Cer- 
tainly it is a normal event for many frog species 
to change colour and markings in the. course of 
their early ontogeny, but the changes reported here 
occurred at a post-reproductive stage, 

In our search tor records of comparable pheno- 
mena m other verlebrales, il has become apparent 
that the term "partial albinism" as employed for 
frogs by Pavesi\ Smallcombe- and Dubois" has a 
different interpretation lo the "partial albinism" 
of mammals exhibiling the Chediak-Higashi Syn- 
drome (CHS) reported by other authors r-ltt, CHS 
is characterized by a paucity of pigment of the 
eyes and skin (or hairs), as opposed to lack of 
pigment in the skin and presence in the eyes. 

We thank Chris Miller for assistance with the 
rearing of the tadpole;, 

■Padgett, G. Am Leadvr, R. W., Corhara, J. R. & 
O'Mary* C. C, t 19641 Genetics 40. 505-12. 

"Taj 1 lor, R. F. & Furretl, R. K. 1 1 973 • Jed. Proc 
Rostand. J. ' L94M- Anourcs RtV So. 84, 5®r$. -^ ^ 2 2 (Abst >. 

<$a<fecc*, F. (1964>. Polls. Bio). (Pratfic) 10, ,, k| . imicr f Wi( Daiis w , D. & rricur. Hi J. 

23 °- j , r , , r 1 -» = -»o 7i (1977). Lah. Invest. 36, 554-62 
OPuYwtf, Pi MK791. Kend. 1st Lomb 2, 528-34. 

uDilhnU A (1979) Mitt. Zoo!. M-js Berlin 55, 'oprieun P. J. & Collier, L. L. [1978}, Am I. 

S o_ 87 ' Pathol. 90. 533-6. 

Mkumi J- TYLER ara» . Davit*, Ocpanmcnl of Zoology. University of Adelaide, Box 498 
O.F.O.. Adelaide S. Aust, 5001. 

VOL. 105, PARTS 3 &4 
11 DECEMBER, 1981 

Transactions of the 

Royal Society of South 



De Deckker, P. Taxonomy and ecological notes of some ostracods from Aus- 
tralian inland waters - - - - - - - - 91 

Beveridge, I. Three new species of Calostaitrus (Cestoda: Davaineidac) from 

the New Guinea wallaby Dorcopsis veterum - - - 139 

Tyler, M. J., Davies, M. & Martin, A. A. Frog fauna of the Northern Territory: 

new distributional records and the description of a new species 149 

Baker, A, N. & Devaney, D. M. New records of Ophiuroidea (Echinodermata) 
from southern Australia, including new species of Ophiacantha 
and Ophionereis - - - - - - - - -155 

Jenkins, R. J. F. The concept of an 'Ediacaran Period' and its stratigraphic sig- 
nificance in Australia --,--- - 179 

Foster, C. B. & Harris, W. K. Azolla capricornica sp. nov. First Tertiary record 

of Azolla Lamarck (Salviniaceae) in Australia - 195 

Bull, C f M. & King, D. R. A parapatric boundary between two species of reptile 

ticks in the Albany area, Western Australia - 205 

Brief communications: 

Von der Borch, C. C. Recent non-marine dolomite from the coastal plain, south- 
eastern South Australia ------- 209 

Kailola, P. J. & Jones, G. K. First record of Promicrops lanceolatus (Bloch) 

(Pisces: Serranidae) in South Australian waters - - 211 

Lange, R. T. & Reynolds, T. Halo-effects in native vegetation - - - 213 

Smith, J. & Schwaner, T. D. Notes on reproduction by captive Amphibolurus 

nullarbor (Sauria: Agamidae) - - - - - -215 

Smith, M. J. & Rogers, P. A. W. Skulls of Bettongia lesueur (Mammalia: Macro- 

podidae) from a cave in the Flinders Ranges, South Australia - 217 

Zeidler, W. A giant deep-sea squid, Taningia sp., from South Australian 

waters - - - - - 218 

Addendum - - - - - - - - - - - - -218 







Two new ostracod genera Alboa and Gomphodella are described and 11 new species: A. worooa, 
Bennelongia barangaroo, B. pinpi, B. nimala, Australocypris dispar, Strandesia phoenix, Reticypris 
clava, R. kurdimurka, Ilyocypris perigundi, Leptocythere lacustris, and Gomphodella maia. 
Ecological notes are presented for these and an additional eight species, some of which are re- 


by P. De Deckker* 


De DiiCKKER, B, ( 1 98 1 1 Taxonomy and ecological notes of some osuacods from Australian 
inland water* Trans. H< Sot: S Anst I05O). 9M3B. 11 December. 1981. 
Two new 0Sira«Kl eencra Alboa and Got7iph<nlr!la are described and II new species: 
A. wafOOQi IhnncUmgia barangAtdOi & p'mpi, B. nimala, AustNklocypris d&pan Strartfltsw 
phoenix, Re(kvp<is ffafttf, R. kunlinwrka, Ityurvftis peiigwsdi, LtptOC$tll*re facwtirfs, and 
Gomj'hotUlia mala. Ecological notes are presented fur these Olid An additional eight species, 
some of which ate rc-descrihed. 


Ostracods arc common inhabitants o! most 
types of watcrbodics in Australia with the. 
halobiont fauna being particularly diverse here 
compared to other parts of tile world (De 
Deckker I9&1 ). However, taxonomic know- 
ledge of Australian ostracods is incomplete, 
as indicated by the numerous ostracods re- 
ferred to in open nomenclature in publications 
dealing with hrnnologieal surveys, The present 
paper sets out to describe some of the common 
ostracod species found in inland waters by 
providing many illustrations of their shell to 
simplify further identifications, 

The specimens used lor the descriptions here 
are deposited at the South Australian Museum 

Systematic descriptions 

Subclass; OSTRACODA Latreille. 180$ 

Order; PODOCOPIDA Muller, 1894 

Su!>i;riamily: CYPRIDACEA Band, 1845 

Famiiv; CYPRIDIOAE Hand, 1845 


Bronstein, 1947 

Alboa n.gcn 

Type species: Alboa worooa u.sp . ponder mas- 

Diagnosis. Asymmetrical valves: in anterior 
view, greatest width of right valve at 0,5 from 
dorsum and o( left value at 0.7 from dorsum. 
Selvage prominent in left valve and placed at 
0.5 pf width ot inner lamella anteriorly. Peri- 
pheral groove on the outside of selvage in 
left valve. 

Department of Zoology, University of Adelaide 
Present &dilfe*5 Deportment of BiagCOsraphy 
& Geomorpboloyy, Australian National Hnivcr- 
sity. P.O. Box 4, Canberra. ACT, 2(>()U. 

Male maxillai palps strongly asymmetrical; 
lateral lobe and distal part of copulatnry sheath 
boot-shaped. Geniculaie joint between 1st and 
2nd thorocopoda I segments with two unequal 

Det'muion u) U&PM! Alboa meaning egg in 
aboriginal language to refer to the egg-shaped 
shell, 1 

Alboa woroua n.sp. 
FIGS 1-2 

l i m Cypris sydruMa King: Chapman, p 27 
Diagnosis As for genus. 

Desciititloft Compare < Cxtci nal) . Pseudo- 
punctate, oval-shaped with ventral area almost 
Mat except in mouth area which is slightly- 
concave at about 0.4 from anterior. In left 
valve, 0thet tfdtii concavity, anterior to mouth 
region at about 0.2 from anterior Greatest 
height and width al about middle. Left valve 
larger and overlapping right one anteriorly, 
posteriorly and ventrally. In anrenoi view, 
valves strongly asymmetrical, left valve, 
broader and larger; greatest width of left valve 
al 0.7 from dorsum and of right value al. 0.5 
from dorsum. Carapace pilose with a few long 
hairs in posterior area. Flange broadest in left 
valve. Normal pore canals simple and rimmed. 
(Internal). Inner lamellae broadest anteriorly 
ant! ventrally but slightly broader in left valve; 
posteriorly, inner lamella of right valve very 
namivv. In left valve, selvage prominent and at 
about 0.75 of inner lamella's width from outer 
margin. Selvage follows euivatuic of shell 
except anteriorly where it is less curved. Pos- 
teriorly, selvage I ess prominent and at about 
.5 of inner lamella's width, Peripheral narrow 
depression on outside of selvage, fn right valve. 

1 words used in this text are from 
Parr'. (1965) and Cooper 1 1 9f.2 ) 



Fig. i. Alboa worooa n.gen., n.sp. a antennula, b mandible — palp, c mandible — coxale, d maxillula, 
e antenna, f thoracopoda II, g maxilla, female, h maxilla, male, i maxilla — endopodite, male, i 
thoracopoda 1, k hemipenis, 1 furca, m furcal attachment, a-e, h-m: holotype adult male e* para- 
type; adult female. Scale; 100 fi. 



selvage peripheral anil faint except anteriorly 
where it is much sharper and extends further 
than edge of shell. Right valve with taint 
tubercles postcroventially along edge of shell. 
Radial pore canals numerous, narrow and 
straight. Central muscle field consisting of 
forec scars in front and two behind; upper and 
lowest scars in front broad and elongated 
whereas other two almost circular Mandibular 
sears below and in front of central muscle 

Ananuny. Anlennula (Fig. la) 7 -segmented; 
Icneth-width ratio of last six secmetits. 2/3, 
3/1, 1/1.4, 3/2. 175 I, 2.5/1 Most plumose 
nautory setae as long as all segments together. 

Anienna; (Fig. 1e) No obvious sexual dimor- 
phism; three lung claws plus a smaller one: 
nar.ttory setae reaching tip q[ claws. 

Mandible: (Figs lb,c) Mandibular coxale with 
seven teeth; lasi tooth longer than penultimate 
and near its base externally with three setae of 
ditferent sizes; longest one pilose. Endopod 
with n bristle short, narrow and barren; fi 
hrisilc thick, sfoui and tufted; y bristle longer 
than terminal segment, thick and with short 
hairs on external side. 

Rake-like organ: Six or seven shon and sioul 
teeth with an additional bifid one on inner side 
of each rake, 

Maxillula. (Fig. Id) 3rd lobe with top toothed 
Zahnborsten; length width ratio of both palps. 

Maxilla; Sexually dimorphic, in male, palp* 
strongly asymmetrical with right one broader 
( Figs Mij). bt femaJc (Fig. tg) endopod 
with (hrec plumose sclae, longest one in 
middle and Iwo others of unequal length. In 
both sexes, cpipod with five long plumose 
Slrahlen and one shorter plumose one near 
base of plate. 

I horacopoda I tFig ljl Penultimate segment 
weakly divided, 'lun seiac on geniculate joint 
helween 1st and 2nd segment- anterior setu 

Thoracopoda II; (Fig. If) End ." last segment 
with two unequal setae and terminal pincers. 
Hcmipenis; (Fig. 1k) Lateral lobe boot-shuped 
with base rather broad and inner distal end of 
enpulatory sheath ot similar shape to lateral 

Zenker organ: Both ends rounded and with 40 

Furca: (Fig. 11) Pectinate claws unequal with 
posterior claw 0.7 of length of anterior one 
and anterior seta 0.17 of anterior claw, and 
shorter than posterior seta. 

Fureal attachment: (Fig, 1m) Long and nar- 
row with distal end bifurcate; ventral and 
dorsal branches of similar width and length. 
Eye: Cups of natiplius eye fused; dark brown. 
Colour of shell: green. 

holotype adult male 

L H I H 

-l.V I540u *MM RV 1460/' 86fM 

paratype fidult female 

I II I. H 

l.V 1700m IOOO.u RV 1640/* YK0," 

Type locality: Pool on the southern side of 
Light River, 20 m from the bridge on the 
Port Wakefield Road, north or Adelaide, S.A. 
(34 d 33'09"S, J38 to 27'20'*E). 

Deiivuiion oj name: From the aboriginal lan- 
guage, worooa meaning green lor the colour of 
the shell. 

EvnlQgy and Jisirfbittion: A. ivorooa inhabits 
i.tkes .md temporary pools. This species is 
found in fresh waters and its highest salinity 
record is 3.5';, at the type locality. A. worooa 
has also been recorded from the following 
localities- roadside pool, 13 km east of Rocky 
River and Duck Lagoon, both on Kangaroo 
Island, S.A , and Granite pool at Ncwmann's 
Rocks, 140 km cast of Norseman, VV.A 
Additionally, valves of this species have been 
found in subsurface sediments at Birchmore 
Lagoon, Kangaroo Island (in those specimens, 
(he asymmetry of the valves is more pro- 
nounced with the right valve forming a broader 
hump dorsally). This species has also been 
described by Chapman (19191 as Cypn* 
sydneta from Pleistocene ( fule Chapman ) 
sediments from Boneo Swamp near Cape 
Sehanck, Vic. 

Rsttuvks: A, worooa is related to the Hetero- 
cypris species as their anatomy is similar, in 
particular the triangular shape of the right 
maxillar palp m (he male and the boot-shaped 
lateral lobe of the hcmipenis. The asymmetrical 
valves and the presence of faint tubercles on 
the edge o( the valve are also similar tn Hetero- 
cypris species. The major difference is the pre- 

- I V, RV left valve, right v>.tve. L. H 

length, height- 



Fig. 2 Alboa worooa n.gen., n.sp. a RV internal, female paratype. b LV internal, female paratype. 
c RV external, female paratype. d LV external, female paratype. e RV internal, male holotype. f LV 
internal, male holotype. g LV external, male paratype. h RV external, male paratype. i C showing 
RV, female paratype. j C showing LV, female paratype. k C dorsal, male paratype. 1 C ventral, 
male paratype. m C showing RV, male paratype. n C showing RV, posterior detail of h. o C 
anterior, male paratype. p C dorsal, anterior detail of k. Scale: 1 — 500m for a-m; 2 — 50m for n; 
3 — 50m for p. LV, RV = left valve, right valve. C = carapace. All views are lateral ones except 
when indicated. 



scnce of a prominent selvage in A. worooa not 
seen in Jleterocypris species nor in the closely 
allied species grouped in Cyprinotus. This fea- 
ture of the shell is sufficient to warrant separate 
generic distinction. Additionally, the right 
valve of A. worooa is not curved outwards in 
the anterior area compared to Heterocypris and 
Cyprinotus species. 

Subfamily; CYPRTDINAE Baird, 1845 
Bennelongia De Deckker & McKenzie, 1981 
Type species; Bennelongia harpago De Deckker 
& McKenzie, 1981 

Remarks: Bennelongia comprises five Austra- 
lian species: B. harpago De Deckker & 
McKenzie, 1981, B. australis, B. barangaroo 
n.sp. ( = Chlamydotheca bennelong (King) 
sensu Sars 1894, 1896), B. nimala n.sp., B. 
pinpi n.sp. It is likely that Strandesia feuer- 
horni Klie, 1932 and Strandesia fiavescens Klie, 
1932, both described from Indonesia, belong 
to Bennelongia. From the original description 
and illustrations of the valves (Klie 1932), if 
appears that these two species are charac- 
terized by the asymmetrical valves so typical 
of Bennelongia species. Two specimens resem- 
bling S. feuerborni have been collected from 
Cauckingburra Swamp, at Lake Buchanan, 
S.W. of Charters Towers, Queensland. One 
specimen is illustrated in Fig. 9r but no further 
identification has been carried out. 

Bennelongia australis (Brady, 1886) 
1886 Chlamydotheca australis n.sp., Brady, p. 91. 
Diagnosis; Area just behind the beak-like fea- 
ture of the left valve strongly concave and 
outline of hemipenis as in Figs. 6f,j. 

Description: Carapace. (External) Adult: 
Pseudopunctate and pilose carapace, oval with 
flattened ventrum and concave mouth region. 
Asymmetrical valves: left beak-shaped antero- 
ventrally with deeply concave depression pos- 
terior to beak. Right valve almost smoothly 
curved antcroventrally except for narrow beak- 
like flange there. This flange overlapped by 
beak-like anterior area of left valve when 
carapace closed. In dorsal view, carapace egg- 
shaped with both valves slightly pinched 
laterally at about 0.17 from anterior. Simple 
type normal pore canals. Bordering edge of 
right valve anteriorly are a number of small 
quadrate tubercles. 

Juvenile: Ellipsoidal to subtriangular in shape; 
symmetrical valves without beak-like feature, 

and surface of shell cither deeply pitted or 
coarsely reticulated with large wart-like tuber- 
cles present mostly anteriorly and posteriorly. 
Reticulation especially varied in smallest 
juveniles with some large sieve-like plates. 
(Internal) Adult: Inner lamellae broader 
anteriorly than posteriorly. In left valve, sel- 
vage narrow but distant from inner margin 
posteriorly and ventrally; anterovcntrally inner 
list forms broad but short lip-like flap which 
is absent anterodorsally. Deep depression 
anterodorsally near edge of shell and following 
curvature of inner margin anteriorly. It is 
absent in vicinity of lip-like flap. No outer 
list in left valve. Right valve with selvage 
peripheral except in anterovcntral area where 
it is slightly broader and further inside inner 
lamella. Behind mouth region, flange thin but 
broad, and curves outward to almost reach 
tubercles. Outer list faint, running parallel to 
curvature of shell bordered with tubercles in 
front of mouth region and posteroventrally. 
Radial pore canals narrow and straight. Cen- 
tral muscle field consists of broad horizontal 
scar above, two parallel ones below and an- 
other broad one behind. Two additional, 
almost circular scars, one behind middle pos- 
terior one and other behind bottom scar. Two 
broad mandibular scars in front and below. 

In juveniles, inner lamellae of similar width 
all around except in posteroventral area where 
they are slightly broader. Selvage broad, fol- 
lowing curvature of shell and with no lip-like 
flap. Flange narrow but obvious in both valves 
and of same width as outer list which runs 
parallel to curvature of shell. Depression 
caused by some wart-like tubercles on outside 
area are seen inside valves. 
Anatomy. Antenuula: (Fig. 6a) Natatory setae 
as long as all the segments together. Length/ 
width ratios of the terminal six segments are: 
1/1, 5/2, 1.8/1, 2/1, 2.3/1, 2/1. 3 
Antenna: (Fig. 6d) Natatory setae extend to 
lip of claws. Three claws plus shorter one in 
both sexes; one of additional short and thin 
claw attached to terminal segment in male 
is longer and denticulated in female. 
Mandible: (Fig. 6b) Mandibular coxale with 
seven teeth; endopod with long, narrow and 
barren a bristle, thick, stout and pilose ft 
bristle and long pilose y bristle which is twice 
as long as last segment. 

3 The ratios of the 6 segments of B. harpago have 
been inverted in the original description. 



Fig. 3. Bennelongia australis (Brady, 1886) n.gen. a LV internal, male, b RV internal male c LV 
internal, female, d RV internal, female, e LV external, female, f RV external female « LV internal 
juvenile, h RV internal, juvenile, i C showing RV, female, j C showing RV, male, k C dorsal female' 
1 C ventral, female, a-1: Creek pool flowing across the road, 2 km N of Leonora, W.A. Scale: 




Fig. 4. Bennclongia austral is (Brady, 1886) a RV internal, detail anterior of Fig. 3d. b LV 
external, detail anterior of Fig. 3e. c LV internal, detail anterior of Fig. 3a. d C showing RV. 
detail anterior of Fig. 3i. e LV internal, detail central muscle scar area of Fig. 3g. g C dorsal, 
enlargement of Fig. 51. h C ventral, detail anterior of Fig. 5k. f C dorsal, enlargement of Fig. 5o, 
i C ventral, detail anterior of Fig. 3p. j C dorsal, detail of Fig. 4f. a-e: Creek pool flowing 
across the road, 2 km N of Leonora, W.A. f-j: Roadside pool, 5 km S of Cunderdin, W.A. Scale: 
1_100m for a-d, h; 2— 50m for e; 3— 100m for f; 4— 50m for g; 5— 20m for i. 



Fig. 5. Bennelongia australis (Brady, 1886) a RV internal, female, b LV internal, female, c 
LV externa], female, d RV external, female, e C dorsal, female, f C showing RV, female, g RV 
internal, juvenile, h LV internal, juvenile, i LV external, juvenile, j RV external, juvenile, k C 
ventral, female. I C dorsal, juvenile, m C dorsal, juvenile, n C dorsal, juvenile, o C ventral, juvenile, 
p C ventral, detail posterior of k. q RV internal, detail anterior of a. r LV internal, detail anterior 
of b. s RV internal, detail posterior of a. a-s: Roadside pool, 5 km S of Cunderdin, W.A. Scale: 
1—250," for a-o; 2— 100m for p-s. 



Fig. 6. Bennetongia australis (Brady, 1886) a antennula, b mandible — palp, c maxillula— palp and lobes, 
d antenna, e rake-iikc organ, f hemtpenis, g thoracopoda I, h maxilla, male, i maxilla — endopodite* 
male, j hemipenis, k thoracopoda 11, 1 maxilla — endopodite. female, m furcal attaebment. n furca. 
a-b, d-k, m-n: adult male — creek pool flowing across the road, 2 km N of Leonora, W.A. c, 1: 
adult female. Scale: 200m. 



Rake-like organ. (Fig. 6c) Five teeth plus 
another bifid PIW on inside of each rake. 

Maxillula: (Fig 6c) Length/ width ratio of 
palp segments: 4/1, 4/1; 3rd lobe with two 
smooth ZahriborMen and tufted thick seta at 
end of 3tx\ lobe near Zahnborsten. About 17 
plumose StrahiOD on ep^pod plate. 

Maxilla: Sexually dimorphic: in IcmaJi* (Fig. 
$|J ihrce unequal plumose setae at tip of 
endopod; in male (Figs 6h,i) grasping palps 
unequal, broadest one on right side — for 
chactotasy sec Fig, bh. 

Thoracopoda T; (Fig. kg) Geniculate distal 
part of 1st segment with two .setae, proximal 
one being almost twice as long as other Pen- 
ultimate segment weakly divided Inner diStal 
seta of 2nd segment shorter than 0.5 length of 
3rd segment and shorter than distal outer seta 
of terminal segment. Inner distal end of penul- 
timate segment with two unequal setae. Inner 
clisial seta on the 4th segment is about 0.33 
ol length of distal claw. 

Thoracopoda II: (Fag. 6k) Distal setae un- 
equal: large one more than twice length of 
other which is hook-shaped Broad pincers 
present distally. 

Hemipcnis; (Figs, fifjj Lateral 3obe broad 
with inner distal end pointed and ciuvcd in 
ward, Copulatory sbealh broadly triangular 
in shape with round inner distal end reaching 
almost curved tip of lateral lobe. Outermost 
point ol sheath forms hump and coresponds 
ki mid-length of inner side of sheath. 

Zenker organ: With 33 rosettes. 

Furca: (Fig. fin) Claws narrow, long and 

unequal and posterior seta longest of the two. 

Fureal attachment (tig. Grn I Bifurcate at 

distal end; median branch thickest and other 

two branches arched inwards 

Eye: Dark brown with two lateral silver 


('ol\>ur nf shell Green to pale green, 


leuniyjur: adult left valve 

i v iysiv zrjr> 

Hiluli male 

I rf in 

LV lSr»U,-i | (J6IV RV 1740*1 il!M)/i 

adiill female 

LV 222tb ftQOs RV ?Uf.iv 122D> 
Rt'nuirk'i FIxaminauori o( the. tvpc specimens 
of Cltfnmyrtothrra Mtffalk Brady, I SX0 in the 

BiUish Museum necessitates the (ollowing 
clarification. One adult loft vaKe (I I98Q ^ J . 
H 1200V), designed here as the Icctotypc, is 
the Valve probably illustrated by Brady [\Wh) 
on Plana I.X.7. The tip of the beak- like feature 
of this left valve is broken olf In the same 
slide, (here is another left valve which probably 
belongs to a tleicrocvprh sp It is likely that 
this valve is the one thought bv Brady (ISXuj 
to be the right vjlve of C. tiuMrnUs and ill LIS 
si rated by him ot) PLuc IXA It dcffnftdj 
lacks (he peripheral posterovenlral tubercles 
and the broad inner lamella anteriorly so 
typical of all SfMiffoftQla species and the 
aniciovemral flange of B, cnt\(rah'\ specimens 
This would explain the incorrect description 
of the right valve of (his species provided by 
Brady (15B&. p. 9l ) rvuminalion of one 
carapace of &, nwdrafh from a slide bearing 
Brady's handwriting in Sars 1 collection in the 
Oslo Museum further confirms Brady's mis 

Finally, in the same slide from the British 
Museum, there are two partly broken cara- 
paces of smaller Uenntfongtn species (length 
1360 ,,. height XOO „ ) with some dried soil 
parts msidc. Their specific identification re- 
mains uncertain .ilthough it is thought they 
belong to B. karatiynr, 

in 1 894i Sars s> rwirryrnizcd C. amrrdia 
Brady. I &8£ with Cyprif hemh-hvig King, i 855 
maintaining that Brady's specimens were the 
same as those of King, and by stating that the 
latter author had described the species from 
juvenile specimens. This was repeated by Sars 
MS9(V) and Henry (1923* Sars' suggestion 
cannot be accepted because King f 1 X55) 
stated on page 63 that C hemwUitig has 
^equal valves', Thk [$ not the ease lor adult 
i(cn/irlofn;hi speci.-s. Sars' ( 1894) argument of 
kmg\ specimens being juveniles cannot be 
accepted here either since juveniles of BfHR&- 
hfng'in (which have symmetrical vatvcs'l are 
cither deeply punctated all over or arc strongly 
ruherculatcd. These features are best seen on 
vpeeimeiiN described by Sars (1-896) ftg Cygrk 
btttratik King 1X55 which arc juvenile* of 
fSsntirh'tmnt >pp. Fither tubercles or pitted 
shell would have surely been diagnosed by 
king otherwise. 

Bats specimens described from Austral 
(Sars 189h) mm] New Zealand (Sars 1894) as 
<\ iHtunrlom: ar'c iruc Bfnntfatrjihi species but 
do not belong to ft, mistfdUS- Since they cannot 
In iJ; iMcd as king's species, they arc there- 



fore renamed here a* B barnn^ifyo n.sp. This 
species is described below 

Ecology and tlLsfrilmtion; B. ctt4\tralis was 
originally collected from Penola, S.A. by Prof. 
R. Tate-. The SpoouWtW were empty shells 
(Brady 18S6). This specie*, inhabits mainly 
temporary pcoK and, w tor; has been found 
alive only in Western Australia and South 
Australia Adult males have been found in a 
permanent hike in Western Australia, suggest 
ing that the mode or reproduction is parthc- 
Oogenetic in ephemeral environments. 
Localities' W A. Roadside pool 2 3 km S of 
Noilheliffe, roadside pool PfeiRer's Road, 8 
km from Mary Pe^ks, lake Sadie, east of 
Wilson Inlei (near Denmark); creek pool flow- 
ing across The road 2 km north Of Leonora: 
roadside pool 5 km S of Cunderdin; roadside 
pool on eastern side of road between Quai- 
nuling and Corhgin (25 *km northwest oT 
( onigin): Lake Biddy S.A .: Roadside pool. 1 
km east of Rocky Rivet, Kangaroo Island. 

Wale i was fresh except in the last two sites 
in Western Australia wTuk- salinity was 4.4 
and 35$, respectively. 

lUniieloikeiia baraucaroo n.sp. 
I IGS?-8 T 9n-q 
1894 Cypfts bvnuttims King San., p. Z4. 

1896 Cyprh bennchmx King: Sari p. 49. 
]8'»n Cyprh luleruria King; S;u".. p, 53 
1923 Cypn\ I'rriiicliitrj KirifJ. WfcnrV. P. Tf$> 
Dfdgtttfte Area just behind heak-like feMurc 
of left valve slightly concave; outline of hemi- 
penis as in Pig. 8j. 

Btesertprlarr. Campace (External) Adult; cir- it) oval carapace smooth or covered with 
faint pustules and pilose. Vcotrum flattened. 
V,.!vi>-, ..yvmnK'tnc.i!' left valve larger, espe- 
cially anteriorly with largest overlap ventrally; 
anteroventral region of left valve slightly cor> 
civi in form beak-like feature whereas right 
valve broadly rounded and with an elongated 
.in- 1 narrow beak-like serrated flange anfero- 
ventrally. Simple type normal pore canals. 
fuvcnilc; More elongated in lateral view, with 
symmetrical, pitted or tuberculattd valves, 
(Internal) Adult ' Inner lamella twice as hroad 
anteriorly in both valves, selvage peripheral 
veiitially, away from outer margin posteriorly 
and especially anteriorly: inner list forming lip- 
like flap far away from edge of shell antcrn- 
ventrally which js preceded by a deep. narrow 
PW*& A ri.ltu- fnllows curvature of inner 
inarL'iu ai'ileiinO\ bU| fades opposite fonguc- 

likc flap. Few small tubercles are visible on 
this ridge above concave depression of shell 
Oulcr lisl often broad posterovenlrally. In 
right valve, selvage peripheral and sharp pos- 
teriorly, broad anterovcnnally and faint antero- 
dorsally. In many specimens, edge of shell 
near flange distorted to form an obvious con- 
cavity which is paralleled by a depression. A 
row of tubercles along periphery of shell pos- 
tcroveiUrally. Outer list also broader postcro- 
venlrally and usually pitted externally near 
edge Of shell; this is best seen in ventral area. 
Juvenile: In both valves* inner lamella of 
almost equal width all around and selvage, 
which follows the curvature of the shell, 

Auofonw: Only the features ol the anatomy 
which differ significantly from ti. aastralts and 
other Benuelongia species are mentioned. For 
other details refer to Fig. S. 

Antentitila: Natatory setae slightly longer than 
all segments together. 

Maxilla; Male grapsing palps (Figs 8g,h) of 
different shape than B. australis: left palp 
shorter and broader and right one with outer 
edge forming 90 a , 

Thoracopoda I: (Figs Sd,c) Inner distal seta 
of second segment at least as long as half of 
3rd segment; longest inner seta tt mid-length 
Of 3rd segment (where it is weakly divided) 
at least as long as 2nd half of 3rd and 4th 
segments together Tnner distal seta on 4th 
segment I '2.3 of length of dislal claw. 

Colour oj shell: Green. 


holotvpe aitllll male 

L H I H 

LV 112(V 700* RV 1100m W.0* 

paraiype adult female 

L n L H 

LV 1 I 60m 700/1 RV IMQji 680/. 

Ncwmann\ ftorfa fctQlttl It-male 

L H I H 

LV 1390" TH0* RV Il20iu 770m 

pprnutrtnn *«/ name: From the aboriginal name 

of Bennelong*s "wife Barangaroo 

Tvpr laeatftt; Lake Bueluman. Old (2T35S, 

145 52'H). 

Lr.oloav and distrduitton, B. barangaroo is a 
common inhabitant of temporary pools z^nd 
usually is only represented by parthenogcnetie 
knulcs. On one occasion, in Lake Buchanan, 
both sexes were Pound. There salinity Was 



Fig. 7. Bennelongia barangaroo, n.sp. a LV internal,, holotype male, b RV internal, holotype 
male, c LV external, paratype female, d RV external, paratype female, e C dorsal, paratype female. 
f LV external, paratype male, g RV external, paratype male, h C dorsal, juvenile, i C showing 
RV, paratype male, j C showing RV, paratype female, k C dorsal, paratype male. 1 C ventral, para- 
type female, m C ventral, juvenile, n LV internal, detail anterior of paratype female, o RV 
internal, detail anterior of paratype female, p C showing RV, detail anterior of j. q RV external, 
detail anterior of d. r RV internal, detail posterior of b. Scale: 1 — 250m for a-m; 2 — 100/* for n-o- 
50m for q; 3— 50m for p; 4— 100m for r. 



Fig. 8. Bennelongia barangaroo, n.sp. a anlennula, b antenna, c mandible — coxale, d thora- 
copoda I, e maxillula — palp and lobes, f mandible — palp, g maxilla — endopodile, male, h maxilla; 
male, i thorucopoda II, j hemipenis, k furca, 1 maxilla — endopodite, female, a-b, d-k: holotype 
aduli male; c 1: paratype adult female. Scale: 100m. 



Fig. 9. Bennelongia barangaroo n.sp. a LV internal, female, b RV internal, female, c LV ex- 
ternal, female d RV external, female, e C dorsal, female, f C dorsal, juvenile, g C showing LV, 
juvenile, h C showing RV, female, i LV internal juvenile, j RV internal juvenile, k C ventral] 
female. 1 C ventral, juvenile, m LV internal, posterior detail of a. o RV internal, anterior detail 
of b. p LV internal, anterior detail of a. q C dorsal, detail of e. a-q: elongated pool in creek bed, 
about 25 km N of Cue, W.A. Bennelongia sp. r C showing RV. Cauckingburra Swamp, at Lake 
Buchanan, via Charters Towers, Qld. Scale: 1 — 500a for a-k; 2 — 100m for m-p: — 20m for a- 
3— 500m for r. 4 ' 



4.l%«. In other localities listed below, water 
w:is known to be fresh. 

I ivc specimens with tubcrculated symmetri- 
cal valves collected m a farm dam at Frorne 
Downs near Lake Prome, S..A. were examined 
in the laboratory. TTiey resembled specimens 
illustrated by Sars (1396a) on Plate VII Hg 
3 and labelled by him as Cyprfy Uiterarsa King, 
1855. After a lew days, these specimens, which 
had traC^ ^ ovaries inside the valves, Wdtfe 
seen lo molt into B. banwgaroo with jts typical 
asymmetrical valves. JSggfl of the latter 
developed inio small highly ornamented ostta- 
cods like C. latcmrith This phenomenon 
requires the two tax a To be synonymized. 
Localities: WA. ; granite pool, Ncwmann't 
Rocks, HO km E of Norseman, roadside ditch 
31 km W of Espcrance (road to Ravcm 
thorpe) and 3.5 km east or Dalyup River: 
small roadside pool about 18 km northeast rjj 
Menzies; elongated pool in bed of Cookarrow 
Creek, about 5 km W of Wikma; pool in creels 
bed about 25 km N of Cue; small farm dam 
on eastern side of road 9 km S of Cunderdin 
o\-\ the Way lo Quairadine. S.A.. Farm dffOI 
at Frorne Downs, near Lake Frorne. Old: 
Creek. 22 km E of Richmond; roadside poo! 
at Miranda: Lake Galilee, near Aramae: L<ike 
Dunn. SOUth of Lake Galilee N.S.W.: Speci- 
mens raised by Sars from sample of dried mud 
collected in walerhotes by Mr Whitelegge itt 
Bourke Street, Sydney (Sars 1806). New Zea- 
land: Specimens raised from dried mud sample 
collccred near Kaitaia in the "North Island 
Chapman (196$) stared that ibis species had 
not been ioumi in New Zealand since Sars -1 
f 1 894) description and no further localities 
arc provided in Chapman it Lewis (1976). 

Remark sv B baron garoo is closely allied to 
B. austrotfs but the species can be separated 
oil the basis ot size (ft, austraJis is much 
larger), on the outline of the hemfpems and 
The cbaclotaxy ol the ihoracopoda I. 

The specimens of B. baranf>an>n described 
here arc identical in morphology to Sars* 
specimens from Bourke Street Sydney and 
from New Zealand. It is assumed here that the 
specimens reported as Cyprh lateraria by Sars 
(1896) from the Sydney site are juveniles of 
B. hanwgnroo 

The shape and width of the antcroventral 
flange on the right valve of B. bcran^aroo can 
vary extensively: in the specimens from Lake 
Buchanan, the edge of the flange is serrated 
(Figs 7p.q). 

Bennelongia nlmala rusp. 

Diagnosis: Oval to subrectangular, pustulose 
carapace with posterior slightly pointed and 
ending with one or two spines. Hump-like 
thickening uf shell anrcrudorsally. Posterior 
sola of furca about 0.7 of length of posterior 
claw and lateral outline of hemipenis as in 
Fie UK 

fh",friptioft: Carapace, (Hxternal) Adult: Oval 
in subreeiangular-srmped with posicrior slightly 
pointed and ending with one or two spines. In 
dorsal view, e ^-shaped with anterior end 
narrow and more pointed than posterior. 
Valves ohiousl} asymmetrical in antcroventral 
area: there, left valve formed like pointed 
beak whereas in right valve, it h broadly 
rounded and there i& a small beak-shaped ser- 
rated flange. Posterior to flange, edge of right 
valve slightly concave. Tip of beak does not 
reach horizontal plane formed by ventral area. 
Greatest height ai about 0.33 from anterior. 
Shell pilose, pseudopunctaled and pustulose 
neatly all over Along edge ot left valve, 
especially, pustules more concentrated and 
some arc pointed, especially anteriorly 
Juvenile. Pseudopunclate subiriangular shell 
with many pointed tubercles scattered all over 
but with greater concentration anteriorly and 
posteriorly Valves symmetrical. Tn dorsal view. 
oval-shaped with hotb ends pointed. 
{ Internal) Adult Inner lamella broadest 
anteriorly in both valves. In left valve selvage 
narrow and peripheral; inner list forming lip- 
like flap antcrovcntrally, Tins flap fades in 
anterodorsal region. In front of flap in ventral 
area, a deep narrow groove Between outer 
margin and just behind beak ventrally, is a 
ridge which is mberculate in some specimens. 
In anterior area, in front of hinge area, is a 
deep groove running parallel to curvature of 
shell which is placed near outer margin. This 
groove absent in beak-like area. In right valve. 
selvage narrow and runs parallel to curvature 
of shell except, at mid-height anteriorly where 
ii ft no* visible. Anteriorly, in region where 
serrated flange present, a small ridge runs 
parallel to, z^nd between, selvage and inner 

Juvenile; Inner lamellae equal in boiTi Valves 

and of same width all along. Selvage broad and 


Anatomy: Same remarks as for & kty0l$$r&o 

lor details ot anatomy, refer to Fig, 11, 



Fig. 10. Bennelongia nimala n.sp. a LV internal, female paratype. b RV internal, female para- 
type, c C dorsal, female paratype. d C showing RV, male, e C showing RV, female paratype. 
f C showing LV, female paratype. g RV internal, male holotype. h C ventral, male paratype. i C 
dorsal, juvenile, j RV internal, anterior detail of g. k LV internal, anterior detail of a. 1 LV internal, 
anterior detail of male paratype. m C showing RV, anterior detail of d. n C dorsal, anterior de- 
tail of c. o LV internal, posterior detail of male paratype. a-c, e-h, j-1, n-o: Georgetown Lagoon, 
near Jabiru, N.T. (type locality), d, i, m: Buffalo Billabong, near Jabiru, N.T. Scale: 1— 500m for 
a-i; 2— 200m for j-m, o; 3—100" for n. 



Fig. 1 1 . Bennelongia nimala n.sp. a antenna, b rake-like organ, c antennula, d maxillula — palp 
and lobes, e mandible — palp, f maxilla — cndopodite, male, g hemipenis, h maxilla — cndopodite, 
male, i thoracopoda I, j thoracopoda II, k maxilla — endopodite, male, 1 maxilla— protopodite, female, 
m furcaJ attachment, n furca. a-c, e-g, i, k-m: holotype adult male; d, h, j, n: paratvpe adult male. 
Scale: 1 — 100m for a, c-n; 2— 50m for b. 



Antennula: (Fig. lie) Natatory setae as long 
as all segments together. 

Maxilla: Male palps asymmetrical (Figs 
llf,k) with right one broader and less arched. 
Two external setae on female palp (Fig. llh) 
of equal length and plumose. 

Thoracopoda I: (Fig. Hi) Inner distal seta 
of 2nd segment almost reaches weakly divided 
area in middle of 3rd segment where another 
seta of similar length occurs. Inner distal seta 
of 4th segment 1/5.6 of length of distal claw. 
Hemipenis: (Fig. llg) Lateral lobe broad: 
greatest length equals its greatest width. Near 
base of lateral lobe on inner side, copulatory 
sheath in form of circular lump. 
Zenker organ: Funnel-shaped at both ends, 
with 30 rosettes. 

Furca: (Fig. 1 In) Posterior seta long, approxi- 
mately 0.7 length of posterior claw. 
Colour of shell: Green to dark green with 
broad transversal white stripes best seen in 
dorsal view. 


holotype adult male 

L H L H 

LV 1500^ 840m RV 1340m 760m 

paratype adult female 

L H L H 

LV 1640> 960m RV 1540m 860m 

Type locality: Georgetown Lagoon, Jabiru, 
Northern Territory. 

Derivation of name: The word nimala in abori- 
language of the Northern Territory means 
lagoon, for the typical habitat of this species. 
Ecology and distribution: B. nimala is a fresh- 
water species found in lagoons near Jabiru, 
east of Darwin, N.T.: Jabiluka Billabong, Buf- 
falo Billabong and Mudginberri Lagoon. Both 
sexes were always found in the collections. 

Bennelongia pinpi n.sp. 
FIGS 12-13 
Diagnosis: Anteroventral area of left valve 
strongly beak-shaped extending below hori- 
zontal plane of flat ventral area. In dorsal 
view, carapace almost circular, with both ends 
broadly pointed. Outline of hemipenis as in 
Fig. 13e. 

Description: Carapace. (External) Adult: tri- 
angular to semicircular with flat ventral area; 
greatest height at about middle. Anteroventral 
area of both valves beak-like but more pro- 
nounced in left valve. Surface of shell smooth 
to pseudopunctate and barren of hairs except 

in mouth region. In dorsal view, carapace 
almost circular with anterior and posterior 
ends broadly pointed. At about 0.17 from 
anterior, shell pinched, being more noticeable 
in posteroventral area. Valves strongly asym- 
metrical anteroventrally where left valve larger, 
elsewhere left valve slightly overlapping right 

Juvenile: Subtriangular with ventrum almost 
flat and greatest height at about 0.4 from 
anterior. In dorsal view, carapace oval to 
almost circular with both ends pointed. Largest 
juveniles with pseudopunctated shell and few 
tubercles. In very small juveniles, shell pitted 
and thinly reticulated and with some sieve 
plates. Most tubercles are cone-shaped and a 
hair protrudes from each of them. Valves 
almost symmetrical with left slightly larger. 

(Internal) Adult: Inner lamella much broader 
anteriorly in both valves. In left valve, inner 
list forms broad lip anteroventrally and is 
absent anterodorsally. Selvage narrow ventrally 
and distant from outer margin and broader 
posteroventrally where it is placed in middle 
of inner lamella. Above concavity behind beak- 
like form in anterior of shell, there are a few 
tubercles between outer margin and lip-like 
inner list. Depression in front of lip-like inner 
list and another near outer margin antero- 
dorsally. In right valve, selvage narrow and 
peripheral. A broad tongue-like flange antero- 
ventrally in some valves which is enclosed by 
broader beak-like left valve when carapace 
closed. An outer list runs parallel to curvature 
of right valve and bordered by small tubercles 
ventrally except in mouth region. 

Juveniles: Inner lamellae twice as broad 
anteriorly and with broad and prominent sel- 
vage following curvature of shell. Outer list 
runs parallel to curvature of both valves and 
bordered inside by faint tubercles in some 

Anatomy: As for other species. For details of 
the anatomy, see Fig. 13. 

Antennula: (Fig. 13b) Natatory setae almost 
as long as all the segments together. 

Maxillula: Male palps (Figs 13i,j) asym- 
metrical with right one broader although both 
are similarly arched. 

Thoracopoda I: (Fig. 13g) Inner distal seta 
on 2nd segment almost reaches level of weak 
division of 3rd segment. There, inner distal 
seta long, extending past 4th segment. Length 



Fig. 12. Bcnnelongia pinpi, n.sp. a RV external, female paratype. b LV external, female para- 
type, c LV internal, female paratype. d RV internal, female paratype. e C ventral, female para- 
type. f RV external, male holotype. g LV external, male holotype. h C dorsal, male paratype. 
i C showing RV, male paratype. j C showing RV, female paratype. k C showing RV, juvenile. 1 C 
showing RV, juvenile, m C dorsal, juvenile, n C ventral, juvenile, o C showing RV, juvenile, p 
RV internal, juvenile, q LV internal, juvenile, r C ventral, juvenile, s C dorsal, juvenile, t C ven- 
tral, enlargement of r. Scale: 1 — 500m for a-p, 1-n, p-s; 2 — 200/* for k; — 300/* for t; 3 — 150/* for o. 



Fig. 13. Bennehngia pinpi, n.sp. a maxillula — palp and lobes, b antennula. c antenna, d man- 
dible — palp, e hemipem's, f maxilla — endopodite, female, g thoraeopoda I, h thoracopoda II, i 
maxilla, male, j maxilla — endopodite, male, k furca, I furcal attachment, a-c, e, g-k: holotype adult 
male; d, f, 1; paralype adult female. Scale: 200m. 



ratio pl ttisla) seta of 4ih Segment and distal 
claw is 1/3.5, 

Hemipenis (Tig. 13c) Outer lobe broad and 
digitate and. near its base on inner side, 
copukuory sheath forms broad trapezoid- 
shaped extension. 
Zenker organ: With 42 rosettes. 
Furea; {Fig, 12k) Setae almost equal-pos- 
terior one longer and about 0.5 length of 
posterior eluw. 

c oloiu of shell. Light green. 
hololypi: adult male 

f II I H 

I V 24BM'i KV ttfbt* MZW 

innaiV(K.' adult female 

L H L H 

LV 3SB0A I 640m RV 2440m |52(.)/i 

Type locality: Pine Tree Creek Lagoon. N of 
Muj-Jicndcn, and exactly \t> km S of Loiusa 
Lake on road to Ktighendeo. Old (2O'W30" 
S. |44 16'44'R). 

Derivation of name: The aboriginal word pinpi 
in Queensland meaning parrot is chosen for 
th« nnteroventral area of the shell which 
resembles the beak of a parrot. 

t.colo&v and distribution: This freshwater 
species is known from three other localities in 
Queensland; Agues Lake, Salt Lake (fresh) 
and Lomsa Lake, all three N of HughendeO 

Subfamily; EUCYPRID1NAE Bronstcin, 


Ativtralocypriv Oe Decided, 1974 

type species: Auslrnhcypris robusta Dc 
Dcekker. V)7A. 

Remarks: The following species are discussed 
in the present work and can be distinguished 
on the outline ot the hemipenis; A dispar 
n sp.. A itisalaris ( — A. hvpersalina) » A , 
rertttnvuiatis and A. robrtsta. All Australa- 
c\ pris species arc lialobiont and pfanktic but 
can also crawl on lake floors. Rarely do two 
species occur together in one lake. 

Austnilocypris dispar n sp. 
FIGS 14, 1 5 a-i 

l>iu,i?no\i\: Lateral lobe of hemipenis digitate 
;md broad at base; long distal seta on 4th seg- 
ment ol ihoracopoda 1, 

Description: Carapace. (External) Pscudo- 
puncUtc, subrcctanyular carapace with an- 

terior end broadly rounded, dorsum slightly 
inclined and posterior steeply inclined. Ven- 
tolin strongly concave past mid-length from 
the anterior. Greatest height at about 0.4 in 
female and 0.33 In male, In dorsal view, cara- 
pace elongated, length more than twice width. 
and both extremities slightly pointed. Lett 
valve slightly larger in female and ventral 
overlap minimal. Both valves extend over one 
artOthCJ in anterodorsal area just before hinge. 
Simple normal pore canals. 
(Jnicrnal) Inner lamella narrow in both valves 
and selvage fainl and between edge of valve 
and selvage which is prominent in that area. 
Radial pore canals numerous and straight. 
Central muscle held consisting of five adductor 
scars and two large mandibular ones in front 
and below. 

Anatomy: Aniennula: (Fig. 14c) Length.' 
width ratio of last six segments. 1/3, 2/1, 

I / 1 . 5/ 3.5. 2/ 1 .4, 4/3. Small waU-hke 
"sensory" organ on side of 2nd segment. Nata- 
toiy setae as long as las! six segments together. 
Antenna: (Fig. 1 4a) Massive and sexually 
dimorphic; lour claws in male and three in 
female; smallest one in male attached lo last 
segment and with long comb-like teeth. Nata- 
tory setae almost reaching tip of claws. 

Mandible: Mandibular eoxale (Fig. I4h) with 
seven teeth, the last one longer, narrower than 
pcnultimale and. at base, three setae occur, 
two of which are pilose Lndopod (Fig. I4d) 
with a bristle short, narrow and barren; fi 
bristle stout and finely pilose y bristle twice 
as long as others and pilose m distal half. 
Distal segment of endopod 0.i3 o( length of 
penultimate segment. 

Rakc-like organ:. Seven short and stout teeth 
plus one inner bifid one on each rake. 

Maxtllula: (Fig- 14b) Endopod with 23 Srrahlen; length widlh ratio of palps: 
.3/1.3, 1.5/ 1.1 with last segment slightly tra- 
pezoidal. Two toothed Zahnborsten on 3rd 


Maxilla: Sexually dimorphic: in male (Figs 
I4j,k), palps almost symmetrical, strongly 
curved and narrow; in female < Fig. Uf ), 
palp faintly divided at its c.\tienut\ and 
with three short pilose setae, the middle one 
being twice as long as others. 

Thoracopoda 1: (Fig. 14c) Penultimate seg- 
ment weakly divided; distal segment with long 
inner sc(a 0.5 length of distal claw. Thoraeo- 



Fig. 14. Australocypris dispar n.sp. a antenna, b maxillula — palp and lobes, c antennula, d mandible — 
palp, c thoracopoda I, f maxilla — endopodite, female, g thoracopoda II, h mandible — coxalc, i 
hemipenis, j maxilla — endopodite, male, k maxilla, male, I furca, m furcal attachment, a, c-e, g-m: 
holoiype adult male; b, f: paratype adult female. Scale: 200«. 



