VOL, 104, PARTS 1 & 2

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

29 FEBRUARY,

Barker, S.

Mawson, P. M.

Plummer, P. S.

CONTENTS

New species and new synonyms of Stigmodera (Castiarina)
(Coleoptera: Buprestidae) - - - - = : :

Some strongyle nematodes (Amidostomum spp.) from Austra-
lian birds) - - - - - 2 : “ H,

Circular structures in a Late Precambrian sandstone: Fossil
medusoids or evidence of fluidization? - - - - -

Tyler, M. J. & Davies, M. Systematic status of Kankanophryne eevet & Liem

(Anura: Leptodactylidae) —- - - 2 . 2

De Deckker, P. New records of Koonunga cursor Saye; 1908 seinea ils, Ana-

Viets, K. O.

spidacea) - - - - : . - -

New Unionicolidae (Acari, Hydrachnellae) from Australia -

17

21

Pa];

1980

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

STATE LIBRARY BUILDING

NORTH TERRACE, ADELAIDE, S.A. 5000

TRANSACTIONS OF THE

ROYAL SOCIETY OF SOUTH AUSTRALIA INC,

PARTS 1 &

Barker, S.

Mawson, P. M.

Plummer, P. S.

CONTENTS, VOL. 104, 1980

2, 29 FEBRUARY

New species and new synonyms of Stigmodera (Castiarina)
(Coleoptera: Buprestidae) - - - - - - -

Some strongyle nematodes (Amidostomum spp.) from Austra-
lian birds - - - - € - 4 - 2 P

Circular structures in a Late Precambrian sandstone: Fossil
medusoids or evidence of fluidization? - = - J -

Tyler, M. J. & Davies, M. Systematic status of Kankanophryne Hever & Liem

De Deckker, P-.

Viets, K. O.

PARTS 3 & 4,

Twidale, C. R.

Breed, W. G.

(Anura: Leptodactylidae) - - - - - 2

New records of Koonunga cursor Sayce, 1908 (Syncarida, Ana-
spidacea) - - - - e e m) nf = E

New Unionicolidae (Acari, Hydrachnellae) from Australia -

30 MAY

The Devil’s Marbles, central Australia - - = - -

Further observations on spermatozoal morphology and male
reproductive tract anatomy of Pseudomys and Notomys species
(Mammalia: Rodentia) - - - - - - -

Flint, R. B., Ambrose, G. J. & Campbell, K. 8. W. Fossiliferous Lower Devonian

Beveridge, I.

Mawson, P. M.

Ludbrook, N. H.

boulders in Cretaceous sediments of the Great Australian Basin

Progamotaenia Nybelin (Cestoda: Anoplocephalidae): new
species, redescriptions and new host records - 4 :

Beveridgea new genus (Nematoda: Strongylida) from the Agile
Wallaby from Northern Australia - - = - x 2

Non-marine molluscs from dolomitic limestones in the north
of South Australia - - - 4 f 7 \, ¥

9

13

41

51

67

81

NUMBERS 5 & 6, 28 NOVEMBER

Robinson, A. C. Notes on the mammals and reptiles of Pearson, Dorothee and
Greenly Islands, South Australia - - - - - -

Williams, D. L. G. Catalogue of Pisecahe wprtghtate fossils and sites in South

Australia - - 4 5 = :
Moore, P. S. Stratigraphy and depositional environments of the Billy Creek
Formation (Cambrian), east of the Flinders Ranges, South
Australia - - - - - - - - - -

Koste, W. & Shiel, R. J. New Rotifera from Australia = - - - - -
McNamara, K. J. Heteromorph ammonites from the Albian of South Australia -

Barton, C. E. & McElhinny, M. W. Ages and ashes in lake floor sediment cores
from Valley Lake, Mt Gambier, South Australia - ms -

von der Borch, C. C., Bada, J. L. & Schwebel, D. L. Amino acid racemization
dating of Late Quaternary strandline events of the coastal plain
sequence near Robe, southeastern South Australia - - ~

Glaessner, M. F. New Cretaceous and Tertiary crabs (Crustacea: Brachyura)
from Australia and New Zealand - - - - - -

van Beurden, E. & McDonald, K. R. A new species of Create Cerio
Hylidae) from northern Queensland - -

Jago, J. B. & Youngs, B. C. Early Papibetan tetas aoe the Olkie Basin,
South Australia - - 3 S

Fitzgerald, M. J. Muckera and Millbillillie—Australian achondritic meteorites -

93

101

167

171

193

197

201

Insert to Transactions of the Royal Society of South Australia, Vol. 104, Numbers 5 & 6, 28 November, 1980

NEW SPECIES AND A CATALOGUE OF STIGMODERA (CASTIARINA)
(COLEOPTERA: BUPRESTIDAE)

BY S. BARKER

Summary

Stigmodera rudis Carter is reinstated in the sub-genus Castiarina. Thirteen new synonyms are given
and two species are resurrected from the synonymy. One name has been wrongly synonymised. A
lectotype of S. rubriventris Blackburn is selected. Seven new species of Stigmodera (aliceae, aquila,
aurea, hateleyi, inflata, kiatae, mimica) are described and illustrated.

NEW SPECIES AND NEW SYNONYMS OF ST/IGMODERA (CASTIARINA)
(COLEOPTERA; BUPRESTIDAE)

by S. BARKER*

Summary

BARKER, S. (1980) New species and new synonynis of Stivmiodera (Castiarina) (Coleoptera,
Ruiprestidae). Trans. R. Soc. S. Aust. W4(1), 1-7, 29 February, 1980.

Stigmodera rudis Carter is reinstated in the sub-genus Costarina, Thirteen new syoonyms
are given and two species are resurrected from synonymy. One nume has been wrongly
synonymised. A lectotype of S. rbrivenriy Blackburn is selected, Seven new species of Stis-
madera (aliceae, aquila, aurea, hateleyi, inflate, hiatee, mintica) ave described and illustrated,

Introduction

Despite the wide occurrence of Stigmmdera
(Castiarina) species in Australia, some have a
restricted distribution, are not represented in
Museums and are difficult to obtain. Since my
catalogue of Stigmodera (Castiarina) (Barker
1979) 1 have borrowed or collected specimens
of a number of species previously unavailable.
From a comparison with types, photographs of
types and eXamination of male genitalia, | now
recognise seven new species and synonyms that
1 had missed previously.

Stigmodera (Castiarina) rudis Carter 1934
FIG. |

| have examined a complete male specimen,
It is a lycid mimic and, except that it has a
hairy head, antennae and pronotum, resembles
all other Stigniodera (Castiarina) and should
be included in the sub-genus. It was collected
on flowers of Leprospernum sp. at Lake Dob-
son National Park, Tasmania, at an elevation
of 914 m an 19.1.1951 by J, R. Cunningham,
Barker’s (1979) key to Stigmodera should he
modified so that the second phrase of the 2nd
couplet appertaining to Caytiarina Teads, “sel-
dom with hair on dorsal surfaces of head and
pronotum”.

Additions to synonymy of Stigniadera
(Castiarina)
australasiae L & G 1837, Mon, Bupr, 2, 32
assimiliy Hope (846, Trans. ent. Soc. Lond. 4,
212 (new synonym)
melbournensis Thomson 1879, Typ, Bupy, App,
la, 34
timida Kerremans 1898, Annals Soe, ent. Lely,
42, 147

* Department of Zoology, University of Adelaide,
Box 498, G.P.O., Adelaide, 8S, Aust. 500],

puerilis Kerremans 1898, Annis Soc, ent. Belg,
42. 147

varmined Saunders 1868, J. Linn, Soe. 9, 474
collivens Kerremans 1890, Bull. Soc, ent, Belg.
1890, 44 (new synonym)
felix Kerremans 1898, Anniy Soc, ent, Bela, 42,
142

dawsonensis Blackburn 1890. Trams. R. Soe. 8.
lust. 13, 155
pulchella Carter 1916, Trans, R. Soc, 8, Aust,
40, 135 (new synonyny)

deuqueti Carter 1927, Proc. Linn. Soe, N.S.W, 52,

225
sutton’ Carter 1932, Proc. Linn, Soc. N.S.W. 57,
104 (new synonym)
palazera Carter 1937, Trans, R, See, 8. Aust.
61, 125 (new synonym )

duaringae Carter 1929, Proc, Linn, Soe, N.S,
54. 6%
bagania Carter 1930, Proe, Lire
55, 534 (new synonym )

inermis Kerremans 1890, Bull. Soe. ent Belg,
1890. 45
nove Kerremans 1902, Getera Miser, 12, 208
rubella Carter 1931, Aust, Zool, 6, 345 (new
synonym)

media Hope (847. Trans. ent. Soe. Lond. 4, 284
seplemnoraa Carter W916, Tras, Re. Soe. §,
Aust. 40, 86 (new synonym)
yeplenemaculate Blackburn 1892, Trais, R. Soe.
S. dust. 1S, 45

piliventris Saunders (868, J. Linn, See, 9, 474
penerasa Kerremans 1898, Aanly Soc. ett. Bele.
42. [50 (new synonym)

rectifasciata Saunders 1868, J. Linn. Soc, 9, 472
vicilans Kerremans 1898, nals Soe. ent. Belg.
42, [43 (recognised by Blackburn 1900 p, 42)

riblounda Carter 1931, Aust. Zeel. 6, 346
Vielatra Denquet 1956, Proc. Linn, Soe. NSW.
81, 156 (new synonym)

rufipenniy USirby) 1818, Trans. Linn, Soe. 12, 456
crocipennis Le & G I837, Man. Bupr, 2, 21
crocipennix Hope 1846, Trans, enn Soe, Land,
1846, 292 Ma
qidadrijavedlard Obenberger 1933, Cay cal, Spel.
ent. 30, 69 (new synonyo)

Soe. NUS A.

2 S. BARKER

scalaris (Boisduval) 1835, Voyage de l’Astrolobe,
89

cyanicollis (Boisduval) 1835, Voyage de l’Astro-
lobe, 91

crucigera L & G 1837, Mon. Bupr. 2, 40
viridis L & G 1837, Mon. Bupr. 2, 46

crucigera Hope 1838, Col. Man. 2, 162
macleayi Blackburn 1892, Trans. R. Soc. S.
Aust. 15, 48 (new synonym)

prudens Kerremans 1898, Annls Soc. ent. Belg.
42,152

suavis Kerremans 1902, Genera Insect. 12, 210

crucioides Obenberger 1922, Arch. Naturgesch.
88, 118

Species resurrected from synonymy

I listed S. gravis Harold, 1869 as a synonym
of S. trifasciata L & G, 1837 (Barker 1979, p-
22). 8. gravis was a replacement name for S.
obscuripennis Saunders, 1868 which is a
homonym of S. (Themognatha) obscuripennis
Mannerheim, 1837. I have re-examined 10
Western Australian specimens in the South
Australian Museum collection. I find that they
are identical to a coloured photograph of the
type of S. gravis and I consider gravis a valid
species.

I also listed S. bucolica Kerremans, 1898 as
a synonym of S. trifasciata L & G, 1837. I
have re-examined 16 unidentified specimens in
the South Australian Museum collection, cap-
tured near Port Lincoln on the Eyre Peninsula,
S.A., and, on the basis of comparison with a
coloured photograph of the type, I consider
that they represent S. bucolica which I now
consider a valid species.

In the catalogue of Stigmodera (Castiarina)
I listed S. subnotata Carter, 1933 as a synonym
of S. subtincta Carter, 1933. It has been
pointed out to me that these are distinct, I
have examined material in the South Austra-
lian Museum and in the Western Australian
Museum and find that the male genitalia are
different; I consider both to be valid species.
In S. subtincta the last four visible abdominal
segments of males are testaceous and of
females metallic green. In S. subnotata all
visible abdominal segments are testaceous in
both sexes.

Selection of lectotype of S. rubriventris
Blackburn
Describing S. rubriventris Blackburn (1900)
p. 47 stated, “In one of the two specimens
before me.” The syntypes are males, one is in
the British Museum, the other in the South

Australian Museum. Most of Blackburn’s types
are lodged in the British Museum collection
and, all other things being equal, I consider
that the type should remain with the majority
of specimens. I hereby select the male speci-
men in the British Museum labelled ‘“W.A.
7556, S. rubriventris Blackburn” as the lecto-
type of the species.

e
=?
*.
».
-
4
»*
..

*
A
<

ney “ee
= b 4
aoe

at

5 mm

Fig. 1. Male Stigmodera (Castiarina) rudis Carter,

NEW STIGMODERA (COLEOPTERA: BUPRESTIDAE) 3

The abbreviations used in the text for
museum and private collections are as follows:
WADA, Western Australian Department of
Agriculture, South Perth; GB, Mr G. Burns,
Mornington, Vic.; ANIC, Australian National
Insect Collection, C.S.1.R.O., Canberra; JH,
Mrs J. I. Harslett, Amiens, Qld; AM, Austra-
lian Museum, Sydney; SAM, South Australian
Museum, Adelaide; WAM, Western Australian
Museum, Perth; MP, Mr M. Powell, Mr. M.
Golding & Mr T. M. S. Hanlon, Perth; NMV,
National Museum of Victoria, Melbourne.

Stigmodera (Castiarina) mimica sp. nov.
FIGS 2A, 3A

Types: Holotype: 3, Goldsborough near Gordon-
vale, Qid, 5.i1.1962, J. G. Brooks, SAM 121, 106.
Allotype: @, same data as holotype, SAM 121,
107. Paratype: | ¢, Marmor, Qld, xi.1946, W. du
Boulay, WAM.

Colour; Head, antennae black with blue reflec-
tions. Pronotum, scutellum black. Elytra red-
brown with black anterior margin, entire apex
black, curving upwards from margin 2/3 along
its length. Undersurface and legs black with
blue reflections. Hairs silver,

Fig. 2. (x3) natural size. A. Stigmodera mimica sp, nov.

inflata sp. nov.
aurea Sp. nov.

D. S. aquila sp. nov.

E. S. aliceae sp. nov.

Shape and sculpture: Head with small close

punctures, median groove between eyes,
muzzle short. Antennae: segments 1—4 ob-
conic, 5-11 toothed. Pronotum with small

close punctures; median basal fovea projecting
forwards as median impressed line to anterior
margin; glabrous basal notch on each side 1/3
from margin to centre; anterior to basal
notches a large irregular depression on each
side; anterior margin projecting forwards in
middle; basal margin bisinuate; laterally
rounded out from base, widest 1/3 from base,
rounded to apical margin. Scutellum: heart-
shaped; without punctures; convex in middle;
both lobes elongate. Elytra: three broad costae
on each side, scutellary, 3rd and 5th intervals;
other intervals punctate-striate and flat at basal
end, convex at apical end, punctured and
rough; laterally angled out from base, rounded
at shoulder, concave until after middle,
rounded to apex which is bispinose; spines very
small, rounded between; apices slightly diverg-
ing. Undersurface with small close punctures
and short close hairs. Last visible abdominal
segment truncate in both sexes. Male with

nn ®

B. S. hateleyi sp. nov. C.,
F. S. kiatae sp. nov. G.

4 S. BARKER

reduced tarsal pads on 2nd and 3rd legs,
pads absent on segments I-3 replaced with
single median spine, pad present on segment 4
only.

Size: Males 11.7 x 4.2 mm (2), Females 12,3
x 4.6 mm (1).

Distribution: North coastal Queensland.
General remarks; A lycid mimic belonging in
the §. sexplagiata group on the basis of male
genitalia and modified tarsal pads in male. An-
other member of the group S. erythroptera is
also a lycid mimic and has the same elytral
colour. S$. mimica differs from that species in
that it has fovea on the margins of the pro-
notum: S, erythroptera does not. S, mimica
has a different elytral pattern with more black
than in S. erythroptera. Elytral marking on §,
mimica is like that of S. nigriventris, also a
lycid mimic without costae on the elytra and
not a member of the S. sexplagiata group.

Stigmodera (Castiarina) hateleyi sp. nov.
FIGS 2B, 3B

Types: Holotype: &, Kiata, Vic., K. Hateley, SAM
121, 108. Allotype: 9, Kiata, Vic., K. Hateley,
SAM 121, 109. Paratypes: 5 3 & 2 9, same data
as holotype, SAM; 2¢' & 2 9, Wurarga, W. Aust.,
28.x.1978, M. Powell & M. Golding, MP, SAM;
1 2, Maranalgo Stn, W. Aust., 10.ix.1978, G.
Barron, MP; 1 9, Lake Grace, W. Aust., 16,x.1970,
K. & E. Carnaby, SAM; 2 ¢ & 1 9, Australia,
Blackburn & White, SAM.

Colour; Head, antennae, pronotum, most of
undersurface and legs dark blue with yellow
and green reflections, last visible abdominal
segment mainly brown in male. Scutellum
black. Elytra red-brown with following black
markings: narrow anterior margin; elongate
angled mark on each shoulder, rounded spot
between on suture all remnant of a pre-medial
fascia; post-medial fascia projecting forwards
in middle of each side and concave backwards,
not reaching margin; pre-apical mark extend-
ing over three intervals, concave forwards;
marks all connected down suture and covering
apex. Undersurface hairs silver.

Shape and sculpture: Head: closely and shal-
lowly punctured; median groove between eyes;

natrow ridge inside each antennal cavity;
muzzle short. Antennae: segments 1-3 ob-
conic, 4-11 toothed. Pronotum: shallowly

punctured; small median basal fovea extending
forwards to middle as glabrous line; basal
notch on either side closer to margin than to
middle; projecting forwards in middle of
apical margin; basal margin almost. straight;
laterally parallel-sided at base, rounded to

apex, widest 1/3 distance from base. Scutel-
lum: heart-shaped, with few punctures. Elytra
punctate-striate, intervals convex and pitted
with shallow punctures; laterally parallel-sided
at base, angled outwards, rounded at shoulder
then concave until after middle, rounded then
tapered to pre-apical area, then rounded to
apex which has no marginal spine, indented to
suture which has minute spines; apices not di-
verging, Undersurface shallowly punctured,
sparsely haired. Last visible abdominal seg-
ment truncate in males, rounded and narrowed
in females,

Size: Males 14.1 = 0.37 x 5.1 + 0.19 mm
(10). Females 14.6 + 0.36 x 5.6 = 0.13 mm
(8).

Distribution: Western Australia and Victoria.
General remarks; The elytral pattern and
colour are like those of §. rubriventris Black-
burn. However, S. hateleyi is a smaller spe-
cies, the male does not have a red abdomen
and male genitalia is different. Named after
Mr K. Hateley.

v" ¥

D E F G

Fig. 3. Photomicrographs of male genitalia of
Stigmodera (Castiarina) species: (A) mimica;
(B) hateleyi; (C) inflata; (D) aquila; (E) ali-
ceae; (F) kiatae; (G) aurea,

NEW STIGMODERA (COLEOPTERA: BUPRESTIDAE) 5

Sligmodera (Castiarina) inflata sp. nov,
FIGS 2C, 3C

Types Holotype: &, Baker's Creek Falls, Armi-
dale, N.S.W., 12.71,1979, T. J, Nuwkeswood, SAM
(21, 110, Allotype: 7, same data as holatype, SAM
T21, (iL. Purntypes: 6 4 & 13 9, Dungar Pally.
Armidale, NS.W.. 22/2401.1978, BS, & TO
Huwkeswood. SAM: 18 do & 3 2, same data as
holotype, SAM; 1 3 & 4 2, Dangar Falls, Arm
dale, NSW. 1O/145),1979, 7, J, Hawkeswood,
SAM,

Colours Head, antennae. scutellim, provotwm,
undersurface and legs bronze. Blytra yellow
with lollowing black markings: anterior tyar-
gin} pre-niedial faseia expanded at outside
ends into vittae Teaching basal and lateral thar-
Bins anteriorly and lateral margin posteriorly.
enclosing basal yellow spot on each side and a
spot on each shoulder, wide post-mechal fascia
reaching margin, concave in piddle of euch
side anteriorly and posteriorly: mark coverilig
whole apex: all marks connected down suture.
Hairs silver,

Shape and sculpture: Head: with close shal-
law puncnires; mediyn groove between eves;
muge shert. Antenme: segments J-4 ob.
conic, 5-t1 toothed. Pronotum closely pune
tured; shallow median basal fovea projecting
forwards to apex as median impressed line; an-
lerior margin slraight; basal murgin bisinviate;
Jaterally rounded froin huse, inflated io middle,
rounded to apex. Seutellum: shield-shaped;
concave in middle: with punctures, Elytra
punctate-striate, intervals convex more so ut
hase than apex; laterally angled out from base,
rounded at shoulder, convex until after the
middle then rountled und tapered to apex
which is bispinase; marginal spine larger than
sutural spine, rounded and indented between;
upices shehtly diverging. Undersurface with
shallow punctiires, covered in moderately long
hair) Last visible ahdominal segment sublrun-
ente in male. rounded in female. Tarsal pads
modified ow feet ef nrales: Ist leg, tarsal pads
absent on segments 1 & 2, reduced on 3; 2nd
Ter, tarsal pad absent on se#ment 1, reduced
on 2; 3rd leg, tarsal pad on segment 7 minute,
reduced an 2 & 4: on each foot central spine
present on undervurfaee of tursal segments |.
2 & 3 getting progressively smaller mm that
order.

Sizes Males 10.9 + 0.12 x 3.9 * 0,04 nim
(26). Females 117 > O17 x 4.4 = 0.0% mm
(211,

Distribulions New England district, N.S,W-

General remarks: Close to § wilsoni on simi
Jaritics in male genitalia, but Js Targer than that
species. The pronotum is More prominently in-
fated in the new species. In S. tei/soni the only
shoulder marking is « single vitea on each siile.
In 5. inflata the vittae are confluent with a pre
medial fascia, All specimens were collected on
the flowers of Bursaria spinosa,

Stigmodera (Castiarina) aquila sp. nov.
FIGS 2D, 3D

fypes; Holotype: ¢@, Gleneagle, W, Aust,
15,1971, K, 7. Richards, ANIC, Paratypes: 4 ¢,
same data as holotype, SAM & WADA.

Colour: Head, antennae, pronotum, scutellum,
undersurface and legs dull green, Elytra yel-
low wilh following black markings: anterior
margin; pre-medial fascia projecting Forwards
to lateral margin from anterior end only, post-
medial lascia reaching maryin, projecting for-
wards On each side from near margin; pre-
apical fascia not reaching margin; marks all
connected down sulure expanded at apex,
Undersurface hairs silver.

Shape und sculpture: Head: closely punctured;
median groove hetween eyes; muzzle sbort.
Antennae: segments 1-3 obconic, 4 4-toothed,
5-11 toothed. Protonum; with close punctures,
median basal fovea; projecting forwards in
muddle of apical margin; basal margin barely
bisinuate: laterally parallel-sided at ase.
rounded at apex, widest before the middle. Seve
tellumm: heart-shaped; with punctures. Elytry:
punctate-striate, intervals slightly wrinkled,
convex at apex; laterally angled out from base.
rounded at shoulder, faintly concave until
after middle then rounded to apex which is bi-
spinose; marginal spine larger than sutural,
wounded and indented between, apices slightly
diverging, Undersurface with close, shallow
punctures; moderately hgiry; hairs lone. Last
visible ubdominal segment truncate in male.
Size: Males 11,0 © 0,34 ¥ 3.9 = 6.19 mm (5).
Distribution; Darling Ratees. Western Austra-
lia.

General remarks: Male genitalia of this species
show similarities to those of 8. xcularis, How-
ever, S. scalariy has alternate intervals on the
elytra slightly raised and the anterior margin
af the pronotum is straight. Also the pre-
medial Fascia is tepresented by two spots on
the shoulder. S, sealaris ts found only io
eastern Australia and nelther species ts repre-
seuted in South Australia. All specimelis were
collected on 4 svertea sp.

6 S. BARKER

Stigmodera (Castiarina) aliceae sp. nov.
FIGS 2E, 3E

Types: Holotype: 3, Montacute, Mt Lofty Rgs, S.
Aust., 3.xi.1963, S. Barker, SAM 121, 129. Allo-
type: °, same data as holotype, SAM 121, 130.
Paratypes: 2 d, Murray Bridge, Lea, SAM; 1
Mt Lofty Regs, 4.xii.1964, N. McFarland, SAM; 5
3 & 5 92, same data as holotype, SAM; 21 3 & § 9,
Athelstone, S. Aust., 22.x.1966, McFarland &
Newberry, SAM; 1 3, Horsnell’s Gully, 12.xi.1967,
S, Barker, SAM; 2 3 & 2 9, Para Wirra N.P., S.
Aust., 7.xi.1971, S. Barker, SAM; 3 ¢ & 1 @,
Uraidla, S. Aust., 23/28.xii.1974, A. Wells, SAM;
1 ¢, Sommerton, S, Aust., SAM; 1 3%, S. Aust.,
SAM,

Colour: Head, antennae, scutellum, undersur-
face and legs blue. Elytra yellow with follow-
ing black markings with purple reflections:
basal margin; pre-medial fascia expanded at
outside ends into vittae reaching basal and
lateral margins anteriorly and lateral margin
posteriorly, enclosing basal yellow spot on
each side and one on each shoulder; wide post-
medial fascia reaching margin; mark covering
whole apex; all marks connected down suture.
Hairs silver.

Shape and sculpture: Head: closely punctured;
median groove between eyes; muzzle short,
Antennae: segments 1-4 obconic, 5-11
toothed. Pronotum: closely punctured; median
basal fovea extending forwards as thin glab-
rous line almost to apex; basal notches on each
side closer to margin than centre; projecting
forwards in middle of apical margin, basal
margin bisinuate; laterally parallel-sided at
base then rounded to apex, widest before
middle. Scutellum: heart-shaped; with punc-
tures, Elytra: punctate-striate, intervals con-
vex, more so at apex than base; laterally
angled outwards from base, rounded at
shoulder, concave until after middle, rounded
to apex which is bispinose; both spines small,
rounded and indented between; apices diverg-
ing slightly. Undersurface shallowly punc-
tured; sparse very short hair, Last visible abdo-
minal segment subtruncate in male, rounded in
female. Tarsal pads modified on feet of male:
tarsal pads absent on segments 1 & 2 on all
feet, single central spine on undersurface of
segments 1-3 on all feet in descending size
from 1-3,

Size: Males 8.3 = 0.1 x 2,9 + 0,04 mm (38).
Females 8.5 = 0.22 x 3.0 + 0.08 mm (14).
Distribution: South Australia.

General remarks: Close to S. cruentata (Kirby)
on similarities in male genitalia. The two spe-

cies differ im the pattern of reduction of tarsal
pads in males. Named after Mrs A. Wells.

Stigmodera (Castiarina) kiatae sp. nov.
FIGS 2F, 3F

Types: Holotype: 3, Kiata, Vic., xi.1967, K. Hate-
ley, SAM 121, 131. Allotype: 9%, Kiata, Vic.,
xi.1967, K. Hateley, SAM 121, 132. Paratypes: 3
3d, Big Desert, Vic., 17/19.xi.1977, G. Burns, GB:
2 3 & 2 9, same data as holotype, SAM: 1 ¢ & 1
Q, Big Desert, Vic., 4.xi.1976, G. Burns, NMV &
GB.

Colour: Head, pronotum, scutellum, undersur-
face and legs purple-bronze. Elytra yellow
with following dark blue markings: anterior
margin; vitta from outer edge of anterior mar-
gin running to lateral margin, enclosing elon-
gate yellow mark on shoulder; rounded mark
on suture remnant of pre-apical fascia; post-
medial fascia expanded on suture and middle
of each elytron not reaching lateral margin;
apical mark covering whole apex, all marks
connected along suture. Hairs silver.

Shape and sculpture: Head: closely punctured;
median groove between eyes; muzzle short.
Antennae: segments 1-3 obconic, 4-11
toothed. Pronotum closely punctured; small
median basal fovea; anterior margin straight;
basal margin barely bisinuate; laterally
rounded from. base, widest }-distance to apex,
slightly tapered to apex. Scutellum: heart-
shaped; without punctures. Elytra punctate-
striate, intervals flat at apex, more rounded at
base; laterally slightly angled out from base,
rounded at shoulder then concave until after
middle, rounded and tapered to apex which is
bispinose; apical spine large, sutural spine
small, rounded and indented between; apices
slightly diverging; sub-serrate lateral margin
from post-medial region to apex. Undersur-
face with close shallow punctures, larger on
pre-sternal sclerites than on abdomen; hairy,
hairs moderately long. Last visible abdominal
segment truncate in both sexes,

Size: Males 9,2 = 0.20 x 3.3 + 0.07 mm (7).
Females 10.0 = 0.21 x 3.6 = 0.07 mm (4).
Distribution: Big and Little Desert areas, Vic-
toria.

General remarks: This species is distinct from
any other on the basis of male genitalia.

Stigmodera (Castiarina) aurea sp. nov.
FIGS 2G, 3G
Types: Holotype: ¢, 18 km E. of Maryborough,

Qld, on Leptospermum sp., 5.xi.1975, S. Barker,
SAM 121, 133. Allotype: 9, same data as holo-

NEW STIGMODERA (COLEOPTERA: BUPRESTTDAE) 7

type, SAM 121, 134, Paratypes: 1 9, same data as
holotype, SAM; 4 2, Maryborough, Qld, xi,1946,
E. Smith, NMV:; L & . Maryborough, 12.xii,
MIM, ANTC; | & Indooroopilly, 11 xi.1933,
ANIC; 2 ¢. Wide Bay, Qld, AM: 1 ¢, Rockhamp-
ton, Qld, 10,1,1946, JH; 2d & 3 2, Qld, du Boulay,
WAM.

Colour, Head, antennae, pronotum, scutellum
black with olive-green reflections, Undersur-
face and legs olive green, Elytra yellow at
basal end, apical end bright orange, with the
following black markings: basal margin; short
vitta on each shoulder reaching lateral and
basal margins; basal elongate rounded mark on
sulure; post-medial fascia expanded on suture
and close lo murgin, reaching margin; pre-
apical fascia not reaching margin; mark cover
ing apices: all marks connected down suture,
the last two broadly, Hairs silver.

Shape and sculpture; Head with small shallow

punctures; median groove hetween eyes:
muzzle short. Antennac; segments 1-3 ob-
conic, 4-11 toothed. Pronotum with small

shallow punctures: basal fovea projecting For-
wards to middle as glabrous linc; basal notch
on each side closer to margin than to middle;
anterior margin projecting forwards in middle;
basal margin bisinuate; laterally rounded from
hase, gradually narrowing to apex, basal angles
turned upwards. Scutellum: heart-shaped,
sides folded inwards from centre; without
punctures, Elytra punctate-striate, the intervals
convex and smooth, the margin flattened; luter-
ally angled out from base, rounded at
shoulder, concave until afler middle, rounded
and narrowed to apex which is bispinose; mar-
ginal spine large, sutural spine very small,
deeply indented between, Undersurface with
close shallow punctures: moderately hairy,

hairs medium Jength. Last vistble abdominal
segment, rounded and pushed in in middle in
both sexes. Males have modified tarsal! pads on
3rd leg. segments | & 2 have reduced pads,
spines not present,

Size: Males 8.9 + 0.9 x 3.2 ~ 0.03 mm (9),
Females 10.5 = 0.09 x 3.9 + 0.05 mm (9).
Distribution: South and central coastal Queens-
land.

General remarks: Close to 8, alternecostata on
the basis of similarities in male genitalia, How-
ever, §. aurea is a smaller species. the prono-
tum 1s continually narrowed from the base
whereas ii S, allernecosiata the pronotum 1s
widest before the middle. Sculpture and colour
pattern of the elylra also differ,

Acknowledgments

1 thank the following for their assistance:
Dr A. Neboiss, National Museum of Victoria;
Mr G. Holloway, Australian Museum; Dr
T. F. Houston, Western Australian Museumy
Mr K. T. Richards, Western Australign
Department of Agriculture; Dr M, W. R, de
V. Graham, Hope Department of Zoology
(Entomology), University of Oxford; Dr M,
Uhlig, Muscum of Natural Science, Hujnboldt
University, Berlin; Miss C. M, H. von Hayek,
British Museum (Natural History); Mr E, EF.
Adams, Edungalba: Mr G. Burns, Morning-
ton; Mr and Mrs K, Carnaby, Wilga; Mrs J,
Harslett, Amiens: Mr K. Hateley, Kiata; Mr
T.J, Hawkeswood, Nedlands; Mr J. Macqueen,
Toowoomba; Herr H, Mihle, Neusaess; Mr
M. Powell, Como; Miss H. Vanderwoude and
Mrs A, Wells, Department of Zoology, Uni-
versity of Adelaide, The Australian Biological
Resources Committee provided a grant-in-aid
of research

References

Barker. S. (1979) New species and a catalogtc
of Stigmodera (Castiarina) (Coleoptera, Bup-
restidae). Trans, R. Soe. 8, Aust. W3, 1-23,

Biackeurn, T, (1900) Further notes on Austra-
lian Coleoptera, with descriptions of new genera
and species, Ibid. 26, 35-68,

Carter, H. J, (1934) Australian and New Guinen
Soleoplera notes and new species No. IL Prec.
Linn. Soc. N.S.W. Sa, 252-269,

SOME STRONGYLE NEMATODES (AMIDOSTOMUM SPP.) FROM
AUSTRALIAN BIRDS

BY PATRICIA M. MAWSON

Summary

Amidostomum acutum is recorded from Anas superciliosa, A. platyrhynchos, Stictonetta naevosa,
Querquedula gibberifrons, Tadorna radjah and Himantopus leucocephalus, A. anseris from Cereopis
novaehollandiae; A. cygni from Cygnis atratus; A. biziurae from Biziaura lobata. Measurements of
most specimens examined and morphological notes on A. cygni and A. biziurae are given. A new
species, A. tribonyx, close to A. acutum, but distinguished by shorter spicules and very large
papillae, is described from Tribonyx ventralis.

SOME STRONGYLE NEMATODES (AMIDOSTOMUM SPY.) FROM
AUSTRALIAN BIRDS

by ParriciA M. Mawson*

Summary

Mawson, P. M.

(1980) Some strongyle nematodes (Amidostomum spp) from Australian

birds. Troms. Ro Soe. 8. duet, WC), 9-42, 29 February, 1980.

Amidastomum acutin is recorded from Anas superciliosa, A. platyrhynchos, Stictonetta
natvasa. Querquedula gibberifrons, Tadarna radjah and Himartopus leucocephalus, A. anseris
from Cereopsis novaehollandiaes A. eyeni from Cypnuy atratis; A. bizlurae from Bizlura
lehuia, Mensurements of most specimens examined and morphological notes on A. eveni and
A. biziurae are wiven, A new species, A. tribonyx, clase to A. acutium, bot distingwished by
shorter spicvles and very large cervical papillae, is described from Tribenyx veniralis,

(nirodection

Neniatedes of the genus Amidostomum
appear to be restricted to waterfowl; almost all
records are from anseriform and ralliform
birds, and a few fram eharadriform birds, ‘Lhe
genus was studied in some detail by Czaplinskt
(1962), Gf the 16 species then recorded, he
synonymised Ll, Pour species have been de-
scribed subsequently,

In the present work three of the species
recognised fy Czaplinski are identified from
Australian birds, one species he considered a
synonym is resurrected and a new species is
described.

Measurements of specimens examined are
tabulated, but those indicating the position of
herve ring, cervical papillae and excretory
pore are omitted in some cases. These
struciures, cspevially the first two, are par-
ticularly obscure in some species. The new
species is noleworlhy because of the unusually
large cervieal papillae.

Types of the new species will be deposited
in the South Australian Museum, and all other
material belongs to the Australian Helmintho-
logical Collection at present in the South Aus-
tralian Museum,

Awidostomum acutum (ftundaht)

Strangylus acutuy Laundahl, 1848.

Hosts und localilies; Anay supereiliosus Gmelin:
Hamley Bridge S.A. (2 cf), Westbury, Tas. (24 3,
(8 2): 4, plaryrhyuehas, Plinders t, Vas, 1 2, t
Qo; Srlerenena naevosa (Gould), Bool Lagoon,
S.A. (2 Bs Querquedulu gibberifrans S. Miiller,
Naracoorte, SA, (1 of); Querquedula sp., Ade-
laide, SA. 01 Sd: Tadorne radjalt Garnot, Humpry
Doon, NT, (1 @); Mimantopus [eucocephalus
Gould, Petermann Ra,, N_T, (1 &, 2 2),

* Department of Zoology, University of Adelaide,
Box 498, G.P.O., Adelaide, 8. Aust. 5001,

In his work on Antidoyromum, Czraplinski
(1962) gives a good account of A, aeurtin,
and referred a number of species fo is syno-
nymy. Except for A. dizivrae Johnston &
Mawson, this synonymy has been adopted
here, Reasons for this exception are given in
the account of 4, biziurae.

Measurements of specimens from <Anay
superciliaxus, which are most numerous and jo
good condition, are given in Table 1.

Amidostomum anseris (Zeder)
FIG. 2

Strongylus anseris Zeder, 1800.
Host and locality: Cereapsis novaehollandiae
Latham, trom Flinders £., Tas. (5 d, 7 9).

These specimens agree generally with the
description, figures and measurements given by
Czaplinski in his Summary of the species. The
only (and slight) difference is that the anterior
lip of the vulva, and in two specimens both
anterior and posterior lips, are enlarged (Fig.
2), Czaplinski describes this region as “vulva
provided with conspicuous hemispherical
bulge”, Measurements of these specimens are
given in Table 1,

Amidostomum biziurae Johnston & Mawson
FIG, |
Amidostomum biziurae Tohnston & Mawson, 1947.
Host and localities: Biziura labata (Shaw) from
Goolwa, S.A. (3 cd, 2 9), Barren Box Swamp,
N.S.W. (5 3, 5 2).

This species, originally described from one
female, was cedescribed from new material of
both sexes (Mawson 1959), The specimens
recorded here from the type host species and
new localities agree with this description.

Czaplinski (1962) placed 4. bizinrae in the
synonymy of A, aeutunr, apparently not having
seen this redeseription, However, he dis-
counted the prajeclions around the mouth,

10 PATRICIA M. MAWSON

TABLE 1. Measurements of Amidostomum spp. recorded here (um unless otherwise stated).

Species A, acutum A. cygni A, anseris A. tribonyx
Anas Himantopus Cygnus Cereopsis Tribonyx
Host superciliosus leucocephalus atrata novaehollandiae ventralis
Male (number) (5) (1) (2) (4) (3)
Length (mm) 9,2-11.4 7.2 15-17 12,3-15.5 7.6-9.7
Oesophagus 640-760 1280-1500 1450 650-856
rods occupy 90-95% 99% 81-84%
A—nr 250-300 350-400 260-280 240-250
—cp 360-410 520-600 350-400 280-350
—exp 300-420 520-600 320-340 -
spicules 130-140 140 190-200 280-312 100-110
gubernaculum 60-70 80 105-110 110-115 30
bucc. caps: length 9 8 12-13 15-16 10
ext. diameter 11-12 13 35 31-35 18-19
Female (number ) (5) ( (2) (4) (3)
Length (mm) 13.5—17.0 13.6-13.7 22.0-22.1 19.5-21.0 7.6-10.0
Oesophagus 760-850 1250-1550 1420-1650 650-730
rods occupy 96% 99% 84-86%
A—nr 250-300 350-390 350-390 210-240
—cp 300-400 500-550 500-550 300-310
—exp 300-490 500-505 505-550 _
bucc. caps: length 10-11 15-16 15-16 10-12
ext. diameter 12-16 17-18 36-40 36-41 18-22
tail 250-280 250-260 250-260 240-330 220-300
vulva—posterior end (mm) 2.4-2.8 2.7-2.8 3.8-3.9 2.7-3.1 1,8-2.5
eggs—L 80-90 91-95 89-90 90-95 90-95
—Br 50-51 42-45 50-51 45-55 50-52

regarded by the authors as one of the main
specific characters, as being enlarged labial
papillae. In fact these are not labial papillae,
but cuticular structures, containing no nerve or
other hypodermal tissue. The four labial papil-
lae and two amphids are seen behind the pro-
jections (Mawson 1959, figs 4-5). The projec-
tions are obvious on all specimens, and arise
in association with the anterior edge of the
buccal capsule.

The other striking feature of the species is
the embossed cuticular annules, which appear
on all specimens, and have not been seen in
any other species. A more detailed figure is
given of the spicules.

Amidostomum cygni Wehr
FIGS 3-5

Amidostomum cygni Wehr, 1933.
Host and locality: Cygnus atratus Latham, Ade-
laide, S.A. (13 3, 16 9).

This species was recorded from C. atratus
(syn. Chenopis atrata) from Tailem Bend,
S.A. by Johnston & Mawson (1947), but that
material did not include the posterior end of a
male. The present material is more plentiful
and a closer study has been made.

The measurements (Table 1) are generally
larger than those given by Wehr (1933) or by

Ryzhikov (1959). Czaplinski (1962) regards
A. similis Freitas & Mendonca (1954) from a
South American swan a synonym of A. cygni:
the only difference being in the greater size and
in that. the authors noted three instead of two
(Ryzhikov) branches to each spicule, Wehr
simply states that the spicule is “similar in
shape to those of other species of the genus”.
In the Australian specimens there are three
branches, one much shorter and thinner, the
other two lying close together except at the
tips where one ends bluntly, the other is
splayed out. In the expanded state all branches
support a membranous structure which appears
to be globular rather than fan-shaped (Fig. 4).

Amidostomum tribonyx sp. nov.
FIGS 6-10
Host and localities: Tribonyx ventralis Gould,
from Swan Reach (3 3, 3 9), and Taperoo (1 6,
19), S.A.
Holotype male, SAM, V1864. Allotype female,
SAM, V1865.

Lateral alae absent, buccal capsule stoutly
built, with single, dorsal tooth reaching almost
to anterior edge of buccal capsule, Oesophagus
widens slightly posteriorly. Cervical papillae
prominent cuticular projections.

Male: Spicules bifid, each branch alate,
rounded at distal end, the more ventral branch

AUSTRALIAN NEMATODES (AMIDOSTOMUM) 1]

100 pm

Fig. 1. Amidostomum hiziurae, spicule.

Fig. 2. A. anseris, vulvar region.

Figs 3-5. A. cygni. Fig. 3, anterior end; Fig. 4, posterior end showing spicules partly everted; Fig.
5, posterior end of female.

Figs 6-10. A. tribonyx. Fig. 6, anterior end; Fig. 7, head; Fig. 8, bursa, ventral view; Fig. 9, dorsal
ray; Fig. 10, posterior end of female.

Figs. 1, 8 & 9 to same scale; Figs 3 & 5 to same scale: Figs 4 & 10 to same scale.

12 PATRICIA M. MAWSON

slightly longer. Gubernaculum poorly sclero-
tized, about + length of spicules. Bursal rays
typical of genus( Figs 8 & 9).

Female: Tail widens about midlength, at level
of phasmids, then narrows to rounded tip.
Vulva at about + body length from posterior
end, with swollen anterior lip.

This species differs from A. acutum in the
smaller spicules and gubernaculum, and from

all congeners by the large cervical papillae.
Measurements are included in Table 1.

Acknowledgments
The birds from which the material used in
this work were obtained were sent to me by Mr
H. Frith (C.S.I.R.O., Canberra), the Northern
Territory Museum, the South Australian
Museum, and Mr R. Green (Victoria Museum,
Tasmania).

References

CZAPLINSKI, B. (1962) Nematodes and acantho-
cephalans of wild and domestic Anseriformes
in Poland. 1. Revision of the genus Amido-
stomum Railliet & Henri, 1909. Acta Parasit.
Polonica 10, 125-164.

Freitas, J. F. T. & Menponca, M. J. (1954)
Novo tricostrongylideo parasito de cisne europen
(Nemaetoda: Strongyloidea). Rev. Brasil. Biol.
14, 397-400.

Jounston, T. H. & Mawson, P. M. (1947) Some
avian and fish nematodes, chiefly from Tailem
Bend, South Australia. Rec. S. Aust. Mus. 8,
547-553.

LUNDAHL, C. (1848) Helminthologische Beitrage.
1. Bemerkungen iiber zwei neue Strongylus
Arten. Notis. sdllsk. Fauna Flora Fenn. Forh. 1,
283-287.

Mawson, P. M. (1959) Some nematode parasites
from Australian hosts. Trans. R. Soc. S. Aust.
82, 151-162.

Ryzuikov, K. M. (1959) Nematodes in the heart
of a swan. [In Russian] Priroda 48, 119.

Wenr, E. E. (1933) Description of two new
parasitic nematodes from birds. J. Wash. Acad.
Sci. 23, 391-396.

ZeDER, J. G. H. (1800) Erster Nachtrag zur
Naturgesichte der Eingeweidewiirmer mit Zu-
faissen und Anmerkungen herausgegeben. 320
pp. Leipzig.

CIRCULAR STRUCTURES IN A LATE PRECAMBRIAN SANDSTONE:
FOSSIL MEDUSOIDS OR EVIDENCE OF FLUIDIZATION?

BY P. §S. PLUMMER

Summary

Small circular structures have been found preserved on the basal surface of a Late Precambrian
sandstone within the Moorillah Formation in the Flinders Ranges, South Australia. Although their
mode of preservation and general appearance suggest a biogenic origin, it is believed that a non-
biogenic process involving gas migration and sediment fluidization was responsible for their
formation.

CIRCULAR STRUCTURES IN A LATE PRECAMBRIAN SANDSTONE:
FOSSIL MEDUSOIDS OR EVIDENCE OF FLUIDIZATION?

by PS. PLUMMER*

Summary

Promune, PS. (1980) Circular structures la a Late Precambrian sandstone: Fossil medusoids
ov evidenve of Mhudization Trans, R. Soc. §. Aust. 104 (1), 13-16, 29 February, 1980.
Small cirenla: structures have been found preserved on the basal surface of a Lale
Precambrian sandstone within the Moorilah Formation in the Flinders Runges, South Australia.
Although their mode of preservation and general appearance suggest o biogenic origin, it is
believed that 8 non-hiogenic protess involving gas migration and sediment fluidization was

responsible for their formation.

Introduction

Recently found within the central Flinders
Ranges of South Australia was a float sample!
of ripple cross-luminated fine sandstone on the
basal surface of which are preserved a number
ot small, roughly circular structures of prob-
lematic origin. The sample was collected about
10 knv southeast of Wilpena Chalet (lat.
31°36'S, long, 138°40°E) near the contact be-
tween the Moorillah Formation and the under.
lying Moolooloa Formation of the Late Pre-
cambrian Brachina Subgroup (Plummer 1978),
Tts lithology is typical of the lower portion of
the Moorillah Formation and, although the
sample 15 from floal, it is considered to have
come fram this horizon,

A passible biogenic origin for these struc-
tures is suggested if their mode of preservation
(i.e, on the basal surface of a sandstone lying
directly above w clay layer) is compared with
that of the many soft-bodied animal fossils
fond within the Late Precambrian Ediacara
assemblage (sce Wade 1968). This, along with
their overall cireular shape and concentric
rings tarking their perimeters, suggests a
resemblance to small medusoid fossils. If this
were so, they would form perhaps the oldest
fossil coelenterale oecurrence known, lying
2500 m helow the level of the Ediacara assem-
blaee. and well beneath the presently knows
oldest occurrence in the Bonney Sandstone
(formerly the “Red Pound") of the Pound
Suheroup where small medusoids ure found in
association with sinuous tracks (Wade 1970).

*Centre for Precantbriun Research, Department
af Geology, University of Adelaide, Adelaide,
GPO, Box 498, 8. Aust, SOOT,

1 Sample in Department of Geology, University of
Adelaide: cat, No, 469/RPB1 LO.

Description

The basal surface of the sample measures
16x 171 em and shows several roughly circular
structures which range 6-20 mm in diameter
with relief up to 2 mm (Pig, 1a). The centres
of these structures are generally shallowly
domed (Fig. lb), although some ate Slat, or
display a central depression (Fig. 1¢). Poorly
ilefined step-like concentric rings are present
toward the edges of some of the structures
(sce Fig. 1b), whilst surrounding them is the
suggestion of a Mat skirting rim up ta 8 mm
wide, Shaly material is patchily preserved om
this basal surface of the sample, indicating that
the ripple cross-laminated sand formed casts
of these structures which were developed in,
or present on an underlying clay bed.

Interpretation

These structures are comparable to the fossil
medusoids classed as Prefelvella Torell 1870,
which includes the taxon Medtsina Sprivg
1949. Protolyella 3s represented by circular
bodies up to 5 cm diameter, comprising a
smooth central area and an outer annular
yone, separated by an annular furrow (Moore
1956; Glaessner & Wade 1966). Although the
central area of these fossils can show faint con-
centric markings, the outer annular zone
usually displays numerous radial grooves not
present on the structures described herein.

Other widely distributed circular organic
structures of Late Precambrian age ate the
planktonic remains known as Chnaria Walcott
L899, These fossils, however, are usually pre-
served as black, discoidul compressions, com-
manly carbonaceous, and having a maximum
diameter of 5 mm (Hofman 1977). As such,
the structures described here are too large and
of the wrong style of preservation ta be
Chuaria.

14 P. S. PLUMMER

Fig. 1. Circular structures. (a) casts on basal sur-
face of sample; (b) shallowly domed centre;
(c) central depression.

Maxson (1940) and Cloud (1960) have in-
terpreted structures comparable to those under
discussion in terms of gas escape and fluidiza-
tion phenomena, Such phenomena are well
known to igneous geologists (e.g. Reynolds
1954; Holmes 1965), but their influence upon
sedimentation has been little discussed. Mills
(1969), however, gives an excellent account
of structures formed experimentally by ‘cold’
fluidization which closely resemble the struc-
tures discussed herein. Although his experi-
ments were concerned with the origin of
craters on the lunar surface, the phenomenon
of ‘cold’ fluidization is equally applicable to
much smaller scale structures.

The process of ‘cold’ fluidization involves
the migration to the surface of gas trapped
within a stationary bed. In a cohesive sedi-
ment, such as a clay, gas migration can cause
the bed to expand and display the flow proper-
ties of a liquid. Often the gas flow finds pre-
ferred channels of escape and, if the flow is
great enough, bubbles may form in the chan-
nels. On reaching the surface the bubbles
either burst, or dissipate gently (depending on
the degree of bed cohesion) causing the bed
to then contract and subside. Ring slumping
and faulting often accompanies the subsidence,

Burial by Capping | Sand

nvr wha

ls

\ I
x ee ee

Gas Migration

Fig. 2. Schematic diagram of crater formation by
‘cold’ fluidization process.

and the net result is the production of circular,
crater-like structures.

Such a process could have operated to pro-
duce the structures found on the sample de-
scribed here. Sedimentologic analysis has indi-
cated that the Moorillah Formation was depo-
sited dominantly within an intertidal environ-
ment (Plummer 1978). During a period of low
tide, gas or air trapped within sandy sediment
could have migrated upward through a cap-
ping clay layer (possibly deposited during the
slack-water stage of the previous high tide) to
burst and form small craters on the surface.
Gas escape phenomena are known to occur on
present-day intertidal flats (Reineck & Singh
1975, Fig. 61), and the burial of such craters
by a layer of sand during the following incom-
ing tide could then preserve them as casts on
its basal surface (Fig. 2). Such gas produced
structures have been called ‘evasion marks’ by
Cloud (1960), who differentiated them from
‘contact marks’ produced by single, or
numerous gas bubbles blown across the sedi-
ment surface; raindrop impressions (Lyell
1851, Shrock 1948); and other ‘pit-and-
mound’ structures produced by compaction-
induced dewatering (Kindle 1916, Schofield &
Keen 1929), or current flow stress (Karez et
al. 1974).

Other organic structures?

Also reported from the Moorillah Forma-
tion are other structures which were initially

LATE PRECAMBRIAN CIRCULAR STRUCTURES 15

believed to be of organic origin, The solitary
trace-like murking BSanyerichnius dalgarnoi
Glacssner 1969 was deseribed as the track of
an animal possibly "related to primitive mol-
lusea without mineralized shells” (Glyessner
1969, p. 379). However, Jenkins (1975, p, 19)
regarded jl “a unigue and accidental set of
markings made by a tethered implement being
moved by a current’ (eg. possibly ribbon-like
algae such as are known from cocks of similar
age in the U.S.S.R.), A close resemblance is
also noted between this marking and certaln
lineations produced by vorticity along lines of
wind flow (Whitney 1978, Fig, 4B). Also.
abundant cylindrical to coni¢al structures
ecelir in this formation that resemble either
certain fossil burrows of other problematic
Precambrian sack-shaped fossil organisms such
us Namalia Germs 1968, the ‘Erniettomorpha’
of Pilg (1972) and Baikalina Sokolov 1972.
These structures, however, are pol-casts pro-

duced by the helical scouring action of water
currents (Jenkins ef al, ih prep.).
Conclusion

Of the two possible origins presented to
explain the structures discussed herein (viz.
the fossilization of small medusoids, or gas
escape and ‘cold’ fluidization phenomena) the
latter, non-biogenie origin is fuvourcd, Simi-
larly, olher circular structures found in depo-
sits of Precambrian age that have previously
been deseribed as problematica or organic
remains (e.g. Bassler 1941, Alf 1959, Shep-
heed & Thatcher 1959, Johnson & Fox 1968)
should be feViewed in lerms of formation by
fluidization phenomena, or other imorganic
processes (as suggested by Cloud 1940, 1973},

Acknowledgments

Dr R. J, F. Jenkins provided much discus-
sion on Precambrian life forms and traces,
whilst both he and Dr V. A, Gostin evitically
read the manuscript.

References

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Proterozoic Buss Formation, Grand Canyon,
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Basser, R, S. (1941) A supposed jellyfish from
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Croup, P. BE. Je, (1960) Gas as a sedimentary
and diagenetic agent. Aim. J. Nei 258A, 35-45.

—~— (1973) Pseudofossils! A plea for caution,
Geolopy 1, 123-127.

Grems, G. J, C, (1968) Discovery of a new fossil
in the Nunia System, South-West Africa, Nutire
259, 53-54,

GLaessvNeR, M, Fb, (1969) Trace fossils from the
Precambrian aud basal Cambrian. Lethaia 2%,
369-393.

GLAgssNek, M. F. & Wank, M-. (1966) The Lute
Precambrian fossils from Edivears, South Aus-
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Hormawn, H, J. (1977) The problematic fossil
CInaria trom ihe Late Precambrian Uinta
Mountain Group. Uiih. Pree. Res, 4. 1-11.

Hoimers, A. (1965) “Principles of Physical
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Jenkins, Ro J F. (1975) The diversification of
early metazoans. Absir Ist Aust. geol. Conv.
weal. Soc. Attist, 19-20,

Jenkins. R$. F.. Plunmier, P. S. & Montarry,
K. C. fin prep.) A South Australian find of
abiindant Late Precambrian pseudofossils fe
thatkubly fesembling |rue fossils of comparable

age.

Jonwsomn, H. & Fox, 8. K.. Jr. (1968) Dipleurozea
from Lower Silurion of North America. Scievice
THz, (19-720,

Kancz, 1, Enos, P. & Lancinrr, G, (1974)
Stuctires generated in fluid stressing of freshly
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2, PRY 290,

Kenoce, E. M. (1916) Small pit and mound
structures deVeloped during sedimentation. Geel,
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Recent, Triassic and Carboniferous periods.
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and possible implications fur the orwgin oF lunar
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vetlebrate Puleuntology. Purt FP, Coelenteraia.”
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Bau und Systematik. Palaeeentupraphica A139,
134-170. /

Prummepr, PS, (1978) The stratigraphy of the
lower Wilpena Group (Late Precambrian),
Flinders Ranges, Sourh Australian. Trans. R. Soc.
S. Ansr 102, 25-38.

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logical process, and its bearing on the problem
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Scnpmieno, RoR, & Kee, B.A, (L824) Rigidity
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Prokl, J, 1t4-123.

16 P. S. PLUMMER

SpricG, R. C. (1949). Early Cambrian ‘jellyfishes’
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— (1970) The stratigraphic distribution of the
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Soc. Am. 89, 1-18.

SYSTEMATIC STATUS OF KANKANOPHRYNE HEYER & LIEM (ANURA:
LEPTODACTYLIDAE)

BY M. J. TYLER & MARGARET DAVIES

Summary

The diagnostic characters of Kankanophryne Heyer & Liem (1976) that distinguish it from the
Pseudophryne Fitzinger (1843) were stated to be in the form of the sacral diapophyses and the
number of slips of the Musculus depressor mandibulae. Our survey of these features in the two
genera fails to support any distinction; hence we propose that Kankanophryne be referred to the
synonymy of Pseudophryne.

SYSTEMATIC STATUS OF KANKANOPHRYNE HEYER & LIEM
(ANURA: LEPTODACTYLIDAE)

by M. J. TyLer & MarGarer Davyies*

Summary
Ty.eR, M. J. & Davies, M. (1980) Systematic status of Kankanophryne Heyer & Liem
(Anura: Leptodactylidae). Trans, R. Soc. 8, Aust. 104(1), 17-20, 29 February, 1980,

The diagnostic characters of Kaukanophryne Heyer & Liem (1976) that distinguish it
from Pseudophryne Fitzinger (1843) were stated to be the form of the sacral diapophyses
and [he number of slips of the Musculus depressor mandibulae, Our survey of these features
in the two genera fails to support any distinction; hence we propose that Kankanophryne be

referred to the synonymy of Psevdophryne.

Introduction

Heyer & Liem (1976) undertook an analysis
of intergeneric relationships in Australian
myobatrachid (leptodactylid) frogs. Amongst
their consequent proposals was the erection of
the genus Kankanophryne for Pseudophryne
occidentalis Parker, described from Western
Australia (Parker 1940) and reported from
South Australia by Tyler (1972). The rationale
for Heyer & Liem’s action was the stated
existence in P. occidentalis of a small series of
character states not shared by congeners. The
authors stated that these were “differences best
reflected at the generic level” (1976, p, 5).

Tyler (1978) failed to adopt the new
generic name, but Cogger (1978) has included
Kankanophryne in the revised edition of his
“Reptiles and Amphibians of Australia”, Be-
cause Our observations on the definitive
characteristics involved are at variance with
those published by Heyer & Liem, we present
these data here to clarify the systematic status
of Katikanophryne.

Our retention of the name Leptodactylidae
instead of adopting Myobatrachidae reflects
the absence of morphological substantiation
for the latter step. This matter is discussed in
detail by Tyler (1979).

Material and Methods

We have examined the external features,
myology and osteology of representatives of
the following species: Pyeudophryne bibroni
Giinther, P. coriacea Keferstein, P. guentheri
Boulenger, P. occidentalis and P. semimar-
morata Lucas. Specimens are deposited in the
collections of the South Australian Museum

* Department of Zoology, University of Adeluide,
Box 498, G.P.O., Adelaide, S.A. S001.

(SAM), Western Australian Museum (WAM)
and Department of Zoology, University of
Adelaide (UAZ).

Fig, L, Augles of sacral diapophyses measured.
SDAR: anterior angle of right sacral diopo-
physis; SDER: gpredtest expansion of right
sacral diapophysis; SDPR: posterior angle of
right sacral diapophysis.

18 M. J. TYLER & MARGARET DAVIES

Muscles were examined with the aid of the
iodine/potassium iodide stain developed by
Bock & Shear (1972). Bones and cartilage
were examined employing either cleared and
Alizarin Red staining or the differential Aliza-
tin Red/Alcian Blue stain (Davis & Gore
1947; Dingerkus & Uhler 1977). The method
of measurement of sacral diapophyseal angles
follows Trueb (1977) and is demonstrated in
Fig. 1.

Generic diagnoses

Heyer & Liem’s diagnoses of Pseudophryne
Fitzinger and Kankanophryne are identical in
the condition of the following features:
separation of cervical cotyles, lack of Musculus
omohyoideus, lack of columella, texture of
belly skin, form of toes, presence of metatarsal
tubercle and mode of reproduction.

The diagnoses differ in the following
respects:
Vomerine bones. Present or absent in

Pseudophryne; absent in Kankanophryne.

Sacral diapophyses: Broad in Pseudophryne;
narrow in Kankanophryne.

Depressor mandibulae:
fascia absent in Pseudophryne;
Kankanophryne.

It follows that the critical characteristics for
the recognition of Kankanophryne are the
conditions of the sacral diapophyses and de-
pressor mandibulae.

Slip from dorsal
present in

Expansion of sacral diapophyses

The definition of character state 12 in
Heyer & Liem’s paper is as follows: State 0:
sacral diapophyses expanded; State 1: sacral
diapophyses rounded uniformly, in at least
some species. They then proceed to score
P, occidentalis as State 1 and the remaining
species of Pseudophryne as State 0. Generic
descriptions of Kankanophryne and Pseudo-
phryne refer to “narrow” and “broad” sacral
diapophyses respectively.

The maiority of anurans have moderately
expanded sacral diapophyses (designated
“dilated” by Trueb (1973) ). Amongst the
Ranidae, however, the sacral diapophyses are
narrow and usually directed posterolaterally
whereas at the other extreme of the spectrum
(in the Bufonidae), sacral diapophyses are
very broadly expanded. Lynch (1971) con-
sidered that any distinction between the
degree of dilation of the sacral diapophyses
exhibited by some of the Australopapuan

leptodactylid genera he examined, is a very
fine one and probably is not defensible.
Trueb’s (1977) investigation into the osteology
of a population of Hyla lanciformis (Cope)
indicated that a degree of uncertainty exists
about the reliability of vertebral characters.
Vertebral anomalies are common amongst
anurans, particularly in the form of bilateral
asymmetry, and presence of additional features
such as transverse processes on the coccyx in
some individuals (unpublished observations).
Trueb (1977) observed that a low coefficient of
variability in the shape and orientation of the
anterior edge of the sacral diapophyses
occurred in her study population, indicating
that some sacral features are reliable for
systematic purposes.

We have carried out Trueb’s measurements
on the species examined by us and the results
are shown in Table 1. The vertebral columns

TABLE 1. Measurements of sacral diapophyses (see
Fig. 1) in individuals of Pseudophryne and Kan-
kanophryne.

Sacral angle measurement in degrees

Species SDAL SDAR SDEL SDER SDPL SDPR
P. bibroni 83.5 75.0 41,0 49.5 55.5 55.5
P. coriacea 68.5 72.0 49.0 39.5 62.5 68.5
P. guentheri 82.5 85.5 50.0 39.0 47.5 55.5
P.semimarmorata 69.5 74.5 53.5 43.0 57.0 62.5
K. occidentalis 80.0 78.5 49.0 52.5 51.0 49.0

SDAL(R): Anterior angle of left (right) sacral dia-
pophysis;s SDEL(R): Greatest expansion of left
(right) sacral diapophysis; SDPL(R): Posterior
angle of left (right) sacral diapophysis.

of the species considered are shown in Fig. 2.
Whilst recognising that the measurements in
Table 1 have been made on individuals rather
than populations, it can be seen that there is
no significant difference in the expansion of
the sacral diapophyses between P. occidentalis
and the other species of Pseudophryne
examined by us, We can find no justification
for Heyer and Liem scoring the expansion of
the sacral diapophyses in P. occidentalis as
“narrow” in comparison with other Pseudo-
phryne.

Depressor mandibulae

Griffiths (1954, 1959) demonstrated the
existence of interspecific divergence in the
form of the M. depressor mandibulae, and the
way in which this divergence could be em-
ployed for systematic purposes. Griffiths

SYSTEMATIC STATUS OF KANKANOPHRYNE HEYER & LIEM 19

Fig, 2. Vertebral columns of (A) Pseudophryne semimarmorata UAZ B536, Koonwarra, Vic,; (B)
P, vecidestaliy, SAM R17522, approx. 100 km S of Balladonia Hotel, W.A. (33°13'S, 123°27'B);
(C) P. gueatheri, UAZ B539, Porrestfield. W.A.; (D) PB. bihroni, UAZ A577, Tandanya Farm, Kan-
garoo Is; (6) P. coriacea, UAZ BS37, Conondale Ra. Qld. Note similarity of shape of sacral dia-
popbyses and the numerous examples of vertebral bilateral asymmetry.

Fig, 3.
muscles. A: Pxevdophrvne semimarmorata; B;

Lateral view of Depressyor miundibulue
PL bibroni; C: PL veeidentilis, A slip of the
muscle occurs upon the dorsal fuscia in each
species. D.M main squamosal/otic element of
depressor mandibulae; O.M.p.fusc.; depressor
mandibulae pars fascialis; D.M.p.tyvmp,: depres-
sor mundibulue pars lympanicus.

recognised three conditions in the origin of this
muscle: (a) arising from the posterior border
of the otic arm of the squamosal, (b) arising
from the squamosal and the dorsal fascia and,
(c) arising from the dorsal fascia alone.

In some respects this descriptive system re-
presents. a simplification because the muscle
commonly comprises three elements: the third
arising from the tympanum and termed the
“pars tympanicus”.

Lynch (1971) examined the condition of the
depressor mandibulae in P. bibroni and P.
corroboree and reported that the genus has
only one slip — the pars tympanicus.

Our studies do not support the observations
of Lynch or those of Heyer & Liem. As illus-
trated in Fig. 3, each of the species examined
by us has Jarge squamosal and dorsal fascial
elements. The relative size of the dorsal fascia
element in P. occidentalis is mtermediate be-
tween that exhibited by P. bibroni and that in
the type species P. semimarmorata. In P,
guentheri the M. depressor mandibulae re-
sembles the condition in P. bibroné except that
the slip to the dorsal fascia is slightly smaller.

20 M. J. TYLER & MARGARET DAVIES

Discussion

Our observations indicate that the form of
the M, depressor mandibulae and sacral dia-
pophyses of P. occidentalis cannot be dis-
tinguished from the condition exhibited by the
type species of Pseudophryne (P. semimar-
morata). Kankanophryne was erected on the
premise that the species differed substantially

from Pseudophryne, but we are unable to sup-
port its maintenance, and we propose that
Kankanophryne be referred to the synonymy
of Pseudophryne.

Acknowledgments
This study was supported by a grant to
M.J.T. by the Australian Research Grants
Committee,

References

Bock. W. J. & SHEAR, C. R. (1953) A staining
method for gross dissection of vertebrate
muscles. Anat. Anz. 130, 222-7.

Coscer, H. G. (1978) “Reptiles and amphibians
of Australia”. 2nd Edtn (Reed, Sydney).

Davis, D. D. & Gore, U. R. (1947) Clearing and
staining skeletons of small vertebrates.
Fieldiana: Technique (4), 3-16.

Dincerkus,. G. & UHLER, L. D. (1977) Enzyme
clearing of alcian blue stained whole small
vertebrates for demonstration of cartilage.
Stain Technol, 52(4), 229-32.

GriFFITHS, J. (1954) On the “otic element” in

Amphibia, Salientia. Proc. zool. Soc. Lond.
124, 35-50.
(1959) The phylogeny of Sminthillus

limbatus and the status of the Brachycephalidae
(Amphibia, Salientia). /bid. 132, 457-87.

Heyer, W. R. & Liem, D. S. (1976) Analysis of
the intergeneric relationships of the Australian
frog family Myobatrachidae. Smithsonian
Contrib. Zool. (233), 1-29.

Lyncu, J. D. (1971) Evolutionary relationships,
osteology, and zoogeography of leptodactyloid
frogs. Misc. Publ. Univ. Kansas Mus. nat. Hist.
(53), 1-238.

Parker, H. W. (1940) The Australasian frogs of
the family Leptodactylidae. Novit. Zool. 42(1),
1-106.

Trues, L. (1973) Bones, frogs and evolution. Jn
J. L. Vial (Ed.) “Evolutionary Biology of
Anurans: Contemporary Research on Major
Problems”. (University of Missouri Press,
Columbia. )

(1977) Osteology and anuran systematics:
intrapopulational variation in Hyla lanciformis.
Syst. Zool. 26(2), 165-84.

TyLer, M. J. (1972) Discovery in the Everard
Ranges of a species of leptodactylid frog new
to the fauna of South Australia. Trans. R. Soc.
S. Aust. 95, 215-7.

— (1978) “Amphibians of South Australia”.
84 pp. (Govt Printer, Adelaide).

—— (1979) Herpetofaunal relationships of South
America with Australia. In W. E. Duellman
(Ed.) “The South American Herpetofauna: Its
origin, evolution and dispersal’, pp. 73-106.
Monogr. Mus. Nat. Hist. Univ. Kansas (7).

NEW RECORDS OF KOONUNGA CURSOR SAYCE, 1908 (SYNCARIDA,
ANASPIDACEA)

BY P. DE DECKKER

Summary

Koonunga cursor is recorded from two new localities, one in southwest Victoria and the other in
northwest Tasmania. Specimens are illustrated with scanning electron microphotographs and
drawings of the appendages.

NEW RECORDS OF KOONUNGA CURSOR SAYCE, 1908 (SYNCARIDA,
ANASPIDACEA)

by P. De DEcKKER*

Summary

Tie Deeckker, FP.

(1980) New records of Keonutga cirsor Sayce,

1908 (Syncarida,

Anuspidacea). Travis. Re Soe, 8, Aust, W4(2), 21-25, 29 February, Los.
Koonunga crrsar is recorded from two new localities, one in southwest Victoria and the
other in northwest Tasmania. Specimens aro illustrated with scanning electron microphoto-

graphs und drawings of the appendages,

Introduction

Most parts of South Australia are too arid
for anaspidaccan syncarid crustuceans to live—
permanent, natural freshwater bodies are rare
exeept in the southeast near Mi Gambier
where rainfall is highest, However, in Victoria
very close lo the S.A, border near Mt Gambier
the living syncarid Koonunga curser has
recently been found and is described here.

Diseussion

Sayee (1908) desvribed K. cursor from
“freshwater reedy pvols beside » ny funnel
joining the Mullum Mullum Creek at Ringwood
near Melbourne’. Drummond (1959) stated
that J, Searle in 1930 failed to find K, curser
in the type locality beeatise it had become a
storm-Wwater drain. However, in an introductory
account of crustaceans from Victoria, Morrison
(1955) mentioned that “Koonunga . ~~ had
been extensively collected by various members
of the Zoology Department of Melbourne
University . . .°. She also stated that “it now
appears that the Koonungidae are widely dis-
tributed, with local abundance, in an area
south of the Great Dividing Range extending
(rom Portland to Wilson's Promontory”. Un-
fortunately no localities were mentioned by
Morriso), Drummond's (1959) short note on
the Australian syneurids said that “Koonunga
lis}... now known to occur sporadically right
ucross the southern part of Victoria’. Never-
theless, he made no reference to particular
localities and none have since been reported,
Even Schminke’s (1978) paper, which tp-
cluded an Ulustration of the telson of a juvenile
specimen of K, eurser given him by FL H.
Drunymond, gave no locality data.

* Deparment of Zoology. Universily of Adelaide,
G.P.O. Box 498, Adelaide, S. Aust. 5001.

Recently K, curser has been collected from
the following localities, (1) Victoria, close to
the S.A. border, under the foothridge over the
rivulet at Bullocky Wells Picnic Area, 1.5 km
E of Greenwald on the main road between
Mt Gambier and Heywood (35°58'42"S,
141°23'09"B) 24, v, 1979; (2) northwestern
Tasmania, from Mowbray Swamp near Mella,
5 km W. of Smithton. This Tasmanian record
is the first to be published, although P. S. Lake
(in Williams in press) has informally reported
its occurrence there, No locality data were
given. At the Victorian locality, 15 females
and seven males were collected from floating
vegetution, The females were larger and
darker in colour than the males, Adult males
were easily recognized by the peculiar globular
organ attached to the antennule (Fig. 4, 9),
The largest female collected was 9.4 mm tong
(anterior tip of head to base of telson) and
ihe largest male, 5 mm long. In Tasmania,
several specimens were collected, but only one,
a female 7,8 mm long, could be examined after
Preservation,

An undissected female and male are
illustrated in Figs |-2, and particular featores
of their anatomy are shown in more detail in
Figs 3-8. To broaden Sayce’s (1908) deserip-
tion, the left appendages of one male specimen
from locality (1) were dissected (Figs 9-34).
These specimens are deposited in the Aus-
trallan Museum, Sydney, with two tindissected
specimens of each sex accompanied by the
Tasmanian specimen.

Acknowledgments

! thank Prof, W. D, Williams, Department
of Zoology, University of Adelaide, for en-
couragement. Dr K. F. Walker brought my
attenlon to the article by Morrison, Dr K.
Bartusek, P, Kempster and J, Wright (Uni-
versity of Adelaide) are thanked for help with
the SEM photography,

22 P. DE DECKKER

Figs 1-2. Koonunga cursor from locality (1). 1, undissected 9; 2, undissected ¢. Both at same mag-
nification. Scale: 500. Specimens frozen dried and coated with gold palladium before SEM photo-

graphy.

Figs 3-8. Koonunga cursor from locality (1). 3, detail of fig. 1 to show cephalon with transverse
sulcus; 4, detail of fig. 2 to show globular organ on ¢ antennule; 5, detail of fig. 2 to show ¢
copulatory appendages in ventral position; 6, detail of fig. 2 to show anterior appendages and globular
organs on antennules; 7, 9 telson viewed posterolaterally; 8, ¢ telson viewed ventrally.

Scale: 500 for figs 3, 6, 7, 8; 50 for fig. 4; 200” for fig. 5.

io
9
A
4
=)
1S)
x
je)
z
=)
z
S
°
wat
=|
~
<
Oo
Zz

SY

E

ON TH

P. DE DECKKER

ON THE SYNCARID KOONUNGA CURSOR 25

References

DRUMMOND, F. H. (1959) The syncarid crustacea, Sayce, O. A. (1908) On Koonunga cursor, a
a living link with remote geological ages. Aust. remarkable new type of malacostracous
Mus. Mag., 13, 63-4. crustacean. Trans. Linn. Soc. London, 11, 1-16.
ie Scuminke, H. K. (1978) Die phylogenetische

Morrison, P. C. (1955) Animal life. In ‘Intro- Stellung der Stygocarididae (Crustacea, Syn-
ducing Victoria’ (Ed. G. W. Leeper) pp. 69-81 carida)—unter besonderer Beriicksichtigung
(ANZAAS: Melbourne). morphologischer Ahnlichkeiten mit Larvenfor-

men der Eucarida. Z. zool. Syst. Evolut.-forsch.,
NicHoLts, G. B, (1931) Micraspides calmani, a 16, 225-39. ,
new syncaridan from the west coast of ‘Tas- Me tae a Bee? sane eee a
mania. J. Linn. Soc. London (Zool.) 37, ‘Biogeographical Ecology in Australia’. Junk,
473-88. The Hague,

Figs 9-34. Koonunga cursor & left appendages except for figs 24, 29. 9, antennule; 10, antenna; 11,
maxilliped; 12, first peraeopod; 13, endopodite p'ate of peraeopod; 14, second peraecopod; 15, man-
dibular coxale; 16, first maxilla; 17, third peraeopod; 18, second maxilla; 19, third peraeopod (claws
missing); 20, fourth peraeopod; 21, first pleopod with endopodite; 22, detail of tip of endopodite;
23, detail of tip of endopodite, different orientation; 24, right second pleopod with endopodite; 25,
detail of fig. 27—tip of endopodite of left second pleopod; 26, detail of fig. 27—-coupling spines; 27,
left second pleopod; 28, median appendage attached to sternum; 29, detail of fig. 24—tip of endo-
podite of right second pleopod; 30, detail of fig. 29—coupling spines; 31, fifth peraeopod; 32,
penultimate pleopod; 33, last pleopod; 34, third pleopod.

Note: not all peraeopod endopodites are illustrated. Top scale: 5004 for figs 9-21, 31-34; middle
scale: 3004 for figs 25, 28, 29; bottom scale: 100 for figs 22, 23, 26, 30.

NEW UNIONICOLIDAE (ACARI, HYDRACHNELLAE) FROM
AUSTRALIA

BY KURT O. VIETS

Summary

Seven new species of the water mite genus Unioncola, subgenus Pentatax (Acari, Hydrachnellae),
are described from Australian freshwater mussels: Unioncola (Pentatax) walkeri n.sp., U. (P.)
procursa n.sp., U. (P.) assimilis n.sp., U. (P.) conjunctella n.sp., U. (P.) ligulifera n.sp., U. (P.)
clipeata n.sp., and U. (P.) scutata n. sp. A further new species is placed in Unionicolopsis
opimipalpis n.g., n.sp.

NEW UNIONICOLIDAE (ACARI, HYDRACHNELLAE) FROM AUSTRALIA

by Kuer O, Viets*

Summary

Viers, KO. (1980) New Unionicolidae (Acari, Hydrachnellue) from Australia. Trans. R.
Soe. §. Aust. 104(2), 27-40, 29 February, L980,

Seven new species of the water mite genus M/nieantcala, subgenos Pentama (Acari,
Hydrachnellne), are described from Australian freshwater mussels: OUniericola (Pentatax)
walkeri nap. U. (P) procursa op. UO. UPL) assimiliy ap, U, CP.) conjunetella nsp,, U.
(P.) ligulifera tsp, Us CP.) clipeata fusp. and U. (PL) seurdta nsp, A further tew species
is placed in Unionirolapsis oplmipalpis n.g., n.sp.

Introduction

The Hydrachnellae are prostigmate Acari,
Within the family Unionicolidae Oudemans,
1909 there are five subfamilies: Encentrito-
phorinae Viets, 1935 (20 spp, from Africa,
Australia, southern Asia), Najadicolinae
Viets, L935 (one sp., North America), Piona-
tacinace Viets, 1916 (many spp., world-wide),
Pollicipalpinae Viets, 1914 (two spp., Africa)
and) = Utonicolinae Oudemans, 1909, ‘The
Unionicolinae include four genera: Umionicola
Hakleman, 1842 (many spp. world-wide),
Vietsutax Uchida & Imamura, 1938 (one sp,
Japan), Hererati® Lamdblad, 1941 (one sp.
Africa) and Atacella Lundblad, 1937 (six spp.,
South America).

The taxonomy of Unionivela and its asso-
ciated subgenera still is somewhat unsatis-
Tavtory, but is based on the numbers of
genital ucetabulae, the appearinee of the
genital ficld. and the sexual differentiation of
the legs of males, The diagnoses of systematic
groups above subgenert) were last summarized
hy Cook (1974), Species of the subgenus
Jionicold 3.8. possess six pairs of genital
acetabulag: (wo species are known fron Aus-
Iralia, Species of the subgenus Pentatas Thor,
1927 have five pairs of acetabulae; three Aus-
tralian species ure known.

Some species of Unionicola are free-living,
others parasitic for part or all of their lite
cycle (ea, im molliises, Viets & Plate 1954;
Mitehell 1955; Hevers. 1978, 1978h). The life
histories of most species, however, are un-
known, In Australia, One L/iinnicola species
only has been reported in association with a
freshwater mussel, namely U. (U/.) clrrosa
Koenike 1914, in the mussel Unio nepeanensiv
(syn, (yridella enstraliy (Lamarek)),
Although the precise locality in Australia is

® Friedensir, 4B, D-2940 Wilhelmshaven, West

Germany:

unknown. the host mussel is likely to have
come from a coastal stream in the southeast
of the continent (cf, MeMichacl & Hiscock
1958),

Recently f obtained collections
mites from various Australian freshwater
mussels (Hyriidae) through Dr Ko Bb, Walker
of the University of Adelaide; [aim grateful to
him and the other collectors. Dr Walker also
adapted this manuscript from the original
German. rom the material seven new species
of Unlonicola, subgenus Pentatax are described,
and Unionicolopsis na, is erected for an
udditional new species.

of Watcr

General Remarks

The types und paratypes deseribed herein
are catalogued i the Viets Collection (Wil-
helmshaven), and ultimately will go to the
Senckenberg-Museum, Prankfurt.

The legs of water mites are 6-scamented: |
trochauler, 2 basifernmur, 3 lelolemur, 4 gent,
§ tibia, 6 larsus. The legs and their segments
usually are symbolized as, for example, first
les tarsus; 1.1.6 and fourth leg tibiar PV,L,5
(see Cook 1974: Fig. 5). The pulps are 5-
segmented: | trochanter, 2 femur, 3 genu, 4
tibia, 5 tarsus. These are denoted P FV (Cook
1974; Pig. 6),

All measurements herein are jim.

Unionicola (Pentatax) walkert rsp.
FIGS 1-8

Holorype 2 prep. 6638; With soft miegument,
In dorsal integument le two pairs of delicate,
oblong plates (muscle attachment sites), an-
terior pair nearer one ahother than the pos-
terior puir; length 35-50,

Coxal area (Fig, 2) 415 long, Third coxae
wonsiderably sherler in mid-line than fourth
coxac; suture between coxal pairs does not

28 KURT QO. VIETS

50m

Figs 1-8, Unionicola (Pentatax) walkeri n.sp. 1. 2 holotype prep. 6638, IV.L.5—6, right; 2. ? holotype,
ventral side: 3. 2 holotype, 1.L.5-6, left; 4. ¢ prep. 6675, I.L.5-6, right, dorsal aspect; 5. @ holo-
type 6638, right palp; 6. ¢ allotype prep. 6642, left palp; 7. d allotype prep. 6642, genital field; 8.
® holotype prep. 6638, genital field.

