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VOL. 105, PARTS.1 & 2 
12 JUNE, 1981 


Transactions of the 


Royal Society of South 
Australia 


Incorporated 


Contents 


McQuillan, P. B. A review of the Australian moth genus Thalaina (repidopiers 
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 - - - - ~ = z = . : 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. Sian she a new pening ostracod 

genus from Australasia - - 53 
Bye, J. A. T. Exchange processes for upper Spencer Gulf, South Australia - 59 
Jenkins, R. J. F., Plummer, P. 8. & 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 


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


TRANSACTIONS OF THE 


ROYAL SOCIETY OF SOUTH AUSTRALIA INC. 


CONTENTS, VOL. 105, 1981 


PARTS 1 & 2, 12 JUNE 


McQuillan, P. B. A review of the Australian moth genus Thalaina (Lepidoptera: 
Geometridae: Ennominae) - - : < 2 = “ 


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


Butler, A. J. & Keough, M. J. Distribution of Pinna bicolor Gmelin (Mollusca: 
Bivalvia) in South Australia, with observations on recruitment 


Pledge, N. S. The Giant Rat-Kangaroo Propleopus oscillans (De aS); (Poto- 
roidae: Marsupialia) in South Australia - - - 


Crook, G. A. & Tyler, M. J. Structure and function of the tibial gland of the 
Australian frog Limnodynastes dumerili Peters - - - 


De Deckker, P. & McKenzie, K. G. Sg anleieeees a new cyprididid ostracod 
genus from Australasia - 5 : = : 


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


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


Womersley, H. B. S. & Sinkora, D. Sonderophycus and the type specimen of 
Peyssonnelia australis (Cryptonemiales, Rhodophyta)  - - 


Brief communication: 
Tyler, M. J. & Davies, M. Partial acquisition of pigmentation in an adult, albino, 
Australian Leptodactylid frog (Limnodynastes dumerili Peters) 


25 


29 


41 


49 


67 


85 


PARTS 3 & 4, 11 DECEMBER 


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


Beveridge, I. Three new species of Calostaurus (Cestoda: Davaineidae) from 
the New Guinea wallaby Dorcopsis veterum - - - - 


Tyler, M. J., Davies, M. & Martin, A. A. Frog fauna of the Northern Territory: 
new distributional records and the description of a new species 


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


and Ophionereis - - - “ 2 . = 3 = 


Jenkins, R. J. F. The concept of an ‘Ediacaran Period’ and its stratigraphic sig- 
nificance in Australia - - . 2 3 5 3 <! 


Foster, C. B. & Harris, W. K. Azolla capricornica sp. nov. First Tertiary record 
of Azolla Lamarck (Salviniaceae) in Australia - - - 


Bull, C. M. & King, D. R. A parapatric boundary between two species of reptile 
ticks in the Albany area, Western Australia - - - - 


Brief communications: 
Von der Borch, C. C. Recent non-marine dolomite from the coastal plain, south- 


eastern South Australia - = . i € Z x 


Kailola, P. J. & Jones, G. K. First record of Promicrops lanceolatus (Bloch) 
(Pisces: Serranidae) in South Australian waters - - - 


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


Smith, J. & Schwaner, T. D. Notes on reproduction by captive Amphibolurus 
nullarbor (Sauria: Agamidae) - - “ 3 2 


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


podidae) from a cave in the Flinders Ranges, South Australia - 
Zeidler, W. A giant deep-sea squid, Taningia sp., from South Australian 
waters - = - - - - 3 = 2 : 
Addendum - - - - = : : : - 7 . r 


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


91 


139 


149 


155 


179 


195 


205 


209 


211 


213 


215 


217 


218 


218 


A REVIEW OF THE AUSTRALIAN MOTH GENUS THALAINA 
(LEPIDOPTERA: GEOMETRIDAE: ENNOMINAE) 


BY P. B. MCQUILLAN 


Summary 


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 
established. 


A REVIEW OF THE AUSTRALIAN MOTH GENUS THALAINA 
(LEPIDOPTERA: GEOMETRIDAE: ENNOMINAE) 


by P, B, McQutLian* 


Summary 
McQuittan, P. B. (1981) A review of the Australian moth genus Vhalaina (Lepidoptera: 
Geometridae: Ennominue), Trans. R, Soc. S. Aust, 105()), 1-23, 12 Sune, 1981. 

The moth genera Thalaing Walker, Thalainedey Lower and Macqueenia Turner are 
reviewed) New evidence has resulted if an expansion of the concept of Thaluina to include 
the other genera in its synonymy, Nine species are reviewed and 7. kimba sp. noy., is described 
from the mallee areas of South Australia aud New Sauth Wales. Immature stages, Foodplants, 
flight activity and distribution of the varfous species are recorded where known, The genus has 
adapted successfully 10 most major habitats in the southern half of Australia wherever (heir 


food plunts (Acacia and Cassia) are established. 


Jntroduction 

This is the first of a proposed series of taxo- 
nomic studies on the ecnnomine geometrids of 
southern Australia, [It reviews the taxononyec 
status and known biology of a distinctive 
group of autumn-flying moths previously re- 
ferred lo Thalaina Walker, Thalalriodes Lower 
and Mecqueenia Turner, 

All the species are large, strikingly patterned, 
nocturnal moths with distinctive larvae, Wot 
surprisingly, all four species found in coustal 
easter Australia were described by 1865. Re- 
siding at Broken Hill O, B. Lower discovered 
and described three inland species between 
1900 and 1902, and his series of two of them 
constitute most of the specimens available tor 
study, An apparently localised species was: dis- 
covered in the 1930's at Millerran, southern 
Queensland by J. Macqueen and is suil very 
poorly known. Jn the 1960's and early 1970s 
au few specimens of a widespread central 
Australian species were collected jind subse- 
quently deseribed in 1972. Similarly, concen- 
tration of collecting in the semi-arid areas of 
southern Australia during late autumn by 
CSIRO collectors over the last ten years has 
mide available a good series of another oew 
species described here. 


Methods 
The following abbreviations are used for 
collections: AM = Australian Museum, 


Sydney; ANIC = Australian National Insect 
Collection, CSIRO, Canberras BMNH == 
British Museum (Natural History), London; 
NMV = National Museum of Victoria, 


+ Division of Entomology. Dept of Agriculture 
Laboratories, St John’s Ave, New Town, Tas. 
TO08. 


Melbourne; PBMcOC ~ FP. B, McQuillan col- 
leetion, Adelaide; QM — Queensland Museum, 
Brisbane; SAM — South Australian Museum, 
Adeluide; TDA = Tasmanian Department of 
Agriculture, Hobart; TMAG Tasmanian 
Museum and Art Gallery, Hobart; UQ = 
University of Queensland, Brisbune; WAM 
Western Australian Museum, Perth. 
Abbreviations of the names of collectors 
are: PA > P, Aitken, PMA = F, M. Angel, 


EA E, Ashby, WBB W. B, Barnard, 
RB - R. Beresford, TR — T. Bluekburn, 
Tac — Ct. G. Campbell, DHC = D. H. 


Colless, IFBC = 1. F. B. Common, JRC — 
J. R, Cunningham, WD J, W, Davies, 
EID~ E£.J, Dumigan, EDE — EB. D. Rdwards, 
BE. — B. Evans, RHF = RK. H. Fisher, CWF 
C. W. Frazier, NG = N, Geary, GCLG 
-G. C. L, Gooding, GHH = G, H. Hardy, 
RJH — R, J. Hardy, JH = J, Harslett, CCL 
C, Cc, Ives, WRK = W. Kleevaj, LEX — 
l,. E. Koch, LRK = L. R, Rurtze, AML = 
A. M. Lea, RL = RK Lewis, OBL — O. B, 
Lower, GL = G. Lyell, NMcF — N. MeFar- 
land, KJMeK = K. J, McKie, JIM — J. 
Maecqucen, PBMcQ = P. B. McQuillan, 
WLM W. L. May, VHM = V. BL Minehin, 


BM — B. Mollison, GBM G, B, Monteith 
JGM ~ J, G. Morris, TN T. Newberry, 
LIN — L. J. Newman, KRN = RK, R, Norris, 


HP — H. Pelz, RIP = R, J. Priest, PR — 
P.. Runford, LMR — L. Mosse-Robinson. VIR 
-V. J. Robinson, ALR ~ A, L. Rovers, RS 
= R. Straatman, JJHS = J. J. A, Szent-fvany, 


IGOT — J, G, O. Tepper, HST = H-. 5S. 
Thirkell, |.BT = L, B, Thorn, NBT — N. B, 
Tindale, HU = H. Udell. MSU = M. &. 


Uptan, MMHW = M. M, H, Wallace, RW 
— R, Went, JOW = J, 0. Wilson, RGW = 
R, G. Winks, FWJ = F. Woad-Sones., 


2 P. B, MeQUILLAN 


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; sce McFarland (1979) for full 
details, 


Genus THALAINA Walker 


Thalaina Walker 1855, p. 659; Type species 
Thalaina klenaea Walker (=Thalaina selenaea 
(Doubleday)), by subsequent designation by 
Fletcher 1979, p. 202.. 

Absyrtes Guenée, 1857, p. 226; Type species 
Absyrtes magnificaria Guenée, by subsequeni 
designation by Fletcher 1979, p. 1. 

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

Macqueenia Turner 1947, p. 101; Type species 
Macqueenia chionoptila Turner, by monotypy; 
syn. nov. 

Adult; Medium sized, moderately robust, noc- 

turnal moths; wing expanse 34—54 mm, Colour 

HON Ab Hee, 

Wine! Za 


pattern basically white with or without darker 
geometrical markings. 


Head (Figs 1, 2) with vestiture of crown 
thick, hair-like slightly roughened; trons well 
rounded (fairly convex), clothed in scales, or 
nearly naked and bearing series of shallow 
transverse ridges beneath large sclerotized 
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; chaetosemata 
present. 

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 R,., 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, 


REVIEW OF THE MOTH GENUS THALAINA 3 


TABLE 1. Comparison of some medn ratios from 
leg measuremenry af Thalainn species tvee fig, 3 


jor abbreviations). 
Fore lez Hind leg 
no F/T TET BT F/T TSBrT ISB/T 
selunaga 3°15! 056 DAS DES 1.66 0.28 
aanyrelinnis 3 150 O53 0.45 0.66 0.67 0.28 
clara f 3 LTS 8.60 O48 TOR UA USS 
inveripta q 3 LT? 0.61 O44 GATE O67 O28 
allgehrad J 3.13!) 0238 O46 O75 867 0.27 
paronyeha 2 #159 0.2 035 O86 G69 ia 
kimba at 2 14% O40 O58 USI 0.73 1.24 
chionepiilla® 2 160) 0.51 o55 068 0.66 0.27 
fepraelailit of 3 146 049 OSI 066 O67 0.28 
macfanlanedi a Lo oWI7T Oa} 0394 O36 O72 O24 


form an areole, R, stalked with Ry, 4, Mz 
often arising nearer to M, than to M.> bind- 
Wing White, sometimes with darker markings, 
cell rather long; femora smooth-scaled, pos- 
terior tibiae in male slightly dilated, fore tibiuc 
with epiphysis, sometimes with apical hook, 
mid and hind tibiae with spurs well developed 
(for lez measurements, see Table L). 
Abdamen with comb on segment 3 in male 
weakly to moderately developed, 

Male genitalia with uncus simple, slender, 
apically acute; socit well developed, gnathos 
simple, slender, with small median recurved 
spine or group of small spines al apex; juxta 
broadly rectangular or $hield-shaped; furca 
usually Jong, bifid; cristate hairs well de- 
veloped; valya elongate, slightly spatulate; 
aedeagus tubular, slightly curved with pointed 
apex and oblique distal orifice, cornuti usually 
a series of stout, often compound spines. 
Female genttalia with papillae anales densely 
hairy, bases of hairs set in conical projections; 
apophyses posteriores about 1.4 times as long 
as apophyses anteriores; sterigma broadly cup- 
shaped; ductus bursae long, with parallel sides, 
weakly sclerotized, in diameter about + that 
of corpus bursae, is posterior part usually with 
ring of longitudinal sclerotized striations; 
corpus bursae ovoid, witheut signa, 

Eee broadly ovate, with or without darker 
marking. 

First lnstar larva with head capsule brown, 
without pattern; body pale yellow; setae ex- 


iremely long and quite stout; setae XD and D 


oo protherax on a lightly sclerolized plate; an- 
terior humps on prothorax lacking. Abbre- 
viated prolegs present on Ad and AS; 
crotchets arranged in complete circle on Ad 
and AS (numbering about 12) and in half 
circles on AG and claspers. 


Final instar larva with head capsule green to 
yellow-green with numerous small brown 
blotches avross vertex; labrum deeply emay- 
ginale; body fairly robust, pale green to dull 
green with at least a whitish Jateral stripe; 
euticle smooth; setae short and fine; prothorax 
somewhat swollen and produced into two an- 
terior dorso-lateral fleshy prajections; ab- 
breviated prolegs present on Ad ond AS; 
erotchets in 3/4 circle on A4, in 5/6 circle on 
AS and } cirele on A6 and claspers, uniot- 
dinal on A4—A6 but tending to be biordinal 
on claspers of same species: numbers of 
erotchets range: 13-21 on A4, 14-29 on .AS, 
25-37 on A6é and 28-48 on claspers; leneth 
23-41 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 und ventral stripes. Those 
species feeding on bipinnate Acacia usually 
have pale dorsal stripes and more mottled 
colouration, 

Pupa dark brown to orange-brown, smoath, 
anterior margin of A1O dorsally with 2-4 
widely rounded teeth: cremasler hooks 2 or 
6-8; length 11-17 mm, width 5-6 mim. Acsti- 
vates in loase Cacoon incorporating detritus, 
beneath litter or in the soil. 

Flight Periad: Mostly Jate summer to fate 
autumn; flight periods for individual species 
range from Jess than four to more than twelve 
weeks. Figure 18 shows flight periods for 
species represented in collections by over 20 
specimens with full data. 

Distribution: Australia, south of about latitude 
24°; see figs 49-51, 


Key tu adults: 
1. FProns more or tess maked, wilh idlefotized 


median projection ..... 2 
— Frons covered with hair-scales, without 
median projection 1.6, alee ahah ie IG 
2, Forewing satiny-white with sharply defined 
orange-hrown or achreous hrown markings 3 
— Forewing reply pale ahah or 
an PE pan cite 5 
3. Forewing with stripe witas: middle one-third 
of costa 4 


— Forewing without stripe along middle ones 
third of costa _ macfarlandi (Wilson) 

4. Forewing with oblique stripe from near mid 
fermen io costal streak at three-quarters 

4 paronycha (Lower) 

_ Forewing without stripe connecting termen 
and costa as above ,.,.,. fetraclada (Lower) 

5. Porewing completely Schteans-gtey, minutely 
speckled with black allochraa (Lower) 


4 P, B, McQUILLAN 


— Forewing white but heavily marked with 
broad, suffused grey markings . kimba sp. n. 

6. Forewing without markings in disc ...,. 7 

— Forewing with markings in dise ......- $ 
7, Hindwing with a large black spot near tornus 


chioneptila (Turner) 
— Hindwing without a large black spot near 
tornus selenaea (Doubleday) . 

: _(form punctilinea Walker) 

8, Foréwing with short; ochreous brown streaks 

on M;, Ms, CuA;y and CuAes near termen 

inscripta’ Walker 

-- Forewing without ‘markings at exlremives of 

Mi, Ma, CuA; and CuAs —.. 2 a) 

9, Forewing with a relatively straight stripe 

from tornus to near mid costa or a litte 

below mid costa... enn FO 

— Forewing with stripe from tornus mot 

reaching mid costa angulosa Walker 

10. Forewing with diagonal stripe from mid 
termen to three-quarters of costa ..,,,,, 

clara Walker 

= Forewing wilhout stripe from mid termen 

lo costa as ubove selenaea (Doubleday) 


Thalaina selenaéa (Doubleday) 
FIGS 4, 14, 18, 19, 29, 39, 49, 52, 53. 
Callimorpha selenuea Doubleday 1845, p. 437, 
pl. 5, fig. 3 
Thalaina selenaea, Meyrick 1892, p, 653; Turner 
1919, p, 387. 
Thalaina klenaea Walker 1855, p, 660, 
Pompeja australiaria Herrich-Schiiffer 1855, pl. 60, 
fig, 333, 
Absyrtes australiaria HSch, (magnificaria Gn.) 
var, fartanara Thierry-Mieg 1899, p, 21, 
Absyvries magnificaria Guenée 1857, p, 226. 
Phalacna magnifica Desmarest 1858 syn. nov, 
(junior objective synonym of magnificaria Gn.). 
Thalaina punctilinea Walker 1865, p. 228. 
Type ot selenaea; not found in BMNH 
(D, S. Fletcher, pers. comm.) or any Aus- 
tralian collections. 
Types of klenaea’ lectotype 2 labelled “Calli- 
morpha yelenuea H. T. [Hobart Town] Aus- 
tralia 44-105" in BMNH, hereby designated: 
14 paralectotype labelled "46-46" in BMNH, 
hereby designated, 


Type of anstraliaria: not found in Zoological 
Institute, Martin Luther University, Witten- 
berg, GDR (N. Grosser, pers. comm.). 


Holotype & of autraliaria var. fortunata 
labelled “Typicum specimen Ex Musaco 
Ach. Guenee : Magn. var A Gn. : Ex Ober- 


thur Coll. Brit, Mus, 1927-3" in BMNH. 

Type of magnificaria: this is the earliest 
scientific name and formal description of text- 
fig. 17 p, 5S of Lucas (1857), colloquially 


named by him as “Phalene magnifique” 
according to Guenée (1857) 4 3, 19 were 
in his collection at the time of description; 
none of these syntypes has been examined. 

Type of magnifica: Desmarest applied this 
name to Lucas’ unnamed figure; the specimen 


R2 R4 
——_\ R5 
Mi 


ey 
— M2 
oo | M3 
-—— | cuat 
| 
—/ CuA2 
/ 


es 28 


Figs 4-5. Venation of Thalaina spp. 4. T. sele- 


naea, 5. T. chionaptila. 


REVIEW OF THE MOTH GENUS THLAINA bi 


on whieh the figure is based haus net been 
cxamincal, 
Type of puncitineas fectoivpe labelled 
“Tasmania S$-60" in BMNH, hereby desig- 
nated; | ¢ peraleerotvpe labelled ~“V_D.L.., 
W.H.S. [or W.W.S.| 43-58" in BMNH, hereby 
designated. 
Adult (figs 52. 53); Head with frons: rounded, 
smoothly huiesealed. whilish with fuseous 
hand helow vertes; vertex. with ereet hair- 
geales, brivht red-brown; labial palp wok 
torminal segment and upex of second seyment 
fuscous, remainder whites antenna of male 
luminute, Thorax above greyish white Gn an- 
(erlor margin, renvainder white, base of Wings 
orange, beneath white; legs white with exterior 
of fore und mid femur and Ubia and all tarsi 
infuseated; forewing (lig. 4) with costa meurly 
straight, apex pointed, termen rounded and 
somewhat sinuate beneath apex, Ry anasto- 
mosed with Se, Ry offen anastomosed with 
R. lo form an areale, ground colour above 
shining white. ced-brown sireak along costa 
from bast to one-half thence angled inwards 
as a narrowly black-margined streak io tarnus: 
similar streak from tornus along inner margin 
to near hase, termen narrowly red-brown, cilia 
red-brown posteriorly fuscous; hindwings shin- 
ing White, moderite fuscous subupical blotch, 
cilia white, forewings beneath shining while 
infuseated near apex and With costal streak 
fuscous; hindwinus beneath shining white, 
subapical bloich much cnlurged, sharply red- 
hrown above My, fuscous helow; wing expanse 
as—S4 mm, 9 §2-36 mo. 
A common variety of this species (fig. 53) 
hus u reduced casial streak, lacks the diagonal 
forewing streuk and has the streak along the 
inner margin poorly developed; the subupical 
hindwing blotch may be reduced or even 
absent butt is Fully developed beneath. 
Male genitalia (ig. 19) with apex of gnathos 
with small spine. furca with two prongs of 
equal length, aedeagus (fig, 29) stout, corauti 
af twa compound spines subequal in lenuth, 


Female genitalia Cig. 39) with corpus biirsac 
elhowed. 

Final tastar hirvas Head capsule 3.3-3.5 min 
wide, pale green wilh small fuscous blotches 
weross vertex; body green with Jine fuscous 
speckling oo aital plate and claspers, fleshy 
whitish Inferal stripe visually present, two white 


yubventral stripes present as blotches adjacent 


to thoracic legs and continuous from micta- 


thoru#s lo A, inconiplete white mid ventral 
stripe on Al to AS: crochets 19-2) on Ad, 
22-29 on AS, 31-37 on AG ond 35-42 on 
anal clispers, crochels on anal claspers unior- 
dinal length 38-4) mm, width 5.S-6 mm. 
Miiterial: G19. 

Pupa (fig. 14) mid to dark brawn in colour, 
cremuster hooks 6-8: dorsal anterior margin 
of AlO with 4 labes. Material: G119, 


Fooddplanis: Acacia amelanaxylon R. Br. A 
retinades Schlecht. 


Specimens evamined: 83 ff 44 9. QUEBNSLAND: 
Brishbune, v. RGW | { UO; Lamington Nt) Park, 
i, v. WK 2 4 UO: Stinthorpe. iv. | of UQ: 
Touwoombi, di, mw. EID 2 3 LIQ; Wyberba, 
iii. JFBC | ¢ ANIC, NEW SOUTH WALES- 
Brown Mountain, i, DHE | 2 ANIC; & km N of 
Bungwahl, ii. [FBC & MSU 1 ¢ ANIC, Church 
Point, Vv. IFBC ( 4+ ANIC: Clyde Mrn 800 m, ti. 
(FBC & MSU | & ANIC. & km W oof Colts 
Harbour. vo MSU 1 ¢@ ANIC; Depol Beach, iv- 
IFBC 1 & ANIC: Kanguroo Valley, iii, DHE | 
@ ANTC: Kaloombs, if, 1 4 SAM; Narara. ty, 
LMR I @ ANIC: 5 km SSE of Parl Macquarie 
v MSU | @ ANIC: 2.7 km NE of Oueuwtbeyan 
670 m, iv. FBC | 2 ANIC: 40 km S of Single- 
ton. fii. [FBO | & ANIC; Tooloom Scrub, ii. | 
4 UG: & km S$ of Wauchope. ili, [PBC & MSU 1 
+1 © ANIC. Wollongong, iii, VIR 1 4 ANIC, 
AUSTRALIAN CAPITAL TERRITORY: 1.6 km 
NE of Lee's Spring 1300 m, i IFBC | & 1 f 
ANIC VICTORIA: Boronia, iv. 1 ¢@ NMYV; 
Castlemuine, ili. | 2 8 km 8S of Gellibrand 230 m, 
ji, IFBC 1 2 ANIC: Gisborne, ii, iv. do 3 9 
ANIC, iv. 68 specimens NMV, i. iii ive GL 
14 4 9 SAM; Meeniian- iv. |? NMY: Mit- 
cham, iv. | &@& 4 9 NMV: Moe, iv, | & NMV, 
ii. GCLG | 2 SAM; Mordislioc, v. COT L oF 
ANIC: Mt. Ditheutt Ry, fi. IFBC 1 & ANIC: 
Springvale, v. 1 2 UQ: Turton's Pass. ti. NBT 
§ 4 1 2 SAM: Wandin, ive | 2 NMYV, TAS- 
MANIA: Condominion Creek. ii, 1 & 2 & NMV; 
Cracroft Crossing, 1i, § 2 NMV-> Devonport, | 
TDA; Freycinet Nt Park, ii. IFBC & MSU | od 
ANIC: Kelso, 1 & SAM; Kingston, fin. iv, IRE 2 
ot ANIC, iv. IRC 2 9 NMV; Leake Leake 661) m. 
iL IPBC & MSU 14 ANIC; Cuunceston. iv. 1 4 
SAM: 16 km W of Muydena, ii. 1 4 9 NMYV; Mt 
Nelson 200 m, iz, tii, PRMcQ 4 9 TDA: Mi Wel- 
lington 280m, i, ih. ii BM 7 o 4 9 ANIC; Ouse, 
ii, ith, iv 341-9 PDA; Parerna, King Island, i, 
PBMcQ 1 2 TDA: Qucefstown, v. 1 2 NMYV: 
Ridseway, lv. JRC 1 &@& 2 2 NMY: Raseherry. ii, 
1 2 ANIC: Sandford, WLM | @ ANIC, 5 km E 
of Wiratah 660 m, ii, IPBC & MSU 2 ¢@ 1 2 
ANIC: 13 km SW of Waratah 600 m. ri TR BO 
& MSU 2 ¢1 9 ANIC; Zeehan, . GHH 1 
ANIC, ji. 1.2 1 9 NMY. SOUTH AUSTRALIA: 
Furner, vo 3 & SAM: Natucoerte, | Y SAM: 
Yorke Peninsula, ii, iy. IGQT 1 f 1 ¥ SAM. 


6 P, B, McQUILLAN 


Figs 6-8. Yenation of Thalaina spp. 6. T, angulosa; 7. T. clara; 8. T- inseripta. 


Distriburion: see fig. 49, Flight Period: see 
fig. 18, 

Canments: T, selenaea is a dimorphic species 
— one torm (var. punctilinea Walker) without 
markings on the forewing and the other with 
a diagonal bar across the forewing; occa- 
sionally a streak along Mg may also be present 
(fig, 52). Tt oceurs in habitats in south eastern 
Australia from open woodland to closed forest 
in areas of greater [han 75 em annual rain- 
fall and at elevations of 0-800 m. Preferred 
hosis afte large phyllodinous wattles such as 
Acacia rélinodes and A. melanoxylon. An in- 
leresting 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 
lo April. 


Thalaina angulosa Walker 
FIGS 6, 13, 15, 18, 20, 30, 40, 50, 54. 


Thaluina angulosa Walker 1865, p. 289; Meyrick 
1892, p, 665; Turner 1919, p. 388. 


Holotype 9 without abdomen, labelled 
"Ss. Aust. 61-104" (FF. Waterhouse) in 
BMNH. 


Adult (fig. 54); Head with frons rounded, 
smoothly hair-scealed, whitish-buff; vertex with 
rough hair-scales, orange: labial palpi with 
terminal segment and apex of second segment 
fuscous, remainder of second segment white: 
untenna of male shortly bipectinate, Thorax 
above greyish-white becoming whiter pos- 
teriorly and on tegulae. orange at base of 


Wings, white beneath; legs with fore and mid 
femora and all tats: mfuscated, hind femora 
and tibiae whilish; forewing (lig. 6) with 
costa nearly straight, apex produced, termen 
strongly arched and sinuate below apex, R, 
anastomosed with Se, ground colour above 
shining white, a narrowly black-margined rust 
coloured streak along costa from base to one- 
eighth costa then extending ta mid-discal area 
where it divides into an upper arm reaching 
mid termen thence to four-fifths costa and a 
lower arm extending to tornus thence along 
the posterior margin to near base, termen 
narrowly rust coloured, cilia rust posteriorly 
fuscous; hindwing shining white, a large fus- 
cous subterminal blotch extending from above 
M, to near apex with a narrower extension 
to anal angle, cilia fuscous on termen from 
anal angle to five-sixths with remainder white; 
forewing beneath shining white with upper 
markings Visible, a subapical blotch extending 
from four-fifths cost to mid termen, orange- 
rust Near costa, Pemnainder fiscaus becoming 
paler apically; hindwing beneath shining white, 
sublerminal blotch as above but less extensive 
and sharply orange-rust above Mj; wing ex- 
panse ¢ 40-48 mm, 9 42-54 mm. Abdomen 
white. 


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


Female genitalia (ig. 40) with ductus bursae 
swollen proximally, 


REVIEW OF THE MOTH GENUS VHALAINA 7 


Ege: Ovoid, greenish, without mackings; 
stimulated ta hateh by rainfall. See McFarland 
1971, p. 242 for full details, Material; G100, 
Final instar larva! Head capsule 3.3 mn wide, 
greenish with namerous small brown blotches 
some of which form suffused band verass 
veriex; body bright green with dark speckling 
on dorsum of prathorax, on anal plate and on 
claspers; yellowish-white lateral stripe often 
heavily edged with fuscous on ils lower margin, 
sametimes four faint parallel whitish dines 
darsaily, two Whitish subventral stripes presen 
ind incomplete mid ventral stripe on Al to 
AG, bases of setae inconspicuous, crotchets 
16-19 on A4, 20-22 on AS, 29-30 on AG and 
33-38 on anal claspers; length 36-40 mm, 
width 45-5.5 mm. Material: G100, 

Pupa (tig. 15) mid brown in colour, cremaster 
hooks 2: dorsal anterior margin of A10 with 
2 lobes. Material: GLOO. 

Foodplantss Predominantly Acacia pyenantha 
Benth.; also recorded from A. brachybarrva 
Benth. and Cassia Prentephita A, Cunn. 
(McFarland 1979), 

Specimens exaniined: 113 f 77%, QUEENS- 
LAND: Stunthorpe. Ww. JE 1 ct SAM: Talwood, 
iv, WBB | specimen QM. NEW SOUTH WALES: 
| km NNW of Goolgowi, v. IFRC & MSU | 9 
ANIC: 10 km SE of Gol Gol, vy. [FBC 1 a tg 
ANIC: Tea Tree Creek ne Armidale, v. CWE 
| 2 ANIC, VICTORLA: Birchip, iv. 8 specimens 
NMV, iv. | 4 SAM; Gippsland, 1 9 NMY; 
Gisborne. ive 1 9 NMY¥, iv. GL 1 9 SAM; 
Hamilton, 1 2 NMV; Haochkirch, iv. LRE I spect 
men 2M; Melbourne, | 2 SAM; Sprngvale, v, | 
® NMV. SOUTH AUSTRALIA; Adelarde, iv., ¥- 
IGOT 1 419 5AM, v. 1 f SAM, v. BA 4 g 
SAM; Aldgate, v, | 2 SAM. Athelstone, iv, v- 
ys 2 of 2 @ SAM: Relair, iv. FMA 1 9 SAM: 
Blackwood. v. NBTV 1 3 1 & ANIC, iii, iv. ¥ 
NMcF 2 #32 ANIC, fil, iv, ve NBT W) ¢ IL 
G SAM, iv. YHM 1 2 SAM, iv. BA | SAM, v. 
FA 19 84M, v. OBL 1 4 5 2 SAM; Bowhili, iv. 
PMA i 2 3AM; Burméide, 1 7 SAM: 100 km E 
of Ceduna, v. JFBRC & MSU 2 ¢ ANIC; Gien 
Osmond. v. FMA | 2 SAM; Highante, iv. 1 4 
SAM; Kadina, v. 1 2 PBMcQC: Lynton, iv, Vv 
RHF 3 ¢ ANIC, iv. RAF 4 3 49 SAM; Mam- 
hruy Creck Nat. Pk. v, TPEBC 1 9 ANIC, Monarta 
Sonth, ve SWB 1 ¢ SAM) 4&8 km ESE of Morgan, 
vy. IFBC & MSU 71 @ ANIC: Ml. Lofty, AML 
1 9 SAM: Parkside, OBL | & SAM, iv, FMA 1 
@ SAM; Port Lingoln, 1 2 SAM; Renmark, ¥: 
PRMcQ 1 2 PBMcQC: Stonyfell ve FMA 1 ? 
SAM: Waikerie, iv, 1 & SAM; Waprradale, iv,. v. 
PBMcQ 4 ¢f 3 9 PBMcQc; 10 km SW of Wil- 
mington, v. TEBC & MSU | 3 ANTC; Whyalla, v 
i © NMYV=4¥ kn § of Whyalla, v, LEBC 3 ¢ 39 
ANIC, WESTERN AUSTRALIA: Erunswick 


Inlel, 1 &@ WAM; Burngup, 1 & WAM; Cape 
Naturaliste, tv, FBC & MSU 2 @ ANIC: Car- 
Jingup, v. | & WAM; Claremont, 1 ¢ WAM, 
26 km N of Collie, iv. [FBC & MSU } d ANIC; 
21 km WSW of Collie, iy. IFBC & MSU 2 d 
ANIC; 27 km SE of Coolgardie, iv. PRC & MSU 
2 4 ANIC, Corrigin, 1 42 WAM; Crawley, vi. 
KRN | ¢ ANIC; Denmark. iii, iv. WBB 2 
specimets QM; Drummond Cove 1] km N of 
Geraldton, v,, yt. NMcF 5 2 ANIC; Dumbleyvung 
1 2 WAM, Dundas, 1 2 SAM, Hamel, vod oc 
NMV; Katanning, v. KRN 1 % ANIC; i d 
WAM: Kojonup, iv.. v., vii ALR 8 ¢ 2 ANIC, 
iv, v. RIP YS J OANIC, iv, vv MMHW 6 3 ANIC, 
| 2 SAM: Lake Grace, iv. | 2 ANIC, iv, 3 speci- 
mens OM; 45 km W of Madura, iv. JEBC & MSU 
2 ft ANIC; 29 km W of Mogumber, iv. [FBC & 
MSU 4 ¢ ANIC, 98 km E af Norseman, iv, 
FBC & MSU 2 4 ANIC. 24 km N of Nor- 
thumpton, iv. JFRC & MSU 3 ¢ ANIC, 11 kin 
§ of Pemberton, iii, IFBC & MSU 1 #19 ANIC; 
Perth, %, Vii, 1 ¢o1 9 NMYV; Bithara, iv. FHC & 
MSU 1 @ ANIC, v. L & WAM, Ravenswood, 
vi. FMA 3 9 2 2 SAM: Stockyard Gully. Jaunen 
Buy, vi. BE 1 9 WAM; Swan River, vii. | @ QM; 
Tammin, ¥., vi. 9 specimens NMV, Yanchep Nail 
Pk, iv, IFBC & MSU 2 & ANIC: Yura, tv. FBC 
& MSU 2 @ ANIC. 

Diseriburion. see fig, 50, Flight Period: see 
lig, 18, 

Comments: T. angulosa is widespread in dry 
sclerophvll forest and mallee areas in South 
and Western Australia extending to wet sclero- 
phyll forest in southwestern Australia. How- 
ever it is rare in this habitat further east. It 
is sympatric with T. rerraclada over a large 
part of its range A favoured habitat is ihe 
open woodland of the Mt Lofty Ranges where 
Acavia pyenaiha is commonly a deminant oF 
co-dominant shrub. Adults emerge after a 
secession of cold nights in mid autumn and 
persist until late May. 


Thalaina clara Walker 
FIGS 7, (6, [8 21, 31. 41. 50, 55, 
Thaletra clara Walker 1853, po 6605 Meyrick 
Meyrick 142, p. fiS4: Turner 1919, pm. 385. 

Holowpe labelled “Australia 52-39" in 
BMNH#I- 

Adult (fig. 55). Head with frons rounded, 
siloothly huir-scaled, -whitish-huff; vertex 
roughly lmir-scaled, rust colour: jabial palpi 
with terminal segment and apex of secand 
seymient fuscouls, remainder while, anterna 
of male laminate. Thorax white, anteriorly 
erey-white. orange it base of wings; legs with 
fore and mid femora and tibiae and all tarsi 
infuiscated, hind femora. and tibiae white: fore- 


8 P, BL McQUILLLAN 


wing (fiz, 7) wilh costa stratehl, gently archest 
apically, tevmen arched und slightly sinuate 
beneath apex, R, anustomosed with Se and 
again wilh Ro, ground colour shining while: 
narrawly hlack-murgined ted-hrewn streak 
trom base to one-third costa the angled to 
mid-discal areca where it bifurcutes emitting 
one siteak lo fornus thenee ulony posterior 
margin to near base and another streak traciig 
My to mid fermen then back to four-fitths 
Costu. fermen warrowly light fuscous, eilla 
orange posteriorly fuscous; hindwing shining 
white with » rounded fiscous subapical blotch 
hever extending to anul angle, cilia white; 
forewine heneath while, costa und a tnangilnr 
subapical blotch fuscaus, costal edge of this 
blotch oranye, hindwing beneath white, suba- 
pical blotch usually larger (han an upperside 
and orange ahove My; wing expanse 4 
38-48 mm, @ 40-48 mm, Abdomen white. 


Male genitalia (fig. 21) with apex of gnathos 
with small spine; aedeagus (lig. 31) with 
vornuti of twe compound spines one noticeably 
longer than other. 


Female genitalia (lig, 41) with selerotised sec- 
tion of ductus bursae longer than colliculuni. 


Final iistar larva: Head capsule 3.2 mim wide, 
blue-green with small fuscous hlatehes on 
upper half; body bright olive green with four 
wayy whitish lines dorsally; enclosed green 
areas irregularly mottled with blue-green, ox- 
treme posterior margin of cach segment whitish: 
fleshy yellowish lateral Tine irregularly edged 
with pinkish and fuscous below, literal areas 
motled with white and finely speckled black, 
two whitish subventral stripes on most seg- 
ments und diagonally extended to thoracic lugs 
and prolegs, ventral areas pile green with 
several whitish parallel lines; erotehets 13-16 
on Ad, 15-19 on AS, 25-28 on AB gna 2R—335 
en anal claspers, length 33-37 mm, width 
43-5. in, Material: G28. 


Pupa (fig. le) dark brown im colours eremaster 
hooks 6: dorsal anterior margin of AIO with 
2 lobes. Materigl: G28, 


Focdlplant: Acavia deeurrens (1, Wendl) 
Willd. 
Specimens examined: 117 4 57 9 QUEENS 


LAN, Blackbnn, 1-2 UO: Millmertin, vo IM 
1? ANIC. y, 2 2 NMYV, iv. TM 1 et UO) Noo- 
woomha, iv. 2d 1 2 NMY, i, iv, v. wii. EID 
9 f4h UO, » IGM 19 UO: NEW SOUTH 
WALES: Audley, vy. HST | @ ANIC. Barrineron 
House via Salisbury, v, GRM 1 9 UQ; Barryrene, 
iv, HST bY ANIC; 8 km N of Bungwahl, ii 


iFEC & MSU 1 6 ANIC; Church Point, v. IFBC 
1 2 ANIC; 7 km SW of Gosford, ii, LFBC & 
MSU 4 J ANIC: Murnshy, iv. 2 cd 2 9 NMV: 
Killara. iv. vo 3 of o1 9 SMV. Marulan, iv 1 ¢ 
SAM: Millagong, iv, 2 ¢ 2 2 NMV; Mulgoa, LMR 
1 o ANIC, Naat, iv, LMR 4 ¢ 6 2 ANIC; 
Nation! Park, iv, 7 ef? NMYV; Orunge, ii, HST I 
do. GANIC: Bing Creek via Coll's Harbor, v, 
GEM | do LQ; Roseville, iv. LMS 5 & ANIC; 
4) km & of Singleton, ii, FBC 1 of ANIC; 
Sydney, iv. 1 NMYV, Tooloon Serub, tii, FID 
19 ANIC: i iil, FID 7 go 18 UO; Tubrubuces 
Creek, Baritnylon Tops, i. RS | od ANIC, iL 
NMYV; 4% km 8 of Wauchope, iii. [FBC & MSU 
) ¢ ANIC. 18 kin NE of Windsor, iii, IFBC & 
MSU 1 dd 8 ANIC. AUSTRALIAN CAPITAL 
TERRITORY: Black Mountain, ii, it, [FBC 5 
ANIC, Ui, TEBC Lt oof UO; Canberra, iv, TPBC 
19 ANIC: Condor Creck 800 m, ik LEBC LY 
ANIC. 3 ko NW of Lee's Spring 1200 to, di. TFBC 
& BEDE 1 @ ANTC, VICTORIA; Bulwyn, xi. 1 od 
NMV; Betku River, Maltacoota, iii, | 9 NMYV; 
Canterbury, iv, Hl specimens NMYV, v, 1 of SAM; 
Castlemaine. tii. 1 a NMVs Cheltenham, iv. 20 
snecemens WMV: Crib Point, iv. CCl 3 4 2 | 
ANIC; Dandenong. ty. 9 specimens NYIVs Gis- 
borne, Wh, iv. GL 20 specimens NMY, iii, Jy, 
aL 5 f@ | & SAM, iv. GLI @ UG: Hazelwood, 
Wo JHO 1 fd ANIM; Macedon, si., i 2 2 NMYV; 
Melbourne. OBL 2 4 | 2 SAM: Mitcham, iv. 
51 2 NMV: Moe, iii, iw. UCLG 1 4 3 2 
ANIC, iv. RW 2 & ANIC, iv, GCLG 2 7 SAM, 
ivy | d UO; ML Donna Buang, i, 1 ad NMV; MI 
frien, ji 1 9 NMVi Mt Wavetley, LL a NMVy 
Myrudeford, iv, 1 & NMV; Oakleigh, 1 9 NMV: 
Otway Ranges. i, PRMcQO 1 1 & PRMLQEC: Sale, 
iv, 2 f NMV> Springvale, iv. vw 6 specimens 
NMYV, vo 1 of SAM, iv. ERY | ¢& UG, Tara 
Valley Natl Park, ii, PBMcQ 2 & PRMeQC; 
Thurra River ov. Cape Eyerard, iii. bot NMY, 
lrurulgon, iv. 2 do 2 2 NMV; Wakroanga, iv, 1 2 
SAM; Wandil, 2 a NMY, Woori Yallock, iis. 
! ff NMV. TASMANIA: Mole Creek, iv, TRE 
2 Ss IMAG. SOUTH AUSTILALIA: Adelaide. 
OBL 2 Gf SAM), Narucoorte, iii, iv. IOW | 
J of ANIC. Waoilyille. ix. TB 1 4G SAM- 
Distribution: see fig, 50. #livhe Period: see 
fiz Ls 

Conunents: T chira ranges over much of the 
same type af habitat as TT. selenwea, however 
Ws presence in Tasmania and near Adelside 
requires confimmution- 


Thalaina daseripta Walker 
FIGS 8, 8, 22, 32. 42. 50, 46. 
Vhaliind tiveripia Wilker 1855, p. 661; Meyrick 
(892, p, 655; Lower (893. po 290 Carvay 
Titrner 1919, py, 388. 
Phisuruyera principéria Herrigh-Sehatler (855, pl, 
7h, fi. 446, 


REVIEW OF THE MOTH GENUS THALAINGA Y 


Absyres principaria Mernich-Sehitler, Guende 
We57. p. 227, 
Thitaina hicreglyphica Lower 1593, p. 289, Gold- 
finch 1944, p, 191. 
Types ot inseripras lectolype 2 labelled 
“V DLL, 51-153" in BMNH_ bereby desiz- 
nated, | @ | © paralveturypey, 1 @ lubelled 
“VDL. 54-9" in BMNH, hereby desigttated, 
L @ Iabelled os for lectolype, in BMNIM, 
hereby designated, 
Type of principarias not in Zoolagicul Lasti- 
tule, Martin Luther University, Wittenberg, 
GDR (N. Gosser, pers, comm), 
Holawpe 4 of hlereglyphica \ybelled “April 
‘93, 906 Blackwood = 906 7. Iiieroglyphica 
Lower” in SAM, 
Adult (Fig, 56): ead with [rons rounded, 
smoothly hairsealed, whitish-butly vertex with 
rough hair-scales, pale fuscous; labial palp with 
terminal scament and most of second segment 
pale fustous, second segment beneath with 
long white seales; antenna of male larminate.. 
Thorax grey-while becoming whiter pasteriorly, 
pule fuscous at base of wings; lorewing (lig. 
8) with costa nearly straight. apex produced; 
termen strangly urched and sinuate beneath 
apex: Ry anastomosed with Sc and again with 
Koy shining white. markings ochreous 10 pale 
fuscous, narrowly margined darker: streak 
from base to one-third costa, then extending ta 
mid dise where it bifureates into an Upper arm 
reaching mid termen thenee to costa at three- 
quarters. and g lower arn reaching to tornus, 
thence thickly ylong posterior margin to base, 
termen with small semi-circular markings at 
the extremities of M.., CuA,, and CuAg and a 
larger triangular blotch below apex, ciliy pale 
fuseous; hindwing shining while with a 
moderate fuscous subapical blotch sometimes 
extending ta termmen and costa; forewing be- 
neath white with upperside martinis visiblet 
coastal half of these being fuinrly outlined with 
pale Tuscous below, hindwing beneath while, 
subapical blotch reproduced und usually larger: 
wing expanse 4 3844 rom, ¢ 40-42 mm, 
Abdomen white with pale luscous shadings 
ahove, 
Male venitilia (iy. 22) with tegamen rather 
nurrows wedeaeus (fig 32) with cornuti of two 
subequal compound spines, 
Fenjule genitalia (lig, 42) with papillae onales 
rather large: ductus bursuc relatively short. 
Final instar larva: (modified frony Lower 
1893), Length 26 mm Head capsule 2.3 mm 
wide, pale green with small fuscus blotches 


gerass upper frons: body green, lightly Necked 
darker, a mid darsal stripe af denise blackish 
speckting; twa line dorsolateral whitish stripes; 
fleshy whitish lateral tue irregularly blotehed 
with fuscous above and below; a thin whitish 
subventral Suipe; ventrally pale green with a 
white mid ventral stripe; erotcbets 13-14 on 
A4, 12 on AS, 25 on AG and 28-29 on anal 
claspers, Muterial: One specimen “Tas., 15 
kim NNW of Buckland, 19 viii, 1980, On 
Aecieia mearnsii De Wild, P. B. MeQ.", in 
TDA. 

Pipe: mid brown in colour, 6 eremaster hooks; 
dorsal anterior margin of ALO with 2 lobes. 
Material; One spevimen labelled as above, but 
“pupa 3 ix, 19KO"> in PDA. 


Foodplanis: Acacia decurrens (I. 
Willd... 4. mearnsii De Wild, 


Specimens examined: 58 & 2b 9. QUEENSLAND; 
Yeppoon, i. [EBC | @ ANIC; NEW SOUTH 
WALES 1.4 km S$ of ML Tinderry 1600 mm, ai. 
IEBC & MSU ] 4 ANIC; 5 km SF of Pilot Hill, 
Buco Forest, Bulow, ili, TGC 1 go ANIC: AUS- 
TRALEAN CAPIEAL TERRITGRY= Blindells 
Creek Rd. 1000 mi ii. TRBC To of ANIC; VIC- 
TORIA: Bendigo, iy. 1 2 NMV; Castlemaine, iv. 
4 9 1 9 NMV: Gippsland, tii, iv. 30 specimens 
NMY¥: Gisborne. iii, av. 4 ¢ 2 9 SAM, iii, iv. 
Mi specimens NMV: Macedon, iii. 2 9 NMYV; 
Melbourne, iv, 1 & NMV; Tuoljnyi, iv, 3 df 1 
NMYV; Wandin, iv. 5 specimens NMV, TAS- 
MANTA: Bellerive. ii, BM 1 4 ANIC, Cressy, 
iv. 1 @ (DA: Fern Tree iii, RIH 1 & TDA; Hell- 
yer Gorge ii. IFBC & MSU | 2 ANIC; Kingston 
iv. IRC 2 7 UO, iv. IRC 2g ANIC, iv. IRC2 4 
49 TMAGs iv, JRO 147 NMYV;L. Leake 1300 m, 
i, JPRC & MSU | G ANIC; 16 km W of May- 
dena, ti. lt & NMV3 Mit Banow 800 m, iii, JFBC 
& MSU 1 @ ANIC: ML Field Natlonal Park 160 
m, ji. TFFRC & MSU 1 @ ANIC: ML Nelson 700 
m, wi. WW. PRMeQ 5 ¢ 3 9 TDA: Mi Wellington 
2711 m, j.. ii ait, iv, BM 13 of SP ANIC, Ouse, 
iit, wv. 3 f TDA: Pyengana 310 m, aii, FBC & 
MSU 2 2 1 9 ANIC: Ridgeway, i. 1 gd NMV~ 
brevallen. iii, RON 1 co TRA; 21 km S of West- 
bury ui. [FBC & MSU 1 7 ANIC. 


DPisrrihurion; see tig. 50. Plight Period: sez hg, 
18, 


Comments T. inseripta ts aun Interesting. species 
with larvae adapted in colour pattern to living 
on bipinnate Acacia food plants. Tt is the 
most colc-tolerant of the genus, being the 
commonest species over much of Tasmania 
where jt occurs in open woodland up to 
TOO0 mm; it also Oceurs in the Australian Alps 
up to 1800 m. ‘The single record from central 
Queensland is noteworthy in view of The ab- 


Wendl.) 


10 P. B. McQUILLAN 


Figs 9-11. Venation of Thalaina spp. 9. T, paronycha; 10. T. allochroa; 11. T. kimba. 


pro meso Al A4 


Fig. 12. Setal map of final instar Thalaina clara. 


sence of records from northern N,S.W. and 
southern Queensland. 

Adult moths were abundant in an open 
eucalypt woodland in which Acacia mearnsii 
was the dominant tall 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 flight 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 al rest on the trees in the dappled 
light when looking against the sky. 


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


A5 


A6 A7_ A8 AQ A10 


12 


Muacgueenia chionoptila Turner 1947, p. 102. 
Types: lectotype ¢ labelled “Milmerran, Q, 
25 APR. 1936 J. Macqueen Macqueenia 
chionoptila Turn. TYPE” in ANIC, hereby 
designated; 4 4 4 @ paralectotypes, 13, 
3.9 “Milmerran, Q. 145-31 J. Macqueen”, 
1 ¢ “Millmerran, Q. 27-4-31 J. Macqueen”, 
1 3 Millmerran, Qld. 27 APR. 1935 J. 
Macqueen”, 1 4 Milmerran, Qld. 8 May 1935 
J. Macqueen”, 1 @ “Millmerran, Q. 20-4—31 
J. Macqueen Genitalia M8iS P.B. McQ. 
1978" in ANIC, hereby designated. 


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


REVIEW OF THE MOTH GENUS THALAINA im 


& Fp 


; fee fs 


17MM 


Fig. 13. Head capsule of final instar Thalaina 
angulosa. 


Thorax greyish-white on anterior third, re- 
mainder white; legs white with exterior of fore 
and mid Icgs infuscated; forewing (fig. 5) with 
costa straight, apex pointed, termen slightly 
sinuate beneath apex and hardly arched, R,; 
often connected with Sc by short bar and some- 
times anastomosed with R.,; ground colour 
above shining white, orange-brown spot at base 
of costa, extreme costal, termen and posterior 
margins orange-brown, cilia orange-brown; 
hindwing shining white; subapical and sub- 
tornal fuscous spot present, cilia white, 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 4 48-52 mm, ¢ 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, furca 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. 
macfurlandi. 


14. T. selenaca: 15, T. angulosa; 16. T. clara; 17. T, 


35 
Oo +—+ selenaea 18 
iw 30 *—+ clara 
Fa =—* angulosa 
& *—e tetraclada 
oO ' inscripta 
chionoptila 
w 
x 
= 
[e) 
= 
Le 
° 
lu 
o 
<q 
- 
z 
Ww 
oO 
ia 
Ww 
a 


JAN —————— FEB ———__——. maR ———_—— apr 


2 P. B. McQUILLAN 


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


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


Immature stages not recorded except that larva 
pupates underground. 

Foodplant: Acacia harpophylla Benth. 
Specimens examined: QUEENSLAND: Millmer- 
ran, iv.. v. JM 5 & 4 2, ANIC; Millmerran, iii., 
iv,, v. JM 28 specimens, NMV: Millmerran, iv.. 
v. JM & specimens, QM; Millmerran, iii., iv., v. 
JM 4 ¢ 8 3, SAM; Millmerran, iii, iv., v. JM 
5 ¢ 4 9, UQ. NEW SOUTH WALES: Trangic, 
iv. RL 1 d, ANIC. 

Distribution: see fig. 49. Flight period: see 
fig. 18, 

Comments: T. chionoptila is mainly known 
from a Jong 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 tetrauclada (Lower) comb. nov. 
FIGS 1, 18, 24, 34, 44, 51, 58. 
Amelora tetraclada Lower 1900, p. 406. 


Thalainodes tetraclada Lower 1902, p. 231: 
Turner 1919, p, 386; Wilson 1972, p. 123. 

Thalainodes nessostoria Turner 1919, p. 386; 
Wilson 1972, p. 123. svn. nov. 

Holotype § of tetraclada \abelled “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. 


Aduit (fig. 58): Head with frons naked and 
bearing rounded projection (fig. 1); 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 tegulac and sides of thorax white; 
legs white, exterior of fore and mid femur 
and tibia and all tarsi infuscated: forewing 
with costa nearly straight, apex produced, ter- 
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: 
faintly black-margined orange or tan streak 


REVIEW OF THE MOTH GENUS 


: 19 


Figs 19-21. Male genitalia, 19. T.. selenaca; 20. T. 


from base to just above mid lermen tracing 
M. and emitting similar streak from about 
one-fifth to one-third tracing CuA., to termen: 
streak along posterior margin from near base 
to tornus, termen narrowly orange or tan, 
cilia orange or yellow; hindwing shining white; 
large fuscous subapical blotch, projection of 
which often touches termen or is continued 
subterminally to anal angle, cilia white; fore- 
wing below white, lightly infuscated basally 
and below CuA., costa and termen narrowly 
oranze or tan: fuscous subapical blotch pos- 
teriorly orange or tan; hindwing below shining 
white: maculation as above; wing expanse d 
38-48 mm, { 38-46 mm. 

Male genitalia (fig. 24) with uneus rather 
long, gnathos with a large apical spine bearing 
numerous smaller ones, furca rather linear and 
both forks of equal length; aedeagus (fig. 34) 
curved basally, cornuti of two groups of about 
3 spines, 

Female genitalia (fig, 44) with apophyses an- 
teriores rather short, colliculum longer than 
sclerotised band of ductus bursae. 

Specimens examined: 37 3 33 2 NEW SOUTH 
WALES: Broken Hill, vy. OBL 2 & SAM; Bourke, 
12 AM: SOUTH AUSTRALIA: 10 km SW of 
Tron Knob, y. [FBC 4 4) 10-9 ANIC, 64 km E of 
Nullabor, iii, IFBC & MSU 1 4 ANTC; 89 km E 
of Nullabor, ve IFBC & MSU 1 d. 1 @ ANIC; 


THALAINA 


angulosa; 21. T. clara. Scale lines | mm. 


46 km SW of Whyalla. v. IFBC 4 9 ANIC; 
Whyalla, v.2 2 NMV; WESTERN AUSTRALIA: 
27 km SE of Coolgardie, iv. IFBC & MSU 3 ¢ 
2 9 ANTC: Dalwallinu, iv. LEK | specimen 
WAM; Dumbleyung, iv. HU 2 specimens WAM; 
Kalbarri NU Park, iv. IFBC & MSU 2 ¢@ ANIC; 
Kojonup, ii, RIP 1 2, iii, RB 1 dy iv. ALR 3 dy 
iv. RIP 6 & 2 9 iv. MMHW 3 461 «&, v. RIP 
[ ow ALR I & ov. MMHW U oy all ANIC; 
45 km W of Madura, iv. IPFBC & MSU 2 @ 
ANIC: Merredin, LIN 1 specimen QM; 46 kn 
W of Merredin, iv. IFBC & MSU 1 ¢@ ANIC, 
29 km W of Mogumber, iv. IFBC & MSU 1 2 
ANIC; 97 km E of Norseman, iv, IFBC & MSU 
1 2 ANIC: 24 km N of Northampton, iv, JFBC 
& MSU 3 4 1 9 ANIC: Pithara, iv. FBC & MSU 
1.442 ANIC; Tammin, v. 2 9 NMV. 


Distribution: see fig. 51. Flight period: see 
fig 18. 

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


14 P, B, McQUILLAN 


Figs 22-24. Male genitalia. 22. 7, inscripta, 23. T. 


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


Thalaina macfarland’ (Wilson) comb. nov. 
FIGS 2, 17, 25, 35, 45, 51, 59. 
Thalainades macfarlandt Wilson 1972, p. 123. 


Types: holotype 2 labelled “NTHN. TERR., 
148 km S of Alice Springs 26 April 1966 N, 
McFarland at uy light’ in SAM; «/lotype and 
paratypes, see Wilson (1972). 

Adult (figs 2, 59) adequutely deseribed by 
Wilson (1972); wing expanse § 38-44 mm, 
Q 38-50 mm. 

Male genitalia (fig. 25) with tegumen and 
valva elongate, apex of gnathos with large 
recurved spine bearing some smaller ones, 
furca goblet-shaped, juxta elongate: acdeagus 
(fig. 35) With apex produced, cornulus a 
sclerotised plate bearing small marginal spines. 
Female genitalia (fig. 45) with very long duc- 
tus bursae and without sclerotisation. 

Eee ovoid, pale green with an elongated brown 
blotch, 1.0 mm long x 0.8 mm wide. Material: 
GI180, 


Pinal instar larva: Head capsule 3,3-3.4 mm 
wide, pale green with white band across vertex 
hearing many small fuscous blotches; hody dull 
green, white lateral stripe strongly developed 
on pro- and mesothorax and again on A6 to 


chionoptila; 24. 7. tetraclada, 


AY but usually poorly developed or absent on 
intermediate segments, spiracles black and 
placed above this line. L setae on A2 to A7 
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 
A8& only; crotehets 13-14 on Ad, 14-15 on 
AS, 28-30 on AG and 45-48 on anal claspers, 
those on hind claspers tending to be biordinal: 
length 27-31 mm, width 5.0 mm. Material: 
G180. 


Pupa (fig 17) pale orange-brown; cremaster 
hooks 2; dorsal anterior margin of ALO with 
2 lobes; length J} mm, width 4.5 mm, 
Material; G180, 


Foodplants: Unknown, but larvae thrive on 
Acavid pycnantha as a surrogate host 
(MeFarland 1979), 


Specimens examined: 14 § 16 &. QUEENSLAND: 
Cunnamulla, ve. NG Lo’ AM; 24 km S of Mill- 
merran, iv. [FBC | 2 ANIC, NEW SOUTH 
WALES: 30.508 146.33E 23 km SSE of Byrock, 
vy. EDE & MSU 1 @ 22 ANIC: Cobar, iv. 
vy, WIR 2 2 ANIC; 29 km E of Vauehan 
Springs HS, vi. KIMcK 1 2 ANIC. SOUTH AUS. 
TRALIA; 27.188 133.25KF Ammoroodinna Creck, 
vy. PBMcQ | & PBMeQC; 160 km NNW of 
Coober Pedy or Wintinna, v. NMcP & TN 1 of 
5AM; 61 km S of Kulgera, v. NMck & TN I 
SAM, NORTHERN ‘TERRITORY; 19 km FE of 
Alice Springs. v. NMcF & TN 3 @ ANIC: 1 2 
19 AM, | 2 BMNH, 1 & 1 & NMYV: 45 km 
WSW of Alice Springs, Vv, NMcF & TN 1 ¥ 
SAM: 148 km S of Alive Springs, iv. NMecF 


REVIEW OF THE MOTH GENUS THALAINA \5 


Figs 25-28. Male genitalia. 25. 7. macfarlandi; 26. T. kimba; 27. T. paronycha; 28. T. allochroa. Scale 
lines | mm, 


16 P. B, McQUILLAN 


| 2 SAM; 24.208 131.35E Amadeus Basin nr 
Reedy Rockhole, vi PR 4 9 ANIC; 23.485 
132.21E 5 km NE of Gosse's Bluff, vy. HP 4 ¢ 
1% ANIC, 

Distribution: see fiz, 51. Flight Period; late 
April to early June. 


Comments: The dispersed nature of the few 
locality records for T. mac/arlandi suggests 
a wide distribution in central Australia. 


Thalaina kimba sp, nov. 
FIGS 11, 26, 36, 46, 51, 62, 63 


Types; holotypé & labelled “32.518 141.376 100 
km S by B of Broken Hill, N.S.W. 3 May 1976 
1LF.B. Common E. D, Edwards: genitalia slide 
M&s46 PBMct) 1978" in ANIC, 10 of 8 @ para- 
types, 2 & same data a8 holotype, in ANIC; | of 
labelled “6 miles S.W. of Iron Knob, S.A. 7 May 
1970 I,F.B. Common: genitalia slide M842 
PBMcQ 1978" in ANIC; | @& labelled “6 miles 
SE. of Gol Gol, N.S.W. 5 May 1970 T.F.B, Com- 
mon” in ANIC; | of labelled “100 km SE of 
Broken Hill, N.S.W. 29 April 1976 1.F.B. Com- 
mon E, D, Edwards” in ANIC; | & labelled “ 1 
km NNW of Goolgowi, N.S.W. 5 May 1976 


ieaian ————_ 1LF.B. Common E. D. Edwards" in NMV; | ¢ 

3 a Same data as for previous specimen in SAM; 2 

“x same data as for previous specimen in ANIC; 

34 _ 1 df 1 @ labelled “33.238 141,40E 82 km NW of 
a Wentworth, N.S.W, 28 April 1976. 1.F.B, Common 
Pa NE OS Ol E. D, Edwards" in BMNH; 1 9 labelled “31,498 
SN, 141.12E Umberumberka Reserve, 9 km NNW of 

\ A Silverton, N.S.W. | May 1976 I.F.B. Common 

—_ EB, DB. Edwards” in ANIC; 1 9 Jabelled "30.508 

35 a 146,.33E 23 km SSE of Byrock, N.S.W. 8 May 
“Se _ tS 1973 E. D. Edwards & M, 8. Upton: genitalia 


slide M845 PBMcQ 1978" in ANIC; 1 @ Jabelled 
“Mambray Creek Nat. Park, S.A. 11 May 1970 
LF.B, Common; genitalia slide M843 PBMcQ 
1978” in ANIC; 1 & Jabelled “30 miles SW of 
Whyalla, S.A. 9 May 1970 T.F.B. Common: geni- 


a ee TH talia slide M822" in ANIC: 1 @ labelled “Kimba 
eal —~]) 12.5,1963 R. E. Harris” in SAM; 1 9 labelled 

: “Minnipa, S.A. May 1970" in PBMcQ; 1 & 

37 - a labelled “Moorunde Wombat Reserve, — nr. 
—— = Blanchetown, S.A, 12 May 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-sealed, greyish 
with suggestion of fuscous transverse bar be- 


a \ . hind antennae; labial palpi with terminal seg- 
i ment white; antenna in male shortly bipec- 
Imm SA linate. Thorax with anterior and posterior 


thirds fuscous-grey, mid-third and tegulae 


Figs 29-38, Aedeagi of Thialaina spp. 29. T- sele- 
naea; 30. T. angulosa, 31, T. clara; 32, T. in- 
seripta; 33, I. chionoptila; 34. T. tetraclada; 35. 
T. macfarlandi, 36. 1. kintha; 37. T. parenycha; 
38. T. allochroa. 


pale grey; legs infuscated, except hind femur 
and tibia; forewing (fig, 11) with costa straight 
in male, slightly recurved in female, termen 
sinuate beneath apex and strongly arched, R, 


REVIEW OF THE MOTH GENUS THALAINA 17 


Figs 39-43. Female genitalia. 39. T. selenaea; 40. T. angulosa; 41. T. clara; 42, T. inscripia, 43. T. 
chionoptila. Scale lines 1 mm. 


18 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. 


REVIEW OF THE MOTH GENUS THALAINA 19 


ve 


selenaea t 


49 


chionoptila © 


angulosa 


50 


Clara 


inscripta 


allochroa 


51 paronycha 


O 
A 
macfarlandi & ny 
tetraclada @ 

| 


kimba 


Fig. 49. Distribution of Thalaina selenaea and T. chionoptila. 
Fig. 50, Distribution of Thalaina angulosa, T. clara and T. inscripta. 
Fig. 51, Distribution of Thalaina allochroa, T. paronycha, T. macfarlandi, T, tetraclada and T. kimba. 


20 P. B. McQUILLAN 


anastomosed with Se, ground colour white, 
markings suffused grey and sprinkled with 
bluish-white scales, base of costa blackish; 
very broud black-margined streak from one- 
third costa dilated posteriorly to mid dise 
where it bifureates, one atm to mid lermen 
thence angled to four-fifths costa and extend- 
ing to half costa, other arm to tornus thence 
along posterior margin to near base where it 
is mixed with black; broad ijnudented streak 
along termen, cilia brownish-grey becoming 
grey below Ma; hindwing transtucent while 
with several irregular terminal und subterminal 
fuscous blurches; forewing bencath white with 
streak along posterior margin ubsent, discal 
streaks narrower and more sharply defined 
with fuscous, termen streak as above; hind- 
wing beneath White, a subapical and a suh- 


tornal blotch only; wing expanse 38-44 
mm, 9 40-404 min, 
Male genitalia (fig, 26) with valva rather 


broud, apex of gnathas with 3 or 4 small 
spines, furca long and sinuate with one branch 
poorly developed; acdeagus (fig. 36) with 
cornull of single stout spine and group of 3 
or 4 spines. 

Female sentalia (tig, 46) with colliculum 
square, sturigma well developed, ductus bur- 
suc very lone its proxinial third wilh thick 
folds. remainder thinly membranous, 
Distribution: see fig. 51. Probably. widespresd 
in mallee habitats from Eyre Peninsula 
through the Murray Mallee to western New 
South Wales. 

Flivht period; late April to early May, 


Vhnlaing paronyeha (Lower) comb. nav, 
FIGS 9, 27, 37, 47, SI, 61, 

Amelora paranyeha Lower 1900, p, 407, 
Vhaluinades parenyeha Tower 1902, p, 2315 
Toner 1919, p, 386; Wilson 1972, p. 123. 
Holotype > labelled “3460 TYPE, Broken Hill 

24.5.98" in Lower's hand, in SAM, 


Adulr (fig. 61): Head with frons caked, 
hearing moderste truncate projection with 
longitudinal rib beneath; vertex wilh ochreous 
hair-scales which extend almost to extremity of 
frontal projection; labial palpi whitish: antenna 
of male strongly bipectinate. Thorax ochreovs 
above, whitish below; legs tinged ochreous, 
fore-ubia very short, beariig apical spines Tore- 
wing (fig, 9) with costa straight, apex pointed, 
fermen Njoderately arched, Ry auastomosed 
with Sc, ground colour shining white, markings 


dark Gehreous or tan finely edged wilh brawn, 
extreme costal edge white slightly broader 
from one-sixth to one-half costa; costal 
sircuk from base te apex narrowly continued 
along termen and slightly extended hut 
broader along, posterior margin) streak from 
One-quarter costa to lermen whove middle trae- 
ing M; thence angled on termen to cesta at 
three-quarters; diagonal streak from half discal 
streak To just wbove torhus, cilia ochreous; 
hindwing shining white tinged oehreous, pole 
fuscous subapical spot, cilia white; forewing 
bencath white, tinged ochreous on basal half 
und on margins, pale fuscous diagonal suba- 
pical spot, hindwing beneath white, subapical 
spot slightly larger and darker than above, 
small faint fusceus spot near fermen between 
CuA, and A,; Wing expanse 4 36-40 min, 
? 42-44 mm Adbomen ochreous. 


Male yenitalia (fig. 27) with tegumen broad, 
apex of gnathos elongate ynd bearing lonet- 
tudinal row of about 6 stout spines. yalva 
relatively long, furca reduced to small lobe: 
wedengus (fig. 37) strongly curved, cornti 
ol single spine and another group of fused 
spines. 