Fig. 15. Australocypris dispar n.sp. a LV internal, female paratype, b RV internal, female paratype. 
c LV internal, male holotype. d RV internal, male holotype. e C showing LV, female paratype. f 
C showing RV, male paratype. g C dorsal, female paratype. h C dorsal, male paratype. i C ventral, 
female paratype. Strandesia phoenix n.sp. j LV internal, female holotype. k RV internal, female 
holotype. 1 C showing LV, female paratype. m C showing RV, female paratype. n C dorsal, female 
paratype. o C anterior, detail of p. p C anterior, female paratype. q C ventral, female para- 
type. Scale: 1—1000^ for a-i; 2— 200/* for j-n, p-q; 3—20^ for o. 


H, l)h OB KKbR 

pOda II: (Fig. I4g) Elongate and narrow seg- 
ttKMiiN «f islul segment with two terminal setae 
and two pincers; shortest setii hook-shaped. 

Hcmipems; (Pjft. I 4i ) Lateral lobe digitate and 
Slightly curved inwards and bioad at its base; 
copulatory sheath broadly heart-shaped. 
Zenker organ; Lung and narrow with both 
ends rounded and usually 67 roseltcs. 

lurea: ([-it;. 141) Unequal claws with anterior 
one 0.66 longer, an J anterior sola slightly 
longer than posterior one. 

I -ureal attachment: (Pig, 14m) Proximal part 
bifurcate; branch strongly arched and 
about same length as ventral one which follows 
curvature of median branch. 

Colour of shi'lh Nacreous brown 

Si. <•; 

bulolypc adult male 

I H I H 

i v ^o-i;-. ::o(v kv \um> m^ 

paralvr»c .Mull frm. it. • 

I H 1. J I 

I V 4 toil/' 2240a RV 4000/' 2060> 

type lottiloy Ephemeral sail lake south of the 
CoOKWIfl Lagoon. .S.A. < 36 " 2b 4S"S, 139 u 47' 
32"E] |ut f 5 in IX- Dcckker &Geddcs, 1M80, 
collected on 1ft XT 978 at I5.5#, salinity. 

QffrtvgtfOt uf name: From Latin <Jnpat mean- 
ing Jittercnt ior the unusual long distal WU 
cm the tiro Segment Ot the thoraeopoda I com- 
pared to congeners. 

Lr,>hr.;\ tuui JiM.'ihttrion: A. dixpav is a halo- 
biont species which occurs in ephemeral salme 
lakes m S.A "Near the Cooroue Lagoon 
iis salinity range is 52',,, De fXvUur & 
Cicddcs (IViSOi provide further details on its 

dJBtrtb ,, there as ".4. n.sp.L" Ir fa also 

recorded on the Voike Peninsula at tv,„ in 
■ui ephtmwai lake ( 35 03 1 ]"S, 137 \35?0"r.! 
and on the Tyre Peninsula m fl swamp south 
i't ihc road near Lake Horn, northwcsl Of Pori 
i incoln &i 32.4*;, 

Rrtnnrk'i; A tit spur is characterj/ed by the 
unusually long seta on the tlistal segment of the 
thoraeopoda I and the faintly divided end of 
the lemale maxillar cudopodile palp These 
characteristics are unknown in other Australu- 
cr/vrv species bill are found in the haJobiont 
nxtracad L-muituwyprix luridnx Shornikov, 
■ I. described from the Kuban Delta in the 
Caspian SC0 region "f (he USSR. Conospon- 
C With D C. Shornikov has led to the con- 
eliiNHHi that other features of the chactotavy of 

many appendages between A. dispar and /.. 
luridus differ loo greatly to justify close rela- 
tionship between the two species. 

Austrutorypri.s nwidwis (Chapman. 1966) 

I'lfih t tfiyfttt itisuhn's; Chapman, p. 373 
/974 .in\tnihu yprfa hviH-txolimt: De Dcckker. p 

I97S dustmlQttyprtu hyffmAtiMti Oc Dcckker, p. 

1978 l-MY'./Aw/-/,;. msuiuri.s: De Deckker % p 17 

f'lt'Huks, Alter examination of many collcc- 
lions of A. inuihinx from Western Australia 
(trom where the species had originally been 
descnbcil), it became clear that A insularis 
and A. hyprrwlinu arc cOjlSpceific. Both taxa 
h;oe very similar hemipencs with Ihc charac- 
teristic tno.ul imd curved lateral lobe and the 
bulbous tuner distal end of the copulatory 
Sheath This iy,.e oi laleral oulline of the hemi- 
pems is not found ill congeners. 

flic examination o{ many specimens of 
Auvffitfttt ■ yf>?i\ has led to Ihe conclusion that 
there art Slight differences m ihc morphology 
of many specimens of ,4, iti\ulu>i\ and A. 
hypct-Mslino Therefore the minor differences 
of the outline of the hemipenis for ihc two 
InXfl originally mentioned by De Dcckkei 
tls»7fi) are here considered insignificant. 

During the revision of all myldocypridmid 

species 1 1 jc Dericker i i >7*>. It appeared that 

I IftStliurt* and A. hyprrsalina differed mainly 
on eharaeiLis of the shell, but it has since 
become appareni that the shape of the shell of 
th jttfSfPaloCjrpfts species is also variable in 
populations taken during different seasons in 
the .same lake. Large specimens collected dur- 
ing winter and early spring months are usually 
more elongated. For the same reason, the 

p'-ciniLns illustrated in De Dcckker, 1978, 
m Figs 18a,b as A itvfu'rsnjina show the 
typical Wllltei form of A hyper\ ( ditta A, 
in\uhni\ not recognized then, as most collec- 
tions described rn lhal publication had been 
taken during summer months. 

ln-oloi'y ntuJ ilistrihMu.n: The distribution of 
A insularis (plus .4. hypctsalina) Was dealt 
with m De Ueckker (1977. I97S ) and is 
Upd&tCd line. The species is commonly found 
ill ephemeral saline lakes near t tie Coorong 
lagoon (see De Dcckker & Gcddcs 1980) 
where it is (bund between 5 and 131',, salinity, 
ir also occurs on the Yorke and Evre Penin- 
sula in simitar ephemeral hikes at the same 
^ahnity range It has also been widely enl\uol 



in 15 Wcsl Australian wtli lakes by Geddes 
(i nl. (1981.) over the range of lM-\22A 

Austmlocvptis rectanetilarix Tte Deckker, 


p)7N An\trtil<tcv!>t'ts'' nciutnuiluri^: De Deekkci. 

P. lf t 
11)80 .■*'iStraIi.>Lvpri.s- rfrwttKittoris: Dc Deckker & 

Geddes. p. 6^1. 
ni<t}:nasix: Lateral lobe l?f hemi penis, "narrow 
and hook-shaped; copulatory sheath almost 
semicircular. Carapace rectangular. 
Remarks: The origins! description of this 
species was incomplete because a number of 
appendages had dried out and had been 
damaged. A number of specimens have .since 
been collected in Saline lakes near the Coorong 
lagoon by Dc Deckker & Geddes (1980), 
They examined the appendages of this species 
and <m morphological grounds referred it with 
confidence to Austrahcyprix 
Ecology and dislri but ion: This species ia a 
true halobiont form which has never been 
tound in salinities below W/',,- Near the 
Coorong Lagoon, its salinity range is 5»V 
195";, and there is evidence there that it even 
hatched above 73% and 115'/, in two dif- 
ferent localities. On the Yorke and Eyre 
Peninsulas, A renew xtdarix occurs in the same 
range o\ : salinities as in the lakes near the 
Coorong Lagoon. 

AusTratocypri\ whnsta Dc Deckker, 1974 
1974 Auttraloeypris mhu^at: Dc Deckker, 1974 
Diagnosis: Chitinous pocket on inside of pos- 
lerodorsal area in female valves; hemipems 
with dieitate lateral lobe which is of about 
same width all along. 

(■colony and distribution: The salinity range of 
A. rahUSt& in Victoria, already available in 
ftaylv & Williams (1966). of 53 .5-93.1 r ,„ 
a, n J of 4-4- I32 r ; , in Geddes (1 976) is 
broadened to the range of 7-145';,, obtained* 
from collections made in many lakes in 
January 1980. This particularly broad range 
was nol reached by specimens of the same 
species in lakes in the Coorong area of South 
Australia. There the range is lS-Jfi^V < Dc 
Deckker & Geddes 1980). The query concern- 
ing the validity of Geddes" (1976) record of 
the salinity for A, rohaxta in Victoria, made by 
De Deckker & Geddes (1980, p. 691) is re- 
solved since further work has demonstrated 
thai the salinity range of A. rolnwa definitely 
differs between Victorian and South Australian 

lakes. Similarly, Geddes (1976) could not 
have misidentified A, robusta since it is the 
only Anstralocyprix species recorded in his 
study area. 

]ri Victorian lakes, A robusia is found in a 
healthy state and in high number* a* salinities 
between 45 and 77.5'/,',. and nearly always 
occurs with Diacypris compacta which is 
found in even higher numbers. At higher 
salinities, these two species are found with 
f'tatycyprtx batten, A. rohuxta occurs in two 
pcrmnncni salt lakes in Victoria, Lakes Gnoiuk 
and Keilamhcte, which have salinilies in the 
vicinity of 60'';^ the whole year round. This 
indicates that this species docs not necessarily 
require a sharp decrease in salinity to hatch 
although Geddes (1976) showed the hatching 
range for A. robusia to be 8.5-108', , 

Soiuamiiv: CYPRICERCINAE McKenzic 

Strandexia Vavra, 1895 
Tvpe species: Stra/tdesia mercatorttm (Vavra, 

Strandesia phoenix n.sp. 
F1GS15 j-q* ffl 
DtegH0S{y. Sfrandesia without shell ornamen- 
tation, with left valve larger than right valve 
all along and overlapping it ventrally, except 
in the antcrodorsal area at the extremity of 
the hinge where it is overlapped by the right 
valve; ellipsoid in lateral view and oval with 
pointed end in dorsal view. Valves asym- 
metrical when viewed from anterior: greatest 
extension of right valve at about 0.33 of height 
from dorsum and of left valve at 0.66. 
Description: Carapace. (External) Pseudo- 
pnnctate ellipsoid shell with dorsum and ven- 
trum gently curved, anterior slightly more 
rounded than posterior which tapers gently; 
in dorsal view oval with both ends pointed, 
in anterior view, valves asymmetrical greatest 
extension of right valve at about 0.33 of height 
from dorsum and of left valve at 0.66. Left 
valve laTger than right all along and over- 
lapping it ventrally except in anlerodorsal area 
at extremity of hinge where it is overlapped by 
right valve normal pores of simple type, some 
rimmed, others funnel-shaped. 
(Internal) Inner lamella broadest anteriorly 
and almost absent posteriorly in both valves; 
in left valve flange broad all along except 
doisallv' inner lamella near outer margin 
perpendicular To flange in anterior of left 



Fig. 16 Stratuicsia phoenix n.sp. a antennula, b maxillula— palp and lobes, c antenna, d mandible— 
coxale, e thoracopoda I, f mandible— palp, g maxilla, h thoracopoda II f i furcal attachment i 
furca. a-j: holotype adult female. Scale: I00X 



vjlve And curved inward in its middle; this flat 
,uv;i met by broad selvage of right valve; 
selvage faint and peiipheral in left valve and 
broad all along in right valve. 

Anatomy, Antcnnulu: (Fig. 16a) 7-segmented. 
length/ width ratio of last six segments: 1/ l .S, 
1/4. 1/1, 16/1, 2/1, 3/1; natatory setae as 
long as last six segments togclhei. 

Antenna: (Fig. I be) Four almost equal claws: 
three on penultimate segment and one on distal 
segment; natatory setae extending past tip of 

Mandible: (Figs 16d,f) Mandibular coxale 
with seven teeth; distal tooth acicular; cpipod 
with fivu long and one short Sirahlen; distal 
segment of palp rectangular and with long, 
slim and barren a bristle, fl bristle stout and 
pilose, y bristle thick and twice length of distal 
see.ment and piiosc on inside of its distal half. 
Rakc-like organ- Six teeth plus one bifid on 
inside ot" each rake. 

Maxillula: (Fig. Ibb) Length/ width ratio of 
palp segments: 3/1, 3/1; two toothed Zahn- 
borstcn on 3rd lobe. 

Maxilla: (Fig. Ing) Middle scfa more than 
twice length of other two which are of almost 
equal length. 

Thnracopoda I: (Fig. lo'c) First segment with 
two unequal setae, distal one pectinate and 
O.S length of other. 3rd segment undivided in 
middle where long seta occurs, 

Thnracopoda lit 'Fig. I6h) Distal pincers 
large and setae unequal : shortest one 
curved, with distal hall comb-like and Od 
length of other smooth one; middle seta on 
lasi segment less than 0.5 length ot all other 
setae which are of similar length. 

Furca: (Fig. 16jl Very long f ureal shaft, twice 
length of longcu claw; interior sela more than 
twice length of other seta iurcal attachment: 
Median branch narrow and gently curved dig 
Ifcj) dorsal branch forming eyelet whereas 
ventral branch straight 

Colour of shell: Purple, 


bo|otypo %dnll fcmnle 

J H ' M 

I v B40> 510m RV 860m 5400 

Type locality: Coastal dune lake at Evans 
Head, southwest of Lismorc, NSW. (29 "06' 

57' S, n2"25-40"E), 

Derivation oj name: From Greek phoenix (= 
purple) for the colour of the shell. 
Ecology and distribution; A phoenix has only 
been collected from Ihe type locality. Water 
was fresh. No males have been found and no 
sperms were noliced in the cvigerous females, 
suggesting that the species was pacthenogenetic 
at that locality 

SUBf \mily. DIACYPRID1NAE McKcnzie. 

Reticyprts McKcnzie, 1978 
Tvpe .xpecies: Rcticypris herbsti MeKenzie, 

Dixmstion: Reticypnx is an Australian en- 
demic genus which groups rhc following halo- 
biont species: R Jwrbxti McKcnzie, 1978, R» 
uathu De Deekker, 1979, R. cluva n.sp. and 
R, kitrditnurka n.sp. R. dedeekktri McKen/ic, 
1978, has been synonymized to R herbsti by 
Dc Oeckker & Geddcs (1980) as berth tnxa 
have an identical hemipenis (see discussion 
below). All Reiicyprix species, which are easily 
distinguishable from one another by the outline 
of the hemipenis, arc discussed below. 

Rttievpris cla^a n.sp. 
FIOS 17-" s 
1980 Rain-prit sp.nov.l, De Deekker & Gcddcs, 
P 6$Z 

Oiavnoxts: Reiieypris with lateral lobe of 
hemipeniy crescentle and broadest at base 
where two lumps are visible on inner side. 
Description: Carapace. (External) Subreetan- 
gular with dorsum gently arched and venirum 
almost flat except in mouth region which is 
concave; anterior broadly rounded and pos- 
terior tapering with posterodorsal area inclined; 
slight depression at extremity ot hinge 
anteriorly; left valve slightly larget all along 
and overlapping right valve in mouth region 
dorsally, especially where slight depression 
occurs; surface of shell with small, dense reti- 
culation usually all over and thickness of reti- 
culation variable; outer flange close to outer 
margin, narrow and thinly denticulated except 
tli mouth region where it is straight: greatest 
height at about 0.4 from anterior; in dorsal 
view like a flattened oval and bolh extremities 
slightly pointed. Normal pore canals of simple 
type and rimmed, 

(Internal) Inner lamella equal in both valves 
and broadest anteriorly, tapcrmg to 0.66 of the 
width posteroventrally. Hinge consists of broad 



Fig. 17. Reticypris clava n.sp. a RV external, b LV external, c LV internal, male paratype. d RV 
internal, male paratype. e C dorsal, male paratype. f C dorsal, male, g LV external, female para- 
type. h RV external, female paratype. i LV external, female paratype. j RV internal, female para- 
type. k C ventral, female paratype. 1 C showing RV, female paratype. m C dorsal, female para- 
type. n C showing LV, male, o LV external, anterior detail of i. p C ventral, anterior detail of k 
a-e, g-m, o-p: Type locality; f, n: Lake Weeranganuk, Vic. Scale: 1— 200m for a-n; 2— 100m for 
o; — 50m for p. 



Fig. 18. fteticvpris ciava nsp. a antennula, b antenna, c mandible palp, d maxillula. palp and lobes, 
e maxilla, male, f maxilla, endopodite, male, g thoracopoda II, h thoracopoda I, i hemipenis, j 
maxilla — endopodite, female, k furcal attachment, I furea, m Zenker organ, a-i, k-m: holotype adult 
male; j: paratype adult male. Scale: 100^. 


p r>fc dlckm r 

groove in left valve m which interlocks right 
valve; radial pore canals numerous and 

Anatomy: Antennula; (Fig. I8a> 7-segmcnted* 
length/width rata- of last UK segments: J 1.5, 
l/I, 1/L 1/1.8, 1/2, 1/1; nai.nory setae 
slightly longer than all segments, together. 
Antenna: (Fig. 18b) Three long "and equal 
claws on penultimate segments with shorter 
claw on distal one which is pectinate in male; 
natatory setae extending past Tip or lI.ivy:.. 

Mandible; (Fig. 18d) Seven teeth on mandible 
coxale last one longer than other '.nice adri- 
ceni to it and. near its base 1wo short pilose 
setae; length/ width ratio of palpv: 4.5/1, 
1.7/1; 3rd Jobe has two smooth Aihnborsten; 
ejpipod with five plumose Strahlcn and a 
smaller barren one. 

Rake-like organ: Fight teeth with an addi- 
lional bifid one on inner side. 
Maxilluta. (Fig. 18c) Distal segment squarish; 
a* ft and y bristle of equal length and slim: 
n smooth, ft pectinate, y smooth 

Maxilla: Sexually dimorphic: in mule, palps 
asymmetrical (Figs, I8e,f); broadest forming 
right angle on outside whereas other more 
arched; in female (Fig. I Si) setae unequal 
with shortest smooth whereas other two 
plumose: for chaeioraxy of pmtopod see Ffa 

Thoracopoda L (Fig. 18h> Penultimate seg- 
ment undivided and all inner setae long and 
of about equal length. 

thoraeopoda It: (Fig. IHgj Distal pincers 
small and distal setae unequal- shorter one 
curved and 0.2S length of other. 

Hemipcnis; (Fig I8i> Lateral lobe uvmxiW 

shaped and broadest at the base where two 

lumps are visible on inner side; inner lobe 

squarish with three sides concave. 

Zenker nrwr.. (Fie;. 18m) Elongated with 16 


Furea (Fig. I SI I Pectinate claws thicl; and 
equal: setae equal, pectinate and R33 length 

til* CjflWfc 

Fureal attachment- (Fig. 18k) Median branch 
straight and of about same length as curved 
inward dorsal branch; ventral branch forming 
obtuse angle with dorsal one and hook-shaped 
distally; short rod-like, extension at right angle 
on hose Of median branch. 

Lye; Cups of nauplius eye fused. 

( oltntt of .ihfll; Light green to while 

• is pc aUuli male 

t. H t 

LV 700M I3i)/i RV 69U/<, 

paratvpe adult female 

I It L 

LV 72Jfo* 445jm 


RV 700M 440> 

Type iomtiry Ephemeral salt lake S of the 
Ccorong Lagoon, S/\ (36' 13 36"S, 139*41' 
29* L) locality 3 of De Dcckker & Gcdde% 
f (VSOi. 

Daivmiott oj name: From Latin cluva (- 
club) for ihc diagnostic shape of (he outer 
lobe of the hemipcnis 

Ecofaxy unci distribution: In collections from 
western Victorian hikes, R, rlava was always 
accompanied hy low salinity oslraeods such as 
\l\/i!ocvftri.v splctuiiihi or M, pravnuncia, and 
occasionally with D. \pinnsa. Salinity for these, 
collections ranged between 4 and 42';',.. In the 
lakes adjacent to the Coorong Lagoon in South 
Australia, the salinity range for R. chiva is 
5-131'/, but if was never tound in high num- 
bers above 68'/,. Only in one lake near the 
Coorong Lagoon (locality 7 of the De Deckker 
& Gcddes |980) was R. clava found together 
with R. herb&th — this co-occurrence persisted 
throughout the year. R. clava has been col- 
lected once 10 Western Australia between 14.6 
and 59.5'.;, (Geddcs ct at 1981). 

Remarks: It is difficult to distinguish R, clava 
and R hcrbsti on features of the shell as reti- 
culation of the shell and even shape and size 
are known lo vary (see Fig. J 7 for R. clava) 
The nutlmc of the hemipcnis is a good diag- 
nostic feature for separation of the Iwo species 

Rvticypris herbsrt 1978 

I'-'7K Rrtu\i'ti\ hn-hsti n.sp.. Mckenzie, p. 1X8 
inagnostic; Renc?pti\ wirh Literal lobe at 
hemipcnis boot-shaped 

Description? $tx McKenzi* (1978), p. 188-9. 
Ectofogy Wlrf iHuttbtitian: R. hcrhstl is tolerant 
to higher salinities th.m R. clava. In western 
Victorian lakes h was found with D eampacia 
at salinities between 99 and 172',, whereas, 
in the takes near the Coorong Lngoon. it 
occurred fll salinities between 12 and HI',,, 
with three supplementary records at 195, 216 
and 2 Is*,,, it \% tound in high numbers be- 
tween 104 and 1 ?4' ,'. salinity, 

In a number of specimens, collected in takes 
near Ihc Coorong Lagoon, specimens with thin 



and family reticulated shell were seen, whereas 
others were thicker with a coarse reticulation. 
Similarly, a ventral ridge was occasionally 
associated with a coarsely reticulated Shell, 
or aKo with juveniles. The ecological signifi- 
cance Of these differences is not known. 

Retlcyprfc knrclitmirka n.sp. 

FIGS 19-20 

t)i(iytto\ix: Rftkypris with oulcr lateral lobe of 

hcmipenis spout-shaped and copulatory sheath 


Lh-MtiphotK Carapace. (External) Subrcetam 
gnlar to squarish in lateral view with both 
valves usually thick; reticulation resembles 
broad punctalion; anterior and posterior simi- 
lar and broadly rounded; ventrum rial except 
in mouth region where it is concave dorsum 
arched or depressed in front of point of 
greatest height, and at 0.33 from anterior, 
and behind it, il slopes gently; outer lamella 
thin and peripheral; left valve slightly larger 
all around and in some specimens overlap 
of left valve over righl one is obvious, at 
both extremities ol the hinge, as it forms 
smooth elongated humus. 
(Internal) Inner lamella slightly broader an- 
teiicirly compared to postcroveiural area, sel- 
vage faint and peripheral in right valve 
whereas at a distance from outer margin in 
left valves, radial pore canals narrow and 
straight; hinge consisls of a broad groove in 
left valve in which right valve interlocks. 
Anatomy Antertnula: Fig. 20a) 7-segmenled 
length/ width ratio of last six segments 1/ 1-25, 
1/1, 1/2.5, 1/2, 1/2, l/l; natatory setae 
longer than all segments together. 
Antenna (Fig 20b) Three equal lone claws 
on pcnulnmaic segment; 4th claw on distal 
segment reaching tip of other claws and pec- 
tinate in male, whereas jt is shorter and with 
smaller teeth in female; natatory setae reaching 
tip of claws. 

Matulihkr (Fig. 20h) Mandibular coxnle with 
seven teeth; palp wilh distal segment squarish. 
a hr tittle short, slim and smooth (i bristle short. 
stout nod pilose, y bristle twice length u{ distal 
M;:mcnr and pilose in its distal half. 
Kake-like organ: Fight Teeth with au additional 
inner one which is bifid. 

MaAillula; (Fig. 20c) Distal palp a&XVQst 
squarish and 3rd lobe with two ttaootft 
Maxilla Sexually dimorphic: m male (Figs 

20l\H almost symmetrical; one slightly more 
arched and narrower than other; in female 
(Fig. 20g) three smooth setae, middle one 
being more than twice length of other two 
which arc equal 

Thoracopoda I (Fig. 20d) 3rd segment un- 
divided; inner setae small and unequal. 

Thoracopoda If: (Fig 20i.) As for R- clava. 
Hcmipenis- (Figs. 20k,li Lateral lobe spout- 
shaped and broadest at mid-length; copulatory 
sheath heart-shaped. 

Zenker organ: (Fig, 20n) Elongated with It 

Furcir (Fig 20m) Claws equal and thick; 
setae equal and small, about 0.25 length of 

Furcal attachment: (Fig. 20j) Median branch 
and long dorsal one gently curving 1 ventral 
branch forming a right angle with dorsal 
branch and forming a loop distally; vertical 
rod-like exlention near base of median branch 

Kye; Cups of nauplius eye fused. 

Colour ol .shell: White when preserved "in 



hololype adult male 









I H 

I.V 565* 3i50W 

panuype adult female 

1 II 

LV 600,u 40<V 

Tyfff locality: Madigao Gulf. Lake Eyre, South 


Derivation of name; Kunlitnarka is an abori- 
ginal name for a legendary creature supposed 
to inhabit the bottom of lagoons and creeks 
in the Lake Eyre district. 

Ecology and rlivfrtburinn: One specimen of R. 
kurdtmurka had been originally collected from 
I ake Fyre N»>rth «.n 2X.IV. 1975 at about 40;, 
salinity when the lake was last Hooded (see 
Bayly 1976, p, 664 where it is referred to as 
"undescribed cypndid genus") Subsequently, 
it has been collected twice from Madigan Gull, 
til lake EytC by W, /culler on I I. XII. 1074 
and 2.XIU975 — no salinity record*, for these 
collections are available. Recently, the same- 
species was collected fiom Lake Annean, 40 
km S oi' Meekaiharra. m W.A., at 21 3 ,, 
salinity (see Gcddcs ct (fi . 'n press), The 
Western Australian specimens had a much 
thinner shell than those from Lake Eyre. 



Fig. 19. Reticypris kurdimurka n.sp. a RV external, male holotype. b LV external, male holotype. 
c LV internal, male paratype. d RV internal, male paratype. e LV external, female paratype. f RV 
external, female paratype. g LV internal, female paratype. h RV internal, female paratype. i C 
showing RV, male paratype. j C showing RV, female paratype. k RV internal, male. 1 C showing 
LV, male, m C dorsal, male paratype. n C ventral, male paratype. o C dorsal, female paratype. 
p C dorsal, male, a-j, n-o: Type locality; k-1, p: Lake Annean, 40 km S of Meekatharra, W.A. 
Scale: 200/*. 



Fig 20 Rcticypris ktttdimurka n.sp. a antenna, b antennula, c maxillula — palp and lobes, d thora- 
copoda I, c' maxilla— endopodite, male, f maxilla— endopodite, male, g maxilla, female, h mandible 
—palp, i thoracopoda II, j furcal attachment, k hemipenis. 1 hemipenis, m furca, n Zenker organ. 
a-b, d-f, h-n: holotype adult male; c, g: paratype female. Scale: 100m. 


p. or nrcKKFR 

Rcticvftis wolbtt De Deckker. 1979 
1979 ftcticypris titbit De Dcckktr, p. 162. 
Diagiu>xi\; K. rnyprt'. with Lifer,:,! lobe of henu- 
penis banana-shaped and inner tobe cudgel - 

tMHftptfan See Do Dcckkct (J'J7'M p. 162-1. 

[)/\nnsiort. H. wallnt was originally described 
from samples collected in mound spring-, be- 
tween Strangways and Cuulmiurka, near J .aJU* 
Eyre South in South Australia. Since then it 
has been recognized from , ( collcen^n made by 
1. A. E. Bayly and W. D. Williams m Ukd 
Buchanan. S.W. of Charters Tower in Queens- 
land, in January 1965. Water sabni'y 
K7.6',;, (Bayly & W.lliams 1^73). It has not 
been fouud in subsequent collections made at 
ihc same lake by B. V, Timms. Similarly, it 
in surprising Lhat R wathu has not been col- 
lected in Lake Byre instead o) U. knnlimnrka, 
Sm h patchy distribution remain* unexplained 

Family ILYOCYPRID1DAK Kanln. ..... 

J 900 

Uyvcyprti Brady & Norman, 1889 
Typ* \f><ri<'\-- flvorvprix $ibba (Ramdohr, 

flvvcvp/U atwt&H&i&i Sais. issv 

1883 flyacypri; australicnsu Sars, p. 4(» 
FIG. 21 

Uiuyjiosis: ]i\t>cypii.s wilh dub-shaped inner 
lobe 09 hemi penis extending to as ruueh ;v 
bh m length oi' trapezoid lateral lobe. 

Remarks: The original description of /. atts- 

frahensn by San (1 889a ) is sutticien*. ami 
does not warrant additional illustrations of 
the anatomy here. The ornamentation and out- 
line of ihc valves at /, umtralietixis. however, 
are extremely variable. Typical variations are 
illustrated in Fig. 21 (all SEM photos are at 
the same scale and only adults are repre- 
sented), The surface oF the shell can be finely 
() : ip 211] to coarsely pitted (Figs 2lc,r',o); 
acicular but small opines occur, mostly pos- 
terodorsally ( Figs. 21h,i) when present, but 
can also COVcr the entire carapace (Fig. 21o) 
The outline of the shell is also variable 
anteriorly and dorsal ly. In dorsal view, 
anteriorly the shell can be pointed (Figs 21j,o) 
or with a squarish blunt end (Figs 2lru,n). 
The latter feature is often accompanied by a 
depression bordering the periphery of the vaJvc 
anteriorly and posteriorly (Figs 2Ja,dl, De 

pressions are also variable dorsally, a circular 
one above the central muscle rlelcl and the 
oiher vertical nfcrove and in Iront C/t the central 
muscle field, 

The ecological significance of all these 
Variations arc not yet understood but this is 
not a surprising phenomenon since t tmsira- 
Ht'ini\- is commonly found in temporary pools. 
which are renowned as the >itcs of hro^l 
variations h\ physical parameters (viz, tern 
peraiure. pH, O..). I <w*iraHen\is can swim 
but is usually found crawling in or above 
muddy substrates in ponds. The species usually 
occurs in ponds which retain water for a few 
mouths and which have a very soft substrate. 
/ uttstnritensw has also been collected in fl 
number of shallow 3nd slightly saline lakes 
in w^tcrn Victoria These lakes are eharac- 
tcri7.ed by small fluctuations m salinity over 
(ho year. This explains the absence oi I. ans- 
(ralietms in the saline lakes near the Coorong 
Lagoon studied by De Deckkcv & Geddes 
M9isT)) where salinity ot some lakes was close 
to fresh m winter, hoi rose rapidly during 
■pving and summer months. In western Vic- 
toria the ranee ot salinity tolerance tij / 
UHsiraiienstfy is usually 4-7%, with one record 
at 10 37% in Lake Kariah. So far. /. ansttA 
iicnsi* has never been collected m deep lake*, 
(fresh or slightly saline) nor in permanent 
fresh lake*. 

/". tiustmlien\i\ has been collected all across 
Australia. Ir is also recorded from North 
Africa (Ciauthier 192S). Asia and southern 
Europe ( Hartmann 1964, p. 148). 

As the morphology of the hemipenis of /. 
i:ti>:nntii'tt\i$ and /, ifeapi,n\ Mjsi. 1906, as 
illustrated in Pcrkovski (1V58). is almost, it is suggested here that these two 
species could be synonymous. Further work is 
necessary la confirm this hypothesis, but ii is 
Important ro be jware of the variation in shell 
ot namenintion of (he llyotyprix species as 
illustrated above, and hy DicbeJ and 
Pietrzenmk (1975) for A brad y i Sar%, IS90, 
since many Jlyocvprix species are separated on 
shell morphology alone. The selection of parti 
cular features of the shell as taxonomieally 
important by van Hartcn (1979) will not 
prove to be useful for /. aitstroliensis speci- 
mens since for example, the "marginal ripplets" 
of van Ifarttn are variable in Australian speci- 
mens, and even occasionally absent. Further, 
distinction of species on lean ires other than 



Fig. 21. Ihocypris australiensis Sars, 1889. a C showing LV, female, b RV external, male, c C 
showing RV, female, d C showing LV at dorsolateral view, male, e C showing RV, male, f C 
showing LV, male, g LV internal, male, h LV internal, female, i RV internal, female, j C dorsal, 
female, k C ventral, male. 1 C dorsal, female, m C dorsal, female, n C dorsal, male, same speci- 
men as d. o C dorsal, female, p LV internal, female, q C showing LV, male, a, d, h-i, m: Pool 
in creek bed at 25 km N of Cue, W.A. b, e, g, j-k: Martin Lake Vic. c, f: in vent to Warrawenia 
Lake, N.S.W.; 1, p-q: Pool in creek bed, 12 km S of Menzies, W.A.; o: Pond very close to Reel 
Inlet (coastside) 19 km S of Mandurah, W.A. Scale: 20(K 



the hemipenis is likely to be unreliable smee 
the morphology o( many appendages of //yo- 
cyptk species are known to vary. For example. 
GauthiL-r (K»28) discussed the variations in 
the anatomy ol /. nu.stnilicn\is and / hiplt'i-at,i 
(kocli, IN3N) fioiu North Afnea and con 
eluded that he was incapable of separating 
the species, although neiihcr was tUS prepared 
lo synonymic them 

n.voe.vpris perigundi n.sp. 

FIGS 22-23 

Oin&nofis: tlyagyprlA with carapace trapezoid, 
aulerodorsal area al hinge extremity com- 
ptessed and forming a hump when viewed 
laterally, three large protuberances Efl dorsal 
area, posterior one being largest, pustulost: 
and ending with a number of wart-like tuber- 
cles, ventrum concave and at O.b from an- 
terior Club-shaped inner lobe of hemipenis 
reaching lip of trapezoid lateral lobe, 

f)<\scrtf>rio/t, Curufuue. ( External ) I rape/oid 
with greatest height 2 from anterior where 
hinge starts; dorsum straight except at point 
of greatest height where bolh valves foim 
flattened hump; ventrum concave at O.n from 
anterior. Surface of shell highly ornarnenled; 
three protuberances dorsally with posterior one 
largest and ending with broad wart-like tuber- 
cles from which seta protrudes; below posterior 
protuberance and above vcutrurn, is an addi- 
tional highly ornamented protuberance: this is 
point of greatest width of shell; deep deprcs 
si* hi dot sally between middle and posterior 
protuberances; shell covered with hroad 
pseudopunetae which are pusrulose inside; 
pointed tubeicles along niacin .interiorly and 
posteriorly. Dorsally, sheH like flattened ellip- 
soid with anterior compressed and pointed 
t..r width o| inner lamella but tapering 

(Internal) Inner lamellae twice gts wide in 
anterior compared lo posteriot in both Valves, 
narrow selvage at O.nb from ouler margin 
anteriorly a\u\ peripheral to inner margin 
posie-i toily: marginal rippIcK broad in jej i 
valve postetoventr^illy. 

Atwumiy; Ametmula. (Fig. 23ct G-scgmcnted* 
length width ratio oi last fjve segments; 5/ J. 
12 1 | # 3/1, 2 1, 4 I; natatory scLi as long 
as all segments together, 

Antenna- I Fig. 2>a) Natatory ^ twice 
length of bst tvs c» segments and claws together; 

three long distal claws on penultimate segment 
plus another as long as other three on last 
segment with a narrower and shorter one. 
Mandible: (Fig. 23d) Length ' width ratio of 
Jasl two segments of mandibular palp: 4/3. 
I, 1. 

Maviliula. (Fig. 23b) Distal segment of palp 
trapezoid .ani ending with three broad long 
setae plus two shorter ones; no Zahnborsten 
iu\ 3rd lobe but setae short and stout on all 
three lobes. 

Mavilla; SOXUaHy dimorphic: m male (Figs. 
23v.h» palps two or three-jointed with one 
seta ai die distal end of the 1st segment; in 
female ( Fig. 23gJ non-segmented, short and 
narrow palp with three unequal setan. For 
chaclokixy see I i -/.h_ epipod plate with 

five Slrahlen, 

Thoracopoda 1: (Fie 23f) Last segment un- 
divided; all setae short 

Thoracopoda II (Fig. 23i) 4-scgmenlcd with 
three disial setae, two long equal ones and 3rd 
about length of others. 

Hemipenis: (Figs 23j,k) Outer lobe trapezoid 
with inner distal end pointed which is nearly 
reached by long club-shaped jnner lobe; outer 
lobe triftngUlur with distal end rounded and 
silently pinched laterally; inner lobe 0.33 
length of others. 

Zenker orgUi: (Fie. 23 I » Both ends of globu- 
lar and with 1 ^ rosclles 

Fuiea: (Fig. 23m) Two lone equal claws: 
posterior setae slightly longer than anterior 
and placed at 0.4 of length of shafr from 

posterior claw. 

CWCW "/ she!!. Transparent white. 

iiotnispc atftfll nude 

t II I. H 

i.v sfm '■ kv sefo assu 

p-uatype udii't female 

L H 1 H 

I V :"K0,m 3B0/1 RV 5fU¥ SStyt 

I Vf& locality Wurrawenia Lake, south ol 
Memndic N.S-W*. (33*WJ0"S, l4l , 44'30*Bj 
(hriv<*n\>n <>f intt>><\- J io m the aboriginal Ian 
I - lu-nymiili meaning meandering lake or 
laeoiin nx this species has been found in a 
lake parr ot ;iu old river system (W;irrnwenia 
1 ' at <d in a lagoon (Katarapko L.). 
Ecnfaftf omJ tlrJtibntiott: This species has Olih 
been collected m ihree localities: Warrawcnia 
L , Katarupko Lagoon, which is a natural billa- 



Fig. 22. Ilyocypris perigundi n.sp. a LV external, female paratype. b RV external, female, c LV 
external, female, d external, male holotype. e C dorsal, female paratype. f RV internal, female 
paratype. g LV external, male holotype. h LV external, juvenile, i LV external, detail of a. j LV 
external, posterior detail of C. k LV external, dorsal detail of a. a,d-g, i-k: Type locality; b-c, h: 
Katarapko Lagoon, near Loxton, S.A. Scale: 1 — 200m for a-h; 2— 50/* for i-k. 



Fig. 23. Jlyocypris perigundi n.sp. a antenna, b maxillula — palp and lobes, c antennula. d mandible. 
e maxilla— endopodile. male, f thoracopoda I. g maxilla, female, h maxilla, male, i thoracopoda II. 
j hemipenis. k hemipenfs. I Zenker organ, m furca, a-f, h-m: holotype adull male, g: paratype adult 
female. Scale; 100m. 



bong converted into an evaporative basin near 
1he River Murray at Loxton, S.A., and from 
Scott Creek, near Adelaide At all three 
localities only a few .specimens have been 
collected; most specimens were coated with 
mud particles. The salinity at Katarapko 
Lagoon was 3';,,. Salinity Of the water from 
the type locality has not been measured, At 
Scott Creek, water was fresh. 
Rcfttarks t periy:uticfi diltcrs from /. austro- 
iienm on the following features; the former 
is smaller and has a very diirereni -shell archi- 
tecture characterized by the three highly orna- 
mented dorsal protuberances, a ventrum placed 
nl 0.66 from the anterior of the shell and a 
Hat dorsal hump in the extremity of the hinge 
imtciiorly. The inner lobe of the hemipems in 
/, perigtfUdi is almost as long as the lateral lobe 
whereas it is only 0.66 the length in / mismh 

St'HRrvMiLY. CVTHERACEA Baird. 1850 

Family LhPTOCYTHFRIDAF Hanai, 1957 

Lcputcythere Saris, 1925 

Type xpecicw Leplncy there pelhicida (Baird 

Leptoe> there lacMslris n.sp. 

FIGS 24-2-1 

1 419 Cvthcrr tubbockmna Brady Chapman, n. 

Diagnoxix: Leptoryrhere with deeply pitied 
external surface of shell, large circular depres- 
sion postcmventrally outside and where inner 
lamella is broadest on inside: posterodorsally 
near termination of hinge, shell is slightly con- 
cave and posterior to il; shell thick and smooth 
especially in left valve. Outline of hemipems 
as in Figs 25j,k. 

t>rnription; Carapace. (External) Subreetan- 
L-iik.r shell, coarsely pitted and with at least 
two: elongated grooves; anterior straight and 
forms an obtuse angle with hinge line and 
other, nt posterior, runs almost parallel to cur- 
vature of shell; a eeeper and broader notch 
oficn visible in posterovemral area in both 
valves; ornamentation nf shell varying from 
fine to eoarsc ribbing; dorsum almost straight 
and inclined except in posiero-dorsal area, just 
before termination of hinge where it is slightly 
COneaY*; behind this, shell '.hick and smooth, 
especially in left valve; greatest height at about 
25 from anterior; mouth region concave and 
at n.4 from anterior, In dorsal view, -shell com- 

pressed and has almost straight sides except 
where grooves occur anteriorly and posteriorly; 
hinge area almost smooth externally. 
(Internal) Inner lamella broad and widest in 
poster ovcntral area opposite external deep 
notch; selvage faint and peripheral in both 
valves; hinge creruilaied all along with broad 
tooth at both ends in tight valve and matching 
sockets in left one; behind anterior socket in 
left valve, are two smaller teeth and in front 
of rhc posterior socket, with a matching de- 
pression m right valve, is also a small tooth; 
central muscle field with vertical row ot four 
scars: two in middle are elongated and parallel 
to hinge line whereas others are almost cir- 

Ajt&t dflty; Anrennula; (Fit; 25a) 6-segmemcd: 
length/ width ratio of last four segments- 
1.6/1, 1/1.4, II to 1.4/ J, 4/1; distal thick 
seta pectinate. 

Antenna: (Fig, 25h) Two smooth distal claws 
on terminal segment slightly shorter than inner 
claw near distal end of penultimate segment, 
Mandible: (Figs 25d,c> Mandibular covale 
with seven teeth and lone ncieulur sela on inner 
side near base of smaller tooth; epipod with 
terminal segment small and square-shaped. 
Maxillula: (Fig. 25c) Epipod with 14 long 
and short Strahlctu terminal segment of palp 
small, rectangular and with one distal seta 

Maxilla: (Figs 25h,i) Distal claw short and 
slightly curved; in female inner seta on 2nd 
sevmenl pilose in its distal half (Fig. 25i I 
whereas barren in male (Fig. 25b). 
Thoracopoda J: (Fig. 25 f) Slightly larger than 
maxilla but with only one distal sela on 1st 

Thoracopoda II: (Fig. 25g) Similar but 
-lightly larger than thoracopoda I and wilh 
diHal claw more slender 
tiemipcnis: See outline in Figs 25j,k 
Genitalia: See outline in Fig. 25nr 
Furea; (Figs 25k.m) One long seta near hemi- 
penis and genital organ. 

End tit body (Fig. 251) With one small seta 
and densely pilose 

Colour of shell: Fight brown. 


holotvpe adult mule 

l n l H 

LV 500.* 2-flttw RV 490* 270m 



Fig. 24. Leptocythere lacttstris n.sp. a RV internal, male holotype. b LV internal, male holotype. 
c C showing LV, female paratype. d LV external, male paratype e LV external female paratype. 
f C external showing RV, male paratype. g C external showing RV, male, h RV external, male 
paratype. i RV internal, hinge posterior detail of a. j RV internal, hinge posterior detail of a. k LV 
internal, hinge posterior detail of b. 1 LV interna), hinge anterior detail of b. m C dorsal, male 
paratype, n C external, detail of g. o RV internal, detail central muscle field of a. p LV external, 
detail of e, e-f, h-m, o-p: Type locality; g, n: Chara Lake, near Robe, S.A. Scale: 1— 200m for a-h, 


-50« for i-1, o: — 25^ for n.p. 



Fig. 25: Leptocythere lacustris n.sp. a antennula. b antenna, c maxillula — palp and lobes, d mandible 
—palp and lobes, d mandible—palp, e mandible— coxalc. f thoraeopoda 1. g thoracopoda II. h 
maxilla, male, i maxilla, female, j hemipenia. k hemipenis. 1 end of body, m genitalia. a-c f g-h, j-k: 
holotype adull male; f, i, m: paratype aduh female; 1: paralype adult male. Scale: 100m. 


J\ |16 OnCKKFR 

pMWlyjifi adult female 

I H L il 

LV 4h u :70m RV J [l 

7 »7>e locttfti] - Fresh Dip Lake near Robe, S.A. 

Dewatton vj name; From Latin htcusirinus 
( of lakes) as I his species, which belongs to 
a typfctllly estuariiic genus, is found in lakes. 
Ecvtp%y and distribution* This truly bcnihio 
species has been collected in only four locali- 
ties near Robe. SA apart from the type 
loealily. A If localities' are eharaclcrizcd by 
petmancnt water only Slight salinity fluctua- 
tions. The salinities were I L > to 2&% v , In addi- 
tion, at the type locality, salinity was 2.S', ( 
So Jar, /,. UiCUStfU has been found in perma- 
nent water around 35 r 7, as no such lakes have 
yet been sampled in ihe search for this ostra- 
eod, Attempts to find it in the permanent and 
saline lakes Keilambete and Guotuk III western 
Victoria (salinity ea. 55-62'/, ) were unsuc- 
cessful, suggesting that the salinity ran^c ol 
L, hictislr'iK docs not reach such values. This 
species, as for all others in the typical 
esluarine uentis Lepocvthere, is indicative Of 
permanent water as d does not produce eggs 
which can withstand desiccation. 

Remarks. The description of L, laett\Ui\ 
corresponds to the specimen identified bv 
Chapman (1919) as C'yrhere labhockiana from 
the fossil site at Bunco Swamp The two 
rounded tubercles on the posterovcniral QfCB 
ol die shell as illustrated by Chapman (1919) 
un Plate IV. V are two sand grains which have 
been removed by me from rhc specimen in tin 
National Museum of Victoria. Ihe shell archi- 
tect nre of L t laenstrix varies from almost 
smooth to coarsely reticulated (Fits 24n.p>; 
at times, reticulation js so thick that the shell 
has a smooth appearance. Such variations hava. 
In. n mentioned for other estuarine ami marine 
species Oi Lvpt\>cytlicre by Shornikov (1966) 
and Haitmaim & Kurd (l l >78). This \$ not 
suipnsms; as environmental conditions, such 
as wafer salinity in salt lakes, can Illicit u \\ 
over time. 

/ . h»-nstrt\ is closely related to / . harittranni 

(MeKouic 19^7). Unforlunalely. no males 
of the latter >pccies have ever been found, as 
analysis d\ die hemipenis morphology would 
httVC eui 1 1 irmed this distinction. The v;t|ves 
t^f (. iiartinanai, however, are not coarsely 
pnied ami reticulation »s more sparse tsce 
Hartmann |9f9, PL Iff; Frgp 3-8) 1 he 

antciioi ami posterior grooves are present in 
both species. The posterovcniral notch is much 
deeper in L. lactistris and ihe smooth postero- 
ior,al hump is absent in £. hartmanni 

r Avniv. LIMNOCyiHLRIDAE Sars, 1925 

Soni amii v; LIMNOCYTHHRiNAE Sars, 



/ v()r \pee/e\ ('uanphodella main n.sp., gender 


DiawtLK-its: < an/pace. Female carapace in 

dorsal view triangular with greatest width at 

about U.ob lo 0.75 from anterior; at about 

4 from anterior, where central muscle field 
occurs, valves are slightly compressed; absence 
^( proiTunenl lateral ridge in ventral area; 
Sieve poie ...mals .md lew broader normal pore 
•.anal-. In iow parallel to posterior edge of shelf 
m bolh vahes. Male, in d.jisal view, like a 
Flattened oval with both en<ls pointed, 

AlWOMy, Female manilla and ihoraeopodac 1 
and If similar, although of different length, 
female furea with three stout, pointed and 
thickly pilose setae. 

i>env<iJiun of name: From a combination of 
the two nanus (?QtfU$<)tyth£re and Cylhcri 
dellu as thi> genus shares diagnostic features 

01 ific twp ^eueia. 

Rtnwrkx; GCWipHodeita is very similar to CV 
thu (delta IXulay, 1005 except that the shell 
ol the former is less compressed in the area 
where the ceutial muscle lield occurs, and as 
thr. ma \ ilia and thoracopodae I attij II arc 
similar in GmnpJu*d<<!}, t (in C^thiiidelhi the 
thorucopoda II is truiMoi nietl into tl prelum 
sile palpi The diagnostic feature which Cam- 
phott&Utt fllltl Cottifdtocvihers Sars, V934, share 
is Ihe presence or Itirec fclflful and ihiekly pilose 
setae ou the female furca — this is m>t seen in 
Cvthendeibt G<>tn,>hodeita lacks the lateral 
veuiral ridge on bulb valves on the outside of 
Mil shell so typical of Gomplwcythere species. 