NEW UNIONICOLIDAE FROM AUSTRALIA 2

reach medial margin. Posterior margins of
fourth coxae rounded, and cach wilh pos-
teriorly-directed process. Medial margin of
third ant fourth coxae 198 long,

Genital field (Fig, &) entirely at poslertor
end of body. It consists of two pairs of
medially-constricted plates; witernor pale each
Wilh two genital acetubulae, and posterior pair
each with three acetabulac, On strongly-
lapered) medial edge of each anterial plate
short, stout sela (stylet), and neac it
smaller, slender seta, On medial edge of each
posterwr plate large. thiek, curved sela, and
alongside two (in some cases three) slender
setae, Gonopore (about 182 long) surrounded
hy pre and post-genital sclerites.

Capitulum (Fig, 2) 162 long, ¢, 104 wide.
Chelicerae similar ti those of other Uaionl
volu species. Dorsal palp segment lengths (left
PI-V): 10, 105.53. 109, 54, P Il on each
side with bwo selne, and PLE one seta, P TV
has no ventral process, apart fram a minute,
often indistinet, peglike seta at its distal end.
PV ends in two strange claws,

Legs LIV with number of ventrally-attached
setae, On seament 5 of all legs, and On segment
4 of legs If and VV, are 3 long. slender swine
iing-setae (el Cook 1974: Big, 5), located
distally, Last segment of leg 1 (Pigs 3-4),
seen fram side, strongly expanded distally (.c.
dorsd-vertrally expanded) to form laree claw
fossa, On dorsal side, at distal end of segment,
slender-based, spnon-shaped, chitinous struc:
jure, In Ply. 40 tuesal claws omitted, Club-
shaped structure situated on distal ead of TV.L.6
(Fig. 1). Small, spoon-shaped seta attached
near tarsal claws on UL.6 and 1.1.6. Tarsal
claws simple in form, having no carpus, hur
with slender, terminal spine. Dorsal segment
lengths af legs Pand Ws LL.3-6: 74. 115 -
97. VIR IV.LGe6: 94. 155, 180, 165,

Alleotype & prep. 6642: Coxal area Clength
410) and dorsal side correspond closely with
& Genital field located terminally as an en-
closed plate 127 long wind |45 wide. Por ar-
rangement of genital acetabulae and hair
pores. see Fig, 7,

Cupitulum 149 long and e. 100 wide. Palps
(Fig. 6) similar to 9, lefi P L-V, measured
dorsally, as follows: 11, 97.. 54. 105 , 53.
Leys similar to 9%) dorsal segmentdensths:
Lbh3a-6; 70, 11). 99. 114: INLD: 92,
146, 17, 152,

Vorivrion- coxal area, leneth 415-453 P27),
480143 4409) gonopore, length incl, selerites

169-188, , genital ficld ¢, length =, T14—
127; genital field ¢, width ,,., 124-165; eapitu-
jum, length 162-177, 143-159; capitulum,
wilth 103-114, 91-101: chelieera, length
(152), (135-152); palp, dorsal segment length
PL K-13, 7-11. P UE 99-113, $2-105: Poot
46-62, 47-59; BP IV 109-120, 100-LI4: P VW
53-58, 44-57; jews, segment length 11.3 74—
84, G9-7h; LI4 114-124, 103-119; LT.5 95-
107, 89-105; [LL.6 110-122, 104-1215 1V.1.3
94-104, 83-99; IV,L.4 155-170, 135-158:
IV.L.5 177-198, 157-182; IV.L.6 165-175,
I46—159; egg, long axis 135-143,

Jats aml lovulities GUNS, Austad: Alathiyria jack
sont Iredale, Lock 3, River Murray, KF. Walker,
19.x.1977 (1 o&\ TL 9, holotypeys Lock 3, R.
Murray, K. F, Watker, 5.411977 (1 @y allotype)>
Velesunio wakicuus (PHU pO, Lock 3, R. Murray,
K. EF. Walker, 19.47.1977 (8 & 7 9)y Point Sturt,
Lake Alexandrina (R. Murray) K.P. Walker,
30.x,1977 (4 9); L. Alexandrina nr Milang,
M, C. Geddes, 20.41,1977 (3 ¢, 2 9): Point Sturt,
L. Alexundrina K, &. Walker, 3.811977 (30, 2 9).

Unionicofas (Pentatax) procursa n.sp,
FIGS 9-14

Nololype & prep. 66712 Dorsal integument and
vosal area (Fig. 9) very similar to Lf. (P,)
Walkeri nap, although posterior process of
fourth coxa somewhat shorter, Coxul area 446
long.

Genital field (Fig, 10, lat, view) at end of
body, In dorsal aspect (Fig. 11) jbdominal
integument at anterior margin of field indented
(although this is not clear fram Fig, 10),
Anterior plites of genital field, eaeh with twa
atetabulue, directed posteriorly, Bach atterior
plate with two stout setae ul hindimost cx-
Iremity, Posterior plates broader, and bear one
seta at the ends of their respective posterior
processes.

Capitulum 150 tong and 14 wide an-
leriorly. Chelicerae like those of congeners,
Long, slender P V (Fig. 12). and, as with re-
lated species, two distinct terminal claws. P
IT) with two setae on outer horder, and three
inner setac. Dorsal sczment lengths (lefr P
VW): 15. 110. 60. 116. 73,

116 similar to that of UL CP.) walkeri nsp,
(Fig. 13; tarsal claws omitted for clarity). It
is, however, smaller and the distal border of
the expanded part of the large chiw fossa is
vonvesly rounded, nat strongly indented as in
other speetes, Large, spoot-shuped appendage

30 KURT QO. VIETS

is

Figs 9-14. Unianicola (Pentatux) procursa n, sp. 9. 9 holotype, prep. 6671, ventral side; 10. 9 para-
type, prep. 6673, genital field, lateral view; 11. 2 holotype, genital field; 12. 2 holotype, left palp;
13. 2 holotype, 1.L.5-6, left; 14. ¢ allotype, prep. 6676, genital field.

on LL.6 like that of U. (P.) walkeri nsp.
Stout seta on dorsal side of IT and III.L.6,
proximal to cach claw; tends to a spoon-shape
on leg Ll, but simple on leg III. No club-
shaped chitinous structure on end of IV.L.6,
as in related species. Otherwise leg setation
shows no significant differences, Dorsal seg-
ment lengths: 1.L.3-6: 95 . 163. 117 . 129;
IV.L.3-6: 104 . 168 . 183 . 167.

Allotype 3 prep. 6676; d and 2 very similar,
Coxal area 365 long. Genital ficld (Fig. 14)
differs from other species in form and
especially number of hair-pores, A small an-
terior projection, 114 and 126 wide dorsally.

Capitulum dimensions 122 : 85. Chelicerae
126 tn length. Palps as in 2; dorsal segment

lengths (left P I-V): 14. 94.50.100. 64,
Legs as in §. Dorsal segment lengths: 1.L.3-6:
78, 124 , 92.. 109; IV.L.3-6: 82.130. 139.
144.

Variation: coxal area, length 365-393 dd (4),
405-446 29 (5); capitulum, length 122-133,
135-150; capitulum, width 83-90, 94-104;
palp, dorsal segment length P [ 12-14, 13-16;
P If 90-100, 98-110; P IIL 44-51, 51-64; P
TV 101-105, 112-119; P V 59-67, 64-73;
genital field (9), length 114-120, |; genital
field (3), width 126-137, . ; legs, segment
length LL.3 75-80, 88-97; LL.4 121-128,
144-164; LL.5 92-99, 105-117; LL.6 105—
109, 115-129; IV.L.3 79-85, 92-108; 1V.L.4
129-139, 149-169; IV.L.5 139-151, 164~-183;

NEW UNIONICOLIDAE FROM AUSTRALIA 31

Figs 15-19. Unionicola (Pentatax) assimilis n.sp. 15. 2 holotype, prep. 6625, ventral side; 16. 9 holo-
type. left palp: 17, 9 holotype, I.L.5—6, left; 18. ¢@ paratype, prep. 6635, dorsum; 19. ¢ allotype,

prep. 6633, genital field.

IV.L.6 137-146, 161-168; egg, long axis,
160-163.

Host and localities (all S, Aust.): Felesraio anm-
biguas, R. Murray nr Mannum, K. F. Walker.
19.x1.1977 (1 o&): Point Sturt, L. Alexandrina (R.
Murray), K. F. Walker, 30.%.1977, (5 @): L.
Alexandrina nr Milung, M, C. Geddes, 20_xi.1977
(74. 1 9. inel. holotype and allotype); Point Sturt,
L. Alexandrina (R. Murray), K. F. Walker,
3.4i1,1977 (5 2).

Unionicola (Pentatax) assimilis n.sp.
FIGS 15-19
Holatype © prep. 6625: Ridged posterior plates
of dorsal integument (Fig, 18, 4) about 130
Jong. Coxal area (Fig, 15) length 406. As in
the other species, suture between the third and
fourth coxae does not reach medial margin,
and is directed transversely. Genital field, capi-

tulum and chelicerae similar to U. (P.) pro-
cursa nsp. P HW (Fig. 16) bears two setae on
inner and outer sides. P V ends in two claws;
length in same proportion to P IV as in other
species. Dorsal lengths of P If and P IV nearly
identical, whereas in U. (P,) procursa o.sp.
P TV is slightly longer than P Tl. Dorsal seg-
ment lengths (left P I-V): 11. 105 . 50.
103. 61.

LL.6 (Fig. 17) expanded distally, and
52.6-62.7% of segment length; it is also
shorter in proportion to I.L.5 (90,5-101.2% )
than is the case for U. (P.) precursa n.sp.
(106.5-118.5% ), 1.1.6 bears a spoon-shaped
distal projection like the other species. On
11.L.6 and HI.L.6, however, only one stout
seta at distal end of dorsal edge, and this is
not spoon-shaped. Dorsal segment lengths:
1.L.3-6: 84 . 136 . 95. 90; IV.L.3-6: 90.
144 . 172 . 149,

32 KURT ©, VIETS

Allorype 3 prep. 6633: Dorsal integument,
coxal area, palps and legs similar to ?, Pos-
terior dorsal plates about 115 long, with strong
chitinous ridge (Fig. 18), Coxul area length
355, Genital field (Fig, (9) about 80 lone and
97 Wile, weakly indented al anterior border,
Capuulum dimensions 132 = 749. Dorsal palp
segment lengths (left P LV): — , — . 42~
85, 54. Dorsal leg segment lengths: I.L3-6:
74. 110, BS. BIS 1V.1. 3-6: 75. 123, 150
134,
Fariation: coxal area, length 344-480 cd (6),
dill 442 29 (6); posterior dorsal plate, length
105-137, 125-140; eapittilum, length 124-138,
143-157: capitulum, width 76-85, 94-100;
cheliverae, length (102), (125-132); palp.
dorsal segment length PT 10-12, 10-13; PH
85—97, 102-112: P TIL 42-48, 50-62; P 1V
82-89, 99-110; P V 50-54, 56-67; genital
field (), length (77-85), 3 genital fleld (4),
width 89-108, + leg, segment length 1.0.3
65-83, 84-98; E.L.4 104-119, 136-145; LS
77-89, 95-105; LL.6 75-85, 89-97; 1V.1.3
7I-84, 89-97; TV.L.4 115-132, 144-155;
IV.L.5 142-160, 172-186; IV.L,A 130-139,
147-165; egg, long axis =, (133-138);
Host and locality: Wextralunia cartert: Tredale.
Murray River S of Dwellinuup, W. Aust, N. M.
Morrissy, [5.xi1.1977 (8a, 13 9, 11 nymphs, ingl
holorype und allotype),
Remarks: Allied to U. (P.) procurya wasp.
Significant differences between the two species
are summarized in Table |.

Unionicola (Pentatax) conjunctella n.sp.
FIGS 20-25
Holotype 2 prep. 6669 Dorsal side (Pig, 23,
a) like that of U (P.) assinills asp. Ridged
plates (c. 135 long) very thin, outline not
always eVident. in contrast to ridge itself,

Coxal area (Fig, 20) 360 long. Suture between
third and fourth coxae rims obliquely, ap-
proximately parallel to anterior margin of third
coxae, and fused with medial edge of posterior
plates. Genital flekt (see Fig. 20) similar to
other species.

Capitulum {42 long and 87 wide, abd, like
the chelicerae, with no distinctive characteris-
tics. Palp (Fig. 21) segment P IL bears two
sefac on cach of its inner and outer sides,
PBorsal lengths of segments (left P I-V): 12.
89, 42.91 .—.

L.L.6 (Fig. 22) similar to that of U, (P.)
assimilix n.sp, Terminal segments of legs I and
Ht each with expanded spoon-shaped seta
dorsally, at distal end, in front of tarsal claws,
Dorsal segment lengths: 1.L.3-6: 64.93. 72.
76; IV.L.3-6: 77.115 , 140, 125.
Allotype 3 prep, 6663: Dorsal side (Pig. 23)
similar to 9, Length of posterior plates 158,
Coxal area 386 long, Unlike 9, first coxae (Fig,
24) extend posteriorly beyond ends of Outer
apodemes. Genital field 94 long and L110 wide,
(Fig. 25) ke U. (P.) asvimilis n.sp.

Capitulum 140 long and 8&7 wide. Palps
similar to 8; dorsal segment Jengths (left P
1-V): 10.93.45. 93. 47, Legs ulso similar
to ?. Dorsal segment lengths: I,L, 3-6; 66 ,
99.76. 80; 1V.L.3—6; 77, 119. 147. 132.

Variation: coxul area, length 340-406 det (7),
320-380 99 (4); posterior dorsal plates, length
130-158, 130-135; capitulum, length 125-145,
124-142; capitulum, width 76-90, 80-87;
chelicera, length (110-125), (115); palp,
dorsal segment length P | 10-13, (12); PW
79-97, 83-89; P [1 37-47, 40-45; P IV &4—
98, 84-93; P V 43-48, 42-47: genital field
(3), length (81-94), | =: genital field (<*),
width 107-129, |; legs, segments length 1.1.3
57-71, 58-64; LL4 85-105, 85-04; LL

TAbies |, Comparison af U. (P.) assimilis and U, (P.) procursa

U. (P.) assimilis nes.

U. (P.) procursa n.sp,

dorsal inteminient
ment sites;
subire between Goxae J and 4 Lransverse

venirul field (4)

P 1 mner side 2 setae
Li.6 => broad
inex, wlth os & segment ;
le twits od: $4.3-83.7%

R9> 52.6-60,0%6

1 pair small ant. musele altach-
Lo piir post. plates
with strony ridge

Without medisl process; post, 4
with many hait-pores

| pair small ant. muscle attuch-
ment sites; L pair post. plates
without strong ridge
oblique, << parallel to
border of 3rd coxa

with medial process; post. 4 with
few hair-pores

3 setae

> slender

ant,

ad: 40,2-41.3%
O- 36,8-40.0%

NEW UNIONICOLIDAE FROM AUSTRALIA 33

\ :

23 st”

Figs 20-25. Unienicola (Pentatax) conjunctella n.sp. 20. 2 holotype, prep. 6669, ventral side; 21, 2
holotype, left palp; 22. 9 holotype, I.L.5-6; 23, ¢ allotype, prep. 6663, dorsum; 24. ¢ paratype,
prep. 6662, coxal area; 25. ¢ allotype, genital field.

66-83, 66-76: LL.6 71-88, 72-77: IV.L.3
69-87, 70-77; IV.L.4 105-129, 108-121;
IV.L.5 131-155, 136-148; IV.L.6 120-143,
117-131; [.L.6, maximum width 43-50, 44—
47: egg, long axis — , (165).

Hasts and localities: Alathyria jacksoni, Lock 3,
R. Murray, S. Aust, K. F. Walker, 19.xi.1977
(1 @): FKelesunio ambiguus, R. Murray nr Man-
num, S. Aust, K. F. Walker, 19.xi.1977 (2 2,
1 2); Lock 3, R. Murray S. Aust., K. F. Walker,
19.1x.1977 (3d, incl, allotype, 1 %); Rocklands
Reservoir, Vict.. P. J. Suter, 20.x1.1977 (1 et);
Point Sturt, L. Alexandrina (R. Murray), S. Aust.,
K. F. Walker, 30.x.1977 (1 ¢ holotype).

Remarks; Distinguished from U. (P.) precursa
n.sp. and U. (P.) assimilis n.sp. in that suture
between third and fourth coxae extends to
medial margin of posterior plates.

P V is considerably shorter than in other
species, as seen in comparisons of P V length
as % PIV length (fo + 2): U. (P.) conjune-
tella n.sp. 46.2—-53.6%; U. (P.) procursa n.sp.
57.1-65.7%; U. (P.) assimilis nsp. 56,0—
64.3%.

Unionicola (Pentatax) ligulifera n.sp
FIGS 26-32

Holotype 2 prep. 6664: As with U. (P.) con-
junctella u.sp.. dorsally two separate, thin
plates with prominent chitinous ridges. Coxal
area (Fig. 26) 373 long. First coxae do not
extend beyond posterior ends of apodemes,
Suture between third and fourth coxae short
and does not reach medial margin, Genital
ficld (Fig. 27) like that of other species except
U, (P.) walkeri n.sp.

34 KURT 0. VIETS

Figs 26-32. Unionicola (Pentatax) ligulifera usp. 26. 2 holotype, prep. 6664, coxal area, left: 27.
4 salons genital field; 28. 9 holotype, left palp; 29. 2 holotype, I.L.5—6, left; 30. 2 holotype,
1.L.5—6, left; 31. ¢ allotype, prep. 6666, coxal area; 32. ¢ allotype, genital field.

NEW UNIONICOLIDAE FROM AUSTRALIA 35

Capitulum dimensions 120 : 81. PTT (Fig.
28) hears two setae on inner and outer sides.
PV long, 57-58%r of PIV length, Dorsal seg-
Ment length Cleft P HV)5 12, 85.44.91,
52.

L.L.6 (Fig, 29) shorter than 1.L.5 and typi-
cal of other species in the Pentatax group, i
and HLA (big, 30), like 1.1.6, auch with a
large spoon-shaped chitinous structure distally.
Species in relaled groups have, on these seg-
inents, Minute setac only slightly expanded
distally, Dorsal segment lengths: 1.1.3-65 72.
TTR, 100. 89: 1V.L.3-6:-85 . 134. 152. 1399.
Allatvpe 3d prep. 6666 (+ paratype a, in tote);
Dorsal integument similar to ?, Coxal afea
(Pig. 31) 332(326) long (paratype dimensions
in parentheses), First voxwe slender in mid-
posterior region and wide and elongate wh ex-
Iremily, Genital field (Fig. 32) 134 wide.
Foremost four adetabulae om cither side
arcanged i rows,

Capitulum 103(102) Jong and 70(69) wide,
Palps as in 2; dorsal segment lengths Clete P
I-W)s WNT2) . TITEL. 34037), 7777)
45(45). Legs similur to Y. Dorsal segment
lengths: 11,346: 59.97 , 87, 78; IV.L,3-6:
73 110, W322. 117,

Aloxsty and ftocaliiwa Alaiiyrta jawksarni, Lock 3,
Ro Murray, So Awash, KR. oR. Walker, 19.97.1977
(1 4th) Velesunia ambiewas, Lock 3, R, Murray, 8.
Aust, Ko B Walker, 1941977 (1 2, 2 of nel,
holotype and ulloty pe)

Remarks: Distinelive ip that UU, and ULL 6
tach bear, on their distal-dorsal mareins. a

large. spoon-shaped chitinous structure similar

to that borne on 1.1.6 Although in the & the
first coxae do not extend beyond the posteriar
upodemes, they are very long in the ef
markedly longer than in the ¢ of U. (P.) cen-
funetella nap, The prajcetion extends to the
suture between the third and fourth coxae, The
suture does not reach the medial margin of
the postermm plates.

Unionicola (Pentatax) clipeata nisp.
FIGS 33-38
Holorpe So prep. 6647 (paratype @ prep.
6649); (Measurements of paratype in paren-
theses). Weakly chitinized, undivided dorsal
shield has dimensions: 202(18B8) 180(171).
Cosal area (Pig. 33) 302(271) long. First
coxae do not extend beyond posterior end of
apodemes, Suture between third and fourth
coxac curved over entire Jenygth, und meets
medial mmirgin of posterior plates. Fourth

coxae have almost right-angled medial-
posterior margin. Genital field (see Fig, 33)
similar to preceding species.

Capitulum dimensions 112(95), 6760),
ehelicerye &7 long, P IL (Fig. 34) has two
sclac on inher and outer sides. Relative length
of P V between that of LL (P.) ceonjunerella
nsp, und &, (P.) proeirsa nsp, Dorsal seg-
ment lengths (left P I-V)> 1108), 60055) .
36(29) . 66(63) — 38(33).

11.6 (Fig. 35) relatively short and wide
(61,3-67.2% of segment lengih), and with
convex distal margin. Distal-dorsal setae on
legs [1 and If only slightly expanded, Dorsal
leg segment lengths: Ef3-6; 51(48)
7O0(68) ~53(52) . 61(59) 5 1V.L.3-62 59055) -
89(86) , 1O7(106) . S4(96),

Allatype & prep. 6648 (+ pararype & prep.
6650): Dimensions of dorsal shield (Fig. 36)
208 : 178, Coxal area 288(280) long, similar
to 2. Genital field (Fig, 37) with many bair
pores, indented anteriorly and 127(132) wide.

Capitulum 99 long and 63 wide, Palp dorsal
segment lengths (left P I-V):—(8) . S8{359) -
32(30) . 62(61) . 36(33); Lews similar to ¥.
Dorsal segment lengths: L1.3-6: S051)
H9(70) . 53054) - 6261), 1IV_L.3-G: 56058) -
R2(86) , 105(103) . 95(96),

Hast and localicv: Welesinia ambieuys, R, Murray
hr Mannum, S, Aust, K. F, Walker, (9.xi,1977
(2 3,2 9) inch holotype and allotype).

Unionicola (Pentatax) scutata n.sp,
FIGS 39-45

Holetype ? prep, 6679; Dorsal shield (Fig. 39)
aboul 280 long and finely pored. On each side
small, weakly chitinized shield, with hair
pore, in front of anterio’ muscle attachment
sites, Coxal area (Pig. 40) 366 long. Suture
between third and fourth coxae extends to
medial margin of posterio’ plates, First coxa
does pot extend beyond end of posterior
apodemes. Genital field (see Fig, 40) similar
to other specics,

Capitulom 114 long and 74 wide. PU (Pig.
41) with 2 setae on inner and outer sides. Bor-
sal segment lengths (left P FeV): 13. 68 , 35
Sl, 40,

1.L.6 (Pig. 42) similar to other species: as
with U. (P.) elipeata n.sp., slightly longer than
penultimate segment. Distal-dorsal setae on ET.
and HhL.6 only slightly broadened (cf, spaon-
shaped), IV.L.6 (Fig, 43) has, in the middle
of its distul extremity, vlub-shaped chitinous
siticture comparable to 0. (P.) walkeri nsp.

36 KURT O. VIETS

+S
36 “ow >

Figs 33-38. Unionicola (Pentatax) clipeata n.sp. 33. 2 holotype, prep. 6647, ventral side; 34. 2 holo-
type, left palp; 35. 2 holotype, I.L.5—6, left; 36. ¢ allotype, prep. 6648, dorsum; 37. ¢ allotype,

ventral side, posterior; 38. ¢ allotype, left palp.

Dorsal leg segment lengths: I.L.3-6: 55 . 88.
65 . 64; IV.L.3-6: 66 . 110. 138 . 123.

Allotype & prep. 6680 (+ paratype ¢ prep.
6681): (Dimension of paratype given in
parentheses). Dorsal side similar to 2. Coxal
area (Fig. 44) 333(375) long. First coxae
extend beyond end of posterior apodemes (in
these characters the ¢ is similar to the d of
U. (P.) conjunctella n.sp.). Genital field (see
Fig. 44) 138(158) wide, and posterior margin
(Fig. 45) indented; large numbers of hair-
pores in posterior area.

Capitulum 109(118) long and 70(75) wide.
Palps are similar to 2; dorsal segment lengths
(eft P I-V): 8(9) . 60(70) . 30(33) .
72(77) . —(41). Dorsal leg segment lengths:
IL.L.3-6: 55(58) . 85(89) . 64(67) . 67(68);

IV.L.3-6: 62(67) .
116(121).

Host and locality: Velesunio angasi (Sowerby),
Ja-Ja Billabong, Magela Creek, Arnhem Land,
N.T., W. D. Williams, xi.1977 (2 o, 1 9, incl.
holotype and allotype).

Remarks: Like U. (P.) clipeata n.sp., this
species has an undivided dorsal shield in which
borders have become fused. Essential
differences between the two species are in
Table 2.

Genus Unionicolopsis n.g.
Diagnosis (based on 2 only): With soft integu-
ment. Coxae in four groups, the posterior
group especially widely separated; anterior
coxal group without posterior apodemes. Geni-
tal field with delicate, weakly chitinized plates,

102(110) . 128(136) .

NEW UNIONICOLIDAE FROM AUSTRALIA 37

Figs 39-45. Unionicola (Pentatax) scutata n.sp. 39. 2 holotype, prep. 6679, dorsum; 40. 2 holotype,
ventral side; 41. @ holotype, left palp; 42, 2 holotype, I.L.5—6, left; 43. 2 holotype. [V.L.5-6, right;
44. ¢ allotype, prep. 6680, ventral side; 45. ¢ paratype, prep. 6681, genital field.

38

body size
dorsum

genital field
IV.L.6

KURT O. VIETS

TABLE 2. Comparison of U. (P.) scutata and U. (P.) clipeata

U. (P.) scutata n.sp.

large (330-380)

dorsal shield strongly chitinized; with a
pair of small shields, each bearing hair-
pore, in front of ant. muscle attachment
sites

with many hair-pores
with distal club-shaped structure

U. (P.) clipeata n.sp.

small (270-300)

dorsal shield weakly chitinized; without
shields in front of ant. muscle attach-
ment sites

with few hair-pores
without distal club-shaped structure

Figs 46-53. Unionicolopsis opimipalpis n.sp. 46. 2 holotype, prep. 6683, ventral side; 47. 2 paratype,
prep. 6684, genital field; 48. 2 paratype, prep. 6682, genital field; 49. 2 holotype, capitulum, lateral

view; 50. 2

left palp; 53. 2 paratype, prep. 6684, leg IV, left.

paratype, pre. 6682, capitulum, dorsal view; 51. 2 holotype, chelicera; 52. 2 holotype,

NEW UNIONICOLIDAE FROM AUSTRALIA 39

anterior plates cach with two, posterior plates
cuch with three, genital acetubulae. Medial
margins of plates without special spines or
setae, Capitulom with broad upper processes,
Pulp stout, without process on P LV.

Type species. Unlonicelepsis opimipalply asp.

Unionicolopsis opimipatpis n-sp,
FIGS 46-53
Holotype ® prep. 6683 (+ paratype 22 preps-
6682, 6684): In soft dorsal integument is pair
of Jong and pair of short muscle atlachment
siles; these are not significantly chitinized, and
not formed us plates, Datu for all three speci-
mens are given (paratypes in parentheses).

Third and fourth coxae (Fig 46) relatively
widely separated in medial line; medial length
about L48(145, 148), Posterior apodemes of
first coxal group absent.

Genital plates (Figs 47-48) very weakly
chiunized, and indistinct in outline; widely
separated, on both sides of gonopore (c. 165
(165, 175) long). As in Pentatay, each an-
terior plate with two genital acetabulae, and
posterior plates with three acctabulae, Nuniber
of avetabulae may vary (Fig. 48),

Capitulum (Figs 49-50) stout, abour 164
(150, 163) lone and —(119, 120) wide, and
ends posteriorly in two wide-lobed, upper
processes. Chelicerae 165 long (Fig, 51),
Palps stuut (Fig, 52), P I with two setae on
inner and outer sides, No process on P IV, P
V ends in two large, dissimilar claws.

Legs very simple in form. On 1.0.3, in about
middle of ventral side, is conspicuous, very
long, strong seta, and beside it a short seta,
On T1.L.3 and 4 is distal seta and another
proxinmully. Sctae on segment 3 dissitnilar int
length, Whereas those of segment 4 are about
equal, and in general longer than those on
seoment 3, At distal extremity of WLL.S paw
of sete (V.L.1 hears three distinet setae (Fig
53); all other segments of leg IV with only
very small spine-like setac. Seements 4-6 on
leg TV elongate, and segment 6 slightly cliirved,
All tarsal claws simple. Dorsal leg segment
lengths: LL, 3-6: 59(59, 62) . 75€67, 75) .
72065, 72) . T5€7S, 79); IV-T.38-6> 109199,
105) _ 169(L58. 173) . 187(179, 199) — —
(154, 164).

Flost and lacalis Velesunio ambleans, Ry Murray
ur Albury. SSW, T, 2. Hillman, xi,1977 (3
incl, holotype).

Remarks’ Only 3 29 of this new species jire
available and, because the 2 remains unknown,

the systematic position of the material is un-
certain, Whether the new species, for which
the Unionicolapyty ng. wow is efected, should
be placed as a subgenus in Uniertieala, must
remain in doubt pending discovery of the ¢.
It appears unlikely ta me.

List of mussels and associated mites

Alathyria jackson] Wedale: U, (P,) walkeri,
conjunctella, ligulifera.

Felesanio ambiguus (Phillipi)s Of CP)
walkeri, conlunctella, lizulifera, precursa, ell-
peala; Unianicolopsis apimipalpis.

Velestinio anger (Sowerby)! U, UP) setitata.

Westralunia carter’ Wedale: U. CP.) avsimiilis,

Key to the species of the Pcoutaisx group
described herein
|. Dorsum with undivided shield, with pair of
distinct chitinous ridges
Dorsum without complete shield 3
Shield strongly chitinized; IV.L.6 with distal
club-shaped chilinous structure; first coxae of
§ falling short of, those of ¢ extending beyond
posterior apodemes seule
Shicld weakly chitinized: 1V.L.6 without distal
club-shaped chitinous structure; first cosas of
Cand @ not extending beyand poslerior apu-

2

13

demes clipeuta
3. Dorsum with two weakly chitinized plates,
euch having disunet chitinous ridge 4

Dorsom without chitinized ridges, hut with
muscle altuchment sites On weakly chitinzed
plates. 6
4. Sulure between third and fourth coxaé Teach-
ing medial murgio of posterior pair af plates;
distul-dorsal edges of IT, and ULL. with only
slightly cxpanded (spounshaped) seta; first
coxue of ® shorter than those of @, longer than
postetior apodemes confunerella
Sature between third and foutth coxae nat
teaching medial murgin of pesteraae pair of
plates a (Wie heetie

5. Th and W1,L.6 with setae Only on distal-dorsal
margin: first coxue of ¢ and % not eatending
beyond posterior apodemes nssiinilis
U. and W.6 with large spoon-shaped chiti-
nous simucture dorsally (as on LLG): first
coxae of 9 shorter (han, and hose of ¢@ extend-
ing bevond posterior apodemes teaching
suture between third and fourth coxae)

livutifera

6. P Vv long (57, 1-65. 19% of P TY Tength); inner
ale of PEL with 3 setae, IV L.6 without elub-
like structure ; procurse
PV short (45,7-41,9% of P TY length): inner
side Of P WE with 2 setae, LV.L.6 with elub-like
structure (? genital field distinct from that of
other species, with 4 similar plates and short,
ifick spines) Pcie nrenKcn . walkers

40 KURT O. VIETS

References

Coox, D. R. (1974) Water mite genera and
subgenera. Mem. Amer. Ent. Inst. 21, 7
860 p.

Hevers, J. (1978a) Zur
Gattung Unionicola (Hydrachnellae,
Zool. Jb. Syst. 105, 33-64.

(1978b) Morphologie und Systematik der
in Deutschland auftretenden Schwamm-und
Muschel-Milben-Arten der Gattung Unionicola
(Acarina: Hydrachnellae: Unionicolidae). Ent.
Generalis, Stuttgart 5, 57-84.

KOENIKE, F. (1914) Neue und _ neubenannte
Wassermilben. Abh. naturw. Ver., Bremen 22,
383-404.

Sexualbiologie der
Acari).

MITCHELL, R. D. (1955) Anatomy, life history
and evolution of the mites parasitizing fresh-
water mussels. Misc. Publ. Mus. Zool. Univ.
Michigan, Ann Arbor 89, 1-28.

McMicuwaEL, D. F. & Hiscock, I. D. (1958) A
monograph of the freshwater mussels (Mollusca:
Pelecypoda) of the Australian region. Aust. J.
Mar. Freshwat. Res. 9, 372-507.

Viets, K. & PLATE, H. P. (1954) Die okologischen
(parasitologischen ) Beziehungen zwischen
Wassermilben (Hydrachnellae, Acari) und
Sufwasser-Mollusken. Z. angew. Ent., Berlin
35, 459-494,

IP

VOL. 104, PARTS 3 & 4 30 MAY, 1980

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED

CONTENTS

Twidale, C.R. The Devil’s Marbles, central Australia - - - - - 41

Breed, W. G. Further observations on spermatozoal morphology and male
reproductive tract anatomy of Pseudomys and Notomys species
(Mammalia: Rodentia) - - - - - - - 51

Flint, R. B., Ambrose, G. J. & Campbell, K. 8. W. Fossiliferous Lower Devonian

boulders in Cretaceous sediments of the Great Australian Basin 57
Beveridge, I. Progamotaenia Nybelin (Cestoda: Anoplocephalidae): new
species, redescriptions and new host records - - - - 67

Mawson, P. M. Beveridgea new genus (Nematoda: Strongylida) from the Agile
Wallaby from northern Australia - - - - - 81

Ludbrook, N. H. Non-marine molluscs from dolomitic limestones in. the north
of South Australia - - + - - - = 2 83

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

THE DEVIL’S MARBLES, CENTRAL AUSTRALIA

BY C. R. TWIDALE

Summary

The Devil’s Marbles consist of castle koppies (catellated inselbergs) and large residual boulders,
both developed in granite. They are exposed in the core of an anticline outlined topographically by
bevelled sandstone ridges. The major granite forms developed in two stages. Fracture-controlled
differential compartment weathering beneath a late Mesozoic land surface was followed by stream
incision and the erosion of the weathered bedrock. In this way the resistant essentially unweathered
compartments and blocks were exposed and left in relief. The landscape in and near the Devil’s
Marbles is almost entirely the result of the erosional exploitation of the differentially weathered
bedrock. The residuals are thus of etch character.

THE DEVIL'S MARBLES, CENTRAL AUSTRALIA
by C. R. PwipaLt*

Summary

‘Twipace, C. R. (1980) The Devil's Marbles, central Australia, Trans. R. Soc. §. Aust. 10403),
41-49, 30 May, 1980,

The Devil’s Marbles consist of castle koppies (catcllated inselhergs) and large residual
boulders, hoth developed in granite, They are exposed in the core of an anticline outlined
topographically by bevelled sandstone ridges. The major granite forms developed in two
stages, Frocture-controlled differential compartment weathering beneath a late Mesozoic land
surface was followed by stream incision wnd the erosion Of the weathered bedrock, In this
way the resistant essentially unweathered compartments and blocks were exposed and left
in relief. The landascupe in and near the Devil's Marbles is almost entirely the result of the
erosional exploitation of the dilferentially Weathered bedrock, The residuals are thus of etch

chirticter,

The massive, simple, granite forms, largely lacking intricate fretting and sculpture, are
typical of humid tropical regions rather than lemperate lands, and stand in marked contrast
with most of the granite landform ussemblages of southern Australia.

Introduction

Though not as dramatic, large or famous as
Ayers Rock and the Olgas, the groups of re-
sidual granite boulders known collectively as
the Devil's Marbles are nevertheless well
known and constitute a considerable attraction
for tourists Visting central Australia. Located
asinde the Stuart Highway about 50 km south
of Tennant Creek, the residuals lie within
200 kin of the geographical centre of Australia
in central Mount Stuart (Fig. [) and with an
average wnnual rainfall of some 350 mm (13.7
itches) per annum are situated within the
tropical semiarid zone of the continent.

Residual granite bowders are by no means
rare, Granite comprises about 15% of the con-
tinental areas and apart from planation sur-
faces, boulders are the most common landform
development of such outcrops, The Devil's
Marbles are however unusual by virtue of ihe
size of the boulders, their arrangement, and
their detailed morphology. Many of the resi
dual houlders are 11-12 m in diameter and at
several sites these giant blocks stand im orderly
arrangements so that they look tike the rem-
nants of Cyclopean masonry (Fig. 2). ‘Vech-
ically they are castle koppics frather than
tors—see Twidale 1971), Other large holders
ure precariously perched either on other blocks
or boulders or on rock platforms: others stand
in pairs, one on the other, fo form cottage
loaves; yet others have split in two us iF sliced
by some gigantic cleaver (Pig. 3).

* Department of Geography, University of Ade
luide, Box 498, G.P.O. Adclue, 5. Ausr. S00),

General setting

The granite on which the boulders are deve-
loped is a coarse-grained grey porphyritic. mus-
covite-biotite adamellite, with large phenocrysts
of feldspar, Radiometric (K/Ar) determina-
tions give an age of about 1510 ma, ic. the
granite is Lower Proterozoic (Smith 1974).
Tt intrudes Hatches Creek Group sedinients,
also of Lower Proterozoic age (Smith 1974)

and is exposed in the core of an asymmetrical,

plunging, anticlinal structure involving these
sedimentary strata (Fig. 1). The structure is
expressed topographically in a V-shaped out-
crop of sandstone Which is part of the Daven-
port Ranges and which borders the granite to
hoth north and south, The sandstone ridge is
bevelled, 1t stands some 490 m above sealevel
and 70-80 m above the eastern lower valicy
floor, though only 30-40 m higher than the
plains at the harrow western end of the low-
land.

On the southwest the granite oaytcrop is
drained by the headwaters of Sutherland Creek
which flows south through a gorge in the sand-
stone fudge, but the greater past of the enclosed
anticlinal valley drains north by way of Dixon
Creek and its tributaries. The axial stream of
the drainage system is situated at the western
end of the Valley und is in fact followed by
the Stuart Highway. The major part of the
granite outcrop lies east of the Highway so
that there is a general decline in the altitude
of the granite surface from east to west, The
main groups of boulders are exposed near the
Highway in the lowest pari of the valley,

42 C. R. TWIDALE

Central _™@
Mt Stuart

8

= AL ge
Devils?”
Marbles

: upland surface (Early Cretaceous)
(probably pre-silcrete)

‘upland ridges in Proterozoic
sediments, with strike shown

Sp
oe

plain in sediments

mesa in weathered granite

gp

plain cut in granite, with
occasional residual boulders

Okm 3

_ high plain cut in granite
“= (slightly weathered)

groups of large residual boulders
(tresh)

Fig. 1. Morphological map of Devil’s Marbles and environs. Location map inset.

though there are others scattered over the
plains (Fig. 1).

The surrounding hills and plains provide as
much evidence concerning the evolution of
these huge balls of rock as do the boulders
and blocks themselves, and for this reason
they are considered first.

Planation surfaces
Plains of low relief are quite extensively
developed on the granite outcrop and indeed
occupy a far greater area than do the residual

boulders. There are also remnants of two
higher surfaces of low relief (Figs 1, 4).

The present plain is rolling or undulating
and is shallowly dissected by numerous stream
channels, It is eroded in granite that, though
hard and cohesive is nevertheless weathered:
the potash feldspar is generally white due to
clay mineral alteration, the biotite is partly
altered to chlorite, and there is abundant frac-
turing both along grain boundaries and across
grains, Iron staining is obvious in thin section.
The cut bedrock surface is covered by a few

THE DEVIL’S MARBLES, CENTRAL AUSTRALIA 43

. ‘ ‘ ; i a. ' _-
fi - i ie _ = —- |
i a te ns 1 ‘ =. ~~

.
s iA

Fig. 2. Castle koppie in Devil’s Marbles complex consisting of massive quadrangular blocks, most in
situ. The residual rises above valley floor beyond which is sandstone ridge with markedly bevelled
crest,

&

>, - =

| <n < : 7
e.. . -

Rahs, i ,

Meee ee ie SN tinned ano b8 .

Fig. 3. Split boulders at Devil’s Marbles. Parallelism of fractures is noteworthy. Fractures can be
traced through several boulders some tens of metres apart.

44 C. R. TWIDALE

high plain (granite) summit surface
sandstone
midslope platform

“ar- Pe ine om - >
Rien
SX SS SANA
RRS - = =
vo sf ‘ses —_ gpm» _— — ——_—___— —_—

met tae as tension
zone of fe
fluctuating ft neutral plane
water level :

compression

7

Fig. 4. Diagrammatic cross-section through Devil’s Marbles and adjacent areas.
Fig. 6, Field sketch of granite high plain and matching platform cut on slope of sandstone ridge,
Fig. 7, Two-stage development of Devil’s Marbles. Dots indicate weathering.

THE DEVIL’S MARBLES, CENTRAL AUSTRALIA 45

centimetres of granite sand (quartz with frag-
ments of feldspar and mica). Near the sedi-
mentary outcrops at the margins of the plains
small plates of sandstone are a_ significant
constituent of the regolith, and in the vicinity
of the few ridges underlain by quartz blows or
reefs, white vein quartz is also abundant.

Small boulders and blocks stand above the
general plain level in several areas. Their lower
slopes are commonly roughened and fretted, or
pitted (see Twidale & Bourne 1976), indicat-
ing recent, possible local, lowering of the soil
surface of 20-25 cm. In addition the major
granite forms known as the Devil’s Marbles
rise from this plain (Figs 1, 4).

Quite extensive remnants of a higher plain
are preserved on the granite (Figs 1, 4).
Bounded by rudimentarily faceted slopes up
to 5 m high, this high plain carries a thin
discontinuous veneer of granite sand. Granite
platforms, with residual boulders and blocks,
are also present. The granite exposed in this
high plain is more weathered than that beneath
the present plains. The potash feldspars are

white due to partial kaolinisation, while the
plagioclase has been strongly altered to an
orange-brown clay, identified as kaolinite prob-
ably discoloured by goethite, The rock is ex-
tensively fractured as a result of weathering
and iron oxide and clays are commonly found
both in fissures and along crystal boundaries.

Standing 25-30 m above the high plain are
three mesas capped by highly weathered
granite. Quartz and muscovite are the only un-
weathered minerals, though the latter is altered
along its cleavage. The rock samples appear
to differ from the main mass of the outcrop
and to be a greisen or primarily altered granite.
The feldspars are completely altered to a
brown orange kaolinite (again probably with
goethite) which also fills intergranular spaces.
The residuals are bounded by faceted slopes in
which coarse blocks of granite are exposed
(Fig. 5).

Both high plain and mesas can be correlated
with planate features preserved on the sand-
stone ranges. Though standing a few metres
lower, the mesa surface with highly weathered,

a 7 ee i 4
96% ae
aT wan ents

Fig. 8. Parted block at Devil’s Marbles,

46 Cc R, TWIDALE

though still cohesive, granite can nevertheless
reasonably be related th the prominent summit
hevel of the Davenport Rasiges (Figs 1, 4, 5).
The tigh plain has ils counterparts in a dis-
Unct platform developed and preserved at
midslope on the inward-facing escarpments of
the sandstone ridges (Fig, 6). Thus the land-
scape around the Devil’s Murbles is multicyclic
in character. Each of the two lower surfaces
is contiguous over wide areas, und the three
display different degrees of hedrock weather-
ing; they are not identified solely on thy basis
of relative elevation.

Mabbutt (1967) has referred the summit
surface of the Davenport Ranges on what he
ealls rather slight palacogeographic evidence.
to the later Mesosme. This supguestion tinds
support in equivalents of the surface in the
Barrow Creck and Alice Springs aves standing
higher, and being therefore older, than silerete
remnants of carly-mid Tertiary ave (Wopfner
& ‘Twidile 1967), The bevelled ridges of the
Devil’s Marbles region also stand higher than
lateritised surfaces to the north, and as these
also are of early-rmid Tertiary age (Stewart
1954; Twidale 1956; Hays 1967), the later
Mesovoic age of the summit surface ix cor
roborated. The summit surface is in fact part
of a later Mesozoic land surface of Jow relief
ihat has been reported from several parts of
tropical and sublropcal Ausiralig (see eg,
Woodard 1955; Twidale 1956, 1966, 197fa,
198%; Wright 1963; Mabbutt 1967; Hays
1967),

This summit surface and tts eqyivalent in
the mesas developed on granite is the datum
to which the formation of the residual boulders
can be referred, for all of the Devil's Marbles
ogeur below it (Fig. 4), It is a weathering
surface that has suffered leaching, strong
kaolinsation above the weathering rimt or
lower limit of significant weathering (Mab-
butt 1961), and precipitation of tron oxtdes
near pnd For some metres below this level.

Major granite forms

hat the summit surlace itplics a period of
relative standstill and weathering of the wader,
lying bedrock is demonstrated by the altered
ytanite preserved in the mesas. Had the granite
been homogencous there would have beer imi
form weathering beneath the plains surface,
but the bedrock is jointed wnd altheush he
cause of erosion it 1s nat possible ty discern
what joint patterns bad developed in the
srinite bencath the contiguous stuninit surface,

there jire indications that fracture spacing
varies both Verteally and laterally. For ex-
ample whereas the joint harks in some of the
Wore prominent residuals are 10-12 m in
diameter the joints exposed in creek beds
nearby are only 20-30 cm apart. Again, the
large residuals stand on plinths that are even
MOre Missy.

Observations in many parts of the world
Suggest very strongly that granite is subdivided
ito massive and well-jointed compartments.
The latter are readily and rapidly westhered
while the former remain intact. As the land
surface is lowered the weathered rock is
eroded, leaving the stillefresh and cohesive
compartments in relief, as residuals the size
and shape of which depends on fracture pat-
tern (Fig, 7). ‘Thus it is argued and has been
urged for almost two centufies (see Hassen-
fraty 1791; Logan 1849, 185): Mennell 1904;
Twidale 19784) that the granite residuals
evolve in two stages, one involving differential
joint-controlled subsurface weathering, the
other the differential crosion of these unequally
weathered compartments (Linton 1955); Biidel
1957; Wilhelmy 1958; Twidale 1971; Ciodard
1977)

In the case of the Devil's Marbles, differen-
tial comparttient weathering look place he-
neath the late Mesozoic fand surface repre-
sented by crestal bevels. preserved on the
sandstone ridges and hy mesa remnants. The
differential subsurface weathering took place
during the later Mesozoic and carly Cainozoic
when this region, like the remainder of central
Australia, was warm aod humid to subhumid
(Kamp 1978), The exposure of the koppies
and boulders, on the other hand, is. a later
Cainozoie event probably related to regional
warping and resultant stream rejuvenation

In these terms the castle koppies arg com-
parltnents characterised by widely spaced sets
of orthogonal joiuts, though the flat-lying joints
afe almost certainly either the Luverklifte of
Cloos (1922) of sheeting joints (Gilbert 1904;
‘l'widule 1973), The residual boulders are large
cerestones or kernels remaining after the
marginal weathered zones of the joint blocks
Wave been ered, The perched blocks and
coltuge loaves are to some extent fortuitous,
though lyecause weathering advanecs down
from the surface it can be expected that more
residuals Will Survive at depth than closer to
the surface. sa that in some places isolated
blocks will come to stand on broader bases
(Pig. 7).

THE DEVIL'S MARBLES, CENTRAL AUSTRALIA +

The present plain ig related to present local
baselevel (Dixon Creek) wnd undoubtedly re-
Meets aw lowering of baselevel compared to
later Mesozore (Surimit surface) times, The
high plain however could cither be an ctch
surface or exposed weathering front (Wayland
1934). or jt could be related to a baselevel
intermediate between the summit surface and
the present plains, The former interpretation
appears the more likely, for the variations in
veanite weathering appear ito be part of a single
profile rather than ovo or more. The platforms
previously noted (Fig. 6) on the inward-facing
sandstone ridges, and toterpreted as extensions
of the high plain stirface, could be of stimtlar
elch type and due to particularly intense
moisture atfuek i ihe then scarp foot zene
Thouth the high plain slopes down to the west,
the weathering front was probably irregular
ii detail because of variations in joint spacing
and glso because of ihe slope of the wuter
lable to the major water courses. The blacky
upstanding koppies and large boulders, which
were presumably once surrounded by well-
jointed weathered rock, also stam on much
broader, plinths or platforms of massive
cranite-compartments thar survived to a inuch
preater extent than thase above them because
they remained beluw the weathering front
(Pig. 7).

Thus. the major Jondforms of the Devil's
Marbles are readily comprehensihle in terns
of the two-stage cofeept. The residual rem-
nants, though angular, probably represent
dome structures developed as a result of com-
nvession in the erust (ef, Denham ef al, 1979),
They have been modified by aggressive
weothering ot depth and at the margins of the
massive conioartnients, Where ground waters
persist. and in contrast with the near-surlace
dner, or seasonally dry, zones,

The minor forms

The boulders, blocks, and large-radius
dornes are fretted and seulptured in detuil,
though the resultant modifications are in most
instances minor, Some blocks and particularly
some of those high on residuals are bath in
Preenated with jron and manganese oxide and
creally fretted and hollowed (alveolar weather-
ing}, Polygonal patterns of surface cracks are
developed in these impregnated zones, These
weathering phenomena argue the presence of
water and the zones of intense attack may be
related to former fluctuating water tables. On
the other hand similar intense fretting is found

at some sites at the bases of blocks and
houlders, close to the old weathering front,
and spain abundant water is suggested as the
reason for the especially aggressive weathering.

Some Jarge boulders are clongate and
streamlined and are called whalebacks or doy
de baleme. A few large blocks have seemingly
moved laterally telalive to one another (Pig.
8), rather like the purted blocks deserihed
front Dartmoor (Worth 1953), Many blocks
are split into two parts (Fig, 3), Some
secondary joints have heen exploited by
weathering to form shallow straight grooves
(Kluftéarren), though there ure on some
domes true Rille (Echikarren) due to stream
erosion, Some of them are coated with a black
compound, consisting of carbon plus oxides of
iron and manganese (Francis 1921). Similar
coatings On granite occur in former pools in
creck beds, indicating that this black material
has been deposited fram water. These black
colours arc however, unusual, for virtnally all
of the residuals catry a patina of reddish-
brown iron oxide which etfeetively masks the
grey colour of the granite seen in ereek beds
and where there has been fire flaking, Sealing
of the rock surface is peneral, and pitting, due
to differential weathering of surfuce crystals
leaving quartz in microrelie! has been noted
on lower slopes, Rock platforms in places dis-
play regular patterns of vertical joints so that
the whole looks like a pavement, Al some sites
the joints are weathered and there are minor
flared slopes developed in the sidewalls of the
clefts so formed,

Basal fretting has already been referred to
and some boulders display poorly developed
flared slopes. A fow basal tafoni have been
noted, but it is because of the virtual ahsence
of this and other forms (flates, enammas or
weather pits, gutters or Rille) that the Marties
stand in such contrast with granite exposures
in southern Australia (see Twidale 1971,
1976b; Twidale & Foale 1972).

The origin of most of these minor land-
forms is well known and ts adequately dis-
cussed elsewhere (Wilbelmy 1958: Twidale
1971, 1976b; Twidale & Poale 1972: Twidale &
Bourne 1975),

Several lines of evidence point to the minor
forms having been initiated by moisture attack,
in the subsurface, at the weathering (rant, Tn-
ciplent pitting, fared slopes, platforms, tafoni,
saucer-shaped depressions and gutters have
been observed already developed on the hed-
rock surface when the rogolithie cover was

48 c. R. TWIDALE

stripped away and some forms, particularly
Rille. can be traced into the subsurfyce
{Logan [851, Twidyle 1962, 1971, 1976),
1978b; Boyé & Fritsch 1973; Twilale & Bourne
1975, 1976, 1977), The cover 1s weathered
granite in situ, not introduced material, so that
there is no question of the minor Jealures
developed in granite having been formed on
exposed surfuces and then buried, The forins
are modified—either developed or destroyed —
aller exposure, but they are initiated in the
subsurface.

Only three of the minor landforms call for
further brie! comment. First, with reward to
parted btteks, Worth (1953) rightly dis
counted gravity, for the blocks stand ow fut
or Very gently sloping inclines. lee vr nivel
action can be discounted at the Devil's
Marbles, and shaking by earthquakes seems
Wolikely, for not only is the area siesmically
quiet hut had there been carth tremors suffi-
cient to dislodge the parted blocks, no perched
hontders would strely have survived.

Second, split rocks have been attributed by
several writers lo healing and cooling under
desert vonditious (e.g, Hume 19275; Hills
1975), Tt is very doubtful whether this pro-
cess alone, or even aided by ihe cooling effect
of rain showers, could achieve the splitting of
such massive bodies of rock. Furthermore split
rucks are even more common in pival revions
fee. the Pyrenees) than they are in arid areas.
It is suggested that three factors are involved,
Though the houlders and bloeks thar are split
we basically defined by orthogonal joints, it
is a matter of Observation that many inclide
ofher, presumably secondary or latent joints
(Fig. 3), These subsidiary [ractures are ex-
Ploited by weathermg processes and notably
by walerio either liquid or solid form, depend
ing oon climatic conditions. Finally arnce

weathering has sufficiently affected the subsr,

diary fracture and provided tt ts vertically ar
near-vertically disposed, the weieht of the twa
halVes will cause the weakened fracture to be
pulled apart, so creating the split rack.
Third, polygonal erackine is clearly net a
primary rock feature ws sugeested by some
writers (Johnson 1927; Leonard 1929), for i
is developed on boulders whieh are Lhemselyves
the product of differential subsurface weather-
ing At Devil's Marbles and iw the Pilbara
of Western Australia it Nas been aoted thal
polygonal cracking is mssocialed with rock that

is heavily impreenated with iron and oian-
ganese oxides, In the Pilbara, near Mount
Magnet and oo Eyre Peninsula the cracking
frequently affects two or Mose superiniposed
Jayers of rock, and some of the plates de-
lineated by the cracks are arched slightly
upwards, Various mechanisms have been cone
sidered and found wanting. The most likely
is thal salts of iron and mangunese which are
mobile are carried in groundwaters and pie-
cipitated out near the weathering front, Le-
near the margins of the boulders whilst they
are sull corestones and beneath the land sul:
face. The continued precipilation of salts
causes the buckling of the rock in the direction
of least resistance, namely outwaris and We
fracturing of the shells into polygonal plites.
Lastly it is emphasised that the Devil's
Marbles are simple groups of residual boulders.
lacking any greal variely or development of
minor forms. With regurd te the latter, the
forms that are absent, or virtually so, are as
interesting ws those that are present. Thus,
flared slopes, tafani, Rifle and rock basins are
rare or poorly developed. In this respect the
Devil's Marbles are more akin ta the tropical
granite cxposures of northwest Qucensland,
the Pilbara and the Darwin area rather than
those of the temperate, particalarly the Medi
lerranean environments of much of South Aus-
tralia, southern Victoria and the Wheat Belt
al Western Australia. This is oot ta sugpest
either that flared slopes for instanee are not
found in the northern areas (though they are
rare and poorly developed there) or that they
have evolved to a spectacular degree on all
southern outcrops, Many factors are mvolved
and in many respects it is not a simple ques-
ion of development or pon-development, but
rather of degree of development und preser-
vation, Nonetheless, zoned contrasts are dis
cernible and the Devils Marbles both in gross
and in detail have more affinities with the
frumid tropics than with the tempetate and
Mediterrancan lands of southern Australia.

Acknowlevlgments

The author wishes to thank Dr A. R. Milnes,
C.S.LR.0. Division of Soils, for notes and
comments on various petrographic samples,
and Liz Campbell for support in the fleld. The
field work was carried out with the aid of an
award from the Austrulidg Research Grants
Crmrieniitec

THE DEVIL'S MARBLES, CENTRAL AUSTRALIA 49

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FURTHER OBSERVATIONS ON SPERMATOZOAL MORPHOLOGY AND
MALE REPRODUCTIVE TRACT ANATOMY OF PSEUDOMYS AND
NOTOMYS SPECIES (MAMMALIA: RODENTIA)

BY W. G. BREED

Summary

Spermatozoa of Notomys cervinus have a head with three hooks, whereas those from N. fuscus are
variable but have only two very short hooks. Spermatozoa from Pseudomys forresti, P. fumeus, and
P. gracilicaudatus all have a head with three hooks. Only one very short hook occurs in P.
novachollandiae, and there are no hooks in spermatozoa from P. delicatulus, P. shortridgei has a
spatulate sperm head with a large acrosome. Insertion of the sperm tail is lateral in novaehollandiae,
offset basal in P. delicatulus and mid-basal in P. shortridgei. Principal and end pieces are shorter in
the last two species.

FURTHER OBSERVATIONS ON SPERMATOZOAL MORPHOLOGY AND
MALE REPRODUCTIVE TRACT ANATOMY OF PSEUDOMYS AND
NOTOMYS SPECIES (MAMMALIA: RODENTIA)

by W, G, Breen*

Summary

Breen, W, G. (1980) Porther observations on spermatozoal morphology and male reproductive
tract anatomy of Psendomys andNolomys species (Mammalia: Rodentia). Trans, 2. Soc. 8,
Aust. W4(3). 51-55, 30 May, 1980,

Spermatozon of Notomys cervinus have a head with three hooks, whereas those from
N. fuseus are variable bul have only two very short hooks, Spermatozoa from Pseucdormys
farrestl, P. fumeus, and P, gracilicaudatas all have a head with three hooks, Only one very
short hook occurs in P. norachollundiae, and there are no hooks in spermatozoa from P-
delicatulus, P. shortridgei has a spatulate sperm head with a large acrosome, Insertion of the
sperm til is lateral in P. novachollandiae, ollset basal in P, delieatulus and mid-basul in
P, shorirideei. Principal and end pieces are shorter in the last two species.

Notamys alexis, N. fuseus, and N, mitchellii have very small testes, whereas lhey are
relatively larger and scrotal in N. cervinus. In the first three species seminal vesicles and
congulating glands are vestigeal or non-existent, whereas in N, cervinus they ore well developed
and weigh up to 150 mg, The possible phylogenetic and functional significance of these

differences is discussed.

Introduction

Phylogenetic relationships between different
species of rodents have been investigated by
a variety of techniques, all of which have at
least some limitations. In several groups of
North American and British =myomorph
rodents spermatozoal morphology and cum
parative anatomy of the male acecssory sex
wands have been used (Friend 1936, Bishop &
Walton 1960, Arata 1964, Linzey & Layne
1969, 1974), The latter uuthors concluded
that in Peromyscuy Variation in accessory sex
glands corresponded well with major taxo-
nomic groupings based on other criteria,
whereas variation in spermatozoal morphology
did not coincide with the subgeneric eroups,
Breed & Sarafis (1979) extended a brief report
by Illison (1971)1; investigated spermatozaal
morphalogy and male reproductive tract ana-
tomy in some species of Australian Todents,
and discussed phylogenetic relationships he-
lween Various genera. They found that within
each genus similar spermatozoal morphology
oceurred apart from Noatamys. Wison (1971)!
reported that Pserdomys shurtridged has a
primilive sperm type, and P. delieatuluy (as
Leeeadina delivatila) a spermatozoal head
with no books, so it appears that Pseudontys

~ Department of Anutomy and Histology, Vni-
versity of Adelante, GPO. Box 498, Adelaide,
S. Aust. S001,

7 HHlison, by Abstract of paper presented at Aust
Martitnal Soc. Meeting val. 2, No. 8, Dee, 1971,

exhibits considerable spermatozoal diversity;
all five species that we investigated had a
sperm head with three hooks. Male accessory
sex glands were generally similar between the
genera investigated, apart from the two species
of Notomys which differed markedly (Breed &
Satafis 1979), Here spermatozoal morphology
of six other Preudwrys and two other Notontys
ypecics is presented, together with an examina-
tion of the male reprodnetive tract from all
four Notoennyis species,

Materials and methods

Spermatozoa were obtained fram single

adult male individuals of the following species
housed ut the Institute of Medical and Veter-
nary Science (1,M.V,S,), Adelaide,
Netomys cervinus: Boru al University pf NLS.W,,
received JWIV.74; N, fuscus; Laboratory bred at
IMVS,, Peevdomys delicatulys! collected at
Shortcut Rd, Nourlangie, NOT. ChM.Y.S. publ,
1017): P, forrestiz received trom University of
N.S.W,, 10.1V.79 (1M,V,S, publ. 953)5 PF. faerneus:
from Arthur Ryluh Institute, Melbourne (1.M_V,S
publ. 952); P. gravilicavdatus: (rom School of
Biological Sciences, Macqtiarie University
(LM.V.S, publ, 950); FP. shortridgei: from
Fisheries and Wildlife Division, East Melbourne
(1.M.Y,S, publ. 951),

In all individuals a small incision was made
into the region of the tail ol the epididymis
under halothane anaesthesia. and a small bi-
apsy taken. Spermatoyea were then squeezed
onl onto microscope slides previously Mouded

52 W. G,
with glutaraldehyde/formaldehyde/ picric acid
fixative (see Ito & Karnovsky 1968). Semi-
permanent mounts were made by placing a
coverslip on top of the sperm and ringing the
mount with DePeX. Subsequently the sperm
were photographed with Nomarski optics at
CSIRO Division of Horticultural Research,
Glen Osmond. Measurements were made of
several spermatozoa from each species under
phase contrast using an eyepiece micrometer
as described by Breed & Sarafis (1979), Al-
though difficulty was experienced in obtaining
accurate Measurements, the mean values for
head length, midpiece, and principal together
with endpiece were determined.

Adult male individuals of Netomys mitehel-
lit, N. alexis, N. cervinus and N. fuscus col-
lected in the field (1960-1978), preserved in
formalin or 70% alcohol, and lodged at the
S.A, Museum, W.A. Museum or 1.M.V.S.,
were Weighed. A single testis, seminal vesicles
with coagulating glands, and ventral prostates

BREED

were dissected out, cleared of adherant fal,
subsequently weighed and, where appropriate,
measured. The testis weight was subsequently
doubled to give the approximate weight of
paired testes. A few laboratory bred adult
Notoniys held at 1.M.V.S. or Medical School,
University of Adelaide, were also killed and
tale reproductive tracts dissected out and
weighed. A single N. cervinus (provided by
Mr R, Briggs) was weighed, dissected and in-
vestigated similarly.

Results

Table 1 and Fig. | present morphological
details of spermatozoa determined by light
microscopy. Sperm of P. forresti, P. fumens,
and P. gracilicawdatusy had a head with three
hooks, although the size of the sperm head
and length of hooks was greater in P, gracili-
equdatus. The sperm head of P. novaehol-
landiae had a single short top hook and a
truncated or non-existent lower hook, That of

Fig. 1. Spermatozoa. A:

principal piece, ac

Pseudams gracilicuudains; B: funteus; C: P. forresti: D: P, novaehollandiae:
E: P. delicatulys; B: P. sheriridgei; G: Notoniys cervinus: A: N. fuses. mp == middle piece, pp =

acrosome, cd = cytoplasmic droplet.

OBSERVATIONS ON SPERMATOZOAL MORPHOLOGY 53

TaBLE 1: head and tail length of spermatozoa from various Notomys and Pseudomys species.

Species Average size of spermatozoa (um)

Principal and
Head Midpiece endpiece Total
Notomys fuscus 7 22 77 106
N. cervinus 9 26 80 115
Pseudomys delicatulus 4 19 65 88
P. forresti 7 26 95 128
P. fumeus 8 20 100 128
P. gracilicaudatus 10 21 98 129
P. novaehollandiae 6 22 78 106
P. shortridgei 5 22 69 96

P. delicatulus had no hooks and was smaller
than the others. P. shortridgei also had sperm
with no visible hooks. It was spatulate in shape,
and a large acrosome occurred over the
nucleus. The principal and endpieces were
shorter in the last two species.

Notomys fuscus had variable sperm head
morphology, but usually there were very short
and truncated top and lower hooks. N. cer-
vinus sperm had a head with a long top hook
and two lower hooks united at their base.

In wild caught N, alexis testes weight ranged
from a mean of 20-37 mg with time of year.
Often no obvious scrotum was discernible.
Seminal vesicles were at most only just visible
in the preserved material and did not exceed
3 mm length. Coagulating glands and dorsal
prostates were vestigeal or non-existent. Large
ventral prostates occurred which varied in
weight during the year; no seasonal trend was
apparent. Laboratory bred adult N. alexis have
similar reproductive tract anatomy (Breed
1979, Breed & Sarafis 1979).

Similar male reproductive tract morphology
occurred in N. mitchellii and N. fuscus (Table
2). By contrast, that of N. cervinus was
markedly different (Table 2, Fig. 2). The
testes were relatively larger and scrotal in posi-
tion, although adult body weights were similar
to the other species. Conspicuous seminal
vesicles and coagulating glands were present;
their average weight being about 130 mg and
length about 10 mm.

Discussion

The suggestion by Breed & Sarafis (1979)
of intrageneric differences in spermatozoal mor-
phology and male reproductive anatomy in
Pseudomys and Notomys has been confirmed.
Most Pseudomys and two of four species of
Notomys have spermatozoa with three hooks.

In P. novaehollandiae there appears to be only
one short truncated hook, and all three hooks
are missing in spermatozoa from P. delicatulus
and P. shortridgei. Preliminary transmission
electron microscopy (TEM) has confirmed a
large, somewhat eccentrally placed, acrosome

Fig. 2. Male reproductive

tracts. A: Notomys
mitchellii; B: N. cervinus. t = testis, sv =
seminal vesicle, vp — ventral prostrate, cg =
coagulating gland, vd = vas deferens,

54 W. G. BREED

TApte 2; Adult reproductive organ weights of male Natomys (mean + S.E.)
Si pe ee a a ag

Seminal
Ventral vesicle &
prostrate coagulating
No. of Bady wt Testis wt wr gland wt
animals (g) (mg) (meg) (mg)
Notomys alexis
Fixed tissue
Month of capture
Jan & Feb 6 20744 22+4 62 + 26 —_
Mar & Apr 16 30+ 0.5 31+6 8426
May & June 19 30+3 37 +4 77+9
July & Aug 18 3043 22+3 5542
Sept & Oct 4 3143 3021 125 +30
Noy & Dec 3 37 tee 31+11 98 + 26
Fresh tissue 7 3 28+ 2.6 38 +3 8948 —_—
ide 4 29+72 3329 97 +20
Noatomys mitchellii
Fixed tissue 13 392+2 50+ 3 162+ 21 —
Fresh tissue 3 48+ 2 Jx47 407 + 223
Notomys fuseus
Fixed tissue 9 2842 4344 5817 =
Natomys cervinus
Fixed tissue 12 3842 133 +8 WoH7 130421
(10 + 0.7) #####
Fresh tissue 1 32 170 103 150
(11) ##4*

* Vestigeal or non-existent: maximum length of seminal vesticlaes — 3 mm,

j¥ From Breed 1979 (80 days of age).
*** Brom Breed & Sarafis 1979,
atch Body wt of only one animal recorded.

*#24> Length of seminal vesicles (mm) given in parenthesis.

in P, shortridgei, and a few cup-shaped cya-
ginations in the apical part of the nucleus, In
P. delicatulus the acrosome appears consider-
ably different and somewhat complex in struc-
ture,

TEM has also shown that the connecting
piece of the tail in P. shortridgei hag a mid-
basal insertion into the sperm head, an offset
basal insertion in P, delicatulus and a lateral
insertion in P. nevaehollandiae and other
species with the more typical sperm head mor-
phology. N, fuscus, N. alexis, and N. miitchellii
all appear to have somewhat variable sperm
head morphology and the former two species
truncated. or nonexistent, hooks (see also
Breed & Sarafis 1979).

Most non-Australian murid rodents inyesti-
gated have a sperm head with a single top
hook (vide Friend 1936, Bishop & Walton
1960), whereas most species of Australian
rodents have sperm heads with at least two,
and usually three. hooks (Illison 1971, Breed
& Sarafis 1979). It seems plausible that some

ancestral, non-Rattus, murid rodent evolved a
sperm head with this complex structure, and
that adaptive radiation of the group within
Australia then occurred, most species main-
taining this specialised sperm head morpho-
logy. The reduction, or lack of, hooks in the
few Notomys and Psendomys species is, there-
fore, probably a recently derived state. Varia-
bility in sperm head morphology in the twa
Notemys species with truncated hooks may
support this contention.

The anatomy of the male accessory sex
organs of Pseudumyys investigated is similar.
regardless of the differences in spermatozoal
morphalogy, There are relatively large ‘leaf-
like’ seminal vesicles 6.5 mm or more in length
in P. gracilicaudatus and P. australis (Taylor &
Horner 1972), P. hermannshurgenesis (Taylor
& Horner 1970), P. apodomoaides, P. forrestt,
P, novaehollandiae, and P. delicatulus (Breed,
unpublished). Taylor & Horner (1972) state
that no coagulating glands follow the pasterio-
lateral contour of the seminal vesicles in P.

OBSERVATIONS ON SPERMATOZOAL MORPHOLOGY as

australis, but that in laboratory-bred animals
such glands were found, and the mixing of
secreuens Of seminal vesicles and coagulating
glands results in the occurrence of hard gela-
fines material. Vaginal plugs, which in com-
mon laboratory rodents result feum a mixture
of secretions from sentinal vesicles and coagu-
laling glands (Mann 1964), have been fe-
corded in laboratory bred PL aistrelis (Smith,
Watts & Crichton 1972),

In Notomy'y considerable inter-specifie dif-
ferences occur in accessory sex gliml and
testicular development. N. cervinus has the
typical complement of male actessary sex
alands, whereas in the other species the seminal
vesicles, couguluting glands. and dorsal pros-
states are vestigeal or non-existent, No cervinns
is also the only species of Netomys for which
a vaginal plug has been treeorded (Crichinn
1974) and it also has typical serorul testes,
wheres they are relatively much stiller and
perisnal in position iw the otbers, Lack of
testicular development presumably results im
the absence of an obvious scrotum thal often
aceurs if these species.

The above findings jndivate, therefore, thal
spermatozoa and the mule accessory sex alanis
are samewhat labile structures, as considerable
variation occurs within Netumys that, on other
morphological and physiological groynds, ap-
pears to be a discrete and closely related group.
Since N- cervinus bas the more vonventional
pattern of male reproductive tract anatomy, the
occurrence of the vestigeal glands, very sriall

testes, and the apparent lack of vaginal plug
formation in the other three species is likely
to be a recently derived state. The tunetional
significance of these differences hus yet to be
elucidated, but it may be significant that rela-
tive testis size appears to correlate with senvinal
vesicle and coagulating gland development
Perhaps in species With relatively small testes,
fewer sperm ure produced, stored, and thus Te-
leased at ejaculation, Less energy, and there-
fore seminal fluid, would thus be required for
ther metaholism in the female ceproduclive
tract, This, in turn, may result in atrophy of
sume of the glands involved in the production
of seminal Huid. Why such divergonee should
evolve in a closely related group of species ts,
al present unknown, bul a study on the sexual
and social behaviour of these animals is being
carried out, and this may shed some light on
the significance of these observed anatomical
differences.

Acknowledgments
1 thank Drs P. Baverstock and C, Watts for
enabling me to obtain sperm from their
animals, Dr D. Kitchener {W.A. Museum)
und Mr P. Aitken (S.A, Museum) for allow-
ing me access ty material, and Mr Ro Rriggs
for the donation of a single Notonrys cervinter,
l also thank Br A. Bird, C.S.LK-0, Division
ol Horticultural Research for permitting me
to use his Nomarski microscope, and Mr R.
Murphy, Mr C, Leigh, Mrs G. Hermuanis, Mr's
|, Brazier and Mrs B, Sheldon for assistance

in the Department of Anatomy,

Referenees

Arata, A. A, (1964) The anatomy und roxena-
mic significance of (he male accessory reprodiic
tive lands of curl rodents. Aull, Florida State
Mus. 9, 1-42-

Baverstoce, P,, Warts, ©, HOS. & Hoar, 1 'T,
(1977) Chromosome evolution in Austeallan
rddenis. | ‘The Pseudomyinae, the Hydromyinue
and the Uromys/Melomys group. Chromosoma
Gi 9§-(25.

Aisin, M. 'W. EL & WaAtron, A. (1960) Sperma-
tovenesis and the structure of mammalian
spermatozoa, Claptor 7 in A. S, Parkes (Fd.
“Marshall's Physiology of Reproduction” Vol,
112) fLongmans, Tondon).

Bheep, W. G, (1979) The reproductive rate of
the hoppine-mouse Nelamys alexis and is ecea-
logical significance. Anse. J, Zaol. 27, 177-94,

& Sansers, Voo61979) On the phylogenetic
significance of spermainzoal morphology nnd
male renraduerve tract anutomy in Australiuin
rodents Trang. 2, Sac S, Anse, U3, 127 035-

Crician, Fo. (1974) Aspects of reproduction
ii the genus Veron (Muridae), Aust, 4, 200,
22, 439.447_

Fauienp, G- F. (1936) ‘The sperm of British muri-
dae. Quart. #, Micros, Sei, 78, 419-443.

Tro. S & Karnovsky, M. 2.17968) Formaklelyde-
wlutaraldehyde fixatives containing dinitra come
pounds, J. Cell Biol. 9, 168A-169A.

Linzky, A, Vo & Layne, I, N. (1968) Compare
tive morphology of the mile reproductive tract
in the rodent genus Perenmyscus (Muridwe)
Aimer, Mus, Navir, (2355), 1-47,

—— (1974) Comparative morphology of sperma-
tozog of the rodent genus Peraeyaruy (Muri-
dace), Thid, (75324 1-20. :

Mann, T, (1964) “The biochemistry and semen
uf the mule reproductive tract.” Methoen, Lune

on.

Suma, TR. Warts. C. HOS, & Cemcrox, EG
(1972) Reprodoction in the Australion desert
rodents Nofomwys alexts and Psencdanive aystea-
lis (Muridne). dusr, J. Mamearol, 1. 1-7,

Tayiorn, J M & Horner, BOR, 11970) Observa-
tions on reoraduction in Leevadinw (Rodentn.
Miridne) SL. Agammral $1, 10-17,

_— (1Y72) Observations on the reproductive
Hulowy of Prydemys (Rodentia) Mirtidac).
(hid 53, 318-428

FOSSILFEROUS LOWER DEVONIAN BOULDERS IN CRETACEOUS
SEDIMENTS OF THE GREAT AUSTRALIAN BASIN

BY R. B. FLINT, G. J, AMBROSE & K. W. S. CAMPBELL

Summary

During 1977-78, 32 fossilferous Lower Devonian quartzite boulders were discovered within
Mesozoic sediments along the southwestern margin of the Great Australian Basin. Previously only
two such specimens had been discovered in South Australia though similar occurrences have been
known in New South Wales since 1898. Fossils not previously recorded in S.A. include the fish
Wuttagoonaspis, the brachiopods Howellella jaqueti and Sphaerirhynchia sp.; the bivalves
Leptodesma inflatum, Sanguinolites sp. and Praectenodonta sp.; the gastropod Strapollus culleni;
and abundant tentaculitids. Similar fossiliferous Devonian rocks are not known in situ in S.A. The
probable source area is the fossilferous Amphitheatre and Mulga Downs Groups near Cobar in
N.S.W. It is suggested that boulders were transported to S.A. during the Permian glaciation and then
reworked into Cretaceous bouldery shales and sands. All but two of the boulders are found within
conglomeratic sediments at the base of the Bulldog Shale. Theories on transport processes during
the Cretaceous ate discussed; it is concluded that conglomeratic sediments at the base of the
Bulldog Shale are reworked submarine debris-flow deposits.

FOSSILIFEROUS LOWER DEVONIAN BOULDERS IN CRETACEOUS
SEDIMENTS OF THE GREAT AUSTRALIAN BASIN

by R. 8 FuIntt, G. J, AMBRose*, & K. S. W. CAMPBELL?

Summary

Font, R. B., Amprose, GP, & Camenuia, K. 8S. W. (1980) Fossiliferous Lower Devonian
boulders in Cretaceous sediments of the Great Australian Basin, Trans, R.. Soc. §, Aust.
104(3), 57-65, 30 May, 1980,

During 1977-78, 32 fossiliferous Lower Devonian guartzite boulders were discovered
within Mesozoic sediments along the southwestern myargin of the Great Australian Basin.
Previously only two such specimens had been discovered in South Australia though similar
ocenrrences have been known in New South Wales since 1898. Fossils not previously recorded
in S.A. include the fish Wuttagoonaspis, the brachiopods Howellella jaquerl and Sphaerivhynchia
sp.; the bivalves Leptodesma inflaturn, Sanguinelites sp, and Praectenodonta sp., the gastropod
Straparallus calleni; and abundant. tentaculitidy Similar fossiliferous Devonian rocks ure not
known in site in S.A. The probable source area is the fossiliferous Amphitheatre and Mulga
Downs Groups near Cobar in N.S.W, Ii is suggested that boulders were transported lo S.A.
during the Permian glaciation and then reworked into Creiacebus bouldery shalos and sutids.
All but two of the boulders are found within conglomerutic sediments at the base of the
Bulldog Shale, Theories on transport processes during the Cretaceous are discussed; |t is
concluded that conglomeratic sediments at the base of the Bulldoy Shale are reworked sub-

marine debris-flow deposits.

Introduction

Fossiliferous Devonian quartzite boulders
from Cretaceous sediments were first de-
scribed from White Cliffs Opalfield in NSW.
by Dun (1898). P, J, Russ collected the first
fossiliferous boulder in S,A. in 1966 Tron) an
opal shaft at the Andamooka Opalfield It was
thought at the time that an opal miner may
have brought the boulder to S.A, from White
Cliffs, However, aller discovery of ja second
fossiliferous boulder near Dalhousie Springs by
M. C, Benbow, the geological implications
Were assessed by Campbell e¢ al, (1977). They
concluded that the fossiliferous boulders, like
those at White Cliffs. were derived from the
Devonian Amphitheatre Group near Cobar in
N.S.W,. and that they were transported to
S.A. during the Permian glaciation, later to be
reworked into Cretaceous steuta.

* Geological Survey of South Austraha, 191
Greenhill Rd, Purkside. S. Aust. 5063.
Department Of Geolegy, Australian National
University,

1 Carr, 8, G., Oliver, J. G,, Conor, C. H. RH. &
Scott D. C. (1978) Andamvooku Opal fields; The
gcology of the precious stones field and the re-
sult of the subsidised mining programme 5.
Aust, Dept Mines & Energy Rept 78/5 (unpub-
lished).

2 Vouk, M. F. (1978) Aspects of the geolory of
the Stuart Creek area, north of Lake Torrens,
South Australia, B.Sc. (Ions.) thesis, Univer
silty of Adelaide (unpublished).

During geological mapping of the BILLA
KALINA 1:250 000 map sheet and subsequent
investigations elsewhere along the margin of
the Great Australian Basin, a further 32 fos-
siliferous boulders were discovered, containing
many species not previously recorded in S.A.
Their occurrence and distribution permit 4
few assessment of their probable ongin and
modes of transport, ‘The earlier concept of
Campbell et al. (1977) is substantiated,

Geological setting

Stratigraphic units in the southwestern Great
Australian Basin include the Algebuckina
Sandstone, Cadna-owie Formation and Mount
Anna Sandstone Member, Bulldog Shale and
overlying younger Mesozoic sediments (Fig |).
The stratigraphic nomenclature adopted here
ix that of Wopfner ef al. (1970) based on
the Oodnadatla-William Creek area (for the
Marree area see Forbes 1966),

The following gealogical summary is com-
piled feam Wopfner & Heath (1963), Lud-
brook (1966, 1978), Wopfner et al. (1970),
Morgan (1977), Carr et al, (1978)4,) Pitt
(1978). Vouk £1978)" and from observations
during geological mapping of the BILLA
KALINA [:250.000 map sheet.