Female genitalia (fig. 47) with apophyses an- 
leriores rather short, » band of selerotisation 
al top of duets bursae similar in length to 
colliculum, ductus bursae very long. 
Specimens examine 11 4 26 2 NEW SOUTH 
WALES: Broken Hill, iv. ve OBL 10 26 F 
SAM, 11 specimens NMV) SOUTH AUSTRA- 
LIA; Ammeroodinna Creek 27.188 133.25E, vy, 
PBMcQ | o& PBMecOc, 


Distribution: sec fig. St. Plight period: mid 
April to lute May- 


Comments: This and the next species ure 
rather isolated fram the rest of the penus by 
siructiiral features such as genitalia, presence 
of @ spine on the fore lihta and the farm of 
the frontal process, 


Thalaina alloelirea (lower) cambh, nov. 
FIGS 10, 28, 38, 48. 51. an 
Thilainedes ullochroa Lower 1902, p, 23%: 

Turner 1919, p 387) Wilson 1972, p, 123, 
Holaivpe labelled "3.5.02 Broken Hill = 
3779 > Amelora allochrou Lower TYPE” in 
Lowers hand, in SAM, 

Adult (fig 60) with head as tor T. paronvelia: 
vertex With Oehrequs-bulf hair-scales whieh ex- 
tend onto Frontal projection; labial palpi white: 


REVIEW OF THE MOTH GENUS THALAINA 21 


61 


Figs 52-63, Adults of Thalaina spp. 52. T. selenaea 3; 53. T. selenaea var. punctilinea 3; 54. T. angu- 


62 


losa ¢; 55. T. clara 3; 56. T. inscripta 3; 57. T. chionoptila 3; 58. T. tetraclada 3; 59. T. mac- 
farlandi 3; 60. T. allochroa 3; 61. T. paronycha &; 62. T. kimba holotype 3; 63. T. kimba para- 


type 2. 


antenna of male strongly bipectinate. Thorax 
ochreous-buft 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 8 2 9. NEW SOUTH 
WALES: Broken Hill, v. OBL 1 ¢ 1 9 NMY, v. 
OBL 6 ¢ SAM; Mootwingee Historical Site 
31.14S 142.18E, v. IFBC & EDE 2 @ ANIC. 
SOUTH AUSTRALIA: Ammaroodinna Creek 
27.188 133.25E, v. PBMcQ 1 o& PBMcQC; 
McDouall Park, v. FWJ 1 2 SAM; Tallaringa 
Well, v. PA 1 9 SAM; 107 km S of Coober Pedy, 
v. NMcF 1 2 SAM. WESTERN AUSTRALIA: 
Giles, v. 2 6 PBMcQC; Skirmish Hill, 1 ¢ SAM. 
Distribution: see fig. 51. Flight Period: May. 


Comments: T. allochroa exhibits a radical de- 
parture from the basic colouration of the 


22 P, B, MoQUILLAN 


genus but structurally it ig virtually indistin- 
fuishable from parenyeha, Wis just possible 
that allaehroa is only a Mendelian sexregute 
of paranyeha but breeding experiments are 
heeded ta conlirm this, Thongh iofrequenily 
collected, both species appear to be widely 
distributed within the 15-25 em isehyets sourh 
of 25° § Jatitude. 
Diseussiun 

‘There are no vonsistent venulional, genitalic 
or other structural differences which justify 
the separation of Thalainodey Lower and 
Mucyueenia Turner from Thaleine Walker, 

The presence of un areole in the forewing is 
not correlated with any other features of 
diagnostic value heyand species level, An 
urcole iS present in one form in selenace, in 
another torm in chionoptlla., clara and in- 
seripla and is absent in the rest, Lowers im- 
pression of a longer cell in the hindwing of 
his Thalaineades is erroneous 4s mdasurement 
will Show. 

Some diversity exists in the male aotennac: 
they are laminate with ventrally produced, 
ciliated sexments in selenwea, inseripta and 
clara, shortly bipectinate in angulasa. mraefar- 
fandi and Ababa. und strongly bipectinate in 
allachroa, paronycha, chionoplila and terra- 
clade. Earlier authors have prevunusly over- 
looked the slight but delinite bipectinate nature 
of the antenmic of aivulexa, 

The corneous frontel process appears to 
have arisen independently several times in the 
genus, as it has in many other arid 2one genera. 
This structure, ii conjunction with the fore 
tibial spines in allechroa, paronyeha and 
kKimba, probably assists the image te fired its 
way to the surface of the soil aller emerging 
fron) the buried pupa. 

A qumber of biological fearures are 
stared, All are late summer to late autumn 


fliers with an annual lifeeyele. Wis very 
likvly that eggs of all species are stimulated to 
hateh by rain as angulesa is (McFarland 
1973). Larvae are associated with Acavia (or 
less often Cassa) and where known, complete 
their feeding in early spring and pupate just 
below the surface of the ground where they 
over-summer. Pupal aestivation is a phenome- 
non shared by many autumn-flying ennomines 
in southern Australia, such ag the species of 
Chlenias (Madden & Bashford 1977). Taa- 
laine has successfully exploited a very wide 
ranve of habitats in the southern half of Aus- 
tralia, a few species having adapted to each 
major ecological zone within the overall range 
of the geous. 


Previous attempts to fragment the genus 
have been based on either incorrect evidence 
Or are Unnecessary, smee several other genera 
(eg one to contain alfochroa and puronychea) 
could be erected on similar evidence. There- 
fore, I feel it is better to slighty expand the 
original definition of Thalaina to contain all 
of the above species, thus reflecting their close 
relationship, 


Acknowledgments 


1 am indebted to Dr fF. B, Common, 
C.S.1LR.0., for helpful discussions and com- 
ments on the manuscript tind to Mr D, §. 
Fletcher, BM(NH), for generous assistance 
with information on type specimens and litera- 
jure. A generous grant from the Royal Society 
of South Australia Research Fund enabled me 
jo visit the Australian National Insect Collec- 
lion, Canberra, for which £ am grateful 
Thanks are also due to the curators who 
loaned material or sent information. to Mr M- 
Poteeiter for photographs, and to Dr J. H- 
Szent lyany and my wife Kathryn who helped 
in various ways, 


References 


Drsmarest, M. B. (1858) Wr Chenu, I. C.. Enev- 
clopddie d'Histoire naturelle —Tuble alphabe- 
tique: Lépidoptéres. ( Paris.) 

Doverepay, E. (1845) Descriptions of some new 
Australian Lepidopterous insects, Jn Bo 7. Ryre. 
dis ypidns Dise, cent. Aust. 12 Appendix pp. 
437-439, pl. 5 

Piercura, DS. (4979) The generic names of 
moths of the world. Vol 3. Geometroiden- 
1. W. B Nye fed.). (British Museum, London) 


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never oder wenig bekannter asserenropaischer 
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pla figs 1-551, 

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243, 

(1900) Descriptions of mew Atstralian Heie- 

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cies Of Australian Lepidoptera. Prune. Ry Sac. 

SN. Aust. 26, 212-247. 


REVIEW OF THE MOTH GENUS THALAINA 23 


Lucas, H. (1857) Im Chenu, J. C., Encyclopedie 
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40 pl., text illustr. (Paris. ) 

McFarianp, N. (1971) Egg photographs depict- 
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Res. Lepid. 10(3), 215-247. 

(1973) Some observations on the eggs of 

moths and certain aspects of first instar larval 

behaviour. [bid. 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. 

Mappen, J. L. & Basurorp, 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. 

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678. 


THIERRY-MieG, P. (1899) Descriptions de lépidop- 
téres nocturnes. Ain. Soc. ent. Belg, 43, 20-21. 

Turner, A. J. (1919) Revision of Australian Lepi- 
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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- 
112. 

WALKER, F. (1855) List of the specimens of lepi- 
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(1865) List of the specimens of lepidop- 
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33, 123-124, pl. 11. 


A NEW SPECIES OF MANAYUNKIA (POLYCHAETA) FROM 
EPHEMERAL LAKES NEAR THE COORONG, SOUTH AUSTRALIA 


BY PAT HUTCHINGS, PATRICK DE DECKKER & MICHAEL C. GEDDES 


Summary 


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. 


A NEW SPECIES OF MANAYUNKIA (POLYCHAETA) FROM 
EPREMERAL LAKES NEAR THE COORONG, SOUTH AUSTRALIA 


by Par HurcHinas*, PArrick Dr Deckkert? & MicnAni C, Gepbesy 


Summary 


Huieninos, P., De Drexerr, P, & Geoors, M. C. (1981) A new species of Munayunkia 
(Polychaeta) from ephemeral likes gear the Coorong, South Australia, Tram, Ry Soe. s. 


Aurs. WSL), 25-28. 12 Sune, L9St 


The polychaete Manaynnhia athdlassia asp. is described from ephemeral lakes adjacent 
ta the Coorong Lagoon, South Australia. This is the first record of this genus from Ais- 
tralia, Manavunkla athalassia iy active over a wide range of salinities (27-95%) and persists 


in dry lake beds during the summer months, 


Introduction 


The (una of athalassic (non marine) saline 
lakes in south-eastern Australia has been listed 
in several studies (Bayly & Williams 1966, 
Bayly 1970, Geddes 1976, Williams 1978, 
De Deokker & Geddes 1980). The only record 
of polychaete worms is from a small lake tear 
Lake Eliza, South Australia where Cerarenc- 
reis eryihiraeensis Fauvel and Capitella capi- 
tate (Pabricius) were collected (Bayly 1970), 
These species are known to tolerate a wide 
range of salinitigs but cannot resist desicca- 
tion. During a recent study of 23 ephemeral 
lakes lear the Coorong Lagoon, ‘South Austra- 
lia (De Deckker & Geddes 1980) another 
polychacte, a mew ypecies of Manayunkia 
was common in several localities. 


Munayunkia athalassta n. sp. 
FIG. | 
Type material 

Holotype (Aust, Mus, WL7671) from ephe- 
meral lake at 36°19'46"S, 139°44'48"E, adja- 
cent tu Coorong, S.A. coll. P. De Deckker & 
M. ©, Geddes 17.vii,1978. 

Paratypes from various ephemeral lakes 
adjacent to Coorong (20 specimens, AM 
W17672), United States National Museum 
(] specimen USNM 63370), British Museum 
(Natural History) (1 specimen ZB L980: 
265), Further material in Australian Museum 
(9 speeimens AM W17677). 

Tentucular erawn colourless, anterior body 
up fo setiver 3-5 darkly pigmented, rest of 
body colourless, Thorax of eight setigers, 


* Australian Museum, P.O, Box A2sS, Sydney 


South, N-S.W_ 2000. 
* Department of Zoolugy, University of Adelaide, 


1 Present address: Depurtment of Biogcowraphy & 
Geomorphology, Australian National Univer- 
sity, Cunberra, A-C.T. 


abdomen of three setigers, All tharacic setigers 
similar in size, abdominal setigers cqual in 
length, slightly shorter than thoracic setigers. 
Tentaucular crown of paired symmetrical halyes 
with semicircular bases, ¢ach hall with two 
short eompact radioles. Outer radiole with 
four short stumpy pinnules and inner radiole 
with three, originating close to the base of the 
radioles, Radioles ciliated and not conneeted 
hy web-like membrane. Ventral paired palps 
vascularised, smooth and with skeleton: palps, 
thicker than tadioles bot similar in length. 
Prostomium bluntly rounded with pair of 
pigment eye spots, visible only after removal 
of tentacular crown. Perisfomial collar well 
developed ventrally, triangular with rounded 
apex: collar developed laterally as narrow rim 
and present dorsally as two small lappets. 
Setiger 1 with notasetae only, subsequent 
setigers with noto- and neuroselae. Notosetac 
of two kinds, basically hroad bladed capillaries 
and hastate setae: thoracic newrosetae long 
handled hooks, abdominal ieurosetac long 
handled uneini with six ac seven harizontal 
rows of teeth, each row with four or five 
tecth, ieeth not perfectly aligned in rows, 
fewer teeth per row towards apex. Number 
of noto- and neurosetae per setiger shown in 
Table 1. Triangular pygidium with no pygidial 
eye spots. 

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


Ecology 

Manavunkia athalassia occurs in ephemeral, 
athalassic, saline Jakes adjacent to Coorong 
Lagoon, South Australia. The lakes, in which 
M-_ athalassia occurs, are not connected to the 
sea, and are characterised hy fluctuating 
salinities und varying walter levels, due to 
the raising and lowering of the saline water 


26 PAT HUTCHINGS, PATRICK De DECKKER & MICHAEL C. GEDDES 


1mm 


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. 


A NEW SPECIES OF MANAYUNKIA (POLYCHAETA) 27 


Number of totesatie anil neureseiue 
per setiger, 
SSS 
Nolusetac Neurosetie 
capillaries hastate Gall uncini) 


TABLE |, 


eliver 


absent 
4 
4 
4 
4 
3 
2 
4 
10 
12 


at 


Thoracic 


Ph fit ed 


Abdominal 


Lad Ld fad Un ed dd a 


1 
2 
3 
¢ 
3 
6 
E 
8 
1 

2 
3 


table and to seusonal precipitation. A descrip- 
tion of the physical and biological features is 
given hy De Deckker & Geddes (1980), The 
polychaetes collecled by Bayly (1970) were 
not found in this stucly. 


Manayunkia athalassia lives in translucent 
gelatinous tubes, in soft clayey carbonate scdi- 
ments which some times. contain shell debris 
(ostracods, gastropods). Occasionally Mf. arfta- 
laysla was found living in a colany 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 
months. Even when the lakes are dry some 
moisture may be trapped by hygroscopie sall 
erystals, and an occasional cover of dead 
aquatic plants such as Lepilaena sp. and 
Ruppia sp. and, less frequently, the alga 
Lamprothamnion papulosum on the surface 
of the mud. However, the summer air tem- 
peratures may exceed 40°C. Tn February 1979 
distilled water was added to samples of mud 
collected from dry lakes where Af. athalassta 
occurred the previous season, Within one day 
active adults were seen at the wide range of 
salinities of 27-95". Further Jahoratory 
studies showed that it could be maintamed in 
wn aquarium of 82: For several months 


Comments 

Manayunkia athalassla differs. from con- 
weners in being found in emphemeral saline 
environments isolated from the seaz the other 
species are found in marine or brackish water 
conditions. Af. brasiliensis Banse, 1956 col- 
lected in mangroves in Canaéa in Brazil may 
experience same hyper-marine salinities m the 


interstitial waler during low tide, bur no 
salinity measurements are given. 

Manayurkia athalussia also differs morpho- 
logically from the other species, M, caspica 
Annenkova, 1928 has six ar seven pinnules 
per radiole and M. speciosa Leidy, 1859 has 
six radioles with ahout 60 pinoules per radiole, 
Southern (1921) suggests that the arrange- 
ment of the filaments, which appear to be 
altached jn groups to a short common stem, 
is un artifact and probably due to the contrac- 
tion af the basal membrane, However. we 
Suggest that the filaments. arise from a short 
commoo stem as in congeners, but we have 
not examined the type. M- polaris Zenkewitsch, 
1935 has equal numbers of pinnules on both 
radioles, whereas M. athulassia has three pin- 
nules on the inner and four on the outer 
radivle. M. aestuarina (Bourne 1883) has 
four pinnules per radiole with the paired palps 
considerably longer than the pinnules, whereas 
in M. arhalassia the palps and the pinnules are 
similar in length. 

Detailed setal counts are given only for M, 
hraviliensis and M. polaris, M4. brasiliensis has 
more setae of all kinds on both abdominal and 
thoracic seligers than MM. athelassta, Ty addi- 
tion Mf, brasiliensis has capillary setae of two 
different lengths whereas AY. athalassia has all 
capillary setae of similar Jength. Ad. poralis 
has considerably more abdominal uncini than 
M. athalaxsin, For these reasons M-_ athalassiea 
is described as a uew species, The specific 
name refers to the type af saline lakes. in 
which this species lives, 

Only three genera of the subfamily Fabri- 
cinae have been recorded from Australia: 
Desdemona, Fabricia and Oriopsis (Day & 
Hutchinus 1979). Hartmann-Sehraeder & 
Hartmann (1979) have described an Oriapsts 
sp. and Fabricivae gen. and sp. indet, from 
Port Hedland. Western Australia. Manayunkia 
dthalassia is the first record of this genus 
froin Australia, but several other species of 
the subfamily Fabricinae are present in Aus- 
tralia and shortly will be desertbed by Hut 
chines, Earlier, general marine collections 
overlooked the Pabricinae because they are 
small and offen have specialised habitat 
requirements, 


Acknowledgments 


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


28 PAT HUTCHINGS, PATRICK De DECKKER & MICHAEL C. GEDDES 


References 


ANNENKOVA, N. (1928) Ueber die pontokaspis- 
chen Polychaeten. 2, Die Gattungen Hypaniola, 
Parhypania, Fabricia and Manayunkia. Ann. 
Mus. Zool. Leningrad 30, 13-20. 

Banse, K. (1956) Beitrage zur Kenntnis der 
Gattungen Fabricia, Manayunkia und Fabriciola 
(Sabellidae, Polychaeta). Zool. Jb. (Syst.) 84, 
415-38. 

Bayty, |. 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. bid. 17, 177-223. 

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

Day, J. H. & Hurcrines, P, A. (1979) An anno- 
tated check-list of Australian and New Zealand 
Polychaeta, Archiannelida and Myzostomida. 
Rec. Aust. Mus. 32, 80-161. 

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


Geppes, 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 Eulitorals der aus- 
tralischen Kiisten unter besonderer Beriicksich- 
tigung der Polychaeten und Ostracoden. (Teil 
2 und Teil 3). Mitt, Hamb. Zool, Mus. Inst. 
76, 75-218. 


Leipy, J. (1859) Manayunkia 
Acad, nat. Sci. Philad. 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, 
563-659. 


WILLIAMs, W. D. (1978) Limnology of Victorian 
salt lakes, Australia, Verh. Internat. Verein 
Limnol. 20, 1165-74. 


ZENKEWITSCH, L, (1935) Uber das Vorkommen 
der Brackwasserpolychaete Manayunkia (M. 
polaris n.sp,) an der Murmankiiste. Zool, Anz. 
109, 195-203. 


speciosa. Proc. 


DISTRIBUTION OF PINNA BICOLOR GMELIN (MOLLUSCA: BIVALVIA) 
IN SOUTH AUSTRALIA, WITH OBSERVATIONS ON RECRUITMENT 


BYA. J. BUTLER & M. J. KEOUGH 


Summary 


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. 


DISTRIBUTION OF PINNA BICOLOR GMELIN (MOLLUSCA;: BIVALVIA) 
IN SOUTH AUSTRALIA, WITH OBSERVATIONS ON RECRUITMENT 


by A. J. Butter* & M. J. Keouau" 


Summary 


Buitea, A, J, & Keoudi, M. Ss 


(1981) Distobulion of Pinna bicalor Gmelin (Mollusca* 


Bivalvia) in South Australia, with observations on recruitment, Vrans. R. See. 8. Ast. 


105(1), 29-39, 12 June, 1981, 


A divite survey wus conducted in January L980 at 43 sites from Port Broughton in 
Spencer Gulf to Ceduna in the Great Australian Bight, to obserye the distribution of the 
bivalve Pina bicolor Gmelin, its density, habital-types and associated species, Eather records 
front Investivator Strait, Gulf St Vincent und Spencer Gulf are also reported, At tl sites 
samples were jaken lo determine distributions of shell length, counts of growth checks and 


wonad slates, 


Although P, bicalor is widespread in suituble habitats throughout South Australia, as 
distribution is ‘patchy’ on Large and small scales. Recruitment is shown to vary m space and 
time and the significance of this in the ecology of the specics is discussed, 


lutroduction 


The ceology of Pinka bicolor Gmelin is af 
intrinsic and practical interest (Buller & 
Brewster 1979) und we huve been studying 
both the population ecology of the bivalve 
(Butler & Brewster 1979) and the epibiota on 
its shells (Kay & Keough 1981, Keough 1981!) 
al a few sites in Gulf St Vincent. However, 
there is po systematically collected intorma- 
tion about the distribution and habitat-types of 
this species throughout the rest of the State. 


It is well known that the ‘recruitment’ of 
many marine organisms, especially those with 
pelagic larvae, is variable in both space and 
lime, By ‘recruitment we mean entry to the 
popolwtion al a size such that they can he de- 
tected or captured—in (his case, sten by a 
diver, This is not the same as ‘settlement’ fram 
the plankton, because newly-settled larvae may 
die before they ure delectable, Varlability in 
reeruitment may be extremely important in 
the ecology of such species (¢.g, Bowman & 
Lewis 1977, Keough’, Sutherland 1974, 
Sutherland & Karlson 1977). Although varia- 
bility in reeruitment is reasonably well docu- 
mented for certain commercially tmportant 
species (e.g. Loosanoff 1966, Andrews 1979) 
there is a paucity of pubished data about its 
occurtenee in a wide variety of organisms, and 
a paucity of detail about the spatial and tom- 
poral scales Of ‘patchiness’ in recruitment, 


© Department of Zoology, University of Adelaide, 
Box d98 GP... Adelaide, S. Aust. 5001. 

1 Keough, M. £ (1981) Dynamies of the epifanna 
of the bivalve Pinna pbicelur Gmelin. PhO 
thesis, University of Adelaide (submilted). 


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, 


Plina bicolor is such a species, and in South 
Australia it is at the southern edge of its 
tropical and subtropical range (Rosewater 
1961), so it 18 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 
thar their density appears to be non-uniform 
and ta vary now-randomly over areas that ap- 
pear to an observer to be uniformly suitable 
(eg. various papers in Coull 1977), Tr is, of 
course, possible that the area is Hot in fact uni 
formly suitable, but also possible that the 
animals ate absent from some habitable sites, 
perhaps as a result of ‘patehy’ recruitment, 
Again, this phenomenon requires documentae 
tion as a first step in its study, ff is important 
fo produce distribution maps showing con- 
firmed absences as Well as records of a species, 


This paper reports a survey dewizned to pro- 
Vide general observations on the distribution 
of P, bicolor in South Australia, the habitat- 
types in which it occurs, the organisms as- 
sociated with it and the regolarity of its re- 
cruitment, Cerlait conelusions can be drawn 
from these general observations made at one 
time; detailed explanations must depend upon 
long-term observatians and expevimental Lests 
ol hypotheses, 


40 A.J. BUTLER & M, J, KEQUGH 


Methorls 

Pinna hes been recorded from depths 9% 
geal as 30 oon the Noor of Gulf St Vincent 
(Shepherd & Sprigeg 19746) but ihis survey was 
confined to areas within 2 km of the share and 
depths no more than 18 m, Our object was to 
Visit us many as possible of those sites where 
Pinta had been reported or where it might 
have been expected to occur fron) the lype 
of botton) add degree of exposure. Our as- 
sumplion was simply that Ping requires a 9of{ 
boitom and no more than moderate wave-ex- 
posure, That part of the South Australign 
coast from the Victorian border ta Backstalre 
Passage 1s not discussed here, Before this str 
vey we had many records from Gulf St 
Vincent and some fram Spencer Gulf and 
Investigator Strait; those records ure sum- 
marised here, and io particuar we discuss Uuta 
from cight sites scored within two months ot 
the main survey and using the same pro- 
cedures (sites 1-8 in Table 2) The survey 
itsel€ cavered 43 sites from Port Brovghton 
in Spencer Gulf to Ceduna in the Great sus- 
tralian Bight in January, 1980, 

Subtidal sites were surveyed using SCUURA 
from an intlatable dinghy. intertidal sites an 
fool, At every site, position was determijied by 
landmarks and bearings, and a map reference 
recorded. Depth was measured in m hy shot- 
line, Surface and bottom water temperatures in 
“C by mereury thermometer, and the shite of 
lide and current voted. An estimate was noted 
of prevailing wave and current conditions, 
judged from aspect, present conditions and 
Weather, hottam lopography and surroundings, 

The diver swam over o variable (but always 
tecarded) wistanee, making notes on a pre- 
pared data-slate, He recorded a qualitative 
statement of bottom type and dominant maero 
biota and estimated pereentage cover for each 
of the seagrass genera Zoviera, Posidanies, 
Ainphibolis and Halophila, The deraity ot P 
Aicolory was estimated separately for cach of 
two size-classes (dorsa-ventral shell height 
<= 7 om, and height > 7 em in No. m-), The 
observer carried an aluminium rod 1 im long 
fo remind him of seale, and had extensive 
prior expenence of measuring density using a 
| me? quadrat, and so the estimates can he 
taken asx sulficrently reliable for use as an 
ingles of density, The smaller size-class is 
likely to have settled within the last year 
(Butler & Brewster (979}> a1 was scored 
separately to give an index of reeruitment at 
sites where samples were not taken, He alse 


noted qualitatively the size distmbulion, spatial 
distribution, morphology and epibiotg of 
P. bicolor. Densities of the animal apeeies oe 
groups listed below were scored on the fol- 
lowing qualitative scale: none seen, rare, com- 
mon, abundaut. These categories were based 
on previous expericnee af ‘typical’ densities for 
these species, and have different meanines for 
each group. The groups scored were> the bi- 
valves Malleus meriilianus, Chlamyy asper- 
rimus, and C. At/rons, gastropods of the genus 
Polinices and Tamily Muricidac, holothurians, 
echinoids, asteroms. fish and ecphalopods, 
Under the headings Muricids, Urehias and 
Asteroids, and commonly under others, it was 
possible fo identify the particuar species 
recorded. 

At certain sites. which are marked with 
asterisks in Tuble |, in adidiion to all the 
ubove observations, random samples of 
P. bicolar Were collected by clearing a Lo m- 
Wide transect in a tandemly-chosen direction, 
and examined in the dinghy or ashore, Por 
each animal antero-posterior shell length and 
dorso-ventral shell height were measured 1s 
described by Butler & Brewster (1979). Scars 
left by the posterior adductor musele in the 
nacreaus layer of the shell were counted: 
these are counted with error, but the number 
of ‘niajar’ sears appears to be aw index of gee 
(Butler & Brewster 1979) and in this study 
they were always counted hy the same oab- 
server, A crude index of aye iw also avuilable 
from the epibiota of the shell, given a know. 
ledge of the biology of the eprbrotic species 
(Keaugh!), which were recarded in (his stud 
on the qualitative seale used by Butler & 
Brewster (1979) with notes on speeres-compo- 
sition, The reprodictive tissue in Pinna 
spreads diffusely under the mantle anterior to 
the posterior adduetor muscle und is not al- 
ways detectable macroscopically, Its deyelop- 
ment is as yet poorly understood, In this study 
i wis scored qualitatively on the following 
scale; 0, nane visible; P, poorly developed, a 
thin layer of what appears to be gonad visible; 
M, moderately developed, uodoubtedly gonad 
Lissue present ohscuriig unverlying organs; W. 
well developed, massive gonad cancesting 
large area of vndertying argans. 

Finally, for each animal we noted shell 
Jamage due to breakage or sponge boring, anu 
the presence of subtuhular spies on the ex- 
terior of the shell. Table | shows the sites 
investigated, In addition to visiting widely 
Spaced logattons, we commonly sampled 


DISTRIBUTION AND RECRUITMENT OF PINNA BICOLOR 31 


TABLE 1. Sites inspected and estimated densities of P. bivolor, sqwrzmer, 1979-80. *Random samples 

of P. bicolor were collected for measurements, ete. at these sites. + Densilies at these sites were meéa- 

sured using a 1 om? quadrat, Density columns are Jeft hlank for sitey where no Pinna was found. 
§: P. bicolor of H <7 em. Ly P. bicolor of H > 7 em. 


. Depth Lat "S/ S 
Site (m) Long. °E No.m* No.m* 
Na, Location (1—intertidal) 
| 1-2 km W of Semaphore jetty 7 34.83/138.45 1.8 2.5 
2* 3km NW of St Kilda 3 34.73/138.48 0 <<01 
3*;+ 2km E of Ardrossan 15 34.43/137.95 1.87 497 
4 Ardrossan: beacon N of bulk loading jetty 7-8 34,43 /137.93 0 0.2 
5’? Stansbury: on intertidal sand-spit I 34.92 /137.83 171 1.41 
6*+ Edithburgh: site of Buller & Brewster (1979) 7 35.11/137.78 0,20 1.54 
7  ‘Troubridge Island: intertidally on SW side I 35,13/137.82 0.30 >I 
& Wallaroo: to 150 m W from site of old jetty 07 34,93/137.6) 2 5 
9 Port Broughton: over 4 km travelled in channel 
und uround mangrove island 3 33.56/137.92 
10 Chinaman Creek 2 km WSW of shacks 10.5 32.23/137.80 
1! Chinaman Creek 4 32.65 137.80 
12 1.5-2 km WSW of Chinaman Creek 4,5 32.65/137,78 
13= 400 m WSW of Chinaman Creek 3-4 32.68 /137.83 10 5 
14. Chinaman Creek 2-3 32.65/137,82 
15 Port Augusta: Playford Power Station jelly 7-10 32.54/137.78 <U.01 4 
16* Port Augusta: first normal channel marker 
§ from Power Station 75 32.55/137.78 OON 0.5 
17 Franklin Harbour: ca, 400 mS of Cowell jetty 3 33,.70/ 136.94 
18 ~~ Franklin Harbour 3-4 33.70/136,94 
19 Franklin Harbour! 500-600 m E. of jetty 4 33.68/136.95 0) <001 
20 Franklin Harbour: Cowell jetty 4-5 33.68/136.94 
21+ Franklin Harbour: (locally called 
Mr Thompson's Reef) 0.5 33.71 /136.94 0.01 0,85 
22. Tumby Bay: 600 m ESE. of jetty 5,5 34.39/136.12 
23% Tumby Bay: jetly 4 34.39/136.11 0.05 ONS 
24 Tumby Bay! 200 m off end of jetty 5 34,39/136.12 
25" Tumby Bay: 100 m off entrance to caravan park i 34.48/136.11 2 7 
26" Port Lincoln: 300 m 6 of caravan park jetty 13.5 34.73 /1395.89 0 0.7 
27. - Port Lincoln: Kerton Point jetty 9 B4,.72/ 135.88 0,001 0) 
78 Port Lincolf; 150 m off caravan park 3-5 34.73 /135.49 
29 Port Lineoln: 300 1 WSW of Ist port 
channel marker 15-18 34.70/195,88 
30) Port Lincoln 13 34.72/135,87 0 0.001 
4) Coffin Bay: between jetty und point to NW 2.5 34.63/135.47 
32. Coffin Bay; in channel leaving Coftin Bay 4 34.62/13546 
33~=— Coffin Bay I 34.62,/135.46 
34 ~= Coffin Bay: between Goat [5, and other 
side of hay 5 34.62/135,47 
45 Coffin Bay: point ut entrance to Coffin Bay 2.5 34.62/135.46 
36 ~~ Kellidie Bay \-2 34,61/135.48 
37. ~—s Kellidie Bay if 34,61 /135.47 
38 © Coffin Buy jetty 4 34.62/135.47 
39 = Elliston: near jetty 5 33.64/134.89 
40 Venus Bay? near jetty 344 33.234134.68 
at Venus Bay ] 43.23 /139472 
42 Venus Bay: downstream from 2nd upstream 
channel marker 3 33.22/ 134.68 
43 Venus Bay: side channel on way back to jetty 4 33.23/134.68 
44 Venus Bay: Ist upstream channel marker 
from jetty 05-4 = 33.22/134.67 
45 Venus Bay: channel SW of Germein Island 05-3 33,22/134.66 
46 Venus Bay 5-1 33,.23/134.64 
47 - Venus Bay: Ist downstream channel marker ; 
Frorn jetty 3 33.23/134.66 
48" Streaky Bay 1 32.80/134.21 14 3.5 
44 = Streaky Bay! 100 m inshore from 48 | 32.40/134,21 3.3 1 
50% Streaky Bay: 200 m § of Ist outgoing 
channel marker, near Crawford Landing 6.5 32.78 134.23 1.5 15 
S| 3 32.13./133.67 1 1 


ee 


Ceduna jetty 


32. A.J. BUTLER & M, J, SEOQUGH 


several sites separated by short distances 
Within one area or embayment, Since one ob- 
ject OF the survey was to invesnigate the smull- 
seale ‘patehiness’ of P, bicolor, and since one 
object of this paper is to allow future workers 
to investigate changes over time, the locations 
of our sites ure given in ag iach detail as pos- 
sible in Table 1. 


Distribution in South Australia 


Pinna bieplor has been recorded by us at the 
locauions for which positive densities are giver 
in Tahle | and fram Fishery Beach (Fleurieu 
Peninsula). American River Inlet near Mustou 
(Kangaroo tsland), Rapid Bay, Aldinga Reef 
und Price {all Gulf St Vincent), Goose and 
Wardang Islands (Spencer Gulf). Shepherd & 
Sprigg (1976) recorded it at many sites on the 
floor of Gulf St Vingent, and Cotton (1961) 
recorded it fram “Beachport to Fremantle’ 
Thus. the species is widespread on sheltered 
shores or in decper water throughout the State. 
However, nole that f2 bicolor was nat found 
at all of the locatians jn Table | and that sites 
¢lose to one another often differ (e.g. China- 
tin Creek, sites 10-L4; Franklin Harbour, 
sites 17-U1; Tumby Bay, sites 22-25; Pori 
Lincoln, sites 26-30). Note also that in same 
embayments which appeared suitable for 
Pinna, we (ound fone (RKellidie Bay, sites 
31-38; Venus bay, sites 40-47: Flliston, site 
39) or very few (Franklin Harbour, siles 
17-21; Por) Lincoly, sites 26-30). Thus. the 
distribution of #, biceler appears ‘patehy'- 


Morphology 


A few shells found af various sites were 
similar to the species which Cotton (1961) 
identified as Sabitapinne virgata, but almost 
all were typical of his Pinna dolabrata. Both 
Of these were referred by Rosewater (1961) 
to the variable species P. biveler. The relation- 
ship between shell length and shell height will 
he discussed elsewhere, but on preliminary 
analysis it appears not to differ significantly 
anongst all the locations sampled, At any 
lovation some Shells hore more sububulae 
spines than others; these were more prominent 
in young individuals; the typical form at «ll 
locations js fairly smooth-shelled (Cotten 
1961, Figs 68 & 69; Rosewater 1961, Pl 147, 
1S5t & 152), 


Density in diflercut habitats 
This survey did not provide data suited for 


powerlul tests af null hypotheses about the re- 
Jationship between P. bicolor density and such 


variables. as hottam type, depth, current and 
the presefice of other organisms. Nevertheless, 
some vaireme posibilities can be eliminated 
from the available data. Table 1 shows esti 
muted densities at lhose sites where P. bicolor 
was found. PL hicalar uveurred in battant 
sediments ranging from very fine sand fa Very 
coarse sand; we could delect no relationship 
belween our qualitative notes on sediment type 
und the presence, or density, of P. bicolor. The 
‘prevailing or ‘average’ conditions of tempera- 
ture aud eurent could only be estimated 
roughly from our imeusurements and notes on 
a single dive, but again we could not see a 
possible explantion for the presence, density, 
or estimated age-distribulion (sce below) of 
P. bicelor in either of these variables, 

There is no significant correlation between 
P, bicolor density and depth (data in Table 1 
for positive PL Aiealar densities; zero densities 
included for all other sites; ¢ — —O,17, n = 
51, P > 0.05), nor between P. bicolor density 
and the percentage cover of stagrasses (ihe 
jatler transformed to wngles. Rohlf & Sokal 
1969, p. 129; r — 0,02, n ~ 51, P > 0.05), 
Since we already had reason to suspect a oega- 
tive correlation hetween P, Aicalor and sea- 
erasses (unpublished datal, this was rechecked 
by exchiding data pertaining ta embayments 
where P- Afeolor was rare or absent, and where 
one might argue larvae have, for some reason, 
failed (@ arrive (namely Pt Lincoln, Kellidie 
Bay, Venus Bay and Elliston), ‘The correlation 
between PF bicolor density and seagrass cover 
remained non-significant (r - —O.L7, n = 29, 
P > 0.05). 

There Was no significant correlation hetween 
P hieolor density and latitude. (Por sites with 
positive densities, r = 0.37, 1 — 22, P > 0.05; 
for nil sites, F ~ O17, n = 51, P > 005,) 

The densitics of other species which migtt 
conceivably influence P. bicolor, or hive 
similar requirements, were seored ou quualita- 
tive scales, The reasons for scoring these 
species were ous follows, The — bivalves 
Mf, mevidlaiits, C. avyperrintas and C. bilrons 
are ecologically similar to Pinna. Certain 
asteroids prey on F /feolor, The pastropods 
Palinices spp. and probably some muricids are 
thought ta do so. Some fish and cephalopods 
may de so, especiuvlly on small Firne., Hole- 
thurians and echinolds may influence the sur- 
vival of recently-seltled posttarvac. We Were 
inferesied im any hiac of assoeiacions (positive 
or ucgative) between the presence of BP. Ai- 
valor, particularly of recent recruits, and the 


DISTRIBUTION AND RECRUITMENT OF PINNA BICOLOR 33 


TABLE 2, 


Assacidatiiy baween qualitative stares for the densities af P. bicolor and three species af 


epibenthic bivalves, Bach figure in the Table is the meither of sites at which that combination ef seares 
necurred. 


Density of Malleus meridianns 


P hiealor low high 
low <= | im 34 5 
high > 1 m-* 5 3 
x= lor2 «x2 
1 contingency table 1.38 
P >0.05 


abundance of any of these species, Inspection 
of a table of these scores showed no obvious 
relationships wih P, biceler density. Most of 
the data do not warrant statistical analysis, but 
the association between the bivalves P.. bicolor, 
M, meridianus, © axperrimiy and C. bifrons 
was examined further. Scores for each species 
Were grouped into two categories—low' (= 
0 + rare) and ‘high’ — (common +> abundant) 
and the scores for P. hicelor were tested 
for independence of each of the other species 
in three 2 *% 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. bieoler, 


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 alsa because the histology of 
gonad development in P, bicolor has not yet 
been established and related to these scores. 
(This work is in progress.) Nevertheless, if 
the populations at different sites were pre- 
dominantly in different stages of the repro- 
ductive cycle, this might he expected to be 
reflected in the scores. whatever their detailed 
hixtolugical meanings, To test this, we first 
determined for cach of the 12 sites at which 
gonads were examined within the lime-period 
December 1979-January 1980. the minimum 
number of adductor muscle scars at which any 
animal was scored “moderately’ or ‘well de- 
veloped’ (M or W), Next, we considered only 
animals with that number of sears or more, 
and calculated the proportion of theny scored 
M or W, This was done because at some sites 
the proportion of the whole sample with de- 
veloped gonads would be depressed by the 
presence of a large number of very small, pre- 
repruductive animals, Scar counts are used 
bere os an index of age (see below) but 
similar results are obtained if shell length, in- 


Chlanws asperrimus Chlanyy hifrons 


low high Jow high 
39 0 7 z 
7 1 6 2 
0.79 1.30 
>0).05 > 0.05 


stead of sears, is used ta determine which 
animals are polential breeders, 

The results are shown in Table 3_ The pro- 
portions scored M and W show highly signifi- 
cant heterogeneity between sites when the 
Whole set is tested as a 2 * 12 contingency 
lable. However, this may possibly be duc to 
the length of time (more than one month) 
between sampling the first and Jast sites, There- 
fore, consider only the seven sites sampled 
over 10 days (S-15.i.80) ond sampled sequen- 
tially from Franklin Harbour to Ceduna so 
that latitude rose and then fell during the 
period, This set is also highly significantly 
heterogenous, 

There ure 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 contingeney table, cach of 
these pairs is homogeneous for the proportion 
M or W. The proportions scered M and W 
(iranformed to angles) are vot significantly 
correlated with latitade (r = 0.03, n = 12, P 
> 0.05). In summary, despite our erude 
method of scoring gonad condition, it 1s clear 
that siles spatially far apart, even if sampled 
ut about the same time, differ in the propor 
tion of animals in breeding condition, Sites 
clase together in space and time receive similar 
seares. 


Counts of adductor miuscle sears 


Adductoe musele scars are counted with 
error, but Buller & Brewster (1979) argued, 
for sile 6, Tuble |, that major scars probably 
represenl winter geowth checks. This awaits 
confirmation from current work on tagyed 
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 diflereut sites. At one 
site, scur counts are prohahly an index af age, 


34 


Table 3. Genad development in Pinna bicolor 
sampled at 12 sites. Sie numbers correspond to 
Table 1, Gandds were Visually scored as '0'—not 
apparent; 'P'—poarly developed; 'M’—moderaiely 
developed und ‘W'—well developed; these have 
heen pooled into two categories here. Only animals 
in reproductive “age’-class, ay determined by ud- 
ductor muscle scars, are included (see text). 


Sile Date Number Number 

Sampled Oo+P M+W 
3 18.x11.79 0 49 
5 20.xii.79 27 131 
6 13,xti,79 20 130 
13 24.13.80 16 39 
16 231.80 19 Ss7 
21 5.1.80 0 74 
23 7.4.80 Il 19 
25 6.1.80 23 42 
26 9.1.80 24 25 
48 13.1.80 11 43 
50 13.i.80 6 22 
51 15.1.80 5 43 


x" tests for homogeneily: 
Whole 2 % 12 table: x2 84.92 (P<0,001) 
" 


Sites 21,23,25,26,48,50,51) v2 =54,13 (P<0.001) 


A.J. BUTLER & M. I, KEOUGH 


but they do not 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 em. We 
have at present no way to test whether inter- 
tidal animals produce more scars per unit time, 
or simply grow more slowly. However, from 
the data available to Butler & Brewster (1979) 
and various oseryations obtained subsequently 
(Butler unpublished) it seems likely that sears 
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 though 
scars are counted with error a comparison of 
the distributions of scars counts from (wo 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 


Sites 13,16 an 0.10 (P>0.05) are highly significantly heterogeneous when the 
Sites 23,25: x! = 0.017 (P>0,05) whole set is tested, or when only the set 
1 sampled in January 1980 is tested (see G-yalucs 
Sites 48,502 x= = 0,01 (P>0.05) in Table 4). Comparing pairs of sites close 
' together in space and sampling date, we find 
TasLe 4. Frequencies of counts of adduetor-muescle scars in sampley of P. bicolor. 
Site Number of Scars ; _ No. of animals 
0 1 2 3 4 5 7 8 9 10 li S12 in 
sample 
2 3 2 WW 3 3 1 1 2 2 56 
3 105 3 4 1 1 1 3 1 3 3. 2% 154 
5 6 6 22 29 21 16 1421 9 1 4 3 3 164 
6 ao 28- 13 27 9 I7 7 8 Ii ] 150 
13 7 3 2 12 10 13 7 6 2 62 
16 3 4 1 21 16 19 7 3 § 2 l §3 
21 1 3 4 15 20 14 12 3 | 1 i 75 
23 1 Ww 2 3 3 1 3 2 1 I 2 31 
25 2 2 i4 7 aS) 32 9 9 2 1 1 84 
26 I I 4 13 30 49 
48 12 9 8 s 6 8 12 8 2 2 75 
50 9 13 1 3 4. 3 3 6 1 2 4 1 SO 
51 a 715 7 7 12 3 4 3 3 3 2 3 1 70 
Log-likelihood ratio tests far heterogencity: 
Whole 13 % 13 table: G 1156 df, = 144 P= 0,001 
Sites 13-51: G = 4999 dt. — 96 P< 0,001 
Sites 13 & 16: G+ 2415 af. 7 Po 0,005 
Sites 23.& 25: G = 28.57 af, = 6 P= 0.001 
Sites 48 & 50: G 14.87 uf. 8 P > 0.08 
Scar classes 0-5 (0 & 1 pooled) 
All sites except 26 G = 463.8 df, — 44 P< 001 
Sites 13-51 (except 26) G = 153.5 M5. = 28 P= 0,001 
Sifes 13. & 16 G 18.42 df. = 4 0.001 < P < 0.005 
Sites 23. & 25 G= 16.73 af. = 4 001 < P < 0,005 
Sites 48 & 50 G= 7,45 df. = 4 P > 0.05 


DISTRIBUTION AND RECRUIEMENT OF PANNA BICOLOR qi 


Taste S$. Miferevers about density vf reeent re- 
erqitnient and regulary ef reoratimenl over pre- 
vious 5-6 years, based on counts of udiduetor 
niscle scars (Table 4) on assinmptiony that sears 
represent winter grawlle checks, and that pest- 
recruitment mortality rales wand dieir year-lo-year 
variations ure sante at all yites. See texte tor 
methods, O, ne reeruilinent: oi, Wat recruitment: 
M, major reerditment, 4 irregulars Ry regular 
Table alsa shaws inferences abour densis of 
recent reeruimnent based only on divers estimate 
of density of Pinna ef A= 7 ent at sites Where 
Pinna density was > 0, (Table 7). 0, deasin 
~ (5, sparse, 0 < density SO.) net) By dense, 


density > OT nee. 
Site 1978-9 Regularity of Recent 
recrul|ment recruitment recruitment 
from scar from scur from density 
counts COUNTS of small 
animals 

1 Dd 

2 m ! 0 

3 M I nD 

4 Oo 

5 m R D 

6 M I b 

7 1B} 

a D 
4 m 1 D 
is s 
16 m f § 
19 Oo 
21 m i S 
23 0 I 8 
25 ” R 0 
26 i?) 1 ) 
27 S 
30 0 
4s M R D 
ag D 
Si) M R D 
5] M R D 


ee” 


that one is homogeneous (sites 48 & 30), the 
other two heterogencouy (sites 23 & 25, sites 
13 & 16). 

The scar distributions were examined further 
to make inferences aboul recruitment, First, we 
considered the density of recent recruitment, 
Because of the difficulty in scoring the first, 
faint scar the categories O 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 cach site 
into one of three eutegories: 0, no animals 
with zero of one scar; minor, O = p = 20 
where p percentage of the sample having 
zero or one sear; major, 20 < p = 100, The 
results ure shown in ‘Table 5, As for most 
animals with planktonie larvae, the density of 
recruitment is fot expected to be coustant from 
year to year, even if some recruitment always 


oveurss this appears to be borne oul hy Table 
4. Further, the fluctuvtions in density of 1e- 
cruitment do not seem io be in phase at all 
sites. Lf fluctuations in recruiiment Were in 
phase, and if subsequent age-specifie mortility 
rates were also the same, the conspicuous 
modes should be in the same scar-classes at all 
sites. They sre not, us ts shown by Table 4. To 
test this, we considered only the first six scar- 
classes (0-5), because for older animuls we 
have less confidence in the assumption of a 
constant schedule of age-specific mortalities. 
The resulis of Lests for homogeneity are shown 
in Table 4, The whole set is highly significantly 
heterogeneous, as is the set of nine sites sam- 
pled in January 1980, More importantly, two 
of the pairs of nearhy sites sumpled close 
together im time are highly significantly 
heterogeneous (sites 13 & 16 and sites 23 & 
25), 

Next. we asked whether recruitment appears 
to be ‘regular at each sife, that is, whether 
some freeriits appewr each year, albeit at 
varying densities. To de this, high sear-classes 
were ignored because mortality may have 
reduced their numbers so much that sampling 
error heeomes important, The first six sear- 
classes (0-5) were examined; recruitment at a 
site was called ‘irregular’ if there were any vero 
frequenvies in the first six classes, otherwise 
it was ‘resulur. Table 5 shows the result, It 
also shows a ranking of the diver’s esitmute 
of density of small Pina, Note that this is 
un cubsolute’ index hased on the number of 
small animals per m2? of bottom, whereas the 
ahoave method is based on the proportion of 
the sample which was young. Also, an animal 
7 em in shell heizht may, at some sites, he 
several years old, The index based on density 
of small animals is thus of limited value, hut 
a6 ineluded beeause it is available Where sam- 
ples Were pot colleered 

The magnitude of the 978-9 recruitment, 
on either index, appears to have differed be- 
iween sites. Many have irregular recruitment 
Im ane ease n pair of sites whieh differed in 
sear-(requency distributions above (sites 23 & 
25) also differ in their 1978-9 recruitment and 
in their regulurity of recruitment. 


The data collected concurrently with the 
samples give no suggestion of explanations for 
these variations. Neither depth nor percentage 
cover of scagrass was significantly associated 
with regularity of recruitment using either the 
Fisher exact fest (assinming seagrass or depth 
yalues (» two categories) or a two-sample runs 


36 AT. BUTLER & M. J. KEOUGH 


test (ordering the depth or seagrass values and 
then counting runs of regularity scores): ip 
both cases, P > 0.05. Similarly if the scares 
for 1978-9 recruitment were grouped into two 
categories (O+m, M) then they were nat sig- 
nificantly associated at the 5% level with either 
depth or seaprasy cover vsing cither test. 

Regularity @f recruitment was not signifi- 
cantly assoviated with latiiude. This was tested 
by #rauping those sites scored LT and those 
scored R and comparing their mean latitudes 
(thy a 1.07, Pp > 0.2)- 

The magnitude of 1978-9 reeruitmeni 
(grouped into two cafegories) was not signi- 
ficantly associated with regularily of reeruit- 
ment (Fisher exact probability test: P = 
0,085), 

When the notes on associated species were 
grouped into two categories (0 + rare; 
common -+ abundant) and tabulated awainst 
the scores for 1978-9 recruitment or for 
regularity, nO posilivé or negative associations 
were apparent on inspection, and certainly 
none was slatistically significant at ihe 5% 
level using Pisher exact tests, 

The density of small animals (hE < 7 em) 
is positively correlated with that of larger ones 
(both estinjated ra sit by the diver). For sites 
where any P. bivolor were found, Pearson's 
tT > 049. P <= 0.05: Spearman's ep = 0.76, 
P < 0.001. This test was repeated, excluding 
sites 5, 7, L3. 21, 23, 25, 48, 49 hecause their 
length-scars relationship showed that animal of 
H = 7 cm may have more than two. scars, 
and thus (he density of small animals may nor 
be ab estimate of the density of recent receuit- 
ment, The conelusion remained the same 
(Pearson's © = 0.54. P < 0.05; Spearman's 
p= 0.75, P = 0.005) 

However, regularity of recruitment was not 
significantly associated with total density either 
by a 2 X 2 contingency table with density 
classified as < 2m? or > 2 m® (Fisher exact 
fesi- P = 0.085) or by a runs test ag used 
above for depths (P > 0.05), 


Discussion 

This survey hos provided a distribution map 
for # Aicoler \n South Australia. Jt is based 
on visits ta many siles apparently suitable in 
having low wave-action with soft hattoms 
There is a temptation to assume that whe a 
species has been recorded at two points, it may 
be expected to occur in guitable habitals in be- 
tween (thus, distribution maps are often 


hatched), but thut seems not to he so in this 
cuse. The distribution ig patchy; P. bicolar 
is absent from some apparently suitable sites. 
The patchiness oceurs on a local scale; &. 
hicolar may be found on some but pot other 
dives On apparently similar bottoms within 
I km or so—eg., sites 10-14, 17-21, 22-24, 
26-30. Rur it js iho evident on a larger 
scale; the species seems lo be absent from cer- 
tain large snd apparently habitable embay- 
ments (Kellidie Bay, Venus Bay, Elliston), 
though present in olhers north and south of 
them. Note that these are well-enelosed em- 
bayments; perhaps the current patterns are 
such that the arrival of planktonic Jarvac there 
from outside is a rare event. If so, then by 
chance a recruitment might occur from ume 
to tiie and establish a temporary ‘population’, 
This seems to hive happened at Port Lincaln 
(site 26). The reason tor giving the details in 
Tuble J is to document this patehiness: later 
warkers might want to check the same loca- 
tions, 


Organisms ure rarely if ever distributed 
evenly. Some of the Unevertiess i their distri- 
hutions wan be explained by an understanding 
of their ecology (Wwe can say why the un- 
occupied sites are unsuitable or have not been 
colonised). hut there may tTemain a compo- 
nent which cannot be explained, even tenta- 
tively, with existing knowledge, The possibility 
remains that the vacant sites are unsuitable 
or ingewessibie, but the reasons ure not at 
present known, We: shall call this “unexplained 
patchiness’; P. lifcelor provides an example. 


There is no detegtable relationship between 
density of P. Aicalor and sediment type, cur- 
rem! regime, water depth (Table 1), cover of 
seagrass, or axsoctaled wunimal — species, 
especially other ecologically similar bivalves 
(Table 2), One might not have expected = 
compelitive interaction of uny importance 
helween these bivalves (Stanley 1977), but 
perhaps their ecological similarity, of even the 
facet that Mallews and ©. ayperrimus use 
P. bicolor for attachment, might have led ta 
#& posilive association Nove is cvident. We 
note in passing that the other three species 
of bivalves were, like P, bicolor, more often 
scored low than high in density even though 
many sites appeared suitable, and any diver 
knoWs ihat they can be abundant. These 
species. too, appear ‘patchy’. 


Species which are tither predators of 
‘malenitiics (Andrewartha 1970) might be 


DISTRIBUTION AND RECRUITMENT OF PINNA BICOLOK 3 


expected a prlori to have most of their in- 
fluence on younger stages of P, bicolor. still, 
We note that they showed no association with 
the density of P, bleeler large enough to be 
seen hy a diver. 

The above is based on imprecise data 
(mostly subjective rankings) and sa there is a 
possibility that real associations exist hut were 
nol dereeted. However, one might have ex- 
pected such associations to be at least 
noticeable in the kind of data we collected, 
even if they were not statistically significant: 
no trends, however slight, were apparent. Thus 
we conclude that with respect to the presence 
and densily of P. bicolor we are observing 
wnexplained patchiness. 


The proportion of the population with 
developed gonads appeats less “patehy’. It 
differs between sites even considering only 
those sampled close together in time. but 
sputially-close sites (pairs of sites in the same 
embayment) do not differ significantly (Table 
3). We cannot infer that these populations are 
in the sume phase (hecause we do not know, 
for example, whether a gonad scored ‘P’ is 
developing or spent) but it seems likely. 


Counts of adductor-muscle scars were 
heicrogeneous between sites, including some 
nearby pairs. These scars probably represent 
checks in the growth of ibe animal, but the 
reasons for the checks, and their periods, are 
nat known with certainty. There is reason to 
vssume that they represent winter growth 
checks and our interpretation of the counts 
was hased on that assumption, On that assamp- 
tion, the ave-distribulions of the standing 
populations of P, bicolor st different sites Cin- 
cluding some nearby pairs) differ, We 
examined those distributions in more detail 
und found that the proportion of the popu- 
lation with lmw scar counts (recent recruits) 
differs between sites inchiding nearby ares 
(Table +4). In other words, the density of 
recriits relative to thal of adults varies. Pur- 
ther, the presence or absence of whole chisses 
(interpreted as ‘regularity’ of recruitment ) 
differs between sites. The absence of an entire 
class is a stringent eriterion of irregularity’, 
given the error in counting rings, It seems clear 
that reeruitment Muctuates fram year ta year, 
and the fluctuations are not in phase at all 
sites, nor necessarily even at nearby  sifes. 
This contrasts with the proportions with deve- 
Joped gonads, which were similar at nearby 
siles, 


Recent recruitment and the regularity of 
recruiiment were not correlated with depth, 
cover of seagrass, nor with each other. “Rexu- 
larity’ was not significantly ussociated with 
toll density of P. bicelor as estimated in No, 
m~. 

No relationships could be detected between 
recruitment and the densities of associated 
species, This is not te swy that the associates 
have no effects. Firstly. it as possible that their 
abundance is Correlated with the recruitment 
af P. hicdlor hat our data are too imprecise to 
detect ik Secondly, they may moye about, 
so that their abundance at a particular place 
and time hears little relationship to thetr effects 
on P, bicelor there at some earher time 
Thirdly, their effects may be masked by other 
variables, especially the density of settlement 
of P, bicelor, The lack of correlations in our 
data does eliminate the grassest hypotheses, 
eg. that dense bolothurians will, by killing 
newly-settled larvae, lead to sporadic recruit- 
ment, 

The above discussion concerns reeruits as 
proportion of the population, Actual densities 
of recruits would be of interest. The oanty 
relevant data we have ore the divers in sid 
estimates of the densities of two size-classes, 
The density of small snimals is positively 
correlated with that of lurge ones. considenng 
all sites Where P. Aicolar were found, How- 
ever. this may be an artefact, because although 
an animal of H = 7 em at sile 3 would very 
likely be under (wo years oht and probably 
under one (Buller & Brewster 1979), this will 
not necessarily he true at all sites (above) - 
However, if we eliminate sites where animals 
huve a large scar count far a given length. we 
sfill find the same canclusion: density af small 
P. hicalar is positively correlated with that of 
large ones. This stems to be rather in con- 
trast to the conclusions. drawn above from the 
scar counts. However, it is consistent with 
them if the events leading fo reeruitment are 
viewed as follows. 

Larvae of Pinnidae can travel long chistances 
in the plankton (R, S$, and A. Secheltema, pers. 
comm.) Thus, the fact that animals breed ar 
all sites docs not guarantee that settlement 
(still Jess, recruitment) will ocenr at all sites, 
and those larvae which settle al a site miry nol 
have been spawned there. Larvae move about 
with the currents and may well be distributed 
patchily within the water (sce reviews in 
Steele 1978), Thus, their probability of suc- 
cesstul recruitment ut a given benthic site 


38 A.) BUTLER & M, F, KEOUGH 


depends firstly on their being carried there on 
a current of suituble strength, etc. for settle- 
ment, ald secondly on subtle properties of the 
hottom (which may vary from time to time), 
the presence or absence of mobile or ephe- 
meral predators, the availability af food fer 
newly settled post-larvae (which itself may 
depend upon planktonic patchiness and on the 
Vagaries of the currents), and so on, We stress 
the term ‘probability’, The mere fact that 
recruitment is partly dependent on currents, 
on the shapes of land-masses and chinnels ond 
on the topography of the bottom, will mean 
that sites differ consistently in the probability 
that larvae will settle. This can account for 
some very —s well-enclosed = embayments 
apparently contumnag few or no P. hivelor, 
and for a correlation between the densities af 
adults and young, but it leaves recruitment as 
a random variable with a large variance which, 
on the present state of our knowledge, we 
cannot explain, Most sites receive Variable, 
and some even irregular, recruitment and we 
cannot explain or prediet this using depth, 
associated species, sediment type or latitude, 
Recruitment is a major “mystery stage” 
(Spight 1975) in the ecology ol many specics 
with pelagic larvae (eg Andrews 1979, Milei- 
kovsky 1971, Sastry 1979, Underwood 1979), 
Tt is important hecause it varies so widely. 


We cin hope to understund the ceolozy of 
the species wfter suceessful reeruitrment, but 
Frecrimtment itsell ix the main event that ceter- 
Wines the density of such w species atu given 
site. Whilst iL muy be possible in some cases 
to predict recruitment from independent 
variables which influence larval survival, our 
data (especially the very small-scule ‘patehi- 
ness') give no encouragement thar 1 will be 
possible in this cuse, For purposes of a general 
understanding of the ecology of the system (ar 
for Jong-lerm planniag. if the species were a 
commercially important one) we may make 
use of a probability distribution for reeruit- 
ment. For purposes of shortterm prediction, 
the only eourse is to monitor recruitment 
directly. as done for commercial species (ec. 
Brand et a/, 1980, Phillips 1972, Phillips & 
Hall 1978). 

The above should not be taken as an asser- 
lion that nothing can be known about the 
reeruitment of Pinna. Knowledge of a probu- 
bility distribution can be powertul, Those 
species which interact with Pinnta—feed on 
its young, live on its shells, ete—must be 
adapted to that probability distribution, lis 
a challenge fo ecology to produce useful 
models for systems in which miny of the 
important events have probability distributions 
with large variances, 


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Anprews, J, D. (1979) Pelecypoda: Ostreidac- 
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Bowman. Ro S. & Liew, FOR. CL977) Annual 
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Burrk. AL J & Brewsrok, FL I. (1979) Sire 
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bicolor Gmelin (Motlosca: Enlamellibranchia} 
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CrLark, G, RK. (1974) Growth lines in invertebrate 
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Corros, B.C. (1961) “South Australian Mol- 
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luide |. 
Court, BO (1977) (EG ‘Beolegy of Marine 
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MiLmkovsky, 5, A. (1971) 
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Fisheries and Oeesnography, Report No. 98 
pp, 1-18 

kobe, Bo, & SakaL, RoR, (196%) “Statistical 
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DISTRIBUTION AND RECRUITMENT OF PINNA BICOLOR 39 


Sastry, A, N. (1979) Pelecypoda (excluding 
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‘Reproduction of marine invertebrates. Vol. V. 
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SIEGEL, S. (1956) ‘Non-parametric statistics for 
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Spicut, T. M. (1975) Factors extending gastro- 
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the fossil record’ pp. 209-50 (Elsevier, Amster- 
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STEELE, J. H. (1978) (Ed.) Spatial Pattern in 
Plankton Communities. (NATO Conference 
Series IV Marine Sciences 3) (Plenum, New 
York) 

SUTHERLAND, J. P. (1974) Multiple stable points 
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& KARLSON, R. H. (1977) Development and 
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THE GIANT RAT-KANGAROO PROPLEOPUS OSCILLANS (DE VIS), 
(POTOROIDAE: MARSUPILIA) IN SOUTH AUSTRALIA 


BY N.S. PLEDGE 


Summary 


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 (rP3); four upper molars : 
tM! (described for the first time), tM?, two of rM? or *; a lower incisor (1],) and four lower molars : 
1M>, rM3 or 4, IMs, rMs. The “deciduous” molar M! is rectangular and quadritubercular, showing 
greater similarity to the M' 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. 


TUE GIANT RAT-KANGAROO PROPLEOPUS OSCILLANS (DE VIS), 
(FOTOROIDAE:MARSUPIALIA) IN SOUTH AUSTRALIA 


by N. S. PLEDGE* 


Sunimary 
Prooce, N.S. (1981) The Giant Rol-Kangarog Propleopes oscillans (De Vis), (Patoraidae: 
Marsupialiad in South Austhilia, Trams, R, Soo. S. Ause, 1O8(1), 41-47, 12 June, 1981. 
Tholated feel peferuble to one juvenile Propleopus oseillans have been found scattered 
in a cave deposit at Naracoorte, They have heen interpreted as a lower premolar (rPs); 


folir Upper molars : 


rM! (deseribed for the first time), rM®, two of rM* or 4) a lower 


incisor (11)) and four Jower molars + Mo, eMy or 4, IMs, ¢M5. The “deciduous” molar Mt 
is rectangular ond quadritubercular, showing greater similarity to the M! of Berongia spp. 
than to Mypsipemnodon. The form ef the incisor indicates that this tooth hud a period 
of continued ‘rootless! growth until maturity. A sarge humerus is referred tentatively to 
P. oxcillans. and body proportions are calculated. P. eveilans was an animal of about ihe 
same bulk us ant Eastern Grey Kangaroo (Macropus giganicus), but much stocker and with 
lonper legs. Apparently il inhabited dense scrub. living on a diet of herhaceous vegetalion, 


ogeasional carrion and small animals. 


fotroduction 


The kangaroos, wallabies and their allies 
{the Macropodoidea), contaihs many specics, 
living and fossil; some are quite commen and 
others exceedingly rare, Modern taxonomists 
(e.g. Archer & Barlholomar 1978) split this 
large group into two Families: the kangaroas 
and wallabies in the family Macropodidie, 
and the rat kungarons (patoroos, bettongs, 
ete.) in the family Potoroidae, The Jatler 
family is divided into the subfamily Potoroinac, 
containing potoroos and bettongs, and the 
subfamily Hypsiprymnodontinae. containing 
the living Musky Rat-kangatoo Hypsiprym- 
nodon nioschatuy Ramsay and the extinct 
Propleapus species, 

Modern A. mnosehatuy is a small, rat-like 
animal living in restricted areas of rain forest 
in northern Queensland, [t ts distinguished 
from other kangaroos by a combination of 
several primitive characters (Ride 1961, 
1964), such as a simple alimentary canal, the 
presence of the halltix of the inner side of the 
foot, ihe presence of an Upper canine and a 
second lower incisor, and relatively simple 
bunodont molars. These fealures are combined 
with a number of specializations. The most 
sirikins: of these are the large ‘plagiaulacoid? 
premolars, ie. secant (bladed) 
which are larger than the adjacent molars, 
with a curved and serrated cuiling edge, and 
faces. strengthened with a number of parallel, 
verticul ridges corresponding to the points of 


+ South Australian Museum, North Terrace. Ade- 


laide, S, Aust, 5000. 


premolars, 


the serrations, Such premolars are, in fact, 
most distincuve tecth (Woods. 1960). 


The fossil species of Propleopus are believed 
to be the closest known relatives of Hypsip- 
rymmodon, A have large secant premolars, 
although in Propleopus these are more than 
three times the size of the modern teeth The 
molars are alsa similar. The Mountain Pigmy 
Possum, Burramys parvuy has similar premo- 
lars but a different molar structure and a jaw 
structure that precludes macrapodoid affinities 
(Ride 1956. 1964), Despite the relatively 
lurge size and robustness of the fossi) teeth 
and jaws, however. P. oscillans is known from 
only a handful of spegimens. By comparison, 
kangaroos of similar size from the same depo- 
sits often are represented by hundreds of speci- 
mens, Nevertheless. occurrences of Propleopus 
are widespread, Woods (1960) 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.S.W_ (Woods 1960) and 
another from 1, Menindee (Tedford 1955. 
1967). A-second Pleistovene species P. ehilla- 
gocnsis has been found in cave deposits at 
Chillagoe. norih Queensland (Archer ef al, 
1978). Archer & Burtholomai (1978) mention 
a specimen from a Pliocene deposit in northern 
N.S.W. This dentary may be conspecifie with 
a single isolated molar (Gill 1953, 1957; Ride 
1964) from a sub-hasaltic Pliocehe depasit 
near Hantilton, Vic. dated al 4.3 million years, 


42 N.S. PLEDGE 


Williams (1980) lists two additional speci- 
mens of #. aseillans, found recently in South 
Australia at Hookina Creek (P22425) and 
fjcal Mi Gambier (Green Waterhole, P20815). 
These have been inspected, but are under 
study elsewhere so only the cheek-tooth jnea- 
surements Will be given in this paper. 

Tooth vomenclature follows the system ol 
Archer ((978)) the total dental formula for 
Propleopus Would probably be dls, 14, dC4, 
C! Pi M/S. The first molar M1 and pre- 
molar P2 ure replaced by the erupting P3, 


New South Australian necurrence 

Rich Pleistocene bone deposits are currently 
being excavated iti caves al Naracoorte, ‘These 
deposits have produced large and varied Los- 
sil faunas of mammals, together with associated 
amphibians, reptiles and birds (Williams 
1980), One of these deposits partially filled 
and sealed a small cave, the Henschke Fossil 
Cave, that Was discovered by quarry opera- 
fions on the edge of Naracoorte township in 
1969. Excavations of the deposit were under- 
taken by me and continue to yield an abun- 
dance of fossil bones, Associated charcoal hus 
been collected, and preliminary radiocarbon 
results give the deposit am age of around 
35.000 years for the upper metre or so of 
sediment, (SUA-I40, depth 105-120 em, = 
35.000 years BP, 8Cl! 997,724.00; SUA 243, 
31-75 om, 33.800 — 7) years BP, scl) 
—985,.1 + 3.9), 

Macropodoids constitule more than balf of 
the Henschke Fossil Cave assemblage, and 
potoroids are well represented. Amongst 


WORTH 


these specimens are several isoluted teeth, 
found over a period of eight years, thal may 
be relerred confidently to P. oscillans, Vhis 
small sumple comprises one lower premolar 
(rPs) and seven molars, one of which | con- 
sider to be a deciduous molar (m!). An un- 
usual lower incisor, by elimination from all 
other taxa in the assemblage, also appears to 
be P, oscillans, Most of the specimens consist 
of enamel crowns only, They show little or no 
wear, indicating a juvenile age for the in- 
dividual(s) represented, There is no evidence 
in the form of duplication to suggest more 
than one individual, but the preservation and 
Spatial distribution of the teeth might indicate 
otherwise (Fig. 1), The teeth with their in- 
ferred serial position, area of occurrence, and 
dimensions are listed in Table |, 


Taste |, Measurements of P. oscillans teeth, 
fAleischke Fossil Cave, 

Excavation ant post 
Yooth Specimen No, urea/level length width width 
m™! P22736 AT/9 #9 74 74 
re p21734 ATI9 10,1 89 8.6 
iMsor4d = P22K15 Abs 11 12.1 Wh 9.2 
iMiort P29826 ALIS /l4+Y 10,7 93° &7 
Uy P22816 All x/ldet 36 4 Ss. 
iP. £22735 A6/10 42 679 
IM. P17692 X3/3-+4 0 82 RD 
Macey  P22814. AT/IT Wad 69,2 9.2 
IM. P227735 AT/9 11.2 1S R.7 
rM- P22813 Alo/12 Wid 93 «8.4 


Measurements in mm. * approximate. + Specimen 
found during cleanup of slumped sediment from a 
large area centred on All, previously excavated tw 
leyel 17, Levels excavated were 15 cm thick except in 
this instance. 