GottiphodeUa ^ to be included in the family 
I unuocvlhennae Sars. 1925. following the 
rrccnl regrouping of subfamiliev in the I imno- 
cythetidac by Colin & Danielopol [1978J> 

<\nmphodelki m;iia n sp, 

FKJS 2u-27 

t>ut£tt<>.\{<. Surface of shell pseudopunctate 

with H(?VC poies. pftfttsrior area of shell 

broadly rounded in lateral view: in dorsal view, 



valves only slightly compressed in female at 
0.4 from anterior where central muscle field 
occurs. Outline of hemipenis as in Fig. 27i. 
Description: Carapace. (External) Adult : 
Subrectangular in lateral view with ventrum 
flat and dorsum arched; posterior broadly 
rounded and antcrodorsat area flatter and 
forming less steep angle; surface of valves 
pseudopunctate with many sieve pores; normal 
pore canals either of simple type or surrounded 
by rim; a few broader normal pore canals 
arranged in row parallel to posterior edge of 
shell in both valves; in dorsal view, carapace 
of male like flattened oval with anterior end 
more compressed and pointed and greatest 
width near middle; in female, greatest width 
at about 0.75 from anterior which is pointed 
whereas posterior almost flat except in middle 
which is slightly convex where both valves 
meet; at 0.4 from anterior, where central 
muscle field consisting of row of four vertical 
scars occurs, valves are slightly compressed in 
female; left valve slightly longer anteriorly 
and posteriorly. In some female specimens, 
occasional faint ridges present ventrally near 
outer margin of each valve and running paral- 
lel to it. 

Juvenile: Almost round in lateral view with 
flattened ventrum; oval in dorsal view; some 
specimens with broad flange sometimes extend- 
ing into one or two spines posteroventrally. 
(Internal) Inner lamellae much broader 
anteriorly; selvage prominent and broad all 
around in both valves; in left valve, however, 
it is placed further away from edge of valve, 
especially anleriorly; in front of mouth region, 
selvages very broad and, when interlocking 
during valve closure, right selvage placed ex- 
ternally; shell perforated internally; radial 
pore canals numeroous, short and usually 
straight. Hinge lophodont: anterior and 
posterior grooves in right valve elongated and 
matched by ridges in left valve. 
Anatomy: Antcnnu\a: (Fig. 27a) 6-segmented; 
length/ width ratio of last five segments: 
1.9/1, 1.2/1, 0.8/1, 1.2/1, 3/1; longest distal 
seta bifid with one side more than twice length 
of other. 

Antenna: (Fig. 27b) Three smooth distal 
claws on terminal segment: pectinate distal 
claw on inner side of penultimate segment 
slightly shorter than other three claws. 
Mandible: (Figs. 27d,e) Mandibular coxale 
with seven teeth: inner one slender and twice 

as long as penultimate; palp with terminal 
segment almost trapezoid and with two thick 
distal setae and a shorter and thinner one. 
Maxillula: (Fig. 27c) Poorly sclerotized palp 
with three unequal and broad setae; epipod 
with 13 long and one small Strahlen. 
Maxilla: (Fig. 27f) Distal claw stout, short 
and curved inward; setae on 1st segment 
smooth, and distal one on 2nd segment finely 

Thoracopoda I: (Fig. 27g) Similar to maxilla 
except for slightly larger size. 
Thoracopoda II: (Fig. 27h) Larger than thora- 
copoda I with distal claw slender, longer and 
almost straight; inner distal end of last two 
segments with a short spine. 
Hemipenis: Strongly chitinized; for outline see 
Fig. 27i. 

Genitalia: For outline see Figs 27j,k. 
Furca: In female (Figs 27j,k) consisting of 
three short, stout and pointed setae densely 
covered with stiff hairs and two longer and 
barren setae; in male (Fig. 27i), two barren 

Colour of shell: Grey to grey brown. 
holotype adult male 

L H L H 

LV 490m 290m RV 460m 300m 

paratypc adult female 

L H L H 

LV 560m 330m RV 520m 320m 

Type locality: Fresh Dip Lake, near Robe, 
S.A. (35°15 ; 42"S, 139°48'42"E). 
Derivation of name: From Greek maia mean- 
ing good mother as some adult females from 
fossil deposits (Pulbeena and Mowbray Swamp 
in Tasmania, sec De Deckkcr 1981)* were 
found with juveniles still inside them. 

Ecology and distribution: This species only 
occurs in permanent waters as eggs cannot 
withstand desiccation (for further details, see 
McKenzie & Hussainy 1969). Brood care 
certainly occurs for this species as quite a 
number of female carapaces have been found 
to yield either one or two juveniles. It is likely 
to be a freshwater species which can tolerate 
slight concentrations of dissolved solids in the 
water as, like G. australica, it has been col- 

DE DECKKER, P. (1981) Taxonomy, ecology 
and pafaeoecology of ostracods from Australian 
inland waters. Ph.D. Thesis, Depi of Zoology, 
University of Adelaide (unpubl.). 



Fig. 26. Gomphodella maia n.gen., n.sp. a RV internal, female paratype b LV external, female para- 
type c C dorsal, male paratype d RV internal, male holotype. e LV external, male holotype. f RV 
internal, juvenile, g LV external, juvenile, h RV dorsal, female, i C dorsal, male, j C dorsal, female. 
k C dorsal, female. 1 RV internal, anterior detail of d. m LV external, detail of e n LV external 
detail of g. a-g, 1-n: type locality; h-k: fossil, Blue Lake, Mt Gambier, S.A. Scale- 100^ for a-k- 
15," for 1; 5fi for m, n. 



Fig 27. GotnphodeUa tna'ta n.gen., n.sp. a antcnnula. b antenna 
dible — coxale. e mandible — palp, f maxilla, g thoracopoda I 
furca. j genital organ and furca. k genital organ and furca. 
b, d, f-i: holocype adult male. Scale: MXK 

c maxillula — palp and lobes, d man- 

h thoracopoda II. i hemipenis and 

a, c, e, j-k: paratype adult female. 


f\ \>h DKCkKbR 

lcctcd ahve once in Fresh Dip Lake < — type 
locality) at 2.3?^ salinity. As a fossil, it has 
been found with C cwsiratica .and other i 
water osiracods in sample*, from a shoa core 
fr©in the Blue lake at Mt Gambier, S.A. It is 
also found in two fossil sites j« northwestern 
Tasmania (De Deckkcr*). In samples from 
both sites, a large number ol' carapaces were 
recovered and some of these were found with 
juveniles msidc them. The significance of this 
phenomenon i.% not properly understood as T 
usually, after death under water, valves of 
osiracods separate prior to or during decay of 
the soft parts. When ponds dry up, nstraeods 
are seen to close their valves tVghtfr and, if 
this period is long enough to cause dehydration 
of the ostraeods, death would occur. Rapid 
sediment aceumulauon is later necessary dur- 
ing a wet phase to prevent carapaces from 
opening. This process is likely to be one 
possible explanation for the high percentage ol 
G. maia carapaces found in samples from the 
two Tasmaman sites. Death caused by changes 
in water chemistry would uot prevent cara- 
paces from dislocation, Another possibility is 
that &, main can in fact burrow in sediment — 
a phenomenon noticed for a closely related 
ostracod Gomphucybthere sp. (with another 
ostracod Darwlnalu sphagna Barclay. 1968) 
found m the interstitial waters of the Rotorua 
lakes in New Zealand by Chapman & Lewis 
(1976)— and, if it was to remain there until 
death, valves could not become easily dis- 

Gomphodelfa nusrralicu (Hussaiviy. 1969) 
i%9 Gomphotythew twstratiea Hussainy, p. 29V 

Diagnosis: Subcectangular shell with both 
dorsum and ventrum almost straight; anterior 
broadly rounded and posterovenlral area more 
pointed with furthest extention of shell at 
about 0.33 from dorsum; shell ornamentation 
consisting of faint but broad reticulation 
especially at anterior and posterior ends, Tn 
dorsal view shell in female like an upside down 
heart and male much narrower with both ends 

Size range: 

6 50-7 5 UM 



adult male carapace- 
iduJL female carap-'LC 

Ecology; G australica is best known from ils 
type locality. Lake Pumimbetc in Victoria. 
This is a permanent lske with salinity of 0.42- 

O.SIV,;,, over the 1 969-1972 period (Tmuns 
1976J and 37-U|4/,,_, over HW-SO. Tt is 
only recorded in that lake (greatest depth > 
40m) at 0.5-1 m hy Tim ins (1973)^. It is 
usually found crawling in among Jtlamcntous 
algae like G main. In South Australia it jh 
also recorded from Scoit Creek near Adelaide, 
and from ihe permanent Fresh Dip Lake near 
Robe. G t;/M//yy//c(/ is considered to be a fresh' 
water species whieh can withstand a slight 
amount of dissolved solids in water, with ils 
highest salinity recorded at Fresh Dip Lake at 
,, . Brooding occurs in this species as de- 
monstrated by Hussainy (1969). Fossil speci- 
mens, with coarse shelJ reticulation have been 
recovered with other fresh water ostracods 
FrotB a short core taken from the fresh Blue 
Lake al Mt Gambier, S.A 

Remark* The transfer of this species to Gom- 
phtnie/la gen nov. is necessary because it does 
not possess the peripheral lateral ridge around 
the H:,t base of the shell of each valve so 
typical ot Gompmny there Sars, 1924. This 
species is easily distinguishable from G. main 
by its larger size, reticulated shell and ihc very 
pointL'd posterior area of the shell which is also 
steeply inclined postcrovcntrally. 


This paper was written during the tenure 
of a Commonwealth Postgraduate Research 
Award under the supervision of Piolessoi 
W. D Williams in the Zoology Department, 
University of Adelaide. 

Many of the specimens described here were 
received from the following; Mr P Bailey. 
Drs T. A. K. Bayly, R. Marchant, R Tail and 
B. V rimms and Mi W. Zeidler. 

I am giatcful lo the following for the 
loan of type specimens; Dr M. Christiansen 
(Oslo Museum), Dr G. Boxshall and Ms A. 
Gumev l British Museum, Natural History), 
Drs I A. E Bayly. R Merchant, R. fait and 
1L V. Jimmsand Mr W Zcidler 
Museum) . 

I would like to thank Ms S. Lawson and C. 

Twang for typing the manuscript and Mr IV 
G. Kempslcr for help m the photography. 

• IJlvtMS. B. V. (jy??) A comparative study of 
the IJfUAOlOK) of three trim laki'B in western 
Victoria. PhD I h^sis. Mcmash Universiiv 



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Ryre. Aost. f. Mar. Freshwa/. Ret- 27: 661-5. 

St Win cams, W D. (1966) Chemical iinJ 

biological studies on some saline lakes of .souib- 
easl Au*li:ilia. ihU 17, 177-228. 

. & fJ97i) "Inland Waters and Their 

fc'cology'V Longman, Melbourne. 

Bfu>Y, G S. iT8S6> Notes on treshwatei Ento- 
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London 54. $M3L 

Chapman, f. (19iyj On an ostracod and shell 
rrmrl of Pleistocene a^c frOfli BflffJ^O Swamp, 
West of Cape Schuncfc Victoria. Prot R. $0& 
\'ht. 32. 24-32- 

CHAPMA^i Nti A. iiyfiJ) A review of the fresh- 
water ostracods o\ New Zealand. H^drohiotnya 
22, 1-40. 

& Li wis. M Hi M978) "An introduction 

to the freshwater Crustacea of New Zealand". 
Collins, Auckland. 

i'(.i in. I H & Daniuopou D. I. 119781 New 
data 00 the systemarics of the Limnocythcridue 
(OMraeoda. Cylbciacac). d-ohios. II. 563-7 

Coopi-.r. H. M. (1962) 'Australian aboriginal 
mimes- 3000 examples and their meanings". 
South Australian Museum, Adelaide. 

Of DrcKtciit, P. (19771 The distribution of the 
"f-'iant" ostracods ( family : Cyprididae Pwrd 
1845) endemic to Australia. In: Loftier. H & 
Danielopol, D. I. (Eds). 'Aspects of Ecology 
and Zoogeography of Recent and Fossil Ostra- 
coda" lank. The Hz.gue. 

(I97X) Comparative morphology and review 

of muilocypndinid ostracods (family Cypri- 
didae). Aust. J. /<>ul Mtppl Sen 58, 1-62. 

f l'>7 ( > > Ostracods from he motmd spring* 

area hclween Sli anyways and Curdimurka. 
South Australia. Trims* #• St>£. S. Aust. 103. 

tl9X|) Oslracods or attfabttttt salt lakes. 

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ceedings of an International Symposium Junk. 
The Hague. 

& GWU*ES. M. C. f 1980) Seasonal fauna of 
ephemeral saline lakes neat the Coorony I a- 
£00h South Australia. Aust J. Mar. Frt'.shwai 
R, a. 31 877»9P 

& McKhN/if:. K. C (19X1) Rr/itu-lotf:!,. 

new cyprididid ostracod genus ftOm Austra- 
lasia. Tram ft. Snr. S. AuM, 105. 5S-R, 

Oliimi, K. & Pimiommuk. F. f 197?) Q&lraco- 
ilen a us dem hotozanen 1 raven in von Bail 
l.tingcrisalza. QuarttirfHdaontolo^ic 2, 1 19-37 

C.MiTitint, H- (192S) OMmcodes et cladocercs de 
I'Afriquc Ju Nord (2* note), flwft. Sot Nisi 
Nut. A jr. Noni. 19, (&#$, 

Ciioms. M. C. [|,976) SfeftSOWd faUM of some 
ephemeral saline walcrs in western Victoria with 
pai ncular rclcrrnce to Pururirmin tU'WottQ 
Siivtc (Critslaccri: Anostraca). Aust. J. Mar. 
FrWhM'tli f^s, 27, I-'/ 

, Ov DtCMcrit. P.. Williams. W. LV. Mob- 

o.n. IX & "Imitin.,, m. ( |9Xi ) On Ihe 
chemistry and hioto of some salmc lakes m 
Western Australia. In: Williams. W. L>. (ed.) 
"Salt Lakes: of an International 
Symposium". Junk, The Hague. 
Martmann, G. 1 1964) Asiati-che Oslivicoden: 
Systemaiische und Zoogeographische UiiIct- 
sitchungen. int. RfVlte #& rlydmbiol., Syxt. 
Beth 3. 2r\$L 

(!')79) Die osiracoden der Ordnuug Podo- 

copiila C. W. Miiller. 1894 der warm-tempeviii- 
ten (antiborcalcn) Wesi -und Sudwcstkiisle 
Auslralicn* (/uisehtn t\'Hi im Noiden und 
El i c fa im Siiden) Mitt. Numb. /'U>l. Ml{£. insr. 
76, 219-101. 

& kiln C. II97SI 7x\\ "Vuriahilitaf der 

Obellathen Oi narueute des Sehalen Ichendcr 
Osttacoden -Popnlnlionen. ibid. 75. 221-3. 

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water entomostraca of New South Wales, Part 
111. Ostracola. Pro* Linn. So<\ NSW 48. 

HussaInv, S. U. M969) A new speties of Com- 
phbttylhM (Lininocythcridae, Osttacoda) from 
Australia. Pkk. H. S<n\ I'Ut, tp> I %\\ 

K'ir. W, (1932.) Oie ostracoden der Dcutsehcn 
limnolugiMchen Suncb Kspediiion Anh jtvtint- 

hn>!., S'lippl. 11. 447-502. 

MrKi sJ/if- K,.C 11978) Osiracoda (Crustacea: 
Popocopida ) from soulhern Australian salt, with the description of Rtttcyprfa new 
genua. Trteib H So* v -<W- 'W, 175-90. 

& j-InssMNY, S. U- (I9n8) Relevance of q 

freshwater cythend (Crustacea, Ostracodu) to 
(he continental drift hypothesis. Nature Land, 
22», XOtS-S. 

Pahps. B. H. 1905 -Aborieinal Words of Aus- 
iraJiu*' Reed, Swlney 

Pi ikov.SKi. r. K- ( I9S8) Siisswasser osiracoden 
aus Jugoslavicn l[ Subfam llyocyprinae. Wav. 
Mac: Scient. Nurar 2(H), 5? -7. 

SABS, <j. O. 118891 On some breshwater Ostra- 
cuda. and Copepod.i ruisud fiom dried Australian 
mud, Fordi, \ Stthk. Krhf. /4M », 3-79. 

(1B94) Contribinions to ihe knowledge of 

Ihe freshwuter JLntomostraca of New Zealand 
as .shown hv arttlkud haleldng from dried mud. 
ihid, r$9j 5, I 6 1 

(1896) On some Freshwater Kntomostraca 

from the Neighbourhood of Sydney, partly 
raised from dt icd mud. Afrit Math. Natarw 18, 


Shokmkju I. I 11966) Sesual dimorphism and 
shell variation in Leptacythert'. lit: Vyaluv. 
O. S. (ed.). bitssil Dsfrauoda — papers from the 
First AH-L'nion Symposium on Fossd Ostra- 
eoda f I vo\, 1963 )r Tr;m%tated by Israel 
Program for Scientific Translation, Jerusalem 
< iy7i ) 


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the limnology of three maar lakes in western ter s to diagnose the species of the llyocypris 

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c * a . j .,, i ******** cal significance. In: Krstic, N. (ed.). Taxo- 

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ostracodes". Serbian Geological Society, 


by Ian Beveridge 


Three new species of Calostaurus Sandars (Cestoda: Davaineidae) are described from the small 
intestine of the macropodid marsupial Dorcopsis veterum from New Guinea. C. dorcopsis sp. nov. 
differs from C. macropus in the size and arrangement of rostellar spines, size of rostellar hooks and 
cirrus sac. C. oweni sp. nov. and C. parvus sp. nov. resemble C. mundayi in features of the 
rostellum, but differ in size, number of testes per proglottis and number of eggs per egg capsule. 


by Tan Bevi:ridgo 


Brveridge, I. (1981) Three new species of Cohstaum.s (Cestoda. Davaineidae) from 
the New Guinea wallaby Dotropsis KOYMM I'rnns. R. Soc. S Ausu it)5(3). I3*J-147, II 
December, \HH\. 

Three new species 61 Catosnutmt Sandal (Cestoda: Davaineidae) are described liom the 
EUJUfl intestine of Hie macrapodid marsupial Doreopstv vetvatm from New Guinea. C 
,i<-nupsi\ sp. nov dittos t um C nnuropiis in the si/e and arrangement of rosteilar spines, 
■ wr of rosteilar hooks and cirrus sac. C. trwmi sp. nov, and C. parvus *P. nov. resemble C\ 
tntttultwi in features ot the roMellum, hut difter in »io, numbei of testes per proglottis unci 
number of eggs per eye capsule. 


Three species of the davaincid cestodc 
genus CaloMiuirus Sandars, 1957 are known to 
occur in Australasian macropodid marsupials, 
namely C. mucropih (Orilcpp, 1^22) originally 
described from Thyhwaie hrunii (Shreber ; 
I77S) (Syn. Maaopits brunti) and redes- 
cribed by Sandars (1957) from Thyhpah 
tfttimaftcat Gould. I860 (syn. 7. *Hcoxi) in 
Queensland, and C, thvlo^tlv Bcvcneigc, I 975 
and C twwdayt from ThytoKo.ic hitlttrtHcru 
(Desmarcst. 1822) and Potorous tftdaC&Uto 
(Kerr. 1**72) (syn P apicah\) respectively. 
both occurring in Tasmania (Bcvcridge, 1975) 
In each instance, a single host species is para 
sitiscd by one eestode species oti!\ By contrast 
in collections of cestodes made recently from 
the wallaby Durcopsis vetcrutn (Lesson. 1K27) 
in New Guinea by Dr I. L Owen, three CBSlOtU 
species may occur in the same nost individual 
The three species are described in this paper. 


Cestodes were stained with Cdesrine blue. 
dehydrated and mounted in balsam. Serial 
sections of each species were cut but the slate 
of preservation oi' the. specimen-, did not allow 
detailed histological examinminn. All measure- 
ments are given in mm us the range followed 
bv the mean of 5 measurcmenls m parentheses 

Calostaurus dorcopsis sp DDi 

FIGS I- 10 

Description based on fragments i\\' several 

specimens; t\o complete cestodes presort 

dorsal system, ramifying across proglottis In 

Institute of Medical <V Veterinarv Science., t'r^me 
ROadi Adelaide-, S. Aust. 5000. 

Moderate sized cestodes. Maximum length of 
fragments 105; maximum width 4. Largest 
fragment contains 350 proglottides Scolex 
large. 0.^0-1.16 (1.00) in diameter, quadran- 
gular in tifi I'uv view, with evcrsible rostellum 
6.42 49 (0.52) in diameter Rostellum 
armed with approximately IhOO hammer 
shaped hooks in iwo rows, arranged in shape 
oi' Maltese cross, with axes extending between 
pans of suckers; hooks 0.012-0.014 (0.013) 
long. Base of rostellum armed with approxi- 
mately 16 transverse rows of rose thorn 
shaped hooks each 0.007-0.011 (0,010) long, 
with awl shaped blade and irregular and highly 
variable bifid base. On everted rostellum. hooks 
extend almost to extremities of Maltese cross; 
never lie between arms of cross. Rostellum 
vanes considerably in shape (Figs 1, 2) de- 
pending upon degree of extension Rosteilar 
spines form narrow band anterior to rostellum 
at opening to rosteilar sac when latter fully 
rwracted, ^Suckers 0.27-0.32 (0,20) in dia- 
meler; margins armed with seven to nine rows 
of tinv awl-shaped spines 0,006-0.008 (0.007) 
long. Neck 38-0.75 (0.52) long. 

Proglottides eraspedotc: velum overhanging 
adjacent proglottis, very narrow. Mature pro- 
glottides greatly extended transversely. 20- 
0.45 (0.36) > 2.4-4.0 (7.9), ratio width: 
length 1:5-1 "20. Genital pores single, uni- 
lateral Genital ducts pass between osmoregu- 
latory canals. Dorsal osmoregulatory canal ex- 
tremely narrow. 001-0.03 (6.02) in diameter, 
lies well internal to ventral canal: ventral 
osmoregulatory canal 0.05-0.09 (0.07) in 
diameter. Transverse osmoregulatory canals 
connect left and right dorsal and venlral canals 
at posterior margin of each proglottis. Com- 
plex, branching network of canals arises from 





Figs 1-10. Calostaurus dorcopsis sp. nov. from the wallaby Dorcopsis veterum. 1. Scolex with partially 
everted rostellum. 2. Scolex with fully evert rostellum 3. Scolex with retracted rostellum en face 
view. 4. Female genitalia, dorsal view. 5. Rostellar hooks. 6. RosteUar spines. 7. Sucker spines 8 
Cirrus sac and distal vagina. 9. Single mature proglottis. JO. Gravid proglottis Scale lines* Fies 5-7 
0.01 mm; Figs 1-4, 8-10, 0.1 mm. • & • 



largest fragment, genital anlage appears in ap- 
proximately 50th proglottis- Male and female 
genitalia mama? in proglottides 200 and 260 
respectively, and involute in proglottides 425 
and J4Q respectively. First gravid proglottis 

Genital atrium small, .situated In anterior 
half of lateral proglottis margin. Cirrus sac 
small, 0.20-0.25 (0.22) X 0.07, elongate, 
muscular wails feeble, cirtus sac not reaching 
ventral osmoregulatory canals. Distal region ol 
emus of greater internal diameter, lined with 
bristles; remainder narrow, coiled. Internal and 
external seminal vesicles absent. Vas deferens 
slightly coiled, narrow, passes medially to- 
wards centre of proglottis, terminating dorsal 
To ovary Vasa effcrcntia not seen. Testes in- 
variably distributed in two lateral groups, ex- 
tending from ventral osmoregulatory canals, 
below dorsal canals, medially. Testes densest 
in lateral regions of each field; testes never 
overlie female genital glands, testes 0.05-0.06 
(0,06) in diameter. Testes number 37-51 per 
pro^lutlis; alwavs more testes in aporal Held; 
H2:i (15) testes in poral field, 21 -V> (31) 
in aporal field. Some differences evident in 
lestis number between strubilae: in one 
Strobila $-13 (101 poral. U 39 (37) aporal: 
in second slrohila 16-23 (20) poral. 22-28 
(24) aporal. 

VaginA opens to genital atrium posterior to 
cirrus" sac. Distal region. OOS^-0.13 (0.10} X 
0,02, dilated, armed with fine bristles. Mid- 
region narrow, uncoiled, leads medially, 
posterior to vas deferens, terminates in fusi- 
form seminal receptacle 0.12-0 18 (0.15) X 
.03 0.07 (0.05) in size, lying dorsal lo poral 
lobe of ovary: sperm duct passes posteriorly 
from seminal receptacle, lined with bristles. 
Ovary bilobed, situated to poral side of pro 
glottis midline. 15 0.20 (0.18) X 0&6-&)! 
(012) in size, with 8-10 elavate lobules hi 
each lobe. Vitellarium ovoid or bean-shaped 
07-0,09 (0.OK1 X 0.12-0.16 (0 IS) in vizt. 
posterior and dorsal to ovary. Melibs* gland 
spherical, anterior lo vitellarium. Uterine duel 
short, passes anteriorly from Mehlis' gland,, 
terminates dorsal to ovaty. Uterus absent Egfeft 
discharged from uterine duct directly Into 
parenchyma, become surrounded by capsule, 
with one egg per egg capsule. Size of egg 
capsule 005-0 06 (6.055): egg 0.01-0.02 
(0.015). Gravid proglottides extended trans- 
verse! v 0*21 0.70 (0.49) X 2.2-3.6 (VI), 
width, length ratio 1 :3-l :7 Terminal pro- 

glottides relatively longer, occasionally as long 
a& (I.03) >\ t .05-1. 55 (I 31 1. 

Uitst; l>onopxis vetvntm (Lesson, 1827) 
(M;irsiipiali;i; Maeropodidae). Locuiiott; Small 
intestine. Type Locality. Veikabu Creek, On 
tral Province, 9' 10'S. I47 09J-. Papua New 
Guinea. Types: In British Museum (Natural 
History), holotype 1 08 1 .6. 1 7. 1 , paratypes, 
1981.6.17.2-10. Eiymolouy; The specific name- 
is derived from I hat of the bos? 

Cutost-aurus owenl sp. nov. 
FIOS 11-19 

Description based on 10 complete mounted 
specimens. Small cc&todcs, 24-^8 (33) long; 
maximum width 1 2-1,4 (13); strohiW con- 
tain 100-147 (120) proglottides. Scole\ rela- 
tively large, 0.62-1.05 (0.99) in diameter, 
quadrangular in en /act 1 view, with rostelkim 
up lo 0.45 in diameter when everted. Rostcllurn 
tinned with approximately 1000 hammer 
shaped hooks in two rows, arranged in lorm 
of open cross, with axes extending between 
pairs of suckers; rostellar cross 0.25-0.32 
(0.29) X 0.18-0.34 (0.2M; rostellar hooks 
0.007-0.010 (0.008) lonir, Base of rostcllurn 
armed with concentric rows id tiny awl shaped 
spines approximately 0.004 long. Spines form 
band posterior to rostellar cross on evened 
rostellum, never lie between arms of cross, 
spines form band anterior lo rostellat cross, 
at opening of rostellar sac when rostcllurn 
lolly retracted. Suckers 0.16-0.23 (021) in 
diameter; margins armed with aboul eight 
r0W8 of tiny, rose thorn shnped spines 0.004- 
0.007 (0.006) lonK- Neck 0.21-0.34 (0.28) 

Proglottides efaspedotc, velum ovcrhaneimi 
adjacent proglottis very narrow. Mature pro- 
glottides wider than W, fo7-Q,33 (011) X 
120-133 (1.25), rntio lenglh:wtdlh 1:3.5- 
1 :4.3. Genital pores single, unilateral; one 
proglottis found with reversed orientation. 
Genital ducts pass between osmoregulatory 
canals Dotsal osmoregulatory canal extremely 
narrow, 005 m diameter, sinuous, lies in- 
ternal to ventral canal; ventral osmoregulatory 
eaiial straight, 02-0.04 (0.03) in diameter. 
Transverse osmoregulatory canals connect left 
and right dorsal and ventral canals at posterior 
margin of each proglottis. Origins of dorsal 
transverse canals sometimes variable, single 
transverse canal occasionally arising from two 
or more points on dorsal longitudinal canal. 



Figs 11-19. Calostaitrus oweni sp. nov. from the wallaby Dorcopsis vcterum. 11. Scolex with retracted 
rostellum, lateral view. 12. Scolex with retracted rostellum, en face view. 13. Mature proglottis. 14. 
Gravid proglottis. 15. Rostellar hooks. 16. Rostellar spines. 17. Sucker spines. 18. Cirrus sac and 
distal vagina. 19. Female genitalia dorsal view. Scale lines: Figs 15-17, 0.01 mm; Figs 11-14, 18, 19, 
0.1 mm. 



Genital anlagc appears iti proglottides 25-40 
C$1); Male and female genitalia mature in 
proglottides 52-73 (64) and 70-98 (81) re* 
spcchvcly, and involute iu proglottides 85-112 
(97) .md 75- 102 (91) respectively. Firsl 
gravid proglottis 89-120 (104), 

Genital atrium very small, situated in middle 
of lateral proglottis margin. Cirrus sac large. 
0.16-0 IX (S0.17) X 0.09-0,11 (0.10), ovoid, 
muscular walls feeble, cirrus sac not reaching 
longitudinal osmoregulatory canals Digftl 
region of citrus of greater interna! diameter, 
lined wilh bristles; remainder narrow, coiled 
muscle bands run from cirrus to internal SU? 
face of cirrus sac, Internal and external 
seminal vesicles .absent. Vas deferens greatly 
coiled, passes medially towards centre of pro- 
glottis, gradually diminishing in diameter, ter- 
minates dorsal to ovarian isthmus by dividing 
into two vasa efferentia each .supplying one- 
group of testes. Testes almost invariably dis- 
tributed in two lateral groups, in posterior half 
of proglottis, extending medially from dorsal 
osmoregulatory canals. Fn very occasional pro- 
glottides, one or two testes present posterior 
to vitellarium joining two lateral testis 
groups. Testes ovelie ovarian lobes and oc- 
casionally vitellarium: testes 0.045-0.055 
(0 050) in diameter. Testes number 15-19 
(16> per proglottis; always more testes in 
aporal Held- 3-7 (5> poral. 10-14 (12) 

ViitMna opens to genital atrium postenor to 
cirrus sac. Distal region. 05-0.08 (0.07) Y 
0.02. dilated Mid-region narrow, sinuous, 
leads medially, posterior tn vas deferens, ter- 
minates in fusiform seminal receptacle, 0.07- 
0.11 (0.09) X 0.03-0.04 (0,04) in si/.e. lying 
anterior and dorsal to pora! lobe of ovary; 
sperm duct passes posteriorly from seminal re- 
ceptacle, dilates, lined with bristles. Ovary 
hilobcd, situated to poral *ide of proglottis 
mid-line. 10-0 IS (0.13) X 0.17-0.20 
(0 IS) with 4-5 clavate lobules in each lobe. 
Viiellurium ovoid or bean shaped, lobniate, 
05-0 07 (0.1)6) X 0.07-0.08 (0.07) in size. 
situated posterior and dorsal to ovary. MehbY 
eland spherical, anterior to vitellarium. Uterine 
duel passes anteriorly from Mehlis* gland ter- 
minated in midline dorsal to ovary. Uterus 
absent. Eggs discharged from uterine diM 
directly into parenchyma, become surrounded 
b> capsule, with one egg per egg capsule. Hue 
Of egg capsule 0.04-0.05 (0.047); egg 0.01- 
02 (0.15). Gravid proglottides extended 

transversely 0.41-0.55 (0.45) X 0.95-1 33 
(1.20), width:length ratio hi 17-1:2 7. 
Terminal proglottides almost as long as wide 
or longer than wide, 0.62-0,75 (0.69) X 
0.52-0.88 (0.70). width:lenglh ratio 1:0 74- 
1 1.3. 

Host: Doixopsh vcren/m (lesson, IS27) 
(Marsupialia: Macropodidac)- location: small 
intestine. Type Locality Vcikabu Creek, Cen- 
tral Province, Papua New Guinea Types: In 
British Museum (Natural History), holotype 
1981.6.17.11, paratypes 1981.6.17.12-21 

Etymology, this species is named after Dr 
I. L. Owen who collected all the material 
described here. 

Cafostaurus parvus sp nov. 
FIGS 20-27 

Description based on live complete mounted 
specimens. Small cestodes, 9.0-11.2 (9.7) 
long; maxrmum width 0.75-1 .20 (0.9rV) ; 
strobilae contain 38-57 (42) proglottides, 
Scolex relatively large, 0.62-1.08 (0.76) en 
diameter, quadrangular in en fact' view, with 
eversible rostellum, Roslcllum armed with 
approximately 1000 hammer-shaped honk* m 
two rows, arranged in form of open cross, 
with axes extending between pairs of suckers; 
rostellar cross 0.29-0.47 (0,35) X 0.28-0.42 
(0.34), rostellar hook* 0,009-0.010 (0.010) 
long. Base of rostellum armed with concentric 
rows of tiny awl-shaped spines, with simple or 
bifid base; spines 0.004-0.005 (0005) long. 
Spines form band anterior to rostcllai cross. 
at opening of rostellar sac, when rostellum 
fully retracted. Suckers 0.19-0.23 (0.20) m 
diameter, margins armed with about 10 rows 
of tiny rose thorn shaped spines 0.002-0.005 
(0.003) long. Neck short. 

Proglottides craspedote; velum overhanging 
jjdjaccnt proglottis veTy narrow. Mature pro- 
glottides wider than long 0.24-0 35 (0.32) < 
0.64-0.98 (0.75), ratio length. width 1*2.0- 
1:3. S. Genital pores single, unilateral Genital 
ducts pass between longitudinal osmoregula- 
torv canals. Dorsal osmoregulatory canal ex- 
tremely narrow, sinuous. (305-0.01 (0.006) 
in diameter lies immediately internal to ventral 
canal; ventral osmoregulatory canal 0.015- 
0.020 (0.017) in diameter. Transverse osmo- 
regulatory canals connect left and right ventral 
canals at posterior margin of each proglottis. 
Genilal antage appears in proglottides 8-28 
(12). Male and female genitalia mature in 




i ' - * ! , 

■ r. ■ ■■■.-;..-*-, ■ . ' ft .U 

:'o / 

-, ■ 

! ' o a ! 

■ i ■■■■ ' 

Figs 20-27. Colostomas parvus sp. nov. from the wallaby Dorcopsis vetoram. 20. Scolex with everted 
rostellum, lateral view. 21. Scolex with retracted rostellum, e/i face view. 22. Mature proglottis. 23. 
Rostellar, hooks. 24. Rostellar spines. 25. Sucker spines. 26. Cirrus sac and distal vagina. 27. Gravid 
proglottis. Scale lines: Figs 23-25, 0-01 mm: Figs 20-22, 26, 27, 0.1 mm. 



proglottides 14-37 (22) and 20-28 (25) re 
spcetively and involute in proglottides J3-36 
(55) and 10-34 (32) respectively. First 
gravid proglottis 34-38 (36). 

Genital atrium insignificant, situated slightly 
anterior to middle of lateral proglottis margin. 
Cirrus sac smalt, 0.07-0,10 (0.09) X 0.04- 
0.05 (0.04). c-lavatc, muscular walls feehlc. 
cirrus Mac almost reaching longitudinal 
osmoregulatory canals. Distal region of cirrus 
of greater internal diameter; armature not seen; 
mid-region narrow, sinuous; proximal region 
form* small, spherical internal seminal 
vesicle, present in most proglottides of all 
strohjlae examined, hut not detectable in every 
single proglottis, internal seminal vesicle 
0.020M)035 (0.025) X 0.025-0.030 (0.026V 
External seminal vesicle absent. Vas deferens 
narrow, greatly coiled, surrounded by pale 
basophilic cells, passes medially towards centre 
of proglottis, then posteriorly towards ovary; 
terminates dorsal to ovarian isthmus; vasa 
elTerenlia not seen. Testes distributed in 
posterior part of proglottis, in single field, con- 
tinent posterior to vitcllarium; testes lie within 
area bounded by lateral osmoregulatory 
canals; usually lie medial to dorsal canals, but 
few testes may overlie dorsal canals. Testes 
not confluent posterior to vitellaria in few 
proglottides. Testes usually overlie ovary and 
vitellarium: testes 0.04-0.05 (0.045) in dia- 
meter. Testes numbei 17-22 (20) per pro- 
grotlrs; poral testes 6-10 f8), invariably less 
numerous lhan aporal testes 11-14 (13). 

Vagina opens to genital atrium posterior to 
cirrus BHC Distal region. 0.06 V. 0.02, dilated. 
Mid-region narrow, sinuous, leads medially, 
posterior to vas deferens, terminating m large 
clavate to pyriform seminal receptacle, 0,08- 
0.13 (0.09) X 0.04-0.05 (0.05X fying 
anterior and dorsal to poral lobe of ovary; 
speiin duct elongate, sinuous, passes posteriorly 
from seminal receptacle Ovary bilobed. ftlftj- 
ated in proglottis midline. 0.09-0.16 f0.I41 
x (1,14-0.16 (0-15) in size with 4-6 clavate 
lobules in each lobe. Vitellarium ovoid or bean 
shaped, Inbulate. 0.05-0.08 (0.06) X 0.04- 
0,05 (0 05) in size, situated posterior and 
dorsal to ovary. Mehlis' gland spherical. 
anterior to vitellarium. Uterine duct passes 
anteriorly from Mehlis' gland. terminates in 
midline, anterior to ovary Uterus absent. 
fciggs discharged from uterine duel directly 
into parenchyma, become surrounded by 
capsules, with one egg per egg capsule. Size 

of egg capsule 0.05-0.07 (0.06); egg 0.015. 
Gravid proglottides longer than wide 0-65-0.75 
(0.69) X 0.67-1.10 (0.83), ratio length; 
width 0.64:1-1 : 1.1. Terminal proglottides 
0.85 x 0.58-0.68 (0.73). ratio length:width 
1.2:1-1.4 I. 

Host: Dorcopsi.f vctetum (Lesson, J X 2 7 ) 
( Marsupialia ; Macropodidae) . Location . 
Small intestine. Type locality: Veikabu Creek, 
Central Province. Papua New Guinea. Types 
In British Museum ("Natural History), holo- 
type 1 981 .6. 19. 1 . paratypes 1981 .6. 1 c >2-6. 
Etymology- the specific name parvus derives 
from the fact that it is the smallest known 
species of the genus. 


Three species of Calostaurux have been des- 
cribed previously, being distinguished prin- 
cipally by the shape of the cross formed by 
the tostellar hooks. C. macropus has the hooks 
arranged in the shape of a Maltese cross, 
C. thv!of>ule in a siK-lobed circle and C tnun- 
dayi m a four-lobcd cross. Of the species des 
erihed here C. dorcopus sp. nov., most closely 
resembles C. macropus in that the hooks are 
arranged in the form of a Maltese cross 
whereas C, oweni sp. nov. and C. parvus sp. 
nov. both have ro.slcllar hooks arranged in the 
form of four-lobed crosses similar to lhat of 
C, mumktyi. The histological anatomy of the 
seoleees of the new species is similar to that 
of C. rh\ locale and C. mundayt and has there- 
fore not been described in detail 

Both C owctn and C. parvus can be dis- 
tinguished from C tmttulayi by size alone, 
since r. mioidayi measures 32.4 to 45. ( cm, 
whereas C oweni and C. parvus measure 2.4- 
3.8 and 0.9-t.<I cm respectively. In addition 
f . mundayt has one to three eggs per egg 
capsule while the other species hove a single 
egg in each capsule, and the range of testis 
number in C. mtmdoyi (25-32) is higher lhan 
in C oweni (15-19) and C. pOTVld (17-22). 

C. ojvefjl and C, parvus differ from one 
another in a number of features. Gtavid speci- 
mens of C. parvus are consistently smaller and 
have fewer proglottides than C, oweni, and 
there is a difference in the rate of development 
and involution of the genital organs in the two 
species. The testes are generally confluent 
posterior to the vitellaria in f . parous, but .ire 
divided into two groups in C. oweni. Some 
variation in this character does occur and 








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occasional proglottid es of C, parvus have the 
testes in two groups, while occasional pro- 
glottides of r. owem have the testes confluent 
posterior to the vitellaria, but in an individual 
strobita, one form or the other very clearly 
predominates. The cirrus sac of f rwnfs 
(0 07-0010 mm) is invariably smaller than ii! Cowetti (0.16-0.18) though the size 
difference may be a reflection of overall 
ceslode size. However, all specimens of 
C. parvus examined possess an internal seminal 
vesicle, a feature which is not present in any 
congener, ll is not detectable in every single 
proglottis, but is invariably visible in some pro- 
L-lotiides of a strobild These various differences 
arc considered adequate foT the erection of 
two new species. 

Comparisons ol C, dorcopsis with C. macm- 
/>//v ire complicated by the fact that the latter 
species has been described on two occasions 
firstly by Ortlepp (1922) from Thylovate 
tffmti and Inler by Sandars ( 1957 V I rent 
7 h vlogale stivmntica ( =Thyfos>ate wilcoxi) , 
the descriptions, differing on some important 
features such as size and the number of the 
rostellar books. 

C. dorcopsis is similar: to both descriptions of 
C. maaopus in the form of the rostellar cross, 
in tlic number and distribution of the testes 
and in the transversely elongated mature pro 
glottis. C dowopsis would appear to ditfer 
I rem all congeners by the distribution of the 
Mies which extend laterally beyond the dorsal 
osmoregulatory canals to the medial edge of 
ihe ventral canaLs. However, this- condition also 
c\isu in the type material of C. macropus in 
the British Museum (Natural History), 
(tboueh not stated by Ortlepp (19221 in hs 
description) and in material collected by me 
from ) hyloRute stipjuaiira in Queensland, con- 
forming in all respects to the specimens des- 
cribed by Sandars (1957) from the same host 
;ind same general geographic region- Sandars 
f 1 957) gffve the locality of her specimens 
incorrectly as *Mt. Tarn bonne m South Aus- 
tralia; Mi. Tambotinc is in Queensland; no 
species of Thyln-:o!r occur in South Australia. 
C dorcopsis drITers from C tnacwpus in a 
number of features. The cirrus sac <n 
C- (/orr.v'vs (0.20-0.25) mm) Is larger than in 
descxiptiote (A C fflBctOPto (0)0, 0.08-0.12 
mm); the roskllar hooks of C dotcopsii (12- 
1 5"m ) are sUghtty longer than these of 

C macropus (9/<m) and the rostellar spines of 
C. dorcopsis (7-1 lMm) are longer than those 
of C macropus (5-6Mm) and are of a different 
shape. The differences in the rostellar spines 
arc very striking indeed as they are very pro- 
minent rose-thorn structures on the rostetla of 
C. dorcopsis but are inconspicuous and awl or 
hook shaped on rostella of C. macropus. Both 
Ortlepp (1922) and Sandars (1957) des- 
cribed und illustrated the rostellar spines of 
C macropus as covering the entire rostcllum 
with the exception of the area inside the 
tostcllar cross. It has not been possible to 
verify this situation since in the type specimen 
examined by me and in the new material from 
1 , \tjonui?ir<i, most of the rostellar spines are 
missing. Assuming the earlier descripiions to be 
correct, C dmcopvis differs markedly from 
t \ macropus in having the rostellar spines 
arranged in about 20 transverse rows, forming 
a distinct band on the everted rostellum but 
never covering the entire surface of it. 

Because of these differences, C. dorcopsis sp. 
nov. has been described. It is evidcnl however 
that the morphology o\ C macropus requires 
re investigation based on new material from 
the type host, T. hrunil The extant type 
material is inadequate for 9 thorough re- 
description and the status of specimens from 
7. si^'aatira, described by Sandars (1957) as 
C macropus requires clarification. 

The principal of differences between species 
of Ctdnstaurus are shown in Table 1- 


Thanks arc due to Dr I. L, Owen. Bnroku, 
New Guinea, for collecting and very carefully 
preserving the material described above, and to 
Mr R. A Bray, British Museum (Natural 
History) for kindly lending the material for 


Bt-vMiiDCC, I. (1975) On two new davaineid cbb- 
todes from Australian marsupials. J. ficlmfn- 
thof 49, 12^-36. 

OftiThPP. R. I. (1922) A new davaineid ccstorte 
— Railliftina ( Parrtniclla) maernpa, sp. n , from 
a wallaby. Ann. map fiat Hht., .SVr 9, 9, 
602- 12. 

SaNOARS, D. F, (1957) Redescriptton of fconic 
cestodes from marsupials (T-Oavaineidae, Hvmc- 
nolcpididue and Anoplocephalidac. Attn. Tr<>p 
Med- Parasitot. 57 330-9. 



by Michael J. Tyler, Margaret Davies & Angus A. Martin 


Uperoleia trachyderma sp. nov. is described from the Northern Territory, and additional biological 
and distributional data are given on U. lithomoda Tyler, Davies & Martin. Ranidella deserticola 
Liem & Ingram is recorded in the Northern Territory for the first time, and Cyclorana cryptotis 
Tyler & Martin is recorded for the second time. 


by Michael J. Tvi .er*, Margaret Davies* & Angus A. Martin! 


'1'VUK, M. L. DAVIES, M & Martin. A. A. (1981) Frog fauna of ihc Northern Territory: 
new distributional records and the description of a new species, Trnns. ft, Soc. S Anst. 105 
Hj, 149-154, M December, 1981. 
Vperoivia itovhydennn sp. nov. is described From the Northern Teiuiory, ;»nd additional 

biological iind distributional data arc gtVfcn on f UihnnujJo Tyler, Davies & Marlin. Rani- 

(h-llo tlesciih (ila licm & Ingram is recorded in the Northern Territoiy for the first time, and 

Cxilonma crvptatix Tyler A Marlin is recorded for the second time. 


Our collaborative sludics on the frogs of 
northern Australia have led to the addition 
of seven species lo the North Territory fauna 
/Martin, et at. 19X0; Tyler, er al. 1978, 1981a; 
Tyler, el aL 1979). Our previous held studies 
in the Northern Territory have been confined 
to the fast Alligator River area. 

However In December 1980 we collected at 
various .sites on the Stuart Highway between 
Darwin and Tennant Creek. These collections 
include a new species of the leptodactylid 
gcitUS Upvruhio, and a second leptodactylicl 
species not previously recorded from the 
Northern Territory. We also provide additional 
information on the distribution and biology Of 
U. Ittlunnothi Tyler, Da vies & Marlin, pre- 
viously recorded in Ihc N.T. only from the 
flood plains west of Arnhem Land (Tyler et 
trl. 1981a). and the hylid Cyrhnvia tfypt&fis 
Tyler & Martin 

Material and Methods 

I he specimens reported here arc located in 
the following collections: American Museum 
o( Natural History i AMNIO; Northern Ter- 
ritory Museum, Darwin (NTM). Museum of 
Natural History, University Of Kansas. Law- 
rence ( KD ) ; South Australian Museum 

Methods of measurement follow Tyler 
(190$) Abbreviations used in the text arc. E 
(eye diameter), IvN (eye to uuris distance), 
IN (interna rial span). S-V (snout to verit 
length), TL (tibia length). 

* Department of /oology, Umversitv of Adelaide 
Huk 4MS G.I'.O., Adelaide, S. Ausl. 5001. 

I IVpaiimenl of /oology, University of Mel- 

Osteological data were obtained from 
cleared and Alizarin Red stained preparations 
using the technique of Davis & Gore (1947)- 
OstcologicaJ descriptions follow Trueb (1979). 

Male mating calls were recorded with a 
Sony TC-510-2 tape recorder and a BeycT 
M 88 dynamic microphone, at a tape speed 
of 19 em/sec. Wet-bulb air temperatures Were 
measured close lo the calling sites of males 
usme, a Schullhers quick-reading thermometer. 
Calls were analysed by means of a UV oscil- 
lograph (San-Ei Visilighl) and a sound spec- 
trograph ( Kay Model 606 1 -B Sona-Graph) 
with the overall response curve maintained 
in the FL-I position. Three calls of each male 
were analysed and mean values calculated. 

b am ilv : Lcptodat'tyliriac 
Iperoletu (raehyderma sp, nov 

pros 1-4 

Ut'fotyp? SAM R20374, an adult male col- 
lected on the Newcastle Creek floodphun at 
the George Redman Causeway ( I7°M'S; 
I3V28'R) L 37 km N of Elliot, N.T. on 
16.xh.80 by M. Davies, A. 4, Martin and 
M. J. Tyler. 

Definition: A small species (males 20.3-22.0 
mm S-V l with small eyes (eye diameter 
equivalent to eye to nans distance), with the 
dorsum covered with small, conical tubercles, 
and with a heavily pigmented ventral surface. 
Maiing call a staccato burst o\ four short 

Description of Iwlotype: Maxillary and vome- 
rine teeth absent. Snout elongate, pointed 
when viewed from above; projecting slightly 
in profile. Rye to naris distance greater than 
internarial span (E-N/ IN 1.27). Canthus 
rostralis straight. Tympanum not visible ex- 
ternally (Fig. I ), 

I Ml 


F'\z. 1. Dorsolateral view of head of VperoU'tu 
\rtii hythrrrif/ 

Fingers long, slender, unwebbed and un- 
tnnged with prominent subarlicular tubercles 
and well developed palmar tubercles (Fig. 
2A». Fingers in order of length 3 > 4 > 2 > 
1 Hindlirnbs very short (TL/S-V 0.35). Toes 
long, slender, slightly fringed and unwebbed 
(Fig. 2B). Toes in order of length 4 > 3 
5 > 2 > I Met alius a I tubercles small and 
poorly developed. Subartieular tubercles small 
but prominent, 

Dorsal surface densely covered with small, 
conical tubercles. Parotoid and inguinal glands 
inconspicuous; coccygeal glands not visible 
externally- Ventral surface finely granular. 
Cloaca! flap narrow but well developed. No 
heel tubercles. 