Vhe Upper Jurassic Algebuckinu Sandstone
consists of fine to medium-grained sandstones
and kaolinitic, conglumeratic sandstone, Clasts
within the conglomeratic sandstones arte chiefly

58 R. B. FLINT, G. J. AMBROSE & K.S. W. CAMPBELL

SIMPSON DESERT

=— —BOGDMADATIA —

a)

TSILLAKALINA 1250. 000
dese” MAP. SH

rounded to well-rounded white quartz pebbles.
However, weathered acid porphyry and quart-
zite pebbles and cobbles are also common, The
unit Wes deposited m a low gradient. fluviatile
environment,

Transgceession tw the Neocomian led to ibe
disconformably overlying Cadna-owie Forma-
tion, consisting of marginal marine very fine to
Medium-peruined. micaceous and oceasionally
conglomeratic sandstones. Clasts within the
sandstunes ure chielly pebble, cobble and
boulder-sized quartzites up to 1 x J x 0.5 m
Later in the Neocomian, partial regression led
to the deposition of the coarser Mount Anta
Sandstone whieh eoasists of medium to conrse-
gmincd. feldspattue ond conglomeratic sand-
stones and micneros sandstones. Clasts of
perphyritic acid volcanics charactertse the con-

\ bARE
— fi wyre )

i ——— =,

lye id
: | camozor

ALEJEN TOAONIAN

Beck: to

CRETACEOUS
SS —s—

Lh}
3 “ a ue ma ' = 3
Ovaly, yO Colm ae am sed Kandy a
Fa
DAWA MWHE FWAA [loty = =
Hye VD (Yaya to ae 3
piualamcucihs Pru «unr eooe =
PLA RHL De Stl’ jlo pip 5
1 ere ee OC en ee Tt Et a =
ao
2
=
= |2 |
Heed tats crt THR Ede idee taueerpt it = 2
tied Abel Ghodt — = =
= &
CHR) Soatemment cceko at or }82 2
Gorl ehe ono 4S
HORA poy Mipwatiivee cian vied meeetere e
1s , q 5
.\ TIBAR DESERT REET RES ida |
STRZELECK! DESERT |

glomeratic sandstones, though white quartz
and quurizite clasts are also common, The
clasts are sub-rounded to well-rounded and in
ihe size range 0.02-0.2 m. Concave and festoon
cross bedding are ubiquitous; forescts are up te
2m high and bedding within the foresets is
graded.

The second Cretaceous marine transgres-
sion, in Aptiaa time, was of much greater ex-
tent and resulted in marine shale deposition
(Bulldog Shale) over a large area of the Great
Australian Basin, Basal lithologies of the Bull-
dog Shale range from bouldery to conglomera-
tic sand to grey shales, cone-in-cone limestones
and sandy limestone, Fossil tree trunks are
common, Clasts within the conglomerate sedi-
ment are predominantly quartzites with minor
acid porphyries and banded chalcedony, and

FOSSILIFEROUS LOWER DEVONIAN BOULDERS 59

occur in either bioturbated grey shales or thin _ silts, and fossiliferous limestones. A subse-
coarse-grained sand lenses (Fig. 2). The sand quent regression and a further two transgres-
lenses vary from only a few centimetres thick sive-regressive cycles occurred in the Albian
to massive lenses up to a metre thick With to Turonian.

boulders scattered in the sandy matrix (Fig. The fossiliferous Devonian quartzite boulder
3). These sediments interfinger with and are from the locality southeast of Oodnadatta is
the only specimen weathered out from mar-

CcuX

oe “
yer

Shale, 60 km north of “Billa Kalina”; 3, Rounded quartzite cobbles near top of coarse-grained sand
lens, and overlying bioturbated marine shales (Bulldog Shale) from 20 km east of Billa Kalina
locality 1; 4, Well-rounded quartzite boulder lag near Billa Kalina locality 2, Many boulders are
fractured due to Holocene weathering; 5, Surface gibber lag east of Coober Pedy: cobbles and
boulders derived from basal sediments of Bulldog Shale. Clasts are dominantly quartzites; less than
1% contain Devonian fossils.

60 R. RB. FLINT, G. J. AMBROSE & K. 8. W. CAMPRELL

ginal marine sediments of the Cadna-owie For-
mation, All other fossiliferous boulders (in-
cluding the original two specimens discovered
at Dalhousie Springs and Andamooka, and
previously thought to be derived Irom the
Cadna-owie Formation) have weathered out
from basal conglomeratico sedimenis of the
Bulldog Shale,

Description of the boulders

Basal conglomeratic sediments of the Bull-
dog Shale crop out poorly but erosion has re-
sulted in Alimerous clasts from the conglo-
merates, ranging in size from pebbles to
boulders, forming a lag on the present day land
surface (Figs -4, 5), Physically-resistant clast
types dominate, mostly quartzites (feldspathic
and/or tithe), with minor porphyritic acid
volcanics and whitish-grey banded chalcedony,
Granite, gneiss, quartz and shale clasts are
rare, hut may be locally more common tear
Proterozoic oliterops.

A high proportion of the quartzite boulders
have abundant clay pellet impressions, » fea-
ture typical of the Upper Proterozoic Arcoona
Quartzite on the Stuart Shelf, The porphyritic
acid volcanics are similar to the Middle Pro-
terozoic Gawler Range Volcanics on the Gaw-
ler Craton (Wopfner et al 1970), while
banded chaleediny clasts ure similar to cherts
and siliceous Goneretions in the Cambrian An-
damooka Limestone,

Less thatt 1% of all boulders contain De-
vonian fossils, The fossiliferous boulders are
siliceous, feldspathic and lithic quartzites. They
consist of quartz-rich, medium-grained sand
(0.2-0.3 mm) and minor (<10% ) potash feld-
spar grams cemented by secondary quartz
overgrowths. The lithic quartzites contain
small Fragments of sericitic schists and ucid
porphyries (Whitehead 1978)". Tt is not pos-
sible (o distinguish |ithologically between fos-
siliferous Devonian quartzites and other
quartzite clasts.

Faunas of the boulders

In the two fossiliferous Devonian boulders
previously recorded in South Australia (Camp>
bell et al. 1977), the specimen from Dalhouste
Springs contained the brachiopod Huwellelly
jaquer’ (Dun) und bivalve Aetinepteria sp.
these were also present in the specimen col-
jected from Andamooka, In the latter sample

" Whitelreasl, S. (1978) Description of quartzite
boulders, Amde!l Rept. No. GS 415/79 Huinpub-
lished).

tentaculitids and brachiopod /yorrhiy sp. were
also present, The fossils found in the boulders
during 1977-78 are documented helow, and
intlude many species not recorded previously
in the boulders. All specimen numbers refer
to the fosstl collection of the Geological Sur-
vey of South Australia,

Billa Kalina locality | Cat.
136°08 00"B) — Specimen
29-34. 39,

Fish plate: Witlageonaspis sp. (Pig, 6)

Brachiopods: Howellella jaqueti (Dun)
Brachiopoda indet.

Bivalves; Lepiodesma inflatem (Dun)
Bivalvia indet.

Tentaculitid; Terdeulites sp. (Fig. 7)

Billa Kalina locality 2 (lat 29°28°00"S, long,
134°06'S0"E} — Specimen numbers 6139 RS
35-36.

Brachiopod: Howellella jayuet? (Dut)

Billa Kalina locality 3 (lat. 29"°58'20"S, long.
134°12'S0"E) — Specimen numbers 6138 RS
7a-T7

Brachiopoda inctet.
Bivalvia indet,
Fish plates and spines

Billa Kalina tloeality 4 Uat. 29°57'30"S, long,
136°18'35"E) — Specimen numbers 6/38 RS
‘78-85,

Brochiopods: Mowellella jaqueri (Dun)
Brachiopoda indet.

Bivalves: Sanguinolites sp,
Bivalvia indet.

Gastropods: Straparollus éullent (Dun) (Fig. 8)
Holapea sp..
Murchisoniidae indet,

Echinodermata indet,

Fish Plates and spines

Billa Kalina locality 5 (lat,
136°2)'0S°E) — Specimen
37
Tentaculitid: Tentaculites sp.
Crinoid ossicles
Bryozoa indet.

Billa Kalina lacality 6 (at 29°12'15"S, long,
136°09'05"E) — Specimen number 6139 RS
38.

Bivalve: Bivalve indet.
Crinoid ossicles

Billa) Kalina lecality 7 Cat. 29°02°05°S, long.
135°12'20"E) — Specimen number 5939 RS 92,
Brachiopod: Brachiopada indet.

Crinoid ossicles

Billa Kalina locality & (lat. 29°55°00"S, long.
135°49'30"E) — Specimen numbers 6038 RS
12-13,

Brachiopods: Stropheodontid (probably Mese-
douvillina or Melearnites) (Fig. 94
Brachiopoda indet,

29°28'10"S, long.
numbers 6139 RS

29") 100"S, long,
fumber 6139 RS

FOSSILIFEROUS LOWER DEVONIAN BOULDERS 61

Tentaculitid: Tentaculites sp.
Crinoid fragments
Bryozoa indet.
Moolawatana (lat. 20°52'12"S, long. 139°38’00" EB)
— Specimen numbers 6838 RS 129-132.

Brachiopods: Howellella jaqueti (Dun) (Fig.
10)
Brachiopoda indet.

Bivalves: Leptodesma inflatum (Dun)
Bivalvia indet.

Tentaculitid: Tentaculites sp.

f- .
a ees
d=? . 38
poe or

au

“Moolawatana” locality; 11, External mould of Sanguinolites sp. from Stuarts Creek locality, speci-

men 6337 RS 21.

62 R. B. FLINT, G. J. AMBROSE & K. S&S W. CAMPBELL

hol HRAS

PRAHD VEY

Amapedia
BASIN. fC

| DFFICER

Lrerno
Major pe

anpertet hip

Ch Ae dros els |

VICTHHIA
erlicmas
T Midi foe |) Adee Geen pnd line
Zoawls Lee = BS White

‘i
7) VASMAMN) A
D5rwn hed G Wieaeen

Loree iw Wee AlN

Fig, 12. Locality plin showing disitibution of
husins incorporating Cretaceous and Permian
sedimen(s in central and eastern Australia.

PTRMLAN oie

Vee Lopraet a lel

Codnadaiia (al. 27°55°30"S, Jong. 135"46'40"B)
— Specimen Numbers 6042 RS 92-93,
Bivalve: Praecienodanta sp,

Tertaculitid, Tevtiuentites sp.

Sivaris Creek (lat, = 3O°US45"S, long,
see se sl — Specimen numbers 6337 RS
21.34.

Brachiopods: Sphivriniyachin sp.
Howellella jaquetti (Dun)
Bivalves: Suaeuinaliters sp. (Pig. it)

Tentaculitid; Venracelies sp.
Crinoid ossicles.

Provenance of the Fossiliferous boulders

Nether the invertebrate not the vertebrate
faunas preserved in the boulders have been re-
cordet trom in situ Devonian sediments in
basins i) S.A. or the adjacent Amadeus or
Georgina Basins, The only possible Devonian
vertebrates known from §.A, are the fish seales
from a mudstone at 817-823 m io Munyarai
No. | within the Officer Basin (Pig. 12), but
these were nol positively identifiable (Gilbert-

Tomlinson 1969)4. Devonian placoderny re-
mains from the Amadeus and Georgina Basins
are bothriolepids (Young 1974) which are nat
similar to the specimens from the boulders, No
Devonian invertebrates are kiown from the
above basins, However, as has been indicated
previously, both the lithologies and inverte-
brate faunas are very similar to those from the
Amphitheatre Group near Cobar in N.S.W.,
described by Landrum (1975), and to those
from the boulders in Cretaccous sediments at
White Clilfs, N.S.W., described by Dun
(1898),

On the other hand, marine Lower Devonian
sandstones with comparable invertebrate
faunas to those at Cobar are also known (rom
the Mt Ida Formation of central Victoria and
the Elden Group of western Tasmania. These
have to be considered as alternative source
ureas, but there are good reasons for rejecting
them, The absence of Notecenchidium trom
the boulders in §,A. is taken as evidence
ugainst either a Victorian or a Tasmaniait
source because this genus js relatively common
in a hard quartzite in both these areas, The
durability of this material is attested by the
fact that Netocenchidium is among the more
common fossils in the Lower Devonian
boulders from Permian diamictites in north-
eastern Victoria. A secand important feature is
Ihe abundance of Howellella jagueti in
boulders from S.A, Although Howellella oc
curs in a variety of forms at Heathcote, ‘Talent
(1965) records that they are poorly pre-
served. ‘The genus is also poorly represented
m the Eldon Group. A. jaqueli is one of the
Most common species in boulders from S.A.,
as it is in several horizons in the Amphitheatre
(Group,

Although there are now many more boulders
with a wider range of species than was known
previously, iL sul] is possible to match the en-
tire invertebrate fauna with that from the
Amphitheatre Group, In the absenee of a
complete account of the Eldon Group fauna,
this evidence of itself can be no more than
suggestive; but taken in conjunction with the
data on Notoconchidiun and Mowellella given
above it is more persuasive of a Cobar source.

! Gilbert-Tomlinson, J. (1969) Fossils from Mun
yarai No. | Well, Officer Basin, South Australia.
‘ “Continental Of: Company of Australia Ltd.
Munyarai No. t, South Austratia,” Well com-
pletion report, S, Aust, Dept Mines & Energy
envy. 979 (unpublished),

FPOSSILIFERGUS LOWER DEVONIAN BOULDERS 63

Further weight is lent to this view by the
discovery of fossil fish fragments in boulders
at three of the Billa Kalina localities, So far
as We are aware no Devonian fish beds ure
known from Tasmania, though they are well
eaposed in central and eastern Victoria and
over much of central N.S,W., where they are
mainly of Late Devonian age, The most im-
portant discovery is the fragnreni referred to
Wuttagounasply Ritehie (1973) from the
Mulga Downs Formation of probable Middle
Devonian age in the Mt Grenfell area wesi of
Cobar, und Mt Jack north of Wileatnia, This
specimen is only an impression of a fragment
of an undetermined bone, but its ornamenta-
tion is distinctive, Its identifieadon has been
gantirmed by Ritchie.

We therefore conclude from the available
evidence that the source for the fossiliferous
boulders is in the Cobar région, the boulders
having been tfafisported at least 1000 km in
a weslerly to vorthwesterly direction.

Transport oF the boulders

A palacoenvironmental interpretation of
JurassieCretaceous sediments in the southe
western Great Australian Basin by Waoplner
et al (1970) indicates that transport of
boulders in this direction and for this distunce
during the Mesozoic Was improbable. How-
ever, Permian ice may have transported the
fossiliferous boulders from the Cobar area to
northern S.A. and the unconsolidated Per-
nian diamictites could then have heen te-
worked iuta Mesozoic sediments (Campbell er
al, 1977), Thus (wo phases of transport would
be involved,

fu northern S.A... Pormian diamictites gre
preserved in Palacozoie basing under the Great
Australian Basin fe.g. Arckaringa, Cooper and
Pedirka Basins) and in stall grubens within
the Gawler Block, These distributions suggest
that such deposits were once widespread but
thit they have been Jargely removed from up-
lifted areas.

Crowell & Frakes (1975) using the distribu-
tion of glacial Hl and Muvial sediments and
palaeveurrent analysis, postulated a larac
Permian continental lee cap over vorthwestern
NuS.W,. with glacial debris being shed east
wards and possibly westwards into the basins
of northeastern S.A, This interpretation differs
from that of Wopther (1990) who concluded
that the composition of erratics in Permian
diamictites of the AfcKartnga Basin indicated

Inval glaciation rather than a continental ice
shect, He suggested that Permian glaciev's
originated on uplaulted highland areas; glacial
Uecbris was dumped along basin margins and
then transported by mudflows alid turbidity
currents inta distal parts of the basins,

Though we prefer the views proposed by
Crowell & Frakes, if must be stressed that to
date no fnssiliferous Devonian boulders have
been discovered in Permiun cdiumietites in S.A.
ice-movement directions during the Permiat
are not known for northern §.A,, and there are
conflicting views on the Permian palacoen-
Vironment and likelihood of long-distance
Lransporl., Nevertheless, ice transport seems to
be the only feasible means for tfansporting
boulders from the Cobar area to northern
S.A.. and the Permian is the only period in
the required iuterval for which glaciation of
an Appropriate magnitude has been demo-
strated,

Final entplacement of the boulders

The processes by which the Cretaceous
boulder heds were formed have been debated
for nearly 100 yeurs, This paper is not in-
tended tw provide a detailed discussion of the
problem, but it does add another feature that
requires explanation — viz. the distant pre-
Venanee of same of the houlders. It has been
thought appropriate that 4 summary of the
issties should be presented,

Beatures requiring explanation are

(a) the scatter of boulders throuyly a sandy
or shaly matrix which is bioturbated in places;

(b) the rounded form olf most of the
boulders, though an occasional fuectted or
striated boulder has been noted (Jack 1915;
Woolnough & David 1926);

{c) the predominunee of quartzites among
ihe boulders, with avid porphyries and chal-
cedony forming the majority of the remainder;

(4) the occurrence of fossiliferous boulders
in Cretaceuus rocks from While Cliffs to Bal-
housic Springs

The shape and eooposition of the boulders
indicates (he operation of processes that have
removed all but the Most divable materials,
and thal these processes were al least in part
physical. Tumbling experiments by Abbott &
Peterson (1978) showed ehert, quartzite and
rhyolite to be the most durable rock types, fol-
lowed by metabreccia, obsidian, mietasand-
stone, gneiss, ‘gaiites’, melabasall, marble and
schist, The first three rocks are also chemically
resistant.

64 kK. B. FLINT, GJ

One possihle source for many of the clasts
is in Proterozoie/Cambrian rocks such as
those of the Gawler Craton and Broken Mill
Block which could provide abundant quart.
zite, acid porphyry (Gawler Ranye Voleanics)
and chalcedony (Andamooka Limestone). A
second source would be the Permian dhamice
tite, mentioned pbove, which is known ta con-
tin clasts of limestone, schist, gneiss, eranite,
acid porphyry, quartzite, quartz, banded iron
formation, chert and shale, The original source
of many of these clasts must have been the
Profterozoic/Cambrian rocks indicated above.
Both the above sourges would have been sub-
jected ty prolonged Weathering between the
Permian and the Early Cretaceous, and trans-
port to the Cretaceous sea with subsequent
shoreline deposition would have resulted in
the removal of the less durable clasts,

Brown (1905), Jack (1915), Woolnough &
David (1926) and Vnuk® considered that ice
rafting was responsible for the final transport
of the boulders and that they were muinly
dropstones. [n our yiew this mechanism ts nae
acceptable. If the proposed ice was culved off
from glaciers there would be oo explanation
for the dominance of resistant clasts, their
rounded shapes, or their abundanee ever su
large w geographical distribution, If sea ie
picking up clasts from 4 boulder-strewn
shore were proposed, it would be possible ta
explain the clast types and shapes, but the prob-
blems of volume and distribution would re-
main. In addition there is no independent evi-
dence of glacial conditions in the Early Cre-
taceous, though the area in question would
have been within 30° of the pole,

Woolnough & David (1926) also con-
sidered, but rejected, tree rafting as a possihle
transport mechanism for the boulders. Much
later Wopfner et af, (1970) reinstated the pro-
posal because of the abundance of fossil
woul in Early Cretaceous sedirients, However

AMBROSE & kK, 8, W. CAMPBELL

because of the abundance and concentration
of boulders within particular horizons, tree rafl-
ing was not accepted as the sole transport
mechanism, Since they considered the boulder
beds to be restricted to margins of basement
highs, they also proposed that the boulders
originales on shorelines and migrated down-
slope by slow sediment creep, [t is this latter
Suggestion that seems to us to provide a clue
io a possible solutiwn — namely that they are
reworked debris-flow deposits, Bouldery de-
bris-flow deposits typically consist of a massive
fine-grained matrix with randomly dispersed
boulders [Fisher 1971; Middleton & Hampton
(973; Carter 1975; Hampton 1975) but the
beds under discussion are not of this type,
Some other processes must have been opera-
tive in addition, We propose, therefore, that
boulders. cobbles and sand were transported
basinwards trom a boulder-strewn shore ling
in a clay-rich, watery matrix over low angle
slopes. The debris flows were episodic events,
permitting time for some reworking of the
debris-flow sediments, and subsequent shale
sedimentation and bioturbation. Winnowing of
muds and fine sands from the debris-flow seci>
ments by currents, and possibly by waves. has
resulted in some of the boulders and cobbles
being concentrated in thin, course-grained sand
lenses, Complete winnowing of the fines and
further shale deposition has resulied in same
boulders being located within bioturbated
shales,

Acknowledgments

Apprecianion is extended ta G, Johansen
(Newmont Pry Led), M. FP Vnuk (Adelaide
Lyiversity) and M, C. Benhow. R, A, Callen
and S. Ro Hewles (Geological Survey of South
Australia) who collected some of the fossili-
ferwus boulders and kindly made then avail-
able for cXamination,

References

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of abrasion durability on conglamerate clust
populations: Examples from Cretaccous anil
Eovene conglomerates of the San Diego urea,
Califomia, J. sedim. Perret. 48, 31-42

Brown, H. ¥. £, (1905) Report on geological
explorations in the west and northwest of South
Australia, Parl. Pap. &. Aast, Th.

Camprers, KS. Wi, Roceas, P. A. & Brapow,
M. C. (10977) Bossiliferaus Lower Devonian
boanlders from the Cretaveous of South Austra
lia, Quart, geal, Notes, geal. Suey. S. Aast, 62,
9-13.

Carrer, ROM. (1975) A discussion ane chissifi-
cliten of subaqoeous muss-fransport with parti-
“ular applicution to erain-flow. slirrytlow, and
Noxotu bidltes, Bart Sel. Rev. Vb, 145-177,

Caowih. 1, ©, & Fraxes, L. A, (1975) The late
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(Eds, “Gondwana Geology.” pp, 313-331
(ANU. Press: Cunberia).

DEN, W.S. (1898) Notes on the fauna of Devon-
jan boulders occurring at the White Cliffs opal:
fields Rec. geol, Surv NSW, 5, 160-174,

FOSSILIFEROUS LOWER DEVONIAN BOULDERS 65

FisHer, R. V, (1971) Features of coarse-grained,
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sedim, Petrol. 41, 916-927.

Forses, B. G. (1966) The geology of the
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Hampton, M. A. (1975) Competence of fine-
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R44

Jack, R. L. (1915) The geology and prospects of
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and the geology of the western portion of the
Great Australian Artesian Basin. Bull. geol.
Surv. S. Aust, 5.

Lupsrook, N. H. (1966) Cretaceous biostrati-
graphy of the Great Artesian Basin in South
Australia. Ibid. 40.

(1978) Australia. 2x “The Phanerozoic geo-
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249. Elsevier.

MippLeTon, G. V. & Hampton, M. A. (1973)
Sediment gravity flows: Mechanics of flow and
deposition, /n G. V. Middleton & A. H. Bouma
(Chairmen) “Turbidites and deep water sedi-
mentation.” Short course lecture notes; pp. 1-38
(Soc. econ, Paleont. Miner.: Los Angeles).

Moroan, R. (1977) New dinoflagellate zones and
a depositional model for the Great Australian
Basin. Quart. geol. Notes, geol. Surv. N.S.W.
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Parkin, L. W. (1956) Notes on the younger gla-
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An Unusual Arthrodire from the Devonian of
Western New South Wales, Australia. Paldeon-
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Tuomson, B. P, (1980) “Geological Map of
South Australia.” 1:1 000 000 Scale. Geol. Surv.
S. Aust.

WooLnouanH, W. G. & Davin, T. W. E. (1926)
Cretaceous glaciation in South Australia. Quart.
J. geol. Soc. Lond. 82, 332-351,

Woprner, H. (1970) Permian palaeogeography
and depositional environment. of the Arckaringa
Basin, South Australia. In “Second Gondwana
Symposium,” pp. 273-291. (Natal Witness
Press: Pietermaritzburg).

, Freytac, L. B. & HEATH, G. R. (1970) Basal

Jurassic-Cretaceous rocks of western Great

Artesian Basin, South Australia: Stratigraphy

and Environment. Bull. Am. Ass. Petrol. Geol.

54, 353-416.

& Heatu, G. R. (1963) New observations on

the basal Creta-Jurassic Sandstone in the Mount

ue region, South Australia, Aust. J. Sci. 26,

57-59.

“YounG, G. C. (1974) Stratigraphic occurrence of

some placoderm fishes in the Middle and Late
Devonian Newsl. Stratigr. 3, 243-261,

PROGAMOTAENIA NYBELIN (CESTODA: ANOPLOCEPHALIDAE): NEW
SPECIES, REDESCRIPTIONS AND NEW HOST RECORDS

BY I, BEVERIDGE

Summary

The following species are described: Progamotaenia spearei sp. nov., from Thylogale stigmatica,
distinguished by its small size, in having paired uteri, a fringed veleum and testes in two groups,
and Progamotaenia johnsoni sp. nov. from Lagorchestis conspicillatus, which has an external
seminal vesicle covered with glandular cells and testes distributed in two elongate groups. P.
bancrofti (Johnston) and P. diaphana (Zschokke) are redescribed, and Lasiorhinus latifrons is
considered to be the usual host of the latter species. P. zschokkei (Janicki) is reported for the first
time from Macropus agilus, Onychogalea fraenata and O. unguifera; additional records of this
species from Petrogale penicillata, Lagorchestes conspicillatus and Thylogale stigmatica are given.

PROGAMOTAENIA NYBELIN (CESTODA:
NEW SPECIES, REDESCRIPTIONS AND

ANOPLOCEPHALIDAE):
NEW HOST RECORDS

by L Beveripce*

Summary

Bevermnce, L (1980) Progamataenia Mybelin (Cestada: Anoplocephalidac): new species,
tedescriptions and new host records. Trans. R, Soc, S. Aus, 104(4), 67-79, 30 May, 1980.
The following species are described: Progamotaenia spearei sp. nov., from Thylogale
stigmatica, distinguished by its small size, in having palted uferil, a fringed velum and testes
in rwo groups, and Progamotaenia johaseni sp. nov. from Lavorchestis conspicillares, which
has an external seminal vesicle covered with glandular cells and testes distributed in two
elongate groups. P. barcrojti (Johneton) and P. diaphana (Zschokke) are tedescribed, and
Lasiorhinus latifrous is considered to be the asual host of the batter species. P_ zschokkei
(Janicki) is reported {or the first time from Macropus agilis, Onyehogalea fraenata and
O, wiguifera; additional records of this species from Petrogale peniciliaia, Lagorchestes con-

spicillatus and Thylogale stigmatica are given.

Introduction

Although the anoplocephalid ecstodes of
Australian marsupials have been the subject of
a recent review (Beveridge (976), continued
collecting has resulted in the discovery of ad-
ditional new species of Progamotaenia (Bever-
idge 1978, Beveridge & Thompson 1979), To
date most collections have been from miar-
supial species inhabiting the south-castern part
of the continent. Recent collecting from mac-
Topodids in north Queensland bas led te the
discovery of two new species of Pragamo-
taenica.

Progamotaenta banerofti and P. diaphana,
were recently redeseribed by Beveridge (1976)
on the basis of very limited or poorly preserved
material only. Both are well represented in
recent Collections, and the opportunity is taken
here to describe them fully,

Materials and methods
Cestodes collected from the small intestines
of macropodids were washed in tap water, re-
laxed in water for several hours and fixed in
10% formalit or Serra's fluid. They were
staimed with Celestine blue, cleared in methyl
salicylate or clove oil and mounted in balsam.
Wand cut transverse sections and serial histo-
logical sections cut in transverse and longi-
tudinal planes were prepared, Drawings were
made with the aid of a cameta lucida, Mea-
surements. are given in mm as the range fol-
lowed by the mean of five measurements
(where available) in parenthesis,
Type specimens have been deposited in the
South Australian Museum (SAM).

* [Institute of Medical & Veterinary Science,
Frome Rd, Adelaide, S. Aust. 5000,

Progamolaenia spearei sp, nov.
FIGS i-8

Types: holotype (slide) from small intestine
of Thylogale stigineitea Gould 1860, Tolga
Queensland, 16 x, 1978, collected by I,
Beveridge, in SAM V1920; 4 pavatype slides,
same data VIG2I-V1924: 1 paratype, spirit
material and serial sections VL925, V1926.

Description; Length 26-30 (28); width 5-7
(6): seolex diameter 131-152 (1.44): sucker
diameter 0.56-0.74 (0.67) x 0.47-0.58 (0.55);
neck 0-0,21 (0,09); no. proglottides 71-85
(78); mature proglottides 3.84.0 (3.9) x
0.21-0.28 (0.23); gravid proglottides 4.1-5.8
{5.4) x 0,56—0.74 (0.65): cirrus sac in mature
proglottides 0.60-0.92 (0,81) & 0.084112
(0.10); cirrus sac in gravid proglottides 0,93-
0.98 (0.94) x 0.11-0.14 (0.12); no, testes per
proglottis 30-40 (34); testis diameter 0.06—
0,09 (0.08); ovary 0.50-0,456 (0.53) x 0.18-
O21 (0.20); vitellarium O.11-0,18 (0.15) x
0.08-0.12 (0.10); dorsal osmoregula-
tory canal 0,02-0.03 (0.021; ventral osmoregu-
latory canal 0,03-0,09 (0.07); cgz 0.06-0.08
(0.07); pyriform apparatus 0,03-0,04 (0.04);
oncosphere 0,02,

Short broad tapeworms with relatively few
proglottides, Scolex squat, almost globose,
prominently four-lobed, each lobe with muscu-
far, cup shaped sucker. Neck absent or, if pre-
sent, very short, segmentation beginning very
close to suckers, Proglottides extended trans-
versely, craspedote with broad, fringed velum
consisting of about 25-35 tongue shaped pro-
Jeclions which are frequently felded and over-
hand 1/2=2/3 of the adjacent proglottis-
Mature proglottides with approximate length:

68 I, BEVERIDGE

Figs 1-8. Progamotaenia spearei sp. nov. 1, scolex; 2. transverse histological section through cortex
showing musculature; 3. egg; 4. mature proglottis; 5. gravid proglottis; 6, cirrus sac; 7, lateral re-
gion of mature proglottis prior to vaginal atrophy and uterine filling; 8, lateral region of mature pro-
glottis at commencement of uterine filling, showing atrophy of vagina. Scale lines 0.1 mm.

SPECIES OF PROGAMOTAENIA (CESTODA: ANGPLOCEPHALIDAB) 69

width ratio of 1:14 to 1:20. Gravid proglot-
tides with ratio 1;6 to 1:10, Cortex thick,
longitudinal muscles developed, numerous, rot
alranged in regular bundles, denser towsirds
cortico-medullary junction, Tratsvetse muscles
well developed, forming thick band runing
along cortico-medullary junction, Dorso-ven-
tral muscles prominent, crossing cortex and
medulla at irregular intervals. Longitudinal
osmoregulatory canals paired, Ventral canal
wider than dorsal canal, situuted medial to it.
Two transverse canals connect left and right
osmoregulatory canals at posterior fnarsin of
each proglottis, Canal connecting ventral ves-
sels of moderate size, readily observed: canal
connecting dorsal canals extremely fie, seen
only with difficulty, Fine accessory canals as-
sociated with ventral canal in some proglot-
tides; accessory canals branch and anastomose
imegularly, Junction of osmoregulatory canals
in seolex not seen. Genital ducts cross osmo-
regulatory canals dorsally, Genital atrium
prominent, very long; walls lined with
thickened tegument, opening in middle of
lateral proglottis margin. Cirrus sac clongste,
musculature of walls strangly developed, cross-
ing beyond osmoregulatery canals. reaching
almost to ovary. Cirrus narrow, distal ‘third
covered wilh numerous prominent spines: mid.
third unarmed, greatly coded when retracted.
vncoiled when cirrus is everted, Internal se-
minal Vesicle prominent, elongate, External
seminal vesicle absent, Vas deferens coils an-
teriorly and medially from cirrus sac. Wasa
efferentia nol seen, Testes distributed in two
compact clusters anterior Un cirrus sac. extend
ing medially from osmoregulatory canals to
poral side of ovary; never extending medially
hevond ovary to form single band. Vayina
tube-Jike, opening to genital atrium posterior
lo cirrus sac. Vagina leads medially lo dimmu-
tive, ovaid, seminal reeepticle situated pos-
terior to median pole of cirrus soe, lateral to
ovary, Ovary fan-shaped, composed of
numerous clavate lobules, on ventral aspect of
mednlla, Vitellarium ovotd, situated posterior
and dorsal to ovary Mehtis’ gland spherical,
medial to vitellarium, Uterus transverse, tube-
like, paired in cach proglottis, anterior to
ovary, Fully developed wferus sacciform with
prominent anterior diverieulay posterlor diver-
ticula very small, Uteri fill space hetween
osmoregulatory canals but do not extend be-
yond canals except in Jast few gravid: progtot-
tides: uteri croxs canals dorsally but da nat
reach posterolateral corners of proglottis, Egg

spherical, thick-shelled. Pyriforny apparatus
conical, terminating in reflexed filaments.
Genital primordia appear In &-10th progh-
tls; genital organs are developed by 15-18th
proglottis: sperm present in internal seminal
vesicle in |5—20th proglottis; (line of seminal
receptacle and alrophy of vagina oceur in 146-
24th provlottis; uterus begins to All in 22—-26th
proglottis, eggs fully formed in 69-73nl pro-
glottis,

Vestigial supernumerary vitellaria, ovaries
or uterine fragments present in proglollides of
some strobilac, of variahle size, arranged be-
Iween Iwo normal sets of genital organs.

Discussion: Pragamolarnia spearei most
closely sesernbles FP, prateregyna (Fubrmann
1932) [tm external features, being a small
worn, with few proglotlides and with a fringed
velum. It differs in the shape of Fringes of the
vellim, the occurrence of testes in two distinet
groups rather than in a continuous band and
in the oecurrenee of vaginal alrophy following
insemination. P eynandroliwariy Beveridge &
Thampson 1974 is another small species, bul
the velum is nol fringed. Po speare? resembles
Po fagorchestis (Lewis 1914) and P. thylogale
Beveridge & Thompson 1979 in having a
fringed velum, paired uteri and testes occurring
in two lateral groups, but differs fram both
species in ils small size, small number of pro-
glattides, the rupidity af development of the
genital organs in the strobila and the small
number of testes.

This species is named after Or R. Speare,
James Cook University of North Queensland,
in appreciation of help given in collecting
material,

Progamotaenia johnseni sp. nov.
HIGS 9-15

Jypess holotype (without seolee) from small
infesting Of Lagerchevtes conspicihitie Gould,
(442, Mingela, Qneensland, 1 4. 1979, col
lected by |. Beveridge. 4 slides, spirit material
aml serial section in SAM V1918; paratype
slide, sane data, VIGTY,

Matevial examined: from Lagerchestes can-
yiicillatas: types; 4 specimens Cuou pray),
Trkemman Station, Olu, Tis d977, BM. tol.
son.

Deseriptions Length (78 Cholotyped; walth 13

(holotype): scolex diameter O74-1.01 (MN RA47
sucker diameter 0,30-40.42 (0.92) ~ O.28-0.36

70 I. BEVERIDGE

eseaien

rrr pitta
qs,2 2
PO. alin

SPECIES OF PROGAMOTAENIA (CESTODA: ANOPLOCEPHALIDAE) TL

(0,31); neck 016-0,36 (0.26); no. proglot-
tides 272 (holotype); mature proglotlides 5,4—
7.0 (6.2) x 0,52-0.56 (0,54); gravid proglot-
tides 9.4-9.8 (9.6) x 0.92-0,99 (0.96): cirrus
sac in mature proglottides 046-1,31 (0.90) %
0.16-0,29 (0.25): in gravid proglotiides 118—
1,18 (£149) x 035440,36 (0.95); no. testes per
proglottis 100-190 (178): testis chameter 0,05—
0,08 (0.06), ovary 0.624180 (0-72) x O44—
O98 (0.36); vitellanitim 0.30040 (O37) &
0.17-0.23 (M18): Meblis’ gland 0,10-1,14
(0.12): dorsul osmoregulatory canal 0.02—-11.04
((\.03); ventral osmoregulatory canal 0,08-
0.29 {O.18i2 egg 0.064-0,072 (0,068); pyr-
form apparatus O0.024-028 (0.025); anca-
sphere O.00960,U16 (0.015),

Laree, broad, ribhon-like worms. Scolex
broad, four-lobed, with cup-shaped sucker al
apex of each lobe, Suckers with anterior inar-
gins cleft, Neck short, Proglotlides greatly ex-
tended transversely, eraspedote, with broad,
fleshy, folded velum covering much of adjacent
proglottis. Mature proglottides with approxi-
mite length:width ratio of 1:10 to ts13. Gra-
vid proglottides with ratio of 1:9 to Is1)-
Longitudinal musculature powerfully — de-
veloped, composed of patlisades of muscle
bundles. Bundles circular or oval in seccian:
ouler biindles smaller, with fewer fibres. Outer
lonoitudinal musculature consists of rng of
individual fibres, inymediately external to
muscle bundles, Transverse muscles well de-
veloped, furminy broad band immediately m-
ternal to lonwitudinal muscle bundles. Few,
scallered transverse muscle fibres present in
ouler cortex. Towards lateral margins ol pro-
slottides, trinsverse muscles from cortex and
cortica-medullary junction fuse into thick band
of muscle encireling cirrus sac and extending
tO genital atrium, Dorso-ventral muscle fibres
well developed, promment, crossing cortex and
medulla at regular Iotervals. Longitadinal nos-
morevulatory canals pulred, Ventral canal
wider than dorsal canal, situated medial to it,
Transverse canal connects left and right ven-
tral canals at posterior margin of cach praglor-
fis. Scolex osmoregulatory canals not seen,
Genital duets cross osmoregulatory canals dor-
sally, Genital atrium shallow, with corrugated
walls, Opening in middle of lateral proglottis

margin. Cirrus sac powerlully developed, wilh
thick muscular walls, extending (o medial mtar-
gin of, of beyond longitudirial osmoregulatery
canals into medulla. Cirrus wide at distal ex-
tremity, heavily armed with spines; mid-region
of narrower diameter, distal part armed, praxi-
mul part unarmed. Internal semimal vesicle
ovoid, with thick, muscular walls. Exteraal
seminal vesicle large, elongate, sometimes
coiled, covered externally with layers of glan-
dular cells, Vas deferens leads medially from
external seminal Vesicle, gradually diminishing
in diameter, Vasa elYereutia not seen. Testes
in two lateral groups anterior to female geni-
falia, each group extending from osmoregula-
tory canals medially beyood medial border of
ovary, accasionally almost to middle of pro-
glottis. Testes in 2-4 longitudinal and 1-3
transverse rows. Vagina tube-like, narrow,
opening to genital atriunt posterior to cirrus
sac, Vagine leads medially, along posterior
border of cirrus suc to ovoid seminal Tecep-
tacle situated lateral to vitellarium. Ovary fan-
shaped, composed of numerous clavate lobules,
on ventral aspect of medulla Vitellarium ovoid
to reniform, dorsal and pasterjor to ovary.
Mchlis’ gland spherical. anterior to Vitellarturn,
Uterus transverse, tube-like, paired in cach
proglottis, dorsal to ovary, Tubular uterus ex-
tends from near centre of proglottis, dorsal ta
ovary, anterior to vitellarium, terminating be-
tween proximal pole of external seminal vesicle
and seminal receptacle, Fully developed uterus
sacciform, with prominent anterivr and pos-
terior diverticula. Uteri crass longitudinal
osmoregulatory canals dorsally, extending to
postero-lateral corners of proglottis. Egg eflip-
saidal, thick shelled, Pyriform apparatus coni-
cal, terminating in reflexed filaments. Genital
primordia appear in «20th proglottis; cirrus
suo developed by 26-30th proglottis; internal
seminal vesicle fills with sperm in 33-47th
Proglottis; insemination oocurs in 40-4461
Proglotiis; Vaginal atrophy not seen; full
maturity of female genitalia reached ip c.SOth
progiottis.

DPiveussiont Although described trom a limited
amount of material, Pragamotaenia Johnson
is sufficiently distinctive to he readily recog-
nised, The gravid holotype is, unfortunately,

Pies 9-15, Pragamotaena folrasoni sp. nov. 9, scolex; 10, eve; 11, mature proglottis; 12, lateral region
of mulure proglollis showing genitalia; 13. transverse histological seetion through lateral region of
mature progtottis showing cirrus sac and muscolature, 14, transverse histological scetion through eor-
tex showing musculature; 15, gravid proglottis, Scale lines, fig. (0, 0.01 mom, figs 9, 11-15, 0 mm.

72 I, BEVERIDGE

without scolex, but the remaining specimens
are not gravid. Mature proglottides of the
specimens from Inkerman are identical mor-
phologically with those of the holotype, and
there is no doubt that they are conspecific.
There is no possibility of confusion with con-
geners that occur in L. conspicillatus, reviewed
by Beveridge & Thompson (1979), since other
intestinal species occurring in this host in east-
ern Australia have markedly fimbriated vela
(Beveridge 1976).

P. johnsoni most closely resembles P. ban-
crofti in being a very large, thick species with
greatly extended proglottides, a broad un-
fringed velum, a powerfully developed cirrus
sac and armed cirrus and an external seminal
vesicle covered with glandular cells. The last
characteristic distinguishes P. johnsoni and P.
bancrofti from all congeners. P. johnsoni is
distinguished from P. bancrofti by a smaller
scolex, a muscular wall to the internal seminal
receptacle, the presence of a pyriform appara-
tus in the egg, and most importantly of all, in
the distribution of the testes, which in P. john-
soni extend from the osmoregulatory canals
beyond the medial margin of the ovary almost
to the centre of the proglottis, but which are
restricted in P. bancrofti to the region lateral
to the ovary. Although in every proglottis of
P. johnsoni examined, the testes were distri-
buted in two groups, in some cases, the dis-
tance between the two groups of testes in the
centre of the proglottis was quite small and
examination of further specimens may well re-
veal instances in which the two groups of
testes fuse in the midline.

An unusual feature of the anatomy of P.
johnsoni is the condensation of muscle fibres
to form a sphincter-like annulus around the
distal extremity of the cirrus sac. A sphincter
surrounding the genital atrium has been des-
cribed in a number of species of Progamo-
taenia (Baer 1927, Lewis 1914), but Beveridge
(1976) considered that the structures reported
by the earlier writers in no way constituted a
sphincter, consisting as they did of an accu-
mulation of parenchymatous elements. The
structure described above in P. johnsoni is a
distinctive muscular structure which is in some

respects sphincter-like. However, its function
is not known at present.

The species is named after Mr P. M. John-
son, National Parks and Wildlife Service, Pal-
larenda, Queensland, in appreciation of help
given in collecting specimens.

Progamotaenia bancrofti (Johnston, 1912)
FIGS 16-23
Material examined: from Onychogalea frae-
nata: 2 specimens, Dingo, Qld, 12.vii.1975,
J. E. Nelson; 1 specimen, Dingo, Qld., (cap-
tive animal) March, 1978, I. Beveridge.

From Onychogalea unguifera: 6 specimens,

Wernadinga Station via Burketown, Qld,
6.viii.1979, R. Speare and P. M. Johnson; 8
specimens, Chadshunt Station via Mt Surprise,
Qld, 7.vili.1979, R. Speare and P. M. John-
son; 2 specimens, ‘Kimberley Ranges’, W.A.,
31.viii.1976, L. Keller.
Description: Length 198-312 (243); width 9-
18 (14); scolex diameter 2.21-2.75 (2.46);
sucker diameter 1.05-1.38 (1.25) x 0.88-1.28
(1.02); no. proglottides 643-672 (660); mature
proglottides 6.6-9.5 (7.9) x 0.34—0.42 (0.38);
gravid proglottides 9.2-10.6 (9.8) x 0.40-0.70
(0.55); cirrus sac in mature proglottides 0.66—
1.52 (1.11) x 0.23-0.34 (0.26); cirrus sac in
gravid proglottides 1.4-1.9 (1.6) x 0.26-0.32
(0.29); no. testes per proglottis c. 200; testis
diameter 0.06-0.09 (0.07); ovary 0.60-0.80
(0,67) x 0.24-0.35 (0.29); vitellarium 0.40-
0.57 (0.47) x 0.14-0.24 (0.18); Mehlis’ gland
0.11—0.17 (0.14); dorsal osmoregulatory canal
0,02-0.06 (0.04); ventral osmoregulatory
canal 0.22-0.35 (0.30); egg 0.036-0.041
(0.039); embryophore 0.017—0.024 (0.018);
oncosphere 0.014—0.020 (0.016).

Large, broad, ribbon like worms, Scolex
large, globular, distinctly demarcated from
scolex. Four cup-shaped suckers embedded
within scolex. Neck absent. Proglottides
greatly extended transversely, craspedote, with
broad, fleshy, folded velum radiating outwards
from strobila, covering + to # of the adjacent
proglottis. Mature proglottides with approxi-
mate length: width ratio of 1:20 to 1:23. Gra-
vid proglottides with ratio 1:16 to 1:20, Ter-
minal proglottides narrower with ratio 1:8.

Figs 16-23. Progamotaenia bancrofti (Johnston). 16. scolex; 17. transverse histological section through
cortex showing musculature; 18, 19. eggs showing variation in shape and in thickness of embryo-
phore; 20. lateral region of mature proglottis; 21. female genital complex, from hand cut trans-
verse section; ventral towards top of page; 22. postmature proglottis showing tubular uterus; 23. gra-
vid proglottis. Scale lines, figs 16, 23, 1.0 mm, figs 18, 19, 0.01 mm, figs 17, 20-22, 0.1 mm.

73

SPECIES OF PROGAMOTAENIA (CESTODA: ANOPLOCEPHALIDAE)

74 L BEVERIDGE

Longitudinal musculature powerfully de-
veloped, composed of pallisades of muscle
bundles. Bundles elongate, arranged radially.
bundles towards periphery smaller, with fewer
fibres, Transverse muscles well developed,
forming a Uccse band along vortico-medullary
junction, Derso-ventral fibres prominent,
crossing corles anid medulla at regular inter:
vals. Longitndinal osmoreguiatary carnals
paired, Ventral canal wider than dorsul canal,
sitvialed medial to at Small accessory canal
associated with ventral canal, on ventral side
of ventral canal, Transverse canal ¢annects
left and nigh! ventral canals al posterior mat-
gin of cach proglottis, Contplex of very
fine branching and anastomosing vessels asst
ciated with dorsal system at posterior mrareins
of proglottides. In scolex, vessels Ivad to con-
necting Ting Yessel in transverse plane at level
of anterior margins of suckers. Genital duets
cross osmoregulatory canals dorsally, Genital
atrium shallow, with corrugated walls, open-
ing in middle of lateral proglottis margin, cutes-
ing interruption af velum. Cirrus suc power-
fully developed with thick muscular walls, ex.
tends beyond osmoregulatory vessels mite
medulla, Cirrus widest at distal extremity,
heavily armed with spines; mid-region coiled,
unarmed, Internal seminal vesicle present,
usuully reflexed distally when filled. External
seminal vesicle large, cloneate, covered exter-
nally with layers of glandular cells, Vas ce
ferens inconspicuous, coils anteriorly tu testes.
Vasa efferentia pot seen, Testes distributed in
two compact groups anterior to cirrus sac, ex-
tending from longitudinal osmuregulatory
canals to lateral margin of OVary. Testes in 3-5
longitudinal and 4-6 transverse rows. Number
of testes in cach group could not be counted
accurately,

Vagina tube-like opening to genital atrium
posterior to cirrus sac, distal Vagina seen only
in serial sections. Vagina lends medially along
ventral aspect of cirrus sac, crosses tu dorsal
aspect of medulla, gradually imereasing i
diameter, Seminal reeeptacle not clearly
separated from yayina. situaled lateral to
vitellarium. Qvary fan shaped, composed of
numerous clavite lobules, on ventral aspeet of
medulla, Vitellarium reniform, dorsal and pos-
terior in ovary, partially enclosing Melis!
wland, which is spherical, anterior to vitel-
larium,

Uterus transverse, tube-like, piited in wich
proglotiis, dorsal to oVaty, Tubular Uterlis ex-
tends from near proglottis midhine, anterior to

Vitellarium and dorsal to ovary, runs along
Ventral aspect of medulla almost to longitu-
dina! osmoregulatery cunaly, Fully developed
uterus sacciform. With prominent «anterior
diverticula, and fewer, smaller posterior
diverticula; crossing longitudinal osmoregula-
tory canals dorsally, extending to postero-
lateral corners of proglottides, Egg elongate
ovoid, outer wall thin. Pyriform apparatus
absent, even in egas from faeces; embryophore
surrounding egg thick, ellipsoidal.

First mature proglottis 165-235rh; filling of
seminal receptacle occurs ut approximately
same time as sperm appears in internal seminal
vesicle; vagina does not atrophy Following in-
semination; ulerine filling commences inc.
Oth proglottis.

Discussion: The descriptions of Progamotaenia
hancrofti by Johnstan (1912), Nybelin (1917)
and Beveridge (1976), are unsatisfactory be-
cause only a limtled number of specimens was
available, The original desetiption by John:
ston (1912) was based on a single non-gravid
specimen from Onvechogalea fraenata, the
description by Nybelin (1917) was based on
four specimens from O. ungul/era, ull of which
were severely contracted, and only one was
gravid, Beveridge (1976) re-examined all these
specimens but was able to add little. The type
host, O. fraenata is now almost extinct, so that
the main source of material for the present re-
description came from the related wallaby. OQ.
lingnifera.

Although ample material was available, the
species is catremely difficult to examine be-
cause of its large size and the thickness of the
longitudinal musculature and velum, Nevers
theless, the present re-deseription supports
earlier descriptions in most respects, dilfcring
only in the features of the uterus panel external
seminal Vesicle, The utertis conimences de-
velopment as i transverse tube, gradually en-
jarginy iv disineter as it fills with cags. la post-
mature proglottides, in which the ovary las
involuted, the Uterus remains tuhe-like, with
Hitle evidence ef anterior allel posterior diver-
cua. ‘The lattur develop subsequently in gra-
vid proglottides. The lack of diverticula on the
Wteri of the holotype deseribed by Beveridge
(1976), presumably reflects the lnpmaturity of
the specimen rather than a significant
difference beiween wv and Lie new material.

A prostate ait the pragimal pole of the cirrus
sue Was dleaerlbed in 2 banevoffl by Nybelin
(1917) and Beveridge (1976). The structure

SPECTES OF PROGAMOTAENIA (CESTODA; ANOPLOCEPHALIDAE) 75

is in fact an external seminal vesicle, diflermg
from comparable structures in congeners in
being elongate rather than ovoid, and sur-
rounded by a mass of glandular cells. This
lattwr feature serves to sepurate PL henere/ti
from all] congeners except P, Jolinsant, How-
ever, here the term “prostate” has been dis-
carded in favour of “external seminal vesicle”
to avoid confusion with a structure (also
termed the prostate) described in certain spe-
cies of ihe anoplavephalid genera Anirva
Railliet 1893 and Ptandrya Darrah 1930,
Rausch (1976) has cast considerable doubi on
the existence of 9 prostate in these genera, sug-
gesting that the orga in question jis the exter-
nal seminal vesiele.

Nybelin (1917) stated that a pyriform ap-
paratus was not present, bot Beveridge (1976)
cautioned that the few specimens available te
Nyhelin may not have heen fully gravid, In
the new material, a pyriform apparatus was
found neither in the terfninal proglottides of
strobilae which were fully gravid, nor in shed
Progloetides collected From the large itestine
of the host. Tt was fot established whether a
pyriform apparatus develops in the external
environment following voiding. The anly can-
gener in which a pyriform apparatus is lack-
ing 1s P. lapgorchestis. In both species the ege
is clonpate rather than spherical ar ovoid,

Serial lonwitudinal sections of the strohila
revealed that the distal vagina is an extremely
narrow duct bul that i remains patent even
m proglotiides in which the uterts is in the
process of filling, The distal vagina was not
s¢en om every proglottis, but in a sulficren!
number bo indicate that earlier descripiions
have been incorrect, and what the distal vagina
is merely dilicull to find, even in sections,
rather than buving atrophied following insem-
ination.

Further collecting has confirmed Onveto-
valea fraenata ind O, wmeniferad as hosts of P,
banerefti, The species has not heen fhynd in
Wallabla bicoler or Serouly braelevis apart
from a single report by Sandars (1957). Her
brief descriptions do nor conform to eutlicr
more detailed descriptions {Beveridge 1976).
The location of Sandars’ specimens is \n-
known, and. their identifications may be i
error.

P, bancrofti was present in seven ot ten €,
unoulferd, with either mine or two cestodes per
wallaby. The cestodes veeutred tu the ileurn.

Progamotaenia diaghana (Zschokke, 1907)
FIGS 24-29

Material examined: types from Lasiorhinus
laiifeons; numerous fragmented specimens,
from bile duets, Swan Reach, 8.A,, 12.v,1977,
I, Beveridge; 14.1%.1978, M, Gaughwin,
Deseription: Leugth up to 102; width to 2;
scolex diameter 0,32-0,46 (0.38); sucker dia-
meter 013-0.16 (O15); neck 0,07-0.16
(O.13)5 mature proglottides 1-1-1.9 (1.5) x
0.23-0,43 (0.31); gravid proglottides 1,7-2.6
(2.0) x 0.28-0.35 (0,34); cirrus sac in matute
proglottides 917-039 (0.29) x 0.06-0.09
(0.07); cirrus sac in gravid proglotlides 0.22-
0.33 (0.28) * U.04-0,09 (0,07); no. testes per
proglottis S$9-64 (S51); testis diameter (.04—
0.05 (0.04); ovary 0,080.14 (11) x 0,04-
0.10 (0,08); vitellarium 0.06-0,08 (0,07) x
O.04-0.06 (0.05); Mehlis’ gland 0.04~0.05
(0.05); dorsal osmoregulatory canal 0.01
0.02 (0,02), ventral osmoregulatory canal
0.02-0.04 (0.03): egg 0.07-0.08 (0.08); pyri-
form apparatus 0.04-0.05; oncosphere 0,03,

Small, fragile worms, almost transparent and
casily broken when fresh, Scolex prominently
four lobed, with each muscular, cup shaped
sucker borne on arm like extentions of scolex,
Seolex frequently, but not invariably dark,
pigmented, Pigment distributed mainly on
arms and at apex of scolex. as masses of small,
brown staining accumulations of granules in
the cortex immediately below the tegument,
Small pigmented granules seatlered around
vieinily of central nervous system. Neck pre-
sent in relaxed specimens. Proglottides ex-
tensed transversely, craspedote, with narrow
fc. 0.05) straight-edyed velum overhanging ad-
jacent proglottis, Mature proglottides With ap-
proximate length: width ratia of 172.5 to 1:8,
Gravid progloitides with ratio 1:55 to 1:75.
Longitudinal muscles poorly developed, com-
posed of 2 pings of fibre butdles with 2 to 8
fibves per bundle. Tyansverse muscle consists
ef individual fibres running along cortico-
medullary jinection, Darso-ventral muscle
filwes sinvle, crossing cortex god medylla at
irregular intervals.

Longitudinal osmoregulatory canals paired,
Ventral canal wider than dorsal canal, situated
medial ti it, Two transverse canals connect
Jeft and right osmoregulatory canals at pos-
terior margin of each proglottis, Canal cone
necting ventral vessels of moderate size,
readily observed: canal connecting dorsal ves-
sels minute, seen only with difficulty, function

I. BEVERIDGE

SPECIES OF PROGAMOTAENIA (CESTODA: ANOPLOCEPHALIDAB) 7

of osmoregulatory canals in scolex not seen.
Genital ducts cross Osmoregulatory canals dor-
sally. Genital atrium of insignificant size,
sometimes situated on small genital papilla,
Atrium in posterior part of lateral proglottis
margin, dividing margin in ratio of 1:2 to 1-3.
Cirrus sac narrow, clavate, musculature of
walls Weakly developed, always reaching ald
usually extending beyond longitudinal os-
moregulatory canals. Cirrus narrow, uncoiled,
armed with minute spines, only clearly visible
on exerted cirri. Prominent internal seminal
vesicle present. External seminal vesicle large,
ovoid, extending to Jeteral margin of, or oc-
casionally anferior to, serninal receptacle, Vas
deferens coiled, passes anteriorly, diminishing
in diameter. Vasa efferentia not seen, Testes
distributed between lateral osmoregulatory
canals anterior to female genitalia, Testes
either in two groups extending from longitu-
dinal canals to medial side of female geni-
talia, or in continuous band across proglottis,
with testes more numerous in lateral parts of
band. Both forms of testis arrangement occur
together in some strobilac, Vagina tube-like,
opening to genital atrimm posterior to cirrus
sac. Vagina leads medially to large ovoid, thin-
walled seminal receptacle. Ovary fan-shaped,
composed of numerous, clavate lobules, on
ventral aspect of medulla. Vitellarium ovoid
or reniform, posterior and dorsal to ovary,
Mchlis’ gland spherical, anterior to Vitellarium,
dorsal to ovary, Uterus transverse, tube-like,
paired in each proglottis, anterior to seminal
receptacle and ovary. Fully developed uterus
sacciform, without diverticula, crossing longi-
tudinal osmoregulatory canals dorsally and
reaching postero-lateral corner of proglottis,
Uteri of proglottis may fuse Jn mid-line on
rare oceasions. Egg spherical, thick-shelled,
Pyriform apparatus cither conical or terminat>
ing in two horns, numerous teflexed filaments
attached to apex of pyrifarm apparatus.
Sperm first appears in external and internal
seminal vesicles in ¢.105th proglottis; seminal
receplacle fills will sperm tn c.1 10th proglot-
tis, after male system commences to function,
Vagina does not atrophy following insemina-
tion. Ovary fully developed from ¢.140th to
150th proglottis, involutes over 2-3 proglot-

tides, Uterus begins to fill with eggs im-
mediately after involution of ovary (¢.1 50th),
Discussion: Progamotaenia diephana was te
described very briefly by Beveridge (1976)
based on a single specimen assumed to be type
Material aod a small number of fragmented
specimens from a captive wombat. Although
the latter collectian suggested that Laxiorhinuy
Jatifrens ovight be the usual host of FP. alu.
phana, no material had been seen from free-
living hosts, and the very closely related ces-
tode, FP. festive, had been collected from the
bile ducts of free-living Vombatus ursinus
(Beveridge 1976), Subsequently, collections of
cestodes fram LL. latifrons in South Australia
were tentatively identified ag P, festiva (Rudel-
phi 1819) as they did not conform exactly to
the description of PF. diaphana, and doubt was
thrown on the status of P. dfuplana itself
(Presidenic & Beveridge 1978). Abundant col-
lections now do hand inilieate thar P, afop hana
is a distinctive form, probably warranting
specific status, and that doubt as tr the identity
of Presidente & Heveridge’s maternal was doc
to a lack of apprectation of the extent of Varia-
bility in the species.

The present redescription differs from that
of Beveridge (1976) in a number of potnts.
A prominent external seminal vesicle was pre
sent in the new nvaterial, but was stated as
being absent in the earlier description, The
number of testes per group was previatisly re
ported as 17-21, but in the new material &
18-33 per group or 39-53 per proglottis. Pan
of this variability is due simply to vanahion
between individupl cestudes, In two srebilwe
examined, the range aml mean number of
testes per proglottis based on examination of
LO proglottides in each strobiln was 32-463
(56) anc 39-53 (46), Number of testes per
Proglottis in the related P. frstiva is FU-140,
so that in spe of the vartubiliny, testis. wuo-
ber is still a uselul uethod of distiuguistung the
species,

In the type specimen redescribed by Bever-
idge (1976) early tasemination of proglottidce
was followed by vaginal atroghy. This pheno-
menin wis not recorded in the orginal des
éription (Zschokke 1907) and was not present
in the ofher material described hy Beveridge

Figs 24-29, Progamotacnia diaphana (Zschokke). 24. transverse histalogical section uf Larcral region of
proclottis. showing cirrus sac, uterus and muscilawire; 25. mature proghottiss 26. mature pravhsides
showing vuriation in proglottis shape and testis distribution; 27, gravid proglottiss 2h, scelux; 29,
mature proglottis showing variation in testis distribution. Scale lines 0.1 mm,

78 I, BEVERIDGE

(1976), In the new material, insemination in
variably occurs after the male reproductive
system of a particular proglottis has stunted to
function and vaginal atrophy sever occurs
following insemination, The distal vagina js
frequently difficult to see in moulited spect-
mens, so fragments of the type material were
serially sectioned, showing thal the vagina re-
mitines? intact in past mature proglottides and
suggesting that Beveridge’s interpretation based
on the single whole mount preparation was in-
correct. BeVeridge’s (1976) suggestion that
vaginal atrophy could vary within a species
should therefore be ignored, The sugrestion
has alsa proven incorrect in the case of P,
laygorchestis and P. tylogaie (Heveridge &
Thompson 1979); species which were initially
confused linder the one name.

Two diagrams of the egg of P. diaphana
were given by Beveridge (1976), one showing
the pyriforin apparatus ending in two horns
and the other showing an undivided pyrilonm
apparatus, Both forms ure present im the sew
material but, being unmounted, it is possible
to rail the eggs under a coverslip and show that
there are indeed two different types and that
they are Wot the same form viewed from dif-
ferent aspects (Beveridge 1976). The undi-
vided form ts identical with that found in P,
festive and the divided form may merely be a
developmental stage (Beveridge 1976), since
it occurs in much lower numbers, This form
of pyriform apparatus was illustrated by Bohm
& Supperer (1958).

The presence of dark pigment in the scolex
of P, diaphana was first reported by Béhm &
Supperer (1953), and ts present in the single
scolex antony the type material as well as in
all lots of new material. The pigment, how-
ever, is not invariably present and cannot
therefore be used as a distinguishing character.

Beveridge (1976) Hsted a number of fea-
tures which distinguished P, diapkana from
the related FL festive, Of these, the presence uf
proterdgvny ad vaginal atrophy should be dis-
carded, while the shape of the uterus requires
a subjective assessment and is therefore of
doubtful value, ‘The two species differ in the
frequent presence of dark pigment jn the sca-
lex of F. diaphane and differences in the num-
her of festes per penglottis, allowing for the
considerable variation which occurs in’ hoth
species. Ly view of the present redescription,
P. digphane probably warrants continued
specific rank, and the specimens described tn-
der the nanic P, festive from L, latifrons by

Presidente & Beveridge (1978) should be re-
named FP. dlaphana. However, Beveridge
(1976) has pointed to the exlensive Variation
in specimens of P. feytiva from various host
species, and it is cyident that a reassessment
of this taxon and P. diaphana will be necessary
when life histories are known and cross infec-
tion experiments can be undertaken.

The data presented above establish 1, lati-
Jrons as the usual host for P. diaphana. Rohm
& Supperer (1958) reported P. didphana from
a captive Vombatus ursinus in Europe, but
subsequent correspondence by a colleague with
Supperer established that this animal was in
fact LL. latifrons (Presidente & Beveridge
1978), and had earlier been misidentified. In
his original description, Zschokke (1907) gave
no details as to how he obtained his speci-
mens, and gave the bost name as Phascolomys
wombet, a name which is now placed as a
synonym of V. ursinas, but which could have
becn applied to any of the species of wombats.
If, as indicated by data associated with the
material considered now to be type material,
the species was initially collected by E. Angas
Johnston, then ZL, Jatifrons may be the type
host. Angas Johnston was a doctor afd ama-
teur naturalist who lived in Adelaide.

lrogamotacnia zschokkei (Janicki, 1909)
Material examined; From Onychogalea frae-
nata: 1 specimen, Dingo, Qld, 12.vit.1975,
J. E. Nelson; 5 specimens, captive atimals
originate from Dingo, Qld, March 1978,
19.i,-1979, T, Beveridge: 8 specimens, same
data, 26.yii,|979, R, Speare. From Onveho-
gelee wnguifera: 4 specimens, “Kimberley
Ranges", WA. 3Lviill976, 6.xi1.1976, L,
Keller; 2 specimens, Chadshunt Stn, Qld,
6 ili 1879, R, Speare and P. M. Johnson.
From Lagorchestes conspicillatus: 5 speci-
mens, Inkerman St, Qld, 7.i%.1979, 141.1979,
P.M. Johnson; 3 specimens. Mingela, Qld,
10.v.1979, | Beveridge. From Thylogale stie-
inaneat 10 specimens, El Arish, Qld, 30, vii.
1978, |. Beveridge, From Petrogale penictllata:
3 specimens. Hervey’s Range, Townsville,
Old, d.iv.1979, 1. Beveridge. From Macropus
agilis: 5 specimens, Marrakai Plains, N.T,
29,jx.1973, L, Corner; |3 specimens, Towns-
Ville. Old, T5.vi,1978, 1, Beveridge,
Discussions Onvehagalea fraenata, O. ungui-
Jera and Meecropus aeiliy are new hosts for
Progamoteenia zchokkei, Part of the material
listed above from M. agilis was identified
erroneously as P. Jagorchestis by Beveridge

SPECIES OF PROGAMOTAENIA (CESTODA: ANOPLOCEPHALIDAE) 7

(1976). M. agilis is not a host of P. lagorches-
tis or of P. thylogale, a species which was con-
fused under the former name by Beveridge
(1976), Further specimens of P. zschokkei
from Thylogale stigmatica and Petrogale peni-
cillata confirm earlier reports from these hosts
based on rather poorly preserved material
(Beveridge 1976).

There was considerable size variation in the
specimens of P. zschokkei examined. Those
from OQ. Jraenata measured 95-120 x 11-12
mm; specimens from Q. unguifera were latger,
measuring 380-460 x 8-12 mm. Specimens
from L, conspicillatus were 160-170 mm long
and varied from 8-13 mm in width, Cestodes
from M, agiliy were long (130-185 mm) but
only 4-6 mm wide while specimens from P,

penicillata and T. stigmatica were the smallest,
measuring 58-61 x 2-3 mm and 50-90 x 2-3
mm respectively. In spite of the great differ-.
ence in size, there were no significant differ-
ences in internal morphology. The size differ-
ence may be due to the influence of the host,
but experimental infections will be needed to
demonstrate whether this is the cause of the
observed differences, or whether a species com-
plex exists. Similar size variation has been
noted in P. festiva and P. macropodis (Beyer-
idge 1976),

Acknowledgments
Thanks are due to M. Gaughwin, R. Speare
and P, M. Johnson for collecting specimens,
or for supplying wallabies for dissection,

References

Baer, J. G. (1927) Monographie des cestodes de
la famille des Anoplocephalidac, Bull. bial. Fr.
Belg. Suppl. 10,

Beveriwor, T, (1976) A taxonomic revision of the
Anoplocephalidae (Cestoda: Cyclophyllidea) of
Australian Marsupials, Aust. J, Zool. Suppl.
(44), pp. 110.

—— (1978) Progamotaenia tuficola sp, nov.
(Cestoda; Anoplocephalidae) from the red kan-
garoo, Macropus rufus (Marsupialia). J. Para-
sitol. 64, 273-276,

& Thompson, R.C. A. (1979) The anoplo-
cephalid cestode parasites of the spectacled hare
wallaby, Lagorchestes conspicillatus Gould,
1842 (Marsupialia: Macropodidae), J. Helmin-
thal, 53, 153-160,

Roum, L. K. & Suppercr, R. (1958) Beitriige zur
kenntnis tierischer Parasiten (1, Zentralbl, Bak-
terlal, Abt, 1, Orig, 172, 298-309,

Jonnston, T, H, (1912) Notes on some entozoa.
Proce. R. Soe. Qld, 24,, 63-91,

Lewis, R. C. (1914) On two new species of tape-
worms from the stomach and small intestine of
a wallaby, Lagarchestes conspicillams, from
Hermite Island, Monte Bello Islands, Proc, zool,
Soc, Lond., 419-433,

Nyse, O, (1917) Australische Cestoden. Re-
sults of Dr E. Mjéberg’s Swedish Scientific Ex-
peditions to Australia, 1910-1913, Kungl.
Svensk. VWetenskapsakad. Hand], 52, 1-48.

Presiente, P, J, A, & Bavertmce, J. (1978)
Cholangitis associated with species of Proga-
molaenia (Cestoda; Anoplocephalidae) in the
bile ducts of marsupials. J. Wildl, Dis. 14, 371-
377.

Rauscn, R. L. (1976) The genera Paranoploce-
phala Lithe, 1910 and Anoplocephaloides Baer,
1923. Ann. parasit, hum. comp, 51, 513-562.

Sanpars, D. F. (1957) Redescription of some ces-
todes from marsupials. IT. Davaineidae, Hy-
menolepididac and Anoplocephalidac, Arn,
Trop, Med, Parasitol, 51, 330-339.

ZSCHOKKE, F, (1907) Moniezia diaphana n. sp.
ein weiterer Beitrag zur Kenntnis der Cestoden
aplacentaler Saugetiere. Zentralbl. Bakteriol,
Parasitenkd, 44, 261-264,

BEVERIDGEA NEW GENUS (NEMATODA: STRONGYLIDA) FROM THE
AGILE WALLABY FROM NORTHERN AUSTRALIA

BY PATRICIA M. MAWSON

Summary

Beveridgea n.g., type species B. corneri, n.sp., is close to Cloacina, differing chiefly in the much
longer buccal capsule, and in the shape of the bursa, which is not joined ventrally. B. corneri has
been taken from Macropus agilus, only on Cape York Peninsula, Queensland.

BEVERIDGEA NEW GENUS

(NEMATODA; STRONGYLIDA)

FROM THE AGILE WALLABY FROM NORTHERN AUSTRALIA

by ParriciA M. MAwson*

Summary
Mawson, P. M. (1980) Beverideea new genus (Nematoda: Strongylida) from the Agile
Wallaby from northern Australia, Trans. R, See, S, Aust. 10404), 61-82, 30 May, 1980,
Beveriigea wg.. type species BL corneri, msp, is close to Cloacina, differing chiefly in
ihe much longer buceal capsule, and m the shape of the bursa, which is not jammed ventrally.
B. corner? bas been taken from Macrapus agilis, only on Cape York Peninsula, Queensland,

Bevyeridgea fg.

Cloacininac: cephalic roll well developed;
submedian cephalic papillae elongate, bi-
seymented, buccal capsule cylindrical, longer
than its diameter, votehed along anterior
border; leaf crown of eight elements arising
near anterior end of buccal capsule; cervical
papillae thread-like; dorsal oesophageal gland
opens into base of buccal caviry-

Male; spicules elongate, alate, bursa not deeply
lobed, ventral lobes almost separate; ventral
rays together, ventro-lateral ray separate from
other faterals, externo-dorsal ray arises
separately, dorsal ray divides twice, Guberna-
eulum present

Female: tail short, conical; vulva near anus,
junction of ovejectors parallel with long axis
of body,

Parasites of stomach of macropodid marsu-
pials.

Type species: B. corneri n. sp.

Beveridgea has been referred to the Cloaci-
ninae sensu Lichtenfels (1980) because of the
type of Jeaf crown and cephalic papillae, and
the fact that the dorsal ocsophageal gland opens
directly into the base of the buccal cavity.
Tt is distinguished from Cloacina Linstow,
1898. mainly by characters of the buccal cap-
sule, which is longer, and of the leaf crawn,
the elements of which arise from the anterior
end of the capsule wally moreover, in Clawelna
the ventral lobes of the bursa are joined.

Beveridgea cormeri o.sp.
FIGS 1-7
Host and loealitiess Macropes agilis Gould, From
Elizabeth Downs Station (1ype locality) and
from Stones Crossing, Wenlock R., Qld. Holo-
type male and allotope female deposited in
South Australian Museum (V19I0),

* Department of Zoology, University of Adelaide,
Box 498, G,P.O., Adelaide, S. Aust. 5001.

This species was taken from at least three
host animals, but very few were present in
each; all are young worms, the females with-
out eggs in the uteri, The species hus not been
found among stomach worms from M, agilis
from more southerly parts of Queenslabd or
from the Kimberley District of W.A.

Length of males 4.7-5.7 mm, females
4.9-5.3 mm. Labial collar well developed,
slightly Jobed anteriorly. Cephalic papillae of
two segments, distal distinctly shorter than
proximal, Buceal capsule 40-50 ym long, its
anterior border notched, its posterior border
lobed, following contour of anterior end of
oesophagus. Fight elements of leaf crown
anise from antetior 1,, of wall of buccal cap-
sule, and end in points around mouth. In
posterior ), of buccal capsule small jeregularly
pluced denticles project into buceal cavity.
Oesophagus 480-520 ym long in male, 440-
$50 pm in female, cylindrical in anterior half,
then slight swelling preecdes position of nerve
rig, and posterior portion widens towards
base, In median swelling small tooth projects
into Jumen. Dorsal oesophageal gland distinct,
openiig on a prominence at anterior end of
dorsal section of oesophagus.

Nerve ring lies just posterior to median
swelling of oesophagus; exeretory pore near
base of oesophagus: thread-like cervical
papiliaeg 125-130 (4) and 120-140 (2) froro
anterior end,

Male: bursa with characters of genus. Dorsal
ray divides just hefore midlength, and final
branches very short. Genital cone short,
flanked by two lateral “balloons” of apparently
inflated cuticle. From base of dorsal ray single
digitiform ‘papilla’ projects into cavity of
bursa (Fig. 6).

Female: tail 150-200 pm long, vulva 220+
310 pm from posterior end, Eggs absent.

82 PATRICIA M, MAWSON

100 jim

Figs 1-7, Beveridgea corneri n.sp. 1 head, lateral view; 2. anterior end, dorsal view; 3. oesophageal
region; 4, 5 and 6, bursa in dorsal, ventral, and lateral views respectively; 7. posterior end of
female. (Figs 4-6 to same scale).

Acknowledgments Dept of Tropical Veterninary Science, James
The material for this study was kindly Cook University of North Queensland). It was
given to me by Dr I. Beveridge (then of the collected by him and by L. Corner.

References

LICHTENFELS, J. R. (1980) Keys to genera in the Linstow, O. von (1898) Nemathelminthen gesam-
superfamily Strongyloidea. CIH Keys to the melt von Prof. Dr F. Dahl in Bismarck Archi-
nematode parasites of vertebrates, No. 7. pel. Arch. f. Naturg. 1, 281-291.

Commw. Agric. Bur., Farnham Royal.

NON-MARINE MOLLUSCS FROM DOLOMITIC LIMESTONES IN THE
NORTH OF SOUTH AUSTRALIA

BY N. H. LUDBROOK

Summary

Non-marine molluscs, from the type section of the Etadunna Formation at Lake Palankarinna
(KOPPERAMANNA 1:250 000 map sheet), and from dolomitic limestones on the BILLA
KALINA and TARCOOLA 1:250 000 map sheets are described and correlated with those occurring
in northern Australia. Etadunna Formation molluscs from Lake Palankarinna are land snails —
Bothriembryon praecursor and Meracomelon lloydi — while those from near “Billa Kalina” and
“Malbooma” and also from Lake Woorong on the COOBER PEDY 1:250 000 map sheet are
freshwater species of Syrioplanorbis, Physastra and Rivisessor. The dolomitic limestones are
considered to be of Miocene age.

NON-MARINE MOLLUSCS FROM DOLOMITIC LIMESTONES IN
THE NORTH OF SOUTH AUSTRALIA

by N. H, Lupprook®

Summary

Luperook, N, H, (1980) Non-marine molluscs from Miocene dolomitic limestones in the
north of South Australia. Trans, R. Soc, S. Aast. L04(4), 83-92, 30 May, 1980.

Non-muriine molluscs, from the type section of the Etadunna Formation at Lake Palan-
karlnna (KOPPERAMANNA 1:250-000 map sheet), and from dolomitic limestones on the
BILLA KALINA and TARCOOLA 1:250000 map sheets are described and correlated with
those occurring in northern Australia. Etadunna Formation mollitscs from Lake Palankarinina
are land snails — Berliriembryon praccursor and Meracomelon lloydi — while those from neat
“Billa Kalina” and “Mualbooma" and also from Lake Woorong on the COOBER PEDY
1.250000 map sheet are freshwater specics of Syrioplanarbis, Physasrra and Rivisesser,
The dolomitic limestones are considered to be of Miocene age

Introduction

In describing fossil pon-marine molluscs
from northern Australia, McMichael (1968)
cited sone of the species as occurring also in
the “Etadunna Formation, Billa Kalina Station,
South Australia.” ‘Tabulating the localiues.
lithology and faunas of samples examined by
MeMichael, Lloyd (1968) similarly attributed
to the Etadunna Formation, Tirari Desert,
three species, two of which came from Lake
Palankarinna and one from Billa Kalina, The
present paper seeks to correct the unfortunate
confusion of two widely separate localities,
sbown on Figure 1, and to distinguish between
those molluses which occur in the type section
of the Etadunna Formation at Lake Palan-
karinna and those in the dolomitic limestones
cropping out north of “Billa Kalina” Home-
stead. The fossil content of dolomitic lime-
stones from near “Malbooma 0.5.” is also
placed on record,

All the material studied is in the Palaeonto-
logical Collection of the Geological Survey of
South Australia, All may references are to the
Geological Atlas Series,

Specimens from both Lake Pulankarinna
and north of “Billa Kalina” were sent to
MeMichael at the Australian Museum in 1963
with references to published data on the Eta-
dunna Formation and a note to the effect that
"the dolomite containing Planerbiy from Billa
Kalina is not necessarily to be correlated with
the Etadunna,” In official correspondence,
McMichael tentatively identified the Etadunna
gastropods respectively us belonging to the
genus Bothriembryen anid similar to Jand

*e/e Deparrment of Mines & Energy, P.O, Box
151, Eastwood, 8S. Aust. 5063.

snails of the genera Meracomelan and Sinn
melon, probably the former, ynd Lhe material
from Rilla Kalina as a large Planorbis-like
shell,

In 1965, Etadynna specimens were again
sent to McMichael to supplement his studies
of the northern Australian fauia, No material
from Billa Kalina was included, McMichael’s
(1968) reference to the occurrence of certain
species in the “Eladunna Formation, Billa
Kalina Station" seems to originate in noles he
made m 1963 when he had specimens from
both localities.

One of the Unfortunate cousequences of the
error is (hat the gastropods of the type seetion
of the Etadunna Formation at Lake Palan-
karinna have been only obscurely recorded,
Moreover, the molluscs from the Etadutna
Formation are land snails, while those from
Billa Kalina are freshwater. The only Iime-
stones containing both land and freshwater
moltluses (tabulated by Lloyd 1968) appear to
be in the Deep Well area of Central Australia
and the Carl Creek Limestone of the Rivers-
leigh area, Queensland

Motluses from the Etadanna Formation

The molluscs from Lake Palankarinna, south
of Cooper Creck, 23 km SW of “Etadunna™
(lut, 28'48'S, long. 138°25°E_ locality sample
number 6540 RS 39, KOPPERAMANNA
1:250 000 map sheet) were collected by R. H.
Tedlord from nodular dolomilic limestone at
the base of member 2e, Etaduijna Formation
(Stirian er al 1961). Additiumal specimens
were collected by 1. M. Lindsay in 1970. They
therefore come rom tow in the formation,
some 5 mi above the base, although gastropods
have been recorded by Stirton and his col-

84

VERTICAL SCALE SECTION AT
("METRES LAKE PALANKARINNA

50-0
WINDRIFT
DUNES

KATIPIRI SANDS

Cross—bedded sands
Gypsum

TIRAR|
FORMATION
TYPE LOCALITY

Red argillaceous
sandstone

PLEISTOCENE

MAMPUWORDU
SANDS
TYPE LOCALITY

Channel sands

ETADUNNA
FORMATION
TYPE LOCALITY

Green arenaceous
® claystone

By Calcareous mudstone

6 Green argillaceous
sandstone

Green arenaceous
claystone

4 Green sandstone

3
ee)

Grewn claystone

Gastropods

Calcsreous mudstone
and dolomitic limestone

Gastropods

Green sraillaceoua
sandstone

WINTON
FORMATION
Grey argillaceous
sandstone

o
Adapted from Stirten.
Tedlord and Miller, |?é!

Bin JPY

w
>
o
tu
o
xt
—
uu
oc
o

FIGURE 2

N, H. LUDBROOK

1797 13)"

|
|

<
=

|
|
|

133" NT

BILLA KALINA

'Malboora 0,

KILOMETRES

LOCALITY MAP

FIGURE 1

139° Ou

——_!

Pe 2a |
KOPPERAMANNA

Lake Patankarinna

SECTION AT
“BILLA KALINA"

Ambrose and Flin, 197)

Dolomite limestones
with dolomite
intraclasts

Clays and shales

Sand lunses with

quartzite cobbles

and boulders
Discenformity

BULLDOG
SHALE

Clays and shale with
sand lenses and
qQuartzle cobbles and
boulders

SJ. Doly, 1979
BO-A4 S.A

SECTION AT
"“MALBOOMA"

Dolomitic Jimestunas
with dolomite
intractasts

Clays and shales

PIDINGA
FORMATION
Garbonaceous
sandy clays

Deportment ol Mines ond Ener

MIOCENE NON-MARINE MOLLUSCS 85

leagues fram calcareous mudstones at both the
basy and top. The measured stratigraphic type
stetion at Lake Palankarinna, redfawn after
Stitton ef al. is shown in Figure 2.