SOUTH! 


<—~ © Nalural — — 
enhance 


Fle 4, 


Cross-section of depasit, Henschke Fossil Cave, Naracorte, showing distvibution of fossil 
Propleopus teeth, (Table 2.) C-14 in Al indicates 


projected position of dated charcoal sample 


SUA-140 (>35 000 years BLP. 56C)! —997, + 4.0), 


PROPLEOPUS OSCILLANS IN SOUTH AUSTRALIA 43 


The chevk-tecth of Propleopuy have been 
described adequately by Woods (1960), Bar 
tholomai (1972) and Archer er al, (1978), 
and will he discussed here only in general 
terms, One of the notable features about the 
teeth is the very slight difference in characters 
that distinguish upper froro Jawer. and indeed 
lefl from right, molars. They have toushly 
ihe same proportions im both upper and lower 
molurs, and differ only in minule features 
such as a small lingual ridge coming 
forwards from the hypocone and a broad an- 
terior cingulum on upper teeth, both absent on 
lower molars. These teeth bear o superficial 
resemblance ta the deciduous molar MM! of 
some short-faced kangaroos, Sthenurus spp. 
(sensu lato), in which the lophs are not as 
well-developed ay in M* However, they are 
distinguished by different development of the 
midlink (mainly on the proteloph in Sterny- 
rus) and of the erest joining the paracone atid 
metacone (straight and more vertical in P, 
ayoillans). The Mi of Stlenuras is also less 
rectangular than are the molars of P. oxcilans. 


One tooth (P22736) is considered here 10 
represent in M! of Propleapus, This tooth is 
quadritubercular, almost rectangular and 
slightly longer than wide, It resembles the other 
molars in general form. bul is smaller and 
relatively shorter, and cannot be matched with 
any desoribed tooth, To some extent it also 
resembles upper molars (e.g, M2 and M") of 
the Koala, Phascolarctas cinereus, which dif- 
fer in being selenodont and Jess rectangular. 


While the molar teeth of P. osciflans are all 
similar (Fig, 2), it has been possible to ten- 
Lify 1wo of them with sorne certainty os last 
lower molars (Ms) by the reduced size of 
the talonid, the pasteriar half of the Tower 
molar (Woods 1960). This is a feature of 
many marsupials. It was thus possible to check 
Whether this tooth (P22736) was the fast 
upper molar (M*) of P.. oveillany by testing 
the occlusion between it and the lower molars 
There was, in fact, mo possible match, because 
of the great size diserepaney, 1 therefare rule 
our the possibility thal the tooth P22736 ts 
a barely crupted M®°, which tooth ts yeh un- 
known. 

The specimen consists only of the enamel 
erawn of the tooth, and is slightly worn on 
the cusps and crests. Tn the Henschke Fossil 
Cave this sort of preservation is typical of 
deciduous teeth, and those barely erupted 
teeth of juvenile individuals where the roots 


and dentine uapparently have oot been fully 
calcified, allowing them to rot.uway- 

[ conelude that the tooth is an M!, despite 
the facet that ie differs su geeatly from the Mt 
al Hypyiprynmaden (Ride 1961), which is a 
rather irregular three-cusped tapth with a 
poorly developed hypacone, This may imply a 
grealer systematic separation from MMypsiprymn- 
nodon than is currently aceepted (eg, Bar- 
tholamai 1972), The condition of the tooth 
ys much more like that of M! in Berrongia, 
where it is small and somewhat trivngular but 
definitely quadri-tubercular, This agrees with 
ihe observations of Bartholomai (1972) on 
the permanent molats of Prapleapts- 


One of the major problems of “cave 
palacontology” 1s 1he ever present risk of Te- 
warking of the fossils as they are moved 
piecemeal from the enirance to theie final 
resting place. Some of the dangers are de- 
scribed hy Archer (1974). In the present case, 
despile the ecansiderable lateral and vertical 
distribution of the specimens, there iy 00 real 
evidence for more than one individual and, 
if P22736 is an M\ the animal was a juvenile, 
This favours the interpretation that P22816 15 
a Jawer incisor of a juvenile P, osri//ans. and 
explains the difference from the only fieured 
specimen (in OM P3302). Woods (1960) and 
Bartholomat (1972) have remarked wpon the 
unique pattern of wear of Ty This is not 
readily eVideat fn P22816 due te its youth, 
bul a similar wear profile (especially at the 
tip) may be seen and the enamel pattern 
corresponds in its ventrolateral distribution to 
that in P. oscillans (Woods 19601), The enamel 
is broken all shart with the tooth, and stain- 
ing indigates that only half the specimen was 
exposed in the jaw. These features and the 
tapering form ol the tooth (whieh is smaller 
in both diameters than P3302) suggest that it 
undergoes considerable open-rooted yrowth 
during anlogeny. 

Dimensions of known specimens of Prap- 
leapus teeth are given in Table 2. While the 
Naracoorte teeth (Table 2) are slightly longer 
and perrower, where comparable, they clearly 
fit (he proportions and descriptian of P. pseil- 
lans belter than P, chillagoensis, 


The natural history af Propleapius 
The toeth, jaw fragments, and associated 
fossils in the same deposits provide cireum- 
stantial eVidence for interpretation of aspects 
of the Habitat, as well as the ceological role 
of (he animal within the habitat, 


44 N.S. PLEDGE 


Fig. 2. lsolated teeth of P. oscillans, Henschke Fossil Caye, Naracoorte. 


PROPLEOPUS OSCILLANS IN SOUTH AUSTRALIA 45 


TABLE 2. Propleopus spp. tooth measurements (in mm), length x width (anterior/ posterior) 
QM F6675 NMYV P15917 
Tooth P. escillans P. chillagoensis 
PS 15.2 x 10.8 21.1% 13.0 
Me 10.5% 9.7 9.7.x 12.5/10.9 
M#* 11.) x 10.3 97x 11,0/ 9.6 
M* _ 10.2x 9.5/ 7.6 
Me? _— 93x 7.5 
QM F3302. UCMP 51697) NMV P15919 UCMP 45171 SAM P20815 SAM P22425 
P.oscillans  P, oscillans  P. chillagoensis Propleopus sp. P. ascillans P. oscillans 
I, 22.9x 69 _ 21.1% 13.4 _ left right _ 
Pr 13.9x 9.7 *14x 10 _ 13.8x 9.8 14.4% 10.2 14.7x7.7 
M, 95x 8.7 * Ox OS 96x 9.7 WO0x B9/ 91 98x 91/92 410.0 x —/8.8 
M, 10.8x 98 *1Lx 95 10.2 x 10.6 11.1 x 10.3/10,2 11.3x10 /10.3 7 —x91/— 
M,) 11.2x 10.3 *11x10 10.6 x 10,7 12.2% 11,.2/10.4 12.0. x 11.5/10.6 441.5 %9,2/8.3 
M. 110x 9.6 1.1% 9.0 11.2 x 10.1/ 8.5 11,.6x 9.9/ 8.6 ~ —x97 — 


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


+ approximate, damaged or in alyeolo. (D. L. G, Williams pers. comm. 1980.) 


' 


|\\ A 
/ X 


/ 


—E | 
0.5cm 


Fig. 3. 
Hypsiprymnodon moschaius, C: 


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


2cm 


Comparison of macropodoid right humeri in anterior aspect. A: Bettongia penicillata, B: 
ef. Propleopus, D: Simosthenuruy maddocki. 


(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, 


46 N.S. PLEDGE 


we may assume they also tad similar hody 
proportions. M4. moschatuy differs from the 
potorvines in having relatively lone forelimbs 
(Woods 1960), apparently a primitive feature 
retained because of some advantage in its 
dense brush habitat, This difference in form 
from the other rat kingaraoy is well shown 
in Troughion (1973, pl, XQ). T calevlated the 
approximate limb-bone ratios for A. maseha- 
us, using the specimen QM JIM2799, as fol 
lows; 


humerus > radius : femur ; tibia 1: 1,17 
| 1.57 = 1,72. This compares with | + 1.19 : 
2.3 2 2.78 for Berongia penjeitlata Gray, 
1837; 1: 1.33 : 1.9 } 3,07 for the Western 
Grey Kunyaroo Macrapus fuligmasuy ( Des- 
marest); and 1 + 1.24 : 148 : 2.09 for the 
extinet shortfaced kangarou, Simosthenuras 
maddacki Wells & Murray, 1979. (SAM 
P17471-82) all of which are, or were, inhabie 
tants of thick serub, Only MW. /uliginasns is a 
grazer, coming out into grassy clearings to 
feed, 

In the Henschke Fossil Cave material there 
is @ large humerus, ft is straighter, more slen- 
der and more cylindrical than that of the 
kangaroos and potoroines, and it has markedly 
reduced deltoid and pectoral mdges and a 
shorter supinator crest. Of the marsupials 
compared with if (including species of Bet 
tongin, [vpsipryninodon, Macrupus, Sthenu- 
rus, Lhylacinus, Thylacelea, Phascelarctos’) 
the fossil fone most closely resembles the 
humerus of A, mreschatus, hut is even 
straighter and more cylindrical (Pig 3), The 
total length of the fossil humerus is estinyated 
a 19S mm, Assuming that it represents /. 
excilans, and applying the Hypsiprvmnadon 
rutios. the radius length is 228 mm, femur 306 
mm, and vibia 335 mm; measurements indi 
cating an dnimal ag bulky as a erey kangaroo 
bit with sherter hind Jess and much longer 
fore-legs, 


2. Fuod: The iweth are relatively simple in 
form, yuadritubercular and bunodont, These 
features afe alse typical of mammals such us 
Man. pigs ond bears, all of which have an 
omnivorous of browsing vegetarian diet, as in 
fact have living peteroids, Hume (1978) con- 
sidered Hy mosehaius to he omnivorous, ane 
Ramsay (1876) teeorded thal it eats “- 
inseels, worms and tuberous roots -,." and 
palm berries {Archontophernix  CPrycho- 
sperma) aexqndrae). The fetediion of rela- 
tively long forcleps may reflect its method of 


food gathering, namely turning over scrub 
debns, and digging like a bandicoot (Trough- 
tow 1973), The large secant premolars also 
seem connected with an omnivorous diet, 
possibly being used to cut flesh os well as 
vegetable matier, P, oscillans probably lived 
on soft herbaceous Vvegelation, carrion, inver- 
lebrates, and meat 


3. Habitat; Aypsipryviunedon aud some of 
the other poteroids live in dense wet serub 
Where they can obtain protection against pre 
gators. Although so much larger than the 
living rat-kangaraos, Prapleopus may have had 
the sume need. Its presumed stocky build 
would be an advantage in thick brush, just as 
it seems to be lo the siocky Kanwaroo Island 
kangaroo (M, f, fulivinoyus). The assaciated 
fauna adds some support to the idea, Although 
(here are a few lossils species present (such as 
Lasiorhinus 3p. atl Preeopladen sp.) that 
night suggest a more open environment, the 
abundance of brush-dwelling animals, such 
a8 potoroines, small wallabies (mainly 
Maecropus rufogrivens) and bandicoots, indi- 
cates the presenee of moderately shrubby, 
ofien forest in the Naracoorte area during the 
late Pleistoeene. The browsing short-faced 
kanearoos (Simosthenurus spp.) are common 
and the cow-sized diprotodontid Zvyomarirus 
frilnbas also is present: these are believed to 
have inhabited moderately thick scrub. Dipro- 
fodon epatum, which | consider to be an open 
scrub or plains tinimal, is rate! only fragments 
of three teeth have heen recognized. Tor- 
toises (Chelodina ef. longicallis) were abun- 
dant in a nearby swamp. The overall picture 
given by the fossil fauna is of an atea at 
Naracoorte more thickly forested und with 
heavier seruh or thicker understory, aml a 
higher rainfall than today. 

Thus Prapleapus may be seen as a latee, 
bulky, relatively short-legeed animal, living in 


alense thickets and scrub, and cating sueculent 


herbaceous vegetation, inseets and other small 
animals and possibly carrion, Ms rarity in the 
fossil record may reflect its strong preference 
for thick serith, where remains seldom beeome 
fossilized. Tt muy be significant that the fossils 
reporicd here represent 4 juvenile individual, 
one (hit was perhaps less cantious than an 
adult, 


Acknowledgments 
fo thank Dr M. Archer for helpful initial 
discussion al the teeth, the Qiteensland 
Museum for providing reference casts of the 


PROPLEGPUS OSCILLANS IN SOUTH AUSTRALIA 47 


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. 


References 


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 reinterpretation 
of the homology of marsupial cheek teeth. [bid. 
18, 157-164. 

ARCHER, M. & BARTHOLOMAL, A. (1978) Tertiary 
mammals of Australia: a synoptic review, 
Alcheringa 2, 1-19. 

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

lagoensis, a new North Queensland species of 

exlinct giant rat-kangaroo (Macropodidae: Poto- 

roinae), Mem. Nat. Mus, Vict, 39, 55-60. 


BARTHOLOMAL, A, (1972) Some upper cheek teeth 
in Propleopus oscillans (De Vis). Mem. Qd 
Mus, 16, 211-213, pl. 8. 


Guz, BF. D. (1953) Australian Tertiary marsu- 
pials. Aust. J. Sci, 16, 106-108. 


(1957) The stratigraphical occurence and 
palacoecology of some Australian ‘Tertiary 
marsupials, Mem. Nat. Muy. Vict. 21, 135-203. 


Hume, I. D. (1978) Evolution of the Macropo- 
one digestive system, Aust. Mammalagy, 2, 
-42. 


Ramsay, E. P. (1876) Description of a new 
genus and species of Rat Kangaroo, allied to 
the genus Hypsiprymnus, proposed to be called 
Hy psiprymiodon moschatus. Proc. Linn, Soc. 
N.S.W. 1, 33-35. 

Ripe, W. D. L. (1956) The affinities of Burramys 
parvus Broom, a fossil phalangeroid marsupial. 
Proc, Zoo]. Soc. Land. 127, 413-429. 

(1961) The check-teeth of Hypsiprymnodon 

moschatus Ramsay 1876 (Macropodidae:Mar- 

supialia). J. R. Soc. W. Aust. 44, 53-60, 

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

Teprorp, R, H. (1955) Report on the extinct 
mammalian remains at Lake Menindee, New 
South Wales, Rec. 8. Aust. Mus, 11, 299-305, 

(1967) The fossil Macropodidae from Lake 
Menindee, New South Wales, Univ. Calif. Publ. 
Geal, Sci. 64, 1-156. 

Trouciron, E. (1973) “Furred animals of Aus- 
tralian’ (Revised and abridged.) Angus & 
Robertson: Sydney.) 

Witwiams, D. L. G. (1980) Catalogue of Pleis- 
tocene vertebrate fossils and sites in South Aus- 
tralia, Trans. R. Soc. S. Aust. 104, 101-115. 

Woops, J. T. (1960) The genera Propleopus and 
Hypsiprymnodon and their position in the 
Macropodidae. Mem. Gd Mus. 13, 199-221. 


STRUCTURE AND FUNCTION OF THE TIBIAL GLAND OF THE 
AUSTRALIAN FROG LIMNODYNASTES DUMERILI PETERS 


BY G. A. CROOK & M. J. TYLER 


Summary 


Limnodynastes dumerili has a large, oval, dermal gland on the tibia; it is perforated with ducts at a 
density of 6/mm7. In horizontal section the gland appears as a series of large horizontal chambers 
filled with secretions. The ontogeny of glands in tadpoles is described. 


STRUCTURE AND FUNCTION OF THE TIBIAL GLAND OF THE 
AUSTRALIAN FROG LIMNODYNASTES DUMERILT PETERS 


by G. A, Crook & M. J. TyLcer* 


Summary 


G. A. Crook & M, J. Tyner (1981) Structure and function of the tibial gland of the 
Australian frog Limnedyaustes diimerili Peters, Trans. Ro Sac. 8. Ause WO5(2), 49-52, (2 


June, 19R1. 


Limnodynastes damerilt has a large, oval, dermal ghind on the tibine it Is perforated with 
ducts at a density of 6/mm*. In ‘horizontal section the ghind appears as a series of Inrge 
horizontal chumbers filled with seercuions, The ontogeny of ghuids in tadpoles: is descriked. 

When (he gland seeretion is expressed manually to. the surface, rats avoid the Frog. In the 
absence of expression the frog is eaten but the glands are left Lntouched. Predators of the 
frog and closely related species with similar glinds aro listed, 


Infroduction 


In the Anura there are diverse kinds of dis- 
erete dermal glands. Examples are the paro- 
toid glands of many species of Bufo (Blair 
1972); the dorsal lipid glands of South Ameri- 
cain phyllomedusine bytids (Blaylock er al. 
1974); the supralabial glands of Rana (Smith 
1954) and the tibial glands of some species 
of Bufo (Blair 1972) and some Australian 
frogs of the genus Limnodynastes (Martin 
1972; Tyler 1976). 


Much attention has been devoted to parotoid 
wlands and their secretions (Low 1972): the 
structure and possible functions of the tibial 
glands have not been examined, Here we 
describe the gross structure and ontogeny of 
these structures in ihe southern Australian 
species Limnodynasres dumerili, and inyesti- 
gate their possible role, 

Linmodynastes dumerili is a robust species 
measuring up to 75 mm snout to vert length 
when fully grown. Tt is widely distribuled in 
southeastern Australia and is a member of the 
t. dorsaliy group of species, whose represen- 
tarives extend over the eastern and south- 
western portions of the continent (Martin 
1972). Tilustrations of the frog appear in 
Barker & Gris (1977), Tyler (1977, 1978) 
und Cogger (1979). 


Material and methods 
The specimens used in this study were col- 
lected at various localities within an 80 km 
radius af Adelaide and were maintained for 
varying periods in vivaria in the University of 
Adelaide. 


* Depariment of Zoology, University of Adcliide. 
Box $98, G.P.0.. Adelaide, S, Aust. SOOT. 


Glands were obtained fram treshty decere- 
brated and spinalised animals. They were 
dissected free from the bia and for histo- 
logical preparations were fixed in 10% 
bultered formalin, Seetions were cut at 7 »m 
ynd stained with haematoxylin aid eosin, For 
studies Where glands were exposed to potential 
predators, minimal time elapsed between the 
death of the donor and such exposure. Itt six 
trials, groups of four rats were Fasted for 48 
hours aud the entire dead frog offered, the 
entire frog minus the glands, ar gland seere- 
tions wiped upon the surface of rac pellets or 
fresh meat. 

Studies of ontogeny were based on tadpoles 
collected at stages 25-32 of Limbaugh & Volpe 
(1957) and reared on o diet of boiled lettuce 
leaves. Samples were taken fram this stack at 
intervals determined by hindlimb development. 


Gross and histological stricture 


The tibial glands of Limnodynastes dumerili 
are single, raised, ovoid, dermal structures 
on the dorsal surface of cach tibia and oecupy- 
ing move than 50% of the dorsal aspeet of thiet 
limb segment (Fig. 1). Dimensions of the 
sland ina 7S om frog are 15 x T1 mm. Ina 
series of nine specimens the length of the 


glund was 51-69% of the lenglh of the tibia, 


The cland is entirely dermal ond does not 
adhere to underlying muscles. 1 is palpable 
and resilient, The dorsal surface of the gland 
is perforated by the apertures of numerous 
ducts at a frequeney of approximately 6/mm-, 
In some frogs the gland is pigmented with a 
metallic sheen contrasting with the renmumder 
of the dorsal skim, however in the majoriy 
there was no such colour differentiation, 

In transverse section (Fig. 3) the gland tas 
a recular, columnar arrangement of eavities 


50 G. A, CROOK & M. J. TYLER 


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 
material. 


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


int 


mn tS 


' 
4 
: 
| 
/ 
i 
3 


- ' 
=. ea 
, i 
‘ 5S) pow MAS 


me 4 


Fig. 2. Horizontal section of tibial gland. Dense areas are the internal secretions of the gland, These 
secretions have contracted slightly, Scale bar = 100 um. Fig. 3. Transverse section of portion of 
tibial gland. Note columnar arrangement of the internal matrix, and duct (arrowed). The smear of 
secretions in the gland is fractured as a result of histological preparation. Scale bar = 1 mm. Fig. 
4. Transverse section of skin of tibial region of tadpole at stage 39 of Limbaugh & Volpe (1957) 
Note occurrence of cell nuclei in most superficial portion of stratum corneum (arrowed). Scale bar 
— 50 um. Fig. 5. Transverse section of skin of tibial region of tadpole at stage 43 of Limbaugh & 
Volpe (1957). Granular glands (G) and mucus glands (M) are well developed and communicate 
with the surface via open ducts (arrowed). Scale bar = 100 um. 


TIBIAL GLAND OF AUSTRALIAN FROG 51 


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 scutatus, 
Tiger Snake (no effect upon the snake); Anas 
platyrhynchos, Mallard (juveniles up to 40 
mm in length eaten readily) and also to 
Rattus norvegicus, rat. Rats ate every portion 


TasLe 1. Predation upon frogs of the Limmodynastes dorsalis species group 


a 


Species of 
frog Predator Details Source 
L. dorsalis* Vulpes vulpes (Red Fox) In 1% of stomachs | 
investigated, 1958 McIntosh 
In 7.7% of stomachs [ 1963 
investigated, 1959 | 
L. dumerilit V. vulpes (Red Fox) 1 in stomach J. D. Croft 


Notechis scutatus ) 
(Tiger Snake) | 
Pseudechis porphyriacus { 
(Black Snake) | 
Litoria raniformis 
(frog) 


L. dumerilit 


L. dumerili 


L. dumerili Owls 


(pers. comm.) 


E. Worrell 
(pers. comm.) 


Fed to captive 
specimens 


M. J. Tyler 
(unpublished) 


Observed swallowing 
L. dumerili on two 
occasions. Predator 
unaffected 

Bones of numerous 
frogs found in 
regurgitated owl 
pellets 


M.J. Tyler 
(unpublished) 


ee 
Pe _  e 

* Frog species identified as “L. dorsalis” were reclassified as a complex of species by Martin (1972). 

+ Identity uncertain: captured in areas of sympatry between L. dwmerili and L. terraereginae. 


t Identity uncertain: locality data unknown. 


52 G. A. CROOK & M. J. TYLER 


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. dumerili 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 
body. 

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). 


Discussion 


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, Roseghini 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 “g/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. 


Acknowledgments 


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, 


References 


Barker, J. & Gricc, G. (1977) “A Field Guide 
to Australian Frogs”. (Rigby: Adelaide). 

Bratr, W. F. (1972) “Evolution in the Genus 
Bufo”. (Univ. of Texas Press: Austin). 

Biaycock, L. A., RurpAL, R. & PLAtTT-Atora, K. 
(1976) Skin structure and wiping behaviour in 
Phyllomedusine frogs, Copeia 1976, 283-295. 

Coccer, H. G. (1979) “Reptiles and Amphibians 
of Australia”. (2nd Edtn) (Reed: Sydney). 

Limpaucy, B. A. & Vopr, E. P. (1957) Early de- 
velopment of the Gulf Coast toad, Bufo valli- 
eens Wiegmann. Amer. Mus. Novit. (1842), 
1-32. 

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


Martin, A. A. (1972) Studies in Australian Am- 
phibia: Ill The Limnodynastes dorsalis com- 
plex. Aust. J. Zool. 20, 165-211. 

McIntosn, 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. 

Biochem. Physiol. 54C, 31-43. 

Situ, 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). 


BENNELONGIA, A NEW CYPRIDIDID OSTRACOD GENUS FROM 
AUSTRALASIA 


BY P. DE DECKKER & K. G. MCKENZIE 


Summary 


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


BENNELONGIA, A NEW CYPRIDIDID OSTRACOD GENUS 


FROM 


AUSTRALASIA 


by P. De Deckker* & K. G. MCKENziET 


Summary 


Dr Drewker, Po & McKenvie. K. G. (1981) Bennelongia, a new evprididid ostracod genus 
from Avstralasia Z'ray. Re Seo. Sy ebast. UWS(2), 53-38, 12 Tune, 1981. 


A vew Australian astracod genus ind species Beinelongia harpage (Paumily: Cyprididae 


Baird, 1845) is described from Queenstatl, 


Introduction 


The ostracod Chlamyrlorheca auyiraliv Brady, 
IBSG awwas deserthed from empty shells col- 
lected at Penola, South Australia, Sars (1894) 
synonymized ©. australis with Cypris henne- 
lane King, 1855 after examination of New 
Zealand specimens but Miller ({912) queried 
their congeneric stutus. Henry (1923) follow- 
ed Sars’ (1894) concept of the synonymy and 
Chapman (1967) later transferred the species 
to € yprinotes. 

MeKenzie (1971) in his review of the 
palaeozvoogeography of freswater Ostracoda, 
referred to the above species ay the “Chlanty- 
dotheea auseraliy species group’. and pointed 
out avatomical differences hetween this group 
and Cypris and Chlamvdethecu ss, This 
therefore indirectly suggested the need Lo erect 
a new genus to include the Australian and 
New Zealand species, 

Since collection of astracods from Queens- 
land, made available to both authors by Mr 
C. Bentley, yielded a new species belonging 
to the new genus. it was decided to describe 
it jointly, and it is here referred to us Benne- 
longia harpige neen, Tsp. 


Systematic description 
Family CYPRIDIDAE Baird, 1845 
Sublamily CYPRIDINAE Baird, 1845 
Bennelongia necn. 
Type species: Bennelongia harpaga n.sp. 
Bennelongia n.gen. 
Diagnosis: Adult: with strongly asymmetrical 
Valves anteriorly: Jeff valve beak-like an- 
teraventrally: selvage displaced inwards with 


* Department of Zoalozy, University of Adelaide, 
Present Address! Department of Biogeouraphy 
and Geomorphology. Australian National Uni- 
versity, P.O, Bax 4, Canberra, AC JT, 2600, 

t School af Applied Science. Riverina College of 
Advanced Educaton, Worga Waga, N-S.W, 
2650), 


inner bist forming broad but short hp-like 
structure which does not extend anterodorsally. 
Antéroveniral area of right valve usually 
smooth-curved bal occasionally with beak-like 
funge. Juvenile with symmetrical yalyes and 
no beak-like structure: external surface cither 
deeply pitted or reticulated, offen with many 
wartlike tubercles. 

Asymmetrical male maxillary palps; thora- 
copoda f Ist segrnent with two unequal setac 
und penultimate segment weakly divided; 
lateral lobe of hemipenis broadly boot-shaped, 


Derivation of name: From Bennelong, the 
first aboriginal to have a long association with 
the carly European settlers of Australia. 


Discussion: MeKenzic (1971) noted uma- 
tamical differences between the new genus 
and the genera Cvpriv QO. F. Miller, 1776, 
Chlamydothecu Saussure, 1858, Riacypris 
Khe, 1935 and Globacyprix Klie, 1939. A 
unique feature of ihe shell distinguishing 
Bennelongia from the otherwise similar South 
African Cypridinag, is the broad hut short 
lip-like inner list which does not extend 
anterodorsally, 

Bennelongia australis (Brady) and eonge- 
ners are described in De Deckker (1981), 


Benrelongia harpago usp. 
FIGS 1-19 
Deseniptions: Carapace (external) oval to sub- 
rectangular 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: 
anteroventrally, left valve “beak"”-shaped and 
extends much further than right valve which 
is broadly rounded in that area, Deep con- 
cavity just posterior to “beak”-like feature of 
left valve. Whereas mouth region only slightly 
inflexted in both valves. Left valve overlaps 
other in anterior region of hinge. posteriorly, 
and yentrally where overlap is broadest. Shell 


54 


Figs 1-6. Bennelongia harpago n.gen., n.sp. 1. 
P31613. 2. Dorsal view of carapace of paratype P31614. 3. LV internal 
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 = 


| 


_ 


for Figs 5 a-b, 3 


— 


P, 


DE DECKKER, 


100« for Figs 6 a-b. LV, RV 


& K. G. McKENZIE 


¢ 

t 

t 

, 
7; 
1 


Lateral view of carapace showing RV of paratype, 
of holotype, P31612. 


150” for Figs 1-4, 2 = 
left valve, right valve, 


NEW GENUS OF OSTRACOD 55 


i rertil 7 


500 y 


Flus 7. Rennelongia harpago n.gen, asp. Internal 
view of LV paratype, PSL615, 


pseudopunctate with simple rimmed type nor- 
mal pore canals and with number of broad 
pusiules anteriorly and posteriorly, some of 
which are perforated by normal pores. Peri- 
phery of right valve tuberculate along ventral 
margin. 

(iiternal) Inner lamella broader anteriorly 
and broadest in left valve, Inner list develops 
into broad lip anteroventrally in left valve in 
“heak’’ region and absent anterodorsally: in 
front of lip is a deep groove; selvage nearly 
peripheral in posterior region, In right valve, 
inner list marrow and nearly peripheral all 
round except anteroventrally where it is broad- 
er; anteriorly, yroove runs parallel to it, Sel- 
yage tuberculate venically. Anteriorly, radial 
pore canals long, numerous and straight. 
Adductor scars consist of two raws of three 
scars, anterior sears being slightly broader, 
un additional small scar behind and below 
hottom scar ef front row, and anterior to 
and below posterior row, The two mandibular 
sears long and narrow, 

Anatomy: Antennula: (Fig, 8) 7-segmen- 
ted: length-width ration of Jast six segments: 
34 24 1 2 OF 1 
a a T Tv" 1 , EW 
slightly longer than all segments together. 
Penullimate segment with one small hrisile 
and four natatory setae and distal seament 
with three natatory setae. For placement of 
other setae and bristles see Fig. 8. 

Antenna: (Fig. 13) with tour claws: three 
equal ones all slightly pectinate on penulti- 
mate segment plus additional claw on last 
segment reaching tip of other three, Distal 
segment with additional short, slim pectinate 
claw. Nalatory setae reaching fips of claws. 


Natatory  selae 


Mandible: (Fig, 12) epipod with five long 
aud plumose Strahlen and a smaller one on 
side of plate; coxale with seven leeth; endopod 
with « bristle long, natrow and smooth, B 
bristle short, stout and tufted all over, y 
bristle almost twice length of distal segment, 
aod pilose in distal half. 

Labrum: For detail see Fig, 9. 


Rake-like organ: (Fig, 17) five teeth and 
alditional bifid one on inner side of each 
rake, 

Maxillula; (Pig. 15) epipod with 17 
Strahlen and four downwards pointing sctac; 
3rd lobe with two smooth Zahnborsten; length 
y S 


=. distal segment 


ratio of palp segments 


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 one; epipod with five 
plumose Stcahlen, For chaetotaxy of proto- 
podite, see Fig. 11. 

Thoracopoda J: (Fig. 14) protoped coxa 
with one lone dorsodistal bristle, geniculate 
basis with one long bristle; 2nd segment with 
one long inner distal bristle; penullimate 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 
pectinate. 

Thoracopoda Il; (Fig. 18) terminal segment 
with two unequal bristles; short one hook- 
shaped and half length of other. 


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

Furcal attachment: (Fig. 19) median brunch 
slightly curved: ventral and dorsal branches 
forming right angle and hook-shaped al tip. 

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

Colour of shell: grcen with beige strips in 
dorsal area. 


lencht. height 

Size: holotype: LV 1020 600 
RY 9200 S80u 

Holowwne: AM P31612;  Paratypes: AM 


P31613—P31615 (Australian Museu) 


56 P. DE DECKKER, & K. G. McKENZIE 


\\ 


\ \ 
7 ‘i 


Figs 8-12. Bennelongia harpago n.gen., n.sp., holotype P31612. 8, Antennula; 9. Labrum; 10. Geni- 
taltay 11. Maxilla; 12. Mandible; ali x 150. 


NEW GENUS OF OSTRACOD 57 


Figs. 13-19, Bennelongia harpago n.gen., n.sp., holotype. P 31612. 13. Antenna, 14. Thoracopoda I, 
15. Maxillula—palps and lobes, 16. Furca, 17. Rake-like organs, 18. Thoracopoda II, 19, Furcal 
attachment. All x 150. 


58 P. DE DECKKER, & K. G. McKENZIE 


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 
found. 


Acknowledgements 


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. 


References 


BrApby, 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, 
267-286. 

McKenzig, 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-XXXII, 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. Foérh. Vidensk. Selsk. Krist. 1894- 
5, 1-62. 


EXCHANGE PROCESSES FOR UPPER SPENCER GULF, SOUTH 
AUSTRALIA 


BY J. A. T. BYE 


Summary 


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. 


EXCHANGE PROCESSES FOR UPPER SPENCER GULF, 
SOUTH AUSTRALIA 


by J. A. T, Bye* 


Summary 


By J. A. T. (1981) Exehange processes for upper Spencer Gulf, South Australia, 7ravts. 
R, Soc, S, Aust, 105(2), 59-66, 12 June, 1L98T. 

Upper Spencer Gulf, which is defined as Jying north of Lowly Paint in Spencer Gulf, 
South Australia, is characterised by a salinity regime in which there is an approximate equi- 
librium between a northward advection 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 evapara- 
lion, and a southward diffusion of salt, probably by the shear eflect induced by stwong 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 ond annual variation of salinity is obtained for the reported diffusion coefficients if 
the uctual evaporation rates for upper Spencer Gulf are about 0.5 that of the regional pan 
evaporation rites. 

‘The response time seale for a dissolved substance ig a function of the departure from 
equilibrium of its concentration, and for the annual salinity cycle, ihe time scale is about 
300 days. Substances introduced into upper Spencer Gulf at approximately its midpoint would 
have o residence time of about 180 days, and transient injections would initially disperse 
on ut time seule uboul 120 days, but this time scale would increase us the Temnant concen- 
tration of substance declines. 

A higher pan factor with proportionately higher diffusion coefficients would also approxi- 
mately reproduce the observed salinity pallerns, and for a pan factor of 0.67 (as reported for 
fake Eyre) the dispersion limes would be rediiced by about 25%. This figure is the suggested 


overall accuracy for the predictions. 
Introduction 


Upper Spencer Gulf may be defined as 
that part of Spencer Gulf to the north of 
Lowly Point, It contains a water volume of 
approximately 4 km, and has 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 transilion 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 oceanoyruphical analysis 
whieh are much simpler than in the general 
situation to be applied, and results on the 
exchange rates belween upper and = Jower 
Spencer Gulf to be obtained, 


Physical Principles 


The dominant currents in upper Spencer 
Gulf are tidal currents (Radok 1978), which 
normally generate sullicient turbulent mixing 
in the vertical to cause dissolved substances 
to become almost uniformly distributed in the 


» Flinders Institute for Atmospheric and Marine 
Sciences, Flinders* University of South Australia. 
Bedford Park, S. Aust, 5042. 


water column, Thus the concentration patterns 
can be deseribed basically using vertically 
averaged values. The observed salinity patterns 


now suggest that a further approximate 
bene 
oh 
u tw on UPPER 
SPENCER ~~ 
OVLF 


a 


ri ' 


aS TAP HS 

<- L i 

Fig. 1. Upper Spencer Gulf. x indicates location 
of sources. 


60 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 of 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 
Gulf. 

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. 


Theory 


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


3 3 a ac 
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 


= P/p 


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


39Q _ an 
ox -~bGp+E-R) 


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


rainfall rate, and $7 is the rate of change of 


water jevel. This pair of equations, together 
with the boundary conditions that the con- 
centration is specified at the open end of the 
gulf (x,) ie. 


yx, a c,(t) 


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


ac 4 a 
KA 5x = Qe), 0, Q)x, 0 


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


pendent of time, and that 2" js 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. 


aA b on 


ot ot 


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 = pbuc 


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


The method of solution of 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 Pt Augusta to Wallaroo. Solutions of 
the steady-state concentration equation for the 
same region have been discussed previously 
in Bye (1976). 


EXCHANGE PROCESSES FOR UPPER SPENCER GULF 


Determination of the diffusion coefficients 
For a steady-stute distribution with P 0, 
the pair of governing equations may be com- 
bined to yield the relation, 
_ 2th - Re 
ac 


K(x) = 


x 
where =(X) if bdsx js the surface area ol 
x, 


the gulf to the north of x, On substituting for 
the quantities on the right haud side of the 
equation from observed salinity data, and 
estimated net evaporation rales, estimates of 
K(x) can be obtained for all x, Details of the 
method, together with cross-sectional area, 
breadth, and surface area data for upper 
Spencer Gulf are given in Holloway (1974), 
The resulls of his calculations indicate that the 
dilfusion cociicients merease from small values 
(~ 5 m/s) at Pt Augusta to an approai- 
mately constant valle (~ BO m/s) south 
of Lowly Point. fn this study, the deduced 
variation of K has been approximated by the 
function, j 
K > (.0TCby) * b= 3.3km 
sOm=/5 b= 3.3 km 


‘the Annual Salinity Cycle 


The purpose of this section is to estiniate 
the net evaporation rates applying Over upper 
Spencer Gulf such that the observed mean 
salinity distribution and annual salinity eyele 
are predicted theoreticully using the diffusion 
coeflicients. It is well known that pan evapora- 
tion rates obtuined from shore stations are 
grealer (han the evaporation rates from neigh- 
bouring water boclies, For example, in a recent 
study of the water balance of flooded L. Eyre 
(Tetzlall & Bye 1976), it was found that the 
evaporation rate over the lake was approni- 
mately 0.67 of the mean evaporation mites 
for Woomera, Qodnadatta and Moomba and 
that the proportionality factor was almost in- 
dependynt of the time of year, No similar 
comparisons between the evaporation rate for 
Spencer Gulf and that of neighbouring sta- 
tions are known, arid hence theoretical predic- 
tions of the salinity cycle have been made 
using u dincar interpolation between pan 
evaporation rates (each adjusted by identical 
factors) al Pl Augusta and Roseworthy 
(Hounam 1961), and rainfall rates (Anon 
1975) at Pt Augusta multiplied by a factor 
of 1.33 to allow for run-off, and at Wallaroo; 
as end points for the section (Table 1). The 


61 


Taare | Evaporaiion and vanifall rates far Pe 
Auzuste and Wallarey 

PT AUGUSTA WALLAROO 

Lviportit- bvapori- 

tion! Rainfall tion? — Rainfall 

(mm/ Month) 
Januiry 371 18 260 {5 
February WY 18 24 1s 
Mareh 274 16 180 19 
April {73 14 112 32 
Muy 109 23 69 46 
lune 71 26 48 AR 
July 74 2) 43 42 
AuEUSI (04 26 58 A) 
Seplember tay 2) &6 34 
Octaber 23) 39 117 au 
November ZRY 1s (78 | 
Deaveniber 43h 17 224 16 
min 

ANNUAL 2507 236 1589 462 


———————— 
PL Augusta pan evaporation 
~Roseworthy pun evaporation 


net evaporation rates were obtained by sub- 
traction, and reasonable agreement with the 
annual mean salinity profile, wnd the ampli- 
tude of the annuul salinity cyeles along the 
sulf 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 eyele of 
salinity at Pt Augusta and Lowly Point, for a 
salinity at Wallaroo of 38%, are shown in 
figures 2 and 3. They are representative of 
salinity data taken at various times and loca- 
tions and reported in Thomas & Edmonds 
(1956), Holloway (1974) and Anon (1980). 

It is interesting thal the volume transports 
(duc to the evaporation in the north) which 
maintain the salinity distributions are ex- 
tremely small, for example the northward 
annual mean volume transport at Lowly Point 


$0 — - qt st a 7) 
i 
r \ 
th \ 
efthet 
pbs 
40 fy - J 
if | : 
es ees. 2 eee ee |e . = 
» 40 too kW 150 


Fig. 2.. Predicted profile of annual mean salinity. 


qT, A, 


4a 
ot) PORT AUGUSTA 
“” 
4h 
-owLy FOV 
40 


dF M A M wv UV &A FS O WN D 


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


is 8 m/s which corresponds to a mean current 
of 0,05 mms, Furthermore, on dividing the 
volume transport into the volume of upper 
Spencer Gulf, we find that a period of ap- 
proximately 15 years is required to renew the 
water volume completely, This time however 
has littile relevance to the adjustment of the 
concentration profiles, since this adjustment 
is controlled by advection and diffusion, II 
would only be the time scale’ for exchange 
of substance instanlanecously at a uniform 
concentration within the gulf, The actual time 
scales for exchange of substances would be 
quite short while high transient nonequilibrium 
concentration profiles persist, and then 
lengthen steadily as the diffusive and adyec- 
live fluxes tend to their equilibrium values. 

These factors are reflected ji a salinity 
variation which responds to the seasonal 
variations in net evaporation only partially, 
with a response factor, R ~ 0.18, where R 
is the ratio of the amplitude of the observed 
salinity cycle to that of a hypothetieal eycle 
In complete adjustment with the annual net 
evaporation cycle. The corresponding lag (L) 
of the observed salinity cycle relative to the 


annilal net evaporation cycle varies between im 


60 days at Pt Augusta aud 80 days at Lowly 
Point, 

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


|The time scale (7) is defined us the non-equili- 

brium mass of substance in upper Spencer Gulf 
divided by the southward flux of substance past 
Lowly Point. Time seule is used rather than time 
constink since ro may vary apprecinbly with t. 
The mass und flus ure positive quantities for loss. 
und negative quantities for gain of substance. 


- BYE 


(R), and the lag (L) are related to the fre- 
quency (9) of the cycle by the expressions, 


L =} iam? or, and R = 1/VF (rye 


one finds that the time scale for salt exchange, 
r~ 300 days. 

The above results indicate that longer 
period climatie variability in net evaporation 
also may be significant in controlling the ob- 
served salinity variation, For example, a net 
evaporation cycle of period 5 years would 
have the parameters, R ~O.7 and L ~ 200 
days, and consequently would be attenuated 
much Jess than the annual cycle, 


Dispersion from within upper Spencer Gulf 


Theoretical predictions for the dispersion of 
substances introduced within upper Spencer 
Gulf can be readily made for the two tnstances 
of a continuous constant production rate and 
an instantaneous injection of a fixed quantity 
of substance. For definiteness the substance 
will be introduced at a location 34 km from 
the head of Spencer Gulf (Fig. 1), and 24 
km south of Pt Augusta. 


(1) A continues production rate of & kyl s 
for a cohservalive (racer 

The concentration pattern for a conservative 
tracer (d — 5 0) is shown in Figure 4. 
The concentration falls sharply soulh of the 
source, and is approximately coustant north of 
the source with a value of 48 ppm; the Lowly 
Point concentration being 15 ppm, The small 
rise north of 34 km mirrors the sulinity dis- 
tribulion since the fields are both source free 
in this region, The total mass of substance in 
the steady-state equilibrium above Lowly Point 


no 


| ~ 
+ 
a 
bis ae 
‘ — 
oon _ 
rs ay oe | a a | 


| 
° eu ion aon isa 
Fig. 4, Predicted concentration profile tor con- 
servative tracer for continuous source of 
ke/s located 34 km from heud of Spencer 
Gulf, 


EXCHANGE PROCESSES FOR UPPER 


{apb) 


a 
o a n J n { a Oe ee nb 
0 5 joo wey 
Fig, 5, Predicted concentration profiles 40) and 
200 days after instantaneous release of 10 L of 
conservative [racer at location 34 kin from 
head of Spencer Gulf. 


isi 


is 90 kt, and hence its residence time in upper 
Spencer Gulf is 180 days, This residence time 
is Jess than the exchange time scale for salt 
of 300 days, due to the concentration gradients 
being relatively greater than for the distribu- 
tion of an introduced substance than for salt, 
and leading to a more rapid dispersion. In 
fact the residence time is controlled almost 
exclusively by lateral diffusion (cf. Table 3). 


(2) An instaittaneous source of 104 of a@ con- 
servative tracer 

The distribution pattern initially shows a 
lateral dispersion approximately symmetrically 
centred on ihe source, After about 150 days 
however the northward dispersion impinging 
on the bead of the gulf has redistributed the 
substance in au similar manner to the steady- 
stale concentration paitern (Fig. 5), Thus at 
Pt Augusta a maximum in concentration of 
5 ppb occurs after about 60 days. 


1b lo potel —————— _ 

Li 
£0} ¥ 
\ 
| 

all 
+ 

| 
fT a ‘ 3 
ol 1 ‘ SS \ —t | 
i ii yoo aes atu Ao 


Fig, 6 Predicted percentage of instiuntaneous re- 
lease located km from hesd of Spencer 
Gulf, nor) of Lowly Point. as function of time 
after release, (  ) indicates loss time scale 
in days. 


SPENCER GULF 63 
‘-. wi Co —T— ———F —- 
+ 
| 
} i 
ar | | 
53 | 
i. te 
| | 
te i ‘ 
/ _\ 
} sre’ \ 
fy ~ 
a 7 a a YU | 
0 su 150 um (40 


Fig. 7. Predicted concentration profile for tricer 
which exchanges with atmosphere for con- 
linuous source of 6 ke/s located 34 km from 
head of Spencer Gulf, 


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). In 
fact, initially the loss time scale is about 120 
days, but this increases to about 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 
upproximate analysis indicates that the loss 
time scale (7) in fact increases exponentially 
wilh time, i.e. 

r=7re@ thr" 
in which 7, and 7 are constants, that for the 
location of the instantaneous source have the 
approximate values, 7, = 110 days, and 1° = 
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 kg/s 
with surface exchange 

The distribution of a substance with a decay 
lime constant of less thin ~ 100 days would 
be expected to be changed significantly trom 
that for the conservative substance, In particu. 
lar, for exchange with the atmosphere a typical 
exchange velocity, / 10% w/s (Flaney 
1972). thus at 34 km the local decay time 


constant (A) would be about 6 days, This 
ue 


short decay time constant causes a large reduc- 
tion in concentration in comparison with the 
conservative tracer (Fig, 4); the maximum 
value of 6 ppm now oceurring at the source 
(Fig. 7), 


64 JA. T. BYE 


The shove result can also be interpreted in 
terms of the lemperature field arising from a 
heat source with subsequent atmospheric ex- 
chinge. For example, for a discharge (q) of 
10" m/day at an excess temperature (AT) 
of 35°C, the apparent source, 


pdx — pgAT 


where x, and X,° are the bounds of the source, 
and the concentrations in Figure 7 correspond 
approximately to excess Water 1emperatures in 
1 100th of °C, ea, the maximum excess on 
the seale nf averaging of the calculation (2 
km) is 0.06°C, and is aligned with the source. 
In the near field however much higher tem- 
perature anomulics would oecur, and the dif- 
fusive heat flux divergence would he negli- 
gible compared with the surface exchange flux, 
so that for a semi-circular temperature distfl- 
bution appropriate to a source on the coast, 
ane obtains, 
(r) AT ef h/r, rsx bin} 
Where vis the rydial co-ordinate, and 4, >> 
(74/1) js the decay length, The travel time 
toy, l, > "ra where h is the thickness of 
ihe surfuee heated layer, and thus approxi- 
mately the hear field jemperature structure has 
a length sealer, = 1 km, and of assuining 
that h = 0-1 m, a time seale 1, = 3 hrs. 
(4) The eect af long period changes in water 
level 

[tis known that long period changes in sea 
leve} distinct from the astronomical tides ogcur 
wong the south coast of Austrilia (Proyis & 
Radok 1979), These fluctuations in sea level, 
which typically have o magnitude of 0.2 m, 
Propagate into Spencer Gull, and those of long 
period (2 1 week) cause the sea levels in 
Wpper Spencer Gulf to change approximately 
uniformly. 

The effect of the level changes is io oscillate 
the water longitudinally with a range of 2a 
=A, where a is the vertical amplitude of ihe 
long period motion. The ratio =/ A is almost 
constant with distince, varying from about 
2500 ut Pe Augusta to 4000 at Wallaroo, Thus 
for 4 =~ 0.2 m, the range of the water particles 
is Im2 km, and this leads to oyelic changes of 
salinity oF omplitude 0.03%. ut Lowly Point 
and OA. at Pe Augusta, Otherwise no impor- 
tut effects on the property distributions are 
expected, 


Conclusions 

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

(1) The accuracy of the estimates 

The main question concerning the estimates 
is of course their accuracy. The principle of 
the analysis has heen to reproduce reasomubly 
well the mean salinity profile, and the annual 
silinity cycle, and then to predict the disper- 
sion of other substances, No data sets are 
known to exist for the distribution of other 
subslances however which evan provide inde- 
pendent checks on the deduced mixing para 
meters, 

In view of this situation recourse must be 
nade to a theoretical error analysis. “Table 2 
shows the variation in mean annual salinity 
levels al Pt Augusta and Lowly Point that 
would occur for diffusion 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 (E-R)/K, 
(cf. Determination of the diffusion coefli- 
cients), Thus the vigour of the exchange pro- 
cess is ot monitored by the mean salinity 
profile. ‘The vigour however is monitored by 
the observed salinity eyele, such that for a 


Taste 2. Predicted mean annual salinities anc 
te seasonal salinity ranee far various (B=R) 
and & 
(A) MPAN ANNUAL SAUNT te)! 
(E-R) 
FACTORS O75 1.90 1.50 
0,75 45,8 48.7 53.1 
42.0 43.4 46.4 
KR L.00 43.7 454 40.2 
41.0 42.0 44.2 
1,51) 417 44a) 45.8 
40.0 40.6 42.0 
(B) SEASGNAL SALINITY RANGE (4y)* 
FACTOR: LOWLY 
(F-R) K PT AUGUSTA POINT 
150 1.50 484 434 4320 4a 
1.00 1,00 47.8 44.) 427 44 
0,75 O75 A7G 444 426 Ald 


17The two values are respectively the mean annual 
salinity at PL Augusta and Lowly Point, 

“the factor 1,00 corresponds to that for the muin 
caleulahons, the other factors ure multipliers fur 
(E-R) or K f 

SThe (wa values ure respectively the maximun 
and minimum salinities during the annual cycles. 


EXCHANGE PROCESSES FOR UPPER SMENCER GULF 65 


TABLE 3, Predicted residence tine th days for a 
continuous conservative source lavated 34 km 


from ihe head ef Spencer Gulf for various 
(L-R) and K 
(be R) 7 
FACTOR! 0.75 1.40 tt) 
75 237 AN) 245 
KL 176 {78 rao 
1.50 Hn 17 119 


(The factor |.00 corresponds to that for the min 
calculations, the other factory are multipliers 
for (lL. -) or K 


constant CE-ROeK the amplitude of the sali- 
nity cyele is reduced as E-R (or K) is in- 
ercased, This trend nevertheless is not well 
marked (Table 2), and hence the true solution 
for E-R and K cannot be defined precisely 
from the ayailable data. In the event thar the 
proportionality factor for the pan evaporation 
is tOO Small, a reduction in the time scales for 
dispersion would oecur, eg. for a pan factor 
of 0.67 (ef, L. Eyre) the residence time for a 
continuous conservative souree would be re- 
duced from 180 days to 130 days (Table 3). 
Similar reductions would apply for other ume 
scales. and the diffusion coefficients south of 
Lowly Point would have the value 110 m*/s 
instead of RO me/s. Thus it is probable that 
the dispersion estimates have an wecuracy of 
about 25. with the likely bias (from econ- 
siderations of the net evaporation) being to- 
wards time scales somewhat less than those 
found using Holloway’s estimates of the diffu- 
sion coefficients. 
(2) Additional factors 

Wind effects have nol heen considered ex- 
plicitly since it as probable that the main 
uvent of the turbulence is the tidal current; 
however @ proportion of the mixing may be 
wind induced, and thus one would expect some 
scusonul variation in the diffusion coellicients, 
It is not anticipated however that indiviedyal 
storms would have a large effeet on the dis- 
persion processes. Periods of culm associated 
with minima in the tidal current ampliludes 
such as occur ufound "dodge tides’ also may 
have u transient effect on mixing by allowing 
a stratified flow regime to exist, 

All these effects have their place, however 
an average carresponding to the salinity dis- 


iribution is perhaps that which is required 
initially in a study of tracer distributions, and 
data suggest that the salinity distribution is a 
reasonahly well defined regime within. the 
recenk 25 years during which measurements 
are available (Thomas & Edmonds 1965; 
Holloway 1974; Anon 1980), 

The special behvyiour of tracers other than 
well vertically mixed conservative and atmos- 
pherically exchanged quantities also hag not 
been considered, fur example, the behaviour 
of surface and bottom seeking substances 
would be significantly different and require 
individual treatment. A comprehensive ireat- 
ment of course must extend to an evaluation 
of the results of exchange of substance for 
marine life and for the almosphere, 

(3) Proposed programmes of observation 

A fuller understanding of the dispersion pro 
cess would of course rely on extended pro- 
frammes of observation, In particular, obser- 
vations of the actual water balance of upper 
Spencer Gulf, including evaporation, rainfall 
and run-oll are required as time series extend- 
ing over several years, These dita could then 
be used fogether wilh observed profiles of 
salinity, and other introduced tracers to specify 
in detail the dispersion provess. In particular, 
attention should be given to the conditions 
just south of Lowly Pomt 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- 
leresting balunee between the net evaporation 
and Jateral diffusion, and are » good example 
of what has been called » ‘reverse estuary’. 
Several similar inlets exist around the Austra- 
lian coastline (notably in Western Australia) 
for which this kind of opalysis would also 
appear fo be applicable 


Acknowledgements 

The ideas presented in this pauper have been 
stimulated by the necessity for obtaining some 
understinding of the dispersion regime in 
upper Spencer Gulf in the event of che pro- 
posed industrial development at Redetif’, 
which is situated 34 km from the bead of 
Spencer Gulf. approximately at the source 
lovation used in the dispersion studies, 


66 J. A. T. BYE 


References 


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


— (1980) Redcliff Project Draft Environmental 
Effects Statement Dow Chemical Co., 178 pp. 


BowbeN, K. F. (1965) Horizontal mixing in the 
sea due to a shearing current. J. Fl. Mech. 21, 
83-95. 


BuLtock, D. A. (1975) The general circulation 
of Spencer Gulf, South Australia in the period 
February to May. Trans R. Soc. S. Aust. 99, 
43-54. 


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


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


Hoiitoway, 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. 
Comm. Aust. Bur. Met. Bull. 44. 

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

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

Rapox, 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. 

Tuomas, I. M. & Epmonps, S. J. (1956) Chlorini- 
ties of coastal waters in South Australia. /bid. 
79, 152-166. 


LATE PRECAMBRIAN PSEUDOFOSSILS FROM THE FLINDERS 
RANGES, SOUTH AUSTRALIA 


BY R. J. F. JENKINS, P. S. PLUMMER & K. C. MORIARTY 


Summary 


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. 


LATE PRECAMBRIAN PSEUDOFOSSILS FROM THE FLINDERS RANGES, 


SOUTH AUSTRALIA 


by R. J. F. JeNkins*, P, S. PLUMMER* & K. C, MoriartyT 


Summary 


Jenkins, Ro J, &, Plummer, P. S. & Moriarty, K, C, (1981) Lale Precumbrian pseudofossils 
from the Flinders Ranges, South Australia. Trans. R. Soc, S. Aust, 105(2), 67-83, 12 June, 
OBL. 

Enigmatic, subeylindrical to conical, sediment-infilled structures occurring in the Trezona 
Tormalion and the lower and upper parts of the Moorillab Formation of the late Precambrian 
ol the central Flinders Ranves, South Australia, resemble certain fossil burrows such 
as Bergaueria Prautl and aso late Precambrian body fossil remains including the sack-shaped, 
solt-bodied form Ernietta Plug from South West Africa (Namibia), The structures occur 
up jo 3,300 m stratigraphically below the well known Ediacara assemblage, ‘The lack of 
disfuption or vents in their matrix seemingly negate the possibility that (hey are waler escape 
structures, but studies of their infilling and literature research indicate that their true origin 
iy ulmost certainly inorganic, their resemblance of Various fossil remains being fortuitous. 
‘Yheir recognition emphasizes the apparent non-occurretice of metazoan remains below the 
Edincara assemblage in the local rock record, lending negative support to recent theories 
postulating either late evolution of the Metazoa or rapid diversification of animal life in the 


latest Precambrian. 


Introduction 

In 1975, a discovery of abundant, peculiar 
eylindrical structures was made by Moriarty 
in the lower part of the Mourillah 
Formation (Plummer {/¥78) i Utanouna 
Creek, within the Bunbinyunna Range, south- 
west of Wilpena Pound in the Blinders Ranges 
(Fig. |), At the time he considered that the 
structures may have been biogenic. During late 
1976, in the course of extensive field mapping 
of the Brachina Subgroup, Plummer found a 
comparable structure in float Within Bunyeroo 
Creek, and later a second specimen only a few 
kilometres from the original find of Moriarty, 
This latter specimen was shown lo Jenkins who 
noticed a radial pattern of markings on the 
base of the (short) column, and longitudinal 
striations on its sides, As such, the form 
showed a resemblance to the Cambrian to 
Jurassic trace fossil Bergaueria Pratl, and also 
to various late Precambrian metazoan remains. 
Subsequent exploration has widened the 
known distribution of the structure and con- 
firmed its constaicy of general form and the 
more or fess stable sedimentological charac- 
teristics attending its preservation. 

Twa specimens of similar structures in 
stratigraphic collections of the University of 


* Centre for Precambrian Research, The Univer- 
sity. of Adelaide, North Terrace, Adelaide, 
§.Aust. S001. 

+ School of Earth Sciences, Flinders University. 
S.Aust, 5042, 


hyestot 


MOUNT chaMBens x 
CREEK / 


! BLINMAN 
PNR ACHILME A ? 
aos “ 
Wire alfa WS 


' 


| 
l 

| 
\ 
| 


‘ 

HRACHINA t---- at ae “A sSttparinng 4s 
HIDING ; t \ 

‘ wax ante CREEK 


‘ 
1 


KEY 
Pseudotess| invcalitres 


* insitw 


% Talus 


wit PENA 
vm HS Homestead 


BLACK oo ia 


fox ones 6 “Sanley 


oo Unsealed 
a hrbaba + 


Fig. Locality map of pseudofossil occurrences 
in artis Flinders Ranges, South Australia, 


Adelaide were found by M. A. Reynolds in 
1950 west of “Bunyeroo Hut" (probably Yan- 
yanna Hut on curren! maps) apparently within 
the Trezona Formation of the Umberatana 
Group, 


Stratigraphy 
The Trezona Formation reaches a thickness 
ef aboul 240 m in the Oraparinna area and 
consists of mud flake rich pelletal limestones, 


68 R. FJ, JENKINS, P. S. PLUMMER & K.-C. MORIARTY 


CAMBRIAN 


RAWNSLEY QZ. 


EDIACARA 
ASSEMBLAGE 


BONNEY 55. 


BUNYER OO FM 


ABC RANGE OTZ. 


POUND SUBGROUP 


» Interval with = 
Pseudotossils © 
~~, \ 


WS 


BAYLEY RANGE FM 


WILPENA GROUP 


MOORILLAH 
FORMATION 


MOGLOOLOO FM_ 


BRACHINA SUBGROUP 


TREZONA FM; 


Pseudotossils 


ENGRAMA SH. 


UMBERATANA GP 
SUBGROUP 


FARINA 


Fig. 2. Charl summurizing regional stratigraphy 
of youngest Precambrian succesyiou in central 
Flinders Ranges, and showing stratigraphic 
intervals in Which pseudofossils are found in 
relation to Edjacara assemblage. Only upper 
part of Umberatana Group shown. Pound Sub- 
eroup aller Jenkins (1975b), 

oryptalgal-laminites und associated stromato- 

lites, oolitic limestones and intraforniational 
limestone conglomerates mterbedded in green 
siltstones (Thomson ef al. 1976). It as of 
shallow water origin and comprises the upper- 
most unit of the Farina Subgroup, within the 

Umberatana Group, and is of early Marinoan 

age in terms of the local late Precambrian 

lime-rock stratigraphy (Thomson 1969), [tis 
succeeded by the approximately 60 m thick 

Elatina Pormation which includes the tillitic 

deposits of the third and youngest glacial phase 

of the Adelaidean (Mawson 1949), and is 
part of the Willochra Subgroup of the 

Umberatana Group. 


The Brachiog Subgroup (Plummer 1978) 
of the Wilpena Group is a coarsening upward, 
dominantly clastic sequence wilh an average 
thickness of 1500 ine It rests mostly conform- 
ubly though locully disconformably on the 
Elatina Formation, Lying within the Subgroup 
approximately 600 m above this glacrogenic 
unit is a 460 m thick sequence consisting 
predominantly of thinly interbedded red ane 
purple shales and siltstones. This sequence 1% 
defined ay the Moorillah Formation and con- 
nionly interposed within it are medium to thick 
beds of purple tuffaccous siltstone which offen 
display intralormational conglomeratic tex- 


tures, and Jenticwlar crossbedded sandstones. 


Soft-sediment deformation structures are ubj- 
quitous. 

The structures which form the subject of 
this paper oecur in two broad intervals within 
the Moorillah Formation. In Bunbinyunna 
Range (Fig, |) several hundreds of specimens 
were observed in a broad zone spanning much 
of the basal half of the Formation. Very few 
specimens were observed near the top of the 
Formation in this area. Specimens were, how- 
ever, moderately numerous near the top of the 
Formation at Mount Chambers Creek, 100 km 
to the northeast, 

Repositoriey 

Representative specimens are lodged in the 
collections of the University of Adelaide; 
catalogue numbers prefixed “A.U,". Studies 
Were also made of a large collection of late 
Precambrian fossils obtamed by Dr Hans 
PHug, Giessen, West Germany, from the Nama 
Croup of Namibia (South West Africa), 
These specimens, catalogued "Pf." are ulti- 
mately to be returned by Dr Pilue to jhe 
appropriate statutory body in Windhoek 


Description 

The shape of the structures in vertical 
section is commonly cylindrical (Pig. 3A & 
E), with either a rounded to hemispherical 
hase (Fig, 4A), or a flat, U-shaped base 
(Pig, 3F; Fig. 4G), which js often lopsided 
and with one or several central indeniations 
(Fig, 4C, D, FE & FP), A sub-conical shape is 
also occasionally displayed (Fiz. 3B & C), 
The upper part frequently expands outwards 
like the mouth of a trumpet. In sceton pari 
Ucl to the bedding the structures display 9 
cifeular to ovoid outline (Fig. 3C & D). The 
width of the structures (disregarding the 
lrumpeilike upper rim) averages 6-5 em. but 


PATE PRECAMBRIAN PSEUDOFOSSILS ay 


varies between 0,8 and 20 em. The expanded 
mouth of the trumpet may reach over 30 cm, 
Their width is frequently less than their depth, 
With Scme speeimeas roughly twice as. deep 
as wile, The depth may reach about 20 em. 
Qlten their surface is nearly smooth, but on 
many a series of fine longitudinal ridges 
and wrooves extends along the cylinder or cone 
ind fans out on the trumpet-like upper rim, 
Occasionally these ridges and grooves eon- 
tinue across the base of the structure as radial 
markings (Big. 4). The significahee of these 
markings is questionable. however, as they are 
often indistinguishable fram, and grade into 
glickensides which also occur commonly 
within the matrix, 

In the Mooriliah Bormation the structures 
are (ound within interbedded shales, siitstones 
and fine sandstones. In the majority of speci 
mens the hose of the structure is either 
sunken down inth, or in close juxtaposition 
with the top of a siltstone or fine sandstone 
bed (Fig. 4©). The upper parts of the struc- 
tures are often enveloped by laminated shales 
which sometimes inchide thin beds of siltstone. 
Where the siltstone below the structures js 
thin it is sarnetimes. bowed downwards and 
there ig a tendency for the shale laminae and 
ihin siltstones subjacent to the sides of the 
siruetures also to be gently cyrved downwards 
(Pig. 3A & Fy Fig. 4C & DB), The uppermost 
part of (he strictures may terminate withils 
shale, or be tcunculed ar capped by another 
hed of siltstone which is often bowed up- 
wards. The capping shyle or siltstone par- 
ticipated in partially tilling the structates. Rare 
specimens occur more or less entirely within 
shale (Fig. 3F). 


Sections of the siructures show that they 
had a complex history of infilling and sub- 
sequent compaction (Piz. 4C, DY B & Gi. 
Ollien the initial infiling consists of fine to 
coarse sand With angular or subrounded mud 
und silt clasts Which came to rest in a subs 
horiyontal attitude, ar sometimes in an edge- 
wise fashion (Pig. 4G). The sand infilling may 
show rather jrregular erosional surfaces within, 
or grade upwards into laminated siltstone still 
conmmining oecasinnal clasts. Tt is apparent that 
during de-watering of the surrounding matrix 
appreciable compaction occurred. The sandy 
material and silly infill of the forms seems 
generully ta have behaved in a more conipe- 
teal fnanner ond compacted less than the 
envelaping matrix, resuluing in the observed 


downward deformation of the external sedi- 
ments below and upward deformation above 
(Fig. 36), The silt laminae of much of the 
infill are frequently bowed upwards (Fig. 4C, 
Dp & 8), The specimens from ihe Trezona 
Formation have a slightly calcareous silty infill, 
but otherwise show similar characteristics to 
the material from the Moorillah Formation, 


Interpretation 
General 

The nature of the infill indicates that the 
structures must have initially been hollows av 
hales within the sedimentation surface. These 
holes thus acted as. traps for any course sedi- 
men! beimg carried in traction before being 
buried by the more turbid finer sediment. A 
syngenetic orgin is therefore envisaged for 
which either an organic origin, or production 
hy inorganic processes, 15 possible, 

Possible Organic Origin 

Two possible orgunie origins could produce 
syngenetic structures similar to those deseribed 
here, Firstly, the burial of dwelling or hiding 
burrows of actinian-like creatures could pra- 
duce trace fossils resembling such forms as 
Berganeria, or Conastichus Lesquereux, Alter- 
natively. the preservation of ecup- or sack- 
shaped animals could yield body fossils com- 
parable to Nawulia Germs. the “Ernietto- 
morpha™ of Plug (1972a) or Baikalina Sako- 
lov. 

Frey (1970) describes the burrows of 
several living aclinians as being considerably 
dovper than wide, or sometimes extremely 
elongate and occasionally branched. Shinn 
(1968) shows that adjacent to such burrows 
it ig Common to see a downwarpiig of the 
sediment amination, In a study which 
Mangum (1970) made on the burrowing be- 
haviour of the anemone Phylluciiy Milne 
Edwards & Haine, she stated that rhythmic 
contractions of the column musculature mani- 
pulate the fluids of the column's hydrostatic 
skeleton to push the pedal dise into the sub- 
strate; “after the pedal dise reaches a hard 
substratum and sand surrounds the colomn 
(burrowing) ceases.” The Lower Cambrian to 
Jurassic trace fossil Bergaueria is normally 
attrihured to aetinians burrowing snd resting 
in this manner (Crimes er al. 1977). Accord- 
ing to Alpert (1973) the depth of Berguieria 
is less thant ar equal to its diameter, although 
Crimes ef al. have observed docper specimens 
in the Lower Cambrian of Spain. Externally, 


70 R. F. J. JENKINS, P. S. PLUMMER & K. C. MORIARTY 


LATE PRECAMBRIAN PSEUDOFOSSILS 71 


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 (Needler 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 11 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 
erosional 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 McAlester (1962) 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 ithe Nama 
Group, Namibia, and Baikalina sessilis Soko- 
lov 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 (Glaessner 
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 (sce 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 
conspecific. 

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) 


nna 


Fig. 3. Field photographs of uncollected pseudof 
Formation, except D and F from upper part; sca 
beds in adjacent matrix curved downwards at sides and thinned near base: 


ossils; examples from lower part of Moorillah 
le divisions in cm. A, subcylindrical form_ with 


Utanouna Creek. 