Dorsal surface dull slate with obscure 
slightly darker mottling; glands faintly 
creamish. Inguinal and post femoral flash 
markings dull red. Ou ventral surface sub- 
mandibular area pale grey; nhdomen with 
numerous small islands of pale grey on a while 

Dimensions; Snout to- vent length 20.75 mm; 
tibia length 7.3 mm; eye to naris distance 
1.9 mm; internarial span 1.5 mm; eye diameter 

1.9 mm. 

Variation; There are four paratypes; all are 
adult males: KU 189561, NTM 9865, SAM 
R20375-6. The scries was collected wilh the 


The range Of body size spans only 1.7 mm 
(20.3-22.0 mm S-V). The hindlimbs are short 
(II S V 0.33-0.36) and the metatarsal 
tubercles (so well developed in congeners) are 
comparatively poorly developed. The eye dia- 
meter is consistently small and approximates 
the eye to nans distance. The eye to naris dis- 

tance is greater than the internarial span 
CE-N/'IN 1.13-MO). 

The unusual tubercular condition of the 

dorsal skin is apparent in all specimens, and 

«n some is more conspicuous than in ttrc 

The dorsal coloration is reasonably uniform 
except thai iu one specimen the parotoid, 
inguinal and coccygeal glands arc brighi 
yellow-cream and stippled with blaek. In all 
specimens the dark mottling on the abdomen 
is more conspicuous than in the holotype. 

In life the dorsum is variegated with 
clearly defined patches of grey on a paler grey 
ground colour, except for the dermal glands 
which are suffused with pale orange. The 
inguinal and post-femoral flash markings arc 
bright carmine. The Mib-mandibular area is 
slate, and the abdomen is marked with islands 
oi' grey upon a crcamish-grey background. 

Osteology Skull (Fig. 3A) moderately ossi- 
licd, sloping antero vent rally. Sphenethmoid 
poorly ossified, not conjoined medially cither 
dorsally or ventrally, Dorsally sphenethmoid 
in tenuous contact with nasals. Ossified portion 
of sphenethmoid extending ventrally for 
about $ length of orbit. Prootic and exocei 
pital not fused dorso- or ventromedially. 
Fenestrae present dorsally and ventrally in 
crista parol ica. Crista parotica short, thick, 
slightly overlapped medially by posterior ex- 
tremities of frontoparietals, widely separated 
laterally from uncxpanded otic rami of 

Fig. 2. (A) Palmar view of hand and (H) plnnta' 
view of foot of U-PtMtek (rachxderma. 



Fig. 3. (A) Dorsal and (B> ventral view of skull of UjwraMtt innhxifcrma. 

Frontoparietal fontanclle extensively ex- 
posed; anterior and posterior extremities being 
Undefined because of lack Of medial ossitka- 
tion of sphenethmoid arid exoccipital regions. 
Nasals extremely well ossified, lying along- 
side each other medially. Maxillary processes 
of nasals short and acuminate, not in bony 
contact with very poorly developed preorbital 
process of shallow pars facialis of maxillaiy. 

Palatines reduced ( Fig. 3B), moderately 
broad, overlying ossified poitfofl of spheneth- 
moid medially at angle of approx. 45*, 

Parasphenoid robust; long, broad cultiform 
process, almost truncate, reaching to level 
of posterior edge of palatines. Alae broad, 
short, slightly angled posteriorly from cultri- 
form process. Pterygoids moderately robust; 
anterior arm slender, in moderately short con- 
tact with poorly developed pterygoid process 
of palatal shelf of maxillary Medial arm 
moderately long, very robust; poslenor arm 
short and moderately sharp. 

Cartilaginous quadrate present between hase 
of squamosal and quadratojugal. Quadrato- 
jugal robust, in linn contact with maxillary. 
Squamosals stocky, no zygomatic ramus, 
long uoexpanded otic ramus. MaxiJIary and 
premaxillary edentate. Alary processes of pre* 
mamillaries moderately broad, perpendicular 
to piema\illar\\ Palatine processes ol' pre- 
maxillanes well developed, not abutting 
medially- Palatal shelf moderately deep with 
poorly developed pterygoid process. Prevomers 
absent. Bony columella present. 

Pectoral girdle arciferal, robust. Omo- 
sternum absent, xiphisternum present. Clavi- 
cles slender, curved, closely applied medially 
Coracoids robust, widely separated medially. 
Bicapitatc scapula considerably shorter than 
clavicles. Suprascapula about i ossified. 

I ight prococlous non-imbricate presacral 
vertebrae. Sacral diapophyses poorly to mode- 
rately expanded. Relative widths of transverse 
processes in > sacrum > IV > II > V = 
VI = VII — Vllf. Ilia extending to anterior 
extremity of sacral diapophyses. Sacrococ- 
cygeal articulation bieondylar. Uiostylc with 
well developed crest extending about J of 
its length. 

Humerus with strongly developed antero- 
proximal crest. Phalangeal formula of hand 
2-2-3-3. No bony prepollex. Palmar sesamoids 
present. Phalangeal formula of foot 2-2-3-4-3. 
Well developed bony prehalltix. Terminal 
phalanges knobbed. 

Comparison with <vhtt sfttcies: Up&roleta 
(rachytLrma is a distinctive species, particu- 
larly in the dorsal skin texture of dense, small 
bid prominent tubercles Other species with 
extensive frontoparietal tontanellcs comparable 
to the condition in U. tunltyderttut {U. rus- 
\etli t U. tirenicofa (J. horealis, V. talpa, and 
[J. orlctuut'is) have moderate to extensive web- 
bing between the toes (except in U, wenicola) . 
Uficroii'ia utvnirola resembles U. irucJtydenna 
in lacking webbing, but the dorsal skin is 
smooth or weakly tubeiculai (tubercular in 
U. trarhydrrma), The call of V. ortontalte 








0-4 0-6 




Fi5 4. Upper: oscillographic tracings of male 
mating calls of: left, Uperoleia lithomodu, 6.4 
km N of (Catherine, NT,; right. Runhleila 
dexerticola. 40 km N of Elliot, N.T. The time 
marker in each case is 10 msec. Lower: audio- 
spectrogram (300 Hz bandpass of two consecu- 
Ijve calk ol Uperoleia trac/ivderma. 37 km N 
of Elliott, N.T. 

(which has moderate toe webbing) is not 
known, but it is probably a larger species 
(males 26-28 mm, compared with 20-22 mm 
in the small sample of U. trachydctma), No 
other northwestern Australian congener is 
known lo have a four-pulsed mating call. 

The similar-sized U, lithomoda taken north 
of the site of V. trachydermn has poorly 
developed dorsal turbercles, a shorter call, a 
very poorly exposed frontoparietal foramen, 
and larger eyes, 

Tabl£. L Physical characteristics of male mating 
Mean values are given with ranges in parentheses. 

Mating roll: Calls of two males were recorded 
37 km N of Elliot, N.T., on 16.xii.S0. An 
audiospeclrogram of a call is shown in Fig. 4 ? 
and physical characteristics of calls are listed 
in Table 1. The call is a harsh "crcak n , con- 
sisting of four short pulses produced in about 
50 msec. The pulse repetition rate is about 
79 pulses/sec and the dominant frequency 
about 3600 Hz. 

Distribution: Known only from the type 

tfsbttAh The lype locality is a Hat iloodplain 
oi adhesive yellow clay. Males were calling 
from i he base oi grass tussocks growing in 
the water The surroundings were flooded to a 
depth of 0.3 m. 

Etymology; From the Greek trachyw 'rough', 
and derma 'skin', in reference to the unusual 
-kin condition. 

Ranideihi dcsertitxda Uem & Ingram, 1977 

This species was known previously from 
localities in southwestern Queensland (I Jem & 
Ingram 1977. Tyler 1978) and northeastern 
South Australia (Brooks 1980). 

We found R, dexerticola upon the Newcastle 
Creek Iloodplain 40 km north of Klliot 
(170' J !4'S; 133°28'E) on 16.xii.80. Five 
males and one female were collected beneath 
debris at the edge of a deep dam (SAM 
R 191 18-23). Calls of two males were recorded 
at a wet bulb air temperature of 25.4"C. 

An oscillographic tracing of a call is shown 
in Fig. 4. Call structure is similar lo that 
described by Licm & Ingram (1977), but two 
clear emphasized frequency bunds are evident 

calls of Uperoleia trachyderma and 17. lithomoda. 

Species and 


No. of 





(pulses/ sec) 




Wet bult> 


U. trachviicrnhi 
37 km N of 
Elliot, N.T. 

U. lithamurfa 
6.4 km N of 
Kathertric, N T. 

If. lithomoda 
1 1.5 km Not 
lake Argylc 
Village, W.A. 






< 13,0-20.0) 





(333-456 J 


(3 150-3400) 


2 ,2-27.0 



ui aboui 3450 and 4050 Hz. The note dura- 
tion, and call duration arc shorter than re- 
corded by Liem & Ingram (1977), hut their 
recording may have been made at a lowci 
temperature: they gave no temperature data. 
Values trom our recordings arc: call duration, 
70*80 msec; first note duration. 24-32 msec; 
second note duration, 33-41 msec 

Examination of Other collections reveals 
that R deserttcola is abundant in the New- 
castle Waters area. M. Gillam found speci- 
mens (SAM RI9184) 24 km W of Newcastle 
Waters homestead <T7 U 32S, 133 , 22E) on 
1 >i.76 in hollows on the mud banks of a 
turkey nest dam. G. A. Crook and W. Zeidk-T 
collected In specimens at Lake Woods, 15 km 
WNW of Elliot o\^ 5.x .77 (SAM R19137-52). 
P. Spalding and W. Ilosmer collected the 
species al Kltioi on II .iv.GO (AMNH 67143- 
45). Newcastle Waters on I3,iv.f>0 (AMNH 
fi7 1 46-52) and Anlhonv Lagoon on 23.iv.60 
(AMNH 67153). 

Snout to vent lengths of our scries and those 
found by Crook and Ze idler arc males 14.6- 
1#.7 mm; females 14.6-17.3 mm. Liem & 
Ingram (19771 cite a range of 13.0-1K4 mm 
for their scries, hut do not indicate the sex 
of the specimens. Presumably il embraces the 
range of both sexes. 

The only congener* recorded from Northern 
Territory are R bfihlffM Martin. Tyler & 
navies (I9K01 and R, remota Tyler & Parker. 
Froin the former R. desertnofa can be dis- 
tinguished bv its smaller st/e fbii'm^w males 
15.5-23 3 mm; females 17.4-20.0 mm S-V) 
and dilTcrenl mating call. We located R, 
bJtfngua calling within 10 km of Katherine 
Gorge. Most of the Northern Territory records 
of rcmnta by Barker & Crigg ( 1 c > 7 7 > and 
Cogger (1979) are presumably based on 
hiHtiittur However reinota occurs m Queens. 
i;uut ami it plight occupy the cos-tern portion 
of Northern Territory, ft fs distinguished most: 
readily bv its long, highly pulsed call (Tvler 
ft Parker" 1974V 

(J fwralcia lithnmoda Tyler, Davic-s & 
Manin, 198 1 

t-'pt'r<>U>ia Hthomada was known previously 
Irom two disjunct populations: one JO the 
east ei ii Kimberley region, W.A., and one on 
the llood plains west of Arnhcm Land, NT 
(Tyler ft f d I9sla). We have now partially 

closed the intervening gap by obtaining speci- 
mens of V. lithotnotht near Katherine, NT. 

On I4.xii.80 we collected seven adult males 
and one adult female (SAM R20440-7) of this 
species within 5 m of the Stuart Highway 6.4 
km N of Katherine (1.4 km N of the northern 
limit of Katherine township) (I4 C 25'S; 132" 
16 "E), and recorded Ihe calls of three males. 
Snout-vent length of the males ranges 19.9- 
22.4 mm, and of the female is 22.1 mm. The 
female is gravid, containing pigmented ovarian 
eggs 1.2 mm in diameter 

In most respects the morphology of these 
individuals is similar to that of other popula- 
tions. However in life the dermal elands arc 
light golden, appearing as continuous stripes 
along the sides. The flash markings in the 
groin and behind the thighs arc scarlet. 

To the ear the mating call is a single, abrupt 
"click"; and fn fact Tyler el al (1981a) 
described Ihe call of the Kimberley popula- 
tion as consisting of a single pulse. However 
oscillographic analysis of the calls of Kathe- 
rine males showed the call to consist of a 
rapid hurst of pulses, Hence calls of the Kim- 
berley populations were rc-analysed, and also 
shown to have multiple-pulsed calls. Fig. 4 
shows an oscillographic tracing of the call 
of a Katherine male, and Table I lists the 
physical characteristics of calls of Kimberley 
and Kalhenne ma1e$- 

The call of U. Hthomoda may be described 
as a short hurst of 4-6 pulses in 9-20 msec, 
with n pulse repetition rate of 250-450 pulses/ 
sec. The dominant frequency is about 3300 

Family: Hylidae 
Cyc/tjiwut crvptoiis Tyler & Martin, 1975 

This species was described from an adult 
nude collected at Waters on 13. Mi. 1971; 
we found a further single adult male upon the 
George Redman Causewa\. 37 km N of Elliot 
on Hurii_l980. Our specimen (SAM R18973) 
was amongst a large number of C, culMpXS 
Parker active on the road surface. (We col- 
lected 41 in 15 mins; many more individuals 
were present, whilst C. rtusfralk and C. mnctt- 
toutv were present hut slightly less abundant) 

In the Kimberley of W.A. C. cryptotis 
breeds in February, and C. caJtripes is en- 
CQunttfed rarely in that period (Tyler ct «/, 
1 9fi I b. 1 982 ) . Our experiences with these 



species permit the interpretation that in the 
N.T. C. cryptotis and C, cultripes have a 
similar breeding sequence. 

The additional record extends the known 
range of C. cryptotis in the N.T. 110 km fur- 
ther south. We note that C. cryptotis is absent 
from the northern floodplain of the NT., and 
the species occupies a narrow latitudinal 
zone illustrated by Tyler et al. (1982). 


We are grateful to the Australian Research 
Grants Committee for the grant to M. J . 
Tyler funding our field study, and to the 

Northern Territory Conservation Commission 
for issuing licence SL54/80, permitting us 
lo undertake the research. Dr R. G. Zweifel 
provided access to M. J. T. to examine speci- 
mens in the collection of the American 
Museum of Natural History. 

We wish to express our gratitude to Mr G. 
Mewett, formerly of Ansett Airlines for 
assistance with transfer of freight. Mr C. 
Miller for expert laboratory assistance, and 
Mr P. Kempster for numerous photographic- 
prints. Call analyses were carried out by Mr 
P. A. Harrison, and the manuscript was typed 
by Mrs J. Russell-Price, 


Barker, J. & Grigg, G. C. (1977) "A field guide 
to Australian frogs." (Rigby; Adelaide.) 

Brooks. J. (1980) First record of the frog 
Ranidella desert kola. Liem and Ingram from 
South Australia. S. A list. Nat. 54. 45-46. 

Cogger, H. G. (1979) "Reptiles and amphibians 
of Australia." (Reed; Sydney.) 

Dams. D. D. & Gore, V. R. (1947) Clearing and 
staining skeletons of small vertebrates. 
firld'mna: Techniques (4), 1-1 ft. 

Liim, D. S. & Ingram. G. J. (1977) Two new 
species of frogs (Anura: Myobatrachidae, 
Pelodryadidaoe) from Queensland and New 
Soulh Wales. Vict* Nat. 94, 255-262. 

Martin. A. A., Tyler, M J. & D.wifiS, M. 
(1980) A new species of Ranidella (Anura: 
Leptodactylidae) from northwestern Austra- 
lia. Copeia 1980(1), 93-99. 

Trleh, L. (1979) Leptodaclylid frogs of the 
genus Tehnatobius in Feuador. with description 
of a new species. Ibid. 1979(4), 714-731. 

Tyler, M. I. (1968) Papuan hylid frogs of the 
genus ffyla, Zoal, Verhattd, Leiden (96), t- 

(1978) "Amphibia of South Australia." 
(Govt Printer: Adelaide.) 

— , Davids, M, k Martin, A. A. (1978) A 
new species of hylid frog from the Northern 
Territory. Trans. R. Soc. S. Aust. 102. 151- 

— -, & (1981a) Australian frogs of 

the leptodaclylid genus Vperoleia Gray. Aust. J. 
ZaoL SuppL Ser. (19), 1-64. 

, & (1981b) New and redis- 
covered species of frogs from the Derby-Broome 
area of Western Australia. Rec. W. Aust. Mus, 
9. 147-172. 

— -. Martin, A. A. & Davies, M. (1979) Biology 
and systematica of a new limnodynastine genus 
(Anura: Leptodactylidae) from north-western 
Australia. Aust. J. ZooL 27, 135-150. 

— , & (1982) Biology, morphology 

and distribution of the Australian fossorial 
frog Cvclorana cryptotis (Anura: Hylidae). 
Copeia 1982(3), fin press). 

— & Parker, F. (1974) New species of hylid 
and leptodactylid frogs from southern New 
Guinea. Trans* R. Soc. 5. Aust. 98. 71-77. 




by Alan N. Baker & Dennis M. Devaney 


Twenty-three ophiuroid species are discussed: nine are new to the region, and the known range of a 
further seven is extended. Ophionereis terba n. sp. and Ophiacantha shaepherdi n. sp., are described 
from Victorian and South Australian waters respectively, and a key to the southern Australian 
species of Ophiacantha is provided. Ophioceres bispinosus, Ophioconis opacum, and Ophioprium 
rosea are new combinations, and the following species are synonymised: Ophiacantha abyssicola 
otagoensis Fell with O. brachygnatha H. L. Clark, Ophiacantha truncata Koehler with Ophioprium 
rosea (Lyman), and Ophiactis laevis H. L. Clark with O. tricolor H. L. Clark. Illustrations are given 
of holotypes of Ophioceres bisponus, Ophiomusium anisacanthum, O. australe, Ophionereis lineata, 
Amphiura trisacantha, Ophriothrix (Placophiothrix) albostriata, O. Placophiothrix) lineocaerulea, 
and O. (Keystonea) hymenacantha. A lectotype is designated for Opiocantha clavigera Koehler. 



by AtAN N. Bak*r* & Dennis M. DevankvI 


HAKf *. A. N & Dgvaney, D. M. ( 1981). New records of Ophiuroidea tCucninode/muta) from 

southern Australia, including new species of Ophiucanftm and OpflioWfttS. TfWtfi # $0F- 

V. -),/•,/ 105(4). 155 178, II December, 1981. 

Twenty-three ophJuroiJ species are discussed: nine are new lo the region, and the known 

range of a hirther seven IB extended. Ofhio'tcreis ttrbn n. sp. and Ophmcu/itha shephenh 

n. sp., are described from Victorian and South Australian waters respectively, and I key lo 

the soulhern Australian species of Ophiacimthn is provided. Ophkurrcs btspittosus, Ophiocoms 

opacUm t and Ophiopriinv ro.wa uic new conihinations, and Ihe following species are 

synonymiscd: Ophhwantha 44hysxico?a oUtKoensix Fell with O. hnn.ltyxiniih<i H. I. Clark, 

Ophiucantho vimcaia Koehlcr with OphioprintH fam (Lyman), and Ophiasis lanis H. L 

(lark with O. tricolor K. L. Clark- Illustrations arc given of hoiotypes of Opluorwb' to- 

plnosus, Qphit'tnutium iinis<uunthum, O. wt\tttih\ Ophionercts iinnuu Amphium inMtattUlia, 

Opliriorhri.x \JHacophiothnx) alhostnuta, O, Witicopluothrlx) tineocttcrHlta. ami (> \hfy\iuntu) 

Itvmenacuniha, A lectotype is designated for Ophiocomhn ctoflgefv Knehlcr. 


The ophiuroid fauna of southern Australian 
waters is known fareely Irom the studies of 
II. L. Clark (1916, 1918, 1928, 1 938, 1946) 
and A. M. Clark (1966). H L Clarke early 
paper covered collections trawled from the con- 
tinental shell by F.I.S. "Endeavour" in 19 10-- 
14. and the later works dealt with specimens 
collected mainly from the Victorian coast by 
Joseph Gabriel, nrnl from Soulh Australian 
waters by Sir Joseph Vcrco and Clark himself 
during a visit in I929, ] n 1946, Clark added 
more specimens in his revision of the- entire 
Australian eehinoderm fauna. The recent 
account by A. M. Clark was based on shallow 
water collections made during the Port Phillip 
Survey, Victoria, 1957-63. The collections 
reported by these authors are in the South Aus- 
tralian Museum, National Museum of Victoria 
Australian Museum, Museum of Comparative 
Zoology, and British Museum (Natural Hfe- 
bfjiy) Seventy-four species of ophiuroids aic 
now known from southern Australia between 
Cape Howe, Victoria < 37*30. 1 S) and Cape 
Naluralisle, Western Australia (3.V30.1 S) 
(Clark 1940, Baker 1981, and this paper) 

In the course of studying Australasian 
Ophiuroidea in ihe past Tew years we have exa 

* National Museum of New Zeulund, Private Batf. 

Wellington. New Zealand. 
I Rcrniee P. Bishop Museum. P.O. Box 19000-A. 

Honolulu. Hawaii. 

mmed recent collections made by trawling ex- 
peditions (H.M.A.S. Diamantina and KimMa, 
M V Aquarius, and M.V Sarda'l and divers 
working along the coasts of Western Australia. 
South Australia, Victoria, and Tasmania. 
Among these collections are a number of spe- 
cies previously unknown in ihe region, includ- 
ing two new species and others hitheito known 
only from oilier Pacific localities, particularly 
New Zealand. Some of these have been dis 
cussed by Baker (1979. 1980). and the re- 
mainder are reported here. 

During visits to the Soulh Australian 
Museum, one of us (A.N.B. ) re-examined 
some of the ophiuroid material recorded by 
H L Clark (1928), and re -illustrated, by 
camera Uicida drawings, several holotypes of 
local species described but not figured in detail 
by that worker. Our .studies have shown that 
some taxonotnic changes are necessary. :md we 
have included such information in this paper. 

Text coiivtiUiotis 

The following abbreviations arc used; fftflff- 

unions, AM — Australian Museum, Sydney. 
BM(NH)— British Museum (Natural Histoiy), 
1 .ondon. BPBM — Be mice P. Bishop Museum, 
Honolulu. Hawaii MCZ. — Museum of Com- 
parative Zoology. Harvard MNB — MttWUffl 
fiiir Naturkunde, Berlin, NMN7- National 
Museum of New Zealand, Wellington NMV — 
National Museum of Victoria. Melbourne. 






SAM — South Australian Museum, Adelaide 
WAM -Western Australian Museum, Perth 
ZMH — Zooloyisches Museum, Hamburg. ZMC 
— Museum, Copenhagen. Morpho- 
' J d —disc diamcler. 

SysteitiHlie Account 

I MtllLYOpMu? due 

Subfamily Ophiolcpidin ic 

Qphfoceftt Koehler, 1922 

Ophiocercs fifr/rfcusus (H 1- Clark) n comb, 

FJGS 1-3, 5, 14-17 

OphiopIotHS i"vtin>Mi\ Hi ' • Oarki l£3fi 33 7 T 

pl- 4, fig. 2; 1946: 216.- A. M. Clark. I96fli 

327 Dlnnall, )*>K0: <tt 
Specimen* examined: Datatype, MCZ 4025. 
Phillip Is. WcsicrnpoiL Vic. no depth. May- 1*115; 
1, NMNZ 2074, Port Macdonnell, S.A. 3-4 m'1; 1, NMNZ 2076, West Id, S.A,. 4 m\ 

Vi.i.vny, :, nmv, U366* Cane Lipwp< vie, 

no depth, 24iil l J7 r >; 2, NMNZ 2075, Fluted 
Cape Brum Id. Tas. 13 m, I0.ii 1972: I, NMN/ 
Z7II2 Deal Id. Trs., 20 nt, 4.V.1974. 

Rr marks: 1 hese specimens range 3~*>.5 nln* 
d.d.. and have arms up to 3.3 X d.d long 
They flf* ftM* similar to the type series in 

Thai (his species belongs in Opluo -crr\ 
rather than Opfttopf&CUs is shown by the pre- 
sence ol one tentacle scale to each pore, short 
cemlal slits originating close to the oral shield 
and hardly extending pasl the first ventral arm 
plotGi and the rcfalively small amount ol frag- 
ment nion of the dorsal arm plates (<lfl 
pieces). In Qphfopfactft Lyman, there are 3"-8 
tentacle scales, genital slils which ore separated 
Irotn ihfi oral shields by small platelets or scales 
and which extend as far as the fourth segment, 
and extensive dorsal arm plate fragmentation 
(<2fi pieces). 

As pointed out by H. t.. Clark (1918), this 
spoors is extremely close to O. huttoni (Fa"»- 
qohat ) from New Zealand. Fxamination of 15 
KpCi lamens ot the latter (NMNZ) shows small 
but consistent differences which we consider 

to be specific Clark's suggestion that the num- 
ber and arrangement of dorsal arm plate frag- 
ments is dilTcjent in these two species is cor- 
ned, but only for the basal and middle part of 
the arms. To confuse matters, however, the 
basic pattern of fragmentation may be altered 
by irregular secondary division of plates on 
sonic or all arms of some- specimens The 
dorsal arm plates consist of three main ele- 
ments— a central plate bordered by two lateral 
plates. The central plate is usually divided 
transversely, and on Ihc extremities of the 
arms, the arrangement is virtually identical in 
both species (Figs 17 & 21). Nearer the disc, 
the lateral plates become separated from the 
proximal portion ot the central plate by an 
oblique plate on each side (Figs 16 & 20). At 
this stage of fragmentation, a difference be- 
tween the two species is evident — in huttoni 
the pair of additional oblique proximal centre*- 
laleral plates (pel) nearly as wide as the distal 
centro-laterals (del) and fully in contact with 
the lateral edge ol the proximal central plate 
tpc). becomes wedged between the proximal 
centro-laterals and the distal central plate; in 
fmpinosus however, the additional pel plates 
are usually less than J the width of the del 
plates and only in contact with the posterior 
lateral portion of the pc plate, On the same arm 
segment, and even more pronounced on proxi- 
mal segments (Figs 15 & 19). the latter species 
does uot develop a third pair of plates distad 
or disto-lateral to the dep as seen in huttoni 
iTigs 16 & 20. 15 & 19). Occasionally, irregu- 
tel longitudinal splits occur on the very basal 
.trni plates of O. hixpinosux (Figs 4& 14). 

The oral shields also differ in proportions — 
in btspinosus they are noticeably wider than 
long (w B 1.44- 1 63L), whereas in huttoni 
they are ahoul as wide as long (w = 1.05- 
1 13F). 

Although there is no noticeable difference 
in the arrangement of the disc scales, the radial 
shields and marginal jntcr-radia| plates are 
larger Jn biSpUlOSus than in httttoni. In fact the 
si2c of the plates rival those in O, marginara 
Fell a second New Zealand species, regarded 

Hgs 1-21, I arm base and adjoining disc, dorsal, of OphtQCtW bUpfaWUS (NMN/ 2074). 2, same, 
ventral. 3, aim bust and adjoining disc, veotral. of Ophiocercs incipient (NMNZ 2715); 4. 6th 
& 7th arm segments of O. Nspinosus (NMV H36tVi; 5, fitb arm segment of O. btspinosm (holo- 
type MC2 4025) 6-^. 2nd. «th. IKth and ditta! arm segments of O. incipient (NMNZ 2715). 
10 IV same segment* of O. marginaia (NMNZ 2705): 14-17, same segments of O. hisptnosus 
(NMN/ 2074),' IK-2?, same .-.^m-nts pj fi tilt Oh. I (NMN7: 1183). Abbreviations: pc — proximal 
central plates; del — distal cen'.ro-latcral plates; pel — proximal centro-laleral plates. Scale lines 0.5 



here as valid (see Pawson 1969). This species 
can be distinguished from bispinosus and hut- 
toni by its shorter arms (2 X d.d.), simpler pat- 
tern of arm plate fragmentation (Figs 10-13), 
shorter genital slits, and 3 instead of 2 arm 
spines on each segment. Unlike the other spe- 
cies, O. marginata is viviparous — a 9.0 mm 
d.d. specimen dissected during this study con- 
tained 18 juveniles 1-1.3 mm d.d. 

The fourth species and type species, O. inci- 
piens Koehler, also has a simple pattern of arm 
plate fragments {Figs 6-9), but has arrowhead- 
shaped oral shields (Fig. 3), 2 very short arm 
spines, and relatively long (3.3 X d.d.) arms. 

The four Ophioceres species are restricted to 
the southern hemisphere antarctic, subantarctic, 
and cool temperate waters. O. incipiens is cir- 
cumpolar in antarctic waters; marginata sub- 
antarctic New Zealand, and southern mainland 
New Zealand, huttoni northern New Zealand, 
and bispinosus southern Australia. Ophioceres 
species are recorded at depths of 0-384 m. 

Ophiomusium Lyman, 1869 

Ophiomusium anisacanthum H. L. Clark 

FIGS 22-24 
Ophiomusium anisacanthum H. L. Clark, 1928 : 
445, figs 133a, b; 1946: 247.- Baker, 1979: 31. 
Specimens examined: Holotype, SAM K254, and 2 
paratypes, K256. Spencer Gulf or Gulf St Vincent, 
S.A.; 4. WAM 896-898-77, 31°00'S, 1 14°51'E, 
W of Lancelin, W.A., 130-160 m, HMAS Dia- 
maniina stations 41, 44. 

Remarks: The Western Australian specimens 
of O. anisacanthum range 11-12 mm d.d., with 
arms 40 mm long, and are thus comparable 
with H. L. Clark's type series of five — the only 
other examples hitherto reported. Clark (1928) 
characterized this species by its single, large 
ventral inter-radial plate and one large arm 
spine in a series of three or four smaller spines. 
In the six specimens examined here, we find 
mostly four spines on the basal seven or eight 
segments. They grade in size from the lower- 
most, which is 0.5-0.75 as long as the segment 
bearing it, to the uppermost, which is 0.2 as 
long as a segment. The two lowermost spines 
are distinctly separated from the uppermost 
pair (Fig. 24). Beyond the seventh or eighth 
segment, only the lowermost pair are present. 
It is this arrangement of arm spines, plus the 
large inter-radial plate and the smooth, closely 
adpressed dorsal disc plates, that distinguish 
the species. 

The type specimens are without exact locality 
or depth, Verco's label bearing no more than 

"Spencer or St Vincent Gulfs". H. L. Clark 
(1946) expressed interest in the depths at 
which anisacanthum might eventually be found, 
for the genus is essentially a deep water one, 
and the South Australian gulfs are less than 
55 m deep. It is thus of considerable interest 
that the species has now been collected off the 
Western Australian coast in 130-160 m, show- 
ing that O. anisacanthum is indeed one of the 
few species in this genus to inhabit the con- 
tinental shelf. 

Ophiomusium australe (H. L. Clark) 
FIGS 25-27 

Ophiomusium simplex var. australe H. L. Clark 

1928: 449, fig. 135; 1946: 274-5. 
Ophiomusium aporum H. L. Clark, 1 928 : 447, 

fig. 134; 1946: 275. 
Ophiomusium australe- Baker, 1979: 30. 
Specimens examined: 9 from Vic, S.A. and W.A. 
(listed in Baker 1979. p. 30), 3, NMV H367 
38°46'S, 141°33'E, 155 m, 26.viii.1975. 

Remarks: This southern Australian species has 
been discussed by Baker (1979). The oppor- 
tunity is taken here to give a detailed illustra- 
tion of the holotype (SAM K256). 

Family Ophionereididae 

Ophionereis Liitken, 1 859 

Ophionereis terba n.sp. 

FIG. 30 

Specimens examined: Holotype NMV H363 & 
paratype, NMNZ 3673, 43 km SSW Portland, 
Vic., 585 m, coll. R. Plant & M. Gomon. 14.v. 
1979; paratypes 4, NMV, W of Cape Nelson, 
Vic, 164-201 m, June 1969; paratypes 2, 
NMNZ 1881, S of Warrnambool, Vic, 220-310 
m, 14.V. 1969. 

Description of holotype: Disc diameter 6.0 mm. 
arms broken but c.9 X d.d. Disc entirely 
covered with coarse imbricating scales, 3-4 in 
1 mm radially; primaries prominent, and 1 
central marginal inter-radial scale with 4 larger 
scales each side. Radial shields 0.9 mm long, 
0.4 mm wide, divergent. Genital papillae small, 
granular, visible near arm bases dorsally. 

Oral shields longer than wide, diamond- 
shaped with incurved disto-lateral margins. 
Adorals meeting within, attenuated distally. 
Distal oral papilla large, triangular, remaining 
3 narrow, blunt. 

Dorsal arm plates widest proximally, with 
evenly curved distal and proximal margins; 
supplementary plates very small and present 
throughout arm. Ventral arm plates widest dis- 
tally, as long as wide, concave laterally, with 



Figs 22-27. 22, arm base and adjoining disc, dorsal, of Ophiomusium anisacanthum (holotype SAM 
K254); 23, same, ventral, 24, 1st & 2nd lateral arm plates of SAM K254; 25, arm base and 
adjoining disc, dorsal, of Ophiomusium australe (holotype SAM K256); 26, same, ventral; 27, 
1st & 2nd lateral arm plates of SAM K256. Scale line 0.5 mm. 


A. N. BAKER & D. M i.ivANt-Y 

P{gs 28-36, arm bases ;»nd adjoining liiSca 2.x & 2^, (>i>hin m >rnx lineata (hoioivpe, MCZ W>\ d»f- 
-siil; Ac ventral; 30. t'p/i/ (J -!rm( /,y/>,/ n, sp 0'mA»Ivpc NWV H?M>. dorsal'. *1 A 32. Ophioionis 
upaatm iNMNZ ww) fowl A IftttUAf; 33 A u th>hit< t ri; trtctttw CtatQiype SAM KZH) ven- 
rr.l ,V dorsal; 35 & 36, Ophiocvmina WUftNltth (fctPBM W1H221 dorsal & ventral Scale tmsa. I \) 



a straight distal margin curved only at the 
corners. Tentacle scales large, oval. Three 
slender, evenly tapered arm spines, middle 
spine longest, up to 2 X length of adjacent 
ventral arm plate. 

Colour pattern (dry): disc mottled brown 
and cream, brown pigment around edges of 
disc scales and radial shields. Dorsal arm plates 
with m-shaped brown band on distal margin 
(Fig. 30); ventral surface uniformly cream. 

Remarks: This species is closely related to O. 
lineata H. L. Clark from north Queensland 
waters. It differs from it (Figs 28-29) in 
having relatively longer and more slender arm 
spines — in lineata they are only slightly longer 
than an adjacent ventral armplate, and arc 
thick for their length. Also, the oral shields of 
O. terba are symmetrically diamond-shaped 
rather than blunt, spearhead-shaped as in O. 
lineata. The colour patterns of these two spe- 
cies provide the most readily observable dis- 
tinguishing features: O. terba has very pro- 
minent m-shaped brown bands across the distal 
part of the dorsal arm plates, and has a uni- 
formly cream ventral arm surface, whereas O. 
lineata has a single longitudinal dark line on 
the arms dorsally, and random dark spots on 
both arm surfaces (Figs 28-29 of the holotype, 

No depth was given for the collection site of 
O. lineata by Clark (1946), but the waters sur- 
rounding Lindeman Island are 37-55 m. The 
bathymetric range of the new species is, in 
contrast, 164-585 m. 

Apart from its striking colour pattern, the 
new species is characterized within Ophionereis 
by the coarse disc scaling the very small sup- 
plementary dorsal arm plates. The trivial name 
terba is an Australian aboriginal word meaning 
pretty, and alludes to the colour pattern. 

Ophiochiton Lyman, 1879 
Ophiochiton lentus Lyman 

Ophiochiton lentus Lyman, 1879: 55. pi. XIV, 
figs 398-400. Baker, 1979: 34. 

Specimens examined: 3, NMV H368, 38"3.4'S T 
I49°23.7'E, E of Flinders Td, Bass Strait, 183 m, 
22. xi. 1973; 1, NMV H360, 39° 32.5'S, 148° 
5L5'E, 274m, 24.xi.1973. 

Remarks: The specimens are 5.0-6.5 mm d.d. 
and differ slightly from the 13 mm holotype: 
the primary disc scales are prominent, there are 
no papillae along the edges of the genital slits, 
and there are two tentacles scales only on the 

first or second pore on each arm. These dif- 
ferences may be attributed to the juvenile 
nature of the specimens, for otherwise they 
agree closely with Lyman's description. 

Ophiochiton lentus is the first representative 
of the genus to be found in Australian waters; 
the species is otherwise known from the type 
locality near the Kermadec Islands (also re- 
ported from there by Baker 1979) and the 
Chatham Rise, east of New Zealand in depths 
of 300-900 m. 

Family Ophiocomidae 

Ophiocomina Koehler, 1922 

Ophiocomina ausiralis H. L. Clark 

FIGS 35, 36, 63-68 

Ophiocomina austraiis H. L. Clark, 1928: fig. 
124; 1946: 188; A. M. Clark, 1966: 327. 

Specimens examined: Holotype, SAM K211, 
outer Gulf St Vincent S.A., no depth: 4 BPBM 
W1822, 2 km W of Outer Harbour, Gulf St 
Vincent S.A., among Posidonia roots, 12 m depth, 
3.iii. 1965. 

Remarks: The additional specimens from Gulf 
St Vincent are the first to be recorded since 
H. L. Clark's original description. They range 
5-8 mm d.d. This rare species has a very 
restricted distribution in the South Australian 
gulfs — extensive collecting along other parts 
of the southern coast by S. A. Shepherd and 
Mrs J. E. Watson have failed to locate other 
examples. The holotype is in poor condition, 
and because the species has not been ade- 
quately illustrated, we give here figures of an 
8 mm d.d. BPBM specimen (Figs 35-36), and 
SEM photographs of the oral plates, dental 
plate, vertebra, and arm spine (Figs 63-68). 

Wilkie (1980) proposed that Ophiocomina, 
with the species austraiis and nigra, be placed 
in the family Ophiacanthidae. It is our opinion, 
that there are better grounds for continuing to 
retain this genus in the Ophiocomidae. Among 
these grounds are the features of Ophiopteris, 
Clarkcoma. and other ophiocomids which are 
also found in Ophiocomina (Table 1). Further- 
more, differences rather than similarities be- 
tween Ophiocomina and Ophiolimna 5pp. in 
terms of the length : height ratio of the oral 
plates, shape of the lateral arm plates and, at 
least in one species of Ophiolimna examined 
(O. cf. perfida), the presence of an oral calci- 
fied ridge to enclose the radial water canal in 
the 2nd rather than the 1st arm vertebra, are 
considered features that do not indicate family 
resemblance between the two genera. 



Tabu- 1. Comparison o\ Ophiocomina with Ophiacanthids and Ophiocomids. 


Hylinated tips 


Hollow I lumen more than half 
wall diameter) 


A. Foramina 

B. Oral end widest 

C. Papillae present 

Ophio- Ophio- Ophia- Other ophio 

comina plerifi Clarkcoma canthids com ids 


A. With radiating indentations on abrudial 
muscle scar 

ft. Abradial muscle scar with flaring 
aboral wing 

C. Adradial muscle sear less than (— ). 
equal to (±), more than t + ) 
half height of plate 1 " 

D. Longer than high ( + ), equal (±L 
higher than long ( — ) 

E. Adradial articular area with upper 
distal edge rounded ( + ) or straight (— ) 


Two condyles and one pit + + 










( + ) 



( + >- 




length == proximo distal axis '( ) for Ophiocotna »(-f) for O, pusiiiu »(-j-J for ophiocoma 
height -= oral-aboral axis vusUla, Ophiop- and Ophiomas- longispina, O. 

sUa, OpnltttQ* ;/v omnia, pusilln and 

'"<'"" Ophiomustix' 


Family Ophiodermatidac 

Oenus Ophioconis LUtken, 1869 

Ophioconis opacum (H. L. Clark) n.eomb. 

FJGS 3K 32 

Ophiurodon opacum H. L. Clark, 1 9>8: 440. 
tigs 132a & b, 1946: 255; A M_ Clark. 1966: 


Specimens examined: Holotypc, SAM K243, "St 
Vincent or Spencer Gulfs", S.A. no depth; L 
NIVfNZ 2122, Upper Spencer Gulf. S.A.. 1 1 m. 
4.ix.l973; 1, NMNZ 2690, Upper Spencer Golf. 
S.A. 18 m, 1l.ix.l973; I, NMNZ 2123. off 
Glenelg, S.A., artificial reef site, 3-4 m, 1472; 
I, NMNZ 2722, W of Outer Harbour Gulf St 
Vincent, S.A-. 5 m. 3.1. 1%5; 2, BPBM W2220 & 
WAM 31-74, Cockhurn Sd, W,A. under stones, 
2 m, I3.lj.t972; 2, WAM 1133-74, 3.2 km 
NW Busselton Jetty, W.A., Pt>,\idoruu. Cyoto- 
dovea & sand, 22-24 m, no date. 

Remarks: H. L. Clark (1946) refctrcd three 
Australian species to Ophiurodon. O. opacum 
was separated from its northern conveners 
(cincfa and permhni) by having blunter, tint 
opaque upper (at least) arm spines. However, 

A. M. Clark (1965) found no generic distinc- 
tion after comparing the type species of Ophio- 
corns (O, forhesi) with pennixta and cincta. 
She did not consider O. opacum at that time 
and, in a 1966 key retained without comment 
opacum in Ophiurodon. 

Comparison of specimens of the three known 
Australian species indicate they are indeed con- 
generic and should all be retained in Ophio- 
ci'tu's. They share the following characters: 
longitudinally striated arm spines; more than 
two tentacle scales on proximal segments; 
broadened hyalinated usually serrated distal 
ends of teeth; adoral shields nearly or actually 
meeting in front of oral shield; disc granulae 
covering all or most of the oral shields as well 
as exposed part of oral plates. 

The six specimens found since 1965 are the 
lust recorded since the type description. They 
range 3.0-6.5 mm d.d., and their characters 
confirm the distinctness of this southern species 
of Ophioronis. It may be separated from the 
two northern Australian species by its flat, 
almost spalulate arm spines. Contrary to the 



type description, O. opm um docs have finely 
striated lateral arm plates, but apart from (hat 
feature, the holotype and new material agrfec 
with H, L Clark's description. bVeause K L 
Ctark\ illustration tacks detail, wc give here n 
camera lucidu drawing flf NMNZ 2690 (Pig* 
31, 12) the holotype being not In a SttittbM 
condition for illustration. 

The records of 9/ttFFfttf ftofll Western 
Au-,U;thu extend the known range of the spe- 
cies westward from the South Australian gulfs: 
this extension and its -.hallow water* habirai 
(0-24 m) indicate Ihnl it ;s probably inure 
widespread on (he southern coasts of Australia. 

Vaihyprctitwru IL ft. Clark. 1909 

Bar hy pet ntmnt hems I Lyman I 

Peetinura furax Lyman. 1 8 7 i> : 48 pi 14. ftjjs 

Huih\pi'<um<ra hrrus: ft. I , Clark, l^OJ 12^ 30. 

Bilker. 1079; Mi 
SpettfiteM 1'AoifinuJ: I, SAM TK 3132, 37 58 - 
.17 47'S, IV.) :t9'-13y : 4.sT_ SW ot tape Button. 
S.A. 54* m. MKVS lifch 2 NMV. 4? km SSW 
of Portland, Vic, 585 1T >, 14.V.I97Q. 
Reitutrkx: This large species (TK 3132 is 62 
mom d.d.) has previously heen recorded from 
Off BalcmatVs Bay, New South Wales and 
south nl Norfolk Hand (Baker |87?) in the 
AasiKitian region. Il has a worldwide distribu- 
tion in 240-2960 m. 

I win v Ophiaeanllmlac 

(JphitiLtinthu Midler & Tioschet. 1 S42 

Ophiacantlui shepberdi n.sp. 

FIGS 37. 38. 69-74 

Spretmenx examined: Holoivpc. SAM K175T! & 

paf*Uy|iC 6P8M VVIH3L Scsl Ruck, Encounter 

Bay, West Id, S.A., 24 m, on al^ae. colt. S.A. 

siilmIiciu. p*T&typ« I, SAM £1749 

« 1 NMNZ 3083, Dcva Inlet. Wc*i III SA, IS 
in. COIL S.A. Shepherd.^67. PauivpL, 
NMN7, 1 1 km art Gknefe. Gulf si vine ni S \ u 
2i m. coll. S.A. Shepherd. 27.Mi.1966. Puratypev 
K, NMN/ 1WI5, 2 BPBM WlBSS, Lands End. 
WcM Id. S.A.. 17-1 « m, coll S.A Shepherd, 
f' t .7 Purnivpes. 2. BPBM 1839, Seal RoeL Wesi 
Id, SA., 24 m. coll. S.A. Shepherd, 5.ii.l'"*7. 
Parut-. f M-H, :, WAM |s*-X1. Omtipit* Pi, VVeM 1,1 
S.A., 15 m, coll. S.A. Shepherd, no date: Puiu- 
typu-s. t. AM, I, BM(NH), 4, NMNZ 3641, 
Middle Pt, C: M u- Nnrthumbt-rliinil. S.A. 13 m, 
in al(?ac. coll. S.A. Shepherd. 19.iii.l974. 

lleseripiion ol lit >}* 'type; Disc diameter 2.5 
mm: arm length 11-12 mm, arms taper grl 
dually. Upper side ol" disc with very low 
conical granules (rarely as high as hroadL 

slightly larger and more prominent inter- 

radially than radially- eranules generally Jtoi 
contiguous, underlying scales evident between 
thein. Radial shields sock-shaped, distal en. I. 
exposed i\ni\ adjacent to Literal edge of first 
exposed dorsal arm plate, also m contact with 
genital plate helow. On lower sal., .a disc. uu 
miles more widely spaced proximalty; disc 
scales evident 

Oral shields approximately 15 ■ hrouder 
than tone, somewhat triangular, with small lobe 
in distal edge, madrcpontic plate much laiger, 
with greater distal prominaiice. Adoral shields 
torfee; m?ctftlg broadly within, encroaching 
distad around most ol ventral shield. 

Three oral papillae on edge of oral plate: 
outer two longer than broad, simibt itl size to 
inner papilla which is more sharply tapered 
and apically directed, The first tooth level with 
oral papillae hul laiger .tnd jnow-head shaped, 
other teeth not visible. 

FJTSI ventral arm pl&tca sliehily broader lhan 
lollowing plales which are tan-shaped, with the 
distal edge convex CFig. 6 ( >); plates separated 
From one another by lateral arm plates, ihe 
separation increasing distally. 

One tcnlacie scale, less than ] a segment 
long, rectangular with rounded up (Fie 70):. 
scales reivunnui'J appro,\nnately same si-'l and 
shape on disltll segments in contrast to dimi 
mshing IjtZe of other arm plates, 

Dorsal aim plates broadly Liu shaped, 1.5 * 
or more broader than long* with exception 
ot ihc first csposcd plate which is in contact 
with second plate, all others separated from 
ttcf) other by lateral plates. Micro- structure 
ot these plates L .. etfc mesh of pores and 
Uabelculae, with smooth raised lumps at inter- 
sections of meshes ('Figs 71, 72). 

Lalcral arm plates meeting broadly above 
>n-\d below ; distal ^c Maircd considerably 
where arm spines attached. Iwo short and 3 
somewhat longer bluntly rounded spines on 
segments one and two respectively; segment 
three (fust free segment) with 4 arm spines, 
the lower three bcine somewhat longer than 
those of \ejjrrient two. minutelv spiculate and 
gently mpering, while upper spine on each side 
is greatly enlarged. 2 •' length of adjacent 
lower spine (750 |ty vs 350 m,» ) and ne.nlv 
iwice .is thick, mu roscopieallv tugose, scarcely 
lapcrinut, and broadly rounded at lip (Fig. 73). 
One of 10 sides with only 3 arm spines, the 
upper enlarged spine being absent. 

Three spines (rarely 4) conrinuc on distal 
segmeius, ihc spines remaining short, blunt, in- 





Pta 49-4S W-54i Ofthif'ihrtv tfiQCQphtOthrtxj tithofttlata (hololype, SAM K215i: 49. rndidl disc 
scymcm. dorsal 5(5, disc Slump; 51. lepicseriitiv;: arm opines, 52, Mh & Mh dorsal arm plates; 
J% 2Uth dorsal arm plate with pigment; M. ffth & 10th ventral arm platen. 55 59. OpftiOlhriX 
(PUtcvphlOtftrlr) MftVflKWtfJta 'hololype, SAM K21K): 55, disc stump; 16, radial disc segment, 
dur-.;>i, M Hfi S Ctrl dorsal arm plates with pigment; 58, 9th & I0!h arm plates with 
pigment; 59, uppermost basal arm spine ftfl ul. QphiOlhrU fJteratftfrwaJ hymemivaruha (holo- 
lype SAM K2I7). 60i ftTOl tow and adjoining disc, dorsal; 61. 5ih * 6th dorsal arm piales; 62, 
9th & M'lh ventral arm plMBfr. St&fc lillCS 1,0 mm. 

creasimrly spieulatc. and near end becoming 
ovulate along one margin (Fig- 74). 
PartOtiOfl, A parutype (HHBM W183I) from 
the same station as the holorype is the largest 
specimen (d.d. ot 2 7 mm). The arms uic con- 
torted and twisted upward making the upper 
side ddheult tO see. However, the lower (oral) 
surface, especially the mouth region, is fully 
CXpOgftl In contrast to the hololype, this speci 
men has the adoral shields separated In front 
of the oral shields., probably due to the flexure 
ni the upwardly directed arms. The teeth We 
visible, there being 2 or 3 below the apical (me, 
somewhat blunl to slightly tapered or. in two 
E3ISCS, the upper hcing elongate and pointed. 
At the inner radial edge Oi each side Of the 
ventral arm shield there is a small papilla Which 
is separate irom either the adoral shield or jaw 
rkite. None &J the other specimens lias this 

feature. An arm spine sequence of 2-3-4-3-3 
U typical o^ the first five proximal segments. 
Four arm spines occur very infrequently iseg 
6, on one side of one arm and seg. 7, on both 
sides of one arm, one side on another) but in 
no case arc these enlarged like those on seg- 
ment 3. I he smallest specimen (d.d. 1.5 mm) 
shows only ;i few sides of segment 3 With the 
lourth spine developed and only on two .sides 
are these enlarged The largest specimen has a 
few enlarged granules about as high as broad 
located in the utter-radial oral region. It is quite 
evident that a distal lobe of each adoral shield 
separates the oral shield from the first lateral 
arm plates in this specimen. In the holotypc 
this lobe is not as well developed and ihc oral 
shields touch the first laterals. 
Remarks-; Ophiacantha shephenli differs from 
congeners in having the combination of low 

f-ips 17—48, 37, mm base :<n<l incoming disc, dorvO oi Ophiacantha shephtrdi n sp. (hololype. 