In the opinion of Stirton et a/, (1961), the
sequence of dolomitic limestones, calcareous
mudstones and claystones with intraforimea-
tional breccias represents deposition in a shal-
low-water lygoon with repeated exposure and
drying. The green claystones and argillaceous
sandstones have yielded the important Ngapa-
kaldi vertebrate fauna which contains lungfish
and Water birds. [ft includes also diprotodonts
and macropodids which appear to have heen
entrapped in hoggy clay. Gastropods in the
dolamitic Iimestones are in the form of moulds
and casis, many of which are freed from the
mateix, They seem to be locally common and
arcearious, but belong to only two species,
Bothriembryon praecursay MeMichael 2nd
Meracomelan lloydi McMichael. Both species
are related to land snails typically inhabiting
and parts of the State — Sothrieimbryen bar-
rei ihe Nullarbor Plains and Mferdeornelon
spp, the Northern Flinders Ranges. As no
freshwater shells have so far been found with
them, they are presumed to represent the
drying-out periods of deposition of the Bta-
dunna Formation postulated by Stirton and his
colleagues,

The possible relationships between land
snails from non-marine deposits in and near
Hobart and species of Bathriembryen und
Meracomelon. occurring in the Etadunna For-
mation were considered by McMichael.
Examination of the Jimited amount of
material now available from Hobart, discussed
in the systematic section, has failed to estab-
lish similarity between the Tasmanian species
in the Geilston Travertine and those fron the
Etadunna Formation,

Mollises from the Billa Kalina area

The following deseriplion of the planorbid-
bearing limestoues from worth of “Brile
Kalina” Homestead (locality ind simple num-
hers 6148 RS 62-73, 87, BILLA KALINA
1;250000 map sheet, tat, 29°53'S, long,
136° 11'E) is modified from Ambrose & Flint
(1979)1,

t Ambrose, G. J. & Flint, R. B. (1979), A regres-
sive Tertiary Tyke system and silicitiod strand
lines, Hilla Kalina area, South Australia S.A.
Dept. Mines & Energy Rept 79/104 (Cunjub-
lished).

Tertiary sediments neat “Billa Kalina” form
a thin capping, maximum thickness 13 m, on
shales and conglomeratic sands of Early Cre-
raceous Bulldog Shale. A fesistan! dolomitic
limestone within the Tertiary sequence over-
lying the more-casily eroded shales results in
flat-topped plateaux and mesas.

An idealised sequence (Fig, 2) comprises a
very thin basal sand horizon containing quart-
zite clasts derived from erosion of the Creta-
ceous sediments, This is overisin hy approxi-
mately 5 m of green dolomitic and oceasion-
ally palyporskite-bearing ¢lays, which are in
turf overlain by 1.5 nt of white fossilifeeous
limestones and dolomitic limestones.

Samples with planorbids, ta which the num-
ber 6138 RS 87 has heen assigned, were first
bollected from these limestones in 1958 by
H. G. Roberts during reconnaissance mapping
tor Clarence River Oil Syndicate. Subsequent
collections (6138 RS 62-73, containing hydro-
bids) were made in 1974 by G. J, Ambrose
and R, B. Flint during mapping of the BILLA
KALINA 1;250000 mup sheet from three
localities within 4 distance of 1.5 km from 3,5-
40 km NNW to N of “Billa Kalina”, The total
amount of material is not large, snd the known
Fauna is limited to four species—an undescribed
species of Roivessor occurring in samples 6138
RS 62-73. Syeioplaterbiy heardmani, Syrio-
plunorhis sp., and Physastra padlagee, eithee in
crowded masseg or scattered throughout the
matrix, These were freshwater inhahitunts of
or were washed into the Tertiary lake postu-
lated by Ambrose and Flint,

The Tertiary sediments on Billa Kalina and
Millers Creek Stations have: been correlated,
on a lithological basis, with the Etadiinna For-
mation of the Lake Eyre Basin and the Namba
Purmanon of the Tarkarooleo Basin (Jessup &
Norris 1971; Ambrase & Flint 1979*)_ Jessup
& Norris divided what they considered to be
the Etadunna Formation in the Billa Kalina-
Millers Creek area into two members — a
jower Bula Kalina Clay Member.and an upper
Millers Creek Dolomite Member, A revision
of this nomenclature is presently being pre-
pared by Ambrose and Flint, and, pending its
publication, the Billa Kalina Clay Member and
the Millers Creek Dolomite Member are here
regarded as units of an unnamed formation.
The molluscs in the dolomites provide the only
direct evidence so far obtained for correlating
them with other formations of known Tertiary
age,

8h N. H, LUDBROOK

Material from “Malbouma Outstation" arca

Tertiary fossils were first collected in [his
area in 1979 by R. B. Flint, S.J. Daly and
A, FB. Crooks (locality and sample nunbers
5736 RS 47-52 TARCOOLA 1:250000 mip
sheet. Jat, 30°39'S, long, 134°05°E) A bret
veological description is provided by S. J, Daly:
‘Possible Jute Teniary sediments west ol
*Malbooma O.S." crap out poorly, and fore
low cises which are veneered by calerete. The
best exposures are in radiway cullings on the
Trans Australia Railway Line. The sequenes,
thought to he approximately 6 4 thick, overs
lies carbonaceous sandy clays and sands of the
Middle Eocene Pidinga Formation” (Fig. 2).

In a railway cutting 15.5 km west of “Mal
booma 0,8." dark olive-green clays with red
and yellow mottling 4re overlain by white-
yellow fossilileraus dolomitic Jimestones which
are fragmenta! at the top. The base of the se
quence is not exposed, No fossils were pre-
viously known an the sequence”.

The limestunes are sparsely fossiliferous with
scatiered impressions, fragments and casts of
Rivisessor sp. in a pelletal matvix, Oceasional
oogonia of a charophyte are also present, The
enviloament was lacusirine, probably similar
to and contemporaneous with that at “Billa
Kalina”

Delomitic limestone from Lake Woorang

ln March 1980. fossiliferous dolomitic
limestone (sumple 5739 RS 23) was collected
by M. C, Benbow, G, W, Kricg and P. A,
Rogers from the southern lake of Lake Woo
rong, 32 km west of Lake Phillipson (lat.
29°36 06"S, long. 134°07'54"E, COORBER
PEDY 12250000 geological map sheet), The
hard dolomitic limestone, with dolomite clasts
find occasional scattered casts and moulds of
small gastropods, is similar ta material val-
lected from near ‘Billa Kalina”, Although pre-
servation is very poor, by analogy with the
Billa Kalina and Malbooma material, the easts

and moulds cam be identified as the freshwater
gastkopodsy Rivisessor sp. and Plrysastrey redin-
eae. The material was collected too recently
for the Jweality to be included uw cither Figure
1 or Figure 2, COOBER PEDY sdjuins TAR-
COOLA on the suwth and BILLA KALINA
on the east.

Age aud correlation of the dolomitic
limestones

The age of the Btadunta Formation has
been determined by W. K, Harris on unpub-
lished palynolagieal data as Middle Miocene
(Callen & Tedford 1974, Callen 1977). ‘The
Etadunta land matluses Bariwiembrven prae-
cursor und Merecemelon llevdi occur, either
separately ar together, in unnamed Tertiary
limestones near “Deep Well” SSE of Alice
Springs and in the Carl Creek Limestone N
and SE of “Riversieigh,” Queensland.

The molluscs in the limestones in the Billa
Kalina, Malhooma and Lake Wooronu areas
ate from a freshwatec environment and do tot
provide direct correlation with the Etudonna
Formation. They are felated to one another
by the presence of Rivivessor sp. The sniall as-
semblage of Syrioplanorhis hardmani and Phy-
saxira redingae permits correlation of the dolo-
nites near “Billa Kalina” with the White Moun-
tain Formation of the Ord Basin in north-
western Australia, the Arltunga Beds, un-
named limestones of Ihe "Deep Well” area, in
the Alice Springs area, ynd the Carl Creck
Limestone, Horse Creek Formation and Brun-
ette Limestone in Queensland, All of these have
been regarded by Lloyd (1968) us of Miocene
age but nol necessarily correlates. ‘The lime:
stones at Billa Kalina and Malbnoma afte there-
fore vonsidered to be also of Miocene age. bul
not hecessarily exact correlates of the lower
part of the Btadunna Formation.

Systematic descriptions
Class GASTROPODA
Subclass PROSOBRANCTUA
Order MONOTOCARDIA

Fig. 3 (a-e) Bothrieubryon proceursur, Btadunna Vormation, Lake Palankiwrinm, (2) M273811) x 1,2,

large specimen, (b, cy M273803), showing fut intergal spiral ribbing. fd, e) Mt74403), murray
specimen with axial ribs, w x 12. (f1) Meracomelon loydi, Fladunna Formation, Lake Palankarinns
(f) M2740. apertural view. (2) M2740), umbiltest view, (h) M27410 11. apical view. (1) M2741(1),
umbilical view. (j) M2743, aperturul view. (ky M2743, apertural view (1) M2743, 1mbilical view,
(m-u) Syrieplanarbis hardinant, Billa Kaling Oreo) M2746(1), broken specimen giving natutal sec-
tion showing usymmetrical Bperlure (p, a) M2402) lawer and upper sides, fr, 4) M2746(3}
lower and upper sides, (6) 4357901), (1) M3S79(3) moulds and casty in limestone, (yw) Syrlas
planorbiy sp.. Billa Kajima tv} M3580(1), showing spiral ribbing. (w) M3SR0(2), shawing syin-
metrical aperture. (x) Plivsestra rodinvac, Billa Kalina, M358107), (¥, 2, ga) RiiLeessar specimens
an Beste (v) 6138 RS 70, Billn Kalina, Ge) 6138 RS 7), Billy Kalina. (4a) 9736 RS 48, Mal.
OMA,

MIOCENE NON-MARINE MOLLUSCS

88 N. H. LUDBROOK

Suborder TAENIOGLOSSA =
TROPODA

Superfamily RISSOACEA

Family HYDROBIIDAE

Genus RIVISESSOR Iredale, 1943

MESOGAS-

Rivisessor sp.

FIGS 3y, z, aa
Material: Numerous specimens scattered through-
out the matrix of samples 6138 RS 62-67 from 4
km NNW of “Billa Kalina”, 6138 RS 68-70 from
3.5 km N of “Billa Kalina” and 6138 RS 71-73
from 4 km N of “Billa Kalina”, casts and moulds
in matrix of samples 5736 RS 47-51 15.5 km W
of “Malbooma O.S.” and 5736 RS 52 9 km W of
“Malbooma O.S.”
Description: Shell small, smooth, thick, solid,
whorls 4 to 5, moderately tumid, suture im-
pressed. Aperture oval, entire, oblique; last
whorl 3/5 height of shell, Height 5, diameter
2.5, height of last whorl 3, height of aperture
1.5 mm. Maximum height estimated from an
imperfect natural section 7, diameter 3 mm.
Distribution: Limestone cappings at Billa
Kalina, localities 6138 RS 62-73 (BILLA
KALINA 1:250000 map sheet) and Mal-
booma, localities 5736 RS 47-52 (TAR-
COOLA 1:250 000 map sheet).

Habitat: Living species of Rivisessor com-
monly inhabit streams, lagoons and ponds.

Order BASOMMATOPHORA
Superfamily LYMNAEACEA

Family PLANORBIDAE

Genus SYRIOPLANORBIS Baker, 1945

Syrioplanorbis hardmani (Wade)
FIG 3 m-u

Planorbis hardmani McCoy. Hardman 1885: 7, 15
(nom. nud.)
Planorbis hardmani (Foord) (sic) Wade, 1924:
29, pl. 1
Planorbis hardmani Wade. Chapman 1937: 61, pl.
6, figs 1, 2
Syrioplanorbis hardmani (Wade).
1968: 141, pl. 10, figs 6-8
Material: 15 internal casts GSSA M2746; numer-
ous internal casts and moulds in matrix, M3579
(locality 6138 RS 87, 4 km N of “Billa Kalina”).
The species was described adequately by
McMichael. All specimens show the slight
asymmetry with a deeply concave upper sur-
face and shallowly concave lower surface typi-
cal of the species, Dimensions of the ten mea-
surable specimens are consistent with those of
the types from White Mountain Hills, Western
Australia and specimens from south of Herr-
mansburg in Central Australia:

McMichael

GSSA Reg. Diam. Height Diam./Height
No. (mm) (mm) ratio
M2746(1) 20.0 7.4 2.70:1
M2746(2) 18.0 7.0 2.57:1
M2746(3) 16.4 5.4 3.04:1
M2746(4) 11.4 5.0 2.28:1
M2746(5) 13.7 6.0 2.28:1
M2746(6) 12.0 5.5 2.18:1
M2746(7) 12.4 5.4 2.30:1
M2746(8) 11.9 6.0 1.98:1
M2746(9) 11.4 5.5 2.07:1
M2746(10) 9.6 4S 2.13:1
Average 13.68 5.77 2.37:1

Type locality: Trig J40, 15 km E of N of “New
Ord River’, White Mountain Hills, lat.
17°15'37"S, long. 128°57'57"E, LISSADELL
1:250 000 map sheet, Kimberley District, W.A.
White Mountain Formation, ?Miocene.

Distribution: The localities cited by McMichael
are here reinterpreted from Lloyd (1968),
Wells et al. (1970) and Playford et al. (1975):
Western Australia-White Mountain Forma-
tion, White Mountain Hills 15 km E of N of
“New Ord River’, LISSADELL 1:250 000
map sheet; Northern Territory—NT 406, un-
named formation, 6 km W of Running Waters,
42 km S of MHerrmansburg, HENBURY
1:250 000 map sheet; NT 409, unnamed for-
mation, 16 km NNE of ‘Deep Well”, 61 km
SSE of Alice Springs, RODINGA 1:250 000
map sheet; NT 417, unnamed formation, 16
km NE of Undoolya Gap, 45 km E of Alice
Springs, ALICE SPRINGS 1:250000 map
sheet; NT 422, Arltunga Beds, 3.2 km SW
of Arltunga airstrip, ALICE SPRINGS
1:250 000 map sheet; NT 423, Arltunga Beds,
Arltunga airstrip, ALICE SPRINGS 1:250 000
map sheet; AS 234, unnamed formation, 24
km SE of “Todd River” H.S., ALICE
SPRINGS 1:250000 map sheet; AS 235,
unnamed formation, 21 km ESE of
“Todd River” H.S., ALICE SPRINGS
1:250000 map sheet; Queensland — Q9,
Horse Creek Formation, 25.6 km SE of
“Springvale” H.S., SPRINGVALE 1:250 000
map sheet; BT 169, Brunette Limestone, 45.6
km N_ of “Rockhampton Downs” H.S.,
ROBINSON RIVER 1:250000 map sheet;
South Australia — 6138 RS 87, unnamed for-
mation, 4 km N of “Billa Kalina”, BILLA
KALINA 1:250000 map sheet.

The genus is recorded from the Late Oligo-
cene of Europe and the Far East, but as liv-
ing only in Lebanon and Syria. No living re-
presentatives are known from Australia.

Habitat: Freshwater.

MIOCENE NON-MARINE MOLLUSCS 89

Syrioplanorbis sp,

FIGS 3y, w
Material; Two internal casts GSSA M3580(1-2),
locality 6138 RS 87, 4 km N of “Billa Kalina’
HS.
Deseviption: Sinistral, discoidal, both upper
and lower surfaces deeply and fairly broadly
umbilicate, Whorls 44, regularly increasing,
laterally somewhat compressed and inclined to
be angulate af upper and tower curvature.
Sutures deeply impressed. Shell unknown, Sut
cast showing three conspicuous spiral ribs on
lateral surface, Aperture more or less sym-
metrical, moderately arched,

GSSA Reg. Diam, Height Diam./Height
Na. (mm) (mm) ratlea
M3580 1) 12.2 63 1.94:1
M3580(2) 10,2 5.0 2.04:1
Average 11,2 St 2.01

Ghservations: Two specimens, although not
well preserved, appear to differ fram Syrio-
planarhis hardmani in having taterally eom-
pressed Whorls, which makes the shell rela-
tively higher than §. Aardmeni, with an average
dizmeter: height catio of 2.001 in contrast with
2Z37c1 average of ten specimens of 8. Aard-
mani, eis not known whether the three spiral
nibs visible on the internal east persist as ex-
lernal feaiures.

There are not sufficient specimens. nor ate
they well enough preserved, to warrant a new
specific name.

Distribution: Locality 6138 RS 87, 4 km N of
“Billa Kalina”, BILLA KALINA |-250 000
map sheet.

Habitat; Freshwater,

Genus PHYSASTRA Tapparone-Canetri,
(883
Physastra rodingae McMichacl
FIG, 3x
fsidara, near ft. peetorosa, Etheriige in

Cameron 1901; I4

Bullinus sp. nov. Chapman 19372 63

fsodora (sic), Whitehouse 1940724

Physavtra rodingae McMichael, 1968: 146, pl.
11, figs 2-5

Material, Four poorly-preseryed casts and

moulds, mostly enthedded in hard limestone,

GSSA M3581 (14), loculity 6238 RS 87, 4

km N oof “Billa Kalina’, They appear to he

casts and external moulds of juveniles, with

about 3 whorls, of the sinistral species whieh

has 4-5 whorls in the adull,

M3581(1), an internal cast freed from the
matrix, has dimensions: height 9.0, diameter
5.3 mm, compared with a height 33.0+, dia-
meter 14.5 mm in the largest paratype.

Type lacality; NT 407, 12 km NE of “Deep
Well", 66 km SSE of Alice Springs, RODINGA
1:250 000 map sheet, Unnamed formation,

Distribution: Western Austealia-White Moun-
tain Hills, White Mountain Formation, 15 km
—B of NS of “New Ord River’, LISSADELL
1.250000 map sheet; Northern Terntory —
NT 407, unnamed formation, 12 km NE of
“Deep Weill", ROPINGA 1;250000 map
sheet; NT 424, Waite Formation. 6.4 km § of
“Alcoota” H,S,, ALCOOTA 1:250000 map
sheet; Queensland—-Q1!, Carl Creek Lime-

stone, & km N of “Riversleigh” H.S., locality

103 on LAWN HILL 1:125 000 map sheet;
QI2, Carl Creek Limestone, 1.6 kro SE of
“Riversleigh” H.S., locality 90 on CAMOO-
WEAL, 1;250000 map sheet; South Australia
— jocality 6138 RS 87, Unnamed formation,
4 km WN of “Billa Kalina” H.S,, BILLA
KALINA [-250000 map sheet, The genus
Physastra is eecatded as living in Indonesia,
Australia, New Zealand and New Caledonia.
Habitat: Freshwater,

Observation. McMichael included in the syn-
onymy Isidera, near 1, pectoresa identified by
R, Etheridye jr, cited by Caumerou (1901) and
by Whitehouse (1940) as being abundant, of
ten in crowded masses, 1n what 1s Now Known
as the Carl Creek Limestone, which overlies
Cambrian limestone near “Riversleigh" from
which Lloyd's material also caine. This syn-
omymy is accepted in the absence of any
material evidence to the contrary,

Order STYLOMMATOPHORA
Superfamily BULIMULACEA

Family BULIMULIDAB

Genus BOTHRIFMBRYON Pilsbry, 1894

Bothriembryon praccursor McMichacl
FIGS 3 a-e
Bothriembryon praccursor McMichael, 1968: 149,
pl, i, figs 7-9
Material: 34 internal casts and several exter-
nal moulds in matrix GSSA M2738 (1-5),
M2739 (1-14), M2744 (1-3), M3582 (1-4),
all from the type section, Etadunna Forma-
tion, dolomitic limestone member 2e. Lake
Palankarinna, §, Aust, (locality 68540 RS 39,
KOPPERAMANNA |:250000 map sheet).
The species is common in dolomitie lime-
stone near the base of the Etadunna Forma-

99 N. H, LIIDBROOK

tion and is here fedeseribed from internal casts
of adult specimens collected from the type
section,

Description; Shell not known, byt from the
appearance of the casis probably fairly thick,
size moderate for the genus, clongate-turbini-
form, with a moderately high spire and large
last whorl: aperture about equal in height to
spire; whorls 5, regularly increasing. Proto-
conth small but fairly high, of two whorls
With Lip immersed, adult whorls 3, slightly to
moderately inflated, suture conspicuous, Im-
bricating, Aperture subovate, outer lip gently
ufeuate, attached less than 4 way towards
adapical suture, parictal lip probably concave,
columellar lip nearly vertical, basal lip ur-
cuate; umbilical chink present. Sculpture, as
shown on internal cast, of axial [olds of
growth ridges fairly evenly spaced, about 17
on jast whorls in oblique light, some speci-
mens appear to have 2 or 3 faint and shalliw
spiral grooves on last whurl, suggesting that
there Inay be some spiral sculpture as well,
Dimensions of Jargest specimen GSSA
M2738(1) heighe 30.5, diameter 17,7 mm,
ratio height: diameter 1.72:1, average of 23
specimens height 22.4, diameter 13,5 mm,
ratio height: diameter |.66:1. Two specimens
M2738(2) and M2739¢1) are conspicunysty
narrower than «Verage, with dimensions —
M2738(2) height 25.1, diameter 14.0 mm,
ratio height: dtimeter |.80;1, M2739(1)
height 28.4, diameter 14.5 mm, fatio height:
diameter 1.96: L-.

Type localiry: Rd 21. 6 km ENE of “Deep
Well” HS., RODINGA 1:250.00) map sheet,
unnamed formation,

Distribution, Northern Territory — NT 409,
unnamed formation, 16 km NNE of “Deep
Well" H.S. 61 kun SSE of Alice Springs, RO-
DINGA 1:250 000 mip sheet; Rd 21 unnamed
formation, 6 km ENE of “Deep Well” H.S.,
RODINGA 17250000 mip sheet; South Aus-
tralia — locality 64540 RS 59, Etadunona For-
mation, Lake Palaskarinna, south of Couper
Creek, KOPPERAMANNA 1:250000 inap
sheet.

The pulmonate land snail Hothelernfevenr is
testricted to Australia, mainly che south west.
but there are represeritatives in Cerra) Aus-
tralia and Tasmania, Bethriembrvon barrett
Iredale ws commonly ound im great Qunmbers
winder bushes in coastal areas of the Nullarbor
Plain, where it survives under dry vonditious.

Habitat: Wis ikely that B, praecursor lived in
a similar enyvironinent.

Observations: MeMichael (1968) distinguished
between B. praecursor und the living B. bar-
rerli, and also the Tasmanian fossil species B,
gunnil (Sowerby), The present study supports
separating these species. The Pleistocene to
Holocene B, harretti is a larger and narrower
shell, of 173 specimens measured from Point
Sinclair, South Australia (Ludbrook 1978),
the largest was 79 mm high and 21 mm in dia-
meter, and the average 31.5 mm high, 17.7
mim in diameter, ratio height: diameter 1.78:1-

Bothriembryon guanii (Sowerby) has been
referred to in the literature as follows:
Bulinas gunnii G, B. Sowerby, 1845, in Strze-

lecki: 298, pl, 19, fig, 6 (not fig, 5}
Bullies gunnit Sowerby, Etheridge 1878; 177,

Johnston 1880: 90. Johnston L&88: 283, pl.

44, fiy, 7
Clparas guint Cunjustified emendation} G. B.

Shy sp. Harris 1897: 3

The specific name has also been attached
to @ living Tasmaniat species thought by some
authvrs to be identical with it:

Suthrieniirven gunnli var, brachysoma Pils

bry, 1900: 18, pl. 3, fig. 53
Beitriembrvin panna! Sowerby. May 1921-92;

1925: pl, 42, fig, 7

This is the speeies referred to in Iredale
(1937; 313) and May revised Macpherson
(1958, pl. 42. fiz. 7) as Tawmdnembryon tas-
manicus Pfeiffer,

Bothriembryon gunni is now represented
solely by the holotype in the British Museum
(Natural History), a internal cast embedded
in matrix aperture down. so thal complete
description is impossible. "The specimen
figured hy Johnston (1888) cannot at present
be found, and no other specimens are known.
Johnston's figure is of an clongate-turbiniform
shell with axial tibs and impressed sutures.
Although both Rulinus and Bulimus have been
widely used for genera in different families, it
may be assumed that in replacing Bulinus (a
sinistral shell, family Planorbidae) by Bulinus
(© Bulimulius) Johnston implied the position of
Bulinns gin iv the Bulimulidae, as did alsa
OG. FF blarris tt placing the holatype in Liparis
fa synonyviy of Bothriembryan), lohnston's
figire immediately invites comparison of
Aiclimus gunniit with Tasmanembryon tasmani-
cum (Pfcitfer), recorded as common on the
vast coast of ‘Tasmania, near the sea On trees

MIOCENE NON-MARINE MOLLUSCS oi

and rocks, but the extreme paucity af material
is a barrier to confirming the identity of the
fossil.

Bulimus guant was desctibed with Nelix
rasmmanlensis Erom travertine limestune quar-
ried near Hobart Town (Strzclecki 1845),
There is some uncertainty whether this was the
quarry visited by Darwin (Banks 1970) or
that at Geilston Bay, Neither quarry ig now
accessible, That in Hobart was iclentified by
Johnston and by Banks as at the western end
om Burnett Sctreet. Johnston's figured speci-
men, occurring alsa with “Helly tasmenierivis
came (rom the Geilston Travertine which was
quarricd at Geilston Bay on the northeast side
of the River Derwent 3 km north of the Tas-
man Bridge. Johnston considered this te he
the locality visited by Darwin and by Strae-
lecki, There seems to be no way of recallect-
ing the material or of confitming that the holo-
type of Bulinus gunnii did in fact come Irom
the Burnett Street quarry and net fram Geil-
ston Bay, but Geilstoy Bay secms the more
Ikely locality, In describing the lwo quarries,
MeCormick (1847) slated Uhat he Found no
traces of shells in the Hobart Town quarry, but
Hela and Bulimus Were embedded in the upper
part of the indurated limestone quarried al
Geilston Bay, Ten specimens of two species of
“Helix” from Geilston Bay were Kindly lent by
the Geology Depariment of the University of
Tasmania, but no specimens of “Aulinus” gua-
nil have been located, One of the specimens of
“Helix” lasmuaniensix, partly embedded in
matrix, is extremely like Sowerby’s holotype,

Direct comparison of Bethriembryon prac-
curser With "Bullnuy” eunnii is therefore im-
possible at present. Moreover, the stratigra-
phic position of the limestone containing
"Bullius’ gunrnil and “Helix” tasitaniensiy can
be stated only as “Tertiary” from present
knowledge, Strzelecki considered it to he of
Pliovene age. A composite section of the Ter-
liary sediments at Gerlston described hy Iahn-
ston (1888) was modified by Tedford ef al.
(1975), Johnston reconled marsupial bones
from yellow and brown maitled caleareous
elay, which, according to Tedford ef al, is in-
terbedded with the travertine, They compared
a diprotodontid from the Geilston ‘Traver-
tine with Neapeknldia from the Etadunna
Formation, of Middle Miocene age. An ap-
Parent age of 22.4 0.5 Ma was obtuined for
basalt overlying the travertine. A sample of
carbonaccons sediments collected below basalt
at Geilston Bay on the west side of the golf

links was sent by the Tasmanian Museum to
W. K, Harris, who has informed me verbally
that “the age of the microflora is ?Pliocene-
Pleistocene; it is not related to mid-Tertiary
microfloras. Widespread in Tasmania”.
ft can only be said that dating of material

fram the sequence at Geilstou Bay is confused,
and any correlation with the Etadunna For-
mation based on molluses out of the question,
Neither “Bulinus” granii nor “Helix” tasmun-
iensiy can be compared with known molluses
from the Etadunna Formation.

Superfamily HELICACEKA

Family CAMAENIDAE

Genus MERACOMELON Iredale, 1937

Meracomelou Woydi McMichael
FIGS 3 f-l
Meracomelon (loydi McMichael, 1968:

Il, figs 10-14
Material; Nine internal casts GSSA M2740-3,
M3582 (162), one external mould,

The species was desenbed by McMichael
from internal casts and so far the Etadunna
Formation has yielded only external moulds
and internal casts. Bladunna specimens are
generally smaller und higher than the holo-
type und paratypes from the Northern Terri-
tory and Queensland, They vary considerably
in their relative height:

151, pl.

GSSA Rep. Diam. Heleht Diam./Helelet
No. (mm) (mm! ralia

M2740 22,0 17,8 1.2421
M2741(1) 185 10.0 135:1
M2741(2) 12.0 6.4 1.8751
M2742(1) 140 10.4 1,342]
M2742(2) 13,5 8.6 L.S7:1
M2742(3) 19 77 1.54;1
M2743 12.0 9.0 133:1
M3S582(1) 13.5 K.7 W553)
M4582 13.3 7,0 1.9021
Average of 9

specimens 14.2 9.5 1.49:
Average of

holotype and §

paratypes 18,9 14.0 1.3521

Type locality; NT 409, 16 km NNE of "Deep
Well’; 6) km SSE of Alice Springs, RO-
DINGA 1:250000 map sheet, unnamed for-
mation,

Divtribulion; Northern Territory — NT 409,
If km NNE of "Deep Well”, NT 407, 12 km
NE of “Deep Well”, NT 408, 1.6 km N of NT
407, and Rd 21,6 km ENE of “Deep Well”, all
unnamed formation, RODINGA 1;250000
map sheet; South Australia — locality 6450
RS 59, Etadunna Formation, Lake Palan-

yd N. H. LUDBROOK

karinna, south of Cooper Creek, KOPPEKA-
MANNA 1:250 000 map sheet.

Acknowledgments

1 am indebted to Dr R, H. Tedford (Ameri-
can Museum of Natural History), and Messrs
Richard Flint, Greg Ambrose, Alistair Crooks
and Mrs Susan Daly (Geological Survey of
South Australia) tor providing both the speci-
tens used in the study and the geological de-
tails of the strata in which they occur, Dr M- R-

Banks (University of Tasmania) provided
valuable assistance in identifying the localities
from which the latd snails were collected in
Hobart and arranged for the loan of speci-
mens, The maps and sections were drawn in
the Drafting and Survey Branch, and assistafice
with photography given by the Biostratigraphy
Division, South Australian Department of
Mines and Energy, The paper is published with
the permission of the Director-General of
Mines and Energy.

References

Baker, F. C, (1945) “The molluscan tumily
Planorbidae” (Univ, Ulinels Press: Urbana),

Banks, M, &. (1970) A Darwin manuscript on
Hobart Town. Pap. Proc. &. Sac. Tas. 105, 3-
19,

Catton, Ro A. (1977) Late Camozoic environ-
ments of part of northeastern South Australis.
J. geal. Soc. Aust, %4, 151-169.

—— & Teprorn, R, H, (1976) New Late Cuino-
zoic rock units and depositional environments,
Lake Fromme area. South Australia, Trans. R.
Soc, 8, Aust. 10, 125-168.

CAMERON, W, E. (1901) Geological observations
in north-eastern Queensland, VIL Post Tertiary
limestones uf the Barklay Tableland. Annual
Progress Report of the Geological Survey for
the year [900. In, Ann, Rep. Dept Mines Qld
for 1900, 189-19),

Cuapman, F, (1937) Cherty fimestone with
Maorbis from the Mount Elder Range, Weaterts
Australia. Proc. RR. Soc. Pict, $0. 59-66, pl, V1

Exureipce, R, jr (1878) “Catalogue of Australian
fossils.” (Cambridge University Press: Cam-
bridge).

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of the Kimberley District, Western Australia
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Harris, G. F. (1897) “Catalogue of Tertiary
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(Natural History)! London),

IREDALE, T. (1937) An annotated check dist af the
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Ausr. Nat. 18, 6-55.

(1943) A basic jist of the freshwater Mol-

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Jessup, R. W. & Norris, R. M. (1971) Cainezoic
Stratigraphy of the Lake Eyre Basin and purl of
the arid region lying 10 the south. J. geal Soe.
Awst, 18, 303-331,

Tonnston, R. M, (1880) Notes on the yellow
limestone firuvertineg) of Geilxton Bay with
other fluviatile and lucasirine deposits. in Tas
otania and Australia, together wiih desenplians
of two new fossil helices. Pap. Prac. R Soc.
Tas, for 1879, 81-90,

—— {I888) “Systematic account of the scology
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Liovm A. RL (1988) Ontline of the Tertiary gear
logy of Northern Austrilia. Aull, Bur Miner,
Resour. Geol. Geophys. Aust. 80-7.

Lubarook, N. AH. (1978) Quaternary molluses of
the western part of the Eucla Masin. Bull, eeol,
Surv. Wo. Austr (25.

MeMiehAta, BD, PF, (1968) Non-marine Mollusca
from Tertiary rocks in Northern Ausiralia, Bull,
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Mar, W. L. (1921) “A check list of the Mollusca
of Tasmania.” (Govt Printer: Hobart) -

(1923) “Lflustrated index of ‘Tasmanian
shell." (Govt Printers Hebart); (1958) (Re
vised Macpherson) hid.

McCormick, R. (1847) Geology of Tasmania. Jn
1, ©, Ross “A voyage of discovery and research
in the southern ana Antarctic regions. during
the years 1839-43." vol. LL Appendix pp 391-
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VOL. 104, NUMBERS 5 & 6 28 NOVEMBER, 1980

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED
CONTENTS
Robinson, A. C. Notes on the mammals and reptiles of Pearson, Dorothee and
Greenly Islands, South Australia - - - - - - 95
Williams, D. L. G. Catalogue of Fivsiecen® Setirbrabs fossils and sites in South
Australia — - - - ~ - - - 101
Moore, P. S. Stratigraphy and depositional environments of the Billy Creek
Formation (Cambrian), east of the Flinders Ranges, South
Australia - ~ - - - - - - - ee ET,
Koste, W. & Shiel, R. J. New Rotifera from Australia —- - - - =. 133
McNamara, K. J. Heteromorph ammonites from the Albian of South Australia- 145

Barton, C. E. & McElhinny, M. W. Ages and ashes in lake floor sediment cores
from Valley Lake, Mt Gambier, South Australia — - = - 161

von der Borch, C. C., Bada, J. L. & Schwebel, D. L. Amino acid racemization
dating of Late Quaternary strandline events of the coastal plain

sequence near Robe, southeastern South Australia - - ~ 167
Glaessner, M. F. New Cretaceous and Tertiary crabs (Genitaces: Brachyura)

from Australia and New Zealand - - - - - 171
van Beurden, E. & McDonald, K..R. A new species of C jd a (Agpra:

Hylidae) from northern Queensland - = 193
Jago, J. B. & Youngs, B. C. Early tS Be pineiNiS —_ Mi SEES vane,

South Australia - - - 197
Fitzgerald, M. J. Muckera and Millbillillie—Australian achondritic meteorites - 201

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

NOTES ON THE MAMMALS AND REPTILES OF PEARSON, DOROTHEE
AND GREENLY ISLANDS, SOUTH AUSTRALIA

BY A. C. ROBINSON

Summary

Collections and observations of the mammals and reptiles of Pearson, Dorothee and Greenly Islands
off the west coast of Eyre Peninsula were made in November 1975 and 1976. Southern Bush Rats
on Pearson Island are smaller, breed earlier, and occur at a lower population density than on
Greenly Island. The difference in population density may be explained by the different stages in the
reproductive cycle on the two islands in November. Population estimates of the Pearson Island
Rock Wallaby and the introduced population of Tammar Wallabies on Greenly Island are given. An
annotated list of all reptiles recorded from the islands is given.

NOTES ON THE MAMMALS AND REPTILES OF PEARSON, DOROTHEE
AND GREENLY ISLANDS, SOUTH AUSTRALIA

by A. C. Ropinson*

Summary

Rosson, Ao C, (1980) Notes on the Mammals and reptiles of Pearson, Dorothee and
Greenly Islands, South Australia, Trans. R. See. 8, Aust, W4(5), 93-99, 28 November, I9KO
Collections and observations of (he mammals and reptiles of Pearson, Dorothee and
Greenly tslands off the west coast of Eyre Peninsula. were made in November 1975 and 1976.
Southern Bush Rats on Pearson Island ure smaller, breed earlier and occur al a lower popula-
lion density than on Greenly Island. The diflerence in population density may be expliined
hy the different stages in the reproductive cycle on the two islands in November. Population
celimates of the Peurson Island Rock Wallaby and the introduced population of Tammar
Wallabies on Greenly Island wre given, An annotated list of all reptiles recorded from. |he

islands is given,

Introduction

In November 1976, a biological survey of
Pearson [sland and Dorothee Island (Investi-
gator Group) and Greenly Island was under-
taken by A, C. Robinson, T, J, Fatchen, A.
Spiers and J, B. Cox (South Austrahan
National Parks and Wildlife Service) and 8, A,
Parker and W. Zeidler (South Australian
Museum).

Pearson Island is located at 34°48, 134°
176, Borothee tsland at 34°0'S, 134°15°E
and Greenly Island at 34°39'S, 134°45'E. Four
days and nights were spent on Pearson Island,
a day and u night on Dorethee Island and
four days and nights on Greenly Island. In
November 1975 a two day trip to Pearson
fsland was made and some small mammal
trapping and observation of the rock wallaby
population was carried out,

This paper presents observations made on
the mammals and reptiles of these islands. The
birds are discussed in Parker & Cox (1978)
while the yegetation wil] be examined in
Fatchen (in prep.). Previous observations
(Summarised here) are available from eXpedi-
tions to Pearson Island in 1914, 1922, 1923
(Proctor 1923, Wood-Joncs 1922, 1923,
1924), 1960. 1968, 1969 (Thomas & Delroy
1971) 1969 (Smyth 1971). 1973 (Gepp
19733), 1974 (Schmitt L975) and Field
Naturalists’ Society Mammal Club unpublished

* National Parks & Wildlife Service, Box 1782,
G.P.0.. Adelaide, South Australia S001,

L Ciepp, B. (1973) The Social Organisation of the
Pearson Island Rock Wallaby (Perragale peni-
cillata pearson’). B.Sc. Hons. thesis. Zoology
Department, University of Adelaide,

observation, from Dorcthee Island im 1969
(Smyth 1971); and from Greenly [sland in
1947 (Finlayson 1948a, b; Mitchell & Behrndt
1949),

The three islands are all essentially granite
based islands with very little of their original
limestone capping remaining, the geomorpho-
logy of Pearson and Dorothee Islands having
been described hy Twidale (1971). Mitchell &
Behrndt (1949) gave a general description of
Greenly Island, while the geology of the
Greenly Island basement rock ix described by
Webb & Thomson (1977).

The vegetation of the three islands is similar,
that of Pearson and Dorothee Islands being
deseribed by Osborn (1923), Specht (1969)
and Symon (1971) while that of Greenly was
described by Fintayson (1948. a, b), Mitchell
& Behrndt (1949) and Cleland (1950).

In addition ta the general biological survey
catricd out on the islands an attempt was
made to determine the effect of fire on the flora
and fauna. An extensive area on the southern
part of the main Pearson Island was burnt in
a fire which is belicved to have started from a
lightning strike on the island peak jin early
April, 1975 (D. Steen pers. comm.). This fire
must have been extremely hot, and total de-
struction of the above ground parts of the
vegetation has occurred over large arcas, The
extent of the fire is shown in Fig. 1.

On Greenly Island a number of fires were
lit by tuna fishermen on @ February, 1974.
The wreas burnt are shown on Fig, 2 from
unpublished records of the fire, prepared by
members of the N.P.A&W.S. who visited the
island on 24 February, 1974,

o4 A, ©, ROBINSON

Fig, 1. Pearson Island showing location of trap-
ping areas, transect courts and walluhy
populations.

4

Methods

An arbitrary grid system with 250 m grid
intervals Was established over the three islands
to be sampled (Figs 1, 2), Collecting and
observation efforts sampled as many of the
grid intersections as possible to obtain good
coverage of the range of habitats on the
islands. Line transects at night using a spot-
light were walked on both central and south
Pearson, and during the day on Greenly Island
to try and estimate the wallaby populations.
Small mammals were sampled by systematic
trapping using Elliot and Sherman aluminium
box traps set either on a grid pattern or in
lines. All Rattus /uscipes trapped were
weighed and individually marked by toe. clip-

ping. The reproductive condition noted by
recording if testes were descended or un-
descended in males and if vaginae were per-
forate or imperforate and if nipples were
enlarged by suckling in females.

(a) Pearson Island: On 2 November, 1975,
two traplilves were set above the northern
bay (Fig. 1). Each had 10 sites at 20 m
intervals with three traps per site. On 24—
25 November, 1976, two traplines were
set on the centre section (Fig, 1). Back
had 12 sites at 20 m intervals with two
traps per site, At the same time a grid was
established on the main island with 80
sites at 25 m intervals and two traps
per site. This grid was positioned to
sample as wide a range of vegetation ax
possible from the low Afriplex shrubland
near the coast through the closed Mela-
leuca halmaturorum scrub along the creek
and into the low Casuarina woodland on
the slopes of the island peak, In addi:
tion the grid sampled as equally as
possible areas burnt in the 1975 fire and
unburnt areas. There were 34 sites in the
burnt area and 46 unburnt sites.

Derothee Island: On 27 November, 1976,
two traplines were set in a WNW and
SE direction from the conservation park
sign on the central eastern shore, Each
had 20 sites at 20 m intervals with two
traps per site.

(b)

Greenly Island; On 29=30 November,
1976, a trapline was sel above the
anchorage (Fig, 2,), It had 12 sites at 20
m intervals with two traps per site, and a
grid was established on the northern
slopes of the main island (Pig, 3) with
48 sites al 25 m intervals and two traps
per site, This grid was positioned to
sample as wide a range of vegetation as
possible from the Pow tussock grassland
on the lower slopes to the low Casuarina
woodland on the upper parts of the island.
The grid sampled areas of grassland burnt
in the 1974 fires. It was not possible to
distinguish boundaries between burnt and
and unburnt sites due to the degree of
regeneration,

All species of mammats and reptiles re-
corded from the islands are discussed. South
Australian Museum registration numbers are
recorded of specimens collected,

MAMMALS & REPTILES OF PEARSON, DOROTHEE & GREENLY ISLANDS 95

Anthon:
Peninsula
\Wandra
Point

_ Nor'West
Crevasse

a! be West rH + ~ Fes depbensyery 6* ber ard |
Peak tm t
a8 . imsppecl 24 Fret oT
|
Comservatvon park Sryr |_|
1°] 400 meres
=
| J
a — + { —— _

ae ae

Tragptinas. iret rect

Wallaby Pour? nensents om

“Eagle de |
Poak
Varn

The Cauldron

yeecwean Mil

MAIN
| ISLAND

_ + 4

yomA

—— abe

Pe eter —

Fipute 2, Greenly Island showing location of trapping areas and wallaby count transects.

Results

MAMMALS
Family MACROPODIDAE

Petrogale lateralis Gould. Pearson Island
Rock Wallaby. This species was collected in
1920 by Wood Jones from Pearson Island
and Thomas (1922) described it as a new
species (P. pearsoni). Since then there has
been considerable confusion about its relation-
ship with other rock wallabies and this is dis-
cussed by Thomas & Delroy (1971). Recent
chromosome and electrophoretic studies (G,
Sharman pers. comm,), place the Pearson
Island animals with western and central! Aus-
tralian populations of FP. lateralis (Poole,
1979). P. lateralis also occurred in the far
north-west of the South Australian mainland
but may now be extinct. Early visitors to
Pearson Island mentioned that the rock walla-
bies occurred only on the northern section
(Wood Jones 1923), This section is separated
from the central and southern section by a
sand spit that dries at low tide, The 1960
expedition also noted the absence of wallabies

on the middle and south sections of the island
and they found no trace of skeletons or of
occupation of several caves on the southern
section. On the 1960 expedition several wal-
jabies Were caught on the northern section and
transferred to a camp on the middle section.
Four females, one male and one animal of un-
known sex escaped. It appears certain that the
present rock Wallaby population on the middle
and southern sections of Pearson Island is
descended from these six escapees.

Thomas & Delroy (1971) estimated that
there were 50-150 wallabies on the middle and
southern sections in 1968 and showed that this
1960-1968 population increase was theoreti-
cally possible from the original six animals, In
1976, transect counts of the middle section
in the evening (24 November) and at night
by spotlight (25 November) revealed 64 and
40 individuals respectively, while a spotlight
transect on the southern section (25 Novem-
ber) revealed 92 individuals. Thus the total
population of the middle and southern sections
is at least 150, The distribution of this popu-
lation is shown in Fig. 1.

on A, C. ROBINSON

{t is much more difficult to estimate the
wallaby population on the northern section of
Pearson Island. The population ts fragmented
(Fig. 1) being confined to areas with suitable
rook crevices and caves and the dense veyeta-
tion makes sighting difficult. Thomas & Delroy
(1971) provide two estimates for ihe northern
section; 1960—500 to 600 and 1968 ib excess
of 500, In November 1975, the author walked
over the whole of the jorthern section and
counted 132 individuals: this was undoubtedly
too law and the total population is probably
iwice this size between 250 and 300, No
estimmles of the population on the northern
section were uttemped in 1976,

The diet of the wallabies was observed to
include Lepidinm faliosum, Atriplex ¢inerea,
A, palidosa, Rhagedia baccata, Enchylaena
tomentosa, Olearia runiulova, Carpohratas
rossii ind Disphyma australe. Og the parthern
section, even around the major colonics of
wallabies there was lime evidence of grazing
of the vegetation. On the middle and southern
sections however, large areas of vegetation
were obviously very heavily grazed. This was
particularly the case with E turmentose, while
D. australe and A, paludosa were also being
affected.

Macropus cugenii (Desmarest) Tammae,
SAM, M9786, An unknown number of Tam-
mars were introdiiced to Greenly Island frarn
Kangaroo Island in about 1905 to act as an
emergency food supply for possible castaways
(Mitchell & Behrndt 1949). They now occur
an both the main central und small south-
Western section of the island but are apparently
absent from the northern section. They spend
the daytime tn the dense teatree thickets on
the south side of the main ridge Of the istund
and also occupy some of the gullies on the
northern slopes of the island, They are ex-
tremely difficult to observe aud the two transect
counts (Fig. 2) resulted in the sighting of
four and 14 individuals, Il is proheble that
the total population is about SQ individuals.

The stability of (he Tammar population on
Greenly Island is hot knowl, bul a eormpari-
son Of the Vegetation of the main island with
the northern section where wallalles are absent
reveals that they have had a sabstantial |mpact
on the island vegetation, Phere his heen signi-
ficant reductions in both species diversity and
ground cover and ih appears that the present
veneration of ihe main section of Creenty

Islan! is a direct result of severe ovet-grazing
by the introduced Tammar population over the
last 70 years.

Family MURiDAE

Ralins Juscipes (Waterhouse). Southern
Bush Rat SAM M9787-91, This species is
comma on both Greenly and Pearson Islands
ful appears absent From Dorothee Island, The
pupulations on both islands Were probably
derived from a population distributed across
most of Southern Australia during the last
ice age When both the islands were connected
to the mainland, Sehmitt (1978) and Schmitt
& White (1979) haVe estimated thal Pearson
and Greenly Islands have been isolated from
the mainland for 14000 years. They have
emphasised the importance of genetic drift
in producing the differences between the island
populations that they studied. The grid trap-
ping carried out on this expedition provides
an opporiunity to obtain additional compara-
tive data, Accordingly, the two islands are
discussed separately below,

Pearson Island
Schmitt (1975) has demonstrated that sig-
nificant genetic differences occur between the

Bush Rat populations on the northern and

southern sections of the island, The grid was

located on the northern section (Fig, 1) and
the following discussion refers to this popula»
fion only,

(a) Pepulation density: Twenty-six individuals

were captured on the 50000 m* of the
grid, Therefore, ignoring boundary effects,
the overall density was 5,2 rats/ha.
As approximately half the grid was in
the area burnt by the 1975 fire it is appra-
priate to examine the differences in popu-
jallon density i the burnt and unburne
areas: Burnt sites 3.8 rats/ha; unburne
sites 6,3 rats/ha.

{b) Ser ratio 1% dc + 13 29

(c) Body weighes do 40.4 » (30-55, mn 13))

9? 49.1 gy (30-70, n = 13)

Reproductive status: All the males cap-

tured were sub-adult with undescended

testes. Five of the females still had im-

perforate yaginas while those with per-

forale vaginac showed no indication that
their nipples had been suckled, This sug-
gests that there had been a spring breed-
ing season with a very high mortality of

(dd)

MAMMALS & REPTILES OF PEARSON, DOROTHEE & GREENLY ISLANDS 97

both male and female adults to result in
this largely immature population im
November.

Greenly Island

Although the part of the island covered by
the trapping grid had been burnt in 1974, it
was impossible to distinguish burnt and un-
burnt areas, so the whole grid was treated
as a single area.

(a) Population density: Thirty-seven in-
dividuals were captured on the 30 000 m*
of the grid, Therefore, ignoring boun-
dary effects the overall density was 14.3
rats/ha,

(b) Sex ratio: 18 des 19 99

(c) Body weight: dd 83.8 g (35-130, n =

18); 22 70.7 g (40-100, n — 19)

Reproduetive status: Seven of the males

were sub-adult but although all the re-

maining males were of adult weight, only
two still had descended testes, OF the
females three still had imperforate vaginae
while four of those with perforate vaginae
had developed nipples, indicating a recent
cessation of the feeding of the young.

This suggests that the breeding season

had just ended and that at this stage in

the population cycle there had heen a high
survival of both males and females from
the parental generation.

A comparison of the Bush Rat populations
of Pearson and Greenly Island indicates that
the Greenly Island population occurs al a
significantly higher density, even considering
the population density on the unburnt portion

(d)

of the Pearson Island afid, In addition the
Greenly Island animals were much heayier
and larger than the Pearson Island animals.
Finally it appears that at least in 1976 there
was a spring breeding season with a possibility
of breeding continuing into early November
on Greenly Island.

Additional comparative data on Bush Rat
population densities oa South Australian off-
shore islands are shown in Tuble 1. The figures
are given as trapping percentages and with
the exception of this present study are based
on (rap lines rather than grid trapping so no
absolute densities can be obtained.

The differences in population densities be-
tween Pearson and Greenly Islands shown
by the grid studies are also apparent in the
line trapping. Schmitt's line trapping figures
obtained in February and March however
showed similar trapping success rates oa hoth
islands and it is possible that the differences
demonstrated in the grid study in November
may be due simply to greater survival of adults
from the breeding season on Greenly Island
than on Pearson Island, The high trapping
success on other South Australian offshore
islands indicates that high population densities
of Bush Rats are a feature common to all
of these islands,

Family OTARIIDAE
Neophoca cinerea (Peron & Lesueur) Aus
tralian Sealion. This species was present on
each of the islands visited.
Pearson island: 20-30 individuals, 4-6 mature
bulls, the major concentration wis On the
beach on the centre section of the island

TABLe 1. Comparison of results of irapping studies of Rattus luscipes en South Australian offshore

ishirils
Trapping
Type of Success
Island Month Trapping 5a Source
Narth Pearson [. Noy line 5 This Study
North Pearson [. Feb line Ai} Schmitt (pers. commt.)
North Pearson I. Nov erid 8 This Study
South Pearson 1, Feb line i Schmitt (pers, comm.)
South Pearson I, wav line 19 This Study
Greenly }. Mar line 22 Schmult (pers, comm),
Greenly I. Nov line su This Study
Greenly 1. Nov grid 21 This Study
Waldegrave L. Feb line 68 Schmitt (pers. comm.)
Williams I Apr line $2 Schmili (pers, comm.)
North Gambier 1. Apr Tine 52 Schmitt (pers. conim.)
Dog I. Jun line 73 Schmitt (pers, comm.)
Goat |, Jun line 62 Schmitt (pers. comm. }

ee __. Ee

98 A. C. ROBINSON

Dorothee Island: 30 individuals, 7 mature
bulls, the major concentration was on a sloping
tock shelf on the north side of the central cre-
vasse and on the saltbush covered slopes
around a wallowing area.

Greenly Island: 30 individuals, 4 mature bulls,
the major concentration was on a sloping
granite shelf on the north face of the main
island.

On all islands there were immature animals
1.5-2 m long associated with females, and
some were observed to be suckling although
they were often abandoned by their mothers
on the higher parts of the island.

Arctocephalus forsteri (Lesson). New Zea-
land Fur Seal. This species was found only on
Dorothee and Greenly Islands.

Dorothee Island: 16 individuals, 1 mature bull,
concentrated in cracks and holes in the rock
around the central crevasse.

Greenly Island: 40 individuals, 4 mature bulls
with the major concentration on the sloping
granite shelf on the north face of the main
island near the blowhole. There was another
small group in the south crevasse.

On both islands there were some immature
animals 1-1.5 m long still associated with their
mothers.

REPTILES

Family GEKKONIDAE

Phyllodactylus marmoratus (Gray). Mar-
bled Gecko SAM, R15800A-—D, R15802, A,
B, R15807 A-D, R15809, R15815 A, B.
Previously reported by Proctor (1923),
Mitchell & Behrndt (1949) and Smyth (1971).
Found on all three islands. Common in areas
of limestone capping but also under exfoliat-
ing slabs of granite. Two clutches of eggs
found on Pearson Island under a large slab
of granite contained 14 and six eggs respec-
tively. As this species lays only two eggs at a
time it would appear that communal laying
occurred at favoured sites.

Underwoodisaurus millii (Borg). Although
not recorded in his paper, Smyth deposited
specimens of this species from Pearson Island
in the S.A. Museum. (R10237 A—B).

Family PyGoropIDAE

Aprasia striolata (Lutken), Although not
recorded in his paper, Smyth deposited a speci-
men of this species from Pearson Island in the
S.A. Museum. (R10232).

Family AGAMIDAE

Amphibolurus fionni Proctor. Peninsula
Dragon SAM R15801, R15803-6, R15820
A-B. Only found on Pearson and Dorothee
Islands and previously reported by Proctor
(1923) and Smyth (1971). Closely associated
with granite out-crops. Further details of its
offshore island distribution are discussed by
Houston (1974).

Family ScrincipAE

Egernia multiscutata Mitchell & Behrndt
SAM R15182 A-B, R15813 A-—B, R15814.
Recorded only from Greenly Island (Mitchell
& Behrndt 1949). It was recorded only from
Greenly Island on this occasion and found
to be abundant in the Casuarina woodland,
living in burrows beneath fallen logs and rocks.
Also trapped in the Poa _ grassland but
appeared much less common in this habitat.

Hemiergis peronii (Fitzinger) SAM R15808
A-C, R15810 A-—B, R15811 A-B, R15816 A,
B. Found on all three islands, previously re-
ported by Proctor (1923), Mitchell & Behrndt
(1949) and Smyth (1971). It was common in
loose soil and accumulated organic matter at
the base of plants and under rocks and fallen
timber.

Leiolopisma_ entrecasteauxii (Duméril &
Bibron). A single specimen was collected from
Pearson Island in 1923 (Proctor 1923). It has
not been collected there since.

Lerista frosti (Zietz). Not found by the
present expedition but reported to be common
on Pearson and Dorothee Island (Smyth
1971), while a single specimen (recorded as
Rhodona tetradactyla) was taken from the peak
of the main section of Greenly Island (Mitchell
& Behrndt 1949).

Lerista picturata’ (Fry). Although not
recorded in his paper, Smyth deposited a speci-
men of this species from Pearson Island in the
S.A. Museum (R10235).

Menetia greyii (Gray). Recorded on Green-
ly Island by Mitchell & Behrndt (1949) but
not seen or collected in 1976,

Morethia obscura (Storr). SAM R15819.
Small fast moving skinks probably of this
species were seen on Greenly Island and one
specimen was collected from Pearson Island.
It has been recorded from Pearson (Smyth
1971) and Greenly Islands (Mitchell &
Behrndt 1949), as M. lineoocellata, but not
from Dorothee Island.

MAMMALS & REPTILES OF PEARSON, DOROTHER. & GREENLY ISLANDS 9

References

Criecanp, J. B. (1950) Patina and Flora of the
Greenly Islands. Part 11 Flora. Rec. S. Aust.
Mus, 9, 349-351.

Fintayson, H. H. (1948a) Greenly Island, South
Australia. §, Aust. Orn. 18, 72-73.

(1948b) Greenly Island, South Australia.
Walkabout. 14, 35-38.

Houston, T. F. (1974) Revision of the Amphi-
holurus decresii complex (Lacertilia: Aga-
midae) of South Australia. Trans. R. Soc. S.
Aust. 98, 49-60.

Mrrciett, F. J. & Benrnot, A. C. (1949)
Fauna and Flora of the Greenly Islands Part
1. Introductory narrative and yertebrate fauna.
Ree. 8S, Aust. Mus. 9, 167-179.

Osnorn, T. G. B. (1923) The Flora and Fauna
of Nuyts Archipelago and the Investigator
Group No. 8 The Ecology of Pearson Islands.
Trans, R. Sac. 8. Aust. 47, 97-118.

Parker, S. A. & Cox, J. B. (1978) Notes on the
birds of Pearson, Dorothee and Greenly Islands,
South Australia. ibid. 102, 191-202.

Pooie, W. E. (1979) The Status of the Australian
Macropodidue. In M. J. Tyler, (Ed.) “The
Stutus of Endangered Australasian Wildlife”,

. 13-27. (Royal Zoological Society of 5.
Aust., Adelaide).

Proctor, J. B. (1923) The Flora and Fauna of
Nuyts Archipelago and the Investigator Group
No. 5—The Lizards. Trans. R. Soc. 8S. Aust.
47, 79-81.

Russett, T. C. (1973) Australia Pilot Vol, L
South Coast from Cape Leeuwin to Green
Point. 6th Ed. Hydrographer for the Navy.

Scumitr, L. H. (1975) Genetic evidence for the
existence of two scparnte populations of Rattus
fuseipes greyii on Pearson Island, South
Australia, Trans. R. Soc. S. Aust, 99, 35-38,

—— (1978) Genetic variation in isolated popula-
tions of the Australian Bush Rat Rattus fuscipes.
Evolution 32, 1-14.

& Wuire. R, J. (1979) A comparison of
metric and protein variation in the Australian
Bush Rat Rattus fuscipes greyii. Aust, J, Zool.
27, 547-559.

SmytTH, M. (1971) Pearson Island Expedition
1969-5. Reptiles. Trans. R. Soc. S, Aust. 95,
147-148.

Specut, R. L. (1969) The vegetation of Pearson
Island: A te-examination—February 1960, (bid,
93, 143-152.

Symon, D. C. (1971) Pearson Island Expedition
1969-3, Contributions to the land flora. /hid,
95, 131-142.

Tuomas, O. (1922) A new Rock Wallaby (Perro-
gale) from the Islands off South Australia, Anu.
Mag, nat, Hist., Ser, 9. 95, 681-683.

Tuomas, I. M. & Derroy, L. B. (1971) Pearson
Island Expedition 1969—4, The Pearson Island
Wallaby. Trans. R. Soc. S. Aust. 95, 143-145,

Twipace, C. R. (1971) Pearson Island Expedition
1969-2. Geomorphology. ibid. 95, 123-130.

Wenn, A. W. & THomson, B, P, (1977) Archaean
basement rocks in the Gawler Craton, South
Australia. Search 8, 34-36.

Woop-Jones, F. (1922) The Flora and Fauna of
Nuyts Archipelago and the Investigator Group
No, 2. The Monodelphian Mammals. Trans, R.
Sac, S. Aust, 46, 181-193.

(1923) The Flora and Fauna of Nuyts

Archipelago and the Investigator Group No. 6.

The Didelphian Mammals. ihid, 47, 82-94,

(1924) The Flora and Fauna_of Nuyts

Archipelago and the Investigator Group No.

15. The Pearson Island Rat and the Flinders

Island Wallaby. ibid, 48, 10-14,

CATALOGUE OF PLEISTOCENE VERTEBRATE FOSSILS AND SITES IN
SOUTH AUSTRALIA

BY D. L. G. WILLIAMS

Summary

The Pleistocene vertebrate fossil sites of South Australia are listed, summarising fossil assemblages
and depositional environments. References to the literature are provided. A list of SAM specimens
is available.

CATALOGUE OF PLEISTOCENE VERTEBRATE FOSSILS AND SITES
IN SOUTH AUSTRALIA

by D, L. G, Witciams*

Summary
Wuaiams, D. L. G. (1980) Catalogue of Pleistocene vertebrate fossils und sites in South
Australia. Tran, KR. Sov. 8. Aust, 104(5), 101-115, 28 Novernber, 1980.
The Pleistocene vertebrate fossil sites of South Australia are listed, summarising fossil
assemblages and depositional environments. References to the literature are provided, A list

of SAM specimens is available.

Introduction

The Catalogue originated as part of a Ph.D,
research project involving Lute Pleistocene los-
sil vertebrates and palacoclimates in the Flin-
ders Ranges area of South Australia, Informa-
tion from museuni records, published work and
the author's own field observations is sum-
marised but discyssion is kept to a minimum,

Sites are listed in numerical order, based an
geographic proximity. An alphabetical index is
provided, Names contorm as far as possible to
the Australian 17250000 Gazelleer (Division
of National Mapping, Dept ef Minerals &
Energy 1975). Referred mup sheets, e.g.
ORROROO, are in the 1:250 000 series, Cave
sites dre numbered according to Lewis (1976),
Sites in the .. Eyre region are distinguished by
numbers (c.g. V5772) assigned by the UCMP,
A key to all sites is given in Figure 1.

lnstitulions housing relevant fossil collections
are abbreviated’ SAM — South Australian
Museum, FUSA—Flinders University of South
Australia, AUGM—Adelaide University Geo-
logy Museum. SADME— South Australian
Department of Mines and Energy, NMVY—
Nalional Museum of Victoria. AM—Austra-
lian Museum, AMNEL—American Museum ol
Natural History. UCMP—University of Cali-
fornia Museum of Paleontology. SI—Smiih-
sonian Institution, BMNH—British Muscum
(Natural History), UM—Hunterian Museum.

In presenting lists of previously published
assemblages, the identifications of other authors
are quoted directly, although in some cases
taxa do not conform to current usage, Pub-
lished lists are avatlable for sites 1-3, 16, 47,
48, 54, 55, 63, G41. 64e. Other assemblages
have been mentioned in the literature, bul were
not necessarily identified formally. These data
reter to sites ¥, 18, 20, 31, 40, 42, 52, 53, 5h,

* School of Biological Sciences, Flinders WUniver-
sity of South Australia, Bedford Park, S. Aust
5042.

57, 60, The majority of sites, however, are
unstudied, Much of the material has been exa-
mined by the author, but not in detail, Tdentifi-
cations rely on museum catalogues and collec-
tions, Sites involved are 4-8, 10-15, 21-24,
28, 32, 38, 41, 43, 45, 49-51, 57-59, G1. 62,
64, 46-68. The remainder of the sites full into
two categories. The first includes sites under
detailed study by the author; 17, 19, 22, 25-27,
29, 30, 33-37, 39, 40, The second includes
assemblages personally communicated by other
workers: sites 44, 46, 641, Specimen numbers
are not given here, but a list of SAM catalogue
numbers, arranged geographically, is available
from the author.

The Catalogue should not be used directly
as a source of biogeographic information, Until
more is known of the ages of the deposits, und
the bias affecting the fossil assemblages, such a
use would be premature, The Catalague is in-
tended os a summary, which will facilitate
access to museum collections, and draw atten-
tion to deposits which may have been over-
looked,

Catalowue

1. WEEKES CAVE (N15)
(COOMPANA),

Nullarbor Plain, near Koonalda Statin, SAM,
BIRDS: Threskiornithidse; Plarihus flavipes.
Accipitridae; Accipiter fasciatus.

Falconidae; Falco cenchroides.

Turnicidac; Turnix sp,

Sylviidae; Cinclorhamphus cruralis, C_ mathewst.
Estrilidae; Poephila exnata.

Artamidae; Artumus leucorlivnetus-

yan Tets (1974b).

2, CALCA 33°02'S, 134°22'E (ELLISTON)

SE of Streaky Bay, near Baird Bay. Red sandy
sediments 5-7 m deep in well excavation. SAM,
MARSUPLALS; Macropodidae; Srhenisus sp,
Pracaptodpn xolialt,

Merrilees & Ride (1965).

35°31°S, 1IYSSE

102

OVewOonf Gry —

KEY

WEEKES CAVE
CALCA
BROTHERS ISLANDS
KYANCUTTA
PANDIE PANDIE
GOYDER'S LAGOON
WARBURTON RIVER
INNAMINCKA
COOTANOORINA
LAKE KANUNKA
COOPER CREEK
LAKE PALANKARINNA
HERGOTT SPRINGS
WELCOME SPRINGS
MURNPEOWIE
LAKE CALLABONNA
BILLEROO GREEK
PERNATTY LAGOON
HOOKINA CREEK
MOUNT EYRE
WILLOCHRA CREEK
BOOLCUNDA CREEK
BUCKALOWIE CAVES
TEETULPA
DEMPSEY'S LAKE
ARNOLDS BANK
HILLPARA CREEK
NECTAR BROOK
PEKINA CREEK
BLACK ROCK GRAVEL PIT
MANUNDA CREEK
PORT P[RIE GRAVEL PIT
WAUPUNYAH CREEK
ULOOLOO
COLLINSVILLE
NEWIKIE CREEK
BALDINA CREEK
BUTE
BURRA CREEK
BUNDEY
MORGAN
CHOWILLA
CURRAMULKA
LAKE FOWLER
PONDALOWIE BAY
ROCKY RIVER
KELLY HILL. CAVES
EMU CAVE; SETON ; MT
HOG BAY RIVER
KAPUNDA
TWO WELLS
GAWLER
YATALA
FROMM’S LANDING
DEVON DOWNS
CHUCKA BEND
ADELAIDE AREA:
BROMPTON
CROYDON
FINDON
KIRKALDY PIT
THEBARTON
HALLETT COVE
ECHUNGA
YANKALILLA
SALT CREEK

Fig, 1. Distribution of Pleistocene vertebrate sites in South Australia.

—

D. L. G. WILLIAMS

South

Australia

TAYLOR

GOOLWA
BLACKFORD DRAIN

HENSCHKE’S BONE DIG usu

JAMES’ QUARRY
NARACOORTE CAVES
PENOLA
TANTANOOLA AREA:
GLENCOE

GREEN WATERHOLE
MILLICENT

MT BURR CAVE
TANTANOOLA CAVE
MOUNT GAMBIER
TANKSTAND CAVE

134

°

i236

PLEISTOCENE VERTEBRATE FOSSILS Ls

3. BROTHERS
(LINCOLN),
Western end of western island, in Coffin Bay
Eroded gave mined for guano for a few yeurs
from 1902. SAM.
MARSUPIALS: Macropudidae;
Sthenurus ef, maddocki,
EUTHERIANS; Otariidae; Aretocephalus sp.
KIRDS’ Dromornithidae; Genyornis newtont,
Unidentified small bird banes.

Jack (1919), Johns (1966), Rich (1979),

ISLANDS 34°35°S, 135°20E

Macropus ap,

4, KVANCUTTA 33° 08'S, 135°33°E (KRIMBA),
Central Eyre Peninsula. Bone fragments sod teeth
labelled “Kyaneutta N.S\W". Specimens possibly
collected in N.S.W., and acquired from the defunct
Kyaneutta Museum. (T. HO Rich pers, comm.
1979). NMYV,

MARSUPIALS: Diprotodontidac; Diprotndon sp,

5, PANDIE PANDIE 26°08'S, 139°23’E (PAN-
DIE PANDIB).

Foy NE. of South Australia. Ineisor and bone frug-
ments. SAM.

MARSUPLALS: Diprotodontidae; Dipretedan sp.

f. GOYDER'S LAGOON 27°0I'S, 138°54'E
(GASON),

Diumuntina River. Bone fragments. SAM,
MARSUPIALS: Diprotodontidae; ?Diprofodon
Sp.

7. WARBURTON RIVER, A number of siles be-
iween L, Eyre and the Birdsville Track. Fossils
(rom Katipiri Sands equivalent. SAM, HM,
UCMP. Note: Lists of fossil assemblages derived
from UCMP collection,

a. Cassidy Locality (V5539) 27°48'S, 138° 12'E
(GASON),

MARSUPTALS: Vombatidacy Phascolonus cf.
Rigas.

Dipratadontidae: Dipretodon sp.

Macropodidae; ?Onyelogalea sp. Pratemnadan

cl, brehas, Srhenurus ef, oecidentalin.
RIRDS: Dromorniihidue.
REPTILES: Varanidue; Mevalania sp,

bh. New Kalamuring (V7205%) 27°44'S, 138°
17°F (GASON),
MARSUPIALS:
vivas,
Diprotedontidac: Dipraradon cf, eptatum,
Muacropodidae; Macropus sp.. Sthenurus sp. Pra-
cepieden ch. raphe.
REPTILES; Varanidae; Meeulunia sp,
Crocodilians
PISH: Unidentificd material.

¢. Marens Locality (V5569) 27°53'S, [37°50 E
(NOOLYEANA)
MARSUPIALS:
laniarius
Diprotodontidae; *Zygomalurus sp,
Macropacidae.

Vormbatidae: Phascolonuy ct.

Dasyuridae; Sareaphilus — cf.

d. Greeo Biufl Loeabity (V5775) 27°53°S, 1377
36'R (NOOLYEANA),
MARSUPIALS: Vombauduer
wivas.

Dipratodontidae; Diproteden cf. oplatum, ZvKe-
mialurus sp.

Macropodidac; Matropus sp., Osphranter sp.,
?Wallabia sp., Protemnodon cf anak, Sthenurts
sp.. 5S. cf. orientally, S. cf. pales, 8. cf, rindalei,
Pracaptodon cf, goliah, P. cf. rapha.

BIRDS: Anhingiaes Anhinga sp,
Phalacrocoracidac; Phialacracorax carbo.
REPTILES: Varanidue. Mevalania sp.

Chelonians and Crocodiljans,

FISH; Siluriformes and Dipnoy.

ec, Lookout Locality (V5776) 27°52'8, 147°55 E
(NOOLYEANA),
MARSUPIALS:
gigas,
Diprotodontidue: Pipraredon cf. aptaium
Macropodidae; Mucropus sp. Sthenurus cf ander-
sont, Pracapladon ef, raph,

BIRDS; Phalacrocoracidue; Phalacrocerax varity.
REPTILES: ?Varanidae..

Chelonians and Croeadilians.

FISH; Siluriformes and Dipnoi.

F. Punkrakadarinna Soakage (V5777) 27°47'S,
137°49'E (NOOLYEANA).
MARSUPIALS: Diprotodontidae: Diprotodan cf.
aptatum.
Tate (1886), Zietz (1899), Etheridge (1894), Sti-
ling (1913). Howchin (1930), Stirton ef al.
(1961), Pledge (1973), Hecht (1975), R. H_'Ted-
ford (pers. comm. 1980).

8. INNAMINCKA 27°45'5,
MINCKA),

NE of South Australia, SAML
REPTILES: Varanidse; Megalania prisca.

9. COOTANGORINA 28°10'S, 135°18'E (WAR-
RINA).
NW of L, Eyre, near The Peake. Jaw found at
depth of “26 to 30 feet’ in well excavation, “al
the head of one of the mound springs” (Chandler
1882).

Phascolonus —cF.

Vombatidae; Phasenlenus cf.

40°44 BE (INNA-

MARSUPLALS: Diprotodontidae; Dipretodun
ansthalix.

Chandler (1882), Brown (1894), Howehin
(1930),

lia LAKE KANUNKA = 28°23'S, [38° 18'F
(KOPPERAMANNA)

Fast of L. Eyre. Kanunka Fauna in Raupin chun-
nel sands and floodplain deposits, ?Rarly Pleisto-
cence. SAM, NMY, AMNH, Si, UCMP V5772Z and
VS5773.

MARSUPTALS: Dasyuridie.

Vombatidaes Pluiscolonus sp., cf. Vembarus ar
Lasierlinus sp.
Thylacoleonidae,
Diprotodontidac, ef,
PaluUrus sp.

Diprotadan sp., ch Zypo-

104 DL. G, WILLIAMS

Macropodidac; Aetrongla sp,, Matrapus spp. Os-
phranter sp., Lagorchestes sp., cl. Wallabia sp,, ef.
Prionwtemmnus sp., Tropasodon kenili, Protentie-
don sp. Sthenurus sp,

EUTHERIANS: Muridae,

BIRDS; Dromornithidae; Genwornis sp,
Phalacrocoracidae; Phalacrocorax spp.

Anatidae; Anas sp., Cyeruy sp.

REPTILES: Varanidae,

Unidentified Chelonians and Crocod)lians.

FISH: Ceratodontidae; Epiceratodus sp.
CRUSTACEANS; Decapads.

Debney (1882), Tate (1886), Stirton ef al.
(1961), Ro H. Tedford (pers. comm. 1980),

0b, MURRAPATERINNA 28°26'S, 138°31'E
(KOPPERAMANNAI.

Between L. Kanunka and Mulka. Fossils from
bore [S m deep, 28 Avy. 1921, SAM Director.
Waite, identified Crocodilian, Lunefish, and
Macropodid teeth and bones.

Also, NMV collection Nov, 1929, “Marree via
Mulka perv Sir Colin MacKenzie” from George
Aistog;

MARSUPTALS: Diprotodontidac; Diproieden sp,.
’Zygomaturns 8p,

Mactopodidae; Macropus sp.. Protemnodon sp.

11. COOPER CREEK. A number of sites between
|. Eyre and the Birdsville Track. Katipiri Water-
hole is the type locality for the Kalipiri Sands
(FP l\o-Pleistocene) from which fossil vertebrates
ure eroded by the creek. SAM, NMV, AMNH,
UCMP, EMNH, HM, Note: Lists of fossil assem-
blages ure derived from the UCMP collection

a, Cannatalkaninna (V5377) 28°40'S, [38°26°E
(KOPPERAMANNAI!-_
East Cooper crossing, 7Sub-Recent.
MARSUPIALS: Dasyuridae.
Muridae; Rattus sp,, Notamys sp,

bh. Unkumilka Waterhole (V5378)
138°19'R (KOPPERAMANNA).
MARSUPIALS; Diprotodontidae; U/preradart sp..
*Zyeomaturns sp.

ce. V5379 28°39'S,
MANNA),
SE of While Crossing.
MARSUPIALS: Thylocoleantdac; ?Thyhicalee sp.
Diprotodontidpe; Diprofodorn sp.
Macropodidae: ?Praceptodan sp.
REPTILES: Varanidae; Megalanin sp.

28°41°S,

MB Sh (KOPPERA-

4d VS5380) 28°37'S, I38°I4E (KROPPERA
MANNA},

MARSUPIALS; Macropodidae; Macropus of.
titer.

PUTHERIANS: Muridae,

& VS38I 28°35'S, J3R°I3E (KOPPERA-
MANNA).

Near Tilla Tilla Waterhole,
MARSUPIALS: Thylacoleonidaes ?Thylacolen sp
Diprotodontidac; Dipratodon cf. aptatum,

Macropodidac; Muerepus sp,, Pretemnedan sp,
Sthenurus sp.
BIRDS: Dromo:nithidae.

f, Malkuni Waterhole (VY5382) 28°34'S, [38°
07 E (KOPPERAMANNA),
MARSUPIALS; Diprotodentidae; Dipratedon sp,
Macropodidae, Macrepus cl. titan, small Macro-
podid, Proremineden ch anak, P. cf. brehue,
Sthenurus sp., Procéptoden cf, goliah.
BIRDS: Dromornithidaec.
Phalacrocoracidae; Phalacrocorax carho, P, varius.
Tytonidae, Tyre cf, novaeliallandiae,

2 V5859 28°33'S. 138°09'E
MANNA).
MARSUPTALS: Dasyuridac; Sarcophilus sp,
Diprotodontidae; Biprotadoan cf. minor.
Macropodidaes *Procopioden sp.
BIRDS: Phalacrocoracidne: Phalacroconay sp,
REPTILES: Crocodihans.
FISH: Cermtodontidae; Epleerarodus sp.

h. VS860 28°35'S, I38°0S°R (KOPPERA-
MANNA).
MARSUPIALS: Dasytridae: Sarcophitus sp.
Vombatidae: Phascolonns cf gigas.
Phalangeridae; Vrithasvrus cf. valpecula.
Diprolodontidae; Diprotedon sp, DB, cf. minar,
?Nototherium,
Mucropodidae; Betrangia cf, lesuene, Macropus cf,
titan, Lagorchestes yp., ?Onychoyalea sp. Protern-
nodan sp. Po cf brehus, Sthenurus sp. Sy cf,
andersoni, 8, cf, brawnei, Proceptadan cf. goliah,
Pict. tapha,
EUTHERIANS: Muridae.
BIRDS; Dromornithidae,
REPTILES: Varanidaes Megalania sp.
Chelonians and Crocodilians.

i. V5861 28°54'S, 1[38°OS'F
MANWA).
Katipiri Waterhole (Cuttapirra Waterhole),

(KOPPERA-

(KOPPERA-

MARSUPTALS; Vombatidac; FPhascelonuy cl
Rigas.

Diprotodontidae; Dipretadan sp,, ?Zyeornaturis
sp

Mucropodidae; °Hallabia sp. Pretemnodun cf,
anak, Sthenuras sp.. 8. ef. rindalei, Procoptodar
ef, goliah,
BIRDS: Dromornithidae.
Anhingidae; Aniinga sp.
REPTILES: Vurunidue; Megalania xp,
Chelonians and Crocodilians.
FISH; Ceratodontidac.

j, VS866 24°34°S, 138°00'E (LAKE EYRE/
KOPPERAMANNA|,
MARSUPIALS: Vombatidae; Phascolanis sp.
Diprotedontidae; Diprotedon sy, ?Z7ygomaturnue
SP,
Macropodidac; Proaremnodon cf. bredins, Sthenuray
cf. andersoni, §, cf, rindalei, Proceprodon sp.
BIRDS: Anhingidae; Anhinga sp.
Phalactocoractdae: Phalacrocoray carbe,
Analidae; Bizivra sp.

PLEISTOCENE VERTEBRATE FOSSILS {Ns

REPTILES: Varunidae; Mezalania sp
Chelomans and Crocodilians,
FISH: Silurifarmes and Dipnoi.

k. VS5868 28°32°S, 137°59E (LAKE EYRE).
MARSUPIALS: Diprotodontidae.
Macropodidue.
BIRDS: Unidentified material,

|, V6147 28°34°S, 138°09E (LAKE EYRE),
MARSUPIALS: Thylacoleonidae: Thylacoles sp.
Diprotadontidae: Diproreden sp.
Macropodidie.
BUTHERIANS: Muridae; Ratiny sp.
BIRDS: Phalacrocoracidae: Phalacrocera. sp,
REPTILES: Varanidae; Megalaniu sp.
Crocodiliuns,
FISH: Unidentified material.
CRUSTACEANS: Decapods.

m. Unnamed focality, approx. 28"00'S, 139°30°R
UNNAMINCKA).
Far NE South Australia, near Cooper Creek.
Tooth recovered hy drilling crew, SAM,
MARSUPIALS: Dipratodontidae.
Tale (L886), Stirton et al. (1961), Rich ev al.

(1978), Rich (1979), Tedford (pers. comm.
19R0)).

12. LAKE PALANKARINNA 28°46’S, 138°25’E
(KOPPFERAMANNA!

Eust of L, Fyre,

a, Channel Sand Locality (V5854)-
Near the top of escarpment about 500 m north of
Turtle Locality. Katipiri Sands (Late Pleistocene),
incising Tirari Formation. UCMP,
MARSUPIALS: Diprotodontidae: Diprotedan sp.
Macropadidae,
Birds, Teleost fish, Chelonian and Crocadilian
remains.

b. Mullet Locality (V71173).
Sinall bluff 25 m NNW of Keane Quarry (V6265),
Katipiri Sands (Late Pleistocene) overlying Eta-
dunna Formation and overlain by Tirari Forma-
tion. FUSA, UCMP,

MARSUPTALS: Diprotodontidae; Zysomalurus.
sp.
Macropodidue; Proteninodon sp, ?Prianotemmnns
Bist

Fish and Crocodilian remains.
Slirton ef al, (1961), Rich ef al
(1979).

13, HERGOTT
IMARREE).

3 km north of Marree. Mandible, SAM.
MARSUPIALS! Diproatodontidae; Diprotedan sp.

14. WELCOME SPRINGS 29°40'S, 137°S0°E
(CURDIMURKA),

15 km SW of Marree, Hard green pebbly clay.
cemented by cileium carbonate from mound
spring. Mandible and bone fragments, SAM,
FUSA.

MARSUPIALS: Diprotodontilue: Diprofodan sp.

(1978), Rich

SPRINGS 29°37'S, 138°04E

15. MURNPEOWIE 29°35'S, 139°07'E
(MARREE).

[00 km east of Murree. Mandible, SAM.
MARSUPIALS: Diprotadonudae: Diprotoden sp.

16. LAKE CALLABONNA 79°50°S. 140° LOE
(CALLABONNA).

NE South Australia. Gypsiferous lacustrine clays
und sands, Millyera Formation. SAM, AMNH,
UCMP, SL

MARSUPIALS: Vombatidse; Phascolonnus gigas,
Diprotodontidae; Diproredon sppt?).
Macropodidae, Macrepus spp., Protemnodan sp.,
Sthenurus sp, nov., S. tindale’ (Wells & Tedford
in prep.}.

BIRDS: Dremaiidae; Dromiainus sp.
Dromarnithidae; Genyernis newtorir.

Unidentified smaller bird remains.

Tate (1893), Brown (1894), Stirling & Zietz
(1896, 1900). Stirling (1900, 1913), Howchin
(1930), Hale (1956), Tedford (1966, 1973),
Callen & Tedford (1976), Callen (1977), Rich
(1979), Wells & Tedford (in prep.)