B, conical specimen: Utanouna Creek. C opening where conical specimen has eroded from matrix; 


upper edge of form apparently extended to line 
atrowed: Black Gap. D, external mould of Jarge bowl-shaped form: 


x-y and striations on mould of its surface are 
Mount Chambers Creek. E, 


subcylindrical specimen showing gently domed ripple bedded sands above and downward deforma- 


tion of beds near base: Utanouna Creek. F, profile of 


Chambers Creek. 


shallow structure within shale: Mount 


R. F. J. JENKINS, P. S. PLUMMER & K. C. MORIARTY 


ae 
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LATE PRECAMBRIAN PSEUDOFOSSILS 73 


and vompased wf several layers of soft hasue 
which could undergo ready deformation. The 
sack is inferred to have been attached at its 
closed end, either to the substrate, or very 
frequently to small clay gulls or same other 
frojection on the surface of the substrate 
(Fig. SD & E; Fig. 6A). The several wall 
membranes of the sack Were interconnected by 
thin, Hexible, radially arranged, longitudinal 
septa whose position on any partieular wall 
layer is marked by either harrow ridges or fine 
longitudinal prooves (Fig, 6B, C & D), The 
clongate tubular spaces se farmed between the 
walls and the septa wre frequently packed with 
silt or sand to form internal moulds, ‘The 
upper and apparently open end of Lrnieha 
terminates in aw crenellate margin ip the few 
rare inwlancey where it is preserved, More fte- 
quently, however, it is found collapsed and 
deformed (lav, SA & BE). This discussian at 
other details concerning the Mructure et this 
extraordinary organism full outside che scope 
of the present wark 

One of the most remarkable characteristivs 
of Ernietia is its preservation, whielr js one uf 
infilliny: compurable to the structures from the 
Flinders. Ranges, Thus, sections of Ernieria 
may show a layered and eraded sand or sill 
infill, with medium-grained sand at the closed 
end of the suck and very fine-grained sand 
near the apparent open end (Fig. SC), Bed- 
ding surfaces indicdte the original depositional 
orientation, Blehs of clay-rich materia] and 
limonite pseudumorphs, presurmahly after 
authigenic pyrite, are frequent in the infillings, 
The few specimens still within their original 
matrix are enveloped by mediuni-gramed san 
(now quartzite), The inflling of Ernicsta dees 
uot, however, show the fine lamination usually 
observed in Gur structures, 

Pile (1972b 1974) aricniated the "Erniet- 
tomorpha” in fhe reverse aspect ta that sug- 


gested here, that is with the closed end of the 
sack upwards, Published data concerning their 
oceutvence in the field (eg. Germs 1968, 
1972a) are ambiguous as to their orientation. 
Nevertheless, eVidence from their distortion 
and layered intill is considered substantialive 
of our interpretation, 

From a stratigraphic Viewpoint it is intrigu- 
ing that the Kuthis Subgroup, which contains 
Ernivila, is succeeded by a sequence contuining 
glacingenic sediments and scraped and grooved 
bedding planes (parts of ihe Schwarzrand 
Subgroup) reflecting several cold phases of 
Juter laie Peeeumbrian glaciation in southern 
Africa (Germs 1472, 1974; Kroner 1977; 
Kroner e/ af 1980) and that our structures 
oceur only celatively short stratigraphic dis- 
tances both below and above the last late Pre- 
camhrian elaciogenic deposits in the Flinders 
Ranges (purts of the Elatina Formation). 
However, the glacial events recorded in the 
Schwarzrand Subyroup are suggested to have 
been of local extent, and radiometric data of 
relevance for the Nama Group (Kroner 1977; 
prelimimary data passed in personal communi- 
cation between Professor Krdner and De Pflug; 
Kroner ef a/. 1980) together with fragmentary 
dating information from Australia reviewed 
by Williams (1975) suggests the likelihood 
that the Nama glaciations are significantly 
younger than the Elatina Formation. 

The Siberian form Balkalina sessilis 16 
apparently closely similar ta Ernietta, but is 
too incampletely deseribed for further cam- 
parisan, 

One objection to a biological origin is that 
if our forms did represent soft-hodied fossil 
organisms then the creatures must be imter- 
preted as having maintained their life func- 
tiong even during the time of their intilling, 
as their presumably soft integument would 
surcly have collapsed flat if death were earlier, 


CECE UU UE UUEEISEtSSE EE 


Pip, 4, Photographs of jnfilling sediment and sections of pseudofossils and field photograph of depres- 


sions cannected by eullere from lower part of Mnorillah 


Formation at Utanowna Creek, except F, 


from same stratigraphic interval at Black Gap and G from loose boulder in Bunyeroo Creek. 


Borescules equivalent to T cm. A, side view of 


plug of mfilimg sediment of pseudofossil eroded free, 
A.U, No, 24731. f, field photograph of irregular 


depressions interconnected by gutters; end of 


ruler (o hinge 50 cm, C-F, rellected light photographs of sections of pseudofoasils; C, erosive base 
(b) of form closely juxtaposed with hedding interface (s-y) beiween siltstone bed (#1) und over- 
lyinw shale (sh), A.U. No. 24729, D, cast infill with erosive surface, arrawed, ind cone-shaped Trac- 
tures (f-f) extending below: shale laminue in matrix down-turried at sides of atruciurc, A.U. No. 
24730, E. specimen with cenirally domed base and domed, finely laminated silt infilling, A-U, No, 
24728 F. undermeath view of part of base of infilling plag of tive pseudafossil with arrows in- 
divating more or tess sadivily arranged murkings, AU, No, 24733. G. broken section through. 
pscudofossil within fine sundstone: inlilling. also fine sandstone, includes numerous shale flakes, some 
ae we ovlentated In edge-wise fashion: overlying bed of lunminaled fine sandstone, AL, No, 


MORIARTY 


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JENKINS, P. 


LATE PRECAMERLAN PSEUDOFOSSILS 75 


Maintenance of life during sedimentation 
scems manifestly impossible. This argument 
makes the explanution of the preservation ol 
Ernicite diteult, but for this form it might 
be presumed that the onset of sedimentation 
wis extremely rapid, perhaps in the form of 
a lurbidity current Which gave rise to the ob- 
served graded infill, In any event. the Ernie/fu 
organisms appear ta haye ultimately suc 
cumbed 10 sedimentation us their tops are 
variably pursed and compressed or overfolded 
(Fig. SA & E), Our structures show na evi- 
dence of (he multiple wall tissues present in 
Evnictia, nor definite evidence of septa or the 
precise and regular organization so chatac- 
teristic of this fossil form, The most conyine- 
ing explanation of the striations on the sides 
of our strueturcs are us slickensides dade ta 
differential compaction. In a nuniber of speei- 
mens the slickensided surfaces project inte the 
matrix below the actual infill in the form of 
a more or less cone-shaped fracture (Fig, 404. 

The systematic posidon of Nanaia, the 
“Erniettomorpha’ and Bafkulina, together with 
several other problematic Precambrian fossil 
organisms, was reviewed hy Glaessoer & 
Walter (1975), who suggested them ta be re- 
laled fo their taxon Aramberia hanks, Vrom 
the Arumbera Sandstone in the Amadeus 
Basin, central Australia. Glaessner & Walter 
considered that While these forms could gene- 
rally be referred io Plog’ (19708, 1970b, 
1971, 19720, 1972b) division of the *Peta- 
lonamac™, the formal application of this name 
was not to be recommended beeatse af 1s 
hypotheticul concept and various speculative 
implications applied to it, Thus they rejected 
the ideas of Pflug (1973, 1974) that the 'Peta- 
lanamae” represent a peculiar branch al Pre- 
eambrian evolution intermediate in posidan 
betwee! the Kingdums of plants and animals. 
Nevertheless. Glaessner & Walter maintained 
that the forms jus! discussed and various Pre- 
cambrian frondtike organisms show a general 


similarity which implies an lnderlying taxo- 
nomie unity, and thal the members of this 
grouping “may be classified provisionally as 
Coclenterata of uncertain systematic position’. 

This viewpoint js criticised by Jenkins and 
Gehling (1/978) who suggested that the origi- 
nal concept of the “Petalonamae” may involve 
the artificial amalgamation of quite distinctive 
classes or even of separate phyla, Varying 
further interpretations of the Petalonamae are 
given by Ford (1979), Brasier (1979), Scrut- 
ton (1979) and Glaessner (1979), but this 
problem falls essentially outside the seope of 
the present study, Based on comparisoyy with 
the geological studies and experimental work 
al Bzulynski & Walton (1965), ~Artanberia” 
is Considered by Brasier (1979) io be “prob- 
ably a pseudefassil cuused by turbid water 
flows". New widespreal discoveries of "Arun 
ferla’ made by Dr B. Daily and Jenkins m 
central Australia (See ulso Kipsehvink 1978), 
where the form is several times found in fear 
agsocigtion with Course grained sandstones or 
conglomerates. provide a body of additional 
evidenee supportive of Beasier’s canclusion- 


Probable lnofvaniec Origin 

In un excellent review of inorganic cylindri- 
cal strictures found within sediments, Bailey & 
Newman (1978) indicate that the usual pos- 
tulated mechanisms for their formation Involve 
early diagenetic upward-lowing springs oF 
escaping pore waters (see also Quirke 1950; 
Hawley & Hart 1934; Gubelman 1955, Cony- 
heare & Crook 1968), Such structures ate {re- 
quently filled with sediment of similar grain 
size, sorting and texture iq the enclosing (silt- 
stone or sandstone) matrix, or show a crude 
coursening in grain size towards the centre of 
the cylinder. Ocvasionally, even an inverted 
cone-in-cone arfuneement of sil) and sand 
Jarnination can weeur (see Dionne & Laver- 
dire 1972: Ganvloff 1974). Often the lower 
end of the cylinder is ipregalar and shows 


| Loe 


Fie, 5. Remuins of Breielia pliteauensis Pug. 1966, from Kuibis Subgroup of lower part of Nurmi 
Group, Namibia: all natural size, A, side of neniy complere specimen Viewed along line at wider 


trinsverse Axis Uppermost purts show gvidlence 


of overfolding, Pf No. 178 B, site of specimen 


viewed normal to wider Transverse axis; at least three separate Wall membranes (WW, W2 andW *) 
shown by intecnal moulds of sand, enigmatic “budding” suture occurs wlong b-b, Ph No, 182. C, 
longitudinal section of sand infilling central cavity showing layers of progressively finer sediment 
and depositional surfaces (2) presumably reflecting iniermiltent stages of infilline and oneniion 


of organism at cach stage; durk square and spots Himonite after aulhigenic pyrite, 


light cotuured 


edee of the infilling duc to weathering PE No. 308. D. view of base of specimen showing deformed 


area (eb) an nippletike prominence 


organisms vipzae sulyire Alongs line x-y, Pf, No. 
ehclasing matrix: infilling indvidial collapsed downwards, itl) 


pursing inwards. Pf. No. 287. 


(pn) which may represent the point of allachment ob the 


99, F, side view of specimen wilh some of 
hase fluttienimg and apper purl 


76 R, F. J, JENKINS, P, S PLUMMER & K. C. MORIARTY 


re Ts 


ee Sa 


tt) 
HM p 


Fig. 6, Ernieuin plateauensis Pflug, 1966, A, reconstruction of group of individuals living on sandy 
Precambrian sea bottom. Several possible alternative, but nol necessarily mutually exclusive, re- 
constructions of individuals shown. Youthful specimens evidently attached to mud galls of othee 
prominences on sea bottom, more muture specimens muy have lived with basal parte buried in 
substrate, Ourermost membrane seems to have been caul-like and only faintly showing underlying 
longitudinal structure. “Mouth” sometimes preserved broadly flared, or more frequently collapsed 
or pursed, Individyals of all sizes may shown enigmatic transverse suture or possible zone of “bud- 
ding’ at about half their length, About .7 natural size. B-D, sketches of sectional cuts through 
aetial specimens: b, traces of bedding in matrix (impule sandstone or quartzite): © adhering thin 
sliver of 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 
x 1.5. B, sublongiludinal section almost normal to zig-zag axis, three wall membranes evidenced. 
PF, No, 120. C, section of another specimen almost transverse to longitudinal surface markings. D. 
section of different specimen cut oblique to longitudinal surface markings, 


LATE PRECAMBRIAN PSEUDOFOSSILS M7 


fissure fillings. In the present structures from 
the Flinders Ranges, bowever, ihe frequently 
fine Jamination of the more silty phases of ihe 
infilling shows no evidence of disruption other 
than slight deformation during compaction, 
and the thin sand beds which often oeeur at 
the base of the cylinder show no signa of 
fissuring or disruption sueh as might be ex- 
pected with the rapid escape of fuids. Thus 
our structures were evidently mot formed by 
the upwurd eseape or flowuge of pore fluids. 
Also, the syngenesis of these structures elimi- 
Hales au epigenetic coreretionsry origin. Such 
syngenctic structures could, therelore, only he 
formed inorgunically as toroids or pal-casts; 
these mames both essentially referring 2 the 
custs of c¢ireular scour pis. 

In past literature, such scour pity are offen 
suguested to have beer formed by whirlpool 
action generated along the boundary belween 
currents lowing in opposite directions. As such 
they might be expected to ovcur in high enerey 
shallow-water environments ig both marine 
and non-marine realms, Aceording to Cony- 
beare & Crank (1968), circular varoids are 
usually wider than deep. The external nor 
phology of their casts 1s suggested to be 
characteristically swirled, like thar of a 
“folded bin”, geiterally with a smooth surface, 
although the formation of roughly concentric 
patterns by scouring is common, Titernally. 
foreid casts may display structural and tex- 
tural homogeneity, 

Our siructures show a marked resemblance 
to the “rippled torouis” deseribed by Dorr & 
Kaullman (1963) from the Mississippian 
Napoleon Saridstane ef southern Michigan, 
They considered these torards to have heen 
“formed by the action of Vorlex currents on 
unconsolidated sand in shallow water of the 
inner sublittoral zone’! and supported their 
findines by inducing vartiees in a water-filled. 
sand-floored box und producing artificial 
laroids. Some of the structures they deseribed 
were elongate, oblique cones, a shape oeca- 
sionally shawn by our material, 

This precise explanarion seems inapplicable 
wok respect to our Structures for several 
reasons Dore & Kaullmann considered that 
the environment of deposition of the Napo- 
leon Sandstone was one of telalively high 
energy and this is censistent wilh the texture 
of the rock (sand), Our strietures oecur 
within @ Variety of lithotypes representing mud 
to fine sand erades, reflecting microenyiron- 


ments of varying energy levels, upd presum- 
ably mostly of cuther low energy. The Vortex 
hypothesis jiivoked by (hese authors implies 
tidal shear ar near-shore phenamend which, 
by their very nature, must be restricted to a 
relutively narfow belt parallel! to the palaeo- 
coast, We, however, have Tocuted our struc: 
lures al geographic separuhons of 20 km 
parallel to the inferred depustitional strike and 
perhaps 60 Km at right angles to it, “Also, our 
forms. cecur sit various levels over a fatal strati- 
eraphie thickness in exeess of 1400 m. 

Aigner & Putrerer (1978) and Aiger (19784 
document and explain inorganic pol-easts and 
sutfer-casty fran the Muschelkath af south. 
west Germany Some of jhe pot-eists which 
they fiaure Ge, Aigner & Fatterer 1978, figs, 
3A. At, D, D! and E) are identical in shape 
and internal layering to the Flinders Ranges 
epecimens. eXeept that the coarse nvaterial ot 
the buse af the “pois consists of shell debris 
rather than ¢lay galls. The Muschelkalk pols 
may be packed within an cdgewise cou- 
glomerate of shells similar to the edgewise 
conulomerate of mud flakes present ia some 
of our specimens (Piz. 4G), 

On the basis of ume experiments Aigner & 
Futterer allribute the Muschelkalk pot-holes 
jn the erosive effects of eddy currents forming 
in shell-filled depressrons due to the influence 
of a steady or increasing unidirectional eur- 
rent over the gencrul surface of the substrate. 
They nole that such pots are unknown on 
present-day tidal Mats and consider that their 
origin is entirely submarine. The implications 
of this ore at some variance with Plummer’s 
(1978) finding that on fhe basis of such struc- 
lures as flat-topped = ripples, interference 
ripples and shallow channels with ebb-orien- 
fated asymmetric ripples, parts of the Mooril- 
lah Formation in which our structures secur 
Frequently aceumulated in-a low intertidal mud 
fiat environment. However, we did not observe 
any sédimentological structures in dipect asso- 
ciation With the pot-casts that would tmely 
emergence, 

An -alicaetive aspect of tis explanation for 
the Flinders Ranges structures is that some- 
times the pat-casts arc peculiarly coalesced or 
joined by uneven trough-like forms (Fig. 4B) 
which are exactly paralleiled jm the Muschel- 
kalk material (Aigner & FPrilterer 1978. figs. 
3B. C and F). In rare instanees there may 
even be evidence of a ripple hollow or ather 
depression in which the clay galls that 


78 R. F. J. JENKINS, P. S. PLUMMER & K. C. MORIARTY 


Number of Readings = 72 
(Level II) 


Level IV 


. 4 
N 
Level Ill + 


Minimum Diameter (mm) 


5 N 
Maximum Diameter (mm) 


Level Il 


Number of 


Readings = 230 : N 
(Levels I-IV) Level aw 


Direction of Alignment 


C 


Frequency 


LATE PRECAMBRIAN PSEUDOFOSSILS Ts 


formed the erosive tools may have originally 
uccumulated (Fig. 3A). 

Measurements of the diameter of the pot- 
custs in ihe Flinders Ranges (Fig. TA & B) 
show a runge of varibility quite comparable 
ta guller-casls studied by Aigner & Futterer 
and the depth of specimens is similar. A histo- 
gram of diameter against frequency (Pig. 7A) 
shows u somewhat skewed distribution with 
the modal diameter at 6-7 om, A suggestion 
of a polymodal conmpunent in itis histogcam 
probably reflects the observation ihal speci- 
mens of the pal-custs on given bedding planes 
tend to be closely similar in size, a likely mn- 
dication of theie Nearly simullancaus genera- 
tion. Karez et wh (1974) deseribe how a field 
of small pis (individually reaching up to 
9mm diameter) may form on freshly deposited 
vluy undet the influesee of a wurrent. and tt 
may be possible that such depressions serve 
46 a nucleus for the subsequent formation of 
pot-holes. 

The study of Avner & Putterer also de- 
onstrates. a Very marked preferred orienla- 
tian fur the Muschelkalk gutter-casts, parallel 
1o the cogstlioe and basin axis, Qur pot- 
structures are usuylly ovoid in outline (Fig. 
7B) and measurements on four bedding-planes 
gn oascending stratigraphic order at Utanounsa 
Creek also d@monsirate 4 marked preferred 
orientation tor any viven bedding plane (Fig, 
70), though this direction changes hy BO" 
between ihe lowest and the third surfaces 
studied. The more or less east-west oricnta- 
tions for Levels T pnd Ul are approxinvately 
normal to the palavodeposmjonal trends for 
the Flinders Ranges area, whilst the N.N,W. 
SSE aliveerions for Levels TH and (V ure 
roughly parallel to che hasin axis and (Wes- 
tern) shoreline (eg. see Plummer 1978), 

The individual pol-custs are spaced on 
bedding plines at intervals of several fens of 
ceniimeters to seven meters. In rare fastances 
specimens are grouped on parr af vu becding 
plane, while pone are present on the remainder 
of the surface, Dorr & Kauffman (1963) 
pointed out dhur ibe “rippled torpids’ which 
they deseribed also show a patchy distribution, 
Similarly Norrman (1964) indivated 


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


Associated pseudofossils and oldest local 

record of metazoans 

Glaessner (1969) described ihe vomplex 
\racetike marking #unyericlinus dalgarnvi 
from the base of the presently recognised 
Moorillah Formation at Bunyeroo Gorge, He 
considered that this marking may have been 
made by an aninial related to a primitive mol- 
Juse Without a shell. Large bedding plane 
exposures uic a persistent feature of miuny 
Outerops of the Moorillah Formation at this 
locality, bul despite extensive searches no new 
finds of B. dalyernen have been made during 
the decade since as discovery, After erlical 
examination of 8. dalvurnei, Jenkins (1975a) 
Sug#ested that i 7s a composite of primary 
impressions occurring on two separate beddme 
planes, und ihus unlikely to be of metazoun 
origin, “The great frequency with which in- 
organic tool markings ceeur in this forovation 
led Jenkins te consider it jo be “a unique 
aevidertal set af markings made by a tethered 
implement moved by the current’, The imple- 
ment may haye been a small toud clast caught 
iu a tassel of twisted algal threads. It is sug- 
vested to have moved in a series af snyall 
jumps in several arcuate swaths fo leave over- 
lapping, curved bands of more or less regular, 
Iransverse imprints marking siccessive bed- 
ding planes. Thalus-bearing, mbbon-like algae, 
which might have provided the tether, appear 
in the Riphean of the US.S.R, apd form ac- 
cumulations grading into saprapelic laminae 
and films in the Vendian (Sakolov. 1977). 

Small circular markings on a bedding plane 
of a single loase bloek of reek found within 
the Moorillah Formation have been considered 
as structures formed by escaping gas 
(Plummer 1980). 

The oldest probable recard of metazoans in 
the Flinders ranges is of markings resembling 
small medusoids and fitte sinuous trails found 
by Dr Mary Wade at Brachina Gorge wilhin 
the Bonney Sandstene (previously the “Red 
Pound Quartzite’) which is the lower forma- 
tion of the Pound Subgroup (Wade 1970), p. 


nn a aaa ass stnSstt ttn 


Fig. 7. Measutenents of pseudofossils at Utenouna Creek. Bunbinyunna Range. A, histogram of 
raaxiniimn transverse diameter of specimens plutled against frequency; specimens occur on_ four 
large bedding planes, levels L-4V im ascending striligraphic order, B. plat of maxinvum and minimum 
Irunsyerse diameters of individual specimens occurring on level Tl, C, direction of orentation of 
Jonger transverse axis of avate specimenes plotted for each bedding plane. 


Bu) RK. FF, JENKINS, BP, 5, PLUMMER & K, C MORIARTY 


92). Other finds of Precambrian Metacoans in 
Australia are apparently either upproxiumately 
equivalent in age bs the Ediscura assemblage 
wasn eric, or are likely al younger ape us 
is Uvferred for che Mt Skinner ~lyuny"” of the 
Amadeus and Georgina Basins (see Wade 
1969: Daily 1972: Kirschvink 1978). It thus 
heeomes timely to consider why the severas 
tholgand metres of Marimoan or youngest Pre- 
cambrian sediments which precede the Pound 
Subgroup mm the Flinders Ranges, aad whieh 
ure well exposed and otherwise oceur exten- 
sively throughout the Adelatde Fold-Belt, are 
appareddy barren af metazaan fossils, Nume- 
rous = stratigraphical and — sedimentolagical 
studies generally imply marine origins for 
mich of this succession (Thomsen L969: 
Thomson ez al. 1976; Plummer 1978). Argue 
ments that the lacies may not have been suit- 
able for the preservation of small soft-bodied 
temams (cf Glaessner 1972) are nat fully 
convincing as finely laminated beds preserving 
the smallest morganic tool-murkinygs (ea. 
0.5 mnt) are frequent, and Wade (1968, 1970) 
has found that the conditions needed Sor the 
preservation of a soft-hodied assemblage were 
Hor particularly stringent, Indeed ihe Ediacarsa 
assemblage sens Jare is now known vo aceur 
in quite variuble facies ar diferent World 
lovalities, 

An extensive literature coneerns supposed 
finds of either body fossils or irace fossils pre- 
ating the Edigesra assemblage serv Jato, but 
subsequent studies have led frequently to a 
questioniig of the aterpretation or sometimes 
|he age of these reported oveurrences (Glaess- 
ner 1969, |979 and pers. comm.) Cloud 1973, 
1978; Cloud et al, 1980; Sokolov 1977; Yakoh- 
son & Kryloy 1977). Other of these finds 
continue to be cited ws possible evidence sup- 
portive of ideas of a gradualistic evolution of 
early metazoans (ee. Glaessner 1472, 1979; 
Beligston 1977; Brasier 1979). The studied re- 
cord of the sequences in the Flinders Ranges 
dacs not provide evidence of can extended 
early histery for the meltazoans. 


Ruther negative evidence and the known 
necutrence of fossils in the Flinders Ranges 
lend suppert to weas such 4s those of Stanley 
(1973, 197ba, 1976b) stigeesting a late radia- 
tio of Melazoans, predicted on a basis of 
ecological modelling, or to an exponential in- 
erease in-animal lite fram ihe late Precambrian 
to the Cambrian, show by analysis of age 
data and counts of recognized taxa {Sepkaski 


1978, 1979), Stanley considered that the early 
rudianon of cukaryetes may have been in- 
hibited by the saturation effect of Preeanbrian 
algul systems und sugwested shut the advent ol 
cell-eating heteratrophy criggered a “kind of 
sell-propagaling Feedback system of diversifi- 
culion” cumulating in the initial major radia- 
tions of both the metuphytes and meltazoans, 

Recently Chouberte & Puure-Muret (1980) 
assigned a middle Riphean age to ihe rocks 
containing Precambrian metazoan tossils on 
the Avalon Peninsula, Newfoundland, and at 
Charawood forest, England, This finding ts 
patently ineorreet, and not only overlooks 
lines of geologigal evidence which suggest 
that these cocks are late Precambrian (Wil- 
liams & King 1979; Patchett et al 1980), but 
ignares a 20 year eyele of research document 
ing Vendian or Edisearan affinitics for the 
genera Charnia Ford and Charntiodiscus Ford 
present in the fossil assemblages (Glaessner 
1977, 1979). 


Conclusions 


The present stiily describes Precambrian 
structures which have a relatively complex 
genesis and show resemblances to certain 
cylindrical to conical fossil burrows, as well as 
to suek-shaped, longitudinally striate body 
fussil remains of comparable age; However. 
these resemblances appear to be fortuitous and 
the Irue avigin of the structures is suggested 
to he a$ erosive pits formed by vortices induced 
hy episedic currenis and the scouring action 
of trapped mud galls. The hydrodynamic con- 
ditions leading to their formation are not fully 
understood and to our knowledge no such 
structures have yet heen deseribed from 
present dav, marine, sub-vidal environments, 
The deeree of resemblance between these 
pscudofossils afd several burrow torms and 
suck-shaped bady fossils emphasizes that a 
yariely of hoth ineregnic and organic pro- 
cesses may lead to the formation of closely 
similar stractures which may even show de- 
grees of gradation. The corollary of this 
finding is that the description of such kinds 
Of Precambrian structures and the assignment 
of a particular genesis, especially ane of hie- 
Jogival origin, should be approached with 
cuution, 

The finding of these structures and reeoe- 
nition of other associated pseudofossily suc- 
gests that true indications of Metazoa are 
restricted itn the highest part of the local 


LATE PRECAMBRIAN FSEUDOFOSSILS a 


Precambrian sequence, thus providing twega- 
live evidence supporting recent theories that 
postulate a late evolution or radiation of 
aninyal life towards the clase of the Precam- 
brian. 


Acknowledgements 

Dr Hans Phug of the Justus Liebig Uni- 
vesity, Giessen, West Germany, is warmly 
thanked for allowing eXamination and study 
of his large eollection of Preeambrian Tossils 
from southern Africa, and for his hospitality 
and numerous valuable discussions on his 
inaterial. The University of Adelaide 1s ac- 
knowledged for making a Study beave Grant 


available towards this purpose, 

Or B. Daily, University of Adelaide, and 
Dr W. ¥. Preiss, S.A. Department of Mines 
and Energy, ure gratefully acknowledged for 
their constructive criticism of che manuseript, 
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 Borch, Flinders 
University, is thanked for discussion. Dr T, D. 
Pord, University of Leicester, England, read 
the manuseript at the “Symposium on Life in 
the Preeambrian, Leicester, April 10-12. 
1980". und passed on the remarks of the 
assembled meeting. 


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SONDEROPHYCUS AND THE TYPE SPECIMEN OF PEYSSONNELIA 
AUSTRALIS (CRYPTONEMIALES, RHODOPHYTA) 


BY H. B. S. WOMERSLEY & D. SINKORA 


Summary 


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. 


SONDEROPHYCUS AND THE TYPE SPECIMEN OF PEYSSONNELIA 


AUSTRALIS (CRYPTONEMIALES, 


RHODOPHYTA) 


by H. B. S. WomerSLEY* & D. SINKORAy 


Summary 


WomersLey, H. B. S. & SinKorA, D, (1981) Sonderophycus and the type specimen of 
Peyssonnelia australis Sonder (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 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. Agardh but a synonym of P. capensis Montagne. 


Introduction 


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 Indien” seems likely to apply to 
a different taxon. The description of Denizot 
was probably based on material such as that 
illustrated by Harvey (1859, pl. 81), and this 
is indeed a distinctive southern Australian 
species, 

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 
isotype material. Sonder later also included 
some Sonderophycus specimens under his 
P. australis. 

P. australis Areschoug (1854, p. 352) from 
“sinu Port Adelaide” (specimens in §) is the 
same as Sonder’s type; Areschoug’s name was 
apparently indepedent of Sonder’s . 


— 


* Department of Botany, University of Adelaide, 


Box 498 G.P.O., Adelaide, S. Aust. 5001. 
+ 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 dates 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. 


Sonderopelta coriacea gen. and sp. nov. 


Peyssonnelia australis sensu Harvey 1859: 
pl. 81. 
NON P. australis Sonder 1853: 685. 
Ethelia australis? (Sonder) W.v. Bosse 
1921: 300. 
Sonderophycus australis (Sonder) Denizot 
1968: 260, 307. 


Sonderopelta 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 


86 H. B. 8S. WOMERSLEY & D. SINKORA 


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 eccentrice extensa, lobata vel 
lacera; Jamina in sectione e filamentis radiatis 
cellularum amplitudinis similaris composita, 
filamenta utrinque extendentia, ad paginam in- 
feriorem cellulis frequentibus terminalibus 
filamentorum haptera efferentibus, ad paginam 
superiorem filamentibus erectescentibus et 
cellularibus brevis cortex formantibus. Repro- 
ductio incognita. 

Type species. §. coriacea sp. nov. 


S. coriacea sp. nov. 


Thallus (Fig. 1)  eccentrically  peltate, 
spreading from a short (to 2 cm long and 


lz cm broad), fibrous and often divided stipe, 
usually growing under overhangs in low light 
intensity; lamina cartilaginous, 4-1 (-14) mm 
thick, radiating eccentrically from the stipe, 
often deeply divided or laciniate with lobes to 
20(-25) cm long and to 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 3-2 cm from the rock substrate. 

Thallus eccentrice peltatus, ab stipe (ad 2 cm 
longo et 13 cm diametro) breve saepe diviso 
patens, plerumque sub petra imminenti in luce 
demissa vivens; lamina cartilaginea, 4-1 (-14) 
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 Jucem superior 
sanguinea, pagina inferior cinerea fibrosa (per 
rhizoidea septata) in plantis magnioribus 
2-2 cm a substrato. 


VONDEROPHYCUS AND PEYSSONNELIA AUSTRALIS aT 


Type locality: Pondalowle Bay, Yorke Penin- 
sulu, S. Aust. (2-3 m deep in shade, 141.1981; 
SM. Clarke). 

Type: ADU, A52035 (Fig, 1), [solypes to be 
distributed as No. 214 in “Marine Algae of 
southern Australia’, 

Sonderepelta is named to commemorate 

Otto Wilhelm Sonder (1812-1881), combined 
with the peltate form of the well-developed 
thallus. 
Distribution: From the Isles of St Francis, 
§. Aust. to Waratah Bay, Vic. and around Tas- 
mania, mainly on rough-water coasts in depths 
of 1-25 m, usually tn heavy shade. 

Sonderopelia differs from Peyssonvelia in 
thallus structure, having longitudinal filaments 
which diverge to both upper and lower sur- 
Faces, Whereas Peyssonnelia bas a distinet basal 
hypothalhial Jayer producing filaments above 
und attachment rhizoids below. Lthelia, in 
Which Sonderopelta was placed with some 
doubt by Weber yan Bosse, differs in having 
upwardly and downwardly directed filaments 
produced from a central, apparently limited, 
layer of distinctly Jarger filaments, it also 
differs in being fully adherent to the subsirate 
but without producing attachment rhizoids. 


Peyssonelia anstralis Sonder 
Peyssonnelia australis is a common subtidal 
alga on southern Australian coasts. Py gine 
niana |, Agardh (1876, p. 387), based on a 
collection of Harvey (3271) trom George- 


town, Tasmania (type in Herb, Agardh, LD, 
27698) which had been earlier referred to the 
European 7. rubra Harvey, becomes a syno- 
nym of P, australix, and P, coccinea J.. Agardh 
(1876, p, 385) from Western Australia (prob- 
ubly near Bunbury) (type in Herb. Agardh, 
LD, 27650) is probably also synonymous. 

Denizot (1968, p. 123) placed P. gunniana 
vs a synonym of P, capensis Montagne (1847, 
p. 177) from South Africa (type in PC?) but 
without detailed comments on their identity. 
Denizot regarded the presence of internal cal- 
ecified granules as well as hypobasal calcifica- 
tion as characteristic of P, capensis (as well 
as other features such as the septate rhizoids), 
and Ausiralian specimens appear to be speci- 
fically identical with ones studied from South 
Africa (e.g. Isaac 307 from Terguiet, near 
Mossel Bay, 13.x.1954; ADU, A40825). 

Hence P. australis Sonder should be re- 
garded ay a synonym (along with P. guaniana 
J. Agurdh) of P. capensis Montagne. 


Acknowledgements 

We are grateful to Dr Paul Silva and Dr 
Jim Ross for comments on nomenclatural 
aspects, and to Mr S. A. Shepherd and Mr 
S. M, Clarke for comments on the growth 
habit and habitat of Sonderopelia. De BE. M. 
Gordon-Mills kindly supplied the latin diag- 
noses. 

Appreciation is expressed by the first author 
to ARGC. tor provision of technical assis- 
tatice. 


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PARTIAL ACQUISITION OF PIGMENTATION IN AN ADULT, ALBINO, 
AUSTRALIAN LEPTODACTYLID FROG (LIMNODYNASTES DUMERILI 
PETERS) 


BY MICHAEL J. TYLER AND MARGARET DAVIES 


Summary 


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. 


BRIEF COMMUNICATION 


PARTIAL ACQUISITION OF PIGMENTATION IN AN ADULT, ALBINO, 
AUSTRALIAN LEPTODACTYLID FROG (LIMNODYNASTES DUMERILI 
PETERS) 


Although there are sporadic reports of the dis- 
covery of albino frogs, the in@ividuals inyolved 
generally are stable in their lack of pigment. 
Exceptions are the observations’ 4 of European 
Rana species in which the tadpoles derived fram 
albino ova acquired norroal pigmentanian gradually 
over i period of approximately twa weeks, We 
have not located the description of pigmentauon 
developing in albine frogs at later stages jm [heir 
ontogeny 

On 22.8.79 we received a gravid female Limno- 
dynastes drmerili collected ot Evanston near 
Gawler, South Australia. The hady was dull pink 
with darker areas on the flanks where the internal 
orguns contd be seen in partivl transparcney 
through the body wall, The eyes were pigmented 
normally and for this reason, conform to the 
description of “partial albino” frogs in the termi- 
nology of Pavesi", Smialtcombe? and Dubois". 

The female was placed with u group) of 
normally pigmented L. dumerili from North Ade- 
lade, On 22.9.79 she mated with one of the 
normal mules and fiid approximately 3.100 pig- 
mented eges of which 1,170 were infertile, The 
tndpolts. Were pigmented normally, and the 
feaultant metumurphosing frogs were perfectly 
normal in external appewrance and behaviour. A 
very high death rate in the offspring following 
metamorphosis was attributed to overcrowding ul 
a fime when competing requirements for other 
research animals limited the space available vor 
them. Ten individuals remained in Oclober 1980, 
but they were relatively small (sfout to vent 
length 33 mm), and they died over the following 
three months. 

In November 1980 the female began to acquire 
a few small (3 om diam.), circular spots of black 
pigment, Initinfly these were confined to the skin 
covering the tibial glands, and the pigment wrews 
expanded there until cach vland was pmented 
entirely. Over a period of several months a dark 
mosaic of pigment then deyclaped upon the 


1Smallcombe, W. A. (1938). Nature 14) (3573), 
753. 

2 (4949), J, Genet, 49, 286-90. 

2Rostumd, J. (L946). Anoures Rey. Se). 84, 564-5. 
iSindecek, F. (1964). Folia Biol (Prague) 10, 
23-9, 

‘Pavesi, P. (1879). Rend, ist Lomb. 2, 528-34, 
“Dubols, A. (1979), Mitt, Zool, Mus Berlin 55, 
59-87. 


Partial albino Limmnodynastes alqeiertit 


Big. 1. 
Pelers. 


dorsum and limbs (Fig, [), so producing a piebald 
uppearance, Unfortunately the femate died before 
pigmentation had extended further, At death her 
snout to vent length was 66 mm which is within 
the normal range for the specics. 


We have been unable to trace in the literature 
other examples of the delayed acquisition of pig- 
mentation comparable to that reported here, Cer- 
tainly it is a normal event for many frog species 
to chunge colour and markings in the course of 
their carly onlogeny, but the changes reported here 
occurred at a post-reproductive slage, 

In our search tor records of comparable pheno- 
mena in other verlebrales, il has become upparent 
that the lerm “partial albinism" os employed for 
frous: by Pavesi®, Smalleombe? and Dubois has & 
different interpreiation to the “partial albinism” 
of mammals exhibiling the Chedigh-Higashi Syn- 
drome (CHS) reported by other authors 1", CHS 
is Characterized by u paucity of pizment of the 
eyes and skin (or hairs), as opposed to lack of 
pigment in the skin and presence in the eyes. 

We thunk Cliris Miller for assistanee with the 
rearing of the tadpoles, 


‘Padgett, G. A. Leader, R. W,, Gorham, §. R. & 
O'Mary, C, €, (1964). Genetics 49, 305-12, 

*Taylor, R, FL & Farrell, R. K, (1973), Ped. Proce 
32, 822 (Abst). 

Kramer, 7. W,, Davis, W, D. & Prieor, D, J, 
(1977), ab. Trivest. 36, 954-62 

\0Prieur, DD. J, & Collier, L. L, (1978), Am. I- 
Pathol. 90, 533-6, 


Micitane J. TYLER and MAnGAner Davins, Depurment of Zoojogy, University of Adelaide, Box 498 


G.P.O,, Adclaide, 8 Aust, 5001, 


VOL. 105, PARTS 3 & 4 
11 DECEMBER, 1981 


Transactions of the 


Royal Society of South 
Australia 


Incorporated 


Contents 


De Deckker, P. Taxonomy and ecological notes of some ostracods from Aus- 
tralian inland waters - - - - - - - - 
Beveridge, I. Three new species of Calostaurus (Cestoda: Davaineidae) from 
the New Guinea wallaby Dorcopsis veterum - - - - 
Tyler, M. J., Davies, M. & Martin, A, A. Frog fauna of the Northern Territory: 
new distributional records and the description of a new species 
Baker, A. N. & Devaney, D. M. New records of Ophiuroidea (Echinodermata) 
from southern Australia, including new species of Ophiacantha 
and Ophionereis - - - - - - - - ~ 
Jenkins, R. J. F.. The concept of an ‘Ediacaran Period’ and its stratigraphic sig- 
nificance in Australia - =. - - - - = 
Foster, C. B. & Harris, W. K. Azolla capricornica sp. nov. First Tertiary record 
_of Azolia Lamarck (Salviniaceae) in Australia - - - 
Bull, C. M. & King, D. R.A parapatric boundary between two species of reptile 
ticks in the Albany area, Western Australia - - - - 


Brief communications: 
Von der Borch, C. C. Recent non-marine dolomite from the coastal plain, south- 
eastern South Australia - - - - - . - 
Kailola, P. J. & Jones, G. K. First record of Promicrops lanceolatus (Bloch) 
(Pisces: Serranidae) in South Australian waters - - - 
Lange, R. T. & Reynolds, T. Halo-effects in native vegetation - - - - 
Smith, J. & Schwaner, T. D. Notes on reproduction by captive Amphibolurus 
nullarbor (Sauria: Agamidae) - - - - 3 - 
Smith, M. J. & Rogers, P. A. W. Skulls of Bettongia lesueur (Mammalia: Macro- 
podidae) from a cave in the Flinders Ranges, South Australia - 
Zeidler, W. A giant deep-sea squid, Taningia sp., from South Australian 
waters - 5 = s = = - - - = 
Addendum - - - = - - - - - - - - - 


PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS 
STATE LIBRARY BUILDING, NORTH TERRACE, ADELAIDE, S.A. 5000 


TAXONOMY AND ECOLOGICAL NOTES OF SOME OSTRACODS FROM 
AUSTRALIAN INLAND WATERS 


BY P. DE DECKKER 


Summary 


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- 
described. 


TAXONOMY AND ECOLOGICAL NOTES OF 
SOME OSTRACODS FROM AUSTRALIAN INLAND WATERS 


by P. De DreckKer* 


Summary 


De Decker, P. (1981) Taxonomy and ecological notes of some ostracods from Austi'alian 
inkind waters, runs. BR, Soc. &. Aust, 198(3), 91-138, 11 December, 198), 

Two new osttacod genern Alboa and Gomplielella are described and 11 few species: 

A. woreda, Bennelongia barangardo, B, pinpi, B, nimala, Australocypris: dispar, Strandesia 

phoenix, Reticypris clava, R. kurdimurka, lyacypris perigundi, Leptoevihere lacustris, and 

Gomphodella maia. Ecological notes are presented for these and an additional eight species, 


some of which are re-descrihed, 


Introduction 


Ostracods ar¢ common inhabitants of most 
types of waterbodies in Australia with the 
halobiont fauna being particularly diverse here 
compared to other parts of the world (De 
Deckker I481). However, taxonomic know- 
ledge of Australian ostracods is incomplete, 
as indicated by the numerous ostracods re- 
ferred to in ppen nomenclature in publications 
dealing with limnological 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 for the descriptions here 
are deposited at the South Australian Museum, 


Systematic descriptions 
Susc.ass; OSTRACODA Latreille, 1806 
OrvER; PODOCOPIDA Miiller, 1894 
SupERFAMILY: CYPRIDACEA Baird, 1845 
Family: CYPRIDIDAE Baird, 1845 


SuaFamity: CYPRINOTINAE 
Bronstein, 1947 


Alboa n.ven 
Type species: Albou wereaa tsp.. gender mas- 
culine, 


Diagnosis: Asymmetrical valves; in anterior 
view, greatest width of right valve at 0,5 from 
dorsum and of left value at 0.7 from dorsum, 
Selvage prominent in left valve and placed at 
0.5 of width of inner lamella anteriorly. Peri- 
pheral groove on the outside of selvage in 
left valve, 


* Department of Zoology, University of Adelaide. 
Present addfess| Department of Biogeography 
& Geomorphology, Australian National Univer- 
sity, P.O, Box 4, Canberra, A.C.T, 2600, 


Male maxilla’ palps strongly asymmetrical; 
lateral lobe and distal part of copulatory sheath 
boot-shaped. Geniculate joint between Ist and 
2nd thorocopoda | segments with two unequal 
sctac, 

Derivation of tame: Alboq meaning egg in 
aboriginal language to refer to the egg-shaped 
shell,’ 


Alboa woroda n.sp, 
FIGS 1=2 
1914 Cypris sydneia King: Chapman, p. 27 
Diagnosis: As For genus. 


Deseripiion: Carapace (External), Pseudo- 
punctate, oval-shaped with ventral area almost 
flat except if mouth area which is slightly 
coneave at about 0.4 from anterior. In left 
valve, other (aint concavity, anterior to mouth 
region al about 0.2 from anterior, Greatest 
height and width al about middle. Left valve 
larger and overlapping right one anteriorly, 
posteriorly and ventrally. In anterior view, 
valves strongly asymmetrical; left valve, 
broader und larger; greatest width of left valve 
at 0.7 from dorsum and of right value at 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 
and Ventraily but slightly broader in left valve; 
posteriorly, inner lamella of right valve very 
narrow. In left valve, selvage prominent and at 
about 0.75 of inner lamella’s width from outer 
margin. Selvage follows curvature of shell 
except anterlorly Where it is less curved. Pos- 
teriorly, selvage less prominent and at about 
0.8 of inner lamella's width, Peripheral narrow 
depression on outside of selvage. In right valve, 


| Aboriginal words used in this text ate from 
Papps (1965) aml Cooper (1962). 


92 P. DE DECKKER 


Fig. 1, Alboa worooa n.gen., n.sp, a antennula, b mandible—palp, c mandible—coxale, d maxillula, 
e antenna, f thoracopoda H, g maxilla, female, h maxilla, male, i maxilla—endopodite, male, j 
thoracopoda I, k hemipenis, | furca, m furcal attachment. a-e, h-m: holotype adult male. g: para- 
type: adult female. Scale: 100 z. 


O8TRACODS. FROM AUSTRALIAN INLAND WATERS v3 


selvaze peripheral and faint except anteriorly 
where it is much sharper and extends further 
than edge of shell, Right valve with taint 
tubercles posteroventtally along edge of shell. 
Radial pore canals numerous, narrow and 
straight, Central muscle field consisting of 
three sears in front and two behind; upper and 
lowest sears in front broad and elongated 
whereas other two almost circular, Mandibular 
scars below and jin front of central muscle 
field. 


Anatamy. Antennula; (Fig, la) 7-segmented; 
leneth-width rutio of last six segments: 2/3, 
B/1, 1/14, 3/2, 1,75/1, 2.5/1, Most plumose 
nalalory setae as long as all segments together, 


Antenna: (Fig. le) No obyious sexual dimor- 
phism; three long claws plus a smaller one: 
Natatory setae reaching tip of claws. 


Mandible; (Figs 1b,c) Mandibulac coxale with 
seven teeth; Jast tooth longer than penultimate 
and near its base externally with three setae of 
different sizes; longest one pilose. Endopod 
with © bristle short, narrow and barren; 8 
bristle thick, stout and tufted; y bristle longer 
than terminal segment, thick and with short 
hairs on external side 


Rake-like organ; Six or seyen short and stout 
teeth with an additional bifid one on inner side 
of each rake, 


Maxillula: (Pig. dd) 3rd lobe with tap loathed 
Zahnborstens length width ratio of both palps: 
BV 1. 


Maxilla: Sexually dimorphic: in male, palps 
strongly asymmetrical with right one broader 
(Figs thi), tn female (Pig. tg) endopod 
with three plumose setae, longest one in 
middle and two others of unequal length, Th 
both sexes, epipod with five long plumose 
Strahlen and one shorter plumase one near 
base of plate. 

Thoracopoda t+ (Fig 17) Penultimate segment 
weakly divided. Two setac on geniculate jomt 
helween Ist and 2nd segment; antertor seta 
longest. 


Thoracopoda 1; (Wig. 11) Bod of last segment 
with two unequal selae and terminal pincers. 


Hemipenis: (Fig, 1k) Lateral lobe boot-shaped 
with base rather broad and inner distal end of 
copulatory sheath of similar shape to lateral 
lohe 


Zenker organo; Both ends rounded and with 40 
rosettes, 


Furcu: (Fig. 11) Peetinate claws unequal with 
posterior claw 0.7 of length of anterior one 
and anterior set 0.17 of anterior claw, and 
shorter than posterior seta. 


Fureal attachment: (Fig, 1m) Long and nar- 
row with distal etd bifureate: ventral and 
dorsal branches of similar width and length. 


Eye: Cups of nauplius eye fused; dark brown. 
Calouwr of shell: green, 
Sizes 
holotype adult male 
L H L H 
“LV 1540 REU RV 14604 Mal 
paratype adult female 
L H L. H 
LV 1700e 000K RV 1640) ORO Ye 
Type locality; Pool on the southern side of 
Light River, 20 m trom the bridge on the 
Port Wakefield Road, north of Adelaide, S.A- 
(34°33°09"S, 138°27'20"B), 


Derivation vf name: From the aboriginal lan- 
guage, worooa meaning green for the colour of 
the shell. 


Ecoloxy and distribution: A. wereoa inhabits 
lakes and temporary pools, This Species 1s 
found in fresh waters and its highest salinity 
record is 3.5%, at the type locality. A. wareea 
has also been recorded from the following 
localities: roadside pool, 13 km east of Rocky 
River and Dueck Lagoon, both on Kangaroo 
Island, S,A., and Granite pool at Newmann’s 
Rocks, 140 km east of Norseman, W.A, 
Additionally, valves of this species have been 
found in subsurface sediments at Birchmore 
Lagoon, Kangaroo Island (in those specimens, 
the asymmetry of the valves is more pra- 
nounced with the right valve forming a broader 
hump dorsally). This species has also been 
described by Chapman (1919) as Cypris 
sydneia {rom Pleistocene (fede Chapman) 
sediments from Boneo Swamp near Cape 
Sehanck, Vic. 

Remarks; A, worvoa is related ta the Hetera- 
evprix species as their anatomy is similar, in 
particular the triangular shape of the right 
maxillar palp in the male and the boot-shaped 
lateral lobe of the hemipenis, The asymmetrical 
valves und the presence of faint tubercles on 
the edge of the valve are also similar to Merero- 
cypris species. The major difference is the pre- 


LLV, RV = left valve, right valve, L 
Jength. height. 


H = 


94 


P. DE DECKKER 


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—500« for a-m:; 2—5Qu for n: 


3—50z for p. LV, RV = left valve, right valve. C = carapace. All views are lateral ones except 
when indicated, 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 95 


sence of a prominent selvage in A. worooa not 
seen in Heterocypris 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; CYPRIDINAE 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, 8B. australis, B. barangaroo 
nsp. (= Chlamydotheca bennelong (King) 
sensu Sars 1894, 1896), B. nimala n.sp., B. 
pinpi usp. It is likely that Strandesia feuer- 
borni Klie, 1932 and Strandesia flavescens Klie, 
1932, both described from Indonesia, belong 
to Bennelongia. From the original description 
and illustrations of the valves (Klie 1932), it 
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 anteroventrally 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 either 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; anteroventrally 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 anteroventral 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. Antennula: (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, 2/18 


Antenna: (Fig. 6d) Natatory setae extend to 
tip 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 @ 
bristle and long pilose y bristle which is twice 
as long as last segment. 


3 The ratios of the 6 segments of B. /iarpago have 
been inverted in the original description. 


96 P. DE DECKKER 


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. g 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-l: Creek pool flowing across the road, 2 km N of Leonora, W.A. Scale: 
500. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 


Fig. 4. Bennelongia australis (Brady, 1886) 


a RV internal, detail anterior of Fig. 3d. b LV 
external, detail anterior of Fig. 3e. ¢ 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: 
across the road, 2 km N of Leonora, W.A. f-j: Roadside pool, 


Creek pool flowing 
I—100 for a-d, h; 2—SOu for e; 3—1004 for f:; 4—50u for g 


5 km S of Cunderdin, W.A, Scale: 
; 5—20« for i. 


97 


98 


P. DE DECKKER 


Fig. 5. Bennelongia australis (Brady, 1886) a RV internal, female. b LV _ internal, female. c 
LV external, 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. 1 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—100 for p-s. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 99 


Fig. 6. Bennelongia australis (Brady, 1886) a antennula, b mandible—palp, c maxillula—palp and lobes, 
d antenna, e rake-like organ, f hemipenis, g thoracopoda I, h maxilla, male, i maxilla—endopodite, 
male, j hemipenis, k thoracopoda I, 1 maxilla—endopodite, female, m furcal attachment, 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: 2004, 


100 


Rake-like organ: (Fig. 6c) Five teeth plus 
another bifid one on inside of each rake. 


Musillulas (Fig Ge) Length/ width ratio of 
palp segments: 4/1, 4/1; 3ed lobe with two 
smooth Zahnborstem and tufted thick seta at 
end of 3rd lobe near Zahnborsten. About 17 
plumose Strahleon on epipod plate. 

Maxilla: Sexually dimorphic: in temale (Fig, 
61) three unequal plumose setae at tip of 
endopod; in male (Figs 6h,i) grasping palps 
unegual, broadest one on right side—for 
chactotaxy see Fig. 6h. 

Thoracopoda T; (Fig. 62) Geniculate distal 
part of Ist segment wilh two setae, proxinnal 
one being ulmost twice as long as other. Pen- 
ultimate segment weakly divided, Toner distal 
seta of 2nd segment shorter than 0.5 length of 
3rd segment and shorter than distal outer seta 
of terminal segment, [nner distal end of penul- 
timate segment with two unequal setae. Inner 
distal sela on the 4th segment is about 0.33 
of length of distal claw, 

Thoracopoda Il: (Fig. 6k) Distal setae un- 
equal; large one more than twice length of 
other which is hook-shaped. Broad pincers 
present distally. 


Hemipenis: (Figs. 6f,j) Lateral lobe broad 
with inner distal end pointed and curved im 
ward, Copulatory sheath broadly triangular 
in shape with round inner distal end reaching 
almost curved tip of lateral lobe, Outermost 
point of sheath forms hump and coresponds 
to mid-length of inner side of sheath, 

Zenker organ: With 33 eoseites, 

Furca; (Fig. 6n) Claws narrow, long and 
unequal and posterior seta longest of the two. 
Fureal attachment; (Fig. 6m) Bifureate at 
distal end; median branch thickest and other 
two branches arched inwards. 

Eye: Dark brown with two Jateral silver 
lenses. 

Colour af shell. Green to pale green, 

Size: 


lecrolype: aduli left valve 
L 


H 
LV 1Y8sde ) 200 
adult tale 
L as Lt it 
LV 186Qu 106th RV [740u inde 
adult female 
IV 22204 1300 RV 06. 122 


Remarks; Examinauan of the type specimens 
of Chlamydatheca anstralls Brady, 1886 in the 


P, DE DECKKER 


British Museum necessitates the lollowing 
clarification. One adult left valve CL 1980 », 
H 1200,), designed here as the lectotype, is 
the valve probably illustrated by Brady (1884) 
on Plate 1X.7. The tip of the beak-like feature 
of thrs left valve is broken off. In the same 
slide, there is anather left valve which probably 
belongs to a Hereroeypris sp. Tt is likely that 
this valve is the oe thought by Brady (1886) 
to be the right valve of ©. auytralis and illus. 
sirated by him on Plate LX.8. It definitely 
Jacks the peripheral posteroventral tubercles 
aid the broad inner lamella anteriorly so 
typical of all Bemmeloneta species and the 
antcroventral flange of &, australis specimens. 
This would explain the incorrect description 
of the right valve of this species provided by 
Arady (1886. p, 91). Examination of one 
carapace of & ansiralis from a slide bearing 
Brady's handwriting in Sars’ collection in the 
Oslo Museum further confirms Brady's mis- 
identification, 

Finally, i the same slide from the British 
Museum, there are Wwe partly broken cara- 
paces of smaller Rennelongia species (length 
1360 jp. Height 800 4) with some dried soft 
parts inside, Their ypeeilic identification re- 
mains Uncertain although it is thought they 
helong to B. barangares, 


In 1894, Sars s\nonvmieed ©. ansrretiy 
Brady, 1886 with Cypris bentrefong King, 1855 
maintaining that Brady’s specimens were the 
sume 4s those of King, and by stating that the 
latter author had described the spevies from 
juvenile specimens, This Wag repeated by Sars 
(1896) and Henry (1923). Sars’ suggestion 
cayinot be accepted because King (1855) 
stated on page 63 that €. bennelong has 
“equal vulves’, This is mot the case lor adult 
Hennelangia species, Sats’ (1894) argument of 
King’s specimens being juveniles cannot be 
accepted here either since juveniles of Benne- 
longia (which have symmetrical valves) are 
cither deeply punctated all over or are stronyly 
tuherculated, These features are best scen on 
speciniens. destribed by Sars (1896) as Cypris 
lateraria King. 1855 which are juveniles of 
Benneclowgia spp. Either thbereleg or pitted 
shell would have surely been diagnosed by 
Kung otherwise, 

Sars’ specimens deseribed from Australia 
(Sars 1896) and New Zeuland (Sars 1894) as 
€\ bennelonp are true Beareloreta species burt 
do not belung to #, australis, Since they cannot 
be identified as King’s species, they are there- 


OSTRACODS FROM 


fore renamed here as 8, barangarea n.sp. This 
species js desenbed below 


Ecology and distribution: B. australis was 
originally collected from Penola, 8,A, by Prof. 
R. Tate. The specunens wete empty shells 
(frady 1886). This species inhabits. mainly 
temporary pools and, so far, has been found 
alive only in Western Australia and South 
Australia. Adull males have been found in a 
permanent luke in Western Australia, suggest- 
ing that the mode of reproduction is parthe- 
nogenetic in ephemeral environments. 
Loculities: W.A.< Roadside pool 2.3 km §S of 
Northcliffe; roadside pool Pfeifler's Road, 6 
km from Mary Peaks; Lake Sadie, east of 
Wilsou Inlet (near Denmark); creek pool flow- 
ing uctoss the road 2 km north of Leotora: 
roadside pool 5 km 8 of Cunderdin; roadside 
pool on eastern side of road betwee Quai- 
rading and Corrigin (25 km northwest of 
Corrigin); Lake Bidéy. 8.A.: Roadside pool, 3 
km cast of Rocky River, Kangaroo Island. 

Water was fresh execpt in the las) twa sites 
in Western Australia where salinity was 4.4 
and 3.3¢/,. respectively. 


Benvelongia barangareo n.sp. 
FIGS 7-8, 9 aq 

1894 Cypriy bennelong King Sars, p. 24. 
1896 Cypris bennelong King: Sars, p. 49. 
1896 Cypris lateraria King: Sars, p, 53 
1923 Cypris beinelons King: Henry, p, 278. 
Diagnosis: Area just behind beak-like feature 
of left valve slightly concave; outline of hemi- 
penis as in Piy. 8), 
Deseviption: Carapace. (External) Adult; cir- 
cular (q oval carapace smooth or covered with 
faint pustules and pilose. Ventrum flattened, 
Vulves asymmetrical: Jeft valve larger, espe- 
cially anteriorly with largest overlap ventrally; 
unteroventral region of left valve slightly con- 
eave to form beak-like feature whereas right 
Vulve broadly rounded and with an elongated 
and narrow beak-like serrated flange antero- 
ventrally, Simple type nornal pore canals, 
Juveniles More elongated in lateral view, with 
symmetrical, pitted or tubereulated valves, 
(Internal) Adult! Inner jamella twice os hroad 
anteriorly in both valves, selvage peripheral 
ventrally, away from outer margin posteriorly 
and espeeially anteriorly: inner jist forming Jip- 
Nike flap far away from edge of shell antern- 
ventrally which ix preceded hy » deep. nurrow 
groove AC ridge follows curvature of inner 
Nurgin anteriorly bul fades opposite tongue- 


MUSTRALIAN INLAND WATERS 11 


like flap. Few small tubercles are visible on 
this ridge above concave depression of shell, 
Outer list often broad posteroventrally, In 
right valve, selvage peripheral and shatp pos- 
teriorly, broad anferoventrally 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 
vow of tubercles along periphery of shell pos- 
teroventrally, Outer list also broader postero- 
ventrally and usually pitted externally near 
edge of shell; this is best scen in ventral area. 
Juvenile: In both valves, inner lamella of 
almost equal width all around and selvage, 
Which follows the curvature of the shell, 
prominent, 


Anatomy; Only the features al the anatomy 
which differ significantly from #. auyrralis and 
other Benneloneia species are mentioned, For 
other details refer to Fig. & 


Antennula: Natatary setae slighly longer than 
all segments together. 

Maxilla: Male grapsing palps (Figs 8g,h) of 
different shape than #&. australis: left palp 
shorter and broader and right one with outer 
edge forming 90°, 

Thoracopoda J; (Figs Sd.c) Inner distal seta 
of second segment at least as long as half of 
3rd scament; longest inner seta at mid-length 
of 3rd segment (where it is weakly divided) 
at least as long as 2nd half of 3rd und 4th 
segments together, Taner distal seta on 4th 
seameol 1/2.3 of length of distal claw. 


Colour of shell: Green. 
Sizer 
holotype adull male 


L L H 
LY 1120« TOD RY LlO0e 660 
puralype adult femole 
L H IF H 
LV 1} 40u TM RV 1110" 6R0u 
Newmann’s Racks adit female 
L H L H 
LV 1390a TRO RV 1320u 770% 


Derivarion of name: From the aboriginal name 
of Bennelong’s wife Barangaroo 

Type locality; Lake Buchanan, Qld (21°35'S, 
145°52'B), 


Feoalogy and distributian; B. harangaroa 1s a 
common inhabitant of temporary pools and 
usually 1s only represented by parthenogenetic 
females. On one occasion, in Lake Buchanan, 
both sexes were found, There salinity Was 


102 P. DE DECKKER 


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 
RY, paratype male. j C showing RV, paratype female. k C dorsal, paratype male. | 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—250» for a-m; 2—100« for n-0; 
50“ for q; 3—50 for p; 4—100 for r. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 103 


Fig. 8. Bennelongia barangaroo, n.sp, a antennula, b antenna, c mandible—coxale, d_ thora- 
copoda I, e maxillula—palp and lobes, f mandible—palp, g maxilla—endopodite, male, h maxilla; 
male, i thoracopoda II, j hemipenis, k furca, 1 maxilla—endopodite, female, a-b, d-k: holotype 
adult male; c 1: paratype adult female. Scale: 1004. 


104 


Fig. 9. Bennelongia barangaroo n.sp. a 


P. DE DECKKER 


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. | 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—500u for a-k; 2 


3—500- for r. 


1004 for m-p; —20- for q; 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 105 


4.1%, In other localities listed below, water 
was known to be fresh, 

Live specimens with iuberculated symmetri- 
cal valves collected in a farm dam at Frome 
Downs near Lake Fronie, §,.A, were examined 
in the Jaboratory. They resembled specimens 
illustrated by Sars (1396a) on Plate VEL; Fly. 
3 and labelled by him as Cypriv fateraria King, 
1855. After a few days, these specimens, which 
had traces of ovaries inside the valves, were 
seen lo moit into B. bararngaroe with its typical 
asymmetrical valves, Kees of the latter 
developed into small highly ornanrented ostra- 
cods like ©. lareraria, ‘This phenomenon 
requires the two taxa to be synofiymized. 


Localities: W.A.; granite pool, Newmann's 
Rocks, 140 km E of Norseman; roadside ditch 
37 km W of Esperance {road ta Ravers- 
thorpe) and 3,5 km east of Dalyup River: 
small roadside pool about 18 km northeast of 
Menzies; clonyated pool in bed of Coakarrow 
Creck, about 5 km W of Wiluna; pool in creek 
bed about 25 km NW of Cue; small farm dam 
oo eastern side of road 9 km S of Cunderdin 
on the way to Quairading. S.A.: Farm dam 
al Frome Downs, near Lake Frome, Old: 
Creek, 22 km E of Richmond; roadside pool 
at Miranda: Lake Galilee, near Arainae: Lake 
Dunn, south of Lake Galilee. N,S.W.: Speei- 
mens raised by Sars from sample of dried mud 
collected in waterholes by Mr Whitelegge in 
Bourke Street, Sydney (Sars 1896), New Zea- 
land: Specimens raised from dried mud sample 
collected near Kaitaia in the North Island, 
Chapman (1963) stated that this species had 
not been found in New Zealand since Sars? 
(1894) description and no further Iocalities 
are provided in Chapman & Lewis (1976). 


Remarks; B. barangaroo is closely allied to 
B. australis but the species ean be separated 
ou the basis of size (#, australis ig much 
larger), on the outline of the hemipens and 
the chactotaxy al the tharacopoda I. 

The specimens of B. barangaroo described 
here are identical in morphology to Sars’ 
specimens from Bourke Street, Sydney and 
fram New Zealand, It is assumed here that the 
specimens reported as Cypris lateraria by Sars 
(1896) from the Sydney site are juveniles of 
B. barangarao 

The shape and width of the anteroventral 
flange on the night valve of B. berangarve can 
vary extensively: in the specimens from Lake 
Buchanan, the edge of the flange is serrated 
(Pigs 7p.q). 


Bennelongia nimala (sp. 
FIGS 10-11 


Diavnosis: Oval to subrectangular, pustulose 
carapace with posterior slightly pomted and 
ending with one or two spines. Homp-ike 
thickening of shell anterodorsally, Posterior 
seta of furca about 0.7 of length of posterior 
claw and lateral outline af hemipenis as in 
Fig. 11g. 

Description: Carapace, (External) Adult; Oval 
to subrecrangular-shaped with posterior slightly 
pointed and ending with one or two spines, In 
dorsal view, eeeshaped with anterior end 
narrow and more pointed than pasterior. 
Valves obiously asymmetrical in anteroventral 
area: there, left valve formed like pointed 
beak whereas in right valye, it is broadly 
rounded and there is a small beak-shaped ser- 
rated flange. Posterior to ange, edge of right 
valve slightly concave, Tip of beak does not 
reach horizontal plane formed by ventral area, 
Greatest height at about 0.33 from anterior, 
Shell pilose, pseudopunctated and pustulose 
nearly all over. Along edge of lett valve, 
especially, pustules mare concentrated and 
some are pointed, especially anteriorly, 
Juvenile: Pseudapunclate. subtriangular stell 
with many pointed tubereles scattered all over 
but with greater concentration anteriorly and 
posteriorly, Valves symmetrical, Tn dorsal view, 
oval-shaped with both ends painted. 
(Taternal) Adult) Inner famella broadest 
anteriorly in both valves, In left valye, selvage 
narrow and peripheral; inner list forming lip- 
like flap anteroventrally, This flap fades in 
anlerodorsal region, In front of flap in ventral 
area, a deep narrow groove. Between outer 
margin and just behind beak ventrally, is a 
tidge which js tuberculate in some specimens, 
In anterior urea, in front of hinge area, ts a 
deep groove running parallel to curvature of 
shell which is placed near oyter margin. This 
groove absent in beak-like area, In right valve, 
selvage narrow and runs parallel io curvature 
of shell except at niid-height anteriorly where 
it is not vistble, Anteriorly, tn region where 
serrated flange present. a small ridge rung 
parallel to, and between. selyage and inner 
margin, 

Juvenile; Inner lameflae equal in both yalyes 
und of same width all along. Selvage broad and 
peripheral, 

Anatony- Same remarks as for B. haranrarde 
Vor details ot anatomy, refer to Fig, 11, 


106 P. DE DECKKER 


a 


wie Me 
ay Y, 
= "| m 


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-l, n-o: Georgetown Lagoon, 
near Jabiru, N.T. (type locality). d, i, m: Buffalo Billabong, near Jabiru, N.T. Scale: 1—S50Qu for 
a-i; 2—200” for j-m, 0; 3—100 for n. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 107 


Fig. 11. Bennelongia nimala nsp. a_antenna, b rake-like organ, c antennula, d_maxillula—palp 
and lobes, e mandible—palp, f maxilla—endopodite, male, g hemipenis, h maxilla—endopodite, 
male, i thoracopoda I, j thoracopoda II, k maxilla—endopodite, male, | maxilla—protopodite, female, 
m furcal attachment, n furca. a-c, e-g, i, k-m: holotype adult male; d, h, j, n: paratype adult male, 
Scale: 1—100 for a, c-n; 2-—50u for b. 


108 P, DE DECKKER 


Antennula: (Fig. llc) Natatory setae as long 
as all segments together. 

Maxilla: Male palps asymmetrical (Figs 
11f,k) with right one broader and less arched, 
Two external setae on female palp (Fig. 11h) 
of equal length and plumose. 


Thoracopoda I: (Fig. 11i) 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. 11g) 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. 11n) 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. 