"SAM K1750 1, &8i nunc, ventral. 39 42, baStt and adjoining niRCf, donut of: J ft , Ophia- 

mmlht ketmryfu fNMNZ 2071); 40. '■'phUuumtha brachyKnatha (NMNZ 2070); 4i T Ophtacanthu 
ilaVi^ru INMNZ 2072) 42, Offhttidinma tiitf+tWta (NMN7 20K7): 43. arm base and adjoining 

djst, dorsal, oj AmpMuna trtitacantha iNMN/ 2073); 44. same, ventral 45j iadiai disc segment, 
dorsal, or 4mphlura frfttfi anthd (hololype, SAM K2I2I: 46, same, ventral: 47 & 4K. h;*sal, & Vth 

<fc 10th arm scymrnts of A, Inwcunifut, ViMy)..- Sci.le line 0.5 mm. 



Figs 63-68. Ophiocomina australis (BPBM W1822): 63, oral plate, adradial view, x81; 64, oral 
plate, abradial view, x72; 65, 1st arm vertebra, x!87; 66, dental plate, xll7; 67, arm spine, x89; 
68, edge of arm spine, x250. 



blunt disc granules and only one pair of 
smooth, elongate and enlarged upper arm 
spines on the segment next to the disc. The 
granules on O. shepherdi are rarely as high as 
broad (spinules are defined as disc elements 
2 X as high as broad, with granules up to 2 X 
or less as high as broad, with blunt tips). 
Whereas O. gracilis (Studer) shows a similar 
pronounced elongation of one pair of basal 
upper arm spines, these and other arm spines 
differ from those of O. shepherdi by being 
acutely tapered to sharp points, and the disc 
covered by delicate rnultifid pointed disc ele- 
ments. O. ahernata A. M. Clark, which has 
been collected with O. shepherdi, differs by 
having short conical granules mixed with 
longer, thicker blunt spinules on the disc, an 
alternating sequence of arm spines, and seg- 
ment 1 with three, or sometimes four, arm 
spines (vs. two, rarely three, on this segment 
in O. shepherdi). O. heterotyta H. L. Clark, in 
contrast to O. shepherdi, carries five rather 
than four spines on the basal free segment and 
the upper arm spine of the first free basal seg- 
ment is much less than twice that of the pre- 
ceding spine in the row, and all the spines taper 
evenly towards the tip. 

O. shepherdi is further characterized by 
having microscopically smooth dorsal arm 
plate trabeculae on fan-shaped plates, oral 
shields which are separated from the first 
lateral arm plates by a distal lobe of the adoral 
shields (at least in the largest specimens), and 
short (except for one pair of upper), few (not 
more than four) smooth arm spines, the rows 
not approximate dorsally. 

This species is named for its collector, Mr 
S. A. Shepherd. 

Ophiacantha heterotyla H. L. Clark 
FIGS 39, 75-78 

Ophiacantha heterotyla H. L. Clark, 1909: 542, 
pi. 52, figs 4-6; 1918: 86; 1938: 209; 1946: 
Specimen examined: 1, NMNZ 2071, off Simp- 
son's Bay, Bruny Id, Tas., 11 m, 18.ix.1972. 
Remarks: About average size for this species 
(3.5 d.d., 13.0 mm arm length). Stout, rather 
club-shaped disc spinelets characteristic of O. 
heterotyla reach 0.5 mm long and are promi- 
nent radially between the radial shields and the 
disc centre. Elsewhere, the disc is covered by 
small, round, glassy plates bearing short, blunt 
stumps. There are 5 or 6 arm spines on the 
first two or three free arm segments, then 4 
spines over the remainder of the arm. The 

spines are stout, cylindrical, and the uppermost 
are initially the longest (Fig. 39), those on the 
first two segments being up to 1\ arm segments 
long. After the 8th or 10th arm segment the 
lowermost spine is noticeably shorter than the 
other three. The micro-structure of the arm 
spines consists of a longitudinal series of 
granule-bearing, divaricating ridges separated 
by grooves and hollows (Fig. 75). The dorsal 
and ventral arm plates are roughly triangular 
and have a micro-structure similar to, but more 
random than, that of the arm spines (Figs 76- 
78). The tiny granules are not present on the 
ventral plates (Fig. 77). 

Ophiacantha heterotyla is closely related to 
O. alternata and O. shepherdi. From the 
former it is distinguished by having a non- 
alternating number of arm spines on successive 
arm segments, and taller disc elements. From 
O. shepherdi, it differs in the greater number 
and size of basal arm spines, and the tall 
spinelets rather than low granules on the disc. 
The microstructure of the arm plates is closer 
to O. alternata than to the other South Austra- 
lian congeners (Figs 78, 89). 

O. heterotyla apparently has a restricted dis- 
tribution from Sydney to Bruny Id, Tasmania 
in 9-1 46 m. 

Ophiacantha alternata A. M. Clark 
FIGS 42, 84-82 

Ophiacantha alternata A. M. Clark, 1966: 328- 

330, Figs 4a-c. 

Ophiacantha clavigera Koehler: H. L. Clark, 

1938: 208 (Koombana Bay, Bunbury, Western 

Australia — part). 

Specimens examined: Paratypes, 3, NMV H 1 6, 
Port Phillip Bay, Vic, 5.5-12 m 21.V.1961; 1, 
WAM 634-78, Arrawarra, N of Coffs Hbr, 
N.S.W., tide pool, 20.iii.1978; 1. NMNZ 2114, 
Port Hacking, N.S.W., 7 m, 30.V.I976; 6, NMNZ 
2379, Erith Id, Kent Group, Bass Strait, 50 m, 
6.V.1974; 1, NMNZ 2116, Westernport, Vic., 3 m, 
2.V.1976; 2, NMNZ 2099, off Pt Marsden, S.A., 
20 m, Jan., 1965; 12, NMNZ 2093-96, West Id, 
S.A. 2-27 m, 19.iii.1966-18.ix.1967; 1, BPBM 
W1810, 8 km N of Western River, Investigator 
Strait, S.A., 45 m, 12.U965; 4, NMNZ 2098. 
11 km off Glenelg, S.A., 25 m, 27.xii.1966; 2, 
NMNZ 2100, Eagle Bay, Cape Naturaliste, W.A., 
1-5 m, 24.ii.1975; 2, WAM 259-71, BPBM 
W2208, Cockburn Sound. W.A., no depth. 28.v. 
1958; 2, MCZ 4900, Koombana Bay, Bunbury 
W.A., no depth, 26.x, 1929 (det. O. clavigera by 
H. L. Clark, 1938). 

Remarks: Hitherto this species was known only 
from Port Phillip Bay, Victoria; the new 



Figs 69-78. 69-74, Ophiacantha shepherdi (paratype NMNZ 3085): 69, 2nd ventral arm plate, 
xl28; 70, tentacle scale xl45; 71, 2nd dorsal arm plate, xl08; 72, surface detail of 71, x435; 
73, uppermost basal arm spine, xl45; 74, distal arm spine, x217. 75-78, Ophiacantha heterotyla 
(NMNZ 2071); 75, uppermost basal arm spine, x72; 76, 2nd dorsal arm plate x72; 77, 2nd 
ventral arm plate, x77; 78, surface detail of 76, x580. 



Figs 79-87. 79-83, Ophiacantha clavigera (NMNZ 2015): 79, 2nd ventral arm plate, xl38; 80, 
2nd dorsal arm plate, xl36; 81, surface detail of 80, x 544; 82, uppermost basal arm spine, x90; 
83, middle basal arm spine, x86. 84-87, Ophiacantha alternata (NMNZ 2097 & 2099): 84, upper- 
most basal arm spine of 2097, x80; 85, uppermost basal arm spine of 2099, x74; 86, surface detail 
of 85, xl50; 87, surface detail of 84, xl50. 



material show* a wide southern Australian dis- 
tribution from ColTs Harbour. N.S.W, (30" 

I8*S) to Cocfcburn sound, w.a [Ji'STS) in 

I-4 1 ) m. This distribution pattern is shored by 
a number ol endemic shallow water ophiuroids 
(Co>iOctodUs (<i4Xi'ali\, Clorkcoma c(Jtuiliculata> 
Ophiupt'iu av:ittv'ti.\ . rumm yt, 
( if/noi/itu. iPiiKophioth'iy) sponsiuulu), ami 
supports the concept of i single southern faunal 
region in Australian coastal waters. 

OphlaxXQtfha ultimata varies remarkably m 
llit shape of its arm spines. The macroscopic 
appearance of the spines ranges from sm. 
to broadly serrate 0Q C&te side (Figs 84, $5), 
I Ik- uppermost basal spines are usually more or 
less straight when smooth, although the tips 
may be Slightly clavate or even bifurcate m rare 
; .iM*v I he laterally serrate spines arc, however, 
nivu/jubty curved <n g $»bre4tke fusbion (Fig 

K5» Microscopically, the -pines are sculptured 

in two way* — the basal portions being an ana- 
stomosing system of smooth longitudinal ridges 
and pores (Tig. 86), whereas the distal por- 
tions :i re linelv rugose, the ridge* havmu hfr 
come finely but irregularly covered with EQJSGd 
lumps hearing small thorns £ Fie.. 87). When 
present the broadly serrate spines occur on aJl 
arms; and rhe adjacent smaller sp$0CS WC simi- 
larly serrate, but bear teeth on both edges 
lather than one (Fig. VI ) Were it net for the 
absence of another constantly different chatae 
ur .in. I the fact that individuals with smooth 
ftplncft occur m samples with those hearing 
SBlTate spines, these two tortus could be mis- 
taken for separate species. In other rfe^pi 
specimens are alike. "The ruicroslnierure pf the 
dorsal arm plates is similar to O hefr-totvtt:, 
with JSoc rUSOSUies scattered over enlarged sec- 
trotlS •>! ihe irabeculae (Figs 88, 90), but Hurt 
species can he distinguished from O. tiUermtm 
because or Ihe laser's alternating sequence 01 
ann spines, a typical sequence being 3-3-7-8- 
5-7-5-6-4-6 on ihe proximal segments |NMV 

I Ik rjdiil shields of O altenmtn are. like 
those of lu ie/i.'tyla unci \hrphenli exposed only 
at men distal lips The shape of these shields 
is unusual — 'hey are "sock-like" I.Fijx- 921 
rather than the long narrow and ' bai Kko" 
shields apparently typical of Ophrotantha 

QpilUtiCttfUhu cluityera Kochlei 
1 JGS 41, 1*M3 
Ophracantha cferiprra Koehler, 1907: 247, \w* 
i-V- H 1, Chirk L&gj tM (m part), i Mrv 
184 (in prut).- A. M. Clark, LBSfi: 330 

Specimens warning- 3, NMNZ 2105. 2089, 2375, 
Seal Rock, West M. SA.. 16-27 m. Mar. & Aug., 
1966, Feb.. 1967; ] NMNZ 2374. II km on* 
GltJiefg, S.A.. 25 m, 27.xii.1966; I. NMNZ 2UF, 
off Wnldengravc id, Greta Australian Biajht, 23 m, 
ltwL97i| I, ZMH E2CM6 (herein iBti&Mi -■' 
lectotype), Stn 51. Cockburrj Sd. S Channel, rocky 
bottom, 6,5-8 m Kt.ix.1905; 3, ZMH E6594, 
' MNfl H5ffl). 1. WAM 4416 (herein designated 
paraleetotypes). Stn 56, Kooinbana Bay, u,6 Off 
11-2 km SW of Bunbary, W.A., rocky bottom 
v\ilh spjrse pl;»nt growth. 14.S-18 m dcplh, 
\90S\ 1, ZMH E6123, Stn 50, Coekburn Sd, 
Suinnern Flats seaweed. 3-4 rn. 30.i\.IVfi5; 5, 
MC/ 4900, Koombana Bav, W.A. no depuY 
26.x |y?9; I. WAM 17-81, it km SW Bunbury. 
W.A. 20 in depth. H.iv.1963; $ NMNZ 3J06. 
OffC&tfi&C lu, W.A., 7.5 nt. 2l.iiU972; I, WAM 
517-7?. 27 km W Ot Cupc Peron, W A.. 35 rn I, WAM lS-81. ta.18 km N of Don- 
gars*. W.A, IS 3 m, I6ii.l976; I, Mt V 4901, 
Broome. W.A., flredgeA no depth. June 192ft. 
Remarks: The specimens range t 5-3.5 mm 
d.d. ( with arms 4-5 X d.d. long, and agree well 
With Koehlers description and rather diagram- 
matic figures. The South Australian records are 
an cvrensioM rjf the species' range eastward 
from Western Australia Xvhere it has been 
reported from Coekburn Sound, Koombana 
tl.v. and Broome (Kochler 1907, H. L. Clark 

New morphological information no Ibis spe 
cies is now avLiikible from ShPvt examination of 
■ urn plates and spines (hies 79-83) The sur- 
face of Ihe ventral arm platen consists of a 
series of bro;id. smooth, transverse "shelves**. 
beneath the edges of which are minute pore« 
(I u- 7'M, The dorsal arm plates have similar 
srieh'es. but they are somewhat crowded ami 
irregular, and <rvir edges bear minute thorns 
(Ties SO, 81). The longest upper arm spines 
art cylindrical mid RITIOPth husally, with two 
longilurJrpal rows of pores runnine_ towards a 
slightly swi'llcii and thorny lip (Fig. 82 t The 
lower arm spines are flattish, with a single row 
oi' poa-.. and strong teeth on their ed^cs i Fie. 
B8'). These fcaures, plus a non-alternatmv 
sequence of arm spines, and forked, thorny disc 
elements, characterize O. rlavtgCtfD. 

Eiehi Of 'he I I specimens reported a-; Ophite- 
i-ntiha c'aiiyrru hy H. L. Clark (l$j&) from 
JCoombana Bay, Bunbury, and one from 
Broome were lent by MCZ Three o\' the 
Roombanj Bay specimens with ^ d I 7 to 3 $ 
mm ci a be assigned to O. alternate! A. M. They .ire easilv distinguished from the 
oiher five by having a well defined alternating 
sequence of arm spiues and conical, tuberculate 



disc elements. Examination of the type speci- 
mens of O. clavigera in ZMH and MNB reveals 
the very characteristic thorny forked disc 
elements as originally shown by Koehler, 
noticeably white curved radial shields, and non- 
alternating sequence of arm spines. These 
characters are evident on the five other MCZ 
specimens reported by H. L. Clark as O. clavi- 
gera from Koombana Bay. 

H. L. Clark's confusion between O. clavigera 
and O. alter nata led to an error in his 1946 
key to Australian Ophiacantha species. He con- 
sidered O. clavigera among those species 
having the disc covered by coarse granules and 
short blunt spinules, and only the first free seg- 
ment with long upper arm spines. O. clavigera, 
as Koehler figured clearly, has delicate, slender, 
forked disc spinules and elongate, often clavi- 
form (towards the tip), upper arm spines on 
at least four segments beyond the disc (Figs 

In addition to failing to note the presence of 
a regularly alternating arm spine sequence for 
several of the specimens from Koombana Bay, 
H. L. Clark (1938) considered that O. clavi- 
gera exhibited "growth changes" with respect 
to the nature of the disc elements, remarking 
that "the disc spinelets tipped with 2 or 3 
glassy teeth gradually lose those tips and be- 
come changed into little sugar-loaf shaped 
tubercles. " In fact, however, one of these speci- 
mens with a d.d. of 1.7 mm, having conical 
tubercles and an alternating arm spine se- 
quence, is smaller than several with forked 
thorny disc elements and non-alternating arm 
spines, and is unquestionably 0. alternata. 
On no individual of O. clavigera examined 
does a mixture of the two kinds of disc 
elements occur. 

Ripe orange gonads were observed protrud- 
ing from a rupture below several radial shields 
in a 2.0 mm specimen (MCX 4900). 

This species occurs mostly on algae in rocky 
areas, from the shallow sub-littoral to a known 
maximum depth of 35 m. 

Lectotype designation. Koehler (1907, p. 
247) reported "deux petits echantillons" from 
Cockburn Sound (Stn 51) and "cinq petits 
echantillons" from Koombana Bay (Stn 56). 
ZMH has 3 specimens labelled "Paratypes", 
MNB has 2 labelled "Types", and WAM has 
one labelled "Holotype", all from Stn 56. ZMH 
also has 1 specimen labelled "Holotype" from 
Stn 51 and 1 from Stn 50 labelled as "Cotype". 
Although Koehler did not select a holotype in 

his original description of Ophiacantha clavi- 
gera, he mentioned the largest example from 
Cockburn Sound was 2.5 mm d.d. and, judging 
from the scale given, his Fig. 1 is of a speci- 
men approximately that size. We have selected 
the ZMH specimen (E2016) from Stn 51 as 
lectotype because with a d.d. of 2.7 mm, it 
conforms closely with the size and original 
description, and remains in good condition in 
alcohol. The other specimens mentioned above 
are all smaller than 2.5 mm, and that from 
WAM is badly decalcified. 

Paralectotype designation is established for 
the ZMH, MNB and WAM specimens from 
Stn 56 although it is not possible to resolve the 
discrepancy in number of individuals from Stn 
56 (5 reported, 6 in collections). One of these 
might be one of the two presently not located 
from Stn 51. Cockburn Sound is now the type 
locality. The specimen from Stn 50 at ZMH 
with d.d. 2.0 mm has no type status. 

Ophiacantha hrachygnatha H. L. Clark 
FIGS 40, 93-98 

Ophiacantha hrachygnatha H. L. Clark, 1928: 

420, figs 123a & b; 1946: 186. 
Ophiacantha abyssicola G. O. Sars var. otagoen- 

sis Fell, 1958: 25 pi. 4, figs, G, L. 
Specimens examined: Holotype SAM K208, St 
Vincent or Spencer Gulfs, S.A., no date or 
depth; paratype, MCZ 4611, same locality data 
as holotype; 6, NMNZ 2084, S of Warrnambool, 
Vic., 220-310 m, 14.V.1969; 3, NMV H359, 
39°44.5'S, 148 P 49'E, Bass Strait, 640 m, 24.xi. 
1973; 1, NMNZ 3570, West Id, S.A., 2 m, no 
date. 1, WAM 844-78, 31°45'S, 115°02'E. NW 
Rottnest Id, W.A., 265-276 m, 18.ifi.1972; 8, 
WAM 235-78, + 3, BPBM W2597, 33°30'S, 
114°31'E, W of Cape Naturaliste, W.A., 250- 
237 m 15.iii.1972; 1, WAM 222-78, 27°18'S, 
114°06'E, NW of Bluff Pt, W.A., 99 m, no date; 
10, NMNZ 2784, 45°46'S, 171°05'E, off Otago, 
N.Z., 660-600 m, 1974. 

Remarks: Hitherto this species was known 
from the two type specimens from Spencer or 
St Vincent Gulfs. The new material, providing 
extensions eastward to Victoria, and New Zea- 
land, and westward to Western Australia, is 
smaller (1.8—4.5 mm d.d., arms <6 X d.d.) 
than the holotype but agrees with it in all other 
details. O. hrachygnatha is characterized by a 
disc cover of thin-stalked, flared multifid spine- 
lets, moniliform arms bearing bell-shaped 
dorsal arm plates, and 5-7 narrow, tapering, 
finely serrated arm spines. The microstructure 
of the arm spines consists of parallel, longi- 
tudinal toothed ridges separated by single series 




■*Ml ^ ' 

» ^ *j v>^^^ 

Figs 88-98. 88-92, Ophiacantha alternate (NMNZ 2099): 88, 2nd dorsal arm plate, x90; 89, 2nd 
ventral arm plate, x90; 90, surface detail of 88. x360; 91, middle basal arm spine, x72; 92, radial 
shield, x 40. 93-98, Ophiacantha brachygnatha (NMNZ 2084 & 2034): 93, uppermost basal arm 
spine of 2084, x96; 94, surface detail of arm spine from 2034, xl50; 95, 3rd dorsal arm plate 
of 2034, xlOO; 96, 2nd ventral arm plate of 20 34, x80; 97, surface detail of 95, xlOO; 98, surface 
detail of 96. xlOO. 



of evenly spaced perforations (Pig. 94). There 
is sometimes one large tooth on the abradial 
side of the largest spjiics, situated at about ,3 
of the spine length (Fig. 93). The external sur- 
faces of the dorsal and ventral arm plates (Figs 
95-96) are covered with perforations and tra- 
becule bearing sharp points and smooth 
lumps, respectively (Figs 97-98), 

Ophiactwtha ahyKsicola var. ataxoensis from 
New Zealami, Is a synonym of this species but 
O. abyssieola s. str. appears to dilTcr mainly in 
the character isUc cioss-shape of it* oral shields, 
compared to the simple triangular shields of O. 

Ophiacamha yatdwyni Fell 
Ophiaeamnha \aLfnvu Fell. 1^58; 23. pi 4, Rgs 
F, H. J. 

Specimen* examined: I, NMV H36I. 38 i 24,5'S, 
149 25.5'E. off Pt Hickb Vic, 823 rn. coral rock 
2l.xi.l973; 2. NMNZ 2658, 42 C 29S, 173 37 t, 
Kaikoura Canyon. N.Z , 1097-1006 m, 18.iii.I976 

Remarks: This species was previously known 
from a single specimen taken in Cook Strait, 
N.Z., at 1005 m (Fell 1958). and the Victorian 
record is thus new for Australia. The specimens 
are slightly smaller (6-9 mm d.d), but agrtR 
very closely with Fell's description. Fell 
recorded an absence of tentacle scales after the 
fourth arm segment in the holotype; the present 
material does not conform in that respect, the 
tentacle scales persist along most of the arms 
as thin spikes with three terminal thorns. O 
yahhvyni is otherwise characterized by u disc 
cover of evenly spaced thorny granules, six 
robust arm spines with asymmetric and largely 
terminal thorns, 5-9 irregularly arranged, club- 
shaped oral papillae, and rhomboidal oral 
shields. The type specimen cannot be located 
and js fenred lost. 

Key to southern Australian species of 


1. Disc covered dorsal! y by slender, tnultilid 

bpinclcts .... 2 

Disc covered dorsully by granules or 

spinules 5 

2, Dorsal arm plates triangular; longest 

armspines clavute OU Wflvfiv/.i 

Dorsal arm plates boll-shaped. longeM 
armspines evenly tapered 

O hravhyxaathn 

V llisc covered dorsally by granules ,. 4 

Owe covered dorsally by spimuVs 5 

4. Disc granules as wide or wider lhan 

high, blunt; first uppcimost armspine 
enlarged . . . O. shepherd; 

Disc granules as high or higher than wide. 
with terminal [horns, uppermost 
armspines of similar size O ynldn-yni 

5, Disc with pointed spinules, enlarged 

inter radially near disc margin; arm 
spines alternating in numher 

O, tdtenwta 

Disc with pointed spinules enlarged 
near disc centre; armspines jtoi 
alternating tn number , O. hercrotyla 

Ophtoprmm rosea (Lyman) n. comb. 

OphiacxHtha rosea Lyman, 1879: 139, pi. X, tigs 
2n7, 268: 1SK2: 184, p|. 2.S, tiGs 1(1, II- Fell. 
1958: 23, pi. 4. fin* J. K.-MeKniRhl, 1907 

Ophiacantha truncato Koehler, 1930: M, pi VI, 
fags 13-14. 

Spe\ imens examined' Holotype, BM(NH) Wilt 
23.175, 50JLVS, 74°42'W, S Indian Ocean. 320 
n). 5i.l876 (Challenger Slh 30KI. 11. NMV 
H364 & |. NMNZ 2654, 3944-S S. 14X^49 1, 
Bass Strait. 640 in, 24.xi.l97}; 2, NiMNZ 265S, 
43*14*$, 173'39'Hv Pegasus Bay, NX. 1006- 
512 m, 27.ix.1976; I. ZMC, ?H"[5S, 349*20'E. 
S Of Cape Lvcrard, Vic. 270 470 m . I^Xv.lVM 
(holotype of Ophiaconthfi tranrota Kochl.:r) 

f<£rnarks; This robust, brightly coloured species 
has been recorded from New Zealand by Fell 
(1958) and Me Knight (1967), and southern 
Chile, Japan, and Cro/ct Is. by Lyman (1879. 
1882), thus its occurrence in Australian waters 
is not unexpected. 

The new material ranges 7-15 mm d.d., and 
has arms about 4 K d.d. At their bases, the 
arms of a 12 mm d.d. specimen are 4 mm high 
and 3 mm wide, with 10 arm spines up to 4 
mm long, The arms Therefore have a very 
robust appearance. The disc i& covered entirely 
with small thorny fftOWpfl >'p to 3 * higher 
than wide. There are 18 oral papillae on each 
jaw, 6 internal and the remainder external on 
each side. The internal papillae are rather spini- 

< Hher s'peciea recorded from southern Australian 
waters and refected To Ophiacamha hy H. L, 
Clark M946) art here regarded as belonging 
|0 different genera, viz: Ophnnanthn trtint&ta 
- Gphioprtnm nwu i t VAltftfl 1 1 QphUl\ •aaihu 
frdelix ~~ Ophiomitrtl/a ftdetis Koehler. Ophia 
tnmha lortzesrtt = Ophialcacn vongesta (Koch- 
let ); Ophiacantha wfermfeHHni Ophivtnut 
i-aU-nciennest \ I.vman ). 



form, whereas (he external ones arc spatulatc. 
The oral shield is of an unusual bell-shape, wilh 
the apex cif ihe hell pointing distallv There is 
one very large paddle- shaped tentacle scale on 
each pore except the lirst. where there are 
occasionally two (as in the holotype), 

The concealed radial shields, numerous oral 
papillae generally of two distinct types and 
position, t'.v.i h;isil tentacle scales, and follow- 
ing single scale of very large si/c\ indicate that 
this species should be placed in Ophinprittm 
H L. Clark rather than Opitiai atitha, Tn other 
respects, such as size, form and sculpturing of 
arm spines, and the nature of the arm plates 

tnscu closely resembles the two lurce species 
of Ophioprium (O, htrissae and <J kapafae) 
known Irom the Sea (Baker 1979)- 
O. rosea is distinguished by its red colour, short 
arms, and the shape of its oral shields and ten- 
tacle scales. 

Ophtacunilm inauato Knehler. known from 
two specimens from just northed*! ol Tas- 
mania, is here regarded as a junior S> uoiiym of 
O. rosea. The holotype has identical disc 
stumps, arm spines, arm plates, and oral 
shields, and up to I 7 oral papillae of two Kinds. 
The five arm bases have only one large Wftl I 
scale to each pore indicating that the presco-.t 
Of more fbun one scale is not a stable, charac- 

Ophtop/mihacn Venill, 1899 
Ophuiplifultuca imisa (Lyman) 

Ophlomtlra imisa I yman, I X83 1 263, pi. VJ. 
fig.s 89, 90. 

OphioptiMhaco itwt.w VerrilL 1899: 351.- H. t 
Ctark, 1915. 211. 

SpechntH* tntnkmh 8. NMV H36t a( k $43 s 

147 25 VR. S of Pt Hlc&b Vic. 923 m. rock Mid 
coral, 21 xi\ 1973. 

Remark*: The discovery of this species in the 
waters of southern Australia is somewhat sur- 
prising considering that all previous records 
have been from western Atlantic Ocean near 
Barbados and Dominica, Its bathyrnetric ranee 
of 610-1572 m. suggests however, that It may 
Well be widespread like some other deep BOD 
tinental slope ophiuroids. 

The largest specimen is 17 mm J *j , and one 
has what is possibly a dwarf male attached to 
its aboral surface. The specimens match 

1 yman's description closely, and the range of 
sizes shows small variations' the arm spines 
vary Irom five to seven, and the disc eovcr 

vanes from smooth stumps at the edge to 
thorny stumps near the disc centre. 

Ophioplintlitutt is cosmopolitan on the con- 
tinental shelf ol deeper, and contains about 30 
nominal species. No representative has hitherto 
been recorded from Australia, although several 
are known from Indonesian waters ( Kochler 
|93Q>. Of Ihose. O. vtnna Koehler from the 
Bands .Sea, 1595 in. is very close to this spe- 
cies. It is known r'roni a single specimen, and 
appears to diltcr mainly in the shape of the 
oral shields and the alignment of the externaJ 
oral papillae. 

Fa Mil v Ophiuctichic 

Ophith fi$ I iitken, 1856 

(iphhu-th tricolor H, L. Clark 

FIGS 33, 34 

Qphtimtto trhvlor If | ClarK. |S2fi: 427 1?>. 
rifi U6; 1938: SB: I94rv 208.- a. m ciark. 
lyfir. i$& en key 1 

Ophimri* teeris H L. Clark, 19*H: 268-270; 

i oar,; 20*. 
OphtBCtirlutffkvni tCortiter HOT: 245 

SpsfTmenx extttrtkted: Holotypc SAM K2I3, no 
locality data or depth; pumtypc SAM K^U, no 
locality data or depth I Res 33 14); 2, NMN/ 
W7& 3 km off West Beach. Gulf Si Vincem, 
S.A.. 9 m, Svi I06R: 2 NMNZ 20X3 otf Pt Mars 
den. S.A., 20 m. Jan. 1965: I, NMN/ 2MXI. 
Yankaldla Bav S.A.. 18 ro. no dnlc. 1 t NMN/ 
207X, West Id, S.A.. 2 m. 20.X.1967. |, NMN/ 
20K2, wreck of KfortMdi Gulf St. Vincent. S.A,. 
J2 m. I.!.t%5; ^ > BPBM W1S06, Investigator 
Strait, S.A. 30 m, no date; 2, BPBM WiytV. 
lrm*Mjpator Strait, 43 m, no date; 3, BPBM 
Wix:s- ofl toen--l.e, Gulf St Vincent. S.A . Cm. 
rock, amnnp sponges, no date, 2. [.muivpes of O 
favii.s WAM 221/2-30, Bunbury. W.A , 26.x. 
1929. no depth; 2, WAM 499-7-1 S\V of Bunbury, 
WW, 20 m. 13aVI9fVs; 4. NMN/ 2069, Ragle 
Nay. Cape Naturalise WA, 1-5 nv, 24ii.l975; 
i. NMN7 IN3K. Yallmgun. W.A„ t-3 m. 72M 
1915: L MNB 4549, Genildlon. W.A.. no depth. 
1. /Mil 15444, Frcmwnlle. Ciuee Road*, W \, 
Ml m. 4.v,I'iMV i. WAM 319-78. W side Gosv 
Passaic, Ahrolho* is, \v \. S3 35 m. no dale, 

Rvmatks: OffMtQTtU "nolor was deseribed btj 
M- I Clsrt from Sir Joseph Vereo's collection- 
dredged in Ciull St Vincent and Spencer Culf. 
Sooih Australia CJark (193S) mentioned a 
1 I imeu of tricolor in (he "Melbourne 
Museum'" collected in 40 fathoms north 01 
Cape. Isorda, Kangaroo Id, S.A. but there 
appear to be no subsequent records of this spe- 
cies m Ihc literature. The new record* listed 



here show il to be distinctly southern and 
southwestern Australia in distribution, 

H. L. Clark (1938) reported a second and 
new species, O. Uteris, from dredged samples 
in Koombana Bay (Bunbmy). and Dongarra, 
Western Australia, a,nd provisionally attributed 
two specimens <MNB 4540 & ZMH ES444) 
from Fremantle and Gcraldton (reported by 
Koehler 1907 as O, lentkenj) to his new spe- 
cies. Clark (1946, p. 207) separated O fa&Vti 
from O, tricolor by differences in the size ot 
the disc scale., and radial shields, as well as by 
colour as follows: 

O. tricolor 
disc scales larger ami 
less smooth. 

O, laevis 
I disc with smooth 

coat of small 

rounded scales. 
2. radial shields very radial shields con 

small, not much Mderably larger 

larger than some of 

disc scales. 
3 disc- usually disc y.ravisb or dirty 

pmplish-rose anus yellowish, arms nor- 

varicgated hut not mally banded with 

banded pinkish white. rosy 

red. and dull blue. 

Both species were characterized in ClarkN 
k(?j by (a) a quite large single oral papilla, 
(b) radial shields small, less than .25 disc dia- 
meter, (c) very small and numerous disc 

The scries of specimens available has shown 
thai the small dilYerences used by Clark to 
separate laevis from tricolor can be attributed 
to normal variation within the one specie-.. 
Consequently, (>. laevix is here- synonyiruzed 
With O tricolor. 

Family Ampbiundac 
Atnphiura chmdifnrmis A M Clark 

Arnptiit/ra elarntiformix A. M Clark, 1966 $31, 
fig. 6 f i.- Dimiudl. 19H0: 40, fig. ]3, 

Specimens examine!: 3. NMNZ 1853, 4V0n'S, 
l4JTIJfVE, off Eagtehawk Neck, Tas., 122 m, 
I3.iil.l973; I, NMNZ 2S| 5 nff Burnie, Tas., 
20 m, 6.ii.J970; 1, NMNZ 2616. Great Taylor 
Bay. Bruny Id, Tas., 14.ii.1972. 
Remarks: This species was previously known 
only from Pt Phillip. Victoria, and with its dis- 
covery in eastern Ta.smaniau waters, it may be 
expected along the entire southern Australian 
coast. The specimens range 5-7 mm d.d., and 
confirm the distinctive features o\' this arnphiu- 
nd; the disc is pctaloid and covered above with 

small scales, and below with naked skin The 
radial shields arc long (2,3 mm on the 7 n>m 
spctHrncrO* narrow, and bowed. The distal oral 
papilla is wide and rounded, there are two ten- 
tacle scales on each pore. and lift ijWCOP4 [m 
lowest arm spine has a prmimalty dncckd 
hyaline book or is bihamulatc. 

Atnphmtu eluniiifortrits is similar to two or her 
southern Australian amphiurids, A, dot in and 
A, muhiretnnfa (both of H. 1 . Cl;uk» and one 
New Zealand species. A. art ret fa Koehler, but 
they can be distinguished by different combina- 
tions oi' the above features. 

Atnphiura fwtltirctntila H L. (lark 

Ampfytwa mNtiitrmnJa H I Clark tflUfc 32B. 

A. M. Clark, ftfifi: 33<\ fie. 7a-c. 
Specimens rjwnine<i: I. NMNZ Jtott* Wttii "' 
S A . 1 m. 29i 197S; |, NMN7 tttt, ftglc 1Uv, 
W.A . 1 m, 24.ti.l975. 

Remarks This HMiphmrid was ImherM known 
Only from 3 Pott Jackson and Lotlg Reef, 
N.S.W. specimens. The present specimens are 
4 <> and 5,5 mm d d . and the largest has 9 Mm 
spines. This large number of arm spines, and 
their Hat, blunt shape help to characterize this 
species, which is otherwise distinguished from 
the many other Australian congeners by Very 
fuic, complete disc sealing, small radial shields 
(0 2-0,33 dd.). and oral shields with a srnu 
circular proximal border 

The holotvpe of A. multirctnnla has hcen 
figured by a" M. Clark ( 1066; 

Amphmni mavcUun'n a Ljunjrman 
Amplvnra magcllanica Ljurjgjnun. 1 867 320.- 

Mortensen, 1924' 132. fig, 14. 
Specimens examined; 2, NMNZ 2fi2(\ S of U'jrr- 
nambool, Vic., 220-310 m. 14 v. 1969. 
f\etnark\: This species is apparent^ tirciuu- 
polar in subantarctic and cold temperate lau 
tudes, having been recorded from the south 
Atlantic Ocean. Pantagonia, southern New 
Zealand and, now. southern Australia. 

The two Warrnambuol specimens measure 
2.4 and 3 7 mm d.d Thev show the charac- 
teristic lengthening of (he lowennost of the 7 
arm --pines, and the large Idf-sbaped distal oral 
papilla and tentacle scales. Amphtura mageU 
fanini is viviparous and hermaphroditic — fea 
tures which separate the species from the simi 
lar A. spinipes Moncnsen from New Zealand 

Amphittnt (tisacantha H. L, Clark 
FIGS 43-48 

Arnphiura trisacantha H, I.. ClarK. |92«: 425. 

figs J 25 a & b. 



Specimens examined: Hoiotype, SAM K2 1 2, 
Spencer or St Vaiecut Cults, SA., no depth ui 
date; 1, NMNZ 2073, Upper Spencw Gnlf, SLA 
9 m. U.UJ973; I, TNMNZ 2619, North Arm, 
Westernport, Vifc, 4 nu ]3-Vi.I976. 

Remarks, Tliis specie* was hitherto known only 
from the hoiotype. 1 he new material measures 
2.5 and 6.5 mm d.d., and although all the amis 
arc broken, fragments with the largest specimen 
Indicate arms 4—5 >' d.d. The arm spines num 
her $ at ihc arm bases, and reduce to 3 within 
si\ segments. The uppermost spines are pointed 
and slightJy flattened, and the laager lowermost 
spines are more circular in cross section, and 
evenly tapered to a sharp poini. 

Tn other respects the specimens arc in close 
agreement with the hoiotype and Clark's 
description. Clark's photograph of the bnlotypc 
docs not illustrate the species adequately, and 
we therefore provide camera iueida drawings 
of the now damaged type specimen 'Figs 45- 
48) and the one from Upper Spencer Gulf 
(Figs 43*44) < 

Family Ophiotrichidae 

QptSathrh MiUler & Troschcl, 1840 

Ophiathrix fPlaraphtetihrix) alhosuuxia H. L 

FIGS 49-54 

Ophfothrix atboxtriata H. I Clark, IV28: 429, 

flit. 127. 
P!arof.<Jih>th,U otbostrim H. L. Clark. 1946: 


Ophiothtn (Ftacoi'hivthrh) MoUrUmt A. M. 
Clark. 1967; 648. 

Specimen examined: Hoiotype SAVf K21J ( 
Great Australian Bight, no depth di date. 

Remarks: This hoiotype td.d. ID mm) remains 
the only known specimen >nd we jrivc here 

drawings of various parts of it to supplement 
Clark's description and figure. The specimen is 
now very faded, but Iwo parallel lines arc 
■visible on both dorsal and veinial surfaces o( 
the arms after about the 20th segment This 
species is similar to O. (Phuophioflin *) SpQngl 
COlo in general appearance, but differs in the 
shape of its dorsal arm plates and oral shields 
The disc cover and overall colour pattern were 
stressed by H ( L Clark as being definitive, but 
in our experience, the variation in these fea- 
tures shown by O. tPlacaphtolhrix). Spongicota 
encompasses those in albostriatH, The disc 
cover of O. athOsttrmta is ulso like thnt of O. 
(Ptacophinthm) ftneocaeriite* H. I. Clark 

(hoiotype SAM K21K) (Fig* 55-59). The 
doisaJ and vernal . m\ plate? | I'ijis 52-54, 57 
& 58 I are. however, quite drslincl hi these two 
specie*, and O, Uneoeuerufeti is known only 
ErrjQfi northern and eastern Australia, from 
Broome to Morcton Bay. 

Ophlothtf* (Keyxtvnea) hymrniKcmtha H I,, 


QphtOfhtiX hymtnacaniha H. L. Chirk, 1928; 431. 

lie t:s 

Ophi<jtrhhoide.\ k>metuzcantha: H 1 C lark, l94(j 

Ophioihrix (Kevsuwv) h\menaeMUiha A. M. 
Clark. 1967: 648. 

Specimen vxarntHrrl Hoioupe. SAM K217, Great 
Australian Bight, no depth or date. 

Remark*: This spL\ k.s is known only from the 

5 mm hoiotype. H. L. Clark (1946) interred 
that the specimen mijrjtl be from the north- 
western coast of Australia, rather than the 
Bight, hot there is no information on the label 
to ttlgg£&J this, and in the absence of new 
material we must assume it is a southern form, 
There are of course other "northern" ophiu- 
roids rn Ihe Great Australian Bight and South 
Australian gulfs (eg. AstrolhKt mute Doderlcin 

6 AwphfopM othroUm-iJ < Brock) ), and it 
may well be that if this is a representative of 
the tropical subgenus K$y$tQnea t it has evolved 
in the relative isolation Of Ihe southern region 

In many ways, however, hyntcttacaniha 
reminds »ls of an aberrant O, (Fhicophtoihn a 
$(HjH$iC0la — h has the same shaped arm spines, 
arm plates, and oral shields, and we have seen 
SpOHgttotp specimens with a naked dorsal disc 
Surface. Wc have as well encountered webbed 
arm spinet in small .-.pecimens o\' the several 
hundred xponxitota examined: Up to a d d of 
3 mm, speeimens of O, sponghohl from the 
Abrolhos ts. W A., and Investigator Strait, 
S.A., reveal, on segments Under ihe disc, pr0jtf« 
mnl arm spine webbing, as well as upper arm 
plate* With scattered low thorny slumps. Larger 
.men*, however- l.<ek these apparent 
juvende characters. In view of that last | 

and the very Jinoml-iivo colon* p <t O. 

spartffkttfa fstC R-ikor I *>S i i, we considci that 
the- colourless, enigrtMic O. hymvnaetmlha 
ahOutd b* retained m Ihe subgenus Krysronea 
b\ virtue erf iis nuked upper disc surface, at 
least until a growth series ■-, lound which indi- 
cates another relationship. Figures 60-62 show 
diagnostic features of the type specimen. 




Wc thank Scoresby Shepherd, Jan Watson, 
and Neville Coleman for allowing us to study 
their collections of ophiuroids, made over the 
past two decades along the southern Australian 
coastline. Our Australian museum colleagues 
Wolfgang Zeidlcr (SAM), Loisette Marsh 

(WAM), Brian J. Smith (NMV), and Frank 
Rowe (AM), also provided material and re- 
search facilities, for which we are very grateful. 
Type specimens and other comparative 
material was examined through the courtesy of 
echinoderm curators at BM ( N H ) , USNM, 
ZMC, ZMH, and MNB. 


Baker, A. N. (1979) Some Ophiuroidea from the 
Tasman Sea and adjacent waters. N.Z. J. Zool. 
6, 21-51. 

, (1980) Euryalinid Ophiuroidea (Echino- 

dermata) from Australia. New Zealand, and 
the south-west Pacific Ocean. Ibid. 7, 11-83. 

, (1981) Brittlestars, Class Ophiuroidea. In 

S.A. Shepherd & I. M. Thomas (Eds), "Marine 
Invertebrates of Southern Australia" (Govt 
Printer, Adelaide). 

Clark, A. M. (1965) Japanese and other ophiu- 
roids from the collections of the Munich 
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, (1966) Echinodermata. Port Phillip Sur- 
vey, 1957-1963. Mem. rutin. Mus. Vict. 27. 

, (1967) Notes on the Family Ophiotrichidae 

(Ophiuroidea). Ann. Mag. nat. Hist. (13) 9, 

Clark, H. L. (1909) Notes on some Australian 
and Indo-Pacific echinoderms. Bull. Mus. comp. 
Zool. Harv. 52, 109-135. 

, (1915) Catalogue of recent ophiurans : 

based on the collections of the Museum of 
Comparative Zooloey. Mem. Mus. comp. Zool. 
Harv. 25, 165-367." 

, (1916) Report on the sea lilies, starfishes, 

brittle-stars, and sea-urchins obtained by the 
F.I.S. 'Endeavour' on the coasts of Queensland, 
New South Wales, Tasmania, Victoria, Souih 
Australia, and Western Australia. Zool. (biol.) 
Results Fish. Exp. 'Endeavour* 4, 1-123. 

, (1918) Brittle-stars, new and old. Bull. 

Mus. comp. Zool Harv. 62, 265-338. 

, (1928) The sea-lilies, sea-stars, brittle-stars 

and sea-urchins of the South Australian 
Museum. Rec. S. Anst. Mus. 3, 361-500. 

, (1938) Echinoderms from Australia, an 

account of collections made in 1929 and 1932. 
Mem. Mus. comp. Zool. Harv. 55, 1-596. 

, (1946) The Echinoderm fauna of Australia, 

its composition and its origin. Publ. Carnegie 
Instn 566, 1-567. 

Dartnall, A. (1980) Tasmanian Echinoderms. 
Fauna of Tasmania Handbook No. 3 Univer- 
sity of Tasmania, pp. 84. 
Fell, H. B. (1958) Deep-sea echinoderms of New 
Zealand. Zoology Pubis Vict. Univ. Coll. 24, 

Forbes, E. (1843) On the Radiata of the Eastern 
Mediterranean Pt I. Ophiuridae. Trans. Linn. 
Soc. Lond. 19, 143-145. 

Koehler, R. (1907) Ophiuroidea. In W. Michael- 
sen & R. Hartmeyer "Die Fauna Sudwest-Aus- 
traliens. Ergebnisse de Hamburger sudwest- 
australischen Forschungsreise 1905" 1(4), 
241-254, Fischer, Jena. 

, ( 1 922 ) Ophiurians of the Philippine 

Seas and adjacent waters. Bull. U.S. natn. Mus. 
100, 5,X + 486. 

, (1930) Papers from Dr Th. Mortensen's 

Pacific Expedition 1914-16. LIV. Ophiures 
recuellis par le Docteur Th. Mortensen dans les 
d'Australe et dans l'Archipel Malais. Vidensk. 
Meddr. dansk naturh. Foren. 89, 1-295, 20 pis. 

Ljungman, A. (1867) Ophiuroidea viventia hue 
usque cognita. Overs. K. danske Vetensk. 
Forhandl. J866 (9), 303-336. 

Lutken, C. F. (1856) Bidrag til Kundskab om 
Slangestjnerne. 11 Oversigt der vestindiske 
Ophiurer. Vidensk. Meddr. dansk. naturh. 
Foren. 8, 1-19. 

-, (1859) Additamenta ad historiam Ophiuri- 

darum. 2. Beskrivelser af nye eller hidtil kun 
ufuldstaendigt kjendte Arter af Slandestjerner. 
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(1869) Additamenta ad historiam Ophiuri- 

darum 3. Ibid. 5, 24-109, 3 figs. 
Lyman, T. (1869) Preliminary report on the 

Ophiuridae and Astrophytidae dredged in deep 

water between Cuba and Florida. Bull. Mus. 

comp. Zool. Harv. 1, 309-54. 
, (1879) Ophiuridae and Astrophytidae of the 

exploring voyage of H.M.S. 'Challenger' under 

Prof. Sir Wyville Thomson, F.R.S. 2. Ibid. 6(2), 

17-83, 9 pis. 

(1882) Ophiuroidea. Rep. scient. Results 

Voy. 'Challenger' (Zool.) 5, 1-386, 46 pis. 
-. (1883) Report on the Ophiuroidea. Reports 

on the results of dredging, under the super- 
vision of Alexander Agassiz in the Caribbean 
Sea in 1878-79, and along the Atlantic coast 
of the United States during the summer of 
1 880, by the U.S. Coast Survey Steamer 
"Blake". Commander J. R. Bartlett, U.S.N., 
commanding. Bull. Mus. comp. Zool. Harv. 10, 
227-287, 8 pis. 
McKnight, D. G. (1967) Additions to the 
echinoderm fauna of the Chatham Islands. 
N.Z. Jl Mar. Freshwat. Res. 1, 291-313. 

Mortensen, T. (1924) Echinoderms of New Zea- 
land and the Auckland-Campbell Islands, II 
Ophiuroidea. Papers from Dr Th. Mortensen's 
Pacific Expedition 1914-16. XX Vidensk, Meddr. 
dansk, naturh. Foren 77, 91-177, 4 pis. 



Muller J. & Troschel, F. H. (1840) Uever die 
Gattungen der Ophiuren. Arch. Naturgesch. 6, 

Pawson, D. L. (1969) Astrothrombus rugosus 
Clark, new to New Zealand, with notes on 
Ophioceres huttoni (Farquhar) , Hemilepis 
norae (Benham), and Ophiuroglypha irrorata 
(Lyman) (Echinodermata: Ophiurodea) N.Z. 
J I Mar. Freshwat. Res. 3: 46-56. 

Verrill, A. E. (1899) North American Ophiu- 
roidea. I. Revision of certain families and 
genera of West Indian ophiurans. Trans. Conn. 
Acad. Arts Sci. 10, 301-371, 2 pis. 