17, BILLEROO CREEK 31°08’S,
(CURNAMONA}).

20 km SE of L. Frome. Red, sandy fluviatile Euri-
nilla Formation (Late Pleistacene). SAM, FUSA,
NMV, AMNHEL.

MARSUPTALS; Vurmbalidue; Lasiorhinus sp.
Thylacoleonidac; Tiyvlacaleo carnifex,
Diprotodontidae; Dipretadon sp.

Macropodidae; Betionyia sp., cf. Propleapus sp.-
Maeroapus sp.. M. cf. forragus, OspAranter sp.,
Sthenurus sp, noy,, 8. tindalet, Sthenuris sp., Pree
coptoden golial.

EUTHERIANS: Muridacy Lepurilluy sp.. Coni-
lurus sp.. Rattus sp. Pyeudomys sp,

BIRDS; Dromaiidue; Dramaius sp.
Dromornithidae; Genyornis newton.

Tate (1886), Callen & Tedford (1976), Callen
(1977), Wells & Tedford fin prep.)-

18. PERNATTY LAGOON 31°37'S,
(TORRENS).

SU km SE of Woomera. Weathered skeleton, Irag-
ments. SAM,

MARSUPIALS: Diprolodontidae; Diprotcdan sp.
Pledge (1974).

19. HOOKINA CREEK
(PARACHILNA)

25 km NW of Hawker. Valley-fill alluvium and
red pver-bank deposits of the Pooraka Formation
(Late Pleistocene). SAM, FUSA,

MARSUPIALS: Dasyuridae: cf. PDasycercuy sp.,
Surcomiilus sp. unidentified small dasyurid,
Peramelidae.

Vombuatidae; Lasiorhinus sp. ct. Wonibatuy sp.
Diprotodontidae; Dipratodan sp,

Macropodidae; cf. Petorays sp., Muacropus ef.
vigantens, M. cf eugenit, Macropus sp. Oy-
pheantwer sp.. Wallabia sp,, Propleopus oscillans,
unidentified small macropou,

40° 1S E

IST IGE

31%44'S, 138°14E

LOG Db, L. G. WILLIAMS

EUTHERIANS; Muridac; Aydromys chryso-
gaster, Leporillus conditor, Pseudamys hermanns-
burgensis, wildentified rodent.

BIRDS: Dromaiidae; Dromrainy sp,
Dromornithidac; Genyornis sp.

Unidentified small bird,

REPTILES: Elapidae; unidentified genus.
Varanidac; Varanis ef. glganteus,

Scincidaes Trachydosanrus sp. unideritified small
lizards.

AMPHIBIANS: Unidentified frogs,

FISH: Unidentified small fish,

Daily (1956), Twidale (1966), Williams & Polach
(1971), Williams (1973), Williams (in prep.).

20, MOUNT EYRE 31°46'S, 138°1'E (PARA-
CHILNA).

26 km NW of Hawker, Western piedinont slope of
ranges, Weathered skeleton on flat of sandy clay
SAM.

MARSUPIALS: Diprotodontidae; Dipretoder sp.
Daily (1954),

21. WILLOCHRA CREEK
(ORROROO).

8 km WS af Quorn, Encrusted skull, SAM.
MARSUPIALS; Macropodidae; Macropus sp.

22a, BOOLCUNDA CREEK 32°13'S, 138°18'E
(ORROROO).

“Lanewarren” 30 km NE of Quorn, Red clay and
gravel of dissected ovtwash fan. Mandible frag.
ment (casis). SAM.
MARSUPIALS: Diprotodontidac:

32°1S'S, 138°OS'E

LY gomalurius

sp.
‘Twidale (1966)-

22h BOOLCUNDA CREEK 32°/3'S, 138°3 °F
(ORROROO).

Unnamed tributary, 18 km south of Craddock.
VUSA,

MARSUPTALS; Macropodidae; Proceprodon sp,
Williams (in prep,).

2Zc HOOLCUNDA CREEK
138°30'S (ORROROO),

Note with specimen: “NE corner Mookra on Bool-
cunda Creek 50 ft below surface in sand with
Wish jn civernous part of munganite lode”
AUGM,

MARSUPTALS; Macropodidae: Pratemnaden sp-

23, BUICKALOWIE CAVES 32°08'S, 138°S5‘E
(ORROROU),

40 km SE of Hawker Maips Cave (B 3), Clara
St Deora Cave (F 4). Bone generally encrusted
with eulcite. SAM.

MARSUPIALS; Dasyuridae, Dasvurns sp., Sarce-
Pilla daeristé.

Thylacinidae; Thylacinus cynocephulas,
Peramelidae.

Thylacolennidae: Thylacolea carnifer.
Mactopodidue; Bellonwia sp,, Petarous sp., Macro-
pus sp.

upprox 32°20'S,

EUTHERIANS: Muridac,
BIRDS: Unidentified.
REPTILES: Unidentified,
Winton (1922),

24. TERTULPA 32°15'S,
MONA).

40 km NE of Yuntu. Note with specimen: “Brady's
Ciully Teetulpa, 14 feet from surface in the drift.
(888, SAM.

MARSUPIALS: Macropodidae; Proceptodon of
rapha,

Brown (1888).

25. DEMPSEY’S LAKE.
(PORT AUGUSTA),

§ km NW of Pt. Augusta. Red acolian sands of
the Pooraka Formation (Late Pleistocene), SAM,
PUSA,

MARSUPIALS: Dasyuridac} Dasyuris of, viver-
rinuy, Sarcophtilaus harrisii.

Vombatidae; Phascalonus cf. gigas, Lasiorhinus
latifrans.

Diprotodontidae; Pipretadoan sp.

Macropodidae; Beltongia lesueur, B. penicillate,
Macrapus sp.. Macropas cf. ferragus, Osphranier
sp., Protemnodon brehus.

BIRDS; Dromajiidac; Praniaius sp.
Dromornithidue; Genvornix sp.
7Anatidac,

REPTILES: Elapidae,
Scincidae;: Trachydosaurus sp.
Cooper (1959), Williams
(1976).

26. ARNOLD'S BANK 32°22'S, 137°46'E (PORT
AUGUSTA),

15 km north of Pt, Augusta, Red sand dine in
area of dunes and salt flats, head of Spencer Gulf.
Pooraka Formation (Late Pleistocene), SAM.
MARSUPIALS: Diprotodontidae: Dipratedan sp-
Williams (1976).

27. YJAILLPARA CREEK
{ORROROO),

30 km NE of Orrarda. Red clayey alluvium asso-
clated wilh creck. SAM,

MARSUPIALS: Diprotodontidae; Piproledon sp,
Macropodidae; Sthenarus sp.

Hale (1956), Daily (1960b), ‘Iwidale (1966),
Williams (in prep,).

28. NECTAR BROOK 32°42’S, (37°56 (PORT
AUGUSTA).

78 km SSE of PL Augusta. Red ulluvium exposed
during dam excavation in 1898, SAM,
MARSUPIALS: Diprotodontidae, Miprotoden sp.
Macropodidve; Maecropus sp.

29, PEKINA CREEK 32°44'S, ]38°37E
fORROROO)

2 km south of Orroroo, Greenish lacustrine clays
exposed on SE shore of reservoir, and poorly-
sorted alluvium below dam wall. Macropus sp. his

139°41"E (CURNA

137°42°B

32°28'S,

(1973), Williams

32°S1'S, |38°S5'E

PLEISTOCENE VERTEBRATE FOSSILS 107

also been recovered from 4 well excavation 18 m
deep on Pekina Creck floodplain. SAM, FUSA,
MARSUPIALS: Vormbatidae,

Diprotodontidae; Dipreteden sp.

Macropodidae; Bellanxia sp.. Macropus sp.
Howehin (1909), Williams (in prep.)

30. BLACK ROCK GRAVEL PIT 32°47'S, 138°
40'E (ORROROO),

8 km south of Orroroo, Distised! gravel pit on allu-
vial plain, Sinuous channel deposit of fine, rouniled
gravel with red sundy matrix. SAM,
MARSUPIALS: Diprotodontidae; Diprotedan sp
Macropodidac; Mucropus sp., Prolemnaden ‘ps
Sthenurus sp.

31, MANUNDA CREEK 32°56'S.
(ORROROO),

40 ko cast of Peterborough, Red clayey overbank
deposit af creek. SAM,

MARSUPIALS: Macropodidae; Proeoptaden sp.
Edwards (1964),

32, PORT PIRIE GRAVEL PIT 32°15'S, 133°
05'B (BURRA).

3 km south of Pt, Pirie, Western flank Of ranges.
Clean coarse quarts gravel, SAM.
MARSUPIALS: Thylacoleonitue: Vhwlacolea sp.
Diprotodontidae, Diprotedar sp-
Macropodidac; = Macopuy sp, M. cf,
Sthenurus sp.

Pledge (1973, 1974).

33. WAUPUNYAH CREEK 93°15, [39°05 'E
(BURRA).

8 km cast of Terowie. Red clayey alluvium of
creck. SAM, TUSA,

MARSUPIALS: Vombatidae: cf, Lastoriinns 5p,
Diprotodontidae: Diprotodon sp,

Macropodidac; Macrapus spp, Osphranter sp.
Pracaptodon sp.

Williams (in prep.).

34, ULOOLOO 33°19'S, 138°53'E (BURRA),
35 km north of Burra, Piedmont and alluvial
depasits of Irwin and Terowie Creeks. SAM,
FUSA.

MARSUPIALS: Diprotodontidae; Diprotadon sp,
Macropodidaes Pracepladon sp.

Williams (in prep.).

35. COLLINSVIELE 33°20'S, [39° 08'E
(BURRA),

50) kin NE of Burra. Red silty floodplain deposit
in Witt Creek valley, expoyed by modern gully-
ing. SAM, FUSA,

MARSUPIALS: Diprotodontidae, Miproradon sp.
Micropodidue; Macropus spp. Procapladen sp.
Williams (in prep.).

46, NEWIKIE CREEK 33°30'S, 139° 10°E
IBURRA)

25 km NE of Burrs, Alluvial fan sands ond
provels. SAM, PUSA.

MARSUPIALS: Vombalidue; Lasiariinis sp-

139°21E

furan,

Diprotodontidue; DBipretocden sp
Macropodidae,
Williams (in prep.)-

37, BALDINA CREEK 33°41'S, 139°U4'E
(RURBRA),

13 km east of Burra, Red silty valley alluvium. anu
fan, SAM, FUSA.

MARSUPIALS: Vombatidaes Lasiorhinis sp,
Thylucoleonidae; Thylacolea carnifex,
Diprotodontidae; Dipreraden sp.

Mactopodidae; Beftongia sp., Macropus sppo Pro-
remnodon sp, Sthenurus cf. atlas,

BLRDS: Dromornithidae; Genyornis newlenit

Vare (1890), Zietz (1890), Stirling & Ziete (1896,
1913) Stirling (1900), Howchin (1930), Hale
(1956), Rich (1979), Williams (in prep.),

38. BUTE 33°52’S, 138°01'R (BURRA).

20 km SW of Snowtown, Material associated With
human remains A25805, from a sand dune, Prob-
ably Holocene, Anthropology Collection. SAM.
MARSUPTALS: Macropodidae; Macrayiis sp,
BIRDS: Dromuaiidae; Yramalus sp,

49. BURRA CREEK 33°52'S, 139°09 E
(BURRA),

30 km SE of Burra, Brown silts, sands, and gravels
of alluvial fan. FUSA,

MARSUPIALS: Diprotodontidae; Diprotedon sp.
Macropodidite.

REPTILES: Unidentified small lizards.
AMPHIBIANS: Unidentified frogs,

Chapman & Mawson (1925), Williams (in prep.),

4), BUNDEY 33°53'S, 139° L8E (BURRA),

40 Km SE of Burra, “Gum Creek”. Red clay ex-
posed by dam excavations in L889 and 1953,
SAM.
MARSUPIALS:
UTNTHUS,
Thylacoleonidae: Thylacalea carnifer,
Diprotodontidae: Diproteden spp(?),
Mucropodidae; ?Sthemurus sp-

Zicté (1890), Stirling £1900), Howchin (1930).
Pledge (1977), Williams (in prep.).

41, MORGAN 34°02'S. 139°40°E (RENMARK)
River Murray flats, north bank, where eliffs are
cut by tribatary fram the NW. SAM.
MARSUPIALS; Vombatidue; Phascolanuy sp.
Diprotadontidae, Bipretodon sp.

Mactopodidse; Macropus sp.

N.S, Pledge (pers. comm, 1979).

Dasyuridae; Surcephilus ef,

42, CHOWILLA 34°01'S, 140°50°R (REN-
MARE),
North of Renmark. Excavation for proposed

Chowilla Dam wall, [8 m deep. SADME
MARSUPIALS;: Vombntidae: Phascolonus sp.
Firtnun (1966), Marshall (1973).

43. CURRAMULKA 34°42'S, [37°44'E (MAITT-
LAND),
a. Town Cave (¥ 2) jn Curramutka. Red, stony

18 D. L, G, WILLIAMS

clay With flowstone. Bone generally enerusted with
calcite. SAM, FUSA, LICMP,

MARSUPIALS: Duasyuridaes Desvurns sp.
Peramelidae; Perameles sp.

Vombatidac: Lasiorhinns sp,

Thylucoleonidac; Tivlacolee carnifex, V. hilli.
Diprotodontidae: cl. Notarherlum sp,
Macropodidae; Mucropus sp., Prateninadon syn,
Wallabia sp.. Sihenurus spp.. Procaptoden sp,
EUTHERIANS: Muridae.

b, Quarry 2 km south of Curramulka, Red cal-
cite-cemented bone breccia filling fissures in Early
Cambrian limestone. SAM, FUSA,
MARSUPLALS: Vombatuae; Lasiverhinus sp.
Macropodidae; Macrapus sp., Procopledon sp,
Pritchard (1491). Howchin (1925), Dally (196Da.
W60b}, Pledge (1977),

44. LARE FOWLER 35°15°S, 137°37'E (KINGS.
COTE),

Southern Yorke Peninsula, Gypsum lunette on east
shore of lake, Fossils exposed by quarrying. SAML
(J, A. MeNamara (974. Undergraduate project,
Zoology Dept, University of Adelaide, unpub,)
MARSUPIALS: Dasyuridve: Sareophilus ef
ursitus,

Peramelidae; Macrotis cf, lavelis,

Vombatidac; Lasorltiinus sp.

Thylacdleonidae; Thylacelee earnifer,
Macropadidac, Belionuia lesueur, Macropuy et.
ferrayus, M. cf evgenli, Onychogalea sp.
EUTHEFRIANS: Muridac; Leparillyus sp.
Howchin (1900), Jack (1921), King (1950),

45, PONDALOWIE BAY 35°14'S, 136°50'F
(KINGSCOTE).,

SW Yorke Peninsula, near one of the Jakes at
Pondalowie, Calearedus chiystone slab with track-
ways. ?Holoceno.. SAM,

MARSUPIALS: Macropodidac: ?Maerapus sp.
BIRDS: Dromaiidaes ?Bromatus sp

46. ROCKY RIVER 35°55'8, 136°47°E (KINGS-
COTE),

Western end of Kangaroo Island. Swamp deposit.
SAM,

MARSUPIALS: Dasyuridae; Sareaplilus sp.
Vombatidae; Unidentified genera,

Phascolarctidaes Phaseolaretos cinereus,
Phalangeridae; Trichosurus valpecule,
Diprotodontidae; Diproieden sp..
trilohius,

Mucropodidae; Macropus fuliginosus, M- engenti,
Pratemnodon sp, Sthenuray spp. S. gilli.
EUTHERLANS: Muridae,

BIRDS: Dromaiidae; Premaluy sp-

Jones (1923) Tindale ef al. (1995), Hale (1956)-
Pledge (1975, 1979), J. BR. Hope (pers. comm.
1980),

47, KELLY HILL CAVES (K 1-4, 14d, 44)
35°S9S, 136°54’E (KINGSOCOTE).
SW Kangaroo Island. SAM,

Zygamatary

MARSUPIALS: Dasyuridac. Dasyurus maculatas,
Sarcaphilus harrisii, Phascogale tapoatafa.
Vombatidue; Lasiorhinus sp,

Phascolarctidacs Phascolarctos cimercus.
Phalangendae; Trichosuras vulpecula.

Petauridae; Pseudocheirns peregrinus,
Macrapodidae; Murrapys fulizinosus, M. eugenil,
Srhenurus ch. ovcidentalis,

BIRDS; Dromaiidac; Uromaius diemenianus.
Howehin (1930), Hale (1956), Hope er al.
(1977), Pledge (1979), R. T. Wells (pers, comny.
1974),

48a. MOUNT TAYLOR CAVE (K&) 35°58'S,
(37°03 E (KINGSCOTE).

SW Kangaroo Island, 5 km east of Mt Stockdale.
SAM,

MARSUPBIALS: Macropodidae: Sihenurns sp.
Pledge (1979),

48b. EMU FOUR HOLE CAVE (K 20) 35°59'S,
136°54'E (KINGSCOTE),

SW Kangaroo tsland, near Mt Taylor, Subfogsil ta
modern hones (Pledge 1979), SAM.
MARSUPLALS: Dasyuridae; Dayvurus viverrinis,
Sminthapsis murina,

Peramelids; Perameles sp.. fsoodon obesutus.
Phalangeridae; Trichosurus vulpecula.

Petauridac; Pyeudocheiruy peregrinus,
Burramyidae; Cercartetus concinnus.
Macropodidae, Poterous platyaps, Macropuy fuli-
rinasus, M. eugenit.
MONOTREMES: Tachyglossidac;
aculeatus,

RUTHERIANS; Muridaez Ranus fuscipes, R,
lutreolus.

BIRDS: Dromnaiidae; Bramualis diemenianus,
Hale (1956), Pledge (1979)

48¢. FOSSIL CAVE (K21t) 35°59'S, 136°S4’E
(KINGSCOTE).

Adjacent to Emu Four Hole Cave. SW Kangaroo
Island, SAM,

MARSUPIALS; Dasyuridae; Sarcophilus cf har
resin.

Vombatidae,

Phascolarctidae; Phascelarctos cinereus
Macropodidac: Maecrapus fuliginosyus, M-. eugenil,
Sthennrus cf. hrowrnel.

EUTHERIANS: Muridae,

Pledge (1979),

48d, SETON ROCK SHELTER (K 30) 35°59'S,
137°03'E (KINGSCOTE),

SW Kanguroo Island, Archaeological; sandy sedi-
ments excavated to a depth of about 27 m. SAM.
MARSUPIALS: Dasyuridace: Oasyuruy — cf,
geaffroi/vivereiins, Bo maculatus, Sarcaphilus har-
risil,

Peramelidae, Perameles borgainviile, tscodon ube
sulas,

Vombatidue;: Lasfarhimey latifrans

Phalangeritae: Trichosueus vulpeewta,
Burramyidae; Cerearterus lepidus,

Tachyglossus

PLEISTOCENE VERTEBRATE FOSSILS 19

Macropadidae; Bevongla penicillata, B. lesueur,
Potorous platveps, Macropas ct. Jaliginosus, M-
greyi. M. rujagrisens, cf. Mepaleia rufa, Lagor-
cheses leporides, Sthenurus cf, gilli,
FUTHERIANS, Mouridae, Afydranys ¢liryse-
waster, Raras pusoipes ereyi, Ry hettecdlas, Mastar
comus fuscus, Pseudoms occidentalis, PB. austra-
lis shortridget.

BIRDS: Procellanidae; Pachyptila cl. salvint, Puf-
finis sp,

Platalwidues Tireskiornis cf, molneca.

Anulidae, ef 4aseranas semipalmaia, Vaderna cf,
nidornoides, Anas v& superciliosa, Anas et, cs-
tanea. Malacarhynelins menbranacers.
Accipitridae; Mieradetus morphinoides.

Falconidae; Falce berivare.

Phasianidaes Coturule cf, pectoralis.

Turnicidacy Turnix varia, T, velox

Rallidae; Ratlus philippensis, R. pectoralis, Par-
sauna ef, fuminea, Gallinula (Tribanyad cf. mor-
lierti, GO, (Tribonyx) ef. ventralis,

Burhinidues Burhinusy mrageinasiris,

Scolopacidae; Gullinugo cf, harilwiekt.

Laridae; Larus noviehollandive, Sterna cf, nereis,
Columbidac: Ocyphups lophates.

Platycercidas; Pezoporus wallicus, Cathamus dis-
colar.

Hirtindinidae; Mirunda cf. taliticn, Petrechelidon
nigricans.

Meliphagicae,

Sylviidae; Cinclerhamphus eruralis.

Grallinidae: Grallina evanalenca,

Cracticidue,; Gyniiorliina dibicur, Slrepent gracu-
lina, 8. versicolor.

Corvidae; Corvus sp.

Unidentified passerines.

REPTILES: Elupidae.

Varanidac: Maranis sp.

Scincidue; Trachydosauras rugexus. Tiliqna niveo-
lutea, cf Egernia whitii.

Agamidaes 4mplibolarus spp.

MOLLUSCS: Marine molluscs of archacologicil
origin, and terrestrial/aquatic molluscs

Hope eral. (1977),

49, HOG BAY RIVER
(KINGSCOTE).

Eastern Kangaroo Island, Found in “Pleistocene
drift’ (Cutulovuc!, SAM.

MARSUPIALS: Mucropodidae, Macrapus sp.

as°49'S, 197"57’E

50, KAPUNDA 34°23'S, 139°00E
(ADELAIDE).

South of Kupnnda. Bones and teeth in matrix.
SAM.

MARSLIPIALS: Vombatidue; Phascolonas sp-
Jack (191%), lohns (1947).

5). TWO WELLES 34°36°S,
LAIDE),

35 km north of Adelaide. Sandpit “near Two
Wells” (notewith mandible), SAM,
MARSUPIALS: Diprotedontidae; Diprvfodent sp.

138°3'ER (ADE-

52, GAWLER 34°35'S, 138°45'E (ADELAIDE)
40 km north of Adelaide. Excavations on banks of
Gawler and South Para Rivers. SAM.
MARSUPIALS: Diprotodontidac: Diprotodon sp.
Stirling (1900). Howehin (1930), Hale (1956)-~

53. YATALA 34°51'S, 138°37E (ADELAIDE).
NE suburb of Adelaide. Jaw “9 feet below the
surfuce in 4 bed of gravel” (Moncrieff 1882). Lo-
cation of specimen unknown.

MARSUPIALS: Diprotodontidae; Diprotoden sp.
Moncrieff (1882),

$4. FROMM’S LANDING 34°46'S,
(RENMARK).

River Murray, near Walker Flat. Archaeplogical;
rock shelter. SAM,

NOTE; The assemblages summarised here do nor
distinguish the numerous straugraphic levels
recognised by Mulvaney er al. (1964).

i, Shelter 2.
MARSUPIALS: Dasyuridae; Duasyurus peoffrai,
Dasyuropy maculatus, Sarcaphilus harrisii,, Dasy-
cercus eristicaudata, Antechinus flavipes, Av swain-
soni, Sminthopsis cf. murina, Myrmecobius fas-
cians.
Thylacinidae; Thylacinus eynacephalis.
Peramelidac; Peramelex bougainville. cf
obesulus, Chueropus ceaudatus,
Vormbatidue; cf. Lasiorhinus latifrons.
Phalangeridae; Trichosurus vulpecula-
Petausidac; Pseudocheirus peregrinuy.
Macropadidac; Retlongiy penicillaia, B. lesueur,
Pororous margani, Macrapus canguru, Thylagale
engenii, Onychogalea lunata, Lagorehestes lepo-
ridvs, Lagastrophus [aseiatus.
EUTHERIANS: Canidae; Canis fanniliaris dingo.
Mutidae; Aydromys chrysogaster, R. greyl, Rattus
luireolus, cf, Psendomys auritus, Thetamyy gp. cf.
Noatemys sp., Contlurus albipes.
Mulvaney er al. (1964), Archer (1971).

b. Shelter 6.
MARSUPTALS: Dasyuridae:
elaius,

Peramelidae; Perameles hougainville, Iyaodon obe-
srlus, Chaeropus ecaucdatiiy.

Phalangeridae; Trieliosurus vulpecula,
Macropodidae; Relrongia penicillata,, Macrepus
canguru, Thylogale ecugenti, Lagorchestes lepo-
rides, Lagostrophus fasciatus,

EPUTHERIANS: Canidae; Caniy familiarts dingo.
Muridae: Hydromys chrysogaster, R. grey’, Rattus
fatrealus:

Mulvaney ef al, (1964),

139°33'E

lsoodon

Myrmecebius as-

55, DEVON DOWNS 34°41'S, 139°37'E (REN-
MARK).

River Murray, nocth of Mannum. Archaeological;
rock sheller, SAM. Note: The assemblages suam-
marised here do not distinguish the numerous
stratigraphic levels recognised by Hale & Tindale
(1930), See also M. Smith, 1977 B.Sc. (Hons)

1d BD. L. G. WILIJAMS

thesis, Avst. Nat. Univ, Dept of Prehistory &
Anthropology (impub,),

MARSUPIALS: Dasyuridae; Dayyvurw viverrinicy,
D. geoffroyi, Sarcophilus harrisii, Phaseagale flax
wipes,
Peramelidae;
sulus,
Vombutidae;, Lasiorhinus sp.

Phalangeridae; Trichosaris Vulpecila,

Petauridae; Psevdocheirus sp-

Macropodidae, FRettongia sp, Poterons $f.
Muacropus sp, Mucropus cf. sigantens, Thylexule
sp.. Lagerchestes leparides,

EUTHERIANS: Canidae; Canis familoriv dinge.
Muridae; Hydromys elhuysegester, Rattus sp.
BIRDS: Dromaiidae; Dromains novuehallanediae-
Anatidae; Querquedula sp.. Chenopis alrwta, Br-
giuvra lobata.

Accipitridae; Urogeruy anedice-

REPTILES: Boidae; Python spilars.

Vuranidae; Varanty cf, eouldii.

Scincidae: Trachysaurus rugosa, Tiliqna sp,
Agamidae; Ampliholurus sp,
TORTOISES: Chelidue; Chelodina
Emydura maequarti,

FISH) Maceullochellidue; Oliguins macquariensis.
Plotosidae: Tyndarus nonelents.

Plectroplitidaes Pleerroplite, ambleuns.
INVERTEBRATES: A range of molluscs ane
crustaceans.

Hale & Tindale (1930).

56, CHUCKA BEND 34°53'S, 139°39'E (REN-
MARK),

River Murray, north of Mannum. Site details and
location of specimens unknown,

MARSUPIALS: Vombatidae; Pliascolonuy sp-
Stirling (1913), Howehin (1930),

57, ADELAIDE AREA 34°55’S, 138°35°E (ANP-
LAIDE)..
A number of sites which appear to be associated
with alluvium of the River Torrens.

a. Allenby Gardens 34°53’S, 138°34°E (ADE-
LAIDE),
Adjacent to NW corner of Adelaide city. Speci-
mens found “Approx. 20 ft, below the surface”
(SAM Catalogue).
MARSUPIALS: Diprotodontidae; Dipraradon sp.,
ef. Nototherivim sp,

bh. Brompton 34°54'S, 138°34°R (ADELAIDE),
Adjaceat to NW corner of Adelalde city: 20, East
Street, Brompton, Note with specimens, "25 feel
below surface" In black clay, SAM,
MARSUPIALS; Diprotodontidae: Diprorodon sp

c. Croydon 54°53°S, |38°34E (ADELAIDE),
Adjacent to NW comer of Adelaide city. Several
localities “within two and three miles of Croydon"
(Tate 1890), "Found on Mr Woodhouse’s pro-
perty (South of Railway line West from Croydon
Railway Station) at Croydon about 4-6 fe helow
the surfuce in an old river bed Of shurp sane and
sravel™. (Note with specimens.) SAM,

Perameles myoyara, Isaode Abie

longicallis,

MARSUPIALS: Vombutidaes cf, Phascolomyy sp-
Diprotodontiduey Diprereadon sp.
Tate (1890),

d. Findon 34°55°S, 138°32'R (ADELAIDE).
Western suburb of Adelaide, One of numerous
gravel pits, SAM,

A Vombatidae; Unidentified man-
(ile,

Dipretadontidae; Diprolodar sp,

HERDS: “Bird bone” catalogued,

Howchin (1913),

c. Kirkaldy Pit
LAIDE).

Western suburb of Adelaide, One of numerous
gravel pits. SAM.
MAKSUPILALS: Diprotodontidae; Diprotadon sp,

{ Thebarton 34°55'S, 138°34'E (ADELAIDE)~
Adjucent 1 west side of Adelaide cily, Rib bone
from “a depth of six feet” (Waterhouse (882)
Origin of P, azael jaw not known. SAM.
MARSUPIALS: Diprotodontidae; Pularehestes
vzuel.

Waterhouse (1882)-

58. HALLETT COVE 35°04'S, 138°30'E (ADE-
LAIDE).

Southern coastal suburb of Adelaide, Waterworn
molar from modern heach gravel, derived from
cliff exposures of Pleistocene sediments, SAM,
MARSUPIALS: Diprotodonmtidae: Diprotodon sp-

54, ECHUNGA 35°06'S, 138°48'E (BARKER).
30 km SE of Adelaide. Bone from "deep plluvium"
{note with specimens), May be Tertiary. SAM.
MARSUPIALS: ?Mucropodidae.

60a. SALT CREEK 35°28'S, 138°20°R
(BARKER)

80 km south of Adelaide, Sulphurous black clays
of swamp deposit. There is a New Salt Creck
“four miles north of Cape Jervis” (Hale 1946),
but Brown (1892) leaves no doubt that the site
is near Normanville. SAM,

MARSUPIALS: Vombatidae; Phaseolonus givas.
Thylacolvonidae: Thvlacalee carnifex,
Diprotodontidae; Diprefedan sp.

Macropodidae, Berfongia sp., Macrepus sp, Os-
plranter sp., Sthenurts sp., Procaptodon sp,
BIRDS: Drompiidae; Drameains sp.
Dromorhithidae: Geayorniy newton,

Unidentified bird remains.

REPTILES: Unidentified material.

FISH: Unjdentified material,

Waterhouse (1880), Brown (842), Stirline &
Zietz (1896). Zictz (1907), Stirling (1913), How-
chin (1930), Hale 11956), Ride (1967), Pledee
(1977)

hb, YANKALILILA 35°28'S, 138°37°E
(HARKER.

KO kin south of Adelaide. Possibly rhe same site as
Salf Creek. SAM.

MARSUPIALS: Diprotodontidae: Plprotedan sp,

34°55'S, 38°30°R (ADE.

PLEISTOCENE VERTEBRATE FOSSILS I

61. GOOLWA 35°31'S, 138°45°E (BARKER).
70 km south of Adelaide. Known as the “Goolwa
Footprints”, specimens actually taken from an
eroding cliff at nearby Middleton Beach in 1938,
Exposed below ubout 7 m of “sands, limestones
ete. above the pipeclay on which the prints were
made,” (Letter by Fenner, 20th June, 1952; copy
wilh originals and casts.) SAM,

MARSUPIALS: Unidentified footprints, approx,
150 mm long.

62. BLACKFORD DRAIN 36°46'5,
(NARACOORTE).

21 km NE of Kingston, $.£. During placement of
bridge pylons, fossils recovered from }i depth of
“11-13 feet", north side of creek, in a bed of
waterworn stones. “Rock bottom” at 13 feet Was
“a hard stone which looked like u flow of black
mud, thickly impregnated with small white shells”.
(Letter from R, V. Flint with specimens.) SAM,
MARSUPIALS: Diprotodontitae: D/preradon sp,
Macropodidue; Macropus spp., Stltenurvs sp. ct,
Procoptodon sp.

1d0°OE

63, NARACOORTE, 36°59'S, [90°45 F 1 NARA-
COORTE),
SE South Australia, Cave Sites, SAM.

a, Henschke’s Bone Dig (U 91/97) Gutskirts af
Naracoorte at Hensehke's Quarry A diverse
assemblage, mostly marsupial. Differs in detail
from Naracoorte Caves deposits. (N. S, Pledge
ners, comm, 1979), SAM.

MONOTREMES: Tachyglossidae; Zaglassus rumt-
Saye,

BIRDS: Megapodiidae, Progura naracoortensis.
AMPHIBIANS: Hylidae; Litaria ewinsi,
Leptoductytidae; Limnadynastes — fasmantlensts,
Ranidella signiferd.

van Tets (19740), Tyler (1977), Pledge (in prep.),

b. James’ Quarry Cave [U 29), Naracoorte
township, UCMP.

MARSUPIALS: Thylacoleonidue: Thylacalee car-
nifex.
Daily (1960a)

fd. NARACOORTE CAVES 37°(1'S,
(PENOLA).

(5 km SE of Naracoorte, Numerous caves, ususlly
with red sundy cave fill containing fossils. SAM,
FUSA,

a, Alexandra Cave (1/3).
Old collections SAM. Recently collected: FUSA-
MARSUPIALS: Thylucoleonidae: Tlrylacolew rur-
nifece.
Macropodidue; Sthenurus cf. oceldentalis, Sthenn-
rus sp., Pracoptodon sp.

b. Brown Snake Cave (U 14),
SAM collection.
MARSUPEALS: Macropadidae, Vrienuries sp,

c. Cathedral Cave (U 12/13),
SAM callection,

14 48'F

MARSUPIALS: Thylacoleanidae, Thylucolea ear.
Nifex.

d, Dogs Prohibited Cave (U —-).
FUSA collection.
MARSUPIALS: Dasyuridac, Dasyirids sf,
Perumelidue.
Macropodidae; Porarons sp, Mucropiy sp,
EUTHERIANS: Muridac,
BIRDS: Unidentified material,

é. Fox Cave (U 22).
SAM collection.
MARSUPIALS: Dasyuridae: Desyverus sp. Sarees
phils sp.
Thylacinidue: Thylacinuy sp.
Peramelidac; Isacean sp,
Vombuatlidac: Varnbaius sp,
Phalangeridae; Trichosurus sp.
Petauridae, Prendocheirus sp.
Thylacoleonidae; Thylacalea earnifex,
Macropodidiue; Beitonyia sp., Pelorous sp, Macre-
pus cf, givantéus, M_ rufegrisens, Sthenueus sp., 8.
willl,
BIRDS; Unidentified material,

f. Haystall Cave (U 23).
SAM collection.
MARSUPIALS: Dasyuridae: Dasvurus sp. Saree-
plulas lantaritas,
Phascolarclidue; Phascolarctas sp,
Thylacoleonidaes Thivlacaled sp.
Macropodidaec: Macropus sp.. Stleaurus sp.
REPTILES: Scincidae; Viliqua sp.

g. Specimen Cave (U 35).
Previously known as Zietz Cave, SAM.

MARSUPIALS: Dasyuridue: Sareophiluy lari-
aris,

Thylacinidae; Thvlactiusy ct, major.

Peramelidae.

Vombatidae.

Thylacoleanidae: Tlvlacelee sp.
Mucropodidac; Macropus sp,, M. ef,
temiriodon ef, anak,

h. Tomato-Stick Cave (U 10/11).
SAM collection.

MARSUPTALS:
Protemnodon sp,

(ilan, Pros

Macropodidae: Marropus sp,

\, Victoria Fossil Cave (U 1).
SAM, FUSA collections,
MARSUPIALS: Dasyuridae: Dasyurns viverrinuy,
PD, maculatus, Sarcophilus sp., Antechinus Havipes,
A. stuartii., A. swainsonii, Sminthopsis crassi-
candata, S. murina.
Thylacinidaes Thylueinus evnocephaluy,
Peramelidue: Perameles sunnii, PP) bodeainville,
Isoodon abesulis.
Vombatidie; Vonthatits sp,
Phascolarctidac; Phaseolaretas sp.
Petauridae; Psendocheirns pereerintus,
hreviceps.
Burramyidae; Ceroarntus nanus,
Thylacoleonidae: Th ylacolee carnifex

Peanuts

112 D. L. G. WILLIAMS

Diprotodontidae; Palorchestes azael, Zygomaturus
trilobus.

Macropodidae; Bettongia penicillata, B. gaimardi,
Potorous apicalis, P. platyops, Macropus gigan-
teus, M. rufogriseus, M. greyi, M. eugenii, M.
titan, Wallabia bicolor, Protemnodon roechus,
Sthenurus andersoni, S. atlas, S. brownei, S. gilli,
S. maddocki, S. occidentalis, Procoptodon rapha,
small macropods.
MONOTREMES:
sp., Zaglossus sp.
EUTHERIANS: At least six species of rodent
(M. J. Smith in prep.).

BIRDS: Dromaiidae; Dromaius novaehollandiae.
Megapodidae; Progura naracoortensis, Leipoa
ocellata, indeterminate species.

Phasianidae; Coturnix pectoralis, C. australis.
Turnicidae; Turnix sp., T. varia.

Pedionomidae; Pedionomus torquatus.

Rallidae; Rallus philippensis.
Charadriidae; Peltohyas australis.
Scolopacidae; Tringa_ glareola,
wickii, Calidris ruficollis.
Platycercidae; Pezoporus wallicus.
Tytonidae; Tyto novaehollandiae.
Grallinidae; Grallina cyanoleuca.
Craticidae; Gymnorhina tibicen.
REPTILES: Boidae; Wonambi naracoortensis.
Elapidae; Pseudonaja cf. nuchalis, Notechis cf.
scutatus, Pseudechis cf. porphyriacus, Unidentified
group,

Varanidae; Varanus varius, V. gouldii.

Scincidae; Trachydosaurus rugosus, Tiliqua nigro-
lutea, cf. Sphenomorphus tympanum, Egernia cf.
whitei.

Agamidae; Amphibolurus cf, barbatus.
AMPHIBIANS: Hylidae; Litoria ewingi.
Leptodactylidae; Limnodynastes cf.
Ranidella signifera, Geocrinea cf. laevis.
Woods (1866), Smith (1971, 1972, 1976), van
Tets (1974a), van Tets & Smith (1974), Wells
(1975), Olson (1976), Wells & Nichol (1977),
Tyler (1977), Murray (1979), Wells & Murray
(1979), Wells (pers. comm, 1979),

65. PENOLA 37°23’S, 140°50°'E (PENOLA).
22 km NNW of Penola. Bones found in sinking a
well on the edge of a swamp. Whereabouts of
fossils unknown.

BIRDS: Dromornithidae; cf. Genyornis sp.
Stirling & Zietz (1896, 1900), Rich (1979).

66a. GLENCOE 37°41’S, 140°37'E (PENOLA).
22 km NW of Mt Gambier. Probably a cave depo-
sit, as fossils are white with red sediment adhering.
Possibly Glencoe West Cave (L77) or Glencoe
East Cave (L108). SAM.
MARSUPIALS: Macropodidae;
Sthenurus sp.

Tindale (1933).

66b. GREEN WATERHOLE
140°32’E (PENOLA).
22 km NW of Mt Gambier. Also known as Fossil

Tachyglossidae; Tachyglossus

Gallinago hard-

dumerili,

Macropus sp.,

(L 81) 37°44’S,

Cave. Fossils from surface of rockpile to a depth
of 15 m in water-filled cave. SAM, FUSA.

MARSUPIALS: Dasyuridae; Dasyurus sp., Sarco-
philus sp.

Thylacinidae; Thylacinus sp.

Phalangeridae; Trichosurus sp.

Thylacoleonidae; Thylacoleo carnifex.
Macropodidae; Bettongia penicillata, Propleopus
oscillans, Macropus sp., Osphranter sp., Protem-
nodon sp., Wallabia sp., Sthenurus gilli, S. mad-
docki, S. occidentalis,

EUTHERIANS: Muridae.

BATS: Unidentified material.

BIRDS: Dromornithidae; ?Genyornis sp.

A diverse assemblage of other birds.

Wells & Murray (1979), Wells & Williams (in
prep.), Rich & van Tets (in prep.), Pledge (in
prep.).

66c. MILLICENT 37°36’S, 140°21’E (PENOLA),
Far SE of South Australia. Fossils found “at a
depth of six feet below the surface, embedded in
peat mixed with shells” (Waterhouse 1882).
Accession card for SI states “18 in. below sur-
face”.

MARSUPIALS: Diprotodontidae; Diprotodon sp.,
Zygomaturus sp.

Waterhouse (1882).

66d. MT BURR CAVE (L 69/70) 37°32’S, 140°
27'E (PENOLA).

Far SE of South Australia. SAM.
MARSUPIALS: Macropodidae; Sthenurus sp.

66e. TANTANOOLA CAVE (L12) 37°43’S,
140°30’E (PENOLA).
Near Tantanoola. Cave
SAM.

MARSUPIALS: Dasyuridae; ?Sarcophilus sp.
Vombatidae; ?Phascolomys sp.
Phalangeridae; Trichosurus sp.
Macropodidae; Sthenurus — sp.,
roechus.

EUTHERIANS: Otariidae; Arctocephalus sp.
Muridae; Rattus sp., Hydromys sp.

Tindale (1933).

in old wave-cut cliff.

Protemnodon

67. MOUNT GAMBIER 37°50’S, 140°47'E
(PENOLA).

Far SE of South Australia,

a. Cave exposed by earthworks in Derrington
Street. SAM.
MARSUPIALS: Peramelidae; Perameles sp.
Phascolarctidae; Phascolarctos sp.
Phalangeridae; Pseudocheirus sp.
Thylacoleonidae; Thylacoleo carnifex.
Diprotodontidae; Nototherium sp.
Macropodidae; Bettongia sp., Sthenurus spp.

b. Cave, location unknown. BMNH.
BIRDS: Dromornithidae; Genyornis sp.
Stirling & Zietz (1896, 1900), Rich (1979),

PLEISTOCENE VERTEBRATE FOSSILS 113

c. Cave exposed by excavation in Gray Street.

SAM.
MARSUPIALS: Sthenurus spp.

d. Moorak 37°52’S, 140°47’E (PENOLA),
5 km south of Mt Gambier. Probably a cave

deposit. SAM,

MARSUPIALS: Thylacoleonidae; Thylacoleo sp.

Macropodidae; Sthenurus sp.

Adelaide area 53, 57, 58.
Alexandra Cave 64a.
Allenby Gardens 57a.
Arnold’s Bank 26.
Baldina Creek 37.
Billeroo Creek 17.
Blackford Drain 62.
Black Rock Gravel Pit 30,
Boolcunda Creek 22.
Brompton 57b.

Brothers Islands 3.
Brown Snake Cave 64b.
Buckalowie Caves 23.
Bundey 40,

Burra Creek 39.

Bute 38.

Calca 2.

Cannatalkaninna | la.
Cassidy Locality 7a.
Cathedral Cave 64c.
Channel Sand Locality 12a.
Chowilla 42.

Chucka Bend 56.

Clara St. Dora Cave 23.
Collinsville 35.

Cooper Creek 11.
Cootanoorina 9.

Croydon 57c.
Curramulka Cave 43a.
Curramulka Quarry 43b.
Cuttapirra Waterhole 11i.
Dempsey’s Lake 25.
Devon Downs 55.
Diamantina River 5, 6.
Dogs Prohibited Cave 64d.
Echunga 59.

Emu Four Hole Cave 48b.
Findon 57d.

Fossil Cave 48c,

Fox Cave 64e.

Fromm’s Landing 54.

Acknowledgements

The author is indebted to Dr R. H. Tedford
for additional data on the L. Eyre region, and
to Ms K. Mackenzie for typing the MS.

68. 72TANKSTAND CAVE (L 65) 37°57’S, 140°
40’E (PENOLA).

3 km west of Mt Shank; ‘Sec 823, Hd Macdon-

nell, Co Sturt” (SAM Catalogue). However, this
should probably be County Grey, where there are

numerous caves (Hundred Map, H. J. Wall, Govt

Photolithographer, Adelaide, 1960),

MARSUPIALS: Macropodidae; Sthenurus gilli.

Alphabetical index to sites

Gawler 52.

Glencoe 66a.

Goolwa 61.

Goyder’s Lagoon 6,
Green Bluff Locality 7d.
Green Waterhole 66b.

Hallett Cove 58.
Haystall Cave 64f.

Henschke’s Bone Dig 63a.

Hergott Springs 13.
Hillpara Creek 27.
Hog Bay River 49.
Hookina Creek 19.

Innamincka 8.
James’ Quarry Cave 63b.

Kapunda 50.

Katpiri Waterhole 11i.
Kelly Hill Caves 47.
Kingston S.E. 62.
Kirkaldy Pit 57e.
Kyancutta 4.

Lake Callabonna 16.
Lake Fowler 44.

Lake Kanunka 10a.
Lake Palankarinna 12.
Lookout Locality 7e.

Mairs Cave 23.
Malkuni Waterhole 1 If.
Manunda Creek 31.
Marcus Locality 7c.
Marree 13.

Millicent 66c.

Moorak 67d.

Morgan 41.

Mount Burr Cave 66d.
Mount Eyre 20,

Mount Gambier 67.
Mount Taylor Cave 48a.
Mullett Locality 12b.

Murnpeowie 15.
Murrapaterinna 10b.

Naracoorte 63.
Naracoorte Caves 64.
Nectar Brook 28.
Newikie Creek 36.
New Kalamurina 7b.
Normanville 60a.

Pandie Pandie 5.

Peake 9,

Pekina Creek 29.

Penola 65.

Pernatty Lagoon 18.
Pondalowie Bay 45.

Pt Pirie Gravel Pit 32.
Punkrakadarinna Soakage 7f.

Reedy Creek 62.
Rocky River 46.

Salt Creek 60a.
Seton Rock Shelter 48d.
Specimen Cave 64g.

Tankstand Cave 68.
Tantanoola Cave 66e.
Teetulpa 24.

Thebarton 57f.
Tomato-Stick Cave 64h.
Town Cave 43a.

Two Wells 51.

Ulooloo 34.

Unkumilka Waterhole 1 1b.
Victoria Fossil Cave 641.
Warburton River 7.
Waupunya Creek 33.
Weekes Cave 1.
Welcome Springs 14.
Willochra Creek 21.
Yalpara Station 27.
Yankalilla 60b.

Yatala 53.

Note added in proof.
More detailed lists of fossil birds are to be

found in P. V. Rich (1975) Antarctic dispersal
routes, wandering continents and the origin of
Australia’s non-passeriform avifauna. Mem. Nat.
Mus. Vict. 36, 63-126.

Id D, -, G. WILLIAMS

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Proceedings, {hid A,

STRATIGRAPHY AND DEPOSITIONAL ENVIRONMENTS OF THE BILLY
CREEK FORMATION (CAMBRIAN), EAST OF THE FLINDERS RANGES,
SOUTH AUSTRALIA

BY P. S. MOORE

Summary

Two new members (the Coads Hill Member and the Erudina Siltsone Member) are defined in the
Billy Creek Formation at Reaphook Hill. The Coads Member consists of a complex sequence of
shale, sandstone, carbonate and tuff which was deposited mainly in a shallow marine environment.
The overlying Erudina Siltstone Member consists of a silty and sandy redbed sequence with minor
dolomite and was deposited on tidal mudflats during a period of regression. A tentative correlation
of the Billy Creek Formation between Reaphook Hill and the type section in the Wirrealpa Basin is
suggested. The Billy Creek Formation also occurs in the Yalkalpo 2 well, east of Lake Frome,
where it comprises mainly green shales and sandstones. Deposition most probably occurred in a
relatively open marine environment, which suggests that the Arrowie Basin extended for a
considerable distance east of the present Flinders Ranges in the late Early Cambrian. The Billy
Creek Formation in the Lake Frome wells Nos 1 and 2, south of Lake Frome, considts of shaly and
silty redbeds with common anhydrite, and was deposited on high tidal mudflats laterally adjacent to
detaic sediments of the Eregunda Sandstone Member.

STRATIGRAPHY AND DEPOSITIONAL ENVIRONMENTS OF
THE BILLY CREEK FORMATION (CAMBRIAN),
EAST OF THE FLINDERS RANGES, SOUTH AUSTRALIA

by P. S. Moore*

Summary

Moort, PS, (1980) Stratigraphy and depositional environments of the Billy Creek Formation
(Cambrian). east of ihe Flinders Ranges, South Australia, Trans. R. Yoo. S$. Alust, 14(5),
117-132, 28 November, 1950.

Two pew members (the Coads Hill Member and the Erudina Siltstone Member) are
defined in the Billy Creek Formation at Reaphook Hill. The Coads Hill Member consists of
a complex sequence of shale, sandstone, curbonale and lui which was deposited mainly ina
shallow marine environment, The overlying Eradina Siltstone Member consists of a silty and
sandy redbed sequence with minor dolomite and was deposited on tidal mudflats during a
period of regression, A tentative correlation of the Billy Creek Formation between Reaphook
Hill and the type section in the Wirrealpa Basin is suggested. The Billy Creek Formation also
sceurs in the Yalkalpo 2 well, east of Lake Frome, where it comprises muinly green shales and
sandstones. Deposition most probably occurred in a relunvely open marine environment, which
suggests (hat the Arrowie Basin extended for a constderable distance cast of the present
Flinders Ranges in the late Early Cambrian, The Billy Creek Formation in the Lake Frome
wells Nos | und 2, south of Lake Frome, consists of shaly and silty redbeds with common
anhydrite, and was deposited on high tidal mudfluts laterally adjacent ta detaic sediments of

the Bregunda Sandstone Member.

Introduction

The Billy Creek Formation consists of an
Early to Middle Cambrian, predominantly
redbed sequence of shale, siltstone and sanc-
stone, with minor limestone, dolomite and tuff.
It crops out sporadically throughout the central
und northern Flinders Ranges of S.A., and has
been identified in the subsurface below the
Cainozoie and Mesozoic of the L. Frome
region (Fig. 1).

The Billy Creek Formation was formally
defined by Daily (1956), Outcrops in the
central and northern Flinders Ranges have
recently been subdivided into three members
by Moore (1979b). after redefinition of the
status of the Edeowie Limestone Member
(Moore 19794). However, these three mem-
bers are not recognised at Reaphook Hill or in
the subsurface oecurrences to the east of the
Flinders Ranges, due to the very individual
wature of the sequences, This paper discusses
the stratigraphy and depositional environments
of the Billy Creek Formation at Reaphook Hill,
and also briefly discusses the subsurface duta,
gathered primarily from the Yalkalpo 2 bore-
core, to the east of L, Frome.

The Reaphook Hill Outerop
The Billy Creck Formation at Reaphook
Hill is subdivided into two members. hereih

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Fig. 1. Quterop locality map, eastern Flinders
Ranges and L. Frome Embayrment.

termed the Coads Hill Member and the Eru-
dina Siltstone Member (Fig. 2). The lower
sandy partion of the sequence (the Coads Hill
Member) was originally considered to be part
of the Hawker Group, comprising Bunkers
Sandstone equivalent and Oraparinna Shale
(Dalgarno & Johnson 1963), However, Dal-
garno (1964) recognised tuffaceous and red

Ls PS, MOORE

Reaphook
4 Hy ll

Erudina Siltstone
Member

“| Goads Hi)!
Member

NM Wilkawillitna
Ps Limestone
Pig. 2, Ourcrop nmap and locution of measured

stratigruphic sections, Billy Creek Pormation,
Reuphook Hill.

Rec setratigraphic
Coon ce ction

ee Fault

Billy. Creek
Formation

Geological
boundary

So Creek ‘ea

silty intervals in the sequence, and redefincel
it as part of the Billy Creek Forjnation, as
shown on the PARACHILNA 1:250 000 geo-
logical sheet (Dalgarno & Johnson 1966).
The upper part of the Billy Creek Formation
at Reaphook Hill (the Erudina Siltstone
Member) is dominated by silty redbeds, with
minor dolomitic atid sandy intervals.

The Reaphook Hill region was mapped in
detail in 197) by Gaunt! and Gehling?. Their
discovery of Emuellid trilobites in che lower
portion af the Billy Creek Formation is of

1Gaunt G. FM. (1971) The geology of the

Kempes Bore urea, eastern Flinders Ranges.
B.Sc. (Huns.) thesis, University of Avelaide
(unpublished }

> Gehling, J. G. (1971) The geology of the Reap-
hook Hill area. Flinders Ranges, South Australis,
B.Sc. (Hons) thesis, University of Adeluide
Uinpublished).

particular interest and the author is indebted
fo these two authors, whase work provides the
basis of the subdivisions presented below,

Stratigraphy of the Coads Hill Member

Introduction

The lower part of the Billy Creek Forma.
tian at Reaphook Hill comprises a sequence of
interbedded, fine to medium-grajined, pale
brown sandstone, dark grey limestone, and
minor red and green shale and shaly sillstone.
Interbeds of calcareous shale, shaly limestone,
dolomite and tuff occur in some units, and a
limestone-boulder conglomerate occurs at the
base of the sequence in the northern outcrops
(Fig. 3), The sequence is herein termed the
Coads Hill Member. The name is derived from
“Coads Hill” which is located approximately
7 km W of Reaphook Hill, Seclion RH-C
is chosen as the type section (Fig. 4).

The hase of the Coads Hill Member

In the north of the Reaphook Hill regian,
the basal 6 m of the Coads Hill Member com-
prise boulder conglomerate, with clasts of
Imestone wp to 30 cm across. The con-
glomerate was mapped as part of the under-
lying Hawker Group by Gebling*, hawever
it rests sharply and uncooformably on pale
grey, fenestral and oolitie Wilkawillina Lime-
stone. Further south, calcareous sandstones and
siltstones af the Coads Hill Member rest clis-
conformably on the Wilkawillina Limestone
(Figs 3 & Sa), A pisolitic calerete horizon,
5-20 ern thick, caps the disconformity surface,

Internal Stratigraphy

The Coads Hill Member is divided into mne
units (Fig. 3), which are deseribed belaw-
Units B to G are essentially the same as unils
recognised by Gaunt! and Gchling®.

Unit A is the basal unit of the Coads Hill
Member in the rorth of the Reaphook Hill
renion. TL comprises cobble to boulder coo-
glomerate, with clasts of pale grey, micritic,
fenestral and oolitic limestone and dolomitic
limestone, up to 30 cm across, The conglo-
merate has a closed fabric, with the matrix
comprising medium to yery coarse sand-sized
quartz and minor carbonate. Clasts are gene-
rally subangular to subrounded. Some are
fossiliterous, containing unidentified fragments
of trilobites and other shelly debris. The clast
lithologies are very similar to underlying Wilka-
willina Limestone

BILLY CREEK FORMATION EAST OF FLINDERS RANGES

Ed fark grey limestone Minor dolomite

Dark arey calcareous siltstone & sity |imestone

Shate imostly green) with calcareous interbeds,

Red, clayey sandstone and clayey selistoang, Minor srale,

Reddish brown taltareous sanastone, Minor shaly intecbeds

herizontal
Scale

Grey, limestone rich, cobble fo houlder congiemerate

Fig. 3. Stratigraphy of Coads Hill Member, Billy Creek Formation, Reaphook Hill.

Unit B is the basal unit of the Coads Hill
Member in the south of the Reaphook Hill
region, and conformably overlies Unit A in the
north, If comprises pale red to reddish brown,
fine to medium grained, feldspathic sandstone
(Fig, 5b). Interbeds of greyish red shale and
shaly siltstone are common in the upper por-
tions of the unit, The sequence is evenly bedded
to ripple laminated on the scale 3-15 cm.
Small to medium seale tabular cross-stratifica-
tian is common in the thicker sections, Desic-
cation cracks, symmetrical ripples, mudstone
intraclasts and small scour-and-fill structures
are common throughout the unit and worm
hurrows, interference ripples and pebbly hori-
zons occur in some outcrops. A 5 cm thick,
bright green tuffaceous interval with devilri-
fied shards occurs in the middle portian of
Section RH-H, in the north of the area,

Unit C comprises greyish red, very poorly
sorted, shaly siltstone to silty sandstone, with

minor granule and pebble-rich bands (Fig.
Sec). Ripple laminated interbeds of moderately
sorted, reddish brown sandstone, 5-20 cm thick,
are common in the lower portion of the
sequence, and define a passage from Unit B.
Mudstone intraclasts and desiccation cracks are
abundant (Fig. 5d). Bedding is poorly defined
in most of the unit, although the upper few
metres are dominated by red silty shale with
well developed, even lamination,

Unit D comprises a thin tongue of pale
yellowish brown, moderately sorted to well
sorted, feldspathic sandstone (Fig 3Se) which
crops out in the southern portion of the area,
where it resis conformably on Unit C (Fig.
3). The sandstones are evenly bedded on the
scale of 3-12 em with some poorly defined
ripple laminations and rare symmetrical tipple
marks, Large scale cross-stratification is absent.
A thin, bright olive green tuffaccous mterval
crops out in the upper portion of the unit in

12) r § MOORK

Section RH-G Green shale intraclasts and
incipient shrinkage cracks wre abundant
throughout the sequence.

Unit B is a sequenee of dark grey, foetid,
silty limestone, caleareous shale and siltstone
(Fig, Sf), In the thicker sections, a well-
developed sequence ranges from buflf-coloured
stromatolitic dolomite at the base, through
burrow-mottled shaly grey limestone and cal-
vareous shale in the lower portion, into dark
grey foctid limestone in the middle and upper
portion. Much of Unit E is bioturbated, and
desiccation cracks occur sporadically through-
our the sequence,

Unit F comprises pale reddish brown to
brown, fine fo medium-grained, feldspathic
sandstone. Interbeds of shale and siltstone are
common in some oulerops. Several bright
olive green tuffaccous intervals up to 30 em in
thickness are present, and are particularly pro-
minent in the wpper shaly portion of the umit.
The sequence is generally evenly bedded to
ripple laminated. Wowever, medium = seale
tabular cross-stratification is Common an the
60 northern outcrops (Fig. Sg). Mud-cracks,

— yuurtz-lined geades, oscillation ripples, current
ripples, mudstone intraclusts, small scour-and-
40 fill structures and worm burraws are common
throughour. Interference ripples occur in some
outcrops, In the north, the sandstones are
relatively mature, and are partly cemented
20 by calcite. Further south, the sandstones are
z interbedded with red shale and siltstone, and
contain rare pebble beds,

Unit G comprises green shale und calcy-
reous shale, with common, thin interbeds of
shaly, dolomitic limestone and dolomite.
Minor red shaly intervals oceur in the lower
portion of the unit, especially in the south,
Bright olive green tuffaceous interhéds, rarcly
up io 1.4m thick, are common throughout the
Type section (RH-C) of Coads Hill Mem- S¢4uenee. Units are generally evenly laminates,
. Billy Creek Formation. See legend Fix 8 although rare asymmetrical ripple marks are

BO

Medion
very
toarse

Wilkawillina
woh

*

Fig
be

=

Fig. 5: a. Red shaly siltstones and sandstones of Unit B, Coads Hill Member, draping irregular discon-
formity surface al top of Wilkawillina Limestone. Hammer: 3] cm Jong. Location: Section RHC,
Reaphook Hill, b. Evenly bedded to cross-siratitied reddish brown siltstones and sandstones of Unil
B, Coads Hill Member. Hommer: 3) cm long. Location: Section RH-D, Reaphook Hill. c. Small
sundy channel in very poorly socted red shaly siltstone of Unit C, Couds Hill Member, Scale: 34
mm diumeter. Location: Section RH-A, Reaphook Hill. d. Incipient shrinkage cracks in fine red sand-
stone of Unit D, Coads Hill Member. Scale; 54 mm diameter, e Typical outcrop of Unit D, Couds
Hill Member, showing prominent ridge of evenly bedded and rarely. ripple Jaminated, mediur
grained satidstone. Locatian: Section RH-A, Reaphook Hill, f. Prominent ridge of grey, fuetid, silty
limestone and culeareous siltstone. Basal unit is lighter°coloured and dolomitic. Location: Unit &,
Conds Hill Member. Section RH-E, Reaphook Will. g. Cross-strutifieation in reddish brown cal
carcous sandstones of Unit Fy Coads Hill Member Note pbundance of mudstone intraclasts con-
conlrated on foresels of cross-strata. Location: Sevtion RH-F, Reaphook Hill, h. Basal view of
yellowish brown shaly dolomite containing abundant halite casts, Scale: 54 mm diameter. Location:
Unit H, Coads Hill Member. Section RH-C, Reaphook Hill.

BILLY CREEK FORMATION EAST OF FLINDERS RANGES 121

122 P. S. MOORE

a pes TY
a7
te

BILLY CREEK FORMATION EAST OF FLINDERS RANGES |23

present in-sitty intervals in the south. Desic-
cation cracks and halite impriatts are econmen
in some sections (Fig. Sh), Abundant trilobites
(Baleoracania dailyi Pocock) and rare; uniden-
tified bracthopod fragments are present in
green Shale overlying dolomite, approximately
43 m above the base of Unit G in Sechoan
RH-A.

Unit H comprises 2 sequence dominated by
dark grey, foetid limestone, Interbeds of shily
limestune are enmmmon, and much of the se-
quence has 9 well-developed nodular (letsoid)
texture (Pig, 6a). The upper and lower par-
tions of Cinit Hoare shaly and dolomitic, and
contain stromatolites (Pig. 6b), desiccation
cracks and Ininor halite imprints. Unidentified
trilobite fragnents are uncommean, but huve
heen recarded From the middle to Upper por-
tien of the sequence

Unit J comprises approximately 3 m_ of
every laminated, khaki shale and tine siltstone,
with minor carbonate bands and nodules, The
trdobite B. deilyé is abundant in the basal
portion, anid is assaciated with care, unidepti-
fied brachiopod fragments. A 0.5 m_ thick
peloidal und aleal, mottled limestone forms a
prominent marker at the top of Unit J,

Age and Palaeontology

Trilobite traeks, worm burrows and mol-
luscan trails oceur sporadically throughout the
Couils Hill Member. At least one type of trace
fossil is present in every unit, with the excep-
tion of Unit C, Emvellid trilobites were first
discovered by Gaunt? and Gehling® in what
is now defined as the basal portion of Unit J
of the Cowds Hill Member, B, Daily (pers,
com jt Gehling’. p. 16) identified the trilo-
bites as B. dati. The species alsa oceurs in the

upper partion af the White Point Conglomerate
on Kangaroo Tsland, where it has been assigned
a late Barly Cambrian age (Pocock 1970),

Depositignal environitent of the Coads Hill
Member

Following deposition of shallow marine
and stpratidal carbonates of the uppermost
Hawker Group. the Adelaide “Geosyneline” in
the vicinity of Reaphook Hill was uplifted and
the Hawker Group eroded. Deposition of the
Coads Hill Member of the Billy Creek Forma-
Lion commenved when the area once again be-
came submerged. Limestone boulders eroded
from nearby areas were deposited in a near-
shore marine environment io the north (Unit
A) while a thin calerete profile developed on
the land surface to the south. Subsequently,
a sequence of shallow murine to intertidal,
caleareous sandstones (Unit B) spread over the
area,

Red, shaly and pebbly sandstones (Unit C)
are considered to be non-marine in origin.
because of a lack of fossils and their traces,
and also the extremely poor sediment-sorting.
Thus, a likely environment of deposition for
Unit © is a muddy alluvial plain, and its
presence in the Reaphoak Hill area indicates
a period of marked local regression.

The origin of the lower, foetid, shaly car-
bonute (Unit B) is of particular interest, since
it is underlain by redbeds which are probably
non-marine (Unit CY, and overlain by shallow
marine to intertidal calcareous sandstones
(Unit F). The lamination and fine grain-size
of Unit E indicates depositian from suspension
in a low energy environment. The lack of body
fossils Sugeests restricted marine conditions,
while the presefree of laminated dolomicrite,

Fie. 6; 9. Nodular (leno), shaly limestones of Unit H, Coads Hill Member, Hammer: 31 ¢m
long. Location: Secrion RH-H, Reaphook Hill b. Interbedded grey calcareous shale aud shaly lime-
stone of Unit HW, Conds Hill Member, Note presenve of weathered-out stromutolites in centre of
photo. Stromatolilte elongation perpendicular 1o outwrop. Hammer 31 cin Tons, Location: Section
RH-F, Reaphook Hill. c. Lurge sealy shuly-curbonate cycle in Unit A. Ertdina Siltstone Member. Red
shales pass grudationally through green shiiles and greyish green calcareous shiles into pule grey
to hyi-caloured dolomftic limestone. Rapid regression buck into red shale is vpical of cycles. Ham-
mer: J} cm long, Location: Section RH-C, Reaphook Hill. d. Thin unit of vellowish brown shaly
dolomite with wavy stromatolitic laminations in niiddle of asymmetrical shale-dolomite-shale cycle.
Scate; 54 mm diameter. Location: Unit A, Erudina Siltstone Member, Section RH-C, Reaphook Hill,
ce Wovy and Jenticulur bedding in shaly siltstones of Unit D, Erudina Siftstone Member, Mudfinke in-
trackisrs are common, Location: Section RH-C, Reuphook Hill. f. Current linented red micaceous
sundscones of Unit C. Erudina Sillstone Member, Scole: 54 mm diameter. Location: Section RH-C.
Reaphook Hill 2. Large lood slirictures with subvertical syfometry, probably associated with
Jewalering. Hammer: 31 em lone Location: Unil ©, Briudina Siltstone Meniber, Section RH-C,
Renphook Hill. bh. Semimetrical wave ripples in coarse red silfstone. Bedding surface dissected by.
large, palvgonul desiceation chucks inflled with red mudstone, Lenscup scule: 54 mm diameter,
Location: Section RH-C. Reaphaok Hr).

124 P. S. MOORE

top Aot exposed

=|

MEMBER

uJ
—_
oO
=
w
—
—!

ERUO |

medium
very

Coads Hill
sno}

Fig. 7. Type section (RH-C) of Erudina Siltstone
Member, Billy Creck Formation, Reaphook
Hill, See legend Fig, &.

stromatolites and desiccation cracks indicates
that af Jeast part of the sequence Was deposited
in shallow water, The foetid odour and dark
colour of the bulk of Unit E is evidence for
restricted circulation and poor oxygenation,
such as occurs in a restricted embayment or
lagoon. Unit D sandstones, which are laterally
equivalent to Unit E, probably represent de-
position in a littoral environment

Shallow marine calearcous sandstones of
Unit F pass gradationally into micaceous
shales of Unit G, The shales contain arthropod
tracks and dolomitic intervals, and thus were
Jeposited on subtidal to intertidal mudtlats
subject to frequent marine inundation and
reducing conditions,

The upper, dark, foetid limestone (Unit H)
is somewhat different from the lower one, for
it contains minimal terrigenous clastic detritus,
and is relatively uniform in thickness and
character throughout the area of outerop,
Thus, Unit H probably accumulated in a semi-
restricted, very shallow marine environment,
subject lo only minor fine-grained terrigenous
influx. A coanection with the open sea is in-
ferred from the presence of trilobites, however
the water was generally quite shallow, as indi-
cated by sporadic desiccation cracks and
stromatolitic intervals, and was also poorly
oxygenated, as indicated by the high organic
eontent of the limestone, its dark colour and
its foetid odour.

The thin sequence of green fossiliferous
shale (Unit J) which occurs at the top of the
Coads Hill Member indicates a period of fine-
grained clastic deposition in a relatively open,
shallow marine environment, A rapid decrease
in fossil fragments towards the top of the unit
indicates progressive restriction and shaflowing
of the basin, prior to the deposition of the
Erudina Siltstone Member.

Stratigraphy of the Erudina Siltstone Member

fatroduction

The upper portion of the Billy Creek For-
mation at Reaphook Hill comprises a sequence
of greyish red siltstones and silty shales, with
minor dolomitic, tuffaceous and sandy iiter-
beds. The sequence is herein termed the Eru-
dina Siltstone Member, The name is derived
from the Erudina homestead, located approxi-
malely 15 km SE of Reaphook Hill.

The Erudina Siltstone Member crops out i
the central portion of a small basinal structure
at Reaphook Hill (Fig. 2), where it conform.
ably overlies the Coads Hill Member, Maxi-
mum measured thickness is 270 m in Section
RH-C, however the original thickness is un-
known since the upper portion of the member
has been removed by erosion, Section RH-C
as chosen as the type section (Fig. 7), The
member is clearly divisible inty four units,
described helow,

BILLY CREEK FORMATION EAST GF FLINDERS RANGES \25

Internal Stratigraphy

The basal unit of the Erudina Siltstone
Member (Unit A) comprises a fine-gcained
sequence of cyclically interbedded silty shale
and carbonale. Halt-cycles, from red shale
through pale greyish green shale info buf
coloured dolomite or dolomitic limestone are
typical (Fig. Ge). The carbonates are cyenly
laminated to wavy laminated, with rare desic-
cation cracks, hahte imprints and stromatolites
(Fig. Gd). Shaly and silty clastic intervals
commonly contain desiccation cracks and
symmetrical ripple marks, Raindrop imprints.
calcareous algal mats, halite imprints, worm
burrows and arthropod tracks and trails are
rare,

Unit B comprises approximately 80 m of red
shale. silty shale and shaly siltstone, with
minor sandy interbeds. The sequence is evenly
laminaled, with minor wavy bedding (Fig. 6e)
and ripple lamination in the coarser units.
Symmetrical and near-symmetncal (wave-
formed) ripples predominate, although highly
asymmetrical (current formed) intetference
ripples, and flat-topped ripples also occur.
Desiceation cracks and mudstone intraclasts
are abundant, whereas halite casts and small
load structures are relatively uncommon. The
redbeds are Weakly calcareous, and crenulated,
carbonale-rich algal mats occur in a few lovali-
ties, Arthrapod tracks and biotucbated inter-
vals are rare. Pink. silty, tuffaceous units ate
also rare,

Unit C comprises a 40 m thick sequence
ol interbedded shaly, silty, and sandy redbeds.
Coarsening-upward cycles are present, and vary
in thickness from a few metres to twenty
metres. The silty units are wavy bedded to
ripple laminated, and are commonly associated
with desiccation cracks, mudstone intraclasts,
trilobite tracks, worm burrows, and symmet-
rical interference and current ripples. Sandy
intervals are commonly ripple laminated, with
rib-and-furrow structures on the upper surface.
Thicker intervals in the upper portion of Unit

hake Frome Se ae
wells

C are horizontally laminated, with well-
developed current lineation (Pig. 6f), current
crescents and rare bounce marks. Trilobite
tracks and scratch marks are common in the
sandy nits, which also contain desiccation
cracks, mudstane intraclasts, load structures
(Fig, 62), rare worm burrows. and molluscan
trails.

Unit D comprises a 100 m thick sequence of
interbedded red silty stiale, shaly siltstone and
silislone very similar in character to Unit B.
Silty intervals ure wavy bedded, flaser bedded,
and ripple laminated, and both symmetrical
and asymmetrical ripple marks are abundant
throughout the sequence (Fig. 6h). Desicca-
tion cracks, halite imprints. and mudstone intra-
clasts are common. A few units contain inter-
ference and flat-topped ripples.

Palaeonfalney
Worm burrows, molluscan trails and tracks
attributed to trilobites occur sporadically

throughout the Erudina Siltstone Member. The
only body fossils found to date are tiny (1-2
mm long) carbonaceous imprints in green
shale in the lower portion of Unit A. These
are interpreted as fossil annelids.

Depositional environment of the Erudina
Siltstone Member

The Erudina Silistane Member consists of
four units which are distinguished primarily
on the basis of grain-size, Unit A is fine-
grained, comprising red shale with dolomtic
interbeds. The shales were deposited in an
oxidising environment on muddy tidal fats
probably as a response to mild tectonism (the
Kangarooian Movements of Daily & Forbes
1969). Carbonate mudstones accumulated in
the lower intertidal to subtidal environment
during periods of relative tectonic quiescence.
Cycles in the shale-carbonate sequence of
Unit A are attributed to local transgressions
and regressions, and reflect the unstable nature
of the basin of deposition and adjacent source
areas during this period,

Mount
Arrowsmich

sea level
-sbom
-1ooom

-1500m

Fig, 8. Simplified cross-section, castern Arrawie Basin, See Fig | for location and legend.

126

Unit B was deposited in response to in-
creased tectonic activity, whereby red shales
and siltstones were deposited on muddy inter-
tidal flats and in the shallow subtidal environ-
ment. The rate of sedimentation was sufliciert
to obscure carbonate accumulation and in-
stead, a sequence of fine-grained redheds with
distinctive tidal stratification (cf. Reineck &
Wunderlich 1968) was developeu..

During the deposition of Unit C, sand was
carried into the basin, forming coarsening-
upward cycles of redbed clastics. The eycles
are attributed to pulses of tectonise which
reached a peak late in the history of deposition
of Unit C. Unit D represents a return to same-
what more stable conditions, as experieticed
during the evolution of Unit B. Fine-pgramed,
shaly and silly redbeds dominate the sequence,
which contains an abundance of poorly defined
simple and wavy flaser bedding.

Subsurface distribution of the Billy Creek
Formation

The Billy Creek Formation occurs sub-
surface in the Arrowie Basin to the east of the
Flinders Ranges, wenerally below the Mesozoic
of the L. Frome Embayment and in some
places below the Cainozoie of the Tarkaroo-
low Basin®, The present limits of the Cambrian
basin in this region, as suggested by Youngs',
are shown in Figure 1. A simplified cross-
section is presented in Figure 8.

* Osborne first introduced the tern “Frome Em-
buyment’ defining it as a aynelinal basin
bounded by (he Flinders und Barrier Ranges, As
presenuly defined (Wopfner 1969). the term
‘Frome Embayment" refers only to the Mesazoic
sedimentary basin. The overlying Ciuinozaic
sediments of the Tatkaroolod Basin! are ungon-
Formuble an the Cretaceous and relute lo u dif-
ferent cycle of events.

Youngs, BC. (1969) Bumburlow |—well com-

pletion report, S. Ausi. Dept Mines & Energy

(unpublished )-

Osborne. N. (1945) Report on ail and gas pos-

sibilities of the Frome Embaoyment, New South

Wales and South Australia, for Zine Corp. Ld.

8. Aust. Dept Mines & Energy open file report
(unpublished).

Callen, R.A. (1976) 1,250 000 geologieal series
explanatory poles, FROME, South Australia.
S, Aust. Dept. Mines & Energy Rept 74/27
funpublistied }.

* Dethi Australin Petroleum bid, and Santos Lid
(1969) Well completion report, Lake Frome
1, 2 & 3, 8S. Aust. Depr Mines & Energy open
‘ile envelope 968 tunpublished),

“Youngs. B.C, (1977) Mudguard t and Yalkalpu
2—well completion reports. S. Aust. Dept
Mines & Energy 77/66 (unpublished).

~

=

PS MOORE

As shown in Figure 8, the Billy Creek For-
mation was intersected by L, Frome strati-
graphic wells Nos, | & 2% and SADME Yal-
kalpo No, 2% The EAR uranium exploration
holes south of L. Frome mostly ended in
weathered and leached, shaly redbeds®, many
of which probably belong to the Billy Creek
Formation. An interpretive subcrop map for
the area south of L, Frame is presented in
Figure 9, hased on ihe aufhar’s own evaluation
of cores and cuttings.

SADM Yalkalpa No. 2

The Billy Creek Formation in Yalkalpo 2
comprises red and green shales and sillstones,
with common reddish brown sandy intervals*.
Approximately 265 m of strata attributed to
the lower portion of the Billy Creck Formation
were intersected (Pig. 10), The original total
thickness of the formation at this locality is un-
known, since the Upper portion of the sequence
has been removed by post-Cambrian, pre-
Cretaceous erosion (Youngs 1978),

Feat oe
ay

|

¥
w
te
rey)
3
2
Fa
m
a

Precambren ceystadine

BS hassmnnt

POLARY aboce
cen SS gy ¢
¢ ‘5 bg :

Tertiary and Quaternary
strata
re
| i ireeh FY Mation

Fa > bate Precambs sah and
“oe stay Gambrien strata

{i SH esa rete ny St 2\ ene
al sasement “yo DS

Fig. 9, Interpretive preCretaccous subcrap mup..
south of Lo Frome, Evidence suggesis only
thin pro-Billy Creek Formation clastic sequence
in this area. with probable faulting against
Olary Block.

—— Geologic eoundaty

—— Faull | ntlerrec!
wt Leke Frome writ
+ FA Ragd welis

™ Giece Dak Wore

BILLY CREEK FORMATION EAST OF FLINDERS RANGES

A moderate to high sand content, and a
dominance of nan-red sediments differentiates
the Yalkalpu 2 sequence from most other
known occurrences of the Billy Creek Forma-
tion, and the absence of carbonates differen-
liates it from the Caads Hill Member, Thus,
the Yalkalpo 2 sequence between 258 m and
523 m depth is defined as. Billy Creek Forma-
lian sensu sirieta.

Fine-grained intervals dominate the se-
quence, particulafly in the upper portions, “A
faeies spectrum is present, from ripple
laminated fine-grained sandstone wih green
shale Nasers, through wavy flaser and lenticu-
lar bedded units (ef. Reineck & Wunderlich
1968, and Reineck & Sinuh 1975) into evenly
Jaminated green shale (Fig. 10), Redbeds com-
prise only abaut 30% of the fine-grained
association and generally consists of wavy
bedded to evenly laminated shales and
coarse siltstones, Worm burrows are common,
and oceur mainly in the green intervals. Mol-
luscan trails and tracks attributed to trilobites
occur both in red and green coloured sediment,
although they are slightly more common in
the latter. Desiccation cracks are abundant in
many of the red shale intervals, and in rare
cases are ussociated with halite imprints. and
patches and veins of anhydrite, Cyclic sedi-
mentation between red and green intervals,
each with its characteristic set of sedimentary
structures and organic markings, is a feature
of the middle portion of the sequence
(especially in the intervals 380-450 m).

Coarse-grained intervals comprise reddish
brown to yellowish brown, very fine to
medium-grained feldspathic sandstone, In
some cuses (e.g. at 497 m) the sandstones rest
sharply on fine-grained sediments, and contain
wbundant mudstone intraclasts in the basal por-
tions of the units. More commonly however,
the sandstones oceur at the top of coarsening-

Rudd, B.A. Pry Lid (1970) Report on mvesliga-
tions, Take Frome Embaymenr, S.A. SMI’s 267
and 268. §. Aust. Dept Mines & Energy open
file envelopes 1109 & 4170 (unpublished).

W Daily, B. (1969) Remarks on the subsurface
strativraphy and palaeontalogsy of the Delhi-
Santos Lake Frome Nos 1-3 Stratigraphic Wells
5. Aust. Dept Mincs & Energy open file
envelope 968 Cunpublished )-

1) Moore, P_S. (1979) Stratigraphy and sediment
ology of the Billy Oreck Formation (Cambrian.
Flinders Ranges) and is equivalents on the
noriheast coast Of Kangaroo Island, South
Australia. Ph.D, thesis, University of Adelaide

Cunpublisheett,

L232

upward sequences, They are generally evenly
bedded, with abundant soft-sediment deforma-
tion i the thicker units. Some of the thinner
sandy units are ripple laminated. and green
und minor red flasers may be present, Mud-
stone intraclasts are common. A feature of the
sandy intervals is the absence of well-
developed large-scale cross-stratifieation.

Delhi-Santos Lake Frome Stratigraphic Wells

The Billy Creek Formation sensu stricto has
been identified from the L, Frame wells Nos.
lund 2 by Daily! This identification is con-
firmed here, Only the upper part of the Billy
Creek Formation was penetrated?, and this
interval is considered to be the approximate
lateral equivalent of the Eregunda Sandstone
Member in the central and nonhern Flinders
Runges (Moore 1979b), However, the interval
is not appreciably sandy and thus should not
be referred to as the Eregunda Sandstone
Member. Rather. it is considered only as
“Billy Creek Formation”.

A lox of the Billy Creek Formation as
intersected in L. Frome No. 1 is presented
in Figure 11. The sequence is rather homo-
genous, comprising fine-grained tedbeds with
Minor green intervals. A transition into the
the overlying Wirrealpa Limestone ts indi-
cated. Cullings were sampled at 10 feet inter-
Vals and cores 5-7 were taken in the Billy
Creek Formation (Fig, 11).

Examination of cored intervals reveals a
spectrum of facies, with increasing silt con-
tent. ranging from evenly laminated claystones,
through wavy bedded mudstanes, inta poorly-
defined flaser-bedded siltstones. Ripple lami-
nated siltstones devoid of clay laminae are
uncommon. Anhydrite and calcite patches,
veins and lenses are present in all cored
intervals, although they are more common in
the finer-grained facies. Secondary reduction,
ussaciated with a colour change fram red to
preen, commonly surrounds the anhydrite’,
Halite imprints oceur sporadically through-
out the cored sections, and typically occur on
rippled bedding surfaces. Desiccation cracks
are common. Mudstone intraciasts and rill
marks are also common in parts of sequence,
particularly in the coatser-grained intervals.
Thin, pinkish igtervale at 780.3 m and 780.7
m in core 7 of L. Frome No. 1 contain
abundant altered. subangular feldspar, and by
anglogy with outerops in the Flinders Ranges,
are considered ta be tuffaceous in origin.