Size: 
holotype adult male 
L H L H 
LV 15002 840" RV 1340u 760" 
paratype adult female 
L H L H 
LV 16404 960u RV 15404 860u 


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 
valve. 


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 
juveniles, 


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 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 109 


Fig, 12. Bennelongia 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—500 for a-p, I-n, p-s; 2—200 for k; —300u for t; 3—150« for o. 


110 P, DE DECKKER 


Fig. 13. Bennelongia pinpi, osp. a maxillula—palp and lobes, b antennula, c antenna, d man- 
dible—palp, e hemipenis, f maxilla—endopodite, female, g thoracopoda T, h thoracopoda II, i 
maxilla, male, j maxilla—endopodite, male, k furca, | furcal attachment, a-c, e, g-k; holotype adult 
male; d, f, 1: paratype adult female. Scale: 200. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS ill 


ratio of distal seta of 4th segment and distal 
claw 1s 1/3.5, 

Hemipeniss (Rig. 13¢) Outer lobe broad and 
digitute and. near its base on inner side, 
copulatory sheath forms broad  trapezoid- 
shaped extension, 

Zenker organ: With 42. rosettes. 

Furea: (Fig. 13k) Sctae almost equal-pos- 
jenur one longer and about 0.5 length of 
posterior cluw. 


Colour ef shell: Light green. 


Sires 
holotype adult’ male 
i} H 1 H 
PV 24800 16008 RY 2340u L420 
puralype adult female 
i H L H 
LV 2580u 1640u RY 24404 1520" 


Type locality, Pine Tree Creek Lawoon. N of 
Hughenden, and exactly t6 km S of Louisa 
Lake on road to Hughenden, Qld (20°00'30" 
S. 144°16'44°B), 

Derivation of name; The aboriginal word pinpi 
in Queensland meaning parrot is chosen for 
the unteroventral area of the shell which 
resembles the beak of a parrot, 

Eeology and distribution: This freshwater 
species is known from three other localities in 
Queensland; Agnes Lake, Salt Lake (fresh) 
and Louisa Lake, all three N of Hughenden, 


SuprAmMity! BUCYPRIDINAE Bronstein, 


1947 
Australacypriy De Deckker, 1974 
Type species: Australecypris robusta De 


Deckker, 1974. 


Remarks: The following species ate discussed 
in the present work and can be distinguished 
on the outline of the hemipenis; A. divpar 
nsp. A. dnsuluris (= A. hypersalinu), A, 
rectaneularis and 4. rabustit. AU Anstralo- 
evariy species are balobiont and planktic but 
can also crawl on lake floors, Rarely do two 
species aceur together in one lake. 


Anstralocypris dispar n.sp. 
FIGS 14, 15 ai 
Diagnayis: Lateral Jobe of hemipenis digitate 
and broad at base; long distal seta on 4th seg- 
ment of thoracapoda I, 


Description: Cuarapace. (External) Pseudo- 
punctate, subrectangular carapace with an- 


terior end broadly rounded, dorsum slightly 
inclined and posterior steeply inclined. Ven- 
trum strongly concave past oiid-length from 
the anterior. Greatest height ut about 0.4 in 
female and 0.33 in male, In dorsal view, cara- 
pace clongated, length more than twice width, 
and both extremitics slightly pointed. Left 
valve slightly larger in female and ventral 
overlap minimal. Both valves extend over ane 
another in anterodorsal area just before hinge, 
Simple tiormal pore canals. 


(Internal) Inner lamella narrow in both valves 
and selvage faint and between edge of valve 
and selvage which is prominent in that area, 
Radial pore canals numerous and straight. 
Central muscle field consisting of five adductor 
sears and two Jarge mandibular ones in fran 
and below. 


Anatomy: Antennula: (Fig. Ide) Length’ 
with ratio of last sik segments; 2/3, 2/1, 
Wt, 5/35, 271.4, 4/3. Small warlike 
“sensory” organ on side of 2nd segment. Nata- 
tory setae us long as last six Segments together, 
Antenna: (Fig. L4a) Massive and sexually 
dimorphic: four claws in male and three in 
female: smallest one in male attached to last 
segment and with long comb-like tecth. Nata- 
tory setae almost reaching tip of claws, 


Mandible: Mandibular coxale (Fig. 14h) with 
seven teeth, the last one longer, narrower than 
penultimate and, at base, three setae occur, 
two of which are pilose, Endopod (Pig, 14d) 
with a bristle short, narrow and barren; # 
bristle stout and finely pilose, y bristle twice 
as long as others and pilose im distal hall, 
Distal segment of endopod 0.33 of length of 
penultimate segment. 


Rake-like organ: Seven short and stout teeth 
plus one inner bifid one on each rake, 


Maxilula: (Pig. 14b) Endopod with 23 
plumose Strahlens length/width ratio of palps: 
3/1.3, 1.5/1.4 with last segment slightly tra- 
pezoidal. Two toothed Zahnborsten on 3rd 
lobe. 


Maxsilla: Sexually dimorphic: in male (Figs 
14j,kK), palps almost symmetrical. strongly 
eurved and narrow; in temale (Fig, '4f), 
palp faintly divided at its extremity aid 
with three short pilose setae, the middle one 
heing twice as long as others. 


Thoracopoda IT: (Fig. lde) Penultimate seg- 
ment weakly divided; distal segment with long 
inner seta 0.5 length of distal claw, Thoraco- 


112 P, DE DECKKER 


. wy Sy 


\ 

Fig. 14. Australocypris dispar n.sp, a antenna, b maxillula—palp and lobes, c antennula, d mandible— 
palp, e thoracopoda I, f maxilla—endopodite, female, g thoracopoda IJ, h mandible—coxale, i 
hemipenis, j maxilla—endopodite, male, k maxilla, male, | furca, m furcal attachment. a, c-e, g-m: 
holotype adult male; b, f: paratype adult female, Scale: 200u. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 113 


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, fernale 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—200u for j-n, p-q; 3—20u for o. 


14 BP DE DECKRER 


poda Il: (Fig. 4g) Elongate and narrow seg- 
ments distul segment with two terminal setac 
ahd two pincers; shortes! seta hook-shaped. 


Hemipenis: (Fig. 141) Lateral lobe digilate and 
slightly curved imwards and broad at its base; 
copulatory sheath broadly heart-shaped. 
Zenker organ; Long and narrow with both 
ends rounded and usually 67 rosettes, 

hurca: (Fig, 141) Unequal claws with anterior 
one 0.66 longer, and anterior seta slightly 
longer than posterior one, 

Furcal uttuchment; (Pig, 14m) Proximal part 
hifureate; dorsal branch strongly arched and 
aboul same length as ventral one which ollows 
curvature of median branch. 


Culour of shelly Nacreous brown, 


Siaey 
holotype adult male 
R H L. H 
LV 4600 22004 RV 4600 = 20604 
puralyoe jdull female 
L- H 1, i 
LV dni 224()u RV 4000u 2060 


Type lacaliiy: Ephemeral salt lake south of the 
Coorung Lagoon, S.A, (36°26°45"S, 139°47' 
32"E) = tne.5 in De Deckker & Geddes, 1980, 
collected on 16.X.1978 at 15.5%) salinity, 


Derivation ef iame: Brom Latin dispar mean- 
ing different for the unusual lon distal seta 
on the list segment of the thoracopoda | com- 
pared to congeners. 


Ecnlogy and distrihutian; A. dispar is a halo- 
biont species which oceurs in ephemeral salme 
lakes in S.A, Near the Coorong Lagoon. 
iis salinity range is 9-527), De Deckker & 
Geddes (1980) provide further details on ats 
distribution there as “4. nsp.l” tr is alse 
recoriled on the Yorke Peniosula at 6%, in 
in ephemeral lake (35°03'L1"S, 137°35°50"B) 
and on the Eyre Peninsula in a swamp south 
of the roud near Lake Horn, northwest of Port 
Lincoln at 32.45; 


Remarks: Ay dispur is characterized by the 
unusialy loog seta on the distal segment of the 
tharacupoda }oand the fyintly divided end of 
ihe female maxillar endopodite palp, These 
characteristics are unknown in other Ansiralu- 
eypriy species bul are found in the balubiont 
ostracad = Lintanacypriy  lvriduy Shornikov, 
INGL, described From the Kuban Delta in the 
Caspian Sea region of the USSR. Correspon- 
dence with De E, Sharnikov has led to the con- 
clusion thal other features of the chavtotaxy of 


many appendages between A. dixypar und L, 

luridus difter too greatly to justify close rele- 

tionship between the two species, 

Australecypris (nsularis (Chapman, 1966) 

1966 Bvevpriy insularis: Chapman, p. 375. 

1974 Anstaloevpris Ipeisalina: De Deckker, p 
101. 

(978 usteuloevpely hypersalina: De Deckker, p- 
16. 

1978 Avs(raloeypris Tavulariy: De Deckker, p- 17, 


Remarks; Alter examination of many collec- 
lions of A. insularity from Western Australia 
(from Where the species had originally been 
described), it became olear that A. insularéy 
and 4. hypervaling ure conspecific. Both taxa 
huve very similar hemipenes with the charac- 
feristic broad and curved lateral lobe and the 
bulbous inner distal end of the capulatory 
sheath, This type of lateral outline of the hemi- 
pens is not found in congeners, 

The cxamination of many specimens af 
Auvtealoeyprix has led to the conelusion that 
there wre slight differences in the morphology 
of many specimens of A, Jnvulariy and A. 
Aypersalina. Therefore the minor differences 
of the oullme of the hemipenis for the two 
faxa originally mentioned by De Deckker 
(197A) are here considered insignificant. 

During the revision of all mytilocypridinid 
species (De Deckkor 1978), it appeared that 
A, (nstlarly and 4, hypersalina differed mainly 
on characters of the shell, but it has since 
hecame apparent that the shape of the shell of 
th Austfaloeypris species is also variable in 
populations taken during different seasons in 
the same lake. Large specimens collected dur- 
ing Winter and carly spring months are usually 
more clongaled. For the sume reason, the 
Specimens illustrated in De Deckker, 1978, 
in Figs 18ab as A. divpersalina show the 
typical winter form af A, Aypersalina = A, 
inyulariy pot recognized then, as most collec- 
tions deseribed im that publicution had been 
taken during summer months, 


Keology and disirihutien; ‘The distribution of 
A. insulariy (plus 4. bvpersalina) Was dealt 
with in De Deekker (1977, 1978) and, is 
Wpdated here, The species is commonly found 
it ephemeral saline lakes near the Coorong 
fLugoon (sce De Deckker & Geddes 1980) 
where it is found between § and 131%,, salinity, 
fr also Oveurs on the Yorke and Eyre Penin- 
suila dit similar ephemeral Jakes at the sume 
salinity range, It has also been widely enllected 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 115 


in LS West Australian salt Iakes by Gerdes 
eral (1981) over the range of 2.9-122.6'y,, 


Australocypris rectangularis De Deckker, 
1978 
LOTS Aastealoevprix? recianeularis: De Deekker 
p. 17, 
L980 Australocy pris reciangularis: De Deckker & 
Geddes. p. 641. 
Piaenesis; Lateral lobe of hemipenis warrow 
ind hook-shaped; copulatory sheath almost 
semicireular. Carapace rectangular, 
Remark: The original deserption of uns 
species way incomplete because a number of 
appendages had dried out and had been 
damaged, A number of specimens have since 
been collected in saline Inkes near the Coorong 
Lagoon by De Deckker & Geddes (1980). 
They examined the appendages pf this species 
and on morphological grounds referred it with 
confidence ta Australoey pris 
Ecology and disiribution: This species is a 
truc halobiont farm which has never been 
found in salinities below 50%,. Near the 
Coorong Lagoon, its salinity Tange is 50- 
195", and there is evidence there that it even 
hatched above 73%, and 115%. in two dit- 
ferent localities, Qn the Vorke afd Eyre 
Peninsulas, 4. rectanguleris oecurs in the same 
range of salinities as in the lakes. near the 
Coorong Lagoon. 


Australocynris robusta De Deckker, 1974 


1974 Anstralaevpriy robusta: De Deckker, 1974 
Diagnosiy: Chitinous pocket on inside af pos- 
terodorsal area in female valves; hemipems 
with digitate lateral Tobe which is of about 
same width all along. 


Ecology and distribution: The salinity range of 
A. rabusta in Victoria, already available in 
Bayly & Williams (1966). of 53.5-93,1%) 
and of 44-192%, in Geddes (1976) js 
hroadened to the range of 7-145, obtained 
fram collections made im many lakes jn 
January 1980, This particularly braad range 
was nol reached by specimens of the same 
species in lakes in the Coorong area of South 
Australia. There the range is 15-38%, (De 
Deeckker & Geddes 1980). The query concern- 
ing the validity of Geddes* (1976) record of 
the salinity for 4, rebysta in Victoria, made by 
De Deckker & Geddes (1980, p. 691) is re- 
solved sinee further work has demonstrated 
that the salinity range of A. robusta definitely 
differs between Victorian avd South Australian 


lakes, Similarly, Geddes (1976) could not 
fave misidentified A, robuyra since it is the 
arly Australacypriy species. recorded in his 
study areca, 

In Victorian lakes, A, rebusta is found in a 
heulthy state and in high numbers gt salinities 
between 45 and 77.5%. and nearly always 
oceurs with Diaeypris compacta which is 
found in even higher numbers, At higher 
salinities, these two specics are found with 
Platyeypris baueri, A. robusta occurs in two 
permanent salt lakes in Victoria, Lakes Gnotuk 
and Kejlambete, which have salinities in the 
vicinily of 60%, the whole year cound. This 
indicates that this species does not necessarily 
require a sharp decrease in salinity to hatch 
although Geddes (1976) showed the hatching 
range for 4. robusta to be 8.5-LO8'-- 


CYPRICERCINAE 
197) 
Strandesia Vavra, 1895 


Type species; Strandesia mercatorunt (Vavta, 
1895). 


SupeaMiny: McKenzie 


Strandesia phoenix nsp. 
FIGS 15 }-q, 16 

Diaenosis: Strandesia without shell ornamen- 
tation, with left valve larger than tight valve 
all along and overlapping it ventrally, except 
in the anteradorsal 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 af right valve at about 0.33 of height 
from dorsum and of left valve at 0.66, 
Deseription: Curapace. (External) Pseudo- 
punctate ellipsoid shell with dorsum and yen- 
trum gently curved, anterior slightly more 
rounded (han postetior which tapers gently; 
in dorsal view oval wilh 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 
yalve latger than right all along and over- 
lapping it ventrally except in anterodorsal afea 
at extremity of hinge where it is overlapped by 
right valve normal pores of simple type, some 
rimmed, others funnel-shaped. 

({nternal) Inner lamella broadest anteciorly 
atid almost absent posteriorly in both valves; 
in left valve flange broad all along except 
dorsally: inner lamejla near outer snargin 
perpendicular to flange in anterior of Jeft 


116 P. DE DECKKER 


Fig. 16. Strandesia 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 I, i furcal attachment, j 
furca. a-j: holotype adult female. Scale: 100x. 


OSTRACODDS PROM AUSTRALIAN ENLAND WATERS 17 


yalve and curved inward in its middle; this flat 
area met by broad selvage of right valve; 
selvage faint and peripheral in left valve and 
brond all along in right valve, 


Anatomy: Antennula: (Fig. 16a) 7-segmented: 
length/width ratio of last six segments: 1/ 1,8, 
(/4. 1/1, 1.6/1, 2/1, 3/1) natatory setae as 
long as last six segmenis together, 


Antenna: (Fig. loc) Four almost equal claws: 
three on penullimale segment and one on distal 
segment} natatory setae extending past tip af 
claws, 


Mandible: (Bigs 16d,f) Mandibular coxale 
with seven teeth; distal tooth acicular; epipad 
with five long and one short Strahlen; distal 
segment of palp rectangular and with long. 
slim and barren a bristle, & bristle stout and 
pilase, y bristle thick and twice length of distal 
segment and pilose on inside of js distal half. 
Rake-like organ: Six teeth plus one bifid on 
inside of each rake, 

Maxillola: (Fig. 16b) Length/ width ratio of 
palp segments: 3/1, 3/1; two teothed Zahn- 
borsten on ard lobe. 


Manilla: (Fig. 162) Middle seta more than 
twice length of other two which are of almost 
equal length. 


Thoracopoda I; (Fig. 16) First segment with 
two unequal setac, distal one pectinate and 
0.5 length of othery 3rd segment undivided in 
middie where long seta accurs, 


Thoracopoda Ul: (Fig, 16h) Distal pincers 
large and distal sctae unequal: shortest one 
curved, with distal half camb-like and 0.5 
length of other smooth one: middle seta on 
last segment less than 0.5 length of all other 
setac Which are of similar length. 


Furca: (Fig. 16j) Very long furcal shaft, twice 
length of longest claw; anterior sela more than 
twice length of other seta furcal attachment: 
Median branch narrow and gently curved (fig. 
16)) dorsal branch forming eyelet whereas 
ventral branch straight. 


Coaleur of shell> Purple, 
Size: 
holotype adult. Female 
lL. if i: H 

Tv §4n S10e RV 860e S40 
Type locality! Coastal dune luke at Evans 
Head, southwest af Lismore, N,S-W. (29°06" 
57°R, 152°25°40"E), 


Derivation of name; Brom Greek phoenix (= 
purple) for the colour of the shell, 

Ecology and distribution; 8. phoenix has only 
been collected from the type locality, Water 
was fresh. No males have been found and ‘no 
sperms were noticed in the ovigerous Cemales, 
suggesting that the species was pacthenogenetic 
at that locality, 


SuBPAMILY! DIACYPRIDINAE McKenzie, 
1978 

Reticypris McKenzie, 1978 
Type species: Rerleypris herbsti McKenzie, 
1978. 
Diseussion: Reticypriy is an Australian en- 
demic genus Which groups the (ollowing halo- 
biont species: R. herbsti McKenzie, 1978, R- 
walbu De Deckker, 1979, R. clave o.sp. and 
R, kurdimurka np, R. dedeckkeri McKenzie, 
1978, has been synonymized to R, herbsti by 
De Deckker & Geddes (1980) as both taxa 
have an jdentical hermipenis (see discussion 
below). All Rericypris species, which are casily 
distinguishable (rom one another by the outline 
of the hemipenis, are discussed below, 


Reticypris clava nsp. 
FIGS 17-18 
1980 Reiicypris sp.noy.1, De Deckker & Geddes, 
p 692. 

Diavnasis: Retievpriy with lateral lobe af 
hemipents crescenti¢ and broadest at base 
where two lumps are visible on inner side. 
Deyeription; Carapace. (External) Subrectan- 
gular with dorsum gently arched and ventrum 
almost flat except in mouth region which is 
concave; anterior broadly rounded and pas- 
lerior tapering with posterodorsal area inclined; 
slight depression at extremity of hinge 
anteriorly; left valve slightly larger 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 clase to outer 
margin, narrow and thinly denticulated except 
in mouth region where it is straight: greatest 
height at about 0.4 from anterior, in dorsal 
view like a flattened oval and both extremities 
slightly pointed. Normal pore canals of simple 
type and rimmed, 

(internal) Inner lamella equal in both valves 
and broadest anteriorly, tapering to 0.66 of the 
width posteroventrally. Hinge consists of broad 


118 P. DE DECKKER 


TTY hom 
Lat Mote es! fs 
te 


aa 


* 
a 
4 


at 


ee 


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—200u for a-n; 2—100u for 
o;—S0u for p. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 119 


V 
StestasteasurnsE 


VANATAVATANANATATAVANAVAN ANNAN 


Fig. 18. Reticypris clava n.sp, a _antennula, b antenna, ¢ mandible palp, d maxillula, palp and lobes, 
é maxilla, male, f maxilla, endopodite, male, g thoracopoda II, h thoracopoda I, i hemipenis, j 
maxilla—endopodite, female, k furcal attachment, | furca, m Zenker organ. a-i, k-m: holotype adult 
male; j: paratype adult male. Scale: 1004. 


120 


groove in left valve in which interlocks right 
valve; tadial pore canals numerous and 
straight, 


Anatamy; Antennula: (Fig, 18a) 7-seamented; 
length/ width ratio of last six segments: 1/15, 
Wd, 1/1, 1/08, 172, 1/15 matatory setae 
slightly longer than. all se@ments together. 

Antenna: (Fig. 18b) Three loug and equal 
claws On penultimate segments with shorter 
claw on distal one which is pectinale in male; 
natatory setac extending past tip of claws. 


Mandible; (Fig. 18d) Seven teeth on mandible 
coxale: last one longer than other three adja- 
cent to it and. near its base, two short pilose 
setae; length/width ratio of palps; 4.5/1, 
1.7/1; 3rd lobe has two smooth Zahnborsten; 
apipod with five plumose Strahlen and 
smaller barre one. 


Rake-like organ: Right teeth with an addi- 
ional bifid one on inner side, 

Maxillula} (Fig, 18c) Distal segment squarish; 
a, § and +» bristle of equal length and slim: 
a smooth, / pectinate, y smooth, 


Muxilla: Sexually dimorphic; in male, pulps 
asymmetrical (Figs, |8e)F): broadest forming 
right angle on outside whereas other more 
arched; in female (Fig. 18j) setae unequal 
with shartest smooth whereas other two 
plumose; for chaetotaxy of protopod see Fie, 
18e, 


Thoracopoda [; (Fig, 18h) Penultimate seg- 
ment undivided and all inner setae long and 
of about equal length, 


Thoracopoda I: (Fig. 18g) Distal pincers 
small and distal setae unequal: shorter one 
curved and 0,25 length of other, 


Hemipenis; (Fig 18i) Lateral lobe erescent- 
shaped and broadest at the base where two 
lumps are visible on inner side: inner Jobe 
squarish with three sides concave. 

Zenker organ: (Fig. 18m) Elongated with 16 
rasettes. 

Furea: (Fig. 181) Pectinate claws thick and 
equal; selae equal, pectinate and 0.33 length 
of claws. 


Furcal attachment; (Fig. 18k) Median branch 
straight and of about same length as curved 
inward dorsal branch; ventral branch forming 
ohtuse angle with dorsal ane and hook-shaped 
Uistaily; short rod-like, extension at right angle 
on base of median branch, 


Eye: Cups of nauplius eye fused. 


P DE DECKKER 


Colour of shell> Light green to white, 
Size 
holowpe adult male 


L " L H 
LV 70H 4A RV 59h 4a 
Purutype adult fenrale 
L H | H 
LV 720K Ags RV 700u aay 


Type locality: Ephemeral salt lake S of the 
Coorong Lagown, S.A, (36°13°36°S, 199%41° 
29"E) = locality 3 of De Meckker & Geddes 
(1980). 


Derivation ef name: From Latin elava (= 
club) for the diagnostic shape of the outer 
lobe of the hermpenis 


Ecology and distribution: in collections from 
Western Victorian lukes, R. clava was always 
accampuniod by low salinity ostracods such as 
Mytilocypris splendida or M. praenuneia, and 
occasionally with D, spinosa. Salinity for these 
collections ranged between 4 and 42%,. In the 
lakes adjacent to the Caorong Lagoon in South 
Australia, the salinity range for R. clava is 
S-131"/, but it was never found in high num- 
bers above 68',. Only in one lake near the 
Coorong Lagoon (locality 7 of the De Deckker 
& Geddes |980) was R, clave found together 
with Rv herbstimthis co-eeurrence persisted 
throughout the year, R. clava has been col- 
lected once in Western Australia between 14.6 
and 59.5%, (Geddes er al, 1981), 


Resharks; Wt is difficult to distinguish R. clava 
and R. herhsti on Features of the shell as reti- 
culation of the shell and even shape and size 
are Known lo vary (sce Fig. 17 for R. clave), 
The ouiline of the hemipenis is 9 good chag- 
nostic feature for separation of the twa species 


Relicypris herbyti McKenzie, 1978 

IS78 Refevpris herbsti rsp. McKenzie, p. 188, 
Diagnostic; Reticypris with lateral lobe of 
hemipenis boot-shaped 

Deseription; See McKenzie (1978), p. 188-9. 
Ecology and disteibation: RB. herhstl is tolerant 
to higher sulinities than R. elava. In western 
Victorian lakes it was found with 2. compacta 
at salinities between 99 and 172%, whereas, 
in the Jakes near the Coorong Lagoon, it 
occurred at salinities between 12 atid 141%, 
with three supplementary records at 195, 216 
and 218%). It is found im high numbers be- 
tween 104 and 1247, salinity. 


Ina humber of specimens, collected in lakes 
near the Coorong Lagoon, specimens with thin 


OSTRACODS FROM AUSTRALIAN INLAND WATERS mi 


and faintly 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 also with juveniles, The ecological signifi- 
cance of these differences is not known, 


Reticypris kurdimurka f.sp. 
FIGS 19-20 

DPiavnosis: Reticypris with outer lateral Jobe of 
hemipenis spout-shaped and copulatory sheath 
heart-shaped. 

Description: Carapace. (External) Subrectan- 
enlar to squarish ip Jateral view with both 
valves usually thick; reticulation resembles 
broad punctation; anterior and posterior simi- 
lar and broadly rounded; ventrum flat execpt 
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, it slopes gently, outer lamella 
thin and peripheral; left valve slightly larger 
all around and in some specimens overlap 
of Jefe valve over right one is obvious, at 
both extremities of the hinge, as it forms 
smooth elongated humps. 


(Internal) Toner lamella shghily broader an- 
teriorly compared to posteroventral area; sel- 
vage faint and peripheral iy right valve 
whereas uf a distance from outer margin in 
jeft valves: radial pore canals narrow and 
straight: hinge consists of a broad groove jn 
left valve in Which right valve interlocks, 
Anatomy: Antenniila! Fig, 20a) 7-segmented 
length/ width ratio of last six segments) 1/ 1,25, 
W/), 1/25, 1/2, 1/2, L/ls natatory setae 
longer than all segments together. 

Antenna: (Fig. 20b) Three equal lone claws 
on penultimate segment; 4th claw on distal 
segment reaching tip of other claws and pec- 
tiuate in male. whereas it is shorter and with 
sutaller teeth in female; natatory setae reaching 
lip of claws. 

Mandible: (Fig. 20h) Mandibular coxale with 
seven teeth; palp with distal segment squarish, 
a bristle short, slim and smooth # bristle short, 
stout and pilose, y bristle Lwice length of distal 
segment and pilose in its distal half. 
Rake-like organ: Right teeth with an additional 
inmee one which is bifid. 


Maxillula; (Fig. 20¢) Distal palp almost 
squarish and 3rd lohe with two sinooth 


Zahnborsten, 
Maxilla! Sexually dimorphic: my male (Figs 


20¢,f) 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 are cqual- 

Thoracopoda |: (Fig. 20d) 3rd segment un- 
divided: inner setae small and unequal. 


Thoracopoda Lf; (Fig, 201) As for R, elava. 
Hemipenis: (Figs. 20k,1) Lateral lobe spout- 
shaped and broadest at mid-length; copulatory 
sheath heart-shaped, 

Zenker organ: (Fig, 20) Elongated with 11 
rosettes. 

Furea; (Fig, 20m) Claws equal and thick; 
setae equal and small, about 0.25 length of 
claws. 

Furcal attachment: (Big. 20)) Median branch 
and long dorsal one gently curving) ventral 
branch forming a right angle with dorsal 
branch and forming a loop distally; vertical 
rod-like extention hear base of median branch, 


Eye; Cups of nauplius cye fused, 


Colour of shells White when preserved in 
alcohol. 


Size: 
holotype adult: male 
| H L 
LV S56Su 35 0h RV S5n0u 330u 
naraiype adult fernale 
) it l. H 
LV 600« aye RV 590" 375u 


Type locality: Madigan Gulf, Lake Eyre, South 
Australia. 


Derivalion ef name: Kurdimurka is an abori- 
ginal name for a legendary creature supposed 
to inhabit the bottom of lagoons and creeks 
in the Lake Eyre district. 


Eevlogy and distriburtan: Qne specimen of R, 
kurdimurka had been originally collected from 
Lake Eyre North an 28.1V_1975 at about AQ. 
salinity when the lake was last flooded (see 
Bayly 1976, p, 664 where it is referred to as 
“andeseribed cypridid genus”). Subsequently, 
it has been collected twice fom Madigan Gulf, 
ut Lake Eyre by W. Zeidler on 11-X11.1974 
and 20X1L.1975—no salinity records for these 
collections are available. Recently, the same 
species was collected from Lake Annean, 40 
km S of Meekatharra, wt W-A., at 21.3% 
salinity (see Geddes ef al, in press), The 
Western Australian specimens had a much 
thinner shell than those from Lake Eyce. 


122 P. DE DECKKER 


as 


a 


x 
2 
7 
ay 
F 
Z 
| 


a 
. <a 


eat HO 
ee 
s 


- 


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-l, p: Lake Annean, 40 km S of Meekatharra, W.A. 
Scale: 200z. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 123 


OS 
ae 
* ANAS 


a 
N 


Fig. 20. Reticypris kurdimurka n.sp. a antenna, b antennula, c maxillula—palp and lobes, d_ thora- 
copoda I, e maxilla—endopodite, male, f maxilla—endopodite, male, g maxilla, female, h mandible 
—palp, i thoracopoda II, j furcal attachment, k hemipenis, | hemipenis, m furca, n Zenker organ. 
a-b, d-f, h-n: holotype adult male; c, g: paratype female. Scale: 100s, 


Retieypris walbu De Deckker, 1979 
1979 Retieypris walhu De Dockker, p. 162. 


Diagnosis: Rerievpris with lateral lobe of hemi- 
penis banana-shaped and inners lobe ecudgel- 
shaped. 

Description; See De Deckker (1979) p. 1624, 
Diseussian: R. walhu was originally described 
rom samples collected in mound springs be- 
tween Slrangways and Curdimurka, near Lake 
Eyre South in South Australia. Since then it 
has. been recognized from a collection made by 
I. A. BE. Bayly and W, D, Williams in Lake 
Buchanan, 8.W. of Charters Tower in Queens- 
land, in January 1965. Water salinity was 
87.64, (Bayly & Williams 1973). Te has not 
been found in subsequent colleetians made at 
the sume luke by B. V, Tinims. Similarly, it 
is Surprising thal R, walbu has not been col- 
lected in Lake Byce instead af &, kurdimurka. 
Such patchy distribution remains unexplained, 


Famiry: ILYOCYPRIDIDAR Kaulmann, 
1900 


Hlyocypris Brady & Norman, 1889 
Type specie; lUyocypris gibba (Ramdohr, 
1808). 

Mveey pris dusttaltensiy Sars, 1889 
[889 flvacypriy australicnsis Sars, p. 46, 

FIG. 21 


Diagnosis: Mvecypris with club-shaped inner 
lobe on hemipenis extending io as much as 
0.66 of Jength of trapezoid lateral lobe. 


Remarks; The original description of J. ers 
traliensis by Sars (18893) is sufficient, and 
does not warrant. additional illustrations of 
the anatomy here, The ornamentation and out- 
line of the valves of J, gustraliensis, however, 
are extremely variable. Typical vartations are 
illustrated in Fiy. 21 (all SEM photos are at 
the same scale and only adults are repre- 
sented), The surtace of the shell can be finely 
(Fig, Z11) to coarsely pitted (Figs 21c,f,o); 
acicular but stnall spines oeeur, mostly pos- 
terodorsally (Figs. 21h,i) when present, but 
can also cover the entire carapace (Fig, 210), 
The outline of the shell is also variable 
antenorly and dorsally. In dorsal view, 
anteriorly the shell can be pointed (Figs 21j,0) 
or with a squarish blunt end (Figs 21mm). 
The latter feature is often accompanied by a 
depression bordering the periphery of the valve 
anteriorly and posteriorly (Figs 2)a,d), De- 


124 PrP. DE DECKKER 


pressions arc also variable dorsally, a circular 
one above the central muscle field and the 
olher vertical above and in tront of the central 
muscle field, 


The ecologicul significance of all these 
variations are not yet understood but this is 
not a surprising phenomenon since J, autytra~ 
liensis is commonly found in temporary pools 
which ure renowned as the sites of broad 
variations in physical parameters (viz. tem. 
perature, pH, O.), #, quvtraliensiy can swim 
but is usually found crawling in or above 
muddy substrates im ponds. The species usually 
occurs in ponds which retain water for a few 
months and which have a very soft substrate. 
f, australiensis bas also been collected in a 
number of shallow and slightly saline lakes 
in western Victoria, These lakes are charac- 
terized by small fluetuations in salinity over 
the year, This explains the absence of 1 aus- 
traliensix in the saline lakes near the Coorong 
Lagoon studied by De Deckker & Geddes 
(1980) where salinity of some lakes was close 
to fresh im winter, but rose rapidly during 
spring and summer months. In western Vic- 
toria, the range ol salinity tolerance of 1. 
itisiraliensiy 13 usually 4-7%, with ane record 
at 10.37%, in Lake Kariah. So far, J. austfu- 
liensiy has never heen collected m deep lakes 
(fresh or slightly saline) nor in permanent 
(resh takes. 


tT. ausiraliensis bas been collected all across 
Australia, It is also recorded from North 
Africa (Gauthiee 1928). Asia und southern 
Europe (Hartmann 1964, p. 148). 


As the morphology of the bemipenis of /. 
ausiralienyis and J, decipiens Masi, 1906, as 
illustrated in Petkovski (1958), is almost 
identical, it is suggested here that these two 
species could be synonymous, Further work is 
necessary lo confirm this hypothesis, but it is 
important to be ywart of the variation in shell 
ornamentation of the thocyprivy species as 
llusirated above, and by Diebel and 
Pietrzeniuk (1975) for #£ fradyi Sars, 1890, 
since many dlvacypirly species sre separated on 
shell morphology alone, The selection of parti- 
cular features of the shell as taxonomically 
important by Van Harten (1979) will not 
prove to be useful for J, aystraliensis speci- 
mens since for example, the “marginal ripplets" 
of van Harten are Vaniable in Australian speci- 
tens, and even oceasioually absent, Further, 
distinction of species on features other than 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 125 


Fig. 21. Ilyocypris 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.; 0: Pond very close to Reel 
Inlet (coastside) 19 km S of Mandurah, W.A. Scale: 200. 


126 


the hernipenis is likely lo be unreliable smece 
the morphology of many appendages of Slya- 
cypris species are known to Vary, For example, 
Gaulttner (1928) discussed the variations in 
the anatomy of /. australiensiy and tf, biplicata 
(Koch, 1838) from North Africa am! con- 
cluded that he was incapable of separating 
the species, yithough nether was he prepared 
to synonymize them, 


Thyocypris perigundi i.sp. 
FIGS 22-23 


Diagnosis: Hyoeyeris with carapace trapezoid; 
anterodorsal area al hinge extremity com- 
pressed und forming a hump when viewed 
laterally; three large protuberances in dorsal 
area, posterior one being largest, pustulose 
and ending with a number of wart-like tuber- 
cles; Ventrum concave and at 0.6 from an- 
terior, Club-shaped inner lobe of hemipenis 
reaching Up of trapezoid lateral lobe, 
Doyeriptions Carapace.. (External) ‘Trapezoid 
with greatest height 0.2 from anterior where 
hinge starts; dorstim straight except at point 
of greatest height where both valves form 
flattened hump; ventrum concave at 0.6 from 
anterior. Surface of shell highly ornamented: 
three protuberances dorsally with posterior one 
largest and coding with broad wart-like tuber- 
cles from which sela protrudes; below posterior 
protuberance and above veotrum, is an addi- 
tional highly ornamented protuberance: this is 
point of greatest width of shell; deep depres- 
sion worsally between middle and posterior 
protuberances; shell covered with broad 
paeudepunectae which are pustulose inside; 
pointed tubercles ulong margin anteriorly and 
posteriorly. Dorsally, shell like flattened ellip- 
soil with auterior compressed and pointed 
for width of joner lamella but tapering 
pasterorly, 


(Internal) Joner lamellae twice as Wide ip 
anteriar campared (o posterior in both Valves: 
facto selyage al 0.66 from ouler margin 
anteriorly and peripheral to inner margin 
posteriorly; inarwinal cipplem broud ip left 
valve posteroventrally, 


Anam: Antemmula: (Fig. 23¢) 6-segmenteds 
length’ width ratio of fast five seenicnts; 5/3. 
12eh, 13/1, 2/9, 4 1; natatory seta as Jone 
as all segments together, 

Antenna: (Fig, 23a) Natatory setae twice 
length of last two segments and claws together: 


FP. DE DECKKER 


three long distal claws on penultimate segment 
plus another as long as other three on last 
segment with a narrower and shorier one, 
Mandible: (Pig. 23d) Length/width ratio of 
Jasi twa seyments of mandibular palp: 4/3, 
tet. 

Maxilfula. (Fig. 23b) Distal segment of palp 
trapezoid and ending with three broad long 
setae plus two shorter ones; vo Zahnborsten 
on 3rd tobe but setae short and stout on all 
three lobes, 

Maaxilla: Sexually dimorphic: in male (Figs. 
23e.h) palps two or three-jointed with one 
seta at the distal end of the Ist segment; in 
female (Fig. 23g) non-segmented, short and 
harrow palp with three unequal setae, Por 
chactotaxy see Figs 23g,.h) epiped plate with 
five Strahlen, 

Thoracopoda 1; (Fig. 231) Last segment un- 
divided; all setae short. 

Thoracopoda [fy (Fig. 231) 4-segmented with 
three distal setae, two long cqual ones and 3rd 
{bout 0.66 length of others. 


Hemipenis: (Figs 23j,k) Outer lobe trapezoid 
with inner distal end pointed which is nearly 
reaehed by long elub-shaped inner lobe; outer 
Jobe thangilur with distal end rounded and 
slighuly pinehed laterally; inner lobe 0.33 
length of others, 

Zenker orgnn: (Fig. 231) Both ends of globu- 
lar und with 13 rosettes, 

Fijteas (Fig, 23m) Two long equal claws: 
posterior setae slightly longer than anterior 
and placed at 0.4 of length of shaft from 
posterior claw, 

Colour of shell: Transparent white, 

Size: 


Holotype adull mile 
t 


' i I i 
LV Stl VOSu RV 560u 35 5u 
purarype wadu't Temule 
L. H 1 H 
LV 380m Tbe RV 580u TS 


Type locality: Wurrawenia Lake, south of 
Menindie, N.S.W, (33°29'30°S, 141°44°30"E) 
Derivation af name: From the aboriginal lan 
fuage perivandi meaning meandering lake or 
lazon. as this species bas been found in a 
lake part of am old river systern (Warrawenia 
1) and ina ligdon (Katarapka b.). 

Ecology and distribution: This species has only 
been collected in three localities: Warrawenia 
L, Katarapko Lagoon, which is a natural billa- 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 127 


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—2004 for a-h; 2—SO for i-k. 


128 P. DE DECKKER 


Fig. 23. Ilyocypris perigundi n.sp. a antenna. b maxillula—palp and lobes. c antennula. d mandible. 
e maxilla—endopodite, male. f thoracopoda I. g maxilla, female. h maxilla, male. i thoracopoda IL. 
j hemipenis. k hemipenis, | Zenker organ. m furea, a-f, h-m: holotype adult male. g: paratype adult 
female. Scale: 100«. 


OSTRACODS FRGM AUSTRALIAN INLAND WATERS 128 


bong converted into un evaporative basin near 
ihe River Murray at Loxton, 8.A., and froin 
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 nol been measured, At 
Scott Creck, water was Sresh, 


Remarks, (, perigunedi differs from J. austra- 
lignsiy on the following features; the former 
is smaller and has a very different shell archi- 
tectuine characterized by the three highly orna- 
mented dorsal protuberances, a ventrum placed 
at 0.66 from the anterior of the shell and a 
fiat dorsal hump in the extremity of the hinge 
anteriorly. The inner lobe of the hemipenis in 
1, perigundi is almost as long as the lateral lobe 
Whereas iL is only 0.66 the length in J. ausztra- 
liensix, 


SUPERFAMILY: CYTHERACEA, Baird, 1850 
Famicy: LEPTOCYTHERIDAE Hanai, 1957 
Lepracythere Sats, 1925 


Type species; Leplocythere pellweida (Baird 
1RSO) 


Lepiocythere lacustris n.sp. 
FIGS 24-25 
1919 Cythere Inbbockiana Brady’ Chapman. p. 
29, 

Diagnosiv: Leptacythere with deeply pitted 
external surface of shell, large circular depres- 
sion pasteroyventrally outside and where inner 
lamella is broadest on inside: posterodorsally 
near termination of hinge, shell is slightly con- 
cuve and posterior to it: shell thick and smooth 
especially in left valve. Outline of hemipenis 
as in Figs 25),k. 

Deseviption: Carapace. (External) Subrectan- 
cular shell, coarsely pitted and with at Jeast 
two clongated grooves; anterior straight and 
forms an obtuse anele with hinge Ine and 
other, at posterior, runs alovist parallel to cure 
yature of shell; a deeper and broader notch 
often visible in posteroventral area in bath 
valves; ornamentation of shell varying from 
fine to coarse ribbing; dorsum almost straight 
and inclined except in pastero-dorsal area, just 
before termination of hinge where it is slightly 
coneave; behind this, shell thick abd smooth, 
especially in left valve; greatest height at about 
0.25 from aaterior; mouth region concaye und 
ut 0.4 from anterior, In dorsal] view, shell eom- 


pressed and hus almost straight sides except 
where grooves occur anteriorly and posteriorly; 
hinge area almost smuoth externally. 
Qnternal) Inper lamella broad and widest in 
posteroventral area opposite external deep 
notch; selvage faint and peripheral in both 
valves; hinge crenulated all along with broad 
tooth at bath ends in right valve and matching 
sockets in left one; behind anterior socket in 
left valve, are two smaller teeth and ip front 
of the posterior socket, with a matching de- 
pression in right valve. is also a Small tooth; 
central muscle field wilh vertical row ot four 
scars! two jn middle are clongated and parallel 
to hinge line whereas others are almost cir- 
cular, 

Anatoiny: Antennula; (Fig, 25a) 6-segmented: 
length/ width ratio of last four segments: 
1.6/1, 1/14, 1/1 to 14/1, 4/15 distal thick 
Sela 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 coxale 
with seven (teeth and long acicular seta on inner 
side near base of smaller tooth; epipad with 
ferminal segment small and square-shaped. 
Musillula: (Big. 25c) Epipod with 14 long 
and short Strahlen: terminal segment of palp 
small, rectangular and with one distal seta 
jointed. 

Maxilla: (Figs 25h,i) Distal claw short and 
slightly curved: in female inner seta on 20d 
segment pilose in its distal half (Pig. 251) 
whereas. barren in male (Fig. 25h). 
Thoracopoda I: (Fig. 25f) Slightly larger than 
maxilla but with only one distal seta of Ist 
sezment. 

Thoracopoda UW: (Fig. 25g) Similar but 
élightly larger than thoracopoda fF and with 
distal claw more slender. 

Hemipenis: See outline in Figs 25),k, 
Genitalia: See oufline in Fit, 251m. 

Fureat (Figs 25f.m) One long seta near hemi- 
penis and genital organ. 

End of body: (Fig. 251) With one small seta 
and densely pilose 


Colour ef shell: Light brown. 
Size: 
holotype adult mule 


L H L H 
LY 500 2701 RV 490i 27 


130 


P, DE DECKKER 


‘ 
© 
+ 
7 
a 
a 
hs 


ec’ 


* 


Fig. 24, Leptocythere lacustris n.sp. a RV internal, male holotype. b LV internal, male holotype, 
¢ C showing LV, female paratype. d LV external, male paratype e LV external female paratype. 
f C external showing RY, 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 internal, 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—200u for a-h, 


m; 2—SO« for i-l, 0;—25m for n.p. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 131 


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—coxale. f thoracopoda 1. g thoracopoda II, h 
maxilla, male. i maxilla, female. j hemipenis. k hemipenis. 1] end of body. m genitalia. a-e, g-h, j-k: 
holotype adult male; f, i, m: paratype adult female; 1: paratype adult male. Scale: 1004, 


132 P. DE DECKKER 


pirilype adult female 
I, H L af 
LV 485u 27 RV 480y 27s 
Type locality; Fresh Dip Lake neac Robe, $.A- 
(35°15'42"S, 139°48'42"R), 


Derivation ef names From Latin Jaeusirinus 
( ~ of Takes) as this species, which belongs to 
a typically estuarine genus, is found in Takes. 
Ecolagy and distribution: This teuly benihie 
speeies has heen collected in only four locali- 
tics Near Robe, S.A, apart from the type 
locality, All lovalitie; are characterized by 
permanent water only slight salinity fMuctus- 
tions. The salinities were 19 to 28%), In addi- 
lion, at the type Juculity, salinity was 2.84%), 
So far, £. fveusiriy has been found in perma- 
nent Water around 35%, ay no such lakes have 
yet been sampled in the search for this ostra- 
cod, Attempts to fild it tn the permanent and 
suline lakes Keilambete and Gnotuk jn western 
Victoria (salinity ea. 55-62';,.) were unsue- 
cesslul, suggesting that the salinity range of 
L, lacustriy does not reach such values, This 
species, as for all others in the typical 
estuarine genus Lepocvihere, is indicative of 
permanent water as tl docs mot produce egps 
which can withstand desicvation. 


Remarks! The deseription of ZL, laeustriv 
corresponds to the specimen identified by 
Chapman (1919) ps Cyrhere lubbockiana ftom 
the fossil site at Boneo Swamp, The two 
rounded tubereles on the posteroventral area 
al the shell us illustrated by Chapman (1919) 
on Plate TV.9 ure two sand grains whieh have 
heen removed by me from the specimen in the 
Nationul Museums of Victoria, The shell archi- 
tectiire of L, feenyiris Varies from almosi 
smooth to coursely reticulated (Figs 241.) 
at times, reticulation js so thick that the shell 
has a smooth appearance. Such variations have 
heen mentioned for other estuarine and marine 
species of Leptoeythere by Shornikeyv (1966) 
and Hartmann & Kuhl (1978). This is not 
suprising as environmental conditions. such 
as water salinity in salt lakes, can Ihieluate 
over te, 


LL. laeustety is closely related to L. harimanni 
(MeRenzie 1967), Unfortunately, no males 
of the Jatter species have ever been found, us 
analysis of the hemipenis morphology would 
have canlicmed this distinction. The valves 
of £. hartinanui, however, are not coursely 
put! and reticulation is more sparse (see 
Hartmann 197% Pl Wf; Bigs 3-8). The 


anterior and posterior grooves are present in 
both species, The posteroventral notch is ouch 
deeper in L. uensiriy and the smooth postero- 
dorsal hump is absent in L. arta, 


Pamiby: LIMNOCYTHERIDAE Sars, 1925 


SusFAMiLy! LIMNOCYTHERINAE Sars, 
1925 


Gomphodella n.gen, 

Type species: Gomphodella maia hap., gender 
masetiline, 
Diagnosis: Carapace. Female carapace in 
dorsal view triangular with greatest width at 
aboul U.66 to 0.75 from anterior; ut about 
0.4 from anterior, where central musele field 
oveurs, vulves are slightly compressed; absence 
of promiment Jateral ridge in ventra) area; 
sieve pore canils and few broader normal pore 
canals in row parallel ty posterior edge of shell 
in both valves, Male, in dorsal view, like a 
flattened oval with both ends pointed, 
Anatomy: Female maxilla and thoracapodae J 
and TF similar, although of different lengthy 
female furea with three stout, pointed and 
thickly pilose setae. 
Pevivation uf mite: From a combination of 
the two names Gontphoevthere and Cytheri- 
della as this genus shares diagnostic features 
of the two geuere. 
Renvirks: Goraphodella js very similar toa Cy- 
theridella Daday, 1905 except ihat the shell 
ol the former iy less compressed in the area 
Where the central muscle field occurs, and as 
the maxilla and thoracopoday 1 and I ure 
similur in Gamphetella (in Cvtheridella the 
tharavopoda ID is traystormed into a prehen- 
sile palp), The diagnostic feature which Gem 
phodella and Gomphoevitere Sars, 1924, share 
is Ihe presenee of three stout and thiekly pilose 
setae wn the female fureimthis is pot seen in 
Cyrheridella. Goniphedella lacks the toteral 
ventral ridge on both valves on the oulside of 
the shell se typical of Gomphoeythere species, 

Gomiphodella is 1a be included in the family 
Limnoecylherinae Sars, 1925. following the 
recent regrouping of subfansilies in the Linno- 
cytheridae by Colin & Daniclopol (1978). 


Gomphodella maia n sp, 
FIGS 26-27 
Diugnosixy Surface of shell pyeudopunctate 
with many Sieve pores: posterior urea of shell 
broadly rounded in lateral view: in dorsal view, 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 133 


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 anterodorsal 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 anteriorly; 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; Antennula: (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 Ist segment 
smooth, and distal one on 2nd segment finely 
pectinated. 

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 27),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. 271), two barren 
setae. 

Colour of shell: Grey to grey brown. 

Size: 

holotype adult male 


L H L H 
LV 490u 290u RV 4604 3002 
paratype adult female 
L H L H 
LV 560u 330u RV 520” 320u 


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, see De Deckker 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- 


4DE DECKKER, P. (1981) Taxonomy, ecology 
and palaeoecology of ostracods from Australian 
inland waters, Ph.D. Thesis, Dept of Zoology, 
University of Adelaide (unpubl.). 


134 P. DE DECKKER 


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. | RV internal, anterior detail of d. m LV external, detail of e. n LV external, 
detail of g. a-g, I-n: type locality; h-k: fossil, Blue Lake, Mt Gambier, S.A. Scale: 100 for a-k; 
15“ for 1; 5“ for m, n. 


OSTRACODS FROM AUSTRALIAN INLAND WATERS 135 


Fig 27. Gomphodella maia n.gen., n.sp. a antennula. b antenna c maxillula—palp and lobes. d man- 
dible—coxale. e mandible—palp. f maxilla. g thoracopoda I. h thoracopoda II. i hemipenis and 
furca. j genital organ and furca. k genital organ and furca. a, c, ¢, j-k: paratype adult female. 
b, d, f-i: holotype adult male. Scale: 100, 


136 P. DE DECKKER 


lected alive once in Fresh Dip Lake {= type 
locality) at 2.3%, sahnily. As a fossil, it has 
been foand with G. ausrralicu, and other fresh- 
water ostracods in samples from a short core 
from the Blue Lake al Mt Gambier, S.A. It js 
also found in twa fossil sites in northwestern 
Tasmania (De Deckker'), In samples trom 
both sites, a large number of carapaces were 
recovered and same of these were found with 
juveniles inside them. The significance of this 
phenomenon is wet properly understond as, 
usually, after death under water, valves of 
osiracods separate prior to or during decay of 
the soft parts, When ponds dry up, ostracods 
are seen to close their valves tightly and, if 
this period is long enough to ¢auise dehydration 
of the ostracods, death would oceur, Rupid 
sediment accumulation is later necessary dur- 
ing a wet phase 10 prevent carapaces from 
openiig. This process is likely to be one 
possible explanation for the high percentage of 
G, mafe carapaces found in samples from the 
two Tasmanian sites. Death caused by changes 
in water chemistry would not prevent cara- 
paces [rom dislocation, Another possibility is 
that G, maia can in fact burrow in sediment— 
4 phenomenon noticed for a closely related 
ostracod Gomphocybthere sp. (with another 
ostracod Darwinula sphagna Barclay, 1968) 
found in the interstitial waters of the Rotorua 
lakes in New Zealand by Chapman & Lewis 
(1976)—and, if it was to remain there until 
death, valyes could fot become easily dis- 
sociated. 


Gomphodella qustralica (Hussainy, 1969) 
1969 Gomphovythere australica Wussainy, p, 294 


Diagnosis: Subrectangular shell with both 
dorsum and ventrum almost straight; anterior 
broadly rounded and posteroventral 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, In 
dorsal view shell in female like an upside down 
hearl and male much narrower with beth ends 
pointed, 


Size range: 


L H 
adult male carapace BUd-YUU) 400-3 00% 
adult female caripace 650-75du 27-3 2 


Ecology; G, australica is best known fram its 
type locality, Lake Purrumbete in Victoria. 
This is 4 permanent lake with salinity of 0.42— 


0.50%) over the 1969-1972 period (Timms 
1976) and 037-0445, over 1979-80, Tk is 
enly recorded in that Jake (greatest depth > 
40m) at O.5-l m by Timms (1973) Tt is 
usually found crawling in among filamentous 
algae like G. maia. In South Australia it is 
also recorded from Scott Creek near Adelaide, 
and from ihe permanent Fresh Dip Lake near 
Robe, G, ausiralica is considered to be a fresh- 
water species which can withstand a slight 
amount of dissolved solids in water, wrth ils 
highest salinity recorded al Fresh Dip Lake at 
2.3%. Brooding oceurs in this species ag ie- 
monstrated by Hussainy (1969), Fossil speci- 
mens. wilh coarse shell reticulation have been 
recovered wiih other fresh water ostracods 
from a short core taken fram the fresh Blue 
Lake al Mt Gambier, $.A. 


Remarks’ The wansfer of this species ta Gom- 
phodella gen. nov. is Necessary because it docs 
nol possess the peripheral lateral ridge around 
the fat base of the shell of each valve so 
typical ot Gomphacythere Sars, 1924. This 
species is easily distinguishable from G. maia 
by its larver size, reticulated shell and the very 
pointed posterior area of the shell which is alsa 
steeply inclined posteroyentrally, 


Acknowledgements 

This paper was written during the tenure 
of # Commonwealth Postgraduate Research 
Award under the supervision of Professor 
W. D. Williams in the Zoology Department, 
University of Adelaide, 

Many of the specimens described here were 
received from the following: Mr P, Bailey, 
Des IT, A. EF, Bayly, R. Marchant, R. Tait and 
BK. V. Timms and Me W, Zeidler. 

IT am grateful to the follawme for the 
loan af type specimens: Dr M, Christiansen 
(Oslo Museum), De G. Bowshall and Ms A. 
Girney (Brilish Museum, Natural History), 
Drs I, A. EO Bayly. Ro Murchant, R. Tait and 
B. V_ Timms and Mr W, Zeidler, 

Museum), 

1 would like to thank Ms S. Lawson and C. 
Twang for typing the manuscript and Mr P. 
G. Kempster for help in the photography. 


“TIMMS, B, V. (1973) A comparalive study of 
the Iimaclogy of three masr lakes im western 
Victoria, PhD ‘Thesis, Monash University 
(unpubl.y. 


OSTRACODS FROM AUSTRALIAN INLANID WATERS 137 


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138 P. DE DECKKER 


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Beograd. 


THREE NEW SPECIES OF CALOSTAURUS (CESTODA: DAVAINEIDAE) 
FROM THE NEW GUINEA WALLABY DORCOPIS VETERUM 


BY IAN BEVERIDGE 


Summary 


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. 


THREE NEW SPECIES OF CALOSTAURUS (CESTODA: DAVAINEIDAE) 
FROM THE NEW GUINEA WALLABY DORCOPSIS VETERUM 


by Tan BEVERIDGE 


Summary 


Bevenwen, IT. (1981) Three new species of Calostaurus (Cestoda: Davaineidae) from 
the New Guinea wallaby Dorcaopsis vetvrum, Trans. R. Soc. S. Aust. 105(3), 139-147, Tl 


December, JY8\. 


Three new specics of Culastanrus Sandurs (Cestoda: Davainewue) are described from the 
small intestine of the racropodid marsupial Dorcapsts veteran from New Guinea, ©. 
dorewpsiy sp. noy. differs from ©, maeropus in the size and arrangement of rostellay spines, 
size of yostellar hooks and cirriis sac. C. owen’ sp, nev, and C. paras 5p. Noy, resemble C. 
mundayi in features of the rastellum, but difler in size, number of testes per proglottis and 


number of eggs per egg capsule, 


Introduction 

Three species of the davaineid cestode 
genus Calostaurus Sandars, 1957 are known to 
occur in Australasian macropodid marsupials, 
namely C, macropuy (Ortlepp, 1922) originally 
described from Tiylogale bruni? (Shreber, 
1778) (Syn. Macropus brunii) and redes- 
cribed by Sandars (1957) from Thylogale 
siigmatica Gould. 1860 (syn. T. wilcoxt) in 
Queensland, and C, thylogale Beveridge, 1975 
and C. mundayt from Thylegale billardierti 
(Desmarest. 1822) and Parordus tridactylis 
(Kerr, 1972) (syn. P. uipicaliy) respectively, 
both occurring in Tasmania (Beveridge, 1975). 
In cach instance, a single host species is para- 
sitised by one cestode species only. By contrast 
in collections of cestode, made recently from 
the wallaby Dorcopsis veteruen (Lesson, 1827) 
in New Guinea by Dr L. L. Owen, three cestode 
species may occur in the same host individual, 
The three species are described in this paper. 


Methods 

Cestodes were stained with Celestine blue, 
dehydrated and mounted in balsam. Serial 
sections of cach species were cut but the state 
of preservation of the specimens did not allow 
detailed histological examination. All measure- 
ments are given in mm as the range followed 
by the mean of 5 measurements m parentheses. 


Calostaurus dorcepsis sp. nov 
FIGS [-10 


Description based on fragments of several 
ypecimens; no complete cestodes presen 
dorsal system, ramifying across proglottis. In 


* Institute of Medical & Veterinary Science, Prome 
Road, Adelaide, S$. Aust SO(M). 


Moderate sized cestodes. Maximum length of 
fragments 105; maximum width 4, Largest 
fragment contains 350 proglottides, Scolex 
large, 0,90-1.16 (1,00) in diameter, quadran- 
gular in en face wew, with eversible rostellum 
042-049 (0,52) m diameter, Rostellum 
armed with approximately 1600 hammer 
shaped hooks in two rows, arranged in shape 
af Maltese cross, with axes extendiig between 
pairs of suckers; hooks 0.012—0.014 (0.013) 
long. Base of rosteilum armed with approxi- 
mately 16 cransverse 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 crass; 
never lic between arms of cross. Rostellum 
varies considerably in shape (Figs 1, 2) de- 
pending upon degree of extension, Rostellar 
spines form narrow band anterior to rostellum 
at opening ta rostellar sac when Jatter fully 
retracted, Suckers 0.27-0.32 (0,29) in dia- 
meter; margins armed with seven to nine rows 
af tiny awl-shaped spines 0,006-0.008 (0,007) 
long. Neck 0.38-0.75 (0.52) long. 
Proglottides craspedote; velum overhanging 
adjacent proglollis, Very narrow. Mature pro- 
glouides greatly extended transversely, 0.20— 
0.45 (0.36) * 24-40 (2.9), ratio width: 
length 1:5-|°20, Genital pores single, uni- 
lateral, Genital ducts pass between osmoregu- 
latary canals. Dorsal osmoregulatory canal ex- 
rremely narrow, 0,01-0.03 (0.02) in diameter, 
jies well internal to ventral canal: ventral 
osmorceulatery canal 0.05-0.09 (0.07) in 
diameter, ‘Transverse osmoregulatory canals 
vonneet left and right dorsal and ventral canals 
at posterior margin of each proglottis. Com- 
plex, branching network of canals arises from 


140 IAN BEVERIDGE 


Figs 1-10. Calostaurus dorcopsis sp. noy. from the wallaby Dorcopsiy vererum. 1. Scolex with partially 
everted rostellum. 2. Scolex with fully evert rostellum 3. Scolex with retracted rostellum, en face 
view. 4. Female genitalia, dorsal yiew. 5. Rostellar hooks. 6. Rostellar spines, 7, Sucker spines. 8. 
Cirrus sac and distal vagina. 9. Single mature proglottis, 10, Gravid proglottis, Scale lines: Figs 5-7, 
0.01 mm; Figs 1-4, 8-10, 0.1 mm. 


NEW SPECIES OF CALOSTAURUS (CESTODA) 14) 


largest fragment, genital anlage appears in ap- 
proximately 50th proglottis, Male and female 
genitalia mature in proglottides 200 and 260 
respectively, and involute in proglottides 425 
and 340 respectively. First gravid proglottis 
345th. 

Genital atrium small, situated in anterior 
half of lateral proglottix margin, Cirrus sac 
small, 0.20-0.25 (0.22) & 6.07, elongate, 
muscular walls feeble, cirrus sac not reaching 
ventral osmorggulatory canals. Distal region of 
cirrus 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 efferentia not seen, Testes in- 
variably distributed in twa lateral groups, ex« 
tending from ventral osmoregulatary carals, 
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 
proglottis; always more testes in aporal field; 
8-23 (15) testes in poral ficld, 21-39 (31) 
in aporal fleld. Some differences evident in 
testis number between strobilae; in one 
strobila 8-13 (10) poral, 34-39 (37) aporal: 
in second strohila 16-23 (20) poral, 22-28 
(24) aporal. 

Vagina opens to genital atrium posterior fo 
cirrus sac. Distal region, 0.08-0,13 (0.10) x 
0,02, dilated, armed with fine bristles. Mid- 
region narrow, unegiled, leads medially, 
posterior to vas deferens, terminates in fusi- 
form seminal receptacle 0.12-0.18 (0.15) > 
0.03.-0,07 (0.05) in size, lying dorsal to poral 
lobe of ovary; sperm duct passes posteriorly 
fram seminal receptacle, lined with bristles, 
Ovary bilohed, situated to poral side of pro- 
glottis midline, 0.18-0.20 (0.18) * 0,26-0,38 
(0.32) in size, with 8-10 clavate lobules in 
cach Inbe. Vitellarium ovoid or beaneshaped, 
0.07-0,09 (0.08) & 0.12-0.16 (0.15) in size, 
posterior and dorsal to ovary. Mehlis’ gland 
spherical, anterior to vitellarium, Uterine duct 
short, passes anteriorly from Mehlis” gland, 
terminates dorsal to ovary. Uterus absent. Rggs 
dischareed from uterine duct directly iota 
parenchyma, become surrounded hy capsule, 
with ane eee per egg capsule, Size of ese 
capsule 0,05-0.06 (0.055): exe 0.01-0,02 
(0,015), Giravid preglottides extended trans: 
versely 0.21-0,.70 (0.49) % 22-3. (3.1), 
widthilength ratio £:3-1;7 Terminal pro- 


glottides relatively longer, occasionally as long 
as wide O0.9O-1,10 (1.03) *% 105-55 (131), 


Host; Dorcupsis veterium (Lesson, |827) 
(Marsupiilia: Macropodidac). Lecation: Small 
intestine, Type Lecaltys Veikabu Creek, Cen- 
tral Province, 9° LO'S, 147709 R, Papua New 
Guinea. Types: In British Museum (Natural 
History), Holotype 1981.6.17.1, paratypes, 
1981.6.17.2-LO. Eivmology: The specific name 
is derived from that of the host, 


Culostaurus owenl sp. nov, 
FIGS 11-19 


Description based on 10 complete mounted 
specimens. Small cestodes, 24-38 (33) longi 
maximum width | 2-1.4 (1.3); sirobilae con. 
tain 100-147 (120) proglortides. Scolex rela- 
tively large, 0.62-1.05 (0.99) in diameter, 
quadrangulac in en Jace view, with Tostellum 
up to 0.45 in diameter when everted. Rostellim 
urmed with approximately {000 hammer 
shaped hooks in two rows, arranged in form 
of oper cross, with axes extending between 
pairs of suckers; rostellar cross 0,25-0.32 
(0.29) * 6.18-0,34 (0,26); restellar hooks 
0.007-0.010 (0.008) Jong, Base of rostellum 
urmed with concentric rows of tiny aw) shaped 
spines approximately 0.004 long. Spines form 
band posterior to rostellar cross on everted 
rostellum, never Jie between arms of crass; 
spines form band anterior to rostellar cross, 
at opening of rostellar sae when rostellum 
fully retracted. Suckers 0.16-0.23 (0,21) in 
diameter; margins urmed with oboul eiaht 
rows of tiny, rose thorn shaped spines 0.004— 
0,007 (0,006) long, Neck 0.21-0.34 (0.28) 
long, 


Proglottides craspedote; velum overhanging 
adjacent proglottis very narrow, Mature pro- 
glottides wider than long, 0,27-0,33 (0,31) * 
120-1.53 (1.25), ratio lenglh:width 1:3.5- 
1:4.3. Genital pores single, unilateral; one 
proglotus found with reversed orientation, 
Genital ducts pass betweetr asmoregulatory 
canals, Dorsal osmoregulatory canal extremely 
narrow, 0.005 in diameter, sinuous, lies in- 
ternal to ventral canal; veotral esmoregulatery 
canal straight, 0020.04 (0.03) in diameter. 
Transverse osmoregulatory canals connect left 
and fight dorsal and veotral canals al posterior 
margity of cach proglottis. Origins of dorsal 
transverse canals sometimes varivble, single 
transverse canal oceasionally arisitg from two 
or more points on dorsal longitudinal canal. 


142 IAN BEVERIDGE 


Figs 11-19. Calostaurus oweni sp. nov. from the Wallaby Dorcopsis veterum. 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. 


NEW SPECIES OF CALOSTAURUS [CESTODA) 


Genital anlage appears in proglottides 25-40 
(31), Male and female genitalia mature in 
proglottides 52-73 (64) and 70-98 (RI). re- 
spectively, and invelute in proglottides 85-112 
(97) and 75-102 (91) respectively, First 
gravid proglottis 89-120 (104), 

Genital atrium very small, situated in middle 
of lateral proglottis margin, Cirrus sae large, 
O.160.18 (0.17) % 0.09011 (0.10), avoid, 
muscular walls feeble, cirrus sac not reaching 
longitudinal osmoregulatory canals, Distal 
region of citrus of greater internal diameter, 
lined with bristles; remainder narrow, coiled 
muscle bands run from cirrms to internal sur- 
face of cirrus sac, Trternal and external 
seminal vesicles absent. Vas deferens greatly 
coiled, passes medially towards centre of pro- 
glatlis, gradually diminishing in diameter, ter- 
minates dorsal to ovarian isthmus by dividing 
into (Wo Vasa efferentia cach supplying one 
group of testes. Testes almost invariably dis- 
iributed in two lateral groups, in posterior balf 
of proglottis, extending medially from dorsal 
asmoregulatory canals. In very occasional pro- 
vlottides, one or tWo testes present posterior 
to Vilellarium = joining two Jateral — testis 
eroups. Testes ovelie ovarian labes and oc- 
casionally vitellarium; testes 0.045-0.055 
(0.050) in diameter. Testes number 15-19 
(16) per praglottis; always more testes itt 
aporal field: 3-7 (5) poral, 10-14 (12) 
aporal, 

Vagina opens to genital atrium pasteriar to 
cirrus sac. Distal region, 0,05-0.08 (0.07) ™ 
0.02, dilated. Mid-region narrow, sinuous, 
leads medially, posterior to vas deferens, ter- 
minates im fusiform seminal receptacle, 0.07— 
0.11 (0.09)  0.03-0.04 (0,04) im size, lying 
anterior and dorsal ta poral lobe of ovary; 
sperm duct passes posteriorly from seminal re- 
veptachs, dilates, lined with bristles. Ovary 
hilobed, situated to poral side of proglottis 
mid-line, O100.18 (0.13) * O17-0.20 
(0.189) with 4-5 clavate Jobules in each lobe. 
Vitellurium ovoid or bean shaped, lobulate, 
0.05-0.07 (0.06) * 0.07-0.08 (0.07) in size, 
situated posterior and dorsal to ovary. Mehlis' 
gland spherical, anterior to yitellarium. Uterine 
duet passes anteriorly from Mebhlis’ gland ter- 
minated in midline dorsal to ovary, Ulerus 
absent. Eggs discharged from uterine duet 
directly into parenchyma, become surrounded 
hy capsule, with one egp per egg capsule. Size 
of eee capsule 0.04-0.05 (0.047): ega 0.01- 
0,02 (0.15), Gravid proglottides extended 


43 
iransyersely 0.41-0.55 (0.45) “ 0.95-),33 
(1.20), widthslength ratio 1;1,17-1:2.7. 


Terminal proglottides almost as long as wide 
or longer than wide, 0.62-0,75 (0.69) ™ 
0.52-0.88 (0.70). width:length ratia 1:0.74— 
171.3. 