Wilkie, i. C. (1980) The systematic position of 
Ophiocomina Koehler and a reconsideration of 
of certain interfamilial relationships within the 
Ophiuroidea, pp. 151-157. In Jangoux, M. (Ed.) 
'Echinoderms: Past and Present'. (A. A. Bal- 
kema, Rotterdam). 


byR. 7. F. Jenkins 


A 20 year cycle of ideas has led to a concept of an 'Edicaran Period' embracing the geological time 
interval characterized by the soft-bodied metazoan assemblages of the terminal Precambrian. 
Continuing finds of such assemblages emphasize their potential value for bio stratigraphy. 
Geochronological studies suggest that the earliest finds may be at about 640-620Ma; metazoans 
appear to have become diverse and relatively numerous during the interval 600-570 Ma. Evidence 
from central England establishes that the Ediacaran does not overlap the Cambrian and that separate 
sequential depositional events represent these time intervals. The Ediacaran is Precambrian in 
aspect, without the significant skeietai-bioclastic component typifying the Palaeozic or Phanerozoic, 
Studies of Harland & Herod (1975) and Glaessner (1977, 1979b) link the Ediacaran with the post- 
glacial part of the Proterozoic Adelaidean succession, and by implication indicate a type area within 
the Flinders Ranges, South Australia. A stratotype for the Ediacaran is nominated at Bunyeroo 
Gorge, and comprises the Wonoka Formation and Pound Subgroup, the age of the Wonoka 
Formation being suggested to approximate the time of appearance of megascopic metazoans. The 
exposure of the Rawnsley Quartzite of the Pound Subgroup in Bunyeroo Gorge is made the 
stratotype of a Late Ediacaran Epoch reflecting the major evolutionary radiation of surviving 
invertebrate phyla. 


sigmik \n< i in AUSTRALIA 

by K. I. F, JknkiW 

JrNKiNN. R. J. F (1981) TH* concept a! EZIt Tdiaearan Period' and its htratierapbic slen|ft._.nKL 
in Australia. fiwiJu KSoi. S Atm UIS(4». I7V-M4, I t December, 198 1. 

A 20 year cycle Off ideas lei * !<> a ojncrpl ot an 'Ediacaran Renod' emhiacing (he 
geological rime Interval characterized by the soft-bodied meta/oan assemblages of the terminal 
f'recambrian. Continuing finds of such assemblages emphasize their potential value, tar hiu- 
StrAjjffrtfpdy, Geoeht-onological studies suggest lhat the earliest finds may be at about 6*10-620 
Ma; mefa/oan:, appear to have become diverge and relatively numerous during the interval 
o()0 570 Ma. Evidence from central England establishes that the Edutearan does not overlap 
the Cambrian and that separate sequential dcpositional events represent these rime intervals. 
Tho Ediacaran is Prccarnbrian in aspect, without the significant skeleial-bioelaslie component 
typifying I he Palaeozoic ov Pbancro/.oic. Studies of Harland & Herod (1975) and Glaessner 
t 1977, l^^b) lirk (be I J» with, the postglacial part of the Proterozoic Adelaide. m 
succession, and by implication indicate a type area within the Flinders Ranee*;. Sooih Aus- 
tralia. A -.trad'tvpc far the Ediacaran is nominated at Hnnyerou Gorge, and comprises the 
Wonoka Formation and Pound Subgroup, the age of the Wonoka Formation being suggested 
to approximate the lime of appearance of megascopic mctazoans. l"he exposure of the Rawns- 
Icy Umm7,re vi the found Subgroup in Bunyeroo Gorge is made the stratotype of a Late 
Fdiucaran I'poch reflecting the major evolutionary radiation ot surviving invertebrate phyla. 


Glaessner (19351 compared finds of early 
soft-bodied mcta/oans al Fdiaeara. South 
•\ustralia, Willi similar fossil remains from 
Charnwood horest, Leicestershire. England, 
and the Nama Group of Namibia (South 
West Africa), and established the age of these 
assemblages as laic Prccambnan. These finds 
are now justifiably famous and the known 
record of comparable fossil remains is 
becoming increasingly extensive with some IK 
or so occurrences widely sited aboui the globe 
(e.g. Glaessner 197'>b. fig 2i Though the 
organisms often atttaiocd large size and show 
a general lack of hard parts, their fossil 
imprints show no evidence of maerophagous 
predation, and this, coupled with rapid burial. 
wa* &i undoubted tiupurianee f-Ol their pre- 
servation (Wade l%S; Glaessner 1 979b). 1 he- 
kinds of organisms represented include meclu- 
soid coclentcrates, a member of the comitate 
ctiiMuirophore hydro/cans, representatives of 
sea pen-like animals, several kinds of worms, 
primitive arthropods and the problematic 
lophophorate Tribntchidium Glaessner Other 
curious sack-shaped organisms and enigmatic 
forms composed of chambered vanes are refer- 
able to the new Phylum Petalonamac PMug. 

* Centra for Precambrittn Research, Univci-silv of 
Adelaide, GIVO Bov 4VK, Adelaide. S, Au-i . 

Amongst students of these fossils there is 
broad consensus that the remains icllect a 
phase of accelerated animal evolution preced- 
ing the Cambrian (Glaessner 1961, 1971 
1972, 1977, 1979a,b; Glaessner & Waller 
19S]; Cloud |97<>a,b; Sokolov I976a»b f 1977; 
Stanley 1 976a. b; Sepkoski 1978, 1979; lord 
1979, l9K0b; Lowenstam 1980). The interval 
during which these organisms lived is variably 
recognized by Western workers as the 
"Ediacarien" Stage (Tcrmier & Tcrmier 
I960! or as either the Ediacarian (Cloud 
1972, I976a,b> or Ediacaran Period (Harland 
& Herod 1975; Glaessner 1977, 1979b), The 
approximately equivalent interval is generally 
termed Vendian by the Soviets (e.g. Sokoiov 
1973; Keller 1979). The broad relationship 
between ihe Ediacaron and Veudiuii i>> indi- 
cated in Glaessner i 1979b, fig. 1), 

Of recent years, diverse new assemblages oi 
Kdiucaran fossils have been located in north- 
ern Russia in parts of the RjsdktttO 9Ub&tagS of 
the Valdai Group, on the Onega Peninsula 
and the Zimnjy Bereu (Winter Coast) of the 
White Sea (Keller & Fcdonkin 1977. 1978, 
1980a, 1981). As well as these having several 
ta\a Ul common, some 10 or II species which 
variably occur in these assemblages arc also 
present at Ediaeara and other U<ssil sites in the 
Flinders Ranges, South Australia (Fig, 1). 
This finding led Keller & Fcdonkin (1976) 




' -l-P'. ••!• 'MM 


, ■ ■ 

( ;l I , 

- ,: 

I |, 

L P 


Flfi, I Fdiacaran localities discussed in text. 
I ai ger map shows greater part of Flindeis 
Ranges with outcrop of Pound Subgroup indi- 
cated by areas of stipple. Occurrences ot EdU 
I :n ,i assemblage arc marked 'F. New occur- 
rcii'.is are from work of Mr Colin Ford, Mr 
lim GehJing and Di Krinn Daily, Base map after 
lenWns St Gehliug I l ( >78). 

and Fcdonkin (1977, 1981) to suggest lhat 
the northern Russian sites and the localities in 
the Flinders Range* arc closely equivalent in 
time. Several of the shared taxa have been 
considered as const it tiling an Oppel /one 
(Rozanov & Sokolov 1980). A number of 
other pairs of sites share either one or more 
species or have taxa showing resemblances 
suggestive of an evolutionary affinity. "Medus- 

oids" are virtually ubiquitous. Thus it is 
apparent that all the finds are broadly related 
m geological lime, and it becomes appropriate 
to consider the potential use of Ediacaran 
fossils as a means of correlating between 
major land areas. 

The recognition of such a bioslratigraphv 
necessitates the nomination of stratotypes for 
the time divisions involved, Time-rock divi- 
sums such as the Sinian System (Grabau 
1922) erected for sequences in northeastern 
China (standard section in Ji Xian County, 
Tianjin, Kao et al. 1934) and extended by Lea 
& Chao (1924) to sections in the Yangtze 
Cioiges, Hubei Province, South China, or the 
Marinoan 'Series' (Mawson & Sprigg 1950). 
measured on rocks cropping out immediately 
south of Adelaide, South Australia (Thomson 
1 966) . have useful regional application for 
purposes of infracralonic correlation. The 
modern usage of the Marinoan in this sense is 
discussed by Thomson (1964, 1969). Historic 
ally, these divisions have priority over subse 
qtiently designated time-rock terms. However, 
the intervals of lime embraced by ihem arc 
incomplete or rather broad and their eorrcla- 
iion between different land areas remains 
essentially arbitrary. 

Current usage of the Vendian is subject to 
controversy as to its stratigraphic beginning 
(Keller 1979) As yet no slratolype has been 
nominated for the Ediacaran, which however, 
is linked geographically with the Flinders 
Ranges in name and in ils associated asscm 
blage. This paper attempts to order geological 
and evolutionary events in the terminal 
Precarnbrian and examines potential strato- 
types for the Ediacaran. 

Concept of an Ediacaran Period 

Tcrmier & Termier's (i960) erection of an 
"Fdiaeancn" Stage met with little support, the 
fossil remains being known essentially at only 
three world localities al that time. Jn a later 
work the same authors (Tcrmier & Termier 
1967, p. 141) discuss "la faune d'Ediacara'\ 
and subsequently they refer to the ditTcrent 
world occurrences of the comparable fossil-, as 
"la faune ediacarienne" or "<Jes fauces cdia- 
eaiienues" (Termier & Tcrmier 1968. p. 74 & 
76) Wade's (1970) notation of an "Ediacara 
fauna'* was widely adopted and used inform- 
ally lo designate assemblages from a then 
increasing number of localities reviewed by 
rdacssiKT (1971). The tacit implication of 
this terminology was that the fossils from 

K. J. h JhNKlNS 


different arca_s showed similarities and were of 
comparable age. that is, late Prccambtian- 

Thc concept of an * Edification" Period to 
embrace the rocks comaming these early soft- 
hod icd meta/oun assemblages was advanced 
by Cloud (1972. 1973. 197600 who also 
reviewed the contentious question as to the 
relationship belwccn this interval and (he sue- 
ivvding Cumbrian characterised principally 
by shellv I'ossiN. Cloud (1976a,b) made several 
proposals concerning either the maintenance 
of the "traditional" Cambrian bouudary, or irs 
lowering to include the Ed'acaran, a posxibi- 
liiv also explored by other authors (eg, J-rcy 
& Seilaeher 1980: Rozunov & Sokolov I9S01. 
Cloud further advanced the notion that the 
etymology of the divisions Phanerozotc and 
Palaeozoic wcte such that these time teems 
BhOUid properly be considered as extending 
downwards to the start of his "EdiacanarT. 

Harland & Herod ( 1 975) nominated an 
Ediacaran Period bounded by suggested age 
limits o\ "u>\)± and ?570^i Ma and indi- 
cated tn comprise the latest part of (he Adel- 
aidean. the time term used (Thomson 19691 
10 embrace the depositions! history of the 
cumulatively measured 24 km or so of Precam- 
brian sediments occurring in the Adelaide 
Inld-Bell (Geosynclinc) Harland & Herod 
placed the Ediacaran \n a sequential division 
of time inteivals from oldest to youngest 
respectively: Sturtwo, Varangian. Ediacaran 
..rid Cambrian. The Sturtian (Howehin I91H. 
p. 34f)', Mawson & Sprigg 1950) is the time 
division associated with the earlier and major 
phases of glaciation in the Adelaidcan The 
Varangian is associated with late ice activity 
in the Precambrian (Harland 1968). and in 
the usage of Harland & Herod (1975) it is 
implicitly equated with tlifl Marinoan phase uf 
glaciogenic deposition in the Adelaidcau This, 
the FJntimi glaciation (Mawson 1*949) • J - ; 
represented by diamichtes and varvile^ in- 
cluded within the Elatiua Format ion. at the 
top of the L'mbcratana Group, in the Flinders 
Ranees. Glaessncr (1977,197%) maintains a 
comparable division of the late Precambrii.n 
to thai of Harland k Herod and in his review ■ 
of pertinent geochronogical data, indicates 
that rocks with EJiaearun assemblages tend to 
fall m the age range 640-575 million years. 
Kc further cautions that the "appropriate 
dating* ... do not support any biosiralt- 
viaphie subdivision at the present state of our 

However, workers such as Sepkoski { 1978, 
1979) and Uraster (1079) extend the studies 
of Soviet students such as Sokolov (1973, 
1976a) and recognise informal subdivisions 
withm the Vendian. tn the work of Sepkoski 
(1978, 1979) the Vendian is -Approximated 
with Cloud's (1972, 1976a.b) usage of the 
"Ediacanan** and is subdivided into an 
informal four fold division of major biottc 
Uiufa* though it is admitted that current bio- 
sn.aigraphic control does not permit accurate 
correlations. Other Soviet reviews of the 
Vendian {Sokolov 1976b. 1977; Yaknbson & 
Krylov 1977; Keller 1979) suggest that these 
divisions reflect an oversimplification. On the 
Russian Platform, Ediacaran fossils occur 
within the Valdai Group which Keller ( 1979") 
recognizes as the Vendian \en\u v/n'r/r/. Keller 
(1979) considers that "there arc no reliable 
radiometric data for dating the Vendian lower 
boundary" and suggests that a previous I v 
accepted c. (SR0-650 Ma K-Ar glauconite 
date for the Vendian basement must he 
corrected On the basis of newer dates he 
considers that that the base of the Valdai 
Group is about 620-bOU million years old. 

The preceding; short review indicates thai in 
the literature the Ediacaran has been attri- 
buted varying slat us. These philosophical 
viewpoints may be summarized as follows* 

1. A majority of studies equate the Ediacaran 
with soft-bodied assemblage?, of sub-Cambrian 
mcttizoans which have been broadly termed 
the Ediacura fauna. Glaessncr (1977, 1979b) 
indicates that the rocjes contain ine these 
remains fall into the approximate age range 
of 040-570 Ma. 

2. A minority of students ntttihute to the 
Ediacaran the significance of a hioslrati- 
graphic ?(age, or suggest that at least in part it 
constitutes an assemblage zone. 

3. A majority ot studies imply or indicate that 
the Ediacaran is Precambrian, and represent- 
ing the latest par! of that Erathem 

4. A minority of authors Lonsider cither that 
the Ediacaran validly comprises part of the 
Cambrian OT 'hat the lower boundary of the 
Cambrian should be extended downwards to 
embrace the Ediacaran. 

5. A minority of students imply that the Edia- 
caran is of Phanero/oic and Palaeozoic aspect 
and advocate its inclusion within these divi- 



»• Several wo ken (Harland & Herod r>/\ 
ttl:i-> ,,.cr 1V79bl allribule Ifw Ldiacaran. 
time-ruck status indicatm: l{ i t p t i w nt \i 
\x\rt cA the Upper Proterozoic and a cit vision '.«l 
the Ailcluidejn Ihcrcbv nominally restnetiuL* 
its iypi>ari:;j to the Adelaide Fold-Belt, By 

impIk^iMHi. (t\t lower boundary of the Bdis- 

wroil Jtcs above the Marmoan glactoyenic 

-vi nt rteogdized in this region* 

7, Some rccncni-._ informal divisions vvithm 
the hdiaearan, these divisions variably 
numbering between two and four 

H. A close equivalence is recot'ni/ed between 
the fidiacatan and Vendian. 



As the Ediacaran has been aiiMhuied 
$evi r.i! different philosphieal nicamues tl*c 
l.nn vi approach in attempting to nominate a 
stratotypc might he to choose a section which 
best illustrates concepts held by the majottfy 
of earlier students. This approach docs not 
imply direct maintenance of priority Certainly 
those students who attribute a time-rock 
mcauine to the term (Harland &. Herod 1975; 
Gl ..Lv,n.-r |977. 197%) come closest to 
I'lovulm? an adequate definition The second 
and major requirement of the stratotypc is 

riutl rl should encompass or be bounded by 

diNiin.-iu c ecological phenomena which pro- 
vide the gro&tctf potential far coi relation, and 
consistent with the broader interpretation of 

the nominate division 

The idea that ihe Ediacaran can be con- 
sidered b* a short downwards extension of the 
lower Cambrian is not juppftftftd by gcitt* 

chronological data A U-T'b /ireon-date 
(■•oncordia upper intercept model) Of 
fi20*20 Ma (Glover & Shlnha 1971) for a 
fejftlc tulT-hreccia in near association with 
worm like impTCWioira from North Carolina 
I Cloud el at 1*376) is petbaps the oldest well 
founded age linked with any of the Ediacaran 
fosstts. Older il-u mt's have been well publicise*.! 
free reviews of Gtncssner 1 077, 1979M but 
arc nor closely consistent with those ohtatnine 
for the majority of fimK- 

Thc BCSI F.tiacaran assemblage actually 
discovered (Gihieh ijofc (933; Germ* 1972. 

1974) occurs in the lower part of the Nama 
Group in Namibia- The Nama Group r« 
place.? in stratigraphic perspective bv Kroner 
(1977) and formally subdivided bv Kroner 

ti al. (UtgQ). Kroner 09 at. (1980) indicate 
that hi part the Nam , Group rests unconform- 
ab|y on a -.equenec containing volcanic rocks 
which give a possible tnctamnrphic rcseuine 
age of h86^32 Via; J, ..where the Nama is 
intruded by granitoid rocks dated at 557 — 1 | 
Ma (Rb-Sr whole rock isochron ages; Allsopp 
f| 4 I 1 -? 1 ' d,.ay constants of Steiner & Jager 
1977>« Kh-Sr dating ftf shales in Ihe lower 
Nama Group eonhims a late Prccantbrian 

and kroner ( 1077 1 brackets the Kuibis 
SUbgr^up, the earliest pari of the Noma and 
major fofsiUfetous interval, between c, 607 - 

and c 636* million years {^superscript mili- 
tates correclion of data to constants of Sterner 
& lUgp\ 1977) R..V1V.W-- of flge data in pre- 
paration by the wnttr ftilggf&t younytr aees 
for other Western occutrences of Ediacaran 
fossils. The possible lower nee ol' 620-600 
Ma for Soviet occutrences (Keller 1979) has 
already been mentioned 

In their assessment of aee data reltflu iu. to 
Ihe Kdiiicanin and Cambrian. Ctiwifl & Cribb 

(197K| bud ibat the base of the Cambrian 
inav be bracketed between 590 and 560 M.i 
Kroner ei al. (1980) suggest that the base of 

the Cambrian be recognized at S6Q Ma. An 

increasing borlv of new radiometric data from 
England SUggeSW that this boundary may 
actually he youneer again- In central western 
fni-land. Early Cambrian strata of the "Non- 
TrilohiiL'" Zone and the "Olencllid" 1 Zone rest 
unconformably on metamorphosed rocks 
dated by Rb-Sr studies at 53n=t8 Ma. with 
related granophvnc intrusion at 533 H "I3 Ma 
(Patched et al. 1 980 k included references). 
A similar Rb Sr isochron age of 540* -58 
M:t was obtained by Crlbb (1975) from 
.li->ritcs intruding the southern part nf the 

1 hamfan Complex. The Early Cambrian Wan- 

shill Formation lies unconlormably above a 
comparable diorfle and the upper part of this 
Formation contains a shelly Hsnernblage con- 
sidered equivalent t.» the Dokt4o<}Un re^u- 
laris /.one ol (he Siberian Tommolian (fJrasier 
t-t aL I978l Brnsier & Hewitt 1979); elements 
of the lowest lomniotuin zone are also present 
'although largely reworked" fPf&SXCT €1 al 
1^»7fli. Further work on the dating of the sub- 
\ a;iibnan ol Enoland is procecdint: (Beckin- 
sale W // 19HH. 

Ihe i*eoehronologteal data |u<t revtewe*^ 
continue to support Glaessuer's t 1977. 1^7'ib) 
finding that rocks with characteristic Pdia- 
cruan melaroans may occur as early as c. 640 



Ma, However, the few items of data full about 
q in. .in of c dZO Ma. which m^ be a slightly 
(inner age for the earliest known mclazonns. 
I Jit* tu initiation of the Hdiaearan may lie at 
c. £60 Ma Off possihly as young as e. MO Ma. 
Tbcsc estimates tndictttc a possible maximum 
-.lutaiion of c. 110 Ma fur the deposition ot 
rocks with Fdiaearan meuuoans. and D ou<u- 
mum duration of c. 60 Ma. Either of these 
estimates ore comparable with the hkelv dura- 

o\ 55 in 95 Via lor the 'traditional* 

Cambrian (based <>,, data Of Cowie A: Crfbb 
I97R) and indicate that in a hivntrchy of 
logical inn. tiiv.'WiHH ihe Ku'iac;uan .md 
i ainbnan musl be accorded equal status 

The occurrence of a reasonably divosc 
Edlocaran assemblage is well documented in 
the middle part of the Charman. or the WOOd- 
hOllfte Beds of the Maplcwell Group ff «m<! 
I '>s lSi ]^8()a: Brasier cr at. 107V). However 
lollowin*! deposition of the Charnian sm 
sion folding, melamorphism, intrusion $m\ 
deep elusion occurred prior to a new nans 
en-vow event heralding the Mart of the Larly 
Cambrian (Brasier 1979, 1 9S0- Brasier & 
Hewnt I 1 )?*}) Any estimate of the Juration 
of this sub-Cambrian oroueuu event and the 
snhsct|uenl i_yelc of uplift and erosion rtTOftJ be 
speculative, but an interval of 10 to 15 Ma is 
reasonable, This tectonic punctuation between 
1hc deposition of rock 1 ; wilh I'diacaran fossils 
and true Cambrian sediments, which can be 
recognizably dated biostratigrapbically as 
Tommolian (Brasier r/ ai lS7fl)i establishes 
beyond reasonable douht that the time interval 
^presented by Ediaearan sediments and Uty 
lime of Cambrian deposits are two quite 
separate geological cycles sequential in Berth 
history This finding ifl also supported by the 
stratigraphy of the Adelaide Fold-Bell 

I am not \r\ agreement with Cloud (1073. 
1070a.h) that the etymology of the names 
Phauerozoie and Palaeozoic provides yn 
initial grounds to alter l he traditional beginning 
of these time intervals and extend them curlier 
t<» include the t -diataran I he comparatively 
rare Ediaearan mota/oan remains occur m 
rocks which are otherwise little different from 
those of curlier times in the Protero/nu\ In 
Lonirasf. sediments of the Early Camhi I 
transgression (Daily 1072; Brasier 107°, 

1980) suggest a filling of iho >eas by mvcrte 
biates, wiih frequent" bioturbation (Btustet & 
Hewitt 1079; Froy & Seilacher 1980; F.dr.nkm 

1981) and an Increasing content of metazoan 

skeletal debris, which in conjunction with 

calcareous higher algae, forms extensive bio- 
genic lmiescones. Comparable character istics 
typify the remaining Palaeozoic, with skeletal 
limestones and reefal deposits abundant 
These epochi.,1 skeletal and lilhie changes 
whit. h arc closely related in lime to the base 
of the "traditional" Cambrian arc surely ol 
greater significance for stratigraphy and his- 
torical geology Ihin the sporadic and frag- 
mentary earlier record of sofl-hodicd animal*, 
intriguing though this may be for our know- 
ledge of evolution. Thus there arc good gpo* 
loeieal reasons for considering ihat the Cdia- 
caran is properly part of the Precambfiau. and 
iis significance for evolutionary studies is that 
if contains the Precambrian precursors of 
major phvla which only later provide the 
spectacular skeletal record characteristic of 
the Palaeozoic and the Pbanerozoic as a whole, 

Vutijally all published concepts of the 
f diaoaran embrace the unique, large, soft- 
bodied melazoan remains which are found in 
its rocks. In this sense, the Ediaearan is 
'v.ntially a hiostratigmphic concept with 
similar fossils in different areas suggesling an 
approximate equivalence in time. Any real- 
fetic stratotypc should reflect this hiostrati- 
graphie aspect as it potentially provides a 
primary framework for cotrclatton. 

The Bdiacafftfl assemblages at the various 
known world sites are not uniform in composi- 
tion and Glaessner (1977) remarked ox\ this 
in relation to the usage of ierm "Hdiacara 
fauna". Jenkins & Gehling (1078) suggested 
that the remains of the eiganisms preserved 
probably represent only part of ihc original 
life community, the barrier t»i Ihe preset vahmi 
of small organisms effectively being the 
grain-size of the enveloping sediment (cl. 
Glac^ncr i'I0?2) I.Mead Jenkins & Gchlina 
prefer to name the incomplete prcM-rven 1 
community the "F.diacara assemblage'. This 
usage is only of strict relevance to the 
remains from the Flinders Ranges or type 
area Assemblages from other *cogM>phic 
localities should be identified hy an appro- 
priate silc or stratigraphic dcsignali»,>n. i e. the 
'Namu assemblage' from Nnmibin or Ihe 
'Charnian a^cmbtage* from England 

The Odiacara assemblage tfiftfft (frfr/p is 
now well documented (Glaessner & Daily 
1959; Glaessner & Wade 1AS61 Ghmfner 
1980) hut the description of the other occur- 



iciaiN is not uniform. Fedonkin ( 1M77, i9Slj 
is extremely active JD the Soviet SfihetC and 
various other studies are proceeding. End 
information is now available to suggest that 
the mefazoan remains can he divided into fttfd 
broad croups. 

One group embraces the remains from ihe 
lower Nama Group, the Charnian, and the 
fossils which Anderson & Misra (1%8K Mfcra 
flfO, 1971) arU j Anderson (19721 ■'■ 
merited from the Conception Group on the 
Avafon Peninsula or Newfoundland. The 
Nama assemblage contains the distinctive 
fossils Ratteen Giitich. Pwridirihum Gitrich and 
Ermeita PHug and new reviews of these enig- 
matic remains are being prepared by the 
writer. Preliminary remarks concerning a 
revised structural interpretation of the sack- 
shaped Emietto are given by Jenkins et af. 
(19X1), Pteridini/tm and t'rnictfn HpU ' 
organisms with ■ unuinc structural organiza- 
tion which qualifies their recognition as a new 
phylum, the Petalonamae PflUg. Tlie N: 
Groirp contains the only known world occur- 
rence of Pteridinium ximphw G ur icb, 7'bc 
structure of Rangea is- quite separate to that ol 
the Petalonamae, but closely similar to a? yet 
undesenbed fossils from the Chsmi.m 
Newfoundland assemblages, and probably to 
Charniii Ford, which is common to bolh tftttc 
areas. These organisms comprise an extinct 
order of primitive anehozoyn codenrerates, 
and their complex, ramifying Structure gtig 
gests specialized adaptation to suspension 
feetling with extreme "scavenging" capacity M 
might be inferred that at this lime either i. 
plankton, or algal or bacterial detritus was 
rare, an interesting possibility in relation to the 
stratigraphic proximity of glaciogcmc deposits 
in Ihc Nama Group, and the potential infer- 
ences this has tu the climate. Age data rvfatin? 
to these assemblages (review In preparation) 
sul'^l'M ;i bracketing hetween c. 6jW)-620 >fa 
and c. «uX)-_S9fl Ma. An aspect of the assem- 
blages of this interval is that associated trace 
tossils are limited in diversity or comparaiiveh 
uncommon The evolution and radiation of the 
RiWL;ea-tike organisms and the Petalonamae 
were evidently concomitant events and may 
relate lo the earliest appearance of meeascopi 

The second grouping comprises 'he better 
known aiid very diverse Ediacarar. Olden blag* 
from the Flinders Ranges and now the new 
localities in northern Russia. The striking 

L-haraetenstK oi (bis grouping is that it 
UKlUcUS many afiiftt&tft which Lin h* more or 
lu>s rvlaUd to divisions and fanubes sail living 
m pie*ent day 94ft (Sefcofov !*J77; Fedonkin 
1981 1. Thus thew w dtvfctte tfnila ol "roedb- 

sotds" chondrophorc hydrozoans, sea pMi -like 
distinctive kinds of worms, 
and pflmhlVe iurhiupods. Other proMcrnutu: 
organisms arc also prcscut. Ihc traec fossils of 
level arc diverse (Glacssncr 1969, 
r-tuunkin r.'77 1 i980a l b l Iftfcl 1. bill art 
restricted to marking made by creatures which 
rnmod over or burrowed horizontally in the 
•ubstrate; the traces lend to show repetitive 
patterns reflecting hehavioural adaptations 
towards the most efficient modes of scavenging 
<cg. thigrtwfcl&tfc pallet!)*) • Characteristic 
genera of tin* time are forms of ' CvWi.<- 
>vt>,t,tw" Spngg, Gliiewurifia Germ-., Dl 
wtiu Spngg and TribrachftHum Glaessner. The 
tn.von Pterhflnttm ttenmi Keller occurs both 
in northern Rn.xa .*,mj the flinders Rancc- 
The appf^vimatc au.e bracketing indicated for 
thTs grouping is V-twccn c. 600-590 Ma and 
C HOVTn. These remains can he considered as 
reflecting an initial radial ion of major 
surviving phyla of invertebrate life (Fedonkin 


Fhfl age data totaling to these two groupings 
Of assemblages arc Dot especially significant as 
the icliahihiy ol Ihe pnmary information is 
probably variable and error factors are signi 
lirmr Certain geological assumptions have to 
be m.iiV. as well in order (o relate igneous 
and dcpositional event* Ruih rhe Newfound- 
land assemblage and the Charnian assembl- 
age include Chntntfnifu us which is a frequent 
genus m the Flinders Ranees, and this alone 
suggests that the possible time separate 1 
between the two assemblage groupings is likclv 
to-be lather short. Stratigraphie. eVrdtfnCe from 
the Soviet lind on Ihe Zironiy Coast < Fedonkin 
i'j?;s, imm ) ilv 1 . that the lime separa- 

tion between the two groupings of assemblages 
19 short Near the /imniyeorsk I ighlhouse a 
116 in thick sequence contains abundant meta- 
zoan remains in both it* lowermost portion 
and near the top of the section Species hi the 

, now pnn include Cychweduw davbii 

SpneiL Ovurosatwtti Glassncr & Wade sp > 

Dkktnsonfa c&tfOM Sprigg, d. tfa&a Wnde and 

Trlknwhxdlum henttdtcim Glaessner. These 
IttXfl arc also present tn the Flinders Ranges 
and are . . ■ pi Mitual time equivalence 

1 Fedonkin t Q 81 \, The lower part ol the 

k i r I FN KINS 


Ziniiiiygoisk section still CCOTlOirtB Oirkttisania 
vostuttt and sorac other forms which .ue u pre- 
sented ifi the Flinders Ranges, these arc 
f' \h lUms bedoukiri, M vduM >■■><-. ■ tn- 
Utdt?\ (Spngg) (= Pattella puteUtformi.s 
Fcdoukin — ? rwnijiphnrrrtnnit; tiityilosum 
Fcdonkin), istiwama flirutcru Spflgg I 
linnitm,! diwitortiuv I'alij, Fcdonkjil I . 'V \ | to- 
nu-ihiMi tic\\<nU" Sprigg and CfnnmoJivnn 
Ford sp. I - RamcUmu pt'twara Fedonkin). 
As well« thiS ■ iwmbliiue includes well pre- 
served examples 0| Churnia nu&oni Ford 
wined has never been located in Ihe Flinders 
Ranges, but is m--t disrinciive form in the 
Chnrnian and in Newfoundland (slides made 
iiv.uiable hy l> B Daily). 

The rapid increase in diversity Ot Ufa '• m-v 
and coeval mulliphylelie skelctah/ation in die 
Farly Cambrian comprise a third metazoan 
r.-JniiMMi ..vent winch is well documented [e.g. 
Daily I97Z lOTfc Bcngtson \9Tf; Braver 
1979; B'asu. & Hewitt 1979). This event 
broadly coincides with the appearance of \\ be 
constructing and lube LrVin^J worms (Glacssnu 
I97n, 197S; Fytyn & Glaessrrer 1979). This 

[i.mjsiiio,i;ii interval between the very tttcw 
Prucambnan and earliest Farly Cambriwri may 
nof be ItJWSB 'ted in the Adelaide Fold-Belt. 
where there in .m unconformity of regional 
extent at about rhis level (Daily 1S»72. !''73. 
1976). An important component of the trace 
lossil assemblages low in I lie Early Cambrian 
of the Flinders Ranges and :dsn benlrafel 
Au-lralia arc- Ktfftaffl dwell nm (uiifowa like 
SL'hrhn: Haldemjn and M>>tio<rui. 
Toiell or upnvht U-shapcd forms audi R> 
Arena ohtcs Salter, ind mi a higher level. 
Piplarmtnfnn TflTtell (Daily 1971 1976. 
Jenkins 1975b-; Ciauld 1 ). In the light of studies 
by Daily and Jenkins in (he ftrCtt Jpst men- 
tioned the trace tossil assemblage described by 
Webby I 1970) from the liottaa Vale Beds in 
ihe Torrowangce Group, wcskmi New South 
WoU's & .-ot.M.jcr.d t.i In; of Cambrian aspect 

in the Fitndeis Ranees the EViinenn,,-! assent- 
hlngc is widely distributed in a thin slrat. 
giuphic interval ruzai the base of the Rawn.sley 
Oti.ur/ite. the uppcimtst formation within the 
Pound Subgroup (Wade 1970. Partes 1971, 

'Gauld. T. D i 1976) Trace fossils and the hM h 
Ihr. Cumbrian ;it AllfftpCna, m.ntlH. mi Flinders 
Ranges. South Australia. B.Sc.llfonsi IlltStti 
Uf0vcfii1> "l AiVhUlr (unpublished) 

lenkins 1975b). The stratigraphy of this major 
occurrence will be amplified later, 

Elements of the assemblage have been 
reported from the northern part of the Officer 
Basin m the possibly late Preeambrian 
Punkcrri Beds of the Punkcrri Hills, north- 
western South Australia (Major 1974 and 
references therein). Re-examination oi the 
material suggests that some specimens are of 
inorganic Origin. A circular marking may 
represent the remain, of an indeterminate 
nicdusoid. Imprints o^ another block of sand- 
"Jene resemble parts of the frond of Chamio- 
ihstnx, but just as likely are disrupted pieces <A 
a crinkled clay film. A possible I race fossil 
icsembles G\rolithc> patonicus Fedonkili from 
the lummotkin o\' East Poland. Thus the finds 
in the Punkcrri Mills do tw\ provide a certain 
indi cation of the Fefiiicanui assemblage, but 
are not ineunsisteiii with a late Prceambrinn 
to Farly Camhrian Bjfc for 'he Punkcrri Beds. 

GUiessner (1969) reported a fragmentary 
impression o( a trond-likc fossil collected in a 
loose block on Ihc Arnmheru Sandstone. 
Ain.idcMs Basin, east of Deep Well Homestead, 
S,S.\ : . t of Alice Springs, central Australia, 
Jenkins & Gchling (I97S) identified Ihc speci- 
men .is an indeterminate species of Clwrnio- 
rfiscuw Further study of the locality confirmed 
ils likefy placement in ihe lower half Of the 
Anmdier/i Sandstone {.sensti siricfo Daily 
i 4 An.niber.i Sandstone I of Daily 1^72) 
but revealed no new material. The Antmbera 
Satidstwe may be brondly correlative with the 
Rawuslcv Quartzite (Daily 1^72) 

The medusae described by Wade 1 1 96*> ) 
lien, ihe Central Mount Stuart Formation of 
the Georgina Basin, near Mt Skinucr^ arc 
-stcd to represent parts of an eaHy 
si\pho/.uan radiation. One form, Hallidaya 
fsruerl Wade, has a rather wide distrbution in 
the GtOrgjrft B.tsin where it is furdier docu- 
mented by Kirschvink (1978) and Walter 
f 1980) from Central Mount Stuart; Wade 
fl%£) also indicated iIn OCCUrrOQCfl in the 
Vm.iiJeLis liasm from probably near Uic top of 
the* Ammbera Sandstone at Faura Creek just 
south of Valley Dam. W.S.W. of Abce Springs. 
These remains nre seemingly younger in age 
thiin the Ediacara assemblage StftSU 0t\\ >•» 
I Daily 1972; Glaessner I«»771, Tlie form 
jdemifted by Glacssncr [In Ftfyn & Gluessner 
197$) is Kutlinyla a(T. conccnirictt. and also 
fmm near LfltirQ Creek, is apparently Early 
Cumbrian, The Mt Skinner assemblage has not 

I ft* 


ii i located In Mic Flinders fe»ng^ »nd »( i* 
ao open ijucMjon aw to whetbci the upper parts. 
ol »h. Ami-'i-M-i Sandsiooe overlap the agi 

QiUll lie or arc slightly voimyer. 

Fine current nils (Dzultriski $ Walton 
1965), identified as the pscudofosul Arntn- 
iutui olaessner & Walter, occur widely in the 
sub-Cambrian 10 Early Cambrian sandstones 
nf CtfUtflJ Australia and art; known at Wd 
■ ■■■■■(> in the tare Preeambrian ol the PTindcis 
Ranges. These inorganic marks arc not con- 
v.i. red to have any straticrapbie significance 

I he preceding iniormahon provides a hasi-s 
far consideration of potential slratoiypes fat 
an Ediacrian System. 


If tir land (1974) made a detailed review of 
!hc extensively discussed problem ol' defining 
an internationally recognised boundary bc- 
Iwccn the Preeambnao and Cambrian, and 
;ilsti outlined major lm.:s of evidence an«.l -. n 
tcria which might serve to identify divisions 
within the Proterozoic. With respect to possible 
sn.'iioiypes he started "Chosen SUCec&iOQfl will 
therefore be those transversmg the boundary 
lhat are most complete and have the richest 
characters, or are correlated directly with 
other rocks Which have critical characters, and 
are well described and easy to of access' 
Compantblc ideas arc embraced in the guide 
to slratigraphie procedure of Holland et al. 
C197S) ( The geology of rhe Flinders Ranges 
satisfies some of Harland's requirements and 
provides Ihick, characteristic successions, 
though not without significant breaks. In the 
Studies of Harland & Herod < 1975) and 
Glaessncr (1977, 1979b) the Odiacuran is 
linked by implication with the post-glacial, 
uppermost part of the Adeluidean of the 
Fliuders Ranges. This pari of the succession 
comprises the Wilpcna Group of Dalgarno & 
Johnson M964). The usaye ot Odiaearan and 
not "Ediacanan" (Cloud 1972, 1973 1976a, 
b) is formalized, as the stem geographic name> 
of other Periods 9rc unbroken and the endimrs 
of Period- names are not uniform (it. Jurassic 
ami not Jirrian ) The locality name "Fdracara" 
is Aboriginal in origin and is repotted by 
Goyder (18601 as "idyakr:.' 

The Wilpcna Group is placed in strati 
graphic and regional perspective by Thomson 
(1969), Thomson et al 11976) and Rutland 

H uJL ir'Ni) Ik poLential correlative* in 

central Australia . ( nd northwestern Australia 
arc indicated m f^gupfi -I W Coats & Pruss 
(19.SCM. The slrati^raphic subdivision of the 
Wilpcna Group in the central-western Flinders 
Ranges is shown in figure 2 ot Jenkins et at, 
1 198] ) and a modified version of this w given 
here | Figure 2 I. 

A boundary between the Preeambrian and 
Cambrian is established by Daily (1972. 1973. 
1976), In the northern Flinders Ranges tin-. 
boundary is the surface of disconformiry at 
fhe top ol rhe Pound Subgroup ( pr.-vioasl\ 

P id Quuiizite, Forbes 1971) and marking 

I he base of I he Uralanna Formation. The 
Bitoa Formation represents a single cycle 
ol Karly Cambrian deposition and locally 
attains a thickness of a little over 0.5 km. 
Uralanna Formation is overlain discon- 
farmahly by the Paraehitna Formation, and 
in the central Flinders Ranges the Parachilna 
r.trr.iatior. rests directly on a surface of 
erosion developed on Ihe Pound Subgroup, 
This erosion surface on Uu Pound Subgroup 
provides an upper boundary for the Ediacaran 

The characteristic Fdtacaran assemblage of 
the Pound Subgroup is principally developed 
in a comparatively thin widespread interval or 
intercalated, line sandstones, flaggy 
quart/itcs and medium to massive bedded 
quart/itcs in proximity to the base of the 
tUwtftJey Quyrtzitc (Wade 1970, Jenkitv 
1975b). This fossililetous interval is termed 
the Ediacara Mcmbei of the Rawnsley Quart 
2i tc and its lithofacies development and 
environmental interpretations arc discussed by 
Goldring & Cut now (1967) and Jenlm. 
(1975b). The stratotype of the Ediftearrj 
Member is nominated as being Unit B of 
G.Mdring & Curnow (1967. tig. 5) |fl hdiacara 
K-mgc at Greenwood ClifT, where it reaches 
a thickness o\' 30 m. The fossil assemblage or 
the Fdiacara Member is the type example of 
the main Ediacaxan bioia Rare impressions 
which mav represent medusoids occur in the 
upper part of the Rawnsley Ouartzite. 

In the western and southern Flinders Ranges 
the base o( the Rawnsley Ouartzite, character- 
ised hy slightly feldspatbic, clean white ortho- 
qnartzites with frequent clay yah 1 conglo- 
merates, is sharply erosive on pinkish felds- 
patbic sandstones included by Forbes (1971) 
In his division now recognised as the Bouncy 

K. J. F. JliNKINS 










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fM_ HDAl 

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< MOuluULCO 

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MROMAfOi if. 



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QD"^ 5 

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^ in E NOR AM A 

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pjrp v/ 

Sandstone, in Parachilna Gorge, 1 km south 
of the type section of the Pound Subgroup, 
the surface of erosion below the Uawnsk-y 
Quart/itc is irregular in form, with undercuts, 
and has a demonstrable relief of at least 10 m 
Small, angular, ripped up blocks of the under- 
lying sandstone are included in the orthoquart- 
zites above the erosion surface and locally, 
chipped fragments o\' the underlying formation 
from minor gravel lenses in small downculs of 
the erosion surface. This evidence suggests 
thai the Bonney Sandstone had undergone 
partial Itlhifieation prior to the deposition of 
the Rawnsley Quart/ite and indicates a tem- 
poral break. The Rawnsley Quart/ite repre- 
sents a transgrcssive cycle of marine deposi- 
tion with the majoi fossililcrous parts of the 
Cdiacara Member being tidal, back-barrier or 
lagoonal deposits (Jenkins 1975b), 

Lower in the Wilpena Group the only 
certain indications of metazoan life are small, 
circular imprints of "medusoids" and simple 
trails (Form B of Glaessuer 1%'.>) located by 
Dr Mary Wade in the Bonney Sandstone 
(Jenkins et ul 1981). These remains indicate 
1 bar the Bonney Sandstone properly belongs 
within the Fdiaearan. 

A major deficiency of the Wilpena Group 
as an Fdiaearan stratotype is that as yet there 
is no indication of the radiation of the Rangca- 
like organisms and early members Of the 
Pctalonomae. However higher parts of the 
Wilpena Group have not been fully explored 
foi meta/oan remains. With reference to the 
Soviet section at Zimniygorsk it seems unlikely 
that the Ran t t>i'(i-\~\ke organisms should be 
fpllnd widely separated stratigraphically below 
the main Ediacara assemblage. 

The base of the Wilpena Group is marked 
by a thin, distinctive widespread dolomite 
which forms a useful marker bed, the Nucca- 
leena Formation. This Formation is mostly 
eon tormable. though loeally discontormable, 
above the partly glaciogenic Elatina Formation. 

Fiy. 2, Chart summarizing regional stratigraphy 
of younger part of Ademidean sueecssion and 
1 atly Cambrian in central and lowei northern 
I liiuters kanyc^. Only upper part of Umhera 
laua GrOUP and lower part of Cambrjun 
shown, Siynilicanl yeological events, some en- 
vironmental indicators, and environmeuial inter- 

pitiiitlojw are summarised adj^eervt to swati* 

graphic column. Parts of section reprcyeminy 
» j'tliaearan System arc shown. 



Shale datmgs Ear ports Ql the Adcluidcan 

occurring in proximity lo (and above I possible 
correlative glaeiogenie deposits In ihe Kjmhei- 
ley Region, northwestern Australia, provide 
ages of 6?Q£B4 Mp and 672^70 Mu (CoaK 
& Prciss 19R0). The same authors review a 
dating of 676±2CM Ma far afl apparent 
correlative of the "Brachina Formation'' on 
the Stuart Shelf, west of the Flinders Ranges. 
Also in Ihe Kimberley Region, a shale dating 
for a possible correlative of the Bunyeroo 
Formation gives the number 639+47 M . 
(Coats & Preiss 1980). Olaessnei (l96 l .M 
described the trace-like marking Bunyt j rh-!wu' 
■lulvarnoi from the *Brachina Formation" but 
the biogcniciiy of this structure is now ques- 
tioned i Jenkins et at. 1 981 ). Au intensive 
search has been carried out in the Drachma 
Subgroup (Plummer 1978a) for megascopic 
bjotic rcmnms, hut structures so far located 
i.n' also considered to be of inorganic origin 
(Plummer 1980, Jenkins rt a! 19*1 > The 
limited available evidence as to the possible 
age of the Branchina Subgroup suggests it to 
h.. older than Hie estimated age of about 620 
Ma for known early metazoans from elsewhere 
in the world. The Brachina Subgroup contains 
abundant sedimentary structures suggestive 
of mostly shallow-water deposition and the 
upper sheet sandstone (ABC Range Quart/ite I 
is considered to reflect regressive conditions 
(Plummer 1978a). It and the overlying shales 
and siltstones of the Bunyeroo Formation are 
frequently reddish in colour, probably reflect- 
ing oxidative environments which are unlikely 
lo be conducive for the preservation of either 
a microflora or higher algal remains, such as 
^re well known at about this level In the 
Vendipn (Sckolov 19771. Some greenish or 
unoxidized intervals arc present (e.g. the 
Bayley Range Formation). In the southern 
Flinders Range* and towards rhc east, the 
Brachina Subgroup cbauges to a thick green 
sfltsrone seqiu-nce. the Ulupa Siltstone. which 
is presumably unoxidized (Thomson I9g91 

A potential placement of the lower boun- 
dary of d>e Fdiacatan b at the base of the 
Nuccaleena Form.iiion. which marks the termi- 
nation of the tlaiina glAttbaJon (Mummer 
1978b). The writer has no intention of debat- 
ing the question as to whether Prceamhtiun 
glaeiogenie events are likely to he either 
synchronous or diachronoim over wide land 
areas and differing parts of Ihe globe Such 
arguments have R CirCUtW eomponem as there 

is- unlikely to be knowledge of the triggering 
mechanism of ancient glacial inns While the 
I tihitMi at (graphic boundaries ot glacrogenie 
intervals may be at once obvious and of great 
value lor local and even regional studies (i .. 
Coats & Preiss I9K0). synchroneity of glacial 
cutis on a World wide hash has not beer, 
established Kroner (1977) marshalled evi- 
dence favouring non-synchroneity for the late 
Preeambnan glaeiations of Alnca, and 
gestcd movmg centres of glacial activity. The 
Sehvvarzrand glaciations of the Nama Group, 
unique in being correlative with section-, 
hracketed by Precambiian metazoao fossils 
(RiVig^ti am} PteUHnium helo\v; trace fossils. 
Cychtmedusa Sprigg sp, and the lost single 
specimen of Paiarncdusitim afrtcamim Giirich 
above; Germs 197 .a et wq I seem to be stgni 
ticantly younger than the local Elatiua glacia- 
!ion (Jenkins et al, I9S1). Placement of the 
stratotype boundary at the base of the Wilpena 
Group negates the underlying biostratigraphic 
implications of rhc Fdiacaran, as there are no 
known unequivocal Titeta?.oan remains in close 
proximity. The time-significance of the Elatina 
glaciation is properly bound in the concepi 
of Ihc Mannoan "Series'' of Mawson & 
Sprigg (1950). Finally au association of the 
Ediacaran boundary with a glaeiogenie event 
eonceptuaPy equates the interval with the 
Soviet Vendian. which has established priority 
(The Marinoun has priority over hoth the 
Varangian and Vcndian). 

A second possible placement for the baa 
nf the stratotype is .it the boundary of the 
Bunyeroo Formation and the Wonoka Forma- 
tion; the Fulter contains glautoriite and stroma- 
tolites and is of presumed marine origin 
(Thomson et at, 1976). One of the stroma- 
tolites is cOr\MdCfCd to he ot Vcndun affinity 
(V el*a 1 977 1. With reference to thee. 640 Ma 
age for a likely correlative of the Bunyeroo 
Formation in the Kimberley Region, it seems 
possible that the time of deposition of the 
Wonoka Formation and succeeding Bonney 
sandstone mav overlap or approximate the first 
appearance of megascopic melazoans elsewhere 
in the world; so far only a token effort has 
been made fo lOCBtC metazoan fossils within 
ihe Wonoka Formation. 