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BILLY CREEK FORMATION BAST OF FLINDERS RANGES 129

<
Qrey moltied limeshone
and argiilateous limestone

‘|

petioles dls yraaayt Wh I,

lamfleted with minor wavy

agal taminge Rare cliibing
\ Tippime Annyurice pate hes
\ane reworked inti qeipscs,

>— Grey #ien minor ced shale &
Vie aredus SiILStOne, Evenly

Greyen Ten ann green onale
ano stestone With ver? Minor
Sandshine Passes qradat onaily
inte the overiying Werreaipa
Loamestone

4

/Greview rad and minge
/greer clalse Gillstone and
snale Evenly laminated ond
flaser bedded Lommon nhatite
\ SastS desigcation crachs,
abhydhif® ahd musstone
Ante aclasle

rore

teh lbhen

Uresien red and itioor green
Shale ano eri stone

1700m
Laswe’

Uy
e :
3

ean’

[Dreyen red ane mingr
Joven shale and fine to

| Mediu eitstone Evenly
‘laminated wort) mince tlaser
[arspny twalite casts,
parsaccarion erarks mudscone
Joyerartasts ane anhyatice

‘ga PS

Beotn
KB!

Fig. 11, Stratigraphic log, Billy Creek Formation
in L. Frome 1,8 of L. Frame-

Tracks attributed to trilobites occur in several
of the cores™, however no body fossils have
been found to date.

foternal correlations

The Pridina Siltstone Member at Reaphook
Hill is considered to be the approximate lateral
equivalent of the Nildottie Siltstone Member
in the central Flinders Ranges (Fig, 12)-
Roth members are interpreted as being the
result of a major phase of tectonism in the
source area which caused basin-wide regres-
sion and promoted the development of red-
bed facies over a wide area.

Similarities between the underlying Conds
Hill Member at Reaphook Hill and the Warra-
gee Member in the central Flinders Ranges
support this correlalion (Fig, 13), In the
Wirrealpa Basin type section, the upper part of

the Warragee Member comprises a semi-
restricted marine sequence with green shales,
several thick dolomite beds, and prominent
tulfaceous intervals. A comparable although
less-restricted sequence is present in the upper
portion of the Coads Hill Member (Units G-J)
at Reaphook Hill and a correlation is made on
this basis. In particular, Unit H of the Coads
Hill Member (an open marine, grey lime-
stone) is considered to pass laterally (shore-
wards) into a sequence of interbedded dolo-
mites and green calcareous shales in’ the
Wirréalpa Basin (Fig. 13). On the basis of this
correlation, it would appear that the strata
containing B. daily/ at Reaphook are essen-
tially the same age as those containing B.
flindersi in the Wirrealpa Basin (Fig. 13),
despite the appearance of B. flindersi in the
type section being partly controlled by sedi-
mentological cather than evolutionary factors
(Moore 1979b).

Correlation af the Billy Creek Formation
between Reaphook Hill and the central Flin-
ders Ranges has been made possible only by
the fortunate coexistence of distinctive tuffa-
ceous and calcareous sequences. Either of
these features on its Own would probably have
been insufficient for a satisfactory correlation,
It was hoped that a similar method could be
employed for the correlation of the Billy
Creek Formation between the outcrops of the
Flinders Ranges and the subsurface occut-
rences to the east, Unfortunately, neither thick
tuifs nor distinctive calcareous beds were
found in the Yalkalpo 2 borecore, and thus
the relationship between this sequence and the
rest of the Billy Creek Formation remains
uncertain, However, since there is no evidence
af a regression in the upper portion of the

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SHON A .
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Fiz. 12, Stratigraphy of Early to Middle Cambrian
Billy Creek Formation,

P. S. MOORE

Ten Mile Creek Reaphook Hill
Section BC-B Section RH-A

metres

aa
Tuff (50cm) UNIT H ow
B. fFlindersi
300 Tuff (4m )—
lJ
Ww UNIT G
a
Bflindersi _|_ _ iii, seasityi
m Tuff (40cm) ge dally! =
pace ratte —_
Sarge al uw
= eat Ban eee Tuffaceous shale
2 Ri eat (15m) =
= 250 Suh dae eee
ee a UNIT G
ee ott
ld e u
oat
acre Vo
(10cm) ‘tos UNIT F
=
ye
UNIT E
UNIT D
lw
Ww
UNIT C
[2
WwW
—
{e)
oO
UNIT B O
<< x
te}
7 “ge {ty
4 ye vu
L x o- €
ao / o* o
joe
/ v= TUFF
/ Limestone
= /
J Dolomite
Shale and fine siltstone
= <

Rippled siltstone and sandstone

Evenly bedded sandstone

SSAARAARASS
SSS
S333

Cross bedded sandstone

vU
=
oO
a)

BILLY CREEK FORMATION EAST OF FLINDERS RANGES (4)

preserved Yalkalpo 2 sequence corresponding
with the development of the Nildottie and
Erudina Siltstane Members further west, it is
likely ihat the Billy Creek Formation in Yal-
kalpo 2 is equivalent to the lower-middle and
upper parts of the Warragee and Coads Hill
Members,

On the basis of isopach and palaeocurrent
trends, Moore (1979¢) considered that the
Ereeunda Sandstone Member was originally
deposited st Reaphook Hill, but has since
been removed by erosion, The apparent ab-
sence of the Eregunda Sandstone Member in
the Delhi-Santos L. Frome wells (Fig, 11) is
interpreted as an effect of lateral facies change,
with deltaie sandstones passing laterally inra
supratidal evaporitic mudflats. A camparable
Recent example is quoted by Thampson (1968,
1975) from the Gulf of California.

Conclusions

In the northern part of the Adelaide “Geo-
syncline’, initial teetonie activity associated
with the Kangarovian Movements (Daily &
Forbes 1969) terminated a major phase of
Early Cambrian carbonate deposition and pro-
moted the development of 4 thick sequence of
red-beds (the Billy Creek Formation). Five
oiemhers are defined within the Billy Creck
Formation (Moore 1979b and this paper),
which crops out in the Flinders Ranges and
oecurs in the subsurface to the east, During
the varly stages of deposition of the Billy
Creek Formation, a broad muddy tidal flac
developed in the west (the Warragee Mem-
ber) while to the cast, a cornplex stratigraphy
(the Coads Hill Member) was evolving, The
great variely of facies in the Coads Hill
Member emphasises the instability of this
pastern region during the late Early Cam-
brian, althouch in general this sequence and the
laterally equivalent scquence in Yalkalpo 2

were deposited in a more open marifte environ-
ment than the Warragee Member, Minor
volcanic activity, probably in the ME Wright
region of N.S.W., is recorded as thin tullaccous
interbeds in the lower half af the formation.

Further uplifts in the source area released
silt and fine sand into the basin, forming the
laterally equivalent Nildottie and Erudina
Siltstone Members. The red-bed facies uf both
members were deposited mainly in the inter-
tidal to supratidal zones undec the influences
of weak wave and current aclivity. while
cyclically interbedded dolomites in the more
easterly outeropping Erudina Siltstone Mem-
ber were probably deposited in sheltered
coastal lagoons in the shallow subtidal zone,

Final uplift further increased topographic
relief in the source trea and a complex of
fluvial dominated fan-delta sands (the Ere-
gunda Sandstone Member) prograded across
(he basin of deposition from the southeast
(Moore 1979c), Palacocurrent and petro-
eraphic data indicate that the main source af
the sediment was the Broken Hill-Olary base-
ment high.

Acknowledgments

| acknowledge discussion and helpful criti-
cism by Dr B, Daily (University of Adetaide),
Dr T. J. Mount (Delhi Petroleum Pty Lid),
Dr G. Necf (W. 5, & L. B, Robmsan Univer-
sity College, Broken Hill}, and Messrs R.
Dalgarno, R. Callen and C. Gatehouse and
Miss B. Youngs (S Aust. Department of
Mines & Energy). The tolk of the work was
completed at the University af Adclaide during
the tenure of a Commonwealth Postgraduate
Scholarship, supervised by Dr Daily and sub-
mitted in extended form as x thesis dissertation
in June 1979. The revised manuscript is pub-
lished with the permission of Delhi Petroleum
Pty Ltd.

References

Daity, B. (1958) The Cambrian tn South Aus-
tralla. Jn J. Rodgers (Fd.), El sistema Cam-
brico, sti palacogeowrafia el problema de su
base, 200 Int. geol. Coner., Meriva, yal. 2, 91-
147.

& Forsrs, B. G. (1969) Notes on the Pro-

terozoic und Cambrian, southern and central

Flinders Ranges, South Australian, Jr B, Daily

(Ed), Geologieas! Excursions Handbook, 23-30.

ANZAAS, Section 3, 1969.

Daccaano, ©, R- (1964) Report on the Lower
Cambrian stratigraphy of the Flinders Ranges,
South Australia, Trans. R, Soc. S. Anst, BR 129-
144.

___ & Jonson, J. E. (1963) Lower Cambrian of
the enslern flunk of the Flinders Ranges. Quart,
cool. Notes, Goal. Surv. suet. 7.

— & Jonnson, §, E. 11966) PARACHILNA
map sheet, Gealogicnl Atlas of South Austraba
1-250 N00 series. Geol. Surv, S. bey?

IS
Fig. 13. Correigtion of Billy Creek Formation between, Renphook Hill and iype section. in Wirrealpu
Basin. 2m (hick dolomite and overlying sequence of calcareous preet shales t Section, BO-B ute
correlated with grey murine limestone (Unit 1) al Reaphook Hill. Both writs are [ossiiiferous, con-

tainins trilobile Lalcorurunia sp. Although sequences ire Lu pceous,

corekuion of individual Tt

heds between oreas iy nol generally possible. und certninly not reliable.

132 P. S. MOORE

Moore, P. S. (1979a) Stratigraphy of the Early
Cambrian Edeowie Limestone Member, Flinders
Ranges, South Australia. Trans. R. Soc. S. Aust.
103, 101-111.

(1979b) Stratigraphy and depositional en-
vironments of the Billy Creek Formation (Cam-
brian), central and northern Flinders Ranges,
South Australia. Ibid. 103, 197-211.

—— (1979c) Deltaic sedimentation—Cambrian of
South Australia. J. Sedim. Petrol. 49, 1229-1244,

Pocock, K. J. (1970) The Emuellidae, a new
family of trilobite from the Lower Cambrian of
South Australia. Palaeontology 7, 458-471.

REINECK, H. E. & SinGu, I. B. (1975) “Deposi-
tional Sedimentary Environments with Refer-
ence to Terrigenous Clastics.” (Springer-Verlag:
New York.)

& WUNDERLICH, F. (1968) Classification and
origin of flaser and lenticular bedding. Sedi-
mentology 11, 99-104.

THOMPSON, R. W. (1968) Tidal flat sedimentation
on the Colorado River delta, northwestern Gulf
of California. Jn R. N. Ginsberg (Ed.), “Tidal
deposits: a casebook of Recent examples and
fossil counterparts.” (Springer-Verlag: New
York.)

Younas, B. C. (1978) Stratigraphic drilling in the
eastern Arrowie Basin, 1975-1976. Quart. geol.
Notes, Geol. Surv. S. Aust. 66, 16-20.

NEW ROTIFERA FROM AUSTRALIA

BY W. KOSTE & R. J. SHIEL

Summary

One hundred taxa of Rotifera, including six species: Lepadella chengalathi, L. williamsi, Lindia
deridderi Koste, 1979, Testudinella walkeri, Filinia australiensis Koste, 1980 and F. hofmanni
Koste, 1980 and five new subspecies are recorded from Australia for the first time. Eight new taxa
in the genera Brachionus, Keratella, Lepadella, Lecane, Dicranophorus and Testudinella are
redescribed and figured. Other species names, with ecological and locality information, are listed
systematically.

NEW ROTIFERA FROM AUSTRALIA

hy W. Kosre* & R, J. Swrery

Summary
Kogru, W. & Sten, Ro J (1980) New Rotifert. from Australia. Trans. R. Soe 5S. Aust. 405).

133-144, 28 November, 1980,

One hundred taxa of Rotifera, including six species: Lepadella chengalathi, L. willtamst,
Lindia devidderi Koste, 1979, Tesnulinella watkeri, Filinta wustraliensis Koste, 1980 and F.
hofmanni Koste, 1980 and five new subspecies are recorded fram Ausiralia for the fipst time.

Right new taxa in the genera Brachrows, Keratella, Lepudella, Lecane,

Dicranepharis and

Testudinella are desenbed and figured. Other species names, with ecological and Jocality infor-

mation. are listed systematically.

Introduction

Rotifera are found in virtually all inland
waters, They are usually the numerically
dominant component of the zooplankton of
lakes, rivers, billabongs and swamps, providing
an important link in aquatic food chains
between the nannoplankton (i.c. less than 60
nm, including bacteria snd micro-algae) and
the carnivoraus zooplankton.

Publication of the first checklist of the
Australian Rotifera (Shiel & Koste 1979)
and a review of the Australian species of
Brachionus and Keratella (Koste 1979) has
resulted in increased interest in the rotifer
fauna. Workers locally and averseas have com-
municated papers and collections, enabling the
faunal fist to he expanded. The first material
from the Northern Territory bas been
examined, and access has been provided to
collections from Tasmania and Weslern Aus-
tralia, Material from the other States has been
collected during a survey of the zooplankton
of the Murray-Darling system (Stel 1978,
1979),

Consequently, while this paper adds to the
known rotifer fauna of the continent, it Tmust
be considered as a preliminary to a more com-
prehensive treatment acu later date, in which
illustrated keys will be provided for assistance
in identifeation of the Australian Rotifera., At
present, taxonomie references in which many
of fhe cosmopolitan taxa are figured include
Barlos (1959), Kutikova (1970) and Kaoste
(1978), none of which is readily accessible to
the English-speaking Australian limmologist,

One hundred taxa of Rotifera are recorded
from Australia for the first time, In addition,
three species (Ascamerpha saltans, Pomphalyx

EE

* Lucwie-Brill-Strasse 5, Quakenbruck, D570,
West Germany.
| Department of Zoology, University of Adelaide.

sulcata and Tesiudinella amphora) are recorded
by Sudzuki & Timms (1980) from farm
dams in N.S,W. These species were recorded
in the present study and are retained in the
listing to provide ecological information. Sud-
zuki & Timms’ record of Filinia maior is con-
sidered synonymous with F. terminalis (see
Koste 1978). A further three species described
by Berzins (1961) were omitted From the ori-
ginal checklist, and are included here, bringing
the recognised Rotifera taxa from Australia
to 437. In some cases, taxa having minor dif-
ferences from the type al the sub-specific or
varietal level are the only representatives of
the species recorded to date from the continent,
and are therefore included in the record as
distinct taxa follawing the principles of Mayr
(1969),

Although it would be premature to give de-
tailed ecological information on the collections,
particularly since material from large areas
af the continent has yet to be examined, we
include information on the habitats from which
the various taxa are recorded. The format
adopted is as follows;

All records are listed systematically (after
Koste 1978). Habitat type is indicated by:

L: lake or impoundment, with name of
locality;

R; river or Stream,
named;

B- billabong (standing waters on the flood-
plain), ‘with nearest town or setiiement
named,

The season of collection is. given as W:
winter: Sp; spring; Su> summer and A:
autumn,

Physico-chentical data js given in the
sequence: temperature in °C (measured with a
Yellow-Springs Instrument Company thermis-
tor); pH (Metrohm portable pH meter);

with nearest town

134

dissolved oxygen (YSI DO meter): turbidity
(Hach turbidimeter, or where a metric mea-
surement is given, by Secchi disc); conductivity
(Radiometer CDM2 conductivity meter). In
the example given below, the units of measure-
ment for each parameter are given in paren-
theses:

“Lecune stichaea Harring, 1913
Loc: R, Mannum, S.A., Wz 11.0 (°C), 8.4
(pH), 10.8

(DO in ppm), 40 (turbidiity in Nephelometric
Turbidity Units), 575 (4S cm),

Several species came from collections
other than by us, For these, physico-chemicyl
data were not available to us.

No distinction 1s made between planktome
and littoral species. Seasonal variation occurs,
particularly when billabong species are flushed
into rivers to constitute the plankton in times
of flood, as occurs in the Murray (Shiel 1979).
Other species become established during algal
blooms in lakes and rivers, using algal
filaments or mats as substrates for attachment,
and are thus seasonal or facultative plankters.

New taxa only are figured, wilh the excep-
tion of Brachionus lyratus Shephard (Fig. 1),
This species was previously considered to be
synonymous with &. cawdatuy (Ahlstrom
1943), however present samples indicate that
the species is valid.

Abbreviations for the locations of type
material are: ZMK, Zoological Museum, Uni-
versity of Kiel, D-2300, F.R.G,; SAM, South
Australian Museum,

Systematics
ORDER BDELLomaA

(See Shiel & Koste 1979, for known Australian
bdelloid rotifers),

ORDER PLoImMipak Hudson & Gosse, 1886
Family Epiphanidae Bartos, 1959
Liliferetracha subtilis (Rodewald), 1940
Loc: B, Thornton, Vie., A; 17-8, 7.1. 8.9, 28.

-,

Rhinoglena frontaliy (Ehrenberg), 1853
Loe; B, Yarrawanga, Vie, Sp; 14,5 8.2. 12.5,
18, 74

Family Brachionidne Kutikova, 1970
Brachionns quadridentatus {. brevispinus
(Ehrenberg) 1832
Log; L, Boort, Vic. Su; 23.5, 7.9, 6.0, 87,

750,

W. KOSTE & R. J. SHIEL

B. quadridentaius 1. cluniortheularis
(Skorikow) 1894
Loc; B, Wodonga, Vic., Su,
B. vartabilis (Hempel) 1896
Loe: B, Wodonga, Vic., Su
B, lyratus Shephard, 1911 (Fig.1)
Trophi preparation, Rot, No. 023, ZMK,

In Koste (1979) a Brachionuy resembling
that deseribed by Shephard (1911) was
figured. Harring (1913) suggested synonymy
of the form with B, anegularis candatus Bar-
Trois & Daday 1894, Voigt (1937) considered
the form insufficiently known, and Afistrom
(1940) considered it synonymous with B. ar-
guleris. Examination of recent collections sup-
ports Shephard’s comments on the peculiar
Morphological characteristics of this distinctive
species. In particular, the presence of sub-
median spines, absent in B. angufaris, and the
“two knobbed projections (posteromedian
spines) which curve outward in a manner
sugeesting the form of a lyre’, Such foot-
opening spines are not described for 8B. angu-
laris or B, caudatus. On the basis of these
laxonomic characteristics the independence of
B. lyratus in the Formenkreis angularis is
recognized, This now comprises the species
groups angularis, charint, dichatomus, dola-
Aratus, forficula and Iratus (see Koste 1978),
Measurements: Loriea length 120-145 um,
lorica width 90-96 pm, foot-opening width
20-24 pm, head-opening width 60-64 jm.
subitaneous egy 65/45 jum.

2(¢)
Figs |-2 |—Frucliouus lvratus Shephard,
191). ah. Loricu, ventral Fig. 2—A&. bidentate
tL testudlinarius (Jakubskit 1912. 9, Lorica,

dormal, 6, Lonea, ventral, c. Lorea lateral.
Lorica Jength 220 am, (ef. Fig. da, b)-

NEW ROTIFERA FROM AUSTRALIA 135

oc: B, Thornton, Vic., A; 17.8, 7.1, 8.9, 28,

B. dimidiatus (Bryce) 193)

B, Wadonga, Vic., A.

B, urceelarisy sessilis (Varga) 1951

Loc: B, Jabiluka, N-T., W: 25,0, 6.15, 3.7, -.
48,

B, dichotomus reduetus Koste & Shiel, 1979
Loc: B, Eskdale, Vic., Su; 29.2, 7.5, 8.0, 6,
85.

RB. hidentata Anderson, 1889 (Figs 2+)

Loe!

B. bidentata minor 0.ssp.
(FIG. 3a-h)

Material: (6 contracled females, sample No. 295,
in formalin.
Holotype: loricate female, sample No. 295, coll.
28.01.77, R. J. Shiel, Rot. No. 025, ZMK,
Paratypes: SAM, V,2092, Koste collection, Qua-
kenbrtick. F.RAG,
Icanatype: Pig, tlostra 3a,b.

Type Jocality: River Murray, Mannum, §.A.
(34°50'S/ 139° 18'E),

Description: Small barrel-shaped lorica with
six dorsal anterior spines. Short, unforked
marginal spines (cf. Fig. 2c). Basal plate pre-
sent. Semicircular dorsal lorica border. An-
terior border of ventral plate slightly indented
medially. Highly placed circular foot-opening.
Posteromedian spines terminally —button-
shaped. Posterolateral spines absent. In larger
specimens slight facettation dorsally (Fig. 3d,
h). Lateral antennae marginally placed,

Measurements: Lorica length 120-156 pm,
lorica height 52-80 pm, greatest Jorica width
84-108 pm, range of marginal spines 72-104
A.

Discussion; The lorica shows comparative
similarity to that of B, bidenrata ft. inermis
(Rousselet) 1906, The latter however has long
murginal spines, absent in minor. Moreover,
this morph is an intermediate to 8. bidentata
f. testudinarius (Jakubski) 1912 (Fig. 2a—c),
which has a short curved posterior spine, In
the new subspecies the posterior border of the
lorica is specifically semicircular. Comparison
with B. bidentata {. jlrove? (Bartos), 1947.
Syn: B. furenlatuy var, jirevci Bartos, 1947
(Hauer 1963; Kutikova 1970) shows that this
intrasubspecies taxon, in contrast to B. biden-
tata miner, bas forked marginal spines, and
also nceasionally posterolateral spines.

3a) 3(b) ~3(c)
rat Ps9 XY . ™.
oO (f
Ning 3a) Sf pp
is. My 50 ym ‘ 4
3(h) 4,0,
uty [
‘iat
a 4 )
wl vy}
-. eS \. ,
Lie" uc air
Figs 3-5. 3—Braciiionus bidentata minor .s5P.

a. Lorica, dorsal. b,c. Loricu, ventral, d.
Anorher lorica, dorsal, e-@. Dillerent loricas,
ventral. f. Lorica with dorsal structure (all From
the same population). Fig. 4—B. bidentata f.
evindinerius. Lorica, dorsal. Length 1704m,
width 140um. Fig, 5—A_ pliecatilis calongula-
ciensis. ssp, Lorica, dorsal, Lorica length 400
um.

Considering the great variability of the
species B. bidentata an exact taxonomic classi-
fication of the different morphs is difficult. For
example, with regard to length of lorica spine
development, Pourriot (1974) showed experi-
mentally that this depended on the presence of
the predator Ayplanchna brightwelli.

The new subspecies is considered here as a
“dwarf-form" sensu Green's (1977) study,
where food-storage, salinity of biotope, etc.,
caused a reduction of lomea-size in crater
luke rotifers.

Therefore the classification of B. hidentata
minor as a subspecies ie. a genetically dis-
tinct farm, is provisionally documented here,
noting However that only a single collection
is involved.

Loc: R. Mannum, S.A,, Su; 17.0, 8.2, 9.7,
86, 1,020.

Brachionus bidentata f.

(FIG, 4)

In one sample from Lake Nillahcootie, Vic.
(36°53'S/146°01'E) on 2741.78 an individual
animal resembling the f. restudinarius was
collected. Forked marginal spines are absent,

? testudinarius

136

however, and the dorsal lorica is unstructured.
The lorica has a resemblance to Brachionus
quadridentatus var. ancylognathus (Schmarda
1859), but the elongated marginal spines, dis-
tinct lorica basal plate and the tube-like foot
orifice indicate that this is a morph of B.
bidentata.

Measurements: Lorica length 164 um, greatest
lorica width 140 ym, marginal spine length
68 ym, posterior spine length 100 pm.

Loc: L, Nillahcootie, Vic., Su; 26.0, 7.9,
11.3, 24, 55.

Brachionus plicatilis colongulaciensis n.ssp.
(FIG. 5)

Type material: 122 females of different ages, in
formalin, sample number 647, collected by W, D.
Williams, Lake Colongulac, Vic. (May 1979).
Holotype: Loricate female, coll. Williams 23.v.79,
Rot.No.024, ZMK.
Paratype: SAM, V.2093; Koste collection, Quak-
enbriick, F.R.G.
Type locality: Lake Colongulac (salinity 10.4%),
W. Victoria (38°10°S/143°10'EB).
Description: Greatly elongated transparent
lorica. Surface unstructured. Anterior spines
much reduced. Anterior border of ventral
lorica with four short, flat, tongue-like projec-
tions. Posterior lorica margin tapered and

rounded. Foot-opening terminal, lowered
ventrally. Dorsal lateral antennae approxi-
mately level with mid-lorica. Ratio lorica

length: width approximately 2:1,

Measurements: Lorica length (adult 2) 400—
440 um, lorica width 210-240 ym, foot-open-
ing width 40 ym, anterolateral spine develop-
ment —75 ym, subitaneous egg 132-150 X 90-
110 pm.
Discussion; In the population examined, all
adult females had an extraordinary large
lorica, corresponding in appearance to that of
B. plicatilis f. decemcornis Fadeev, 1925. The
latter has a different foot-opening (see Koste
1978) and is smaller. Hauer (1925) describes
an elongated form from Bad Oldesloe with a
cross-sectioned lorica length of only 260 pm.
A lorica of approximately corresponding out-
lines was figured by Ahlstrom (1940) with a
length of 275 ym and a greatest width of 210
pm. Neither of these is in accord with the
Australian find. The greatest lorica length
known to date is 315 ,m,

It is apparent that the new ssp. population
has arisen in Lake Colongulac in response to
specific ecological pecularities of the habitat.

W. KOSTE & R. J. SHIEL

For the anatomy of this Brachionus, see
Koste (1980a). For distribution of the species
group see De Ridder (1960).

Loc: L, Colongulac. Vic., A; 12.7, 8.2, -,
42, 10.4 gm 1-4

Keratella procurva (Thorpe) 1891
(FIG. 6a—k)

This pantropical species, known previously
only in its typical form (see Ahlstrom 1943,
Berzins 1955 and Koste 1978), shows con-
siderable variability in posterior spine develop-
ment and overall lorica size. Paggi (1973)
figures and reports on a series of different
lorica forms (Fig. 6h) from the Parana River,
Argentina.

In the many Australian samples examined,
this species occurred also with variable lorica
forms. The most abundant forms are figured
(6a,f,g,i). A previously unknown form of the
species group (Fig. 6b-e,j,k), notable in
respect to its smallness and form of posterior
spines, is documented below.

Keratella procurva robusta n.ssp.
(FIG. 6k)

Holotype: Female, coll. R. J. Shiel, Keepit Dam.
N.S.W., 24.v.78, Rot. No, 026, ZMK,
Description: Short, squat, relatively wide and
opaque lorica, ornamented with large pustules.
Facetted as in f. typ., although with minor
differences in populations from widely
separated localities. All lorica spines, particu-
larly short posterior spines, with wide bases
(cf. Figs. 6b,j with 6d,h,i). Latter curved
backwards in lateral view. Forms without
caudal spines or with only a single postero-
lateral spine rudiment as in K. tropica (Apstein
1907) not recorded for K. procurva. Compara-
tive morphological characteristics and measure-
ments of known forms of K. procurva are as
follows. In common forms (Figs 11la,f) the
lorica surface is smooth or weakly granulated.
Total lorica length from 155-220 ,m; lorica
width 67-80 pm; anterior spines 19-20, 17-
22 and 30-40 um and posterior spines 22-24
pm (left) and 27-64 pm (right). K. procurva
robusta has a strongly studded lorica surface,
total length 148-150 ym; lorica width 82-90
pm; anterior spines 15-17, 10-15 and 22-23
um; posterior spines 14-15 ,.m (left) and 20-
23 um (right).

Loc: L,B,R, Darling catchment, Su; 10.5—
25.0, 7.4-8.7, 6.4-10.8, 270-1000, 15-1000,
15-115,

NEW ROTIFERA

6(b) 8(c) 6(d)

Fig. 6a. 1891.

(Thorpe)
Lorica, dorsal. Morph wilh dorsolaterally curv-
ing posterior spines. S. Australia. Lorica length
220 nm. b. K. procurva robusta mssp. Lorica,

Keratella procurve

dorsal, length 1S0em. c. Lorica, lateral. d.
Another specimen. e. Subitaneous egg of A. pro-
curva robusta. T-g. K. procurva type, common
form from Australian waters. h. A. procurva,
variations from Argentinian waters, 8S. America
(afer Paggi, 1973). i. K. precurva from
Australian warm waters, N.T. Lorica length to
230 um. j,k. Two forms af K, pracurva robusta
from Darling River waters.

Notholea squamula (Muller), 1786

Loc: R, Donald, Vic., Ay 11.0, 8.0, 11.2, -— =
Anuraeopsis navicula Rousselet, 1910
Loc: B, Yea, Vie. A; 10.5, 7.2, 10.1, 33, -.

A. coelata De Beauchamp, 1932
Loc; B, Jabiluka, N.T,, W; 25.0, 6.2, 3.0, —
Ag.

Family Euchlanidae Bartos, 1959
Euchlanis dilatata *. univetata (Leydig), 1854
Loc: B, Seymour, Vic., W; 10.0, 7.2, 11.6,
2, 330.

E, praxima Myers, 1930
Loc: R. Wyangala, N.S.W., Su; 15.5, 7.3,
10.2, =, =

FROM AUSTRALIA

\37

Dipleuchlanis propatula (Gosse), 1886
Loc: B, Jabiluka, N.T., W; 25.5, 6,2, 2.9, — 62.

Family Mytilinidae Bartos, 1959

Myrilia macracaniha (Gosse), 1886

Loc: B, Jabiluka, N.T., Ws 24.5, 6.3, 5.8, -
59.

M. bisulcata Lucks, 1912
Loc: B, Jabiluka, N.T,, Wi 25.5, 6.2, 2.9, -,
62.

M. crassipes (Lucks), 1912
Loc: B, Jabiluka, N.T., Wz 25.0, 6.2, 3.0, -,
48,

Family Trichotridae Bartos, 1959

Wolga spinifera (Western), 1894
Loc; R, Wentworth, N.S.W,, A; 10.0, 8.0,
10.2, 12, 675,

Family Colurellidae Bartos, 1959

Colurella adriatica Ehrenberg, 1831
Loc: L, Boort, Vic., Su; 27.5, 7.9, 6.0, 67,
750, R. Benalla, Vic., A; 8.5, 7.0, 11.2, 21,
575.

Sguatinella mutica (Ehrenberg), 1832
Loe: B. Yea, Vic.. Ay $9.9, 7.2, 8,5, 0.8 m,
85.

Lepadella ehrenbergi (Perty), 1850
Loc: B, Jabiluka, N.T.. W; 24,0, 6.25, 5.1, -,
42.
L. elliptica Wulfert, 1939
Loc: B, Jabiluka, N-T., W; 25.5, 6.2, 2.9, —
62.

L. rhontboides rhaniboides (Gosse), 1886
Loc: B, Jabiluka, N.T., W; 24.5, 6.3, 5.8, -,
59.

L. rhomboides f curinata (Bonner), 1934
Loc: R, Benalla, Vic., Su; 27.0, 7.6, 7.0, 52,
L. dactyliseta (Stenroos), 1898
Loc: R. Benalla, Vie,. Su; 27.0, 7.6, 7.0, 52,
L. henjamini Harring, 1916
Loc: L. Boort, Vic., Su: 20.0, 7.4, 8.8, — =
B, Jabiluka, N.T., W; 24.0, 6.25, 5.1, -, 42.
L. heterodactyla Fadeew, 1925

Loe: L, Boort, Vic., Su; 20.0, 7.4, 8.8, — —
L. apsicora Myers, (934
Loc; L, Boort, Vic., Su; 23.5, 7.9, 6.0, 67, 750.
L, nehoissi Berzins, 1960
Loc: R, Kinglake West, Vic., Sp.
L, ptilota Berzins, 1960
Loe: R, Clunes, Vic., W.

138 W. KOSTE & R, J. SHIEL

50pm
Fig ?

) 50 ym
Fig 8
8(c) ad
Figs 7-8. 7—Lepadella chenvoluthi nsp- a.

Lorica, yentral, 6. Distal foousegment with
toes enlarged. c. Lorica cross-section. d. Neck
aperture, dorsal. ¢. Ventral aspect, Fig, 8—L,
williaiisi 1.sp. ab, Ventral uspecls of two
loricas. ¢, Lateral view. d. Lorica cross-section.

L. angusta Berzins, 1960
Loe: R, Morang, Vic, Su,
L, marnedactyla Berzins, 1960
Loc} B, Jabiluka, N-T.. We 25.5. 615, 2.9, -
62.
L, rererostyla (Murray), 1913
Loe: L, Boort, Vic., Su; 23.5, 7.9, 6.0, 67, 750,
L.. latusinus (Hilgendort), 1899
Loc; R Benalla, Vic), Su: 27.0, 7.6, 7,0, =, -

Lepadella chengalathj sp.nov.
(FIG. Ta-e)

Type ninterial: 3 lavieate 23, sample number 307,
Holautype: Pemile, sample number 307, call. Ref.
Shicl, Rot No, 021, ZMK.

Uvpe locality: Lake Boort, endorheic (internal)
drainage, near loddon River, Vieloria (36°
US'S/143°45'B),

Deseriptions Rhomboidul lorica (Fig, 7a) with
rounded corners, widest in first third, Mead
opening small, not drawn down ventrally.
Foot-opening semivireular at widest point of
cuudal boundary of dorsal lorica, which is
weakly arched towards interior, Three flexible
foot-urticulations, distalmost slender and
elongated. Toes symmetrical, long, needle-like
and straight (Fig. 7b). Lorica vross-section

shows highly vaulted dorsal lorica with slender
downward-drawn lateral wings. Ventral plate
with delicate double convex keel (Fig. 7e).
Measurements; Total length 135-140 jum,
lorica height 48 jum, lorica length 96 ,m,
lorica width 92 ym, head-opening width 28
wim, foot-opening 28 % 28 um, distal foot-
articulation 19-20 pm, toe-length 33-36 pm.
Discussion; The new species has similarities
With Lepadella benjamini Haring, 1916. In this
species, however, the head-opening is cap-like,
more or less Ventrally drawn down, and the
distal foot-articulation is not elongated (12-
13 pm). CYL. also an Amazonian form described
by Koste (1972, 1978) L, benjamini f. brasi-
liensiv with rounded side rims. The genus is
reviewed by Chengalath (1976).
Etymology; Named after Rama Chengalath,
National Museum of Canada, Ortttiwa, in
uppreciation of years of scientific cooperation
and friendship with W.K.

Loc: L, Boort, Vic,, Su; 23.5, 7.9, 6,0, 67,
420,

Lepadella williamsi n.sp,
(FIG. Sa-d)

Holotype; Locicate female, sample namber 657,
coll, R, J, Shiel, 13.vi,79, Rot. No. 19, ZMK,
Paratype, SAM V,2904; number 657 Koste col-
lection, Quakenbriek, D-4570 FRG.
Type lovality: Ja-Ja Billabong, Jabiluka, N,T,
(12°40'S/133°00'R).
Peseriptions Smoothly oval, highly-vaulted
loriva (Pig. 8d) With ventrally-direeted head-
opening, Dorsal loricu tapers caudally to a
variuble-length dorsal and laterally curved point
(Figs 8a,b), Foot-opening narrow and ends
at base of dorsal lorica projection. Only three
flexible, approximately equal length foot sex.
ments visible. Last segment with long, dorso-
laterally curved sharp toes.
Measurements, Total lorica length 112-116
pm, lorica height —56 »,m, lorica width 60-64
pm, toe length 36-40 am, foot-opening 17-20
* 19-20 am, head-opening 33-36 * 28-30
win, distal foot-segment length 10 jm, caudal
lorica projections 19-28 jum.
Diveussion: The new species resembles the
Species group L, acuminata, Which also occurs
in Australia (Figs 9,10), and which is charac-
icrised by a more or less clongated and ovea-
sionally split lorica point, However the margin
of the foot-opening of this projection is closed
(Fig. Yb, 10b). The lateral antennae, which are

NEW ROTIFERA FROM AUSTRALIA

dorsal to the basis of the caudal projection, in
contrast to L. williamsi are very narrowly
placed. In L. acuminata and its various modi-
fications (Koste 1978) the dorsal margin of the
head-opening is more or less concavely cut out.

The new species was previously seen in a
sample from a tank near Kuala Lumpur,
Malaysia collected by C. H. Fernando, Uni-
versity of Waterloo, Ontario, Canada.
Emmolagy: Named after W. D, Williams, Dept
of Zoology, University of Adelaide for assis-
tance in collecting rotifer material and support
toward this work.

Loc: B, Jabiluka, N.T., W; 25.5, 6.15, 2.9.
5, 62.

Heteroleapdella ehrenbergi (Perty), 1950
Loe: B, Eskdale, Vic. Su; 29.2, 7.5, 8.0, 6,
85,

Family Lecanidae
Lecane hernemanni (Ehrenberg), 1834

Loc: B, Yea, Vic., A; 19.9, 7.2, 8.5, .8 m, 85.
B, Jabiluka N.T., W, 24.5, 6.3, 5.8, — 59.

L. ludwigi (Eckstein), 1893
B, Jabiluka, N.T., W; 25.5, 6.15, 2.9,

L. aculeata (Jakubski), 1912
B, Jabiluka, N.T.. W; 24.0, 6.25, 5.1,

L. stichdea Harring 1913
Loe: R. Mannum, S.A. W: 11.0, 8.4, 10.8,
40, 575.
L. curvicornixy (Murray), 1813
Loe: B, Jabiluka, N.T.. W; 25.0, 6.2,
48,
L, furcéta (Murray), 1913

Loe: B. Jabiluka, N.T., W: 24.5, 6.3, 5.8, -,
59.

L. grandis (Murray), 1913
Loc: B, Jabiluka, N.T., W; 25.0, 6.2, 3.0, -,
48.

L. doryysa Harring, 1914
Loc: B, Jabiluka, N-T., W; 24.5. 6.3, 5.8, -
59,

L. crepida Harring, 1914
Loc: B, Jabiluka, N.T., W; 25.0, 6.2, 3.0, -
48.

L, tenuiseta Harnng, 1914
Loc: B. Jabiluka, N.T,, W; 25.0, 5.85, 2.2,
-, 29,
L. lunaris crenata (Harring), 1923
Loc: L, Yarrawonga, Vic,, Su; 24.2, 7.7, 8.6,
—, 60.

9 (a)

50 ym

I (a) l(b) tI {c)
Figs 9-Il. 9—Lepadella acuminata (Ehren-
berg), 1834 from N.T. a. Lorica, dorsal,

b. Foot-opening and toes, ventral, Fig. 1O—L.
wcwninata form with abbreviated caudal loricu
point from N_T., Australia, a, Dorsal. b, Foot-
opening and toes, ventral. Fig. 11—Lecane
(Monostyla) hamata Vietariensis nassp. a. Ven-
tral view, lorica contracted, b. L. hamata
hamata Stokes: 1896, ¢, L. sinueata Hauer, 1938.

L. unguitata (Fadeew), 1925
Loc: B, Jabiluka, N.T.. W; 25.0, 6.2, 3,0, -
48,

L. acronycha Harring & Myers, 1926
Loc: B, Jabiluka, N.T.. W: 25.0, 6.2, 3.0, —,
48.

L. inopinata (Harring & Meyers), 1926
Loc: L, Wyangala, N.S.W., Su; 27.0, 7.4, 7.5,
, 245,

L. pertica Harring & Myers, 1926
Loc: B, Jabiluka, N.T., W; 24.5, 6.3, 5.8. -,

L, elsa Hauer, 1931
Loc: B, Alexandra Vic., Sp; 20.0, 7.2, 9.1, —

L, nodosa Hauer, 1938
Toc: B, Jabiluka, N.T., W: 24.5, 6.3, 5.8. -,

L. ruttneri Hauer, 1938
: L. Boort, Vic., Su; 20.0, 7.4, 8.8.

L. hamata victoriensis n.ssp,
(FIG, 11a)
Type material: 2 loricute 99. sample number 263,
Holotype: loricate 2, sample number 263, coll. R.
J. Shiel (14.x.77) Rot. No. 029, ZMK,

140

Type locality: Billabong “Goulburn View”,
near Alexandra, Victoria (37°13’S/145°42’E)
from Juncus bed, 30 cm depth.

Description: Both anterior points of dorsal
lorica at head aperature are congruent with
ventral lorica, therefore not visible. Whereas
this aperature in type flat or deeply curved,
in spp. it is a deep trapezoid notch, basally
8 um wide, bordered by light, curved lines.
Lorica outline widely oval. Ventral plate
smaller than dorsal plate. Toe gradually taper-
ing from point of insertion at second foot
segment.

Discussion: Morphologically, this ssp. shows
several characteristics in agreement with L.
(M.) sinuata Hauer, 1938 (Fig. 11c). A simi-
lar rotifer also was described from India by
Wulfert (1966) as L. arcuata (Bryce), how-
ever the notch between the anterior spines in
this latter form are otherwise shaped, and sig-
nificant differences are seen in the measure-
ments.

Measurements (in pm) are given in the
following sequence: overall length, dorsal plate.
ventral plate anterior transverse width, toe
length, form of neck opening for each form of
L. hamata, f. typica 103-136, 61-88 xX 52-
66, 77-98 X 40-60, 18-30, 26-38 double
semicircular. v. sinuata 108-110, 74-76 X53-—
55, 80-82 X 41-46, 21, 28, curved. v. arcuata
100, 52 X 56, 70 X 40, 25, 28, semicircular.
v. thienemanni 113-123, 64-70 X 62-64,
77-88 X 46-55, 26-35, 33-34, ventral flat-
concave, dorsal straight. victoriensis 124, 84
xX 64,92 X 56, 24, 32. trapezoid. Fig. 11a
shows morphological differences of the new
spp. from that described by Stokes, 1896 (Fig.
11b). In contrast to L. h. victoriensis, L. h. var.
thienemanni has a characteristic horizontal
dorsal lorica margin, and the toe is distended
in the middle.

Loc: B, Alexandra, Vic., Sp; 20.0, 7.2, 9.1,
2, 115.

Family Proalidae Bartos, 1959

Proales fallaciosa Wulfert, 1937
: B, Jabiluka, N.T., W; 24.5, 6.3, 5.8, -,

Family Lindiidae Dujardin, 1841
Lindia torulosa Dujardin 1841
: B, Yea, Vic., W; 11.0, 7.2, 6.1, 17, 170.

L. deridderi Koste, 1979
: B, Wodonga, Vic., W; 10.2, 7.2, 9.0, 4,

W. KOSTE & R. J. SHIEL

Family Notommatidae Remane, 1933
Itura myersi Wulfert, 1935
Loc: B. Wodonga, Vic., Sp; 14.7, 7.1, 4.1, 5,
240.

Eothinia elongata (Ehrenberg), 1832
Loc: B, Wodonga, Vic., Sp; 14.7, 7.1, 4.1,
5, 240.

Monommata grandis Tessin, 1890
Loc: B Jabiluka N.T., W; 24.5, 6.3, 5.8. -,
59,
M. actices Myers, 1930
Loc: B, Jabiluka, N.T., W; 25.0, 6.15, 3.7, -,
48.
M. arndti Remane, 1933
Loc: B, Alexandra, Sp; 20.0, 7.2, 9.1, -, -.
Notommata glyphura Wulfert, 1935
Loc: B, Alexandra, Vic., A; 17.8, 7.2, 8.9, 8,
Cephalodella eva (Gosse), 1886
Loc: B. Eildon, Vic., A; 17.9, 7.2, 9.2, 0.5,
87.
C. mucronata Myers, 1924
Loc: R, Mungindi, N.S.W., A; 16.0, 8.1, 9.2,

C. panarista Myers, 1924
Loc: B, Jabiluka, N.T., W; 25.5, 6.2, 2.9, -,
62.
C. ventripes Dixon-Nuttall, 1901
Loc: B, Wodonga, Vic., Sp; 14.7, 7.1, 4.1, 5,
240.

Family Trichocercidae Remane, 1933

Trichocerca cavia (Gosse), 1889
Loc: B, Eildon, Vic., A; 18.0, 7.1, 11.0, 0.5,
70.
T. collaris (Rousselet), 1896
Loc: B, Jabiluka, N.T., W; 24.5, 6.3, 5.8. -,
59.

T. chattoni (De Beauchamp), 1907
Loc: L, Wyangala, N.S.W., Su; 27.0, 7.4, 7.5,
0.5, 245.

T. myersi (Hauer), 1931
Loc: B, Jabiluka, N.T., W; 24.0, 6.25, 5.1,
—, 42.

T. flagellata Hauer, 1937
Loc: B. Jabiluka, N.T., W; 18.0, 6.1, 6.7, —, 63.

T. mus Hauer, 1938

Loc; L, Nagambie, Vic., Su; 15.0, 7.4, 8.9,
1.3m, -.

T. jenningsi Voigt. 1957
Loc: B, Jabiluka, N.T., A; 29.0, 6.5, 2.9, —
28.

NEW ROTIFERA FROM AUSTRALIA 141

T. similis erandis (Hauer), 1965
Loc! B, Wodonga, Viec.. Sp; 15.5, 7.0. 10,8.
16, 43,
YT. agnatha Wulfert, 1939
Loc: R, Echuea, Vic., Su; 15.0, 7.6, 9,0, — ~.
Ascomorpheila valvovicola (Plate), 1886

Loc: B, Thornton, Vie. A; 12.4, 7.1, 8.6, 2,5.
11S.

Family Gastropodidac Remane. 1933
Ascoimorphe saltany Bartsch, 1870
Loc: L. Burtinjuck, N.S.W,, Su; 26,0, 7.6, 8.0,
1, 190,

Family Synchaetidae Remane, 1933
Synchaeta tavina tavina Hood, 1893
Loc: L, Kerang, Vie. W.
3. litoralis Rousselet, 1902
Loc! B, Wodonga. Vic., W; 10.2, 7.2, 9.0, 4
154,
Family Dicranophoridue Remane, 1933
Dicranepherus uncinatus (Milne), 1886
Loc: B, Wodonga, Vie, A,

D, aquilus (Gosse), 1887
Loc: R, Mungindi, N.S.W., Ai 16,0, 8.1, 9.2,

Dicranophorus claviger australiensis n.ssp.
(FIG. 12a.b)

Type material; 14 92, sample number 669,
Hototype: female, coll. 13.vi.79, R. J. Shiel, Rot.
No. 027, ZMK.

Type locality: Ja Ja Billabong, Magela Creek
floodplain, confluent of Alligator River near
Jabiluka, N.T. (12°40'8/132°50"))

Description: Elongate spindle-shaped body.
Integument rigid, longitudinal lines from neck
to short horizontal line on dorsal part, Head
purt cylindrical with moderate concave out-
lines. Small prolongation of dorsal lorica over
foot. Toes extremely Jong and slender, curve
somewhat outward, end in acute pointed claws
seen Only in lateral view. Corona with two
frontal eyes and paired knobbed palps, Trophi
large. Rami terminate wilh smal) tecth as in
the trophi of D. caudatus (Fig, 12c) but have
broad lamellar alula, which are apically clon-
gated, ending in pai of double short teeth.
Unci each have only a tooth. Short fulerum
has broad triangular form. Manubria shghtly
curved, spatulate ended, Beneath unci contact
point is lamellar triangular clement. Preserva-
tion Ww formalin precludes discussion of internal
organization.

Discussion: This species is related to D. cau-
datuy, from Victoria, and D. claviger, from
South America by virtue of ils morphology,
Comparative information on the related forms
is given below. Affinity of this ssp. with D.
claviger (Hauer) 1965 is suggested by the
palp number and similar trophi structure,
although D. claviger (= Itura claviger Hauer,
1965) has a different shaped fulcrum, shorter
toes and a significantly shorter overall length,

Measurement (in pm) and comments are
given in the sequence overall length, greatest
width, palp number, palp length, toe length,
trophi length, manubrium form, uncus, ful-
crum, rami, manubrium, teeth on ramus point.
supra-rami teeth, supra-rami teeth length, dis-
tribution: D, caudatusy Ehrenberg, 180-310,
90, 1, 9-10, 69-77, 34-36, sticklike, 13. 6,
21, 25, — absent, —, cosmopolitan; D. caudatus
braziliensis Koste, 1972, -330. — 2, 2, 84-88,
48. terminally crutch-like, 32, 8, 26, 32, 4.
absent, — Amazon. S. America; 2. claviger
(Hauer) 274, 100. 2, 16, 24-44, 41-46, ter-
minally broad, 17, ti, 32, 24, 4-5, oralplate
(2), =, Amazon, S. America; 0 claviger i.ssp.,
—532, -120, 2, -21, 155-164, 50-53, termin-
ally broad, 34, 13. 40, 39, 6, present, 18, N,
Aust.

Loc: B, Jabiluka, N.T., W; 23.5. 3.8, 2.4,
=, 30.

Aspelta pitta Harring & Myers, 1928
Loc: B, Wodouga, Vic,, W; 10.2, 7.2, 9.0,
4, 154,

Encentrum gibbosum Wulfert, 1936
Loc: R, Wangaratta, Vie, A; 12,0. 7.7, 10.0,

ORDER Gwesiotrocia De Beauchamp, 1965
Family Testudinellidae

Testudinella parva (Ternetz), 1892
Loc; B, Jabiluka, N.T.. W; 250, 6.2, 3.0,
-, 48.
T. emarginula (Stenroos), 1898
Loc: B, Yea, Vic., A; 12.0, 7.3, 9.8, —-.

R. Benalla, Vic., Su: 27.0, 7.6, 7.0, -, —

T. tridentata Smirnov, 1931
Loe: L, Yarrawonga, Su, 24.2.. 7.7, 8.2, 22,
160.
B, Jabiluka, N.T., W: 24.5, 6.3, 5.8. -, 59.

T. amphora Hauer, 1938
Loc: B, Jahiluka, N-T., Wy 24.5, 6.3, 5.8, -,
59,

142 W, KOSTE & R. J, SHTEL

Fig. 12, Dicranophorus claviger anstraliensis ssp.
a. Dorsal view, total length 532 4m. b. Trophi,
dorsal view, length 53 um, c. Comparative view
of trophi of DB. caydums (Ehrenberg, 1834),
length 35 um,

Testudinella walkeri n.sp.
(FIG. 13a,b,c,d,e)
Type material: 6 9°, preserved in formalin, sample
number 659.

Holotype; Lovicale female, sample number 659,
coll. 13.vi.79, R. J. Shiel, Rot. No. 020, ZMK.

Type locality; Mine Valley billabong, Mageia
Creek floodplain, tributary of Alligator River
near Jabiluka. N.T. (12°40'S/132°50°E).

Description: Juyenile lorica (Fig. 13b,e) widely
oval, strongly circular outline in adults (Fig.
13a). Dorsal lorica slightly concave at widest
point. Ventral Jorica under neck-aperature with
keel-like folding, two edges of which curve
outwards to lower lorica rim. Foot-opening
slit-like under middle of ventral lorica, Dorsal
anterior rim of head-aperature with rounded,
bulge, medially lightly notched (Fig. 13b). In
poor preparations this can appear concave
(Fig. 13¢c). Lorca end with shallow indenta-
tion, Lateral antennae located slightly above
lorica midline. Lorica cross-section shallow,
triangular (Fig. 13d).

Measurements; Lorica length overall 100-112
pm. greatest lorica width 75-98 pm, neck
aperature width 40-44 ym, greatest Jorica
height in midlme 20-25 ym, foot-opening 20
pM Over the posterior Jorica rim, foot-opening
width 20-24 jm.

Discussion: The new species belongs on the
Formenkreis ineisa (see Koste 1978), A
typical form from this group from the same
sample is shown in Fig. 14, All are described
with oval or egg-shaped lorica outlines, as are
the adult individuals of the brycei-amphora
group (Koste 1978). With the exception of T’,
amphora Hauer, 1937, neither of these groups
has the ear-shaped lateral elevation of the
dorsolateral rim of the neck aperture.

The new species is characterised by this
projection on the upper rim of the wide lorica,
by the presence of the two prominent ventral
divergiiig lines which begin at the keel-like
crease under the ventral margin of the neck
aperture,

15(b) Se

15 (a)

Figs 13-15. 13--Yestudinella walker? nsp. a.
Ventral. b, Anterior lonca detail, contracted,
c Juvenile Jorica, ventral. d. Lorica cross.
section, e¢. Species from Malaysia, lorica,
ventral, Single individual collected, Fig, 14—T.
incisa Yar. emarginula (Stenroos), 1898, Lorica
length 110 2m, lorica width 85 em, Fig. 15—
TW ainphora Hauer, 1937 from N.T. a. Dorsal
b. Ventral, oblique. Lorica length 96 am, Joricu
width 78 wm,

NEW ROTIPERA FROM AUSTRALIA 143

A lorica of similar form but lacking the
wide ventral keel was. found in a sample trom
Malaysia (coll, C. H. Fernando, University of
Waterloo, Canada),

Comparative measurements (in pm) are
given in the sequence lorica length, lorica
width. fout-opening, neck aperture, lorica
cross-section: T. walkeri nsp, 100-112, -98,.
slitlike, With lateral “ears” 40-44, shallow trian-
gular, 7. sp. fram Malaysia, 136, 84, angular
16 ™ 10, with “ears” 64, shallow triangular;
T. amplhera, 90-95, 60-78, slittike 18-20,
with “ears”, triangular, T. bryce’, 85, 58, slit-
like, medially tongued-shaped, triangular; T,
incisa ve emarginula, 80-130, 65-82, stitlike,
medially shallow, tongue-shaped shallow tri-
angular,

Loe: B, Jabiluka, N-T., Wy 24.5, 6.3, 5,8,

=~, 59,
Erymalogy: Named after Dr K, BF, Walker,
Department of Zoology, University of Ade-
laide, in appreciation of support during a Ph.D,
program by RIJS,

Pymily Flosculariidae Harring, 1913

Piygura furcillate (Kellicolt), L889
Loc: L, Boort, Vic., So, 23.5, 7.9, 6.0. 47,
750.

Po meliceria vy mucicola (Kellicott), 1889
Loe: B, Wadonga, Vic., Su.

P. tacita Edmondson, 1940
Loc: Bo Yea, Vic., Az 19.9, 7.2. 8&5, 8 m. 35,

Family Hexarthridae Bartos, 1959
Hexurthra polyodonta (Hauer), 1957

Loc: B, Seymour, Vic., Sp: 18.0, 7,2, 9.4, —
265.

Family Filiniidae Barts, 1959
Filinia holimarini Koste, 1980

Loe: R, Manoum, S.A., Su A; —28,0, 7.7-8.4,
-10,.8, —]35, =1080,

F australiensiy Koste, 1980

Loc: R. Mannum, S.A., Su; 17.0, 8.2, 9.7. 88,
1020.

Acknowledgments

The following are thanked for making collec-
tions or intormation available; Dr B. Berzins
(Sweden); De T. J. Hillman (Wodonga); Ms
I, J. Powling (Melhourne) and Mr R. D. Tait
(fabluka), Pancontinental Mining is thanked
far use of facilities and for hospitality during
fieldwork at Jabiluka, N,T. The material from
which these records were derived was collected
by RIS during a Ph.D. programme at the
University of Adelaide supported by a Com-
monwealth Postgraduate Research Grant. Prof
W. D. Williams und Dr K. F. Walker are
thanked for their support during this project,
Microscopic work by WK was made possible
with the assistance of the Deutschen For-
schungsgemeinschaft, Bad Godesberg, Bonn,
F.RG.

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HETEROMORPH AMMONITES FROM THE ALBIAN OF SOUTH
AUSTRALIA

BY K. J. MCNAMARA

Summary

The heteromorph ammonite Protanisoceras Spath is described from Australia for the first time. Five
forms are recognised from the Albian Oodnadatta Formation in South Australia, including P.
auriculum sp. nov. and P. gracile sp. nov. By comparison with species from the Albian of Europe, a
Middle Albain age is suggested for part of the Oodnadatta Formation in which they occur. In
addition Appurdiceras cordycepoides (Etheridge 1905) is redescribed and the relationships of the
genus reassessed; a species of Hamites is described; and the Late Albian species Anisoceras sweeti
sp. nov. described.

HETEROMORIM AMMONITES FROM THE ALBIAN OF SOUTH AUSTRALIA

by K. J. McNAMARA®

Summary

McNamara, K, J, (1980) Heteromorph ammonites from the Albian of South Australia. Trans.
R. Soc. §. Aust. 104(6), 145-159, 28 November, 1980.

The heteromorph ammonite Pratanisoceras Spath is described from Australia for the first
lime. Five forms are recognised from the Albian Oodnadatta Formation in South Australia,
including P. auriculam sp, nov, and P. gracile sp, nov. By comparison with species from the
Albian of Burope, a Middle Albian age is suggested for part of the Oodnadalta Formation
in Which they occur, In addition Appurdiveras eardycepoides (Etheridge 1905) is redeseribed
and the relationships of the genus reassessed; a species of Hayites is described; and the Late
Albian species Anisoceras sweeti sp. noy. described,

Introduction

Ammonites form a much less important part
of the marine Cretaceous fauna of the Great
Artesian Basin in South Australia than ip
Queensland; hence few species have been de-
scribed, Albian ammonites were first described
from S.A, by Etheridge (1905) who de-
seribed “Haploceray sp,, Atiisaceras. (2?) sp.,
Ancyloceras cordycepoides and Criveeras
flindersi”® {rom the area of Dalhousie Springs.
The “Haploceras” is probably Falciferella (sce
helow). Whitehouse (1926) proposed Appur-
diceras to accommodate A. cordycepoides,
while “C.' flindersi belongs in Myloceras
(Reyment 1964a; McNamara 1978), The only
other Albian ammonites to have been de
scribed from S.A. are Faleiferella breadeni and
F. reymenti Brunnschweiler (1959) and a Late
Albian fauna including species of My/oceras
und Labeceray described by Reyment (1964
a,b). Ludbraok (1966) illustrated specimens
which she referred to Myloceray axongiides
(Etheridge) and Lebeceray papularum While-
house.

The present study is based primarily on col-
lections from the Albian Oodnadatta Forma-
lion housed in the palacontolagical collections
of the S.A, Department of Mines & Energy
(GSSA). The specimens were collected from
caloarcous nodules, which occur within a soft
#ypscous shale. mainly from Algebuckina 17
(Ludbrook 19646), 32 km SSE of Oodnadatta
at lat. 27°499°S, lone. 135°344°E. Additional
specimens are from Toodla 8, which is 3 km
SW of MI Arthur at lat, 27°31'S and long.
145°41'B (Fig. 1).

Specitnens identified from these localities
(OODNADATTA 1.250000 geological map
Sheet) are:

Street,

* Western Australian Museum, Francis

Perth, W.A, 6000.

Protaniseceras auriculum sp. noy.. P,
gracile sp. nay., P. aff. gracile sp. nov.,
Hamites cf. attenuatus J. Sowerby 1814,
Falciferella sp. (Ludbrook 1966, p. 44).
The second collection on which this work
is based was obtained by G. Sweet frora the
Maree Subgroup and presented to the National
Museum of Victoria (NMV) in 1914. The
specimens arc recorded as coming from “Prim-
rose Springs, Peake Station”. Ludbronk (1966)
considered that this locality may be near
Primrose Hill at 28°10'S, 136°25°E. From an
carly description, she now considers that
Primrose Hill and Springs 1s a mound spring
with seepage at the base and that the various

Stratigraphy of the | nore

|| Qodnadatta region carr

| c. z “Mount Senin
Kin Sarelstane Mn

DALHOU SLE a_i 4

SPINES

S'5,_0 ~ 500m
|| >| Woanitnd ge
oO, S| Sone Mer =

Ls
| = FORMATION
2 Goatkiana Member
rt ONT
ue
a | Jj)S 2) surtooc
r x
\ a
r= |

200m

SHALE
250m

“CADNA.OWIE "300m
P| “OS ey bt FORMATION
Ovni |

\

VEE CRIN AG «
BY wo
r ' ar Fe
WUMROSL
SURINGS

— «= ailyiy
ah Brwal Paw ality

Fig. |. Map showing fossil localities mentioned in
text. Aptian/Albian boundary after Ludbrook
(1966, Map 2). Stratigraphic information after
Freylag (1966, Fig. 1).

146 kK. J.

specimens collected from *Primrose Springs’
are frorn the general area along he Neales
in the NE of the WARRINA 1:250 000 map
sheet where both Aptian and Albian strata
occur (Ludbrook 1966. Map 6: Thonwon
1980). Specimens identified from this locality
are:
Ajlisoceras sweeti sp. nov. Prorant-
soceras sp. A, P. sp. B, Hamiter cl,
alfenuaiks J, Sowerby 1814, Sarvmartine-
ceras (Singevia) fontinale (Hudleston
1890),
The age of the rocks at this locality is dis-
cussed below,

Finally, Appurdicerus cordycepoides (Ethe-
ridge 1905) is redeseribed and reinterpreted
an the basis of Etheridge’s syntypes from the
Oodnadatta Formation al Dalhousie Springs
{Fiu, 1). one of which is housed in the S.A.
Muscum (SAM), whilst the other has been
located in the NMY, All material was collected
from surface outcrop,

Systematic palacontology
Family ANISOCERATIDAE Hyate 1900
ANISOCERAS Pictel 1854
Type species: Hamites saussureanty Pictel in
Pictet & Roux 1847,
Anisoceras sweeti sp, nov
FIGS 2, 6D

Eyvitelogys Named aller G. Sweet whe
ohtzined the specimeie
Holetype> NMV_ P52328, an incomplete

phragmocone from “Primrose Springs. Peake
Station”.

Paratypes; NMY P52325, an inconiplete
phragmocone, and NMY P52326, an incam-
plete body chamber; both from same locality
as holotype.

Age; Late Albiaiy.

Diagnasis: Species of Anisoeeras in which
lateral and ventro-lateral ribs may cover up to
three ribs; ribs broad and strongly prorsira-
diate on phragmocone, Lateral Jobe of suture
three mes width of umbilical lobe,
Deseription; Complete shell form unknown;
description based on straight fragments, Whorl
height slightly greater than whorl width, Phrag-
mogone known from whorl height of 20-24
mov Ornament consists of coarse, prosiradiate
ribs which arise from Jateral tubercles in twos
or threes, and form loop with circular to
clongate ventrolateral tubercles. ‘The lewped

McNAMARA

ribs form a swollen bundle between which
usually lics one relatively depressed inter
calated ib between the looped ribs, On dorsum
are 12 ribs in a distanee equal to the wharl
height; ribs well developed on dorsum. Maxi-
mum known height of body chamber 27 mm.
Ornament similar to that on phragmocone,
though tubercles a little more widely spaced.
Ventrolateral tubercles more elongate than
on the phragmocone. Lateral lobe of suture
bifid and large, covering more than bulf flank
of phragmocone; three times width and about
twice height of bifld umbilical lobe,

Discussion: A. sweeti is similar to A. armatum
(J. Sowerby 1817) from the Late Albian
Stolicrkia dixpar Zone (Cooper & Kennedy
1979: Scholz 1979) in possessing prominent
looped ribs with usually one intercalated rib.
However, the ribs are only looped in pairs in
A, armatum. Furthermore the ribs are more
strongly prorsiradiate in A. sweerl. The two
species can most easily be distinguished by the
relative sizes of the lateral and umbilical lobes
of the suture. In 4. armatum they are of
approximately equal size or the lateral is just
slightly larger than the umbilical (Scholz 1979,
Fig. 8), In A. sweeti the lateral Jobe is very
large, being three times the width of the
umbilical lobe

In A. heasi Cooper & Kennedy (1979)
three, or four, cribs may be looped between the
tubercles, Mowever, the nbs are much finer
than in A. sweeti.

APPURDICERAS Whitehouse 1926

Yvpe speciexr Ancyloceras cordycepoides
Etheridge 1905 p, 14, Pl 1, figs 3-5, Pl, 2,
fig. 4; by original designation of Whitehouse
1926, p. 229.

Emended diagnosiv: Coiling ancyloceratid,
Ribs strongly developed, widely spaced; bifur-
cule laterally and at umbilical tubercles on
shaft. Ventrolateral spines regularly developed,
bases covering one to three ribs. and forming
‘loop and button’ ornament across venter-
Ventrolateral spines on different ribs from
umbilical tubercles, Umbilical lobe of suture
bifid.

Remarks: Whitehouse (1926) — proposed
Appurdiceray lo accommodate Aneyvloceras
cordycepoides Etheridge, distinguishing it from
other Australian heteromorphs by its
proninent ventrolateral spines. Whitchouse
Yuestioningly placed a single labeceratid speci
men from Old in thes genus, calling it 4.(?)

ALBIAN HETEROMORPH AMMONITES 147

Fig, 2. Anisoceras sweeti sp. nov.; NMV P52328, holotype, A, lateral view, B, ventral view; C, NMV
P52326, paratype, dorsolateral view; from “Primrose Springs”, Peake Station; Oodnadatta Formation,
Late Albian; all x 1.

etheridgei, with the result that Appurdiceras
itself came to be thought of as a subgenus of
Labeceras (Wright, in Arkell et al. 1957;
Reyment 1964a; Klinger 1976) which bears
ventrolateral tubercles. A.(?) etheridgei is
quite unlike A. cordycepoides, being a true
labeceratid which has ribs which do not bifur-
cate on the flanks of the shaft, as they do in
A. cordycepoides, and which do not form the
characteristic anisoceratid looped ribs across
the venter. The umbilical lobe of labeceratids
is trifid, whereas in Appurdiceras it is bifid.

Consequently, Appurdiceras is herein
regarded as an anisoceratid and not a sub-
genus of Labeceras. Whitehouse (1926)
included Jdiohamites spiniger (J. Sowerby)
from the English Late Albian in Appurdiceras.
He also noted a similarity with Hamites
nodosus J, Sowerby. This latter species was
made the type of Heteroclinus by Casey
(1961). However Klinger (1976) places this
genus in synonymy with Protanisoceras (see
below).

Unlike species of Anisoceras and Protani-
soceras, which may bear both ventrolateral
and lateral tubercles or spines, Appurdiceras
possesses ventrolateral and umbilical tubercles.
It can also be distinguished from these two
genera by the umbilical bifurcation of the ribs
on the shaft at tubercles which do not form
by the coalescence of the ribs bearing the
ventrolateral tubercles or spines, but by the
coalescence of an intercalated rib with one
which does bear a ventrolateral tubercle, The

ribs may also occasionally bifurcate laterally.
Species assigned to /diohamites tend to have
numerous, fine ribs between the tuberculate
ribs (Spath 1939), though as Klinger (1976)
has noted, /diohamites grades into Anisoceras.
Anisoceras and Protanisoceras differ from one
another in the nature of the suture (see
below). Whereas the umbilical lobe is trifid
in Protanisoceras it is bifid in both Appurdi-
ceras and Anisoceras.

Appurdiceras cordycepoides (Etheridge 1905)
FIG. 3
1905 Ancycloceras cordycepoides Etheridge, p.
14, Pl. 1, figs 3-5, Pl. 2, fig. 4.
1909 Crioceras cordycepoides (Etheridge);
Etheridge, pp. 142, 156, 159, 160.

1926 Appurdiceras cordycepoides (Etheridge);
Whitehouse, p. 230.

21964 Labeceras (Appurdiceras) cordycepoides
(Etheridge) Reyment, p. 25, ?PI. 1, figs
6 & 7.

1969 Appurdiceras cordycepoides (Etheridge);
Day, p. 156.

non

1966 Labeceras (Appurdiceras) cordycepoides

(Etheridge); Ludbrook, p. 190.
Lectotype: Herein designated: NMV P30032,
from Dalhousie Springs (DALHOUSIE
1:250000 geological map sheet) figured by
Etheridge (1905, Pl. 2, fig. 4).

Paralectotype: SAM P2990 from Dalhousie
Springs; figured by Etheridge (1905, Pl. 1,
figs 3-5).

Age: Late Albian.

148 K. J, MCNAMARA

Diagnosis: As for genus,

Description: Phragmocone initially ovoid in
cross section and forms an open criocone for
one whorl, then opens into a shaft. Whorl of
lectotype has maximum radius of 22.8 mm. At
a quarter whorl, whorl height is 4.5 mm; at
commencement of shaft it is 8.5 mm. Early
part of whorl damaged, and it is not known
whether ribs are tuberculate. Ribs are known,
however, to be rectiradiate and not bifurcant.
Whorl section of shaft sub-circular. Ornament
changes on shaft such that ribs become more
widely spaced than on coiled, early part of
phragmocone. Early ribs on shaft infrequently
bifurcate on flanks. Bifurcations more common
along shaft toward crozier. At these bifurca-
tions tubercles are not developed. Bifurcations
more commonly occur umbilically between a
rib which bears a ventrolateral spine and an
intercalated non-spinose rib. Ribs more
strongly developed along shaft. Ventrolateral
spines appear as rounded tubercles on internal
mould. Across venter ribs which connect
spines form a raised swollen band. A ‘button
and loop’ ornament is formed across ventor by
coalescence of three ribs at ventrolateral
spines. Initially ribs on shaft rectiradiate, but
become increasingly prorsiradiate adapertually.
On dorsum ribs less well-developed, but do not
disappear. Shaft almost twice length of maxi-
mum diameter of coiled phragmocone, Ada-
perturally the body chamber recurves to form
crozier. Paralectotype reaches a maximum
whorl height of 16 mm and a maximum whorl
width of 14.5 mm. Suture incompletely
known: umbilical lobe bifid and narrow;

internal lobe trifid.

Discussion: Since the original description of
the two type specimens the only specimens
ascribed to this species are two from the Late
Albian of Fossil Creek, Wooldridge Limestone
Member, Oodnadatta Formation, locality
5/550/1, 42 km NW of Oodnadatta (Reyment
1964a) and specimens from Oodnadatta For-
mation, Algebuckina 17 (5/571/17), 13 km
W of Mt Dutton (Ludbrook 1966). However,
it is not possible for me to assign Reyment’s
fragmentary specimens to A. cordycepoides
with any certainty as, although seemingly pos-
sessing an ovoid to subcircular whorl section
and ventrolateral tubercles, there is no indica-
tion of bifurcation on the flanks of the shaft,
In his description of these specimens Reyment
does suggest the presence of occasional bifur-
cations. If these specimens are true members
of A. cordycepoides their occurrence with
species of Myloceras and Labeceras indicates
that Appurdiceras may be a Late Albian form.
Specimens referred to A. cordycepoides by
Ludbrook are species of Protanisoceras (see
below).

Etheridge’s specimens were collected by
H. Y. L. Brown from the region of Dalhousie
Springs. Ludbrook (1966) has recorded both
Albian and Aptian molluscs from this area.
In addition to describing A. cordycepoides
from the Dalhousie area, Etheridge described
and illustrated other ammonites which have a
Late Albian, rather than Aptian, affinity, in-
cluding the Late Albian Myloceras and
Labeceras,

The development of ventrolateral spines is
seen in other anisoceratids, such as Anisoceras
(Klinger 1976), Idiohamites (Spath 1939)

, lectotype, lateral view; SAM

P2990, paralectotype, B, lateral view, C, ventral view; from Dalhousie Springs; Oodnadatta Forma-

tion, Late Albian; all x 1.

ALBIAN HETEROMORPH AMMONITES 149

and Protanisoceras (Spath 1939). The two cordycepoides, differ in lacking the bifurcation
species from the English Late Albian, Idio- of the ribs on the flank of the shaft and having
hamites spiniger and Protanisoceras nodosum, lateral tubercles developed on the same ribs as
with which Whitehouse (1926) compared A. the ventrolateral spines.

Fig. 4. Protanisoceras auriculum sp. nov.; GSSA M2416, holotype, lateral view; from Algebuckina 17,
32 km SE of Oodnadatta; Oodnadatta Formation, Middle Albian; x |.

150 K. J. MCNAMARA

PROTANISOCERAS Spath 1923

Type species: Hamites raulinianus d’Orbigny
1842, p. 546, Pl. 134, figs 5-8; by original
designation of Spath 1923, p. 75.

Remarks: Spath (1939) distinguished Protani-
soceras from the morphologically similar Ani-
soceras by its smaller size, the more regularly
planar coiling and simpler suture line which
has a trifid umbilical lobe. Klinger (1976) has
noted that this lobe is smaller than the lateral
lobe in Protanisoceras, whereas he believed
that it was of similar size to the lateral lobe in
Anisoceras. The species of Anisoceras de-
scribed here shows that even in some members
of this genus the umbilical lobe may be much
smaller than the lateral lobe. However, the
trifid umbilical lobe of Protanisoceras and
bifid umbilical lobe of Anisoceras are
diagnostic.

Klinger (1976) has further noted that the
coiling and ornamentation are variable within
Protanisoceras. Both ventrolateral and lateral,
or just ventrolateral tubercles or spines may
be present; they may appear on all, or only
some, ribs. The tubercles may be connected
by one or more ribs, Furthermore, the orna-
mentation on the recurved crozier may vary
considerably from the shaft.

Casey (1961) regarded P. nodosum as
belonging within a separate genus, which he
named Heteroclinus, on account of its ‘button
and loop’ ornament. Klinger (1976) has pre-
ferred to regard Heteroclinus as a synonym of
Protanisoceras as species such as P. parce-
tuberculatum Collignon show intermediate
characters between Protanisoceras and Hetero-
clinus in possessing both single and looped
ribs. This situation also occurs in one of the
Australian species described below.

Protanisoceras auriculum sp. nov.
FIGS 4, 5, 6B, 9A-—C
Etymology: Latin—auriclum—ear; pertaining
to the shape of the shell.

1966 Labeceras (Appurdiceras) cordycepoides
(Etheridge) (pars.); Ludbrook, p. 190.
1966 Myloceras axonoides (Etheridge); Lud-

brook, pp. 44, 190 (pars.), Pl. 28, fig. 1.
Holotype: GSSA M2416, part of the phragmo-
cone and body chamber (Figs 4, 5A); from
Oodnadatta| Formation, Algebuckina 17
(5/571/17) (Ludbrook 1966), 27°492’S,
135°344’E, 32 km SE of Oodnadatta, S.A., on
North Creek, near its junction with Neales
River (OQODNADATTA 1:250000 geological
map sheet).

Paratypes:; GSSA M2444, 3546, from the
same locality as the holotype and M3547 from
Toodla 8 (5/561/8), 3 km SW of Mt Arthur,
27°31°S, 135°41E,

Other material: In addition to the type speci-
mens, 12 further specimens are known: GSSA
M2446, 2454, 3548-3556 from Algebuckina
17 and M3061, 3557 from Toodla 8.

Age; Middle Albian.

Diagnosis: Coiling aspinoceratid. Ventrolateral
tubercles occur infrequently on single ribs on
early phragmocone and late body chamber;
occur more frequently on later phragmocone
and early body chamber where may cover two
or three ribs. Ribs broad and widely spaced on
early phragmocone; becoming finer and more
closely situated adaperturally.

Description: Phragmocone forms open coil
throughout. Whorl section ovoid throughout,
whorl height slightly greater than whorl width.
At earliest known part of phragmocone (Fig.
SE) at whorl height of 4 mm, ornament con-
sists of simple, non-tuberculate, slightly pror-
siradiate ribs. At this stage there are four ribs
in a distance equal to whorl height. At slightly
greater whorl height every fifth or sixth rib is
flattened across venter and small tubercles
occur ventrolaterally. Up to a whorl height of
9 mm ribs become increasingly prorsiradiate;
ribs more strongly inclined toward dorsum.
Tubercles at this whorl height occur more
frequently and become elongate, spreading
across two ribs; up to two intercalated non-
tuberculate ribs. Across venter ribs form loop
between tubercles and on latter part of phrag-
mocone form swollen band. Tuberculate ribs
may also form swollen band on flanks. At
whorl height of 10 mm tubercles may cover
three ribs and be separated by only one or two
intercalated ribs. Ribs on dorsum pass straight
across and are weaker than on flanks, Between
whorl height 14 mm and 26 mm ribs become
more rectiradiate. At whorl height of 30 mm
there are 11 ribs in a distance equal to whorl
height.

Body chamber commences at whorl height
of 32 mm. Tubercles become more widely
spaced, covering 2-3 ribs being separated by
up to 7 intercalated ribs. Last large tubercles
occur at whorl height of 41 mm. Tubercles
may become spinose on early body chamber,
but progressively decrease in size adapertur-
ally, covering only one rib but occurring on
every rib. On body chamber ribs become almost
rectiradiate; they become increasingly sinu-

ALBIAN HETEROMORPH AMMONITES 151

Fig. 5. Protanisoceras auriculum sp. nov.; A, GSSA M2416, holotype, ventral view; B, GSSA M2444,
paratype, lateral view; GSSA M3547, paratype, C, lateral view, D, ventral view; E, GSSA M3550,
lateral view. A, B, E from Algebuckina 17, 32 km SE of Oodnadatta; C, D from Toodla 8, 3 km
SW of Mt Arthur; all Oodnadatta Formation, Middle Albian; all x 1.

soidal adaperturally. Body chamber reaches a
maximum known whorl height of 50 mm.
Suture line with broad, bifid lateral lobe and
small, trifid umbilical lobe, half width of
lateral lobe; internal lobe trifid (Fig. 6B).

Discussion: Klinger (1976) has expanded the
concept of Protanisoceras to include those
forms which have tubercles spreading across
more than one rib. This looped ornament, so
well developed in P. auriculum, is seen in the
European Joricatus Zone (mid-Middle Albian)
P. nodosum and P. flexuosum (d’Orbigny). P.
auriculum can be distinguished from these
species by its more elongate tubercles which
extend over a greater number of ribs on the

phragmocone, more prorsiradiate ribs and less
ornamented body chamber. The Madagascar
species P. parcetuberculatum Collignon (1962),
like P. auriculum, possesses both single and
looped ribs. It can be distinguished from P.
auriculum by its narrower ribs and smaller
tubercles.

Protanisoceras gracile sp. nov.
FIGS 6A, 7, 9D, E
Etymology: Latin
1966 Labeceras (Appurdiceras) cordycepoides
(Etheridge) (pars.); Ludbrook, p. 190.
1966 Myloceras axonoides (Etheridge) (pars.);
Ludbrook, p. 44.

eracilis—slender.

(52 kK. 1. McNAMARA

Hololype; GSSA M2455, from Oodnadatia
Formation, Algebuckina 17 (5/571/17) (Lud-
hrook 1966), 27°494’S, 135°344°E, 32 km SE
of Oodnadatta, on North Creek, near its
junction with Neales River (OODNADATTA
1:250 000 geological map sheet),

Paratypes: GSSA M3558 from same locality
as holotype and M3059 from Toodla 8, 3 km
SW of Mt Arthur,

Other material: GSSA M2449, 3560-3567
from Algebuckina 17 and M3568-3569 from
Toodla 8.

Ape: Middle Albian.

Diagnosis; Coiling ancyloceratid; whorl see-
tion ovoid, Ventrolateral tubercles infrequent,
occurring only on shalt and restricted to single
ribs. Ribs prorsiradiate throughout, except on
recurved hook where become rectiradiate
adaperturally.

Description; Earliest known part of phrag-
mocone with whorl helght of 3 mm; mod-
crately strongly coiled; with single, non-tuber-
culate, gently prorsiradiate ribs. Tubercles first
uppear al whorl height of 9 mm as small
prominences either side of siphonal line. On
early part of shaft tubercles occur on about

A
U L E
fo 3s animes
Ves Soe at : r
UP Ww) aa
\ 2 vA
aN
C.
| i |
70mm

every sixth rib. At Whorl height of 10 mm ribs
become sinuously prorsiradiate, more strongly
inclined toward dorsum. Ribs thicker and
more widely spaced along shaft. At whorl
height of 12-14 mm small tubercles present on
all ribs in some individuals, but Jess frequently,
or not at all, on others, On recurved body
chamber tubereles absent. Adaperturally ribs
on body chamber wider and rectiradiate.
Whorl section ovoid throughout shell with
width §2% whorl height, with maximum whorl
height on recurved body chamber uf 19 mm,
Suture line like that of P. auriculam.

Discussion; P, gracile can be easily distin-
guished from P. evriculum by its smaller size:
possession of a shaft; weaker tuberculation;
absence of looped nbs and. slightly smaller
lateral lobe. P. gracile is most similar to P.
nodaxum from the Middle Albian /ericatus
Zone in southern England in lacking lateral
tubercles and having ventrolateral tubercles
which do not occur on all ribs, P. gracile
differs in its lack of looped ribs, tubercles
Which are not spinose, possession of more
strongly prorsiradiate ribs and absence of
luberculation on the body chamber. #&
flexuosum, also from the Joricatuy Zone, simi-

Fig. 6, Suture lines of: Ay Protunisoecras gracile 8p. noy., GSSA M2455, holotype, at whorl height of

12.5 mms BL P. avricuham sp. noy.,

GSSA M3546, paratype, at whorl height of 16 mm; C, Proiani-

soceray sp. B, NMV_ P52327, at whorl height of 14 mm; D, Anisoceras sweeti sp, nov. NMV

P52328, holotype, at whorl height of 23 mm, 1 — inlernal lobe; U = umbilical lobe: L

E © external lobe.

lateral lobe:

ALBIAN HETEROMORPH AMMONITES 153

Fig. 7. Protanisoceras gracile sp. nov.; GSSA M2455, holotype, A, lateral view, B, ventral view; C,
GSSA M3059, paratype, lateral view; GSSA M3558, paratype, D, lateral view, E, ventral view;
A,B,D,E, from Algebuckina 17, 32 km SE of Oodnadatta, C from Toodla 8, 3 km SW of Mt Arthur;
all from Oodnadatta Formation, Middle Albian; all x 1.

larly lacks lateral tubercles and possesses only
small ventrolateral tubercles like P, gracile;
however in P. flexuosum they occur more
frequently.