Hosts Dorcopsiy vererum (Lesson, 1827) 
(Marsupialia: Macropodidar)- Location: small 
intestine. Type Localiryy Veikaby Creek, Cen- 
tral Province, Papua New Guinea, Typesr Jo 
British Museum (Natural History), holotype 
1981.6.17.01. paratypes 1981,6,17.12-21, 

Etymology. this species is named alter Dr 
t. L. Owen who collected all the material 
described here. 


Calostaurus parvus sp. nov. 
FIGS 20-27 
Deseription based on five complete mounted 


specimens. Small cestades, 9O-11.2 (9,7) 
long; maximum width 0,75-1,20 (0.96); 
strobilae contain 38-57 (42) proglattides. 


Scolex relatively. large, 0.62-1.08 (0.76) in 
diameter, quadrangular in en face view, with 
eversible rostellum, Rosiellum acmed with 
approximately 1000 hammershaped hooks tn 
two rows, arranged in form of open cross, 
with axes extending between pairs of suckers; 
rostellar cross 0.29-0.47 (0,35) & 0.28-0.42 
(0.44); rostellar hooks 0,009-0.010 (0010) 
long. Base of rostellum armed with concentric 
rows of tiny awl-shaped spines, with simple or 
bifid base; spines 0,004-0.005. (0.005) long. 
Spines form band anterior to rostellar cross, 
at opening of rostellar sac, when rostcilum 
fully retracted. Suckers 0,19-0,23 (0,20) 1n 
diameter, margins armed with about 10 rows 
of tiny rose thorn shaped spines 0,002-0.005 
(0.003) long. Neck short. 


Proglottides craspedate; velum overhanging 
adjacent proglottix very narrow, Mature pro- 
gtottides wider than long 0.24-0.35 (0.32) 
0.64-0,98 (0.75), ratio length.width 1°2.0- 
1:3.8. Genital pores single, unilateral) Genital 
ducts pass between longitudinal osmaregula- 
tory canals. Dorsal osmoregulatory canal ex- 
tremely narrow, sinuous. 0.005—0,010 (0.006) 
in diameter, lies immediately internal to ventral 
canal; ventral osmoregulatory canal O.0i5— 
0.020 (0.017) in diameter. Transverse osmo- 
regulatory canals connect left and right ventral 
canals at posterior marein of cach praglottis. 
Genital anlage appears in proglottides 8-28 
(12). Male and female genitalia mature in 


144 IAN BEVERIDGE 
Sa nt, 
CONE MOON 


Maat 


Figs 20-27. Calostaurus parvus sp. nov. from the wallaby Dorcopsis veterum. 20, Scolex with everted 
rostellum, lateral view. 21. Scolex with retracted rostellum, en 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. 


NEW SPECIES OF C4ALOSTAURUS (CESTODA) 


proglottides 14-37 (22) and 20-28 (25) re- 
spectively and involote in proglottides 33-36 
(35) and 30-34 (32) respectively. First 
gravid proglottis 34-38 (36). 

Genital atrium insignificant, situated slightly 
anterior to middle of lateral proglottis margm. 
Cirrus sac small, 0.07-0.10 (0,09) ® 0,04- 
0.05 (0,04), clavate, muscular walls feeble, 
cirrus sac almost reaching — longitudinal 
osmoregulatory canals. Distal region af cirrus 
of preater infernal diameter; armature not seen; 
mid-revion narrow, sinuous; proximal region 
forms small, spherical internal seminal 
vesicle; present in most proglottides of all 
strobilac examined, but not detectable in every 
single proglottis; internal seminal vesicle 
0.020-0.035 (0.025) ™ 0.025-0.030 (0.026). 
External seminal vesicle absent, Was deferens 
narecow, greatly coiled, surrounded by pale 
basophilic cells, passes medially towards centre 
of proglottis. then posteriorly towards ovary; 
terminates dorsal te ovanan isthmus; vase 
efferentia not seen. Testes distributed in 
posterior part of proglottis, in single field, con- 
fluent posterior to vitellarium; testes lie within 
area bounded by lateral osmoregulatory 
canals; usually lie medial to dorsal canals, but 
few testes may overli¢ dorsal canals. Testes 
nol confluent posterior to vitellaria in few 
proglottides. Testes usually overlie ovary and 
vitelarium: testes §.04-0,05 (0,045) in dia- 
meter. Testes number 17-22 (20) per pro- 
glottis; poral testes 6-10 (8), invariably less 
numerous than aporal testes }1—-J4 (13). 

Vagina opens to genital atrium posterior to 
cirrus sac, Distal region, 0.06 % 0,02, dilated, 
Mid-region narrow, sinuous. leads medially, 
posterior ta vas deferens, terminating in large 
cluvale to pycifarm seminal receptacle, 0.08- 
013 (0.09) & 0.04-0.05 (0.05), lying 
anterior and dorsal ta poral lobe of ovary; 
sperm duct elongate, smuous, passes posterjarly 
{rom seminal receptacle, Ovary bilobed, sity- 
ated in proglottis mid-line, 0.09-0,16 (0.14) 
“~ 0.14-0,16 (0.15) in size with 4-6 clavate 
lobules in each lobe. Vitellarium ovoid or hean 
shaped, Inbulate, 0.05-0.08 (0.06) * O,04— 
1.05 (0.05) in size, situated posterior and 
dorsal to ovary. Mehlis’ gland spherical. 
gaterior to vilellacium. Uterine duct passes 
anteriorly from Mebhlis’ gland, terminates in 
midline, anterior to ovary. Uterus absent, 
Fives discharged from uterine duct directly 
into parenchyma, become surrounded hy 
cupsules, with one egg per egg capsule. Size 


145 


of egg capsule 0.05-0.07 (0.06); egg 0,015, 
Gravid proglottides longer than wide 0,65-0.75 
(0.69) ™ O.67-1.10 (0.83), ralio length; 
with O.64:1-1:1.1- Terminal  proglottdes 
0.65 © 0,580.68 (0.73), ratio length:width 
1.2:1-1.4¢1. 

Host: Dercopsiy veterum (Lesson, 1827) 
(Marsapialia:  Macropodidae), Location: 
Srnall intestme. Type locelity: Veikabu Creek, 
Central Province, Papua New Guinea. Types! 
Jn British Museum (Natural History), holo- 
type 1981.6.19.1. paratypes 1981.6.19,2-6. 
Enology: the specific nant parvus derives 
from the fact that it is the smallest known 
species of the genus, 


Discussion 


Three species of Calostaurus have heen des- 
cribed previously, being distinguished prin- 
cipally by the shape of the cross formed by 
the rostellar hooks, C. macrapuy has the hooks 
arranged in the shape of a Maltese cross, 
C. thvlopale in a six-lobed cirele and Cy nuin- 
dayi in a four-lobed cross, OF the species des- 
cribed here, C. dereapsis sp. nov., most closely 
resembles ©. macropys in that the hooks are 
arranged in the form of a Maltese cross 
Whereas C, ewenf sp, nav. and C. parvus sp. 
nov. both have rostellar hooks arranged in the 
form of four-labed crosses similar to that of 
C, mundayi. The histological anatomy of the 
scoleces af the new species is similar to that 
of C. rhylagele and C. mundayi and has there- 
fore not been described in detail, 


Both C. oweni and C. parvus can be dis- 
tinguished from C, mundayi by size alone, 
since C. mucmdayvi measures 32.4 to 45.1 em, 
whereas C. oweni and C. parvus measure 24— 
3.8 and 0.9-1.1 em respectively. In addition 
C. munday? has one to three eggs per egg 
capsule while the other species have a single 
ege in cach capsule, and the range of testis 
number in C. miundayi (25-32) is higher than 
in ©, oweni (15-19) and C. parvus (17-22)- 

C. oweni and C. parvus differ from one 
another in ia number of features. Gravid speci- 
mens of C. parbuy are consistently smaller and 
have fewer progloitides than C, ewert, 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 vitellana in C. parvias, but are 
divided into two groups in C. owen?. Some 
variation in this character does occur and 


NEW SPECIES OF CALOSTAURUS (CESTODA) 


146 


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IAN BEVERIDGE 


occasional proglottides of C, parvuy have the 
testes in two groups, while occasional pro- 
glottides of C, aweni have the testes. confluent 
posterior to the vitellaria, but in an individual 
strobila, one form or the other very clearly 
predominates. The cirras sac of C. parvus 
(007-0010 mm) ts invariably smaller than 
that al C,owent (0.16-0.18) though the size 
difference may be a reflection of overall 
ceslode size. However, all specimens of 
C. parves examined possess an internal seminal 
vesicle, a feature which is not present in any 
congener, Et is not detectable in every single 
proglotts, bul is invariably visible in some pro- 
vlotlides of a strobils. These various differences 
are considered adequate for the erection of 
two new species, 

Comparisons af C, dorcopsis with C. macro- 
pus re complicated by the fact that the latter 
xpecies has been described on two occasions, 
ficsly by Ortlepp (1922) from Thylogale 
bruit and Jater by Sandars (1957) from 
Thylogale stiematica (=Thylegale wilcoxt), 
the descripiions differing on same important 
features such as sizé and the number of the 
rostellar hooks. 


C. dercapsiy is similar to hoth descriptions of 
C. macropus in the form of the rostellar cross, 
in the number and distribution of the testes 
and in the transversely elongated mature pro- 
glottis, ©. dorcepsis would appear to differ 
from oll congeners by the distribution of the 
testes Which eXtend laterally beyond the dorsal 
osmoregulatory canals to the medial edge of 
the ventral eanals.. However, this condition alsa 
exists in the type material af C. macrepus in 
the British Museum (Natural History), 
(thourh not stated by Ortlepp (1922) in his 
description) and in material collected by me 
from ViyvJegale stigmatica in Queensland, con- 
forming in all respects to the specimens des- 
cribed by Sacdars (1957) from the same host 
and same eeneral geopraphic region, Sandars 
(1957) gave the locality of fer specimens 
ineorreetly as ‘Mt. Tamborine in South Aus- 
tralia’; Mit. Tamborine is in Queensland; no 
spocies of Thylogale occur in South Austeali:. 

©. dorcapsis differs from ©. macropus in a 
number of features. The cirrus sac in 
C_ dorcapsis (0.20-0.25) mm) is larger than in 
descriptions of ©. macropus (0.10, 0.08-0.12 
mm); the rostellar hooks of C, dercapsis (12- 
lSum) arte slivhily Jenger than these of 


14) 


C. maeropus (9m) and the rostellar spines of 
C. doreepsis (7-114m) are longer than those 
of ©. macrapuy (5-G4m) and are of a different 
shape, The differences in the roslellar spines 
are very striking indeed as they are very pro- 
minent rose-thorn structures an the rostella of 
C. dercopsiy but ace inconspicuous and awl or 
hook shaped on rostella of C. macropus, Both 
Orilepp (1922) and Sandars (1957) des- 
cribed and illustrated the rostellar spines of 
©, maeropus as covering the entire rostellum 
with the exe¢eption of the area inside the 
rostellar cross. Tt has nal been possible to 
verify this situation since in the type specimen 
examined by me and in the new material from 
T. stigmatica, most of the rostellar spines are 
missing, Assuming the earlier descriptions to be 
correct, C. doreupsiy differs markedly from 
C. muacropus in having the rostellar spines 
arranged in about 2Q transverse rows, forming 
a distinct band on the everted rastellum but 
never covering the entire surface of it. 

Because of these citterences, C, dorcepsis sp. 
nov. has been described. It is evident however 
that the morphology of ©, macrapus requires 
re-investigation based on newW material from 
the type host, 7. brunil. The extant type 
material is inadequate for a thorough re- 
description and the stalus of specimens from 
T. sthematica, described by Sandars (1957) as 
C. macrepus requires clarification. 

The principal of differences hetween species 
of Calostaurtis are shown in Table 1- 


Acknowledgements 


Thanks are due te Dr I, L, Owen, Boroka, 
New Guinea; for calleeting and very carefully 
preserving the material described above, and to 
Mc k. A. Bray, British Museum (Natural 
History) for kindly lending the material for 
study, 


References 


RBevermec, f. (1975) On two new davaineid ces- 
todes from Australian marsupials. /. Aelmin- 
thol, 49, 129-36. 


OerLerr, R, J, (1922) A new duvaineid cestode 
—Ruillietina (Pareniella) macropa, sp, 0, from 
a wallaby. aa. «nag nat Hist, Ser 9, 9, 
402-12, 


SanpArs, D. F, (1957) Redescription of some 
castodes from marsupials 1-Davaineidae, Hyme- 
nolepididae and Anoplucephalidae. Aan. Trep. 
Med_ Parasital. 87, 330-9. 


FROG FAUNA OF THE NORTHERN TERRITORY: NEW 
DISTRIBUTIONAL RECORDS AND THE DESCRIPTION OF A NEW 
SPECIES 


BY MICHAEL J. TYLER, MARGARET DAVIES & ANGUS A. MARTIN 


Summary 


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. 


FROG FAUNA OF THE NORTHERN TERRITORY: NEW DISTRIBUTIONAL 
RECORDS AND THE DESCRIPTION OF A NEW SPECIES 


by MicwAse J. TyLer*, MarGarer Davies* & Ancus A. Martint 


Summary 


Tyirr, M. J. Davies. M, & Martin, A. A. (1981) Frog fauna of the Northern Territory: 
new distributional records and the description of a new species, Trans, KR. See, §. Auyt, WS 


(3), 149-154, 11 December, 1981. 


Uperoleia rachyderma sp. nov, is described from the Northern Territory, and additional 
biological and distributional data are given on C, lithomeda Tyley, Davies & Martin, Ranti- 
della desertivala Liem & Ingram is recorded in the Northern Tertitory for the first time, and 
Cyclorand cryploriy Tyler & Martin is recorded for the second time. 


Tatroduction 


Our collaborative studies on the frogs of 
northern Australia have led to the addition 
of seven species to the North Territory fauna 
(Martin, ef al. 1Y80; Tyler, er al. 1978, 1981a5 
Tyler, ef al, 1979), Our previous. field studies 
in the Northern Territory have been confined 
to the Fust Alligator River area, 

However in December 1980 we collected at 
various sites on the Stuart Highway between 
Darwin and Tennant Creek, These collections 
include 4 new species of the Jeptodactylid 
venus Uperoleia, and a second leptedactylid 
species not previously recorded from the 
Northern Territory. We also provide additional 
information on the distribution and biology of 
U. ltthamada Tyler, Davies & Martin, pre- 
viously recorded in the N.T, only from the 
flood plains west of Arnhem Land (Tyler e¢ 
al, 198la). and the hylid Cyclorqna ervprotis 
Tyler & Martin 


Material and Methods 
The specimens reported here are located in 
the following collections: American Musetint 
of Natural History (AMNEH); Northern Ter- 
ritery Museum, Darwin (NTM); Museum of 
Natural History, University of Kansas. Law- 


rence (KU); South Australian Museum 
(SAM). 

Methods of meustirement follow ‘Tyler 
(1968), Abbreviations used in the text are: EB 


(eye dimmeter), E-N (eye to naris distance), 
IN (internarial span). SV (snout ta vent 
length), PL (tibia length). 


* Department of Zoology, University of Adelaide. 
Box 498 G.P.O., Adelaide, S. Aust. SOOT, 

' Deparment of Zoalogy, University of Mel- 
hourne, 


Osteological data were obtained from 
cleared and Alizarin Red stained preparations 
using the technique of Davis & Gore (1947). 
Osteological descriptions follow Trueb (1979), 

Male malting calls were recorded with a 
Sony TC-510-2 tape recorder and a Beyer 
M 88 dynamic microphone. at a tape speed 
of 19 em/sce. Wet-bulb air temperatures were 
mensured close to the calling sites of males 
using a Schultheis quick-reading thermometer. 
Culls were analysed by means of a UV oscil- 
lowruph (San-Ei Visilight) and a sound spec- 
(rovraph (Kay Model 6061-B Sona-Graph) 
with the averall response curve maintained 
in the FL-) position, Three calls of each male 
were analysed and mean values calculated, 


Famicy: Leptodactylidac 
Uperoleis trachyderma sp, nov. 
FIGS 1-4 
Holeiype SAM ®20374, an adult male coal- 
lected on the Neweastle Creek floodplain at 
the George Redman Causeway (17°14'S; 
133°28'F) 37 km N_ of Elliot, N-T. on 
16.41.80 by M. Davies, A. A. Martin and 
M.. J. Tyler, 
Definitions A small species (males 20.3-22.0 
mm S-V) with small eyes (eye diameter 
equivalent ta eye to narig distance), with the 
dorstim covered with small, conical tubercles, 
and with a heavily pizmented ventral surface. 
Mating call a staccato hurst of four short 
pulses. 
Description of halotype: Maxillary and vome- 
rine teeth absent. Snout elongate, pointed 
when viewed from ubove; projecting slightly 
in profile. Fye to naris distance greater than 
internirial span (E-N/TN 1.27). Canthus 
rostralis straight. Tympanum not visible ex- 
ternally (Fig. 1), 


su 


Fig. 1. Darsolateral view of bead of U/perelvia 
traclryelertna, 


Fingers long, slender, unwebbed snd un- 
fringed With prominent subarticular tubercles 
and well developed palmar tubercles. (Pig. 
2A), Fingers in order of length 3 > 4 > 2 > 
1, Hindlimbs very short (TL/S-V 0.35), Toes 
long, slender, slightly fringed and unwebbed 
(Fig. 2B). Toes in order of length 4 > 3 > 
5 > 22> |, Metatarsal tubercles small and 
poorly developed. Subarticular 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. 
Cloacal fap narrow but Well developed. No 
heel tubercles. 


Dorsal surface dull slate with obscure, 
slightly darker motiling; glands faintly 
creamish, Inguinal and post-femoral flash 
markings doll ted. On ventral surface sub- 
mandibular area pale grey; abdomen with 
numerous small islands of pale grey on a white 
background, 


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, 


Fariajion’ There are four paratypes; all are 
adult males: KU 189561, NTM 9865, SAM 
R20375-6. The series was collected with the 
holotype. 


The range of body size spans only 1,7 mm 
(20,3-22,0 mm S-V). The hindlimbs are short 
(TL/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 naris distance. The eye to naris dis- 


MICHAEL J, TYLER, MARGARET DAVIES & ANGUS A. MARTIN 


tance is greater than the internarial span 
(E-N/TN 1,13-1.40), 


The unusual tubercular condition of the 
dorsal skin is apparent in all specimens, and 
m some is more conspicuous than in the 
holotype. 


The dorsal coloration is reasonably uniform 
except that in one specimen the parotoid, 
inguinal and coeccygeal glands are bright 
yellow-cream and stippled with black. 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, excepi for the dermal glands 
which are suffused with pale orange. The 
inguinal and post-femoral flash markings are 
bright carmine, The sub-mandibular area is 
slate, and the abdomen is marked with islands 
of grey upon a ereamish-grey background. 


Ovtcolagy; Skull (Pig, 3A) moderately ossi- 
fied, sloping anteroventrally, Sphenethmoid 
poorly ossified, not conjoined medially cither 
dorsally or ventrally, Dorsally sphencthmoid 
in tenuous contact with nasals, Ossified portion 
of sphenethmoid extending ventrally for 
about 4; length of orbit. Prootic and exocel- 
pital mot fused dorso- or ventromedially- 


Fenestrae present dorsally and ventrally in 
crista parotica, Crista parotica short, thick, 
slightly overlapped medially by posterior ex- 
tremities of frontoparietals, widely separated 
unexpanded — otic 


laterally from rami of 


squamosals, 


Fig. 2. (A) Palmar view of hand and (B) plantar 
view Of foot of Uperaleia trachyderma. 


FROGS OF NORTHERN TERRITORY 


Vig. 3. (A) Dorsal and (B) ventral view of skull 


Frontoparietal fontanelle extensively  ex- 
posed; anterior and posterior extremities being 
undefined because of lack of medial ossifica- 
tion of sphenethmoid and exoccipital regions, 
Nasuls 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 maxillary, 


Palatines reduced (Fig, 3B), moderately 
broad, overlying ossified portion 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, Alac broad, 
short, slightly angled posteriorly from cultri- 
form process, Plerygoids moderately robust; 
anterior arm slender, in moderately short con- 
tact with poorly developed pterygoid process 
of palatal shelf of maxillary, Medial arn 
moderately long, very robust; posierior arm 
short and mwioderately sharp. 


Carlilaginous quadfate present between base 
of squamosal and quadratojugal, Quadrato- 
jugal robust, in firm contact with maxillary. 
Squamosals stocky, no zygomatic ramus. 
Jong ubexpanded otic ramus. Maxillary and 
premaxillary edentate. Alary processes of pre- 
maxillaries moderately broad, perpendicular 
to premaxillary, Palatine processes of pre- 
maxillares well developed, not abutting 
medially. Palatal shelf moderately deep with 
poorly developed pterygoid process, Preyorners 
absent, Bony columella present. 


of Lperaleia tracivderma, 


Pectoral girdle arciferal, robust. Orno- 
sternum ubsent, xiphisternum present. Clavi- 
cles slender, curved, closely applicd medially, 
Coracoids robust, widely separated medially, 
Bicapitate scapula considerably shorter than 
clavicles. Suprascapula about } ossifled, 


Light precocious non-imbricate presacral 
vertebrae, Sacral diapophyses poarly to mode- 
rately expanded. Relative widths of transverse 
processes UE > sacrum > IV > Il > V = 
Vi = VIL — VIIT. Ilia extending to anterior 
extremity of sacral diapophyses. Sacrococ- 
eygeal articulation bicondylar. Urostyle with 
well developed crest extending about 4+ of 
its length. 


Humerus with strongly developed antero- 
proximal crest. Phalangeal formula of hand 
2-2-3-3, No bony prepollex. Palmar sesamoids 
present. Phalangeul formula af foot 2-2-3-4-3. 


Well developed bony prehallux. Terminal 
phalinges kKnabbed, 
Camparison with arher species: Uperoleia 


trachyderma is a distinctive species. particu- 
larly in the darsal skin texture of dense, small 
but prominent tubercles, Other species with 
extensive frontoparictal tontanelles comparable 
to the condition ia LL rrachydertna (UW. ris- 
velli, LU. arenicola, U. herealis, U. talpa, and 
U. arientaliv) have moderate to extensive Web- 
bing between the toes (except in U. arenicela), 
Uperaleia arenivala resembles U/. trachyderma 
in lacking webbing, but the dorsal skin 1s 
smooth or weakly tubercular (tubercular in 
UU, traclyderma), The call of U, orientalis 


{52 MICHAEL J, TYLER, MARGARET DAVIES & ANGUS A, MARTIN 


ee Mm 


02 0-4 06 08 10 


Fig. 4, Upper: oscillographic tracings of male 
mating calls of: left, Uperoleia lithomoda, 6.4 
km N of Katherine, N.Ty; right, Ranidella 
deserticola, 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- 
uve calls of Uperoleia trachyderma, 37 km NW 
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. trachyderma), No 
other northwestern Australian congener is 
known to have a four-pulsed mating call, 

The similar-sized U, lithomoda taken north 
of the site of U. trachyderma has poorly 
developed dorsal turbercles, a shorter call, a 
very poorly exposed frontoparietal foramen, 
and larger eyes, 


Mating call: Calls of two males were recorded 
37 km N of Elliot, N.T., on 16.xi1.80, An 
audiospectrogram of a call is shown in Fig. 4, 
and physical characteristics of calls are listed 
in Table 1, The call is a harsh “creak”, 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: 
locality. 
Habitats The type locality is a Mat floodplain 
of adhesive yellow clay, Males were calling 
from the base of grass tussocks growing in 
the water, The surroundings were flooded to a 
depth of 0.3 m. 


Etymology: From the Greek trachys, ‘rough’, 
and derma ‘skin’, in reference to the unusual 
skin condition. 


Known only from the type 


Ranidella deserticola Liem & Ingram, 1977 


This species was known previously from 
localities in southWestern Queensland (Liem & 
Ingram 1977, Tyler 1978) and northeastern 
South Australia (Brooks 1980), 


We found &, deserticola upon the Neweastle 
Creek floodplain 40 km = north of Elliot 
(170°14°S; 133°28'R) on 16.xii1.80, Five 
males and one female were collected beneath 
debris at the edge of a deep dam (SAM 
RI9L18-23). 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 to that 
described by Liem & Ingram (1977), but two 
clear emphasized frequency bands are evident 


Taste |. Physical characteristics of male mating calls of Uperoleia trachyderma and U. lithomeda. 
Mean values are given with ranges in parentheses, 
Species and No. of Duration Pulse 
locality N pulses (msec) repetition Dominant Wet bulb 
rate frequency temp. 
(pulses/ sec) (Hz) (°C) 
U. trachyderma 2 4 51.4 78.5 3600 24.2-27.0 
37 km N of (4) (48,0-54,7) (73.0-84.0) (3500-3700) 
Elliot, N.T, 
U. lithomoda 3 5.3 16,3 342 3250 26,5 
6.4km N of (5-6) (13,0+-20.0) (250-462) (3150-3400) 
Katherine, N.T. 
U. Wthamoda 5 4.6 11.6 401 3420 26.0) 
Lt.S km N of (4-5) (9.0)-13.0)) (333-456) (3200-3600) 


Lake Argyle 
Village, W.A, 


ON kwvwvwn— SSS 


FROGS OF NORTHERN TRRRITORY 183 


at about 3450 and 4050 Hz. The note dura- 
tion, and call duration are shorter than re- 
corded by Liem & Ingram (1977), but their 
recording may have been made at a lower 
lemperalure: they gave no temperature data. 
Values trom our recordings are: call duration, 
79-80 msec; first note duration, 24-32 msec) 
second note duration, 33-41 msec, 


Examination of other collections reveals 
that R. deserticola is abundant in the New- 
castle Waters area, M. Gillam found speci- 
mens (SAM KI9f84) 24 km W of Newcastle 
Waters fiomestead (17°32'S, 133°22’E) on 
1.41.76 in hollows on the mud banks of a 
turkey nest dam. G. A. Crook and W, Zeidler 
collected 16 specimens at Lake Woods. 15 km 
WNW of Elliot on Sx%-77 (SAM R19137-52). 
P. Spalding and W. Hosmer collected the 
species at Elliot on L1.iv.60 (AMNH 67143- 
45), Neweastle Waters on |3,iv.40 (AMNE 
67146-52) and Anthony Lagoon on 23,iv.60 
(AMNH 67153). 


Snout to vent lengths of our series and those 
found by Crook and Zeidler are: males 14.6- 
18.7 mm; females 14.6-17.8 min, Liem & 
Ingram (1977) cite a range of 13.0-18.4 mm 
for their series, hut do not indicate the sex 
of the specimens. Presumubly it embraces the 
range of both sexes. 


The only congeners recorded from Northern 
Territory are R, bilineua Martin, ‘Tyler & 
Davies (1980) and &, remota Tyler & Parker. 
From the former R. deserticela can be dis- 
tinguished by its smaller size (hilingia males 
15.5-23.3 mm; females 174-2010 mm S8-V) 
and different mating call. We located R, 
hilingua ealling within 10 km of Katherine 
Garge. Most of ike Northern Territory records 
of remota by Barker & Grigg (1977) and 
Cogeer (1979) wre presumably based on 
Ailineue. However remeta oveurs in Queens- 
land avd it might occupy the eastern portion 
of Northern Territory. Ut is distinguished most 
readily by its long, highly pulsed call (Tyler 
& Parker 1974). 


Uperaleia lithhameda Tyler, Davies & 
Martin, 1981 


Wperoleia lithomeda was known previously 
from two disjunct populations: one jn the 
easter) Kimberley region, W.A., and one on 
the Mood plains west of Arnhern Land, N.T. 
(Tyler et al. \98iad. We have now partially 


closed the intervening gap by obtaining speci- 
mens of (. lithomoda near Katherine, N-T. 


On 14.xi1,80 we collected seven adult males 
and one adult female (SAM R20440-7) of this 
species within 5 mi of the Stuart Highway 6.4 
km N of Katherine (1,4 km N of the northern 
limit of Katherine township) (14°25'S; 132° 
16°E), and recorded the 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 
fernale 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- 
lions. However in life the dermal elands are 
light golden, appearing as continuous stripes 
along the sides, The flash markings in the 
groin and behind the thighs are scarlet. 


To the ear the mating call is a single, abrupt 
“olick™: and in fact Tyler et al. (1981a) 
described the call of the Kimberley popula- 
tion as consisting of a single pulse. However 
ascillographic analysis of the calls of Kathe- 
tine males showed the cail to consist of a 
rapid burst of pulses, Hence calls af the Kim- 
berley populations were re-analysed, and alsa 
shown to have multiple-pulsed calls, Fig, 4 
shows an oscillographic tracing of the call 
of a Katherine male, and Table | lists the 
physical characteristics of calls of Kimberley 
and Katherine males. 


The call of U. ittermoda may be described 
as a short burst of 4-6 pulses in 9-20 msec, 
with a pulse repetition rate of 250-450 pulses/ 
sec. The dominant Frequency 15 sheut 3300 
Hz. 


FAminy: Hylidae 
Cyclorana ervpiatiy Tyler & Mariin, 1975 


This species Was described from an adult 
mate collected at Daly Waters on 13.xi1..1971; 
we found a further single adult male upon the 
George Redman Causeway. 37 km N of Elliot 
on 16.xiL L980, Our specimen (SAM R18973) 
was amongst a large number of €, cultripes 
Parker uctive on the road surface. (We col- 
lected 41 in 15 mins: many more individuals 
Were present. whilst C. enstraliy and ©. mracu- 
lass’ were present but slightly less abundant)- 

In the Kimberley of W.A. ©. ecryptotis 
breeds in February. and C_ evilfripey is. en- 
countered rarely in that period (Tyler et al. 
1981b, 1982). Our experiences with these 


154 MICHAEL. J. TYLER, MARGARET DAVIES & ANGUS A, MARTIN 


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. eryptotis in the N.T. 110 km fur- 
ther south. We note that C, erypteris is absent 
from the northern floodplain of the N.T., and 
the species occupies a narrow latitudinal 
zone illustrated by Tyler ef al. (1982). 


Acknowledgements 
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. 


References 


Barker, J. & Gricc, G, C, (1977) “A field guide 
to Australian frogs.” (Rigby; Adelaide, ) 

Brooks, J. (1980) First record of the frog 
Ranidella deserticola, Liem and Ingram from 
South Australia. 9. Aust. Nat. 54, 45-46, 

Coccer, H. G. (1979) “Reptiles and amphibians 
of Australia,” (Reed; Sydney.) 

Davis, D. D, & Gore, V. R. (1947) Clearing and 
Stuining skeletons of small vertebrates. 
Fieldiana: Techniques (4), 1-16. 

Liem, D. S. & Tncram, G. J. (1977) Two new 
species of frogs (Anura: Myobatrachidae, 
Pelodryadidaoe) from Queensland and New 
South Wales, Vicr. Nar, 94, 255-262, 

Martin, A. A., TYLeR, M, J. & Davies, M. 
(1980) A new species of Ranidella (Anura: 
Leptodactylidae) from northwestern Austra- 
lia, Copeia 1980(1), 93-99, 

True, L, (1979) Leptodactylid frogs of the 
genus Telmatobius in Ecuador, with description 
of a new species. /hid, 1979(4), 714-733. 

Tyter, M. J. (1968) Papuan hylid frogs of the 
gon Ayla, Zool, Ferhand. Leiden (96), 1- 
203. 


(1978) “Amphibia of South Australia,” 
(Govt Printer: Adelaide.) 

. Davies, M, & Martin, A. A. (1978) A 
new species of hylid frog from the Northern 
Territory. Trans. R. Soc. §. Aust. 102, 151- 
157. 


‘ & —— (1981u) Australian frogs of 
the leptodactylid genus Uperoleia Gray. Aust. J. 
Zool, Suppl. Ser. (79), 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 systematics 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 Cyclorana cryptetis Hylidae). 
Copeia 1982(3), (in press). 

& Parker, F. (1974) New species of hylid 

and leptodactylid frogs from southern New 

Guinea, Trans, R. Soc. S. Aust. 98, 71-77. 


(Anura: 


NEW RECORDS OF OPHIUROIDEA (ECHINODERMATA) FROM 
SOUTHERN AUSTRALIA, INCLUDING NEW SPECIES OF 
OPHIACANTHA AND OPHIONEREIS 


BY ALAN N. BAKER & DENNIS M. DEVANEY 


Summary 


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. 


NEW RECORDS OF OPHIUROIDEA (ECHINODERMATA) FROM 
SOUTHERN AUSTRALIA, INCLUDING NEW SPECIES OF 
OPHIACANTHA AND OPHIONEREIS 


by ALan N. Baker* & Dennis M, DEVANEY? 


Summary 


Baxue, A, N. & Devaney, D. M. (198L). New records of Ophiuroidea (Euchinedermata) from 
soothern Australia, including new species of Ophiacantha and Ophionereis. Trans, RB. Soc- 
S. Aus, 105(4). 155-178, (1 December, 198], 


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 shepherd 
i. sp. are described from Victorian and South Australian waters respectively, and a key to 
the southern Australian species of Ophiacanthe is provided, Ophioveres bixpinosus, Ophioconis 
opacum, avd Ophieprium raxea ure new combinations, and the following species are 
synonymised: Ophiacantha abyssicola otagoensiy Fell With Q. brachygnatha H. &. Clark, 
Ophiacantha truncata Koehler with Ophiaprium rosea (Lyman), and Ophiaetiy laevis HL Lb. 
Clark with O- pricdfor H. Lb. Clark, Mlustrations are given of holorypes of Ophioreres his- 
pinesus, Ophiomusium anisacanihum, O. australe, Ophionereis lineata, Amphiuria trivacaniha, 
Ophrioathrix (Placophiothrix) albostriata, O. (Placophiothrix) lineovacrulea, atid OG. (Keystonea) 


hvmenacantha, A lectotype is designated for Ophiecanrha clavigera Koehler. 


Introduction 


The ophiuroid fauna of southern Australian 
waters is known largely from the studies of 
H, L, Clark (1916, 1918, 1928, 1938, 1946) 
and A. M. Clark (1966), H, L, Clark's eurly 
paper covered collections trawled from the con- 
tinental shelf by F.1S. “Endeavour” in 1910- 
14, and the later works dealt with specimens 
collected mainly from the Victonan coast by 
Joseph Gabriel, and from South Australian 
waters by Sir Joseph Verco and Clark himself 
during a visit in 1929, In 1946, Clark added 
more specimens in his revision of the entire 
Australian echinoderm 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 Compurative 
Zoology, and British Muscum (Natural His- 
tory). Seventy-four species of ophiuroids are 
new known fron southern Australia between 
Cape Howe. Victoria (37°30.1'S) and Cape 
Naturaliste, Western Australia (33°30,1'S) 
(Clark 1946, Baker }981, and this paper), 


In the course of studying Australasian 
Ophiuroides in the past few years we huve cxa- 


* National Musctin of New Zeulind, Private Buy. 
Wellington, New Zealand. 

+ Bernice P. Bishop Museum, P.O. Box |900D-A, 
Honolulu, Hawaii. 


mined recent collections made by trawling ex- 
peditions (H,M.A,S. Diamantina and Kimbla, 
M.Y, Aquarius, and M,V, Sarda) and divers 
working along the coasts of Western Australia, 
South Australia, Vietoria, and Tasmania. 
Among these collections are a number of spe- 
cies. previously Unkiown in the region, includ- 
ing {Wo NeW species and offers hitherto known 
only from other Pacific localibes, particularly 
New Zealand, Some of these have been dis- 
cussed by Baker (1979, 1980), and the re- 
mainder are reporicad here, 


During visits to the South Australian 
Museum, one of us (A.N.B.) re-examined 
some of the ophiuroid material recorded by 
H, L. Clark (1928), and re-illustraled, by 
camera lucida drawings, severul holotypes of 
local species described but not figured in detail 
by that worker, Our studies have shown that 
some taxonomic changes ure necessary. and we 
have included such information in this paper. 


Text conyentions 

The following abbreviations are used: Irsti- 
nitions; AM—Australian Museum, Sydney, 
BM(NH)—B8ritish Museum (Natural History), 
London. BPBM—BSernice P. Bishop Muscum, 
Honolulu, Hawaii, MCZ—Museun of Comte 
parative Zoology, Harvard. MNB—Museum 
fiir Naturkunde, Berlin, NMN#—WNational 
Museum of New Zealand, Wellington. NMY— 
National Museum of Victoria, Melbourne. 


156 A. N, BAKER & D, M. DEVANEY 


SOUTHERN AUSTRALIAN OPHIUROIDEA \S7 


SAM—South Ausirolian Museum, Adelaide 
WAM-—Western Anistralian Museum, Perth. 
ZMH—Zoolavisches Museum, Humburg. ZMC 
—Zoologisk Museum, Copenhageu. Morpha- 
logy: d.d.—disc diameter, 


Systematic Account 


Pamity Ophiundae 
Subfamily Ophiolepidinac 
Ophiaceres Koehler, 1922 
Ophiaceres hispinesus (HO 1, Clark) nm camb, 
FIGS |-3, 5, 14-17 


Oplioplocns biypiiasus Hy, Le Clark, IB. 347, 
pl, 4, fig. 2; 1946: 278- A. M, Clark, 1966: 
327.- Darnall. 1980: 43. 

Specimeris entmined; Holotype, MCZ 4025, 

Phillip ts. Wosternport Vie. no depth. May. 115: 

2, NMNZ 2074, Port Macdonnell, $3.4. 3-4 m 

29,n7-1964; 1, NMNZ 2076, West Id, S.A, 4 m, 

29.1,1975; 2, NMY, H366, Cape Liptrap, Vie, 

no depth, 24.1979; 2, NMNZ 2075, Blurei 

Cape, Bruny hl. Tas, 13 om, 10.1972: 1, NMNZ 

2702. Deal Td. Vas, 20 m, 4-v, 1974. 


Remarky; These specinens range 3-9.5 mm 
d.d.. and have arms up ta 3.3 & did long. 
They are thus similar to the type serics in 
dimensions. 

That this species belongs in Ophiacerey 
rather than Opfiiuplocus ig shown by the pre- 
senee of one tentacle scale to each pore, shart 
genital slits originating close to the oral shield 
ind hardly extending past the first ventral arm 
plate, wd the relatively small amount af Frae- 
Mentation of the dorsal arm plates (<1) 
pieces), In Ophioplecuy Lyman, there are 3-6 
tenlacle scales, genital slits which are separated 
from the oral shields by small platelets or scales 
and Which extend a§ fay as the fourth scament, 
and extensive dorsal arm plate fragmentation 
(<20 pieces). 

As pointed our by H.C, Clark (1918), this 
species js extremely close to O, fyttan{ (Far- 
quhac) from New Zealand. Examination of 15 
specimens of the latter (NMNZ) shows small 
but consistent differences which we consider 


Figs 1-21. 
ventral; 
type MCZ 4025)- 6-9, Ind. Sth, 


(NMNZ 274); 


18th and distal army segments of O. 
(0.13, sume seyments of O. marginata (NMNZ 2705): 


to be specific, Clark's suggestion that the num- 
ber and arrangement of darsal arm plate frag- 
ments is different in these (wo species is cor- 
rect, but only for the basal and middle part of 
the arms, To confuse matters, however, the 
basic pattern af fragmentation may be altered 
hy irregular secondary division of plates on 
gome 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 the 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 portian of the central plate by an 
oblique plate on each side (Figs 16 & 20). At 
this stage of Tragmentation, a difference be- 
tween the two species is evident—in Auton 
the pair of additional oblique proximal centro- 
lateral plates (pel) nearly as wide as the distal 
centro-laterals (del) and fully in contact with 
the loteral edge of the proximal central plate 
(pc), becomes wedyed between the proximal 
centro-lateralg and the distal central plate; in 
hispinosus however, the additional pel plates 
are usually less than 3 the width of the del 
plates and only in contact with the posterior 
lateral portion of the pe plate, On the same arm 
scement, and even more pronounced on proxi- 
taal segments (Figs 15 & 19), the latter species 
does not develop a third pair of plates distad 
or disto-lateral to the dep as seen in Auttani 
(Figs 16 & 20, 15 & 19), Occasionally, irregu- 
lar longitudinal splits occur on the very basal 
arm plates of O. hivpinosus (Pigs 4 & 14). 

The oral shields also differ in proportions— 
in Aispinosus they are noticeably wider than 
Jung (Wo 1.44-1.63L), whercas tn Aurtoni 
they are whout as wide as long (w = 1.05- 
1.138). 

Although there ig no noticeable difference 
in the arrangement of the dise scales, the radial 
shields and marginal inter-radial plates are 
larger In bispinesuy than in Auttoar, In fact the 
size of the plutes rival those in O, marginata 
Fell. a second New Zealand species, regarded 


\, arm hase and adjoining disc, dorsal, of Ophioceres hispinasys (NMN?Z 2074): 2, same, 
3, arm base und adjoining dist. ventral, of Ophiocerey incipiens (NMNZ 2715): 4. 6th 
& 7th arm segments of QO. bispinosus (NMV H3n6}; 


5, ith arm segment of O. hispinosys (hola- 
inciptens (NMNZ 2718)- 
1417, same segments of O. Aispinosux 


1821, saroe Resanents of O. fiiitone (NMNZ 1183). Abbreviations: pe— proximal 


ceolral plates: del—distal centro-lateral plates; pel—proximal centro-lalera) plates. Scale lines 0.5 


nim, 


158 A. N. BAKER & D. M. DEVANEY 


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, 114°51'E, 
W of Lancelin, W.A., 130-160m, HMAS Dia- 
mantina 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, 1928: 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, 1859 
Ophionereis terba n.sp. 

FIG. 30 
Specimens examined: Holotype NMV H363 & 
paratype, NMNZ 3673, 43 km SSW Portland, 
Vic., 585m, coll. R. Plant & M. Gomon, 14.y. 
1979; paratypes 4, NMV, W of Cape Nelson, 
Vic., 164-201m, 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 


SOUTHERN AUSTRALIAN OPHIUROIDEA 159 


Figs 22-27. 22, arm base and adjoining disc, dorsal, of Ophiomusium anisacanthum (holotype SAM 
K254); 23, same, ventral, 24, Ist & 2nd lateral arm plates of SAM K254; 25, arm base and 
adjoining disc, dorsal, of Ophiomusium australe (holotype SAM K256); 26, same, ventral; 27, 
lst & 2nd lateral arm plates of SAM K256. Scale line 0.5 mm, 


160 


A. WN, BAKER & D, M, DEVANEY 


mi 


a Ki 
Se Secu, 
A ey 


Figs 2836. arm bases and adjoining dises. 28 & 24, Cpldonercis lineata (holotype, MCZ $829) der- 
sal, & yeotrul; 30, Gphianereis terha n, op. (holotype, MMV W363), dorsal. 31 & 32, Opliivconis 
upacien UNMNZ 2690) docsal & youll; 33 & 44, Ophinvriy tricelor (holotype SAM K213) ven- 
trol & dorsal) 33 & 36, Gphiecomina. wustraliy (BPAM W822) dorsal & ventral, Scale lines 10 


enna, 


SOUTHERN AUSTRALIAN OPHIUROIDEA 161 


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 are 
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, 
MCZ 5829). 


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, pl. XIV, 
figs 398-400. Baker, 1979: 34. 

Specimens examined: 3, NMV H368, 38°3.4’S, 

149°23.7’B, E of Flinders Id, Bass Strait, 183 m, 

22.xi.1973; 1, NMV H360, 39° 32.5’S, 148° 

51.5°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 australis H. L. Clark 
FIGS 35, 36, 63-68 


Ophiocomina australis 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 australis 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 spp. 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 Ist arm vertebra, are 
considered features that do not indicate family 
resemblance between the two genera. 


162 ALN. BAKER & D. M. DEVANEY 


TABLE 1. Comparison of Ophiocomina with Ophiacanthids and Opltiiocomids. 


TEETH 


Ophio- Ophio- Ophia- Other ophio- 
coming pleris Clarkcoma  canthids comids 

ORAL PLATES 
A. With radiating indentations on abradial 

muscle scar —e — a +(—)) 
RB. Abradiatl muscle scar with flaring 

aboral wing _ a oS +f 
C, Adradial muscle scar less than (—), 

equal to (+), more than (-+-) 

half height of plate — - x4 +, — 
D. Longer than high (+), equal (+), 

higher than long (—) ++ + — + + 
2. Adradial articular area with upper 

distal edge rounded (++) or straight (—) + - +. — 

Hylinated tips — = + = + 
ARM SPINES 

Hollow (lumen more than haif 

wall diameter) + — + + (+)2 
DENTAL PLATE 
A. Forantina = ++ + — 
B, Oral end widest + + + = —(+)# 
C, Papillae present +- + 4 (+) 
RADIAL SHIELD AND GENITAL PLATE ARTICULATION 
Two condyles and one pit + + + -,— + 


“length = proximo-distal axis 
height = oral-aboral axis 


mella 


FAMILY Ophiodermatidae 
Genus Ophioconis Litken, 1869 
Ophloconis opacum (H. L, Clark) n.comb. 
FIGS 31, 32 


Ophiuroadon opacum WH, LL, Clark, 1928: 440, 
figs 1324 & b, 1946 255; A. M. Clark, 1966: 
327. 

Specimens examined; Holotype, SAM K243, “St 

Vincent or Spencer Gulfs", S.A. no depth; 1, 

NMNZ 2122, Upper Spencer Gulf, S.A., LJ m, 

4.1x.1973; 1, NMNZ 2690, Upper Spencer Gulf, 

S.A, 18 m, 11,ix.1973; 1, NMNZ 2123, off 

Glenelg, S.A,, artificial reef sile, 3-4 m, 1972; 

|, NMNZ 2722, W of Outer Harbour Gulf St 

Vincent, SAL, 5 m, 3.1.1965; 2, BPBM W2220 & 

WAM 31-74, Cockburn Sd, W.A. under stones, 

0-2 m, 13,1,1972; 2, WAM 1133-74, 3.2 km 

NW Busselton Jetty, W.A., Posidonia, Cymuo- 

docea & sand, 22-24 m, no date. 

Remarks: H. L, Clark (1946) referred three 

Australian species to Ophiurodon. O. opacumn 

Was separated from its northern congeners 

(cinela und permixta) by having blunter, flat 

Opaque upper (at least) arm spines. However, 


1(~-) for Ophiocoma *(+) for O, pusitle 
pusilla; Ophiap- 
sila, Ophiaco- 


4(--+) for ophiocoma 
longispina, O. 
pusilla and 
Ophiomastix 
variahilis, 


and Ophiomies- 
ix Ornata, 


A. M. Clark (1965) found no generic distinc- 
tion after comparing the type species of Ophio- 
conis (O, forbest) with permixta and cincta. 
She did not consider O. opacum at that time 
and, in u 1966 key retained without comment 
epacium in Ophinradon. 

Comparison of specimens of the three known 
Australian species indicate they are indeed con- 
generic and should all be retained in Ophio- 
conis, 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 shiclds neatly 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 
first recorded since the type description. They 
range 3.0-6.5 mm d.d., and their characters 
confirm the distinctness of this southern species 
ol Ophiacenis, Tt may be separated from the 
two northern Australian species by its flat, 
almost spatulate arm spines. Contrary to the 


SOUTHERN AUSTRALIAN OPHIUROIDLA 163 


type deseviption, O. opacnm does have finely 
stciated lateral arm plates but apart from thal 
feature, the holotype and hew material agree 
with H, L. Clark's description, Because H. L 
Clark's illustration lacks detail, we give bere a 
camera lycida drawing of NMNZ 2690 (Pigs 
31, 32) the holotype being not in a suitable 
condition for illustration. 

The records of OG. epachim front Western 
Australia extend the known range of the spe- 
cies westward from the South Australian guifs: 
this extension and its shallow water habitat 
(0-24 m) indicate thal a! ts probably raare 
widespread on fhe southern coasts of Australia, 


Bathypectinura Hy, 1, Clark, 1909 
Berhypercrinure heros {Lyman 
FPeetinira heros Lyman. 187%: 48 pl 
489-91, 
Barhypectinura feros: H, L, Clark, 1909 129-30, 
Baker, 1979; 34, 
Specimens examined: |, SAM TK 3132, 37°58'- 
37°47'S, 139° 49'-139°45°E. SW of Cupe Buffon, 
S.A, 548 m, May, 1981; 2 NMV, 43 km SSW 
af Portland, Vic., 585 m, 14.¥.)979, 
Reimuirks; This larue species (TK 3132 is 62 
nim did.) has previously been reeorded from 
off Bateman'’s Bay, New South Wales, unc 
south of Norfolk Island (Baker 1979) in the 
Austratian region. Tt has a worldwide distribu- 
tion in 240-2960 m. 


14, figs 


Famity Ophiacanthmae 
Ophiacantha MiNer & Vroschel. 1842 
Ophiacantha shepherdi n.sp- 

FIGS 37, 38. 69-74 
Specimens examined: Holotype. SAM KAI750 & 
paralype BPBM WHS3), Seal Rock, Encounter 
Alay, West td, S.A. 24 m, on algac, coll. S.A, 
Shepherd, 18.ix.1967, Paratypes, |, SAM [i749 
& 1 NMNZ 3083, Deva Inlet. West Id. SAL, Lo 
m. coll. S.A. Shepherd. 24.97.1467. Paratype, 
NAMINZ, 11 km off Glenelg, Gulf St Vincent, SA., 
25 m. coll. S.A. Shepherd. 27.411-1966, Purutypes, 
8, NMNZ 3085, 2, BPBM W1835, Lund’s End. 
West Id, S.A. 17-18 m, coll, S.A, Shepherd, 
1967. Paratypes. 2, BPBM 1839, Seal Rock, West 
Id, SA, 24 m, coll, S.A. Shepherd, 35.11.1967, 
Paralypes, 2, WAM, 19-81, Gedipus Pr, West Tel. 
S.A. 18 m, call, S.A. Shepherd. no date: Puri 
types, I. AM, 1, BM(NH), 4, NMNZ 364), 
Middle Pt, Cape Northumberland, S.A, 13) m, 

in algae, coll. S:A. Shepherd, 19.1974. 

Description of healetype: Dise diameter 2.5 
mov arm teneth [!-!2 mm, arms taper gre 
dually, Upper side of dise with very low 
conical granules (rarely us high as broad), 


slightly larger and more prominent inter- 
radially than radially; eranmules generally nol 
contiguous, underlying seales evident between 
them, Radial shields so¢k-shaped, distal ends 
exposed and adjacent to lateral edge of first 
exposed dorsal arm plate; also in contuet with 
genital plate below, On lower side of disc, vra- 
nules more widely spaced proximally; disc 
scales evident 

Oral shields approximately 1,5 > brogder 
than Tong, somewhal triangular, with small lobe 
in distal cdye, madrepontic plate much larger, 
with greater distal prominance. Adoral shields 
large, meeting broadly within, encroaching 
distad uround most of ventral shield. 

Three oral papillae on edge of oral plate: 
ouler (wo longer than broad, similar in size to 
inner papila which is more sharply tapered 
and apically directed, The first tooth level with 
oral papillae bul larger and arrow-head shaped, 
other veth not visible. 

Pirst ventral arny plates slithtly broader than 
following plates which are tan-shaped. with the 
distal edge convex (Fig. 69): plates separated 
fram one another by lateral arm plates, the 
separation increasing distally, 

One tentacle seale, less than 4 a segment 
lang, rectangular with rounded tip (Fig. 70); 
scales remuimiig approximately saute sie and 
shape on distil segments in contrast to dimi- 
nishing size of other arn plates, 

Dorsal arm Plates broadly fan shaped, 1.5 « 
or more hroader than long: with exception 
of the first exposed plate which is in contact 
with second plate, all others separated from 
euch other by Jateral plates. Micro- structure 
of these phites ts a lose mesh of pores and 
trabelculae, with smooth raised lumps at mier- 
sections of meshes (Figs 71, 72). 

Lateral arm plates meeting broadly above 
and below; distal edge flaired considerably 
where arm spines attached. Two short und 3 
somewhat longer bluntly counded spines on 
segments oue und two respectively: segment 
three (first free segment) with 4 arm spines, 
the lower three being somewhat Jonger than 
those of segment two, ininutely spiculate und 
penily fupering, while upper spine on each side 
is greaty enlarged, 2 * length of adjacent 
lower spine (750 fi, vs, 350 my) and nearly 
twice as thick, microscopically rugose, seurcely 
tapering, and broudly rounded al lip (Fig. 73), 
One of TO sides wilh only 3 arm spines, the 
upper enlarged spine being absent, 

Three spines (rarely 4) continue on distal 
segments, the spines remaining short, blunt, in- 


164 A“. N, BAKER & D. M. DEVANEY 


SOUTHERN AUSTRALIAN OPHIUROLDEA {ie 


Pigs 49-62. 49-54, Ophiothrix (Placephiotirix) alhosteiata (holotype, SAM K215): 49, radial disc 


sexment, dorsal: 50, disc slump; 51, reptesentative arm spines, 32, Sth & 6th dorsal arm plates; 
53, 20th dorsal arm plate with pigment; 54, 9th & 10th ventral arm plates, 54-59, Ophiothrix 
(Placophiailrix) (Nwocaerulea (holotype, SAM K218): 35, dise stump: 46, radinl disc segment, 
dorsal; 57, Sil & 6th dorsal arm plates with pigment, 58, 9th & lth ventral arm plates with 
pigment; 59, uppermost basal arm spine, G0-62, Qphiolhrix (Keystonea) frymenacantha (holo- 
type. SAM &217)) 60, army base and adjoining disc, dorsal; 61. Sth & 6th dorsal arm plates; 62, 
Sth & COth -veniral arm plates. Seale dines 1,0 mm- 


creasingly spiculate, and near end becoming 
denlate along one margin (Fig, 74), 

Varlation, A paratype (BPBM W1831) From 
the sare station as the holotype is the largest 
specimen (d.d. of 2.7 mm), The afms ale con- 
torted and twisted upward making the upper 
side difficult to see. However, the lower (oral) 
surface, especially the mouth region, is fully 
exposed. In contrast to the holotype, this speci 
men has the adoral shields separated In front 
of the oral shields, probably due to the fiexure 
of the upwardly directed arms, The teeth are 
visible, there being 2 or 3 below the apical one, 
somewhat blunt to slightly tapered or, in two 
cases, the upper being elongate and pointed. 
At the inner radial edge of cach side of the 
venteal arm shicld there is a small papilla Which 
is separate from vither the adoral shicld or jaw 
plate. None of the other specimens has this 


feature, An arm spine sequence of 2—3-4-3-3 
is typical on the first five proximal segments. 
Four arm spines occur very infrequently (seg, 
6, on one side of ove arm and seg, 7, on both 
sides of One arm, one side on another) but in 
no case are these enlarged like those on seg- 
meut 3, The smallest specimen (dd. 1.5 mm) 
shows only a Few sides of segment 3 with the 
fourth 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 jnter-radial oral region, It is quite 
evident that a distal tobe of each adoral shield 
separates the oral shield from the first lateral 
arm plates in this specimen. In the holotype 
this lohe is not as well developed and the oral 
shields touch the first laterals. 

Remarks; Ophiacantha shepherdi differs from 
congeners in having the combination of low 


Fives 47-48, 37, arm base and adjoining disc, dorsal, of Ophiacantha shepherd’ nm sp._thololype, 


SAM KITS; 38, same, ventral; 3942, any bases and adjoining discs, dorsal, of: 39, Ophia- 
einha hereromla (NMSZ 2071); 40, @phiacuntha brachygnatha (NMNZ 2070); 41, Ophtacanihy 
avigera (NMNZ 2072): 43, Ophlucantha aliernita (NMNZ 2087); 43, arm base und adjoining 
disc, dorsal, of Arnphlune teewcandie (NMNZ 273). 44, same, ventral, 45, tadial disc segment, 
dorsal, of Amphiara trisdeantha (holotype, SAM E212): 46, same, ventral: 47 & 46, basal, & Din 
& With am segments of 4, trivacunihe, Nolotype, Scale line 5S mm. 


166 A. N. BAKER & D, M. DEVANEY 


Figs 63-68. Ophiocomina australis (BPBM W1822): 63, oral plate, adradial view, x81; 64, oral 
plate, abradial view, x72; 65, Ist arm vertebra, x187; 66, dental plate, x117; 67, arm spine, x89; 
68, edge of arm spine, x250. 


SOUTHERN AUSTRALIAN OPHIUROIDEA 167 


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 multifid pointed disc ele- 
ments. O. alternata 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. heterotyla 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, 
pl. 52, figs 4-6; 1918: 86; 1938: 209; 1946: 
184. 

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 24 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 9-146 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 H16, 
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.iii1.1978; 1, NMNZ 2114, 
Port Hacking, N.S.W., 7 m, 30.v.1976; 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.ii1.1966—-18.ix.1967; 1, BPBM 
W1810, 8 km N of Western River, Investigator 
Strait, S.A., 45 m, 12.1.1965; 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, 24.11.1975; 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 


168 A. N, BAKER & D, M. DEVANEY 


io 


‘ 

ze 4 ? 

P Me F 

: ; 4 5 
* V 

_ 

4 a 7 
oF ae 

Koay 

> 


Figs 69-78. 69-74, Ophiacantha_ shepherdi (paratype NMNZ 3085): 69, 2nd ventral arm plate, 
x128; 70, tentacle scale x145; 71, 2nd dorsal arm plate, x108; 72, surface detail of 71, x435; 
73, uppermost basal arm spine, x145; 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. 


SOUTHERN AUSTRALIAN OPHIUROIDEA 169 


Figs 79-87. 79-83, Ophiacantha clavigera (NMNZ 2015): 79, 2nd ventral arm plate, x138; 80, 
2nd dorsal arm plate, x136; 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, x150; 87, surface detail of 84, x150. 


170 


material shows a wide southern Australian dis- 
wibutian from Colfs Harbour, N.S.W, (30° 
18'S) to Cockburn Sound, W.A. (31°S7'S) in 
1-45 m. This distribution pattern is shored by 
a number at endemic shallow water ophiuroids 
(Conecladus australis, Clarkcoma canaliculata, 
Ophiopeza assimilix, Ophiaracknella ramyayl, 
Ophiorhrix (Placophjothrix) spongicola), and 
supports the concept of a single southern taunal 
region in Ausirslian coastal waters. 


Ophiacantha alternata yaries remarkably in 
the shape of ils arm spines, The macroscopic 
appedrance of the spines ranges from smonth 
10 broadly serrate on ofe side (Figs 84, 85), 
‘The uppermost basal spines are asually more or 
less straight when smooth, although the tips 
may be slightly clavate or even bifurcate in rate 
cases, The laterally serrate spires ure, however, 
juvaniubly curved in a sabre-like {wshion (Fig. 
8S), Microscopically, the spines are sculptured 
in (wo Ways—the hasal portions being an ana- 
stomosing system of smooth longiudinal ridges 
and pores (Fig. 86), Whereas the distal por 
lions are finely rugose, the ridges having he- 
come finely but irregularly covered with raised 
Jumps Wearing small thorns (Fig. $7). When 
present, the broadly serrate spines oeeur on all 
aris; and the adjacent smaller spines are simi- 
larly serrate, but bear teeth on both edges 
father than one (Fig. 91). Were it nol for the 
absenee of another constantly different charuc- 
ter, and the fact that individuals with smooth 
spifies oecur in samples with those hearing 
serrate spines, these two forms could be mis- 
taken for separate species. In other respects 
specimens are alike, The micrasinmicture of the 
dorsal arm plates is similar to O- hereranyle, 
with line rugosities scattered over enlarged sec 
lione of the traheculae (Pigs 88, 90), bul that 
species cun be distinguished from O, glrernata 
becatise of the latter's alternatiny sequence of 
amn spines, a typical sequence being 3-3-7-8- 
5--5-)-4-6 on the proximal segments (NMV 
HI6), 

The radial shields of ©, alternata are. like 
those of Arierntyla and srepherdi, exposed only 
at theic distal tips. The shupe of these shields 
is tnustigl—they sre “sock-like” (Fig, 92) 
rather thay the log narrow and “bar-Tike” 
shields apparently typical of Ophiarantha 

Ophiacwitha clavivera Koehler 
FIGS 41, 79-83 
Ophiacantha clayigera Koehler, 1907; 247, fies 
1-3- HET, Clork 1938: 208 Cin part), 1946: 
184 (in pare). A. M. Clark, 1968: 330- 


AN, BAKER & D, M, DEVANEY 


Specimens examined: 3, NMNZ 2105, 2089, 2375, 
Seal Rock, West Id, S.A., 16-27 m, Mar. & Aug., 
1966, Feb, 1967: 1. NMNZ 2374, 11 kin off 
Glenelg, SA, 25 m, 27.01.1966; 1, NMNZ 2085, 
olf Wuldengrave [4, Gregt Australian Bight, 23 m, 
Tiw.t87l) 1, ZMH E2016 {herein designated 
lectolype), Str St, Cockburn Sd, S Channel, rocky 
bottom, 65-8 m, 30hix, 1905; 3, ZMH E6594, 
2, MNH 4540, |, WAM 4416 (herein designated 
Puralectotypes), Sta 56, Koommband Bay, 9,6 or 
11.2 km SW of Bunbury, W.A., rocky hottom 
with sparse plant growth, 14.5-L8 mi depth, 28-vi. 
1905; 1, ZMH E6123, Sty 50, Cockburn Sd, 
Southern Flats, seaweed, 3-4 m, 30.74.1905; 5, 
MCZ 4900, Koomhana Bay, W.A, po depth, 
26,.%.1929 1, WAM 17-81, 17 kim SW Bunbury, 
W.A,, 20 m depth, |3.iv.1963; 5, NMN2Z 3106, 
off Carnac Id, WA. 7.5 m, Za. 1972; 1, WAM 
$17-77, 27 km W of Cape Peron, W-A,, 35 m, 
29.V1.1977, |, WAM 18-81, ca.18 kai N of Don- 
ara, WA, 18.39 m, 16401976; 1, MCZ 4901, 
Broome, W.A., dredyed, no depth. June 1929. 

Remarks: The specimens range 1S—3.3 mm 
d.d,, with arms 4-5 * dd, Jong, and agree well 
with Koehler’s description and rather diagram- 
matic figures. The South Australian records are 


uh cxrensian of the species’ range eastward 


from Western Australia Where it has been 
reported from Cockhum Sound, Koombana 
Kay, and Broome (Koehler 1907, H. L. Clark 
1938), 

New morphological information an ibis spe 
cies is now availuble from SEM examination of 
arm plates and spines (Figs 79-83). The sur- 
fuce of the ventral arm plates consists of a 
series of browd, stoooth, transverse “shelves”, 
beneath the edges of which are mite pores 
(Pig. 74), The dorsal arm plates have similar 
shelves, but they are somewhat crowded and 
irvepular, and their edges bear minute thorns 
(Figs SO, 81). The longest upper arm spines 
am cylindrical and smooth basally, with two 
longiludinal rows of pores running towards a 
slightly swollen aud thoroy tip (Frye. 82). The 
lower unm spines sre flattish, with a sinule row 
of pores, and stroajig tecth on their cdges (Fig. 
83). These feaures, plus a non-alternatin 
sequence of arm spines, and forked. thorny dise 
clements, charseterive GO. elavigeru- 

Ejght of the 1) specimens reported ag Ophia- 
curha elavigera by BR. L. Clark (1938) from 
Koombanu Bay, Bunbury, and one from 
Broome were tent by MCZ Vhree of the 
Koombana Bay specimens with d.d, 1,7 to 3.$ 
mm can be assigted ta GO. allernala A. M- 
Clark. They are easily distinguished from the 
other five hy having a well defined alternating 
sequence of arm spines and conical, tuberculate 


SOUTHERN AUSTRALIAN OPHIUROIDEA 


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 QO. alternata 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 
41, 82). 

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 O. 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 


171 


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 dd. 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 brachygnatha H. L. Clark 
FIGS 40, 93-98 
Ophiacantha brachygnatha H. L, Clark, 
420, figs 123a & b; 1946: 186. 
Ophiacantha abyssicola G. O. Sars var. ofagoen- 
sis Fell, 1958: 25 pl. 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°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.iii.1972; 8, 
WAM 235-78, + 3, BPBM W2597, 33°30’S, 
114°31°E, W of Cape Naturaliste, W.A., 250- 
237 m 15.11.1972; 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. brachygnatha 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 


1928: 


172 A. N. BAKER & D, M. DEVANEY 


> 


5 
} A 
Y 


ee ee a 


oo 


g 


4 


a | 
vetlel teen : 
é eo “a 


Figs 88-98. 88-92, Ophiacantha alternata (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, x150; 95, 3rd dorsal arm plate 
of 2034, x100; 96, 2nd ventral arm plate of 2034, x80; 97, surface detail of 95, x100; 98, surface 
detail of 96, x100. 


SOUTHERN AUSTRALIAN OPHIUROQIDEA 


of evenly spaced perforations (Fig. 94). There 
is Sometimes one large tooth on the abradial 
side of the largest spines, situated at about 3 
of the spine length (Fig, 93). The external sur- 
faces of the dorsal and ventral arin plates (Figs 
95-96) ure covered with perforations and Lra- 
beculae beating sharp points and smooth 
lumps, respectively (Figs 97-98), 


Ophiacantha abyssicola Yar, otagoensis from 
New Zealand, is a synonym of this species, but 
O. abyssicola s. str, appears to differ mainly in 
the characteristic cross-shape of its oral shields, 
compared to ibe simple triangular shields of O. 
brachygnatha, 


Ophiacantha yaldwyni Fell 
Ophiacaninha yaldwyni Fell, 1958: 23, pl, 4, flgs 
F, HJ, 

Specimens examined: 1, NMV H361, 38°24,5°S, 
149°25,5'B, off Pt Hicks Vic. 823 m, coral rock 
21,x7,1973; 2, NMNZ 2658, 42°29°S, 173°37'E, 
Kaikoura Canyon, N,Z,, 1097-1006 m, 18-11-1976. 
Remarks: This species was previously knowo 
from a single specimen taken in Cook Strait, 
N.Z., at 1005 m (Fell 1958), and the Victorian 
record js thus new for Australia, The specimens 
are shghtly smaller (6-9 mm dd,), but agree 
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, 0 
yaldwyni is otherwise characterized by a 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 1s fegred lost. 


Key to southern Australian species of 
Ophiaeantha* 

{, Dike covered dorsally by slonder, multifid 

spinelets 


TDise covered eae bY granules 0 or 
spinules ive 3 


w 


2, Dorsal arm plates triangular; Jongest 
armspines clavate QO. elavirera 


Dorsal arm plates bell-shaped, longest 
armspines evenly tapered 
PACES iia tear? fe be z O. hrachygaatha 

3. Oise covered Jorsully by granules... 4 
Disc covered dorsally by spinules  . 5 


173 


4, Disc granules as wide or wider than 
high, blunt; first uppersaost armispine 
enlarged ' O. shepherd 
Disc granules as high or higher ihan wide, 
with terminal thorns, uppermasi 
armspines of similar size. QO. yaldwyni 


§, Disc with pointed spinules, enlarged 
inter-radially wear disc margin; arm 
spines ae inallte in number 


0. alternata 


Dise with pointed spindles enlarged 
néur dise centre; armspines sat 
wlternating in mumber OF hererotvla 


Oplioprium rosea (Lyman) n. comb, 


Ophiacantha rosea Lyman, 1879: 139, pl. X, digs 
287, 268) L882: 184, pl, 25, figs 10, 11— Fell, 


1958: 23, pl, 4, fies J, K.-McKnight, 1967: 
308, 

Ophiacantha truncata Koehler, 1930: 62, pl VI, 
figs 13-14. 