Over wide areas the Bunyeroo Formation 
passes gradalionally up into the Wonoka 
Formation. However, in the northern Flinders 
Ranges abnormally thick intervals of Wonoka 
sediments occur in several discordant troughs 

R. i. J. Jt.NKINS 


which involve erosion of silicones ol Lbt 
upper Hunycroo Formation and sometimes as 
much as 1,070 ID of prc-Wonoku deposits 
(Coats 1964), These troughs, described by 
I homson (1969) as "submarine canyons . 
have a width of .several km and show marginal 
brCOCia /.ones attributed to Skimping. Compar- 
able geological talwres have been studied by 
ihe wrucr in [he vicinity ot Pichi Richi Pass, 

near Qiiom. Here the base of the Wonoka 
Formal ion ic associated with a downcut 
surface and locally the Bunyeroo Formation 
is missing. The downeutting surface may 
evidence sleep slopes and at one point trun- 
cates a major pari of Hie ABC Range Quart 
zitc. A megabreccia developed m the adjacent 
Wonoka Formal urn contains boulders wiih 
Buityeioo Forma! ion and ABC Range Quart 
zitc lithologics. Individual hoilldcft reach 3 tit 
in maximum diameter and their size suggests 
lithilication prior to their erosion. Boulders 
with a silty litbology show a repetitive frac- 
turing at a steep angle to then mrcrnal bedding 
and suggestive of an incipient fracture cleavage, 
I his fracturing appears to be at chlTerent 
nuentations in adjacent blocks and not related 
W the fracturing Of the matrix maleiial 
(sillslonc). Thme may be some local discord- 
ance between the folding oi the ABC Range 
Quartzite and the Wonok-a Formation but fa 
significance is enigmatic The ecological fea- 
tures described surest litbifreation ot rhe ABC 
Range Quartzite and Dunvcroo Formation 
sediments prior to the time of Wonoka Forma- 
tion deposition, and possible deformation 
before th^- strong erosive downcutiioy pfl 
look place, Professor C C, von tier Borcb Is 
studying comparable geological textures else 
where in the Flinders Ranges These charac* 
ifTivlits suggest that ihe ^normal' oinl.i.i 
between the Bunyeroo Format ton and the 
Wonoka Formation may represent a paracon- 
formily (cf. Coals 1964). 

Ihe Wonoka Formation consisis of green- 
grey calcareous siltstone. minor greenish or 
purple shales, intercalated thin-bedded to 
massive-hedded grey limestone, and some 
HtodstonCi Pclletoids forming a component of 
the hmesiones arc considered as possibly 
being of faecal origin (von der Boreh, perw 
COttUJl.) In the upper parts o| the \Vuiii4... 
Formation frctjucnt intraformanonal congto 
meratcs unit oolitu limCfltOflCS attest a shallow- 
water origin. Spectacular sole markings 
formed by currents aie a persisicni feature 

Sandy interbeds near the top of the Wonoka 
evidence tidal deposition and there are intct- 
bedded dolomitic sh.iles, (SoJOtOftcs and sand- 
stones at the passage into the intercalated 
pink, tctdspathic sandstones and red-coloured, 
micaceous siltstoncs of the Bonney Sandstone. 
Ilieie is no evidence of anv bn.-ak in deposi- 
tion between these two formations. Well 
rounded grave] l..g a associated with thin inter- 
calated silts and sands in ihe lower parls ol 
the Bonney Sandstone in .some areas suggest 
a or deltaic biHttcncc Sandstones with 
large-scale cros:sbedding may reflect channel 
development Mud-cracked hatrzaos, possible 
rain-drop markings ..nd other sedimentarv 

structure Indicate fftqutrit emergence and 

suggest a lidal environment for major parts of 

the Bonney Sandstone (Forbes I 47 I | 

The potential placement of the lower 
boundary of the Fdiaearan ;il the b-w ot ih. 
Wonoka Formation is .idvaiUagcous fOl the 
lollowing reasons; 

J This boundary reflects a renewed phase of 
sedimentation following possible localised 
reelonism and associated enisivc down 

2 The Wonoka Formation is almost certainly 
marine and is known to include biogenic 
remains in the form of stromatolites. Jts 
frequent greenish colour indicates that it is 
onoxidized anil the variety of terrigenous 
and calcareous lit holy pes offer maximum 
potential for the tinding of a significant 
merofiorii or megascopic algal remains. 

3. Fluctuating Change* in sad level ami sub- 
sequent hransgrrafiitMI associated with the 

channelling :n the- base of the Wonoka may 
bO reflected it) other regions. Discovery uf 
i jmh'cam uucrciflora may provide a 
biological maker for these events. 

4. The boundary between thr Wonoi-.. 
I'onnation and the Ronnev S&ntfftOltt is 
transitional, suggesting continuous dc 
lion. The Bonney Sandstone includes frag- 
mentary evidence of metazoan* 

1 Several lugmcniary lines of evidence 
silgge,' tfK tirtW Of dcpoMii-M, 0j Ihe 
Wonoka t f oi nah.'ii may be inure or less 
•-•quoalenl to the first recorded appearance 
of well preserved metazoan icnvains J- 
where in the world. The Wonoka h.. 
yet been intensively explored for mega- 
scopic fossils. 



Re i. Locality ntup showing EdfacarflM strtrtu- 
type, between ntrt>v\\ and nc»>:*;% (racks. 

6. Placing the Ldiaeaian boundary at the 
beginning of a formation which is conti- 
guous with the Pound Subgroup preserves 
the concept which equates this Period won 
the occurrence of early tossil mctazoanv 

Accordingly the stratotypc for an Ediaeai fen 
System is formally nominated as comprising 
[be whole of the Wouoka Formation anil the 
Pound Subgroup. The type action is nomi 
nattod aa bjring al Bunyeioo Gorge (Figii' 
& \) t where the ahovc stratigraphic intervals 
ate excellently exposal and have a combined 
thickness o\ about 1320 m. Bunyeioo Gor-e 
is also the type section of the Ronney Sand- 
stone and Rawnstev QuartzitL iicspccuvcly 
305 m and 508 m thick. Forbes 197 1), and 
contains a fossilifcrous exposure of the Edia- 
carn Member (52 m ihickt of the Rawn*ley 
Qnarizitc. In the noithern Flinders Ranges 
the Wonoka Formation and Pound Suhgroup 
reach a cumulative thickness nt about 
47(10 rri. 

The Rawnslcy Qmirt/itc in Bunyeroo Gorge 
is nominated as the stratotvpc tor a L ale EJia- 
earati Series characterised by a divot • -i 
of mcumiau fossil Tcmain* probabty reftetttiing 
an initial radiation of majui surviving antn.d 
phyla, The index assemblage fot the 
carlv part -A this epoch comprises (\*-t,.> 
Htedttsa tfttvTttl mntu -irieto. C, plana Gtacft 
nor & Wade. "Mtnfi<;aniti" atwitlatu Sptigg, 
AffvV/v/nuv.v a si m tides, Ptttegid stellate, [ 
curia fhrnie*Ki, GIae\incri.)f) vnindis. McftiB 
soma coxfato. O. lisiu, Trlbmchidi'*f>\ twr<it- 

ilicu.ft, and Pteriditiinm nenoxa. The Kd:, 
Member of the Rawnslcy Quailzilc ts estab- 
lished to be biostratigraphieally equivalent to 
pari of the Redkino subsiage of the Valdai 
Group In tlie Vendian ot nurlhern Russia tcf. 
I cdnnkin I98l). Futher correlation of dif- 
ferent world Fdiaear.ui sequences requires 
ci'iiMdciahon nf locali/cd geological informa- 
tion outside the scope of the present study. 


I Over 20 years evolving ideas have deve- 
loped concerning the geological lime 
interval characterised by the soft -bodied 
mctazoan assemblages o\ the terminal Prc- 
cambtian. Continuing finds of such assem 
Mages emphasize their potential value fbl 
Itiostratlgraphy and of iccent years there 
have been several informal designations of 
an Edi^caran Period. 

2, Cunt in umy: gcochronological studies associ 
atcd with different known Ediacaran sites 

establish that early rnctnztan assemblage? 

tirst ;tppear at 6 h4(M»20 Ma and lhat 
soft-bodied mcta/oans became diverse and 
frequent during the interval .. 6Un— 570 Ma. 
The total time rrtlefval iftvolwd is eompar- 
able with that of the Cambrian, lite base 
ot which may be placed at c 5£0 Ma or 
possibly even younger, 

I videncc Irom eenlral Lngland e\\ ablishes 
thai the EJiacaran does not overlap the 
(. aiobrian, and that separate, ggquenrta 
deposilional events hSpfi lenl these lime 
divisions Fdiaearan deposition in the 
English Charniaii i- terminated hy folding, 
metarnorphtsm, intrusion and cTO&ion 
to a transgression beginning Ihe liuily 
Cumbrian, I he mci imorphism and intrusive 
everts arc dated at c. J lO 530 Ma. 

4. The rocks containing the E<fl4€4rat1 fnssiU 
are Prccambriun in aspect, WitflOUl IhC 
significant skeletal-bioelastie COlttpOflCM 
typifying the Palaeozoic and the Phanero 
zoii. as a whole. 

;■ Fnr studies of Hnrland & HefOd (1975) 
and ClacxMier < 1**77, !97xU->> link |ht 
Tdiacaran with the posr-glaua! part of the 
Pioleii'/oie \delaidean succession and bv 
implication indieaie a type area withm the 
Flinders Ranges, The appropriate strata - 
e-iiiplue i. nerval is the Wilpcna Group. 

ft Olooingcmk deposits (Flalina Foiinai 

immediately preceding the Wilpena Group 
;oe argued to be of unecitain significance 



for international correlation. Instead (he 
base ot the Ediacaran is placed at the lower 
boundary of the Wonoka Formation, the 
time of deposition of which is suggested 
to approximate (hat of the first known ap- 
pearance of well preserved mctazoans, 'I he 
Wonoka Formation reflects marine trans- 
giession and possible later discoveries ot a 
microflora, megascopic algae or even of 
Met;t/oa may ofler a means of correlation. 
7. 'I he Fdiacaran sttutotype comprises the 
Wonoka Formation, Bonney Sandstone arid 
Rawnsley Quarlzite in Bunyeroo Gorge 
I he Wonoka Formation represents a shal- 
lowing upward cycle after an initial trans- 
gression; the Bonney Sandstone is generally 
pnralic or regressive and Eg separated from 
the Rawnsley QuarlzilL* by a widespread 
disconformity. The Rawnsley Quartzitc 
represents a renewed cycle of transgression, 
The diverse metu/oan assemblage in the 
Fdiaeara Member, near the base ot the 
Rawnsley Quartzitc, is considered to reflect 
the major evolutionary radiation oi surviv- 
ing invertebrate phyla. The exposure of the 

Rawnsley Quanzite in Bunyeroo Gorge is 
nominated as the stratotype of a Late Edia- 
caran Epoch. An unconformity of regional 
extent separates the Ediacaran sequences in 
the Flinders Ranges from the succeeding 
Early Cambrian. 


Professor Preston Cloud, University of 
California* read an initial version of parr ol 
the manuscript, and offered constructive 
criticism and posed significant questions. Dr 
Victor Ciostin, University of Adelaide, read 
and criticised the linal manuscript. Professor 
C C. von dcr Borch, Flinders University, Dr 
W. V, Preiss. South Australian Department of 
Mines and Energy, and Drs B. McGowran 
and B. Daily, University of Adelaide, arc 
thanked for stimulating advice and discis- 
sion. Drs Gostm and Daily also made invalu- 
able assistance with literature and without Ihe 
help of Mrs Mjma Stojanovie in translating 
pans of new Soviet works, this study would 
not have been possible. 


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P^VN, S. * GiAtvs.Niv:. M, F t!97'>'l ffrtyw 
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I i i iittbrUn transition In • No/tt 

gcviy Tithskr. 59. 25 4't. 
1'Ntv, R. W. & Si'HAtHLk. A. (I'^iOy Lin- uritULJ 
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2«4, 71* 7H 

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I 1M76.I Parly Phatiero/pic annclnl well I 

i'u! (hen geologleflJ and biological significance. 

/, Ctoh Sot iContiA 132. 255»-a?i 

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POl '" " -d' the Me1.'6>;i. trt A. V Ni.lo- 
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v-l. ]'\ pp. :57-2( l N. (Nauka. Moscow | 

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MWa) Precarnbn «n pah:obiolopv: Middle 

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— — f iQStti Ptirvaf\\ mine «.. i; ihropnd from 

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'•'■ ■ ff, ■!,,./ A/;/. 13. h, '1-40 1 

— A WadBi M. (I96A) Ow late Ptccambrlon 

fossils from Kb/icara, South AttsUab'a hihi< I 

- A Wat oil \1 R. tt9Sl ) Ai.-.lrahi.n 

mifnttrtnn ptitecoWolojy Fn i>, R. Hnttui * t- it i 

k. i. r, jp.nkins 


'" of II - S rfl Hemisphere" pp. 

l$l 396 Developments in Precarnbri.-m <ieu 

i.M't, 2 Ml.sevieT- Ncthciland". ) 
iK. I .. Ill A Simii-v, A K. (1*73) The Vm- 

L-iiiiitu d''lu/i>i.i!!ini, ,. Ink Piecarnbriiiri U) t-ulv 
i ri. in fVJ orogenfc tfvepi In Ihc Bcntrul 

Kicrtmonl »T Vinzitiia ami Nottrr C',-. fw, 

J. to 27 V 23«5I 
Coloring R .v<"i'Ru..w r_ n f ift67) rbewofr 
inhy .TiH Taci.;-. Ol Inc I He Pre^unbrun at 

iiliaeaia V.nrh Australia /. Kcol, ,V'r, .-|//\f 

14 !'>-.. •< 4 
{",..>,>,,», Ci W. [J6350] Mf. Goyilcr's report on 

UOi thci ti n MinflltHlRHl VarVtamc tinny t % ui>ns. 

South AutityMo 177. |h6 
CiMftAU, A. \v. UV2ZI The SflJUTJ SvMeiii. Hull. 

few! svv. cji/iy/4 i as bs, 

GvKfMi n. ri-929-) Die allesien fotfil&n Siiii 
rfnW i • Pr&hl CtaV 37, W. 

(I9VM Die Kiihis-Fossilicii dct Nanaa— 

Fbrtnotksn von Stldwrrtofrilea. FttlUonr. Z. 
15. 137-15- 

Hakmm> W. Ti. i 19811 On Ihc pniteipie o| - 
I mc Pre-t ambnun Srruiipraphieal .Standard 
Scale. fnt K \ fn{ ',r->(. Com: 23rd Sesx. Prague 
ISA8, 4, I53-2M. 

<lQ?-i: The fte-C&mtiriaTM ambri.™ boim- 

.i.nv r>< < H, lloll ,nd (ltd ) *( iinb^inn ni the 
RriiUii i-!l-.. Nonien, and Spii.-brrgen"' ilobn 
Wilcv & Sons: London ) 

$ Ht uon. K- N. i I9TS] Glactettons through 

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O^A Jftu tctul Sfc(\ Hip l\ MB- 

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(|97M^ An environmental Mady o{ ihc 

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& CiiiiiiNU I G | IV7K) A review or ilic 

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Hws S. .ht\>, Mam n 347-234 

■, Ptl'MMUt, f. S- & Mok \KIV. K. C ||[>8!') 

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( hin.. ftfAf? r,Y, ( / .SV;* f/.^ui- 13. :43-2«S. 

ku.ii*. B. M I1979I Precatrtbrian ^tratlgraphk 
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& liiM.NKtN, VI. A H&7S) New Ifcitft o1 

Ims,i)-, iq Vnia.M sr.u, .-i P^ecombriftfi rfoiTS 
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KiRstnviNK, I I ( 1 1 >7M > TIk- PfCiiivibrinii C mil- 
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Kkonhi. A. (10771 Nnn-Mnviioncity pf Late 
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289 300 

, McWtlLiAM^ M. O., GhKMS. Q. J. B., 

Ktil>. A. 15. Al StHMK K. E; U (1^80) PtfcO 
maynetism ot l.iie PreOBPlhriam to eat ly Palto- 
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BI.Hibcker Fonnnnon. iffffi J A h 280. 942-968. 

! i \ J. S. &. CH.\o. Y. T. ( 1924) Gculoey of the 
<_ior£c tiivirteJ of 1be Yangtze (from Ichang to 
TVekitei) wiih speeinl rcferenee to the develop- 
ment of ibe Corpex. iHtli. WOf, Soc. ('hunt 2>, 

I...AVIN ! ;m H. A. (1V80) Whut, if anything, 
h;»|'p(Mie.l ji the tr.m^ilion rium Ihe Preeum- 
hiinn to Ihc Phanen>/oie. f'rrcum. Avcv, 11. « l >- 


Major. R. B. * 1 S>7-4 > The Piir.kerri Reds Qmui. 
,^v//. Notify ,i?ro/_ ,V//n\ .S. 51, 2-5. 

MaW^on. D. fl'949 1 ) The Hlatino C.lacialion A 
tliiiJ recurrence of glHpTatTqri evidenced in the 
Adelaide Svslnr.. Trans. /?. S<>r S, ,1^7. 73, 
U 7-1*1 

& S;-Hi.a,, k C. (1^50) SubdiviMon of the 

■ Sysleni. Ausi. J, Set, 12. <i9-72. 

Mi<ira. S. B. U'X>y) LatC Preeambnan (?» fossils 
from Soulfieastern Newfoundland. Cff^/. .V"e 
timer. Butt 8<i, 2133-2141 

H97U Slra(i.v.ruphV and depo^jfionaj history 

of late Preeamhrian eoelenterate-bearine rocks, 
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Pa rem n, I P. Oaif., N, H.. Goodwin, R. & 
IIumm. Ivl. I. I 1980) Rb-Sr wbole-tock i^ehron 
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rocks from southern Brilain. /. Geo!. Site. 
iLanaf.) 137. 649-656. 

Pi tJMVH;K, P. S. ( I97«a) Straiiyiaplu of (he lower 
Wilpeua Group ( I ote Piccambrian), Flinders 
Ranges. South Australia. Trans. R. Soc. S. Aust. 
102. 25 38. 

(I97S1*) Note on the palacoenvironmental 

significance of the Nuccalcena rormation 
(Upper Precarubrian), central Flinders Ranges, 
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(1980) Circular stiuctuics in a Late Pre- 
cambiian sandstone: Fossil medusoids or evi- 
dffnee of rlindizalion? Trans, if. Soc. S. Aust. 
104, IVI6. 

Piuass, W. V. M977t Ihc hiostrutigraphic poten- 
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Riv.anov, A Vu & Sokotnv. B S. (19803 The 
problem of Ihc Precambnan-Cambrian boun- 
Jjiv. Gtol Mas 117. 23-27 

RfTiANO. K. W. R . PARKBft, A J . PrtT, O, M-, 
Pripss. W. V. & Mlhrku. B. I 1981) The Pre- 
^amhriiin of South Australia //> D. K. Hunter 
(I'd.) ' "Precambrian of the SoHlhctii Mrnn- 
sphere", pp. j09-?h(l, Developments in Pre- 
cambrian Cicology. 2. I'Llscvicr: Netherlands.) 

Sm'koski. .1. J. ( 1978) A kinelic model ot 
Phaneauoie t;i\otioniie diversity I. Analysis of 
marine orders. FeiffChtOtogy 4. 223-251. 

(1979) A kinematic model of Plumcnvoic 

ta.\onomic diversity If, Early Phanerozoic fami- 
lies and multiple equilibria, IhUt 5, 223 251 

»,, B. a (197^) Vendifln or northern 

hir.^iH. in M. G. Pitcher (Ed.) "Antarctic 
Geoloev" pp. 204-2IH Mrrn Am A ss, f'ettol. 
(hot. Yk ( Tulsa: Oklahoma.) 



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1976, 1, 126-143. 

(1976b) Precambrian Metazoa and the Ven- 

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(1977 ) Perspective on Precambrian bk>- 

stratigraphy. [Translated from Russian.] Gcol, 
Geofiz. 18, 54-70. 
Stanley, S. M. (1976a) Fossil data and the 
the Precambrian-Cambrian evolutionary transi- 
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(1976b) Ideas on the timing of metazoan 

diversification. Paleobiology 2, 209-219. 

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premier etage paleontologique. Revue gen. 
Sci. pur. appL 67, 79-87. 

(1967) "Formation des continents et pro- 
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(J 968) ''Biologic el ecologic des premiers 

fossiles," (Ibid.) 

Thomson, B. P. (1964) General outline. Quart, 
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(1969) Precambrian basement cover — the 

Adelaide System. /// L. W. Parkin (Ed.) "Hand- 
book of South Australian Geology", pp. 49-83 
(S.A. GeoL Surv.: Adelaide.) 

, Daily, B., Coats. R. P., & Forres, B. G. 

(1976) "Late Precambrian and Cambrian geo- 
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Shelf, South Australia." 25th International 
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33A. (Progress Press: Canberra.) 

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(1969) Medusae from uppermost Precam- 
brian or Cambrian sandstones, central Austra 
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(1970) The stratigraphic distribution of the 

Ediacara fauna in Australia. Trans. R. Soc. S. 
Aust. 94, 87-104. 

Walter, M. R. (1980) Adelaidean and Early 
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Webby, B. D. (1970) Late Precambrian trace 
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lower boundary of the Vendian in its stratotype 
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7, 59-70. 


by C. B. Foster & W. K. Harris 


Megaspores and massulae of Azolla capricornica sp. nov. are the first recorded Tertiary examples of 
the water fern Azolla Lamarck 1783 from Australia and are the oldest known representatives of the 
genus in the Southern Hemisphere. Extensive TEM and SEM studies show that the megaspore 
apparatuses have randomly distributed vacuolae, rather than defined float structures. It is this 
feature which readily distinguishes this species from the widespread extant species, Azolla 
filiculoides Lamarck. Columellae (sensu Fowler & Stennett-Willson 1978) are not discernible. The 
microspore massulae possess glochidia with anchor-shaped tips, and each contains as many as 12 
laevigate to finely granular microspores. 


by C B. Fost M* & W. K, Harris! 


i nsii:R, C. B. A: Harris, W, K. (IV8I) AzolUt iopriot/tica sp. nov. First Tertiary record of 
4Z0fhl Lamarck (SMvutimvyc) In Australia. Trans K, Sac. S, Aitst. 105 (4). 195 204. tl 
December, 198 1. 

Mcgaxpores and massuJue or AZolk MpilCOMtGd sp. nov. are the first recorded Tcniary 
examples of ihe water fern A:ollfi Lamarck 1783 from Australia and are the oldest known 
representatives of the genus in the Southern Hemisphcic. Extensive TEM and SEW 
studies show that the meeaspore apparatuses have riindomly distributed vacuolae, rather than 
iierined float structures. \\ is this feature which readily distinguishes this species fiom the 
widespread extant species, At.oIUi filtciilottfea Lamarck, Lolumellae (ffftSfl Fowler & Stennett- 
Willson 197S) are mil discernible (he microspore massulac possess glochidia with anchor- 
shaped rips, and each contains as many as 12 laevigale to linely granular microspores. 

The fossils arc of Middle to Late Eocene age, and were recovered from The Narrows 
Hods, Narrows Groben, Queensland. Their presence suggests a relatively still fresh water en- 
vironment of deposition. Known occurrences of A:,olla i .aprUomU a sp, nov, are confined to 
presently tropical area* of Queensland, Its phylogcnetic relationship to the Quaternary and 
extant species, A. fUitttloitics, found in temperate and sub-tropical regions of Australia, 
remains obscure. 


The first Australian fossil occurrence of 
members of Ihe frec-fioaling heterosporous 
turn /Italia Lamarck 1783 was repot led by 
Duigap & Cookson (1957) from Quaternary 
sediments in Victoria. They identified their 
specimens with an extant species. A. filiado'uks 
var ntbrtu which is common and widely distri- 
buted in Australia !t grows on areas of ret;, 
lively slil! freshwater. Although records of ihe 
genus are widespread in I. ale t reta.eous and 
Ternary sediments of the northern hemisphere 
(see Collinsoo t980)i A:.olla has not been re- 
ported previously fum southern hemispheie 
sequences which are older than Quaternary, 

The species described in this paper is readily 
distinguished from A, ftUcuhleieSt and con- 
geners. However, we assume Ihut the ecologi- 
cal requirements of A. cupricornica were 
similar to that of extant Azolla, although its 
geographic range seems !o be confined to the 
presently tropical areas of Australia. Despiic 
apparently suitable biolaeies in the Tertiary 
coastal and intracratonic basins of southern 
parts of the continent, no examples of Azolla 
have been found in the many areas studied. 

v Ceological Survey of QU6en3j&ll<t, 41 George St 

Brisbane, Old 4000. 
t Western Mining Corporation Ltd, I6K Greenhill 

Rd. Parkside, S. AUSt, 5063. 

Geological Setting 

The Narrows Beds, defined by Ktrkegaard. 
ci &l (197U), occur in a northwest-southeast 
trending, grabcivlike structure which is 
bounded by Curlis Island and the Rundle- 
Mt Larcom Ranees (Fig. 1), The Beds in- 
clude a thick sequence (500+ in) of oil shales, 
known as the Rundle Oil Shale deposit, cur- 
rently being assessed for commercial hydro- 
carbon exploration. Lindner & Dixon (197b) 
have given a detailed description ot the Rundle 
deposit, which includes at least six oil shale 
units or seams (of the Rundle Formation ) 
which are inlercalated with green-grey mont- 
morillinmc claystoncs, mudstoncs, carbona- 

Fig. L Position of drill-hole KDD66. fa the Nar- 
rows Grahcn 





Karoatna Creak mm 


Tol*«*«»ny Cr««k unit 



Mintlutin dot. a * h hi 


M»m»» Cr«*k %iam 


Bilck K'ln ■•■m 


Rnrntsy CrOftfting ••Am 



Fig. 2. Stratigraphy of the Narrows Bcdv N.^r 
rows Grabcii. Queensland. Units arc presented 
in descending %tr:iiigruphic order. 

ccons lhfrle&, iiniioi impure carbonates and 
lignites. Figure 2 depicts the local stratigraphy. 
The lithostxatigraphie names have been re- 
served with the Australian Central Register 
pi Stratigrnphit Names and will he formalised 
mi a paper hy Henstridge & Missen (1981). 

Although the stratigraphie boundaries of 
The Narrows Beds have not been established 
With certainly, the oeeum-ncc of rreshwaUt 
osi/aeods, often in enormous numbers »" mud- 
stones and oil shales, suggests that the Beds 
were deposited in lakes that were largely 
shallow but probably of* a permanent nature' 
(Beaslcy |"94i). As discussed below, the pre- 
sence of Azolla within the Curfew Formation 
and Kerosene Creek scam (in ROD 66, 1 39. 43 
ml further supports a wholly non-mar iue 
environment of deposition. 

Foster (1979) suggested that a penologi- 
cal assemblage from a core ol the Curlew 
Formation (designated at that time -is the 
Wattle Creek scam) intersected in hnriMiott.' 
ROD 66 at 37.40 m was of Late Eocene to 
1 iitc Oliuoccne age. This range can he nar- 
rowed co Middle lo Late Eocene using; com- 
parative range* o£ ia\a from southern Alto 
tralia -W. K- Hun. 1 , unpublished data). 

Material and Methods 

Azolla gloehidta were first recognised rn 
sirt'w slides prepared hy standard procedures 
Foi routine palynologreal examination al the 
Geological Survey of Queensland. Further 
specimens of massulae and megaspores were 
isolated by washing the disaggregated sedimenl 
iftcr digestion in 70% hydrofluoric acid 
through a 200 mesh/ men sieve. Individual 

rrtctiasporcs and massulae were picked from 
tlli coarse traction and either mounted for 
SIM CM bleached in [% sodium hypo- 
chlorite suluturn aritl mounted in glycerine jelly 
on microscope slides for examination hy trans 
mured light Those selected for SEM study 
were gold coated -md examined with a Cam 
bridge S€0(] SEM al the University of Queens 
land or with tin ISI Super Mini SEM at the 
Geological Survcv pl South Australia. 

Unbleached specimens were selected for 
shuly hy both conventional transmitted light 
and transmission electron microscopy (TEM) 
TEM sections were prepared and studied at 
die I'lcctron Microscope Unit, University ot 

Specimens are retained in the palynological 
Collection at the Geological Survey of Querns 
laud, Catalogue numbers prefixed Q are those 
of that Institution, All photomicrographs are 
from unretouched negatives and prints, Mag 
mlicaiions arc given for each figure. Co- 
ordinates of specimens, given after slide 
numbers are from a Zeiss Photomicroscope 
III No 1369 housed al the Geological Survey 
of Queensland. SEM negatives prefixed MIS 
We held in the above repository; all TFM 
negatives are stored al the Electron Micro- 
seme Unit. University of Queensland. All 
specimens arc from the Type Material 

Systematic Description 

Family SALVIMACEAE Dummtici 

Genua AZOLLA Lamarck 1783 

A/nlta laprivornica Foster Sl Harris sp. nov. 

FIGS 3-8 
Diagnosis! M ll 1 . ^p.uL apparatus without Hon' 
structures and lacking discernible columella 
| TeftSU Fowler & Slennelt-Willson 1978), 
Proximal hemisphere of megaspore proper 
surmounted by dense, hairy, conical perinal 
structure, within which vacuolae are randomly 
developed and distributed | vacuolae best seen 
in TFM sections, barely discernible using con- 
ventional light microscopy). Cone commonly 
larger than remainder of megaspore apparatus; 
towaids cone apex, outermost hairs are com- 
monly fused lo delimit a sfc smooth apical 
area (best seen using SEM). Basal perimeter 
of cone defined hy o ± well developed collar. 
Below collar, and covering distal hemisphere 
of megaspore, pcrinc (differentiated into en- 



Fig. 3. A zolla capricornica Foster & Harris sp. nov. a-c. Microspore massulae. a. Holotype (X170); 
slide 9884, 9.5 98.0, Q130. b. Paratype (X205); slide 9884, 21.5 119.9, Q131. c. Paratype (X209), 
note microspores; slide 9884, 11.2 120.8, Q132. d, e. Microspore (X500), proximal and distal foci re- 
spectively; slide 9884, 3.2 96.6, Q133. f-h. Megaspore apparatuses, f, g. Holotype (X139) at differ- 
ing foci; slide 9883/9, 11.7 108.2, Q134. h. Paratype (X118), showing folded megaspore exine; 
slide 9883/6, 10.5 107.5, Q135. 

doperinal and exoperinal layers) less hairy and 
coarsely reticulate. Reticulum perfect to im- 
perfect; lumina 7—30 ^m wide; muri of fused, 
intertwined, irregular regulae-like elements, 
5-12 juin in overall width, up to 20 ^m high. 
A central canal, extending from proximal 
megaspore to apex of cone may occur in 
some specimens. Microspore massulae irregu- 
lar mostly spherical to ovoid, dispersed or 
attached to megaspores; often sterile or with 
as many as 12 microspores in each. Micro- 
spores, spherical, trilete; laesurae simple or 
with narrow, low labra, almost reaching to 
equator; exine 1.5-2 ^.m thick, laevigate to 

finely granular. Surface of massulae bearing 
3-20+ glochidia. Each glochidium has anchor- 
shaped tip, mostly with two recurved hooks 
(each 10-14 ^m in length from apex of tip), 
extremely rarely with single recurved hook at 
terminus. Glochidia mostly aseptate, very few 
with one or two septa; each ca 0.5 ^m thick, 
strap-like; 3-15 ^.m wide, constricted at point 
of origin and immediately below anchor-shaped 
tip, maximum width about midway along 
length (42-93 //m). Exine of megaspore (in 
surface view) minutely punctate or granulate, 
with irregularly distributed cavities in section, 
5-8 jirni thick; perine 13-28 ^m thick. 



, ' 


- ■ 


Fig. 4. 





lf:"'-v ^^ 


'* J 


.^ l > " ~ 


Fig. 5 



O m % 

Fig. 6 



Dttnmstom. Mcgasporc appanilus: ■ 
410-560 fan: bn [wirJftO MO-400 

(50 Specimen v) i Microspore massulac: 103 
240 ,,,mi (maximum diameter V\ specimens! 
microspores 24 JO " m fequtttartnl dfiiiimiei ). 
No/ufvpf: Mts4sparc ..ppu.aiuv Fig- ?. f,g; 
single mount, slide 9883/9. coordinates 11.7 
108,2, OM4 i. ".i length ins ,,m. maximum 
width, iit collai J32 wir. Mcgasporc wtfll 5-1<1 
,,m thick minutely ta Pen 

distal hemisphere d megaspore 10.5-1 ! c 

|M im thick! imperfectly rcticutei lina 7 MJ 
,,m wide*, rauri 3- i2 .<rn Vide 

Microspore m-i'-Miiii, Ffgurd 3a! strew 
mount, slide 9884, coordinate* 9.5 98.0. Qim 
Outline ' cjr I ; ' ^m to diameter. Ftmr 

nucmsenpt". fctt visiM.. • , (2 /.m in 
meter, cxinc I 7S „m Ihfcl {I 

arc visible, rr»U Irnyttl tawh dlsocnriblfl 63 
,jm. w iilllt 3-5 &.S imi. 

/v.v locality Diamond drill hole RDD 
at 11)0.25 m Grid rcfcttncc 738ZIN i 
G I \DS1 ONE l 1 00 000 shed no. 9150; 
khamp'.on 1:250 000 Shed Area. Oueem- 

horizon- The Narrows Beds, Curlew 
Fi»rma!i-:-i. 5l I " kitfrotagy; grey -brown car* 
ludfitone. Age: Middle to ' 


Derivation p/ name; f loin l lie I *upie of 
Capricorn; present known occurrence-. of -4. 
capticottuat arc confined to lis which 

straddle the tropic, between £2 artUl 2 s 

Comparison and reworks: A simplified ter- 
minology has been used to describe Specimens 

i . -olhi capritaffuCa sp. nov. and the accom- 
panying illustrations i Pigs W!) clarify the 
m0ipholo;;v ol thi: species. Floats, as currently 
rccoi'ii -.1 (sec Martin 1976. Fowler & Sten- 
ttcit-WilT&on |»7% CeUtnson I9S0) have not 
been recognised in the presently described 
Specjes ;ind il is their absence which separates 
A. cajnii-oinica horn the vaiious members of 
Azoltfl which have 3-24 floats (see Collinson 
|9Kf), lor summary") A, rapricarnica is there- 
fore readily distinguished from the Australian 
Quaternary and extant species of A. filkulouies 
it has a three float uiegosporc apparatus 
(sec Fowler & Stcnnctl-Willson 1978, for 
recent description). 

Comparison with taxa which are considered 
lu possess ,mly a single float-like Structure or 
viM.imeJIak'-ilout. namely, Arolfa ximplex Hal! 

■ .4 prltnaeVtx (Arnold) Penhallow 1955: 
and A mrteana Hills & Weiner I9$5 must 
hi ;ii ! relatively superficial morphologic level 

&USC of the lack of comprehensive strue- 
Urro] information about them. A. simplex is 
distinguished by rlj$ finer reticulum with lumina 
of 2-6 ..m and iruiri at 1 „m wide, and its 
loosely attached float-like structure; A. 
primtwa has a less conical, proximal pcrmal 

eterrc, add s non reticulate distal perine, 
and massulac detail* differ (gluehidia aic 

h narrower)! a. ^eneseatw is distinguished 

by. AmoDgM other ctetatts lis slender glochidia 
with irregularly shaped rips. Also, because of 
thej lack nf structural detail, A. cupricornica 
has not been assigned to Section Simplicispora 
(Hull 1970) which was created to accommo- 
date the above three species. 

Fig, •>. .i SI M i • I ' m I nmcis|i,.,p- nmsMih. ftftflched to megaspore apparatus; 

specimen hdI recoverei I S W [K120K ut WIS ' microspofuftftn "Wing cluster of 

mfcrospoit ina*sul i> m imh ...t.-m rr:i c SEM (Xl72)j nog. np. MIS |3;meRaspor© uppftpa- 

Iny nolo smooth :.pi< flrefl oil COHC; *pccin\ft. n , pc<3 •! <"iiorhidnini ( X 1 000 I with single 

recurved book aU^nrtinti li 17 * 136. e IsoOJaied jBlOcMdlutn, (DIC, >C<55SK$haw- 

mi: evnviriel!i>n*( aI point of I i i i*&ul.i and immedifttely m\ b;isc *»f up, wide 9SM.s, 1^.2 

91.9. 0l37. f St M i E30) nea no. :()'' 1 ' CKaapwre apparatus Wltli 3 m&6«ulae .itr ; .ched; speci- 
men not rceovcred n, SJ M * I 1 ■ i j Mis ii, meg^sporc upparatDn showing MAOQfb epical 
area, eoarsely r«hcuMn« ptrlnc, well Jefined cotUr; specimen ndi recovered. 

Fig. S. ii STM i>s00),nc*p nn_ MIS \2: dftail of COiTui V«a, (cj Willi rcliculntc dtskil pcinif (D) 
and ti;niy perlne Oi proximal OOfl' LP)i*P" '"' recovered b SUM r x5000). ncg. no. MIS 26; 

gtochi^ium wiOi itnchoi ^itfped tip ami mi urved hooks, note cxKisUtelloai at 6940 of tip, speci- 

inrn httQUhed i>' i'U-i-.i,|H)rc ;.pp.irit1i(s, HO 'Cd, c. Sf*.M I >■ I500), ncg no. 200/14*i section Of 

tneg&xpare appttrutixs; 1 M l nusgaspore woll feo) erickmeriAe and (ex> exopcrinc; specimen not rc- 
covctt*0. il. SI M |.-M)), n=g (!(■• rCltl eticulttte ilntal dist:;I purine, showing muii of fuse*!, 

IntertWmed (Uftula«IJk< ' '"' , • 51 M I ■ -' ; --M, nep no. 200/28; mega- 

spore npp«rotua slrowmg vmooih nren jWrmited bv fused hairs at apex (A I; specimen not recovered) 

Rj fi LS Of nettUStiArc BpttaratOR, section en I um thick. b;u- scale 50 am Mega&pore wall CM), col- 
l.tpsc:t and intol.K-it:, .-rutoptrinr (mi ;tn0 exOperiOc {e\) well defined. Randomly developed va- 

cuolae if v >ns,,*v,>\ cone tndleatcj bv ivj; slide 5»8»3/ tO, (2.9 77.2, qijs, 



• - •» (L,/ • i A 


Fig. 7. TEM (X1050) neg. nos 7813, 7814, 7815. LS of megaspore appartus. Megaspore wall (M) 
infolded as in Figure 6, note irregularly shaped, randomly distributed cavities in wall; (en) endo- 
perine; (ex) exoperine; (V) detail of vacuolae developed in proximal perine. 


Collinson (1980) has reviewed the possible 
phylogenetic implications attributed to the 
various structural differences or morphotypes 
(which are manifest in float number, mode of 
attachment of floats; microspore massulae, 
shape of glochidia) within Azolla. She con- 
curred with Hall (1969), and Martin (1976) 
that four lineages may be recognisable within 
the genus. Members of the lineages respec- 
tively possess 1,3,9 and 9+ floats. In this con- 
text it is worthwhile emphasising that A, 
filiculoides and A. capricornica are separable 
on the basis of 'float* number, 3 and l(o?) 
respectively. The term 'float' is used here in 
the sense of Fowler ( 1 975 ) . Usi ng this 
criterion it would be imprudent to suggest 
that A. capricornica was the ancestral form 
of the Australian Quaternary and extant 

Azolla capricornica is presently known in 
oil shale deposits in the Narrows Graben and 
to the northwest in the Dnaringa and Yaamba 
Basins (Foster unpublished). These deposits 
straddle the Tropic of Capricorn, between 22° 

and 24° (for localities see Swarbrick 1974). 
Members of the genus have not been recorded 
from coeval sediments from more southern 
coastal Tertiary basins, or from intracratonic 
sediments of central Australia. On present 
evidence, it seems that Azolla entered Austra- 
lia from more northern and tropical latitudes 
at least during the Middle to Late Eocene. 
Should the hypothesis of northern entry be 
correct, even earlier occurrences of Azolla may 
yet be found in the late Cretaceous of northern 


We draw attention to the value of Azolla 
as an environmental indicator in present day 
environments. It is mostly a warm-temperate 
to tropical genus inhabiting freshwater. Ideal 
growing conditions for Azolla are still water 
where the effects of turbulence and periodic 
flooding will not fragment the colonies. Should 
intermittent turbulence occur, the fern has 
the ability to regenerate rapidly once favour- 
able conditions are restored. The fine grained 
nature of the enclosing sediment at Rundle 
and the high sapropel content of the acid 



Fig. 8. a. TEM (X1000), neg. nos 7895, 7896. 
TS section taken about midway between collar 
and apex of cone. Large randomly developed 
vacuolae (V) of (?) endoperine surrounded by 
denser exoperine. b. TEM (X3000), neg. no. 
7812. LS section of megaspore appartus; (M) 
megaspore wall, (en ) endoperine, (ex ) exo- 

insoluble residue together with the abundance 
of Azolla megaspores, indicates that sedimen- 
tation occurred in a freshwater environment 
with little turbulence and low detrital sediment 

Under such conditions any water flow would 
be slow, and it is likely that mats of Azolla 
colonised surface waters. Such mats would 
largely reduce available light in the water 
column, leading to anaerobic and reducing 
conditions; nevertheless blue-green algae and 
certain other plant life could survive under 
these conditions. Ashton & Walmsley (1976) 
for example, have drawn attention to the 
importance of the blue-green algal symbiont 
{Anabaena azollae) in the life cycle of Azolla 
filiculoides where it is responsible for nitro- 
gen fixation, allowing the fern to colonise 
nitrogen-deficient environments. 

The palaeo-ecological setting proposed is 
restricted to those sections of the Narrows 
Beds which contain Azolla. Nevertheless the 
overall environmental setting is suported by 
petrographic studies of Hutton, et al. (1980) 
who described the dominant maceral of the 
Rundle oil shales as a finely bounded lamella 
alginite (Alginite B) interbedded with matter 
in well-laminated sediments. They used the 
term 'iamosite" for this maceral type and 
further, suggested that it was derived from 
algal mats. All the evidence is consistent 
with Beasley's (1945) suggestion that the beds 
were deposited in shallow and probably per- 
manent lakes. 


We are grateful to Southern Pacific Petro- 
leum N.L. for providing the core material 
for study and approving this publication. C. B. 
Foster publishes with the approval of the 
Under Secretary, Department of Mines, 
Queensland. The skilled assistance of J. V. 
Hardy and staff of the Electron Microscope 
Unit, University of Queensland is gratefully 


Ashton, P. J. & Walmsley, R. D. CI 976) The 
aquatic fern Azolla and its Anabaena symbiont. 
Endeavour 35, 39-43. 

Blasley. A. W. (1945) Ostracods from some 
Queensland Tertiary basins and their bearing 
on the correlation of strata. Proc. R. Soc. Qd 
56, 95-124. 

Collinson, M. E. (1980) A new multiple-floated 
Azolla from the Hocene of Britain with a 
brief reveiew of the genus. Palaeontology 23. 

Duigan, S. L. & Cookson, I. C. (1957) The 
occurrence of Azolla filictdoides L. and 
associated vascular plants in a Quaternary 
deposit in Melbourne. Australia. Proc. R. Soc. 
Vict. 69, 5-13. 

Foster, C. B. (1979) Report on Tertiary mio- 
spores from the Rundle oil shale deposits. 
Qd Govt Min. /. 80, 442-443. 

Fowler, K. (1975) Megaspores and massulae of 
Azolla prisca from the Oligocene of the Isle 
of Wight. Palaeontology 18, 483-507. 



& Stennett-Willson, J. (1978) Sporo- 

derm architecture in modern Azolla. Fern Gaz. 
11, 405-412. 

Hall, J. W. (1969) Studies on fossil Azolla: 
primitive types of megaspores and massulae 
from the Cretaceous. Amer. J. Bot. 56, 1173- 

(1970) A new section of Azolla. Taxon 

19, 302-303. 

Henstridge, D. A. & Missen, D. (1981) The 
geology of the Narrows Graben near Glad- 
stone, Queensland, Australia. 51st ANZAAS 
Conf. Brisbane 1981. (in press). 

Hutton, A. C, Kantsler, A. J., Cook, A. C. & 
McKirdy, D. M. (1980) Organic matter in 

oil shales J. Aust. Petrol. ExpL Assoc. 20, 



C. G. (1970) Geology of the Rockhampton and 

Port Clinton 1 :250 000 Sheet Areas. Rept 

Geol. Surv. Qd 38. 
Lindner, A. W. & Dixon, D. A. (1976) Some 

aspects of the geology of the Rundle oil shale 

deposit, Queensland. J. Aust. Petrol. ExpL 

Assoc. 12, 165-172. 
Martin, A. R. H. (1976) Some structures in 

Azolla megaspores and an anomalous form. 

Rev. Palaeobot. Palynol. 21, 141-169. 
Swarbrick, C. F. J. (1974) Oil shale resources 

of Queensland. Rept GeoL Surv. Qd 83. 


by C. M. Bull & D. R, King 


The distribution of populations of a tick morphologically similar to Aponomma hydrosauri was 
mapped near Albany in Western Australia. This taxon occupies an area of approximately 60 x 40 
km. To the west its boundary appears to coincide with the edge of of the jarrah forest belt. To the 
north and east its distribution abuts that of the more widespread reptile tick. Amblyomma 
albolimbatum. There is no obvious ecotone associated with this parapatric boundary. It is suggested 
that the boundary is maintained by an interaction between the two species. The stability of the 
boundary is not yet known, but one explanation of the present pattern of distribution is that Amb. 
Albolimbatum is gradually expanding its range into areas previously occupied by Ap. Hydrosauri. 


by C, M BULL* & D R KtNC.t 


Bill, C. M. <Sr King, O. R. (I9KU A p;ir.t r»atric boundary between two species of reptile 
lieJB ii. [be Albany arc;.. Western Australia, I runs. A\ Sue. S. Attst. 105 (4), 205 BOft, 11 
necember. 1981. 

The distnbulion of populations of a tick morphologically similar 1o Aponvttunn hy<im- 
sttttri was mapped near Albany in Western Australia. This laxon occupies an area uf 
:t|)pio\iniiitcly M) X 40 km. To the west its boundary appears to coincide wiih the edge of 
the iarrah lores! belt. To the north and cast its distribution abuts that of the mote widespread 
repiile tick, Amblyttmnm nUuihmhutnm There is no obvious ccolonc associated with this 
paiapatric bouiidaiy. It is suggested that the boundary is maintained by an interaction between 
the two species. The stability of the boundary is not yet known, hut one explanation of the 
present pattern Ol' distribution is that Amb. albolimbcllum is gradually expanding its range 
into areas pieviously occupied by A p. hydvosouri. 


Smyth (4973) first described parapalry in 

the tick species, Aponomma hvdruMiiiri, Am 
blvoninut lintlnmun and Atnh. all>o{imhtttmn in 
South Australia. All three species infest the 
same reptile hosts, In any one place only one 
of the three species is found, and their dis- 
tributions abut at common boundaries. Ifi- 
vestigalions have failed to provide a satisfac- 
tory explanation of the mechanisms which 
prevent range overlap at the boundaries (Bull 
& Smyth 1973, Sharrad 1 , Bull, Sbarrad & 
Pctney I9SI). Smvlh (1973) suggested two 
hypotheses. One hypothesis is that the boun- 
daries follow environmental ecotones, and 
that at least one of the contacting species K 
poorly adapted lor conditions across the boun- 
dary. This hypothesis was supported by the 
results of a detailed study near Ml Mary. 
South Australia, where the boundary between 
Ap. hydrosauri and Amb. Hwhatittn coincides 
with the sharp vegetattonal change from 
niallec scrub to open woodland; however trans- 
plant experiments 1 - have failed to demon- 
strate reduced fitness of either species across 
the ecotone (Bull et al. 1981), At other 
boundaries hclween pairs of these species 
environmental changes arc less obvious (Smyth 
1973. Bull et at. 1981), 

The second hypothesis proposes thai para- 
patric boundaries arc maintained by ecological 

interactions between the tick species, resulting 
in the exclusion of one of them from the 
range of the other. The outcome of the iniet- 
aclion may be reversed where there js an 
environmental change, such as at an ecotone. 
I bus stable boundaries will form at ecotones, 
hut boundaries will be established also 
wherever the ranges of Iwo of the species con- 
tact, independently ol the environmental 
conditions. A prediction is that where the 
ranges of two species meet in tit her areas 
parapatry would be cxpecled also 

fn southwestern Australia, Atnb. albaltm- 
hiihtm is the common reptile tick but another 
species, morphologically similar to Ap. hydro- 
sattvi, was found hy CM.B. near Albany in 
southern Western Australia (Smyth I97 J . ) 
The tavonomic status of this population i- 
uuder investigation but here it will be referred 
to as "Ap. hydrosami". Sharrad 1 and Sharrad 
ei King (1981) confirmed its presence in at 
least four small isolated areas along the south 
coast of Western Australia. Thetr collections 
were sufficiently detailed to show that there 
\\a\ a parapatric boundary between Amb 
aihohmbatiim and Ap. hydrosauri at Cape 
Naturalistc (Sharrad & King ]98l)< which 
may coincide with a vcgetalional ccotunc 
( Bridgewater & Zarnmit 1979). We describe 

1 School of Biological Sciences. Fhndcts Univer- 
sity of South Australia, Bedford Park, S. AuM. 

f Ai'iicuhuie Protection Board, rorresttield. \VA. 

'Shunad. R. D. (1980) Studies of Ihe foojore 

which determine the distribution*- of Hi tec 
species of Smith Australian ticks. Ph.D. Thesis, 
University of Adelaide. limpnh|ishcd ). 
- Pelney, T. N. (1981) The interaction of two 
parapatric tick species with their off liosl en- 
vironment. .PhT> Thesis. Flinders University, 
l unpublished). 



Uui h> . 

1 IT' 30' 

118 E 
Stirling Fi- 

ll 8 U 30 




34 n 30 

■\\ Jar rah m 


Ti Forest (Li||||ii£B 


o " r 



t?° ° a , if ,Pia . • . 