P. gracile is similar to some species from the

English Early Albian mammillatum Zone,
such as P. raulianum (d’Orbigny), P. canti-

anum (Spath) and P. blancheti (Pictet & Cam-
piche). These species, like P. gracile, possess
only ventrolateral tubercles and lack looped
ribs, However, P. gracile can be distinguished
from these species by its more strongly pror-
siradiate ribs. P. gracile resembles P. gradatum
Collignon (1963, p. 40, Pl. 256, fig. 1101)
from the Early Albian of Madagascar, but it
is smaller and possesses more inclined ribs.

Protanisoceras aff. gracile sp. nov.
FIGS 8, 9F

Material and locality: One specimen, GSSA
M2441, from Oodnadatta Formation, Alge-
buckina 17 (5/571/7), 32 km SE of Oodna-
datta (OODNADATTA 1:250000 geological
sheet map).

Age: Middle Albian.

Remarks: This specimen, consisting of the
latter part of the shaft and the recurved body

chamber, differs from P. gracile in the pos-
session of tubercles which extend across two
ribs on the latter part of the phragmocone.
Like P. gracile the body chamber lacks tuber-
culation; however, the tighter recurving of the
body chamber has resulted in the umbilical
bifurcation of some ribs. In terms of whorl
section and size this specimen conforms with
P. gracile, but its more strongly developed
tuberculation and tighter recurving are dis-
tinctive.

Protanisoceras sp. A
FIGS 9G, 10A, B

Material and locality: An incomplete shaft,
NMV P60543, from “Primrose Springs, Peake
Station”.
Age: Middle Albian,
Description: Whorl section circular with dia-
meter of 16 mm. Ribs slightly prorsiradiate;
each rib bears small ventrolateral and lateral
tubercle. Tubercles cover only single rib. There
are eight ribs in a length equal to whorl dia-
meter,
Discussion: This form can be easily distin-
guished from P. sp. B by the absence of looped
ribs with tubercles covering more than one rib,

154 Ke Ts

McNAMARA

Fig. 8. Protanisoceras aff, gracile sp. nov.; GSSA M2441, A, lateral view, B, ventral
view; from Algebuckina 17, 32 km SE of Oodnadatta; Oodnadatta Formation, Middle

Albian; both x 1.

The absence of suture and the impression of
the dorsum of the phragmocone on the matrix
attached to the specimen indicate that the
specimen is the recurved body chamber. It is a
much longer body chamber than possessed by
other Australian species of Protanisoceras of
similar size. The ribbing is finer and more
closely spaced than seen in English Early-
Middle Albian species. The Middle Albian P.
cantianum Spath (1939) from southern
England has a similar whorl section and tuber-
culation, but it possesses thicker, more widely
spaced ribs.

Protanisoceras sp. B
FIGS 6C, 9H, 10C, D

Material and locality: A single, incomplete
phragmocone, NMV P52327, from ‘‘Primrose
Springs, Peake Station”.

Age: Middle Albian.

Description: This specimen, an incomplete,
largely internal mould of the latter part of the
phragmocone, is characterised by possession of
large ventrolateral and lateral tubercles of
similar size. Tubercles connected by looped ribs
and extend across three ribs; separated by 0—2
intercalated non-tuberculate ribs. Whorl section
is semicircular. This form particularly charac-
terised by large, bifid lateral lobe (Fig. 6C)
which is almost half as wide again as first
saddle; it is four times wider than_ trifid
umbilical lobe.

Discussion: Specimens of Protanisoceras
described by Klinger (1976) from Zululand

also possess both ventrolateral and lateral
tubercles. However, in all the forms
Klinger described the tuberculate ribs are

much more widely separated by non-tuber-
culate ribs than in the S.A. forms. P. sp. B
compares with some English Late Albian
species of Anisoceras, such as A. armatum,
in the nature of the tuberculation; however,
whereas species of Anisoceras possess a bifid
umbilical lobe it is trifid in P. sp. B. The
ornamentation is like that of A. sweerti. The
two species can be distinguished by the nature
of the umbilical lobe and the smaller size of
P. sp. B. This species differs from P. auriculum
and P. gracile in its possession of lateral
tubercles.

Family HAMITIDAE Hyatt 1900
HAMITES Parkinson 1811
Type species: Hamites attenuatus J. Sowerby
1814, by subsequent designation of Diener
1925, p. 65.

Hamites cf. attenuatus J. Sowerby 1814
FIG, 11

Material and localities: A complete body
chamber, NMV_ P52336 from ‘Primrose
Springs, Peake Station”, and an incomplete
body chamber, GSSA M2447, Oodnadatta
Formation, Algebuckina 17 (5/571/17).
Age: Middle Albian.

Remarks: The well preserved body chamber
has an almost circular whorl section. Like H.
attenuatus from the Middle Albian dentatus-

ALBIAN HETEROMORPH AMMONITES 155

Fig, ¥, Diagrammatic representation of variation in distribution of ribs and tubercles in species of
Protanisoceras. A-~C, P. auriculum sp. nov., A, GSSA M3550, B, GSSA M3547, paratype, C, GSSA
M2444, paratype; D-E, P. gracile sp. nov., D, GSSA M3558, paratype, E, SADM M2455, holo-
type, F, P. aff, gracile sp. nov., GSSA W2441, G, PF. sp. A, NMV P60543; H, P. sp. B, NMYV P52327,

All natural size.

loricatus Zones of southern England, the
S.A. form possesses ribs which are slightly
prorsiridiate prior to the hook; rectiradiate
on the hook, becoming initially rursiradiate
on the recurved shaft, then finally rectira-
diate. Similarly it has 7-8 ribs in a length
equal to the whorl diameter. The whorl section
of the English form is said to be slightly
laterally compressed (Spath 1939) whereas it
is slightly dorsoventrally compressed in the
S.A. form.

In both forms the adapertural part of the
body chamber curves away from the phragmo-
cone, The impression of the dorsum of the
phragmocone against the body chamber (Fig,
111A) shows that the body chamber was
deflected away from the phragmocone during
growth as the two shafts came into contact.

The suture line of the $.A. form is of similar
proportions to the English form, with a bifid
lateral Jobe and small trifid umbilical lobe,
Whitehouse (1926) described a torm from the
Albian of Qld which he called A. aff. maxi-
mus J. Sowerby. This specimen, as can be seen
in’ Whitchouse’s figure, possesses a trifid

lateral lobe, whereas in Hantites it is bifid:
it iy thus not a species of Hammites.

Age of the ammonites

The Marree Subgroup comprises the Aptian
Bulldog Shale and the Albian Oodnadatta
Formation (Freytag 1966, Ludbrook 1966,
1978, 1980). Since Ludbrook’s (1966) bio-
stvatigraphical study was submitted for pub-
lication, the OODNADATTA 1:250 000 geo-
logical map sheet has been published (Freytag
et al. 1967) and the rock units comprising the
Aptian-Albian sequence named in some detail
(Freytag 1966). It is now possible to relate the
ammonites to the rock units, as mapped on the
OODNADATTA and adjoining sheets, from
which most of them were collected.

The ammonite Sanmartinvceras (Sinzovia)
/ontinale from the Marree Subgroup at *Prim-
rose Springs, Peake Station’ was described by
Hudleston (1890), Thomson (1974) discussed
the range of Sanmartinoceras and believed
that it is, by and large, an Aptian form, The
oecurrence of S. (Sinzevia) in association with
Tropaeum in Qld led Day (1969) to conclude

156 K. J. MCNAMARA

Fig. 10. Protanisoceras sp. A; NMV P60543, A, lateral view, B, ventral view.
Protanisoceras sp. B, NMV P52327, C, lateral view, D, ventral view. Both from “Primrose Springs,
Peake Station”; Oodnadatta Formation, Middle Albian, both x 1.

Fig. 11. Hamites cf. attenuatus J. Sowerby 1814; A, lateral view, B, ventral view;
from “Primrose Springs, Peake Station”; Oodnadatta Formation, Middle Albian;
x 1. Arrow indicates impression of dorsum of part of phragmocone.

a Late Aptian age for the genus in Australia.
A Late Aptian age for part, at least, of the
Bulldog Shale is further indicated by the
presence of Tropaeum in S.A. (Howchin &
Whitehouse 1928).

In her biostratigraphical study of the
Cretaceous rocks of the Great Artesian Basin
in S.A. Ludbrook (1966) concluded that
there was no reliable means of establishing a
detailed biostratigraphic zonation of the

ALBIAN HETERDMORPH AMMONITES 157

Marree Suheroup between the Late Aptian
and Late Albian an the evidence of animonites.
Part of the reason for this hus been the aftribu-
tion of any small heteromerph ammonite
{ound in the formation to the Late Albian
genera Myloceray or Lubeceras (eg. Lod-
hrook T8966), although it has naw been shown
Unat a numbec of other geners ave present. Ut
wiso stems from the inaccurate stalement of
Brunnschweiler (1959) who, in deseribing
species of Falcitereiia tram the Santos Oodna-
datta Wo. 1 Well, stated, “The heds from the
surface dowh to about 375 fect contain also
Myloceray, Labeceras, Appurdiceras, Bolire-
cerus ete, and are to he regarded as early
Upper Albian - -) Palciferella he recorded
hetween 71,5 m (235 ft) and 113.3m (372 ft)-
Vhis icd Bninnschweiler to conclude that
fuleiferclla was of Late Albian age in
Australia (even though it is resericted ta the
Mickle Albian interinedtus afl niobe Sub-
zones of the favicatas Zone in England (Owen
1971)) as the Lebeecrays—-Myluceras faana in
Old, Madagascar and Zululand has been
established as Late Albian (Whitchouse 1926,
Klinger 1976, McNamara 1978). However, as
Ludbrook (1966) has noted, this Late Albian
latina does not occur with Fuleiferella in the
Oodnadatta No. | Well. Brunnschwelller was
probably referring to the Late Albian
ammonite fauna of Fossil Creek (Reyment
YA aly).

Ludhrook (19667 ploced the “unnamed green-
sand member” (now the Coorkiina Member
of the Oodnadathe Formation), which ovenrs
in Santos Oodnadatta Na, 1 Well between 13)
and 137 m, ar the base of the Albian, She
records Falciferclla in the well between this
member and the lop of the Oodnadatta Forma-
tion, Which is thoughe to be of Late Albian age
(Ludbrook 1978), at bout 91 om, This is sug-
veslive of a Midle Athian age for Faleiferelfa
in Australia as in Eneland,

Cooper & Kennedy (1979) have recently
placed the two deseribed Australian species
F, hreadent and F. reymenti, im the binneyitid
Norissiakareras, which, apart from an tmacom-
plete specimen feom the uppermost Albian
of Angola, is restricted to the Late Cero-
manian to Early Turonian (Wright ta Arkell
+f al. 1957), Cooper & Kennedy consider that
jhere is a qlircet phylogenetic relationship
belween the Middle Albian Falciferclia, with
its Faleoid growth lines and ribs. and the almost
smooth Berissiakeceras. Following Brunn-
schweiler's assignment of 4 Late Albian age to

BP. hreadent and FP. reymenii, Cooper &
Kennedy suggested that these species may
helons in Borisstakoceras, However, these
species, like the type species of Fulciferclla, F,
millbaurnei Casey (1954), have tine ribs on
the body chamber. and a trifid lateral lobe,
which in Borissigkoceray is bifid (Cobban
1961), Examination of the type and topotype
material of F. breadeni fram the Qodnadatra
Formation, reveuled the suture line to be more
erenulate than shawn by Brunnschweiller
(1959), being very close to that af £, mill-
bourne’. Furthermore the occurrence of F,
hreaden’ and FF. reymenti with the Middle
Albin species af Pratanivaceras (see below)
and a Middle Albian species of Marites, and
possessing Tibbing which 35 more akin to Fal-
ciferella than to Borissiakoceray, Sugeest that
Brunuschweilers ariginal emplacement in
Falciferella is) more appropriate. Obviously
there is a close relationship between these two
genera, sufficient for Kennedy & Juignet
(1973) to have placed Fale‘/eretla in the
Binneyitidae anc not the Oppeliidac os did
Wright (in Arkell e¢ al 1957),

At Algebuckina 17 Faletferella occurs (Lud-
hrogk 1966) with Protanisaceras auriculiin
und P. yraete. Speeies of Protanisacerey alse
occur at “Primrose Springs, Peake Station’.
Protanisoceras was considered by Wright (in
Arkell et ei. 1957) io range from the Early-
Middle Albian. “The yougest species in the
English Middle Albian aceurs in the inivr-
inediny Subzone of the foricaray Zone (Qwen
1971), Klinger (1976) extended its lower
range into the Lale Aptian. Casey (1861) has
shawn thal the vurliest English Albian speeies
of Pretanisoeeras, whieh appear im the
flortdion Subzone of the manmilldtian Zoe,
compare closely with cantemporancous species
ul Mamites, differing only in the develop-
ment of small vertrolateral tubercles which
cover single ribs, In the succeeding rau/fanun
und pucianus Subzones there is a general trend
toward inereasing tuberculatian, the species
possessing more Frequent and prominent ven-
trotateral tubercles and the development of
lateral tubercles! these species coexist with
poorly tuberculute species, In the early Middle

Albian dentures and early Jorfearuy Zones
(Spath L939) species with lateral tubercles

predominate. and an increasing number of
species, sich as FP. nodes and P, flexvasum
oF the foricatis Zone (Owen 1971), have
looped rihs. In these Tnfer species there is a
loss of lateral tubercles. as oceurs in PL auricu-
lum and P grecile.

158 kK. J, McNAMARA

The occurrence at Algebuckina 17 of species
of Protaniseceras morphologically closest to
the Jericatis Zone species, coexisting with
species of Falciferella, which oeeurs only tn
the first two subzones Of the loricarus Zone
(Owen 1971), and with Mamites cf artenu-
alas, Which ranges from the late deratus Zone
through the /oricatus Zone, suzeests that this
part of the Godnadatta Formation may cor-
relate with the mid-Middle Albian Jertcunes
Zone of the Angzlo-Paris Basin-

The locality cited as “Primrose Springs,
Peake Station” in the old literature probably
includes more than one locality, ws stated
above, Until the WARRINA 1:250 000 geo-
logical map sheet is mapped in detail, it will
not be possible to identify the precise localitics
From which the Late Aptian Sanmerrinoceras
(Sinzevia) fontinale, and the Middle Albian
species of Pretanixoceras with looped cibs aud
Hanites cf. atretuatus, were collected In
audition, the occurrence of Anisocéras sweetl

indicates that younger strata also outerop in
this region. ws dhe genus ranges from the Late
Albian to the Late Turonian (Wright in Arkell
et al. (957). As the Blanchewater Formation
Which o¥erlies the Marree Subgroup in the
Marree area is thought to be latest Albian in
age, possibly extending into the Cenomanian
(Ludbrook 1966, 1978), the upper part of the
Oodnadatla Formation is early Late Albian in
age. This has been established at Fossil Creek
by the presence of My/oceras and Labeceras,

Acknowledgements

1 thank Mr J. M. Lindsay (GSSA), Mr N-
§. Pledge (SAM) and Mr T. A. Darragh
(NMY) for the loan of specimens; Dr N, H,
Ludbrook for drawing to my attention the
existence of the GSSA specimens and provid-
ing information on the sttatigraphy; Or W, J.
Kennedy (Oxford) for kindly assisting me
with literature; and Miss Val Ryland for
photographing the specimens,

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AGES AND ASHES IN LAKE FLOOR SEDIMENT CORES FROM VALLEY
LAKE, MT GAMBIER, SOUTH AUSTRALIA

BY C. E. BARTON & M. W. MCELHINNY

Summary

A set of our cores from Valley Lake shows a sequence of fresh water organic muds above a band of
argonite 8-10 cm thick, overlying a graded calcareous tuff. Magnetic remanence and susceptibility
measurements indicate the absence of volcanic episodes since the onset of deposition of the organic
muds 5000 to 6000 years ago. Radiocarbon ages of ~14 000 years and ~38 000 years for the
aragonite band and the tuff respectively are not considered to reflect their ages of formation.

AGES AND ASHES IN LAKE FLOOR SEDIMENT CORES FROM VALLEY
LAKE, MT GAMBIER, SOUTH AUSTRALIA

by C. E. Barton*+ & M. W. McELHINNY*

Summary

Barton, C. E. & MCELHINNy, M. W. (1980) Ages and ashes in lake floor sediment cores
from Valley Lake, Mt Gambier, South Australia, Trans. R. Soc. S. Aust. 104(6), 161-165, 28

November, 1980.

A set of four cores from Valley Lake shows a sequence of fresh water organic muds above
a band of aragonite 8-10 cm thick, overlying a graded calcareous tuff. Magnetic remanence
und susceptibility measurements indicate the absence of volcanic episodes since the onset of
deposition of the organic muds 5000 to 6000 years ago. Radiocarbon ages of ~14 000 years
and ~38000 years for the aragonite band and the tuff respectively are not considered to

Teflect their ages of formation.

Introduction

Valley Lake (37° 51'S, 140° 46’E) is the
second largest of the four lakes in the Recent
volcanic craters at Mount Gambier, South
Australia. Chemical and biological aspects of
the lake have been described by Bayly & Wil-
liams (1964), and the morphology and benthos
by Timms (1974). The lake is now fresh and
at a level considered to be controlled by the
ground water table.

The geology of the Mt Gambier complex
has received considerable attention, the most
recent being a detailed study by Sheard (1978)
which includes a review of the previous work.
In Sheard’s reconstruction the two larger lakes,
Blue Lake and Valley Lake occupy open
craters called maars, formed by volcanic ex-
plosions, as do the smaller lakes, Brownes
Lake and Leg of Mutton Lake.

Two charcoal samples picked from soils
beneath tuff layers have been radiocarbon
dated. The first was collected by C. G. Stephens
in the township of Mt Gambier and dated
at 4830 + 70 BP (Gill 1955; Fergusson &
Rafter 1958), and the second was collected
4 miles away to the SSE and dated at 1410
+ 90 BP (Blackburn 1966). Blackburn was
of the opinion that these dates may represent
separate volcanic episodes, and this view has
also been adopted by Sheard (1978).

Four cores were recovered in 54 mm (class
12) PVC tubes using a 6 m Mackereth corer
(Mackereth 1958): VB in 16 m of water
from the deepest part of the lake, and VA, VC

* Research School of Earth Sciences, Australian

National University, P.O. Box 4, Canberra,
A.C.T. 2601.
* now at: Geophysics Department, Edinburgh

University, U.K.

and VD in 14.5 m of water from the flattish
area about 50 m NE of the deepest part (see
Timms (1974) bathymetric chart). Echo
soundings showed small scale variations in
bottom topography not resolved in Timms’
chart; cores were collected as close as possible
from the hollows.

Core descriptions

All four cores displayed the same features:
approximately 1 m of black fresh-water organic
muds, above a very clearly defined 8-10 cm
band of extremely fine grained creamy white
aragonite, overlying a graded column of cal-
careous tuff. VD achieved the maximum
penetration through the tuff and is pictured in
Figure 1.

Shells are abundant in the organic muds
and were identified by B. J. Smith as “. . .
assemblages of ostracod shells and the small
freshwater bivalve Pisidium sp. These are
found in fresh waters low in dissolved salts
and usually permanent.”

X-ray analysis of the creamy white band
performed by J. Caldwell of A.N.U., gave a
composition of 95% aragonite + 5% low
magnesium calcite. Under the microscope
much of the material consisted of rods ~1 pm
in length. Diatoms were common and, in
contrast to the underlying tuff, there was a
marked absence of quartz and ferromagnesian
(opaque) minerals,

The graded tuff was rich in carbonates
with some euhedral rhombs (calcite/ dolomite)
but mainly irregular fragments. Volcanic
glass, angular with inclusions of crystallites
and gas bubbles, was common. Quartz grains,
usually fairly well rounded, were present to-
gether with a scattering of microfossils.

(50

160

240°

70

260

(90

270

200

210

Fig. 1. Photocomposite section through Valley
Lake core VD. Distances from top of core
given in cm.

Evidence of horizontal bedding occurred in
the top 10 cm of the tuff column, but other-
wise the material appeared to have been rapidly
deposited under water. The boundary with the
aragonite band was less abrupt than that
between the organic muds and the aragonite,
but nevertheless quite sharp, as can be gauged
from the photograph.

J. R. Dodson examined samples from the
aragonite band and the upper calcareous tuff
for pollen grains. In the aragonite there were
plenty of grains, particularly Casuarina and
Eucalyptus with some aquatic taxa, whereas
the tuff contained very few grains: a scattering
of Casuarina and Eucalyptus but no Com-
positae, Although insufficient material was
examined to provide a definitive conclusion,
these assemblages are consistent with a Holo-
cene age for the aragonite and preclude an

. BARTON & M. W. McELHINNY

age greater than 15 000 BP for the deposition
of the tuff, which must have occurred rapidly
(Dodson pers. comm.).

Magnetic results

Measurements of the horizontal natural
remanent magnetisation (NRM) were made
at 1 cm intervals along the length of each
complete core using an automated version
(Barton 1978)! of the “Digico” whole core
spinner magnetometer (Molyneux et al. 1972).
VD was sliced open, subsamples were extracted
in adjacent pairs of perspex cube shaped pots
(volume 5,3 cm") every 2.5 cm, and measured
on a _ cryogenic “SQUID” magnetometer
(Goree & Fuller 1976).

All cores yielded mutually consistent results,
showing a large contrast in NRM_ intensity
between the organic muds (typically 0.5—1.5
mA.m!) and the calcareous tuffs (typically
100-180 mA.m-) as illustrated in Figure 2.
A well dated magnetic secular variation pat-
tern exists for SW Victoria covering the last
1000 years based on the magnetic remanence
of sets of cores from L. Keilambete, L. Bullen-
merri and L. Gnotuk (Barton!). Unfor-
tunately, the directional results from Valley
Lake (Fig. 2) are too scattered to permit
magnetic dating. Magnetic cleaning in an alter-
nating magnetic field (AF) of peak value 15
mT failed to reduce the scatter. Median de-
magnetising fields (i.e. the peak AF required
to halve the initial remanence) for 8 specimens
distributed throughout core VD ranged from
2 to 6.5 mT, which is too low to sustain a
stable primary remanence. The high water
content in the organic muds (85-90% by
weight) certainly contributes to this instability.

Initial susceptibility measurements on core
VD (Fig. 3) reflect a contrast of about 3
orders of magnitude between the organic muds
and the calcareous tuffs. Much of the organic
mud is weakly diamagnetic, i.e. the suscepti-
bility is negative.

Radiocarbon dating
An initial radiocarbon age of 38400 BP
on the total organic fraction from VD 107—
117 cm prompted a more detailed investigation
into the chronology of these cores. The results
are given in Table 1. Both the calcareous tuff

1 Barton, C. E. (1978) Magnetic studies of some
Australian Lake Sediments. Ph.D. thesis (un-
publ.), Australian National University,
Canberra.

LAKE FLOOR SEDIMENT CORES MT GAMBIER

OF

(cm)

CORE

IN

DEPTH

Ww DECLINATION E

INCLINATION

163

CORE VDNRM

INTENSITY (MA /M)

Fig. 2. NRM results for core VD. Vertical grid interval (bar spacing) is 20 cm; pairs of measurements
at same stratigraphic level linked by lines. Above 20 cm core was 100 wel lo be subsampled; whole
core measurements showed that intensity remained low in this zone.

and the aragonite band yielded barely sufficent
organic carbon to obtain a date, hence the
large counting uncertainties. A carbonate date
was obtained for the aragonite band in VA.

Radiocarbon ages are plotted against the
equivalent distance from the top of VB in
Figure 4, Correlation between YD and VB
for sample ANU 1809 is based on equal
sedimentation rates within the tuffs and may
therefore be in error by up to say + 5 cm;
there can be no uncertainty about the equiva-
lent position in VB of the aragonite band,
sample ANU 2051.

Within the organic muds, the monotonic
MC age sequence is consistent with uniform
deposition since 6000 BP, The fact that the
MC ages within the organic muds plotted in
Figure 4 extrapolate to near the origin is taken
to indicate the absence of any major systemutic
increase in ages due to the incorporation of
ancient carbon (from say, the Miocene lime-
stone basement which outcrops around parts
of the lake). Although further age determina-
tions are really requred to confirm this, it is
worth noting that no systematic age increases
in excess of a few hundred years have been

164

OF CORE VDODSUSC

TOP

E \oo

u

w —t a

c .
Go

o hE
= a"
x

a

oa

ve

a

200

| ! 10

SUSCEPTIBILITY

loo 1000
(MICROGAUSS/ GE }

Fig. 3. Initial susceptibility results for core VD.
Format of plot is as for Fig. 2 except that
iegative (diamagnetic) susceptibilities plotted
us positive with X jnstead of square. Much of
upper 80 cm of organic muds diamagnetic.
Volume susceptibility of 1 Gauss Oersted-! in
ces equivalent to 47 in SL system and
dimensionless.

C. E. BARTON & M. W. McELHINNY

found in sequences from L. Keilambete, L.
Bullenmerri and L. Gnotuk in SW Victoria
(Bowler & Hamada 1971; Barton & Polach
1980). These lake sediments have been inten-
sively dated by radiocarbon and are from simi-
lar geological environments to Valley Lake.

Ages of eruption

The graded calcareous tuff has every appear-
ance of having been rapidly deposited in a
single episode into a lake containing at least
1-2 m water, Excluding the possibility that
this occurred 38 000 years ago, which is incon-
sistent with previous age determinations, the
pollen data, and the morphology of the Mt
Gambier complex, there must be considerable
amounts of ancient organic carbon (charcoal)
incorporated in the tuff to account for such
an age.

Intensive radiocarbon analysis of magneti-
cally correlated cores of organic mud from
Bullenmern, 38°15'S, 143°06'E (Barton!;
Barton & Polach 1980) indicates 4 25% praba-
bility of >20% anomalies in radiocarbon
ages, These muds are not dissimilar from those
in Valley Lake, Although these figures over-
estimate the dating uncertainties in many
lacustrine sequences, e.g. Keilambete, 38°13'S,
142°52’E, (Bowler & Hamada 1971; Barton
& Polach 1980), they should nevertheless be
regarded us a guide in assessing the significance
of isolated "'C. determinations. Hence the 6180
+ 80 BP age at the bottom of the organic
muds is not necessarily inconsistent with an
eruption age of 4800 BP.

Magnetic intensity and susceptibility con-
trasts between the organic muds and the tuffs
provide a sensitive measure of the presence
of yoleunic ejecta within the sequence, At no
point within the organic muds do either of these

Tati lL. Conventional radiocarbon ages for samples from three Valley Lake cores, All determinations,
with the exception af that marked ©, are made on total arganie [ractians.

Equivalent.
ANU Sample Depth in Depth in MWC age
number Core (cm) VB (em) *1SD Comment
ANU 2125 VB 40-50 40-50 2960 —& 90 Organic mud
2126 VB 80-90 73-83 3960 + 80 Organic mud
2052 VB 110-120 103-113 6180 80 Organic mud
2051 VA 117-122 114-122 13900 + 370 Aragonite (organic)
15450 > L600" Arigonite (inorganic!
1809 VD 107-151 fd 151 38400 + 2070) Calcareous tuff

1640)

" total inorganic fraction

LAKE FLOOR SEDIMENT CORES MI GAMBIER 165

REOIOCSABON AGE (Kyi

G \ i 3 + 9 6
rT
of
an ’
e tb
5 KI ’ |

'

PS 7aH5 PLM CLP OINVEND HaLeMHSaH4

fa)
a
>
rh wf view +
us
awit
4) 16 +
- Jumeamy
= mele el | te ing WHITE
Ciypy abled AMEGUNITE
"7 wm | &gnk of
FE Deeetnhiray
wit ong ee
Lae 22 -
wy 2un aKad sinh ‘¢ 40)
E
= ec
+ o
thi
wh >
| ®
i” ’

Fig 4, Conventional radiocarbon age-depth plot
for core VB together with simplified log of core,

Ages expressed in units of 1000 years (Ky).

Vertical error bars denole sample lengths and
horizontal ones, | standard deviation counting
Uncertainties in ages. Sumples ANU 2051 und
ANU 1809, taken from cores VA and VD
respectively, murked al their equivalent dis-
lunces from top of core VB,

parameters even remotely approach the high
values within the tuffs, nor is there any eyi-
dence to the naked cye of a volcanic interlude.
It is therefore concluded with some confidence

that no eruption has occurred near the lake
since the deposition of the calcareous tufts.

The problem remains as to the significance
of the organic and inorganic ages of 14.000
years for the aragonite band, Sufficiently slow
deposition of aragonite could explain the 6000
year lime break at the upper boundary iodi-
cated by the average age of the whole band.
However, since the organic muds indicwle
fresh water conditions throughout the lust
5000 to 6000 years, it is considered improbable
that conditions under which only 95% pure
aragonite was deposited could have existed
within the lake for many thousands of years
previously, The preferred conclusion, and one
which is more consistent with the geologesl
evidence, is that the aragonite was produced
fairly rapidly at the end of the eruptive phase
at 5000 BP or possibly 6000 BP, and that both
ages reflect the presence of dead carbon derived
from the volcanic ejecta or from stirring of
the original lake floor,

Acknowledgments

We thank the Corporation of the City of Mt
Gambier for granting access to the lake and
for providing storage tacilities and information,
Dr B. J, Smith (National Museum of Vic-
ioria) for shell identification, and Dr J. R.
Dodson (University of New South Wales) for
pollen identification,

References

Baviy, | A. BE. & Witiiams, W. D. (1964)
Chemical and biological observations on some
volcanic lakes in S.E. South Australia, dust J.
Mar. Freshw, Res, 15, (23-132,

Barron, C, BE. & Potach, H. A. (1980) 14C ages
and magnetic stratigraphy in 3 Australian
muars. Radiocurbon 22 (in press).

BLAckBuRN, G, (1966) Radiocarbon dates relating
to soil development, coustline changes, and
volcanic ash deposition in south-east South
Australia, Aust. J. Sei, 29, 50-52,

Bow er, J, M, & Hamapa, T, (1971) Late Quater-
nary stratigraphy and radiocarbon chronology
of water level fluctuations in L. Keilambete,
Victoriu. Nature 232, 330-332.

Ferousson, G, J, & Rarrer, T. A. (1957) New
Feuland C age meusurements 3. NZJ. Sci.
Tech., 38B, 732-749.

Gne & OD, (1955) Radiocarbon dates for
Australian archuevlogicul and — geological
samples. Aust. J. Sei. 18, 49-52.

Goren, W. 8. & Futtor, M. (1976) Magneto-
meters using R-F driven SQUIDS and their
applications in rock magnetism und palueo-
magnetiant Rev, Geophys. Space Phys, U4, 591-
OX.

MackeretH, FP. J, H. (1958) A portable core
amet for luke deposits, Lininel. Oreanes. 3,
[81-191.

Movynnux, L., THoMson, R., Orpereco, F. &
McCartan, M. B. (1972) Rapid measurement
of the remanent magnetisalion of lung cores of
sediment. Narre 237, 42-43.

Surarp, M. J, (1978) The geological history of
the Mount Gambier volcanic camplex, soul
east South Australian. Trans. R. Soc. §. Aust
102, 125-139.

Times, B. V, (1974) Morphology and henthos
of three voleanic lakes in the Mt Gambier
district, South Australia, Anst, J. Mar. Freshy.
Rey, 25, 247-297,

AMINO ACID RACEMIZATION DATING OF LATE QUATERNARY
STRANDLINE EVENTS OF THE COASTAL PLAIN SEQUENCE NEAR
ROBE, SOUTHEASTERN SOUTH AUSTRALIA

BY C. C. VON DER BORCH, J. L. BADA & D. L. SCHWEBEL

Summary

The amino acid racemization dating technique has been applied to three selected mollusc samples
collected from the Quaternary strandline sequence of southeastern South Australia. Results of the
study are consistent with previous uranium-series age determinations in the area and imply that at
least the uppermost component of the Woakwine Range barrier-estuarine sequence was emplaced
during the last interglacial sealevel maximum around 125 000 years ago.

AMINO ACID RACEMIZATION DATING OF LATE QUATERNARY
STRANDLINE EVENTS OF THE COASTAL PLAIN SEQUENCE NEAR
ROBE, SOUTHEASTERN SOUTH AUSTRALIA

by C. C. von per Borcn,* J, L, Bava,7 & D. L. ScinwepeLt

Summary
VON pEt Boren, ©. C., BApa, J, 1. & Scuwnnet, D. L. (1980) Amino.acid racemization dating
of Late Quaternary strandline events of the coastal plain sequence near Robe, southeastern
Soulh Australia. Trans. R, Soe. S. Aust. 104(6), 167-170, 28 November, 1980,

The amino acid racemization dating techniqne has been applied to three selected mollusc
samples collected from the Quaternary strandline sequence of southeastern South Ansteulia.
Results of the study ave consistent with previous Yranium-series ave determinations in the area
and imply that at least the uppermost component of the Woakwine Range barrier-estuurine
sequence wag emplaced during the lust interglacial sealevel maxinrum around 125 OOU years

uO.

Introduction

Oxygen isotope studies of deep-sea pelagic
sediments, combined with magnetostratigraphy
and other dating techniques (Shackleton &
Opdyke 1976; Hays et al, 1976), have estab-
lished a relatively detailed chronology of
Quaternary yvlacial and interglacial stages. Re-
lated custatic sealevel oscillations recorded as
stranded shoreline deposits on continental
margins are currently under scrutiny, Although
more difficull to date, they serve as an inde-
pendent check on some of the deep sea data.
In addition, the establishment of an weceplable
chronology for Quaternary and older shoreline
sequences is of foremost interest from a geo-
dynamics point of view. A correctly dated
succession of terraces can reveal the temporal
variation in uplift rate of convergent plate
boundaries such as island ares, ad mid-plyic
tectonie movemerits such as regional warping
on passive margins. It is of interest to establish
acceptable chronologies from coastal strandlinc
sequences from a Variety of tectonic settings
and areas,

The coastal plain of southeastern South
Australia (Fig. 1) is characterized by what
may be one of the most complete and best
preserved gequenees of Quaternary strandiines
in existence. At least 20 emergent shorelines,
consisting of stranded calcarcous sand barriers
and associated lagoonal and lacustrine Ueposits,
occur in a region 9U km wide by about 400
km long (Fig. 1). A sequence of less obvious

* School pf Earth Sciences, Flinders University
of South Australia, Bedford Park, S. Aust. 5042.

+ Seripps fastinion of Oceanography and Lnstl-
tudor of Mating Resourees, Cahfornia,

+ Esso Australia Lud, Sydney,

siliceous sand beach ridges of Plio-Pleistocene
age (not shown in Fig. 1) extends for a further
100 km east of the Naracoorte Range into the
State of Victoria (Hills 1960; Blackburn et al.
1967).

The calcareous strandlines shown in Figure
| owe their preservation to a combination of

guaieaas

Pender begeonel-
Lacuttiine Factor (Fletey

Bench Dune Facies triduest

ath AWery

Uy) Revita Send

ore

Gambrer Linve erane

Volcan(Gs- 05h & babat®
jl bate Comogare vue

Grarilic juchs Early Petaeadois

9 10 20 soni

Fly, 1. Coastal Plain, southeastern South Australia,
showing Quaternary strandlines; section X-Y
refers to Figure 2.

168 Cc. GC yon der BORCH, |. L, BADA & D. L. SCHWEBEL

factors, the dominant of which has been gentle
regional upwarping of the coastal plain
throughout the Quaternary, centred on the
volcanic region in the extreme southeast of the
state (Hossfeld 1950; Sprigg 1952). This up-
warping has been responsible for the stranding
of the sequence, in which oldest shorelines in
yeneral lie furthest inland. Preservation of
these strandline features has been due largely
to rapid “case-hardening™ of the calcareous
barrier facies sands by extensive calcrete
development which generally begins imme-
diately the sands become stabilized by vegeta-
tion. Only high sealevels are represented in the
record, due to a combination of relatively slow
uplift rate and the dynamics of sediment trans-
port as sea Jevel rises from # low stillstand.

In common with other Quaternary shoreline
successions of this type, the establishment of
chronological sequence and absolute age of
individual strandlines is fraught with difficul-
lies, Palacomagnetic studies of cores from
recent stratigraphic drilling suggests that the

oldest component of the complex Naracoorte
Range barrier, shown in Figure 1, is older than
the Bruhnes-Matuyama magnetic reversal at
720 000 years; all ridges to the southwest are
younger (Cook et al. 1977). Limited radio-
carbon dating of the youngest deposits in the
sequence reported by Blackburn (1966), von
der Borch (1976), Cook er al, (1977) and
Schwebel (1979)! has established a preliminary
chronology of Holocene and late Pleistocene
sediments from lagoonal and lacustrine areas
near the present coast. Uranium-series dating
techniques have been applied to aragonitic
lagoonal sediments and molluscs dating back
to the [ust interglacial high sealevel (Schwebel
1979)),

This paper reports an initial application of
the amino-acid racemization (AAR) dating
technique (Masters & Bada 1978) to the prob-
lem of deciphering the chronology of some
aspects of the Woakwine strandline region
shown in Figure 1, It serves as an independent
check on uranium series dates obtained from

—_
x WOAKWINEYVRANGE Y
ROBE
RANGE LAKE
HAWODON
LAKE B
SICLAIR =
=z
10 7
x
o
a oy a
QF Mere ll eer RRE See So S05 s tase cP 232s geet esse ®, Ae mele |e Teele. | #3 oto ed
4
=O
wo
= Fivaipagecs 6
10
20
Beach—-Lune Caleareous Sand
i) Amino Acid Dating Sample
QUATERNARY P| Estuarine-Lagoondal Calcgrequa @ Location
SEQUENCE Muddy Sand
Boundaries of Quaternary
Lacustrine Calcareous, Muds \ Transgressive~Regressive
Cycles
mt

LL

locations of samples A, B and €,

Fig. 2. Section X-Y (Fig. 1), after Schwebel! showing Flinders University stratigraphic borehole und

AMINO ACID DATING OF LATE QUATERNARY STRANDLINES

samples beyond the range of ™C techniques
(Schwebel 1979)",

Saniple localities

Maternal used for AAR dating was obtained
from localities A, B and C (Fig. 2) on the
transect X-Y (Pig, 1), ‘The molluse Katelysta
yedlurina, (Lamarck) waa selected as the
gpecies most suitable for the raccinzation
analysis since it has relatively thick. nen-
porous. vailves,

Sample A is a surface sample from a shallow
pit at Lake St Clair where articulated speci-
mens were selected, This locality lies on the
first interdune flat inland from the present
coast, where dates ou molluscs have been
reported by Blackbur (1966) at 4330 = 100
years using MC. Stratigraphic observations hy
Schwebel (1979) are in accord with the above
date, and show the sediments to have been
deposited during the peak of the Holocene
sealevel transgression prior to final separation
of the Mat from the ocean by modern barrier
accretion and possible sea level decline. Be-
cause of the established radiocarbon dates, this
sample was used us a reference point far the
calculations involved in determining the ages
of samples Band C.

Sample B ig from the side of a dram locally
known as Drain L which interseets the Woak-
wine Range harrier and ils related estuarine-
lagoonal strata to the east, Shells comprising
sample B, many of which are in-situ, occur
withm a 20-30 em thick indurated layer, on
an erosion surface which is onlupped by Holo-
cene Jucustrine calcareous muds of the mast
recent lake Hawdon phase.

Sample C. interpreted by Schwebel (1979)!
to come from the same horizon as that of
sample B, was collected fram a stratigraphic
horchole (Fig, 2) from a molluse-rich layer
280 em below the sediment surface. Articulated
samples of Kavelysia sealarina (Lamarek)
were sampled and used for dating,

Dating methods and procedures
Approximately 5-10 grams of a single
eleaned Katelysia valve were processed accord-
ing to the procedures desenbed for the “total”
fraction by Masters & Bada (1977). The
alloigoleucine/isoleueine (alleu/iso) ratio was
determmed on a Beckutun-Spinco Model 118

__ Sa rrrrovr'"—

7 Schwebel, DA. (1979) Quaternary stratigraphy
of the southenst of Soulh Australia. PhD.
thesis (unpubl). Flinders Univ. of S, Aust.

has

TanLe J. Baten of siting avid pacemeiation ty
iKarelysin shelly fren mranite terrace depestta dry
angrherne otistraliy

vt glutumie Del alleu!
Sample alanine = acid ~—s leucine isa
A 0.29 O17 028 O11
B DA 0.30 O35 028
Cc O73 N37 O46 036
Modern Katelysia 0,13 0.08 O09 0.01

i

automatic amina acid analyzer, The enantio-
meric ratios of ihe other amino acids were
determined by gas chromatography of the
N-Irilluaroacetyl-L-prolyl peptide methyl esters
(Hoopes et al. 1978).

Results and discussion

The racemization results for the various
samples are given in Table 1. The extent of
AAR in sample A ts consistent with a Holo-
cene age for this sample, Substututiag the
measured alleu/iso ratio and an age of 4330
years (Blackburn 1966) in eq. (2) of Mas-
ters & Bada (1977) yields k,, = 2.3 * 10%
yori. The value of K,,, in this equation was
assumed to be ~1 3, This kj, value is in close
agreement with that calculated using Holocene
Chiene molluscs from Southern California
coastal archacological sites, This is the expected
resull due to the similarity of the mean annual
cir temperatures of the Califarman and South
Australian localities (Felton 1965: Floegel
1972). and since the Katelysia and Chione
species have similar shell morphologies.

The extent of AAR i sample © is nearly
idendeal to that measured in Chinne (Masters
& Bada 1977) and Pretathaca molluses (Weh-
miller 1977) from a terrace deposit in San
Dicyo, California. This terrace has been dated
al 120000 = |G000 years by vranium-serics
uating of corals (Ku & Kern 1974). Since the
Holocene saniple suggests that the rate of
racemization is similar at the Californian and
Australian sites, the sinvlariny of the extent
of racemization in sample C and the 120 :000-
year-old Californian terrace supports the con-
clusion that sample C corresponds in age to the
maximum high-sea level stand dunng the last
interglacial period (ic. Stage 5e in the 04/01
palacotemperature curve, ~125 000 B,P,).

In comparison ta sample C, the extent of
AAR is consistently slightly less in sample B.
On this basis, fi would appear thal sample B
may come from one of the other episades of

170 C. C. von der BORCH, J. L. BADA & D. L. SCHWEBEL

high sea level, tentatively dated at ~85 000
and 105000 years B.P., which occurred in
the vicinity of the last interglacial period
(Bloom et al. 1974). Substituting the measured
alleu/iso ratio for sample C and an age of
120 000 years for this sample into eq. (2) in
Masters & Bada (1977), yields Kis. = 2.9 X
10-"yr-". Using this ki,» value to date sample
B yields an age of ~92 000 years.

The AAR dates for samples B and C given
above imply that the uppermost portion of the
Woakwine Range strandline complex was
formed during the last interglacial high sea
level. This is in accord with the uranium series
data of Schwebel (1979).

Sample B with an age of 92000 years
appears slightly younger than C, which is about
120 000 years old. In fact, the age of B lies
approximately midway between the 85 000
and 105 000-year-old sealevel highs described
by Bloom ef al. (1974). If the assumption is
made that C actually correlates with the

125 000 year sealevel high (i.e. that the date
used to calculate the k;,,, value given above is
a few thousand years too young), then B could
possibly be correlated with the established
105 000-year-old sealevel high. On the other
hand the small number of samples, and the
resolution of the amino acid dating technique
as applied to the study area, may imply only
that the two samples B and C were laid down
in response to some stage or stages of the last
interglacial sealevel maxima, of the order of
120 000 years ago.

Acknowledgments

We thank D. Darling and E. Hoopes for
assistance with the racemization analyses. The
racemization work was partly supported by a
grant from the U.S. National Science Founda-
tion (Grant EAR 77-14490). Manuscript
typing was done by Nena Bierbaum and
diagrams were drafted by Gail Jackson.

References

BLACKBURN, G. (1966) Radiocarbon dates relating
to soil development, coast-line changes and
volcanic ash deposits in south-east South
Australia. Aust. J. Sci. 29, 50-52.

——. Bonn, R. D. & Crarke, A. R. P. (1967)
Soil development in relation to stranded beach
ridges of County Lowan, Victoria. CSIRO Soil
Pub. No. 24.

BLoom, A. L., BROECKER, W. S., CHAPPEL, J.,
MatTHews, R. S. & MESOLELLA, K. J. (1974)
Quaternary sealevel fluctuations on a_ tectonic
coast: new Th*80/U284 dates from Huon Penin-
sula, New Guinea. Quaternary Res. 4, 185-205.

Cook, P. J., Cotwetr, J. B., Firman, J. B.,
LinpsAy, J. M., SCHWEBEL, D. A. & VON DER
Borcu, C. C. (1977) The late Cainozoic sequence
of southeast South Australia and Pleistocene sea-
level changes. Bur. Miner. Resour. J. Aust. Geol.
Geophys... 2, 81-88.

Fetton, E. L. (1965) “California’s many
climates”. (Pacific Books: Palo Alto, Cali-
fornia).

Fiorcer, H. (1972) The position of the lower
Tertiary artesian aquifer within the hydrology
and hydrogeochemistry of the Gambier Embay-
ment area. Ph.D. thesis (unpubl.), Facultat fur
Allgemeine Wissenschaften der Technischen
Universitat, Miinchen.

Hays, J. D., Imprie, J. & SHACKLETON, N. J.
(1976) Variations in the earth’s orbit: Pace-
maker of the ice ages. Science 194, 1121-1132.

Hitts, E. S. (1960) “The physiography of
Victoria: An introduction to geomorphology”.
4th Ed. (Whitcombe and Tombs: Melbourne).

Hoopes, E. A., PeLTzer, E. T. & Bana, J. L.
(1978) Determination of amino acid enantio-
meric ratios by gas-liquid chromotography of
the N-trifluoroacetyl-L-prolyl-peptide methyl
esters. J. Chromat. Sci. 16, 556-560.

HossFetp, P. §, (1950) The late Cainozoic
history of the southeast of South Australia.
Trans. R. Soc, S. Aust. 73, 232-279.

Ku, T. L. & Kern, J. P. (1974) Uranium-series
age of the upper Pleistocene Nestor Terrace,
San Diego, California. Geol. Soc. Amer. Bull.
85, 1713-1716.

Masrers, P. M. & Baba, J. L. (1977) Racemiza-
tion of isoleucine in fossil molluscs from Indian
middens and interglacial terraces in southern
ete ae Earth and Planet. Sci. Lett. 37,
73-183,

—— (1978) Amino acid racemization dating of
bone and shell. Jn G. F. Carter (Ed.) “Archaeo-
logical Chemistry”. Advances in Chemistry
Series, No, 171, pp. 117-138 (American
Chemical Society).

SHACKELTON, N. J. & Oppyke, N. D. (1976)
Oxygen isotope and paleomagnetic stratigraphy
of Pacific core V28-239 late Pliocene to latest
Pleistocene. Geol. Soc. Amer. Mem. 145, 449-
464,

Spricc, R. C. (1952) The geology of the south-
east province, South Australia, with special
reference to Quaternary coast-line migrations
and modern beach developments. Bull. Geol.
Surv. S. Aust. 29,

VON DER Borcu, C. C. (1976) Stratigraphy and
formation of Holocene dolomitic carbonate
deposits of the Coorong area, South Australia.
J. sedim. Petrol. 46, 952-966.

WEHMILLER, J. F. (1977) Amino acid studies of
the Del Mar, California, midden site: apparent
rate constants, ground temperature models and
chronological implications. Earth Planet. Sci.
Lett. 37, 184-196,

NEW CRETACEOUS AND TERTIARY CRABS (CRUSTACEA:
BRACHYURA) FROM AUSTRALIA AND NEW ZEALAND

BY §. BARKER

Summary

From the Cretaceous of Australia and New Zealand three new genera of Brachyura, one new
subgenus, and seven new species are described and one new family is proposed. Four new species
from the Eocene are described, one from Australia and three from New Zealand, and changes in
taxonomy and nomenclature are made. The new classification of the Brachyura (Guinot 1977) is
applicable to the new material which contributes significantly to the clarification of taxonomic and
phylogenetic relations at an early, critical stage in the evolution of the Brachyura. Modified Tethyan
palaeobiogeographic relations of the Cretaceous and Palaeogene faunas are recognised.

NEW CRETACEOUS AND TERTIARY CRABS (CRUSTACEA: BRACHYURA)
FROM AUSTRALIA AND NEW ZEALAND

by M, F. GLAgSSNER™

Summary

GLAgssNER, M, F, (1980) New Cretaceous and Tertlary crabs (Crustacea, Brachyura) from
Australia and New Zealand, Trins. 8. Soc. 8. Aust. 104(6), 171-192, 24 November, 1980.

From (he Cretaceous of Australia und New Zealand three new genera of Brachyura, one
few subgenus, und seven new species are described and one new family ts proposed. Four new
species from the Bocene are described, one from Australia and three fram New Zealand, and
changes in taxonomy and nomenclature are made, The new classification of the Brachyura
(Guinet 1977) is applicable to the new material which contributes significantly to the clan
ficalion of taxonomic und phylogenetic relations at an early, crilical stage an the evolution of
the Brachyura, Modified Tethyan palacabiogeographic relations of the Cretaceous und Paluea-

gene faunas ure recognised.

Classifivation

A new classification of the Decapoda
Brachyura praposed by Guinot (1977, 1978)
is of particular interest to students of the
evolution of these ¢éruistaceans, Based on
generally sound and stated principles, and on
a re-examination of a very large amount of
zoological material as well as literature, it
takes into consideration conclusions Treached
hy palacontologists, questioning some of these
conclusions specifically. New material from
Australia and New. Zealand provides a suitable
starting point for the task of answering some
of the queries raised, and of testing the
suitability of the proposed new system of the
Decapoda Brachyura, The following tabulation
places the ncw finds in the framework of
Guinot’s classtfication and indicates their age
and oceurrence.

Section Podotremata Guinot, 1977

Subsection Dromiacea de Haan, 1533
Superfamily Homolodromioidea Alcock, 1899
Family Prosopidac von Meyer, 1860
Oanoton woodsi nov, gen,, nov. sp. Upper

Albian, central Queensland and South
Australia.

Subsection Archaeobrachyura Guinot, 1977

Superfamily Homoloidea White, 1847

Family Homolidae White, 1847

Heamelapsis etheridee? (H. Woodward, 1892).
Upper Albian, central Queensland.

Homolopsis spinulosa nov. sp. Upper Ceno-
mianian, northern Australia.

= Department of Geology, University of Adelaide,
Box 498 G.P.O,, Adelaide, 5. Aust, 5001.

Superfarmly Raninoidea de Haan, 1833

Family Raninidae de Haan, 1833

Notepocorystes (Cretacoranina) exiguus Twv,
sp, Lower Cenomanian, Northern Australla,

Hemioon nevezelandicum nov. sp. Upper
Albian, New Zealand.

Ranilia porerariensis nov. sp. Upper Bocene,
New Zealand.

Lyreidus waitakiensis nov, sp. Middle to Upper
Eocene, New Zealand.

Superfamily Tymoloidea Alcock, 1896,

Family Torynommidae nov, fam,

Torynomma (Torynomma) flemingi nov. sp.
Upper Senonian, New Zealand,

Torynomma (Paratarynamma) dentatyim nav,
subgen., nov. sp. Upper Cenomanian—
Lower Turonian, northern Australia.

Diorationus salebroxsus Woods, 1953 Upper
Albian, Queensland and South Australra.

Dioratiopus sp. Upper Cenomanian, northern
Australia,

Eodorippe spedeni nov, gen., Noy. sp. Upper
Senonian, New Zealand.

Section Heterotremata Guinot, 1977,

Superfamily Portunoidae Rafinesque, 1815

Family Portunidae Dana, 1852.

Subfamily Psammocarcininae Beurlen, 1930.

Rhachiovoma granalifera (Glaessner, 1960)
Upper Eocene, New Zealand.

Family uncertain.

Pororaria eocenica noy. gen, nov. sp. Upper
Eocene, New Zealand.

Superfamily Xanthoidea McLeay. 1838,

Family Panopeidae Ortmann, 1893.

Panopeus whittenensts nav. sp, Upper Eocene,
South Australia.

The most distinctive innovation alfecting ihe
classification of the Brachyura discussed here

172

is the demotion of the “Section Dromiacea”
which has dominated considerations on the
origin and evolution of the Mesozoic Brach-
yura for more than a century. Guinot recog-
nises three sections: the Podotremata, Hetero-
tremata and Thoracotremata, named accord-
ing to the position of the gonopores, This looks
rather like a single-character classification to
which I objected (Glaessner 1969) when the
peditreme-sternitreme distinction (Bouvier
1897) was used by Gordon (1963) to remove
the former group from the Brachyura. How-
ever, Guinot (1978) has amply demonstrated
that it is a distinction by grades, the use of
which she finds inevitable for taxa of high
rank while following to some extent “pré-
occupations d’ordre cladistique” for lower-rank
taxa. This taxonomic innovation involves
recognition of two subsections of the Podotre-
mata, the Dromiacea and the Archaeobrach-
yura. The former comprise the Superfamilies
Homolodromioidea and Dromioidea, the latter
the Homoloidea, Raninoidea and Tymoloidea.
This classification is significant for the present
investigation. There is ample morphological
evidence for close links between the Tymo-
loidea and the Homoloidea, The Dromioidea
(Families Dromiidae and Dynomenidae) have
taken a different evolutionary path. With
reference to the descriptions and discussions
which follow, it is sufficient to note here that
Homolopsis and Torynomma (with related
genera) are morphologically close and, as
Wright & Collins (1972) have indicated,
appear to have had Prosopidae, hence Homo-
lodromioidea, as ancestors in Jurassic time.
The Dromioidea differ significantly from this
group in many morphological, embryological
and ethological characters. For the question
of the evolution and systematic position of the
Raninoidea there is no significant new material
under discussion here (see Forster 1968,
Stevcié 1973). I had previously (Glaessner
1969) assigned the Tymolinae to the Dorip-
pidae, following the latest comprehensive clas-
sification available at that time (Balss 1957).
This is now unnecessary and unacceptable,
since Guinot’s work has shown that the oxy-
stomatous condition (which has to do with the
direction of the inhalent and exhalent currents
of water under the carapace) was reached
independently by very different groups of
crabs. Thus there is no justification for
retaining the artificial taxon Oxystomata.
With this demonstration most of the major
classification of the Brachyura, of long stand-

M. F. GLAESSNER

ing but often questioned, had to be abandoned.
The origin of all or some of the older Hetero-
tremata (Dorippoidea, Calappoidea, Cory-
stoidea (+Cancriformia), Portunoidea and
(questionably) Xanthoidea (see Wright &
Collins 1972) from Cretaceous Tymoloidea or
their ancestors is possible or even probable,
but these investigations would lead beyond the
limits set by the material here described.

Summary of stratigraphic distribution
(New Zealand species marked with asterisk)
Upper Albian: Oonoton woodsi, Homolopsis
etheridgei, *Hemioon novozelandicum, Diora-
tiopus salebrosus,

Lower Cenomanian: Notopocorystes (Creta-
coranina) exiguus,

Upper Cenomanian: Homolopsis spinulosa, D.
sp.

Upper Cenomanian to Lower Turonian: Tory-
nomma (Paratorynomma) dentatum.
Campanian-Maastrichtian: *Torynomma (T.)
flemingi, *Eodorippe spedeni.

Middle to Upper Eocene: *“Lyreidus waita-

kiensis.
Upper Eocene: *Ranilia pororariensis, *Rha-
chiosoma granulifera, *Pororaria eocenia,

Panopeus whittenensis.

Remarks on palaeobiogeography
While we know only a small sample of the
Brachyura of the Cretaceous and Eocene of
Australia and New Zealand, theoretical con-
clusions are unwarranted. It is worth noting
that known relations are dominantly with

European genera. Homolopsis, Notopoco-
rystes (which is almost cosmopolitan),
Hemioon, Dioratiopus, Rhachiosoma and

Panopeus (which is also Atlantic) are well
known from Europe. As far as the Austra-
lasian region is concerned, the origin of these
genera can be considered as Tethyan. Tory-
nomma has its range extended from Queens-
land to the north of Australia in the
Cenomanian, and to New Zealand in the
Campanian-Maastrichtian. Its close relative
Dioratiopus, a genus shown to include many
European species, is recorded, in addition to
Queensland, from northern South Australia
and from Melville Island north of Darwin.
The undescribed macruran and thalassinacean
decapods from the Cenomanian of the island
(Paraclytia, Hoploparia, Trachysoma and
Protocallianassa) are well known from the
European Upper Cretaceous. The Tethyan

NEW CRETACEOUS AND TERTIARY CR VBS

relations of the Brachyura from Bathurst
Ishind are in agreement with Lhe character of
its Cenemanian ammonue fauga (Weight
1963), Eodorippe trom the Upper Senenian of
New Zealand is endemic, and the endemic
Recent. Nectocarcinuy may have had an ances-
tor dating back to the Eacene in the same
region, the new genus Porotarta.

Descriptions
Family Prosopidae
Oonotim nov, gen,
Derivation of name: From Greek oon:
eee, noton; back, with reference to shape.

Diagnosis: Carapace ovoid, without sharp
lateral margins, posterior margin short, ros-
trum triangular, truneated, with transverse
groove and row of graniiles at bases eye sockets
close-set with strong supra- and infraorbital
spines; mesogustrie lohe long and narrew.
uragastric lobe indistinct, carapace surface
granulated,

Thad intended to assign this fossil to Fectls
Withers, 1946, but Mr C. W. Wright kindly
informed me in July 1980 of his disagreement
with such an extension af this taxon, | accept
his view that the differenees between the
Australian ¢fab and the three English species
ure of generic significance, They leave Gono-
ton closer to Veerlsy than to other genera,

Oonoto woodsi nov, sp.

Derivation ef names After Dr J, T, Woods,
Director of Mines, Queensland who has made
Valuable contributions to the knowledge of
Cretaceous Decapoda from Queensland,

Material: 1. Holotype—An almost complete
carapace. Qld Mus, K 2876. 2. One ffag-
mentary carapace, Geological Survey of
South Australia No, Cr 1.

Localities: 1. Currane Station, 16 ki N of
Dartmouth, central Queensland. 2. 14 kin W
of Mt Dutton (Loe, 17, Miap Sheet 5/571,
see Bull 40, Geol, Survey S. Aust. p. dd).

Preservation; The Queensland specimen is
well preserved in a smooth, round, concre-
tionary nodule, similar to those which contain
other crabs from the Queensland Cretceous,
The South Australian specimen consists of
internal moulds of two angular fragments of
the posterior portion of the carapace, partly
overlapping, in a concretion containing
numerous mollusea, The mode of preservation
and the assactalion suggest that the carapace
may have been broken hy a predator, probably
a belemiite or an anymenite

173

Ages Upper Albian, Tambo Formarion of
Queensland and Oodnpadatta Formation ot
South Australia,

Descripiun: Curapace of holotype avoid,
27 mm long, 224 mm wide, about 8.5 mm
high Convex antero- aid posterolaceral mar-
gins of ahoul equal length; lateral margins
almost parallel, longitndinal and transverse
profiles of carapace (Pigs, 1A, B) strongly
and smoothly convex. Apically truncated
triangular rosttul plate directed forward, set
off from (interior slope of curapave by trans-
Verse groove afd row of granules in front of
mesovastric region. Eye sockets deep and
small, with strong, conical, supraorbital and
long infraorbital spines Which are twice as
long and set helow. Anteralateral outlines
diverge 10 crossing points of cervical grooves
and widen only slightly to metubranchial
regions. Posterior margin apparently not as
long as in Vectly, Regions well marked by
smooth, shallow furrows. Mesogastric lobe
not subdivided, Almost entire surface of cara-
pave fairly evenly covered with large granules
Additional small tubercles on metabranchial
regions, and 5 distinctive, lurger, granulate
elevations. Two are symmetrically arranged
on mesogastric lobe, 2 of metragastric lobe
which is divided by deep median furrow, |
on cardiac lobe, sy distance from posterior
margin 4 that from hase of rostrum, Cardiac
erades into narrow intestinal lobe,

This species differs from two Aptian and
nne Albian species of Vectiv previously des-
cribed from Envland in details of shape and
surface sculpture, Tt has all the significant
characters of their carapace regions but they
ure uot as tum and therefore less con-
spicuous in the new species, The characters
which are preserved in the smaller specimen
from South Australia do not differ from those
af the holotype from Queensland,

Family Homolidae
MHomalopyis Bell, 1863
Homolopsis etheriduct (Woodward. 1892)
FIG. 2
Presapon elleridee’ Woodward 1892, p, 301, pl
4) Erhevidge 1917. po 5, pl 1, fix, 1-4,
Harmolopyiy etheridgels Vian Sraelen, 1928, p, 6195
Woods, 1953, p, 30, pl. 2, fig, 1-3, text fig 1.
Material: One almost complete specimen
(P22945),
Lacality: “Currane’, 14 km WN of Dart-

mouth. central Queensland, Coll, J, T, Woods
1955.

SSNER

E

<
I
QO

, Bo a

N

NEW CRETACEOUS AND TERTIARY CRABS \72

Aye: Upper Alan, Tambo Formation.

Remarks: This specimen is figured here for
comparison for completeness of the record of
currently known Australian Cretaceous crabs,
ft hus a significantly more convex carapace
than the species figured by Wright & Collins
(1972) and its Upper surface ws certainly not
“mare or less flat” as their generic diagnosis of
Haomolopyis requires, Tt may be appropriate
to place the Australian species in a new sub-
genus unless species with traditional shape
Characters exist.

Homplopsis spinulosa nov. sp.
FIG. 3, 3A

Holotype material: Once small, somewhat
crushed specimen with counterpart, P22934.
Coll. B. Daily 1954

Locality: South coast of Melville Esland,
N-f., about 10 km NW of Cape Gambier.

Ave: Upper Cenomanian, Bathurst Island
Formation, about 8 m above “Tapara Bed"

Preservation: Preserved in a slightly
weathered yellowish clay is the cepbalothorax
with two abdominal somites, the left cheliped
and parts of all other perciopods. Ferruginised
and affected by compaction and weathering;
most of the carapace margins defective. Arti-
ficial moulds of the counterpart show details
fot recognisable in the damaged specimen,

Description; Carapace about }2 mm tong
and td om wide, fronto-ocbital width about
8 mm} anterolateral margins diverging, Pos-
(erolateral margins slightly converging;
posterior margin about 5 mm long, curved,
with granulated edge, Ornamentation consists
of distinct, granulated tubercles; at least |
epigastric, 2 protogastric, | hepatic, 2 ept-
branchial and 3 metabranchial tubercles
present on each side and also metagastric and
cardiae tubercle-bearing ridves. Surface finely
and evenly eranulated. Left cheliped short and
robust, with a spindse carpus; P2 and F3
strong. long, with spinose edges, the P4 appear
near the posterolateral angles of the carapace,
thin, smooth P5 near the hase of the abdomen,
First two abdominal somins about 5 num wie,
rectangular in outline, subequal, about 0,75

ne Eee E EEE SenS Sn

grooves and cleyations on a blunt transverse
ridge on gach somite, Abdomen extending
horizontally backward.

This species appears to differ from other
species of Momolopsiy and also from “Glaess-
nerie” depressa (Carter) as described by
Wright & Collins ((972) mainly in details of
ornamentation, The linea homolica ts partly
visible and partly obscured by conipression and
fracturing of the only known carapace,

Family Raninidae
Notopacorystes NVCoy, 1849
Natopocorystes (Cretacoranina) exiguus
nov, sp.

FIG, 4, 4A

Derivation of name: From Latin exiguus;
small,

Material: One carapace, with counterpart,
in w bore core. Forwarded by Mr PF. Bollen,
P22929,

Locality’ Bathurst Island Ojl| Development
Well No. 2, about 4 km N of the mouth of

Pipanyamilr Creek, about 30 km W_ of
Bathurst Island Mission, Care from 280 m
depth,

Age: Lower Cenomanian, Bathurst Island
Formation, grey clay.

Preservation: Carapace undistarted but most
of the extremely fragile shell attached to the
counterpart and anterolateral and orbital mar-
gins damaged.

Description: Carapace ovoid, 15 mm long,
10 mm wide. Width of the fronto-orbital
margin about 5.4 mm, Carapace gently convex
transversely and longitudinally, greatest height
in the centre. Median ridge very faint. Rostrum
hifid, medially grooved, slightly detlexed. Two
supruorhital fissures. Extraorbital teeth could
riot have been large. Orly a very weak lateral
tubercles, 1 hepatic, | cpibranchial and 1
extremely weak mesobranchial, followed by a
finely granulated ridge along posterolateral
margin. Posterior margin about 6 mm long
but not well preserved. Cervical furrow, gently
curved, runs from a notch in front of the epi-
branchial luberele in a forwardly convex curve,
then follows @ sinuous course to distinct pos-
terior gastric pits. Epibranchial turrows short,

mm Jong. with two longitudingl shallow

Pig. |. Qanoren waedst nev. sp. x2.

Fie, 2. Momolopsiy ethertitge’ (A, Woodward).

Fig. 3. Mamolapsis sninulasus nov. sp, x3,

fie. 4. Netopocorysies (Cretacoranina) extyvuy nay. sp. x3.

Fig. 4. Memioen nevoselandicun nov. sp, x3.

Pip. 4. 6a. Ranilia porerariensis nay, sp. x2, a—dorsal view, b—ventt wl view

176

10mm

10 mm

M. F. GLAESSNER

10mm

10 mm

NEW CRETACEOUS AND TERFIARY CRABS 7

iransverse, do not reach the cervical groove.
Branchiocardiac furrows more distinct, particu-
larly their posterior edges, merging with the
deep epimeral grooves. Epimeral and posterior
gastric. muselé attachments well marked on
ihe inside of the carapace, The described Cur-
rows, together with hepatic and Weaker
Mesogastric furrows form # peculiar radiating
pattern on the outer surface which Is finely
pitted and granulated, Two elongate, tubercu-
late protuberances on the anterolateral parts
of the carapace: outer ones smaller .and
prominent on the fepalic lobes, inner ones
situated on outer portions of the protogastric
lobes Which are partly separated from the main
paris by shallow grooves,

Remarks: The weak marginal spines and the
surface sculpture plave this species in the sub-
genus Cretacoranifa Mertin, 1941, It does not
appear to he particularly close to any of its
known species but detailed coniparison must
await the discovery of further specimens.

Hemioon Bell, 1863
Hemloon novezelandioum mov, sp.
FIG. 5, SA
Holotype material. Gne specimen in a hurd

conerction. Geologival Survey of New Zeuland
GSI 1845,

Loealiry: Swale Stream, Coverham, South
Island of New Zealand (P30/f 36}.

Age: Latest Albian, Lower part of Swale
Siltstone, Ngaterian locul Stage, Dr |, Speden
(in fitt, 1977) suggests lower Ngateriau, from
near the base of the Worthuceray parvum
Assemblage Zone of Henderson (1973).

Preservation; Cephalothorax preserved as an
internal mould, eff central position, in a Very
hard concretion which does not split evenly.
Sternum not visible hut fragments of Jegs und
chelae preserved im slightly displaced positions
and much of the inside of the carapace shell is
visible. Although the rostrum remained th
the Counterpart matrix when the coneretion
was split, it was possible to extruct its tip and
rejoin it so that the length of the rostrum could
he determined (Fig, SA).

Peseription: Cephalothorax ovoid, flattened
jonwitudivally and convex, transversely, lateral
margin discontiniious, weakly developed, ‘Two
weak anterolateral spines aml un acicular
extraorbital sping with yranulated surface.
Orbits large, well marked, with two supra-
orbital fissures and a stout supraorbital tooth.
Rostrum about 3 mm long, straight, with two
jateral basal spines. Markings on carapace sur-
face very weak, Internal mould shows median
parts of cervieal groove curving laterally from
posterior gastric pits, the anterior tip of the
Mesagasinc lobe, epimeral grooves, and relics
of the ‘branchiocardiac grooves, Narrow
median ridge marking cardiac und mlestinal
lobes, Strong muscle pits in the hepatie regions.
Surface of carapace finely pitted where it can
be seen, Posterolateral margins granulate, The
greatest width (16.5 mm) is found hetween
postermlateral (marging Which are separated
trom the anterolateral margins by a slight
constriction behind the second anterolateral
teeth, Marked, nirrow, pterygostomial ridge
and a wide depression alone brachiostegite.
Posterior margin missing. Carapace was al least
25 mim long, greatest width is ac about 0.6 of
its length.

Remarks: The new species differs from A,
clangatvin (A. Milne Edwards 1862) in the
shape of the carapace, the weak lateral tecth
und the absence of surface tubercles, A, etr-
cvmniatay Wright and Collins 1972 also has
strung unterolateral spines and convergent
lateral margins on the posterior half of the
carapace. These differences remove the new
spevics further from Raninella ws revised by
Wright & Collins than those eansidered by
these authors {t differs more in shape and
carupace sculpture from Netopacarystes (Cre-
facoranina),

Raailia H, Milne Edwards, 1837
Ranilia pororariensis noy. sp,
PUG, 6, 6a
Helorype eaterianl One carapace. slightly
Uistortect by flattening, most of Fronto-orbital
margin missing, Chelipeds and some displaced
oy broken pereciopods preserved, sternum not

Fin 1A. Danoten woadst nov, sp.. left side view: IB front view. SO—position of supreurbital spine,
To-infraorbital spine. Stippled parts of campate are missing or concealed.
Fig. 3A. Hemvlopsis spinulosus Nov, so. Partial reconstruction Dotted line indicates outline of erushed

caramice,

Vig. 4A. Netopocoryaiey (Creiacorunina) exigins poy, sp. Reconstruction of carapace showing pattern

us scen on inner surliee.

Fig. SA. Henioon novezelancdiedia ney. sp. Reconstruction of carapace. Stippled paris are missing
Fig. (2A, Diorwiopus solebrasis Woods, Reconstruction of carapace.
Fig, I3A. Epdorippe speden? oov, sp, Diagrammatic recanstriction of carapace pattern and rostrum.

178

visible, first abdorainul somite preserved, Can-
terbury Museum, Christchurch, New Zealand,
No.zfe 7, Coll, RK. S, Allen, Feb. 1935.

Localirys Coastal cliffs at Pororari, 2.4 km
N of Punakaikt. Sheet No, $37/723, Grid ref.
C852318.

Age; Kaiatan—Runangap, Late Eocene.

Description: Carapace ovoid, strongly con-
vex transversely; height about equal to half
the width; gently convex longitudinally
Distance between the extraorbital and lateral
acicular teeth equals that between Jateral teeth
and point of greatest width which is at the
level of the posterogastric pits; distance from
this point to posterior margin almost 3
times longer, Posteriorly convergent postcro-
lateral margins marked by a smooth ridge
which is convex in dorsal view for first half of
their length, then straight, Posterior margin
straight, A very faint median ridge on posterior
half of carapace, Two supraorbital fissures
separated by ~ small tooth, only base preserved,
Surface of carapace uniformly pitted, marked
only by weak epimeral muscle impressions.
First abdominal segment trapezoidal, surface
pitted but otherwise smooth. Propodus of
cheliped has sharp, denticulated upper edge
and strongly deflexed fixed finger, Dactylus
gently curved, with narrow dorsal groove
hetween two ridges,

Remarks; The new species differs m shape
and ornamentation and in the deflexed fixed
finger from living species, [t is distinguished
by the weak longitudinal ridge and the antero-
lateral teeth from the Eocene R, (“\Noatopella”)
vareolata (Lorenthey), Further specimens in a
better state of preservation and preparation are
required for complete description and com-
parison,

Lyreidus de Haan 184
Lyreidus waitakiensis nov. sp.
FIG. 7, 7a

Halotype material? One carapace, front
mostly missing, Canterbury Museum, Clirist-

ehureh, New Zealand, No. zfe 30,
Loculity: Black Point, Waitaki Valley.
§127/368, Grid reference c 364 916. Another
less well preserved specimen (Canterbury

M. FF. GLAPSSNBR

Museum, No, 2fe 8, coll. R. S. Allen Feb,
1935, frotn coastal cliffs at Pororari, 2.4 km
N of Punakaiki) probably belongs to this
species.

Age; Middle to Upper Eocene, Tapui glau-
conitic sandstone, Bortonian Stage. This may
he the oldest kiownh species ef Lyreidus. The
Other specimen, tentatively mamed L. ef-
waltukiensis, is from the Upper Evcene,
Katatan-Runangin.

Preservation; Inner layer and parts of outer
ol shell of holotype present. Complete
branchiostegites, part of the left antennal base
and smull, displaced sternal fragments also
preserved,

Description: Carapave ovate; Pronto-orbital
region 6,5 mm wide, with 2 supraorbital fis-
sures; anterolateral margins diverging from the
extfaorbilal ta obliquely pointed lateral tecth.
Distance between their tips is about 3,5 nim,
equal to greatest width of carapace measured
about 3 mm behind them, Abterolateral mar-
Bins tounded, uv bluntly conical tooth on each
side halfway between extraorhital ynd laceral
teeth, A blunt edge extends a short distance
behind the lateral teeth, replaced Irom below
by a distinct, sharp, posterolateral ridge with
amall granules. Posterolateral margins con-
verge 10 afelale posteriie margin which
equals fronta-orbilal margin in length, Cara.
pace strongly convex transversely, gently con-
vex Jonvitudinally, pterygostomial regions
inflated. Pleural suture sinuous anteriorly.
parallel fo lateral margits and a shore distance
below then, Surfice of carapace shows pose
terogastric pits 2 mm behind tevel of the
lateral teeth, und weak epimeral stiractor
muscle markings sume distanee behind them:
smooth hut may have beet faintly pitted and
weakly granulated jf) some places, Two weak
epigastric yuhercles. Pterygostomial regions
aranwilated. Orbits only slightly oblique in
frontal, view. witht small infraorbital spines
separaled by fissures from extraohital weth
Anteunal base has taleral, longitudinal, eranu-
lated mupe with channel along its cuter side,
helween it and edge of carapace; jt may lead
tn the proximal side of the infraorbital tooth.

Remarks: This species can be distinyulshed
by its avoid shape and its ornamentation, ‘Che

ee eeSFSFSSSSSSSSSSSSSSSSSSSS——eaieseseseseFh

Fig. 7, 7a, Lweidnus woinekiensis nov. sp. x2. a—dursul view, b—ventral View,
Fig. & Toryronima guadratum Woods. Plaster cast of holotype, x2. Rostrum drawn from anuther spe-

cimer.

Figs. 10, 11, Tarvnammea (Paroterynemmad) dentetum ney sp

')—holoryye. “15, dersal view. Tp

Paratype P2231. x2. Ventral view with atener stermum and abdumen of 9 sind perciopaids.

NEW CRETACEOUS AND TERTIARY CRABS 179

180

shape of the carapace appears to be of slightly
more generalised raninid type than in the
Miocene and living species which are
anteriorly more sharply narrowed and strongly
elongated,

Family Torynommidae nov. fam,

The genus Torynomma was originally placed
in the family “Prosoponidae” (recte Prosopi-
dae), together with Dioratiopus (Woods 1953,
p. 52). These genera have hardly more than a
few primitive (plesiomorphic) characters in
common with the Prosopidae but they do not
have any of the distinctive, diverse develop-
ments of shape of the carapace and rostrum
or the dominance of the transverse carapace
grooves which characterise this family. The
placing of Torynomma, Dioratiopus and other
extinct genera in the subfamily Tymolinae
Alcock, 1896, as proposed by Glaessner
(1969) is considered inappropriate by Wright
& Collins (1972). Concerning the placing of
the Tymolinae in the Dorippidae, Gordon
(1963, p.56) stated: “Certainly the so-called
Tymolinae with sternal furrows and coxal
genital pores should not be placed in the same
family as the dorippids without sternal furrows
and with the genital openings of the female
sternal”. I rejected (Glaessner 1969, p.R440)
Gordon’s further conclusion that it seems
logical to exclude all peditreme crabs “from
the Brachyura, restricting the term to the vast
majority of crabs with the female genital
openings sternal’. I noted that the Dromiacea,
Raninidae and Tymolinea which have coxal
female gonopores are “exceptional” and that
their separation “on the basis of an obviously
primitive character is an extreme application
of ‘horizontal classification’ which is not
acceptable, particularly as the steps in the
evolution from the primitive peditreme to the
advanced sternitreme condition have not yet
been studied on fossil material”. Hence I fol-
lowed the earlier systematists and Balss (1957)
who placed the Tymolinae in the family
Dorippidae. Guinot’s work (1977, 1978) has
now removed the traditional major subdivi-
sions of the Brachyura such as the Oxystomata,
which had been hampering the development
of systematics in this group since the middle
of the last century. It showed that Gordon
had been remarkably far-sighted in recognising
the peditreme-sternitreme evolutionary transi-
tion as a fundamental change which can
provide a firmer basis for the major classifi-
cation of the Brachyura than the diagnostic
characters of the traditional major taxa, How-

M. F. GLAESSNER

ever, this reclassification also rejected Gordon's
“logical conclusion” that the peditreme crabs
are not Brachyura. The steps in the evolution
from the peditreme to the sternitreme grade
(Guinot’s Sections Podotremata and Hetero-
tremata) are now better documented both in
living and in fossil forms including those
described or reconsidered here. Earlier
erroneous classifications on family and sub-
family level (Balss, Glaessner) must now be
corrected in the light of these data. However,
they are still incomplete as far as palaeonto-
logical material is concerned,

Wright & Collins (1972) assigned to the
family Cymonomidae (erroneously ascribed
to Ihle 1916 but actually named by Bouvier
1897 as Cymonomae) the fossil genera
Glaessneria Wright & Collins, 1972 (re-named
Glaessnerella in 1975) and Dioratiopus J. T.
Woods, 1953, considering them as “closely
allied”. Tt will be shown below that they are
synonymous. Also included was Mithracites
Gould, 1859, but Withersella Wright & Collins,
1972 and Binkhorstia Noetling, 1881 were
omitted and rather unconvincingly placed in
the Carcineretidae. The fossil genera included
by Wright & Collins range from Lower Aptian
to Cenomanian; Binkhorstia is Maastrichtian.
I have included in the subfamily Tymolinae
the Upper Eocene Falconoplax Van Straelen,
1933. Its sternum is flat and wide, with a deep
abdominal depression and well marked sternal
grooves between sternites 4-8 and a deep
furrow in the anterior portion of sternite 8 of
the female. On the criteria used by Guinot
for suprageneric taxa it would seem necessary
to exclude this genus. The Tymoloidea, accord-
ing to Guinot (1978) with one family Tymo-
lidae, comprise 2 subfamilies Tymolinae
Alcock, 1896 and Cymonominae Bouvier,
1897, in which the living genera Cymopolus
A. Milne Edwards, 1880 and Cymonomus A.
Milne Edwards, 1880 are included, They have
a square, rugose, granulate or spinose cara-
pace, a narrow, triangular, pointed rostrum,
reduced eyes or fixed eyestalks without
cornea, in addition to other characters which
are not discerned in fossils. They live in
deeper water, from 134 to 1269 m (Thle 1916)
and some are abyssal. It seems undesirable to
attach a varied and widely distributed group
of shallow-water crabs with a spatulate ros-
trum, large, apparently normal eyes and a
tymoline sternum to a minor group of small
deep-water crabs which, as we shall see, are
not their only descendants. I propose to

NEW CRETACEOUS AND TERTIARY CRABS 181

separate the Cretaceous genera Torynomma,
Divratiopus, Mithracites, Binkhorstia and pro-
visionally Eodorippe as a new family Tory-
nomidae which appears to be a more
“natural” taxon for them in the sense that its
recognition helps the discussion of tts rela-
tians, Those with Homoloidea and Prosopidae
remain to be clarified after farther studies of
their Jurassic representatives and, if possible,
af the Hauterivian “?Glaexsieria gteneuxi
(Van Straclen) mentioned by Wrght & Collins
(1972, p35). However, this single. frag-
mentury specimen, Of uncertayy provenance,
can no longer be found (R, Forster, pers,
comm. July 1980). Relations to Dramiidac
and Dynomenidac with which their living
descendants have been campared do not
appear to be close. The Torynammidae are
probably ancestral to Tymolidae (Cymono-
minae+Tymolinae) and possibly also Dorip-
pidae and Palicidae. similar to relations
depicted jn Bouvier's phylogenetic diagram
(Bouvier 1897, A. Milne Edwards 1902,
p,106) where their place is taken by unspeci-
fied “Dyhomeniens”.