Specimens examined: Holotype, BM(NH) 82,12 
23.175, SQ? ID'S, 74°42'W, S Indian Ocean, 320 
m, S5J.1876 (Chalieneer Sth 308); 11, NMV 
H364 & 1, NMNZ 2654, 39°44.5'S, [48°49'F, 
Bass Strait. 641) m, 24.x1.1973; 2, NMWNZ, 2655, 
43°14'S, 173"39R. Pegasus Bay, NZ, 1006- 
SIZ m, 27.1x.1976; |, ZMC, 38° 15'S, 149°20'E, 
S of Cape Everard, Vic,, 270-470 m, 19.7x.1914 
(holotype of Ophiacantha truncata Koehler), 


Remarks; This robust, brightly coloured species 
has been recorded from New Zealand by Fell 
(1958) and McKnight (1967), and southern 
Chile, Japan, and Crozet 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 % did At their bases, the 
arms of 4 12 mm d.d, specimen are 4 mom high 
and 3 mm wide, with 10 arm spines up to 4 
mm long, The arms therefore have a very 
robust appearance. The dise is covered entirely 
with small thorny stumps up to 3 ™ higher 
than wide. There are 18 oral papillae on each 
jaw, © internal and the remainder external on 
each side, The internal papillac arc rather spini- 


* Other species recorded from sowlhern Australian 

waters und referred to Ophiacuntha hy WoL 
Clark (1946) are here regarded ay belonging 
ta different genera, viz: Ophiacantha twrhneata 
- Ophioprium rosea (Lymant; Opliticaritha 
fidelis — Ophiomitrella fideliy Koehler @plhia- 
caniha coneesta = Ophialeaea congesta (Koeh- 
ler); Ophiacanthe valenciennesi Ophioireta 
valeneiennest (Lyman )- 


174 ALN, BAKER & D. M. DEVANEY 


form, Whereas the external ones are spatulate. 
The oral shield as of un unusual bell-shape, with 
the apex of the bell pointing distally, There is 
one very large paddle-shaped tentacle scale on 
each pore excep! the first, where there are 
ooeasionally two (as tn the holotype), 


The concealed radial shields, numerous oral 
papillac generally of two distinct types and 
position, two basal tentacle scales, and follow- 
ing single scale of very large size, indicate that 
this species should be placed in Ophlopricwn 
H, L. Clark rather than Ophiacaniha, To other 
respects, such as size, form and sculpturing of 
arm spines, and the nature of the arm plates, 
O. rosea closely resembles the two large species 
of Ophioprium (OQ, larissae and O. kapalae) 
known (rom the Tasman Sea (Baker 1979). 
O. rosea is distinguished by its red colour, short 
arms, and the shape of its oral shields une ten. 
tacle scales, 


Ophiacantha truneate Koehler, known from 
two specimens from just northeast of Tas- 
Mania, is here regarded us a junior synonym of 
QO, rosea. The holotype hus identical dise 
stumps, arm spines, arm plates, and (ral 
shields, and up to 17 oral papillae of two kinds. 
The five arm bases have only one large tentacle 
scale to cach pore indicating that the preseace 
of more than One scale is not aw stable charae- 
ler, 


Ophioplinthaca Vervill, 1899 
Ophioplinthaca ineisa (Lyman) 
Ophiomitra inevisa Lyman, 1883; 263, 

fizs 89% 90. 

Ophioplinthaca ineisa Verrill, 1899: 351— H. L- 

Clark, I9f5: 211, 

Specinens examined: B NMV Hi6l, 38°24.5'S, 
149°25.5°F, Sof Prt Hicks, Vie, 823 m, rock and 
coral, 21-x1.1973. 

Remarks; 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 bathymetric range 
of 610-1572 m, suggests however, that it may 
well be Widespread like some other deep con 
linental slope ophiuroids. 

The largest specimen is 17 mm d.d,, and ope 
has what is possibly a dwarf male attached to 
its aboral surface, The Specimens match 
Lyman's description elosely, and the range of 
sizes shows small variations; the arm spines 
Vary from five to seven, and the dise enver 


pl V4, 


Varies from smooth stumps at the edge to 
thorny stumps near the dise centre, 


Ophluplinihaca is cosmopolitan on the con- 
tinentul shelf or deeper, and contains ahout 30 
nominal species. No representative has hitherta 
been reeerded from Australia, although several 
are known from Indonesian waters (Koehler 
1930). Of those, QO. vicina Kochler from the 
Banda Sea, 1595 m, ts very close to this spe- 
cies, It is known from a@ single specimen, and 
appears to differ mainly in the shape of the 
oral shields aud the alignment of the external 
aral papillae, 


VAMILY Ophiactidae 
Ophiuctis Liitken, 1856 
Ophiactiy tricolor H, L, Clark 
FIGS 33, 34 


Ophiactris tricolor We 1 Clark. 1928: 427-428, 
Fig, (26, 1038: 262; |946- 20R— A.M. Clark. 
166: 326 (in key), 

Ophiacns laevis BW, L, Clark, 
14h: 208. 


Opliqetiy lnetken? Koehler, WOT: 245. 


Specimens examined: Holotype SAM K213, no 
locality dala or depth: paratype SAM K214, no 
locality data or depth | Figs 93, 34). 2, NMNZ 
2070. 3 km off West Beach, Golf St Vincent, 
S.A. 9m, 8.v. 1968: 1 NMNZ 2083, off Pr Mars- 
den, S.A, 20 m, Jan. L965: 1, NMNZ 2081, 
Yankalilla Bay, S.A. 18 m, no date; 1, NMNZ 
2078, West Id, S.A. 2 m, 20.x%.1967. 1, NMNZ 
2082, wreck of Norma, Gulf St, Vincent, S.A,. 
12 m, 14.1965; 2, BPRM W1806, Investigaior 
Strait, S.A. 30 m, wo date; 2, BPRM WHS09, 
Investipator Strait, 43m, mo date; 3, BPRM 
W825 off Glenelg, Gulf S! Vincent. S.A. 12 m, 
rovk, among sponges, no date, 2, paratypes of O. 
larvis WAM 221/2-39, Bunbury, WA, 26x. 
1929, no depth; 2, WAM 499-74 SW of Bunbury, 
WA, 20 m, 13.iv.1963; 4, NMNZ 2069, Fagle 
Ray, Cupe Naturaliste. W.A,, 1-5 m, 24ii.1975; 
1, NMNZ 1838, Yallingup, W.A,, 1-3 m, 22.1. 
1975: |. MN&G 4549, Geraldton, W.A., na depths 
|, ZME S444, Fremantle, Guge Rods, W.A., 
7-18 mm. ev, 1905; 1, WAM 319-78. W side Goss 
Passage, Abrolhos Is, W,A,, 33-35 m. no date, 


1948: 268-270); 


Remarks; Ophlactis tricolor was described by 
H-1. Clark from Sie Joseph Verco's collections 
Sredged in Gulf St Vincent and Spencer Gulf, 
South Australia. Clark (1938) mentioned a 
spesimen of ¢trivalor in) the “Melbourne 
Museum” collected in 40 fathoms north of 
Cype Borda, Kangaroo Id, S.A. but there 
appear to be no subsequent records of this spe- 
cies in the literature. The mew records listed 


SOUTHERN AUSTRALIAN OPITUROIDEA 


here show it to be distinctly southern and 
southwestern Australia in distribution, 

H. L. Clark (1938) reperted a second anc 
new species, O. laevis, from dredged samples 
in Koombana Bay (Bunbury), and Dongarra, 
Western Australia, and provisionally attributed 
two specimens (MNB 4549 & ZMH E5444) 
from Fremantle and Geraldton (reported by 
Kochler 1907 as Q. lentkeni) 10 his new spe- 
cies. Clark (1946, p, 207) separated O. laevis 
from O, tricolor by differences in the size of 
the dise scales, and radial shields, as well us by 
colour as follows; 


O. tricoler 


dise seales larger and 
less smooth. 


0, laevis 
1. dise with smooth 
coat of small 
rounded seales. 

2. radial shields very 
small, nat much 
larger than some of 
disc scales, 

_ disc usually 
purplish-rose arms 
variegated hut not 
banded. 


radial shields con. 
siderably larger, 


Ww 


dise grayish or dirty 
yellowish, arms nor- 
mally bunded with 
pinkish white. rosy 
red, and dill blue, 


Both species were characterized in Clark’s 
key by: (a) w quite large single oral papilla, 
(b) racial shields small, less than .25 dise dia- 
meter, (c) very small and numerous disc 
scales. 

The series of specimens available has shown 
thal the small differences used by Clark to 
separate laevis fram tricolor can be aliributed 
to formal variation within the one species. 
Consequently, ©. laevis is here synonyimized 
with QO. rricolar. 


Famity Amphiuridae 


Amphiura elandiformis A. M. Clark 


Amphiura elandiformis A, M. Clark, 19662 331, 
fig. 6 f-i- Bartmall, 1980; 40, fig, 13, 

Specimens examined: 3, NMNZ 1853, 43°00'S, 
148°13.6'R, off Baglehawk Neck, Tas., 122 im, 
13.11.1973; 1, NMNZ 2615, off Burnie, Tas., 
20 m, 611.1970; 1, NMNZ 2616. Great Taylor 
Bay, Bruny Id, Tas., 14.77.1972, 

Remarks: This species was previously known 
only from Pt Phillip, Vietoria, and with its dis- 
tovery in ¢astern Tasmanian waters, it may be 
expected along the entire southern Australian 
coast. The specimens range 5-7 mm dd, and 
confirm the distinctive features of this amphiu- 
rid: the dise js petaloid and covered above with 


175 


small seales, and below with naked skin. The 
radial shields are long (2.3 mm on the 7 muy 
specimen), narrow, and bowed, The distal oral 
papilla ts wide and rounded, there are two ten- 
tacle scales on each pore, find the second to 
lowest arnt spine has a proxinvally-directed 
hyaline hook or is bihamulate, 

Amphiura elandiformis is similar to two other 
southern Australian amphiurids, 4, delin and 
A, multiremula (both of H. 1... Clark) and one 
New Zealand species, A, correcta Koehler, but 
they can be distinguished by differen! combina- 
tions of the ahove features, 


Amphiura muultiremula AH. L. Clark 

Amphivra muttiremula HL, Clark, (938: 228.- 

A, M, Clark, (966: 336, fig, Ta-c. 
Specimens examined: 1, NMNZ 2617, West Id, 
S.A,, 2m, 29,11975; 1, NMNZ 2618, Eagle Ray, 
WA, 1 m, 2411-1975, 
Remarks: This amphiurid was hitherta kriews 
only from 3 Port Jackson and Lone Reef, 
N.S.W_. specimens. The present specimens are 
4.0 and 5,5 mm did, and the largest has 9 arm 
spmes, This large number of arm spines, and 
their Mat, blunt shape, help to characterize this 
species, which is otherwise distinguished From 
the many other Australian cofigeners by very 
fine, complete dise sealing, small radial shields 
(0.2-0,33 did.), and oral shields with 9 semi- 
circular proximal harder 

The holotype of A. muliremiula has been 
figured by A. M. Clark (1966), 


Amphiura magellanica Ljungman 
Aniphinra magellanica Liywagman. L867 

Mortensen, 1924* 132. fig, 14, 

Specimens examined; 2, NMNZ 2620, 5 of Warr- 
nambool, Vic, 220-310 m, 14.¥,1969. 
Reinarky: This species is apparently circum- 
polar in subantarctic and cold temperate Jati- 
tudes, having been recorded from the south 
Atlante Oceat, Pantagoma, southern New 
Zealund and, new. southern Australia. 

The two Warrnambool specimens measure 
2.4 and 3.7 mm dd) They show the eharae- 
tense lengthening of Lhe lowermost of the 7 
arm spines, and the lurge leaf-shaped distal aral 
papilla and tentacle scales. Amphinra magel- 
lanica 18 Viviparous and hermaphroditic—feu- 
tures which separate the species from the simi- 
lar A_ ypinipey Martensen from New Zeuland- 


320 


Amphitra trisacantha H, L, Clark 
FIGS 43-48 


Amphiura trisacanthie H, L. Clark, |928: 
figs 125 a & b, 


415, 


176 


Specimens examined: Holotype, SAM K2}2, 
Spencer or St Vineet Gults; S:A., no depitly ur 
date; 1, NMNZ 2073, Upper Spencer Golf, 5A, 
9 m, [144.1973; 1, NMNZ 2619, North Ann, 
Westernport, Vic, 4 m, 13.VL1976. 


Remarks: This species was hitherto known only 
from the holotype. The new material measures 
2.5 and 6.5 mm d.d,, and although all the arms 
are broken, fragments with the largest specimen 
indicate arms 4—5 * d.d, The arm spines num- 
ber 5 at the arm bases, and reduce to 3 within 
sid Segments. The uppermost spines are pointed 
and slightly flattened, and the longer lowermost 
spines are more circular if cross section, and 
evenly tapered to a sharp point. 

In other respects the specimens are in close 
agreement with the holotype and Clark's 
description. Clark's photograph of the holotype 
does not illustrate the species adequutely, and 
we therefore provide camera lucida drawings 
of the now damaged type specimen (Figs 45— 
48) and the one from Upper Spencer Gulf 
(Figs 43 & 44). 


FAMILY Optnotrichidae 
Ophiothrix Miler & Troschel, 1840 


Ophiothrix (Placophiothrix) albestriata Hy b 
Clark 


FIGS 49-54 


Ophiathrix atbostriata H, 1 Clark, 1928: 429, 
fig, 127, 


Placophiothrix albestrigga AH. L. Clark, 1946; 
227. 
Ophiothric (Placoplilothria) alhosirlann AL MM. 


Clark, 1967; 648. 


Specimen examined: Holotype SAM K213. Cireat 
Greal Australian Bight, no depth or date. 


Remarks: This holotype (d.d. | mm) rernnins 
the only known specimen and we ive here 
drawings of various parts of it to suppiement 
Clark's description and figure. The specimen |s 
now very faded, but two parallel Iines are 
visible on beth dorsal and vewtral surfaces of 
the arms ufter about the 20th segment. This 
species is similar to O, (Placophiothrix) spongi- 
cola ja general appearance, but diflers m the 
shape of its dorsal arm plates and oral shields. 
The dise 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 @. (Placuphiolhrix) spongicota 
encompasses thase im albestriata, The disc 
cover of OQ. albostriata is wiso like that of O. 
(Placephiothrix) lineocaerulea H, L, Clark 


A. N, BAKER & BD. Mo DEVANEY 


(holotype SAM K2L8) (Figs 55-59). The 
dorsal and ventral army plates (Figs $2-54, 57 
& 58) are, however, quite distinct 1 these two 
species, and ©. lineorwerulee is known only 
from northern and eastern Australia, from 
Broome to Moreton Bay, 


Opliethrix (Keystonea) ymenacansha H. 1, 
Clark 
FIGS 60-62 
Ophiorlirix hymenacantha H, L. Clark, 1928; 434, 
fig. 128, 
Ophlotrichoides hymenacantha: H, L, Clark, 1946; 
233, 
Ophiothrix (Keystened) fhymeneeanthia A. Me 
Clark, 1967; 648. 
Specimen examined Holotype. SAM K2L7, Great 
Australian Bight, no depih ar date. 


Remarks: This species is known only from the 
8 mm holotype. H, L, Clark (1946) inferred 
that the specimen might he from the north- 
western coast of Ausiralia, eather than the 
Bight, but there is no information on the label 
to siggest this, and in the ahsence of new 
material We must ussume it is a southern form, 
There are of course other “northern” ophiu- 
roids in |he Great Australian Bight and South 
Australian gulfs (e.2, Astrebod ernae Déderlein 
& Amphloplus eclraleucea {Brock)), and tt 
imiv Well be that if this is a representative of 
the tropical subgenus Keyyronea, it has evolved 
in the relative isolation of (he southern region, 
In many ways, however, Ayrmenacantha 
reminds us of an aberrant O, (Placophiorhrix) 
spungicola—it has the same shaped arm spines, 
arm plates, und oral shiclds, and we have seen 
sporngicola specimens with a naked dorsal disc 
surface, We have as Well encoulitered webbed 
arm spines in small specimens of the sevcral 
Ntindred! \pongicola examined: Up ta.a dd. of 
3 mm, specimens of G, spongicola from the 
Abrolhos [s, W.A.. and Investigator Strait, 
S.A., reveal, on segments under the disc, proxi- 
mal arn spine webbing, as well as Upper arm 
plates With scattered low thorny slumps, Larger 
specimens, however, Jack these apparent 
juvenile characters, In view of that last aet, 
and the very distinctive colour pattern of O, 
spongicale (see Baker 19881), we consider that 
the ecolotirless, enigmatic @. hymenaeanita 
should be retained in the subgenus Kepyronee 
by virtue of its naked upper disc surface, at 
least until a growth series is found which indi- 
cafés another relationship. Figures 60-62 show 
diagnostic features of the type specimen, 


SOUTHERN AUSTRALIAN OPHIUROIDEA 177 


Acknowledgments 
We 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 Zeidler (SAM), Loisette Marsh 


(WAM), Brian J. Smith (NMY), 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(NH), USNM, 
ZMC, ZMH, and MNB. 


References 


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). 


Ciark, A. M. (1965) Japanese and other ophiu- 
roids from the collections of the Munich 
Museum, Bull. Br. Mus. nat. Hist. Zool. 13, 
37-71. 


——, (1966) Echinodermata. Port Phillip Sur- 
vey, 1957-1963. Mem. natn. Mus. Vict. 27, 
289-384, 


—, (1967) Notes on the Family Ophiotrichidae 
(Ophiuroidea). Ann. Mag. nat. Hist. (13) 9, 
637-655. 


Crark, 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 Zoology. Mem. Mus, comp. Zool. 
Harv. 25, 165-367. 


——, (1916) Report on the sea lilies, starfishes, 
brittle-stars, and sea-urchins obtained by the 
F.LS, ‘Endeavour’ on the coasts of Queensland, 
New South Wales, Tasmania, Victoria, South 
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. Aust, 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. 


Fei, H. B. (1958) Deep-sea echinoderms of New 
Zealand. Zoology Publs Vict. Univ. Coll. 24, 
1-40. 

Forses, E. (1843) On the Radiata of the Eastern 
Mediterranean Pt I, Ophiuridae. Trans. Linn. 
Soc. Lond, 19, 143-145, 


KOEHLER, R. (1907) Ophiuroidea. Jn W. Michael- 
sen & R. Hartmeyer “Die Fauna Siidwest-Aus- 
traliens. Ergebnisse de Hamburger siidwest- 
australischen Forschungsreise 1905” 1(4), 
241-254, Fischer, Jena. 


. (1922) 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 pls. 


LuuNGMAN, A. (1867) Ophiuroidea viventia huc 
usque cognita. Overs. K. danske Vetensk. 
Forhandl. 1866 (9), 303-336. 


LUTKEN, C. F. (1856) Bidrag til Kundskab om 


Slangestjnerne. II 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. 
K, danski Vidensk. Selsk. Skr. 5, 179-271, 5 pls, 


, (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 pls. 


, (1882) Ophiuroidea. Rep. scient. Results 
Voy. ‘Challenger’ (Zool.) 5, 1-386, 46 pls, 

. (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 
1880, 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 pls. 


McKnicut, D. G. (1967) Additions to the 
echinoderm fauna of the Chatham Islands. 
N.Z. Jl Mar, Freshwat. Res. 1, 291-313. 


MortTeENSEN, T. (1924) Echinoderms of New Zea- 
land and the Auckland-Campbell Islands, U 
Ophiuroidea. Papers from Dr Th. Mortensen’s 
Pacific Expedition 1914-16. XX Vidensk, Meddr. 
dansk, naturh, Foren 77, 91-177, 4 pls. 


178 A. N. BAKER & D. M. DEVANEY 


MULLER J. & TROSCHEL, F. H. (1840) Uever die Verritt, A. E, (1899) North American Ophiu- 
Gattungen der Ophiuren. Arch. Naturgesch. 6, roidea. I. Revision of certain families and 
327-330. genera of West Indian ophiurans. Trans, Conn. 

: Acad. Arts Sci. 10, 301-371, 2 pls. 
Pawson, D, L. (1969) Astrothrombus rugosus Witkie, I, C. (1980) The systematic position of 


Clark, new to New Zealand, with notes on Ophiocomina Koehler and a reconsideration of 
Ophioceres  huttoni (Farquhar), Hemilepis of certain interfamilial relationships within the 
norae (Benham), and Ophiuroglypha irrorata Ophiuroidea, pp. 151-157. Jn Jangoux, M. (Ed.) 
(Lyman) (Echinodermata: Ophiurodea) N.Z. ‘Echinoderms: Past and Present’. (A. A. Bal- 


Jl Mar. Freshwat. Res. 3: 46-56. kema, Rotterdam). 


THE CONCEPT OF AN ‘EDIACARAN’ PERIOD AND ITS 
STRATIGRAPHIC SIGNIFICANCE IN AUSTRALIA 


BY R. J. F. JENKINS 


Summary 


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 biostratigraphy. 
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 skeletal-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. 


THE CONCEPT OF AN ‘EDIACARAN PERIOD’ AND ITS STRATIGRAPHIC 


SIGNIFICANCE IN AUSTRALIA 
by R. I. Fy JenkIns* 


Sunimary 
Joneis, RoI. B. (1981) The concept of an ‘Edisearan Period’ and its stratigraphic sjeniticance 
in Australia. Trans Ay Soe, 8. Ast W504), 179-194, 11 December, (Y¥81, 


A 20 veor cycle of ideas bas led to a concept of an ‘Ediacaran Perjod’ embracing [he 
geological time interval characterized by the soft-bodied metazoan assembliges of te ferminal 
Precambrian, Continuing finds of such assemblages emphasize their potential value far biw- 
stratigraphy, Geochronoloyicul studies suggest thal ihe earliest finds may be at about 640-620 
Muy metazouns appear to have become diverse and relatively numerous during the interval 
600.570 Ma, Evidence from central England establishes that the Ediacaran does pot overlap 
ihe Cambrian and that separate sequential depositional events represent these rime ilervats. 
The Edivcuran is Precambrian in aspect, without the significant skeletal-bioclustie component 
typitying Ihe Palagozoic or Phanerozoic. Studies of Harland & Herod (1975) and Gluessner 
(1977, (979) liok the Ediaciran with the postylacial part of the Proterozoic Adelailecan 
succession. und by implication indicate a type area withity the Flinders Ranges, South Aus- 
tralia, A Strutetype for the Ediacaran is nominated at Bunyeroo Gorge, and comprises the 
Wonoka Formation and Pound Subgroup, the age of the Wonoka Formution being sugested 
LO approximate the lime of appearance of meguscopic metazouns. The exposure of the Rawns- 
ley Quartzite of the Pound Subgroup in Banyeroo Gurge is made the stratntype of a Late 
Ediscaran Ppoch reflecting the mujor evolutionary radiation of surviviog invertebrate phyla. 


Introduction 

Gilaessher (1959) compared finds of carly 
soft-hodicd metazoans al Ediacara, South 
Australia, with) similac fossil remains rem 
Charnwood Forest, Leicestershire, England, 
and the Nama Group of Namibia (South 
West Africa), and established the age of these 
assemblages aS late Precambrian. These finds 
are now justifiably famous and the known 
record af compurable fossil remains is 
becoming inercasiligly extensive with some 1% 
or so occurrences widely sited about the globe 
(eg, Glaessner 1979b, fiz, 2). Though the 
organs often atttained lange size and show 
a general lack of hard parts, their tussil 
imprigts show no evidence of macrophagous 
predation, and this, coupled with rapid burial, 
was of ugdoubled importance for their pre- 
servation (Wade 1968; Glaessner 1979b). The 
kinds of organisms cepresented include mecdu- 
soid coclenterates, a member of the conulata, 
chondtophore hydrozoans, representatives of 
sea-pen-like animals, several kinds of worms, 
primitive arthropods and the problematic 
lophophorate Tribrachidiun: Glaessner. Other 
curious sack-shaped organisms and enigmatic 
forms composed of chambered vanes are reter- 
able to the new Phylum Petalonamae Pflug. 


* Centre for Preciimbrian Research, University of 
Adelaide, G.P.O, Box 498, Adeluide, 5, Aust. 
soo). 


Amongst students of these fossils there is 
broad consensus that the remains rellect a 
phase of accelerated animal evolution preced- 
ing the Cambrian (Glaessner 196], 1971 
1972, 1977, 1979a,b; Glaessner & Waller 
1981; Cloud |976a,b; Sukolay 1976a,b, 1977; 
Stanley 1976a.b; Sepkoski 1978, 1979; Ford 
1979, 1980b; Lowenstam 1980), ‘The interval 
during Which these organisms lived is variably 
Tecogmized by Western workers as the 
“Ediacarien” Stage (Termicr & Termier 
1960) or as either the Ediacarian (Cloud 
1972, 1976a,b) or Ediacaran Period (Harland 
& Herod 1975; Gluessner 1977, 1979b). The 
approximately equivalent interval is generally 
termed Vendian by the Soviets (e.g. Sokoloy 
1973; Keller 1979). The broad relationship 
between the Ediacaran and Vendian is ini- 
cated in Glaessner (1979b, fig. 1), 


Of recent years, diverse new assemblages of 
Ediacaran fossils have been located in north- 
ern Russia mm parts of the Redkino substage of 
the Valdai Group, on the Gnega Peninsula 
and the Zimmy Bereg (Winter Coast) of the 
White Sea (Keller & Fedonkin 1977. 1978, 
1980a, L981). As well as these having several 
taxa in commoan, some 10 or 11 species which 
variably occur in these assemblages are alsa 
present at Edjacara and other fossil sites in the 
Flinders Ranges, South Australia (Fig, 1). 
This finding fed Keller & Fedonkin (1976) 


180 
x GH » 
‘ LHEER ~~ 
t eh 
a | ‘ 
, r 
= u j 
he, 
e' = , 
iil Sen = f 
a (pF —h Se oe 
DR | 
wy j a 
A al 
Fil 
i 
} ‘ f 
ii F% 
; — 
3 
up Seer . ; i | 
tr FA, \ 
hy . 
ry \ 
4 | 
n MA, 
= in r i= \ 
ny EDIAC AT AN I Dey -\ 
So - STRATOTYPE. 4 ee 
“ = ~ .—- ~ 
«| Fa f 
r + 
< ' , f 
2 
~- ~ £ 
of 2 
" t | 
- AN ah 2p HORTAFRA ter onion 
SD ety — Hr roRy 
ball TT aAWREM —— Pr kis 
fl ff "Ff ——— 7 at sa") 
h a: a 
j f 
b ; ‘ =H Vpetear 


/ eb ® 
rQi! ; —_ . “ fi 4 


f t i H 
| 


} ‘a 
* GUOPN ' toon nas var iat q i 
i ed Ab a a \ f 
i 4 —— 
Fie, | Edisearan localities discussed in text. 


Larger map shows greater part of Flinders 
Ranges with outcrop of Pound Subgroup indi- 
toted by areas of stipple. Occurrences of Edia- 
cara assemblage are marked ‘F’. New occur- 
renees ure from work of Mr Colin Ford, Mr 
Jim Gehling and Dr Brian Daily, Base map alter 
Jenkins & Gehling (1978). 


and Fedonkin (1977, 1981) to suggest that 
the northern Russian sites and the localities in 
the Flinders Ranges are closely equivalent in 
time. Several of the shared taxa have been 
considered as constituting an Oppel zone 
(Rozanov & Sokolov 1980). A number of 
other pairs of sites share vither one or more 
species or have taxa showing resemblances 
suggestive of an evolutionary affinity. “Medus- 


THE CONCEPT OF AN EDIACARAN PERIOD 


eids” are virtually ubiquitous. Thus it is 
upparent that all the finds are broadly related 
in geological (ime, 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 biostratigraphy 
necessilates the nomination of stratotypes for 
the time divisions involved, Time-rock divi- 
sions 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 
Gorges, Hubei Province, South China, or the 
Marinoan ‘Scries’ (Mawson & Sprigg 1950). 
measured on rocks cropping out immediately 
south of Adelaide, South Australia (Thomson 
1966), have useful regional application for 
purposes of intracratonic correlation, The 
modern usage of the Marinoan in this sense is 
Jiscussed by Thomson (1964, 1969), Historic- 
ally, these divisions have priority over subse- 
quently designated time-rock terms. However, 
the intervals of time embraced by them are 
incomplete or rather broad and their correla- 
tion between different land areas remains 
essentially arbitrary, 

Current usage of the Vendian is subject 10 
controversy as to its stratigraphic beginning 
(Keller 1979), As yet no stratotype has been 
nominated for the Ediacaran, which however, 
is linked geographically with the Flinders 
Ranges in name and in its associvted ussem- 
blage. This paper attempts to order geological 
and evolutionary events in’ the terminal 
Precambrian and examines potential strato- 
types for the Ediacaran. 


Concept of an Ediacaran Period 

Termicr & Termier’s (1960) erection of an 
“Ediacarien” Stage met with little support, the 
fossil remains being known essentially at only 
three world localities at that time. In a fater 
work the same authors (Vermier & Termier 
1967, p, 141) discuss “la faune d’Ediacara”, 
and subsequently they refer to the different 
world occurrences of the comparable fossils as 
‘Ja faune ediacarienne” or “des faunes edia- 
cariennes” (Permier & Termier 1968, p. 74 & 
76), Wade's (1970) notation of an “Ediacara 
fauna’ was widely adopted and used inform. 
ally to designate assemblages from a then 
increasing number of localities reviewed by 
Glacssner (1971). The tacit implication of 
this terminology was that the fossils from 


R. J. FP. JENKINS 


different arcas showed similarities and were of 
comparable age, that is, late Precambrian. 


The concept of an “ERdiacarten” Periad to 
embrace the rocks containing these carly saft- 
bodied metazoan assemblages was advanced 
by Cloud (1972. 1973. 1976a.b) who also 
reviewed the contentiaus questian as to the 
relationship between this interval and the suc- 
ceeding Cambrign, characterised principally 
by shelly (ossils, Cloud (1976a,b) made several 
proposals concerning either the maintenance 
of the “traditional” Cambrian boundary, or its 
lowering to include the Edjacaran, a passibi- 
lity also explored by other authors (e.g. Prey 
& Svilacher 1980: Rozanov & Sokolov 1980) 
Cloud further advanced the nation that the 
elymology of the divisions Phanerozor and 
Palaeavoie were such that these time terme 
should properly be considered as extending 
downwards to the start of his “Ediacarian’, 


Harland & Herod (1975) nominated an 
Ediscaran Period bounded by suggested age 
jimis of 76502 and YS702 Ma and indi- 
cated to comprise the latest part of the Adel- 
aidean, the time term used (Thomsan 1969) 
to embrace the deposrional history of the 
cumulatively measured 24 km or so of Precam- 
brian sediments occurring in the Adelaide 
Fold-Belt (Geosyncline), Harland & Herod 
placed the Ediacaran in a sequential division 
of ume intervals from oldest to youngest 
Tespectively: Sturtian, Warangian, Edhacaran 
and Cambrian, The Sturtian (Howchin 1918, 
p. 346; Mawson & Sprigg L950) js the time 
division associated with the eaclier and major 
phases of vlaciation m the Adelaidean. The 
Varangian is associated with late ice activily 
in the Precambrian (Harland 1968), and in 
the usage of Harland & Herod (1975) it is 
implicitly equated with the Marinoan phase of 
glacrogenic deposition in the Adelaidean, This, 
the Elation glaciation (Mawson 1949), is 
represented by diamichtes and Yaryiles in- 
cluded within the Elatina Formation. at the 
top af the Umbcratana Group. in the Flinders 
Ranges. Glaessner (1977,1979b) maintains. a 
comparable diviston of the late Precambrian 
to that of Harland & Herod and m tis reviews 
of pertinent georhroanogical data, tndieates 
that rocks with Ediacarun assemblayes tend to 
fall in the age range 640-575 million years. 
We further cautions that the “appropriate 
datings . . . do not suppoert any biosiral- 
graphic subdivision at the present state of our 
knowledge”, 


al 


However, workers such as Sepkoski (1978, 
1974) and Brasier (1979) extend the studies 
of Soviet students such as Sokolov (1973, 
1976a) and recognise informal subdivisions 
within the Vendian. In the work of Sepkoski 
(1978, 1979) the Vendian is approximated 
wilh Cloud's (1972, 1976a,.b) usage of the 
“Fidiacanan” and is subdivided into an 
informal four fold division of major biotic 
units, though it is admitted that current bio- 
stratigraphic control does not permit wecurate 
correlations, Other Soviet reviews of the 
Vendian (Sokolov 1876b. L977: Yuakobson & 
Krylov 1977; Keller 1979) suggest that these 
divisions reflect an oVersimplification. Oh the 
Russian Platform, Ediacaran fossils oceur 
within the Yaldai Group which Keller (1979) 
recognizes as the Yendian sensu stricto. Kellet 
(1979) considers that “there are no reliable 
radiometric data for dating the Vendian lower 
boundary” and suggests that a previously 
acvepted co, 680-650 Ma K-Ar glauconite 
date for the Wendian basement must he 
corrected. On the basis of newer dates he 
considers that that the base of the Valdai 
Group 1s about 620-600 million years old. 


The preceding short review indicates thal in 
the literature the Edjacaran has heen attri- 
huled varying sistus. These philosophical 
Viewpoints may be summarized as Follows: 


1, A majority of studies equate the Ediacaran 
with soft-bodied assemblages of subCambrian 
metuzoans which have been broadly termed 
the Ediacara fauna, Glacssner (1977, 1979b) 
indigates thal the rocks contaiming these 
remains fall into the approximate age range 
of 640-570 Ma. 


2. A minority of students attribute to the 
Fdiacaran the significanee of a hiostrati- 
graphic slage, or sugges! What at least in part it 
canstitutes an assemblage zone. 


3. A majority of Studies imply or indicate that 
the Ediacaran is Precambrian, and represent- 
ing the latest part of that Erathem. 


4. A minority of authors consider either that 
the Ediaearan validly comprises part of the 
Cambrian or that the lower boundary of the 
Cambrian should be extended downwards to 
embrace the Ediacaran, 


5, A minority of students imply that the Fdia- 
caran is of Phanerozoic and Palaeozoic aspect 
and gdvocate its inchisian within these div)- 
sions. 


182 THE CONCEPT OF AN 
6. Several workers (Harland & Herod 1975; 
Glaessner 1979H) allribuie the Edinearan 
lime-rock status, indicating it ie represent (t 
part of the Upper Proterozoic and a division af 
the Adelaidean therehy nominally restricting 
its lype-area to the Adelaide Fold-Belt, By 
implication, the lower boundary of the Edia- 
varan dies above the Marinoan placngeni¢ 
event recognized in this region. 

7, Some recognise informal divisions within 
the Ediacuran, these divisions — variably 
numbering between two and four, 

8. A elpse equivalence is recognized between 
the Ediacaran and Vendian, 


Discussion 
General 


As the Edisearan hay been atirihvired 
several different philosphical meanings the 
fairest approach in altempting to nominate a 
stratetype might be to choose & section which 
best illustrates concepts held by the majority 
of ecarher students. This approach does not 
imply direct maintenance of priority Certainly 
those students who altmbute a time-rock 
meaning to the term (Harland & Herod 1975; 
Glaessner 1977, 1979b) come closest to 
peoviting an adequale definition, The second 
and major requirement of the stratotype is 
that it should cneompass or be bounded by 
distinctive geological phenomena which pro- 
vide the greatest potential for correlation, and 
gonststent with the broader interpretation of 
the nominate division, 


The idea thal the Ediacaran can be con- 
sidercd as a short downwards extension of the 
Lower Cambriati is pot supperted by eeo- 
chronological dala A U-Pb zircon-date 
froncordia upper interkept model) of 
620420 Ma (Glover & Shinha 1973) for a 
felsic qdulf-hreceia in near association with 
worm-like impressions from North Carolina 
(Cloud ef a/, 1976) is perhaps the oldest well 
founded age linked with any of the Ediacuran 
fossils. Older datings have been well publicised 
(see reviews of Glucssner 1977, 197%) hut 
are not closely consistent with those obtaining 
for the majority of finds. 


The first Ediacaran assemblage actually 
discovered (Gijrich 1929, 1933; Germs 1972, 
1974) occurs in the lower part of the Nama 
Group in Namibia. The Nama Group is 
placed in stratigraphic perspective by Kroner 
(1977) and formally subdivided by Kréner 


EDIACARAN FPERION 


el al, (1980), Kroner er al, (LY80) indicate 
that i part che Nama Group rests unconform- 
ably on & sequence containing voleanic rocks 
which give a possible metamorphic resetting 
age of HA6S32 Maj elsewhere the Nama is 
intruded by granitoid rocks dated al 553213 
Ma (Rb-Sr whole rock isachron ages; Allsopp 
etal, 1979) decay constants of Steiner & Jager 
1977), Rb-Sr dating of shales in the lower 
Nama Group confirms a late Precambrian 
age and Kroner (1977) brackets the Kuibis 
Subgroup, the earliest pant of the Nama and 
major fossiliferous interval, between c, 607% 
and ¢, 636% inillion yeaty ¢*superscript iuli- 
cales correction of data to constants of Stemmer 
& fuger 1977) Reviews of age data in pre- 
paration by the wriler suggest younger aves 
for other Western ocecurrenees of Edigcaran 
fossils. The possible lower age of 620-600 
Ma for Soviet occurrences (Keller 1979) has 
already been mentioned, 


In their assessment of ape data relating bo 
the Edivearan and Cambrian, Cowie & Cribb 
(1978) find that the base of the Cambrian 
may be bracketed between 590 and S60 Ma 
Kroner er al. (1980) suggest that the base of 
the Cambrian be recognized at 560 Ma. An 
increasing hody of new radiometric data from 
Englund suggests that this) boundary may 
actually be younger again, In central western 
England, Early Cambrian strata of the “Non- 
Trilobite" Zone and the “Olenellid” Zone rest 
unconformably on  inetamorphosed rocks 
Jaled by Rb-Sr studies at 536-8 Ma. with 
reluted granophyric intrusion at 533413 Ma 
(Patehett et al. 1980 & included references), 
A similar Rb-Sr isochron age of 540°+58 
Mu was obtained hy Cribb (1975) from 
diarites intruding the southern part of the 
Charnian Complex, The Early Cambrian Hart- 
shill Portiation ties uncontormably above a 
comparable diorte and the upper part of this 
Formation contains a shelly assemblage con- 
sidered equivalent to the Dokidecvathuys revu- 
lariy zone of the Siberian Tommotian ( Brasier 
eral, 19785 Brasier & Hewitt 1979); eloments 
of the lowest Tommotiun zane are also present 
“although largely reworked" {Brasicr et al 
1978), Further work on the dating of the sub- 
Cambriin of England is proceeding (Beckin- 
sale eral, JORV). 


The yeochronological data just reviewed 
continue ta support Glaesstier’s (1977. 19796) 
finding that rocks with characteristic Edia- 
caran melazoans may occur as early as c. 640 


R, J, F, JENKINS 


Ma, However, the few items of data fall about 
aomenn of © 620 Ma. which may be « slightly 
finner uve for the earliest known metuzouns. 
The terminstion of the Ediacuran may lie at 
c. 560 Ma or possibly as young as ¢ 530 Ma. 
These estimates indjcute a possible maximum 
uration of c 110 Ma for the deposiion ot 
cocks with Ediaewran metazeuns, and a ovini- 
mum duration of ¢, 60 Ma, Either of these 
estimates are comparable with the likely dura- 
von uf 55 to 95 Ma for the ‘traditfonul’ 
Cambrian (based of) data of Cowie & Cribb 
1978) and indicate that in a hierarchy of geo- 
logical time divisions the Eqjacuran and 
Cambrian musi be accorded equal status. 

The pecurrence of a reasonably diverse 
Ediacaran assemblage is well docunrented in 
the middle part of the Charnian, or the Wond- 
house Beds of the Maplewell Group (Ford 
L958, 1980a; Brasier er al. 1979). However 
following deposition of the Charniun succes: 
sion, folding, metamorphism, intrusion anc 
deep crosion oecurred priar to a pew ans- 
gressive event heralding the start of the Early 
Cambrian (Brasier 1979, 1980: Brasier & 
Hewitt 1974). Any estimate af the duration 
of this sub-Canibrian orogetic event and the 
subsequent cyele of uplift and erosion must be 
speculative, but an interval of 10 to 15 Ma is 
reasonable, This tectonic punctuation between 
the deposition of rocks wilh Ediacaran fossils 
and true Cambrian sediments, which can be 
recognizably dated biostratigraphically as 
Tommotian (Brasier er al, 1978), establishes 
beyond reasonable doubt that the time interval 
represented by Ediacvaran sediments and the 
time of Cambrian deposits are lwo quite 
separate geological cycles sequential in Barth 
history. This finding is also supported by the 
stratigraphy of the Adelaide Fold-Bell, 


Jum nol in agreement with Cloud (7973, 
19764.) that the ctymology of the names 
Phanerozoic and Palaeozoic provides un 
initial grounds to alter the tracitianal beginning 
of these time intervals and extend them carlier 
to include the Edticaran, The comparatively 
rare Edjacaran metazoan Temains occur im 
rocks which are otherwise little different from 
those of earlier times in the Proterozoic. Th 
contrast, sediments of the Early Cambrian 
transgression (Daily 1972; Brasier 1979, 
1980) sugvest a filling of the seas by inverte- 
brates, woh frequent bioturbation (Brasier & 
Hewitt 1979; Frey & Sejlacher 1980; Fedonkin 
1981) and an increasing content of metazoan 


183 


skeletal debris, which in conjunction with 
calcareous higher algae, forms extensive bio- 
genic linestones, Comparable characteristics 
typify the remaining Palaeozoic, with skeletal 
limestones and reefal deposits abundant, 
These ¢pochiql skeletal and lithie changes 
which are ¢losely related in time to the base 
of the “traditional” Cambrian are surely ol 
ereater significance for stratigraphy and his- 
torical geology than the sporadic and frag- 
mentary earlier reverd of soft-bodied aninals, 
intriguing though this may be for our know- 
ledge of evolution, Thus there are good veo- 
logical reasons for vonsilermmg that the Edta- 
catan is properly part of the Preeambrian, and 
its significance for evolutionary studies ts that 
it contains the Precambrian precursors ef 
major phyla which only later provide the 
spectacular skeletal record characteristic of 
the Palaeozoic and the Phanerozoic as a whole, 
Biaxtraligraphy 

Virtually all) published coneepts of the 
Edjacaran embrace the wnigue, large, soft- 
bodied metazoan remains which are found in 
iis roeks. Ino this sense, the Ediacaran ts 
essentially a bjostratigraphic concept with 
similar fossils in different areas suggesting an 
approximate equivalence in Gme. Any real- 
istic stratotype should reflect this biostrati- 
graphic aspect as it polentiilly provides a 
primary framework for correlation. 

The Ediscaran assemblages at the various 
known world sites are not uniform in composi- 
tion and Glaessner (1977) remarked on this 
in relation to the usage of term “Ediacara 
fauna”, Jenkins & Gehling (1978) suggested 
that the remains of the erganisms preserved 
probably represent only part of the original 
life community, fhe barrier to the preservation 
of small organisms cffectively heing the 
grain-size of the enveloping sediment (cf. 
Glavesner (1972), Instead Jenkins & Gehling 
prefer (9 name the incomplete preserved 
community the “Ediacata assemblage’, This 
usage is only of sinict relevance to the 
remains from the Flinders Ranges or type 
area, Assemblages from other gcographic 
localities should be identified by an appro- 
Priate site or stratigraphic designation, ie. the 
‘Nama assemblage’ from Namibia ar the 
“Charnian assemblage’ from Hngland. 

The Ediacara assemblage sens stelere ts 
how well documented (Glaessner & Duily 
1959; Glaessner & Wade 1966; Glaessner 
T4980) but the description of the other occur- 


1&4 


fences is not umform. Fedonkin (1977, (981) 
is extremely active in the Soviet sphere and 
various other studies are proceedina, Enough 
information is now available to suggest that 
the metazoan remains can he divided inte twa 
broad groups. 


One group embraces the remains trom the 
lower Nama Group, the Charnian, and the 
fossils which Anderson & Misra (1968). Misra 
(1969, 1971) and Anderson (1972) docy- 
mented from the Conception Group oan the 
Avalon Peninsula of Newfoundland. The 
Nama assemblage contains the distinctive 
fossils Rarigea Gurich, Preridiniunr Gairich and 
Ernietra Plug and new reviews of these oniz- 
matic remains ate being prepared by the 
writer. Preliminary remarks concerning a 
revised structural interpretation of the sack- 
shaped Ernietta are given by Jenkins ef al. 
(1981), Preridintiunm and Ernietia tepresent 
orgamisms with a umique structural organiza- 
tion which qualifies their recognition as a new 
phylum, the Petalonamae Pflug. The Naina 
Group contains the only known world ooccur- 
rence of Preridinivm simplex Giurich, The 
structure of Rangea iy quite separate to that of 
the Petalonamae, but closely similar to as yet 
undescribed fossils from the Charnian and 
Newfoundland assemblages, and probably to 
Charnia Ford, which 1s common fo holf these 
arcas. These organisms comprise an extinct 
order of primitive anthozean coclenterates, 
and their complex, ratifying structire sug. 
gests specialized adaptation to suspension 
feeding with extreme “scavenging” capucity, It 
might be inferred that at this time either the 
plankton, or algal or bacterial detritus was 
rare, an interesting possibility in relation to the 
stratigraphic proximity of glaciogentc deposits 
in the Nama Group, and the potential infer- 
ences this has to the climate. Age dala relating 
to these assemblages (review in preparation) 
suggest a bracketing helween ¢. 64{+620) Ma 
and ¢. 600-590 Ma, An aspect of the assem- 
blages of this interval is that associated trace 
fossils are limited in diversity or comparatively 
uncommon, The evolution and radiation of the 
Rangea-like organisms and the Peralonanae 
were evidently concomitant events and may 
relate to the earliest appearance of megascopic 
metazoans. 


The second grouping comprises the better 
known and very diverse Edjacaran assembtave 
from the Flinders Ranges and now the new 
localities jn oorthern Rossia. The striking 


THE CONCEF'T OF AN EDIACARAN PERIOD 


eharacteristie of this grouping is that it 
includes many annals which cin be more or 
fess reluled to divisions aud Families still fiving 
in present day sees (Sokolov 1977; Fedonkin 
JO8L). Thus there are diverse kinds of “medn- 
soids”, chondrophore hydrozoans, sea-pen-like 
animuls, several distinctive kinds of worms, 
and primitive arthfapods. Other preblematic 
organisms are also presest. Whe trace fisyits of 
this level are diverse (Glacssner 1969, 
Fedonkin 1977, 1980ab, IYStb, bot ore 
restricted to markings made by creatures which 
moved over or burrowed horizontally im the 
substrate; the traces lend to show repelilive 
fatteros reflecting hebavioural adaptations 
towards the most ¢fficient modes of scavenging 
(ee, thigmotactic patterns), Characteristic 
genera of this time are forms of “Cvelu- 
medusa” Sprigy, Glaessnerina Germs, Diekin- 
sonia Sptige and Tritrachidium Glaessner. The 
timion Preridinum nenoxa Keller oecurs both 
in northern Russia and the Flinders Ranges 
The approximate age bracketing Indicated for 
this grouping ts between ec. AOO-590 Ma und 
¢ A70Ma. These remains can be considered a3 
reflecting an anita! radiation of — major 
surviving phyla of invertebrate life (Fedankin 
1981), 


The age data relating to these two groupings 
of assemblages are pot especially significant as 
the rehahihty af the primury information is 
probably variable and error factors are signi. 
ficunt. Certain geologicsl assumptions have te 
he made, as well, m order fo relate igneous 
und depositional events, Both the Newfound- 
land assemblage and the Charnian assembl- 
age include Chrarniedivcus which is a frequent 
genus in the Flinders Ranges, and this alone 
suggests that the possible tite separation 
between the (wo assemblage groupings is likely 
to be rather short. Stratigraphic evidence from 
the Soviet find on the Zimmniy Const (Fedonkin 
1978, TO8L) also Suggests that the time separa- 
tion berween the two groupings of assemblawes 
is short, Near the Zimniygorsk Lighthouse a 
116 on chick sequence contains abundant meta- 
zoan remains jo both [ts lowermost portion 
and wear the top of the section. Species iu the 
topmost part include Cyclomediisa davidi 
Sprige, Ovarescutue Glossner & Wade sp. 
Dickinseia costata Sprigg, D. lissa Wade and 
Tribrachidium keraldicum Glacssner, These 
Taxa are also present in the Flinders Ranges 
and are suggestive of yitthal time equivalence 
(Fedonkin 1981). ‘The tower part af the 


R, J, F& IENKINS 185 


Zimmiygorsk section still contwins Dickyrsornia 
rasta and some other forms which ate tepre- 
sented in the Flinders Ranges; these are 
Pinegia stellaris Fedonkin, Medusintites aster- 
oidey (Sprigg) (= Paliella —patelliformix 
Fedonkin --? Pretedipleirosama ruuulasnm 
Fedonkin), &diacaria flindersi Spiigg (= 
Vraviana divcijarmiy Valij, Pedookin),“Cyclo- 
medusa david’ Sprigeg and Charniodiseny 
Ford sp. {— Ramellina pennata besdankin). 
As well, this assemmblave fncludes well pre- 
served cAamples of Chernin meson’ Ford 
which hus never been located in the Flinders 
Ranges, bur is most distinctive form in the 
Charnian and in Newfoundland (slides made 
availible by Lr B. Daily). 

The rapid merease in diversity of life forms 
and coeval mulliphyletic skeletalization in the 
Early Cambrian comprise a third metazoan 
radiation event which is well documented (e.g, 
Daily 1972, 1976; Bengtson 1977; Braswwr 
1979; Brasier & Hewitt 1979). This event 
broadly coieides with the appeyrance of tbe 
constructing and tube living worms (Glaessner 
1976, 1978: Fayn & Glaessmer 1979). This 
trefisitional mterval between the very 1Aatest 
Precambrian and earliest Farly Cambrnan may 
not be represented in the Adelaide Pold-Helt. 
where there is an iniconforrmity of regional 
extent at about this level (Dinly 1972, 1973. 
1976). An important component of the truce 
lossil assemblages low mm the Early Cambrian 
wl the Flinders Ranges and also central 
Australia are vertical dwelling burrows like 
Skolithas Haldeman and  Monacrateriary 
Torell, or upright U-shaped forms such as 
Avenicolites Salter, and alo a higher level. 
Piplocraterion Torrell (Daily 1972, 19765 
Jenkins 1975a; Gauld'), dn the light of studies 
by Daily abd Jenkins in the areas just men- 
tioned the trace fossil asscriblage desenbed by 
Webby 1970) from the Lintiss Vale Beds in 
the Torrowangce Group, western New South 
Wales, is considered to be of Cambtian aspect 


Avsteiliah acchirences of Edlacaran fossils 


In the Flinders Ranges the Edincaran asserm- 
blage is widely distributed in a thin strati- 
gtuphic interval near the base of the Ruwnsley 
QQuurtzite, the uppermrest formation within the 
Pound Subgroup (Wade $970, Forbes 1971, 


AGuuld, ‘TD. (1976) Trace fossils and the huse of 
the Cumbrian at Angepensa. verthernn Flinders 
Ringes, South Australia B.Se.(Hons) ‘Thesis, 
Unveraty of Adelaide (unpublished ), 


Basin in’ the 


Jenkins L97Sb). The stratigraphy of this major 
occurrence will be amplified later, 


Elements of the assemblage have been 
reported (rom the northern part of the Officer 
possibly late Precambrian 
Punkerri Beds of the Punkerri Hills, north- 
western South Australia (Major 1974 and 
references therein), Reexamination of the 
inaterial suggests that some specimens are of 
iNorganic origin, A circilar marking may 
represent the remains of an indeterminate 
medusoid. Imprints on another block of sand- 
stone resemble parts of the frond of Charnio- 
discus. but just as likely are disrupted pieces of 
a erinkled clay film. A possible trace fossil 
resembles Gyreliihey polonicus Fedonkily from 
the Tommotian ef East Poland, Thus the finds 
in the Punkerri Hills do not provide a certain 
indication of the Ediaearaqn assemblage. but 
are not inconsistent with a late Precambrian 
to Early Cambrian age for the Punkerri Beds. 


Gluessner (1969) reported a fragmentary 
impression of a frond-like fossil collected ina 
lwose block on the Arumbera Sandstone, 
Amadeus Basin, east of Deep Well Homestead, 
S.S.E, of Alice Springs, central Australia, 
Jenkins & Gehling (1978) identified the speci- 
men as an indeterminate species of Charnio- 
discus. Further study of the locality confirmed 
ils likely placement in the lower half of the 
Arumbera Sandstone (yensv .xiricto Daily 
1974 = Aruntbera Sandstone I of Baily 1972) 
but revealed no new material. The Arumbera 
Sandstone may be broadly correlative with the 
Rawnsley Quartzite (Daily 1972), 


The medusae deseribed hy Wade (1969) 
from the Central Mount Stuart Formation of 
the Georgina Basin, near Mt Skinner, are 
Stigveyted to represent parts of an early 
seyphozoun radiahon. One form, Afallidaya 
brver? Wade, has a rather wide distrbution in 
the Georgina Basin where it is further docu- 
mented by Kirschvink (1978) and Walter 
(1980) from Central Mount Stuart; Wade 
(1469) also indicated ify occurrence in the 
Amadeus Basin. from probably near the top of 
the Arumbera Sandstone at Laura Creek just 
south of Valley Dam, W.S.W_ of Alice Springs. 
These remains are seemingly younger in age 
than the Ediacara assemblage sensu srrleta 
(Daily 1972; Glaessner 1977), The form 
identified by Glaessner (/n Fieyn & Glaessner 
1979) us Kullingia all. concentrica, and alsa 
from near Laura Creek, is apparently Early 
Cumbrian, The Mt Skinher assemblage has not 


[RG THE CONCH PT OF AN EDIACARAN PERIOD 


been located in the Flinders Ranges amt it os 
ao open question ay to whether the upper pacts 
of the Arumbera Sandstone overlap the age of 
the Rawnsley Quartaite or are slightly younger, 


Fine ctirrent ollx (Dzulinski) & Walton 
1965), identified as the pscudofossil Ariim- 
beria Glaessner & Walter, occur widely in the 
sib-Cambrian to Early Cambrian sandstones 
nf central Austraha and are known at two 
levels in the fate Precambrian of the Flinders 
Ranges. These inorganic marks are oot con- 
sidered to have any stratigraphic sinificance 

The preceding information provides a basis 
for consideration of potential stratotypes for 
an Ediacrian System, 


Stratotypes 


Marland (1974) made a detailed review of 
the extensively discussed problem of defining 
an internationally recognised boundary be- 
fween the Precambrian aod Cambrian, and 
also outlined major tines of evidence and eri- 
teria Which might serve to identify divisions 
within the Proterozoic. With respeet te possible 
stratotypes he started “Chosen successions will 
therefore be those transversing the houndary 
that are most complete and have the richest 
characters, or that are correlated directly with 
other rocks which have critical characters. and 
are well «described and casy to of avecss™, 
Compariible ideas are embraced in the gulde 
to stratigraphic procedure of Holland ef al. 
(1978), The geology of the Flinders Ranges 
satisfies some of Harland's requirements and 
provides thick, characteristic successions, 
though not without significant breuks. In the 
studies of Harland & Herod (1975) and 
Glaessner (1977, 1979b) the fidiacuran is 
linked by implication with the post-glacial, 
uppermost part of the Adelaidcan of the 
Fliuders Ranges. This part of the succession 
comprises the Wilpena Group of Dalgarna & 
Johnson (1964). The usage of Ediacaran and 
nol “Ediacanan” (Cloud 1972, 1973 1976a, 
b) is formalized, as the stem geographic names 
of other Periods sre unbroken and the endiniss 
of Perjod-names are tot uniform (ec. Jurassic 
and not Surian). The locality name “Ediacara” 
is Aboriginal in origin and is reported by 
Goyder (1860) as “Idyakra™, 


The Wilpena Group is placed in strati 


graphic and regional perspective by Thomson 
(1969), Thomson ef af. (1976) and Rutland 


et oat, CIYRly, Tes potential correlotives in 
central Australia and northwestern Australia 
are indicated in figure 4 of Coats & Preiss 
(1980), The stratigraphic subdivision of the 
Wilpens Group in the central-wesjern Flinders 
Ranges is shown in figure 2 af Jenkins e¢ ul, 
(198) ) and a modified version of 1his is given 
here (Figure 2). 


A houndary between the Precambrian and 
Cambrian is established by Daily (1972, 1973, 
1976), Ip the northern Flinders Ranges this 
boundary is the surface of disconfourmity at 
(he top ef the Pound Subgroup (previously 
Pound Quamzite, Forbes 1971) and marking 
the base of the Uralanna Formation. The 
Uratanna Formation represents a single cycle 
of Early Cumbrnin deposition and locally 
attuins a thickness of a litthe over O.5 km, 
The Uratanna Formation is overlain discon- 
formahly by the ParachiiIna Formation, and 
in the central Flinders Ranges the Parachilow 
Formation rests directly on a surface of 
efasion develaped on the Pound Subgroup. 
This erosion surface on the Pound Subgroup 
provides an upper boundary for the Edjacaran 
Stralotype. 


The characteristic Ediavaran assemblage of 
the Pound Subgroup is principally developed 
in a comparatively thin widespread interval of 
interealated silistones, fine sandstones, flaggy 
quartzite; and mediim to massive hedded 
guartzites in proximity to the base of the 
Rawnsley Quartzite (Wade 1970, Jenkins 
19756). This fossiliferous interval is termed 
the Ediacara Member of the Rawnsley Quart- 
zie snd its lithofacies development and 
environmental interpretations are discussed by 
Goldring & Curnow (1967) and Jenkins 
(1975b), The stratotype of the Ediacaru 
Member is nominated as heing Unit B of 
Goldring & Curnow (1967, fig. 5) [n Ediacara 
Range at Greenwood Cliff, Where it reaches 
a thickness of 30 m. The fossil assemblage of 
the Ediavara Member is the type example of 
the main Ediacauran biota Rare impressions 
which muy represent medusoids occur in the 
upper part of the Rawnsley Quartzite, 


In the western and southern Flinders Ranges 
the base of the Rawnsley Quartzite, character. 
ised by slightly feldspathic, clean white ortho- 
quartzites with ffequent clay pall conglo- 
merates, ts sharply erosive on pinkish felds- 
pathic sandstones included by Forbes (1971) 
in his division now recognised as the Bonney 


R. J. F. JENKINS 187 


MAL Sandstone, Jn Parachilna Gorge, 1 km south 

AJAX LIMESTONE DOLOMITES of the type section of the Pound Subgroup, 

= wat the surface of erosion below the Rawnsley 

| Quartzite is irregular in form, with undercuts, 

and has a demonstrable relief of at least LO m, 

Small, angulat, ripped up blocks of the wnder- 

SHEE lying sandstone are included in the orthoquart- 

channel sANDA| == Zites above the erosion surface and locally, 
SYNE Peds | ONAL : = “ « 

FALILTING chipped fragments of the underlying formation 

from minor gravel lenses in small downcuts of 

the erosion surface, This evidence suggests 


CAMBRIAN 


1iat 


CAMBRIAN 


= oa 

= 2 SHbUy that the Bonney Sandstone had undergone 
we & aterm partial lithification prior to the deposition of 
= a TAGOONAL the Rawnsley Quartzite and indicates a tem- 
aye) a TITLAL poral break, The Rawnsley Quartzite repre- 
3 rial sents a (ransgressive cycle of marine deposi- 
> 3 BONNEY SS LE LTAL tion with the major fossiliferous parts of the 
a Ediacara Member being tidal, back-barrier or 

ps HAL lagoonal deposits (Jenkins 1975b), 

STRAMATOLITES 

re WONDKA =u : vie Lower in the Wilpena Group the only 
< CARBONATE certain indications of metazoan Jife are small, 
o circular imprints of “medusoids” and simple 


=. |SHARNIN, SANDE trails (Form B of Glaessner 1969) located by 
Dr Mary Wade in the Bonney Sandstone 
(Jenkins ef a/, 1981), These remains indicate 
that the Bonney Sandstone properly belongs 
Within the Ediacaran, 


MEGABRERCIA 


MNCANYOINS A major deficieney of the Wilpena Group 
as an Ediacaran stratotype is that as yet there 
LTwiribatich, is No indication of the radiation of the Rangea- 


like organisms and early members of the 
Petalonomae. However higher parts of the 
Wilpena Group have not been fully explored 
for metazoan remains. With reference to the 
Soviet section at Zimniygorsk it seems unlikely 
that the Rangea-like organisms should be 
ASC RANGE G12.) MMH Hal found Widely separated stratigraphically below 
BAVLEY RANGE FM,| INTERTIDAL the main Ediacara assemblage. 


MULL AT 
The base of the Wilpena Group is marked 
by a thin, distinctive widespread dolomite 
MOORILLA EM. TIDAL Which forms a useful marker bed, the Nucca- 
PSENDO OSS leenny Formation. This Formation is mostly 
conformuble, though locally discontormable, 
ahove the partly glaciogenic Elatina Formation, 


BUNYERUO FM 


WILPENA 


metres 
1090 


SUBGROUP 


MARINOAN 


MOULUULOO FM, SUBTIDAL 


BRACHINA 


_JNUCCALEENA FM Fig. 2, Charl summarizing regional stratigraphy 

of younger part of Adelnidean succession and 

Early Cambrian in central aud lower northern 

STROMATDLITES Flinders Ranges. Only upper part of Umbera- 

TREZONA FM. SHALLOW tana Group and lower part of Cambrian 

MARINI shown, Signilicunt geological events, some en- 

vironmental indicators, and environmental inter- 

ENORAMA SH SHEL pretations are summarized adjacent to stati- 

graphic column, Purts of section represeniing 
Ediacaran System are shown, 


Lott 
ELATINA FH, UPPER bACIALS 


UMBERATANA 
GROUP 
SUBGROUP 


RT WwW 


188 


Shale datings for parts of the Adeluidean 
occuring in proximity lo (and above) possible 
correlutwe glactogenic deposits in the Kimber- 
ley Region, northwestern Australia, provide 
aves of 670484 Ma and 672270 Ma (Coals 
& Preiss 1980). The same authors review a 
dating of 676+204 Ma for an apparent 
correlative of the “Brachina Formation’ on 
the Stuart Shelf, west of the Flinders Ranges- 
Also in the Kimberley Region, a shale dating 
lor a possible correlative of the Bunyeroa 
Formation gives the number 639247 Ma 
(Coals & Preiss 1980), Qlaessner (1969) 
described the trace-like marking Burweriehunus 
delearnet from the “Brachina Formation” but 
the biogenicily of this structure is now qgues- 
tioned (Jenkins er al. 1981). An intensive 
search has been carried out in the Brachina 
Subgroup (Plummer !978a) far megascopic 
biotic nemains, but structures so far located 
are also considered to be of inorganic origin 
(Plummer 1980; Jenkins ef al 1981). The 
limited available evidence as to the possible 
age af the Branchina Subgroup suggests ir to 
he older than the 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 shect sandstone [ABC Range Quartzite) 
is considered to reflect tepressive conditions 
(Tlummer 19782). It and the overlying shales 
and siltstones of the Bunyeroo Formation are 
frequently reddish in colour, probably reflect- 
ing oxidative environments which are unlikely 
to he conducive for the preservation of either 
a microflora or higher algal renvains, such as 
are well known at about this level In the 
Vendian (Sekoinv 1977). Some greenish or 
unoxidized intervals are present (eg. the 
Bayley Range Pormation), In the southern 
Flinders Ranget and towards the east, the 
Brachina Subgroup changes to a thick green 
siltstone sequence, the Ulupa Siltstone, which 
is presumably unoxidized (Thomson 1969), 


A potential placement of the lower boun- 
dary of the Ediscaran ts at the base of ihe 
Nuccaleena Formation, which marks the termi- 
nation of the Blating glactatlon (Plummer 
1978b), The wrifer has no intention of debat- 
ing the question ax to whether Precambrian 
glaciagenic events are likely ta he either 
evichrofous or diachronoys over wide Jand 
areas and differing parts of the globe, Such 
arguments have @ cirevlar component as there 


THE CONCEPT OF AS EDIACARAN PERIGD 


is unlikely to be knowledge of the trigecring 
mechanism of ancient glactations. While the 
lithostratigraphic boundaries of glacrogernic 
intervals may be at once obyious and of great 
value for local and even regional studies fi.c. 
Coats & Preiss 1980), synehroncity of glacial 
events on a World wide basis has not been 
established. Kroner (1977) marshalled evi- 
dence favouring non-synchroneity for the late 
Precambrian glaciitions of Africa, and sug- 
gested moving centres of glacial activity. The 
Schwarzrand glaciations of the Nama Group, 
nique in being correlative with sections 
hracketed by Precambrian metazoan fossils 
(Rengea and Preridininm below; trace fossils, 
Cyclomedusa Sprigg sp, and the lost single 
specimen of Puremedusium africanum Girich 
above, Germs 19724 ef veg.) seem to be signi- 
ficantly younger than the local Elatina glacia- 
tion (Jenkins e al, 1981). Placement of the 
stratotype boundary at the base of the Wilpena 
Group negates the underlying biostratigraphic 
implications of the Ediacaran, as there are no 
known unequivocal metazoan remains in close 
proximity. The tine-significance of the Elatina 
glacistion is properly bound jn the concept 
of [he Marinoan “Senes’ of Mawson & 
Sprige (1950). Finally am association of the 
Ediacaran boundary with a glaciogenic event 
conceptually equates the interval with the 
Soviet Vendian. whieh has established priority. 
(The Marinoan has priority over both the 
Varangian and Vendian), 


A second possthle placement for the base 
of the stratotype is ac the boundary of the 
Bunyeroo Formation and the Wonoka Forma- 
tion; the Iutler contains glauconite and strama- 
tolites and is of presumed marine origin 
(Thaimson er al, 1976). One of the stroma- 
tolites ig considered to be of Vendian affinity 
(Preiss 1977). With reference to the c. 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 
sundsione may overlap or approximate the first 
appearance of megascopic melazoans elsewhere 
in the world; so far only a token effort has 
heen made to locate metazoan fossils within 
the Wonoks Formation, 


Over wide areas the Bunyeroo Formation 
pusses grudafionally up into the Wonoka 
Formation, However, in the northern Flincers 
Ranges abnormally thick intervals of Wonoka 
sediments oceur in several discordant troughs 


R. J, F. JENKINS 18u 


which Involve erosion of siltstones of the 
upper Bunyeroo Formatien and sometimes as 
much as 1,070 m of pre-Wonoka deposits 
(Coats 1964), These troughs, described by 
Thomson (1969) as “submarine canyons”, 
have a width of several km and show marginal 
breecia zones attributed to slumping. Compar- 
able geological features have been studied by 
the writer in the vieinity of Pichi Richi Pass, 
near Quorn. Here the base of the Wonoka 
Formation 1s associated with a downcut 
surface und locally Use Bunyeroo Fornyation 
is missing. The downcttting surface may 
evidence sleep slopes and al one point trun- 
vates a major part of the ABC Rane Quart- 
zitc, A megabreccia developed in the adjacent 
Wonoka Formation contains boulders with 
Bunyeroo Purmation and ABC Range Quurt- 
zite lithologies, Individual boulders teach 3 m1 
in maximum diameter and their size suggests 
lithification prior to their erosion, Boulders 
with a silty lithology show so repelilive frac- 
turing at a steep angle ta thei mrernal bedding 
and suggestive of an incipient fracture cleavage. 
This fracturing appears to be at chilerent 
nrientations in adjacent blocks and not related 
to the fracturing of the matrix material 
(sillstone), There may be some local discord- 
anee between the folding of the ABC Ranwe 
Quartzite and the Wonoka Fermation but its 
significance 1s enigmatic, The geological fea- 
tures deseribed suyvest lithification of the ABC 
Range Quartzite and Bunyeroo Formation 
sediments prior to the time al Wonoka Forma- 
tion deposition, and possible deformation 
before the strong erosive downculling process 
look place, Professor ©. C, von der Boreh is 
studying comparable geoltoyvical features else- 
where in the Flinders Rafiges. These charac- 
feristic¢s suggest that adhe ‘normal’ contact 
beiween the Bunyeroo FPermation and the 
Wonoka Fornyition may represent a paracon- 
formity (cf. Cants 1964), 

The Wonoka Formation consists of green- 
grey calcareous siltstone, minor greenish or 
purple shales, intercalated thin-bedded to 
massive-hedded grey limestone, and same 
sandstone, Pelletoids forming a component of 
the limestones are considered as possibly 
being of faecal origin. (Von der Borch, pers. 
comm), In the upper parts of the Wonoka 
Formation frequent intraformational vongle- 
merates anid Golitic limestones attest a shallow. 
water origin, Spectacular sole markings 
formed by currents are a persistent feature, 


Sandy interbeds near the top of the Woroka 
evidence tidal deposition and there are inter- 
bedded doloniitic shales, dolumites and sand- 
stones at the passage into the intercalated 
pink, feldspathic sandstones and red-coloured, 
micaceous siltstones of the Bonney Sandstone. 
Vhete is no evidence of any break in deposi- 
tion betweett these two formations. Well 
rovinded gravel lugs associated with thin inter- 
calated silts and sands in the lower parts of 
the Bouney Sandstone in some areas suggest 
a Nuvial or delttic influence. Sandstones with 
large-scale crossbedding aay reflect channel 
development. Mud-cracked hotizous, possibte 
rain-drop markings and Gther sedimentary 
structures indicate frequent emergence and 
suggest a lidal environment for major parts of 
the Bonney Sandstone (Forbes 1971). 