/- 1 



35 J S 

Fig. li Distribution of Aponommu hydrosauri I open squares and Amblyomma alkoUmbatum (closed 
circles) in the study area. Lizards with both species attached are shown with open circles, and 
those with no ticks attached, by open triangles. 

investigations of the prediction that there is 
parapatry between the species near Albany. 

Materials and Methods 

Earlier records from a general survey of 
reptile ticks in Western Australia (Sharrad & 
King 1981 ) were supplemented with additional 
data collected in 1979 and 1980, and a de- 
tailed survey made in October 1980 of the 
area around Albany (Figure 1 ). Lizards, 
mainly Tracliydosaurus rugosus, were captured 
as they crossed the roads or basked on the 
roadsides, and were examined for attached 

Only Ap. hydrosauri and Amb, albolim- 
batum were found on T. rugosns. Adult ticks 
but not larvae and nymphs could be easily 
identified in the field and the immature stages 
were removed for later identification. A simple 
distinguishing feature is that Amhlyomnui 
species have eyes but Aptmomma species 
lack them (Roberts 1970). 

Other data came from road-killed hosts- 
Individuals of T. rugosus are commonly run 
over by vehicles, and some ticks remain 
attached to their dead host for several days 
(unpubl. observ.). 

Other reptile species were examined when 
encountered. One blue tongue skink (TUiqua 
occipitalis) and six freshly-killed brown snakes 
(Pseudonaja nuchalis) were examined, but had 
no ticks attached. Two snakes (no identifica- 
tion provided) and six goannas (all Varanus 
rpsenbergi) from the area previously had been 
found infested by the tick species Aponommu 


The distributions in the study area of the 
tick species. Amb. albolimbafttm and Ap. hy- 
drosauri, attached to 7*. rugosus are shown in 
Figure I. The range of Ap. hydrosauri is 60 
km (north-south) X 40 km (east-west). To 
the north and east its distribution abuts that 



of Amb. alboi'unbainm which is widely distri- 
hutcd in southern Western Australia (Sharrad 
& King 1981). Amb. albolimhautm was noi 
found within (he distribution of Ap hydro- 
Kauri, except in u narrow boundary zone 
(Figure I). where both species were found 
together. Five host individuals collected in 
the boundary zone had both tick speeic> 
attached lo them. The width of the overlap 
zone is not known, but at the northern boun- 
dary there were less- than lf> km separating 
hosts with only A p. hydrosauri from hosts 
with only Amb. albnlimhafttm. 

lo the west, the distribution of Ap. hydro- 
xauri abuts with the edge of the uncleared 
jarrah forest ( fcucalxptus marxitwta) . Road- 
killed T. rw^nvwv were found regularly owr 
most of the study area, but on the roads 
through the jarrah forest neither live nor dead 
T. ntgosus were seen. Seven live 7\ rugasux, 
captured on the southeastern edge of the jarrah 
belt in Country still containing extensive un- 
cleared areas of forest, had no ticks attached 
to them. In the rest of the study area only two 
other uninfected T Tttjjotttl were found 
(Figure ! ). 

Amb. albolimbatum Ls distributed along 
the south coast to the east and west of Albany 
(Figure I), and abuts with, and may just 
overlap, Ap. hyttroxtwrf about 6 km west and 
about 3 km east of Albany. 


It »y not clear which factors prevent Ap. 
iiyitrosanri from spreading beyond its very 
narrow range around Albany. The morphologi- 
cally similar species in South Australia occupies 
a wide range of environmental conditions 
(Smyth 1973) One environmental change 
which seems to have an important influence 
on the distribution of Ap. hy drosawi near 
Albany is the Jarrah iorcst to the west. The 
density of T. rut>osus appears ta be lower Lhere. 
and those found on the margins of the forest 
had no ticks on them- This suggests lhal some 
characteristic of the forest makes it unsuitable 
for occupancy by ticks 

The northern and eastern boundaries do 
not follow macrocjirnatic elines as does the 
boundary between Ap. hydrosauri and Amb. 
hmbaium in South Australia (Smyth 1973). 
Near Albany ihc tick boundary crosses rain- 
fall isoclines (Sharrad & King' 1981). More- 
over, Ap hydrnsauri is found in Western 

Australia in areas receiving higher rainfall, 
such as south of Cape Naturaliste, and in 
-ires receiving lower rainfall, such as Bremer 
Bay (Sharrad & King 1981) 

I here are no obvious vcgctational changes 
at the northern and eastern boundaries, Much 
of the area has been elearcd for farming, but 
wide roadside verges still maintain native 
vegetation classified as malice heath (Beard 
1976). There may be a subtle change, bur if so 
it is not significant enough to show on the 
vegetation maps of the area (Beard 1976). 
In fact, Ap. hydrosmiri does survive in the 
distinctly different and less mesic coastal scrub 
on drift sand dunes at Bremer Bay. (The 
Cape Riche and Bremer systems of Beard 

Topographic barriers are also unlikely lo ht 
important in maintaining the boundary. The 
area is dominated by two mountain ranges. 
the MiiaU Poroitgorup Ranges starting about 
35 km north of Albany, and the more exten- 
sive Stirling Ranges starting about &S km 
north of Albany (Figure 1 ) These ranges 
rise abruptly from the plains and there is a riat 
mtermontanc area, about 20 km wide, between 
them. At one point the boundary i? located 
on this plain- The Woogenellup Road runs 
northeast from Ml Barker and along the 
southern edge of the Stirling Ranges; only 
Amb. afbolhnbiitum is found along it. The 
Porongorup Road runs east from Ml Barker 
along the northern edge of the Porongorup 
Kangcs; most lizards found along this road 
carried Ap. hydrouniri Thus the boundary 
region must occur on the plain in between 
these roads, and the mountain ranges are not 
barriers to the extension of the range of either 
species. There are no topographic features qf 
any significance along the eastern boundary 
of Ap hvdro\attrL 

An alternative explanation is that the ptefc 
lion of the boundary is independent of environ- 
mental gradients, but is maintained by an 
interaction between the species. The strength 
of this hypothesis is the lack of any other 
obvious options, although attempts to demon- 
strate interactions of ticks at other parapatric 
boundaries have not been successful (Bull et 
ai 19*1 >. Nevertheless, the frequent occur- 
rence* of parapatric boundaries within this 
group of ticks (Bull ei id. 1°81) is circum- 
stantial evidence in favour of the interaction 


C. M. BULI & D. R. KING 

Perhaps Ap. hydrosauri was once distributed 
continuously from South Australia to southern 
Western Australia, but Amb. alholimhatwn has 
subsequently spread and displaced it, to leave 
geographically isolated populations in eastern 
South Australia, in the southern part of Eyre 
Peninsula, and in a number of locations along 
the southern coast of Western Australia. 


S. Habel, R. D. Sharrad and J. D. Roberts 
helped with field collecting and R. H. Andrews, 
T. N . Petney and R. D. Sharrad provided 
useful comments. Field trip funding came 
partly from the Flinders University Research 
Budget, and partly from an ARGC grant for 
analysis of genetic variation in reptile ticks. 


Beard, J. S. (1976) The vegetation of the Newde- 
gate and Bremer Bay areas of Western Austra- 
lia (2nd ed.). (Vegmap Publications: Perth). 

Bridgewater, P. B. & Zammtt, C. A. (1979). 
Phytosociology of S.W. Australian limestone 
heaths. Phytocoenologia 6, 327-43. 

Bull, C. M., Sharrad, R. D. & Petney, T. N. 
(1981) Parapatric boundaries between Austra- 
lian reptile ticks. Proc. Ecol. Soc. Aitst. (in 

& Smyth, M. (1973) The distribution of 

three species of reptile ticks, Aponomma hydro- 
sauri ( Denny ) , A mhlyomma athotimhatum 

Neumann, and Amb. limbatum Neumann. IT. 
Water balance of nymphs and adults in rela- 
tion to distribution. Aust. J. Zoof. 21, 103-10. 

Roberts. F. H. S. (1970) "Australian Ticks". 
C.S.I.R.O.: Melbourne). 

Sharrad, R. D. & King, D. R. (1981). The geo- 
graphical distribution of reptile ticks in Wes- 
tern Australia. Aust. J. Zool. (in press). 

Smyth. M. (1973). The distribution of three 
species of reptile ticks, Aponomma hydrosauri 
(Denny), Amblyomma albolimbatum Neumann. 
and Amb. limbatum Neumann. I. Distribution 
and hosts. Ibid. 21, 91-101. 


by Christopher C VonDer Borch 


The coastal plain of southeastern Australia has been the site for formation of a variety of carbonate 
minerals throughout the Quaternary. Dolomite and prodolomite (approximate chemical composition 
of dolomite but lacking in ordered reflections), magnesite, hydromagnesite, magnesian calcite and 
aragonite are presently forming in Holocene ephemeral lakes on portion of the coastal plain which 
is in close proximity to a modem coastal lagoon known as the Coorong. Radiocarbon dating has 
verified a modern age for at least some of these coastal dolomite occurrences. Pleistocene dolomitic 
carbonates have been detected in sediments from many interdunal depressions which lie between 
Pleistocene barrier islands remote from the present coast. These areas have been stranded by a 
combination of slow regional upwarp combined with eustatic sealevel fluctuations. 




Ihe coastal plain of southeastern South Aus- 
tralia has been the site fc* for/nation ot a variety 
ol carbonate minerals throughout the Quaternary. 
Dolomite and protodolomile (approximate ehcm> 
eal composition of dolomite bur lacking in ordered 
reflections), mugncsilc. hydromaimesite, m&Sut&Jftn 
cajcitc and aragonite arc presently forming in 
Hnlocene ephemeral lakes on iiortion of the 
coastal plain which is in close proximity to a 
modern coastal lagoon known as the Coorong'- . 
R;<. luteal hop dating hay verified a modern age for 
at least some of These consul dolomite oecur- 
tences". Pleistocene dolomitic carbonates have 
been detected in sediments from many interdunal 
depictions which lie between Pleistocene buoir-r 
islands remote from the present coasts Thcae 
areas have been stranded by a combination of 
slow regional upwarp combined with euslatic sea- 
level nucliialions 1 - 1 ". 

■ he OCCUirdlce Of the majority uf active dole- 
milc-forming lakes imrnediuLely adjacent lo the 
pteseni coastal lapoon suggested that proximity 
of n marine environment is a prerequisite for 
Coorony type dolomite formation, It Was con- 
sidered ibid the bulk fjf Ihe nccess;uy [oftS could 
be derived from a nearby marine reservoir, either 
from wind-borne cyclic salt- or by entrapment of 
acawater in coastal groundwater circulation, Sub- 
sequent hvdroloejcal and Mralijuraphic studies, 
however, h.ive east doubt on the necessity of a 
marine ionic source 11 - 4 . To resolve the enigma. 
which has an important bearing on theories of 
dolomite cent-sis in the Coo -on e reeaon, strait- 
"u.phie studies were initiated on Lake Ormerod, 
an inland, ephermeral, carbonate lake. 

A series of shallow, ephemeral, carbonate lakrs 
including I -ake Ormerod occurs within a broad 
interdunal depression west of the town of Nara- 
eooiie The lakes arc situated approximately 85 
km from the present coast at an elevation of 50 
m above present svalevcl. Based on palaeomannetic 
da ling" 1 this depression was ar least in parr a 
coastal tiuioon about 600,000-700.000 years ago, 
but has since been separated from a marine 
environment by the regional upwarping and conse- 
qenl regression, 

In most aspects the inland lakes appear identical 
to their more numerous coastal counterparts. They 
arc cmphcmeial in nature, filling with water dur- 
ing winter months by groundwater seepage and 
direct rainfall, and evaporating to dryness during 
ensuing arid summer-,, When full of water they 
are about I m deep and like their coastal counter- 
parts support an abundant agnatic biota, including 
Ihe aquatic grass Ruppia marltima and the Mirall 
gaslrojjod Om/W/u a'ttfusu. Some of the Jakes 
are floored with fine-grained (U.2-UO dolomite 
mud. Ihe purpose of this reconnaissance study Fs 
lu deline the age of the dolomite in these jnlurid 
lakes usiny radiocarbon dating, in order to deter- 
mine whether it could be forming tit present or 
whether it is relict from times when a Pleistocene 
murine shoreline existed in the area. 

A J m sediment core collected I'rom Lake 
Ormerod comprises an uppermost 50 cfn unit of 
white, tine-grained carbonate, shown by x-ray 
diffraction to be protodolomile (dolomite unit), 
overlying a dark grey organic-rich rnud composed 
oi calcile and subsidiary dolomite (calcile anil). 
Assuming that the dolomite is a primary form, 
this vertical mineralogical variation implies an 
environmental change from a pre-existing per- 
manent swamp oi lake to the present alkaline 
ephemeral lake, posibly in response to a decrease 
in rainfall durrny the past few thousand years'. 

t "wo samples from die top and bottom of the 
dolomite unit were selected for reconnaissance 
ladioearbon dating The uppermost shows a U C 
ape ot 1.300 "^ 60 years whiNt the lowermost is 
5770 =£ yo years (Table l>, proving that this 
lake at its surface contains modern and not Pleis- 
tocene dolomite, The age of the surface mud 
should be considered a maximum one. This 
anomalously old age could be due to a variety 
of factors, including dilution by reworking of oldeT 
dolomite or formation from an admixture of "old" 
cartoon derived by groundwater leaching of the 
underlying Pleistocene and Tertiary carbonate 
aquifers. The possibility also exists of a sluw 
reaction iav, ot i T><;- OCtt&l ot thousands Of vears, 
in the 'penccomemporaneous" formation of dolo- 
mite in such a situation. 

Table 1- Data tabulation py- Lake Ormerod carbonate sample* from rmvHftaissnttce hpn'hrtic. 


depth (cm) 




age. years BP. 



— J.O *fc 0.2 

—3.4 * 0.2 

48.S ± 0.5 

-149.8 ± fi.4 
-512.2 = 5.2 

1300 ± 60 
5770 ± 90 


The modern age of dolomites in Lake Ormerod aquifer, and that proximity to a marine environ- 

suggests that this mineral was formed a con- 
siderable time after sealevel regressed from the 
region. This implies that it must have formed 
from seasonally discharging Ca and Mg bicarbon- 
ate groundwaters of the unconfirmed regional 

ment is not a necessary pre-requisite for genesis of 
this type of dolomite. 

Radiocarbon dates were made at the ANU 
Radiocarbon Laboratory by Henry Polach. 

i Alderman, A. R. & Skinner, H. C. W. (1957). 

Am. J. Sci. 255, 561-67. 
-'Skinner, H. C. W. (1963). Ibid. 261, 449-72. 
;:Von Der Borch, C. C, (1965). Geochim. Cosmo- 

mochim. Acta 29, 781-99. 

4 (1976). J. Sedim. Petrol. 46, 952-66. 

5 & Lock, D. E. (1979). Sedimentology 26, 


o Rubin, M., & Skinner, B. J. (1954). Am. J. 

Sci. 263. 1116-8. 
-Hossfeld, P. S. (1950). Trans. R. Soc. S. Aust. 
73, 232-79. 

N Sprigg, R. C. (1952). Geol. Surv. S. Aust. Bull. 


•Cook, P. J., Colwell, J. B., Firman, J. B., 
Lindsay, J. M., Schwebel, D. A. & Von der 
Borcb, C. C. (1977). Bur. Min. Resour. Aust. 
Geology & Geophysics 2, 81-8. 

< f, Idnurm, M. & Cook, P. J. (1980). Nature 
(2867), 699-702. 

ii Von der Borch, C. C, Lock, D. E., & Schwebel, 

D. (1975). Geology. May, 283-5. 

Christopher C. Von der Borch, School of Earth 
Bedford Park, S. Aust. 5042. 

Sciences, Flinders University of South Australia, 


by P. 7. Kailola and G. K. Jones 


Promicrops lanceolatus (Queensland Grouper or Sea Bass) is widespread through the tropical 
waters of the Indo-Pacific region, inhabiting lee and seaward sides of reefs and lagoons, as well as 
estuaries. The limits of its distribution have been reported as Algoa Bay (East Africa), India, 
Taiwan, Gilbert Islands, Fiji and Hawaii. In Australia it has been recorded from northern N.S.W., 
Queensland, Western Australia and "presumably N.T." 




Prornicrops Uinceolatus (Queensland Grouper 
or Sea Bass) is widespread through the tropical 
waters of the Indo-Pacific region, inhabiting lee 
and seaward sides of reefs and lagoons, as well as 
estuaries 1 *-. The limits of its distribution have 
been reported as Algoa Bay (East Africa), India, 
Taiwan, Gilbert Islands, Fiji and Hawaii 3 " 4 . In 
Australia it has been recorded from northern 
N.S.W., Queensland, Western Australia and "pre- 
sumably N.T." r, -°. 

This account extends the southern distribution 
of the species and its presence in South Australia 
waters adds further support for a tropically- 
derived current occurring in southern Australian 

On 28. ix. 1980, a fresh specimen of P. lance- 
olatus was washed up on the ocean beach of The 
Coorong (approx. 36°00'S, 139°30'E). It was 
taken to the South Australian Museum in Adelaide 
and a cast was prepared for the Museum. The 
specimen was retained there to recover its skele- 

A description of the fish follows: 

Family Serranidae. Prornicrops lanceolatus 
Holocentrus lanceolatus Bloch 1790, Nat.. 

Ausland. Fische 4, p. 92, pi. 242, fig. 1 

(reference copied), 
S. Aust. Mus. cast, F.4562. Total length 

(TL) 2120 mm; standard length (SL) 1775 

mm; Weight (fresh, intact) 153 kg. 

D.Xr, 16. A.III,8. P.ii,15,i. L.Iat. about 61. 
Tr. 14 or 15 + 43 or 44. G.R. 8 + 1+8. The low 

rakers anteriorly are difficult to distinguish from 
the rough platelets covering the gill arches. (Most 
authors credit P. lanceolatus with a gill raker 
count of 10+14+16. The anomaly here is remark- 
able. Randall 7 found a lower-than-normal gill 
raker count for a 574 mm specimen of Serranus 
fuscoguttaius (Forsskal) and surmised that there 
is a loss of gill rakers in larger fish (Serranidae) 
because of fusion). 

An irregular double series of stout teeth along 
each jaw is succeeded by bands of curved sharp 
teeth; bands of teeth also occur on vomer and 
palatines. The maxillary extends well past the eye, 
which is contained 3.7 times in the broad convex 
interorbital space. The middle opercular spine is 
slightly nearer the lower spine than to the npper. 
The 8th is the longest dorsal fin spine. The pre- 
opsrculum is highly angular; soft fins are 

Body colouration was dull dark brown, the fins 
somewhat darker and charcoal at their margins. 

Gut contents were sand and grit, with several 
teleost vertebrae near the end of the intestines. 
Gonads could not be located. 

Prornicrops lanceolatus has been reported to 
grow to nearly 4 m long 1 ' 4 -' 5 , and Schultz? has 
investigated the authenticity of records of large- 
sized Prornicrops. Specimens above 200 kg are 
known from the Hooghly River, India (TL 2.3 m, 
209 kg)*; Thailand (TL 1.8 m, 216 kg)"; Bikini 
Atoll, Micronesia (TL 2.3 m, 214 kg; photograph 
at U.S. National Museum). The largest Queens- 
land specimen reported weighed 288 kg 10 ; Four- 

Fig. 1. Queensland Grouper Prornicrops lanceolatus. Total length 2120 mm. 


manoir & Laboute- state that Pramtcrops can 
reach 400 kg. 

The South Australian specimen is believed to 
be the most southerly record for the species. 
Algon Bay (approx. 3<rOO'S, 26 OOF) was the 
previous southerly record and this location is 
close t" the southern-most limit of the tropical 
"Agulhus Current" which extends down the east 
const of South Africa. Similarly, its discovery here 
can be explained by the presence of a current 
system originating from the tropical waters of the 
Indian Ocean. This current (the "Leeuwin Cur- 
rent") has been described 1 -- 1 ' 1 as an eastward 
flowing current during autumn and winter months 
along the southern coast of Western Australia and 
across the Great Australian Bight. Satellite obser- 
vations of sea surface temperature fronts 1 "' show 
that this current of relatively warm, low salinity 
water flows completely across the Great Australian 
Bight, It is also interesting to note that Markina 1 ' 1 
classified the phylo- and zooplanton in the Great 
Australian Bight as of tropical origin. 

We lhank Messrs. D. Wankie, A. Billing, D> 
bivan* and D. Roberts who brought the fish to our 
attention and assisted with its transportation and 
measurements; a^o to Messrs. R. O. Ruchle and 
W. Head for the photograph and museum assis 

Table 1. Measurements (in mm) of P. lanceolatus 

(obtained jmm the fresh specimen). 

Total length 2120 

Standard length 1775 

Head length 730 

Body depth 670 

Snoiu length 170 

Eye diameter 5 I 

Interorbital width 190 

Lower jaw tip to vent 1320 

Maxillary length 390 

Head depth 435 

Caudal peduncle depth 215 

Caudal peduncle length 345 

Periorbital head length 510 

Longest pectoral ray 360 

Longest ventral ray 285 

Longest anal ray 280 

Third anal spine 1 11 

Longest dorsal ray 250 

Eighth dorsal spine 99 

First dorsal spine 49 

Last dorsal spine 80 

Longest gill filament 69 

Gill raker at angle 25 

'Roughly, T. C. (1951). 'Fish and Fisheries of 

Australia/ 1 (Angus & Robertson: Sydney). 
-Foiirmanoir, P. & Laboute, P. (1976). Toissons 

dc Nouvelle Oaledonie et des Nouvelles 

Hebrides." (Lcs Lditions du Pacifique: Tahiti). 
•ScbullA L P. (1966). Addenda, p.p. 145-165. 

///: Sehultz, L, P. & collaborators. U.S. natn. Mus. 

Bull. 202, 3. 
ISmith, J* L. B. (1965). "The Sea Fishes of 

Southern Africa." 5th edln (Central News 

Agency; South Africa). 
•McCulIoch, A. R. (1929). Mem. Aust. Mas. 

512)' 145-^29 
"Munro, I. S. R. (1961). Fish. News). 20(8> : 


7Randall, J. E. (1964). Pacific Sci. 18(3): 281-96. 
*A!coek, A. M905). Nature 71: 415. 
"Smith, H. (1933). T Siam Soc. Nat. Hist. 
Suppl. 9(1): 85. 
^Gnmt, E. M. (1978). "Guide to Fishes" (4th 

ed.) Dcpt Harbours & Marine, Qld. 
"Darbyshire, J, (1964). Deep-Sea Res. 11: 781- 

1 2Rochford, D. J. M975L Proc. Ecol. Soc. Aust. 

8: 57-83. 
"•tPashkin, V. N. (1968). Tr. Atl. Nauchno-Issled. 

Inst. Rybu. Khoz. Okeanogr. 28 (transl.). 
"Crcsswell, G. R. & Golding, T. J. (1980). Deep- 

Sea Rts. 27A: 449-66. 
i:, LegcckJS, R, & CressweU, G. (in press). J bid. 
T'Markina, X. P. f)976>. Oceanology 15; 602-4. 

P. I Kailoi-A and G. K. Jones. Department of Fisheries, 25 Grenfell Street. Adelaide, S. Ausl. 5000. 


by Robert T. Langeand Timothy Reynolds 


Circular zones of suppressed growth, called halos, surround individuals and clumps of some plants, 
particularly trees such as Casuarina cristata, in South Australian vegetation. These halos extend well 
beyond the canopy spreads of the plants at their foci and show up clearly on aerial photographs. 
Suggested explanations of the suppression include water shortage induced by the central plant, 
allelopathy, or grazing and seed-harvesting animals harboured by the central plant, but field 
observations indicate that none of these provides a satisfactory explanation on its own. 




I ifVuhir BCMP8 of suppressed growth, called 
hnlos 1 *", surround individuals and clumps of some 
pl;iiit\. particularly trees such as Cusuarina 
t rixtdta, in South Ajsiralian vegetation. These 
halos extend well beyond ihc canopy .spreads of 
the plants ui their loci and show up clearly on 
aerial photographs, Suggested explanations of I lis 
siippie\xion include w;iter shortage induced by (he 
central plant, allelopathy, or grazing and seed hat - 
vesting animals harboured by the central plant, but 
held observations indicate thai none of these pro- 
vides a satisfactory explanation on its own-. 

/ones ot enhanced growth also occur, pailicu- 
larly under trees such as Aaic'ut papytocarpa and 
shrubs such as Maircana sedifolia. These zones 
usually do not spread beyond the canopies of their 
producers, and do not show up on aerial photo- 
graphs, but are obvious to the ground observer. 
Suggested explanations for the enhancement in- 
clude soil* and microclimate 1 conditions induced 
by the central plant and, under intricate shrubs, 
protection from the grazing of large vertebrates 4 , 
yielding contrasts with unprotected interspaces. In 
general field observations support the view that 
these are very plausible hypotheses, but Ihere are 
complications, b*»r instance. A. papyrocarpa cano- 
pies in the Whyalla region usually enhance the 
likelihood of finding species beneath them, and 
lack an cxIc.tJed grass suppression halo, hut 
sometimes the reverse is observed. Both categories 
of zones are well known elsewhere 1 *'". The most 
recent Australian publication about them concerns 
I iicuhpmx htntert suppression zones in Vieionart 
Irec-luath 1 ". 

We have detected a fun he category of such 
plant centred cffcOtS which, so ~"ar as wc can estab 
lish, is unreported, It differs from the preceding in 
thai it is not expressed as overt genet al suppression 
or enhancement of plant growth around particular 
tree* or shrubs, and cJinnot be proved by inspect- 
ing any single tree or shrub for halo effects. The 
elfcct is proved only by considering die species 
population collectively and consists of tendencies, 
within small circles centred on individuals, for 
halos of shifts in Ooristie composition of suuound- 
ing veeeiation. i.e. for shifts in the probabilities of 
incidence of other species, from those applying 
within small circles centred on individuals of a 
species to different probabilities applying in the 
interspaces between the circles, T his note sum- 
maizes our investigation and its findings. 

Traverses in native vegetation were marked out 
separately between individuals of each species 
under lest for halo-elfect. Our technique required 
successive individuals to be separated by a least 
twice then canopy diameter. 

Cuele* of radius n, r± — r tu tr in the range 
ft3-3-0 in) were marked oul around each indivi- 

dual of a test species and searched for incidence 
of all other species. This was repeated at a ran- 
dom point in each midlocation between test indi- 
viduals. Data for each quadrat size separately were 
assembled as 2 '>; 2 homogeneity tables for each 
combination of test and orher species in turn, and 
tested for significance by v- or fishers exact 
lest", using a computer. Results were graphed as 

3 J 

E. microcarpa 

• A 


1.0 1.5 2-0 

Fig. I HjlO signatures of Eucalyptus microcarpa 
and /:'. leiwoxylnn in native vegetation at Belair. 
individuals of these species were intermingled 
is tree cover over dense, low sclerophyllous 
icnih. 3 = the significance level of p ^J 0.001. 
-b and — signify positive and negative associa- 
tions respectively. A = Acroiriche scrrulata, B 
Gtmocarpus elatus, C — Olea eutopca. 


in Figure 1 to display the "signature" of the halo 
effect. We also examined association patterns aris- 
ing from 4 m square quadrats distributed randomly 
through the same vegetation. 

We applied these techniques in local relict native 
vegetation of the sort that the National Parks and 
Wildlife Act aims to preserve. Findings were as 

( 1 ) Various native species had halo effects ex- 
pressed only as shifts in the probability of inci- 
dence of other species within the halo of the test 
species. These shifts involved both increased and 
decreased probabilities simultaneously. Various 
other species lacked such effects. 

(2) Most of these effects were not detected by 
large quadrats scattered randomly in the vegeta- 
tion, which explains why they seem to have been 
overlooked in earlier plant sociological studies. 

(3) Some halo-producing species intermingled in 
the one habitat have equivalent habit (e.g. Euca- 
lyptus microcarpa and E. Icucoxylon in layered 
woodland at Belair) but have different signatures 
(Figure 1 ) , indicating species-specificity beyond 
single-cause explanations such as bulk inhibition 
or enhancement by waterstress or shade. 

(4) Perennial weeds, which are a main cause of 
degradation in heritage vegetation, are caught up 
in these effects. In woodland near Belair, Aspara- 
gus asparagoidcs (bridal creeper) was positively 
associated with small clumps but not separated in- 
dividuals of Exocarpus cupressiformis. Rhamnus 
alaternus (buckthorn) and Olea europea (olive) 
seedlings were highly associated with the halo of 
Eucalyptus leucoxylon (blue gum) (Figure 2), 
but not of E. viminalis (manna gum), E. camal- 
dulensis (river gum), E. fasciculosa (pink gum) 
or E, microcarpa (peppermint). To that extent 
blue gum represents a "weak link" in such heritage 
vegetation, and the replacement of some foothills 

E. leucoxylon 



3 H 
1 - 


- 1 — 



— I — 



Fig. 2. Halo effect of Eucalyptus leucoxylon on 
Olea (C) and Rhamnus (D) in native vegetation 
at Black Hill. Other legend as in Figure 1. 

native woodland by olive woodland probably was 
fostered by this link, for which no explanation is 
available. Once established, olive harbours its own 

(5) There is evidence that some native sub-shrubs 
interact negatively with some weeds. This encour- 
ages the view that thorough screening might reveal 
natives with halo suppression of weeds sufficient to 
warrant their evaluation for reclamatory and buf- 
fer planting. 

In conclusion, we think that this evidence points 
to a neglected but important feature of patterning 
in native vegetation, with scope for practical appli- 
cations, and which demands the fomulation of 
convincing explanations. 

1 Andrew, M. H. (1978). The initial impact of 
depasturing sheep on arid chenopod shrublands. 
Ph.D. Thesis, University of Adelaide. 

-Johnson, A. W. (1978). Vegetation suppression 
in the Whyalla region, with particular reference 
to the phenomena around Casuarina cristata 
Miq. B.Sc. (Hons ) Thesis, Department of 
Botany, University of Adelaide. 

3 CorrelI, R. L. (1966). Studies on the nitrogen 
economy of semiarid vegetation at Yudnapinna 
Station, South Australia M.Sc. Thesis, University 
of Adelaide. 

4 Barker, S. (1972). Effects of sheep stocking on 
the population structure of arid shrublands in 

South Australia. Ph.D. Thesis, University of 

•>Muller, C. H. (1953). Amer. J. Bot. 40, 53-60. 

'HVluller, W. H. & Mullen C. H. (1956). ibid, 43, 

7Jaksic, F. M. & Fuentes, E. R. (1980). J. Ecol. 
68, 665-70. 

^Storey, R. (1967). Aust. J. Bot. 15, 175-87. 

9 Nat. Acad. Sci. (1971). "Biochemical Interactions 
among plants." Environmental Physiology Sub- 
Committee. (Washington D.C.). 

"'Del Moral, R. Willis, R. & Ashton, D. (1978). 
Aust. J. Bot. 26, 203-20. 

"Sokal, R. R. & Rohlf, F. (1969). Biometry. 
W. H. Freeman Co., N.Y. 

Robert T. Lange and Timothy Reynolds, Department of Botany, University of Adelaide, Box 498, 
G.P.O., Adelaide, S. Aust. 5001. 


by Julia Smith and Terry D. Schwaner 


Two Amphibolurus nullarbor Badham were collected 20 km E of Nullarbor Homestead, S.A. 
(31°28'S, 131°12'E), amongst bluebush (Maireana sedifolia) on the Nullarbor Plain (above the 
cliffs) by S. Doyle on 30.viii.1980. Abdomens of both females were distended and oviducal eggs 
were easily palpable. One specimen (S.A.Mus. R18170, SVL=140 mm) was preserved; dissection 
revealed six shelled eggs in the right oviduct and eight in the left. Because reproduction in this 
species has not been reported, the other lizard (SAM R18581, SVL=135 mm) was kept alive until 
parturition to document clutch size, egg sizes during incubation, hatching times and hatching sizes 
under laboratory conditions. 





Two Amphibolurus nullabor Badham were col- 
lected 20 km E of Nullarbor Homestead, S.A.. 
(3 1 °28'S, 1 3 1° 1 2'E) , amongst bluebush (Mai- 
reana sedifolia) on the Nullarbor Plain (above the 
cliffs) by S. Doyle on 30.viii.l980. Abdomens of 
both females were distended and oviducal eggs 
were easily palpable, One specimen (S.A.Mus, 
R] 8170, SVL— 140 mm) was preserved; dissec- 
tion revealed six shelled eggs in the right oviduct 
and eight in the left. Because reproduction in this 
species has not been reported, the other lizard 
(SAM R18581, SVL=135 mm) was kept alive 
until parturition to document clutch size, egg 
sizes during incubation, hatching times and 
hatching sizes under laboratory conditions. 

On 1-2. x. 80, 12 eggs were found scattered in 
the vivarium enclosure. Each egg was measured, 
marked for identification, and placed on damp 
aquarium gravel in a clear plastic container 
loosely covered with plastic wrap. The container 
was placed on top of a refrigerator, near the back, 
where the temperature was 27-29°C 1 * 3 . Water 
was sprayed on the eggs weekly to prevent dessica- 
tion. Three eggs became mouldy within the first 
month of incubation and were discarded. Two 
additional eggs were laid on 8.x. 80; these were 
preserved in formalin (R18581~eggs). 

Egg sizes measured at different times during 
incubation indicated an average increase in egg 
volume of 83% (computed as the volume of an 
ellipsoid from differences between initial and 
maximum egg sizes, Table 1 ) . Most eggs de- 
creased slightly in length and/or width just prior 
to hatching (compare size data for 11/13 and 
12/14, Table 1). 

Hatchlings emerged 18-27.xii.80, after 79-80 
days incubation. Neonates remained in the eggs 
for 2-16 hr with only their heads protruding. 
Three lizards left the eggs with parts of the yolk 
sac still visible but the yolk was absorbed 2-3 
hr after full emergence. Hatchlings ranged 33.6- 
37.3 mm (X=35.9±l,l mm) SVL and 75.6- 
87.5 mm (X=81.9±4.3 mm) total length. There 
was no significant correlation (r=.04, n— 9) be- 
tween hatching SVL and maximum egg volume 
(computed as before from Table 1). Colour and 
pattern of the young (Fig. 1) were similar to 
that of adults 4 " 5 . 

Fig. 1. Hatchling Amphibolurus nullarbor, SVL- 
37.3 mm. 

Table I. ffgg sizes (length and width in mm), hatching dates, and hatchling sizes (SVL and total length 
in mm) for a clutch of Amphibolurus nullarbor eggs. 

Votes Measured (1980) 





Egg no. 





(Dec. 1980) 




x 13.9 

24.7 x 17.4 

26.0 x 18.6 

26.1 x 19.1 


36.3 85.3 



x 14.6 

26.1 x 17.0 

27.1 x 18.3 

26.6 x 17.9 


37.0 87.5 



x 15.8 

26.6 x 17.9 

27.6 x 19.4 

26.9 x 20.0 


36.3 83.2 



x 15.0 

26.2 x 18.4 

27.6 x 19.3 

24.9 x 19.2 


37.3 80.0 



x 15.3 

233 x 18.1 

26.1 x 19.1 

25.1 x 19.1 


35.9 86.1 



x 15.6 

26,3 x 18.7 

27.0 x 19.3 

26.0 x 18.2 


33.6 75.6 



x 14.4 

26,0 x 18.0 

26.7 x 18.7 

26.6 x 18.9 


35.6 76.0 



x 16.1 

24.8 x 17.9 

25.6 x 18.7 

24.2 x 18.9 


35.0 80.2 



x 14.2 

24.8 x 17.9 

26.4 x 18.6 

25.8 x 18.4 


36.1 82.8 

X ± SD 



25.0 17.9 

26.7 18.9 

25.8 18.9 

35.9 81.9 



±1.1 ±0.5 

±0.7 ±0.4 

±0.9 ±0.6 

±1.1 ±4.2 

f Dates when eggs laid. 

* Note that all eggs except No. I decreased in length and/or width just prior to hatching. 


These observations are similar to those reported 
for A. barbatus f which most resembles A. nullar- 
bar, morphologically 4 . Eggs of A. barbatus in- 
creased 90% by volume from parturition to 
maximum size, under incubation conditions simi- 
lar to those described here* 1 . Also A . barbatus 
eggs shrank slightly just prior to hatching, and 
hatchlings remained motionless in ruptured eggs 
for several hours 11 . Furthermore, hatching times 
(76-84 days) for a clutch of A. barbatus eggs 
from Queensland, incubated at a similar tempera- 
ture, were similar to the A, nuflarbor data fi . 

In S.A. populations of both species are allo- 
patricP, Parturition seems to occur slightly earlier 
for A. nullarbor (early October) than for A. 
barbatus (late October, with most clutches re- 
ported in November-December 2 * "). This difference 

may be related to the more northerly distribution 
of A. nullarbor populations in S.A., which ex- 
perience the seasonal effects of exogenous stimuli 
(longer photoperiod and increasing temperatures) 
favourable to the onset of gonadal cycles* earlier 
than populations of A. barbatus. Clutch sizes arc 
much smaller for A. nullarbor (14-16 eggs for 
the two females examined) than for A. barbatus, 
which lay 15-35 eggs per clutch (X=25±=6 for 
six observations from S.A.-' s , and often have two 
clutches per season 2 * & *. Snout- vent lengths for 
hatchling A. barbatus average 5 mm larger than 
those of A. nullarbor. These differences arc be- 
lieved to be related to differences in body size of 
adults (maximum SVL 141 mm for A. nullarbor 
compared with 220 mm in A. barbatus)". 

Roman Ruehle photographed the halchling A. 

'Bustard, H. R. (1979) Australian Lizards. Col- 
lins, Sydney, 162 pp. 

-Smith, J. (1974). S. Aust. Herpetol., 2(1): 10. 

sSmith, J. (1979). Hcrpetofauna, 19(2): 12-14. 

'Uadham, J. A. (1976). Aust. J. Zool., 24: 423- 

"Houston, T. F. (1979). 'Dragon lizards and 

goannas of South Australia.' S. Aust. Mus. Spec 

Ed. Bull. Ser., 84 pp. 
'-Bustard, H. R. (1966). Brit. J. Herpetol., 3: 

•Licht, P. (1973). Comp. Biochem. Physiol. 45A: 

*MitchelI, F. J., unpublished data. 

Julia Smith, 7 Jeffrey Road, Vale Park, S. Aust. 5081 and Terry D. Schwaner, South Australian 
Museum, North Terrace, Adelaide, S. Aust. 5000. 


by Meredith J. Smith and P. A. W. Rogers 


Many Australian mammal species suffered a drastic reduction of range and some became extinct 
soon after European colonization. Information on the former distribution of these species can be 
obtained only opportunistically by the discovery and recognition of their remains in protected 
locations such as caves, dunes and middens. 




Many Australian mammal species suffered a 
tlru.stic reduction oF range and some became 
extinct suou after Furopean colonisation [nformu- 
tion on the former distribution of these ftpeCfeS 
can be oblairted only opportunistically by the dh- 
eovcrv and recognition of their lemnins in pro- 
tected locations &0ch as eaves, dunes and middens. 

During the exploration of Yellowfooted Rock 
Wallaby Case (lat. 31 J 20'S. long. |3X 40 RV 
about 30 km noiih of Wilpena in the Flindets 
Range?, Saudi Australia one of us (l\R.) found 
two macropodiil skulls and a pelvis in a low tight 
passage sume distance from the entrance to the 
cave. The topography of the cave indicated thai 
there was little likelihood of the bones having been 
earned to their resting place by a water current. 

T{w more complete skull (now registered South 
Australia* Museum (SAM MJQ76<fl lacks left V 
and a few foments of the pre maxillae. The 
lower jaw and the anterior cervical vertebrae are 
held tightly in place by dry flesh and skin, and 
fragments of dry sMn bird fur adhere to the skull. 
The oiher skull (SAM Ml 0770) lucks the incisors 
and anterior patts of the nasal bones. It has no 
skin adhering but is greasy. The pelvis and 
ailactied lumber vertebrae have ligaments and 
dried flesh adhering; they seem to have been 
gnawed by rndenw 

Both skulls are of adult Rettoncio tesnettr 
(Quoy & Oamiard) and the peU IS is almost cer- 
tainly of this species B. t£&fUr is characterixed 
by verticallygroomL veiy lone premolars, low- 
cmwned himodimi molars, steep si/e gradient of 
molars WJjtl W »W small. a>d by inflated .-indi- 
lory bullae. The P 4 length is about 14% of basal 
leneth of kkuN, the bulla length ihout 259; ot 
hasaf length 1 For the two cave skulls the measure- 
ments in mm relevant to diagnosis arfc; basal 
length f>3.0, appro*. r»;<; p< length R.9, $.9; bulla 
length I7.n, 17.8 respectively. 

If. It.tHtur was widespread in Australia and as 
laic as I'MO was abundant in the Musgrave- 
Gverard Ranee area'-' although it exists now only 
on offshore islands in Western Australia' 1 . Fintay- 
son- found "abundant evidence to show that Qt p 
tanutu lewettr was *'ormerlv one of the most 
numerous and universally distributed mammals ol 
South Australia, finding and colon b .trig siti tabic 
habitat /ones in all the tlisli acts ol the Stale with 
the possible exceplion of the deeper Malice and 
(In- flooded portion of 'be lower South-Fast afl<J 
the inner portions of the MulUrbOl Plain". Hii 
broad outline of the distribution included all nf 

\SakeUiuJ, V A. i IUft7 |, yict. Mat K4, N->2 
'frt*la>son f It II. UK5B). Kee. S. Ausl, Mus 
I312J, 235^T02 
Kitle, W. O. L <I'>70K "A guide to Ihe tiatfW 

mtimnuds of Ausiiaha". Oxt'oid University Press 

lie. I Map of South Australia showing location 
of origin of specimens of /?. lesuettr in the South 
Australian Museum mammal collodion. v\rrow 
shows Yellowfoolcd Rock Wallaby Cave, 

South Australia except the far northeast However, 
few specimens were retained in museums. In the 
South Australian Museum there are no specimens 
o^ B. h'stiair from the Hinders Ranges die nearest 
locality being Pi Augusta, whei'e one specimen 
(K9Q931 was taken in 1880 (Pig. It and living 
specimens have not been recorded from the Flin- 
ders Ranges'- A mandible fragment oi' H. Icsucur 
has been found in an owl pellet deposit in 
Chambers Gorge, northern Flinders Ranges (Ci. C. 
Medlin, personal communication) 

The rocky habitat around Yellowfoolcd Rock 
Wallaby Cave is not typical for B. te.saetir, indi- 
viduals of which are burrowers, and live in war- 
rens dug in sandy soil 1 . Whilst ihe small areas of 
alluvial sand deposited in the gorges by Fnorama 
and Brachina Creeks may have provided some 
stnlable substrate for but rowing, the position of 
the bones deep wUhin a small cave suggests that 
Li i ej as well as burrows were used for shelter Tt 
is unlikely that the specimens from the cave lived 
' than in (he Ranges because the plains 
to the west are about |y km distant and lo the 
east hlflber away. 

Whilst dried flesh may survive on bones in a 
cave for thousands of years-"' the skulls from Yel- 
low footed Rock Wallaby Cave provide a positive 
record of ft- trvatir I mm the Filmier* Ranges in 
the late Holocene. 

'Vilken, P. F. (19H0I Mammals, pp. 288-314 in 

Cornell, D. ed., 'A Field Guide to the Flinders 
Ranges." Rigby, Adelaide 

-''Partridge, J. ' I'W). J. R. 


Soc, w. AusL 50, 

Mini mm I. Smith :iM d I 1 A W. KootRsN 'Department of Zoology, University of Adelaide, G. P.O. 
Box 4s>x. Adelaide, S. Aust. 5001. [Department of Human Morphology. Flinders Medical Centre. 
Hetlford Park, S. Aust. 5042. 



by Wolfgang Zeidler 


In April 1980 fishermen from Port Lincoln reported the capture of a giant squid bearing hooks 
instead of suckers on the arms. The specimen, estimated to be 2 m long x 1.5 m wide, was floating 
on the surface 120 km offshore, south of Port Lincoln. Unfortunately the boat from which the 
discovery was made did not have the facilities to keep the specimen whole, and so it was cut up and 
subsequently sold for bait. However, two arms and the beaks were recovered, thus permitting 
generic identification. 




In April l^SO fishermen from Porl Lincoln 
reported the capture of a giant squid bearing 
hooks instead of suckers on the arms, The speci- 
men, estimated to be 2 m lony a 1.5 m wide, was 
floating on the surface 120 km offshore, south of 
Port Lincoln. Uufotumately the boat from which 
the discovery wa& insole did oat have facilities to 
keep ihe specimen whole, and so it was cut up and 
subsequently sold for hail. However, nvo ann^ DTU3 
the bea\s were recovered, thus permitting yenerie 

Dismay at the destruction of such a scientifically 
valuable specimen prompted fishermen to look out 
for more and. in early May, 1980 two were re 
covered from the same general area. One of these 
had Ihe head and arms missing, but still weighed 
93 kg and had a dorsal mantle length of 158 cm 
The other specimen weighed in excess ot I Hi fcf 
and measured 2.1 m total length. It was in almost 
perfect condition and is currently frozen awaiting 
.i more detailed examination 

Preliminary Investigations have shown that tbv 
specimens represent the genus Tanin^h ( Family: 
Oclopoteuthidae), but as they have not been exa- 
mined in detail a specific determination could not 
be made. However, they are most likely T datmr 
Joubin, 1931, as the genus is mouotypic and the 
material at hand does not seem to vary markedly 
from published descriptions 1 - 2 . 

There arc a number ol distinctive features which 
characterize this species. The arms are relatively 
Short, about i of the manrle length and arc armed 
with a series of hooks instead of suckers, The 
second arms each bear a taree light organ at then 
tip which was bright yellow in the present speci- 
mens and measured about 5 X 3 cm. The tentacles 
which account for most of the length in the hue 
giant squids, ArrhUnKhis, ate absent m mafurc 
specimens and vestigial in juveniles of Tunlngta. 
The fins extend for most of the manile and arc 

very large, making the body of the animal almost 
SI broad as the manile length. The mantle wall 
and fin Art extremely thick and gelatinous and the 
body is relatively more bulky than any other spe- 
cies of squid 

Nearly Ml of the known specimens of Juninxht 
have been collected ftom sperm whale stomachs, 
and it is unusual to encounter them floating: on the 
surface. It J9 possible thai they were regurgitated 
by sperm whales, and this may he the case for the 
specimen lacking a head, but the other two were 
found in relatively good condition and the fisher- 
men estimated that they had died only very 
recently. According to the Porl Lincoln fishermen 
dead squid often i:ome to the surface near the edge 
of the continental shelf, but they arc quickly eaten 
by seabirds and it is difficult to find a complete 
Specimen* T, danar forms an important part of 
the diet of sperm whales and by weight \h the most 
Significant part of sperm whale dicrs a . Clarke* 
recorded beaks and portions of flesh of this species 
from the stomachs of sperm whales caught near 
Albany, W.A., but this is the first definite record 
of Taningia from Australian waters, tfec pre 
viously known distribution of Tanhtxia, mainly 
From sperm whale stomachs, includes Ihe Azores, 
Madeira* the equatorial Atlantic and the eastern 
coast of South Africa". 

I intend to publish the results of a more detailed 
examination of Ihe specimens in the near future, 

My sincere thanks lo Dinko Lukin and the crew 
ot the "Karina G' 1 and to .luergen 2cnkc and the 
crew of the "Rosalind Star" for collecting and 
donating the specimens to the South Australian 
Museum. My thanks also go to S.A.F.C.O.T. who 
transported the specimens to Adelaide Iree. ot 
charge. A special thanks is due to Neville Wanklyn 
of the 'Tort Lincoln Times" who alerted the fisher- 
men to the importance of the specimens, and 
to his Editor for bein? so understanding. 

'.louhin. I . MW. Annh Inst i^o.-mogr, Monaco "CtaHie, R. '1956). Discovery Rep. 28, 237-98. 

^Liut*l!'R. ,1.67) Symp ,ool &*. Loud. ***** ** & "*»** A* *, 124. 
(3*1, 127-43 

Woi.i-gan.; ZiiOLtn. South Museum North Terrace, Adehide, S. Atist. 5000. 


BifTLUK & Keough— Trans. R. Soc. $ Aust- 105(1), 29-39. 

The authors apologise for ihe omission from rhc manuscript of Acknowledgements. We 
intended to include the following: 

We are indebted lo Tony Oliver and Dcudrc Cooofbc for assistance rn ihe held. The work 
was supported by a grant from the University ttf Adelaide Research & Publications Committee 
and byARGC Grant D1 78 15083. 





D. W. P. CORBETT, B.Sc, Ph.D. 


M. DA VIES, M.Sc. 


M. J. TYLER, M.Sc. 

Assistant Editor: 
A. R. MILNES. Ph.D. 


B.Sc, Dip. Lib. 

Programme Secretary: 
P. J. SUTER, Ph.D. 

Minute Secretary: 
P. M. PEARSON, B.Sc, Dip.Ed. 

Membership Secretary: 

Members of Council: 

K. H. NORTHCOTE, B.Ag.Sc I. A. T. BYE, Ph.D. 

D. C. LEE. M.Sc J. SELBY, B.Sc 

C. R, TWIDALE, Ph.D., D.Sc 


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