Diagnosis of the family Torynommicdac:
Carapace square, rectangular or pentagonal in
outline, convex, front spatulale, projecting,
nal strongly deflexed; regions well marked by
grooves including the braachiocardiac; lateral
margins not sharp, side walls steep, eyes well
developed, retractable into shallow orbits.
Sternum triangular anteriorly, oval in outline
posteriorly, with the last sternites vertical and
chevron-shaped, It resembles the sternum of
the Tymolinac and Homolidae rather than the
configuration of the sternum in the Dromiaces.
Where known, the gonopares are coxal (pedi-
treme). Chelipeds subequal, secand and third
pereiopods long and strong, fourth and fifth
Cor the fifth only) in dorsal positioly and much
reduced, Abdomen with first somites exposed
dorsally. Cretaceous (Aptian to Maastrich-
rate).

Toryanmora Woods, 1953
Torynomma (Torynamma) flemingi nov. sp.
FIG. 9

Derivation of name: After Sir Charles
Fleming F.R.S,, Who has made outstanding
contributions to the palaeentological know-
ledge of New Zealand.

Holotype material:
Wellington, No. VA122.

Locality: Koutu Pojnt, Hokianga Harhour
(Loe. V2114). New Zealand.

Vietoria University,

Ave: Upper Senonran-Maastrichtian, Mata
Series.

Description: Carapace slightly convex,
approximately square in general outline, about
30 mm long and wide, Anterolateral margins
almost straight, posterolateral margins which
are damaged were slightly convex, width of
carapace across antero- and posterolateral
murgins was about equal but reduced about
middle of its length. Orbits large, very shallaw;

orbital margin struight, transverse, Extra-
arbital tooth coniesl, pointed, directed
diagonally forward, outward and upward.

Regions und lobes well marked, One small
tubercle on anterior mesogastric lobe, one
painted tubercle on its centre. Pasterior gastric
pits clearly marked. All other spinoge tubercles
arranged symmetrically: } on each side of the
hepatic, metagastri¢ and cardiac Johes, 2 on
cach protogastric and epibranchial lobe. Bran-
chial regions granulate. Carpus and merus of
the cheliped have sharp dorsal ridges. Cheli-
peds robust, subequal, Fourth and fifth pereio-
pads very weik.

Remarks; This species differs from T. quad-
rainm Woods (Fig. 8) in its size and the shape
ef the carapace which is flatter and has a
squarish outline. The spines on the surface
are more Hume;rous and much more prominent,

Paratoryuomima nov. subgen,

Type species. Torynamma (P.) denratum

nav, sp,

This subgenus differs from Terynomrna
Woods in its flatter and wider carapace with
a less deflexed rostrum and a strongly and
evenly granulaled surface, Its most obvious
difference i4 scen in the prominent antero-
lateral extraorbital teeth which are triangular,
directed forward and outward, and mark the
ercatest width of the carapace, The chelae
are more slender and elongate and the
chelipeds are longer, It differs trom Biora-
tiapuy by its almost straight fronto-orhital
margin and slightly converging posterolateral
margins hut resembles it in its granulate surface
sclupture und clongate chelae.

Torynomma (Paratoryuomma) dentatam nov,
5p
FIGS 10-11, 20
Derivation of naine fram the conspicuously
dentate Fronte-orbital margin.

Marerial) Holotype P22930 and Paratypes
P2793) (tacality 1), P2293, P22941,

182

P22943 (Lue, 2); P22944 (Loe, 3), P22942
(Loc, 4), Collected by H, Daily 1954,

Localities: South coast of Bathurst Island,
vorth of Darwin, Northern Territery, Beach
cliffs and shore plattorm, Loc. I-3 are 16-25
km SW of Bathurst Island Mission. Loe, 1:
2.5 km E of Moonkinu Creek, Loe, 2:
Meadinga, E of Moonkinu Creek, Loc. 3:
Pouplimadurie Point, about 4 km E of Moon.
Kinu Creck; Loc. 4: Palliamundery Creek,
ceutral south coast, 35 km W of Bathurst
Island Mission,

Age: Upper Cenamanian, Bathurst Island
Formation, Moonkinu Member, from 3 tm
above to 5 fm below “Tapara Bed” with
Acanthoceray etc. (see Wright 1963, p,612)
at Loc. 1-3. Also From Lower Turonian, upper
part of Bathurst Island Formation with
Collignoniceray cf woolvari (Mantell); Lov.
4, All from glauconitle sands and elays.

Preservation. Most of the & specimens
examined are exceptionally well preserved aud
most were almost complete when embedded
in the sediment but the shell is in various stages
of decortication, The fragile shell and forma-
tion of incipient concretions uround Lhe bedies,
a common feature of the preservation of
decapod crustaceans in clays and sills, makes
complete preparation difficult There is little
distortion and most specimens show little
maverent between the carapace ai the rest
of the body, In several specimens a wap of
a few mm belween carapace and abdomen and
opening of the pleural suture tndivates that
they are probably moults, Seven specimens
are preserved In grey silly clay, one in glau-
conic sand, There is no doubt that they
lived where they were buried,

Deseriptions Carapace rectangular in outline,
wider than long, with a straight fronte-orbital
margin; gently convex longitudinally and
almost flat transversely, No disinct lateral
edge, side walls vertical; posterior maryin
sinuous. Front projecting only 4 short distiinee
forward but defected downward, with median
groove and pair of short basal spines, Pointed.
conical, supraorbital spine ahout eqnidistani
fron) front and conspicuous extraarbital~
anterolateral tooth. This is triangular im outline,
with flat surface, drawty out inte a long,
sharp, anterolaterally directed spine. Its
anterior margili bears 3-4 small lubereles ani
its tip is cranulated, Surface of carapace uni-
formly granulated. with only few more peo-
minent tubercles and few smoother arcas such

M. k GLATSSNER

os furcows and surfuce of anterolateral teeth-
Cervienl, hepatic and greater part of the
branchiocardiac lirrows about equally well
marked, Another posteriolaterally directed
groove vwonees ¢ervicd) and branchiocardiac
irrows and divides epibranchial lobes Meso-
ahd melabranchial lobes divided medially by
a longitudinal groove extending through the
faintly dclimited urogasirié lobe into the
cardiac region. Meso- and metrahranchial
tohes confluent. Orbits very larye and shallow,
vyestalks unusaully robust with smooth sur-
face. In one specimen a part of what appears
to be an antennal stalls projects forward from
helow the supraorbital tooth far about half the
distance separating front from anterolateral
tooth, Its proximal portion cannot be freed
withoul Gamage to the anterior pun of he
carapace.

The shape of the sternum (Fig. 11) 1s
distinelive. It is generally flat bur with sternite
7 (and consequently also 8 which js not pre-
served) turned upward, All sulures eacepl Ty 2
and 4/5 complete and crossing the midline
End of female abdomen extends lo a ridge on
sternite 4-5. Maly abdomen unknown, Gono
pores could jot be fecoenised with certainty
in this species. A third maxilliped preserved
in one specimen shows the ellipticul outline
of the merus, with subterminal articulation of
the carpus. Chelae of the P! subequal, with
clongate rectungulur propodus, covered wilh
spiny yerinylitions which tend to be aligned
in 2 rows Of ils Upper edge, Fingers as long
as the propodus and gently curved. Fixed finger
deflexed slightly dawnward, Merus and carpus
spinose, P2 and P3 about cqually strong and
long, With upper and lower rows of spmes on
the podomeres, Their length exceeds 20 mm,
with a diameter wf up te S mm, P4 and PS
thin, eylindcieal, with a diameter of ahour 1
in. smooth, articulated above the bases of
the Ps.

Divratiopus Waods, 1953
Type species PD salebrosay Woods
Dioratiopas Woods, 1953, p, $2) Wright & Collins

1972, p. 44. 34. 42
Yoritiopas Woods, Glacsener 1969, p. RAV? [erro-

Hioous spelling 1
Ghicawnerta Weight & Collins 1972 (non Takeda &

hiyake 1964), p. 34 17.

Glacswerellt Wright & Collins 1975, p, 441.

As noted hy Writht & Collins (1972, p, 33),
“the penera Glaessneria nov. and Dieratiopay
Woods are closely allied, The authors state
that the species of their new. genus ure dis-

NEW CRETACEOUS AND TRRTTARY CRABS tks

linguished from the Australian Dioratiopus
by being “much flatter in both transverse and
longitudinal section” They “lack the infer
oblique sulcus on the mesobranchial area
and have more strongly apinose frontal areas”.
The diagnostic validity of these characters must
be questioned, While some of the English
species are much flatter than the Australian
type species, this is not correct for the Euro-
pean type species “G.’ spinosa (Van Stfaelen
1936) which is more convex transversely and
unly insiynifizantly less so longitudinally,
except for the raised sides of its rastrum, The
oblique furrow on the branchial area is present
in Dioratiopuy, though possibly less pro-
noutived Jaterally where iL joins the branchto-
cardiac sulcus. ‘The generally weaker
development of carapace furrows in Diora-
tiopus cannot be considered as a reliable
generic character, and the same applics to the
less “spinose frontal areas” which are almost
without spines in several European species.
Small basal rostral, supraorbital and extra-
orbital/anterolateral spines are present in
Dioratiopus. The generic identity of Europea
und Australasian mid-Cretaceous decapod
crustaceans 1s not anomalous but is found in
Homolopsis, Notopocorystes and Hemioon
among the Brachyura and in Macrora. The
speeies. Homolopsis dawasonensis Bishop,
1973, which does not have a linea homolica
was considered by its authar to resemble
Homolopsix depressa Carter but to differ “by
being even less ornamented (haying no
ureoles), having a continuous sagittal ridge.
having more inflated branchial regions, and a
broader cardiac-intestinal region” (Bishop
1973, p, 20). These are specific differences
from “Glaexsnerina” depressa which is a
Dioratiopus, Bishop's species extends the range
of this genus to the Maastrichtian Pierre Shale
of Montana.

Diorariopus salelrasus Woods, 1953

FIG, 12, 12A

Diavationts salebrosus Woods, 1953, p. 53, nl 2.
fix, 4, 5, lext-fig 2.

Doratlapay salebroves Woods, Cilaessner 1969, p-

R492, fig, 704 12)

Materialy One specimen collected ancl
peesunted by H, Wopfner and DB. Scott on
hehall of Geosurveys of Australia No, (22953.

Leeallty: Wooldridge Creek ( =Fossil
Crevk), a tributary of the Alberga River, about
40 km NW of Oodnadatta, South Austraha.

Age: Upper Albian, Marree Formation (See
N. H, Ludbrook, Bull, 40, Geol. Survey 5,
Aust, 1966, ». 38, Map 4).

Remarks: The specimen is preserved ji part
ol # hard coneretian as an exlernal mont,
showing the almost complete carapace and
fragmentary pereiopods, lt agrees completely
with the holotype in type in size, shape aiid
ornamentation,

Dioranapus sp,
FIG. 21

Material; One specimen, P22928_ collected
by B. Daily 1954,

Locality: South coast of Bathurst Island N
of Darwin, Northern Territory, Poupanderi
Point, about 16 km SW of Bathurst Island
Missican

Age: Upper Cenomanian, Moonkinu Mem-
ber of Bathurst Island Formation, within
about | m below the base of the “‘Tapara Bed"-

Preservation, A poorly preserved carapace
in a fereuginised concretion containing alse
scattered remnants and external moulds of
perelopods,

Deseription: Carapace sibrectangular, sides
slightly converging toward the front. About 21
mm long, 16 mm wide and 7 mm high, Sur-
face gently rising, for about S mim from byse
of rostrum, generally fit longitudinally, gently
convex transversely. Greatest width appears to
be at level of posterior end of cardine revion.
Regions of carapace as in lype species; spines

apparently placed as in Mumnoalopsiy edwardsi

Bel] ‘but almost obliterated hy erusian and
probably originally less prominent, Surface
finely granulated. Orbital depressions below
the heputie lobe, stnailer than in the type
species. Chela with rows of granules on pro-
podius and carpus and with slender, curved
dactylus.

This species differs from AL ypinalese an
some details of outline and Jn its weaker
ornamentation, ‘The specimen is insufficiently
preserved for a specific diagnosis hut the
presence of an additional species in the
Bathurst Island Formaticn is of interest,

Eadorippe nov ger
Type species E. spedeni nov, sp,

Diagnosix: Characters as described for the
type species.

EFodorippe spedent noy. sp.
FIG, 14, 13A

Derivation ef tame After Dr [. Speden,
Geological Survey of New Zealand, whe sug-

12. Dioratiopus salebrosus Woods, Artificial cast of carapace of specimen P 22933. x2.

13. Eodorippe spedeni nov. sp. x2.

14. Rhachiosoma granuliferum (Glaessner). x1.

15, 15a. Pororaria eocenica nov. sp. Holotype, 15—dorsal view, 15a
16. Pororaria eocenica nov. sp. Paratype DC 361, x2.

frontal view. x2.

NEW CRETACEOUS AND TERTIARY CRABS 185

10 mm

{O mm

Fig. 14A. Rachiosoma granuliferum (Glaessner), Diagrammatic reconstruction of carapace.
Fig. 20, Torynomma (Paratorynomma) dentatum nov. sp, Paratype P 22936, sternum and appendages,
showing tip of mandible, basal parts of Mx3, P 1-3; right side reconstructed in outline.

Fig. 21, Dioratiopus sp. Carapace reconstructed.

gested the investigation of specimens from
New Zealand and assisted with information,

Material: One specimen, N.Z.G.S., AR 675.
Collected by Mrs J. Wiffen.

Locality: Stream boulders from bed of
Mangahouanga Stream, a tributary of the Te
Hoe River, from between bridge and waterfall.
Loc. No, N 104/f 909, Grid ref. N 104/
261088.

Age: Campanian-Maastrichtian (Pripauan-
Haumurian).

Preservation: Carapace showing dorsal
aspect and left flank almost completely pre-
served, with fragments of shell adhering to
surface of internal and partially preserved
external mould.

Description: Shell thin, carapace transversely
oval in outline, very moderately convex longi-
tudinally and transversely. Greatest width
across mesobranchial and cardiac lobes. Ros-
trum long and narrow, spatulate, pointing
forward; small granules on each side of its

18h

base, upper surlace with a longitudinal
Wepression, lower surface ridged, Fronto-
orbital aargin long, transverse. [ts mer
portion, occupying Jess thin half its wadth
ends laterally in a small, gutter-like depression;
Guler portion has i smeoth rounded,
prontinent ¢die. Infraorbital margin projects
hevond upper surface, orbital depressinns
large wnd shallow aud no suborbital tooth
was observed, Extraorbital-aiterolateral teeth
turn sharply forward, ending in short, pointed
spines, Rounded anterolateral margin bears x
small hepatic spine and leads to a pronounced
sinus mW) whith cervical and hranchiocurdiac
furrows ineet, continuing down the think 4s a
single straighe graave. Behind the incision there
is u shore, sharp ridge. It beoins with a shor
spine and is difected in an outwantly concave
curve backward and outward, slightly above
the rounded true later) carapace pefiphery
Carapace narrows Slightly behind end of ridge,
Posterolateral anil posterior margins sinuous
and toarked by a smuoth rilee Concave
median part of the posterior margin
short, Surface of catapace prominently marked
by transverse furrows aml few tubercles.
Cervical furraw sinuous and medially inter
rupted; hranchiocardiac firrows straight, con.
vergent bul become uaclesr neir the cardiac
region. Between ¢ervical and branchlocardine
furrows a pair of lnterimedlate oblique grooves
are the Most distinctive charaeter ol his cari-
pace pallern. They are literal extensions of a
groove heiween the meso ansl metagastrie
lobgs which are bisected by a weaker median
(lonwitudinaly furrow. Anterior and posterior
portions at the deesal surface are undivided by
gronves and are convex on éach side. Proto-
Bastfic lehes bear three tuhereles cach, spaced
whowl equally along an ure in front of the
cervical groove. leading to hepatic spine,
Mesogastrie Jobe, hounded by straight lines,
extends a long and very narrow tongue for-
ward, An elongate anterior and a shorter
posterior cpibronchial lebe on tach side, the
interior ending in a small marginal tubercle
iv the junetion of the main transverse grooves,
the posterior if shorter, obtiquely triangular in
outing, and sharply delimited from the meta-
gastric lobe, A faint, medially interrupted,
Transverse ridge and sume granulation on the
anterjor part af the i}l-defined cardiac regan
Surface minutely eranulated, with seme coarser
griinules on the Wesebranchial lobes.

Kemarkes There are some similarities in
stapes of the carapace anil the orbits between

M, F. GLAESSNER

(he New species and Milhracites vectensiy
Ciould from the Lower Aptian of England,
The differenees in the pattern of the transverse
furrows, the shape of the mesogastric lobe,
the hartow fostrum and the lack of the coarse
omamentation in Eodorippe justify a generic
istiniticon,

Svitemtatle position; Withers (1951) placed
Mithracites in the Prosopidac, a view rightly
criticised by Wright & Collins (1972, p, 40).
They point with some justification to certain
similarities With “Glaessneria”, here placed
in the synonymy of Dioratiapuy, and conse-
queauy include it in the family Cymonomidae,
The Late Cretaceous mew genus bas some
similarities with Cymonominae, The lack of
information on the morphology of the ventral
side of the cephalothorax of Eoderippe makes
its placing in a modern classification difficult.
In the «bsence of more complete specimens we
cannut decide whether this new genus ix pedi-
freme or sternitreme ancl whether it is oxysto-
matous, Despite the absence of such informa-
lion the striking resemblance between the cara-
pace of Eodorippe and the genus Dorippe
cannol he disregarded. ft is seen not only in
trivial generalities of shape and ornamentation
which can be due to common convergences
bul in details which are unique and cannot
he dismissed. The sinudsity of the wide
posterjor-posterolateral margin can be inter-
preted a8 indicating a position of the third and
fifth perciopods similar to that in Dorippe (the
hases uf the third pereiopods do not affect
the shape of the carapace margin), The lateral
convergence of the cervical and branchio-
cardiac grooves resembles closely the pattern
in several species of Dorippe (D. Jacchine
Herbst, 8. japonica von Siebald, OD. granulate
deo Hain) while the intermediate transverse
xroove appears to be present in D, dars/pes,
The position and shape of the shallow orbits,
the divided supraorhital margia and the extra-
orbital teeth ave also similar. If the classifi.
cation of Guinot (1978) is followed and the
Pulicidae are placed with the Dorippoidea,
some resemblances between Eodorippe and
Pulicus Philippi, 1838 become significant: the
great width across the (nesobranchial lobes,
the unusual multiplication of the transverse
froaves, afd the development of a sapittal
Incsogastnic and metagastrie furrow, Significant
differences ave the appurent absence of the
praminenc infraorbital teeth of Derippe and
the shape of the rostrum, ‘This shape ovakes it
Unlikely that Evderippe had resched the oxy-

NEW CRETACEOUS AND TERTIARY CRABS

stomatous condition of Dorippe with — its
dorsally visible exhalent opening. It agrees with
this structure in the Tymolidac and Torynom-
midae. The known characters of Eodorippe
suggest a derivation of the Dorippoidea from
Tymoloidea, This hypothesis remains ti be
tested by » study of the still unknown ventral
structures of the cephalothorax of Bodorippe.
It js tentatively attached to the Family Tory-
nommidae,

Family Portunidae
Rhachiovema eranuliferim (Glaessner, 1960)

FIG, 14, 14A

Portunites granulifer Glaessner 1960, p. 21,
pl. 3, fig, 7 text fig, 9,

New locality: Coustal cliff at Pororari, 2.4
km N of Punakaiki, Grid ref $37/723,
e85231 8.

Material and preservation: The incomplete
left half of a carapace, broken from the right
half of the front to the posterior margin above
the: fifth left pereiopod, The anterolateral teeth
2-5 are damaged, Mast of the shell surface is
preserved. Canterbury Museum, Christchurch,
New Zealand, Nozfe 9, coll, R, S. Allen,
Feb, 1935.

Age: Upper Focene, Kaidtan-Runangan.
The holotype was from the Middle Eocene,

Remarks; Purther studies of the type species
of Portunites (P. incertus Bell) suggest that
the Original generic assignment of the species
eranulifera was inuppropriate. Prominent
lateral spies are equal to almost half the
width of the carapace in Rhachiosoma bixpino-
yum Woodward, 1871 from the Lower Eocene
London Clay, but less than a quarter of
that width in R, granulifera. However, the
orbits and carapace sculpture are similar, The
new, fragmentary specimen has weaker and
probably fewer tubercles on the carapace
surface than the holotype bur until new and
more complete material is found, these
differenees are insufficient for the establish-
ment of a new taxon, The classification of these
fossils should be reviewed when the ventral
aspect and pereiopods are known,

Family uncertain
Pororaria nov. gen.
Type species P, cocenica nov, sp.

Diaenosis: Characters as described for the
Type species.

Pororaria cocenica sp. nov
FIGS 15, 15, 16

Marerlal; Two almest complete carapaces-
Holotype (including right chela). New Zealand
Geological Survey collecitan DC 360, paratype
NC 361, Also fragments of carapaces, chelne
aid walking legs and one carapace with both
chelwe: Canterbury Museurn, Christehurch,
New Zealand, Na_ 2fe 31-38, coll R. 8, Allen
Feb, 1935,

Locality: Cliffs al Porotari, 24 km Not
Perpendicular Point, Punakaiki (fe 71-38);
800 m S$ of Perpendicular Point, Map Sheet
S 37/735, NZ Geol. Survey locality
GS (0490 (DC 360-361)

Axe-> Upper Eocene. Kaittun-Runangan,

Description; Carapace as long as wide. tut
line oval ta hexagonal, very slightly convex
to flat surface smooth with localised concen-
trations of granules. Pront with a median
fgtch aid on cach side ) strong and 2 weak
teeth, Orbits as Wide as the front, with supra-
orbital yeauulations and 2 nolehes, with a gap
between the large antennal base and an tfra-
orbital tooth. Antennular bases folded back
obliquely, About 5 anterolateral teeth, obscured
by marginal granulaticns, Posterolateral
margin starts from a ridge behind last tnteral
loath hut not delineated on the inflated meso-
branchial lobes. Cervical groove well defined,
starting from posterogastric pils and
Uelimiting on cach side in three forwardly
eonvave arcs the mesogastric, protogastric and
heputic lobes, These are equally well defined
by hepatic grooves, There ure couieal tubercles
on hepatic lobes; groups of spinose granules
on meso-, proto- and metagastric. efi-, Meso-
and metabranchial, und eardiac lobes. Epi-
branchial ficld of granules has an unuswal
U-shape which resembles similarly placed
granulations in Dakelicaneer Rathbun, As
in this genus, a branchwcardiac groove
crosses rounded lateral margin behind
this granulated lobe. Urogastcie lobe finely
pitted and rectangular, cardiac region broadly
hexagonal. Posterior margin has a strong,
granulated ridge wilh » sieht median forward
bend, Chelipeds are sirong and heterochelous.
Carpus with spine near its inner distal angle.
Inflated propodus has smooth twner and
Pranulatcd outer and upper surfaces, Granules
tend to be arranged in longitudinul rows.
Fingers shorter than the upper length of the
propodus. strong minutely granulated, grooved

188 M. F. GLAESSNER

but otherwise smooth. Walking legs preserved
as scattered fragments which are thin and long.

Systematic position: In the absence of com-
plete pereiopods and of the sternum it is diffi-
cult to reach a definite conclusion about the
assignment of this new genus to an existing
higher taxon. It resembles the less advanced
Portunidae such as some Carcininae (Necto-
carcinus, Carcinus) in the general shape of
the carapace and in some characters of its
fronto-orbital region, Nectocarcinus shows a
similar transition from anterolateral teeth to
groups of spiny granules. The chelipeds are
also similar. Thin, long legs do not occur in
Carcininae and the lobes of the posterior half
of the carapace are different. There are also
resemblances with Atelecyclidae among the
Corystoidea but the relevant details of the
fronto-orbital region are not well enough pre-
served in the genus to allow a definite
conclusion about portunoid or corystoid
affinities; the configuration of the postero-
lateral portion of the carapace does not favour
the latter but the chelae are cancroid, A
similarity of the carapace with that of Avitel-
messus Rathbun does not apply to its median
portion which shows a corystoid pattern.
Avitelmessus is very close to Dakoticancer.
As Guinot (1978) remarked, these genera
should not be assigned to the Dromiacea.
Whether Pororaria is related to Corystoidea
or Portunoidea remains to be elucidated. Until
further evidence is found, the new genus is
tentatively placed in the superfamily Por-
tunoidea.

Family Panopeidae
Panopeus H. Milne Edwards, 1843
Type species P. herbsti H. Milne Edwards

Panopeus whittenensis nov, sp.
FIGS 17-19

Material: More or less complete carapaces,
chelae, fingers, and fragments of legs, Holo-
type P22935, paratypes P22938, 22939,
22946-22950.

Localities: Whitten Bluff, S of. Christies
Beach, 28 km SSW of Adelaide (type locality
and most common occurrence). Also from

Fig. 17. Panopeus whittenensis nov. sp. Holotype,
x3. Posterolateral margins restored.

Fig. 18. Panopeus whittenensis nov. sp. Paratype
P 22938, x2.

Fig. 19. Panopeus whittenensis nov. sp. Right
chela, specimen P 22939, x2.

NEW CRETACEOUS AND TRRTIARY CRABS

Maslin Bay. about I5 km turther south,
Coastal cliffs on the E shore of Gulf St
Vincent (P22438, coll, Dr M, Buonajuia)

Age: Upper Eocene, upper Tortachilla
Limestone 4nd hase of Blanche Point Porma-
tion (= “Transitional Marl"). Planktanic
foraminiferal zone 15.

Preservation; Some complete and same
broken carapaces without branchiostegates,
sternum, abdomen, or ativehed appendages,
The propodus of the cheliped may have the
dactylus and parts of the carpus attached but
more offen it is disjointed, Small fragments
of perciopods apparently belong to this species,
The only other decapods found with Panopets,
in a rich shallow-water fauna of cirripeds,
bryozoans, brachiopods, echinoderms, serpulid
worm lubes (some attached to carapaces and
chiws of Periopeds) and numerous mollusca
are one incomplete dromiid carapace (Drurni-
fires? sp.J, a few claws of an oxyrhynchous
erab, and chelae of a callianassid.

Description: Carapace slightly wider than
long, convex. Frontal margin straight, with
median indentation and transversely grooved
and finely granulated edge. Orbits well
developed, with inner supra- and infraorbital
teeth, evenly granulated edges and 2 supra-
orbital fissures. Anterolateral margins with 5
conical teeth of which the fourth is the
strongest and mast prominent and the fifth the
weakest, Posterolateral margins converging
hackward. Posterior margin wide, with raised,
faintly granulated rim, Main regions of the
carapace surface well marked by shallow
grooves, Posterior gastric pits clearly visible,
Surface appears to be smooth when well pre-
served, becoming granolated with decortication
by weathering, Some mdividual variability: in
omamentation. No remarkable features in the
pattern of regions except perhaps strong
development of the epigastric and metagastric
lobes, Right chela strong, propodus smooth,
with a very faint depression below the upper
edye and two rows of pits above and along
lower cdge of fixed finger. A tow of irregu-
larly sized and blunt teeth on its upper edge,
Lower edge of the propodus and finger strongly
curved to fit snugly along fhe anterior margin
cephalothorax. The carpus fs square.

Measurements (in mm): Lengths and widths
17.8 and 22.5, 22 and 26, 20.5 and 23, 22,5
and 28. In the carapace which is 20,5 mm tong,
the frontal margin has a half-width of 4 mm
and the orbit was 4.5 mm wide In a cara-

isy

pace which wis about 20 mm long and 24 mn
Wide. the orbit was about 5 mm wide A
large propodus wilh fixed finger is about 35
mm long and 18 mm high.

Remarks: In the absence of any parts of
the ventral surface of the cephalathorax or
appendages other than the large claw, the
precise systematic position of this ctab remains
in some doubl, The absence of transverse
raised lines ou the carapace would distinguish
it from the living species of Panepeus for
which their presence ts a Convenient diagnastic
character. tt would not necessarily exclude it
from the evolubonary lineage of this genus
which has been reported from the Paleocene
and Eocene. The new spocies as far as known,
16 morphologically closest ta Parnapeus and
therefore assigned to it, pending diseovery of
other still missing parts.

Phylogenetic conclusions

The phylogenetic significance of several of
the new Cretaccous and Tertiary crabs here
described has been briefly mentraned, It can
now be considered in the framework of the
adaptive radiation of the Brachyura (Fig. 22,
see also Glaessner 1980). This updated version
of a phylogenetic diagram which & first puh-
lished 50 yeurs ago (Uilaessner 1930; L960,
Fig, 22) incorporates some of the results of
the monogeaphic work of Wright & Collins
(1972) on British Lower Cretaceous crabs
and reflects the new classification of ihe Brach-
yura by Guinot (1977, 1978) and other
important studics which are considered jn it,
Certain modifications and ce-acrangements of
this classification are suggested ss a result of
the present investigation.

The new family Torynommidac may have
evalved ultimately from the family Peosepidae
of the Homolodromividea, as implied by
Wright & Collins (1972) and probably from
archacobrachyuran descendants, the Homo-
loidea, iv Late Jurassic-Eacly Cretaceous or in
Mid-Cretaceous time, Their living descendants
are related fo them in a manner depicted with
remarkable foresight by Bouvier as early as
1897. They also may have given rise to the
Late Cretaceous Carcineretidae which acquired
an early adaptanon ta swimming. From similar
origins in the podotrematous Homoloidea the
well docurnented Necrocarcininac-Calappinae
lineage (Férster 1968) and possibly Orith-
yidae (sce Guinot 1978, p.255) evolved and
and eventually attained the fieterotrematous
trade. Their Mid-Cretacemus descendants, the

190 M. F. GLAESSNER

HETEROTREMATA. ssi THORACOTR.
PODOTREMATA |
ARCHAEO | BRACH. DROMIACEA

\ Cc
er rerogoRO*_co®

a 2
4X nh abgey, p>

50
U
Cc
100
F HABITS &
c HABITATS s] m2
B: BURROWING at Sr csk
150-7 C: CONCEALMENT Fram cape ot re
L:LAND
z S: SWIMMING

200

Fig. 22. Adaptive radiation of the Brachyura.
Superfamilies (and lower taxa marked with *): LEU Leucosioidea, MAJ Majoidea, PAR Partheno-
poidea, COR Corystoidea, POR Portunoidea, DOR Dorippoidea, CARC* Carcineretidae, TYM
Tymoloidea, RAN Raninoidea, ORI* Orithyidae, CAL* Calappidae, NEC* Necrocarcininae, HOM
Homoloidea, H/DR Homolodromioidea, DROM* Dromiidae, DYN* Dynomenidae, XAN Xanthoi-
dea, GEC Gecarcinoidea, GRAP Grapsoidea, PIN Pinnotheroidea, HEX Hexapodoidea, OC Ocypo-
doidea. Geochronometric (in million years, my) and stratigraphic scales at left. N Neogene,
P Palaeogene, UC Upper Cretaceous, LC Lower Cretaceous, UJ Upper Jurassic, MJ Middle Jurassic,
LJ Lower Jurassic, Note that top of Paleocene is shown within Palaeogene, and Albian and Ceno-
manian are marked below and above line dividing Cretaceous. Dotted lines mark the three periods

of major diversification of the Brachyura.

Raninidae, adapted to burrowing and remained
at the podotrematous grade, The discovery of
Eodorippe strongly suggests a Late Cretaceous
origination of the heterotrematous Dorippidae
from Tymoloidea, as foreseen by Bouvier. This
is still subject to confirmation by more com-
plete material.

The origins of the remaining Heterotremata
are still unknown, probably diverse, and dated
mostly Late Cretaceous to Early Tertiary.
Wright & Collins (1972) conclude that the
Xanthidae were derived from Dynomenidae
in Late Jurasic-Early Cretaceous time, This
derivation cannot apply to other Heterotre-
mata which show no signs of origination from
Dromiacea or indeed of pre-Tertiary existence.
It is possible that further studies of Pororaria
may clarify relations with or between Por-
tunoidea and/or Corystoidea (a prior synonym
of Cancriformia). Some members of these
two superfamilies show a dichotomy between

habits and adaptations for swimming (as in
Portunus) as opposed to burrowing (as in
Corystes). The portunoid adaptations for
swimming do not seem to be derived from but
rather heterochronously convergent to those of
the Carcineretidae. The burrowing habit of the
Corystidae is generally considered as a con-
vergent or possibly parallel adaptation com-
pared with that of the Raninidae. The origins
of the Leucosiidea and Majidea remain
obscure; their adaptations diverge distinctively
from those of any other group of crabs.

The Thoracotremata, at the highest grade,
include the land crabs and various successful
strand dwellers with a variety of habits, They
resemble the Xanthoidea more than other
Heterotremata but their origin, in the Late
Cretaceous or Early Tertiary, is cryptic.

As in all phylogenies, many questions
remain, but new material, particularly of
Mesozoic and Early Tertiary age, can now be

NEW CRETACEOUS AND TERTIARY CRABS ina

fitted more easily and rationally into the
framework of existing knowledge and classifi-
cation, Most importantly, the replacement of
the traditional but mostly ill-conceived “Scec-
tions” with a horizontal classification at the
highest level by grades, has improved our
understanding of the history of the Brachyura.

Acknowledgments

My thasks are duc fo Dr B. Daily, Depart-
ment of Geology and Mineralogy, University
af Adelaide, who collected the fossil ¢rabs
{rom outcrops on Melville and Bathurst Island,
Northern Territory, and established their
stratigraphic position; to Dr 1. Speden, Geo-
logical Survey of New Zealand; to the
authorities of the Victoria Museum, Welling-
ton and the Canterbury Museum. Christchureh,

New Zealand, and to Dr Mary Wade, Queens-
land Museum, Brisbane. for the loan of speci-
mens from collections in their institutions, to
Dr N. H. Ludbrook for ihe loan of a
specimet from South Australia and for infor-
mation, und to Mr ©. W. Wright for valuable
discussions on British Cretaceous crabs, Dr
Daniéle Guinot, Muséum noational dHistaire
naturelle, Paris, discussed relevant aspeets of
her work with me and provided essential
literature.

All specimens with numbers prefixed P have
been deposited in the Sovth Australian
Museum, Adelaide, The photographs were
prepared by Mr R. Barrett, The skilful
drafting of Fig, 22 is the work of Misy A.M,
C. Swan. Geology Department, University of
Adelaide.

References

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Histor, G. A. (1973) Homolopsis dawisenensise
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Bouvien, E. L. (1897) Sur ta classificutian. Jes
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192 M. F. GLAESSNER

Postscript

Via Boada (1980) reviewed the evolution of the
Ocypodoidea, tracing their origin to the Creta-
ceous genera Ophthalmoplax and Archaeopus. I
had previously (Glaessner 1969) referred the first
to the Carcineretidae and the second, doubtfully,
to the Palicidae. Subject to further studies, Via’s
views suggest interesting possibilities of exploring
evolutionary links between Ocypodoidea, certain
Dorippoidea, and Carcineretidae. When I received

his new data it became possible to identify a
Middle to Late Eocene crab from a limestone core
from the Ashmore Reef No. 1 Bore off the NW
coast of Australia (sent to me in 1968 by Dr P. J.
Coleman and the Burmah Oil Company) as at
least very close to Retropluma as described from
rocks of similar age in Spain, It lives now in deep
water in the Indian Ocean.

A NEW SPECIES OF CYCLORANA (ANURA: HYLIDAE) FROM
NORTHERN QUEENSLAND

BY ERIC VAN BEURDEN & K. R. MCDONALD

Summary

Cyclorana manya sp. nov. from Cape York Peninsula, northern Queensland, is described. It is small
compared with known congeners and has close affinities with C. brevipes and C. longipes.

A NEW SPECIES OF CYCLORANA (ANURA: HYLIDAE) FROM
NORTHERN QUEENSLAND

by ERIC VAN BEURDEN* & K. R. MCDONALD?

Summary
VAN BEURDEN, E. & MCDONALD, K. R. (1980) A new species of Cyclorana (Anura: Hylidae)
from northern Queensland. Trans. R. Soc. S. Aust. 104(6), 193-195, 28 November, 1980.
Cyclorana manya sp. nov. from Cape York Peninsula, northern Queensland, is described.
Tt is small compared with known congeners and has close affinities with C. brevipes and C.

longipes.

Introduction

In a partial revision of frogs of the genus
Cyclorana Steindachner, Tyler & Martin
(1977) described five new species and raised
the total to 10, highlighting the diversity
within the genus. We describe a further species
from northern Queensland. It resembles C.
longipes Tyler & Martin but is considerably
smaller than this or any other congener.

Methods

The type specimens are lodged in the
Queensland Museum (QM), and South
Australian Museum (SAM). Measurements
were obtained using a pair of Helios dial
calipers (to nearest 0.1 mm). Eye to naris dis-
tance (E-N) is the distance from the posterior
margin of the external naris to the anterior
margin of the eye. Internarial span (IN) is the
distance between the medial margins of the
external nares. Other measurements are as
defined and abbreviated by Tyler and Martin
(1975): HL = head length, HW = head
width, S-V = snout-vent length, TL = tibia
length.

Mating calls of two individuals were
recorded on a Uher 4000 report tape recorder
at a tape speed of 19 cm/sec using a Grampian
DP4 microphone. Air temperature was
recorded within 3 cm of the frog using a
Schultheis mercury thermometer. One call of
each individual was analysed using a Kay
Model 6061—A Sono—Graph audiospectrograph
set on a narrow-band (45 Hz band-pass), and
a response curve setting of FL—-1. This pro-
vided measures of call duration, band spacing,
lowest and highest frequency. Pulse repetition
rate and fundamental frequency were taken

* Department of Zoology, University of New Eng-
land, Armidale, N.S.W. 2351.

* Queensland National Parks & Wildlife Service,
Pallarenda,

from photographs (Polaroid B & W type 107)
of oscilloscope traces using a time scale of 2
msec/cm,

Cyclorana manya sp. nov.
FIGS J-2

Definition: A small species (male adults 27.2—
29.9 mm; female 27.5 mm) with short hind
limbs (TL/S-V_ 0.32-0.38). The head is
bluntly rounded and the eyes are large (E/ E-N
= 1.71). There is no dark head stripe from
the tip of the snout to the eye or from the eye
to the axilla.
Description: Holotype (QM J34886), an adult
male collected between Coen airport and Deep
Creek crossing 25 km N. of Coen township
(13°52’S, 143°12’E), Cape York Peninsula,
Queensland, by E. van Beurden, M. Sabath,
B. Easteal, M, Robinson and J, Sparkes on
17.1.1979.

Habitus stout, body globose (Fig. 1). Size
small. Head broadly triangular when viewed
from above, flattened and broader than long.

Fig. 1, Paratype of Cyclorana manya shortly after
capture near Coen.

194 ERIC VAN BEURDEN & K. R. McDONALD

Fig. 2. Palmar and plantar aspects of hand and
foot of holotype of Cyclorana manya (QM
J34886).

Snout rounded when viewed from above and
bluntly rounded in profile (HL/HW 0.89).
Nostrils inconspicuous and dorsally inclined.
Canthus rostralis slightly curved but not
prominent. Eye prominent, its width more than
3 times eye to naris distance. Upper part of
iris golden, lower part, silver/grey and an-
terior and posterior portions black.

Tympanum distinct except for upper and
posterior margins which are obscured by
supra-tympanic fold.

Tongue ovate, about 4+ free behind and just
over 4 as wide as gape, Choanae large and
widely spaced. Vomerine teeth present on
obliquely converging elevations, between pos-
terior margins of choanae.

Fingers short and unwebbed (Fig. 2) with-
out lateral fringes. Subarticular tubercles
prominent; paired nuptial pads on each thumb.

Toes slightly webbed. Webbing on medial
side of 5th toe does not reach subarticular
tubercle at base of penultimate phalanx.
Relative toe lengths 4>3>5>2>1. Prominent
inner metatarsal tubercles, twice length of first
toe (Fig. 2). Hind limbs short (TL/S-V 0,34).

Skin of anterior portion of thigh smooth;
that on posterior pitted.

Dorsal body markings of preserved specimen
consist of dark grey-brown irregular blotches
on lighter brown background. These blotches
diminish in size and intensity on flanks and

posterior parts of dorsum. A pale, unbroken
vertebral stripe runs from tip of snout to tip of
urostyle. A broad, light-coloured post-orbital
bar crosses head posteriorly at level of tym-
pana. Top of head dark grey-brown while
sides, extending down to upper lip, are pale
brown.

Ventral surfaces of body and limbs pale
cream, as is lower lip. Skin beneath vocal sac
darkly pigmented, and skin of abdomen trans-
lucent. Posterior region of thigh and flanks
pale brown with darker brown blotches.

Dimensions: S-V 29.9 mm; TL 9.5 mm; HL
9.2 mm; HW 11.2 mm; E-N 2.1 mm; IN 1.5
mm; E 3.8 mm; T 1.2 mm.

Variation; Two paratypes: a mature female,
QM J34888, and a mature male QM J34889,
were collected with the holotype. Twelve fur-
ther paratypes SAM R17420-R17424; QM
J36894—900) including nine mature males, and
three immature individuals were collected by
R. G. Atherton and K. R. McDonald at the
southern end of Coen air strip (13°46'S,
143°07’E) on 6-8.iii, 1979.

The S-V range is 27.2—29.9; the largest
specimen being the male holotype. The head
is consistently broader than long (HL/HW
0.82-0.96) whilst the nostrils are narrowly
spaced (E-N/IN 1.25-1.69), The hind limbs
are consistently short (TL/S—V 0.32-0.38).

Colour and pattern are variable. The colour
of the dark blotches on the dorsum varies from
light grey to a rich, dark brown-grey, The pro-
portion of the dorsum covered by these
blotches varies from about 55-85%. The
vertebral stripe varies in its conspicuousness
and the variation is not consistent with the
intensity of blotches.

All specimens larger than 25 mm S—V,
except the female, were males with pigmented
vocal sacs and nuptial pads. The single female
was gravid and contained about 100 eggs, each
about 1 mm diameter. Specimens less than
25 mm lacked eggs or male secondary sex
characteristics.

Mating Call: The call is a short plaintive bleat
of 1.1-1.3 sec. duration (Fig. 3). The domi-
nant frequency is about 2500 Hz and the call
consists of S—8 distinct bands separated by 328
Hz, and ranging 400-3000 Hz. Pulse repetition
rate is 294 pulses/sec., and the call is
repeated 10-16 times per minute. Calling males
were recorded and collected while floating,
legs extended, at the surface of shallow

A NEW SPECIES OF CYCLORANA 195

ENCY (kL!

FREOU

TIME anc}

Fig. 3. Audiospectrogram of mating call of Cyclo-
rana manya vecorded during rain at Coen Air-
port. (Air temp. 24.8°C, band pass = 45 Hz.)

temporary puddles,
during heavy rains.
24.8°C.

Breeding: The presence of juveniles in March
and the calling activity in heavy rainfall sug-
gests that this species is an opportunistic,
summer breeder, Little calling was heard on
nights when rain was light, and there was none
on the dry nights following that on which
recordings Were made.

Comparison with other species; Comparisons
are based on descriptions provided by Tyler &
Martin (1977), The most obvious difference
between C, maya and congeners is the S=V of
adult males which is 4.8 mm shorter than the
smallest adult previously reported, It also lacks
a head stripe from the snout to the eye and eye
to axilla, The range of B-N/1N overlaps only
C. longipes. Call characteristics are most
similar to those of C, brevipes Gunther, The
call duration of the tWo species is about 1 sec.,
und the dominant frequency has 3-4 side

in grassy depressions
Air temperature was

bands above and below it. The call of C,
manya differs from that of C. brevipes in
that the dominant frequency i higher and
band spacing is wider. Coloration most closely
resembles that of C. longipes except that dark
head pigmentation terminates at a broad, pale
transverse bar extending around the tympanum
and down to the axilla.

Four frogs (DT-D 0737-40) taken at
Lower Archer River on Cape York Peninsula
by J. Thompson in 1933 may represent a (orm
intermediate between C. Jongipes and C.
manya. Their E-N/IN ratios (1.22-1.25) and
bold head coloration are similar to C, lon-
gipes, S-V (32,2 and 40.7 for the two adults)
is intermediate between C. manya and the
larger C. longipes.

Etymology: ‘Manya’ is the Aboriginal word for
‘small in the local dialect (Wik-munken) at
Coen. This is appropriate for ihe smallest
‘water-holding frog’ described.

Acknowledgments

We thank J. D. Miller and Michael J. Tyler
for reviewing the manuscript. Drs Michael
Sabath, J. W. Winter and Janet A. Taylor also
provided valuable comments. E.V.B. was funded
by the Australian National Parks Wildlife
Service (grant for the study of Bufo marinus)
and COMALCO, and Ian Potter Foundation
trants to M. Sabath and S. Basteal. K.R.M.'s
work was carried out in the course of a
Melllwraith Range fauna survey (Dr J. W.
Winter and R, G. Atherton co-ordinators)
partly funded by the Australian National Parks
and Wildlife Service. We are also grateful to
the Donald Thompson Collection Administra-
tion Committee, and the National Museum of
Victoria for allowing us to examine the speci-
mens from Lower Archer Raver.

References

Tyrer, M. J. & Martin, A. A. (1975) Australian
leptodactylid frogs of the Cyclorana australis
complex. Trans, R. Soc. S. Aust. 99, 93-99,

(1977) Taxonomic studies of some Austra-
lian leptodactylid frogs of the genus Cycloraru
Steindachner. Rec. 8. Aust. Mus., 17, 261-276.

EARLY CAMBRIAN TRILOBITES FROM THE OFFICER BASIN, SOUTH
AUSTRALIA

BY J. B. JAGO & B. C. YOUNGS

Summary

Fragmentary trilobites, probably of early Early Cambrian age, occur in the Observatory Hill Beds
from the eastern part of the Officer Basin. The trilobites were recovered from limestone core at a
depth of 87.85 m in Marla-1, a stratigraphic hole drilled by the South Australian Department of

Mines and Energy. Their discovery allows the first reliable macrofossil dating of the Officer Basin
sediments in South Australia.

EARLY CAMBRIAN TRILOBITES FROM THE OFFICER BASIN,
SOUTH AUSTRALIA

by J. B. JAGo* & B. C. YOUNGs*

Summary
Jaco, J. B. & Younes, B. C. (1980) Early Cambrian trilobites from the Officer Basin, South

Australia. Trans. R. Soc. S. Aust. 104 (6),

197-199, 28 November, 1980.

Fragmentary trilobites, probably of early Early Cambrian age, occur in the Observatory
Hill Beds from the eastern part of the Officer Basin, The trilobites were recovered from lime-
stone core at a depth of 87.85 m in Marla-l, a stratigraphic hole drilled by the South Austra-
lian Department of Mines and Energy. Their discovery allows the first reliable macrofossil
dating of the Officer Basin sediments in South Australia.

Introduction

The Officer Basin is a large intracratonic
depression in western South Australia and
southeastern Western Australia (Fig. 1). The
main part of the S.A. portion covers an area
of approximately 100 000 sq. km. and contains
sediments ranging in age from Late Proterozoic
to Devonian in surface and near-surface out-
crops (Fig. 1). Subsurface outliers exist to the
east and south of the main basin (Pitt et al.
1980). The deep northern parts of the Officer
Basin may contain at least 6000 m of
sediments.

This paper reports the first trilobites recorded
from sediments in the S.A. portion of the
Officer Basin. The only previously recorded
macrofossil from this part of the Basin is a
single specimen possibly representing Biconu-
lites (Gatehouse 1976). Prior to the present
report, the only palacontological evidence for
ee ——

* School of Applied Geology, South Australian
Institute of Technology, P.O. Box 1, Ingle Farm,
S. Aust. 5098.

+ Geological Survey of South Australia. Present
address: South Australian Oil and Gas Corpora-
tion.

1 Harris, W. K. (1968). Continental-Sun-Exoil-

Transoil Munyarai No. 1 well palynological exa-

mination of cores. S.A. Dept Mines & Energy Rept

Book 754, Appendix 4 in envelope 979 (Cunpub-

lished).

2 Gilbert-Tomlinson, J. (1968). Fossils from Mun-

yarai No. | Well, Officer Basin, South Australia.

Appendix 7 in S.A. Dept Mines & Energy Rept

Book 979 (unpublished).

3 Vlierboom, F. W. (1973). Palynology and source

rock potential of core samples from the Conoco

exploration well Munyarai-1, Officer Basin, South

Australia. In S.A. Dept Mines & Energy Rept

Book 979 (unpublished),

4+Muir, M. D. (1979). Palynological examination

of microfossils from the Observatory Hill Beds,

Wilkinson No. 1 DDH, Officer Basin, South Aus-

tralia. Appendix 2b in S.A, Dept Mines & Energy

Rept Book 78/88 (unpublished ).

the age of any of the basin’s sediments came
from Devonian microfossils recovered from
cores in Munyarai-1!* and Early Cambrian
microfossils from the Observatory Hill Beds in
Wilkinson—I+, The specimens figured herein
are housed in the palaeontology collection of
the S.A. Museum; the catalogue numbers refer
to this collection.

Stratigraphy

The stratigraphy of the eastern Officer Basin
is summarized in Figure 2. The fauna discussed
in this paper was recovered from cores in
Marla—I (lat. 27°28.1'S, long. 133° 44.8’E), a
stratigraphic well drilled by the S.A, Depart-
ment of Mines and Energy in 1974 (Thornton
1978). A more complete section through these
beds was drilled during 1979 in Marla—1B
(Fig. 3): this well, located near the site of

==

+, +

OFFICER BASIN
(Area of outcrop and

shallow suberop} OBSERVATORY HILL A
(iyps, soeuson)

Wilkinson—1¥y

OFFICERY DI
BASIN SA jm~ienoms)
peat

KILOMETRES
o 100

0 yono

Lt MELBOUIINE
YILOMETRES:

Was cADME

Fig. 1. Locality map, Officer Basin.

198 !,

Boorthanna Formation

Isufsurtace Arckarnngs
Basin only)

Minsabie and Wartaona Beds
‘Munyarat Sequence’
(Subsurface only)

Carn Beds

Blue Hills Sandstone
Indulkana Shale

Mount Chander Sancstane

Sequence

Munda
PALAEOZOIC

Traior Hill
Sandstone

Mi Johns
Dongomerate

Cambrian

OBSERVATORY ‘Doves Anre

HILL. BEDS

Conglomerate
Gan)

3 Marit Sequefice

Marinoan
LATE <=
PROTEROZOIC

>

Fig. 2. Simplified stratigraphy, Officer Basin, S.
(modified after Pitt ef al, 1980),

Marla—1, intersected nearly 300. m and was
still in che Observatory Hill Beds when drilling
was stopped.®> Marla-1 well intersected
22,58 m of Observatory Hill Beds unconform-
ably beneath the "Early Permian Boorthanna
Formation, The specimens desoribed herein
are trom 87.85 m depth, Other, at present
indeterminate, tnlobite fragments have been
found at a depth of 333 m in Marla—-1B.

The Observatory Hill Beds in Marla-IB are
a flat lying sequence of predominantly fine-
grained, buff to grey, silty carbonates with
minor thinly interbedded calcareous siltstones
and sandstones, Above 176 m, caleite und
dolomite mudstones and boundstones are coni-
mon. with chert and fluorite occurring
throughout. Below 176 m to the total depth of
379 m, the section is predominantly dolomitic
and evaporitic mudstones with some bound-
stones, wackestones and packstones, Fine- to
medium-grained calcareous «and dolomitic
siltstones aod sandstones are interbedded
throughour, Fluorite is rare, bul secondary
chert and sulphides are common

" Benbow, M. C. (1980). Marla-1A, 1B well com-
pletion report. S.A, Dept Mines & Energy Rept
Book 80/22 (unpublished )_

"Lydyard, A. J, (1979). A petrographic study of
the sediments in seven Officer Basin stratigraphic
wells. S.A. Dept Mines & Energy Rept Book 79/55
(unpublished ).

B, JAGO & B,C. YOUNGS

Apart from algal mats and stromatolites
throughoul the sequence, the only fossils are
trilobite fragments. These are found oily at the
top of the drilled section and were recorded at
levels similar to those in Marla—l-

Unbke the Observatory Hill Beds in Byil:
Kagoora-l (Fig. |) which are considered non-
Marine (White & Youngs 1980), those in
Marla-!, —1B are interpreted as marginal
marine, The basal 200 m were deposited on
mudflats and in tidal channels which may have
been subjected to periodic exposure. The beds
above 176 m show an increase in marine
influence and were probably deposited in the
shallow, quiet waters of a lavoon on a broad,
shallow platform,

Valaeontology

Several trilobite fragments were recovered
from limestone core at a depth of 87.85 ni in
Marla—|, after having been discovered during
pelrographic work,” However, there are only
three small, fragmentary specimens on which
reasonable comment can be made. The small-
ext available specimen, P22981, is very small
(Fig. 4a) and may be wn immature form of the
speeies described below. However, the glabellar
furrows of P22981 seem to be shorter and less
distinct aud the anterior border is narrower
than in that species, Until more and better
material is available, it will not he possible to
identify this specimen.

QUATERNARY AND EARLY CRETACEOUS AND
2 EARLY PERMIAN |BOORTHANNA FORMATION)
(See Bantow, 1980)

EARLY CAMARIAN—OBSERVATORY HILL BEDS

f Carpandtis (predaminaniy miudstonesy with
f imerbedderl silty camanaias, salearcnus
seirstunes and sandstones

Algal bounderanes inroughout
Cheriy Urroughout

Flubtite Predum ates above 170in
and sulphides helow (Oi

Berumindus oratetial in laminae and 4tylatiles

ae
ia 329 60m

1 et

Fig. - Simplified log, Maria 1-B, Officer Basin.

‘

EARLY CAMBRIAN TRILOBITES FROM OFFICER BASIN 199

Fig. 4. A. P22981, immature cranidium, X9. B.
P22982, part of left hand anterior part of crani-
dium, X8. C. P22983, partial cranidium, X7.5.
All photographs are of silicone rubber casts
whitened with magnesium oxide.

The other two specimens, P22982-3 (Figs
4B-C), represent a single species. This species
has a glabella which apparently has a well
developed occipital furrow and three pairs of
lateral glabellar furrows. The lateral glabellar
furrows are directed inwards and backwards,
with the anterior furrows being shallower than
the posterior furrows. The position of the
glabellar anterior cannot be determined. The
wide palpebral lobes extend from opposite the
occipital furrow to opposite the 3p glabellar
furrows; the wide eye ridges meet the glabella
immediately forward of the 3p glabellar fur-
rows, There is a suggestion of the presence of
a para-frontal band.

The wide, almost flat border widens
abaxially; it stands well above the preglabellar

tions of the facial suture commence opposite
the 3p furrows and diverge markedly to the
marginal furrow, from where they converge
gently. The almost flat palpebral areas of the
fixigenae are separated from the palpebral
lobes by narrow shallow furrows.

The short, postocular sections of the facial
sutures diverge markedly and enclose small
posterolateral limbs. The marginal furrows are
broad. The forward part of the glabella has a
low reticulate ornament. The remainder of the
cranidium visible is finely granulose.

The shape of the facial suture, and the shape
and position of the palpebral lobes suggest
affinity with Pararedlichia, Eoredlichia, Wutin-
gaspis, Chaoaspis or a related genus of the
Redlichiidae. Such genera are found in lower
Lower Cambrian rocks of China (Chang 1966,
Lu et al. 1974, Li 1980), France (Courtessole
& Jago 1980), Vietnam and Morocco (Hupé
1953). In South Australia Eoredlichia has been
recorded from Faunal Assemblage II in the
Flinders Ranges (Daily 1972). Daily cor-
related this assemblage with the early part of
the Atdabanian of Siberia, This suggests that
the specimens figured herein are also of early
Early Cambrian age. However, until more
material is available a more precise age cannot
be given, The presence of further trilobite
fragments at a depth of 333 m in Marla 1B
indicate that the fauna figured herein cannot
be of earliest Early Cambrian age.

Acknowledgments
B. C. Youngs publishes with the permission
of the Director-General, S.A. Department of
Mines and Energy. Dr B. Daily (University of

field from which it is separated by a narrow, Adelaide) is thanked for useful advice and
shallow marginal furrow. The preocular sec- criticism.
References

Cuana, W. T. (1966) On the classification of Red-
lichiacea, with description of new families and
new genera. Acta Palaeont. Sinica 14, 135-184
[Chinese].

CourtTEssoLe, R. & Jaco, J. B. (1980) Biostrati-
graphie du Cambrien inférieur du Cabardes
(versant sud de la Montagne Noire, France
méridionale. Geobios 13 (in press).

Daity, B. (1972) The base of the Cambrian and
the first Cambrian faunas. Univ. Adelaide
Centre for Precambrian Res. Spec. Pap. 1, 13-
42.

GateHouse, C. (1976) A fossil in the Observatory
Hill Beds, South Australia. Quart. geol. Notes,
Geol. Surv. S. Aust. 60, 5-8.

Hupé, P. (1953) Contribution a l'étude du Cam-
brien inferieur et du Précambrien III de l’Anti-
Atlas marocain. Notes Mém. Serv. Geol. Maroc,
103, 402 pp.

Lt, S. J. (1980) Trilobites from the Chiulaotung
Formation (Lower Cambrian) in Emei Area,
Western Sichuan. Acta Palaeont. Sinica 19, 42-
50 [Chinese].

Lu, Y. H., Cuu, C. L., Curren, Y. Y., Lin, H. L.,
Cuow, T. Y. & Yuan, K. S. (1974) Bio-environ-
mental control hypothesis and its application to
the Cambrian biostratigraphy and palaeozoogeo-
graphy. Nanking Inst. Geol. Palaeontol. Mem.
5, 27-110 [Chinese].

Pitt, G. M., Bensow, M. C. & Younes, B. C.
(1980) A review of recent geological work in
the Officer Basin, South Australia. J. Aust.
Petrol. Explor. Assoc. 20, 209-220.

THoRNTON, R. C. N. (1978) The geological results
of the drilling of Manya No. 1 and Marla No. 1.
Miner, Resour. Rey. S. Aust. 143, 47-65.

Wuite, A. H, & Younas, B. C. (1980) Cambrian
alkali playa lacustrine sequence in the north-
eastern Officer Basin, South Australia. J. Sed.
Petrol. (in press).

MUCKERA AND MILLBILLILLIE —- AUSTRALIAN ACHONDRITIC
METEORITES

BY M. J. FITZGERALD

Summary

The chemistry, mineralogy and petrographic features of two Australian achondritic meteorites are
described in detail. The previously undescribed South Australian meteorite, Muckera, found on the
Nullabor Plain in 1951, is classified as a howardite. One of the masses of the Millbillillie meteorite
from central Western Australia is shown to be chemically and mineralogically similar to another
mass presumably from the same fall. This new mass is classified as an eucrite.

MUCKERA AND MILLBILLILLIE—AUSTRALIAN ACHONDRITIC
METEORITES

by M. J. Firzcnracn*

Summary

Erezoeraco, M. J, (1980) Mueckera and Millbillitiie—Australian achondritie meteorites. Trams.
R. Sac. S. Aust, 04(6), 201-209, 28 November, 1980,

The chemistry, minernlogy and petrographic features of two Australian achondritic
meteorites are described in detail, The previously undescribed South Australian meteorite.
Muckera, found on the Nullarbor Plain in 1951, is Classified as a howardite. One of the masses
of the Millbillillie meteorile from central Western Australia is shown to be chemically and
mineralogically similar to another mass presumably from the same fall. This new mass i¥

classified 65 an euerite.

Introduction

Meteorites can readily he subdivided into
two groups on the basis of bulk chemistry,
with the individuals of one group having
compositions resembling the sun. These undif-
ferentiated meteorites are known as chandrites
as most contain chondrules—small spheroidal
bodies commonly having diameters of the
order of one millimetre which typically contain
varying amounts of erystalline material in
dendritic or skeletal form, and glass, which
may Or may not be devitrified. Of the three
major chondrite groups — carbonaceous,
enstatite and ordinary—the carbonaceous
chondrites. best approximate pristine solar
material even though they have undergone
some alteration, albeit isochemical (McSween
1979).

Conversely, an achandrite is a meteorite
whose chemisity is unlike that of the sun.
Traditionally this term has been restricted to
differentiated meteorites containing little or to
free metal, but this restriction appears ta
conceal important genetic relationships,

The differentiated and undifferentiated
meteorites can be distinguished on the basis of
their chemistries relative to the Cl (type i
carbonaceous) chondrites by cvaluating the
following ralios (atomic abundances nor
malized to 100 silican atoms):

K/0.42, Al/8.5, Ca/ 7.21, P/0.96, Ti/0,2775

and Fe/83.0

with the divisors being the corresponding
normalized abundances for the C1 chondrites
from Cameron (1973), An undifferentiated
meteorite is then ucfined as one for which
these ratios generally fall in the range 0.4 to
1.3 while the achondrites fall outside,

Centre for Precambrian Research, University of
Adehode, G.P.O, Box 498, Adelaide S007.

The achondrites can be subdivided into nine
eroups on the basis of selected atomic ratios
(on a percentage basis) with almost every
member of two of the groups, the eucrites and
howardites, being characterized as follows:

Eucrites Howardites
AV/Si S10
Fe/5i <= 100
Ca/Si > 20 <20

These two groups encompass a large pro-
portion of the silicate-rich achondntes and
because of their chemistry and mineralogy
are often collectively referred to as the basaltic
achondrites, They consist essentially of plagio-
clase and pyroxene with pigeouite generally
predominating in the euerites and hypersthene
in the howardites. This predominance of
calcium-rich pyroxene im the eucrites is
reflected in the Ca/Si ratio used to charac-
terize them,

Most of the ecuerites and howardites are
breceiated with the former being deseribed as
manomict breccias (tc. they contain clasts
Which are similar in composition), although
some of these meteorites show little evidence
of brecciation, On the other hand = the
howardiles, almost without exception, contain
clasts which vary widely in texture and com-
position but which nonetheless appear to be
closely related to cach other and are thus
deserlbed as polymict breccias (cf, Duke &
Silver 1967, Wasson 1974, Mason er al. 1979).
The basaltic achondrites resemble breceiated
materials recovered from the lunar surface
and it is likely that all are surface breccias
produced by similar processes.

These chemical and petrological definitions
are generally complementary and in most cases
produce a consistent classification, However,
there seems to be a genetic relationship

202

between the two proups which renders the
distinction between them somewhat unneces-
sary, Furthermore, this relationship extends
to other achondrite groups as presently defined
and it ts suggested that they should all be
grouped together under the generie heading
of “cognate achondrites”.

Muckera Meteorite

Mistavical Background

On either May 1 or 2, 1951, while walking
some 3 km SW of a group of rock holes on
the northern edge of the Nullabor Plain about
70 km north of Cook and known as Muckers
Rock Hole (30°05'S 130°02’E), Police Con-
stable C, P, Doyle kicked a rock and. as it
seemed different from the country rock, picked
it up. Mr Doyle removed much of the cover-
ing crust in an attempt to identify the abnor-
mally heavy rock but was not successful and
so took it back to Cook. Ultimately the
meteorite Was brought to the Geology Depart-
ment of the University of Adelaide in Feh-
ruary 1972 where it was identified as a
meteorite hy Dr J. B, Jones, The meteorite
was named Muckera. This hame is sufficiently
different from Mukerop, a rarely used synouym
for the Giheon meteorite, to avoid confusion,

This is the first published account of the
Muckera meteorite other than the author's
classification given by Hutchinson er al.
(1977) and mentioned by Mason et al.
(1979).

Deveription

The meteorite is approximately ellipsoidal
in shape (Fig. |), and about 10 * 8 ® 5 em.
Part of the fusion crust which is commanly
red-brown in colour due to terrestrial weather-
ing is still preserved. This crust also contains
occasional white mineral grains.

Internally the meteorite consists of sub-
rounded to angular fragments, varying 01-3
cm. set in a dark grey to black eroundmass,
Small grains of fickel-iron and troilite, sur-
rounded in mast cases hy weathering haloes,
ure present. Commonly the nickel-iron is
confined to the clasts, while the dark colour
of the matrix is due to the presence of fine-
grained tarnished sulfides and olass. Oliyine
and calcium-rich pyroxenes are generally con-
fined) to the dark matrix whereas most of the

M. §, FITZGERALD

calcium-poor pyroxencs are in the clasts.
Numerous fractures and vughs, many of which
are lined with, and commonly filled by, a
brown mineral, ace present. In same cases
this filling is layered parallel to the cavity
walls.

Many of the silicate clasts are white to
cream. others are colourless. Many are aggre-
gates of smaller, sub-rounded clasts which in
turn are composed of angular and sub-rounded
mineral fragments (Fig, 2),

Petrographie examination emphasises. the
brecciated nature of the meteorite (Fig, 3).
The grotindmass, containing abundant glass,
is dark grey to black while the fragments are
eammonly light grey. Several different types of
clast are present, ranging from mimeral frag-
ments through monomineralic aggregates ta
polymineralic lithic clasts,

The monominerali¢c fragments include both
angular pyroxene and Jarge plagioclase prains,
many of which are strongly shocked and
braken but not disaggregated. The rmono-
mineralic aggregates include fragments. which
in terrestrial racks would be referred to as
orthopyroxeniles and anorthosites,

The lithic fragments include rock types such
as hasalts as well as numerous fine, equi-
granular breceia clasts containing angular
to sub-rounded grains, Pyroxene is more
abundant than feldspar in these pre-existing
breccias anda plassy mesostysis is commonly
present. There is generally littie miatris
material separatiny the small clasts whiely have
been Incorporated into the larger ones.

Muekera is thus a palymict breecia with a
clast to matrix ratio of about S:1, the latter
being composed nainly of glass and small
grains exhibiting a wide variation in grain size
and composition. Such textures can be readily
imagined as having been produced by repeated
impacts on the surface layers of the meteorite
parent body so producing fragmental material
which was subsequently compacted and
lithitied.

Milthillitlie Meteorite
Historical Background
At about | p.m, one day in October 1960,

Messrs F. Quadrie and F. Vicenti, two stution
hands working near the boundary fence

Om. re 8

Fig. |, External view of Muckera meteorite: Absence of fusion crust reveals Ir@mental nitive of

meleoriie, Scale 2 cm,

Fig. 2, Polished slab of Muckera meteorite showing brecciated texture. Scale | em
Fiy. 3. Thin section of Muckera meteorile showine diverse nature of clasts, Seale 2 mr.

ILLIE METEORITES 203

-RA AND MILLBILL

CKEI

MU

204 M. J.

Fig. 4. External surface of Millbillillie No. 3
meteorite showing fusion crust and radiating
bubble trains, Scale 2 cm.

between Millbillillie and Jundee Stations in
the Wiluna district, central W.A. (26°27’S
120°22’E), saw a fiery object trailing sparks
fall on a spinifex-covered area of the plain
to the north. No search was made but Mr D.
Vicenti found a piece of meteorite on the
plain in 1970. This mass weighed about 20 kg;
another mass of some 600g was found the
next year by Mr M., Finch (Clarke 1972).

Further material has been recovered in the
area by local aborigines. The third recorded
recovery was purchased by the late Mr Clem
Latz of Adelaide thereby precluding its export
from this country. This 368g individual was
completely covered with a fusion crust when
purchased, and from it a mass of some 26g was
acquired by the University of Adelaide. A
portion of this mass was used for the analyses
described in this paper. Subsequently, the
main mass of 233g was also acquired by the
University. The designation No. 3 was
appended to this mass to indicate that it is
the third documented recovery from the area
(cf. Graham 1980).

Oher specimens of the meteorite include
those lodged in Museums in Perth, Sydney,
Canberra and London and in private collec-

FITZGERALD

tions in Wiluna. All told it appears that at
least 26 kg of material has been recovered.

Some results have been reported for the
first mass found, Passing references to its
petrography were made by McCall (1973)
and several trace elements have been
determined (De Laeter & Hosie 1978; McCul-
loch et al. 1976, 1977; Rosman & De Laeter
1978; and Smith et al. 1977). At least two
bulk analyses have been carried out—one by
Dr J. A. Nelen (Mason et al. 1979), and
another by Dr R. A. Binns (Stolper 1977)
which is yet to be published in full, This
paper presents the first published results for
the third mass.

Description

The No. 3 mass resembles a flattened
ellipsoid with dimensions about 4 X 7 X 10
cm (Fig. 4). It is completely encrusted with
a black glassy layer about 0.3 mm_ thick
underlain in turn by dark crystalline material
and a further layer of a light coloured crystal-
line material. Much of the crust, which is
patterned with numerous lines radiating from
a central point on one of the flat surfaces,
is now covered with an orange film. The sur-
face lines consist of trains of numerous dark
brown to black glassy bubbles about 0.5 mm
in diameter; other bubbles are randomly
distributed over the remainder of the crust.
Furthermore, the crust exhibits a series of
polygonal fractures commonly forming quad-
rilaterals with dimensions about 2-5 mm. The
individual cracks are less than 0.1 mm wide
and are filled with the orange material which
covers part of the crust surface. In places
this filling stands above the level of the
surface and in other cases below. In general,
the fractures do not extend through the
fusion crust. However, in rare examples where
the fractures do pass through the crust, the
secondary orange mineral occurs along the
interface between the crust and the interior
mass of the meteorite. In some places the
orange mineral is, in its turn, overgrown with
an apple-green one.

A cut surface shows numerous sub-angular
particles from 0.1—3 cm set in a grey to white
crystalline matrix, A sub-ophitic texture can
be seen in some of the coarser-grained clasts
even at low magnification. Clast-rich and clast-
poor regions define a layering. Finer-grained
clasts generally possess very sharp boundaries
whereas the coarser ones commonly merge into
the matrix, the grain size of which increases
in that area, resulting in the average grain size

MUCKERA AND MILLBILLILLIE METEORITES

of the clast-rich regions being greater than
that of the clast-poor regions. No metal or
sulfide grains were seen on the surfaces
examined,

The two clast types are readily seen in thin
section (Fig. 5). The finer consist of extremely
fine-grained granoblastic aggregates contain-
ing many opaque grains while the coarser
clasts are ophitic and subophitic basalts, that
is, feldspar and pyroxene aggregates. The feld-
spar laths in these basaltic areas vary in
length up to 0.5 mm while occasional equi-
dimensional grains achieve similar dimensions.
In addition to these clasts there are also
numerous feldspar fragments and spherulitic
aggregates of devitrified glass. The silicate
grains contain numerous opaque inclusions.
The merging of the coarse-grained clasts and
matrix is even more obvious in thin section
and some of the matrix is also basaltic in

205

texture. The layering seen on the macro scale
can also be seen in thin section. There is no
evidence of brecciation either within the
basaltic clasts or in the host matrix,

The matrix consists of tiny (0.02—0.03 mm)
grains of pyroxene and feldspar set in glass
with an overall texture of “feathery” quench
material. Much of the matrix has been devitri-
fied and recrystallized. There is no evidence of
weathering in the form of iron-staining but
the matrix appears dark due to the presence
of numerous grains of opaque minerals and
regions of opaque glass.

It is concluded that the fine-grained clasts
may have been incorporated into the matrix
prior to crystallization. There are several
ways of achieving this—one possibility is that
pre-existing clasts (possibly impact-derived
fragments of a rapidly cooled lava flow) were
caught up in a later flow which _ initially
cooled less rapidly perhaps as a more extensive
lava flow or blanket. During this cooling stage,
gravitational setting could have brought about
the layering. Subsequent rapid crystallization
arising from changes in the cooling regime or
the attainment of conditions of supersatura-
tion in the liquid phase could have produced
the glass in the matrix. Regardless of the exact
mode of formation of the texture, subsequent
shock events must have been minimal as any
such — significant reworking would have
destroyed this layering.

Bulk chemical compositions

X-ray fluorescence analysis was used for
the determination of all elements except
sodium for which a flame photometric
method was employed. A modified version of
the Norrish & Hutton (1969) technique for
XRF analysis was used. Mineral compositions
were determined with a Technisch Physiche
Dienst microprobe fitted with a lithium drifted
silicon detector, the method of Reed & Ware
(1975) being used to reduce the data, Full
details of all methods are given in Fitzgerald
(1979).

The bulk chemical compositions of the
meteorites are listed in Table 1 along with the
results of normative mineral calculations. The
assumptions made in the modified form of
the CIPW calculations used to calculate the
norm are fully explained in Fitzgerald (1979).
Phosphorus has been allocated to the mineral
merrillite, (@-Ca,(PO).), as Dowty (1977)
has shown this is the principal meteoritic
phosphate. For comparative purposes, Table 2
also lists the results of Mason ef al. (1979).

206
TABLE |, Buelk chemical composition and narmative
minerqlogy.
Elemental
abundances
(weight Millbillillie Millbillillie
percent) Muckera! No. 3! No, 1
Fe 13.85 15,30 14.22
Mn 0.40 0.46 O47
Ti 0.25 0.44 (4)
Ca 5.06 7.42 7.29
K 0.040 0.071
P 0.029 O.034
Si 23.20 23,84 24,3]
Al 4,32 7,03 6.77
Mg 8.95 4.42 4.05
Ni 0.090 0.018
S 0.27 0,003
Cy 0.37 (21 0,23
Na (2) (33 0.29
Normative
mineralogy
(weight percent)
Nickel-iron ().7 0.4
Troilite 0,7 0.01
Merrillite 0.1 0,2
[lmennte 0.8 3
Chromite O# O4
Feldspar 23.9 34.3
Diopside 1S 13.0
Orthopyroxene 67.0) 41.9
Olivine as
Quartz 8.9
Molar percent
composition
Ah 11 It
An 88 88
Ps 35 40
Fa 3
tT his work *Masan etal. (1979)

As Mason (1962) pointed out, there is an
almost perfect balance between silica and the
basic oxides in the basalGe achondrites so
that a slight excess results in quartz being
present or a slight deficit appears as olivine.
The excess silica generally occurs in the
eucrites and the olivine in the howardites, The
normative caleulations of Table 1 jlustrate this.

Mineralogy

Eucrite and howardite fteldspars commonly
fall in the compositional range Any, o9;. with
most compositions lying in the narrower range
Angs-an (Duke & Silver 1967). Feldspar
compositions measured in this work (Table 2)
lic within these limits and are accordingly
consistent With the classifications proposed
below

M. J. FITZGERALD

The pyroxene Variation in Muckera (Fig, 6)
is very similar to that of the Kapoeta and
Malvern howardites (Desnoyers & Jerome
1977; Duke & Silver 1967; Dymek ef al.
1976; Simpson 1975), The majority of the
analyses fall within the field En, Fs. ,-Eny ,,
sz y—Eny 4 F854, Woo Eng yFsyq Won, This
Variation is in distinct contrast to that of the
eucriles.

In these cases Duke & Silver (1967) found
a continuous range in composition from
pigeonite through sub-calcic ferroaugite to
ferroaugite with the magnesium silicate com-
ponent remainifig relatively constant, Figure
7 shows such a situation for Millbillillie and,
exeept that the most calcium-rich analyses
show about 5 mole percent more of the
calcium silicate component than the results of
Duke & Silver, the distribution most closely
resembles that of Juvinas as figured by these
authors. A more fecent determination
(Takeda et al. 1978) shows an even closer
resemblance between Juvinas and Millbillillic.
The Millbilllilie results are also yery similar
fo those obtained by Desnoyers & Jerome
(1977) for a clast in the Malvern howardite.
Simpson (1975) and also Wilkening & Anders
(1975) have suggested that this apparent
linear series of compositions, which is charac-
teristic of eucritic pyroxenes, results from the
analysis of differing proportions of two
exsolyed phases, the lamellae being too fine to
resolve with the microprobe. Mason ef al,
(1979) came to a similar conclusion, Micron
sized lamellae visible in many of the Millbil-
lillie pyroxene grains are probably thus
responsible for the trend seen in Figure ‘7.

Modal olivine was not observed in Millbil-
lillie und miner amounts only in Muekera, In
the latter it was confined almost exclusively
to the dark groundmass with only uo few
prains being found in lighter clasts, The com-

TABLE 2. Mineralogical comiposilions,

ML i ay

Muckeru! No, 3! No
Feldspar
(% An)
Runge T4 lo 04 78 lo 2 &2to4l
Average 88 84 Ra
Pyroxene —_— Fisuo Wax Fsiy Won
lo, to
Fsoy Woy, san Woy

‘This work *Mason ef al, (1979)

MUCKERA AND MILLBILLILLIE METFORITES 207

rad —

Shs

Fig, 6. Pyroxene compositions in Muckera meteor-
ite,

\

f m An
“ \
neal a. .

Fig. 7. Pyroxene compositions in’ Millbillillie No.
3 meteorite.

positional range Fasy 95, as measured by
microprobe, is consistent with the results of
Desnoyers & Jerome (1973). These modal
results are in accord with the normative calcu-
lations of Table 1.

The iron-manganese correlation in lunar,
meteoritic and terrestial olivines and pyroxenes
is well documented with the three groups
being fairly readily distinguished on this basis
(Desnoyers & Jerome 1973; Dymek et al.
1976; Simkin & Smith 1970). The correlation
arises as a result of the ability of Mn*+ to
replace Fe+* in olivine and pyroxene lattices
due to the similarity of their ionic radii
(Wanke et a/, 1973). Intergrain constancy of
this ratio indicates the grains have been
derived from related source materials (Dymek
et al. 1976).

The Muckera pyroxenes and olivines show a
wide range in iron and manganese contents.
That the pyroxene data is somewhat non-
cohesive is shown by the correlation coefficient
of 0.91 for 118 determinations, The average
Fe/Mn ratio for the pyroxenes is 28.8, while
the value for the bulk analysis is 33.9 (Fig. 8).

A relatively narrow range in iron composi-
tion in the olivine, coupled with a considerable
spread in manganese values, results in the
relationship between these elements in the
olivine grains being less obvious (Fig. 8), with
the spread being indicated by the lower cor-
relation coefficient of 0.34 for the 50 analyses,
The average value of the ratio for the olivine

is 50.3, thereby explaining the difference
between the bulk ratio and that of the
pyroxenes.

In Millbillillie the pyroxenes also exhibit
a wide range in Fe and Mn values but the
data is even more cohesive as shown by the
correlation coefficient of 0.97 for the 103
analyses (Fig. 9). The value of the ratio for
the bulk analysis ts 33.1 while the average
value for the pyroxene analyses is 32.9. Simi-
lar agreement of pyroxene and bulk chemistry
is shown by eucrites such as Moama (Lover-
ing 1975) which has a bulk ratio of 29,3 and
one of 27.5 for the host hypersthene.

Classification

Table 3, giving values for several atomic
ratios discussed previously, clearly shows that
both Muckera and Millbillillie are differen-
tiated meteorites while Table 4 shows that,
on the basis of the ratios listed here, Muckera
is a howardite and Millbillillie an eucrite,

Feldspar, pyroxene and olivine contents and
compositions are consistent with these classi-
fications as are the petrological descriptions
showing Muckera to be a polymict breccia
and Millbillillie monomict. Muckera is thus

L 7 —
= 0
reo

Fig. 8. Plot of MnO versus FeO in Muckera
meteorite olivines und pyroxenes. X: bulk com-
position,

7] red 3

Fig. 9, Plot of MnO versus FeO in Millbillillie No.
3 meteorite pyroxenes. X: bulk composition.

208

Tasre 3. Evaluation of “differentiation” ratios on
atomic hasis.

Ti’
K/0.42 AWS Cal7-2b P/0.96 0.2775 Fe/83 0

RA eT oil 2 D4

Millbil-
lillie No.3 05 36 3.0 Of 39 D4

TABLE 4. Classifications based on atomic ratios
(an a percentage hasis),
_ Ratios Classilica-
AL/Si Fe/Si- Cu/Si tion
Muckera 19.4 30,2 15.3 Howardite
Millbillillic
No. 3 30.6 32.4 21.8 Fuerite

the first recorded example of a polymict brec-
ciated achondritic meteorite to be recovered
in Australia,

M. J. FIIZGERALD

On the basis of chemical and textural fea-
tures it has been suggested (c.g. Stolper 1977)
that most of the known eucrites lie together
in a cohesive group with the others in iwo
smaller groups with one containing eucrites
deseribed ay cumulates and the other meteorites
considered to be residual fractions. Applica-
hon of the technique of Principal Component
Analysis by the author (Fitzgerald 1979)
to a data set including analyses of 32 eucrites
has shown that Millbillillig belonys to the
main body of the cucrites, an assignment
Which is supported by the textural relation-
ships described above.

Acknowledgments

This work was carried out during the tenure
of a University of Adelaide Postgraduate
Research Grant and the receipt of both this
and the constructive help wad criticism of Dr
J. B. Jones and an anonymous referee is
gratefully acknowledged,

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