The potential placement of the lower 
boundary of the Edivesran at the base al the 
Wohoka Formation is advantageous for the 
following Teasons; 


1. This boundary reflects a renewed phase of 
sedimentation following possible localised 
Tectonism and associated enmive dowu 
cutting. 


te 


The Wonoka Formation is almost certainty 
murine and is known to include biogenic 
remains in the form of stromatolites, Its 
frequent greenish colour indicates that it is 
unoxidized and the variety of terrigenous 
and calcareous lithotypes offer maximum 
potential for the finding of a significant 
Mitroflara or meguseopic ilgal remains, 


3, Fluctuating changes in sea level and sub- 
sequent transgression associated with the 
channelling at (he base of the Wonoka may 
be reflected i other regions, Discovery of 
a sigmficant microflora may provide a 
biological maker for these events, 


4. The boundary between the Worneaka 
Formation and the Bonney Sandstone is 
transitional, suggesting continuous depost- 
tion, The Bonney Sandstone inchides frag- 
mentary cvidenee of metazaans- 


4. Several fragmentary lines of evidence 
suggest that the time of deposition of the 
Wonoka Formation may be mure or less 
equivalent to the first recorded appearance 
of well preserved metazoun remuins else 
where io the world, The Wentka has not 
yet been intensively explored for mega- 
scopic fossils. 


190 THE CONCEPT GF AN 


SS MT. RUPERT AA 


_ 


Fie. 3. Locality mup showing Edijacaran strito- 
iype, between arrows, and pccess trucks, 


6. Placing the Ediacaran boundary at the 
beginning of a formation which is conker 
euous with the Pound Subgroup preserves 
the concept which equates this Period with 
the occurrence of early fossil metazoans. 


Accordingly the stratotype for an Ediacaran 
System is formally nominated as comprising 
the whole of the Wonoka Formation and the 
Pound Subgroup, The type section is nomi- 
nated as being at Bunyeroo Gorge (Figures | 
& 3), where the above stratigraphic inlervals 
are excellently exposed and have a combined 
thickness of about 1320 m, Bunyeroo Gorge 
ig also the type section of the Bonney Sand- 
glane and Rawnsley Quurtzite (respectively 
305 m and SOS m thick, Forbes L971), and 
contains a fossiliferous exposure of the Edia- 
cart’ Member (52 m thick) of the Rawnsley 
Ovarizite. In the northern Flinders Ranges 
the Wonoka Formation and Pound Subgroup 
reach a cumulative thickness al about 
4700 m-. 


The Rawnsley Quartzite in Bunyeroo Gorge 
is nominated as the stratotype for a Late Edia- 
earan Series characterised by a diverse suite 
of mitazoan fossil temains probably reflectine 
an initial radiation ef major surviving animal 
phyla, The index assemblage for the 
early part of this epoch comprises Cyelo. 
medusa davidi sensu stricta, C, plana Glaess- 
ner & Wade. “Madigania" annnlaia Sprige, 
Medusinites avreroides, Pinegia stellaris Baio- 
raria flindersi, Glaéessnerina grandis, Diekiu- 
sonia cestala, D. lista, Trlbrachidium bheral- 


EDIACARAN PERIOD 


dicum, and Pleridinium nenoxe, The Ediacara 
Member of the Rawnsley Quartzile tm estah- 
lished to be biostratigraphically equivalent to 
part of the Redkino substage of the Valdai 
Group in the Vendian of northern Russia (ef, 
Fedonkin 1981). Futher correlation of dif- 
ferent world Edigcaran sequences requires 
consideration of localized geological informa- 
tion outside the scope of the present study. 


Conelusions 


1, Over 20 years evolving ideas have deve- 
loped concerning the geological Lime 
interval characterised by the soft-bodied 
metazoan assemblages of the terminal Pre- 
cambrian. Continuing finds olf such assem- 
blages emphasize their potential value for 
biostratlgraphy and of reeent years. there 
have been several intormal designations of 
an Edigearun Pertod, 

Continuing geochronological studies associ- 

ated with different known Ediacaran sites 

establish that ¢arly metnzoan assemblages 
first appear al «, 440-6020 Ma and that 
soft-bodicd metazoans became diverse and 
frequent during the interval e, 60-570) Ma. 

The tatal time interval invelyed is compar- 

able with that of the Cambrian, the base 

of which may be placed at ve, AGO Ma or 
possibly even younger, 

3, Evidence from ceniral England establishes 
that the Ediacaran does not overlap the 
Cambrian, and that separate, sequential 
depositional events represent these time 
divisions. Ediavaran deposition in the 
English Charnian is terminated by folding, 
metamorphism, intrusion and erosion prior 
io 9 transgression beginning the lurly 
Cambrian, The metamorphism and intrusive 
events are dated at ¢. 540-330 Ma. 

4. The rocks containing the Ediaearan fossils 
are Precambrian if aspect, without the 
significant skeletal-bioclastic component 
typifying the Palacozoic and the Phanero- 
zo 4s a whole, 

_ The studies of Marland & Herod (1975) 
and Glaessner (1977, 19796) link the 
Faiacaran with the post-glacial part of the 
Proterozoic Adelaidean snecessien, uml hy 
implication indivate a type area within the 
Flinders Ranges, The appropriate strati- 
graphic interval is the Wilpensa Group. 

6, Glieingenic deposits (Elalina Tormatioa) 

immediately preceding the Wilpena Group 
gre argucd oy be of uncertain significance 


Ht 


Rk. J. Fo JENKINS 191 


for international correlation, Instead the 
base of the Echacaran is placed at the lower 
boundary of the Wonoka Formation, the 
time of deposition of which is suggested 
to approximate that of the first known ap- 
pearance of well preserved metazoans, The 
Wonoka Formation reflects marine trans- 
gression and possible later discoveries of a 
microflora, megascopic algae or even of 
Metazoa may offer a means of correlation, 
7, The Ediacaran stratotype comprises the 
Wonoka Formation, Bonney Sandstone and 
Rawnsley Quarizite in Bumyeroo Gorge, 
The Wonoka Formation represents a shal- 
lowing upward cycle after an initial trans- 
gression; the Bonney Sandstone ts generally 
paralic or regressive and is separated from 
the Rawnsley Quarlzile by a widespread 
disconformity. The Rawnsley Quartzile 
represents a renewed cycle of transgression, 
The diverse metazoan assemblage in the 
Ediacara Member, near the base of the 
Rawnsley Quartzite, is considered to reflect 
the major evolutionary radiation of surviy- 
ing invertebrate phyla. The exposure of the 


Rawnsley Quartzite in Bunyeroo Gorge is 
nominated as the sirutotype of a Late Edia- 
caran Epoch. Au unconformity of regional 
extent separates the Ediacaran sequences in 
(he Flinders Ranges from the succeeding 
Early Cambrian. 


Acknowledgments 


Professor Preston Cloud, University of 
California, read an initial version of part of 
the manuscript, and offered constructive 
criticism and posed significant questions. Dr 
Victor Gostin, University of Adelaide, read 
and criticised the tinal manuscript, Professor 
C, C. von der Borch, Flinders University, Dr 
W. V, Preiss, South Australian Department of 
Mines and Energy, and Drs B, McGowran 
and B. Daily, University of Adelaide, are 
thanked for stimulating advice and discus- 
sion. Drs Gostin and Daily also made jnvali- 
able assistance with literature and without the 
help of Mrs Mina Stojanovic in translating 
parts af new Soviet works, this study would 
not have bee possible, 


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AZOLLA CAPRICORNICA SP. NOV. FIRST TERTIARY RECORD OF 
AZOLLA LAMARCK (SALVINIACEAE) IN AUSTRALIA 


BY C. B. FOSTER & W. K. HARRIS 


Summary 


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. 


AZOLLA CAPRICORNICA SP, NOV. FIRST TERTIARY RECORD OF 
AZOLLA LAMARCK (SALVINIACEAE) IN AUSTRALIA 


by C. B. Fosrer* & W. K, Harrist 


SUMMARY 


Fostne, C. BL & Elyrris, W, K. (1Y81) Acel/la capricopica sp. noy. First Tertiary record of 
Acelfa Lamarck (Salviniacese) Im Australia, Trans. RK, Soc, §, Aust. 105 (4), 195-204, 11 
December, 198), 


Megaspores and massulae of Asolla capricornicd sp. nov. are the first recorded Tertiary 
examples of ihe water fern droll 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 yacuolae, rather than 
defined float structures, It is this fealure which readily distinguishes this species from the 
widespread extanl species, Azella filiculoides Lamarck. Columellae (sensy Powler & Stennett- 
Willson L978) are not discernible, The microspore massulac possess glochidia with anchor- 
shaped tips, und eich contiins as many as 12 laevigale to linely granular microspores, 

The fossils are of Middle to Late Eocene age, and were recovered from The Narrows 
Beds, Narrows Graben, Queensland, Their presence suggests o relatively still fresh water en- 
vironment of deposition, Known occurrences of Azella capricornica sp, nov. are confined to 
presently tropical areas of Queensland, Its phylogenetic relationship to the Quaternary and 
extant species, 4. filieuloides, found in lemperate vod sub-tropical regions of Australia, 


remains obscure. 


Introduction 


The first Australian fossil occurrence of 
members of the free-loaling heterosporous 
fern Azolla Lamarck 1783 was reported by 
Duigan & Cookson (1957) from Quaternary 
sediments in Victoria, They identified their 
specimens with an extant species. A. filiculoides 
var. rubra, which is common and widely distri- 
buted in wAustralia, It grows on areas of rely- 
tively still freshwater. Although records of ihe 
genus are Widespread in Late Cretaceous and 
Tertiary sediments of the nerthern hemisphere 
(see Collinson 1980), Acella has not been re- 
ported previously from southern hemisphere 
sequences which are older than Quaternary, 

The speeres deseribed in this paper is readily 
distinguished from A, filicu/otdes, and eon- 
geners. However, we assume that the ecologi- 
cal requirements of A, capricornica ‘were 
similar to that of extant Azel/a, allhough its 
geographic runge seems to be confined to the 
presently tropical areas of Australia, Despite 
apparently suitable biofaciey i the Tertiary 
coastal and intracratonic basins of southern 
parts of the continent, no cxamples of A zollu 
have been found in the many areas studied. 


* Geological Survey of Queensland, 41 George St, 
Brishane, Old 4000, 

+ Western Mining Corperation Ltd, 168 Greenhill 
Rd. Parkside, S. Aust, S063, 


Geological Setting 

The Narrows Beds, defined by Kirkegaard, 
et af. (1970), occut in a northwest-southeasl 
trending, graben-like structure which — is 
bounded by Curtis Tsland and the Rundle- 
Mt Lareom Ranges (Pig. 1), The Beds in- 
clude a thick sequence (500+ mi) of oil shales, 
known as the Rundle Oil Shale deposit, cur- 
rently being assessed for commercial hydro- 
carbon exploration, Lindner & Dixon (1976) 
have given a detailed description of the Rundic 
deposit, which includes at least six oil shale 
ujils or seams (of the Rundle Formation) 
which are intercalated with green-grey mont- 
miorillinitic claystones, mudstones, carbona- 


ROCKHAMPTON 
% 


CURTIS 


ISLAND 


Fig. 1. Position of drill-hele RDD66, in the Nar- 
rows Graben. 


196 


CURLEW FORMATION 


Keroaens Cresk seam 
Téelogtapny Craak unit 
Manduran Creek eoam 
Mempy Creek soam 
Grick Kilo aeam 
Ramsay Crossing seam 


WORTHINGTON FORMATION 


Fig, 2, Stratigraphy of the Narrows Beds, Nar 
rows Graben, Queensland, Units are preseited 
in descending stratigraphic order. 


RUWOLE 
FORMATION 


o 
o 
uw 
a 
” 
= 
o 
ec 
c 
« 
= 
wi 
x 

- 


ceous shales, minor impure carbonates and 
lignites. Figure 2 depicts the local stratigraphy, 
The Jithostratigraphic names have been re- 
served with the Australian Central Register 
of Straligraphic Names and will be formalised 
in a paper by Menstridge & Missen (1981). 


Although the stratigraphic boundaries of 
The Narrows Beds have not been established 
with certainly, the occurence of Freshwater 
ostracods, often in enormous numbers in mud. 
stones and oil shales, suggests that the Beds 
were deposited in lakes that were ‘largely 
shallow but probably of a permanent nature’ 
(Beasley 1945). As discussed below, the pre- 
sence of Azgella within the Curlew Formation 
and Kerosene Creek seam (in RDD 66, 139.45 
m) further supports a wholly jon-marine 
environment of deposition. 

Foster (1979) suggested that a palynologi- 
cal assemblage from a core of the Curlew 
Formation (designated al that time as the 
Wattle Creek seam) intersected in borehole 
RDD 66 al 37,40 m was of Late Eocene to 
Late Oligocene age, This range can be nar- 
rowed to Middle to Late Bocene using com- 
parative ranges Of taxa from southern Aus- 
tralia (W. K. Harris, unpublished data), 


Material and Methods 


Asolla glochidia were first recognised im 
strew slides prepared by standard procedures 
for routine palynological examination at the 
Geological Survey of Queensland, Further 
specimens of massulae and megaspores Were 
isolated by washing the disaggregated sediment 
aller digestion in 70% hydrofluoric acid 
through a 200 mesh/inch sieve. Individual 


C, B. POSTER & W, K. HARRIS 


Megaspores and massulae Were picked from 
the coarse fraction and either mounted for 
5EM study or bleached in 1% sodium hypo- 
chlorite solution and mounted in glycerine jelly 
on microscope slides for examination by trans 
mitted light, Those selected for SEM study 
were gold coated! and examined with a Cam- 
bridge 5600 SEM at the University of Queens- 
fand or with an IST Super Mini SEM at the 
Geologicul Survey of South Australia. 


Unbleached specimens were selected for 
study by both conventional transmitted light 
and transmission electron microscopy (TEM). 
TEM sections were prepared and studied at 
ihe Electron Microscope Unit, University of 
QOucensland, 


Specimens ure retained in the palynological 
collection at the Geological Survey of Queens- 
fand; catalogue numbers prefixed Q are those 
of that institution, All photomicrographs are 
from Unretouched negatives and prints, Mag- 
nifications are given for cach figure. Co- 
ofdinates of specimens, given after slide 
numbers are from a Zeiss Photomicroscope 
MHL No, 1369 housed al the Geological Survey 
of Queensland, SEM negatives prefixed MIS 
are held in the above repository; all "TEM 
negatives ure stored af the Electron Micro- 
scope Unit, University of Queensland. All 
specimens are from the Type Material, 


Systematic Description 
Famity SALVINIACEAE Dumorticr 
Genus AZOLLA Lamarck 1783 
Azolla cupricormica Foster & Harris sp. nov. 
FIGS 3-8 
Divenosiss Mevasporé apparatus without flont 
structures 4nd lacking discernible columella 
(sensi Fowler &  Stennett-Willson 1978), 
Proximal hemisphere of megaspore proper 
surmounted by dense, hairy, conical perinal 
structure, within which vacuolac are randomly 
Weveloped and distributed (vacuolue best seen 
in TEM sections, barely discernible using con- 
Ventional light microscopy). Cone commonly 
larger than remainder of megaspore apparatus: 
towards cone apex, outermost hairs are com- 
monly fused to delimit a © smooth apical 
area (best seen using SEM). Basal perimeter 
af cone defined by a = well developed collar. 
Below collur, and covering distal hemisphere 
of megaspore, perine (differentiated into ¢n- 


TERTIARY AZOLLA (SALVINIACEAE) 197 


Fig. 3. Azolla capricornica Foster & Harris sp, nov. a-c. Microspore massulae. a. Holotype (170); 
slide 9884, 9.5 98.0, Q130. b. Paratype (205); slide 9884, 21.5 119.9, Q131. c. Paratype (209), 
note microspores; slide 9884, 11.2 120.8, Q132. d, e. Microspore (500), proximal and distal foci re- 
spectively; slide 9884, 3.2 96.6, Q133. f-h. Megaspore apparatuses. f, g. Holotype (139) at differ- 


ing foci; slide 9883/9, 11.7 108.2, Q134. h. Paratype 


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 ,»m in overall width, up to 20 pm 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 ym thick, laevigate to 


(X118), showing folded megaspore exine; 


finely granular. Surface of massulae bearing 
3—20+ glochidia. Each glochidium has anchor- 
shaped tip, mostly with two recurved hooks 
(each 10-14 wm 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 ym thick, 
strap-like; 3-15 ym wide, constricted at point 
of origin and immediately below anchor-shaped 
tip, maximum width about midway along 
length (42-93 ym). Exine of megaspore (in 
surface view) minutely punctate or granulate, 
with irregularly distributed cavities in section, 
5—8 »m thick; perine 13-28 »m thick. 


B. FOSTER & W. K. HARRIS 


C. 


198 


_ 
_ — 
ie 
.. | 


ab 
ii 


7 


_ 
_ 
- 


Fig. 4. 


99 


1 


SALVINIACEAE) 


TERTIARY AZOLLA ( 


Fig. 5 


C. B. FOSTER & W. K. HARRIS 


200 


<a 


a 


cil 
cs ae 


a 


a 


Fig. 6 


TERTIARY AZULL4 (SALVINIACEAR) 


Dimensions: Megaspore apparatus: Cota) length 
410-560 pm; breadth (widest) 340-100 pm 
(50 specimens), Microspore massylac: 105- 
240 ym (maximum diameter 2) specimens}; 
microspores 24-36 pin (equatariqh dhameer)- 
Holotype: Mevaspore apparatus, Fig. 3, fey 
single mount, slide 9883/9, onordinales 11,7 
108.2, Q134. ‘Total length 465 oy maxienun 
width, at collar, 332 win, Megaspore wal] 5-10) 
wm thick, minulely pramlafe, Petine over 
distal hemisphere of megaspore 10.5-13,5 
wii thick: imperfectly reticulate, Jumina 731) 
pm wide, muri 3-12 vin wise, 

Microspore mmussuli, Piguré Sa; strew 
mount, slide 9884, wourdinates 9.5 98.0, Q130), 
Outline } circuit, 176 pm in diameter. Fuur 
microscopes are Visible, up 1 32 pm in dia- 
meter, exine 1.75 wm thick, Eight glichishii 
tre visible, amiaatmurnt length discernible 63 
wt, width 3.5-8,5 wih, 


Type Joeality! Diamend drill hole RDD 66 
at 100.25 m. Grid relerence 73SZPN 29950E, 
GLADSTONE 15100000 sheet no. 9150; 
Rockhampton |:250000 Sheet Arca, Queens. 
Jand. 


Type horizon: The Nutfows Beds, Curlew 
Formation, Sean lithology: grey-brown car- 
bobaceaus mudstone. Age, Middle to Laty 
Eocene. 

Derivation ef name; From the Tropic of 
Capricory; present kKoown occurrences of A. 
capriconmica are confined to depasits which 
straddle the Tropic, between 22° atid 24°S, 
Comparison aiid remarks. A simplified ter- 
minology has been used to desertbe specimens 


201 


wf! Azelly capricurnice sp. aov, and the accom- 
panying illustrations (Figs. 3-8) clarify the 
morphology of the species. Floats, as currently 
recognived’ (see Martin 1976, Fowler & Sten- 
nen-Willsorn 1278, Collinson 1980) have not 
heen recognised in the presently described 
species and it is their absence which separates 
A. capricorniea Jeom the various members of 
A2olla which have 3-24 floats (see Collinson 
1980, for summary). 4. capricornica is there- 
fore readily distinguished from the Australian 
Quuternary and extant species of 4, filiculoides 
which has a three float megaspore apparatus 
(see Fowler & Stennett-Willson 1978, for 
recent description). 


Compariso) with taxa which are considered 
ly possess only a single Moat-like structure or 
columellate-float, namely, Azalla yimiplex Hall 
1969. A. prinaeva (Arnold) Penhallow 19355: 
anu 4. geneseana Hills & Weiner 1965 must 
he at a relatively superficial morphologic level 
beeause of the lack of comprehensive struc- 
tural information about them. A. simplex is 
distinguished by ifs finer reticulum with lumina 
of 2-6 wm and muri ca 1 um wide, and its 
loosely attached float-like structure; A. 
primjaeva has a less conical, proximal perinal 
sinicture, und a non-reticulate distal perine, 
ai omassulae details differ (glochidia are 
Much narrower); A. geleveata is distinguished 
by, amongst olher delarls, ils slender glochidia 
with irregularly stiaped Ups. Also, because of 
the Tack of structural detail, A. capricornica 
has wot been assigned to Section Simplicispora 
(Hall 1970) which was created to accommo- 
date the above three species. 


Fig. 4. 0, SEM (675), hep, nO MES 29) micraspore Mmassula attached 16 mewaspore apparatus; 


specimen npt recovered, b. SEM (* 120), nex. no, MIS 22: microsporahwurm, showing cluster of 
Micruspare imassulicy specimen fol recovered, o SEM (172), neg, na, MIS 13: meguspore appara. 
tus; note smoath apical grea on cones specimen no} recovered, d, Gtochidium (& 1000) with single 
recurved hook al ferminus; slide S884, 18.0 107.5, QO136. & Isolated glochidium, (DIC, *635), show- 
ing constrictions at point of altachment io massula und immediately at base of tip; slide 9885, 14.2 
919 OI37.f SEM (150) neg no, 200/19; mepaspore apparalus with 3 massulae attached; speci- 
men mot recovered 9, SEM (20D) neg, no, MUS 34; megaspore apparatus showing smooth upical 
area, coarsely periculace perinc, well defined collar; specimen mot recovered, 


Fig. Sw SEM (S500), neg no. MES 32; detail of collur area, (o) with reticulate distal perine (D) 
aid hairy periic of proxinal Gone (P 1, specimen not recovered, b. SEM (5000), neg. no, MIS 26; 
glochidium with anchorshaped tip and two recurved hooks, nole consirietion at base of tip, Speci- 
men attached to megaspere apparatys, nat Tecovered,.c, SAM (1500), neg. na. 200/14; section of 
megusxpore upparatus; (M) meguspore wall, (en) endoperine and (ex) exoperine; specimen not re- 
covered, d. SUM (20750), pep. ne, 200/17; reticulite distal distal perine. showing muri OF flised, 
intertwined rugula-like elements, specimen not recovered, © SEM (525), neg no. 200/28; mega- 
spore upparalus showing smooth area delimited by fused hairs at wpex (A); specimen nol recovered, 


Fig 6, LS of meguspore apparatiis, section ca | am thiek, bur seule 50 um. Megaspore wall (M), col- 
lapsed and infolded; endoperine (en) pnd exoperine (ex) well defined. Randomly developed va- 
cuolae ti proximal cone indlgated by (VY), slide 9883/10, 12.9 77.2, QUSS, 


202 


C. B. FOSTER & W. K. HARRIS 


Fig. 7. TEM (1050) 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. 


Discussion 


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 1(0?) 
respectively. The term ‘float’ is used here in 
the sense of Fowler (1975). Using this 
criterion it would be imprudent to suggest 
that A. capricornica was the ancestral form 
of the Australian Quaternary and extant 
species, 


Azolla capricornica is presently known in 
oil shale deposits in the Narrows Graben and 
to the northwest in the Duaringa 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 
Australia. 


Palaeoecology 


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 


TERTIARY AZOLLA (SALVINIACEAE) 


a 


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 (3000), neg, no. 
7812. LS section of megaspore appartus; (M) 


203 


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 
influx. 

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, ef 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 “lamosite” 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. 


Acknowledgements 

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 


megaspore wall, (en) endoperine, (ex) exo- Unit, University of Queensland is gratefully 
perine. acknowledged. 
References 


AsuTon, P. J. & Watmsiey, R. D. (1976) The 
aquatic fern Azolla and its Anabaena symbiont. 
Endeavour 35, 39-43. 

BEASLEY, A. W, (1945) Ostracods from some 
Queensland Tertiary basins and their bearing 
on the correlation of strata. Proc. R. Soc. Qd 
56, 95-124. 

Coxtuinson, M, E, (1980) A new multiple-floated 
Azolla from the Eocene of Britain with a 
brief reveiew of the genus, Palaeontology 23, 
213-229, 


Dutcan, S. L. & Cooxson, I. C. (1957) The 
occurrence of Azolla_ filiculoides 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. 
Od Govt Min. J. 80, 442-443. 

Fow er, K, (1975) Megaspores and massulae of 
Azolla prisca from the Oligocene of the Isle 
of Wight. Palaeontology 18, 483-507, 


204 C, B. FOSTER & W. K. HARRIS 


& STENNETT-WILLSON, J. (1978) Sporo- 
derm architecture in modern Azolla. Fern Gaz. 
11, 405-412. 

Hatt, J. W. (1969) Studies on fossil Azolla: 
primitive types of megaspores and massulae 
from the Cretaceous. Amer. J. Bot. 56, 1173- 
1180. 

—— (1970) A new section of Azolla. 
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. & 
Mckirpy, D. M. (1980) Organic matter in 


Taxon 


oil shales, J. Aust. Petrol. Expl. Assoc. 20, 
44-65. 

KIRKEGAARD, A G., SHAW, R. D. & Murray, 
C. G. (1970) Geology of the Rockhampton and 
Port Clinton 1:250000 Sheet Areas. Rept 
Geol, Surv. Qd 38. 

LinpNerR, A. W. & Drxon, 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, 

Swarprick, C, F, J. (1974) Oil shale resources 
of Queensland. Rept Geol. Surv. Qd 83. 


A PARAPATRIC BOUNDARY BETWEEN TWO SPECIES OF REPTILE 
TICKS IN THE ALBANY AREA, WESTERN AUSTRALIA 


BY C. M. BULL & D. R. KING 


Summary 


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. 


A PARAPATRIC BOUNDARY BETWEEN TWO SPECIES OF REPTILE 
TICKS IN THE ALBANY AREA, WESTERN AUSTRALIA 


by C, M. BuLt* & D. R, Kina? 


Summary 


Butt, C. M. & Kina, B. R. (1981) A parapatrie boundary between two specics of reptile 
ticks in the Albany areca, Western Australia, Trany. R. Soe, S. Aust. 10S (4), 205 208, 11 
December. 1981 


The distribulion of populations of a tick morphologically similar to 4 peremvng hve 
sauri Was mapped near Albany in Western Australia, This taxon occupies an area of 
approximately 60 * 40 km. To the west its boundary appears to coincide with the edge of 
the jarrah forest belt. To the north und east ils disuabution abuts that of the more witespreud 
replile tick, Anthlyamma alholimbuium, There is no obvious ecolone ussocialed with this 
parupatric boundary, 1 is suggested that the boundary is miinfained by un interaction between 
the two species. The stability of the boundary is not yet Known, but one explanation of the 
present pallera of distribution is that Amb. albolimbatum is gridually expanding its range 
into areas previously occupied by Ap. hydrosauri. 


Introduction 

Smyth (1973) first deseribed parapalry in 
the tick species, Aponoinina hydrasauri, Am- 
blyamme limbatunt and Amb, albolimbatany i 
South Australia. All three species infest the 
sume replile hosts, In any one place only one 
of the three species is found, and their dis- 
tributions abut at common boundaries. [n- 
vestigalions have failed to provide a satisfac- 
tory explanation of the mechanisms which 
prevent range overlap at the boundaries (Bull 
& Smyth 1973, Sharrad', Bull, Sharrad & 
Petney 1981). Smyth (1973) suggested two 
hypotheses. One hypothesis is that the boun- 
daries follow environmental ecotones, and 
that at least one of the contacting species is 
poorly adapted for conditions across the boun- 
dary. This hypothesis was supported by the 
results of a detailed study near Mit Mary, 
South Australia, where the boundary between 
Ap, hydrosauri and Amb. liinbatunr comncides 
with the sharp vegetational change from 
mallee serub ta open woodland; however Wans- 
plant experiments'* have failed to demon- 
strate reduced fitness of either species across 
the ecotone (Bull ef al. 1981), At other 
houndaries helween pairs of these species 
environmental changes are less obvious (Smyth 
1973, Bull et al. 1981), 

The secand hypothesis proposes that para- 
patric boundaries are maintained by ecological 


" School of Biologicul Sciences, Flinders Liniver- 
sity of South Australia, Bedford Park, S, Aust 
5042, 

} Agriculture Protecuon Board, Forrestfield. W.A- 
GOS8. 


interactions between the tick species, resulting 
in the exclusion of one of them from the 
range of the other, The outcome of the inter- 
action may be reversed where there is an 
environmental change, such as at an ecotone. 
Thus stable boundaries will form at ecotones, 
but boundaries will be established alsa 
wherever ithe ranges of lwo of the species con- 
tact, independently of the environmental 
conditions. A prediction is that where the 
ranges of two species meet in other areas 
parapatry would be expected also, 

In southwestern Australia, Amb. albolim- 
balun is the common reptile lick hut soother 
species, morphologically similar to Ap. hydre- 
sauri, was found by C.M.B. near Albany ia 
southern Western Australia (Smyth 1973), 
The taxonomic status of this population is 
ulider investigation but here it will be referred 
ta as “Ap. Aydrosauri’. Sharcad' and Sharrad 
& King (1981) canfirmed its presenve in at 
least four small isolated areas along the south 
coast of Western Australia, Their collections 
were sulficiently detailed to show that there 
was © parapatric boundary hetween Arb. 
albolimbatunt and Ap. hydroxauri at Cape 
Naturaliste (Sharrad & King 1981), which 
may coincide with a vegetational ecotone 
(Bridgewater & Zammit 1979). We describe 


1Sharrad. R. DB. (1980) Studies of the factors 
which determine the distributions of three 
species of South Australian ticks. Ph.D. Thesis, 
University of Adelaide, (unpublished), 

*Peiney, T. N. (1981) The interaction of two 
parapairic tick species with their off host en- 
vironment. -Ph.Bo Thesis, Flinders University, 
(unpublished). 


C.M. BULL & D, R. KING 


Pe 


Fig, 1. 


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 4 de- 
tailed survey made in October 1980 of the 
area around Albany (Figure 1). Lizards, 
mainly Trachydosaurus rugosus, were captured 
as they crossed the roads or basked on the 
roadsides, and were examined for attached 
ticks. 


Only Ap. Aydrosquri 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 Amblyomma 
species have eyes but Aponomma species 
lack them (Roberts 1970). 


MT. BARKER 
o 
r=) 


7 uv s a 

We E 18 30 
Stirling R- . 

s 
*% * e g4° 30 
e 
® 
®) ". 
bed s 


Distribution of A ponomma hydrosauri (open squares and Amblyomma albolimbatum (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. 


Other data came from road-killed hosts. 
Individuals of 7. rugosus are commonly tun 
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 (Tiliqgua 
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 Paranus 
rosenbergi) from the area previously had been 
found infested by the tick species Aponomma 
fimbriatum., 


Results 


The distributions in the study area of the 
lick species. Amb. albolimbatum and Ap. hy- 
drosauri, attached to T. ragosus are shown in 
Figure 1. The range of Ap. hydrosauri is 60 
km (north-south) * 40 km (east-west). To 
the north and east its distribution abuts that 


PARAPATRY IN REPTILE TICKS 


of Amb, albalimbainm which is widely distri- 
huted in seuthernn Western Australia (Sharrad 
& King 1981). Amb. albolinbatum was not 
found within the distribution of 4p. hydra- 
sauri, except in a narrow boundary zane 
(Figure 1), where both species were found 
together. Five host individuals collected in 
the boundary zone had both tick species 
allached to them. The width of the overlap 
zone is not known, but at the northern boun- 
dary there were less than 10 km separaling 
hosts with only Ap, hydresauri from. hosts 
with only Anh. alhalimbarunt. 


To the west, the distribution of Ap. hydro- 
veut abuts with the edge of the uncleared 
jatrah forest (Kucalypius marginata). Road- 
killed 7. regosvs were found regularly over 
most of the study area, bul on the roads 
through the jartah forest neither live nor dead 
T. ragasus were seen. Seven live TP. riugasus, 
captured on the southeastern edge of the jarrah 
belt in country sull containing extensive un- 
cleared areas of forest, had no ticks attached 
to therm. In the rest of the study area only two 
other uninfested T  rugasas were found 
(Figure 1). 


Amb. albolimbarum is distributed along 
the south coast to the cast and west of Albany 
(Figure 1), und abuls with, and may just 
overlap, 4p. hydrovaur? about 6 km west andl 
about 3 km east of Albany. 


Discussion 


It is not clear which factors prevent 4p- 
hydroysauri from spreading beyond its very 
narrow range around Alhany. The morphaologi- 
cally similar specres in South Australia occupies 
a wide range of environmental conditions 
(Smyth 1973), One enviranmental change 
which seems to have an important influence 
on the distribution of Ap. ladroyaur? wear 
Ajbany is the jarrah forest to the west. The 
density of T. rugasus appears to be lower there, 
and those found oo the margins of the furest 
had no ticks on them. This suggests that some 
characteristic of the forest makes it unsuitable 
for occupancy by ticks. 


The nerthern and casterh boundaries do 
not follow macroclimalic clines as does the 
houndary between Ap. hydrosanri and Amb. 
limbarum in South Australia (Smyth 1973). 
Near Albany ihe tick boundary crosses rain- 
fall isoclines (Sharrad & King 1981). More- 
over, Ap. fydrasauri is found in’ Western 


107 


Australia. in areas receiving higher rainfall, 
such as south of Cape Naturaliste, and in 
areas receiving lower rainfall, such as Bremer 
Bay (Sharrad & King 1981). 


There are no obvious vegetational changes 
at the northern and eastern boundaries, Much 
of the arca has been cleared for farming, bin 
wide roadside verges still maintain native 
Vegetation classified as mallee heath (Beard 
1976). There may be a subtle change, but if so 
if is not significant enough to show on the 
vegetation maps of the area (Beard 1976). 
In fact, Ap. fydrasauré 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 
(1976).) 


Topographic barriers are also unlikely to be 
important in maintaining the boundary, The 
arca is dominated by two mountain ranges, 
the small Porogotup Ranges starting about 
35 km north of Albany, and the more exren- 
sive Surling Ranges starting about 65 km 
north of Albany (Figure 1) These ranges 
rise abruptly from the plains and there is a Hat 
intermontane area, about 20 km wide, between 
them, At one point the boundary is located 
on this plain. The Woaogenellup Road runs 
northeast from Mt Barker and along the 
squihern edge of the Stirling Ranges; only 
Amb. albolimbatum is found along it. The 
Porongorup Road runs east from Mt Barker 
along the northern edge of the Porongarup 
Ranges; most lizards found along this road 
carried Ap, hydrevauri Thus the boundary 
region must occur on the plain in between 
these roads; aid the mountain ranges are not 
barriers to the extension of the range of cither 
species. There are na topagraphic featires of 
any significance along the eastern boundary 
of Ap. hydrosauri. 


An alternative explanation is that the posi- 
lion of the boundary is independent of enyiron- 
menial 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 fickys al other parapatric 
boundaries have not been successful (Bull er 
al, 1981). Nevertheless, the frequent occur- 
rence of puarapatric boundaries within This 
group of ticks (Bull er al. 1981) is circum 
stantial evidence in favour of the interaction 
hypothesis, 


208 C, M. BULL & D. R. KING 


Perhaps Ap. Aydrosauri was once distributed 
continuously from South Australia to southern 
Western Australia, but Amb. albolimbatum 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. 


Acknowledgements 

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. 


References 


Bearp, 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, & ZAM™iT, C. A, (1979). 
Phytosociology of S.W. Australian Jimestone 
heaths, Phytocoenologia 6, 327-43. 

BuLL, C. M., SHARRAD, R, D. & Perney, T. N. 
(1981) Parapatric boundaries between Austra- 
lian reptile ticks. Proc. Ecol. Soc. Aust. (in 
press). 

& SmytH, M. (1973) The distribution of 

three species of reptile ticks, Aponomma hydro- 

sauri (Denny), Amblyomma albolimbatum 


Neumann, and Amb. limbatum Neumann, II. 
Water balance of nymphs and adults in rela- 
tion to distribution. Aust. J. Zool, 21, 103-10. 

Roperts, F. H. S, (1970) ‘Australian Ticks”. 
C.S.1.R.0.: Melbourne), 

SHARRAD, R. D. & Kine, 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. Jbid. 21, 91-101. 


RECENT NON-MARINE DOLOMITE FROM THE COASTAL PLAIN, 
SOUTHEASTERN SOUTH AUSTRALIA 


BY CHRISTOPHER C. VON DER BORCH 


Summary 


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 modern 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. 


BRIEF COMMUNICATION 


209 


RECENT NON-MARINE DOLOMITE FROM THE COASTAL PLAIN, 
SOUTHEASTERN SOUTH AUSTRALIA 


The coustul plaia of southeastern South Aus- 
(ralia has been the site for formation of a variety 
of carbonate minerals throughout the Quaternury,. 
Dolomite und protodolomite (approximate chemi 
cal composition of dolomite bur Jacking in ordered 
refleclions). mugnesile, hydromuagnesite, mugnesian 
calcite and aragonite are presently forming in 
Holocene ephemeral lakes on portion of the 
coastal plain which is in close proximily to a 
modern coastal lagoon kKnewn as the Coorong. 
Radiocarbon dating has verified a modern age for 
at least some of these coustal dolomile oceur- 
rences", Pleistocene dolomitic carbanztes have 
heen detecled in sediments from many interdunal 
depressions which [ie between Pleistocene barrier 
islands remote from the present coast’, These 
ureus Huve been stranded by a combination of 
slow regional upwarp combined with eustalic sea- 
level (lucluations'-™", 

The oectrrence: of the majority of active dolo- 
mite-forming lakes immediately adjacent to the 
fitesent coustal lagoon sugested That proximity 
of a marine environment is a prerequisite for 
Coorong type dolomite formation, Tt was con- 
sidered that the bulk of the necessary ions could 
be derived from a nearby marine reserveir, either 
from wind-borne cyclic sale or by entrainment of 
scuwater i coastal groundwater circalation, Sub- 
sequent hyelrologicnl unc stratigraphic studies, 
however, have cast doubt on the necessity of a 
marine idnic source’ ‘To resolve the enigmi- 
which has an jmportant bearing on theories of 
dolomite genesis in the Coorang region, sirati- 
graphie studies were initidted on Luke Ormerod, 
an inland, epbermeral, carbonate lake, 

A serics of shallow. ephemeral carbonate lakes 
inching Luke Ormerod occurs wilhin a broad 
interdinul depression west of the town af Nara- 
coorte. The takes are situated approximately &5 
km from the present coast at un elevation of 50 
m above present senlevel. Based on palaeomagnetic 
dating! this depression Was. ar least an pare a 
voustal fagoon about 600,000—700,000 years aga, 
bul hus since heen separated fram a marine 
environment by the regionyl! upwarpimg and conse- 
gent pegression, 


Tn most aspects the inland lakes appear identical 
to their more numerous coastal counterparts, They 
are emphemeral in outure, filling with water dur- 
ing winter months by groundwater seepage and 
direck rainfall, and evaporating 10 uryness during 
ensuing arid simmers, When full of water they 
dre ubout J m deep and like their coastal counter- 
paris suppert un ubundant aquatic biota, including 
ihe aquatic grass Ruppia maritima and the small 
gastropod Cuxiella confusa, Sone of the Jakes 
ure floored with fine-grained (0.2-L) dolomite 
mud, The purpose of this recannuissance study is 
(o deline the age of the dolomite in these inland 
lakes Using radiocarbon dating. in order ta deter- 
mine whether it coold be forming at present or 
Whether it is relict From times when a Pleistovene 
marine shoreline existed in the area. 

A Jm sediment core collecfed [rom Lake 
Ormerod comprises an Uppermost SU em unit of 
white, fine-grained carbonate, shawn by x-ray 
diffraction to be protodolomite (dolomite unit). 
overlying a dark grey organic-rich mod compased 
ot caleile jind subsidiary dolomile (ealetle unit). 
Assuming that the dolomite is a primary form, 
this vertical mineralogical variation implies an 
environmental change from a pre-existing per- 
manent swamp of lake to the present alkaline 
ephemeral lake, posibly in response lo a decrease 
in rainfall during the past few thousand years, 

lwo samples from the top and bottom of ihe 
dolomile unit were selected for reconnaissance 
radiocarbon dating, ‘The uppermost shaws a TC 
age of 1,300 + 60 years whilst the lowermost 73 
5770 + 90 years (Table 1), 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 af older 
dulomite or formation from an udmoixture of “old” 
carbon derived by groundwater leaching of the 
underlying Pleistacene and Tertiiry carbonate 
aquifers: The pessibility also exists of a slow 
reaction jute, of the order of thousands of years, 
int the “penecantemporaneaus” formation of dolo- 
mite in such # situation. 


TABLE 1. Data tabitlation for Lake Ormerod carbonate sanipley from reconnaissance horehote. 
ANU Sample Measured % ANC 4, Correcied 
Sample depth (cm) sly, modern age, years BP. 
(419 a —1.0+0.2 65.0 + 0.6 —149.8 + 6.4 1300 + #0 
1420 Su 34202 48.8 = 0.5 —412.2= 52 5770 = 90 


210 


The modern age of dolomites in Lake Ormerod 
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 


1Alderman, A. R. & Skinner, H. C. W. (1957). 
Am, J. Sci. 255, 561-67. 

“Skinner, H. C. W. (1963). Ibid. 261, 449-72. 

2Von 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, 
813-24, 

Rubin, M., & Skinner, B. J. (1954). Am. J. 
Sci. 263, 1116-8. 

THossfeld, P. S. (1950). Trans. R. Soc. S. Aust. 
73, 232-79. 


6. 


CHRISTOPHER C, VON DER BorcH, School of Earth 
Bedford Park, S, Aust. 5042, 


aquifer, and that proximity to a marine environ- 
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. 


SSprigg, R. C. (1952). Geol. Surv. S. Aust. Bull. 
29, 


"Cook, P. J., Colwell, J. B., Firman, J. B., 
Lindsay, J. M., Schwebel, D. A. & Von der 
Borch, C. C. (1977). Bur. Min. Resour. Aust, 
Geology & Geophysics 2, 81-8. 


1%dnurm, M. & Cook, P. J. (1980). Nature 
(2867), 699-702. 


11Von der Borch, C. C., Lock, D. E., & Schwebel, 
D. (1975). Geology. May, 283-5, 


Sciences, Flinders University of South Australia, 


FIRST RECORD OF PROMICROPS LANCEOLATUS (BLOCH) PISCES: 
SERRANIDAE) IN SOUTH AUSTRALIAN WATERS 


BY P. J. KAILOLA AND G. K. JONES 


Summary 


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.” 


BRIEF COMMUNICATION 


21) 


FIRST RECORD OF PROMICROPS LANCEOLATUS (BLOCH) PISCES: 
SERRANIDAE) IN SOUTH AUSTRALIAN WATERS 


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!-2, The limits of its distribution have 
been reported as Algoa Bay (East Africa), India, 
Taiwan, Gilbert Islands, Fiji and Hawaii3-4. Jn 
Australia it has been recorded from northern 
N.S.W., Queensland, Western Australia and “pre- 
sumably N.T."9-, 

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 
waters. 


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- 
ton. 


A description of the fish follows: 
Family Serranidae. Promicrops lanceolatus 


Holocentrus lanceolatus Bloch 1790, Nat. 
Ausland. Fische 4, p. 92, pl. 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.XI, 16. A.IIL8. P.ii,15,i. L.lat. 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? found a lower-than-normal gill 
raker count for a 574 mm specimen of Serranus 
fuscoguttatus (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 upper. 
The 8th is the longest dorsal fin spine. The pre- 
operculum is highly angular; soft fins are 
rounded, 


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, 


Promicrops lanceolatus has been reported to 
grow to nearly 4 m long!-4:6, and Schultz? has 
investigated the authenticity of records of large- 
sized Promicrops, 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! Four- 


Fig. 1. Queensland Grouper Promicrops lanceolatus. Total length 2120 mm. 


212 


manoir & Laboute? state that Promicrops can 
reach 400 kg. 

The South Australian specimen is believed to 
be the most southerly record for the species. 
Aleon Bay (approx. 34°00'S, 26°00B) was. the 
previous southerly record and this location is 
close to the southern-most Jimit of the tropical 
“Aculhus Current” which extends down the east 
coast 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!*-4 as an custward 
flowing current during autumn and winter months 
along the southern coast of Western Australia snd 
across the Great Australian Bight. Satellite obser- 
vations of sea surface temperature fronts!5 show 
that this current of relatively warm, low salinity 
wiler flows completely across the Great Australian 
Bight, It is also interesting to note that Markinat® 
classified the phyto- and zooplanton in the Great 
Australian Bight as of tropical origin. 

We thank Messrs. D. Wankie, A. Billing, D, 
Evans and D. Roberts who brought the fish to our 
allention and assisted with its transportation and 
measurements; also to Messrs. R. O. Ruchle and 
W. Head for the photograph and museum assis- 
lance. 


IRoughly, T. C. (1951). ‘Fish and Fisheries of 
Australia,’ (Angus & Robertson: Sydney). 
*Fourmanoir, P. & Laboute, P, (1976). “Poissons 
de Nouvelle Caledonie et des Nouvelles 
Hebrides.” (Les Editions du Pacifique: Tahiti). 
SSehultz, L, P, (1966), Addenda. p.p. 145-165. 
Jn: Schultz, L, P, & collaborators. U.S, natn. Mus. 
Bull, 202, 3. 

ISmith, J. L. B. (1965). 
Southern Africa.” Sth 
Agency; South Africa). 
5MecCulloch, A, R. (1929). Mem. Aust. Mus. 


5(2): 145-329 
tMunro, LS. R. (1961). Fish. Newsl. 20(8); 


161-4. 


“The Sea Fishes of 
edin (Central News 


Pd 


Tas.e 1. Measurements (in mm) of P, lanceolatus 
(obtained Jram the Jresh specimen), 


Total length 2120 
Standard length 1775 
Head length 730 
Body depth 670 
Snoult length 170 
Eye diameter 5) 
Interorbital width 190 
Lower jaw tip to vent 1320 
Maxillary length 390 
Head depth 435 
Caudal peduncle depth 215 
Caudal peduncle length 345 
Postorbital head length 510 
Longest pectoral ray 3460 
Longest ventral ray 285 
Longest anal ray 280 
Third anal spine lil 
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 


TRandall, J. E. (1964). Pacific Sci. 18(3): 281-96. 

S8Alcock, A, (1905). Nature 71t 415. 

‘Smith, H. (1933). J. Siam Soc. Nat. 
Suppl. 9(1): 85. P 

10Grant, E. M. (1978). “Guide to Fishes” (4th 
ed.) Dept Harbours & Marine, Qld. 

11Parbyshire, J, (1964). Deep-Sea Res. 11: 781- 
815. 

hNochbort, D. J, (1975). Proc. Ecol. Soc. Aust. 

: S7-83. 

itPashkin, V. N, (1968). Tr. Atl, Nauchno-Issled. 
Inst. Rybu. Khoz. Okeanogr. 28 (transl.). 

‘Cresswell, G. R. & Golding, T. J. (1980). Deep- 
Sea Rts. 27A: 449-66. 

lLegeckis, R. & Cresswell, G. (in press). Jhid. 

MWiMarkina, N. P. (1976), Oceanology 15: 602-4. 


Hist., 


KaAILoLa and G. K. Jones, Department of Fisherics, 25 Grenfell Street, Adelaide, S. Aust, 5000, 


HALO-EFFECTS IN NATIVE VEGETATION 


BY ROBERT T. LANGE AND TIMOTHY REYNOLDS 


Summary 


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. 


BRIEF COMMUNICATION 


HALO-EFFECTS IN NATIVE VEGETATION 


Circulur zones of suppressed growth, called 
halos'@, surround individuals and clumps of some 
plants, particularly trees such as Caswarine 
cristata, in South Australian vegetation. These 
halos extend well beyond the cinopy spreads of 
the plants at their toci and show up clearly on 
werial photographs, Suggested explanations of the 
suppression inclide water shortage raduced by the 
central plant, allelopathy, or grazing and seed-har- 
Vesting unimals harboured by the central plant, but 
fieldl observations indicate that none of these pro- 
vides a satisfactory explanation on ils own*. 

Zones of enhanced growth also oceur, particu- 
lurly under trees such as Acdeks papyrecarpa and 
shrubs such as Maireana sedffalia, These zones 
usually do nol 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’ conditions induced 
by the central plant and, under intricate shrubs, 
protection from the grazing of large vertebrates, 
yielding contrasts With unprotected interspaces. In 
general field observations support the view that 
these ure very plausible hypotheses, but there are 
compleations, For iastance 4, papyrecarpa cance 
pies in the Whyalla region usually enhance the 
likelihood of finding species beneath them, and 
luck an extended grass suppression halo, but 
sometimes the reverse is Observed, Both culegories 
af zones are well known elsewhere’. The most 
recent Austrian publication about them concerns 
Eucalyptus baxter’ suppression zones in Victorian 
tree-heath!", 

We have detected a further category of such 
plant-centred effects which, so var ws we can estab- 
lish, is unreported, It dillers from the preceding in 
Uhal it is Nol expressed us Overt weneral suppression 
ur enhancement of plant growth around particular 
trees or shrubs, and cannot be proved by inspect- 
ing any single tree or shrub for halo effects. The 
effect is proved only by considering the species 
populution collectively and consists of tendencies, 
within small circles centred on individuals. for 
hulos of shifts in floristic composition of sulround- 
ing vegetation, Le. for shifts in the probubilities of 
incidence of olher species, from those applying 
within small circles centred on individuals of a 
species fo different probabilities applying in the 
interspaces. belween the circles, This note sum- 
marizes our Investigation and its findings, 

‘Traverses in native vegetation were marked out 
separately between individuals of each species 
onder test for halo-elfect, Our technique required 
successive individuals to be separated by a least 
twice their canopy diameter, 

Circles of raditis rj, meen ry, Ur im the range 
0.5-3.0 m) were murked out around each indivi- 


dual of a test species und searched for incidence 
of all other species, This was repeated at a ran- 
dom point in exch midlocation between test indi- 
viduals. Data for each quadrat size separately were 
assembled as 2 % 2 homogeneity tables for each 
combination of test and other species in turn, and 
tested lor significance by y+ or Fishers exact 
test!'!, using a computer. Resulls were graphed as 


E.microcarpa 


E.leucoxylon 


3 0.5 1.0 1.5 20 2.5 
r(m) 
Fig. |. Halo signatures of Eucalyprs microcarpa 


and £. leuneexylon in native vegetation at Belair: 
Individuals of these species were intermingled 
as tree cover over dense, low sclerophyllous 


scrub, 3 = the sigmficance level of p < 0,001. 
+ and — signify positive and negative assacia- 
lions respectively. A = Acrorriche serrulata, B 


= Gonocarpus elatus, C = Olea eurapea. 


214 


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 
follows: 


(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. leucoxylon 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 asparagoides (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 


1Andrew, M. H. (1978). The initial impact of 
depasturing sheep on arid chenopod shrublands. 
Ph.D. Thesis, University of Adelaide. 

2Johnson, A. W. (1978). Vegetation suppression 
in the Whyalla region, with particular reference 
to the phenomena around Casuarina cristata 
Mig. B.Sc. (Hons) Thesis, Department of 
Botany, University of Adelaide. 

3Correll, R. L. (1966). Studies on the nitrogen 
economy of semiarid vegetation at Yudnapinna 
Station, South Australia M.Sc. Thesis, University 
of Adelaide. 

4Barker, S. (1972). Effects of sheep stocking on 
the population structure of arid shrublands in 


E. leucoxylon 


+n 
) 


15 2:0 25 3:0 


r(m) 


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. 


05 1:0 


native woodland by olive woodland probably was 
fostered by this link, for which no explanation is 
available. Once established, olive harbours its own 
seedlings. 


(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. 


South Australia, Ph.D. Thesis, University of 
Adelaide. 

5Muller, C. H. (1953). Amer, J, Bot. 40, 53-60. 

6Muller, W. H. & Muller, C. H. (1956). ibid, 43, 
354-61. 

TJaksic, F. M. & Fuentes, E. R. (1980). J. Ecol, 
68, 665-70. 

8Storey, R. (1967). Aust. J. Bot. 15, 175-87. 

9Nat. Acad, Sci, (1971). “Biochemical Interactions 
among plants.” Environmental Physiology Sub- 
Committee. (Washington D.C.). 

10Del 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. 


Rosert T. LANGE and TIMOTHY REYNOLDs, Department of Botany, University of Adelaide, Box 498, 


G.P.O., Adelaide, S. Aust. 5001. 


NOTES ON REPRODUCTION BY CAPTIVE AMPHIBOLURUS 
NULLARBOR (SAURIA: AGAMIDAE) 


BY JULIA SMITH AND TERRY D. SCHWANER 


Summary 


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. 


BRIEF COMMUNICATION 


NOTES ON REPRODUCTION BY CAPTIVE AMPHIBOLURUS NULLARBOR 


(SAURIA: 


Two Amphibolurus nullabor Badham were col- 
lected 20 km E of Nullarbor Homestead, S.A. 
(31°28’S, 131°12’E), amongst bluebush (Mai- 
reana 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=140mm) 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). 


AGAMIDAE) 


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+1.1 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. 
37.3 mm. 


TaRLe |. Egg sizes (length and width in mm), hatching dates, and hatchling sizes (SVL and total length 
in mm) for a clutch of Amphibolurus nullarbor eggs. 


Dates Measured (1980) Dates Hatchling 
hatched sizes 

Egg no. 10/1-2+ 10/28 11/13 12/14* (Dec. 1980) SVL TL 
1 22.9 x 13.9 24.7 x 17.4 26.0 x 18.6 26.1 x 19.1 19 36.3 85.3 
2 23.4 x 14.6 26.1 x 17.0 27.1 x 18.3 26.6 x 17.9 21 37.0 87.5 
3 24.2 x 15.8 26.6 x 17.9 27.6 x 19.4 26.9 x 20.0 22 36.3 83.2 
4 23.5 x 15.0 26.2 x 184 27.6 x 19.3 24.9 x 19.2 18 37.3 80.0 
5 23.0 x 15.3 23.2 x 18.1 26.1 x 19.1 25.1 x 19.1 21 35.9 86.1 
6 23.8 x 15.6 26.3 x 18.7 27.0 x 19.3 26.0 x 18.2 25 33.6 75.6 
7 23.8 x 14.4 26.0 x 18.0 26.7 x 18.7 26.6 x 18.9 27 35.6 76.0 
8 22.9 x 16.1 24.8 x 17.9 25.6 x 18.7 24.2 x 18.9 24 35.0 80.2 
9 24.0 x 14.2 24.8 x 17.9 26.4 x 18.6 25.8 x 18.4 20 36.1 82.8 
X+SD 23.5 15.0 25.0 17.9 26.7 18.9 25.8 18.9 35.9 81.9 
+05 +408 #11 +405 40.7 +04 +09 +0.6 #11 +42 


+ Dates when eggs laid. 


* Note that all eggs except No. 1 decreased in length and/or width just prior to hatching. 


216 


These observations are similar to those reported 
for A. barbatus, which most resembles A. nullar- 
bor, morphologically+. Eggs of A. barbatus in- 
creased 90% by volume from parturition to 
maximum size, under incubation conditions simi- 
lar to those described here. Also A. barbatus 
eggs shrank slightly just prior to hatching, and 
hatchlings remained motionless in ruptured eggs 
for several hours®, 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, nullarbor data‘. 


In S.A. populations of both species are allo- 
patric’, 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*: 7), This difference 


1Bustard, H. R. (1979) Australian Lizards. Col- 
lins, Sydney, 162 pp. 

2Smith, J. (1974). S. Aust. Herpetol., 2(1); 10. 
®8Smith, J. (1979). Herpetofauna, 19(2): 12-14. 

Ss al J. A. (1976). Aust. J. Zool., 24: 423- 


JuLia SmirH, 7 Jeffrey Road, Vale Park, S, Aust. 


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 are 
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.%: 5, and often have two 
clutches per season? 65, Snout-vent lengths for 
hatchling A. barbatus average 5 mm larger than 
those of A. nullarbor. These differences are 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 hatchling A. 
nullarbar. 


‘Houston, T, F. (1979), ‘Dragon lizards and 
goannas of South Australia.” S. Aust. Mus, Spec 
Ed, Bull. Ser., 84 pp. 

SBustard, H. R. (1966). Brit. J. Herpetol., 3: 
252-259, 

‘Licht, P. (1973), Comp. Biochem. Physiol. 45A: 
7-20. 

SMitchell, F. J., unpublished data, 


5081 and Terry D, ScHWANER, South Australian 


Museum, North Terrace, Adelaide, S, Aust. 5000. 


SKULLS OF BETTONGIA LESUEUR (MAMMALIA: MACROPODIDAE) 
FROM A CAVE IN THE FLINDERS RANGES, SOUTH AUSTRALIA 


BY MEREDITH J. SMITH AND P. A. W. ROGERS 


Summary 


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. 


BRIEF COMMUNICATION 


217 


SKULLS OF BETTONGIA LESUEUR (MAMMALIA: MACROPODIDAE) FROM 
A CAVE IN THE FLINDERS KANGES, SOUTH AUSTRALIA 


Muny Australian mammal species suffered a 
drastic reduction of range and seme became 
extinci svon after Puropean colonizalion, Informu- 
tion on the farmer distribution of these species 
can be ublianed only opportunistically by the dis- 
covery atd recognilian of the remains in pro- 
tected loentions such as caves, dunes and middens, 

Puring the exploration of Yellowfooted Rock 
Wallnby Cave (lat. 31°20°S, jong. 18°40 BY, 
about 30 km narth of Wilpena in the Flinders 
Ranges, Sauth Australia one of us (PLR-) found 
two macropodid skulls and a pelvis in a low tight 
passage sume distance from the enlrance to the 
cave. The topography of the cave indicated that 
there wis little likelitaod of the bones having been 
carried to their resting place by a water current 

The more complete skull (now registered South 
Australian Museum (SAM MI0769) lacks left U 
and a few fragments of the premaxillac. The 
lowet jaw und the anterior cervical vertebrae are 
held tightly in place by dry flesh and skin, and 
fragments of dry skin and fur adhere to the skull. 
The other skall (SAM M10770) lacks the incisors 
und untenor putts of the nasal benes. Nh has no 
skin aehering but is greasy, The pelyts and 
allached lumber vertebrae have leaments and 
dricd flesh adhering; they seem to have been 
gnawed by rodents. 

Both skulls are of adult FReltengia lesneur 
(Quoy & Gaimjard) and the pelvis is almost cer- 
tainly of this species. B. leseur is characterized 
by vertivally-crooved. very Jong premolars, low- 
crowned bunodsm| molars, sleep sice gradient of 
molars with M? very small, and by inflated «audi. 
tory bullue, The P+ length is abour 14% of basil 
length uf skull. the bulla length about 25% of 
basal leneth'!, Por the two cave skulls the tneasure- 
ments in mm relevant t diagnosis are: basal 
Tength 63.0, approx, 43; Pf length 8.9, 8.9: bulla 
length 17.6, 17.8 respectively. 

A. lesuenr was widespread in Anstralia and as 
lale as [940 wus abundant in ihe Musuraye- 
Everard Range area? althotligh it exists now only 
on offshore islinds to Western Australia’. Pinloy- 
son found “abundant eviclerice to show that @er- 
fiona lesvenr Was formerly one of the most 
numerous und universally distributed mammals at 
South Adisiralia. finding and colonizing sitable 
hubitat zones in all the disuicts of the State with 
jhe possible exception of the deeper Mallee and 
the flooded partion of the lower South-Bast and 
the Winer porttans of the Nullurbor Plain". His 
broad aulline of the distribulion included all af 


'Waoketicld, N. A, 11967). Viel. Nat, BA, 8-22 
“BiWayson, HE Fh (1958). Rec, S. Aust, Mus 
[h(Q), 245-302 

‘Ride, W. DL, (19700. ‘SA guide to the ative 
miumnials of Australia”, Oxfurd University Press 
Melbourne, 


Meribrri dt, Survie and PAW. Rogerst, 


‘ 


T= 


' 
' 
4 
' 
‘ 
, 
‘ 
‘ 


Fig. |; Map of South Australia showing location 
of origin of specimens of A. lesveur in the South 
Australian Museum mammal collection. Arrow 
shows Yellowfooled Rock Wullaby Cave, 


South Australia except the far northeast, However, 
few specimens were reliined in museums. In the 
South Australian Mbscum there are no specimens 
of B. lesticar from the Flinders Ranges, the nearest 
locality being Pr Augosta, where one specimen 
(M8093) was taken in 1880 (Pig. 1) und living 
Specimens have not been recorded from the Flin- 
ders Ranges! A mutdible fragment of A. levueur 
has been found in an owl pellet deposit in 
Chambers Gorge, northern Flinders Ranges (G. C. 
Medlin, personal communication), 

The rocky habitat around Yellowfooted Rock 
Walluby Cuve ds not typical for B. lesveur, indi- 
viduals of which are borrowers, and live in war- 
rens dug in sundy soill. Whilst the small areas of 
alluvial sand deposited in the gorges by Pnorama 
and Brachinu Creeks may have provided some 
sinlible substrate for burrowing, the position af 
the hones deep within a small cave suggests that 
caves us Well as burrows Were Used for shelter Tt 
is unlikely that the specimens from the cuve lived 
elsewhere than in the Runges because the plains 
to the West are about 19 km distant and to the 
east Further uway, 

Whilst dried flesh may survive on bones in a 
cave for thousands of years? the skulls from Yel- 
lowfooted Rock Wallaby Cave provide a positive 
record af B. lesueur Tram the Flinders Ranges in 
the late Holocene. 


'Ailken, BP. F. (1980)), Mammals, pp. 288-314. in 
Corben, D. ed., “A Field Guide to ihe Flinders 
Ranges.” Rigby. Adelside. 


‘Partridge, # (1967). J, R- 
57-59 


Soe, W. Aust. 50, 


~Department of Zoulogy, University of Adelaide, G.P.O, 


Rox 498, Adeliide, S. Aust. SUO1. {Department of Human Morphology, Flinders Medical Centre, 


Bedford Park, S. Aust. S042, 


A GIANT DEEP-SEA SQUID, TANINGIA SP., FROM SOUTH AUSTRALIAN 
WATERS 


BY WOLFGANG ZEIDLER 


Summary 


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. 


D8 


BRIEF COMMUNICATION 


A GIANT DEEP-SEA SQUID, T4ANING/A SP. FROM SOUTH AUSTRALIAN 
WATERS 


In April L980 fishermen from Port Lincoln 
reported the capture of u giant squid bearing 
hooks instead of suckers on the arms, The speci- 
men, 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 dic nor have facilines to 
keep the specamen whole, and so jt was cut up and 
subsequently sold for bait. However, two arms and 
the beaks were recovered, thus permilting generic 
identification. 

Dismay at the destruction of such a scientifically 
valuable specimen prompted fishermen to look out 
for more and, in carly May, 1980 two were re- 
covered from the same general area, Oue of these 
had the head and arms missing, but still weighed 
95 ke and had a dorsal mantle length of 158 cm, 
The other specimen weighed jn excess of [10 ke 
and meusured 2.1 m total length. {t was in almost 
perfect condition and is currently frozen awaiting 
a more detailed examination, 


Preliminary investigations have shown that the 
specimens represent. the genus Taningia (Family: 
Detepoteuthidae), but as they have not been exa- 
mined in detail a specific determination could not 
be made. However, they are most likely T dariae 
Joubin, 1931, as the gents is monotypic and the 
Material at hand does not seem to vary markedly 
from published descriptions?~, 


There are a number of disunetive features which 
characterize this species, The arms sre relatively 
short, about } of the mance length and are urmed 
with a series of hooks instead of suckers, The 
second arms each bear a large heht organ at their 
tin which was bright yellow in the present speci- 
mens and mewstired about 5 * 3cm. The tentacles, 
which account for most of the lengli in the true 
giant squids, Archifeuthis, are absent m matire 
specimens and vestigial in juvermles of Jan/ngia- 
The fins extend for most of the mantle and are 


iJoubin, L, (1931). Annls Inst, opeanogr , Monaco 
10, 169-211, 
“Clarke, M. R, (1967) 
(19), 127-43. 


Symp. gaol, Soc. Lond. 


very large, making the body of the animal almost 
us broad as the mantle length. The mantle wall 
ond fin are extremely thick and gelatinous and the 
body is relatively more bulky than any other spe- 
cles of squid, 

Nearly all of the known specimens of Tuningia 
have been collected from sperm whale stomachs, 
and it is unusual to encounter them floating on the 
surface. If i8 possible thal 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 Port Lincoln fishermen 
dead squid offen come to the surface near the edge 
of the continental shelf. but they are quickly eaten 
by seabirds and it is difficult to find a complete 
specimen. T, danae forms an important part of 
the diet of sperm whales and by weight is the most 
significant part of sperm whale dicts’. Clarke 
recorded heaks 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, The pre- 
viously known distribution of Taningia, mainly 
From sperm whale stomachs, includes the Azores, 
Madeira, the equatorial Atlantic and the eastern 
coast of South Africa®. 


T intend to publish the results of a more detailed 
examination of the specimens in the near future, 


My sincere thanks to Dinko Lukin and the crew 
of the “Karina G7 and to Juergen Zenke and the 
crew of the “Rosalind Star” for collecting and 
donating the specimens to the South Australian 
Museum, My thanks also go to 5.A.F.C,0.L, who 
transported (he specimens to Adelaide tree of 
charge. A special thanks is due to Neville Wonklyn 
of the “Port Lincoln Times” who alerted the fisher- 
men to the importance of the specimens, and 
to his Editor for being sc understanding, 


*Clarke, R. (1956). Discovery Rep, 28, 237-98, 
‘Clarke, M, R, (1980). hid. 37, L-24. 


WOLFGANG ZripLer, South Australian Museum, North Terrace, Adelaide, 8, Aust. 5000. 


ADDENDUM 


BurTLer & KEouGHR—Trans. R. Soe. 3, Aust. 105(1), 29-39. 
The authors apolovise for the omissian from the manuscript af Acknowledgements. We 


intended to include the following: 


We are indebted to Tony Oliver and Deitdre Coambe for assistance in the field. The work 
was supported by o grant from the University of Adelaide Research & Publications Committee 


and by ARGC Grant DI7815083. 


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